Dodota JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALBION REED HODGDON, Editor-in-Chief ALBERT FREDERICK HILL STUART KIMBALL HARRIS RALPH CARLETON BEAN ROBERT CRICHTON FOSTER Associate Editors ROLLA MILTON TRYON RADCLIFFE BARNES PIKE LORIN IVES NEVLING, JR. VOLUME 68 1966 The Nem England Botanical Club, Ine. Botanical Museum, Oxford St., Cambridge 38, Mass. FARLOW REFERENCE LIBRARY APR cc Arn å JOO Todora JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALBION REED HODGDON, Editor-in-Chief ALBERT FREDERICK HILL y STUART KIMBALL HARRIS RALPH CARLETON BEAN ROBERT CRICHTON FOSTER \ Associate Editors ROLLA MILTON TRYON RADCLIFFE BARNES PIKE LORIN IVES NEVLING, JR. Vol: 68 January-March, 1966 No. 773 CONTENTS: A Multivariate Analysis of the Pinus chiapensis-monticola- strobus Phylad Jobn W. Andresen ............ een 1 Cytogeography of Orontium aquaticum (araceae) John W. Gream, JT. wcccccooccssscrsscsssersssssssssscscssccsssccscssesessacesesssenes 25 Authors of Plant Genera and the International Plant Index (Review) Reed C. Rollins ................ esee 35 Studies in the Capparidaceae VIII. Polanisia dodecandra CL.) DC.: Hugh H. Iltis ................. eren 41 Polanisia dodecandra in New Hampshire Edward J. Hehe ..........soososssoseocsoosescscecssussovossosossoscoensocscososcoseesos 48 | (Continued on Inside Cover) Cee tr P 1X0. The Nem England Botanical Club, iuc. Botanical Museum, Oxford St., Cambridge 38, Mass. ^. > " S ~ D CONTENTS: — continued Chromosome Numbers in Vernonia (Compositae) S. B. Jones, Jr. and W. H. Duncan ees 49 Natural Hybrids of Lady's Slippers (Cypripedium) in Manitoba H. H. Marshall, A. T. H. Gross and G, A. Stevenson ........ 58 A Study of Variation in the Cytotypes of Dryopteris spinulosa Rolla Tryon and Donald M. Britton ess 59 Varieties of Ballota nigra in the Eastern United States C. V. Morton esset 93 Paspalum minus (Gramineae) in Louisiana and Mississippi Donald J. Bankes ..sss e 94 Studies in the Flora of Bolivia, — IV. Gramineae Robert C. Foster sss 97 Errata lrer ett dl 120 — RHODORA.—A quarterly journal of botany, devoted primarily to the flora of North America and floristically related areas. Price, $6.00 per year, net, postpaid. in funds payable at par in United States currency in Boston; single copies (if available) $1.80. Back vol- umes 1-58, with a few incomplete, can be supplied at $5.00 per volume, Volume 59— available at $6.00. Somewhat reduced rates for complete sets can be obtained upon application. Scientific papers and notes. relating directly or indirectly to the plants of North America, will be considered by the editorial com- mittee for publication. Articles concerned with systematic botany and cytotaxonomy in their broader implications are equally accept- able. All manuscripts should be double-spaced throughout, Please conform to the style of recent issues of the journal. Illustrations can be used only if the cost of engraver's blocks is met through the author or his institution. Forms may be closed five weeks in advance of publication. Extracted reprints, if ordered in advance, will be furnished at cost. Address manuscripts and proofs to Albion R. Hodgdon, Dept. of Botany, Nesmith Hall, University of New Hampshire, Durham, New Hampshire. Subscriptions and orders for back issues (making all remittances payable to RHODORA) should be sent to Albert F. Hill, Botanical Museum, Oxford Street, Cambridge 38, Mass. Second Class Postage Paid at Boston, Mass. Manufactured by THE LEXINGTON PRESS, INC. Lexington, Mass. QTRbooora JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Vol. 68 January-March, 1966 No. 773 A MULTIVARIATE ANALYSIS OF THE PINUS CHIAPENSIS-MONTICOLA-STROBUS PHYLAD* JOHN W. ANDRESEN As considered here, Pinus chiapensis (= Pinus strobus var. chiapensis Martinez) is the southern member of the North American pine phylad Pinus chiapensis-monticola- strobus. Phytogeographically, this complex forms a broad discontinuous triangle with apices of species distribution in British Columbia, Newfoundland, and Guatemala (Fig. 1). Earlier, the taxonomic recognition of Pinus strobus var. chiapensis Martinez was questioned by several authors (Braun 1950: 483; Martin and Harrell 1957; Sharp 1946) who regarded it as a weakly-differentiated geographic form of P. strobus L. undeserving separate taxonomic rank, but others (Loock 1950: 117-119; Soto, Barrett, and Little 1962: 52-53; Standley and Steyermark 1958: 55-56) agreed with its varietal status. Gaussen (1960: 91, 198) proposed it as a distinct species but his new combination was not validly published because he omitted citation of the basionym (Art. 33, LC.B.N., Lanjouw, et al. 1961). These divergent opinions prompted the present biometric study to provide a statistical basis for a determination of the proper taxonomic disposition of this controversial taxon. In a companion paper (Andresen 1964), it is recommended that P. strobus var. chiapensis be elevated from varietal to This study was supported in part by funds from National Science Foundation Grant No. G-15879. bo Rhodora [Vol. 68 Pinus chiapensis è ipinus monticola ERE 4 — ' Pinus strobus NN m" SCALE Fig. 1.— Distribution of the phylad Pinus chiapensis-monticola- strobus in Central and North America. Adapted from unpublished map supplied by U. S. Forest Service. Plotted on Goode's Base Map No. 202 by permission of the University of Chicago Press, specific rank with the binomial Pinus chiapensis (Martinez) Andresen. The proposal was predicated on evidence ob- tained from the following study of morphologic and progeny data that were not available to previous authors. Following the analysis of this data, it became apparent that there was a much wider genetic and morphologic divergence than was previously suggested between P. strobus var. chiapensis and typical P. strobus. Recent conceptual advances in the field of biosystematies (Heslop-Harrison 1963, Heywood and Love, 1963), that presented a clearer perspective of the problem at hand, led 1966] Pinus — Andresen 3 Rogers (1963), and Sokal and Sneath (1963) to stress the value of reliable statistics that incorporate fundamental quantitative characters to determine taxa affinities or dif- ferentials. They coined the terms “taximetrics” and “nu- merical taxonomy” respectively, to focus attention on the statistical techniques designed to solve problems associated with experimental taxonomy. In this respect it is of pri- mary importance to the taxonomist that contemporary statistical analyses and those yet to be derived can be pro- grammed, especially with the aid of electronic computers, to assess the taxonomic value of any taxon trait or group of characters. As an example, Davidson (1963) developed an “Itemized frequency distribution" and set of association indices based on the F statistic, to discriminate Cirsium altissimum from C. discolor. In a study of variability of the Pacific Coast and Rocky Mountain forms of Pinus contorta, Jeffers and Black (1963) used three forms of multivariate analyses which incorporated the “Q-technique,” discriminant anal- ysis, and component analysis, all of which rely on various manipulations of the correlation coefficient and its components. In this study, to show the degree of discrimination within the Pinus chiapensis-P. monti- cola-P. strobus complex, I employed the coefficient of divergence (C.D.) technique (Clark 1952) which is based on the sum of mean differentials of multivariate characters. To evaluate and assign relative importance values to the 12 leaf and cone characters used in the C.D. analysis, the F Statistic, intraclass correlation (expressed as a reliability index, R,), and coefficient of variation were also used. Data for the multivariate character analysis of cones, leaves, seed, and seedlings were obtained from herbarium and living specimens. Sample sizes, characters examined, statistical techniques, and pertinent examples to illustrate a particular method used are included as footnotes to data tables or in the discussion of the characters. 4 Rhodora [Vol. 68 In the following text, Pinus chiapensis is coded as A; P. monticola Dougl. as B; and P. strobus as C, except where the Latin binomials are more useful for clarity or emphasis. COEFFICIENT OF DIVERGENCE VALUES Before the importance of individual characters and their component roles in species delimitation within this phylad are evaluated, an explanation of the simultaneous treatment of fundamental characters is in order. A useful measure that incorporates this technique is the coefficient of divergence which is used in association with F statistics and reliability indiees to summarize leaf and cone characters Q, to Q, species pairs AB, AC, and BC. The method (Clark 1952, Klauber 1940) of arranging or ranking taxa in relation to the ascending magnitude of their C.D.’s is especially ger- mane to studies utilizing objectively scored traits. Although Clark (1952) referred to herpetological data in his example, the extension of Klauber's (1940) one-dimensional coeffi- cient by Clark is adaptable to any numerical or coded array of biologic parameters computed as a series of multiple character means. In the present example (Table 1) four results are ap- parent: (1) About the same degree of information was provided by either 39 random samples or the total number of herbarium specimens examined (see Tables 1 and 2 to compare means of characters Q, to Q,. for the two sample sizes) ; (2) The C.D.’s derived from cone characters alone were greater in pairs AB and AC: (3) Leaf characters were more important than cone traits in computing the C.D. for BC; (4) With the incorporation of all 12 characters, BC is less than one-half as divergent as either AP or AC. This magnitude of divergence, or lack of it, is of special significance — for according to the C.D. ranking, two long- recognized species, P. monticola and P. strobus, are much more closely aligned to one another than the species P. strobus is to its supposed variety chiapensis. This evidence, primarily the strength of the following F and R, statistics 1966] Pinus — Andresen 5 for the leaf and cone characteristics, plus other pertinent data, motivated the author to elevate P. strobus var. chi- apensis to specific level (Andresen 1964). LEAVES Leaf length (Q,) was greatest for A, but leaf width (Q;) was least (Table 2). Thus, the resultant W/L value (Qs) for A was smallest. Conversely, the shorter but wider leaves of B had a high W/L ratio that was twice as great as the value for A. An intermediate value of Q; for C is a reflec- tion of the interjacent values of Q, and Q. The interpreta- tion of this value, however, should be tempered by the lack of significance in the F statistic and the low reliability index of .497 for Q, in species pair AC; thus the component of greater relative importance for Q, in C is leaf length. The role of leaf length in the ratio is emphasized because numerous authors (Loock 1950: 117; Martinez 1948: 133; Schwerdtfeger 1953; Sharp 1946; Soto, Vazquez, and Little 1962: 53) describe the leaves of P. chiapensis as “more delicate," “more slender," "finer," or “thinner” than P. strobus. These comparatives are misleading, however, when Q, and Q, are examined objectively. As shown earlier, there is a highly significant difference between A and C in regard to length but the width differential is non-significant. With these and other facts in mind, the following thoughts are offered to attempt an explanation of the reasoning of previous observers. In the field, the illusion of angustifoli- ation is probably created when observers familiar with leaf length in P. strobus subconsciously compare the much longer (and seemingly thinner) leaves of P. chiapensis and infer that the leaves of A are more delicate. But more important, because most descriptions are based on dry material, the gracile appearance of desiccated specimens is related to the herbarium artifact caused by severe de- hydration and subsequent collapse of the leaf mesophyll. This condition was suggested when dried leaf specimens of A were noted to be deeply concave on all three surfaces (turgid leaves bear slightly convex surfaces), were curled 6 Rhodora [Vol. 68 or twisted, and seemed lighter in weight than dried leaves of either B or C. To advance this postulation I (1) dried 125 leaves of A, B, C, and B X C grown under similar conditions near Placerville, California, and (2) calculated the proportionate weight loss of each set of samples by comparing fresh weight (weighed immediately after col- lection) to oven-dry weight. The differential percentages (Table 4) illustrate a much greater weight loss in A than the other three which were not significantly different from each other. Although tabulated at the 5% level of signifi- cance A was actually significant at the 2% level of con- fidence. The loss of 69% of the original weight in A is attributable mainly to the removal of fluids (mostly water) from the abundant intercellular space within the leaf meso- phyll. A comparison of intercellular spaces within the leaves of the four taxa (Table 4) illustrates the significant differential between A and the other three. Another feature which has previously been used to sep- arate A from C is the supposed higher number of foliar resin canals in the former (Gaussen 1960: 91; Loock 1950: 117, Martinez 1940, 1947: 134; Soto, Vasquez and Little 1962: 53). This group of authors contends that the more common number of resin canals in A is 3 and more often 2 in C. The analysis (Table 4) however, provides data to the contrary. Based on 500 observations, ¢ values at the 5% and 146 levels shows no significant differences between A and C. Mean values of 2.52 and 2.57 respectively, low standard deviations, and ranges of 2 to 3 resin canals for both species invalidate this character as a trait of differ- entiation. Although there is an overlap of range in number of resin canals between B and the former two, there is a significant difference between the means, so this feature could be used to separate B from A or C. Of interest is the broad range of numbers of resin canals in the hybrids of B and C and the large standard deviation, but this is an established pattern with numerous hybrid pines ( Keng and Little 1961). 1966] Pinus — Andresen zi Even though the original description (Martinez 1940) of A was predicated on the criteria of thinner leaves and a greater number of foliar resin canals than C, the data indicate that both characters are of little or no value in separating the two taxa. On the other hand, a character apparently overlooked in the past that has high diagnostic utility is the degree of leaf serration, Q, (Table 2). In fact, the F value for this character for species pairs AB or AC was the most sig- nifieant of any leaf parameter. Twice as many serrations per 5 mm interval were found on A when compared to C. The high reliability index of .995 for Q, of pair AC also demonstrates the dependability and high level of confi- dence to be expected when using this trait for species sep- aration. Also, a low coefficient of variability of 14.3 for A in contrast to 25.4 for C and 57.5 for B is added measure of reliability for this character in A. In a progeny study of juvenile forms of P. strobus, Mergen (1963) found an in- crease in serration number in relation to progression from southern to northern sources of origin. We found just the opposite trend in mature field specimens, and observed a random pattern in juvenile specimens. The weakest leaf character of the five examined (Table 2) was the number of stomatal rows borne on the ventral (adaxial) surface, Q.. Although there was a significant difference between A and C for Q., the reliability index was lowest in comparison to the other significant features of leaf characters. Stomata on the dorsal (abaxial) surface, although not included in the character analysis, were absent on all 440 leaves examined of A and all 3408 leaves exam- ined of C. Dorsal stomata were present, however, on B leaves with a mean of 0.28 and a range of 0 to 4 based on a 1279 leaf-count. When present, this is a reliable character to separate B from C (Harlow 1947, Sargent 1897: 23, Shaw 1914: 34). In two studies of pine leaves however, (Doi and Morikawa 1929, Sutherland 1934) reported an absence of dorsal stomata in P. 8 Rhodora [Vol. 68 Fig. 2. — Comparison of cone characteristics of Pinus chiapensis (A), P. monticola (B), and P. strobus (C). PLATE 1318 CONES Since the value of the mean for ovuliferous scale number (Q;) was highly dissimilar between the three species and C.V.'s were relatively small, large F statistics and strong reliability indices were derived. The highest number of scales were found on B (Fig. 2, Table 2) and even though there were about 25% more scales per cone in A than in C, cone lengths were not significantly different. Precise cone- phyllotaxis of the three species was very difficult to deter- mine in mature open cones, so this trait was ignored. 1966] Pinus — Andresen 9 Shaw (1914: 12) observed that this pattern of cone scale arrangement presented an indefinite phyllotaxy for the sub- genera Diploxylon and Haploxylon and the only importance of phyllotactic differential is that it separates the two sub-genera on the basis of a higher order-fraction in the former group. Nevertheless he commented that in cones of equal size, B had an obviously higher phyllotaxy than C. a phyllotaxy of A was also higher than C but lower than Apophysis width (Q;) in both A and C was similar, but length (Q.) was more divergent, (Table 2, Fig. 2). Even though Q; and Qs were dimensionally greater in B than in the other two taxa, the resultant ratio (Q,) of the three species was largest for A because of its higher propor- tionate width. A high F statistic and reliability index of .959 of Q, for AC indicate the potential diagnostic value of this trait. In pair BC, the ratio differential was non- significant with a low R,, and so was of little value in separating the cones of B and C. Length of peduncle (Qı) was greatest in A, but shorter and about the same for B or C. Although F and R, values of Q, were high for pairs AB and AC, the low values in association with BC indicate the similarity of this trait in B and C. Character Q.» is, at times, difficult to measure for the peduncle is brittle and tends to fracture easily. Caution must be exercised to carefully collect and store the cones to avoid negation of this character. Another reliable cone trait to isolate A from B or C is the degree of scale margin undulation (Q;, ). In A, the apo- physis margin with inflexed umbo-tip forms several involu- tions (Fig. 2) rarely found in B or C. As with peduncle length, scale margin bore differentially high statistics for pairs AB and AC, but was useless to segregate B from C in which the umbos were simply concave. Martinez (1940) and Soto, Barrett, and Little (1962: 53) described the umbo as having ". - - undulating edges turned inward.” but in their work this conspicuous feature was not com- 10 Rhodora [Vol. 68 Fig. 3. above — Basal scale configuration of Pinus chiapensis (A), P. monticola (B), and P. strobus (C). Fig. 4. below — Ovuliferous scales (adaxial surface) of Pinus chiapensis (A), P. monticola (B), and P. strobus (C). PLATE 1319 pared to other species. Also, the scale apex of A is truncate versus the rounded apices of P and C (Fig. 4). The most outstanding difference in cone morphology and the character that was related to the highest F and R, val- ues of all leaf and cone characters for pairs AB and AC was the relative number of basal reflexed scales (Qi.). This curving of scales contiguous to the peduncle of taxa A and B is obvious in numerous illustrations (Gaus- sen 1960: 199; Harlow and Harrar 1958: 57; Sargent 1897: plates 539, 541; Sudworth 1908: Fig. 2). In Fig. 3, 1966] Pinus — Andresen 11 a comparison of the number and presence of these scales in B and C is made with the lack of reflexing in A. In our samples of A, none of the cones bore basal scales that were fully reflexed, and only a few were weakly curled. Since F and R, values were high for all three pairs, an exam- ination of Q,. alone would suffice to determine individual members of the phylad. In addition to the foregoing values obtained from leaf and cone measurements, the data following the discussion of sample size also add confirming evidence that P. chia- pensis is strongly divergent from P. strobus. SIZE OF SAMPLE In biometric analyses, achievement of the highest degree of precision commensurate with judicious use of time is a goal of primary importance. In Table 1, it is demonstrated that approximately the same coefficient of diver- gence can be calculated by measuring relatively large num- bers of specimens or by a less time-consuming random sample of 39. The question then is how many samples are required to provide a desired level of accuracy in separat- ing these taxa? Numerous models and examples are offered by Cochran and Cox (1950), Dixon and Masey (1957), and Snedecor (1956), but I prefer the technique derived by Wright and Freeland (1960). Their method, in which it is advantageous to use small sample numbers, employs stand- ard deviations and means of the components of any species pair. When substituted in the formula in Table 3, these values multiplied by an appropriate “t” provide a predict- able sample size for any required level of accuracy. Since the minimum number of complete samples within A, B, or C was 39 and a recommended minimum sample (Freese 1959) should approximate 30, we used a set of 39 observations for the 12 characters to obtain the s, X, F, and R, data in Tables 2 and 3. Also, from past experience with the group Strobi, it seems that 30-40 random samples are required for an initial character survey. Estimates of sampling (Table 3) for further studies of characters Q, to Qi: for taxa pairs 12 Rhodora [Vol. 68 AB, AC and BC indicate that a relatively small number of specimens is required to exhibit statistical significance for most traits. A maximum of ten samples of each species within a pair would be required to detect a difference at the 5% level of confidence in 28 of the 36 possible character- taxa pair combinations and 20 samples to detect differences at the 1% level for 28 out of 36. These are reasonable sample sizes with which to work, and the number of char- acters, if definitive, could be expanded to give a more pre- cise evaluation of a total difference. Note the direct rela- tionship (Table 3) between the reliability index and the number of samples required. Low or negative R, values correspond to higher sample numbers. Large sample sizes, e.g. 1552 and 227 for BC-Q,, and Q,, would require a prohibitive amount of observation time, so these characters are best not measured.. The time saved could be better spent on other observations such as those which follow. SEED AND SEEDLING CHARACTERISTICS Differences in seed weight of the three species used in the progeny tests are given in Table 5. To determine the accurate weight of sound seed each lot was first floated in 95% ethyl alcohol to separate the denser, sound seed from the buoyant, partially filled seed. After winnowing, the seeds were floated and it was discovered that about 10% of the seed were blank. Based on this unit of malformed seed and experiences with the other group Strobi seed, we con- verted the reported weights (Anon 1948: 269) of P. monticola and P. strobus to arrive at 53,460 per kg which compared very closely to our data for the two species. The large number of 60,000 seed per kg for A is related to the smaller size of the individual seed. The smaller seed may be linked to the ability of propagules with reduced endo- sperm to survive in the favorable growing and germinating conditions within the natural range of A. The seed of all three species, in either the stratified or unstratified condition, began to germinate within 10 to 16 days after sowing (Table 5). In contrast to B and C, the 1966] Pinus — Andresen 13 germination period for A had ceased 37 days after initiation of the test— also, the germination values for either treatment were identical. On the other hand, un- stratified seed of B and C had considerably lower values than those stratified and some seed of both treatments had not germinated at the end of the test. The marked con- trast in germination values between A and B or C is related to a major physiological difference within the phylad ger- mination regime. Under uniform growing conditions in a greenhouse the seed sources of A produced seedlings with longer hypo- cotyl and cotyledon length. A was significantly larger than B and C, but B was smallest. Larger numbers of samples are required to determine the variability within B and C, but the 30 sources of A seemed adequate to assess some in- fraspecific variation. Cotyledon numbers were not signifi- cantly different from one another although C had a slightly higher range and mean. Englemann (1880) reported 6-9 for B and 7-11 for C. Of all the seedling characteristics examined in this study, cotyledon number was most uni- form and of least diagnostic value. Shaw (1914: 1) suggested that cotyledon morphology and number are unreliable for species identification unless there are distinct numerical differences between species. In a recent study of the Pinus flexilis - P. strobiformis com- plex (Andresen and Steinhoff 1965), a significant differ- ence was found in numbers of cotyledons with a mean of 8.8 for P. flexilis and a mean of 11.5 for P. strobiformis. In addition to the above seedling characteristics, there was a difference in growth of secondary leaves and bud set- ting. As might be expected, because of its southern origin (Mirov 1962), A produced secondary leaves much earlier than either B or C. No secondary leaves were formed on either B or C during the first 200 days of observation. Also, A did not produce any dormant buds, for as soon as a bud was formed it produced continuous growth of leaves. Definite dormant buds did form on B and C. In the course of my 14 Rhodora [Vol. 68 field work in December 1962, I observed numerous P. chia- pensis south of Sola de Vega, Oaxaca, Mexico, which had produced several increments of cones the past growing season and with a few trees undergoing anthesis. Mr. Boone Hallberg (personal correspondence) has noted this phenomenon for several years and reports that an average stand in Oaxaca has the following periodicity of cone matu- ration: Percent of Trees Date of Maturation 20 20 July + second crop about 20 November 10 1-20 August 60 25 August - 15 September 10 15 September - 10 October Mirov (1962) also observed that P. oocarpa in Nicaragua and other tropical pines are characterized by accelerated and uninterrupted growth rhythms. Only one year is re- quired for seed set after pollination. CONCLUSIONS Coefficient of divergence values indicate a much closer morphological affinity between Pinus strobus and P. monti- cola than between P. strobus and the former variety chiapen- sis. The wide divergence between P. chiapensis and P. strobus provides one form of biometric evidence to argue against the opinions of earlier writers advocating that the two taxa should be conspecific. With the available C.D. values and the sustantiating data that follow, little doubt re- mains that P. chiapensis is a distinct species. An analysis of the more statistically significant characters that discrimi- nate P. chiapensis from P. strobus reveals that P. chiapensis differs by having: (1) 25% more leaf serrations per unit length, (2) leaves that are 31% longer, (3) cone-scale apo- physes that are truncate and 20% shorter, (4) extremely thin and wavy apophysis margins, (5) no reflexed scales contiguous to the peduncle, (6) 13% more seed per kg, (7) seed that germinates rapidly and uniformly without after- 1966] Pinus — Andresen 15 ripening, and (8) continuous growth in the primary seed- ling stage without formation of dormant buds. Ecologically, P. monticola and P. strobus grow within similar sub-boreal to cool temperate habitats. Although there undoubtedly are clinal or ecotypic variants (Hanover 1962; Mergen 1963; Wright, Lemmien and Bright 1963) within the geographic distribution of the two species, three omnipotent growth factors are present: (1) both species are influenced by annual photoperiodic cycles of short and long days that occur between 34° and 52° N. Lat., (2) pre- cipitation ranges from 500 to 2000 mm (with up to 3450 in Coastal Washington) with pronounced dry periods during the summer months, and (3) temperatures are low enough to induce marked winter dormancy of three to six months. On the other hand, P. chiapensis is subjected to sub-equa- torial insolation from 15° to 20° N. Lat. and is under the influence of a humid, warm-temperate climate associated with abundant precipitation concentrated in the summer months. In addition, this effective precipitation is further enhanced by frequent mountain fogs. Leopold (1950) de- scribed the P. ckiapensis phyto-association as a tropical cloud forest while Beard (1944) termed it a temperate rain forest. Even though most P. chiapensis forests are limited to subtropical or warm-temperate climates there are excep- tional isolated stands that occasionally are exposed to freez- ing temperatures. In addition to the foregoing morphological and ecological divergences there also are ancient phytogeographical divari- cations that are extremely important when considering the origin of Pinus chiapensis. I believe that the original dis- persal of the phylogenitors of P. chiapensis, P. monticola, and P. strobus was accomplished at least by early Eocene and that later but similar forms, morphologically and phy- siologically allied to the three contemporary taxa, were isolated from one another by late Pliocene. My reasoning follows: Substantial evidence (Gausen 1960) of an ancient Hap- 16 Rhodora [Vol. 68 loxylon pine flora richer and more diverse than the sub- genus now extant has been revealed in the fossil record of the late Mesozoic and Arcto-Tertiary forests. With the advent of accelerated climatic change in the pre-Pliocene (Good 1953: 263), xeric climatic bands evolved (Schwartz- bach 1963) that were unfavorable to the mesophytic plants of the southern portion of the Arcto-Tertiary biome. These latitudinally oriented zones contributed to the formidable hiatus which first isolated the early Quaternary progenitors of such contemporay pine relicts as Pinus ayacahuite, P. chi- apensis, P. griffithü, P. parviflora, and P. peuce. In the early Pliocene of North America pronounced orogenesis con- tributed to a secondary discontinuity that further divided the forests above 259 North Latitude into eastern and west- ern disjuncts. It was then, in the forested and mountainous areas of the Appalachians, the Pacific North West, and the southern Mexican-Guatemalan highlands, that the immedi- ate ancestors of P. chiapensis, P. monticola, and P. strobus persisted in competition with their associates. Thus, prior to the continental and montane glaciation of the Pleistocene, isolation of the phylad segregates was com- plete and to this day final. The waves of subsequent Pleis- tocene ice with their periglacial climatic regimes undoubtedly caused areal concentration of both P. monticola and P. strobus. However, the cooler temperatures and neo- pluvial conditions that were contemporary with glaciation did not, as Deevey (1949), Dressler (1954), and Sharp (1953) suggest, create an ameliorated environment across the semi-arid Texan hiatus that would have permitted the mesophytic phyto-associations of the southeastern United States to reach and penetrate the montane regions of east- ern and southern Mexico. Martin and Harrell (1957) observed that further de- tailed ecologic and taxonomic revisions should be made of the vicariads of these two biotas. For only with new evi- dence can the theory of mid-Cenozoic disjunction be ex- panded, 1966] Pinus — Andresen 17 Finally, the divergence patterns and biometric differen- tials within the Holocene Pinus chiapensis-monticola-strobus phylad provide one such example of floristic confirmation of pre-Pleistccene isolation of the montane forests of south- eastern Mexico, as suggested by McVaugh (1952) ; Martin and Harrell 1957; and Steyermark 1950. DEPARTMENT OF FORESTRY, SOUTHERN ILLINOIS UNIVERSITY CARBONDALE, ILLINOIS ACKNOWLEDGEMENTS I am grateful to the personnel of the Western Institute of Forest Genetics and the Moscow (Idaho) Field Service Unit of the U. S. Forest Service, to Drs. John C. Genys, University of Maryland, How- ard C. Kriebel, Ohio State University, and Mr. Boone Hallberg, Oaxaca, Mexico for fresh foliage and seed. The curators of the following herbaria kindly provided specimens for the study: A, F, MEX “F” (Instituto Nacional de Investigaciones Forestales), MEXU, MSC, NA, MO, US. In addition I am grateful to Drs. John H. Beaman, J. L. Ruby, and J. W. Wright, and Mr. W. Rubel for their helpful comments. Special recognition is given to Dr. Nicholas T. Mirov who originally suggested this investigation. Mr. Nathan Shier, laboratory technician, Michigan State University, performed the majority of the leaf and cone measurements, LITERATURE CITED ANDRESEN, J. W. 1964. The taxonomic status of Pi Phytologia, 10: 417-421. . and R. J. STEINHOFF. 1965. The taxonomy of the Pinus flexilis-P. strobiformis complex. [Manuscript] ANONYMOUs 1948. Woody-plant seed manual. U. S. Dept. Agr. Misc. Pub., No. 654, vi 4- 416 pp. illus. BEARD, J. S. 1944. Climax vegetation in tropical America. Ecology 25: 127-158. BRAUN, E. L. 1950. Deciduous forests of eastern North America. Philadelphia, Blakiston Co. xiv + 596 pp. CLARK, P. J. 1952. An extension of the coefficient of divergence for use with multiple characters. Copeia 2: 61-64. CocuRAN, W. G. and G. M. Cox. 1957. Experimental designs. ed. 2. New York, John Wiley and Sons, Inc., 611 pp. Davipson, R. A. 1963. Initial biometric survey of morphological varia- tion in the Cirsium altissimum-C. discolor complex. Brittonia 15: 222-241. nus chiapensis. 18 Rhodora [Vol. 68 DEEVEY, E. S. 1949. Biogeography of the Pleistocene. Bull. Geol. Sco. Amer. 60: 1315-1416. Dor, T. and K. MoRIKAWA. 1929. An anatomical study of the leaves of the genus Pinus. Kyushu Imp. Univ. Jour. Dept. Agr. 2: 149- 198. Dixon, W. J. and F. J. Massey, JR. 1957. Introduction to statistical analysis. ed. 2. New York, McGraw-Hill Book Co., Inc. xiii + 488 pp. DRESSLER, R. L. 1954. Some floristic relationships between Mexico and the United States. Rhodora 56: 81-96. FREESE, F. 1959. A guide book for statistical transients. U. S. Dept. Agr., Southern For. Exp. Sta. 77 pp. ENGELMANN, G. 1880. Revision of the genus Pinus and a description of Pinus elliottii. Trans. St. Louis Acad. Sci. 4: 161-189. GAUSSEN, H. 1960. Les Gymnospermes actuelles et fossiles. Chapitre 11. Généralités, Genre Pinus. Trav. Lab. Forestier Toulouse, tome II, sect. 1, vol. 1. pt. 2, 272 pp. illus. Goop, R. D. 1953. The geography of flowering plants. ed. 2. New York, Longmans, Green. 452 pp. HaRLow, W. M. 1947. The identification of the pines of the United States, native and introduced, by needle structure, New York State Coll. For. Tech. Publ. No. 32, 19 pp. illus. . and E. S. Harrar. 1958. Textbook of den- drology. ed. 4. New York, McGraw-Hill Book Co, Inc. xi + 561 pp. illus. HESLOP-HARRISON J. 1963. Species concepts: theoretical and practical aspects. pp. 17-40, In: Swain, T. (Ed). Chemical plant Tax- onomy. New York. Academic Press ix + 543 pp. HEYwoop, V. H. and A. LOVE, (Eds). 1963. Symposium on biosyste- maties. Regnum Vegetabile 27: 1-72. HANOVER, J. W. 1962. Clonal variation in western white pine. U. S. Dept. Agr. Intermountain Forest and Range Exp. Sta. Res. Note 101, 4 pp. JEFFERS, J. N. R. and T. M. BLACK. 1963 An analysis of variability in Pinus contorta. Forestry 36: 199-218. KLAUBER, L. M. 1940. Two new subspecies of Phyllorhynchus, the leaf-nosed snake, with notes on the genus. Trans. San Diego Soc. Nat. Hist. 9: 195-214. KENG, H. and E. L. LiTTLE, JR. 1961. Needle characteristics of hybrid pines. Silvae Genetica, 10: 131-146. LANJOUW, J.et al. 1961. International code of botanical nomenclature. Utrecht, Netherlands, International Bureau for Plant Taxonomy and Nomenclature, 372 pp. LEOPOLD, A. S. 1950. Vegetation zones of Mexico. Ecology 31: 507-518. 1966] Pinus — Andresen 19 Loock, E. E. 1950. The pines of Mexico and British Honduras. Union South Africa, Dept. Forestry Bull. No. 35, x + 244 pp. Martin, P. S. and B. E. HARRELL. 1957. The Pleistocene history of temperate biotas in Mexico and eastern United States. Ecology 38: 468-480. MaRTINEZ, M. 1940. Pinaceas Mexicanas, An. Inst. Biol. Mex. 11: 57-84. McLemore, B. F. and F. J. CZBATOR. 1961. Length of stratification and germination of loblolly pine seed, Jour. For. 59: 267-269. McVaucu, R. 1952. Suggested phylogeny of Prunus serotina and other wide-ranging plylads in North America. Brittonia 7: 317- 346. MERGEN, F. 1963. Ecotypic variation in Pinus strobus L. Ecology 44: 716-727. Mirov, N. T. 1962. Phenology of tropical pines. Jour. Arnold Arb. 43: 218-219. Rocers, D. J. 1963. Taximetrics — new name, old concept. Brittonia 15: 285-290. ScuwARTZBACH, M. 1963. Climates of the past; an introduction to paleoclimatology. London, D. Van Nostrand Co., Ltd. xii + 328 # pp. SCHWERDTFEGER, F. 1953 Informe al gobierno de Guatemala sobre la entomolgia forestal de Guatemala. Vol. I. Los pinos de Guate- mala. United Nations-Rome, informe FOA/ETAP No. 202, 59 pp. SARGENT, C. S. 1897. The silva of North America. Vol. XI Coniferae. New York, Houghton, Mifflin and Co. 163 pp. illus. Suaw, G. R. 1914. The genus Pinus. Harvard University, Arnold Arb. Publ. No. 5, 96 pp. illus. SHARP, A. J. 1946. Pinus strobus south of the United States. Elisha Mitchell Sci. Soc. Jour. 62: 229-230. . 1953. Notes on the flora of Mexico: world distribution of the woody dicotyledonous families and the origin of the modern vegetation. Jour. Ecology 41: 374-380. SNEDECOR, G. W. 1956. Statistical methods. ed. 5. Ames, The Iowa State College Press, xii + 534 pp. SoKAL, R. R. and P. H. A. SNEATH. 1963. Principles of numerical taxonomy. San Francisco. W. H. Freeman and Co. xvi + 359 pp. Soto, J. VASQUEZ, W. BARRETT, and E. L. LITTLE, JR. 1962. Botany pp. 1-64. In: Anon. Seminar and study tour of Latin-American conifers. FAO of UN, English Edition No. 1, x + 209 pp. STANDLEY, P. C. and J. A. STEYERMARK. 1958. Flora of Guatemala. Fieldiana: Botany, Part 1. ix + 478 pp. STEYERMARK, J. A. 1950. Flora of Guatemala. Ecology 31: 368-372 SupworTH, G. B. 1908. Forest trees of the Pacific Slope. U. S. Dept. Agr., For. Serv. 441 pp. illus. 20 Rhodora [Vol. 68 SUTHERLAND, M. 1934, Microscopical study of the structure of the leaves of the genus Pinus. Trans. Proc. New Zealand Inst. 63: 517-568. WRIGHT, J. W. and F. D. FREELAND. 1960. Plot size and experimental effieiency in forest genetic research. Michigan Agr. Exp. Sta. Tech. Bull. 280, 28 pp. Table1. Computation and Comparison of Coefficients of Divergence within the Pinus chiapensis — P. monticola — P. strobus Complex. Based on Means! of Characters Q, to Qe for Taxa A to C. Taxa Parameter Means Qi Q. Q: Q: Q: Q: Q: Qs Qo Qi Qu Q: A 104.6 0.65 0.0062 12.9 4.3 87.7 17.7 9.9 1.79 22.4 2.5 0.04 B 68.1 0.92 0.0135 3.1 4.0 110.8 20.0 14.2 1.41 16.1 1.0 13.10 C 72.7 0.67 0.0092 6.1 3.4 67.6 16.4 12.4 1.32 16.4 1.1 7.00 Coefficients of Divergence, C. D., derived from the above data and means of 39 randomly selected samples from same populations values: Taxa with all samples with 39 samples species pair agegre-| aggre- Leaf cone} gate ||eaf | cone gate BC .87|.155| .169 |.189 | .145 |.165 P. monticola vs, P. strobus AC .206 .414 | .343 | -207 | 418 .346 P. chiapensis vs. P. strobus | | AB 343/424|.392 ||.348 | .423 |.393 P. chiapensis vs. P. monticola ' In the above summary of means let: A denote 60 foliage samples and 40 cone samples of Pinus chiapensis collected from individual trees found in southern Mexico, and northern Guatemala. B denote 120 foliage samples and 45 cone samples of Pinus monticola collected from individual trees found in southwestern Canada and the northwestern United States. C denote 300 foliage samples and 47 cone samples of Pinus strobus collected from individual trees found in the Lake States, and the northeastern and southeastern United States, and let following Q, to Qi» represent: Qı total leaf length in mm Q: leaf width in mm Q: ratio of leaf width to leaf length Q./Q, Q. number of serrations per 5 mm interval of leaf edge at leaf center 1966] Pinus — Andresen 21 Qo Qio Qu Qu To number of stomatal lines on vertral leaf surface total number of ovuliferous scales of mature cone width of scale apophysis in mm Length of scale apophysis in mm ratio of apophysis width to length Q:/Qs length of cone peduncle in mm involutions of edges of central scales, range from 0 to 4 number of reflexed basal-scales contiguous to peduncle; range from 0 to 18 calculate any C. D., the following general formula was used: IC BIET tas — bot Laie = bt CD. -4/ Babee 2) +... + (am ELEM k Ay ; ur Bi where: a, = AB) b= A. 4- B, for Q, etc. k — no. of characters, or 12 in this study Table 2. Means and standard deviations of characters Qi. . . Qu: for Pinus chiapensis (A), P. monticola (B), and P. strobus (C), derived from 39 random samples. F statistics for species pairs AB, AC, and BC incorporate 1 and 38 d.F. Char. chi- acter Pinus apensis Pinus monticola Pinus strobus F Statistics’ X g X s X s AB AC BC Qı 106.51 14.44 69.21 16.42 77.97 12.29 113.51 88.30 7.138 Q: 0.66 0.06 0.95 0.12 0.68 0.08 203.47 2.97 142.09 Q: 0.0062 0.0008 0.0148 0.0042 0.0089 0.0013 154.83 121.90 69.46 Q: 12.92 1.84 3.15 1.84 5.69 1.45 548.54 370.60 45.61 Qs 4.33 0.83 4.09 0.89 3.46 0.77 1.46 23.20 11.41 Qe 87.72 9.08 114.95 29.68 68.36 12.90 30.03 58.76 80.85 Q: 17.74 1.82 20.10 3.14 16.49 1.50 16.53 11.08 42.17 Qs 9.95 1.15 14.33 2.14 12.54 1.74 126.87 60.02 16.44 Qs 1.80 0.22 1.41 0 23 1.33 0.17 55.03 106.02 2.75 Qi 25.10 5.84 15.77 4.66 16.10 3.10 60.82 72.26 0.14 Qu 2.43 0.72 1.10 0.31 1.05 0.22 113.66 132.28 0.71 Qu 0.03 — 12.54 2.39 7.41 1.68 1064.13 794.19 122.13 ‘Values under 2.98 are non-significant. Value of 7.13 is signifi- cant at 5% confidence level. Values greater than 11.07 are significant at 196 confidence level. "The coefficient of variation (C. V.) discussed in the text is cal- culated from the formula C. V. = sX (100). For example in A — Q C. V. = (14.44)?/106.51 (100) = 13.6 22 Rhodora [Vol. 68 Table 3. Character reliability indices (R:)* and number of samples? of each taxon required to detect significant differences at the 5% and 1% level within species pairs AB, AC, and BC. Char- Reliability index Samples needed to detect differences acter AB AC BC AB AC BC 5% 1% 5% 1% 5% 1% Qı 983 978 754 3 5 4) 6 22 39 Q, .990 497 986 2 3 50 82 3 4 Q. 987 984 972 2 4 3 4 5 6 Q, 996 995 957 2 3 2 3 7 8 Q: 189 .917 .839 99 157 8 14 15 26 Qs 936 965 976 7 1 6 7 5 6 Q: 883 835 954 10 17 16 27 7 8 Q. 984 967 885 3 4 5 T 10 17 Qs 964 959 288 6 7 4 5 68 108 Qio 968 973 —.758 6 7 5 6 1852 2160 Qi: 983 985 —169 4 5 3 4 227 854 1 1 1 1 3 4 12 .998 | .997 .984 'Although Ri, as the intraclass correlation, is more often used to compare characters between genetically related individuals, it also serves here as a statistic to evaluate character weight, It is com- puted from the model R: = o o + g where oc? is the be- b B Ww B tween taxa component of the total variance and c? is the within wW component of the total variance for any one character. In AB—Q;, .983 means that 98.3% of the proportionate variance is related to differences between taxa rather than within taxa. High R+! values indicate “strong” characters of differentiation. ?Sample sizes were determined by solving for n in the formula n = 2 Vt/(X, — X.) which was derived from the model of “stu- dent's" t. In our sense, n — degrees of freedom or sample size, V — pooled variance of a species pair, t —estimated value from "student's" t-distribution, X: + X. = means of character X in species 1 and 2 respectively. "To calculate the number 4 proceed as follows: The variance (o?) of Q: for A is 207.36 and for C is 151.29 (pooled Vi + V; should be divided by 2), the mean leaf length for A is 106.5 and 78.0 for C; these values substituted in the n formula give: n — 2/2 (358.65) t?/ (28.5)? = .442t. Since .978 is a high R; and indicative of a low n, substitute n — 2 to find an empirical t; (t — 2n — 2): 1966] Pinus — Andresen 23 with 2 df so try 6 df so try 3 df 2 = .442U 6 — .442U 32> 4470 t for 4—2 df t for 12—2 df for 6—2 df at 5% — 2.78 at 5% = 4.30 at 5% — 2.23t 3 = .442 (1.13) 2 — .442(18.50) 6—.442(4.97) 3— 83.42 rounded = 4 2-—S8 S 6 — 2.20 so the conservative n — 4 but 218.18 but 612.20 with 4 df the “t” side was only 2.63, so the appropriate n is reached the first time the “t” side of the equation exceeds the “n” side. Table4. Internal leaf characters: number of resin canals and suc- culence represented by percents of moisture and intercellular space of P. chiapensis, P. monticola, P. strobus, and hybrid P. monticola X strobus (D). Resin Canals Succulence Taxa Mean Std. %o % No. dev. Range | Taxa Moisture Intercellular sp. D 1.89" 581 0-4 A 67.75** 407E B 2.25 .266 2-3 B 58.89 10 A 2.52 .207 2-3 C 58.76 10 C 2.57 176 2-3 D 58.26 10 ‘Taxa D and B or A and C not significantly different from each other in number of resin canals but former pair is significantly dif- ferent from latter at 1% level using the value of t. **Taxa A values significant at 1% level, B to D non-significant. Table 5. Seed and Seedling Characteristics of Pinus chiapensis, P. monticola, and P. strobus. Taxa Seed per kg Seed Germination Germination rs rate in days values' X Range Unstrat.* Strat. Unstrat. Strat. A 61,100 46,900 to 104,200 | 10 to 37 10to 37 10.35 10.35 B 52,300 39,400 to 67,800|10t01864- 13 to186+ 0.04 0.48 C 53,800 47,000 to 66,700|16t0186-- 13 to186+ 0.48 1.28 1Unstratified seed stored dry at 4°C. for 60 days. ? Seed stratified in moist peatmoss at 4°C. for 60 days. °Germination value = greatest cumulative number of seed that germinated in fewest days multiplied by total percent germinated, divided by total number of days in test period (McLemore and Cza- bator 1961). Rhodora [Vol. 68 Seedlings Hypocotyl Cotyledon Length mm Length mm Number Xx Range X Range X Range 39 20 to 70 28 20 to 40 8 6 to 10 20 15 to 35 18 14 to 25 8 7 to 10 31 18 to 44 25 15 to 30 10 8 to 12 CYTOGEOGRAPHY OF ORONTIUM AQUATICUM (ARACEAE)! JOHN W. GREAR, JR.* The Golden Club, Orontium aquaticum L., constitutes a monotypic genus of subfamily Calloidae Engl., an alliance also including the genera Symplocarpus, Lysichitum and Calla. These genera are interpreted as temperate or sub- arctic extensions of the predominantly tropical family Ara- ceae. All but Orontium are represented in eastern Asia and in North America, and thereby lend strength to the affinities existing between the floras of these regions. Morphologically, Orontium is regarded as a derived, highly modified genus without any near relatives. The geo- graphical distribution of the genus and its allies lends con- siderable support to this view. Orontium aquaticum is a perennial herb possessing 4 deep-seated, firmly-anchored, vertical rhizome. Its distri- bution (Fig. 1) is restricted to the eastern United States, where it occurs on the Coastal Plain in aquatic or semi- aquatic habitats such as bogs, marshes, streams and pools, and in similar habitats of the inland physiographic prov- inces, sometimes up to 2800 ft. in the mountains. However, lack of suitable habitats has localized its distribu- tion in the Appalachian Highlands. To date there has been no comprehensive morphological study of Orontium. Harshberger (1916), in his account of the vegetation of the New Jersey pine barrens, described and illustrated the anatomy of the leaf in consider- able detail. In a meticulous study of the floral structure, *Present Address: New York Botanical Garden 1Condensed from a thesis submitted to the Department of Biology in partial fulfillment of the requirements for the degree of Master of Science in the Department of Biology, Vanderbilt University, January 1963. 25 26 Rhodora [Vol. 68 Fig. 1. Map showing distribution of Orontium based upon records documented by herbarium specimens. The southernmost extent of glaciation is shown by the dotted line. The heavy broken lines sepa- rate respectively the Coastal Plain, Piedmont and Appalachian high- lands, including the Blue Ridge, Ridge and Valley and Appalachian Plateaus provinces. Schaffner (1937) pointed out inaccuracies of previous descriptions and reported a wide range of variation among the flowers on a single spadix. In a taxonomic study of the temperate North American Araceae, Huttleston, (1953, unpublished) found the chromosome number of Orontium to be 2n—28, a number different from that previously found by Malvesin-Fabre (2n—24, 1945, un- 1966] Orontium — Grear 27 published) see Darlington and Wylie, 1956. In the treat- ment of the Arales for the generic flora of the Southeastern United States, Wilson (1960) summarized the existing in- formation concerning Orontium, indicating that the mor- phological and biological details of this plant warranted further investigation. The present investigation was prompted by conflicting statements about the chromosome number as well as the relationship between this data and the distribution of the plant in all major physiographic provinces of the Eastern United States. It was further felt that by correlating cyto- logical with geographical data, some insight into migra- tional patterns might be gained. Through the correlation of past geological history with cytological, morphological, taxonomical and ecological evidence, it is sometimes possible to postulate an earlier distribution and possible migration route, which would ex- plain the present distribution of Orontium. The Araceae are a predominantly tropical family and during the Ter- tiary period, when tropical conditions prevailed farther northward, a wide range of plants such as temperate coni- fers, deciduous hardwoods and warm-temperate plants, including araceous plants, ranged to high northern lati- tudes, far beyond the limits of the present tropics. In Symplocarpus and Lysichitum, the two genera most closely related to Orontium, the migration route to present ranges appears to have been from the region of the Arcto- Tertiary flora of the then tropical boreal regions of west- ern North America and eastern Asia (Rosendahl, 1911). Subsequent climatic changes forced them to migrate south- ward. Symplocarpus apparently migrated to eastern North America while Lysichitum remained in the West. As with Orontium, plants in both of these genera require a very moist or wet habitat. Symplocarpus foetidus (L.) Nutt., constituting a monotypic genus, occurs in two widely dis- junct areas — eastern Asia and eastern United States. Lysichitum consists of two closely related species: L. cam- tschatcense (L.) Schott, the Asiatic one, occurs in Japan 28 Rhodora [Vol. 68 and Kamtschatka, and its American counterpart, L. ameri- canum Hultén and St. John, is distributed in western North America between California and Alaska (Hultén & St. John, 1931). Fossil evidence substantiating a more extensive dis- tribution for araceous plants has been found in Spitsbergen (presumably as imprints) referable to extant species. An aquatic aroid from the Tertiary flora, Acorus brachysta- chys, has been described recently by Schloemer-Jager from the Brogger Peninsula of the same island (Andrews, 1961). Cockerell (1926) gives evidence supporting a once more westerly distribution of Orontium in North America. The relevant specimen consisted of a spadix, incomplete at the distal, fertile end, and its sterile stalk. Although the identity of the specimen is uncertain, it was sufficiently characteristic, according to the author, to be referred to this genus. The only character found to separate the fossil plant from the modern species was a lack of thickening of the scape below the spadix. Cockerell states that “this fossil may take the name Orontium fossile n. sp.; it adds one more to the numerous examples of genera now existing in the Eastern and Southern states, but found in the Rocky Moun- tain Region only in the fossil state.” If Orontium and related genera migrated from the re- gion of the Arcto-Tertiary flora, then they were probably distributed over a wide range during the late Cretaceous and early Tertiary, from the east coast of the continent to the region of Colorado or perhaps even further west, under the influence of the equable climate of the era. At that period, the Appalachian Uplands were essentially a pene- plain, affording ideal conditions for coastal-plain types such as Orontium in the streams and swampy areas. Uplift of this old Cretaceous (Schooley) peneplain during the Mio- cene epoch, and the accompanying climatic changes and erosion cycles resulted in greatly altered conditions. The climate to the west became much more arid, eliminating possible populations of Orontium west of the Mississippi River. A more restricted distribution was placed upon the 1966] Orontium — Grear 29 species in the Appalachian Region and only relict popula- tions survived in the favorable mesophytic and hydro- phytic habitats of the newly elevated highlands. With the later recession of the sea, the newly-emergent Coastal Plain became available to the migrating species from the up- lands, which found optimum conditions in the bog-marsh habitats. An account of the Coastal Plain element in the flora of the Appalachian uplands, correlated with physio- graphic history of the provinces concerned, has been de- scribed thoroughly by Kearney (1900), Harshberger (1903), Fernald (1931, 1937), Braun (1937), Carr (1938), Core (1938), et al. Observation of the map (Fig. 1) shows that the present range of Orontiwm has extended into the formerly glaciated region in some of the Northeast. Peattie (1922) reports the occurrence of Orontium from as far inside the glacial boun- dary as Lake Oneida, New York. During the glacial and interglacial stages of the Pleistocene, the climatic fluctua- tions probably caused a retreat of the species to refugia farther south of the advancing ice. From these refugia, when the ice receded for the final time, probably it migrat- ed to its present northernmost limits. As stated earlier, varying chromosome numbers for Or- ontium have been reported. Huttleston (1953) has counted 28 somatic chromosomes from root tip squashes of four plants, two from Duval County, Florida, and two from Middlesex County, New Jersey. Both of these locations are in the Atlantie Coastal Plain. A different count, of 24 chromosomes, has been reported by Malvesin-Fabre (1945) in a thesis submitted to the University of Bordeaux, France. As this work was unobtainable, the source and the state of the studied plant material are not known to the present writer. However, the possibility of different numbers sug- gested that some correlation might exist between chromo- some number and geographical distribution. It was planned originally to work with meiotic material, but flowers from plants collected in early April of 1961 30 Rhodora [Vol 68 Fig. 2. Chromosomes of Orontium. First meiotic metaphase, Ma- terial from Fentress County, Tennessee. X approx. 500. had already undergone meiosis. In collections made in early September of the same year, meiosis had already occurred in young flowers that would not make their appearance until the following spring. It appears that the greater part of the meiotic process probably occurs sometime during the summer season, more than eight months in advance of of anthesis the following spring. Only one determination of chromosome number was made from meiotic material. This was obtained quite by chance from the tip of an inflorescence collected in Fent- ress County, Tennessee, April 1961, in which meiosis was complete. A few pollen mother cells of the young sporo- genous tissue showed 26 chromosomes at the first meiotic metaphase, the number not yet having been reduced. The inflorescence was collected in Newcomer’s solution and the squash was made using acetocarmine. Some of the chromo- somes exhibited a tendency to stick together, as evidenced by strand-like connections between them (Fig. 2). Because of the uncertainty relating to the exact time of meiosis, all other work was done with root tip chromosomes. Living samples of Orontium, variously consisting of fruits, seedlings and mature plants, were obtained from various sources for root tip material. The samples were maintained in the green house in concrete water tanks dur- ing the course of investigation. Fresh, vigorously-growing 1966] Orontium — Grear 31 root tips about 1 cm. in diameter were excised immediately before treatment. The live tips were pretreated with a-bromonapthalene to inhibit spindle formation, shrink the chromosomes and render the centromeres more evident. The method of Tijo and Levan (1950), utilizing 8-hydroxyquinoline as a pre- treatment agent, proved less effective in fixing the meta- phase than a-bromonapthalene even though other conditions were constant. The root tips were then killed and fixed with aleohol-acetic acid (3:1) and stained with aceto-orcein after the method of LaCour (1941). The meristematic tip of the root was then excised and squashed in a drop of 45% acetic acid. Mitotic chromosome counts were obtained from 24 dif- ferent populations and in every instance the diploid number was 26. Neither of the previosly reported numbers (2n — 24, 28) was ever found, although attempts to verify Huttleston's Duval County report were made using material collected from the same site. The chromosomes are very large, intensely-staining and easily observed where the squash preparation is good. The position of the majority of the centromeres is median (iso-brachial), a feature re- garded by some workers (e.g. Levitzky, 1931) as represent- ing the generalized or primitive condition. The chromosomes of one pair have clearly discernable satellites (Fig. 3). A tendency toward fragmentation at the centromere may ac- count for Huttleston's report of 2n = 28. However, this was a random occurrence in the present study. Because the chromosome number appeared the same in all plants examined, regardless of source, consideration was given karyotypic variation among populations. Graphic portrayal of chromosome arm length was attempted, but, because of differential shrinkage and swelling of the chromo- some arms it was not possible to match homologous chromo- somes. Therefore, no correlation was found between quantitative chromosomal data and the geographical source of the samples studied. The only two chromosomes that could be paired with any degree of certainty were the two 32 Rhodora [Vol. 68 = Ó Ea Dm Z e c 3 Fig. 3. Chromosomes of Orontium. Mitotic chromosomes. Material from Cameron Parish, Louisiana. X approx. 700. with satellites. Until a more reliable procedure is developed, permitting assessment of differences in linear dimensions, the quantitative information must be considered insignifi- cant. It appears that Orontiwm chromosomes would be par- tieularly suitable for differential staining (see Fukuda and Kozuka, 1958, for work on Trillium), and that the homolo- gous pairs of chromosomes might thereby be identified. There was some indication of differential stainability of the chromosomes of Orontium during the present investigation and this suggests the possibility of relating the populations to distribution. SUMMARY Information obtained from this study contributes to knowledge of the cytology of Orontium, a monotypic genus of the Araceae. The diploid chromosome number of all populations studied is 26, irrespective of geographical 1966] Orontium — Grear 33 source. Two satellites or secondary constrictions are evident in every karyotype and the chromosomes are extremely large (ca. 204), exhibiting essentially median centromeres. DEPARTMENT OF BIOLOGY VANDERBILT UNIVERSITY, NASHVILLE, TENNESSEE ACKNOWLEDGEMENTS Appreciation is due the following persons, who supplied living ma- terial as well as valuable information regarding distribution: Dr. Wade T. Batson (University of South Carolina), Dr. Harold D. Ben- nett (West Virginia University), Dr. Edward Davis (University of Massachusetts), Mr. J.B. Ebert (Pembroke State College), Dr. Robert Kral (Virginia Polytechnic Institute), Dr. Carl Monk (University of Florida), Dr. Herman O'Dell (University of East Tennessee), Dr. James D. Ray Jr. (University of South Florida), Dr. William Reese (University of Southwestern Louisiana), Drs. Elsie Quarterman, R. B. Channell, Howard F. L. Rock, Messrs Yoshimichi Kozuka, H. Oliver Yates and William Brode (Vanderbilt University). I am particularly indebted to the late Dr. Howard F. L. Rock, supervisor of this work, and to Dr. R. B. Channell, who suggested the problem, for their advice and encouragement, and to Dr. Howard S. Irwin for critical evaluation of the paper. The writer wishes to acknowledge the curators of the following herbaria for the loan of material: ALU; BUS; CINC; DUKE; FSU; GA; GEO; GH; IND; KY; MASS; MISSA; NCU; NO; NY; SMU; TENN; TEX; UARK; US; USF; VDB; WVA; UNIVERSITY OF SOUTHWESTERN LOUISIANA. LITERATURE CITED ANDREWS, H. N. 1961. Studies in Paleobotany. New York: John Wiley & Sons. p. 381. BRAUN, E.L. 1937. Some relationships of the flora of the Cumberland Plateau and Cumberland Mountains in Kentucky. Rhodora 39: 193-208. Carr, L.G. 1940. On the “residual” interpretation of coastal floras in the Appalachian uplands of eastern United States. New Phytol. 39: 129-132. COCKERELL, T.D.A. 1926. A miocene Orontium (Araceae). Torreya 26: 69. Cork, E.L. 1938. Plant migrations and vegetational history of the southern Appalachian region. Liloa 3: 5-29. 34 Rhodora [Vol. 68 DARLINGTON, C.D. & A.P. WYLIE. 1956. Chromosome Atlas of Flower- ing Plants. New York: The Macmillan Co. p. 374. FERNALD, M.L. 1981. Specific segregations and identities in some floras of eastern North America and the old world. Rhodora 33: 25-63. —————————— 19387. Local plants of the inner Coastal Plain of southeastern Virginia. Rhodora 39: 266-321, 379-415, 438-459, 465-491. FUKUDA, I. & Y. Kozuka. 1958, Evolution and variation in Trillium, V. A list of chromosome composition in natural populations of Trillium kamtschaticum Pall. Jour. Fac. Sci. Univ. Hokkaido Ser, V. 6: 2779-319. HARSHBERGER. J.W. 1903. An ecological study of the flora of moun- tainous North Carolina. Bot. Gaz. 36: 241-258, 368-383. 1916. The vegetation of the New J ersey Pine Bar- rens. Philadelphia: Christopher Sower Company. pp. 265-260. HULTEN, E. & H. St. JoHN. 1931. The American species of Lysichitum. Sv Bot. Tidskr. 4: 453-464. HUTTLESTON, D.G. 1953. A Taxonomic Study of the Temperate North American Araceae. Cornell. (Thesis). KEARNEY, T. H. 1900. The lower austral element in the flora of the southern Appalachian region, Science 12: 830-842. LEVITZKY, G. A. 1931. The karyotype in systematics. Bull. Appl. Bot. Genet. Plant Breed. 27: 220-240. MALVESIN-FABRE. 1945. Contribution a la caryologie des Araceae. Bordeaux. (Thesis). PEATTIE, D.C. 1922, The Atlantic Coastal Plain element in the flora of the Great Lakes. Rhodora 24: 57-70, 80-88. ROSENDAHL, C.O. 1911. Observations on the morphology of the under- ground stems of Symplocarpus and Lysichiton, together with some notes on geographical distribution and relationship. Minn. Bot. Stud. 4: 137-152. SCHAFFNER, J. H. 1937. The flowers of the golden-club. Am. Bot. 43: 99-103. WILson, K.A. 1960. The genera of the Arales in the southeastern United States. Jour. Arnold Arb. 41: 47-72. AUTHORS OF PLANT GENERA AND THE INTERNATIONAL PLANT INDEX REED C. ROLLINS An impressive volume printed with the aid of computers and assembling in one place a lot of information heretofore somewhat scattered has recently appeared.’ However, most botanists will not find the information to be of great in- trinsic value and an occasional consultation of the work will probably suffice for their purposes. The bulk of this paperbound book is devoted to an alpha- betical listing of the “authors” accompanied by four columns under the headings, AUTHOR CODE, BORN, COUNTRY, and DIED. This is followed in double-column format by 62 pages of “author” listings, giving the author code in each instance. The final section gives a three-column listing of name ab- breviation, author name and author code in a double-column format, followed by an addenda to each of the three lists. If botanists did indeed take up and use the author code- order index for citation of author names of plants, as urged by Mr. Gould, it would be a radical departure from past practice. It would also involve giving up to the machine the essences of familiarity which link plant names with the men who gave them to the plants. We now have no diffi- culty in remembering that Schlecht. stands for Schlechtendal or Schott for Schott but how many botanists will remember that SCH794 is Schlechtendal and SCO794 is Schott? How can one be expected to remember that SHO904 is A. J. Sharp and SHA904 is W. M. Sharp? These unnecessary abbrevi- ations reflect the choice by the authors of an inadequate system to handle the information they are trying to present. :Authors of Plant Genera, by Sydney W. Gould and Dorothy C. Noyce. International Plant Index 2: 1-336. 1965. The New York Botanical Garden, New York; The Connecticut Agricultural Ex- periment Station, Box 1106 (IPIx) New Haven, Connecticut 06504; $6.00 in U.S.A. and Canada, $6.50 in other countries. 35 36 Rhodora [Vol. 68 It can be argued that having the names of the men of historical botany closely associated with the names of the plants they helped reveal to the public is of no great moment. However, it is not as simple as that. The name of the author, if it is known and recalled, helps put the name of the plant into its historical setting, which in turn helps with the geography and other matters concerning the plant itself. The richness of the presently used name content, with ci- tations that are well understood and appreciated, will be greatly denuded by the device offered in the book under review. In addition, one will be forced to look up, on every occasion, the correct author code. This will be a nuisance for taxonomists; for non-taxonomists, it will make author citations both incomprehensible and ridiculous. Are citations such as Rhododendron canadense (LIN707)BSP859 or Buchloé dactyloides (N UT786) ENG809 to the advantage of either taxonomy or plant sciences generally? many persons who have not named any plant taxon what- ever. Many are not botanists at all. One finds librarians, zoologists, friends of the authors and the authors themselves, none of whom named genera, Is this the kind of capricious- ness that should characterize an accurate scholarly work? Do the authors think they are playing games with the ma- 1966] Plant Index — Rollins 37 terials of science? What then are the criteria for the in- clusion of a name in the “Authors of Plant Genera”? ACCURACY AND COMPLETENESS: One reliable and readily available source that should have been a major check-point for “Authors of Plant Genera" is Index Nominum Generi- corum, published by the International Association for Plant Taxonomy. Apparently this recent authentic work, now only partially published but having over 21,000 cards in it, was hardly utilized at all. This circumstance permits us to use it as a check for the accuracy of the book under review. The alphabet from Aa to Af, amounting to approximately 400 cards of Index Nominum Genericorum, was checked against “Authors of Plant Genera” by a colleague. This number turned up six errors Or omissions, of which five were complete omissions of botanists who had described genera of plants. If this rate of omission holds when checked against the rest of the Index Nominum Genericorum presently available, it means that over 250 omissions of authors who named plant genera were carelessly left out of the book being billed as “the most complete work of its kind.” If Index Nominum Genericorum were complete, one wonders how many thousands of names that legitimately should have been included in “Authors of Plant Genera" would be shown up by a check against it. Checking for accuracy is a time consuming and unin- spiring enterprise and one tends to turn to the quickest way of seeing whether the data given in à book stands up or not. We noticed that Leslie Andrew Garay and Clarence Garay are the same in the author code, page 86, and that he is alleged to be from Canada. Leslie Garay, who works nearby, tells me that he has never published under the name of Clarence Garay and has not been known under that name. Furthermore, he has been in the United States since 1957. and was born in Hungary. Although he did spend some years in Canada, the information on Garay is both inac- curate and misleading. We know that Thomas Coulter was born, lived and died in Ireland, but on page 69 he is listed as England. On the 38 Rhodora [Vol. 68 other hand, N. L. Bor is given as Ireland, although he Specialized on the grasses of southeast Asia, spent much time there, and for many years he has been at Kew in England, where he has been professionally identified right down to the present. Oddly enough, although Edward Perceval Wright was born, worked and died in Ireland, ap- parently on the basis of a visit to the Seychelles Islands for six months in 1867, "Seychell. has become his "country of principal work or home." In short, every entry under this column must be checked by the user to determine its meaning and accuracy, if these are important to him. Under the column "Country" the strangest designations appear. For example, S. America is given as a country many times, but so also are Argentina, Brazil, Colombia, etc., the true countries of South America. It must seem strange to the well known Argentinian botanists, Professors Arturo Burkart and Lorenzo R. Parodi, to read that their country is "S. America." I am Surprised to see listed as countries: Patagonia, Sahara, Transvaal, South West Africa, etc. Certainly these subversions cannot be read as a sign of erudition on the part of the authors. One is led to ask, how can such sloppiness be supported as a crusade ostensibly to help librarians, taxonomists and plant scien- tists ? THE INTERNATIONAL PLANT INDEX: The two volumes so far published by IPIx, as poorly prepared as they are, can scarcely be said to have done any real harm. This is so because the content of each of them is readily found else- where in reliable sources and the real scholar will soon discover that the volumes are not worth bothering with, in spite of the exorbitant claims to the contrary. However, I am alarmed by the forecast that the third volume will be "Genera of the Plant Kingdom." If such a work is published with the same lack of understanding of botanical nomen- clature and taxonomy, and the same disregard for the In- ternational Code of Botanical Nomenclature that is evident in the published work so far done by the people at Inter- national Plant Index, it will be a disaster much worse than 1966] Plant Index — Rollins 39 the appearance of the infamous volumes of Otto Kuntze? in the 1890's. We have no quarrel with the use of computers or other data-processing machines to help organize and produce an index of plant names. Properly handled, such a single com- plete index would not only be extremely useful in itself but it could be the basis for encoding a wide range of botanical information that could then be made readily available. We part company with the organizers of IPIx when they insist on going beyond a simple index, yet use only books (and secondary sources, at that) as the basis for value judge- ments concerning the taxonomy of the plants whose names they are handling. In Family Names of the Plant Kingdom,’ it is a simple fact that a value judgement was made every time a name was selected for use as as operative name. A taxnomic judgement was made every time a name was listed as a synonym. These are facts even though the authors specifically state that the contrary is the case. The hundreds of value judgements made by the author of *Family Names" were completely unsupported and evi- dently were made without any real knowledge of the plants involved or of any reasons why one course of action should be taken in preference to another. The use of the word "operative" or some other designation such as “correct” or “in use” does not lessen the responsibility for making an acceptable choice. To be acceptable the name chosen has to stand up to the requirements of the science. It is not the business of an indexer to make choices where taxonomic judgements are involved, unless he can produce the scientific evidence to back up the choice. Quite obviously, Family Names of the Plant Kingdom is not an unbiased index and it cannot, therefore, be reliably used as an index. As à HE orcum ?Revisio Generum Plantarum. Pars I-III plus supplement. 1891- 1898. *Family Names of the Plant Kingdom, by Sydney W. Gould. Inter- national Plant Index 1: 1-111. 1962. For a review see Botanical Nomenclature, Punched Cards, and Machines, by C. E. Wood, Jr., R. S. Cowan and G. Buchheim. Taxon 12: 2-12. 1963. 40 Rhodora [Vol. 68 reliable source of the proper name of a family, it is com- pletely hopeless. In his writing, Mr. Gould has lectured to taxonomists, quoting his favorite authors and giving naive analogies to telephone communication and space science. Unfortunately he has kept himself immune from even an elementary under- standing of the basic ideas and tenets of taxonomy and nomenclature, the subjects in which he is apparently trying to attain an authoritative position. The real danger that lies ahead arises from the fact that those most deeply committed to IPIx are untrained for the task they are trying to perform and will not listen to any- one knowledgeable enough to help them. Their supporters and advisers are equally lacking in an understanding of the problems involved. If the International Plant Index continues in the path so far laid down, taxonomists and botanists generally will be saddled with a nomenclatural mess, a heritage largely from those who have willfully or unwittingly given their support to IPIx. GRAY HERBARIUM HARVARD UNIVERSITY STUDIES IN THE CAPPARIDACEAE' VIII. POLANISIA DODECANDRA (L.) DC.: HucH H. ILTIS? FURTHER NOTES ON ITS TYPIFICATION. In two studies in this series (Iltis 1954, 1958) it was maintained that the common American Polanisia graveolens Rafinesque should properly be called P. dodecandra (L.) DC., the latter based on Cleome dodecandra of Linnaeus's Species Plantarum (1753). However, in a recent generic review of the Caper family for the Southeastern United States, Ernst (1963: 90-91, footnote 6) casts serious doubts on the validity of this disposition, suggesting instead that the C. dodecandra L., based as it is on a description in the Flora Zeylanica (No. 242, p. 109), a work based in turn on Paul Hermann's Ceylon plants and drawings, is actually a hitherto unrecognized or misunderstood Asiatic species which should not be equated with the well-known American Polanisia graveolens Raf., the two being presumably totally different taxa. The present paper attempts to show that on the basis of the critical collections housed in London (and 1[Ed.: Published as Capparidaceae rather than Capparaceae at the request of the author.] ?I am much obliged to the Botanical Society of America for a grant- in-aid to attend the Xth International Botanical Congress in Edin- burgh, and thus be privileged to visit London and its herbaria, and to Dr. Wallace Ernst, U. S. National Herbarium, Smithsonian Insti- tution, Washington, for the careful presentation of his arguments in correspondence. The staffs of the Linnaean and British Museum herbaria, especially J. E. Dandy, Keeper of Botany, N. K. B. Robson, and D. Hillcoat have been most generous with their help, as has been my colleague John W. Thomson and particularly Mr. Dandy for con- structive and instructive criticisms of the manuscript. To them, my very sincerest thanks. This research was supported in part by Re- search Committee of the University of Wisconsin on Funds from the Wisconsin Alumni Research Foundation. 41 42 Rhodora [Vol. 68 thus on somewhat different grounds than previously), and on a careful reexamination of the texts involved in this argument, my original conclusions must still be considered valid: namely that Cleome dodecandra L. and Polanisia graveolens Raf. are synonymous. Therefore, P. dodecandra (L.) DC. is still the correct name for the American taxon. The original Linnaean citation (Sp. Pl. ed. 1, 2: 672) of Cleome dodecandra reads as follows: "dodecandra. 5. Cleome floribus dodecandris, foliis ternatis. Fl. Zeyl. 242*". As was fully discussed in earlier studies (Iltis 1954, Ernst 1963), the two other following citations refer to other species. The star after Flora Zeylanica indicates the pres- ence of a good description in that work. As to the origin of the species, he reported it as “Habitat in Indiiis" meaning by that the warm regions of both Asia and America, as shown by the inclusion in synonomy of a Sloane plant from Jamaica. As was pointed out previously (Iltis 1954), the short de- scription in the Flora Zeylanica is nevertheless a vivid. and. excellent one (apparently having been made from living plants), and, with its reference to 8 stamens, 3-foliolate leaves, emarginate white petals, red pistil, unilateral gland and thick, hispid capsules, fits exactly only one taxon in the world, the American Polanisia graveolens. Thus, while Lin- naeus' total conception of his species was confused if his diverse synonomy is considered, his own personal descrip- tion of the plant in question is brilliantly clear. "While the third and last of these Linnaean polynomials clearly refers to the New World C. serrata Jacq., the second polynomial (and reference to “Burm. Zeyl. 216, t. 100 f. 1") was thought to refer to perhaps C. burmannii W. and A., or to C. rutidosperma DC., though, truly, the drawing is unidentifiable. In the Geneva Herbarium are several "Herbier Burmann" collections, including one that matches this drawing — a depauperate miserable specimen, about 15 cm tall, with one hidden flower, labelled “Cleome dodecandra" [by Burmann?], a plant that conceivably could be a young C. viscosa or C. aspera, or [doubtfully], as Briquet had annotated it, C. burmannii. What- ever taxon this specimen belongs to, the plant is much too poor for Linnaeus to have used in drawing up the excellent Flora Zeylanica description, and can thus not be considered type material. 1966] Polanisia — Iltis 43 However, the Flora Zeylanica is not based on American collections, but on collections of plants and drawings made in Ceylon in ca. 1670-1677. But how can we reconcile a tem- perate American plant in tropical Ceylon? Fortunately, these drawings and plants of Hermann’s are still preserved in the British Museum of Natural History, where they may be studied. They are thus of immense scientific value. In the Flora Zeylanica, to quote Stearn (1957 "1183"... only phrase-names are used but, as pointed out by Trimen (1887), when dealing with Ceylon plants in the Species Plantarum, ‘Linnaeus was careful to quote under them the number of the Fl. Zeylan., and thus the specimens of Her- mann’s herbarium become types for many of Linnaeus’s species . . . especially as the large majority of the species in Hermann’s herbarium are unrepresented in Linnaeus’s own collection.’ " Thus, when typifying a species described in the Flora Zeylanica, one must consult the Hermann her- barium! Recently an opportunity presented itself to make a search for the type of Cleome dodecandra, and, with the generous help of Miss D. Hillcoat, the four volumes of the Hermann herbarium were examined page by page, specimen by specimen. Of the Cleome species cited in the Flora Zey- lanica, the following were found to be represented by speci- mens: C. gynandra, Fl. Zeyl. No. 239 (in vol. I, p. 1, and vol. Hb: 754 C. viscosa, Fl. Zeyl. No. 241 (in vol. I, p. 75, and vol. III, p. 2); C. monophylla, Fl. Zeyl. No. 243 (in vol. I, p. 52 and 57, and vol. III, p. 2). There were no specimens for C. icosandra, Fl. Zeyl. No. 240, and, more importantly, there were none for C. dodecan- dra Fl. Zeyl. No. 242! Next, an examination of Hermann's Icones volume showed but two Cleome illustrations, one on page 421 (numbered 242 — was this number entered there 44 Rhodora [Vol. 68 by Linnaeus.'), the other on page 24, neither of which could be C. dodecandra. Both appear to represent normal plants of Cleome aspera, the latter figure (p.24) but a small- scale copy of the first, and matching very well such Cleome aspera collections as Beddome 203 from Madras, India (BM!) or the C. aspera type (K!). Thus, neither Her- mann's drawings nor his herbarium contain any material referable to Linnaeus's Cleome dodecandra. Futhermore, we can follow neither Trimen (1887:146), who equated No. 242 of the Flora Zeylanica (i.e. C. dodecandra, with the first of these drawings (No. 242), as “C. viscosa L. var.", nor Ernst (1963), who, following Trimen, suggests that this drawing should be taken as a type of C. dodecandra. For these draw- ings, both evidently of the same plant, represent C. aspera, a 6-staminate, glandless, slender-fruited species, and in no way agree with Linnaeus's excellent descriptions of C. dode- candra. As a matter of fact, how could Linnaeus have obtained all this detail, as well as flower and pistil color, from this simple drawing? At this point Mr. Dandy very kindly (and literally) came to the rescue, for in examining the Flora Zeylanica text (p. 109) he immediately noted a very significant omission in the Cleome dodecandra description, one that neither I nor Ernst was aware of: namely that Linnaeus does mot cite (as he usually does in the Flora, Zeylanica) any collec- tion or plate of Hermann’s Musaeum Zeylanicum following the description of C. dodecandra! This can only mean that Linneaus did not have a Hermann specimen in front of him when he drew up this description, but rather some plant of different origin, a plant which he for some reason must have believed should belong in a book on Ceylon plants. Very possibly, he equated one of his garden plants with Burmann's Ceylon drawing which, being based on a Ceylon plant, was to be included in Flora Zeylanica whether or not ‘According to Trimen (1887:130), “In the [Hermann] herbarium itself he [Linnaeus] has added to Hermann's labels a reference to the number of the species in his own ‘Flora Zeylanica. " 1966] Polanisia — Iltis 45 Hermann had collected it. In any case, he must have had his reasons, for the inclusion in the Flora Zeylanica of plants other than those collected by Hermann in Ceylon is a rare (though not unknown) occurrence. That an error on the part of Linnaeus is involved here was deduced earlier on somewhat different grounds (Iltis 1954: 67, footnote). Since Linnaeus evidently had a plant available when writing the description of C. dodecandra, we are now again forced to look for a specimen, and the logical place to seek is in the Linnaean Herbarium. There, the one plant that clearly fulfills all requirements is sheet 850.12 (Savage 1945). In my earlier studies I could only guess at the iden- tity of this plant from the poor photograph then available. Now, however, after an examination of the actual speci- men, there is no doubt that this plant is indeed the same as Polanisia graveolens Raf., with the emarginate petals, and at least 8 stamens (inflorescence very young and not vigor- ous) clearly visible. Since Linnaeus mentions “capsula crassa, hispida”, he must have had in hand (or in memory) the characteris- tically thick and hispid-fruited plants of this species, prob- ably from his Uppsala garden, where the species was evidently in cultivation, the specimen in question (850.12) having a small tag glued to its stem that reads “H. U. 12 andr” [Hortus Upsaliensis dodecandra]. This cultivated plant, therefore, we must take as the type of Cleome dode- candra L.: it is clearly the only specimen that agrees (and then exceedingly well) with the original description, and is furthermore labelled by that name with a tag that we have reason to believe is contemporaneous with the publication of the Flora Zeylanica (1747) or earlier. As to the country of origin, we can only guess the Northeastern United States or adjoining Canada. Linnaeus evidently did not know the source, but believed it to be identical to that of the Burman and Boerhaave plants from tropical Asia cited in Flora Zeylanica. Complicating this story are the whole series of errors that occurred in the labelling (contemporaneously or subse- 46 Rhodora [Vol. 68 quent to 1753?) of the Cleome collections in Linnaeus's herbarium as well as in his texts. Thus: 1) he was mistaken in later writing on sheet 850.12 the name “viscosa” (though the older tag label says *H. U. 12 andr"!). Whoever scrib- bled this reidentification on the bottom of this sheet (and presumably it was Linnaeus himself ), he must have suffered a lapsus calami, for this specimen is so unlike any of the other sheets labelled “viscosa” that the identification ap- pears absurd; 2) he was mistaken when he wrote ''dode- candra" on sheet 850.14 which is clearly C. viscosa, and which in every respect matches the other C. viscosa sheets in the Linnaean Herbarium (e.g. 850.11, 850.13) ; 3) he was mistaken when he wrote “dodecandra” on sheet 850.7, which carries a good specimen of C. serrata Jacq. (C. polygama L.), only later to scratch this name out, and to substitute the correct name “polygama” after it; 4) lastly he made the curious error of misquoting the word octandris as dodecan- dris when copying the original definition (polynomial) of No. 242 from the Flora Zeylanica to the Species Plantarum, and then taking the specific epithet dodecandris from this miswritten adjective. It is an error easy to understand, however, for the young specimen (850.12) that he studied, as young specimens of that species are apt to, had 8 stamens, while no doubt on the somewhat older plants in his garden he observed the commoner higher stamen number, hence the later and very descriptive epithet “dodecandra.” SUMMARY Cleome dodecandra L., though published in Flora Zey- lanica, is based on cultivated American materia] grown in Hortus Upsaliensis by Linnaeus and now preserved in the Linnaean Herbarium. This material is conspecific with Polanisia graveolens Raf. The correct name of this taxon is, therefore, as was maintained earlier (Iltis 1954) on less certain evidence, Polanisia dodecandra (L.) DC., or, if re- tained in Cleome, C. dodecandra L. 1966] Polanisia — Iltis 4T POSTSCRIPT THE SUBSPECIES OF POLANISIA DODECANDRA. A reinterpretation of evidence presented earlier (Iltis 1958, esp. Figs. 6 and 9-21), and observations of some of these taxa in the field, demands that the Mexican and South- western P. uniglandulosa (Cav.) DC. should be considered a perfectly distinct species on morphological as well as geo- graphic grounds, and not as a subspecies of C. dodecandra,” though the two are evidently closely related. On the other hand, P. dodecandra and the western P. trachysperma be- have like geographical subspecies, the former evidently a taxon of more recent evolutionary origins, restricted to the glaciated northeastern United States and southern Quebec, but intergrading with the latter in a broad belt from Mis- souri to Minnesota. The species is therefore composed of P. dodecandra (L.) DC. ssp. dodecandra and P.d. ssp. trachysperma (T. & G.) Iltis stat. nov. (— P. trachysperma T. & G. Fl. N. Am. 1: 669. 1840). HERBARIUM, DEPARTMENT OF BOTANY UNIVERSITY OF WISCONSIN, MADISON, WISCONSIN LITERATURE CITED BURMAN, J. 1737. Thesaurus Zeylanicus. Amsterdam. EnNsT, W. R. 1963. The genera of Capparaceae and Moringaceae in the Southeastern United States. Jour. Arnold Arb. 44: 81-95. Iuris, H. H. 1954. Studies in the Capparidaceae I. Polanisia dodecan- dra (L.) DC., the correct name for Polanisia graveolens Rafin- esque. Rhodora 56: 65-70. Pl. 1201. . 1958. Studies in the Capparidaceae IV. Polanisia Raf. Brittonia 10: 33-58. LINNAEUS, C. 1747. Flora Zeylanica. Amsterdam. . 1753. Species Plantarum. Holmiae. SAVAGE, S. 1945. A Catalogue of the Linnaean Herbarium. London. STEARN, W. T. 1957. Introduction, in Linnaeus's Species Plantarum. Facsimile of first edition. Ray Society 140: 1-176 (esp. p. 119). TRIMEN, H. 1887. Hermann's Ceylon Herbarium and Linnaeus's Flora Zeylanica. Jour. Linn, Soc. London, Bot. 24: 129-155. 5T am obligated to Dr. W. L. Brown, Jr., Museum of Comparative Zoology, Harvard University, for calling some of the pertinent rea- sons for this interpretation to my attention. POLANISIA DODECANDRA IN NEW HAMPSHIRE" Polanisia dodecandra (L.) D. C. of the Capparaceae, not previously recorded in New Hampshire, has been found growing in Durham, New Hampshire. In New England, this species was reported only from the Lake Champlain region of Vermont by Bean, Hill, and Eaton (Rhodora, 63:53, 1961). The records, however, are based essentially on specimens in the Harvard Herbaria. In Connecticut, Weatherby reported it from the banks of the Hockanum River in East Hartford, and Driggs, from the vicinity of the Connecticut River at Hartford. These reports are cited in The Catalogue of Flowering Plants and. Ferns of Con- necticut, State of Connecticut Geological and Natural His- tory Survey, Bull. 14:211, 1910. I have been unable to locate any specimens collected in Connecticut in the her- baria at the University of Vermont, University of Con- necticut, Yale University, and the Connecticut Botanical Society. I found the plants growing in crushed granitic rock of the B&M railroad bed about two hundred yards north of the old Durham depot, on September 4, 1965. The sandy soil beneath the rock was fairly moist despite a dry super- ficial appearance. On September 23, 1965, Dr. A. R. Hodgdon, Mr. Wendell Berry, Jr., and I examined the station more closely and estimated that betwen two and three thousand plants were growing in an area about three hundred feet long and thirty feet wide. The plants ranged in height from less than an inch to two feet. The collections from this station have been de- posited in the University of New Hampshire Herbarium and in the New England Botanical Club Herbarium at Harvard University, Cambridge, Massachusetts EDWARD J. HEHRE DEPT. OF BOTANY UNIVERSITY OF NEW HAMPSHIRE 'I wish to acknowledge the assistance of the following: Mr. Frank Seymour, Mr. Leopold Charette, Dr. George Torrey and Dr. James Neale. Their help in my search for herbarium Specimens is much appreciated. 48 CHROMOSOME NUMBERS IN VERNONIA (COMPOSITAE) S. B. JoNES, JR. AND W. H. DUNCAN The genus Vernonia is à member of the tribe Vernonieae of the Compositae, which is not as well known cytologically as the other tribes in the family. It is the largest genus in the tribe with about 1000 species of temperate and tropical America, Asia and Africa. The members of the genus are herbs or shrubs, and nearly all are perennial. The North American species of Vernonia were monographed by Gleason (1906) who also treated them in “North American Flora" in 1922. Chromosome numbers reported in Vernonia are summarized in Table 1. This list includes both Old and New World species. Gametic numbers of 9, 10, 16, 17, 18, 20, and 27 are indicated. The first known determination was in 1933 by G. W. Bohn, who counted n=18 in V. bald- winii (personal communication to the senior author, April, 1962). Grant (1953) suggested that the basic number for the genus may be 9 based on his record for V. cinerea, a spe- cies considered by Gleason to be one of the more primitive Vernonia. Additional work is needed, however, to establish Grant's hypothesis firmly. Hunter (1964), ina recent paper, has presented chromosome counts of n—1'7 for seven spe- cies and one hybrid. In connection with taxonomic studies in the genus, the chromosome numbers were obtained for 13 taxa and these are reported in this paper. Meiotic chromo- some counts were obtained from pollen mother cell squashes. Buds were collected in the field or from transplants, and killed and fixed in modified Carnoy's (4 parts chloroform : 3 parts: absolute alcohol: 1 part glacial acetic acid). All material was squashed and stained in acetocarmine. Voucher specimens were made and deposited in the University of Georgia Herbarium, Camera lucida drawings and photo- micrographs were used to record the observations which were made from fresh mounts. 49 50 Rhodora [Vol. 68 All plants studied had a gametic number of n=18. Differ- ences in chromosome size and morphology were not discern- ible. Meiotic figures appeared normal. This was true even in those individuals suspected of being of hybrid origin. Hunter (1964) reports a gametic number of n=17 or ca. 17 for three of the taxa reported in this paper. The reason for, or the significance of this variation is not known at this time. It is possible that both n—17 and n=18 exist within the populations and taxa of this genus. Hybridization frequently occurs in Vernonia (Gleason, 1906) and those that we have studied are no exception. Hy- bridization studies by Jones (1964a) demonstrated that all combinations of taxa attempted were cross-fertile. Certainly chromosome number would not be a limiting factor pre- venting hybridization. The taxa are usually ecologically or geographically isolated; however, these external barriers are not completely effective, producing much of the varia- tion we find in populations of Vernonia. Field cllected in- termediates, as well as artificial F, hybrids, appear to undergo normal meiosis during pollen formation. Since chromosome numbers are known for only about 30 of the possible 1000 species, it is obvious that additional cytotaxonomic work is needed in Vernonia before a realistic concept of cytological relationships in the genus may be established. Since the genus is widespread in temperate and tropical America, Asia, and Africa, geographic and taxo- nomic correlations of chromosome numbers might prove of interest. DEPARTMENT OF BIOLOGY, UNIVERSITY OF SOUTHERN MISSISSIPPI, HATTIESBURG DEPARTMENT OF BOTANY, UNIVERSITY OF GEORGIA, ATHENS, Table 1. Chromosome Numbers in Vernonia (Compositae). Taxon Chromosome Author(s) or Number Vouchers* V. acaulis (Walt.) Gleason n—17 Hunter 1964 7,—18 22394, Duncan, Elbert Co., Ga. —— ———— — . . *Vouchers collected by senior author unless otherwise noted. 1966] SS S NEXSAEN . angustifolia Michx. . angustifolia Michx. var. angustifolia . angustifolia Michx. var. mohrii S. B. Jones . angustifolia Michx. var. scaberrima (Nutt.) A. Gray . altissima Nutt. . anthelmintica (L.) Willd. . baldwinii Torr. . blodgettii Small . cinerea (L.) Less. . colorata Drake X concinna Gleason (pro sp.) . conferta Benth. . X dissimilis Gleason (pro sp.) . glabra Vatke greggii Gray . guadalupensis Heller . interior Small . karvinskiana DC. . kotschiana Sch. Bip. V V V. V V. guineensis Benth. V V V . lindheimeri Gray Vernonia — Jones & Duncan 51 n—ca.17, 17 Hunter 1964 n—18 n—=18 n—18 n—18 n—27 n—18 n—18 2n—18 2n—18 9, —18 2n—18 n—18 2u—20 n—=17 n—17 2n—20 1055, McDuffie Co., Ga.; 1089, Camden Co. Ga.; 1119, Tel- fair Co., Ga.; 1112, Lowndes Co, Ga.; 1116, Coffee Co., Ga.; 1130, Wilkinson Co., Ga. 1091, 1092, Duval Co., Fla.; 1059, Bibb Co., Ala.; 1060, 1061, Autauga Co., Ala.; 1054, Talbot Co., Ga.; 1096, Putnam Co., Fla.; 1102, Lake Co., Fla.; 1105, Alachua Co., Fla.; 1111, Lowndes Co., Ga. 1075, Emanuel Co., Ga.; 1122, Hancock Co., Ga; 1420, D. J. Banks, Claredon Co., S. C. 2595, Perry Co., Ala. Prameshwar 1960” Bohn 1933 1099, 1100, Charlotte Co., Fla. Mangenot et al. 1957”; Miege 1960"; Grant 1953 Mangenot et al. 1957” 1093, Putnam Co., Fla.; 1104, Lake Co., Fla. Mangenot 1957” 1063, Lee Co., Ala.; 1066, Harris Co., Ga. Riley and Hoff 1961” Hunter 1964 Hunter 1964 Mangenot 1958" n— ca.171, 17 Hunter 1964 n—ca.17 2n—18 Turner et al. 1962" Miege 1960” n—ca.17,17 Hunter 1964 V. marginata (Torr.) Raf. n=17 Hunter 1964 "Cited in “Index to Plant Chromosome Numbers," M. S. Cave, Editor. 52 Rhodora [Vol. 68 V. noveboracensis (L.) Michx. n=18 2604, Gwinett Co., Ga. V. nudiflora Less. n—=16 Covas and Hunziker 1954” V. ovalifolia T. & G. »—18 1095, Putnam Co., Fla.; 1104, Lake Co., Fla. V. perottetii Sch. Bip. 2n—18 Miege 1960” V. platensis Les. n—=20 Cabrera 1944 V. primulina O. Hoffm. n—20 Mangenot et al. 1958" V. pulchella Small n—18 1084, Tattnal Co., Ga.; 1085, Long Co., Ga. V. X recurva Gleason (pro sp.) n—18 1077, Emanuel Co., Ga.; 1083, Tattnal Co., Ga. V. texana (A. Gray) Small »—17 Hunter 1964 n=18 2292, Franklin Co. Miss.; 2293, Amite Co., Miss. LITERATURE CITED CABRERA, A. L. 1944. Vernoniae Argentinas (Compositae). Darwiniana 6: 265-379. CAVE, M. S. (Editor). Vol. I, 1956-1959. Index to plant chromosome numbers. Univ. of N. C. Press, Chapel Hill, N. C. Vol. II, 1960-62. Index to plant chromosome numbers. Univ. of N. C. Press. Chapel Hill, N. C. GLEASON, H. A. 1906. A revision of the North American Vernonieae. Bull. N. Y. Bot. Gard. 4: 144-243. . 1922. Vernonieae. North American Flora. N. Y. Bot. Gard. 33: 52-95. GRANT, W. F. 1953. A cytotaxonomie study in the genus Eupatorium. Am. J. Bot. 40: 729-749. HUNTER, G. E. 1964. Chromosome Numbers in Vernonia: Section Lepidoploa: Subsection Paniculatae Verae. Southwestern Natu- ralist, 9: (4): 289-244, JONES, S. B. 1964a. Taxonomy of the narrow-leaved Vernonia of the Southeastern United States. Ph. D. dissertation, Univ. of Georgia. 1964b. Taxonomy of the narrow-leaved Vernonia of the Southeastern United States. Rhodora, 66: 382-401. . In Press, Experimental hybridizations in Vernonia (Compositae). Brittonia. NATURAL HYBRIDS OF LADY'S SLIPPERS (CYPRIPEDIUM) IN MANITOBA H. H. MARSHALL, A. T. H. GROSS AND G. A. STEVENSON A small yellow lady's slipper, Cypripedium Calceolus L. var. parviflorum (Sal.) Fern., occurs frequently in low meadows in southern Manitoba, Canada, while C. candidum Muhl., a tiny white lady’s slipper, is more restricted and has been reported from only three Manitoba locations, Wood- lands, Brandon and Aweme (Scoggan, 1947). In 1954 an in- vestigation of cream colored lady's slippers found in associ- ation with C. Calceolus var. parviflorum and C. candidum, was conducted by the authors. From 1954 to 1964 annual surveys of three areas of 5 to 20 acres between Brandon and Aweme revealed similar lady’s slippers but profuse flower- ing was recorded only in 1954. These hybrids were similar to those described by Fuller (1932) and Curtis (1932) as C. X Andrewsii Fuller. Seedlings of C. Calceolus var. parviflorum are influenced greatly by local environmental conditions (Curtis, 1943). He reported that plants grown from seed may bloom in 9 years under favorable conditions but may take 15 years to bloom and C. candidum probably requires 12 or more years to bloom. Plants of both species are long-lived. Lady’s slippers in nature are pollinated by bees of the genus Andrena (Darwin 1890). The sticky pollen does not permit wind pollination and the location of anthers and stigma precludes self pollination. Hybrid lady’s slippers can therefore occur naturally if suitable pollinators are present and cross-compatible species occur in the same area and have overlapping periods of bloom. Fuller (1932) states that it is very likely wherever C. candidum and C. Calceolus var. parviflorum grow in the same vicinity, hy- brids will be found. Stebbins (1957) describes a method of indexing plant characteristics to identify hybrid populations. Each trait 53 54 Rhodora [Vol. 68 is indexed on a scale of from 1 to 5 although the range may be either reduced or extended. Plants with the lowest total for indexed characteristics are considered of the one pa- rental type and those with the highest total for indexed characteristics are of the other parental type. This paper reports characteristics of natural hybrids of C. candidum X C. Calceolus var. parviflorum in Manitoba and presents an hypothesis to explain the origin of the hybrids. In June 1954, 55 specimens of Cypripedium were collected from a low lying meadow in section 11 township 10 range 18 west of the principal meridian southeast’ of Brandon, Manitoba. Measurements or descriptions of 10 character- istics,, i.e. height, leaf number, length and width of the largest leaf, length and width of lateral petals, color and length of lip, and length and width of the upper sepal, were recorded. Plant characteristics were indexed according to the method described by Stebbins (1957) to identify hybrid populations. The smallest measurement, fewest number and least intensity of color Was assigned the lowest index number. The specimen plants with the lowest total for indexed characteristics are most similar to the C. candidum parent and those with the highest totals are most similar to the C. Calceolus var. parviflorum. parent. In 1963, controlled crosses were made between C. candi- dum and C. Caleeolus var. parviflorum to establish whether the purported parents were cross-compatible and could give rise to hybrids. Viable hybrid seed was readily produced by hand polli- nation of C. candidum and C. Calceolus var. parviflorum thus indicating that they are cross-compatible and could have given rise to hybrids. Native C. candidum commences bloom two weeks earlier than C. Calceolus var. parviflorum at Brandon, Manitoba, hence, their bloom periods seldom coincide and rarely overlap for more than a few days. This is at variance with Fuller (1932) who stated that they bloom at the same time. The bloom period of C. X Andrewsii commences 10 days after C. candidum, thus it overlaps the bloom period of C. Calceolus var. parviflorum to a greater 55 Table 1. Plant measurements (cm) of the two parental species, C. candidum and C. Calceolus var. parviflorum. 1966] Lady’s Slippers — Marshall, Gross & Stevenson Plant C. candidum C. Calceolus var. parviflorum Characteristic E 2. 1. 2. Plant height 15 - 40 14 - 22 20 - 70 17.2 - 32.5 Leaf-length - 9.5 - 12.5 11 - 20 12 - 15.5 -width 1-4 2.0- 3.5 4-12 3.8- 6.7 Sepal-length 2-3 21- 2.8 4-7 3.8- 4.8 -color Petal-length -color Lip-length -color greenish spotted with purple 2.5 -3.5 greenish spotted with purple 1.8 - 2.5 waxy white, purple veined green with purple streaks 2.2- 3.2 green with purple streaks 1.6- 2.0 white, purple veins greenish yellow streaked purple 5-9 3-5 golden yellow greenish yellow streaked purple 4.8- 6.1 2.9- 3.5 yellow 1. Data taken from Seoggan (1957) and Fernald (1950. 2. Brandon data. 56 Rhodora [Vol. 68 extent than that of C. candidum. Bloom appears to be influ- enced greatly by local environmental conditions as indicated by Curtis (1943), Only in 1954 was profuse flowering observed. In 1964 late spring frosts on May 30 (25°F), May 21 (27°F) and June 1 (26°F) destroyed flower buds and bloom was observed in only the most sheltered places. Spring frosts of 27°F or lower were recorded after May 20 in 1958, 1959, 1961, and 1963 which could account for the restricted amount of flowering observed in those years. Measurements of C. candidum and C. Calceolus var. parviflorum (author’s collection No.’s 884 and 883) are similar to those published by Scoggan (1957) and Fernald (1950). The C. candidum plants were smaller than the C. Calceolus var. parviflorum specimens in all 6 of the measure- ments recorded (Table 1). Indices for the naturally occurring Cypripedia indicated 8 plants resembled C. candidum, 8 resembled C. Calceolus var. parviflorum and 39 were intermediate between these two species and resembled C. X Andrewsii (Table 2). This classification was corroborated by the identifications re- ported by Boivin (1960). The index totals for the 8 C. candidum plants ranged from 10 to 14 and did not overlap with the 39 C. X Andrewsii specimens which ranged from 19 to 38. The 8 C. Calceolus var. parviflorum specimens index totals ranging from 30-47 formed a continuous series with C. X Andrewsii. The parental species were not readily distinguishable on the basis of leaf number and width of lateral petals, hence, these two characteristics were of no value in indexing the specimens. Lip color of the hybrids ranged from white of the C. candidum parent to yellow of the C. Calceolus var. parviflorum parent with many creamy intermediate types. The seven other plant characteristics revealed a complete overlap of the parental species by the hybrids. The hybrids were readily separated from the C. candidum. plants using either the indexing method of Stebbins (1957) or the stand- ard taxonomic bases of specimen identification, but they were less readily distinguished from the C. Calceolus var. 1966] Lady's Slippers — Marshall, Gross & Stevenson 57 parviflorum parent. This is in agreement with Stebbins (1957) that hybrids tend to resemble one parent more than another. It is postulated that the hybrids are of relatively recent origin. The largest plants are the parental types and the hybrids are either single stemmed or small clumps. The slow growth and infrequency of bloom of Cypripedia in- dicates that the hybrid swarms could have developed since 1900. This would explain why J. H. Macoun and H. Criddle (Scoggan 1947), who observed the parental species in the Brandon — Aweme district did not report hybrid swarms of creamy colored Cypripedium. These hybrid swarms probably developed as a result of a change in the environment in which the parent species grew. Hybrid swarms usually appear in a disturbed habitat but this is not possible with Cypripedia as they are never found in cultivated land, rarely in pastured areas, and are confined to areas that have been undisturbed. It is suggested Table 2. Frequency of plant measurements of C. candidum, C. Calceolus var. parviflorum and C. X Andrewsti indexed according to Stebbins (1957). l C. candidum C. X Andrewsii C. Calceolus var. parviflorum ^. Index No. > 7 mcd ess d n Plant characteristic frequency frequency frequency Plant height CERE RU b 812' 9 5b 0.0.1 93 4 Largest leaf-length bun2 un go 0. NT 050 2 2 4 -width (ed Ub t & 8 7 15 b 00 0 8 5 No. of leaves’. Si. i. 727 5 — — 0 6 2 — — Upper sepal-length 7 1 0 9 11-10-81 0290770::35* 0 -width 8 0 0 6 6 14 10 3 000 2 6 Lateral petals-length 7 1 0 2 13-14 8 4 QOO 1-1 -width* 3 5 0 5 17 16 1 — oe 0 2 4 — Lip-length 8 0 0 312 7 8 9 0008 5 -color 8 0 0 3 6 9 3 18 0.0008 Range of totals 10 to 14 19 to 38 39 to 47 Plant average 12.6 29.1 42.4 Indexed on a 1 to 3 scale. ?Indexed on a 1 to 4 scale. 58 Rhodora [Vol. 68 that although native Andrena bees were the probable natural pollinators, the introduction of the honeybee made the hy- brid swarm possible. The introduction of honeybees in the early 1900's coincides with the estimated time when the hybrids originated and they could have been the insect vector which brought about the development of hybrids. CANADA DEPARTMENT OF AGRICULTURE EXPERIMENTAL FARM, BRANDON, MANITOBA REFERENCES BorviN, B. 1960. Centurie de plantes Canadiennes III. Naturaliste Canadien, 87: 25-49. Curtis, J. T. 1932. A new Cypripedium hybrid. Rhodora. 34: 239- 242. Curtis, J. T. 1943. Germination and seedling development in five species of Cypripedium L. Am. J. Botany, 30: 199-206. DARWIN, C. 1890. 2nd ed. The various contrivances by which Orchids are fertilized by insects. John Murray, Albemarle Street, London, England. FERNALD, M. L. 1950. 8th ed. Gray’s Manual of Botany. American Book Co., New York. FULLER, A. M. 1932. A natural Cypripedium hybrid from Wisconsin. Rhodora 34: 97-101. GARAY, L. A. 1953. A natural Cypripedium hybrid from Ontario. Can. J. Bot. 31: 660-662, SCOGGAN, H. J. 1957. Flora of Manitoba. Canada Department of Northern Affairs and National Resources. National Museum of Canada. STEBBINS, G. L. JR. 1957. Variation and evolution in plants. Colum- bia University Press. New York. A STUDY OF VARIATION IN THE CYTOTYPES OF DRYOPTERIS SPINULOSA. ROLLA TRYON! AND DONALD M. BRITTON?” Among the groups of ferns in which cytological and evo- lutionary studies are being pursued most vigorously is the genus Dryopteris (sens. strict. of North America and Europe. In eastern North America (see Bibliography) thirteen fertile cytotypes have been recognized, and among these six are diploid (n—41), six are tetraploid and one is hexaploid. Sixteen sterile hybrids of these fertile cytotypes have been examined cytologically and four of them are di- ploid, nine are triploid, two are tetraploid and one is penta- ploid. These previous studies have been concentrated on chromosome counts of the taxa and hybrids to determine the level of ploidy and on chromosome pairing analyses of natural and synthetic hybrids as evidence of their genome relations. There have been no extensive cytological surveys of any of the cytotypes nor analyses of their variation. It was against this background, and our belief that cytological data can be fully exploited in evolutionary studies only when accompanied by data on morphological variation, that the present work was initiated. The data that we have obtained on variation of the cytotypes of Dryopteris spinulosa is pertinent to the problem of the origin of the tetraploid var. spinulosa and to the problems of the evolutionary potential of triploids and the variation of derived hexaploids. Dryopteris spinulosa var. intermedia (2 X) and var. spin- ulosa (4X) were chosen as especially suitable taxa for investigation because of their abundance and because of the abundance of their sterile triploid hybrid. These taxa pre- sented an opportunity to study and compare variation, in a related diploid and tetraploid and their hybrid, on a broad "Gray Herbarium, Harvard University. "Department of Botany, University of Guelph. This study was sup- ported, in part, by grants from Sigma Xi-RESA to D. M. Britton. 59 60 Rhodora [Vol. 68 basis. Dryopteris spinulosa is a rather common Species in northeastern United States and adjacent Canada where it usually grows on wooded hillsides, or in wooded swamps and less often in shrubby swamps, or along damp roadsides. Variety spinulosa (figs. 9, 10) grows more commonly in wet, often water-logged sites, while var. intermedia (figs. 5, A 3 PLATE 1320 Figs. 1-4. Illustrations of characters c to f. Fig. 1 — character c, basal pinnae of intermedia, each inferior basal pinnule, A and B, is shorter than the adjacent one, C and D. Fig. 2 — character d, basal pinnae of the hybrid, five basal tertiary segments, A to E, on the acroscopic side of the inferior pinnules, are parallel to the pinna-rachis. Fig. 3 — character e, basal pinnae of the hybrid, three of the inferior pinnules, A to C, are longer than the basal superior pinnule, D. Fig. 4—character f, central pinnae of intermedia, on each of the four pinnae, the basal inferior pinnule, A to D, overlies the rachis, E. 1966] Dryopteris spinulosa — Tryon & Britton 61 PLATE 1321 Figs. 5-6. Lamina of Dryopteris spinulosa var. intermedia, diploid, x 14. Fig. 5— from Plant 110. Fig. 6 — from Plant 357. o bo Rhodora [ Vol. AAAS M > AYA PLATE 1322 Figs. 7-8. Lamina of Dryopteris spinulosa var. intermedia, X var. spinulosa, triploid, X 4. Fig. 7 — from Plant 487. Fig. 8 — from Plant 268. 6) is most often in moist but well drained sites, but both exhibit considerable latitude in their habitat. The two varieties frequently grow intermixed or in close proximity to each other. In our experience, their hybrid (figs. 7,8) is found whenever the two parents are in some abundance (and sometimes when one is rare) and it is usually frequent or even more abundant than either parent. 1966] Dryopteris spinulosa — Tryon & Britton 63 PLATE 1323 Figs. 9-10. Lamina of Dryopteris spinulosa var. spinulosa, tetraploid, x 14. Fig. 9— from Plant 272. Fig. 10— from Plant 294. The classifieation and nomenclature of this group Is sub- ject to considerable uncertainty and disagreement. While this paper does not intend to deal with either of these matters in a definitive manner, we do present some reasons later for the classification adopted. It is well established that var. spinulosa and var. intermedia are taxa (major evo- lutionary units) and disagreements about their taxonomic rank is of secondary importance. Some of the difficulties involved in the correct nomenclature are so involved and technical that we wish to do no more than mention their 64 Rhodora [Vol. 68 existence, The following synonymy will serve to orient the reader with other names currently used for these plants. Dryopteris spinulosa (O. F. Muell.) Watt, var. spinulosa (Dryopteris spinulosa; Dryopteris spinulosa, “typical”; Dryopteris austriaca (Jaeq.) Woynar, var. spinulosa (Muell.) Fiori). In the remainder of the paper this will be referred to as spinulosa. Dryopteris spinulosa var. intermedia (Willd.) Underw. (Dryopteris intermedia (Willd.) A. Gray; Dryopteris austriaca var. intermedia (Willd.) Morton). In the remainder of the paper this will be referred to as intermedia. Dryopteris spinulosa var. intermedia X var. spinulosa (Dryopteris intermedia X spinulosa; Dryopteris spinulosa var. fructuosa (Gilbert) Trudell; Dryopteris austriaca var. fructuosa (Gilbert) Morton; Dry- opteris X triploidea Wherry). In the remainder of the paper this will be referred to as the hybrid. This study was initiated in 1959 and has continued through June of 1965. Rolla Tryon has made the analyses of the characters, Donald M. Britton the cytological determi- nations and the collections of specimens; both have con- tributed to the discussions and conclusions. MATERIALS AND METHODS OF STUDY. Most of the plants used in this study (155) were obtained from southern Ontario (west of 799 W. Long. and south of 46? N. Lat.). Twenty-five others were from the Algoma District of Ontario (east of Lake Superior) and eight were from Pennsylvania and Virginia. The material for cyto- logical study was usually collected in late May to the middle of June. At that time a voucher leaf was taken from the plant and prepared as an herbarium Specimen. In many cases, the plant also was marked with the collection number and additional leaves were collected from some of these plants in September of the same year and also in the follow- ing year. Individual plants, therefore, have been repre- sented by one to sixteen leaves and often by sets of leaves collected in successive years. Eighty-nine plants of inter- media were determined as diploid ( n—41); forty-seven plants of the hybrid. were determined as triploid (n—123) ; 2 and fifty-two plants of spinulosa were determined as tetra- ploid (n=82). Voucher specimens of these cytologically 1966] Dryopteris spinulosa — Tryon & Britton 65 determined plants have been deposited in the Herbarium, Department of Botany, University of Guelph and a selection of them in the Gray Herbarium, Harvard University. Not all of the 188 plants and 746 leaves available have been used in this study. Some leaves were defective and others were too small (less than 25 em. long) to be included in case they would bias the results. About 175 plants and 550 leaves were available for most characters; the number studied for each is given as part of Charts 1 and 2. Thirty-four characters were considered during the course of our work. Eleven of these were studied in some detail and the results of these analyses are presented. Preliminary studies were made of eighteen other characters but these were not continued because the characters were either too variable to be of particular value or because no significant differences between spinulosa and intermedia were observed. Among these, a few of the variable characters were the following: complexity of the lamina, shape of the pinnae, tapering of the apex of the pinnae and size of the indusium. The variation of four other characters was not analysed principally because of technical difficulties in their study. An initial survey of these indicated that they probably be- haved in the same manner as some of the characters ana- lyzed in detail. These characters were the orientation of the rhizome, the color of the petiole scales, the evergreen quality of the leaves and the kind of habitat. Finally, the apex of the spines of the spores was not studied simply because of the small size of the character, the spines them- selves being only about 1 micron long. The illustrations have been prepared from a selection of specimens of the cytologically determined plants. These specimens were photographed and the negative printed as a silhouette. The specimens were relaxed in warm water, prior to photographing, so that they were substantially restored to their fresh condition. This is of some importance because minor shrinking, wrinkling and curling of the teeth and ultimate lobes can alter the total aspect of a pinna or lamina to a considerable degree. 66 Rhodora [Vol. 68 ANALYSIS OF CHARACTERS, The following comments on the individual characters ex- plain the character, when necessary, and the manner in which it was studied. The original measurements or counts for six of the characters are presented in Charts 1 and 2 and in Graph 1. In order to compare the range of variation of the different characters and their means, this data was ad- justed to a scale of 0 to 100 for each character (the minimum count or measurement—0, the maximum- 100). For ex- ample, character e varies from a minimum count of 1 to a maximum of 8. Then 1—0 and 8—100 and counts of 2 to 7 —14, 28, 43, etc., respectively. For characters b and ec the score was then reversed (0—100, 100—0) so that the scores for all of the characters would be high for intermedia and low for spinulosa. These adjusted scores are presented in Chart 3 and in Graph 2. Selections of the data are presented in Charts 4 and 5 and in Graph 3 to illustrate special features of the variation of the cytotypes. a. Glandularity of the base of the pinna. The lamina of intermedia is glandular while that of spin- ulosa is usually glabrous. The glands are small, stalked and have a more or less Spherical cap cell. They are usually most abundant at the base of the pinnae. There is some tendency for them to be deciduous as the leaf ages, although they will persist on leaves that have lived over the winter. In order to obtain a measurement of this character, the cytological voucher leaves (collected in early summer) were used and a single central pinna was chosen for observation. The glands that could be seen on the lower surface on one linear millimeter of the pinna stalk were counted. The data obtained from sixty plants are presented in Chart 1. b. Length of the guard cells of the stomata. This character was studied only in a preliminary manner although the results indicate that extended data would be of much interest. A portion of a leaf collected in late summer was relaxed in warm water and then placed in a lactic acid mount. Measurements were made, in micra, of the maxi- 1966] Dryopteris spinulosa — Tryon & Britton 67 mum length of the guard cells from stomata between the veins on the lower surface and the mean from ten measure- ments was obtained for each leaf. The data obtained from 27 plants are presented in Chart 1. c. Length of the inferior basal pinnules on the basal pin- nae in relation to the adjacent ones. Each basal pinna of intermedia usually has the inferior basal pinnule shorter than the adjacent one (fig. 1), while spinulosa usually has the inferior basal pinnule longer than the adjacent one (fig. 16). CHART 1. Analysis of variation in Dryopteris spinulosa, original counts and measurements of characters a and b. (See Graph 1.) Range of variation of Mean of all leaves. all leaves. (Total number of leaves.) a. Glandularity of the base of the pinna: number of glands on one linear mm. of pinna-stalk, each leaf from a different plant. intermedia 10-75 45.6 (20) hybrid 9-55 29.2 (20) spinulosa 0-9 1.6 (20) b. Length of the guard cells of the stomata: mean of ten measure- ments for each leaf, each leaf from a different plant. intermedia 34-53 micra 42.5 micra (9) hybrid 40-58 micra 49.5 micra (9) spinulosa 44-68 micra 55.7 micra (9) The character was measured by taking the average of the length of both basal pinnules and both adjacent ones because the relative length of these two pinnules may differ on each basal pinna. The measurements are in millimeters and the average is expressed as the basal pinnules being longer, or shorter, than the adjacent ones. The data ob- tained from 546 leaves are presented in Chart 2; illus- trations of the variation are given in figs. 11-16. 68 Rhodora [Vol. 68 16 PLATE 1324 Figs. 11-16. Basal pinnae, X 1/3. Figs, 11-12. Dryopteris spinulosa var. intermedia, diploid. Fig. 11 — from Plant 28, the inferior basal pinnules are 15 mm. shorter than the adjacent ones. Fig. 12 — from Plant 110, the inferior basal pinnules are 7 mm. longer than the adjacent ones. Figs. 13-14. Dryopteris spinulosa var. intermedia x var. spinulosa, triploid. Fig. 13 — from Plant 38, the inferior basal pinnules are 7 mm. shorter than the adjacent ones. Fig. 14 — from Plant 502, the inferior basal pinnules are 10 mm. longer than the adjacent ones. Figs. 15-16. Dryopteris spinulosa var. spinulosa, tetraploid. Fig. 15 — from Plant 68, the inferior basal pinnules are equal in length to the adjacent one. Fig. 16 — from Plant 36, the inferior basal pinnules are 15 mm. longer than the adjacent ones. d. Number of parallel tertiary segments. In intermedia there is a Strong tendency to have many of the basal tertiary segments, on the acroscopic side of 1966] Dryopteris spinulosa — Tryon & Britton 69 the pinnules, disposed more or less parallel to the pinna- rachis. This character is illustrated (by the hybrid) in fig. 2. In spinulosa (fig. 15) there is a strong tendency to have rather few such segments parallel to the pinna rachis. This character has been counted on one arbitrarily chosen basal pinna and on the pinnules on its inferior side, as illustrated in fig. 2. The data obtained from 565 leaves are presented in Chart 2. This character may be correlated with the size of the pinna and for this reason small leaves were not used in its study. In leaves of moderate or large size there is no abso- lute correlation with the size of the pinna as the following counts wil show. In a plant of spinulosa, a basal pinna of one leaf was 9.5 cm. long and had 1 parallel tertiary segment on its inferior side. A basal pinna, from another leaf of the same plant, was g cm. long and had 2 parallel tertiary segments on its inferior side. e. Number of longer inferior pinnules. The pinnae in spinulosa tend to be relatively broad at the base and to be abruptly reduced in width. In intermedia there is a tendency for the pinnae to be hardly, or not at all, broadest at the base and to be gradually reduced in width. These tendencies have been reduced to an easily observed character by using one basal pinna and comparing the length of the inferior pinnules with the basal superior one. The character was taken from the basal pinna with the shortest basal superior pinnule and, in reference to this pinnule, the number of inferior pinnules that were longer was de- termined. The data obtained from 546 leaves are presented in Chart 2; and the character is illustrated (by the hybrid) in fig. 3. As in the study of the previous character, small leaves were not used because of the possibility that small pinnae would bias the data. In leaves of moderate to large size, there is no absolute correlation with the size of the pinna as the following counts will show. A basal pinna of one leaf of a plant of intermedia was 11.5 cm. long and had 5 70 Rhodora [Vol. 68 longer inferior pinnules. A basal pinna from another leaf of the same plant was 10 cm. long and had 6 longer inferior pinnules. CHART 2. Analysis of variation in Dryopteris spinulosa, original counts and measurements of characters ¢ to f. (See Graph 1.) Range of variation Mean of all leaves. All leaves. Plant means from (Total number of leaves.) plants with 4 or more leaves. (Total number of plants: total number of leaves.) c. Length of the inferior basal pinnules of the basal pinnae in relation to the adjacent ones. intermedia 16 mm. shorter to 10 mm. shorter to 4.5 mm, " mm. longer. 2 mm. longer. Shorter (255) (31: 172) hybrid 7 mm. shorter to 1.5 mm. Shorter to 1.7 mm. 11 mm. longer. 5 mm. longer. longer (119) (16: 87) spinulosa equal to 16 mm. 3.7 mm. longer to 6.4 mm. longer. 9 mm. longer longer (108) (11:60) d. Number of parallel tertiary segments. intermedia, 3-10 4.0-6.8 5.7 (296) (19: 107) hybrid 1-8 2.5-4.7 3.4 (137) (18: 95) spinulosa 1-5 1.6-3.5 2.5 (132) (14: 82) e. Number of longer inferior pinnules. intermedia 2-8 3.4-6.2 4.8 (278) (34: 212) hybrid 2-5 2.0-3.6 2.9 (135) (17: 96) spinulosa 1-4 2.0-3.1 2.1 (133) (14: 84) f. Number of pinnules that overlie the rachis. intermedia, 0-10 1*-2.9-10 7.8 (299) (30: 162) hybrid 0-7 0-3.0 1.0 (155) (20: 107) spinulosa 0-3 0-0.75 0.07 (138) (17: 97) *Plant 357 with two leaves. 1966] Dryopteris spinulosa — Tryon & Britton 71 f. Number of pinnules that overlie the rachis. In intermedia, the pinna, the basal inferior pinnule and its basal segment nearest the rachis are disposed so that the basal segment is relatively close to the rachis and most often so that it overlies it (figs. 4 and 5.). In spinulosa these same structures are disposed so that the basal segment is relative- ly far from the rachis (figs. 9 and 10). This character was analyzed by the number of basal inferior pinnules that overlie the rachis on the lowest ten pinnae. Fig. 5 shows a lamina with 10 overlying pinnules and figs. 9 and 10 show laminae with no overlying pinnules. Only 10 pin- nae were used in order to avoid bias due to high counts in unusually large leaves. Bias due to low counts in small leaves was eliminated (or reduced) by taking only those leaves with a lamina at least 30 cm. long. In order to confirm that this character was not related to changes in orientation due to pressing of the leaf, counts were made on the same leaves when they were fresh and after they were pressed. No difference was observed. The data obtained from 592 leaves are presented in Chart 2. g. Position of the pinnae in relation to the rachis. The pinnae may be patent (more or less at right angles to the rachis) or they may vary to ascending (at an acute angle to the rachis). This character was originally studied because it has been one that has been prominently used for the identity of the taxa, and especially of the hybrid. How- ever, so much variation was soon observed in both inter- media and spinulosa that the character was neither studied further nor refined. Figures 17 and 18 of intermedia and figs. 31 and 32 of spinulosa illustrate some of the variation observed in the pinna-angle. Ninety-seven leaves of inter- media had patent pinnae, 55 leaves had them slightly ascend- ing and 23 had them definitely ascending. The same range of variation, but in somewhat different proportions, was seen in spinulosa in which 13 leaves had patent pinnae, 47 leaves had slightly ascending pinnae and 23 leaves had definitely ascending pinnae. The character was also studied on plants from which several leaves were available but these 72 Rhodora [Vol. 68 PLATE 1325 Figs. 17-23. Central pinnae of Dryopteris spinulosa var. intermedia, diploid, X 1/3. Fig. 17 —from Plant 105. Fig. 18 — from Plant 357. Fig. 19 — from Plant 335. Fig. 20 —from Plant 108. Fig. 21 — from Plant 29. Fig. 22 — from Plant 104. Fig. 23 — from Plant 111. 1966] Dryopteris spinulosa — Tryon & Britton 73 results confirmed the previous data, i.e., although there was a slight statistical difference between intermedia and spin- ulosa it was not a sufficient one to invite further analysis. E LO pis 1 9 C " `. ^ Nd B S 24 È 27 PLATE 1326 Figs. 24-29. Central pinnae of Dryopteris spinulosa var. intermedia X var. spinulosa, triploid, X 1/3. Fig. 24 — from Plant 86. Fig. 25 — from Plant 84. Fig. 26 — from Plant 562. Fig. 27 — from Plant 87. Fig. 28 — from Plant 358. Fig. 29 — from Plant 32. 74 Rhodora [Vol. 68 30 33 35 PLATE 1327 Figs. 30-35. Central pinnae of Dryopteris spinulosa var. spinulosa, tetraploid, X 1/3. Fig. 30 — from Plant 33. Fig. 31 — from Plant 370. Fig. 32 — from Plant 79. Fig 33 — from Plant 305. Fig. 34 — from Plant 488. Fig. 35 — from Plant 289, h. Aspect of the lamina. The aspect of the lamina is usually different in spinulosa and intermedia. This may be readily observed in fig. 5 of intermedia and fig. 9 of spinulosa which are characteristic of each taxon. One of the initial objectives of this study was to analyze the general character of aspect into its individual components. This objective was partially but not wholly 1966] Dryopteris spinulosa — Tryon & Britton 15 realized. Characters c to g are individual characters of as- pect and our analyses of these, therefore, also represent an analysis of aspect. Other individual characters have not been analyzed and among these may be mentioned shape, symmetry, spacing and complexity of the pinnae, of the pinnules and of the tertiary segments, and the length, shape and orientation of the marginal teeth. Figures 17 to 35 illustrate the central pinnae of leaves that were selected to present the variation in aspect of the three cytotypes. Common aspects of intermedia, are illus- trated by figs. 21 and 23, of the hybrid by figs. 25 and 27, and of spinulosa by figs. 31 and 35. The other figures of 17-35 represent less common variations. The greatest resemblance of intermedia. to spinulosa. is seen in. figs. 20 and 34. The aspect of the hybrid. seems to intergrade with that of both of its parents. Figure 17 of intermedia is close to fig. 29 of the hybrid and fig. 19 of intermedia is close (except in the orientation of the pinnae) to fig. 24 of the hybrid. The aspect of the hybrid and spinulosa is similar in fig. 27 of the hybrid and fig. 33 of spinulosa, and in fig. 26 of the hybrid and fig. 34 of spinulosa. i. Glandularity of the indusium. Glands of the same type as those on the lamina also occur on the indusium of intermedia and the hybrid. The indusium of spinulosa is glabrous. Eighty-one per cent of the 89 plants of intermedia had the indusia definitely glandular and nineteen per cent had them slightly glanduiar; eighty-five per cent of the 47 plants of the hybrid. had the indusia defi- nitely glandular and fifteen per cent had them slightly glandular. These observations were made only on the cyto- logical voucher leaves, collected in early summer, since the glands tend to be deciduous with age. However, in both intermedia and the hybrid, glands were also observed on the indusia of leaves that had lived over the winter. j. Spores. Our observations of the spores of numerous plants ot intermedia and spinulosa showed them to be normal in ap- 16 Rhodora [Vol. 68 pearance and presumably viable. Only rarely was an abor- tive spore seen. The spores of the hybrid, on the other hand, were uniformly shriveled (abortive). No normal appearing spores were seen among those of the hybrid al- though some effort was made to find them. k. Chromosome number. The chromosome counts, already mentioned, may be sum- marized here for the sake of a complete list of the characters studied, intermedia: diploid, n = 41 (89 plants). hybrid: triploid, n = 123 (47 plants). 2 spinulosa: tetraploid, n = 82 (52 plants). DISCUSSION. Variation in the cytotypes. — In relation to the numerical as well as the geographical limitations of our sample, it is desirable that some assess- ment be made of the parameters of the variation that have been established in our study. An effort was made to obtain a cytological analysis of as many variants as possible and we believe that most of the variation was obtained in the areas sampled. Nevertheless, it is patent from a study of materials in the Gray Herbarium and the Herbarium of the New England Botanical Club that there is greater vari- ability in the Dryopteris spinulosa group than in our cyto- logically determined samples of it. For this reason we are reluctant to place reliance on our parameters as those of the taxa and their hybrid. We feel that most of the param- eters can be extended by further cytological sampling, especially in other regions. The means of the characters, however, and the mean of the means, can probably be ex- trapolated with some confidence as representative of the taxa and the hybrid because of the inertia of the mean. The range of variation of the original counts and measure- ments is presented in Graph 1. It is portrayed more clearly in Graph 2 where the original data has been adjusted for comparative purposes. Considering the data from all leaves, 1966] Dryopteris spinulosa — Tryon & Britton 77 CHART 3. Analysis of variation in Dryopteris spinulosa, adjusted scores (see text) of characters a to f. (See Graph 2.) Character. a to f *Plant 357 with two leaves. intermedia, hybrid spinulosa intermedia hybrid spinulosa intermedia hybrid spinulosa intermedia hybrid spinulosa intermedia hybrid spinulosa intermedia hybrid spinulosa intermedia hybrid spinulosa Range of variation. All leaves. 13-100 12-62 0-12 47-100 29-82 0-71 28-100 16-72 0-50 22-100 0-77 0-44 14-100 14-57 0-43 0-100 0-70 0-30 Mean range of variation. 21-100 12-70 0-42 Mean of plants with 4 or more leaves. 44-81 34-55 22-38 33-64 17-41 7-28 34-74 14-37 14-30 10*-29-100 0-30 0-7.5 Mean of all leaves. Mean of means. 64 94 Lf it will be seen in Graph 2 that the range of variation (mini- mum to maximum score) of each cytotype overlaps that of the others in characters b to f. In character a, the range of variation of intermedia and the hybrid overlap while that of spinulosa is contiguous, or nearly so, with. the range of those two. The range of variation of the mean of plants with four or more leaves is presented for characters c to f. In 78 Rhodora [Vol. 68 o co 30 20 10 -10 -20 eG — O Oo Qo 4 GRAPH 1. Analysis of variation in Dryopteris spinulosa, original counts and measurements of characters a to f, from Charts 1 and 2. White dot: mean of all leaves. Bars indicate range and extent of variation of a character. Solid bar: all leaves. Outline bar: plant means from plants with four or more leaves (extension by dash of character f in intermedia includes Plant 357 with only two leaves). I: intermedia, H: hybrid, S: spinulosa. 1966] Dryopteris spinulosa — Tryon & Britton 79 X [^4] Uu -— M [ 2] C) = - C) (0) le o T € = = era BS oO me} OQ "TO o GRAPH 2. Analysis of variation in Dryopteris spinulosa, adjusted scores of characters a to f, from Chart 3. White dot: mean of all leaves. Bars indicate range and extent of variation of a character. Solid bar: all leaves. Outline bar: plant means from plants with four or more leaves (extension by dashes of character f in intermedia includes Plant 357 with only two leaves). I: intermedia, H: hybrid, S: spixulosa. 80 Rhodora [Vol. 68 the means of plants the range of variation of spinulosa and intermedia is discrete in each character, while that of the hybrid. overlaps the variation of both parents in each of them. The extent of variation (maximum score minus the minimum) of the means of plants is about half that of the extent of the variation of single leaves. The mean of each character is strikingly different in spinulosa and intermedia. The mean of the hybrid falls between those of the parents. In characters a to c it is more or less midway between the parental means, while in charac- ters d to f it is closer to the mean of spinulosa. The mean of the means of the six characters demonstrates that, in these, the hybrid falls somewhat closer to spinulosa than to inter- media. We do not wish to imply, however, that this may represent the actual genetic position of the hybrid for in other characters it is closer to intermedia. It will be noted also in Graph 2 that in most cases the mean of a character is more or less centrally located between the extremes, while in a few cases it is notably close to one extreme. This is especially true of character a in spinulosa and of character f in all of the cytotypes. The genetic factors which might control these characters would be worthwhile to investigate. At the moment nothing is known of the inheritance of any character in Dryopteris. Information concerning numbers of pairs of genes, dominance relation- ships and factor interaction is lacking. It can be seen in Graph 2 that the extent of variation in a character is usually greater in intermedia than in spin- ulosa. In characters a, e and f the extent of variation in intermedia is twice as great, or more, than in spinulosa. In characters ¢ and d it is about half again as great in inter- media than in spinulosa. Only in character b is the extent of variation greater in spinulosa than in intermedia. In each of these characters the extent of variation in the hybrid is equal to or greater than that in the less variable parent and smaller than that in the more variable parent. This com- parison of variation is summarized in Graph 2 by the bars that indicate the mean range (and extent) of variation of 1966] Dryopteris spinulosa — Tryon & Britton 81 CHART 4. Analysis of variation in Dryopteris spinulosa. Range and extent of variation between extreme leaves and plants in the means of adjusted scores (see text) of characters a and c to f. (See Graph 3.) Character and score. Mean of Range and scores. (extent) a c d e T of variation of mean. Leaf from inter- Plant..viv 13 48 33 57 40 40 media Leaf from 40-79 (39) Plant 104 66 75 78 86 90 19 Leaf from Plant 126 19 . 99 338 144 0 16 hybrid Leaf from 16-47 (31) Plant 95 44 471 18 B: Ge Leaf from spinu- | Plant 91 0 8 22 14 0 9 losa Leaf from 9-24 (15) Plant 370 6 84 33 48." I0 724 Character and mean of Mean of plant means. a c d e f Plant 101 inter- 4 leaves 13 52 33 64 57 44 media Plant 113 44-71 (27) 4 leaves 55 80 56 74 92 "1 Plant 284 6 leaves 12 38 30 29 3 22 hybrid Plant 116 22-40 (18) 4 leaves 13 50 33 36 10 40 Plant 36 spinu- 13 leaves 3 27 13 14 (-- 11 losa Plant 43 11-19 (8) 8 leaves 5 38 21 30 09 all six characters (mean of the minimum scores to mean of the maximum). The extent of variation is almost twice as great in intermedia (79) as in spinulosa (42), while in the hybrid it is quite intermediate (58) between the two. 82 Rhodora [Vol. 68 | | l | l | | | 0 10 20 30 40 50 60 70 80 90 GRAPH 3. Analysis of variation in Dryopteris spinulosa, range and extent of variation in means of adjusted scores of characters a and € to f, from Chart 4. Solid bar: variation between extreme leaves. Outline bar: variation between extreme plants. I: intermedia, H: hybrid, S: spinulosa. The greater extent of variation of the characters of inter- media is consistent with the opportunities for recombination and mutation to be expressed phenotypically at the diploid level and the smaller extent of variation of spinulosa is con- sistent with the restrictions on their expression at the tetra- ploid level. The variation as represented by extreme leaves and ex- treme plants and the extent of variation between them is given in Chart 4 and in Graph 3. It will be seen that the extent of variation among extreme leaves and extreme plants is more than twice as great in intermedia as in spin- ulosa while in the hybrid it is about midway between that of the parents. The fact that the extent of variation in individual characters, in extreme leaves and in extreme plants of the Aybrid, is so nearly intermediate between that of the parents implies that the hybrid plants we have ana- lyzed have come from the whole spectrum of variation of the parent plants as we have seen it. One of the most interesting aspects of the variation of the cytotypes is their intraplant variability. Data on this variation, from selected plants, are presented in Chart 5. 1966] Dryopteris spinulosa — Tryon & Britton 83 The last column of the chart gives the extent of the intra- plant variation as a per cent of the total extent of variation in the cytotype. Most commonly the intraplant variation is rather small and this is illustrated by plants 27 and 301 which show the variation in the leaves of a plant in one year (one crown) and plants 36 and 38(1959, 1960) which show the variation in the mean of crowns in successive years. The frequent extent of intraplant variation of a character is about 15%, or less, of that of the cytotype. Rarely the intraplant variation may be great and this is illustrated in plants 38(1959) and 504 which show the variation in the leaves of a plant in one year and plants 32 and 117 whick show the variation in the mean of crowns in successive years. These examples illustrate that rarely the extent of variation of a character in one plant may be 70%, or more, of that of the cytotype or, as in plant 117, may even exceed that of the cytotype. (This is due to the fact that in Chart 3 the mean of plants is derived from all availa- ble leaves of a plant rather than only those of a single crown.) The genetic implications of the great intraplant vari- ability suggest that the genes controlling these characters are not markedly different in intermedia and spinulosa. They also suggest that the expressivity of the genes is extremely variable, providing that no unusual mitotic divisions are involved and that the cells of a plant all have a characteristic genotype. Similar variability in gene expression is well known in other plants, for example, the leaves of Morus alba. The taxonomic implications are that it vitiates, to à degree, the differences between the taxa brought out by the difference in their means of these characters. The only character that we have studied in which no vari- ation was observed within a cytotype was the chromosome number. Although both the spores and the indusia have been grouped into two discrete classes: normal-abortive and glabrous-glandular respectively, there is in fact variation in both of these characters. We have not studied the proportion of normal to abortive spores in spinulosa and intermedia, 84 Rhodora [Vol. 68 CHART 5. Intraplant variation in Dryopteris spinulosa. Range and extent of variation of single characters in selected plants. Data on individual leaves are in roman type, data of means of plants are in italic. Plant, cytotype Range of Variation. Extent of Variation and material, In example as 96 of Character. cytotype. S [^ Original — Adjusted, . E data. scores. " 8 HS A > ov o Plant 301, 0.5 mm. longer hybrid, 5 c to 41-48 7 56 12% leaves, 1960 3 mm. longer Plant 27, intermedia d 5 to 6 44-56 12 78 15% 4 leaves, 1960 Plant 504, 4 mm. longer spinulosa, c to 3-38 35 50 70% 4 leaves, 1961 15 mm. longer Plant 38, 6.5 mm. shorter hybrid 5 c to 20-70 50 56 90% leaves, 1959 9.5 mm. longer Plant 36, 7.2 mm. longer spinulosa, 4 c in '59 to 7.4 27-28 1 16 6% leaves, 1959, mm, longer in 5 in 1960 60 Plant 38, 1 mm. longer hybrid 5 c in '59 to 1.8 44-47 3 21 14% leaves, 1959, mm. longer in 3 in 1960. '60 Plant 32, 2 in 1959 hybrid, 4 f to 20-43 23 30 77% leaves, 1959, 4.3 in 1960 3 in 1960 Plant 117, 3.66 in 1959 hybrid, 3 e to 9-38 29 23 126% leaves, 1959, 1.66 in 1960 3 in 1960 1966] Dryopteris spinulosa — Tryon & Britton 85 being content with the observation that the abortive spores are rare. The glandularity of the indusium of intermedia and of the hybrid is clearly variable and although this has not been studied in detail it seems to show a range of vari- ation similar to that of the pinna base in intermedia and the hybrid. Identification. — Identification is most readily made during the rather brief time when mature spores are present in abundance. There is a possibility of contamination when there are only a few spores. During this two or three week period, the hybrid is readily identified by its abortive spores, intermedia by its normal spores and glandular indusium and spinulosa by its normal spores and glabrous indusium. Before, or after, this period fertile leaves of spinulosa may be identified by their glabrous indusium but intermedia. and the hybrid cannot be distinguished from each other by the indusium. Sterile ma- terial of spinulosa or intermedia cannot always be dis- tinguished from the hybrid. The difficulties of identification, then, center on the many herbarium specimens and plants in nature that are collected or observed at a time when uncontaminated mature spores are not present. Other characters must then be employed, if possible, for identifi- cation and it is with these that this study has largely been concerned. We have mentioned our conclusion that the parameters of the characters we have studied are undoubtedly greater in the taxa and their hybrid than in our sample of them. For this reason we do not believe that any of the data on single charaeters (Charts 1, 2 and 5, Graphs 1 and 2) afford a basis for the identification of all plants of the cytotypes. In all of the characters, except a, the range of variation of one cytotype overlaps that of another (or both) and in character a, the range of spinulosa is very close to that of the other two. The data on extreme plants (Chart 4, Graph 3), derived from the mean of 5 characters, shows that the range of variation of each cytotype is distinct from that of the other 86 Rhodora [Vol. 68 two. However, the degree of distinction is small: a differ- ence (in the mean of the adjusted scores) of 3 between plant 43 of spinulosa (19) and plant 284 of the hybrid. (22) and a difference of 4 between plant 116 of the hybrid (40) and plant 101 of intermedia (44). We do not believe that these small differences afford a basis for the identification of all plants of the cytotypes. It seems worthwhile to emphasize the variability of character c, the relative length of the inferior basal pinnules on the basal pinnae and the adjacent ones, because this character has been much relied upon in the literature for purposes of identification. Chart 2 and figs, 11-12, 15-16 demonstrate the variability in this character. Intermedia usually has the inferior basal pinnules shorter than the ad- jacent ones (fig. 11), but they may be longer (fig. 12) and spinulosa, usually has the basal inferior pinnules longer than the adjacent ones (fig. 16), but they may be equal in length (fig. 15). The great variation among the leaves of one crown can be seen in Chart 5, Plant 38 (1959) in which one of the five leaves had the inferior basal pinnules 6.5 mm. shorter than the adjacent ones and another leaf had them 9.5 mm. longer. The evergreen quality of the leaves has also been empha- sized in the literature and it is generally true that leaves of spinulosa are deciduous and those of intermedia are ever- green. However, our limited observations have indicated that this character is a variable one especially influenced by the environment of the individual plant. In most of the plants of the hybrid that were studied the leaves were de- ciduous or only partially evergreen but in some they were evergreen as is typical intermedia. Other characters employed in the identification of the taxa and their hybrid that we have also found to be too vari- able to be utilized are the position of the pinnae in relation to the rachis (see Analysis of Characters, g.) and the as- pect of the lamina (see Analysis of Characters, h). Con- sidering the variability discovered in all of the characters mentioned, we feel that others such as the orientation of 1966] Dryopteris spinulosa — Tryon & Britton 87 the rhizome, the color of the petiole scales and the type of spines on the spores cannot be accepted as affording a dis- tinction of the taxa until their constancy has been confirmed by an extensive study of plants of known identity. There remain then as wholly applicable characters for identification of the cytotypes only the glandular or glabrous indusium, the abortive or normal spores and the chromo- some number. Certain extremes of other characters do seem sufficiently distinctive to allow some confidence in their use and these have been added to the key. It will be obvious that the key will not allow the identification of all material. Key. Indusium glandular. Spores normal; n—41. (Any leaf with 9-10 basal inferior pinnules on the lowest 10 pinnae overlying the rachis, or any plant with four or more leaves averaging 6-10 overlying pinnules probably belongs here.) .......— eene intermedia Spores abortive; n—123. (Any plant with a glandular in- 2 dusium and four or more leaves averaging 0 cverlying pin- nules probably belongs here.) ...- Hmmm hybrid Indusium glabrous; spores normal; n—82. (Any leaf with the pinna-bases glabrous probably belongs here.) .....- spinulosa Classification. — The authors are in amiable, but firm, disagreement about the proper taxonomic status of intermedia and spinulosa. Donald M. Britton is of the opinion that, in these taxa, the cytogenetic evidence must be decisive. Intermedia is a basic diploid taxon genetically isolated from spinulosa which is a derived polyploid taxon. Accordingly, the status of species is most appropriate for each of them. Rolla Tryon is of the opinion that the taxonomic status of intermedia and spin- ulosa must be determined by a comparision of the two to each other and to other taxa in their alliance on the basis of all evidences of similarity and difference. He is of the opinion that the similarities and differences brought out in the pres- ent study make the category variety most appropriate, at the same time admitting that this status is tentative pending adequate comparative studies of other allied taxa. 88 Rhodora [Vol. 68 It is difficult to decide whether or not the hybrid. should be treated as a taxon and then be designated by a binomial. Rather than explore this complex question here we prefer not to recognize it as a taxon but to formally refer to it by the appropriate formula. This is advantageous because it directly conveys the parentage which is the most im- portant information about a hybrid. The triploid hybrid. — There are several aspects of the natural history of the sterile hybrid. that deserve some additional discussion. Al- though its distribution is not well known, its presence has been cytologically confirmed in Ontario, Massachusetts, New Jersey, Virginia and Michigan. It is probably to be found throughout northeastern United States and adjacent Canada within the range of the two parents and in most localities where they both grow. We have found it at all (seven) localities where cytological sampling revealed the presence of both intermedia and spinulosa. In Massachusetts, it has been found at each of several localities where a search was made for it. Most of these plants were identified by their abortive spores; a few were identified by their chromosome number. The data in Chart 6 suggest the abundance of the hybrid in a few localities and also demonstrate how it may be more abundant than either parent. We have observed in the same locality that the triploid plants often have larger leaves and more leaves than those of the parents. While this might be due to environmental conditions that are more favorable for it than for either parent, it seems more likely that it is due to the phenomenon of hybrid vigor. The hybrid plants may arise in two ways. The plant may come from the fertilization of a gametophyte of one parent by one of the other parent, or it may come from an existing plant by vegetative reproduction. Although we have seen some evidence of clonal development in the hybrid we have not been able to assess the relative importance of this in the formation of populations, The principal reason being that although a close similarity of plants that are near each other may be taken as evidence of a clone, it does not follow 1966] Dryopteris spinulosa — Tryon & Britton 89 CHART 6. Relative abundance of Dryopteris spinulosa at four locali- ties, from random samples of the populations. % of Locality. Number of plants. population. ‘High and Dry’, Wellington Co., Ontario. intermedia 16 19 hybrid 56 65 spinulosa 14 16 Cockburn Swamp, Wellington Co., Ontario. intermedia 3 11 hybrid 14 52 spinulosa 10 37 Sodom, Halton Co., Ontario. intermedia 16 31 hybrid 20 38 spinulosa 16 31 Littleton, Massachusetts. intermedia 26 2T hybrid 62 64 spinulosa 9 9 that dissimilar plants always represent independent hybrids. The data in Chart 5 on variation in a crown from one year to the next (plants 32, 117) are sufficiently suggestive of the variability that may be present in a clone to make its identification in the field most uncertain. Some estimate of the extent of clonal development may be made by measuring the growth of the rhizome apex. The growth of the adult rhizome extends the apex some two or three centimeters a year and when the apex occasion- ally branches two separate plants are soon produced. Later- al stolons are also commonly produced from between the persistent leaf bases, on the older portions of the rhizome. These are slender and grow three or four centimeters in length before the internodes shorten and a crown of small leaves is produced. The base of the stolon soon dies and the rhizome of the small separate plant then may spread away from the parent. The spread of a clone by these two methods is rather slow and it would not occupy more than fifty to a hundred square meters of area in a few hundred years. In many localities where the hybrid. grows, it occupies an area of several thousand square meters. Even allowing 90 Rhodora [Vol. 68 for maximal clonal development, each of these large popu- lations must represent a few dozen, or even a few hundred, original hybrid plants. Such populations must occur fre- quently over an extensive area, at least from Virginia to New England, Ontario and Michigan, and they must repre- sent several hundred thousand F, plants. This frequency of crossing of the parental gametophytes can be evidence of the breeding behavior of the parents themselves. Since the gametophytes of the two taxa can cross with some frequency, those from different plants of the same taxon should do so with equal or greater frequency. Although we do not have sufficient data on the population Structure of intermedia and spinulosa to bring to bear as evidence of predominant outcrossing, our data on variation are consistent with the idea that outcrossing occurs. It seems appropriate to bring out this indirect evidence for relatively frequent outcrossing since so little is known about the natural breeding behavior of ferns. The relative greater abundance of the hybrid than either parent (Chart 6) is unusual, perhaps unique, in sterile hy- brid ferns. Most of these in Dryopteris are rare and occur only as an occasional plant or small colony in a very few of the places where the parents grow together, for example, D. marginalis X spinulosa var. intermedia and D. Goldiana X marginalis. A few are more common and occur in many localities where both parents grow, for example, D. cristata X spinulosa and D. cristata X spinulosa var. intermedia. The latter, perhaps more familiar as D. X Boottii, is the most common hybrid, except for intermedia X spinulosa, but it is never as abundant as its parents. We believe that the great abundance of intermedia X spinulosa is due to the large number of hybrid sporelings that are formed and to their intermediate physiological traits. Their physiology may adapt them to a large number of niches favorable for immediate survival. It may also provide them, and older plants, with the capacity to survive drought and pluvial periods, that might diminish the numbers of spinulosa, which prefers wet conditions, and also of intermedia, which prefers drier ones. 1966] Dryopteris spinulosa — Tryon & Britton 91 When we first became aware of the frequency of the hy- brid, we fully expected that further sampling would reveal a hexaploid. Although this expectation was not realized there are some aspects of the triploid hybrid that have a pertinence to studies of naturally occurring hexaploids. One of these aspects is the proximity of extreme plants of the hybrid. (Chart 4 and Graph 3) to certain plants of the parents. A hexaploid produced from one of these plants would be much closer to one parent than to the other and if the triploid became extinct (or was unknown) the rela- tions of the hexaploid might be correspondingly obscured. Further diffieulties in the analysis of the hexaploid would result if it were produced more than once and from plants at the opposite ends of the spectrum of variation of the triploid. The origin of the tetraploid. — The hybrid of spinulosa and intermedia has approximate- ly 41 pairs of chromosomes and 41 single chromosomes and on the basis of this evidence, Walker (1961) has concluded that the parents share a common genome. Therefore, he considers spinulosa to be an allopolyploid with intermedia as one of its ancestral parents. Among the known diploids only intermedia is a credible parent for spinulosa and, if spinulosa is an allopolyploid, it is evident that the other parent is either extinct or is as yet undetected. It must also be considered, however, that spinulosa may be an autopoly- ploid between two members of an original diploid spinulosa taxon which was closely related to intermedia. There are two characters of spinulosa that are not con- sistent with Walker’s hypothesis. These are the glabrous indusium and the only slightly and infrequently glandular lamina. A hybrid with typical intermedia would be ex- pected to have a glandular indusium and a more gland- ular lamina as, for example, in D. cristata X spinulosa var. intermedia and in D. cristata var. Clintoniana X spinulosa var. intermedia. However, it must be admitted that we can not be certain that the gladularity of intermedia, is always strongly dominant in its hybrids and so the slight 92 Rhodora [Vol. 68 glandularity of spinulosa can only be a qualified objection to Walker's hypothesis. Our data on the breadth of variation in intermedia, and in the hybrid, suggest that Walker's interpretation may be correct under certain cireumstances. The present data would be sufficiently consistent with his hypothesis if one of the parents of the original hybrid spinulosa were a plant of intermedia that was rather distant from the central varia- tion of the taxon in the direction of spinulosa. It would also be consistent if one of the parents of the original hybrid spinulosa was a member of an ancestral intermedia that was closer to the present spinulosa and which has since diverged from it. The variation that we have observed in spinulosa and in intermedia suggests that the origin of spinulosa may be a complex problem and that further studies are desirable to substantiate the claim of intermedia as one of its parents. BIBLIOGRAPHY. BRITTON, D. M. 1953. Chromosome studies in ferns. Amer. Jour. Bot. 40: 575-583. 1961. The problems of variation in North American Dryopteris. Amer. Fern Jour. 51: 23-30. 1962. Dryopteris dilatata (Hoffm.) A, Gray in North America. Rhodora 64: 207-212. and J. H Soper. 1966. The cytology and distribution of Dryopteris species in Ontario. Can. Jour. Bot. 44: 63-78. MANTON, I. and S. WALKER. 1953. Cytology of the Dryopteris spin- ulosa complex in eastern North America. Nature 117: 1116-1118. WALKER, S. 1959. Cytotaxonomic studies of some American Species of Dryopteris. Amer. Fern Jour. 49: 104-112, . 1961. Cytogenetic studies in the Dryopteris spinulosa complex, II. Amer. Jour. Bot. 48: 607-614. — ————. 1962. The problem of Dryopteris Leedsii. Amer. Jour. Bot. 49: 971-974. WAGNER, W. H., JR. 1963. Pteridophytes of the Mountain Lake area, Giles County, Virginia, including notes from Whitetop mountain. Castanea 28: 113-150. and D. J. HAGENAH. 1962. Dryopteris in the Huron Mountain Club area of Michigan. Brittonia 14: 90-100. VARIETIES OF BALLOTA NIGRA IN THE EAST- ERN UNITED STATES — The Black Horehound, Ballota nigra L., common in western Europe both native and as a weed, is an uncommon weed in the eastern and south- eastern United States, probably originally introduced mostly on ballast. Although no varieties or forms are mentioned in the eighth edition of Gray's Manual, it does occur in two rather easily distinguishable forms, which are called in the New Britton and Brown Illustrated Flora var. foetida Vis. and, var. ruderalis Koch, but these names are not correct. The former has the calyx lobes short and abruptly cuspidate at the apex, whereas the latter has them long-triangular and gradually acuminate. Dr. A. Patzak, of the Naturhistorisches Museum, Wien, published some time ago a revision of Ballota in which seven subspecies of B. nigra L. are recognized. Recently, a number of plants introduced into the United States were submitted to Dr. Patzak for study, and came back annotated partly as subsp. foetida Hayek and partly subsp. nigra. There can be no question as to the correctness of the latter name; as a variety it would be called var. nigra, of which var. ruderalis (Swartz) Koch is a synonym. The former name is not a new entity with Hayek (Repert. Sp. Nov. Fedde Beih. 30 (2): 278. 1929), but was based ultimately on Ballota foetida Lam. (Fl. Franc. 2: 381. 1778), which was merely a change of name for B. nigra L., and was thus superfluous when published and illegitimate. However, an originally illegitimate epithet like foetida can be validated in another category by a later author, who in this instance appears to be Koch, who published the variety B. nigra L. var. foetida Koch (Syn. Fl. Germ. Helv. 572. 1837), which must be considered as a new variety and not a transfer of B. foetida Lam. The authority for the combination B. nigra subsp. foetida should therefore be (Koch) Hayek, rather than merely Hayek. “Revision der Gattung Ballota Section Ballota," Annal. Naturhist. Mus. Wien 62: 57-86. 1958. 94 Rhodora [Vol. 68 When this plant is considered to be a variety, as such plants commonly are in floras of the eastern United States, the epithet will not be the same. Hayek was the first (ap- parently, according to Patzak’s synonymy) to propose a sub- specific epithet, and foetida thus has priority in this rank, but there were earlier varietal names proposed for this plant. Linnaeus himself realized, after the publication of B. nigra in the Species Plantarum of 1753, that there were two entities involved, and for the second he proposed Ballota aiba L. (Fl. Suec. ed. 2, 206. 1775), which is the plant with short, cuspidate calyx lobes that is called subsp. foetida by Patzak. As a variety its correct name will be B. nigra L. var. alba, (L.) J. E. Smith (Brit. Fl. 2: 635. 1804). C. V. MORTON SMITHSONIAN INSTITUTION, WASHINGTON, D. C. PASPALUM MINUS (GRAMINEAE) IN LOUISIANA AND MISSISSIPPI: Recently I reported Mobile, Alabama as a new location for Paspalum minus Fourn. (Banks, 1964). In the United States the species had been known previously in eastern Texas, near Fannett, Beaumont, and Anahuac. According to Jones et al. (1961), P. minus was collected by Silveus in Aransas county in southeastern Texas. My Mobile collec- tion extended the range of the species considerably but ap- peared to be isolated from the other locations. Last summer, while I was collecting plants for my Pas- palum studies, I found P. minus growing in Louisiana and Mississippi. The Mississippi collection was made two days after the Louisiana one during a field trip with Dr. Thomas Pullen, Department of Biology, University of Mississippi. — Contribution No. 64 from the Stephen F. Austin State College Department of Biology. 1966] Paspalum — Banks 95 Dr. Pullen arranged for lodging facilities at the University Forest Lands and provided local transportation to the collec- tion site for which I am grateful. These new locations help to bridge the gap between the Texas and Alabama stations and indicate that the species is probably more widespread in the Gulf Coastal Plain than we have realized. I contacted the following curators in Texas, Louisiana, and Mississippi to determine if other locations for the spe- cies might be known: Joseph Ewan (No), Frank Gould (TAES), G. W. Johnston (MISSA), Thomas Pullen (UNIV. OF MISS.) , Claude McLeod (SHST), Lloyd Shinners (SMU), John L. Strother (TEX), Clair A. Brown (LSU) and John Thieret (LAF). I wish to thank the above persons for checking the locations of P. minus in their herbaria and for allowing me to include the information in this paper. This correspondence did not reveal additional locations in Louisiana or Mississippi. Additional locations in Texas are, according to Strother’, Smith Point, and according to MeLeod?, Weldon, Lovelady, Diboll, Huntsville, and Grove- ton to Trinity. I do not know how P. minus was introduced into the United States. It is possible that hurricanes carried seeds from the West Indies. Another possibilty is that it may have been introduced in foreign imports of seeds of Pas- palum notatum Fligge (Bahiagrass). P. notatum is be- coming widespread in its distribution in the southeast, especially along highways and in pastures where it is often planted for erosion control and for forage. P. minus closely resembles P. notatum, the former being only slightly smaller in most characteristies, and probably accounts for its hav- ing been unnoticed. Closer observation may reveal addi- tional specimens of P. minus in colonies of P. notatum. Cytological examination of the Louisiana and Mississippi material using the aceto-carmine squash method revealed 10 chromosomes in the microspores and meiosis appeared normal. See Figure 1. Gould (1958) reported n=20 in the Huntsville material which was collected by McLeod?. My ?Personal communication. 96 Rhodora [Vol. 68 Fig. 1. Meiotic chromosomes of Paspalum minus (Banks 3500). n = 10. report therefore represents a new chromosome count for P. minus and indicates that some races are diploid. My collection data for P. minus in Louisiana and Missis- sippi and the disposition of specimens are as follows: LOUISIANA. ST. TAMMY PARISH: Edge of Pine woods along a wet ditch about 1.6 miles east of Abita Springs along Louisiana State Highway 36. Scattered clumps. D. J. Banks 3395. July 16, 1964. (ASTC, GH, LSU, MO, NO, NY, SMU, US). MISSISSIPPI. JACKSON COUNTY: In low pinelands in a wet ditch about 8 miles south of Vancleave. Plants erect. D. J. Banks 3500. July 18, 1964. (ASTC, GH, MISSA, MO, NY, SMU, University of Missis- sippi, University of Southern Mississippi, Us). DONALD J. BANKS STEPHEN F. AUSTIN STATE COLLEGE NACOGDOCHES, TEXAS LITERATURE CITED BANKS, D. J. 1964. Notes on Paspalum minus and Andropogon per- tusus in the United States. Rhodora. 66: 273-274. GOULD, F. W. 1958. Chromosome numbers in southwestern grasses. Am. Jour. Bot. 45: 757-767. JONES, F. B., C. M. ROWELL, JR. and M. C. JOHNSTON. 1961. Flower- ing plants and ferns of the Texas Coastal Bend counties. Welder Series B-1. Sinton, Texas. STUDIES IN THE FLORA OF BOLIVIA — IV. GRAMINEAE RoBERT C. FOSTER To anyone who has made use of Hitchcock’s treatment of the central and northern Andean grasses (Contrib. U. S. Nat. Herb. 24, pt. 8 [1927]), my indebtedness will be obvi- ous. I have followed it in the arrangement of genera and species, with such interpolations as time and the accretion of further material have made necessary. Since Hitchcock’s work was published almost forty years ago, it is not sur- prising that genera and species not known to him from Bolivia have now been found there. In the matter of species- concepts and synonymy, too, Hitchcock has been followed, almost without exception. This is conservative, perhaps ultraconservative, but that seems preferable to over-recogni- tion of categories of possibly uncertain value. It also means that if Hitchcock made errors they are perpetuated here. A non-agrostologist is perhaps too greatly impressed by the number of exceptions and qualifications made necessary in writing keys if the genera are first divided into tribes. Consequently, a continuous key is provided for the 109 genera treated in this paper. It is based, like the generic descriptions, on the species found in Bolivia, not on extra- territorial species. Certain points as to its terminology should be noted. A fertile floret may be either perfect or pistillate; when necessary, specific use is made of one term or the other. A sterile floret may be staminate, neuter, or a rudiment. Most often, the term is applied to an empty lemma. Again, a primary division is made on the basis of the disarticulation of spikelets — above the glumes and not above the glumes. This phraseology has been deliberate- ly used rather than above the glumes and below the glumes. It is made necessary by the fact that in some genera both glumes are lacking. Disarticulation may also occur in the pedicels or in the rachis itself, which is somewhat too far 97 98 Rhodora [Vol. 68 below the glumes to permit even a broad use of that ex- pression. Once again, this paper is based on the collections of the Gray Herbarium (GH). These have been supplemented by material borrowed from the incomparable collection of Gramineae in the United States National Herbarium (US). I am much indebted to Dr. W. L. Stern and Dr. T. R. Soder- Strom for their generous coóperation. In some cases, where Bolivian material has not been seen, collections seen by previous workers have been cited, with that worker's name in parentheses. For corrections and additions to this paper, I shall be most grateful. KEY TO GENERA 1. Inflorescences not surrounded by a hard bony involucre, 2. Disarticulation above the glumes. 3. Spikelets 1-flowered. 4. Inflorescence a solitary, terminal, pedunculate spikelet. MM 44. Aciachne. 4. Inflorescence not a solitary, terminal, pedunculate spike- let. 5. Rachilla prolonged as a bristle behind the palea. 6. Inflorescence not digitate. ........ 37. Calamagrostis. 6. Inflorescence digitate. occ... 57. Cynodon. ct Rachilla not prolonged behind the palea, except in the central spikelets only in Hordeum. 7. Inflorescence a terminal spike. 8. Spikelets 3 at a node, not unilateral on the rachis. sessen. 26. Hordeum. 8. Spikelets solitary at nodes, unilateral on the rachis, sees 58. Microchloa. 7. Inflorescence a panicle. 9. Spikelets paired, 1 pistillate and subsessile, 1 staminate and pedicellate, ................ 67. Pharus. 9. Spikelets not paired. 10. Awn of lemma excentric (in side view). 11. Lemma enclosing the membranous palea. sse 46. Nassella. 11. Lemma not entirely enclosing the coriaceous bicarinate palea. ................ RR 47. Piptochaetium. 10. Awn of lemma not excentric, or lemma awnless or cuspidate. 1966] Flora of Bolivia — Foster 99 19. Awn of lemma trifid. ........ 49. Aristida. 12. Awn of lemma, if present, undivided. 13. Lemma shorter than glumes. 14. Lemma neither fusiform nor indurate. ..............-. 38. Agrostis. 14. Lemma fusiform, indurate. ....-... si a Uo 48. Stipa 13. Lemma equal to glumes or longer than at least 1 glume. 15. Lemma awnless. s.es. EU RPM E ir e a 45. Sporobolus. 15. Lemma awned or cuspidate. 16. Awn dorsal „e.e. mE e MARO on 43. Triniochloa. 16. Awn terminal, if present, otherwise the lemma cuspi- date. ........ 42. Muhlenbergia. 8. Spikelets more than 1-flowered. 17. Spikelets with fertile florets only. 18. Rachilla villous. 19. Lemma with the true awn 1.1-2.2 cm. long. .....- NNNM 16. Cortaderia. 18. Rachilla glabrous. 20. Florets distant, glumes minute. ...Áná8 NNNM 7. Aphanelytrum. 20. Florets not distant, glumes not minute. 21. Spikelets and glumes over 1 cm. long. .......—.« NECS te 14. Arundo. 21. Spikelets and glumes less than 1 cm. long. 29. Glumes less than 2 mm. long. erer MM 8. Puccinellia. 29. Glumes more than 2 mm. long. 23. Glumes shorter than lemmas. ......-- vecececeausesecsanecececeseesseeesseeeess 19. Anthochloa. 23. Glumes longer than lemmas, seee NNNM 22. Triodia. 17. Spikelets with fertile and sterile florets, or the florets all pistillate or all staminate. 24. Plants dioecious. 25. Rachilla villous. ....MMÁ 16. Cortaderia. 25. Rachilla glabrous. 26. Leaf-blades about 1 mm. wide; inflorescence a rather condensed panicle. .... 13. Distichlis. 26. Leaf-blades to 4-6 cm. wide; inflorescence a plumose panicle. „e.e 17. Gynerium. 100 Rhodora [Vol. 68 24. Plants not dioecious. 27. Fertile florets above the sterile florets in the Spikelet. 28. Lateral nerves of first glume with a row of tubercles bearing long golden-brown hairs. RM 70. Tristachya. 28. Lateral nerves of first glume not tuberculate, not with long golden-brown hairs. 29. Spikelets strongly flattened and imbricate. np MM 64. Phalaris. 29. Spikelets not strongly flattened and imbri- cate. 30. Spikelets with more than 1 fertile floret. 31. Inflorescence a plumose panicle. RR 15. Phragmites. 31. Inflorescence not a plumose panicle. 32. Leaf-blades linear, to 6 mm. wide. ........... 1. Arthrostylidium. 32. Leaf-blades lanceolate, to 3 em. wide. ................ 2. Arundinaria. 30. Spikelets with only 1 fertile floret. 33. Fertile branches fascicled at nodes of main plant-axis; terminal floret perfect. sees 3. Chusquea. 33. Fertile branches not fascicled at nodes of main plant-axis; terminal floret pistillate, 34. Lemma of pistillate floret glab- rous or scabridulous. eee. TTT etn 68. Arundinella. 34. Lemma of pistillate floret densely long-pubescent. sess. HM 69. Trichopteryz. 27. Fertile florets below the sterile florets in the spikelet. 35. Inflorescence condensed into small glomerules more or less hidden by leaf-sheaths. ................ 5LIMMMEA 63. Munroa. 35. Inflorescence not condensed into glomerules. 36. Inflorescence digitate or paniculate (in one instance a panicle with whorls of digitately arranged branches). 37. Inflorescence digitate. 38. Rachis prolonged beyond the upper- most spikelet as a short subulate point, ................ 56. Dactyloctenium. 1966] Flora of Bolivia — Foster 101 38. Rachis not prolonged beyond the uppermost spikelet. 39. Spikelets with more than 1 per- fect floret. .........- 55. Eleusine. 39. Spikelets with only 1 perfect floret. .. RH 60. Chloris. 37. Inflorescence a panicle. 40. Lemma wing-margined. ... 40. Lemma not wing-margined. 41. Rachilla glabrous. 42. Glumes or lemmas awned. 43. Spikelets unilateral in 2 rows on the rachis. .......-- Et 61. Trichloris. the rachis. 44. Lemmas longer than glumes. 45. Inflorescence a spi- cate, dense, inter- rupted panicle. ...... Rude 31. Trisetum. 45. Inflorescence an open racemose pani- cle. .... 53. Gouinia. 44. Lemmas shorter than glumes. eA "— 59. Gymnopogon. 42. Glumes and lemmas awnless. 46. Spikelets unilateral on the the rachis in 2 rows. — 52. Leptochloa. 46. Spikelets not unilateral on rachis in 2 rows. 47. Glumes longer than lemmas. 48. Spikelets paired, 1 sessile and 1 pedi- cellate. .. 34. Holcus. 48. Spikelets not pair- ed. 49. Inflorescence spicate, dense. .. .... 29. Dissanthe- lium. 102 Rhodora [Vol. 68 49. Inflorescence not Spicate, the branches dis- tant, spreading. ee 18. Melica. 47. Glumes equal to or shorter than lemmas. 50. Leaf-blades long- petiolate, lance- ovate. eese EM 21. Zeugites. 50. Leaf-blades not long-petiolate ^ nor lance-ovate. 51. Paleas persist- ent on rachilla after fall of lemmas. ........... . 12. Eragrostis. 51. Paleas not per- sistent. .. 10. Poa. 4l. Rachilla not glabrous. 52. Lemma not bidentate. 53. Lemma apically dissected into numerous awns. ........ MM 23. Pappophorum. 53. Lemma not apically dis- sected, 54. Paleas persistent on rachilla after fall of lemmas. nsere ee 12. Eragrostis. 54. Paleas not persistent. 55. Spikelets 1.5 cm. long or longer, ... — 4. Bromus. 55. Spikelets rarely longer than 1 cm., usually shorter. 56. Glumes longer than lemmas. .. serere 18. Melica. 56. Glumes equal to or shorter than lemmas. 1966] Flora of Bolivia — Foster 103 57. Lemmas awnless. ......- S 10. Poa. 57. Lemmas AWE, ........- .. 6. Festuca. 52. Lemma bidentate or 4- toothed. 58. Lemma-awn dorsal. 59. Spikelets 1.5-2.5 cm. long. s 33. Avena. 59. Spikelets 4-5 mm. long. 2d 32. Deschampsia. 58. Lemma-awn from base of sinus between teeth. 60. Spikelets about 2.5 cm. long. ienn 4. Bromus. 60. Spikelets 1.5 cm. or less. 61. Glumes shorter than lemmas. .......- . 81. Trisetum. 61. Glumes longer than lemmas, 62. Spikelets 1.5 cm. long. 63. Leaves glab- TOUS, nenne 9. Amphibro- mus. 63. Leaves papil- late - hirsute. . 85. Dan- thonia. 62. Spikelets about 7 mm. long. ...... . 96. Lampro- thyrsus. 36. Inflorescence neither digitate nor pani- culate, but a solitary terminal spike or raceme. 64. Inflorescence a raceme with few, distant, ultimately spreading spikelets. MEM 5. Brachypodium. 104 Rhodora [Vol. 68 65 Spikelets placed edgewise to the rachis, sesse. 27. Lolium. 65. Spikelets not placed edgewise to the rachis. 66. Glumes awned. ........ 25. Elymus. 66. Glumes awnless. 67. Glumes shorter than lemmas. — HR 54. Tripogon. 67. Glumes not shorter than lemmas. 68. Plants 1-2 dm. high; spikes 1-3 cm. long. ... -~ 62. Bouteloua simplex. 68. Plants to 1-2 m. high; spikes 7-20 em. long. ...... MEM 24. Agropyron. 70. Spikelets with unisexual or neuter florets. 71. Inflorescence a spicate panicle or raceme, 72. Inflorescence a raceme. 73. Spikelets in whorls around the rachis. .......... mE 28. Pariana. 73. Spikelets not in whorls around the rachis. .... IM 51. Aegopogon. 72. Inflorescence a Spicate panicle. ........ 41. Lycurus. 73a. Pistillate and staminate spikelets in separate inflorescences on the same plant. ...... 66. Luziola. 73a. Pistillate and staminate spikelets not in separate 74. Spikelets not paired. co.cc, 94. Olyra 74. Spikelets paired. ................ 100. Hyparrhenia. 70. Spikelets with perfect florets. 75. Inflorescence a raceme or spike. 76. Inflorescence a raceme, ............... 51. Aegopogon. 76. Inflorescence a Spike, eee 50. Tragus. 75. Inflorescence a panicle. Ti. Glumes lacking. Jess 65. Leersia. 77. Glumes present. 78. Spikelets paired. 79. One spikelet perfect, the other staminate or neuter. arcen. 41. Lycurus. 79. One spikelet perfect and sessile, the other pistillate and pedicellate. 97, Eriochrysis. 78. Spikelets not paired. 80. Spikelets awnless. ............ 20. Orthoclada. 1966] Flora of Bolivia — Foster 105 80. Spikelets awned. 81. Inflorescence not interrupted. .............- ERE A DER OREL 39. Alopecurus. 81. Inflorescence interrupted. ............- MUI EEUU 40. Polypogon. 69. Spikelets with 2 or more florets. 82. Fertile floret below the sterile floret. 83. Inflorescence a panicle; spikelets usually solitary. Codb LER ements 20. Orthoclada. 83. Inflorescence of pectinately arranged spikes; spike- lets not solitary. ..... MM 62. Bouteloua. 82. Fertile floret above the sterile floret. 84. Spikelets subtended by bristles, these separate, or at least partially united and forming a bur. 85. Spikelets and bristles forming a BUTS e terre devesccssvscassscsccscccscesenauacnscessonscacossssscsonasees 93. Cenchrus. 85. Spikelets and bristles not forming a bur. 86. Bristles persistent, ....... 91. Setaria. 86. Bristles deciduous. ...........--- 92. Pennisetum. 84. Spikelets not subtended by bristles. 87. First glume absent. 88. Spikelets without unisexual florets. 89. Spikelets not unilateral on the rachis. .... E E 71. Leptocoryphium. 89. Spikelets unilateral on the rachis. 90. Second glume and fertile lemma turned away from the rachis. .... 80. Axonopus. 90. Second glume and fertile lemma turned toward the rachis (first glume rarely present but minute). 91. Fruit flexible, cartilaginous, usually dark. HM 13. Digitaria. 91. Fruit rigid, chartaceous, usually light. ........m HH 81. Paspalum. 88. Spikelets with unisexual florets; pistillate and staminate florets in separate spikelets. ........ vcunueeuusensecencnaneceesseceeeussssessseeeeeeeeeeeseeeeeeee 94. Olyra. 87. First glume present. 92. Second glume ventricose and uncinately spiny at maturity. ee 75, Pseudechinolaena. 92. Second glume not ventricose nor uncinately spiny at maturity. 93. Spikelets paired, with 1 spikelet sessile, the other pedicellate. 94. Both spikelets, or only 1, with a per- fect floret. 95. Both spikelets with a perfect floret. 106 Rhodora [Vol. 68 96. Spikelets awnless. sesse. — ÉE— 96. Saccharum. 96. Spikelets awned. .. 98. Erianthus. 95. One spikelet with a perfect floret. 97. Perfect spikelet pedicellate. aravassonaiesereseressenne 104. Trachypogon. 97. Perfect spikelet sessile. 98. Disarticulation immediately below glumes. ............. een —— 99. Andropogon. 98. Disarticulation not immedi- ately below glumes, but in rachis or pedicel. 99. First glume concealing the rest of the spikelet. ........ TE 107. Hackelochloa. 99. First glume not conceal- ing spikelet. 100. Inflorescence a soli- tary raceme (occa- sionally more than 1). 101. Spikelets awned. .. . 103. Hetero- pogon. 101. Spikelets awnless. 102. Glumes pubes- cent, hirsute or nurus. 102. Glumes glab- YOUS. secssssivecessss suris, 100. Inflorescence panicu- late. 103. Pedicellate spike- let staminate. ...... .. 101. Sorghum. 103. Pedicellate spike- let reduced to a pedicel, .................. eene 102. Sorghas- trum. 94. Neither spikelet with a perfect floret. —ÓÉORRE 108. Tripsacum. 93. Spikelets not ‘paired, or if in pairs, not 1966] Flora of Bolivia — Foster 107 with 1 spikelet sessile and the other pedi- cellate. 104. Inflorescence of solitary, or mostly so, racemes. 105. Spikelets paired. ..... 74. Thrasya. 105. Spikelets unpaired, but sometimes in 2 rows. 106. Spikelets (pistillate only) sunken in rachis. e.i iinsert. NMERRUNUR-: Su 108. Tripsacum. 106. Spikelets not sunken in rachis. 107. Sterile lemma and second glume with tufts of hairs. .. NEUE BIET 77. Mesosetum. 107. Sterile lemma and second glume glabrous. ............ MEE 79. Brachiaria. 104. Inflorescence a panicle. 108. Spikelets paired. 109. Glumes awnless. 110. Sterile lemma and second glume alike. ...... 72. Trich- achne. 110. Sterile and fertile lemmas alike, unlike second glume. ——Ó 95. Imperata. 109. Glumes awned or mucronate. 111. Ring-callas on rachilla at base of spikelet. ................- NEMPE 78. Eriochloa. 111. Ring-calus absent from rachilla. ...... 89. Oplismenus. 108. Spikelets unpaired or clustered. 112. Spikelets clustered. 113. Spikelets awnless, not plano- convex. .... 86. Hymenachne. 113. Spikelets awned, plano- COnVeX. ...... 90. Echinochloa. 112. Spikelets unpaired. 114. Spikelets placed obliquely on pedicels, ............ 84. Lasiacis. 114. Spikelets not placed obliquely on pedicels. 115. Spikelets awnless. 116. Second glume in- flated, saccate. .......... M eibi 85. Sacciolepis. 108 Rhodora [Vol. 68 116. Second glume not in- flated nor saccate. 117. Lemma stipitate. .. . 83. Ichnanthus. 117. Lemma not stipi- tate. 118. Spikelets with 2 perfect flo- rets, subglo- DOSE, ................ .. 88. Isachne. 118. Spikelets with 1 perfect floret, not subglobose. 119. First glume as long as Spikelet, ...... ..87. Homo- lepis. 119. First glume not as long as spikelet. ... 82, Pani- cum. 115. Spikelet awned or mucro- nate. 120. Ring-callus on rachil- la at base of spikelet. NEN 78. Eriochloa. 120. Ring-callus absent. 121. Glumes very un- equal. .............. ees sen 76, Rhynchely- trum. 121. Glumes equal or subequal. ................ ... 89. Oplismenus. 1. Inflorescences, at least the pistillate, surrounded by a hard bony involuere. essre 109. Coix. 1. Arthrostylidium Rupr. Arthrostylidium racemiflorum Steud. Syn. Pl. Glum. 1: 336 (1854). Perennial, climbing or sprawling, to 5 m. long; main axis to 5 mm. thick, the fertile branches clustered at the nodes. Leaf-blades linear, to 6 mm. wide, often short-pubescent above and beneath, with scattered longer hairs. Inflorescence racemose, the racemes to 8 cm. long, the 1966] Flora of Bolivia — Foster 109 rachis pubescent. Spikelets few-flowered, the lowest lemmas empty; disarticulation above the glumes. Lemma to 6 mm. long, 7-nerved, awned, the awn 1-2 mm. long. Palea longer than the lemma, the keels ciliolate. SANTA CRUZ: ICHILO: Río Yapacani, 350 m., Steinbach 7587 (GH). This collection is a sterile sheet identified by Mrs. Agnes Chase as “probably” this species. 2. Arundinaria Michx. Arundinaria Herzogiana Henrard in Meded. Rijks Herb. Leiden, no. 40: 75 (1921). Shrubby perennial. Leaf-sheaths long-fimbriate at the apex, the fimbriae to 2 cm. long; blades lanceolate, acute, to 25 cm. long and 3 cm. wide, glaucous, glabrous above, scabrous and more or less hirtellous beneath. Inflorescence a terminal, lax, racemose panicle to 3 dm. long, hardly, if at all, exserted from the uppermost leaves, the pubescent branches distant on the axis. Spikelets 2-3 cm. long, pedicellate, about 5-flowered; disarticulation above the glumes. First glume absent; second glume about 1 mm. long, 3-nerved, the margin ciliate, with a 3-4 mm. awn. Sterile lemmas 2, awned, the first pubescent on the upper portion. Fertile lemmas to 1 cm. long, 7-nerved, shortly puberulent, with a 2-3 mm. awn. LA PAZ: INQUISIVI [?]: Río Saujana, 3200 m., Herzog 2396 (US; type-number). There is no information available as to the height of the plant. 3. Chusquea Kunth Shrubby perennials, sometimes low-growing, or the culms 2-7 m. long, slender to stout, woody. Leaf-blades lanceolate to linear- lanceolate, 2-18 cm. long, acute, disarticulating from the persistent sheaths. Fertile branches loosely to densely fascicled at the nodes, or the inflorescences terminal on the culms; inflorescence an open to rather crowded panicle, or spicate in appearance, spikelets few to numerous in an inflorescence. Spikelets pedicellate, small; glumes present or absent; sterile lemmas 2, below 1 perfect floret; disarticu- lation above the glumes. a. Glumes absent or less than 0.5 mm. long. b. Glumes absent. ....... eem 2. C. delicatula. b. Glumes present, less than 0.5 mm. long. c. Culms 2-7 m. long, 1 cm. thick; fertile stems densely clustered at nodes, with numerous spikelets... 4. C. peruviana. c. Culms much shorter, 4 mm. thick; fertile stems few at nodes, spikelets few. eene emen 1. C. uniflora. a. Glumes present, 1 mm. long, or more. d. Rachis, branches and pedicels glabrous. „eee 5. C. serrulata. d. Rachis, branches and pedicels puberulent. 110 Rhodora [Vol. 68 e. Spikelets mostly distinctly pedicellate, the panicle somewhat open; lemmas distinctly awned. ......... 6. C. scandens. e. Spikelets rarely pedicellate, the inflorescence densely spicate in appearance, lemmas shortly mucronate. ............ 3. C. spicata. 1. Chusquea uniflora Steud. Syn. Pl. Glum. 1: 827 (1854). C. longipendula O. Ktze. Rev. Gen. 3(2): 348 (1898). Leaf-blades pu- bescent or glabrous beneath. Spikelets to 7 mm. long, sterile and fertile lemmas strongly carinate, the keel prolonged as a distinct scabrous awn. LA Paz: NOR YUNGAS: Bella Vista, Hitchcock 22753 (GH). 2. C. delicatula Hitchcock in Contrib. U. S. Nat. Herb. 23: 309 (1927). Leaf-blades glabrous beneath. Spikelets 5-6 mm. long; sterile lemmas ending in a short subulate point. LA PAZ: NOR YUNGAS: Bella Vista, Hitchcock 22748 (GH; type-number). 3. C. spicata Munro in Trans. Linn. Soc. 26: 60 (1868). Leaf- blades glabrous beneath, the margins densely ciliolate. Spikelets about 6 mm. long. LA PAZ: LARECAJA: Mapiri, Rusby 196 (Hitchcock). 4. C. peruviana E. G. Camus, Bamb. Monogr. 88, t. 53B (1913). Leaf-blades sparsely pilose beneath, the apex long-setaceous. Spikelets glabrous, to 10 mm. long, the lemmas shortly mucronate. LA PAZ: NOR YUNGAS: Bella Vista, Hitchcock 22746 (GH), 22744 (GH). 5. C. serrulata Pilger in Engler, Bot. Jahrb. 25: 719 (1898). Leaf- blades glabrous beneath. Spikelets about 7 mm. long, the lemmas mucronate. LA PAZ: MURILLO: La Paz, Rusby 10 (Hitchcock); NOR YUNGAS: Bella Vista, Hitchcock 22750 (GH) ; Coroico, Bang 2348 (GH) ; SUR YUNGAS: Colaya, 1810 m., Mexia 4298 (GH). 6. C. scandens Kunth, Syn. Pl. Aequin. 1: 254 (1822). C. quitensis var. patentissima Hack. in Fedde, Rep. Sp. Nov. 6: 161 (1908). Leaf- blades somewhat pilose beneath, the apex setaceous. Spikelets about 8 mm. long. LA Paz: NOR YUNGAS: Unduavi, 3300 m., Buchtien 4185 (GH); Bella Vista, Hitchcock 22747 (GH). 4. Bromus L. Annual or perennial, the culms erect or spreading, sometimes prostrate. Inflorescence a terminal racemose panicle, the spikelets several- to many-flowered, the puberulent rachilla disarticulating above the unequal glumes, the lemmas awned or awnless, glabrous or pubescent, the uppermost floret a sterile lemma or a rudiment. a. Glumes scabridulous to scabrous, but not puberulent nor pubescent. b. Lemmas pubescent, awn twisted. sss. 1. B. Trini b. Lemmas not pubescent or only very slightly so, awn not twisted. c. Leaf-blades to 1 cm. wide; lemma tuberculate-scabridulous. .... uM 2. B. segetum. c. Leaf-blades to 5 mm. wide; lemma not tuberculate-scabridu- lous. Meme eene S. B. catharticus. a. Glumes puberulent to pubescent. 1966] Flora of Bolivia — Foster 111 d. Inflorescence-rachis and pedicels short-pubescent. .... 4. B. lanatus. d. Inflorescence-rachis and pedicels scabridulous but not pubescent. rue d evotn ceklestesevi cobobuis TTT 5. B. pitensis. 1. Bromus Trinii Desv. in Gay, Fl. Chile, 6: 441 (1858). Annual, to 6 dm. high, the culms, inflorescence-rachis and pedicels glabrous. Leaves sparsely pilose on the sheaths, the blades pilose, at least when young. Spikelets to a little over 2 cm. long, about 7-flowered, the uppermont floret reduced to a narrow sterile lemma. First glume about 1.1 em. long, 1-nerved; second glume about 1.3 cm., 3-nerved, the nerves green. Lowest lemma about 1 cm. long, 2-toothed, the teeth very evident, the awn about 1.4 em. long. Palea about 9 mm. long, bicarinate, the keels ciliate near the apex. LA PAZ: LARECAJA: Sorata, Mandon 1370 (US). 2. B. segetum HBK. Nov. Gen. & Sp. 1: 151 (1816). Perennial, the panicle to 2 dm. long, culms, rachis and pedicels glabrous. Leaves glabrous, the blades scabridulous beneath. Spikelets to 1.5 em. long, 2.4-flowered, the uppermost floret reduced to a rudiment. First glume 5 mm. long, 1-nerved, the second glume to 8 mm. long, 3-nerved. Lowest lemma about 1.2 cm. long, shortly bidentate, awn about 2 mm. long. Palea as long as lemma, bidentate, bicarinate, the keels ciliolate. COCHABAMBA: CHAPARE: La Aduana, 3000 m., Steinbach 9533 (GH). 3. B. catharticus Vahl, Symbol. Bot. 2: 22 (1791). B. unioloides HBK. Nov. Gen. & Sp. 1: 151 (1816). B. angustatus Pilger in Engler, Bot. Jahrb. 25: 719 (1898). Annual or perennial, the culms glabrous, the rachis and pedicels scabridulous, the open panicles to 3 dm. long. Leaf-sheaths glabrous, blades scabridulous beneath. Spikelets about 2.5 cm. long, with up to 9 florets, the uppermost a rudiment. First glume to 9 mm. long, 3-nerved, the second glume to 1 cm., 5-nerved. Lowest lemmas to 1.4 cm. long, not toothed, awn about 5 mm. long, upper lemmas awnless. Palea to 1.3 em. long, bidentate, bicarinate, the keels long-ciliate. LA PAZ: MURILLO: La Paz, 3700 m., Buchtien 8533 (GH). COCHABAMBA: CERCADO: Valle de Cochabamba, Steinbach 8798 (GH). TARIJA: CERCADO: Calderillo, 3200 m., Fiebrig 2904 (GH). Very variable in size, spikelet-length and awn-length. Many lemmas awnless. 4. B. lanatus HBK. Nov. Gen. & Sp. 1: 150 (1816). B. Pflanzü Pilger in Engler, Bot. Jahrb. 49: 189 (1912). Perennial, to 6 dm. high, the leaf-blades somewhat villous. Panicle very lax, 10-20 cm. long. Spikelets to 2 cm. long, with about 6 florets, the uppermost a rudiment. First glume to 1 cm. long, 3-nerved, the second glume to 1.3 cm. long, 5-nerved. Lowest lemmas about 1.2 cm. long, densely villous along the margins, shortly bidentate, the awn 4-5 mm. long. Palea about 1 cm. long, bicarinate, the keels ciliate. LA PAZ: INGAVI: Comanche, Asplund 6459 (Hitchcock). TARIJA: CERCADO: Calderillo, 3200 m., Fiebrig 2904 in part (GH). 5. B. pitensis HBK. Nov. Gen. & Sp. 1: 152 (1816). B. Buchtieniüi ssssssesssesses 112 Rhodora [Vol. 68 Hack. in Fedde, Rep. Sp. Nov. 11: 30 (1912). Perennial, to 1.5 m. high. Leaf-blades glabrous or somewhat pilose, Panicles lax, up to 3 dm. long. Spikelets to 2.5 cm. long, up to 9-flowered. First glume about 6 mm. long, l-nerved, the second glume about 9 mm. long, 3- nerved. Lowest lemmas to 1.3 cm. long, bidentate, sparsely pubescent or villous on the lower half, the awn 4 mm. long. Palea about 1.1 em. long, bidentate, bicarinate, the keels ciliate. Anthers villous at base. LA PAZ: LARECAJA: Sorata, 2600-2800 m., Mandon 1365 (GH); MURILLO: La Paz, 3800 m., Buchtien 518 (GH). 5. Brachypodium Beauv. Brachypodium mexicanum Link, Hort. Berol. 1: 41 (1827). Peren- nial, straggling, to 6 dm. high. Leaves glabrous, the blades usually much shorter than the inflorescence, scabridulous beneath, acute, to 1.2 cm. wide, usually narrower. Inflorescence a solitary, terminal raceme of a few distant spikelets, these at first appressed, ultimately spread- ing or deflexed, the raceme to 10 cm. long. Spikelets to 2.5 cm. long (excluding the awns), subterete, to 10-flowered, the glabrous or upper portion and downward along the lateral nerves, awned, the minutely puberulent rachilla disarticulating above the glumes; glumes unequal, the first about 6 mm. long, the second to 7 mm. long, both awnless; lemmas about 9 mm. long, 5-7-nerved, puberulent on the awn about 4-6 mm. long, the 2 uppermost florets reduced and sterile; palea as long as the lemma, bicarinate, the keels densely ciliate. LA PAZ: LARECAJA: Sorata, Holway 544 (US). COCHABAMBA: CERCADO: Cochabamba, Tequifia, Hitchcock 22855 (US), Holway 409 (vs). 6. Festuca L. Annual or perennial, of varying heights, Leaves flat or involute, shorter than or exceeding the inflorescence. Inflorescence a terminal racemose panicle, the spikelets few. to several-flowered, the puberu- lent rachilla disarticulating above the unequal or subequal glumes. Lemmas firm to somewhat indurate in texture, awned, awn-tipped or awnless, the uppermost reduced and sterile, a. First glume 1 mm. long or less; lemmas ciliate at apex, tuber- culate-scabridulous. ...........ssssseee 1. F. megalura. a. First glume at least 1.5 mm. long; lemmas not ciliate at apex, not tuberculate-scabridulous. b. Lemmas with awns at least 5 mm. long. ............ 2. F. ulochaeta. b. Lemmas awn-tipped (about 1 mm.) or awnless. c. Rachis and pedicels scabrid to pubescent; lemmas not pubes- cent along margin. d. Lemmas (at least some) awn-tipped. e. Leaf-blades flat. sss 3. F. procera. e. Leaf-blades involute and pungent. f. Lemmas glabrous or puberulous at apex. .. 4. F. sublimis. 1966] Flora of Bolivia — Foster 113 f. Lemmas scabridulous at apex. seese 5. F. rigescens. d. Lemmas awnless ........../"7 DEA ES 1. F. dolichophylla. c. Rachis and pedicels glabrous; lemmas pubescent along margin. vosueceussssecensssscsnsccnccsensceascscsesssconscsassnecsonssonsesnasseeeees 6. F. orthophylla. 188 (1848). Annual. Panicles narrow, usually over 10 cm. long, often exceeded by the uppermost flat glabrous leaf-blades. Spikelets (excluding awns) about 1 cm. long, with 4-5 florets. Glumes unequal, the first about 1 mm. long or less, the second, 2.5-3 mm. long. Lowest lemmas to 5 mm. long with a 1.5 cm, scabrid awn, upper lemmas long-ciliate near the apex, most lemmas tuberculate-scabridulous (at least in dried material). Palea as long as lemma, bicarinate, the ciliate keels nearly marginal. LA PAZ: LARECAJA: vic. Sorata; Ullon- tiji, 2650-2900 m., Mandon 1363 (GH). COCHABAMBA: CHAPARE: Inca- chaca 2250 m., Steinbach 9498 (GH); CERCADO: south side of Mt. Tunari, 3300 m., Steinbach 9850 (GH). Potosi: CERCADO: Potosi, Cár- denas 203 (GH). TARIJA: CERCADO: Calderillo, 3200 m., Fiebrig 3159 (GH). 2. F. ulochaeta Steud. Syn. Pl. Glum. 1: 305 (1854). Perennial, up to 1 m. high, the culms, rachis and pedicels scabrid. leaf-blades sometimes exceeding the inflorescence, to 5-8 mm. wide, scabrid be- neath. Panicle fairly lax, to 2 dm. long. Spikelets (excluding awns) about 8 mm. long, 4-5 flowered, Glumes 1.5 and 2.5 mm. long. Lowest lemmas 5-6 mm. long, convex to rounded, 3-nerved, the central nerve scabridulous at apex, awns very filiform, flexible, 5-8 mm. long. Palea as long as lemma, bicarinate, the keels ciliolate. LA PAZ: MURILLO: Unduavi, Buchtien 6415 (Hitchcock). COCHABAMBA: CHA- PARE: Incachaca, 2300 m., Steinbach 8976 (GH). 3. F. procera HBK. Nov. Gen. & Sp. 1: 154 (1816). F. Fiebrigü Pilger in Engler, Bot. Jahrb. 37: 510 (1906). Perennial, to 1-2 m. high, the panicle to 3 dm. long; rachis and pedicels short-pubescent, leaf-blades narrow, flat or somewhat folded, scabrid beneath. Spike- lets about 8 mm. long, 3-5-flowered. Glumes 4 mm, and 5 mm, long, the first 1-nerved, the keel scabrid near the apex, the second 3-nerved, at least the keel scabrid above. Lowest lemmas to 6-7 mm. long, rounded, several-nerved, scabrid to scabrid-puberulent, especially on the upper portion, awn-tipped. Palea as long as lemma, bicarinate, the keels ciliolate, bidentate, the apex puberulous. TARIJA: AVILES: Pinos, 2800 m., Fiedrig 3117 (GH; type number of F. Fiebrigii), 3118 (GH). 4. F. sublimis Pilger in Engler, Bot. Jahrb. 25: 718 (1898). Perennial, caespitose, to 1 m. high, the involute, glabrous to scabrid, pungent leaf-blades sometimes exceeding the inflorescence. Panicle fairly lax, to 2.5 dm. long, rachis and pedicels scabrid to short- pubescent, Spikelets about 1 cm. long, 4-flowered (in most). Glumes 4 mm. and 5 mm. long, the second minutely ciliolate and sometimes 114 Rhodora [Vol. 68 scabridulous. Lowest lemmas about 7 mm. long, awnless, the upper ones awn-pointed (1 mm.), glabrous to puberulous. Palea as long as lemma, bidentate, puberulent, bicarinate, the keels ciliate. LA Paz: LARECAJA: vic. Sorata; Chileani, Mandon 1362 bis (GH). 5. F. rigescens (Presl) Kunth, Rév. Gram. 1: suppl. XXXI (1830). Caespitose perennial, to 4 dm. high, leaf-blades involute, pungent, glabrous, sometimes exceeding the narrow 5-20 cm. panicle, the rachis and pedicels puberulent. Spikelets to 1.1 cm. long, about 4-flowered. Glumes 6 mm. and 7 mm. long, the second scabridulous near the apex. Lowest lemmas to 8 mm. long, rounded, the apex scabridulous, some- times slightly 2-toothed, awnless to awn-pointed. Palea shorter than lemma, puberulous on both surfaces, bicarinate, the keels ciliolate. LA Paz: YUNGAS, Bang 173 (GH); MURILLO: Palca, 3700 m., Hitchcock 22560 (GH). Potosi: CERCADO: Potosi, 4000 m., Cardenas 208 (GH). 6. F. orthophylla Pilger in Engler, Bot. Jahrb. 25: 717 (1898). F. orthophylla var. boliviana Pilger in Engler, Bot. Jahrb. 37: 508 (1906). Caespitose perennial, to 5 dm. high, leaves involute, glabrous, firm, pungent, often equaling or exceeding the inflorescence. Panicle narrow, the branches ascending, rachis and pedicels glabrous. Spike- lets about 1 cm. long, 4-6-flowered. Glumes about equal, 6 mm. long, the first l-nerved, the nerve scabrid, margins slightly ciliate, with an apical tuft of hairs, the second 3-nerved, broader than the first but otherwise similar. Lowest lemmas about 7 mm. long, awnless, pubescent along the margin. Palea as long as lemma, pubescent at apex, bicarinate, the keels ciliate, at least on the uppper half. TARIJA: AVILES: Puna Patanca, 3700 m., Fiebrig 3192 (GH; type-number of F. orthophylla var. boliviana). 7. F. dolichophylla Presl, Rel. Haenk. 1: 258 (1830). F. laeteviridis Pilger in Engler, Bot. Jahrb. 37: 510 (1906). F. Buchtienii Hack. in Fedde, Rep. Sp. Nov. 6: 160 (1908). F. Pflanzii Pilger in Engler, Bot. Jahrb. 49: 188 (1912). Caespitose perennial, 4-10 dm. high, the involute pungent leaves glabrous or scabrid, occasionally equaling the inflorescence. Panicle narrow, the branches appressed or spreading, to 15 em. long, the rachis and pedicels scabridulous. Spikelets to 1.1 cm. long, 3-4-flowered. Glumes unequal, the first to 4.5 mm. long, apically scabrid, the second to 7 mm. long, ciliolate at base, apically scabrid. Lowest lemmas to 7 mm. long, awnless, scabrid toward apex, but otherwise glabrous. Palea as long as lemma or longer, bidentate, bicarinate, the keels minutely ciliolate. LA Paz: LARECAJA: vic. Sorata; Millipaya, 3700-4200 m., Mandon 1361 (GH) ; MURILLO: Palca, 3700 m., Hitchcock 22568 (GH); La Paz, 3300 m., Bang 33 (GH), 4100 m., Buchtien 428 (Gu). COCHABAMBA: CERCADO: Cerro Molle-Molle, Stein- bach 4093 (GH), 3800 m., Steinbach 4062 (GH); AYOPAYA: Tabacruz, 4200 m., Steinbach 9760 (GH); south side of Mt. Tunari, 3600 m., Steinbach 9787 (GH), 9784 (GH). POTOSÍ: SUR CHICHAS: Tupiza, 2700 m., Fiebrig 2955 (GH; type-number of F. laeteviridis). 1966] Flora of Bolivia — Foster 115 7. Aphanelytrum Hack. Aphanelytrum procumbens (Hack.) Hack. in Oesterr. Bot. Zeitschr. 52: 13 (1902). A straggling perennial, culms decumbent, sometimes stoloniferous, to 8 dm. high. Leaves glabrous, not scabridulous, much shorter than the inflorescence, the short acute blades about 3 mm. wide. Inflorescence a lax racemose panicle up to 2 dm. long, the very slender branches distant and bearing 2-3 spikelets at their ends. Spikelets about 1.2 cm. long, pedicellate, 2-flowered, the naked rachila disarticulating above the glumes, the second floret distant from the first; glumes greatly reduced, minute; lemmas to 7 mm. long, long-acute but not awned, more or less hyaline, several-nerved, the nerves green; palea similar to the lemma but shorter, 1-nerved, the nerve green. LA PAZ: MURILLO: Unduavi, 3300 m., Buchtien 4268 (US); NOR YUNGAS: Bella Vista, Hitchcock 22756 (us). 8. Puccinellia Parl. Perennials, low-growing, the more or less decumbent culms up to 20 em. high. Leaves glabrous, short, very narrow, the uppermost blade on a culm equaling or sometimes exceeding the inflorescence and often enveloping its base. Inflorescence a short, dense, race- mose panicle up to 12 cm. long, the branches erect, appressed, the pedicels often (or usually) short, thick, sulcate. Spikelets 2-3- flowered, all florets perfect, glumes present, the naked rachilla dis- articulating above the glumes; lemma rounded, the nerves obscure, glabrous, but sometimes puberulent at the base, awnless. Spikelets about 2 mm. long; lemma about 1 mm. long. .... 1. P. parvula. Spikelets 4-5 mm. long; lemma about 2.5 mm. long. .... 2. P. oresigena. 1. Puccinellia parvula Hitche. in Contrib. U. S. Nat. Herb. 24: 325 (1927). Spikelets 2.3.flowered. Glumes unequal, the first about 0.5 mm. long, the second, 1 mm. long. Lemmas elliptic-oblong, the apex obtuse or lacerate; palea about as long as the lemma. PoTosi: SUR CHICHAS: Atocha, 3300 m., Hitchcock 22878 (US; type). 2. P. oresigena (Phil). Hitche. in Contrib. U. S. Nat. Herb. 24: 326 (1927). Spikelets 3-flowered. Glumes unequal, the first about 1 mm. long, the second, 1.5 mm. long. Lemmas acute. PoTosi: SUR CHICHAS: Atocha, Hitchcock 22879 (GH). 9, Amphibromus Nees Amphibromus scabrivalvis (Trin.) Swallen in Amer. Journ. Bot. 18: 413 (1931). A rhizomatous perennial, the culms erect (the base sometimes decumbent), to 1 m. or more in height. Leaves glabrous, the lowest blade to 40 cm. long, the uppermost reduced, 2-6 mm. wide. Inflorescence a rather lax racemose panicle to 20 cm. long, the slender branches often spreading or drooping. Spikelets pedicellate, up to 7-flowered, the florets distant, the uppermost floret sterile, to 1.5 cm. 116 Rhodora [Vol. 68 long (excluding the awns), the rachilla unilaterally short-villous, with an apical ring of hairs below each floret, disarticulating above the glumes. Glumes unequal, the first about 4 mm. long, the second, 5 mm. long, the apex erose. Lemmas about 7 mm. long, bifid to the middle or below, the 1.2 em. twisted geniculate awn arising from the base of the sinus, up to 9-nerved, scabrid, the apices erose; palea shorter than the lemma, bicarinate. LA PAZ: MURILLO: La Paz, Buchtien 6422 (us). 10. Poa L. Annual or perennial, one species dioecious. Culms usually but not always exceeding the leaves. Leaf-blades flat or involute, usually scabridulous beneath. Inflorescence a terminal racemose panicle, narrow and condensed or more open and spreading, the rachis and pedicels glabrous, puberulent or scabridulous, sometimes sparsely so. Spikelets 2-6-flowered, the glumes usually unequal, the glabrous or puberulent rachilla disarticulating above the glumes; fertile lemmas below the sterile lemma or rudiment. a. Plants dioecious. l, 1. P. Buchtienii. a. Plants not dioecious. b. Dwarf plants, about 4 cm. high; glumes and lemmas subquadrate. Tete tte e eee e tere eH MILLIA d. Spikelets to 6 mm, long, 4-5-flowered; annual. .... 9. P. annua. d. Spikelets to 3 mm. long, 3-flowered; perennial. nes. 10. P. denticulata. c. Rachis and pedicels puberulent or scabridulous. e. Spikelets subquadrate, as broad as long. ...... 5. P. Hieronymi. e. Spikelets longer than broad (but note no. 11). f. Spikelets 2-flowered. sss 12. P. candamoana. f. Spikelets almost always more than 2-flowered. g. Lemmas not villous on any part. h. First glume about 2.5 mm. long. ............ 6. P. Lilloi. h. First glume about 4.5 mm. long. ........ 4. P. gymnantha. g. Lemmas with at least basal villosity. i. Rachilla puberulent. ........ s 8. P. asperiflora. i. Rachilla glabrous. j. Spikelets to 7 mm. long, often almost as broad as long. sese 11. P. horridula. j. Spikelets 4-5 mm. long, definitely longer than broad. k. Glumes equal or subequal. ........ 7. P. pratensis. k. Glumes unequal. ss 2. P. scaberula. 1. Poa Buchtienii Hack. in Fedde, Rep. Sp. Nov. 11: 29 (1912). P. Buchtienii var. subacuminata Hack. in Fedde, Rep. Sp. Nov. 11: 30 (1912). Perennial, to 8 dm. high, the leaf-blades flat, to 3 mm. wide. 1966] Flora of Bolivia — Foster 117 Rachis and pedicels scabrid. Staminate spikelets: about 5 mm. long, 3-5-flowered, the rachilla glabrous. First glume 2.5 mm. long, 1- nerved, the nerves scabrid, at least on the upper portion, the second glume about 3.5 mm. long, broader than the first, obscurely 3-nerved, the keel scabridulous. Lowest lemmas about 3.5 mm. long, obscurely nerved, the mid-nerve scabridulous on the upper part, the apex erose. Palea 3 mm. long, the 2 keels ciliolate. Pistillate spikelets: about 6-7 mm. long, 5-flowered, the rachilla glabrous. First glume about 25 mm. long, 1-nerved, the keel scabridulous, the second glume 3 mm. long, 3-nerved, the keel scabrid. Lowest lemmas about 4.5 mm. long, densely long-pilose at base and on the lower half of the keel, the apex erose to acute. Palea shorter than the lemma, the keels long-pilose below and ciliate above. LA PAZ: MURILLO: La Paz, 3700 m., Buchtien 2468 (GH) ; Palea, 3700 m., Hitchcock 22559 (GH), 22570 (GH). One culm on the Gray Herbarium sheet of Buchtien 2468 bears staminate spikelets with the basal villosity of the pistillate spikelets. 2. P. scaberula Hook. f. Fl. Antarct. 378 (1847). Annual, to 5 dm. high, the leaf-blades to 2 mm. wide and scabrid beneath, the rachis and pedicels densely scabrous. Spikelets to 4 mm. long, about 4- flowered, the rachilla glabrous. Glumes 2 and 2.5 mm. long, the keels scabrid. Lowest lemmas 2.5 mm. long, villous at base and halfway up the keel, the lateral nerves often somewhat villous basally. Palea much shorter than the lemma, the keels ciliate. LA PAZ: LARECAJA: vic. Sorata; Lancha de Cochipata, 2000-3700 m., Mandon 1336 (GH); MURILLO: La Paz, 3670 m., Buchtien (GH). 3. P. humillima Pilger in Engler, Bot. Jahrb. 37: 378 (1906). Very dwarf, caespitose perennial, the leaf-blades hardly 2 cm. long, involute, glabrous, the panicle not over 1 em. long, rachis and pedicels puberu- lent. Spikelets to 5 mm. long, 3.4-flowered, the rachilla glabrous. Glumes equal, 3.5 mm. long, subquadrate, apex coarsely and erosely toothed, obscurely 3-nerved. Lemmas about 4.5 mm. long, subquad- rate, 5-nerved, the apex erosely toothed. Palea shorter than lemma, bicarinate. LA PAZ: LARECAJA: vic. Sorata; near Anilaya, 4500 m., Mandon 1353 (GH). 4 P. gymnantha Pilger in Engler, Bot. Jahrb. 56, Beibl. 123: 28 (1920). P. perligulata Pilger in Notizbl. 11: 779 (1933). P. gym mantha var. aperta Pilger in Notizbl. 11: 780 (1933). Caespitose perennial to 20 cm. high, leaf-blades flat or involute, glabrous, 5-10 em. long, the panicles 3-7 cm. long, rachis and pedicels scabridulous- puberulent. Spikelets 5-7 mm. long, 2-3 (-4)-flowered. Glumes un- equal, the first to 4.5 mm. long, 1-nerved, scabridulous at apex, the second to 5 mm. long, 3-nerved, scabridulous at apex. Lemmas acute to subacute, 5.5 mm, long, 5-nerved, the keel scabrid-ciliate, the rest 118 Rhodora [Vol. 68 scabridulous. Palea as long as lemma, scabridulous, the 2 keels scabrid- ciliolate. LA PAZ: LARECAJA: vic. Sorata; Llachisani, 4200 m., Mandon Mandon 1351 (GH); MURILLO: vie. La Paz, 5000 m., Mandon 1352 ma COCHABAMBA: AYOPAYA: Tabacruz, 4200 m., Steinbach 9761 GH). 5. P. Hieronymi Hack. in Oesterr. Bot. Zeitschr, 52: 380 (1902). P. myriantha Hack. in Stuckert in Anal. Mus. Nac. Buenos Aires, 13: 517 (1906). Perennial, to 2 m. high, leaf-blades Scabrous, the flat blades 8-10 mm. wide, scabrous beneath, the very lax open panicle to 4 dm. long, the ultimate branches extremely filiform, the rachis and pedicels sparsely scabridulous. Spikelets, subquadrate, to 5 mm. long and broad, 5-6-flowered, the rachilla glabrous. Glumes equal, about 5 mm. long, the first l-nerved, the nerve scabrid, the margin scabrid- ciliolate, the second 3-nerved, the nerves scabrid. Lemmas acute, to 3.5 mm. long, the nerves Sscabrid, the margins ciliolate, a small tuft of tangled hairs at base of central nerve. Palea decidedly shorter than the lemma, the 2 keels ciliolate. COCHABAMBA: CHAPARE: Inca- chaca, 2700 m., Steinbach 8956 (GH), 2500-3000 m., Steinbach 9517 (GH). 6. P. Lilloi Hack, in Stuckert in Anal. Mus. Nac. Buenos Aires, 21: 153 (1911). Perennial, 10-20 (-30) em. high, leaves glabrous, the very narrow blades involute, panicle 2-4 cm. long, rachis and pedicels Scabrid, sometimes sparsely so. Spikelets about 5 mm. long, about 4- flowered, rachilla glabrous to slightly puberulent. First glume 2.5 mm. long, 1-nerved, the keel scabrid, the second glume to 3 mm., 3- nerved, scabridulous. Lemmas to 3.0 mm. long, 3-5-nerved, scabri- dulous, especially on the nerves. Palea nearly as long as the lemma, the 2 keels ciliolate, PoTosí: sUR CHICHAS: Atocha, Hitchcock 22931 (Hitchcock). The description given is based on an isotype, Lillo 5619 (GH). 7. P. pratensis L., Sp. Pl. 67 (1753). P. boliviensis Hack. in Fedde, Rep. Sp. Nov. 11: 25 (1912). Rhizomatous perennial, to 5 dm. high. Leaf-blades to 3 mm. wide, glabrous or scabridulous beneath. Panicles to 10 em. long, the rachis and pedicels scabridulous. Spikelets about 5 mm. long, 3-5-flowered. The rachilla glabrous or sparsely and minute- ly puberulent. Glumes subequal, about 3 mm. long, the keels scabri- dulous, the first l-nerved, the second 3-nerved. Lowest lemmas about 3 mm. long, 5-nerved, the keel basally long-villous, the upper half scabrous, marginal nerves basally villous. Palea nearly as long as the lemma, the 2 keels ciliolate. LA PAZ: MURILLO: San Jorge, 3550 m., Buchtien 236 (GH). 8. P. asperiflora Hack. in Fedde, Rep. Sp. Nov. 11: 28 (1912). P. Pflanzii Pilger in Engler, Bot. Jahrb. 49: 187 (1912). Caespitose perennial to 5 dm. high, the involute leaf-blades sometimes exceeding 1966] Flora of Bolivia — Foster 119 the inflorescence, scabrid. Panicles fairly open, generally 10 cm. or less in length, the rachis and pedicels scabridulous. Spikelets about 7 mm. long, 4-flowered, the rachilla puberulent. First glume about 3.5 mm. long, 1-nerved, the second glume about 5 mm. long, 3-nerved, the keel slightly scabridulous. Lowest lemmas about 5 mm. long, 5-nerved, the keel basally short-villous, with scattered pubescence on the lateral nerves. Palea about 4 mm. long, the keels ciliolate. LA Paz: LARECAJA: vic. Sorata; Lacatia, 3360-4200 m., Mandon 1341 (GH); MURILLO: Pilaya, 4000 m., Hitchcock 22587 (GH). 9. P. annua L. Sp. Pl. 68 (1753). Annual, the leaf-blades to 3 mm. wide, sometimes scabridulous beneath, the panicles open, to 6 cm. long, the rachis and pedicels glabrous. Spikelets to 6 mm. long, 4-5- flowered, the rachilla glabrous to sparsely puberulent. First glume about 2.5 mm. long, 1-nerved, faintly scabridulous near the apex, the second glume 3 mm. long, 3.nerved. Lowest lemmas 3 mm. long, 5-nerved, the keel with few to many hairs at base and about halfway up, but not forming a basal cobweb. Palea nearly as long as lemma, bicarinate. LA PAZ: LARECAJA: Vic. Sorata; Achacache, 2650-3950 m., Mandon 1335 (GH). COCHABAMBA: CERCADO: Cochabamba, 2600 m., Steinbach 8787 (GH); CHAPARE: Incachaca, 2250 m., Steinbach 9496 (GH). SANTA CRUZ: VALLEGRANDE: Cerro de San Mateo, Comarapa, 3400 m., Steinbach 8486 (GH). 10. P. denticulata Hack. in Fedde, Rep. Sp. Nov. 11: 27 (1912). Perennial, to 25 dm. high, leaf-blades 2-3 mm. wide, scabrid beneath. Panicles lax, to 8 cm. long, primary branches whorled, rachis and pedicels glabrous. Spikelets 4 mm. long, 3.flowered. Glumes 2 and 2.5 mm. long, the first 1-nerved, the second, 3-nerved. Lowest lemmas to 3 mm. long, ovate, obtuse, the apex erosely toothed, 5-nerved, the basal hairs reaching about one-third the length of the lemma. Palea as long as the lemma, bidentate, the keels scabridulous. LA PAZ: MURILLO: Unduavi, 3200 m., Buchtien 2584 (US; type-number). 11. P. horridula Pilger in Engler, Bot. Jahrb. 37: 506 (1906). P. androgyna Hack. in Fedde, Rep. Sp. Nov. 6: 159 (1908). P. dume- torum Hack. in Fedde, Rep. Sp. Nov. 11: 27 (1912). P. dumetorum var. unduavensis Hack. in Fedde, Rep. Sp. Nov. 11: 27 (1912). Perennial, to 1-1.5 m. high, leaf-blades to 8 mm. wide, scabridulous beneath. Panicle to 2 dm. long, the rachis and pedicels scabridulous. Spikelets to 7 mm. long, often about as broad as long, 4-5-flowered, the rachilla glabrous. First glume to 2.5 mm. long, 1-nerved, the nerve scabridulous, the second to 3 mm. long, 3-nerved, the nerves scabridulous. Lowest lemmas to 3.5 mm. long, 5-nerved, pilose on lower half of keel and along lower marginal nerves, the keel scabridulous on upper half. Palea as long as the lemma, the keels ciliate. LA PAZ: LARECAJA: vic. Sorata; Cochipata, 2700-3900 m., Mandon 1339 (GH) ; MURILLO: La Paz, Buchtien 846 (GH; type-number of P. androgyna), 3800 m., Buchtien 168 (GH) ; Unduavi, 3200 m., Buchtien 2583 (GH; 120 Rhodora [Vol. 68 type-number of P. dumetorum var. unduavensis) ; NOR YUNGAS: Bella Vista, Hitchcock 22754 (GH). 12. P. candamoana Pilger in Engler, Bot. Jahrb. 37: 381 (1906). Perennial, leaf-blades to 3 mm. wide, scabridulous beneath. Panicle to 15 em. long, but usually much Shorter, rachis and pedicels scabrid. Spikelets to 5 mm. long, 2-flowered, the rachilla sparsely villous. Glumes 3 and 4 mm. long, acute, the second much broader than the first. Lowest lemmas about 4 mm. long, villous on the lower half, the first floret sometimes not perfect but staminate. Palea shorter than the lemma, the keels ciliate. LA PAZ: MURILLO: La Paz, 3800 m., Buchtien 8536 (GH); NOR YUNGAS: Pongo, 3640 m., Hitchcock 22769 (GH). Errata: Volume 67 - p. 351 line 22 Daura should read Datura - p. 356 line 12 Gleichena should read Gleichenia / p. 868 Fig. 4 Caption should read Mean temperature for January in ^F, (after Hare 1952) - p. 372 line 27 sessiliflora should read sessilifolia p. 373 line 13 macrocarpa should read macrocarpon p. 409, 411 running head thyoites should read thyoides Volume 68, No. 773 including pages 1-120, was issued April 6, 1966. “wo OW REFERENCE LIBRARY ^. JUL 1 X 1966 LA TES — Dodova JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALBION REED HODGDON, Editor-in-Chief ALBERT FREDERICK HILL > STUART KIMBALL HARRIS RALPH CARLETON BEAN ROBERT CRICHTON FOSTER | Associate Editors ROLLA MILTON TRYON RADCLIFFE BARNES PIKE LORIN IVES NEVLING, JR. Vol 68 >`- April-June, 1966 No. 774 CONTENTS: New Data on North American Oak Ferns, Gymnocarpium Warren H. Wagner, Jm. ssssesrssessssesssesronsseereesesoessovereeeesereeseeeveese 121 Lactuca muralis in New England James P. Poole ................ 138 Cuchumatanea A New Genus of Compositae (Heliantheae) C. E. Seidenschnur and J. H. Beaman ............ eere 139 Seed Germination of Shortia galacifolia T. & G. Under Controlled Conditions Mary H. Rhoades ..................... 147 The Application of the Linnaean Names of Some New World Species of Euphorbia Subgenus Chamaesyce Derek Burch 1... eerte eee eee ee eese ta on ase tetra eee eP Pase eene eee eaae sepan ea en 155 (Continued on Inside Cover) The Nem England Botanical Club, Inc. Botanical Museum, Oxford St., Cambridge 38, Mass. iini BEREN 1 CONTENTS: — continued Identity of Witchweed in the Southeastern United States C. Earle Smith, Jv. ceccccscccsssssssssssscsssssescsssescsesesescscsescscseseseseseees 167 Bromus mollis and Allies in New England Frank C. Seymour |... eerte eher eren 168 Contribution to the Fungus Flora of Northeastern North America. IV Howard E. Bigelow and Margaret E. Barr |... 175 Notes on Rafinesque's Species of Lechea (Cistaceae) Robert L. Wilbur |... esee eren enne etat 192 Gentiana puberulenta sp. nov., A Known but Unnamed Species of the North American Prairies James S. Pringle ............ eee ettet ettet ntn tntntnta 209 New Chromosome Numbers in Zinnia and Sanvitalia Andrew M. Torres vicccccccssssssssssssssscsssssssssssessecsscecssssessccssseesssassce 215 Two New Naiads from Illinois and Distributional Records of the Naiadaceae Paul L. Fore and Robert H. Mohlen- brock csssiceranensncetscsantensseocivusiasisossscsabovssnnevosensesspeazsacaesmenmerserseecsvienes 216 Aquatic Vascular Plants New for Illinois Glen S. Winterringer crsccccsscscsssssessscscsssscsscsssssscsssscssasescscsssscesece 221 Studies in the Flora of Bolivia-IV (Continued) Robert C. Foster... eterne then 223 QTRbooora JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Vol. 68 April-June, 1966 No. 774 NEW DATA ON NORTH AMERICAN OAK FERNS, GYMNOCARPIUM WARREN H. WAGNER, JR.' The oak ferns, Gymnocarpium Newman, include few taxa. Except for one or more separate and endemic elements in eastern Asia (cf. Pichi-Sermolli, 1965. p. 148), there are only two basic species over most of the circumboreal range. Nevertheless, there are controversial points regard- ing both the generic treatment and the species taxonomy. The common oak fern, Gymnocarpium dryopteris (L.) New- man has broadly triangular leaf blades with the axes so sparsely glandular that they are usually designated as “glabrous.” The “limestone” oak fern, G. robertianum (Hoffm.) Newman, is far rarer and more sporadic in most of its range. It has narrower leaf blades, and the blade axes (and the lamina) are densely glandular. Both of these ferns are variable, both in habitat and morphology. Dwarfed plants of G. robertianum, especially those growing on dryish rock cliffs, may tend to mimic the leaf outline of G. dry- opteris. Fully developed, however, the characteristics of "Research supported by National Science Foundation Grant GB-2025. I wish to acknowledge the aid of Miss Sandra Smith and Mrs. Katherine Lim Chen, and the suggestions of Doctors E. T. Wherry, Eric Hultén, and Conrad V. Morton. The following herbaria kindly lent materials: Cranbrook Institute of Science, Gray Herbarium, Missouri Botanical Garden, U. S. National Museum, and the Natur- historiska Riksmuseet, Stockholm. Dr. J. H. Soper made it possible for me to study the oak ferns in the herbarium at the University of Tcronto. 121 122 Rhodora [Vol. 68 OWesernvess op Horass, Tu Uswis ot Michasaw GF WASHINGTON Gymnocarpium dryopteris New, Kl > Deer conirereua woods: at Creek turnorf, 9.5 ig, Sneg * OnRt'l, Forest, Abundant in we June 25, 1957, M. i . Vagner 53057 å, By Eruickeberg sna PLATE 1328 Fig. 1. Gymnocarpium dryopteris var. disjunctum. Voucher speci- men, n — 40, Wagner 63087 and Kruckeberg (MICH). 1966] Oak Ferns — Wagner 123 Eoo eL £ * 4 atej 1 ‘ Con -— a hit Bg Coinimetey, He c d. WARREN H WRONER, fa te 28 iiem afa fl We, (PH! PLATE 1329 Fig. 2. Gymnocarpium heterosporum n. sp. Type specimen. Wagner 283 (MICH). 124 Rhodora [Vol. 68 these plants are distinctive and most authors have upheld them as separate species. Good keys and brief descriptions are found in familiar manuals and guides (e.g., Tutin et al., 1964; Wherry, 1961); and the taxonomic characters are fairly well established. At least two recent writers (Lawalrée, 1950; and Boivin, 1962) have questioned whether the two widespread taxa are specifically different. This question will be discussed further below. Also, there has been some disagreement about the generic placement of the oak ferns, but this matter will not be dealt with. Suffice it to say, the combi- nation of characters does seem very well-marked, and in the past few years there has developed a widespread tendency to recognize Gymnocarpium as a distinct genus. ne question is what its relationships are. The base number of x — 40 is quite different from such thelypteroid ferns as Phegopteris (x = 30), Cyclosorus (x = 36), and such species of Thelypteris as T. palustris (x = 35) and 7. "l 3 1 | Fig. 3. Distribution map. Stars = very small spored G. dryopteris (less than 23.9 X 1.18 microns). Dots = G. heterosporum. 1966] Oak Ferns — Wagner 125 noveboracensis (x = 27). The chromosome number of Gymnocarpium resembles that of typical aspidioid ferns (e.g., Dryopteris, Cystopteris, Polystichum, Tectaria, Cteni- tis, etc.) with x = 40, 41, or 42. Elizabeth Root (1961) studied intermediates between G. dryopteris and G. robertianum and pointed out, for the first time, that “the majority of the spores of all intermediate specimens were found to be abortive." Since her study we have learned that the “hybrid” Gymnocarpium is much more frequent and widespread than had been heretofore realized, as is illustrated in the map (Fig. 3, dots). We have also discovered that there are three ploidal levels present in North American Gymnocarpium — the diploid, triploid, and tetraploid. These facts will be examined below in connection with the following questions: Are G. dryopteris and G. robertianum distinct species? Should the western representative of G. dryopteris be recognized as a separate variety or subspecies? What is the nature of the plants which are morphologically intermediate between G. dry- opteris and G. robertianum? THE DISTINCTNESS OF G. DRYOPTERIS AND G. ROBERTIANUM Boivin (op. cit.) is the most recent worker who en- deavored to interpret taxon robertianum as merely a variety of G. dryopteris. He did not find it possible to uphold taxon robertianum as a species because “Tous les caracteres donnés chevauchent largement, sauf celui de la glandulosite qui est assez net si l'on excepte 6 ou 8 spécimens (sur en- viron 200 examiné). Cette glandulosité est si fine et ces deux espéces sont si semblables que, sur une cinquantaine de spécimens glanduleux examinés, un bon quart étaient mésidentifiés." I should point out that herbarium collections of ferns are often misidentified. The fact that certain species (e.g., in Gymnocarpium, Dryopteris, and Botrychium) are commonly misidentified should not be used as a basis for taxonomic judgments. The reasons for upholding G. robertianum as a distinct species, rather than a variety of G. dryopteris, fall into two 126 Rhodora [Vol. 68 classes: (1) The morphology of well-developed specimens is unquestionably distinct, including characters of the over- all blade outline, the details of segment form, and the glandularity. Small, ecologically depauperated or juvenile, specimens may of course cause difficulties if only gross characters are used. But the glandularity of G. robertianum is evidently always a dependable character. (Some of the problems of detecting the extent of glandularity in these plants could be alleviated by eliminating the unfortunate herbarium practice of pasting down fronds, thus coating the delicate glands with a thin “varnish” which makes them practically invisible.) The only really intermediate plants in respect to glandularity are those to be discussed below, but these plants are intermediate in other respects also between G. dryopteris and G. robertianum and they are probably of hybrid origin. (2) The second reason for upholding G. robertianum is simply the behavior of the natural populations. The two species grow together in numerous localities in the Great Lakes area, in exactly the same habitats and growth conditions, and their distinctness is immediately evident to the field worker. The fact that these two taxa can exist together sympatrically without merging, and the fact that there is no genomic difference (both have n — 80 in this region) in ploidal level, shows that using the varietal rank to express their relationships would be stretching that category too far (cf. Wagner, 1960). The differences between these ferns are best seen in the rich, shady swamps of northern Michigan where they develop to the maximum size in the luxuriant, moist con- ditions. Mixed populations are well known in several locali- ties near the University of Michigan Biological Station (Cheboygan, and nearby Presque Isle Cos., Michigan). GYMNOCARPIUM DRYOPTERIS VAR. DISJUNCTUM The western North American representatives of G. dry- opteris, especially in Idaho, Oregon, Washington, U.S.A. and British Columbia, Canada, tend to develop unusually large fronds. The area is roughly that shown by the stars 1966] Oak Ferns — Wagner 127 Fig. 4. Tracings of spores. Inner line — exospore; outer line — perispore. R. Gymnocarpium robertianum, Mich., Voss 4709 (MICH). X. G. heterosporum, Pennsylvania, Darling in 1956 (MICH). D. G. dryopteris, Mich., Voss 4708 (MICH). of especially small-spored plants in Figure 3. In the most extreme form these plants become thrice-pinnate and the larger segments are more toothed than in typical G. dry- opteris. Under the guidance of Dr. Arthur R. Kruckeberg of the University of Washington, I had an opportunity to study this form in the field. The best plants (Fig. 1) were seen in the deep coniferous woods at Denny Creek turnoff along U.S. 10, in the Snoqualmie National Forest, King Co., Washington. Here the "giant" G. dryopteris flourishes in damp, mossy forest with such other ferns as Dryopteris dilatata. and Athyrium filix-femina, the latter two species also becoming remarkably large. Chromosome materials were obtained and squashes of meiosis showed clearly n = 40 pairs (Figs. 6 and 7, top illustrations). This is the first discovery of the diploid condition in the genus Gymno- carpium, all previous counts for which indicated the tetra- 128 Rhodora [Vol. 68 x A ; x G. ROBERTIANUM x x x x x x x x x x x x x x x x x x x x x x x X E x x x x | & e x x x x * t t t t t ———— G. DRYOPTERIS V. DRYOPTERIS +k RK KAR RAR KK RARER KK EK RA TA AX OO X OO XXX TERK AK AK XOOD X X OX ROC X FARK KX RA KKK PRM Ke 3. DRYOPTERIS V. DISJUNCTUM KE X X X x x x x + D G. DRYOPTERIS -- ALASKA AND YUKON 20 25 3o Conversion: n X 1. 18 microns Fig. 5. Frequency histograms of spore lengths (exospore only). Each cross equals a 10-spore sample from a separate collection (Dots = spore samples of G. continentale). A. G. robertianum, all col- lections. B. G. dryopteris, all collections from Colorado east to Europe. C. G. dryopteris, all collections from British Columbia and Oregon east to Alberta and Montana. D. G. dryopteris, all collections from Alaska and Yukon. (Conversion: n X 1.18 microns). 1966] Oak Ferns — Wagner 129 ploid condition of n = 80. (According to spore size, as shown in Fig. 5, A, the chromosome number of the Siberian G. continentale may also turn out to be diploid.) For several reasons — the diploid chromosome number, the smaller spores (Fig. 5), the larger, more divided leaf (Fig. 1) and the distinctive western range — I choose (with some reluctance, however) to uphold the plants with this combination of characters as a separate variety. Morton (1941, p. 217) pointed out that the name disjunctum was based upon specimens from Sitka, Alaska, “and refers to certain large, lax forms that are essentially tripinnate, with the lower tertiary pinnules somewhat spaced out." At the time of his writing, Morton was of the opinion that “they do not seem to be worth nomenclatural recognition . . . " Boivin (op. cit.), on the contrary, upheld the large form as a distinct variety. Although I am inclined to agree now that var. disjunctum (Led.) Ching should probably merit recognition, I must point out that the situation becomes very complicated in Alaska and the Yukon, where both small- and large-spored types occur in the same general area (note the frequency histograms in Fig. 5, D). In this region also, we find numer- ous dwarfed individuals of both types, so that the gross appearance typical of *'var. disjunctum” becomes modi- fied and indistinguishable in many specimens from *'var. dryopteris," using the respective spore sizes as the criterion. Thus many specimens from the far northwest of North America cannot be readily identified. Dwarfing obscures the gross characters of the fronds; and the overlap of spore sizes between the diploid and tetraploid conditions is such that many specimens could not be identified without chromosome counts. Our spore surveys have uncovered still other compli- cations, the full meaning of which must await further study. It turned out that a number of collections of what I had interpreted to be ordinary "var. dryopteris" have abortive spores which in appearance match those of the presumed hybrid fern to be discussed below (like those in Fig. 4, X). 130 Rhodora [Vol. 68 PLATE 1330 Fig. 6. New chromosome conditions in Gymnocarpium. Top: G. dryopteris var. disjunctum diploid, n — 40 pairs, Wagner 63087 and Kruckeberg (interpretation top left in Fig. 7). Bottom: G. hetero- sporum triploid, 3 x = c. 120, Wherry s.n., from Blair Co., Penn- sylvania, in 1956 (bottom left in Fig. 7). 1966] Oak Ferns — Wagner 131 f T ^ y% wis 2 * M w Pr `a ^ x é Ps » x 4». e ? .* $ à % > 4, oc em % ane Ny uw V. fe a> » "m na E" ^ 4 è wet f $ b a ef ut ox D Iw ro ow ap o Ly Use Pixs 4 48. y ; JJ ww ^ ett Y 4 n me Ap ata + + 9F $3 s, | 4 a * °F D "A . rl » d. » * 5 19 S ¢ o a ` o » 4^2? % 9796 o2 * g A uem ek OS $m e^ $$ k ` 45 E E h ene > 090 of r A = Y | 3. Rr i 1.3 ^. € eu We o A PES M Y 9, EN ò s oe P 4 oo T v vw & $ , RAE M 4o. ake. Ere d 2 ¥ de rom $90 o, ql s Fig. 7. Camera lucida interpretations of chromosome squashes. Top: Diploids, same locality as in Fig. 6. Bottom: Triploids, same locality as bottom photograph in Fig. 6. Miss Virginia M. Morzenti kindly examined these collections and confirmed that the morphology of the abortion is very similar in the “sterile” G. dryopteris and the putative cross, G. heterosporum. It seems possible that at least some of these collections may be apomictic derivatives of G. dry- opteris var. disjunctum (the diploid) X var. dryopteris (the tetraploid), assuming here, as elsewhere in this dis- cussion, that the type of var. disjunctum corresponds to the diploid, large-leaved western plant. We failed to find a single example of G. robertianum in all those that we studied which showed a similar abortion of spores. Thus, if one argued that these "sterile" G. dryopteris plants might re- present mutations that are propagated locally by rhizomes, we wonder why similar mutations were not found in the related G. robertianum. Those collections we found of pre- sumably "sterile" G. dryopteris will not be enumerated in detail here, but their geographical distribution is extremely wide as shown by the following records: Alaska (2 col- lections), Alberta (1), British Columbia (1), Wisconsin (1), Michigan (5), New York (4), Vermont (2), Maine 132 Rhodora [Vol. 68 (1), New Hampshire (1), Quebec (1), Newfoundland (2), and Nova Scotia (1). Further research on these curious “sterile” oak ferns with the morphology of G. dryopteris is greatly to be desired, especially to determine their cytologi- cal features and the mode of their reproduction. THE APPARENT CROSS OF G. DRYOPTERIS AND G. ROBERTIANUM This is the plant discussed by Root (op. cit.). As stated above, the apparent cross is much more frequent and wide- spread than we had previously realized. Its occurrence, especially in the western Great Lakes area, indicates that it is more than a rare hybrid, and I have concluded therefore that it should be given a binomial designation as an apomic- tic species, as follows: Gymnocarpium heterosporum n. sp. W. H. Wagner. Planta frondis segmentique forma et glandularitate intermedia inter G. dryopteridem et G. robertianum; spori atro-brunnei, magnitudine et forma valde irregulares. Type: PENNSYLVANIA: BLAIR CO., Canaan Station, limestone slope, W. H. Wagner 283 (transferred from University of Pennsylvania to MICH — 2 sheets, one fertile, the holotype (Fig. 2), and one sterile). Representative Collections: U.S.A.: PENNSYLVANIA: BLAIR CO, 2 mi. n. w. of Hollidaysburg, among limestone rocks, T. Darling, Jr. in 1956 — same locality as the type (MICH). MICHIGAN: MARQUETTE CO., ca. 6 mi. n. w. of Ishpeming, near Ropes Gold Mine, n. w. % sect. 29, T 46 N, R 27 W, in woods on igneous rock cliffs, E. G. Voss 4707, W. H. Wagner and D. J. Hagenah 9415 (MICH). WISCONSIN: BAYFIELD co., Orienta Falls, G. H. Conklin and M. F. Somerville 1127 (WIS); BARRON CO., Barron Hills, just n. w. of Lehigh at the base of quart- zite talus, N. C. Fasset 15817 (wis), R. M. Tryon, Jr. 4154 (WIS, GH, MO). MINNESOTA: CARLTON CO., Carlton, Carlton-Thompson Gorge, St. Louis R., M. F. Somerville in 1928 (MICH); ST. LOUIS CO., obser- vation tower at Ash River, Wagner 9034.5a (MICH); LAKE CO., Gooseberry Falls State Park, 15 mi. n. e. of Two Harbors, in crevice of diabase cliff, damp and shady, R. M. and A. F. Tryon, A. C. Faber 4889 (MO); 45 mi. n.e. of Two Harbors, Manitou Falls near Lake Superior, under ledge of diabase, damp and shady, R. M. and A. F. Tryon, A. C. Faber 4895 (M0). CANADA: ONTARIO. ALGOMA DISTRICT, rocky cliffs at edge of Soulier Lake, vicinity of Michipicoten Harbor, R. C. Rosie, H. M. Harrison, E. O. Hughes 1092 (TRT, GH). ONTARIO, THUNDER BAY DISTRICT, 1 mi. n. of Little Pigeon Bay, Crooks Twp., talus slope, mostly shale, common, C. E. Garten 1933 (TRT); ca. 19 mi. n, of Nipigon, spruce-fir thickets on damp moss-covered talus 1966] Oak Ferns — Wagner 133 at foot of cliff, E. G. Voss 10335 (MICH, TRT) ; 15 mi. e. of Port Arthur, 2 mi. w. of Silver Islet, shady sandstone talus, R. M. and A. F. Tryon, A. C. Faber 4963 (MO). U.S.A.: ALASKA: WISEMAN, on Middle Fork of the Koyukuk River, ca. 67° 30' N, 150° W, Edith Scamman 913 (Gn). This fern, being intermediate between the two familiar species of Gymnocarpium, and having obviously peculiar spores might more appropriately be designated as merely a sporadic hybrid (e.g, as G. dryopteris X robertianum). There are several reasons why this was not done. (1) The plants develop large colonies locally, and behave as an apomictic species. (2) As shown on the range map (Fig. 3) plants conforming to this description run from Alaska southeast through the western Great Lakes area (the region, evidently, of its greatest abundance) down to central Penn- sylvania. (3) Cytogenetically (see below) at least one population, the type, is triploid, and not tetraploid as would be expected from our present knowledge of the parents as they occur in the eastern United States. The nearest dip- loids (judging from spore sizes and assuming correlation with the single diploid count) seem to be limited to the western part of the continent. (4) Finally, the intermediate is evidently to be found in at least some localities where one of the putative parents (G. robertianum) is unknown. The holotype, but not the cotypes, of Dryopteris linnaeana C. Chr. f. glandulosa R. M. Tryon, Jr. (Amer. Fern Jour. 29: 5, 1939) is evidently the same as the plant described above. The author wrote of his new “form” that the “rachis varies in glandularity, some specimens having only a few glands, while others have relatively many and rarely the blade is slightly glandular.” Judging from the specimens cited, the author included two kinds of populations — the form of G. dryopteris in which the glands are more frequent and conspicuous than usual, and the plant described above. As to the glandular form of G. dryopteris itself, I am not sure that it is worthy of recognition. In her survey (1961, p. 18) Mrs. Root concluded that “It is generally stated that ‘Additional collection localities not cited here but plotted in Fig. 3 were supplied by Dr. Erie Hultén. 134 Rhodora [Vol. 68 the fronds and rachises of G. dryopteris range from glabrous to rarely slightly glandular. However, upon close exami- nation of specimens from various localities it was found that every individual of this species examined has glands.” In synonymizing “forma glandulosa” with G. robertianum (as “G. dryopteris var. pumilum (DC) Boivin”), Boivin (1962) was in error. Neither the type nor the cotypes of Tryon’s form conform to the characters of G. robertianum. Specimens of G. heterosporum in herbaria have previously been determined as one or the other of the putative parents, but most commonly as “G. robertianum" or “G. dryopteris f. glandulosa.” The recognition of G. heterosporum is usual- ly fairly easy in herbarium material, so long as the speci- mens have ripe spores. These tend to be discharged and to lie around the axes of the frond, sticking to the sheet. If the spores appear blackish and strongly irregular in size and shape, this is an excellent diagnostic feature, in combination with the presence of the intermediate con- dition of glandularity. The spores can be seen even with a dissecting microscope or a high power hand lens; and once their appearance is learned it becomes easy to pick out other collections with the same spore conditions, so a compound microscope is unnecessary. The spores of typical specimens of G. robertianum and G. dryopteris seem always to have a “glassy,” tan or brownish color, and are strikingly regular. Both the soriferous specimens, as well as those which lack sori, of the intermediate oak fern will show the intermediate glandularity of the axes, especially on the upper parts of the rachis (if they have not been imbedded in herbarium paste). The glands of G. heterosporum are fewer and more widely spaced than they are in the densely glandular G. robertianum. In the field it is helpful to hold up the living fronds in the sun's rays and examine them with a good hand lens. The glands will glisten and thus be conspicuous, so that even small, sterile fronds of all three taxa, if growing together, may be separated. Wherry (1942) described the habitat of the type popu- lation as follows (italics mine): “... the base of a steep 1966] Oak Ferns — Wagner 135 northwest-facing wooded slope, above a brook, the under- lying rock being limestone which breaks into small angular blocks, from the crevices of which moisture oozes out, and by evaporation keeps the rocks cool." The Michigan station near Ishpeming, Marquette Co., is on steep shaded rock cliffs. In the latter area, the rocks contain serpentine. Garten took his specimens on a talus slope comprising most- ly shale. The Lake Co., Minnesota, material was recorded from crevices of a “diabase cliff," while that from Barron Co., Wisconsin, came from “quartzite talus." The speci- mens from east of Port Arthur, Ontario, were found on *shady sandstone talus." Thus, ecologically, the substratum itself does not seem to matter much. The constant features of the habitats of G. heterosporum seem to be (a) shadiness, (b) moisture and humidity, (c) presence of rock substratum of some kind, the plants growing either directly from crevices of cliffs or upon the fallen talus. One or both of the presumed parental species may be present at the same locality. CHROMOSOMES AND SPORES The chromosomes and spores seem to have significance in this group of plants because of their variability and their correlation with each other and with other systematic data such as distribution and morphology of the fronds. There are three chromosome conditions. The tetraploid with n = 80 or near that number is characteristic of G. robertianum as found by Manton and myself (Chiarugi, 1960; Fabbri, 1963). Judging from spore sizes (see Fig. 5) all populations of G. robertianum are probably tetraploid and the curve is normal. In Siberia, however, the rather similar G. con- tinentale (Petrov) Pojark. has small spores (indicated by dots in the graph) and may bea diploid. For G. dryopteris we first plotted all spore size averages and produced a broad, apparently undifferentiated curve. However, when we selected specimens from the area of presumably typical var. disjunctum (where we know there are diploids) an entirely different curve separated out. This area extends from British Columbia and Oregon east to 136 Rhodora [Vol. 68 Alberta and Montana, and the spore sizes average between 23-25 X 1.18 microns’ (Fig. 5, C) in maximum exospore diameter, in contrast to between 27-29 X 1.18 microns for var. dryopteris east of Colorado and including Europe (Fig. 5, B). Thus in G. dryopteris there is a correlation between spore size and polyploidy, as has been found in a number of other fern groups. The “sterile-spored” G. dryopteris commented upon above may represent triploid populations, derived by interploidal hybridization and spreading by some means of spore dis- seminule not yet fully understood, The question is where the diploid and tetraploid may meet geographically. The graph of spore samples from Alaska (Fig. 5, D) shows values all the way from 21 to 29 micrometer units, strongly indicating that in this area, G. dryopteris is a mixture of ploidal levels. It was extremely interesting to find that the type popu- lation of G. heterosporum (lower illustrations in Figs. 6 and 7) is triploid. Even though the pairing is very irregular, by totaling the estimated bivalents and univalents re- spectively of four sporocytes the following numbers re- sulted: (29) + 63, (33) + 50, (33) + 55, and (39) + 42. Their average was (33.5) + 52.5, or 119.5 chromo- somes in all. Assuming that G. heterosporum is truly of hybrid origin and that both G. dryopteris and G. roberti- anum in the eastern U. S. are tetraploid, then it is possible that the type population originated as 2x G. dryopteris var. disjunctum X 4x G. robertianum. Further cytological sur- veys of G. heterosporum in other areas are highly desirable. It is possible that 4x forms of it exist. I must alert collectors to the need of obtaining fertile materials of these plants of which the sori contain ripe spores. Out of 500 specimens that we examined, 150 proved to be sterile or too young for spore observations. Also, I should point out that it is not correct that the spores of ‘Conversion factor: one micrometer unit — 1.18 microns. Fig. 5 is plotted in micrometer units only, from spores taken from herbarium specimens and mounted directly in diaphane. 1966] Oak Ferns — Wagner 197 Gymnocarpium lack perispores. Ching’s description of “spores bilateral, warty, and without perispore" (quoted in Morton, 1941) surely does not conform to our observations. As shown in the spore tracings in Figure 4, there are two clearly defined layers — the outer, perisporial covering which is irregular, and the inner, exosporial boundary which is smooth. This is not different from other members of this group of ferns associated with Thelypteris and Dry- opteris. The perispores of Gymnocarpium show up especial- ly well in the abortive spores of “sterile G. dryopteris" and of G. heterosporum, where they actually are more or less exaggerated in development. This excessive perisporia] thickness is probably the factor which produces the darker, nearly black, appearance of the spores as seen en masse. UNIVERSITY OF MICHIGAN, ANN ARBOR REFERENCES CITED Borvin, BERNARD 1962. Etudes ptéridologiques. Il. Gymnocarpium Newman. Bull de la Soc. bot. France 109 (nos. 5-6): 127-128. CHIARUGI, ALBERTO 1960. Tavole cromosomiche delle Pteridophyta. Caryologia 13 (no. 1): 27-150. FABBRI, FERNANDO 1963. Primo supplemento alle Tavole cromoso- miche delle Pteridophyta di Alberto Chiarugi. Carologia 16 (no. 2): 287-335, LAWALRÉE, ANDRE 1950. Ptéridophytes. In Flore Générale de Bel- gique. Jardin Bot. de l'Etat, Brussels. Morton, C. V. 1941. On the name of the oak fern. Rhodora 43: 216-219. PICHI-SERMOLLI, RODOLFO E. G. 1965. Index filicum, suppl. IV. Intern. Bur. Pl Tax. and Nomencl., Utrecht. Root, ELIZABETH EICHSTEDT. 1961. Hybrids in North American Gymnocarpiums. Amer. Fern Jour. 51 (no. 1): 15-22. Tryon, ROLLA M., JR. 1939. Notes on the ferns of Wisconsin. Amer. Fern Jour. 29 (no. 1): 1-9. TUTIN, T. G. et al. (eds.) 1964. Flora europaea. Vol. I. Lycopodi- aceae to Platanaceae. Cambridge Univ. Press. WAGNER, W. H. JR. 1960. Evergreen grapeferns and the meanings of infraspecific categories as used in American pteridophytes. Amer. Fern Jour. 50 (no. 1): 32-45. 138 Rhodora [Vol. 68 WHERRY, EDGAR T. 1942. The ferns and lycosphens of Pennsylvania. Bartonia, no. 21: 11-63. 1961. The Fern Guide. Northeastern and Midland United States and Adjacent Canada. Doubleday and Co., Garden City, N.Y. LACTUCA MURALIS IN NEW ENGLAND Recently a number of duplicates from a collection of New Hampshire plants by Andrew P. N elson, Assistant Professor of Biology at Dartmouth College, were sent to Professor Albion R. Hodgdon at the University of New Hampshire. Among these was a specimen of Lactuca muralis (L.) Gaertn. (A. P. Nelson 1059) that was collected by the side of a gravel road between Cornish Mills and Plainfield in the town of Cornish, Sullivan County, N. H. on August 6, 1964. Professor Hodgdon states that this is the first report for this species in New England. Gray's Manual, 8th edition, describes the species as growing on roadsides and in waste places, local, w. Que., and e. N. Y. to Mich. (Adv. from Eu.). In Gleason and Cronquist Manual of Vascular Plants (1963) it is listed as a native of n, Eu. now known from N. Y. and Que. Voucher specimens have been deposited in the her- barium of the New England Botanical Club, Cambridge, Mass. and in the Pringle Herbarium at the University of Vermont in addition to the specimens in the University of New Hampshire Herbarium and the Jesup Herbarium. JAMES P. POOLE, CURATOR. JESUP HERBARIUM, DARTMOUTH COLLEGE CUCHUMATANEA A NEW GENUS OF THE COMPOSITAE (HELIANTHEAE)' C. E. SEIDENSCHNUR AND J. H. BEAMAN Cuchumatanea Seidenschnur & Beaman, gen. nov. Herba annua, minuta, depressa. Folia opposita, spathulata, obscure trinervia, basi leviter connata. Rami oppositi ad nodos. Capitula homogama, sessilia vel subsessilia, terminalia, solitaria, cum pare foliorum clavato-spathulatorum cireumclusa. Phyllaria 2, late oblongo- obovata, basi subite attenuata, membranacea valde concava, laxe flosculos involventia. Receptaculum conicum, paleaceum, paleis mem- branaceis flosculos non amplectentibus, exterioribus oblongo-ovatis, interioribus gradatim angustioribus usque ad centrales lineares. Flosculi 5-10 in capitulo, hermaphroditi actinomorphi. Corolla anguste tubuloso-campanulata, 3-4 lobata, infra flava, sursum purpurea. Sta- mina 3-4, antheris connatis basi sagittatis, apice cum appendice late ovata vel suborbiculata ornata. Stylus infra ramos contractus, ramis oblongo-ellipticis, acutis, lato margine dorsoque dense papilloso. Achaenia ellipsoideo-oblonga deorsum paulo attenuata levissime com- pressa, striolata, brunneo-nigra. Pappus nullus. Minute, depressed annual. Leaves opposite, spatulate, inconspicu- ously 3-veined, the bases slightly connate. Each node with two axillary branches. Heads homogamous, sessile or subsessile, solitary, terminal, surrounded by a pair of spatulate-clavate leaves. Phyllaries 2, broadly oblong-obovate, subpetiolate, membranaceous, strongly concave, loosely enclosing the florets, the outer oblong-ovate, the inner becoming pro- gressively narrower, the innermost linear. Florets 5-10, hermaphrodi- tic, actinomorphic. Corolla tubular, narrowly campanulate, 3-4 lobed, yellowish below, purplish above. Stamens 3-4, anthers connate, sagit- tate, with broadly ovate to suborbicular apical appendages. Style constricted just below the branches, the branches oblong-elliptical, acute, papillose to the base. Achenes ellipsoid-oblong, weakly com- pressed, brownish-black, striate. Pappus absent. Cuchumatanea steyermarkii Seidenschnur & Beaman, sp. nov. Herba annua, minuta, depressa usque ad 1 cm. alta, radice palari, saepe caespitosa. Caulis plus minusve sparse hirtellus vel glabratus, "The genus was discovered in the course of field work supported by National Science Foundation Grant G-9045. The research has been conducted in connection with NSF Grant GE-4051 for Undergraduate Research Participation. We are indebted to Dr. José Cuatrecasas for helpful suggestions and for preparing the Latin diagnoses and to Dr. D. C. D. De Jong for making the cytological preparations. 139 [Legend for figures 1-11] Fig. 1-4, 6-11. Cuchumatanea steyermarkii, based on Beaman 3962. Fig. 5. Jaegeria hirta, based on Beaman 4532. Fig. 1-3. Pales (X 15). Fig. 4-5. Stamens ( 40). Fig. 6. Head and subtending leaves (X 7, L, leaf; Ph, phyllary). Fig. 7. Phyllary (X 15). Fig. 8. Floret with achene (X 20). Fig. 9. Style branches (X 75). Fig. 10. Diagram of a terminal portion of the branching system (B, bud; H, head; L, leaf). Fig. 11. Diagram of a head with subtending leaves (B, bud; F, floret; L, leaf; Pa, pale; Ph, phyllary). 1966] Cuchumatanea — Seidenschnur & Beaman 141 viridis vel purpurascens composite dichasiale ramosus. Folia opposita decussata, basi paulo connata, saepe in pare inaequilonga, 3-7 mm longa 0.8-3 mm lata, spathulata, integra, margine parce revoluta sparseque ciliata, 3-nervata sed nervis plerumque inconspicuis, supra glabra, subtus praecipue in nervo medio sparse pubescens. Capitula homogama, sessilia vel subsessilia, terminalia, solitaria, campanulata, circa 2 mm alta 1 mm lata, cum duobus foliis brevibus clavato- spathulatis amplectentibus, 2-3 mm longis 1.2 mm latis, adpresse subtendentia partimque tecta. Phyllaria 2 circa 2.5 mm longa 1.5 mm lata plerumque inaequilonga, oblongo-ovata basim angustata, mem- branacea sed apice recurva herbacea leviter lacerata, 8-venulosa, dorso ad venulam mediam parcis pilis, valde concava, laxe flosculos in- volventia. Receptaculum 0.2-0.5 mm altum, conicum, paleaceum. Paleae membranaceae persistentes flosculos non involventes, 1.5-2 mm longae, venula media conspicua aliquando 2-3 venulis lateralibus fere inconspicuis, glabrae, sursum margine minute laceratae deorsum integrae, exteriores oblongo-ovatae, ceterae gradatim angustiores usque ad centrum lineares. Flores omnes hermaphroditi, 5-10, plerum- que 7 in capitulo. Corolla tubulosa 1 mm longa, limbo subcampanulato, . 3-4 lobato, lobis ovatis acutis purpureis glabrisque, tubulo 0.5 mm longo flavo, sparse pilosulo. Stamina 3-4 non semper pari quam corollae lobi, antheris basi sagittatis, apice appendice ovata vel subor- biculata, thecis 0.4 mm longis parietibus crassis. Stylus infra lobos constrictus, ramis oblongo-ellipticis, acutis, marginibus dorsoque papil- loso, uno saepe quam altero longiore. Achaenia 1.1 mm longa, ellip- soideo-oblonga, basim versus attenuata, levissime compressa, basi obtusa, brunneo-nigra, striata, glabra. Pappus nullus. Chromosoma m = 8. Minute, depressed, taprooted annual herb, 1 em or less high, oc- curring in patches and tending to form open mats. Stems glabrate to sparsely hirtellous, greenish or purplish, with two axillary branches at each node, one branch system arising from each leaf axil. Leaves opposite, decussate, 3-7 mm long, 0.8-3 mm wide, one member of each pair usually larger than the other, spatulate, entire, the bases slightly connate, margins sparsely ciliate, somewhat revolute, with 3 inconspicuous veins, glabrous adaxially, sparsely pubescent mostly on the midvein below. Heads homogamous, sessile or subsessile, solitary, terminal, campanulate, ca. 2 mm high and 1 mm wide, closely subtended and partly covered by a pair of small spatulate-clavate leaves 2-3 mm long and 1.2 mm wide. Phyllaries 2, ca. 2.5 mm long, 1.5 mm wide, one usually larger than the other, oblong-obovate, narrowing basally, mostly membranaceous but with herbaceous reflexed tip, lacerate near the apex, 3-veined, with a few hairs abaxially on the median vein, strongly concave, loosely enclosing the florets. Receptacle 0.2-0.5 mm. high, conical, paleaceous. Pales membranaceous, not sur- rounding the florets, 1.5-2 mm long, 1-veined, sometimes 2-3 incon- 142 Rhodora [Vol. 68 Spicuous lateral veins present, glabrous, minutely lacerate above, entire below, the outer oblong-ovate, the inner becoming progressively narrower, linear at the center of the head, persistent. Ray florets absent, Disk florets 5-10, commonly 7, hermaphroditie, Corollas tubular, narrowly campanulate, ca. 1 mm long, with 3-4 ovate acute lobes, tube 0.5 mm long, yellow, sparsely pilose, limb and lobes purplish, glabrous. Stamens 3-4, usually of the same number as the corolla lobes, anthers connate, sagittate, with broadly ovate to suborbicular apical appendages, thecae 0.4 mm long, with thick walls. Style constricted just below the branches, style branches oblong-elliptical, acute, papillose to the base, one branch often longer than the other. Achenes 1.1 mm long, ellipsoid-oblong, weakly compressed, with an obtuse base, brownish-black, striate, glabrous. Pappus absent. Chromosome number n = 8, based on Beaman 3962. Fig. 1-11. GUATEMALA. HUEHUETENANGO: Sierra de los Cuchumatanes, ca. 3 kms south of road between Llano de San Miguel and Todos Santos, from a point 2.5 miles west of Llano de San Miguel, near the highest point in the Cuchumatanes, collected from about the 3,680 to 3,740 meter level, in open Pinus rudis forest, occurring in patches in black soil near limestone outcrops, frequent in a local area, 2 August 1960, J. H. Beaman 3962 (Msc 171943, holotype; F, GH, K, TEX, UC, US isotypes) ; vicinity of Chémal, summit of Sierra de los Cuchumatanes, 3,700-3,750 m, Steyermark 50260 (F). The existence of this remarkable tiny plant was first pointed out by Dr. Julian A. Steyermark to Beaman in con- nection with the planning of a collecting expedition to Guatemala in 1959. Dr. Steyermark found it on August 8, 1942, in the vicinity of Chémal, Sierra de los Cuchumatanes, Department of Huehuetenango, Guatemala. In his field book he noted that it was annual, with the corolla yellow, the leaves green and shining, fleshy, fairly membranaceous, growing in bunches and patches. His collection was subse- quently examined by Dr. S. F. Blake who suggested to Steyermark that it probably represented a new genus near Jaegeria or Schkuhria. Blake pencilled on the sheet: “Jae- geria? Material too scanty & depauperate." In early August, 1959, and late July, 1960, Beaman made unsuccess- ful searches for the species in the vicinity of Chémal. But on August 2, 1960, it was discovered a few kilometers south- west of Chémal in one of the highest and most remote regions of the Cuchumatanes. In publishing the species it is a pleasure to associate the name of Dr. Steyermark with 1966] Cuchumatanea — Seidenschnur & Beaman 143 that of the Sierra de los Cuchumatanes. He has been the leading contributor to the understanding of the exceptionally diverse and unusual flora of this mountain range. Cuchwumatanea steyermarkii is an extremely reduced species, especially characterized by reduction of the plant body and of the florets. There is also slight asymmetry of the pairs of axillary branches, leaf and phyllary pairs, and style branches, with one member of each pair generally somewhat smaller than the other. In spite of reduction in size, however, the species has retained a complex branching system (see fig. 10). Two branches generally arise from each of the 3-4 nodes (except the first which is often un- branched). Additional axillary branches or buds are also present at the nodes of the primary branches. A solitary head terminates each branch. The heads are enclosed and somewhat concealed by sub- tending pairs of leaves. The two phyllaries follow the decus- sate pattern of the leaves (see fig. 6 and 11) and resemble immature leaves. In immature heads the subtending leaves are differentiated from the phyllaries mainly by the ciliate margins of the former and by their position. The two phyllaries partially surround the head and each subtends a pale and a marginal floret which is more advanced in de- velopment than other florets of the head. The number of corolla lobes varies from 3 to 4, which is also true for the stamens. While the number of corolla lobes and stamens is often the same, there are sometimes florets with 4-lobed corollas and 3 stamens. Florets with 3.]obed corollas and 4 stamens have not been found. In one floret with two normal stamens a third stamen was found with the filament of twice the normal width and two fused anthers. Such a condition may support the contention that the 3- and 4-merous florets are the result of a reduction process which is still in progress. Our findings confirm Blake's suggestion that the genus is related to Jaegeria. Among species of this genus it ap- pears to be closest to J. hirta (Lag.) Less. The habit of Cuchumatanea is very similar to that of a collection of V. [Vol. 68 Rhodora 144 SUME snonproop c-e snoiqu[d ‘Buorqo -prosdio ‘poj3ue -G ‘ƏY ‘ystumorg sesepuodde ojeAo Á] -peoiq YIM *'o39e3318€s Aysys sioqjue ‘e sdi} poyjnj *ie[n3uve -L3 yy” 192urT] po3orijsuoo JON gU ÁA[MO.L1€U ‘pajaay JON S9LIoS p jnoq* ur payeo -Hqui f'peoiq Alaa ayeIpey quesqy snoiqe[s ‘suo[qo-plosdyja — *oje -H3S ‘yorlq-ystumorg sogepuoedde aqyvao YUM ‘UIII 'ojejjrdes Apu3is sioqgjue ‘e əseq oui 0} oso[ -ded ‘aynow ‘avout U | pojorijsuoo 40N OF "8D pə JON iouur Jo S9LI9$ g pue i2jno c ojerpeqr quesqy snoiqes *Buo[qo-prosdi[[o *oje -H8 ‘youlq-ystuamorg sodup -uedde .e[norq.i0 q1r« '93e331d€s SloyjuR ‘e aseq a} 03 oeso[[rd -ed 'ojnow *oje[ooouv' po3orrjsuoo JON p9[99xM g ƏFeIpey juosqy snoiqeys "Suo[qo-prosdi[|p ‘ale “13S *xoe[q-usiua oq Sooup -uedde .1epnorq.toqns 0} 97840 A[peoiq YB 'ojej1pLdvs SIIYJUL *p-g aseq əy} 0} oso[[rded '[enboun uo3jo *o3noe 'eordi[[o-Suo[qo) sououe.tq əy} MOTE pojorijsuo;) p-e OT-S pooox 40N Z proostq snddeq sausyoy Suaure1g Saqout.q ƏS 9[ As Soqo[ e[[0.100 ysIp Jo ioquinN S]2.10g XSIp jo ioquinN sored serte[Auq SpeaH (0fzf ununag ‘naurbpjunjd 'g) D0112 Ş (6698 unwunoq ‘suadat 'g) D1322082D218 (GEST unuinag ‘pga p) Dia han Sf (96& umwv»og ‘yumu '7)) DauDJDUNYIND ""w1ouo3 poje[o1 YPM p2wpjpumion;) jo uosrieduro? T AQEL 1966] Cuchumatanea — Seidenschnur & Beaman 145 hirta (Beaman 4532) from La Cima, Federal District, Mexico. This specimen also has sagittate anthers with orbieular appendages which correspond closely to those of Cuchumatanea, although the latter has shorter thecae with thicker-walled cells along the margins. In another specimen of J. hirta (Bourgeau 1553 from the Valley of Cordoba, Mexico) the anthers seem to differ from those of Cuchuma- tanea mainly in not having thick-walled marginal cells. The genera are also similar in achenial structure, both having ellipsoid-oblong, slightly compressed, striate, glabrous achenes without pappus” Although the features noted above indicate a close re- lationship of Cuchumatanea with Jaegeria, they differ by characters of a magnitude equal to those by which other Verbesinoid and Galinsoginoid Compositae are distin- guished. The most important characters which separate the genera are as follows: (1) The heads of Cuchumatanea are discoid. Only J. gracilis Hook. f. from the Galapagos Islands is without ray florets (the type of J. discoidea Klatt, as noted by Robinson (1900), is radiate). (2) Cuchumatanea has only two phyllaries which do not surround the marginal florets but partially enclose the head. Jaegeria has 5 or more phyllaries each of which encloses an ovary or achene of a ray floret with slightly overlapping marginal membranes. (3) The tubular corollas of Cuchumatanea have 3-4 lobes rather than 5 as in Jaegeria, and the number of stamens in the former is also reduced. (4) The pales are unkeeled in Cu- chumatanea and keeled in Jaegeria. (5) The chromosome number in Cuchumatanea is n = 8, while counts reported for four species of Jaegeria have all been on the base of x = 9 (Turner et al., 1962; Beaman and Turner, 1962; Turner 2After submitting the manuscript for publication we sent a copy of it and some of the material of Cuchumatanea to Dr. A. M. Torres who is currently monographing the genus Jaegeria. He notes that the specimen of Jaegeria hirta (Beaman 4532) which we have used for comparison is at the extreme end of variation (a reduced form) in this species and that more typical specimens appear less similar to Cuchumatanea. He considers J. axillaris Blake, known only from the type collection from Colombia, to be closer to Cuchumatanea. 146 Rhodora [Vol. 68 and King, 1964). A further comparison of similarities and differences of Cuchumatanea and Jaegeria, in addition to the characters of two other related genera, is given in Table 1. We have not found any features to support Blake’s tenta- tive suggestion that Cuchumatanea might be related to Schkuhria (Helenieae). Because of its alliance with Jaegeria and other genera placed by Hoffmann (1894) in the subtribe Verbesininae, we tentatively assign Cuchumatanea to this subtribe next to Jaegeria. A more satisfactory classification may ultimately bring certain genera of the Galinsoginae into closer as- sociation with this group of the Verbesininae. DEPARTMENT OF BOTANY AND PLANT PATHOLOGY, MICHIGAN STATE UNIVERSITY, EAST LANSING, LITERATURE CITED BEAMAN, J. H., and B. L. TURNER 1962. Chromosome numbers in Mexican and Guatemalan Compositae. Rhodora 64: 271-276. HOFFMANN, O. 1894. Compositae in Engler & Prantl, Natürl. Pflan- zenfam. 4°: 87-391. ROBINSON, B. L. 1900. Synopses of the genera Jaegeria and Russelia. Proc. Amer. Acad. 35: 315-321. TURNER, B. L., A. M. POWELL, and R. M. KING. 1962. Chromosome numbers in the Compositae. VI. Additional Mexican and Guate- malan species, Rhodora 64: 251-271. TURNER, B. L., and R. M. KING. 1964. Chromosome numbers in the Compositae. VIII. Mexican and Central American species. South- west. Nat. 9: 27-39. SEED GERMINATION OF SHORTIA GALACIFOLIA T. & G. UNDER CONTROLLED CONDITIONS' Mary H. RHOADES” Until 1936 it was belived that Shortia galacifolia could not reproduce by seed because the scapes decayed and fell over before the seeds were mature (Kelsey, 1902). Observa- tions by Ross (1936) disclosed that Shortia seeds do reach maturity and germinate, but in an unusual manner. Germi- nation occurs in the capsule, the seeds being held against the placenta by the firm, partially opened capsule walls. In this position the seeds do not readily fall out when ripe, and are kept fairly moist because the capsule and cup-like calyx retain water. Seedlings reach the ground when the scape decays and falls. There they have little chance of survival because of predation by insects and because of the dense mass which Shortia plants form by vegetative reproduction. Attempts by Ross to germinate seeds on soil in the labora- tory were unsuccessful, and he concluded that there was little possibility for the spread of Shortia plants by seed. Ross's work suggests that seedlings of Shortia would be rare in nature, and that most of them would be near mature Shortia plants. On the contrary, several people have re- ported finding abundant seedlings in the field, both under- neath (Crandall, 1956) and several feet from patches of mature plants (Crandall, 1956; Vivian, 1965). However, it was not determined whether these seedlings were from seeds which germinated in the capsule or on the ground. Since it is more likely that seeds are transported rather than seedlings, it appeared worthwhile to investigate the possibility of seed germination outside the capsule. Capsules of Shortia galacifolia were collected on May 15, 1964, at an elevation of 1440 feet on the northwest side of "Part of a thesis submitted in partial fulfillment of the requirements for the M.A. degree at Duke University, Durham, North Carolina “Present address: Department of Biology, Virginia Polytechnic Institute, Blacksburg, Virginia 147 148 Rhodora [Vol. 68 Crossroads Mountain in Transylvania County, N.C. The capsules were open and most of them contained many seeds, but no seedlings were found in them. Immediately when picked they were wrapped in paper towels and put in plastic bags to prevent drying. A previous experiment showed that freezing the seeds did not impair germination, so the capsules were kept frozen until the seeds could be removed and counted. When the seeds were removed from the capsules, those with any visible defect, such as mold, discoloration, or extreme flatness, were discarded. The sound seeds were then put into envelopes of 50 seeds each and refrozen until all seeds were sorted. To prevent the seeds from drying out, only about six capsules were taken from the refrigerator at a time for removal of the seeds. Eight environmental factors were tested for their effects on germination. Temperature, moisture, and light were combined into one experiment, so there were six experiments in all. For all but the first experiment, germination was tested in an air-conditioned room with a temperature range of 20° to 24°C and a light intensity of 30 ft-c. Germination was tested in all treatments, except those involving soil, by sprinkling the seeds on moist Whatman No. 1 filter paper in a clean 9 em petri dish. The paper was moistened once a day with distilled water from a medicine dropper. Each treatment included six replications of 50 seeds each. The seeds were assigned to a particular treatment by numbering the envelopes of seeds and then using a table of random numbers to select the envelopes for a prearranged order of treatments. The number of seeds germinated was recorded every 4 days up to 32 days, and then once again after 50 days. The first appearance of the radical was the criterion for germination. To determine the effect of the interaction of temperature, moisture and light, germination was tested at 10°, 20°, and 30°C, in the light and the dark, and at three moisture levels. The moisture levels were “abundant,” in which the filter paper was saturated with water; “sufficient,” in which it 1966] Shortia — Rhoades 149 was thoroughly moistened but not saturated; and ‘‘de- ficient,” in which it was never thoroughly moistened. Both incandescent and fluorescent lamps provided illumination. The combinations of three temperatures, two light con- ditions, and three moisture levels made 18 treatments. Two other treatments tested the effect of a changing temperature regime on germination. In one treatment seeds were placed alternately between 10°C and 20°C every 48 hours. In the other treatment seeds were moved pro- gressively from 10°C, to 20°C, to 30°C. After 5 days at each of these temperatures, the seeds were put at 22°C for the remaining time. For both of the changing temperature treatments the seeds were kept in the light and given “suf- ficient” moisture. Since Shortia seeds germinate within the capsule, it was decided to test the effect of capsule parts on germination. For example, the calyx from about 10 capsules was cut up and the pieces were sprinkled on the filter paper along with the seeds.. The placenta, calyx, and ovary wall were tested for their effects in this way. To obviate the effect of moisture retention by capsule parts, a control treatment was set up in which pieces of filter paper were sprinkled in the dishes. The cumulative effect of the capsule parts was tested by two treatments: putting seeds into empty, intact capsules and putting pieces of all capsule parts with the seeds. Three other experiments were designed to test the effect on germination of substrate type, acid, and soaking. Steam sterilized sand, sphagnum, and a mixture of two parts sandy loam, one part sand, and one part sphagnum were the sub- strates tested. Petri dishes were half filled with the sub- strate and the seeds were sown on the surface, which was kept moist. The acid solutions tried were glacial acetic acid of pH 3.5 and two leachates of pH 4.5, one of moss and the other of leaf litter collected at the site where the seeds were obtained. Soaking periods used were 1, 3, 6, and 9 days in distilled water. Seeds kept at 22°C in the light and with “sufficient” moisture were used as the control for all tests. 150 Rhodora [Vol. 68 In the final experiment six lots of 200 seeds each were put in petri dishes and allowed to air dry for 1 to 6 days before they were transferred to moist filter paper. Dupli- cate sets of 100 seeds each were also air dried, then weighed and oven dried at 105°C to a constant weight. In this way moisture content of the seeds, as percentage oven-dry weight, was determined for each of the air drying periods. Although eight tests were designed for statistical treat- ment of the results, for some tests the great variation in number of seeds germinated among replications or the low total number of seeds germinated made a meaningful sta- tistical analysis impossible, Even the mean and the standard deviation cannot be given for some treatments because they indicate the possibility of a negative number of germina- tions, which is meaningless. Table 1 gives the results for the effect of temperature, moisture, and light on germination. Many more seeds germi- nated at 20°C than at 30° or 10°C, and more germinated in the light than in the dark. No germination occurred at the “deficient” moisture level in any temperature regime. At 20°C more seeds germinated in the “abundant” moisture condition than in the "sufficient." The difference is signifi- cant at the 2.5% level using a Student's t-test (Abundant, mean number of germinations/50 seeds = 4.33 + 1.51; Suf- ficient, mean = 2.17 + 1.33). The effect of the changing temperature treatments on germination is also shown in Table 1. Almost identical re- sults were obtained for the two treatments. A constant temperature of 20°C was as good for germination and per- haps better than, the two changing temperature regimes. The presence of the placenta, the calyx, the ovary wall, or all parts improved germination about fivefold over that of the control. An analysis of variance using the 2.5% significance level showed that there was no difference among the treatments involving capsule parts. Adding filter paper instead of capsule parts did not improve germination. Sand and sphagnum were superior to the sandy loam mixture or filter paper as substrates for seed germination. 1966] Shortia — Rhoades 151 Table 1. Effect of Temperature, Moisture, and Light On Seed Germination of Shortia galacifolia Temperature Number of Seeds Germinated After and Moisture 50 Days (300 seeds/treatment Light Dark 10° C Abundant 0 0 Sufficient 0 1 Deficient 0 0 20° C Abundant 26 2 Sufficient 13 0 Deficient 0 0 30° C Abundant 1 0 Sufficient 0 0 Deficient 0 0 Changing temperature Alternating 10^, 20° 4 Increasing 10° to 20° to 30° to 22° 3 Constant 20° 13 Using water with leaf litter in it instead of plain distilled water may have improved germination slightly, but using water with moss or glacial acetic acid added did not. Better germination was obtained for all soaking treatments than for the control, and the period of soaking had no effect. The average moisture content of Shortia seeds before they were air dried was 12% oven-dry weight. No loss of germi- native capacity occurred for seeds air-dried for up to 6 days. However, no loss of moisture occurred, because the vapor pressure deficit of the air remained low over the whole period. A separate set of 100 seeds was made up to be left until a significant amount of moisture was lost. The seeds were allowed to air-dry for about 3 days. Since no moisture loss had occurred after this time, the seeds were put in an oven at 30°C for 12 hours. Their weight was reduced 4% during this treatment, which was three times the weight lost by any of the air-dried lots of seeds. After removal from the oven, the seeds were left in the air for 20 days. 152 Rhodora [Vol. 68 They had regained their original weight when they were put on moist filter paper to test. germination. None of the seeds germinated. For 21 treatments the number of seeds germinated was recorded every 4 days. For most of the treatments the first germination occurred between 13 and 16 days from the start. Soaking caused germination to begin 4 to 8 days earlier. The distribution of germination over the 32 day period followed no particular pattern. A final count for all treat- ments was made after 50 days, and a few germinations were recorded at this time. Most germinations, however, had oc- curred before 32 days. Seeds of Shortia galacifolia germinated in the laboratory under a wide variety of conditions, indicating that the seeds do not have to be in the capsule to germinate. The factors which appear to be most significant in influencing germi- nation of Shortia seeds are temperature, moisture, light, and presence of capsule parts. The optimum temperature for germination is probably around 20°C, because substantially more germination oc- curred at this temperature than at either 10°C or 30°C. An abundant moisture supply is necessary for good germi- nation. No seeds germinate when moisture is always de- ficient, but some may germinate when moisture is alternately available and unavailable. Although the evidence is not conclusive, it also appears that Shortia seeds lose their germinative capacity if their moisture content falls below a certain critical level, even if they later regain moisture. Soaking seems to improve germination slightly, and even 9 days of soaking does not impair germination. Light appears to be beneficial for germination, since many more seeds germinated with 900 ft-c illumination than in the dark or with 30 ft-c. Although any capsule part improved germination when present with the seeds, combining parts did not produce a cumulative effect. Using leachate from leaf litter to water the seeds did not improve germination as much as the 1966] Shortia — Rhoades 153 presence of capsule parts, indicating that there is some or- ganic substance peculiar to Shortia which aids germination. From the results of the various tests it might be predicted that the best germination of Shortia seeds would be obtained by soaking the seeds at least 1 day, using sand as a substrate with capsule parts scattered over the sand, providing an illumination of about 900 ft-c, and supplying abundant moisture. Combining the two treatments which gave the best germination, “addition of placenta" and *209C-light- abundant moisture," was tried, but no increase in percentage of germination resulted. Although germination of Shortia seeds in nature outside the capsule has never been observed, the results of the labo- ratory experiment indicate that it undoubtedly occurs. Fac- tors which were found to be important for good germination in the laboratory are present in the natural habitat. Micro- environmental measurements made in several different stands of Shortia show that the air temperature during the growing season ranges from 13°C to 32°C, that the relative humidity never falls below 50%, and that the light intensity ranges from 20 to 1000 ft-c (Vivian, 1965). Decaying capsule parts and leaf litter are present on a sandy soil. In 1936, Ross thought the answer to the extreme locali- zation of Shortia was the fact, that the seeds are held tightly in the open capsule and must germinate there, thus limiting the spread by seed. Although Ross noted that some seeds fall from the capsules, he did not mention how many. The author has observed that 50% or more of the seeds may be lost within a month after the capsules open. Rain is proba- bly the primary agent in releasing the seeds, which could easily be carried far from the mature plants by rivulets or streams. It does not seem that lack of seed dispersal can be used to explain why Shortia fails to colonize new, apparently suitable habitats. Low seedling survival may be much more important in limiting the spread of Shortia by seed. Obser- vations suggest that the seedlings grow very slowly in nature and are quite subject to winter kill (Vivian, 1965). Seeds germinated in the laboratory produced seedlings about 154 Rhodora [Vol. 68 2 cm high after 8 months of growth. The amount of competi. tion that seedlings can tolerate is probably small. It seems that the problem of the restricted and discontinuous distri- bution of Shortia galacifolia has not been solved, but that requirements of mature plants, competition, and seedling survival, perhaps along with a low percentage of germi- nation, are more significant factors than lack of seed dispersal, DEPARTMENT OF BOTANY, DUKE UNIVERSITY, DURHAM, NORTH CAROLINA. LITERATURE CITED CRANDALL, DoRoTHY L. 1956. Shortia galacifolia in Gray’s Manual range. Rhodora 58: 38-40. Kersey, H. P. 1902. Shortia, p. 1664. In L. H. Bailey, Cyclopedia of American Horticulture. Macmillan Company, New York, 4 York Bot, Gard. 37: 208-211. VIVIAN, V. E. 1965. Department of Biology, Glassboro State College, vols. Ross, M. N. 1936. Seed reproduction of Shortia galacifolia. J. New Glassboro, New Jersey. Personal communication. THE APPLICATION OF THE LINNAEAN NAMES OF SOME NEW WORLD SPECIES OF EUPHORBIA SUBGENUS CHAMAESYCE DEREK BURCH In 1960 Wheeler resurrected a subject which most workers in the Euphorbiaceae must have thought long since settled. His proposal, in 1939, to change an established and well understood usage of the name Euphorbia maculata L. stirred up a controversy which had died down in print by 1948 (Svenson 1945; Fosberg 1946, 1947; Croizat 1947, 1948), and which Fosberg's rejustification of his earlier stand failed to revive in 1953. In 1962 Croizat answered Wheeler's case with what should have been the final word on the subject, but in the course of a revision of the genus Chamaesyce in the Caribbean it has been brought home to me that confusion stil exists in American herbaria, and that some collectors are following Wheeler’s suggestion even though the case for the change was never proved. If this were the only Linnaean name whose application to a New World species of Euphorbia sensu lato had been questioned the matter would best be allowed to lie, but there are other situations to clarify and, since some of them de- pend on the use of E. maculata, I have reluctantly decided to drag the old bones of the argument out once more. The other areas which must be discussed are the names for a group of erect, large-leaved plants including E. hyperi- cifolia (the name which Wheeler refers to as a *bandora's box" — presumably with some knowledge since he was the last to force its lid closed); the confusion between the Caribbean E. prostrata and E. chamaesyce of the Old World, and finally a note on the application of the name E. thymi- folia L. In the interests of brevity, only the relevant parts of earlier papers will be discussed. Most of the confusion in applying the names proposed by Linnaeus stems from a difference in interpretation of the relative importance of the parts of the material on which 155 156 Rhodora [Vol. 68 his description was based. Both Svenson (1945) and Stearn (1957) have published very lucid accounts of his descriptive method, pointing out the importance of the phrase-name, not only as a partial characterization of the plant but also as a key to the genus when taken in conjunction with the polynomials of the other species. These phrases were re- vised to maintain this differentiating function as new material or information was received, and the superseded phrases were put into synonymy. A mistake commonly made in deciding the application of a Linnaean name is to confuse the need to establish the concept of species which was in the author's mind with the present-day wish to have a single specimen as a representa- tion of the name given to the species. Linnaeus did not work with "type specimens", but based his species on an aggre- gate of earlier descriptions, illustrations, and actual ma- terial. It must be left to a worker familiar with the group, and having a clear understanding of the Linnaean method, to decide just what elements were included by Linnaeus in composing his polynomial phrase-name. Only when this has been done should the specimens now in the herbarium, which may or may not have been present when the differ- entiae were drawn up, be considered in the light of the International Code for designation as types of the names. 1. The application of Euphorbia maculata L. The species to which Linnaeus gave the trivial name “maculata” is number 21 in Species Plantarum edition 1. The full entry is made up of the usual polynomial phrase- name and references to earlier works (in this case a descrip- tion and illustration by Plukenet), an indication of habitat, and also a supplementary description including characters such as color which Dandy suggests to mean that it was taken from fresh material (personal comm. 1964). The author’s concept, then, we know to be based on a plant which Plukenet described and illustrated, with the phrase-name rewritten to be comparable with others in the genus, and probably to include information taken from a specimen. 1966] Euphorbia subgenus Chamaesyce — Burch 157 These are the facts in the case, and from this point we move into conjecture. For many years the name E. maculata has been applied to a prostrate plant, a specimen of which is in the Linnaean Herbarium on sheet number 630-11. The sheet is marked “21 maculata", which agrees with the entry in Species Plantarum. Most authors have taken the number to be one written on the sheet by Linnaeus, although Savage (1945), in his invaluable catalogue of the herbarium, ascribes it to Sir J. E. Smith. When I examined the speci- mens last summer I was unable to decide between these two viewpoints, but, in any case, there is general agreement that the epithet “maculata” on the sheet was added by Smith. There is another sheet, 630-4, on which Linnaeus wrote “17 Euphorbia maculata". In Species Plantarum number 17 is the species to which the trivial-name “hypericifolia” was given, and the specimen on the sheet is of one of a group of superficially similar erect species to which Linnaeus applied this name. It is clear that the number and the name on this sheet are contradictory, and that Linnaeus was mistaken in writing one or the other. The choice of which is in error is the first matter for conjecture, but in reaching a decision it should be borne in mind that the number was part of a major work which was in progress or had recently been completed (for it is not certain when the annotation on the sheet was made), while the name was, literally, a trivial name — a new, “shorthand” way of referring to a species, the use of which may still have been unfamiliar to Linnaeus. Wheeler, however, chooses to regard the trivial name as correct, the number as wrong, and on this builds the whole case for his application of the name. In effect, he first selected his type specimen, and then showed that it is feasible that it might have fitted Linnaeus’ concept of the species. Let us instead examine the entry in Species Plantarum, keeping in mind the two entities which have been offered as candidates for the name. The polynomial specific name, which was designed to separate the species from all others 158 Rhodora [Vol. 68 rather than as a description, contains no features which positively identify or disqualify either plant. Wheeler was correct in omitting this as a factor in his argument. Con- sider, however, the position of the species in the genus. Linnaeus was a methodical worker, and his purpose was to distinguish between the discrete groups that he considered species. What better way is there of doing this than by arranging similar things together, and then pointing out the features in which they differ? It is not suggested that Linnaeus followed this plan with complete consistency, but Croizat (1962) has shown in convincing detail that his genus Euphorbia is laid out in this fashion. The succulent species fall in the early part, followed by the shrubby types, while in the section Dichotomae, species 16-26, the arrange- ment is even more systematic. Species 16 through 19 are of erect or ascending habit with serrate leaves, species 20 through 24 are prostrate and have various leaf margins, and species 25 and 26 are erect with entire leaves. The placement of 21 maculata between 20 thymifolia and 22 prostrata, is strong evidence that the traditional application of the name to the prostrate plant is the correct one to follow. The descriptive phrase of Plukenet is noncommittal for our purpose of deciding between the two species, and I would agree with Wheeler that the illustration must also be re- jected as a decisive factor. However, I feel that he is rash in seizing on features of the italicized description that follows the note on the habitat in Species Plantarum. to support his choice of a type specimen on which to base the name. In his most recent article (1960) he persists in assuming that one of the two now in the Linnaean Her- barium must be treated as a type specimen in the modern sense. Even if these sheets were in Linnaeus' possession at the time of writing Species Plantarum, Wheeler's choice of one over the other because it agrees with parts of the description may be countered by reference to other charac- ters which would support the opposite choice. The shape of the leaves, and the description *subpilosa", for example, 1966] Euphorbia subgenus Chamaesyce — Burch 159 are each more appropriate to the specimen of the prostrate plant on sheet 630-11 than to the other. Nothing in the original entry in Species Plantarum, then, gives grounds for changing the application of the name, and there is sufficient doubt about Linnaeus' view of the speci- mens now in the herbarium for them to form an unreliable basis for such a drastic step. On the other hand it has been shown that the layout of Species Plantarum gives a positive reason for not making such a change, and the application to the prostrate species, affirmed by Jacquin's plate in 1772 and adopted by the vast majority of subsequent authors, should be maintained. There is one further point that must be discussed before this matter may finally be laid to rest. Both Wheeler (1939, 1960) and Fosberg (1946, 1953), who correctly apply the name E. hypericifolia to an erect plant, take the statement “Euphorbia maculata similis E. hypericifoliae" by Linnaeus in the Mantissa Altera (1771) to be selection of the erect element from the two which they feel he had earlier com- bined under the name E. maculata. Croizat's dismissal of this as an irrelevancy (1947, 1948) was not satisfactory, but it was not until I had the opportunity to examine some of the books in the Linnaean Library that I found a reason- able explanation. One of the first books to use differential phrase-names in the manner of Linnaeus was Patrick Browne's Civil and Natural History of Jamaica (1756). Stearn (1957) reports that this delighted Linnaeus, who went through the book providing (for his own use in later publications) the nomina trivialia that Browne had thought it unnecessary to give. Browne’s “Euphorbia 2. Minima reclinata, foliolis ovatis denticulatis . . ." gives as synonym the same “Tithymalus erectus acris . . ., Sloane Cat. 82, & H. t. 126" that Linnaeus included in his 17. hypericifolia. In recognition of this, “E. hypericifolia?” has been written in the margin of this entry, probably by Linnaeus himself. Browne goes on to describe his plant *. . . it is a slender weakly creeper and seldom runs above three or four inches from the root; its branches are smooth and slender and the 160 Rhodora [Vol. 68 leaves small and oval". Here, then, is the basis for the statement in the Mantissa. Linnaeus knew “maculata” to be a prostrate plant, and, on reading Browne's description of the plant based on the same element as his “hypericifolia” as a creeper, he simply drew attention to this similarity of habit. 2. The application of the name Euphorbia hypericifolia L., and the names for the group of morphologically similar species. This group, with strong, spreading to erect stems and leaves which are usually toothed, includes at least four species in America. They are distinguishable on pubescence, size of capsule, and on stipule and seed-coat characters, and have somewhat distinct geographical distributions. The names under which they have been known may be sum- marized thus: Entity A — northern plant with pubescence only in lines on stem; cyathia never glomerulate; capsules large; seed coat rippled . . . Euphorbia nutans Lag.; E. preslii Guss.; E. hypericifolia L. sensu Michaux, Torrey; E. maculata L. sensu Wheeler. Entity B — plant of the southern United States to South America; rarely and sparingly pubescent; cyathia never glomerulate; capsules large; seed coat transversely ridged . . . E. hyssopifolia L.; E. brasi- liensis Lam.; Chamaesyce nirurioides Millsp.; C. jenningsii Millsp. Entity C — tropical or subtropical plant; rarely pubescent; cyathia glomerulate; capsules small; seed coat wrinkled . . . E. hypericifolia L.; E. pilulifera L.; E. glomerifera (Millsp.) L. C. Wheeler. Entity D — tropical plant with all herbage close-pubescent; cyathia never glomerulate; eapsules large; seed coat transversely ridged ... E. lasiocarpa Klotzsch; E. hypericifolia L. sensu Wheeler. The central problem here is the application of the name E. hypericifolia L., which has been widely used for both Entity A and Entity C, and which Wheeler proposed in 1939 for Entity D. Entity B need not be involved in this discussion. It was described by Linnaeus in Systema Naturae edition 10 (1759) with the trivial name hyssopifolia, and this has been con- sistently applied by all authors except those who divided the taxon on the basis of leaf shape. Examination of a long series of this highly variable species has so far failed to 1966] Euphorbia subgenus Chamaesyce — Burch 161 substantiate any such division, and the name £F. hyssopifolia L. should probably be applied throughout. Wheeler’s case for the application of the name E. hyperi- cifolia to the pubescent West Indian species, Entity D, was made at a time when neither the Linnaean specimens nor Savage’s catalogue was available for study. He examined photographs of the two sheets in the herbarium labelled “hypericifolia,” and, having disposed of sheet 630-4 as the type of E. maculata L. in the fashion discussed above, he selected the second sheet, 630-3, as the type of the epithet hypericifolia. He correctly identified the specimen on this sheet as an example of Entity D, and proposed that the epithet be used for this species. The sheet, however, is marked “Br,” indicating that it came from Patrick Browne, most of whose collections were not added to the herbarium until 1758 (Stearn, 1957), and, thus, could not have been considered by Linnaeus when he described his E. hyperici- folia. Wheeler's proposal may be rejected on these grounds alone. Entity D was first described by Klotzsch in 1843 under the specific name E. lasiocarpa. Grisebach, in his Flora of the British West Indian Islands (1864), reduced the taxon to varietal level under E. hypericifolia L., but there has been general agreement that it merits specific rank, and that the name used by Klotzsch should be maintained. The question of the application of *hypericifolia" should also be approached by considering the entry in Species Plantarum. The polynomial phrase-name is taken without alteration from Wiman's Euphorbia, ejusque Historia . . ., a dissertation published under the supervision of Linnaeus in 1752. The entry here includes even more synonyms than are carried over into Species Plantarum, and also a descrip- tion which makes it clear that a variety a, with almost soli- tary “flowers”, and another 8, with “flowers” collected into heads, were recognized and included in the concept of the species. Variety « is probably Entity A, which appears to be the plant described in most of the earlier works of Lin- naeus given here as synonyms. Variety B is easily recog- 162 Rhodora [Vol. 68 nisable as Entity C, and this was the plant described and illustrated by Sloane as “Tithymalus erectus acris . . ." and by Commelin as “Tithymalus Africanus, seu Peplis major Brasiliensis . . ." (fide Tab. 10, “Tithymalus Ameri- canus, flosculis albis") to which reference is made in the synonymy. ]t is clear that Linnaeus held a broad concept of E. hypericifolia at the time of publication of Wiman's work and Species Plantarum, since the polynomial phrase- name of his earlier works was expanded in these to include features of Sloane's plant. In a case such as this, where the elements included by Linnaeus are now considered to form more than one species, Stearn (1957) suggests that the choice as to which is to bear the name should take into account not only all parts of the type material, but also the author's intent, and, if possible, subsequent and current usage. We have seen that both Entity A and Entity C are represented in the synonymy, and that while Entity A was the first to be described, this is balanced by the later broadening of the description to include Entity C. There is little in the entry in Species Plantarum to guide the choice, and no indication of the author's intent. The usage of the names, however, is another matter. Sir J. E. Smith, who acquired the herbarium in 1783, favored Entity A (Torrey, 1843), and his example was followed by Michaux (1803), Torrey, and a few early American authors. With these exceptions there has been an overwhelming use of E. hypericifolia as the name for Entity C, including Crantz' Institutiones Rei Herbariae (1766), Miller's Gardener's Dictionary edition 8 (1768), Aublet's Histoire des Plantes de la Guiane Francaise (1775), Humboldt's Nova Genera... (1817), Hooker's Exotic Flora (1823), and Boissier in DC. Prodromus (1862). In fact, the usage is so firmly established that it would be very much against the interests of stability of nomenclature to consider making any change, and the name E. hypericifolia L. should continue to be applied to this subtropical, erect plant with small capsules usually borne in glomerules. Sloane's illus- tration, cited by Linnaeus in Species Plantarum, is an ap- 1966] Euphorbia subgenus Chamaesyce — Burch 163 propriate lectotype for the name used in this fashion, and the specimen on which the illustration was based, if it still exists in the Sloane Herbarium in the British Museum, would constitute what Dandy has referred to as a “typo- type" (Stearn, 1957). The name for Entity A remains to be decided. The varietal status accorded it by Linnaeus was not questioned for a number of years, but in 1816 Lagasca described a taxon under the specific name E. nutans. Wheeler (1941) reports that he examined a specimen which probably repre- sents the plant that Lagasca described, and concluded that the name was apparently based on unusually vestite plants of Entity A. On these grounds the name E. nutans Lag. will be adopted for this species, with the more commonly used but later name E. preslii Guss. passing into synonymy. In summary, then, the names for this group of species are as follows: Entity A (Northern plant with large capsules) — E. nutans Lag. Entity B (Southern plant with large capsules) — E. hyssopifolia L. Entity C (Tropical plant with small capsules) — E. hypericifolia L. Entity D (Tropical plant which is densely pubescent) — E. lasio- carpa, Klotzsch. 3. Euphorbia prostrata Ait. and E. chamaesyce L. Wheeler suggested in 1941 that these two species are con- specific, basing his case on the fact that one of the two sheets labelled *chamaesyce" in the Linnean Herbarium (sheet 630-17) is actually a specimen of the plant which has been known as E. prostrata Ait. This sheet, however, is another of Brown's specimens, and was not in Linnaeus' possession until long after his description of E. chamaesyce in Species Plantarum. The other sheet (630-15) is a good example of the European species to which the name has usually been applied, and nothing in the original entry sug- gests that Linnaeus intended the limits of his species to be set broad enough to include the West Indian plant. The two species are, in fact, quite distinct, and the anno- tation by Linnaeus on sheet 630-17 is a simple misidentifi- cation. Croizat (1945) has documented the consistent use of the two names for the distinct entities, and there seems 164 Rhodora [Vol. 68 little room for argument against maintaining E. chamaesyce L. as the name for a Mediterranean plant not so far found in the New World, and E. prostrata Ait. as the name for a weed of the Old and New World tropics. Wheeler accepts this usage in a supplement to the second edition of Kearney & Peebles Arizona Flora published in 1962, but his correc- tion has been overlooked by the authors of other floras which have appeared since that date. 4. Application of the name E. thymifolia L. Wheeler (1941) drew attention to a suggestion by A. Gray that the specimens 630-10 “Euphorbia 20 thymifolia" and 630-11 in the Linnaean Herbarium are of the same entity. An examination of the sheets confirmed this, and that both are the prostrate E. maculata L. There is no specimen in the Herbarium of the plant which has tradi- tionally been called E. thymifolia L., and the protologue in Species Plantarum is of little direct help in deciding the application of the name. The polynomial phrase-name and descriptions by Burmann and Plukenet indicate that the plant is prostrate and pubescent, with serrate leaves and "flowers" somewhat collected into groups, and the illustra- tions cited fit these descriptions but show no characters which are diagnostic. Burmann, however, includes in his synonymy *'Cha- maesyce, Sloane Cat. 83". This was based on a Specimen from St. Jago de la Vega, now in the Sloane Herbarium, which is cited by Fawcett and Rendle in Flora of Jamaica (1920) as an example of E. thymifolia L. in the sense in which the name is used by most authors. This gives a posi- tive link between the name and a well-understood taxon, and since no other such link can be made, and since nothing in the protologue contradicts this usage, it seems reasonable to accept it even though Sloane’s plant was probably not seen by Linnaeus (Stearn, 1957). The plant to which this name has been applied is found in most tropical regions, and is distinguished from similar species by the shortness of the stalk of the female flower, which forces the cyathium to split down one side as the 1966] Euphorbia subgenus Chamaesyce — Burch 165 capsule develops. Bernard, Icones Bogorienses, 4, t. 315. 1910, illustrates this character well. Grisebach, in his Flora of the British West Indian Islands (1864), treats the taxon as a variety thymifolia of E. maculata L., but Boissier in DC. Prodromus (1862), Drury in Handbook of the Indian Flora (1869), Urban in Symb. Antill. (1910), and most subsequent authors consider that it merits specific rank. This usage is now so widespread and so well understood that nomenclatural stability will be best served by maintaining it, unless some contrary intention on the part of Linnaeus can be shown. MISSOURI BOTANICAL GARDEN, ST. LOUIS, MISSOURI LITERATURE CITED CRoIzAT, L. 1945. “Euphorbia chamaesyce" in the United States. Bull. Torr. Bot. Club 72: 312-318. 1947. Euphorbia maculata L. Bull. Torr. Bot. Club 74: 153-155. 1948. Euphorbia maculata: A rejoinder. Bull. Torr. Bot. Club 75: 188. 1962. Typification of Euphorbia maculata L. A Re- statement and a Conclusion. Webbia 17: 187-205. FosBERG, F. R. 1946. Application of the name Euphorbia maculata L. Rhodora 48: 197-200. 1947. Euphorbia maculata again. Bull. Torr. Bot. Club 74: 332-355. 1953. Typification of Euphorbia maculata L. Rhodora 55: 241-243. JacQUIN, N. J. 1772. Hortus Botanicus Vindobonensis 2: 87, t. 186. Vienna. KLoTzsCcH, J. F. 1843. Nov. Act. Nat. Cur. 19: (supp. 1): 414. La4GASCA, M. 1816. Genera et Species Plantarum... 17. Madrid. LINNAEUS, C. 1753. Species Plantarum ed. 1. Stockholm. 1771. Mantissa plantarum altera. Stockholm. MicHaUX, F. A. 1803. Flora boreali-americana. Paris. SAVAGE, S. 1945. A catalogue of the Linnaean Herbarium. London. STEARN, W. T. 1957. An introduction to the Species Plantarum and cognate botanical works of Carl Linnaeus, (Introduction to Ray Society facsimile of Species Plantarum ed. 1). London. 166 Rhodora [Vol. 68 SVENSON, H. K. 1945. On the descriptive method of Linnaeus. Rhodora 47: 273-302, 363-388. TORREY, J. 1843. Flora of the State of New York, Albany. WHEELER, L. C. 1939. A miscellany of the New World Euphorbia- ceae — II. Contr. Gray Herb. 127: 48-78. 1941. Euphorbia subgenus Chamaesyce in Canada and the United States exclusive of southern Florida, Rhodora 43: 97-154, 168-205, 223-286. (Reprinted as Contr. Gray Herb. 136). 1960. Typification of Euphorbia maculata. Rhodora 62: 134-141. IDENTITY OF WITCHWEED IN THE SOUTHEASTERN UNITED STATES In 1956 witchweed was found to be parasitizing field plantings of corn (Zea mays L.) in North Carolina and South Carolina. Because potential loss from this infestation is severe, immediate steps were effectively taken to prevent the spread of the parasite. The original Specimens were tentatively identified as Striga asiatica (L.) O. Kuntze. The botanical nomenclature of this genus has been subject to interpretation by the various botanists who have dealt with it and only recently has an effective review been written (Saldanha, C. J. 1963). In late 1964, Dr. V. P. Rao, a visitor from India, ques- tioned the identification of the witchweed in the United States. He kindly furnished a copy of the Saldanha paper and a suite of specimens identified by Saldanha with which the local Striga could be compared. No doubt remains that this plant is Striga lutea Lour., which includes one of the elements of S. asiatica, a name now abandoned as a source of confusion. However, Rao writes (in a personal communication) * ... the species found in the United States has red flowers which I have never seen in the common witchweeds which we come across in this country [India]. While it is true that the bulk of the population in the United States has very bright chinese-red flowers, individuals with lemon yellow flowers occasionally appear. Rarely an individual plant is found in which the flowers are yellow with red marks or stripes. Perhaps the flower color is due to the original introduction of red-flowered plants from Kottayam, India, where a population is known, or from South Africa, where the bulk of the plants of this species are said to be red-flowered. C. EARLE SMITH, JR. BOTANIST, CROPS RESEARCH DIVISION, AGRICULTURAL RESEARCH SERVICE, U.S. DEPARTMENT OF AGRICULTURE, BELTSVILLE, MARYLAND. 167 BROMUS MOLLIS AND ALLIES IN NEW ENGLAND FRANK C. SEYMOUR A problem like that in many other species is presented by Bromus mollis L., in which confusion has arisen because of unrecognized species, varieties, or forms. Apparently this species has been interpreted as including all specimens with short peduncles and hairy lemmas. In its form with glabrous lemmas, it has often been confused with B. race- mosus L., and the form of B. commutatus Schrader with hairy lemmas has often been mistaken for typical B. mollis. To Mr. Peter S. Green of the Arnold Arboretum I am indebted for calling my attention to the presence, in New England, of more than one species commonly included in Bromus mollis, for lending me a key which set apart ne- glected or little recognized taxa, and for making very val- uable suggestions regarding this article. To him I express my hearty thanks. I thank Dr. Richard A. Howard, Direc- tor of the Arnold Arboretum, and Dr. Reed C. Rollins, Di- rector of the Gray Herbarium, and Mr. Richard J. Eaton, Curator of the Herbarium of the New England Botanical Club, for the privilege of referring to specimens deposited in their institutions. I appreciate very much the help of Dr. Robert C. Foster, of the Gray Herbarium, and Dr. Lily M. Perry and Mr. Green, of the Arnold Arboretum, in finding references for original descriptions. I thank Mrs. Lazella M. Schwarten for the facilities of the Harvard Herbarium Library, and Dr. C. E. Hubbard of the Royal Botanic Gardens, Kew, England, for the reference for B. commutatus Schrader var. pubens Wats. Appreciation is expressed also to Dr. Howard W. Pfeifer, of the University of Connecticut and Mr. Leopold A. Charette, until recently of Yale University, for citations of specimens in their re- spective institutions. In 1833, Hardouin (Congr. Sci. France 1: 59. 1833) pro- posed a new name, Bromus Thominii, for a grass which had previously been described and named B. arenarius 168 1966] Bromus mollis — Seymour 169 Thomine-Desmasures (Mem. Soc. Linn. Calvados 1: 40. 1824) because Labillardiére had used the name B. arenarius for another species in 1804. However, many botanists have considered this species not sufficiently distinct from B. mol- lis; and in America, where it has been introduced, it has commonly been overlooked or ignored to the present time. In 1924, another segregate of B. mollis was recognized at specific rank, namely B. lepidus Holmberg (Botaniska Noti- ser 1924: 326.). Plants differing from the typical by having, or lacking, pubescence on the lemmas occur in these species and have often been treated as varieties. Here, the example of Fern- ald and Holmberg is followed by reducing them to forms. When a taxon differs from the typical variant in only one character and does not seem to have a distinct geographic range, it seems more appropriate to recognize it as a form rather than a variety. Much of the confusion in these species arises from the fact that all three of them and also Bromus commutatus Schrader and B. secalinus L. may have either glabrous or hairy lemmas. Consequently, pubescence on the lemmas, or lack of it, does not separate these species. In an effort to clarify this group and some others to which they are closely related, the following key is presented, dealing with all the New England species in the sections Bromium Dum. in the Manual of Grasses of the United States by A. S. Hitchcock, ed. 2 by Agnes Chase, 1951, and Zeobromus Griseb. in Gray’s Manual, ed. 8, by M. L. Fernald, 1950. A. Branches of panicle short; peduncles all or mostly shorter than spikelets; panicle erect; margins of lemmas overlapping the lemma next to them. B. Lower lemmas 8-11 mm long; lower glume (6-)7-9 mm long; upper glume (7-) 8-9 mm long. ...m 1. B. mollis. C. Lemmas hairy. «HH 1a. B. mollis f. mollis. C. Lemmas glabrous. ......... 1b. B. mollis f. leiostachys. B. Lower lemmas 5.5-7.5 mm long; lower glume 4.5-6 mm long; upper glume 5.5-6.6 mm long. D. Lower lemmas 6.5-7.5 mm long; lower glume 4.5-6 mm 170 Rhodora [Vol. 68 long; upper glume 6-6.6 mm long; grain shorter than palea. —— HERRERA 2. B. Thominii. E. Lemmas glabrous, except minutely scabrous margins. —— RE 2a. B. Thominii f. Thoméanii E. Lemmas hairy. .................. 2b. B. Thominii f. hirsutus. D. Lower lemmas 5.5-6.5 mm long; lower glume 4.5-5 mm long; upper glume 5.5-6 mm long; grain longer than DEI MM 3. B. lepidus. F. Lemmas glabrous, .................. 3a. B. lepidus f. lepidus. F. Lemmas hairy. .................... 3b. B. lepidus f. lasiolepis. A. Branches of panicle longer; peduncles all or mostly longer than spikelets; panicle erect or drooping; margins of lemmas not always overlapping the lemma next to them. G. Margins of lemmas inrolled at maturity, leaving axis visible; lemmas 6.5-9 mm long. ......... eee 4. B. secalinus. H. Lower lemmas 7-9 mm long; sheaths glabrous. ................-- RM 4a. B. secalinus var. secalinus. I. Lemmas awnless. .....cccccsssecscesesssssesessssceeccescsececccecsenscesceceete MEME 4c. B. secalinus var. secalinus f. submuticus. I. Lemmas with awns 1-6 mm long. J. Lemmas glabrous. .......... eese — 4b. B. secalinus var. secalinus f. secalinus. J. Lemmas hairy. ...... sese 4d. B. secalinus var. secalinus f. hirtus. H. Lower lemmas 6.5-7 mm long, glabrous; lower sheaths hairy. ...eeeeseeeeeeteee nre 4e. B. secalinus var, hirsutus. G. Margins of lemmas concealing axis, not inrolled; lower lemmas 6.5-12 mm long. K. Lower lemmas 6.5-8(-9) mm long. L. Anthers 3-4.5 mm long; upper lemmas about as long as lower; panicle open, its branches spreading, lower branches more than two at a node, each bearing two or more spikelets; lower sheaths hairy, upper sheaths glabrous. .......eeeeeeree eee eere 7. B, arvensis. L. Anthers 2-2.5 mm long; upper lemmas distinctly shorter than lower; panicle rather dense. its branches erect or ascending, lower branches 1-2 at a node, each bearing 1 (rarely 2) spikelets; sheaths all hairy. ...... RR 5. B. racemosus. K. Lower lemmas 9-12 mm long. M. Spikelets drooping, 0.8-1.3 em wide, awnless or nearly so; anthers 0.8-1 mm long; branches of panicle usually bearing 1 spikelet; lower lemmas about as wide as long; palea much shorter than lemma, about 6 mm long. .......eeeeeeeeene 8. B. brizaeformis Fischer & Meyer 1966] Bromus mollis — Seymour 171 M. Spikelets erect or ascending, 4.5-6 mm wide; awns usually 4-10 mm long; anthers 1.5-2 mm long; branches of panicle usually bearing 2 or more spikelets; lower (and other) lemmas much longer than wide; palea about as long as lemma. ..............- 6. B. commutatus. N. Lemmas glabrous except minutely scabrous mar- EIE esie 6a. B. commutatus f. commutatus. N. Lemmas hairy. ........ 6b. B. commutatus f. pubens. To assist further in identifying these species, a few rep- resentative specimens are cited. Bromus mollis is too familiar to require citations; likewise the typical forms of some other species. It will be evident from the list that in America B. Thominii f. hirsutus also is frequent whereas the other species and forms most closely related to them are less common. 1. Bromus mollis L. Sp. Pl. ed. 2. 112. 1763. 1a. Bromus mollis f. mollis. 1b. Bromus mollis f. leiostachys (Hartman) Fernald, Rhodora 35: 316. 1933. In this article Fernald makes clear the reasons why this infraspecific epithet must be used instead of leptostachys of Persoon. REPRESENTATIVE SPECIMENS. MASSACHUSETTS: Roxbury, 1878, E. Faxon (YT); Lancaster, 1 July 1943, F. C. Seymour 7240 (NEBC). CONNECTICUT: Fairfield, 12 June 1914, E. H. Eames 8753 (GH). NEW JERSEY: Camden, 3 June 1877, Parker (GH). DISTRICT OF COLUMBIA : Washington, 31 May 1905, A. S. Hitchcock, Am. Grasses 891 (GH), distributed as B. racemosus L., see Rhodora 35: 316. 1933. MICHIGAN: near Lansing, 22 June 1886, L. H. Bailey, Jr. (GH). IDAHO: New Plymouth, 21 May 1910, J. F. Macbride 94 (GH). OREGON: without precise locality, Elihu Hall 1871 (GH). 2. Bromus Thominii Hardouin, Congr. Sci. France 1: 59. 1833. 2a. Bromus Thominii f. Thominii. REPRESENTATIVE SPECIMENS. MASSACHUSETTS: Stockbridge, 25 July 1914, R. Hoffmann (NEBC). RHODE ISLAND: Block Island, 12 Aug. 1919, C. B. Graves (NEBC). CONNECTICUT: Bridgeport, 24 June 1915, E. H. Eames 9022 (NEBC). BRITISH COLUMBIA: Vancouver Island, 20 June 1907, C. O. Rosendahl 1887 (GH). 2b. Bromus Thominii f. hirsutus (Holmberg) Hylander, Uppsala Univ. Arskr 1945 (7): 85. 1945. REPRESENTATIVE SPECIMENS. MAINE: Seal Harbor, Mt. Desert Island, 9 July 1928, G. L. Stebbins, Jr. 327 (NEBOC) ; N. Berwick, 4 July 1892, 172 Rhodora [Vol. 68 J. C. Parlin (GH, NEBC). MASSACHUSETTS: Salem, 26 May 1890, W. P. Conant (NEBC) ; Cambridge, 10 June 1891, M. L. Fernald (NEBC) ; Lincoln, 16 June 1913, M. L. Fernald (NEBC) ; Boston, Parker's Hill, June 1877, C. E, Faxon (Gu); Centerville, 17 June 1896, E. F. Wil- liams (NEBC) ; Nonamesset, 29 May 1928, J. M. Fogg, Jr. 3394 (NEBC) ; Penikese, 3 July 1928, J. M. Fogg, Jr. 3412 (NEBC); Chilmark, 23 Aug. 1895, Sydney Harris (NEBC); Nantucket, 15 June 1909, Mrs. N. F. Flynn (VT). RHODE ISLAND: Block Island, 30 June 1938, K. P. Jansson (CONN). CONNECTICUT: New London, 1891, C. B. Graves (GH); 16 June 1932, K. P. Jansson (CONN); Hartford, 2 June 1901, A. W. Driggs (GH). NEW YORK, Long Island, Orient Point, 26 July 1940, K. P. Jansson (CONN). MICHIGAN: Meridan Twp. T4N, RIW sec. 7, Ingham Co., G. W, Parmelee 231 (VT). ILLINOIS: Peoria, July 1904, F. E. McDonald (GH). NEVADA: King's Canon, C. F. Baker 1104 (VT). CALIFORNIA: Mather, 30 May 1931, D. D. Keck 1126 (GH); Redding, Shasta Co., 4 July 1914, L. E. Smith (GH); Webb Ranch, San Mateo Co., 5 May 1960, D. M. Porter 678 (GH). WASHINGTON ; Pullman, 8 July 1950, L. C. Hulbert 499 (CONN). 3. Bromus lepidus Holmberg, Bot, Not. 1924: 326. 1924. 3a. Bromus lepidus f. lepidus. REPRESENTATIVE SPECIMENS. MASSACHUSETTS: Stockbridge, 21 June 1915, R. Hoffmann (NEBC). CONNECTICUT: Bridgeport, 24 June 1915, E. H. Eames (vT). 3b. Bromus lepidus f. lasiolepis Holmberg, Bot. Not. 1924: 326. 1924. Syn. f. micromollis (Krósche) Holmberg in Hartman, Handb. Skand. Fl. ed. Holmberg, 265. 1926. REPRESENTATIVE SPECIMEN. MASSACHUSETTS: Jamaica Plain, 19 June 1963, P. S. Green 49/63 (NEBC). 4. Bromus secalinus L. Sp. Pl. 76. 1753. 4a. Bromus secalinus var. secalinus. 4b. Bromus secalinus var. secalinus f. secalinus 4c. Bromus secalinus var. secalinus f. submuticus (Reichenb.) Ascher- son & Graebner, Syn. Mittelleurop. Fl. 2(1): 604. 1901. REPRESENTATIVE SPECIMENS. MAINE: E. Auburn, 1 Sept. 1897, ex. herb. E. B. Chamberlain (NEBC); N. Berwick, 26 June 1896, J. C. Parlin (NEBC). MASSACHUSETTS: Jamaica Plain, 6 Aug. 1925, E. J. Palmer 28164 (NEBC); New Bedford, s.d., E. W. Hervey (NEBC). CONNECTICUT: West Haven, 30 June 1908, R. W. Woodward (NEBC). 4d. Bromus secalinus var. secalinus f. hirtus (F. Schultz) Seymour, stat. nov. Basionym: B. mutabilis F. Schultz var. hirtus F. Schultz, Flora 32: 234. 1849. Synonym: B. secalimus L. var. hirtus (F. Schultz) Ascherson & Graebner, Syn. Mitteleurop. Fl. 2(1): 604. 1901. 1966] Bromus mollis — Seymour 173 REPRESENTATIVE SPECIMENS. MAINE: Bethel, 25 June 1938, L. A. Wheeler (NEBC). CONNECTICUT: Southington, 26 June 1903, C. H. Bissell (NEBC). 4e. Bromus secalinus var. hirsutus Kindberg, Bot. Not. 1887: 32. 1887. REPRESENTATIVE SPECIMENS: VERMONT: Townshend, 23 June 1912, L. A. Wheeler (NEBC). MASSACHUSETTS: Revere, 1878, H. A. Young (NEBC); Needham, 30 July 1883, T. O. Fuller (NEBC) ; Milton, 15 July 1918, N. T. Kidder (NEBC); Hingham, 11 July 1926, C. H. Knowlton (NEBC). RHODE ISLAND: Edgewood, 26 June 1913, T. Hope (NEBC). CONNECTICUT: Waterbury, 14 July 1910, A. E. Blewitt 160 (NEBC). 5. Bromus racemosus L. Sp. Pl. ed 2. 114. REPRESENTATIVE SPECIMENS: MAINE: Foxcroft, 25 June 1894, M. L. Fernald 565 (NEBC) ; Seal Harbor, 9 July 1928, G. L. Stebbins, Jr. 326 (NEBC) ; 29 June 1899, E. L. Rand (NEBC). MASSACHUSETTS: Charles- town, 19 Aug. 1879, C. E. Perkins (NEBC) ; Brookline, 14 June 1883, C. W. Jenks (NEBC); Nantucket, 21 June 1910, E. P. Bicknell 9773 (NEBC). 6. Bromus commutatus Schrader, Fl. Germ. 1: 353. 1806. Synonym: B. pratensis Erhart ex Hoffm. Deutschl. Fl. ed. 2. 2: 52. 1800, non Lamarck 1783. 6a. Bromus commutatus f. commutatus. 6b. Bromus commutatus f. pubens (Wats.) Seymour, stat. nov. Basionym: B. commutatus Schrader var. pubens Wats., Phytologist 1: 1062. 1844. REPRESENTATIVE SPECIMENS: MAINE: N. Berwick, 27 June 1897, J. C. Parlin 849 (NEBC). VERMONT: Ely, 27 June 1935, A. A. Beetle 37 (NEBC). MASSACHUSETTS: Milton, 8 June 1924, N. T. Kidder (NEBC). CONNECTICUT: North Haven, 30 June 1962, R. J. Eaton 2 June 1909 & 18 June 1910, E. P. Bicknell (NEBC). 7. Bromus arvensis L. Sp. Pl. 77. 1753. REPRESENTATIVE SPECIMENS: MASSACHUSETTS: Arnold Arboretum, 8 June 1942, E. J. Palmer 46002 (NEBC) ; Grafton, 30 June 1951, B. N. Gates 27059 (NEBC); Worcester, 10 July 1946, B. N. Gates 20499 (NEBC). CONNECTICUT: North Haven, 30 June 1962, R. J. Eaton (NEBC) ; Bridgeport, 1 July 1912, E. H. Eames 8543 (NEBC) ; Bridge- port, 10 July 1912, E. H. Eames 8568 (NEBC). 8. Bromus brizaeformis Fischer & Meyer, Ind. Sem. Hort. Petrop. 3: 30. 1837. REPRESENTATIVE SPECIMENS: VERMONT: Charlotte, 27 June 1880, 174 Rhodora [Vol. 68 C. G. Pringle (VT). MASSACHUSETTS: South Boston, 28 May 1879 & 8 June 1879, C. E. Perkins (NEBC) ; Charlestown, 23 July 1881, Herb. C. W. Swan (yu); New Bedford, 15 July 1903, Herb. E. W. Hervey (NEBC). CONNECTICUT: East Haddam, 5 July 1903, H. S. Clark (NEBC); New Haven, 1884, R. W. Woodward (NEBC). WASHINGTON: Pullman, May 1898, A. D. E. Elmer 1330 (vT); Whitman Co., Karniak Mt., Willamette Mer. 20 July 1951, L. C. Hulbert 2326 (CONN). ONTARIO: comte d'Haldimand, 2 July 1936, F. Marie-Victorin, F. Rolland-Germain, F. Dominique 46200 (CONN). PRINGLE HERBARIUM UNIVERSITY OF VERMONT, BURLINGTON CONTRIBUTION TO THE FUNGUS FLORA OF NORTHEASTERN NORTH AMERICA. IV" HowARD E. BIGELOW AND MARGARET E. BARR? The following continues a report of unusual or rare species of higher fungi that we have collected in New Eng- land during recent years. In the summer of 1963 our field work was done in the White Mountains of New Hampshire, and during the summer and fall of 1964 we collected in the Mt. Mansfield area of Vermont. Although both years were generally dry in New England, sufficient moisture did ac- cumulate at these two areas to produce a number of interest- ing species. The senior author is responsible for the information on agarics, while the junior author has provided the data on ascomycetes. The colors within quotation marks are from Ridgway, R., 1912. Color Standards and Color Nomencla- ture. Washington, D. C. The collections are deposited in the Herbarium of the University of Massachusetts, Amherst, unless otherwise noted. Recent research has been possible through the support of National Science Foundation Grant G-19534. We are grate- ful also to Mr. Stanley J. Smith, Senior Curator of Botany, New York State Museum, and Dr. I. MacKenzie Lamb of the Farlow Herbarium, for the opportunity to examine the type specimens cited. Mr. R. Goodrich, District Ranger, White Mountains National Forest, New Hampshire, and Mr. Perry Merrill, Commissioner, Department of Forests and Parks, State of Vermont, kindly extended permission for our col- lecting in New Hampshire and Vermont. We also wish to acknowledge the Research Council, University of Massa- chusetts for financial assistance with the cost of plates and reprints of this article. T. Rhodora 62: 186-198. 1960. II. Rhodora 64: 126-187. 1962. III. Rhodora 65: 289-309. 1963, Contribution from the Department of Botany, University of Massachusetts, Amherst. "Mrs. Howard E. Bigelow. 175 176 Rhodora [Vol. 68 * Plate 1331 Callistosporium luteoolivaceum (Berk. & Curt.) Singer. x 1. BASIDIOMYCETES Callistosporium luteoolivaceum (Berk. & Curt.) Singer, Lloydia 9: 117. 1946. Plate 1331. Agaricus luteoolivaceus Berk. & Curt. Ann. Mag, Nat. Hist. III, 4: 286. 1859. Agaricus coloreus Peck, Buffalo Soc. Nat. Sci. Bull. 1: 46. 1873. Agaricus rubescentifolius Peck, N. Y. State Mus. Rep. 39: 38. 1886. Collybia luteoolivacea (Berk. & Curt.) Sacce. Syll. Fung. 5: 215. 1887. Collybia colorea (Peck) Sace. Syll. Fung, 5: 230. 1887. Collybia rubescentifolia Peck, N. Y. State Mus. Rep. 41: 83. 1888. Tricholoma rubescentifolium (Peck) Sace. Syll. Fung, 9: 15. 1891. Collybia colorea var. rubescentifolia (Peck) Peck, N. Y. State Mus. Rep. 49: 51. 1896. Callistosporium psilocybe Murr. & Singer, Mycologia 36: 363. 1944. Pileus 1-4 cm. broad, convex at first with an inrolled and slightly incurved margin, expanding to broadly convex, finally plane, margin slightly inrolled for some time, exceeding gills at times, sometimes sulcate but not pellucid-striate, surface glabrous when moist, matted 1966] Fungus Flora — Bigelow and Barr TUE fibrillose or innately radiate-fibrillose when faded, dull yellow at first becoming cinnamon to rufous brown (“orange cinnamon”, “Hays russet”, “brick red”, “Brussels brown”, “cinnamon rufous”), hygroph- anous, fading to yellowish or buff (near “antimony yellow”, “deep colonial buff”, “light ochraceous salmon”, “light ochraceous buff”), disc darkest; flesh thin, rather brittle, watery yellowish fading to pale yellowish or whitish, no odor and taste or taste rarely bitter. Lamellae adnate or adnexed, finally emarginate then seceding, close, broad (up to 7 mm.), not forked or intervenose, arched at times, yellow (“empire yellow", “yellow ochre”, “antimony yellow”, “cha- mois”) not fading, edges crenate or merely uneven, at times marginate with reddish brown in age. Stipe 2.5-5 em. long 1.5-5 mm. thick at apex, equal or the base en- larged (up to 7 mm.), and tapering upward, solid at first (interior concolorous with pileus flesh), becoming hollow, at times eccentric, often curved, surface innately fibrillose-striate, concolorous with lamel- lae, darkening to dingy “ochraceous buff", “cinnamon”, or “dresden brown”, base with white tomentum and sometimes a few white rhizoids. Spores 4.5-7 X 2.5-4 n, elliptical, sometimes elliptic-oblong, smooth, not amyloid, often with vinaceous droplets or contents in KOH, white in mass; basidia 16-28 X 4.5-7 u, 2- or 4-spored, often with vinaceous droplets or granules in KOH; pileus tissue: fresh material yellowish in water mount, revived material pale vinaceous in KOH, pigment in cell contents and often punctate-encrusted, surface at times with protruding cystidioid end cells, cuticular hyphae cylindrical, 2.5-5.5 & in diameter, tramal hyphae cylindrical to slightly inflated, 2-8 u in diameter, clamp connections absent; gill trama subparallel, broad, hyphae cylindrical to slightly inflated, 2.5-7.5(-9) & in diameter. Scattered to gregarious or occasionally cespitose, On conifer logs and stumps. Material examined: Bigelow 10788, near Katahdin Stream Camp- ground, Baxter State Park, Maine, August 6, 1962; 7309, 7310, 7311, Amherst, Massachusetts, August 4, 1958; 7411, Amherst, August 9, 1958; 7515, Amherst, August 17, 1958; 7600, Mt. Toby, Sunderland, August 27 1958; 7739, Mt. Toby, September 15, 1958; 7802, Mt. Toby, September 20, 1958; 8264, Amherst, July 25, 1959; 8956, Wahconah Falls, Dalton, July 26, 1960; 9007, D.A.R. State Forest, Goshen, August 2, 1960; 9234, D.A.R. State Forest, September 1, 1960; 9624, Pittsfield State Forest, Pittsfield, July 26, 1961; 9683, New Salem, August 2, 1961; 11990, near Sawyer Rock, White Mts. Nat, Forest, New Hampshire, July 27, 1963; 12246, Oliverian Brook Trail, White Mts. Nat. Forest, August 13, 1963; C. H. Peck, North Greenbush, New York (type of Agaricus rubescentifolius, nys), August; C. H. Peck, Croghan (type of Agaricus coloreus pro parte, NYS). This agaric can be very confusing in the field as the 178 Rhodora [Vol. 68 colors of the pileus and stipe vary markedly according to the age of the carpophore or its moisture content. In my field work the brown stage of the pileus seemed to be encountered much more frequently than the yellow stage. I have not had the opportunity to study other species in this genus, but the carpophores of C. luteoolivaceum at least turn a rather bright vinaceous red when dilute KOH solu- tion is applied to fresh material. In drying, specimens be- come a dark vinaceous color over all. Clitocybe albimontana Bigelow, nomen novum Omphalina oreades Singer, Pap. Mich. Acad. Sci., Arts & Letters 32: 123, 1946. Pileus (2.5-)4-10 mm. broad, convex with an incurved margin at first, expanding to broadly convex, dise rarely slightly depressed, glabrous and striate moist, disc brown, striations dark brown (“bister”), paler in between striations (near “snuff brown”), hygro- phanous, fading to sordid alutaceous, fibrillose and opaque faded, fibrils often erect near margin; flesh thin, pliant, concolorous with pileus moist or faded, no odor, taste mild. Lamellae decurrent, usually evenly and forming a collar on the stipe apex, narrow to moderately broad (+ 1 mm.), distant, rarely forked, not intervenose, arched, color a pale “buffy brown", edges even. Stipe 6-9 mm. long, 0.5-1.5 mm. thick, equal or tapering downward from a slightly enlarged apex, pruinose at first becoming glabrous, solid, rather soft, darker brown than lamellae ("buffy brown", *Sac- cardo's umber”). Spores 7.5-10 X 3-4 a, shape variable: elliptical to obovate or sub- cylindrical in face view, sublacrymoid or lacrymoid in side view, some- times slightly bent at either end, smooth, not amyloid; basidia 24-31 AX 5.5-7 u, 4-spored, sterigmata up to 7 u long, basidioles somewhat irregular in shape at times; cystidia not differentiated; pileus tissue: cuticle brown in KOH, pigment finely but distinctly encrusted, hyphae cylindrical or inflated, 3-8.5 u in diameter, trama pale brown, hyphae smooth or encrusted, cylindrical or inflated, 3-11 4 in diameter, clamp connections absent; eill trama interwoven, pale brown, hyphae smooth or encrusted, cylindrical or inflated, 1.5-7(-11) 4 in diameter. Scattered or gregarious. On moss or sandy soil in alpine zone. Material examined: Bigelow 11918, 11919, Lake-of-the-Clouds Trail from peak, Mt. Washington, New Hampshire, July 23, 1963; 12339, 12340, trail to Alpine Garden from peak, Mt. Washington, August 19, 1963; R. Singer, Mt. Washington (type of Omphalina oreades, F). 1966] Fungus Flora — Bigelow and Barr 179 O% Plate 1332 Hygrophorus lilacinus (Laest.) M. Lange. X L The preceding records are apparently the first of this species since Singer found it twenty years ago in the same locality. C. albimontana is easily overlooked because of its small size and a tendency to grow under overhanging moss where water or paths have cut down into the soil. A number of carpophores were found on both east and west sides of the mountain below the peak. In transferring Omphalina oreades to Cline rape. a new species epithet is necessary because of Clitocybe oreades Murrill. Clitocybe asterospora (Lange) Moser apud Gams, Kleine Kryptogamen Flora II, p. 52. 1953. Gregarious. On wet moss, in depression between rocks. Material examined: Bigelow 14223, top of Mt. Mansfield, Vermont, September 24, 1964. This agaric appears to be rare in North America. I know of fruit- ings only in Michigan and Ontario besides the specimens found on Mt. Mansfield. Hygrophorus lilacinus (Laest.) M. Lange, Meddelelser om Gr¢nland 148(2): 68. 1957. Plate 1332. 180 Rhodora Plate 1583 Hygrophorus monticola Hesler & Smith. X 1. 1966] Fungus Flora — Bigelow and Barr 181 Agaricus (Omphalia) lilacinus Laestadius, Lapp. Torn. p. 45, n. 156. 1860. Hygrophorus violeipes M. Lange, Meddelelser om Grénland 147 (11) : 18. 1955. Material examined: Bigelow 13656, Mt. Mansfield, Vermont, August 29, 1964. A small group of carpophores was found in a mossy de- pression near the “upper lip” on top of Mt. Mansfield. The bright luteous colors of cap and gills and the violet stipe make this species easy to recognize in the field. In micro- scopic characters as well as all field characters, my speci- mens agree exactly with Lange’s (1955) description of H. violeiipes. Apparently Bigelow 13656 is the first record of H. lilacinus in North America. In 1957, Lange placed Omphalia luteolilacina Favre in synonymy with H. lilacinus. While there is certainly a resemblance between the two species, I do not agree with Lange’s treatment. As described by Favre (1955), O. luteo- lilacina lacks clamp connections, has elliptic-cylindric spores and a hispid stipe. These characters are unlike H. lilacinus and certainly sufficient to maintain O. luteolilacina as a separate taxon. Hygrophorus monticola Hesler & Smith, North American Species of Hygrophorus, p. 359. 1963. Plate 1333. Pileus 1.5-16 em. broad, at first convex with an incurved and inrolled margin, not striate, slowly expanding to plane with a decurved margin, elevated in age, surface dry and unpolished, finely matted-fibrillose on disc under a lens, glabrous at the margin, cracked at times in age or dry weather, color dingy "cinnamon" to *vinaceous cinnamon” or “sayal brown", margin whitish at times, “russet vinaceous" over all when dry; flesh thick on dise, tapering abruptly to margin, whitish (near “cartridge buff"), firm, brittle, odor strong, sweet like that of cherries, taste mild. Lamellae decurrent (evenly at times and forming a collar on the stipe apex), distant, thickened, broad (3-11 mm.), occasionally forked, sides venose at times, arched, color “cream buff" to “ochraceous buff", edges even. Stipe 2-12 em, long, apex 0.8-4.2 cm. thick, equal in smaller car- pophores but tapering downward strongly in larger ones, solid (white 182 Rhodora [Vol. 68 within), no veil, surface dry and unpolished, velvety to pubescent under a lens, appressed in age, whitish. Spores 9-11.5(-13) X 6-7.5 4, elliptical, smooth, not amyloid; basidia 46-68 X 5.5-8 u, 4-spored; cystidia not differentiated; pileus tissue: cuticle pale yellowish in KOH, pigment dilute in the contents of some surface hyphae, cuticle an ixocutis, hyphae cylindrical, 2.5-4 4 in diameter, walls thin, tramal hyphae mostly cylindrical, 3-9 (-14) u in diameter, walls often slightly thickened, clamp connections present; gill trama: divergent in young carpophores but appearing more interwoven in large caps, hyphae cylindrical, 4-7.5 u in diameter. Gregarious to cespitose, In soil and needle beds under spruce. Material examined: Bigelow 13917, 13984, 14124, 14204, 14278, 14323, 14350, 14373, Lamoille and Washington counties, Vermont, September and October. In Hesler and Smith's (1963) monograph of Hygro- phorus, H. monticola is described from specimens collected in Idaho and eastern Canada. The pilei measured 2-5 cm. broad, and the stipes 3-6 em. X 3-25 mm. In view of these dimensions, the Vermont specimens are most impressive in size. The odor of cherries was very pronounced as well and could be detected without crushing the flesh, sometimes from several feet away. Of further note, and perhaps worthy of emphasis in diagnosis, was the tendency of the pileus to become more intense in color with drying. Hygrophorus nitiosus Blytt, Norges Hymenomycetes. Videnskab Selskabets Skrifter. Math. Naturv. Kl. 1904, No. 6. 1905. Gregarious. On soil of dirt road in hardwoods. Material examined: Bigelow 13132, Stowe, Lamoille Co., Vermont, July 28, 1964. Hygrophorus tahquamenonensis Smith & Hesler, Sydowia 8: 831. 1954. Gregarious to subcespitose, on moss by stream, near hardwoods. Material examined: Bigelow 13242, Bingham Falls, near Stowe, Lamoille Co., Vermont, August 3, 1964. Mycena atkinsoniana A. H. Smith, North American Species of Mycena, p. 144, 1947. On leaves and humus under beech. Material examined: Bigelow 13936, Bingham Falls, near Stowe, Lamoille Co., Vermont, September 9, 1964; 14059, near Lake Mans- field, Lamoille Co., September 14, 1964. 1966] Fungus Flora — Bigelow and Barr 183 Paxillus curtisii Berkeley, Ann. Mag. Nat. Hist. II, 12: 423. 1853. On hemlock logs. Material examined: Bigelow 11906, South Conway, Carroll Co., New Hampshire, July 22, 1963; 12092, same locality, August 3, 1963; 12398, 12399, same locality, September 4, 1963. Phaeocollybia kauffmanii (Smith) Singer, Rev. Mycol. 5: 11. 1940. Naucoria kauffmanii Smith, Mycologia 29: 52. 1937. Pileus 1.3-5.5 em. broad, obtuse-conie or convex at first with mar- gin slightly incurved and narrowly inrolled, not striate, expanding somewhat to broadly convex with margin remaining incurved and inrolled, sometimes lobed, surface glutinous or viscid, smooth, gla- brous, color watery “cinnamon” to “sayal brown", fading to “cinnamon buff", at times with a few dark reddish flecks; flesh thick, firm, whitish, odor and taste farinaceous. Lamellae emarginate (notch small and abrupt, superficially ap- pearing '"free"), close, narrow to medium broad (2-7 mm.), forked at times, white at first then “vinaceous buff" or “pale pinkish buff", edges slightly uneven. Stipe 4-13 cm. long, apex 5-11 mm. thick, often subventricose (up to 16 mm. thick), attenuated and radicating (about half of stipe above soil), at times twisted, usually curved below, surface innately fibrillose, moist but not viscid, solid (cortex thick and brown in sec- tion, interior firm, whitish to 'pale ochraceous buff"), color watery *cinnamon buff" above, rufous stained below, especially on lower por- tion embedded in soil, Spores 7.5-9 X 5.5-6 uw, ovate in face view, inequilateral in side view, with a minute apical pore but no conspicuous beak, rugulose- roughened, fulvous in KOH; basidia 23-31 X 7.5-10 u, 4-spored, pro- truding beyond the hymenium at maturity; cystidia: cheilocystidia abundant, pleurocystidia rare, both basidioid to clavate in shape, sometimes rather irregular and with broad lump near apex, rarely 2-celled, often with dilute brownish contents, smooth, thin-walled; pileus tissue: surface layer a distinct gelatinous pellicle in KOH, hyaline, hyphae 1.5-2.5 u in diameter, trama brown in KOH, pigment finely but distinctly encrusted on some hyphae in subpellicular zone, in smooth and thickened walls below, hyphae cylindrical to somewhat inflated, 4-13 u in diameter, clamp connections absent; gill trama parallel, brownish in KOH, pigment in thickened but smooth walls, hyphae cylindrical, 3-8 u in diameter. Seattered or subcespitose. On soil of river bank near hemlock. Material examined: Bigelow 13570, Bingham Falls, near Stowe, Vermont, August 20, 1964. P. kauffmanii is well known in the Pacific northwest 184 Rhodora [Vol. 68 although apparently not common. Hesler (1949) has re- ported its occurrence in Tennessee, but I have not found any other record from the east. My specimens from Ver- mont fit Smith’s (1957a) description except in dimensions of the carpophore. Western specimens of P. kauffmanii are considerably larger in size. Pleurotus elongatipes Peck, Jour. Myc. 14: 1. 1908. Pileus 2.5-14 cm. broad, convex to broadly convex or nearly plane, margin narrowly inrolled and incurved for some time, slightly pubes- cent when young, sometimes with short suleations, surface glabrous, mottled about the dise with small watery-appearing spots, white to pale avellaneous or pale buff ("cartridge buff", *light buff"); flesh usually rather thick on dise, firm, brittle, odor often Spicy and fra- grant when flesh crushed, taste not distinctive, Lamellae at first adnexed or adnate, becoming sinuate or emar- ginate, seceding at times, moderately broad (5-11 mm.) near stipe but narrowed at margin of pileus, usually close but sometimes crowded, not forked or intervenose, white or whitish, edges uneven and brittle in age, Stipe 3-10 em. long, apex 4-20 mm. thick, base usually somewhat enlarged and tapering upward, sometimes ventricose, solid (interior white and soft), cavernous in age, often curved, eccentric at times, surface with a thin fibrillose coating, becoming innately fibrillose- striate, apex pruinose to pubescent at times, white or nearly so. Spores 4-5.5 X 4-4.5 u, globose to subglobose, smooth, not amyloid, white to slightly cream tinged in deposit; basidia 16-24 X 4-6 u, 4-spored, carminophilous granules absent; cystidia not differentiated; pileus tissue: surface subgelatinous to gelatinous in KOH, cuticular hyphae cylindrical, 1.5-3 u in diameter, tramal hyphae cylindrical to slightly inflated, 2.5-13 u in diameter, clamp connections present, walls thin or slightly thickened and refractive; gill trama regular to subparallel, broad, hyphae mostly cylindrical, (1.5-)4-9(-11) 4 in diameter, wall thin or slightly thickened and refractive. Solitary or cespitose. On hardwood logs and stumps, sometimes in wounds of living maple. Material examined: Bigelow 8665, North Pond, Savoy Mt. State Forest, Florida, Massachusetts, September 22, 1959; 9362, Beartown State Forest, Monterey, September 27, 1960; 10509, Katahdin Stream Campground, Baxter State Park, Maine, August 6, 1962; 11146, near Norcross, Penobscot Co., August 18, 1962; 11259, Cedar Lake, Penob- scot Co., August 24, 1962; D. E. Fischer, Detroit, Michigan, October 5, 1905 (type, Nys); Bigelow 12524, Oliverian Brook Trail, White Mts. Nat. Forest, New Hampshire, September 7, 1963; 13842, near John- son, Lamoille Co., Vermont, September 2, 1964; 13937, Bingham 1966] Fungus Flora — Bigelow and Barr 185 Falls, near Stowe, Lamoille Co., September 9, 1964; 14048, Stowe, September 13, 1964; 14110, Ranch Brook Trail, Mt. Mansfield State Forest, September 16, 1964; 14116, Monroe State Park, Washington Co., September 17, 1964. Pleurotus elongatipes has proven to be quite common in New England. I have found it growing on birch and poplar debris as well as on dead or living maple. A description based upon specimens from Mt. Rainer, Washington, has been published by A. H. Smith (1957b). This species is a Pleurotus in the traditional sense of the genus, but the globose spores exclude it from modern inter- pretations. According to Singer's (1962) classification, the closest affinities may be with Hypsizygus, but some emenda- tion of this genus would be necessary to admit P. elonga- tipes. ASCOMYCETES Keissleriella caudata (Müller) Corbaz, Phytopath. Zeitschr. 28: 411. 1957. Figure 8. Material examined: Barr 2896, on cultivated rose stems, Conway, Massachusetts, May 7, 1961. K. caudata was described from Triticum stalks in Europe. The collection on rose stems agrees with the description of the species, although the ascospores are slightly longer, 15.5-17.5 X 3.5-4.5 y rather than 10-13 x 4-4.5 y. This species differs from K. cladophila (Niessl) Corbaz by smaller, thinner-walled ascostromata, lack of a clypeus, and narrower ascospores. Keissleriella is included in the Pleosporaceae. Lasiosphaeria chrysentera Carroll & Munk, Mycologia 56: 83. 1964. Material examined: Barr 3338, Abol Field, near Baxter State Park, Maine, July 6, 1962. The species, described from Costa Rica, is closely related to L. ovina (Pers. ex Fr.) Ces. & de Not. L. chrysentera differs in golden-yellow pigmentation of the hymenium, bright yellow globule at apex of ascus, and yellowish asco- spores. One additional collection has been examined from 186 Rhodora [Vol. 68 Idaho, A. W. Slipp, U.LF.P. Herb. 3834, on decorticated Alnus tenuifolia, 1943. Lasiosphaeria is a member of the Sordariaceae. Leiosphaerella falcata Barr, sp. nov., figures 1-4. Perithecia 180-300 yu diametro, 130-250 wu alta, immersa, gregaria, ostiola eccentrica vel lateralia, 60-120 u diametro, 70-150 & alta, per- iphysata; peridium 13-16.5 x crassum, brunneum, hyphis brunneis vestita. Asci 55-100 X 11-14.5 u, elliptica, unitunicata, ad apicem poro amyloideo, paraphysati. Ascosporae 46-86 X 3-5 4, fusiformae vel faleatae, hyalinae, uniseptatae, fasciculatae in ascis. Specimen typicum in caulis Aceris pensylvanicum L., a M.E. Barr lectus, n. 3460, prope “Katahdin Stream Campground, Baxter State Park, Maine, July 23, 1962", in herbario Univ. Mass. depositum. Perithecia 180-300 & in diameter, 130-250 u high, depressed globose, immersed beneath epidermis and raising it slightly, thickly scattered, ostiole eccentric to lateral, 60-120 u in diameter, 70-150 u high, pierc- ing the covering layer of substrate, canal periphysate, perithecial wall 13-16.5 & wide, of polygonal, brown, compressed layers of cells, inner layers hyaline, with brown hyphae surrounding perithecium and penetrating substrate tissues. Asci 55-100 X 11-14.5 à, elliptical, apex rounded or truncate, narrowed abruptly below spores into rounded base, wall single, thickened above with refractive pore, amy- loid in Melzer's, paraphyses deliquescent and forming an amorphous mass around asci. Ascospores 46-86 X 3-5 u, greenish hyaline, fusi- form to faleate, tapered to pointed ends, 1-septate in or near the middle, not constricted at the septum, wall thin, smooth, contents homogeneous or globular, in two overlapping fascicles in the ascus. In small dead branches of Acer pensylvanicum and A. spicatum. Material examined: Barr 3267, Norcross, Penobscot Co., Maine, June 30, 1962; 3460 (type), Katahdin Stream Campground, Baxter State Park, July 23, 1962; 3644, Norcross, August 18, 1962; 3995A, Stony Brook Trail, near Gorham, New Hampshire, July 26, 1963; 2098, Lake Munroe, Mt. Tremblant Park, Quebec, July 24, 1957; 4179, Underhill Camp Area, Mt. Mansfield State Forest, Vermont, June 30, 1964; 4334, Mt. Elmore State Park, July 16, 1964. L. falcata is closely related to L. praeclara. (Rehm) von Hóhnel, the type of the genus, and L. phoenicis (Chona & Munjal) Miiller & Ahmad in spore shape. The other mem- bers of the genus have shorter, relatively broad spores. The eccentric to lateral position of the beak of L. falcata and the longer spores distinguish it from the other species. This fungus shows affinities to members of Ceriospora but differs by lacking spore appendages and by simpler apical appara- 1966] Fungus Flora — Bigelow and Barr 187 Legend for Figures 1-8 Figs. 1-4. Leiosphaerella falcata Barr, sp. nov.: 1, perithecia be- neath bark, 2, ascus, 3, ascus showing amyloid apical ring, 4, asco- spores. Figs. 5-T. Mycosphaerella linnaeae Barr, sp. nov.: 5, asco- stroma, 6, ascus, 7, ascospores. Fig. 8. Keissleriella caudata (Müller) Corbaz: ascospores. Figs. 1, 5, X 80; figs. 2-4, 6-8, X 750. tus of the ascus. In the classification of Müller & von Arx (1962), Leiosphaerella is considered to be a member of the Amphisphaeriace2e. Mycosphaerella linnaeae Barr, sp. nov., figures 5-7. Ascostromata 80-100 & diametro, 72-100 & alta, immersa, gregaria, epiphylla; peridium 11-14.5 4 crassum, brunneum, hyphis brunneis. Asci 26-45 X 7.5-9 p, oblonga, bitunicata, aparaphysata. Ascosporae 16.5-21 X 2-3 y, fusiformae, hyalinae, uniseptatae, in duobus fasci- culatae in ascis. Specimen typicum in folia Linnaeae borealis L. var. americanae (Forbes) Rehd., a M.E. Barr lectus, n. 3831, prope “Millbrook Trail, Jefferson, White Mts. Nat. Forest, New Hampshire, July 6, 1963", in herbario Univ. Mass. depositum. Ascostromata 80-100 u in diameter, 72-110 » high, globose, slightly depressed or conical, immersed in leaf tissue, with broad blunt apex 188 Rhodora [Vol. 68 erumpent through upper epidermis, scattered thickly and evenly over most of leaf, wall thin, 11-14.5 u wide, two to three layers of dark brown, polygonal cells, compressed hyaline cell layers toward interior, brown hyphae copious in leaf tissues, Asci 36-45 X 7.5-9 u, oblong, apex rounded, base foot-like, wall double, thickened above, numerous in a fascicle, aparaphysate. Ascospores 16.5-21 X 2-3 u, greenish hyaline, fusiform, ends pointed, straight to curved, one-septate in mid- dle, not constricted at septum, wall thin and smooth, contents guttulate or with two globules in each cell, overlapping in two fascicles in the ascus. On overwintered leaves of Linnaea borealis var. americana. Material examined: Barr 3831 (type), Millbrook Trail, Jefferson, White Mts. Nat. Forest, New Hampshire, July 6, 1963; 1906, woods near chalet, Mt. Albert, Gaspe Prov. Park, Quebec, July 6, 1957. A species of Cercospora is intimately associated with M. linnaeae, and I suggest that it may form part of the life cycle of this species. This Cercospora has conidiophores short, brown, septate, erumpent through upper leaf epider- mis, often arising from apex of young ascostromata ; conidia 24-54 X 2 m, greenish hyaline, cylindrical, ends rounded, several-septate, contents guttulate. I have found no records of Cercospora on Linnaea (Chupp, 1953). At least twenty-four species of M ycosphaerella have been described on members of the Caprifoliaceae, but only two were reported on Linnaea. M. minor (Karst.) Johans. has smaller ascostromata, broadly saccate asci and clavate as- cospores 11-12 X 4 u. No measurements have been given for Sphaerella leightonii (Berk. & Br.) Cooke, but ascospore shape and septation is quite different from that of M. lin- naeae. As described in the literature, the other species of Mycosphaerella on members of the Caprifoliaceae do not appear similar to Barr 3831 and 1906. M. crepidophora (Mont.) Rehm and M. ebulina Petrak have elongate asco- spores and seem most closely related to M. linnaeae. M. crepidophora develops in spots in living leaves of Viburnum: the ascospores are cylindrical or oblong and are somewhat broader than those of M. linnaeae. M. ebulina occurs on dead leaves of Sambucus. The ascostromata are densely gregarious to form angular black spots, asci are larger, and ascospores broader than in M. linnaeae. The broad apex of 1966] Fungus Flora — Bigelow and Barr 189 the ascostromata and elongate, narrow fusoid ascospores are distinguishing characters of M. linnaeae. Plagiostigme petrakii Müller, Sydowia 18: 90. 1965. Plagiophiale petrakii (Müller) Petrak, Sydowia 18: 387, 1965. Material examined: Barr 4094, on overwintered leaves of Polygo- num viviparum, Alpine Gardens, Mt. Washington, New Hampshire, August 19, 1963. This rare and distinctive fungus has been illustrated and described recently by Müller (1965), who made several col- lections of it on Polygonum viviparum from the Swiss and Italian Alps. Petrak (1955) was first to collect P. petrakii, but thought it to be identical with Sphaerella eucarpa Karst. At that time he erected the genus Plagiophiale for the species. However, Sphaerella eucarpa Karst. is a species of Wettsteinina (Barr 1959, Müller and von Arx 1962), and therefore Plagiophiale Petrak is a nomenclatural synonym of Wettsteinina. Miiller (1965) has discussed in detail the generic position of the species on Polygonum. The North American material appears to be identical in all respects with that described from Europe. Plectosphaera salicis (Fckl.) v. Arx & Müller, Beitr. Kryptogamenfl. Schweiz 11(1): 204. 1954. Material examined: Barr 3928, Salix twigs, South Conway, New Hampshire, July 18, 1963. This appears to be the first record of P. salicis in North America. According to the literature, this pyrenomycete has been found previously only in Europe. My collection agrees with the description and figure in von Arx and Müller (1954). Podosphaera clandestina (Wallr. ex Fr.) Lév. Ann. Sci. Nat. Bot. III, 15: 86. 1851, var. clandestina Material examined: Barr 4578, Vaccinium angustifolium, top of Mt. Mansfield, Verment, August 28, 1964; 4583, V. uliginosum, data as above; 4585, Ledum groenlandicum, data as above, This minute powdery mildew has been recorded only rarely on members of the Ericaceae, chiefly species of Vac- cinium, and never, as far as I could determine, on Ledum. 190 Rhodora [Vol. 68 The specimens are in close accord with the description of P. oxyacanthae var. oxyacanthae in Salmon (1900). Thaxteria fusca (Fckl.) Booth, The Naturalist, London, p. 90. 1958. Sphaeria phaeostromoides Peck, N. Y. State Mus. Rep. 28: 77. 1876. Chaetosphaeria phaeostromoides (Peck) Sacce. Syll. Fung. 2: 93, 1883. Chaetosphaeria phaeostroma var. phaeostromoides (Peck) Ell, & Everh. North Amer. Pyreno., p. 160. 1892. Material examined: Barr 4041, on fallen beech, near Sawyer Rock Picnic Area, White Mts. Nat. Forest, New Hampshire, August 10, 1963; C. H. Peck, on decorticated Acer, North Greenbush, New York, September 1874 (type of Sphaeria phaeostromoides Peck, NYS). The collections cited above agree with Booth’s (1958) description and figures of T. fusca, and bear both conidial and perithecial stages. Ellis and Everhart (1892) cited a collection from Missouri, but otherwise the fungus appears to be collected rarely. The genus Thaxteria is a member of the Coronophoraceae. LITERATURE CITED ARX, A. J, VON and E. MÜLLER. 1954. Die Gattungen der amero- sporen Pyrenomyceten. Beitr. Kryptgamenfl. Schweiz 11(1): 1-434. Barr, M. E. 1959. Northern Pyrenomycetes I, Canadian Eastern Arctic. Contrib. Inst. Bot. Univ. Montreal 73: 1-101. BoorH, C. 1958. The genera Chaetosphaeria and Thaxteria in Bri- tain. The Naturalist, 83-90. CHUPP, C. 1953. A monograph of the fungus genus Cercospora. Ithaca, N. Y. 667 pp. ELLIis, J. B. and B. M. EVERHART. 1892. North American Pyreno- mycetes, Newfield, N. J. FAVRE, J. 1955. Les Champignons Supérieurs de la Zone Alpine du Pare National Suisse. Rés. tech, scient. entr. Pare Nat. Suisse 5: 1-212. Hester, L. R. 1949. Notes on Southern Appalachian Fungi, VIII. Jour. Tenn. Acad. Sci. 24: 81-93. HESLER, L. R. and A. H. SMITH. 1963. North American Species of Hygrophorus. Knoxville, Tenn. 416 pp. 1966] Fungus Flora — Bigelow and Barr 191 LANGE, M. 1955. Macromycetes Part II. Meddelelser om Grénland 147 (11) : 1-69. LANGE, M. 1957. Macromycetes Part III. Meddelelser om Grénland 148(2): 1-125. MÜLLER, E. 1965. Beobachtungen an Ascomyceten. Sydowia 18: 86-105. MÜLLER, E. and A. J. von ARX. 1962. Die Gattungen der didymo- sporen Pyrenomyceten. Beitr. Krypt. Fl. Schweiz, 11(2) : 1-922. PETRAK, F. 1955. Plagiophiale nov. gen., eine neue Gattung der sphaerialen Ascomyceten. Sydowia 9: 585-587. SALMON, E. S. 1900. A monograph of the Erysiphaceae. Torrey Bot. Club Mem. 9: 1-292. SMiTH, A. H. 1957a. A contribution toward a monograph of Phaeo- collybia, Brittonia 9: 196-217. 1957b. Additional new or unusual North American agaries, Sydowia, Beih. 1 (Festschr. f. Franz Petrak): 46-61. SINGER, R. 1962. The Agaricales in Modern Taxonomy, 2nd ed. Weinheim. 915 pp. NOTES ON RAFINESQUE'S SPECIES OF LECHEA (CISTACEAE) ROBERT L. WILBUR" It was suggested in an earlier paper (Wilbur & Daoud, 1961) that some of the twenty species of Lechea published by Rafinesque (1836) would eventually prove identifiable and that their names would replace binomials then cur- rent. This may have seemed an unwarranted assumption since three careful students of this technical genus had diligently studied Rafinesque’s monograph and concluded that the descriptions were not adequate to identify the species intended with any degree of confidence. Leggett (1878) republished much of Rafinesque’s monograph and suggested the possible identity of several of the taxa. Brit- ton (1894) indicated that he had worked closely with Leg- gett in attempting to unravel the problems of the genus and together they had “pored over Rafinesque’s mono- graph”. They finally concluded that “unless type specimens could be found, there could be no certainty” in applying any of the names. Hodgdon (1938:30) suggested that Rafinesque’s “perplexing” monograph left the genus in a “deplorable state" and claimed to have been able to identify only four of Rafinesque’s twenty new binomials. I believe however that the majority of Rafinesque’s species of Lechea can be identified with considerable cer- tainty even in the absence of types by comparing his descrip- tions with the species known to occur in the areas mentioned in the original publication. The results of this analysis are presented in the following paragraphs presented in the same order as Rafinesque’s monograph (1836) and followed chronologically by his other papers dealing with the genus. Unfortunately, for stability’s sake, three of Rafinesque’s ‘Grateful acknowledgement is made to the National Research Foun- dation for a grant of research funds to Duke University (NSF- G18799) which were used in part for this study. 192 1966] Lechea — Wilbur 193 binomials must replace those of species long known by later names. 1. Lechea pulchella Raf., New Fl. N. Am. 1: 91. 1836. Rafinesque definitely mentioned L. pulchella only from the pine barrens of New Jersey although suggesting that its range probably extended to the south. The species was described as being "quite smooth" and, as a member of Rafinesque's subgenus Menandra, it can be assumed that its outer sepals were noticeably shorter than the inner, à fea- ture which helps to narrow the list of possible species to four. Of the seven species of Lechea known from New Jersey, only L. maritima Legg., L. intermedia Legg. ex Britt., L. leggettii Britt. & Holl. and usually L. racemulosa Michx. have external sepals exceeded by the inner. Lechea maritima may be safely discounted as a suspect as it could never be described as “quite smooth” even upon the most superficial observation and its cauline leaves tend to be whorled and not “scattered.” Rafinesque described the leaves of L. pulchella as “scattered long linear" and its capsules as *obovate." These features make it possible to exclude L. racemulosa from consideration for its principal or lower and median cauline leaves are commonly opposite or even whorled and elliptic to lanceolate or even oblanceo- late and perhaps more significantly it possesses a barrel- shaped or cylindrical capsule. Rafinesque’s subgenus Menandra was described as having seeds 1-3 in number which agrees with the number characteristic of L. leggettü but not with the 4-6 seeds characteristic of L. intermedia. Furthermore L. leggettii is known from the New Jersey pine barrens while L. intermedia apparently is not. Leggett (1878: 247) questioningly suggested that Rafinesque's L. pulchella was the same as the species then called “L. minor Lam.” which is what has recently been called L. leggetti. Leggett (1878: 250) twice encountered in the pine barrens *an abnormally smooth Lechea" which pretty well matched Rafinesque's description, These smooth plants were what is now called L. leggettii. Accompanying the original publica-. 194 Rhodora [Vol. 68 tion of L. leggettii the name var. pulchella was appended with L. pulchella Raf. questioningly listed in synonymy indicating Britton’s earlier similar suspicions. The evidence seems sufficiently convincing that I do not hesitate to take up Rafinesque’s binomial for the species most recently known as L. leggettii, New combinations for the varieties recognized by Hodgdon are not made for I am not convinced that these tendencies represent biological varieties or subspecies. Rafinesque (1836: 91) named three varieties based upon the luxuriousness of the plants and the Shape of the inflorescence. These varieties were var. minor Raf., var. elegans Raf. and var. pyramidalis Raf. 2. Lechea cinerea Raf., New Fl. N. Am. 1: 91. 1836. Leggett (1878: 250) felt that Rafinesque's description was a "pretty correct" one for the species he had previously distributed as L. torreyi. He noted however that there were 3-seeded and 6-seeded forms of the species and hoped that his proposed L. torreyi would prove distinct. Hodgdon (1938: 106-107) discussed Rafinesque's description and concluded that it was apparently based upon two or more species. The original description includes nothing that would suggest to me that Rafinesque had both L. torreyi and L. patula as seemed likely to Hodgdon but the point is academic. Rafinesque parenthetically placed L. thymifolia Michx. in synonymy of his own L. cinerea and again in closing mentioned the possibility that it was “the real thymifolia.” Rafinesque’s binomial hence is illegitimate as nomenclaturally superfluous when published according to the International Code (Article 63). The fact that Michaux’s binomial is actually a synonym of L. minor L. does not alter the fact that Rafinesque first placed Michaux’s name unequivocally in the synonymy of his own species. 3. Lechea ternifolia Raf., New Fl. N. Am. 1: 91. 1836. Rafinesque stated that his L. ternifolia from “Virginia, New Jersey and probably elsewhere” “must be the real 1966] Lechea — Wilbur 195 L. racemulosa” of Michaux. Both Rafinesque and Michaux quote the same Gronovian polynomial. And Rafinesque's description itself (“stem . . . adpressed pubescent, leaves and branches mostly ternate cuneate acute ciliate nearly smooth; racemes paniculate lax nearly naked, peduncles elongate, capsules oblong’’) indicates that Lechea ternifolia Raf. can be placed with confidence in the synonymy of L. racemulosa Michx. Leggett (1878: 247) with some ques- tion and Grosser (1894: 135) unequivocally placed Rafi- nesque’s binomial in the synonymy of Michaux’s species while Hodgdon (1938) did not mention it. 4. Lechea furfuracea Raf., New Fl. N. Am, 1: 92. 1836. Leggett stated (1878: 250) that he had “never seen any- thing to which this description would apply.” No one else has been able to suggest what species Rafinesque had from “Kentucky and Illinois” with “mealy pubescence and globose capsules,” very short external calyx-lobes, and rounded internal and “narrow linear scattered nearly obtuse" leaves. According to the most authoritative listings of Lechea in Illinois (Jones & Fuller, 1955) and in Kentucky (Braun, 1943) there are in fact no species known to be common to both states possessing external calyx-lobes shorter than the inner. Lechea racemulosa Michx. is the only species from Kentucky whose calyx approximates Rafinesque’s descrip- tion and its capsule and foliage would definitely exclude it as a possible synonym of L. furfuracea. The three species from Illinois with appreciably shorter external than inner sepals are L. stricta, Legg., L. intermedia Legg. and L. leg- gettii Britt. & Holl. Rafinesque's description is not detailed enough to permit me to determine which species he had. 5. Lechea laxiflora Raf., New Fl. N. Am. 1: 92. 1856. Rafinesque's description seems certainly to apply to the species long known as L. leggettii. Leggett (1878: 247) thought it probably was “L. minor Lam." which was the name by which L. leggettii was known in the later nine- 196 Rhodora [Vol. 68 teenth century. Rafinesque found this species in “the New Jersey Pine Woods with L. pulchella” to which he felt it closely related. The more significant points in Rafinesque's description are as follows: “smooth ... branches lax, leaves scattered linear cuneate acuminate ciliate; racemes scattered lax, flowers remote naked, pedicels elongate, calix and capsules ovate.” I recommend (see No. 1 above) that L. pulchella be adopted as the earliest name for this species with L. laxiflora placed in its synonymy. Rafinesque also distinguished a var. brevifolia with reddish rather than greenish flowers and shorter, more cuneate, less ciliate leaves. Rafinesque indicated that the principal difference between this species and his L. pulchella was chiefly in the leaves which in L. pulchella were “scattered long linear acute” and by inference eciliate in contrast to the acuminate, ciliate leaves of L. laxiflora. 6. Lechea tenuifolia Michx., Fl. Bor.-Am. 1: 77. 1803. Rafinesque’s account is largely a translation of Michaux’s original description. It is placed by Rafinesque with those species with short external sepals while the species to which we apply the name has external calyx lobes equaling or more commonly longer than the inner. Rafinesque cited no locali- ties other than the Santee River which had been cited in Michaeux’s original account so it is not certain whether Rafinesque had actually seen specimens of this species in spite of his statement that he had “specimens of all of the described species.” Perhaps Rafinesque placed it with the species of Lechea with short calyx-lobes since Elliott (Sk. Bot. S. C. & Ga. 1: 185. 1816.) implied that this species had no external calyx and this statement was included by Rafinesque. Elliott in turn had been misled by Walter (FI. Car. 83. 1788) who reported there was no outer calyx in his L. juncifolia, a species which has never been satisfacto- rily identified. 6[a]. Lechea verna Raf., New Fl. N. Am. 1: 92. 1836. This binomial was published as a substitute name (*6. L. 1966] Lechea — Wilbur 197 tenuifolia Mx. Elliot or L. verna Raf, L. juncifolia Walter.” ) and is therefore illegitimate. 7. Lechea recurvata Raf. New Fl. N. Am. 1: 93. 1856. This species was placed by Rafinesque in his subgenus Menandra possessing short external calyx-lobes. It was said to possess globose capsules and to occur "in Carolina and Virginia." I believe that this is the species referred to in the most recent monographs of Britton (1894) and Hodgdon (1938) as L. leggettii. If my interpretations of Rafinesque's species are correct, Lechea pulchella, L. laxiflora and L. recurvata are all prior names for the species now called L. leggettii. There are only two Lecheas common to both Virginia and “Carolina” possessing external calyx-lobes shorter than the inner; these two are L. racemulosa and L. leggettii. The described globose capsules of L. recurvata would seem to exclude L. racemulosa as a possibility al- though, judging by the following features mentioned *branches opposite and ternate, leaves broad oblong acute at both ends ciliate," it might seem a better match for Rafinesque's description than L. leggettii. 8. Lechea mucronata Raf., Précis des Découvertes Somiol. 37. 1814, To my knowledge only House (New York St. Mus. Bull. 254: 497. 1924) has taken up this name although Robinson (Syn. Fl. N. Am. 1: 192. 1895) and Grosser (1903: 135) both placed it unquestioningly in the synonymy of L. major in the sense of Michaux but not of Linnaeus. Lechea major sensu Michaux, Robinson and Grosser is synonymous with L. villosa Ell. Hodgdon, the most recent monographer of the genus, concluded that there was not enough evidence presented in Rafinesque’s original description to warrant the displacement of Elliott’s L. villosa, a “well-characterized and clearly typified name.” Rafinesque's original description is presented below in full so that others may form their own conclusions independent of my own interpretation. I am 198 Rhodora [Vol. 68 indebted to Mrs. Lazella Schwarten, Librarian of the Har- vard Herbaria, for making a copy of it available. 120. Lechea mucronata. Poilue, tige droite et simple, feuilles oblongues-cunéiformes mucronées, fleurs en grappe composée, bractéolées, bractées oblon- gues aigues. Dans les bois du N. Jersey. Hodgdon stated (1938: 56) that the "only possible diag- nostic features of Rafinesque's description are “feuilles oblongues-cunéiformes mucronées, fleur en grappe com- posée." He concluded that the statement concerning the arrangement of tne flowers “might apply to most members of the genus" and that the described leaf shape would apply "equally well to L. minor as to L. villosa." The description's most definitive word in my opinion is “poilue,” and in addi- tion the habitat and geographical location certainly con- siderably narrow the number of possibilities. "Poilue" meaning “shaggy, hairy or pilose” best describes the species recently called L. villosa. The features men- tioned in the brief description single out the species long incorrectly known as L. major and more recently known as L. villosa. There are seven species of Lechea known from New Jersey: L. minor, L. tenuifolia, L. racemulosa, L. mari- tima, L. leggettii, L. intermedia and L. villosa. Of these seven the last mentioned alone seems indicated by Rafines- que’s description of 1814. Lechea maritima, L. leggettii and L. intermedia all possess external sepals conspicuously shorter than the internal which would exclude them from consideration. Of the seven species of Lechea mentioned above found in New Jersey the term "poilue" would seem to me to exclude all but L. villosa and perhaps L. maritima. The description of the leaves, although not definitive, would seem to exclude L. maritima but include L. villosa whose leaves are often mucronate. It is true that the leaves of L. minor sometimes would fit Rafinesque’s description as indicated by Hodgdon but certainly “poilue” would scarcely be used to describe the subappressed pubescence of that species. Rafinesque (1836: 93) more fully described his L. mucro- 1966] Lechea — Wilbur 199 nata indicating it to be the same as the L. villosa of Elliott from whom he had received specimens. Included are state- ments puzzling even to a casual student of the group (e.g. "capsules ovate villose") but sufficient, I believe, to convince one that Rafinesque was indeed describing, although perhaps poorly, the species called L. villosa by Elliott. In passing it may be noted that the capsules of all species of Lechea are glabrous. There actually seems to be nothing other than prejudice why we should not take up Rafinesque's name for this species since it has two years priority over Elliott’s L. villosa. Rafinesque (1836: 93) briefly described three varie- ties all of which came from the extensive area of Hodgdon's var. typica. The three varieties were var. simplex, var. ramosa and var. sessiliflora. New combinations have not been made for Hodgdon's var. macrotheca ranging from central Nebraska south into Texas or his var. schaffneri from northeastern Mexico since I have not seen enough material to form an opinion as to the merits of these pro- posed taxa. 9. Lechea heterophylla Raf., New Fl. N. Am. 1: 93. 1836. Rafinesque described this species from Kentucky and Illinois as having an appressed pubescent stem and acute, carinate inner sepals about equaling the external. Neither Leggett (1878: 248) nor Grosser (1903: 140) would hazard a guess as to what it might be. The three Species common to Kentucky and Illinois possessing external sepals at least occasionally equaling the inner are Lechea villosa Ell., L. tenuifolia Michx. and L. minor L. The shaggy pubescence of the stem of L. villosa would exclude it from further consideration. The leaves were described as “commonly ternate, lower obovate, subsessile, upper cuneate and linear petiolate acute smooth.” I þelieve this a better approxima- tion of the greater variation shown between the lower and upper leaves of L. minor than the comparatively uniform condition exhibited by L. tenuifolia. The acute inner sepals on the other hand are a better fit for the “subacute” condi- 200 Rhodora [Vol. 68 tion of L. tenuifolia than the “obtuse” inner sepals of L. minor. I suspect Rafinesque was describing L. minor since the conspicuous difference in the foliage was less subject to error by Rafinesque than the shape of the apex of the inner sepals. Rafinesque placed “L. minor Smith” in the Synonymy of his L. heterophylla but his description would exclude the villous species that was apparently considered L. minor by J. E. Smith. Britton (1895: 248) has demon- strated that Smith’s L. minor was the species which Elliott called L. villosa. Rafinesque published the following three varieties of this species: var. major Raf., op. cit. 94. 1836, var. minor Raf., l.c. and var. parviflora Raf., Lc. and only the last of these was even briefly described. 10. Lechea glomerata Raf., New Fl. N. Am. 1: 94. 1836. Rafinesque's description of this species from the " Apla- chian mts." strongly points to L. mucronata Raf. (= L. vil- losa) except for his characterization of it as “smooth.” The carinate sepals, short pedicellate, glomerate flowers, linear external sepals about equaling the internal, the trig- onous calyx surrounding the subglobose capsule and the mucronate, petiolate leaves are all indicative of L. villosa but that species is apparently always densely villous to spreading pilose. Leggett (1878), Grosser (1903) and Hodgdon (1938) have all found it impossible to identify Rafinesque’s description and I claim no greater success. ll. Lechea corymbosa Raf., New Fl. N. Am. 1: 94. 1836. The description of this species from the “Alleghany” Mountains, belonging to Rafinesque's subgenus Lechea, a group characterized by the external and internal sepals equaling one another, is most suggestive of L. villosa. EL., a later synonym of L. mucronata Raf. (see No. 8 above). The original description of L. corybosa (“stem . . . pilose . , leaves petiolate broad oblong nearly obtuse, . . flowers corymbose, pedicels equal to flowers, sepals round concave, ext. linear, capsules globose .. . 6 to 12 inches high 1966] Lechea — Wilbur 201 ... capsules large with 3 to 6 seeds... ") is not especially definitive but I believe it is most suggestive of L. mucronata of the species occurring in the Alleghenies. 12. Lechea surculosa Raf., New Fl. N. Am. 1: 94. 1856. Leggett (1878: 248) questioningly suggested that this description might be Lechea minor [in the sense of] Lam. which is the species most recently called L. leggettii Britt. & Hollick. This seems an unlikely identification since L. leggettii is characterized by its external sepals being pro- nouncedly shorter than the inner while L. surculosa, as a member of Rafinesque's subgenus Lechea, would be expected to have the inner and outer sepals of approximately equal length. It is certainly true that one can not place the species of Lechea into three groups depending on whether the ex- ternal sepals are shorter, equaling, or longer than the in- ternal since there is too much variation in the relative length of the sepals in several species but the external sepals of L. leggettii are always clearly appreciably shorter than the inner. Rafinesque's species was described as possessing “acute” inner sepals while those of L. leggettii are clearly obtuse. There are other discrepancies, but the two men- tioned are sufficient to discount completely Leggett's sug- gestion. Rafinesque knew this plant from *Pennsylvania on dry hills" and indicated it was the L. minor of some authors. I believe the description of L. surculosa best matches the characteristics of L. racemulosa, a species whose external sepals range from obviously shorter than the inner sepals to equaling them. The inner sepals of L. racemulosa, accord- ing to Hodgdon's description, vary from “subacute to obtuse" while Rafinesque described those of L. surculosa as “acute.” The capsule of L. racemulosa was described in Hodgdon's monograph as “slenderly ellipsoid to slenderly ovoid" while Rafinesque said those of L. surculosa were “ovate.” Rafinesque described the sterile stems of L. surcu- losa as “pilose with leaves ternate ovatoblong acute ciliate" 202 Rhodora [Vol. 68 which is a reasonably close approximation of Hodgdon’s description of the basal shoot of L. racemulosa: ‘basal leaves frequently verticillate, . . . oblong-ovate to elliptic- lanceolate, acute to slightly mucronulate . . . prominately Spreading-pilose to villous on the margins." The other species of Lechea from Pennsylvania with external lobes approximately equaling the inner at least occasionally (L. villosa, L. tenuifolia and L. minor) have characters which compare unfavorably with those delineated by Rafinesque. The “smooth” branches and “linear” leaves of Rafinesque’s L. surculosa are sufficient to discount it as a possible syno- nym of L. villosa (= L. mucronata). The basal leaves of L. tenuifolia were described by Hodgdon (1938: 87) as “narrowly lanceolate or oblanceolate to linear” which cer- tainly implies a much narrower leaf than does Rafinesque’s description and the cauline leaves of L. tenuifolia are described by Hodgdon as “scattered” while Rafinesque described those of L. surculosa as "opposite and alternate." I believe then that L. surculosa Raf. is an ecologically modi- fied form of L. racemulosa Michx. and that it should be placed in its synonymy. This binomial first appeared three years earlier in Rafi- nesque's Herb. Raf. 68. 1833 but without description. 18. Lechea revoluta Raf., New Fl. N. Am. 1: 94. 1836. Rafinesque stated that this species is the Lechea “minor of Lin. not of Smith nor other Authors” and that the ‘‘de- scription is taken from Linnaeus altogether, and agrees very well with some specimens [that Rafinesque had] from the Alleghenies of Pennsylvania." Nomenclaturally the bi- nomial is a synonym of Lechea minor L. Rafinesque pro- vided a new name since J. E. Smith had indicated that Linnaeus had confounded several species within his L. minor, 14. Lechea virgata Raf., New Fl. N. Am. 1: 95. 1836. This Alleghenian species was described as possessing’ 1966] Lechea — Wilbur 203 pubescent stems, opposite, whorled or even scattered, petio- late, linear, cuneate, ciliate, acuminate leaves with acute, carinate inner sepals, pedicels equaling the flower-length and ovate capsules. As a member of subgenus Lechea, it can be safely assumed that its external sepals equaled the inner. There are four species within the area of the Alle- gheny Mountains whose external sepals may commonly or occasionally equal the internal ones. These four species are Lechea minor L., L. racemulosa Michx., L. tenuifolia Michx. and what has been most recently referred as L. villosa Ell. The description is an unusually full one for Rafinesque and the above mentioned characters seems best to describe L. tenuifolia Michx. of the four species listed above from the Alleghenies whose external sepals commonly or occasionally equal the inner. The linear leaves, which are the only type mentioned for L. virgata, are sufficient to exclude L. villosa and L. minor from further consideration. The inner sepals of L. racemulosa are non-carinate while those of L. virgata were described as carinate. Two varieties were appended to this species with brief description: var. breviflora and var. bracteata. 15. Lechea floridana Raf., New Fl. N. Am. 1: 95. 1836. There are eight species of Lechea known from Florida (Hodgdon, 1938; Wilbur & Daoud, 1961). Five of these (L. deckertii Small, L. leggettii Britt. & Holl., a later synonym of L. pulchella Raf. as is pointed out in no. 1 above, L. tor- reyi Legg. ex Britt., L. divaricata Shuttlew. ex Britt. and L. cernua Small) possess external sepals conspicuously shorter than the internal. The three others (L. villosa Ell., a later synonym of L. mucronata Raf. as is discussed in no. 8 above, L. minor L. and L. patula Legg.) all possess external sepals conspicuously shorter than the internal, as was described by Rafinesque for his L. floridana. None of these three species is “smooth” as mentioned by Rafinesque but certainly the copious, spreading villosity of L. mucro- nata, (— L. villosa) together with its oblanceolate leaves ex- cludes it from consideration. The appressed pubescence of 204 Rhodora [Vol. 68 L. patula and L. minor could easily have been overlooked, especially if the specimens were aged, by an observer as hasty and superficial as Rafinesque often gives evidence as having been. The leaves of L. floridana were said to be "scattered, minute linear" which is more suggestive of L. patula than of L. minor. The external calyx of L. floridana was implied to equal the inner which is sometimes true of both L. patula and L. minor. The external sepals of L. patula most frequently are a little shorter than the inner but range up to slightly longer while those of L. minor rarely are as short as the inner and most frequently are conspicuously longer. The capsules of L. floridana were said to be ovate which is not a very apt description for the capsule of either species but, interpreted as “ovoid,” not impossibly bad for either. I suspect this to be a form of L. patula Leggett but due to the discrepancies mentioned would not feel its identity completely certain. 16. Lechea secundiflora Raf., New Fl. N. Am. 1: 95. 1836. This species came from “the glades of West Kentucky" making it impossible that its binomial could ever displace that of any of the four species definitely known from that state since each has an older name. The known Species are L. villosa Ell., a later synonym of L. mucronata Raf., L. minor L., L. tenuifolia Michx. and L. racemulosa Michx. Rafinesque’s description fits none well and there are men- tioned features that would exclude each of the geographical possibilities. Grosser (1903: 135) placed Rafinesque’s name in the synonymy of L. minor but his description of “leaves scattered, lax narrow linear, . . . flowers remote secund" would, as pointed out by Hodgdon (1938: 60), make that an unlikely choice. Hodgdon (1938: 88) placed Rafinesque's binomial in the synonymy of L. tenuifolia Michx. although the description of the capsules as "oblong" and the inner calyx as "lanceolate" causes considerable doubt as does Rafinesque's description of the plant as “smooth.” I am unable to identify the binomial and believe the mentioned discrepancies are of such magnitude as to prevent one from 1966] Lechea — Wilbur 205 assigning it with confidence to the synonymy of any species known from Kentucky. 17. Lechea pauciflora Raf., New Fl. N. Am. 1: 96. 1836. The identity of the plants described by Rafinesque under this binomial from “Near the Sea Shore in Long Island and New Jersey" was questioningly suggested by Leggett (1878: 250) to be L. thymifolia in the sense of Pursh which is L. maritima, Legg., à species whose external sepals are much shorter than the heavily appressed-pilose internal calyx. Rafinesque placed L. pauciflora in his subgenus Eudiexa, characterized by outer sepals exceeding the inner, and in the diagnosis of the species specifically stated “sepals smooth, external a little longer.” Grosser (1903: 135) assigned Rafinesque’s binomial to the synonomy of L. minor L., a disposition to which Hodgdon (1938: 60) objected, feeling the described 3-inch specimens to be an “ecological variant of some other species” whose identity was undecipherable because of the incompleteness of the original publication. The only two species apparently known from both Long Island and New Jersey with outer sepals longer than the inner and “adpressed pubescent” stems are L. tenuifolia Michx. and L. minor L. Rafinesque described the capsules of his L. pauciflora as “globose” which among other features mentioned in the description would eliminate L. minor. The leaves of Rafinesque’s species were described as “rather obtuse” which together with the “smooth” calyx makes it difficult to place this name in the synonymy of L. tenuifolia with which the description agrees in most other respects. Therefore I am unable to suggest the identity of this bi- nomial. 18. Lechea brevifolia Raf., New Fl. N. Am. 1: 96. 1836. Leggett (1879: 249) was "quite at a loss" as to what species Rafinesque's description might apply while Grosser (1903: 135) placed the binomial in the synonymy of L. minor L. without discussion, Hodgdon (1938: 60), while 206 Rhodora [Vol. 68 admitting that much of Rafinesque’s description applied to L. minor, felt that the diagnosis of “flowers secund, pedicels equal” indicated confusion on the author’s part. Hodgdon’s objection on these grounds are difficult to understand for the flowers and fruit are sometimes secund as his own description of L. minor L. admits (p. 59 “fruit mostly clustered, at times scattered or even secund") and the 1-2 mm. long pedicels are often equal to one another or to the flowers. The species came from the “Mts. Apalaches" (Ap- palachian Mts.) and I believe it most probable that this binomial belongs in the synonomy of L, minor. 19. Lechea uniflora Raf., New Fl. N. Am. 1: 96. 1836. Leggett (1878: 249) was unable to suggest what species Rafinesque was describing based upon 3-4 inch specimens from the mountains of Maryland possessing few leaves or flowers and longer outer sepals than inner and with a 6- seeded, globose capsule. Grosser (1903: 135) placed the binomial in the synonymy of L. minor L. but Hodgdon (1938: 61) disagreed with this placement concluding that it was “hopeless to try to identify this particular one of Rafinesque’s Lecheas.” I am in total agreement with Hodg- don for I too am unable to suggest the identity of L. uniflora. There are too many features mentioned in its original des- cription which are at complete variance with the character- istics of L. minor for us to accept Grosser’s identification. To my knowledge a “smooth” specimen of L. minor with globose 6-seeded capsules has not been reported. 20. Lechea stellata Raf., New Fl. N. Am. 1: 96. 1836 I am in complete agreement with Leggett’s decision (1878: 251) that this description is "undoubtedly the L. thymifolia" of Michaux which is a synonym of L. minor L. The more diagnostie portions of Rafinesque's description seem to be as follows: “Pilose . . leaves 3-4 nate petiolate elliptic mucronulate, upper leaves alternate lanceolate; racemes foliose, . . . ext. sepals very long, capsules ovate- 1966] Lechea — Wilbur 207 oblong.” Both Grosser (1903: 135) and Hodgdon (1938: 61) considered L. stellata to be synonymous with L. minor, although the latter fails to place it formally in the synonymy of that species. 21. Lechea sessiliflora Raf., New Fl. N. Am. 1: 97. 1836. Leggett (1878: 251) thought that the description ac- companying this binomial might “possibly be L. patula" but no species known to him had sessile mature flowers. In spite of this, Index Kewensis equated Rafinesque's binomial with L. patula Legg. and Grosser (1903: 138) unquestion- ingly placed it in the synonymy of that species. Hodgdon (1938: 63) felt that Rafinesque's description could not be positively identified but that its sessile flowers and 3-9- seeded capsules obviously excluded the pedicellate, 1(2)- seeded L. patula as a possibility. However, since no Lechea is actually sessile, it would perhaps be only reasonable to interpret Rafinesque's description as “subsessile” and con- sider short-pediceled species as possibilities. Also, since no known species of Lechea possesses more than 6 seeds, we had best treat the 3-9-seeded condition attributed by Rafines- que to his subgenus Hudiera as an obvious error. It is well to note that Rafinesque did not mention seed-number in his diagnosis of L. sessiliflora ; seed number was stated only in conjunction with the characterization of the subgenus. There are five species known to be common to both Florida and Alabama: Lechea villosa Ell., L. minor L., L. patula Legg., L. leggettii Britt. and L. torreyi Legg. ex Britt. The two last-mentioned of these can be discounted as possibilities since their external sepals are considerably shorter than the internal and the reverse is clearly described as the state of L. sessiliflora. Lechea villosa can be eliminated from con- sideration since its shaggy-villous stems could never be described as “adpressed pilose” while both L. minor and L. patula both fit that characteristic. Rafinesque's character- ization of the species “branches diffuse virgate, leaves scat- tered ... linear... ext. sepals linear not much longer..." all point most clearly to L. patula rather than to L. minor. 208 Rhodora [Vol. 68 I believe that Leggett's suspicion as to the identity of L. sessiliflora has been fully confirmed. There is hence no reason not to take up L. sessiliflora Raf. which has forty-two years priority over Leggett's L. patula. The remaining names proposed by Rafinesque are listed below: Lechea linifolia Raf., Atl. Jour. 1: 18. 1832. nom nud. Lechea paucifolia Raf., Atl. Jour. 1: 18. 1832. nom. nud. Lechea mexicana Raf., Sylva Tell. 133. 1838. Rafinesque provided this as a substitute name for Heli- anthemum tripetalum of Mexico no doubt because tripetalum was a pointless epithet in a genus characterized by three petals. The name is nomenclaturally superfluous and there- fore illegitimate (Art. 63). DEPARTMENT OF BOTANY, DUKE UNIVERSITY DURHAM, NORTH CAROLINA LITERATURE CITED BRAUN, E. Lucy. 1943. Lechea in An Annotated Catalog of Sperma- tophytes of Kentucky. p. 93. BRITTON, N. L. 1894. A Revision of the Genus Lechea. Bull, Torrey Club 21: 244-253. GROSSER, W. 1903. Lechea in Engler's Das Pflanzenreich 14 (IV, 193) : 133-140. HopGDoN, A. R. 1938. A Taxonomic Study of Lechea. Rhodora 40: 29-69, 87-131. (Reprinted as Contrib. Gray Herb. 121.) JONES, G. N. & G. D. FULLER. 1955. Lechea in Vascular Plants of Illinois. pp. 328-329. LEGGETT, W. H. 1878. Rafinesque's Lechea. Bull, Torrey Club 6: 246-252. RAFINESQUE, C. S. 1836. Monograph of Lechea, New Flora and Bot. of N. Am. pp. 89-98. WILBUR, R. L. & H. S. Daoup, 1961. The Genus Lechea (Cistaceae) in the Southeastern United States. Rhodora 63: 103-118. GENTIANA PUBERULENTA SP. NOV., A KNOWN BUT UNNAMED SPECIES OF THE NORTH AMERICAN PRAIRIES: JAMES S. PRINGLE The species which for the past century has been known as Gentiana puberula is one of the showiest of the North American gentians; Curtis (1959) called it “by all odds the most beautiful member of this famed genus in Wis- consin," and described its flowers as “unrivalled in the clarity and intensity of their deep blue color." It is a characteristic species of mesic and xeric prairie commu- nities from Ohio west to Manitoba and Arkansas, and has been collected as far east as western New York and Mary- land, as far south as Louisiana, and as far west as the Black Hills of South Dakota. Some authors have designated this species the “Prairie Gentian” ; others have used the translation “Downy Gentian”, although the puberulence is so minute that it is not a conspicuous feature to the casual observer. Gentiana puberula Michx. Fl. Bor.-Am. 1:76. 18083 was based on a specimen Michaux had collected at “Fort Chero- quis" (Fort Massac) in present-day Massac County, Illinois, where he had spent part of the autumn of 1795. This speci- men is preserved in the herbarium of the Laboratoire de Phanérogamie, Muséum National d'Histoire Naturelle, Paris, France. A photographic negative of this specimen was made available through the kindness of Prof. A. Aubré- ville and J. Leandri. This negative and enlarged prints have been placed in HAM. Duplicate prints have been deposited Tn addition to those persons whose contributions are noted in the text, I should like to express my appreciation to Dr. A. J. Sharp of the University of Tennessee and Dr. J. M. Gillett and Mr. W. J. Cody of the Plant Research Institute, Canada Department of Agriculture, for their reviews of the manuscript, and to Dr. A. H. Moser of the University of Tennessee, for his checking the Latin description. Contribution No. 5 from the Royal Botanical Gardens, Hamilton, Ontario, Canada. 209 [Vol. 68 Rhodora 210 —— —————^P—————É—————————————" 000 7 7 E '6'0 €» X ‘dA, 10g pojoo[os Ways uo doAO[ "p ‘DIA '&'0 BO X 'spoqe[ yya ‘pys umnrrieq -19u uo uaurreds ejo[duio)) *g ‘DIA Bung Djuojm4oqnd ^r) Jo uauitoods odÁ] 'p pu? g “SDIA ‘6'0 BOX ‘SLOMO[ ST Z 7014 gO €» X ‘Sfeqe, pue ueurroedg '[ D ‘XII Djm4oqnd Dupiuor) Jo sueunoeds ƏdAL 'g pue [ 'S5I4 me) emp, ty) 08 igh prse aluo gong: S ee fe 4 wy ag TID W jose ao Ao peat T Y7 fedt y sez ving ama "^5. 1 — "y megy i npo d + Hiker vpn eye pce wage TUVI NOU LEVEY. 1966] Gentiana puberulenta — Pringle 211 in DAO, GH, ILL, and WIS, and are reproduced here as Figures 1 and 2. The status of this specimen as the type is confirmed by the label in Michaux's handwriting on which both the collection data and the description correspond closely to those in Michaux's (1803) Flora Boreali-Americana. The type specimen of G. puberula Michx. is clearly not a specimen of the species described under that name in recent manuals (e.g. Fernald, 1950, and Gleason & Cronquist, 1963), but is, instead, a specimen of G. saponaria L. which falls well within the range of variation commonly found in populations of that species. Gentiana saponaria is usually described as having glabrous stems, but individuals with puberulent stems are occasionally encountered throughout most of its range. Table 1 lists some of the conspicuous differences between the type specimen of G. puberula Michx. and the species hitherto commonly known as G. puberula, here designated G. puberulenta. The characters selected for use in Table 1 are those readily discernible in the photo- graph and ones which have been found valuable in the dis- tinguishing of Gentiana species. No scientific name appears to be available for the Prairie Gentian. Authors of floras of North America and treatises on the North American gentians published prior to 1848 seem not to have known this species (with the possible exception of Rafinesque, as noted below). Grisebach (1838, 1839, 1845), in his works on the Gentianaceae, tentatively assigned the name G. puberula Michx. to the synonymy of his G. saponaria 3 linearis. (The latter name was based on G. linearis Froel., the name of a distinct species with which Grisebach had confused narrow-leaved specimens of G. saponaria.) In so doing, Grisebach was following Pursh (1814), but he (1838) added that this interpretation might be incorrect, and that “it is desirable that this question should be determined by the inspection of Michaux's her- barium." From labels on mid-nineteenth-century collections of the Prairie Gentian, it appears that the first botanists to collect it called it G. rubricaulis Schwein. (a name properly applied to a more northern species), and that it is the basis 212 Rhodora [Vol. 68 for the inclusion of “Gentiana rubricaulis" (without de- scription) in some plant lists of that period (see Jones and Fuller, 1955). Although many names for gentians were published by Rafinesque, it does not appear that any can be applied to this species. In certain parts the brief descriptions of Gentiana shortiana, G. torreyana, G. rigida, and G. gracilis (all in his Medical Flora, 1828) suggest the possibility that they were based on specimens of this species ; however, each of the descriptions, when considered in its entirety, appears much more probably to have been based on G. saponaria L. In Rafinesque’s (1837) Flora Telluriana these specific epi- thets appear in his genera Pneumonanthe and X olemia; the descriptions of both genera contain material which is in- applicable to the Prairie Gentian, although applicable to G. saponaria. Probably Rafinesque never encountered this species, which was rare in the area of his field work and had been collected by few botanists at the time of his pub- lishing. It is highly unlikely that any of these Rafinesque taxa will ever be typified by a specimen, since very few of the plant specimens collected by Rafinesque are extant. Dr. Ronald L. Stuckey, who has studied the herbarium (now in PH) of C. W. Short, to whom Rafinesque gave some speci- mens, reports that none of the specimens bear any of these names. John Torrey’s Gentiana specimens (now in NY) were annotated by Rafinesque, but I found none of these names among the annotations. Probably the earliest published description of the Prairie Gentian is that by Gray ( 1848) in the first edition of his Manual, in which it was designated G. saponaria B puberula (Michx.) Torr. & Gray. In later editions Gray recognized this taxon at the rank of species, as G. puberula Michx. Gray’s treatment has generally been followed by later authors. Thus, even though a number of descriptions of the Prairie Gentian have been published in the past, it is necessary to describe it here as a new species: 1966] Gentiana puberulenta — Pringle 213 Gentiana puberulenta Pringle, sp. nov. Herba perennis singulis vel pluribus caulibus suberectis longis 1-6 dm puberulentis subtiliter abundanterque. Folia lanceolato-oblonga interdum paene linearia prope basem latissima uninervia marginibus ciliatis saepe revolutis. Folia superiora longa 1.5-7 (-10.5) em lata 4-18 (-30) mm. Flores paene sessiles 2-bracteati aggregati apud sum- mum caulem et interdum de axillibus pedicellati. Tubus calycis longus 7-18 mm. Lobi calycis lineares longi 4-18 (-25) mm lati 1-3 (-4) mm. Corolla atrocyanea (raro violacea vel rubro-violacea) vasi- formis certe se aperiens longa 3-6 cm. Lobi corollae ovati patentes longi 6-14 mm lati 4-10 mm marginibus saepe erosis. Partes liberae appendicium corollae longae 2.5-6 mm bifidae praeterea segmentibus laciniatis. Antherae non cohaerentes, Semina alata. Type: Chase 1276, 8 mi. N.E. of Princeville, Peoria County, Illinois, 7 Oct. 1906 (ILL; isotypes DAO, ILL, KSC, MIN, NY, US). A more extensive description of this species, along with a discussion of its distribution and probable relationships to other species, will be included in a forthcoming mono- graph by the author on Gentiana, section Pneumonanthae, in eastern North America. No infraspecific taxa have been recognized within this species. The two named hybrids involving it are both typi- fied by specimens, so the applications of the names G. X bil- lingtonii Farw. (pro sp.) (Pap. Mich. Acad. 1:96. 1923) and G. X curtisii Pringle (Trans. Wis. Acad. 53: 277. 1964) remain unchanged. Their formulae, however, should be emended to G. puberulenta X G. andrewsii and G. puberul- enta X G. alba, respectively. LITERATURE CITED Curtis, J. T. 1959. The Vegetation of Wisconsin: An Ordination of Plant Communities. The Univ. of Wisconsin Press, Madison. FERNALD, M. L. 1950. Gray’s Manual of Botany. ed 8. American Book Co., New York. GLEASON, H. A., and A. CRONQUIST. 1963. Manual of Vascular Plants of Northeastern United States and Adjacent Canada. IDE Van Nostrand Co., Princeton. Gray, A. 1848. A Manual of the Botany of the Northern United States . . . James Monroe & Co., Boston. Rhodora [Vol. 68 214 GRISEBACH, A. H. R. 1838. Gentianeae Juss. In Hooker, W. J., Flora Boreali-Americana 2:54-71. Henry G. Bohn, London. [Re- printed 1960 as Hist. Nat. Class. 5. H. R. Engelmann (J. Cramer), Weinheim, and Wheldon & Wesley, Codicote.] 1839. Genera et Species Gentianearum. J. G. Cotta, Stuttgart and Tubingen. 1845. Gentianaceae. In Candolle, A. P. de, and A.L.P.P. de Candolle, Prodromus Systematis Naturalis Regni Vegetabilis . . . 9:38-141. Victor Masson, Paris. JONES, G. N., and H. J. FULLER. 1955. Vascular Plants of Illinois. The Univ. of Illinois Press, Urbana, and The Illinois State Mu- seum, Springfield, Sci. Ser. vol. 6. MICHAUX, A. (L. C. M. Richard, ed.)1803. Flora Boreali-Americana -.. Fratres Levrault, Paris PursH, F. T. 1814. Cochrane, & Co., London. RAFINESQUE, C. S. 1828-1830. Alexander, Philadelphia. 1836-1837. Flora Flora Americae Septentrionalis . . . Medical Flora and Strasburg. White, Atkinson & Telluriana, Publ. by the author, Philadelphia. (Photolith. 1946. Arnold Arboretum, Cambridge.) ROYAL BOTANICAL GARDENS, HAMILTON, ONTARIO, CANADA TABLE 1. THE TYPE SPECIMEN CONTRASTED WITH G., Type specimen of G. puberula Leaves few and widely spaced. Leaves widest at or above the middle. Calyx lobes oblong, widest above the middle. Corolla lobes comprising about 10% of the total corolla length. Corolla lobes widest at the base, less than twice as long as the appendages. OF GENTIANA PUBERULA MICHX, PUBERULENTA PRINGLE G. puberulenta Leaves many and crowded. Leaves widest near the base. Calyx lobes linear. Corolla lobes comprising 20-25% of the total corolla length. Corolla lobes ovate, more than twice as long as the appen- dages. NEW CHROMOSOME NUMBERS IN ZINNIA AND SANVITALIA ANDREW M. TORRES! In recent systematic papers concerning Zinnia ( Torres 1962, 1963) and Sanvitalia (Torres 1964) the author was unable to provide chromosome counts for 1 of the 17 species of Zinnia and for 3 of the 7 species of Sanvitalia. The Zinnia species, Z. purpusii, had been known only from the state of Chiapas in Mexico. Because the collection site, *Hacienda Monserrate", was not, and still is not, located and because the species was known only from the type col- lection, it was surprisingly pleasant to find a population on the hillsides of gravelly soil along Mexican Highway 190, 1.6 miles east of the Oaxaca-Chiapas border (1739) and again 12 miles east of the same border (1743). From the first collection the chromosome number of n = 12 was determined. Meiosis was regular. Its chromosome number, although predictable, was of interest because its nearest phenetic ally, Z. maritima, is also n = 12. Yet several other near allies are n = 11. The count is significant also since chromosome numbers of all species are now known. A few more collections of Sanvitalia tenuis had been made than of Z. purpusii yet its habitat too was relatively in- accessible. It apparently grows only on the eastern side of the Sierra Madre Occidental in western Chihuahua. Some 19 miles west of Mifiaca (near Ciudad Guerrero and La Junta) in Chihuahua near the edge of oak-pine hills, I chanced upon a population of some 20-25 young plants just beginning to flower. Interestingly, all my collections were from the base of young oak trees on gentle slopes in gravelly soil. During the rainy season when S. tenuis is in flower the area is accessible by logging roads and only in high- center vehicles. An anomalous chromosome number pattern had already been established for Sanvitalia because the species are n — 8 or 16, except for S. abertii which is TField work was supported by NSF grant GB-3842. 215 216 Rhodora [Vol. 68 ^ = 11. It therefore became of especial interest to obtain the number of S. tenuis to see if it formed a link in a pos- sible aneuploid series or if it was also n = 8, 16 or 11. Its chromosome number is n — 9; meiosis appeared regular. Perhaps one of the two species as yet uncounted, Mexican S. fruticosa and South American S. versicolor, will fill the n = 10 gap. Voucher specimens of both species will be housed at KANU with duplicates at Msc. DEPARTMENT OF BOTANY, UNIVERSITY OF KANSAS, LAWRENCE, KANSAS LITERATURE CITED TORRES, A. M. 1962. Cytotaxonomy of cespitose zinnias. Amer. Jour. Bot. 49: 1033-1037. 1968. Taxonomy of Zinnia. Brittonia. 15: 1-25. 1964. Revision of Sanvitalia (COMPOSITAE-HELIAN- THEAE). Brittonia. 16: 417-433. TWO NEW NAIADS FROM ILLINOIS AND DISTRIBUTIONAL RECORDS OF THE NAIADACEAE: PAUL L. FORE? AND ROBERT H. MOHLENBROCK? In the process of identifying Illinois species of Naias and in comparing fresh material with herbarium specimens, it was discovered that considerable misidentifications of speci- mens of this genus existed in Illinois. In order to present an accurate account of Naias in Ilinois, and to up-date ‘An additional species Najas marina L. is reported from Illinois by Winterringer in this number of Rhodora (p. 221). *Cooperative Fisheries Research Laboratory, Department of Zoology, Southern Illinois University, Carbondale. "Department of Botany, Southern Illinois University, Carbondale. 19661 Illinois Naiadaceae — Fore and Mohlenbrock 217 distributional data of the species, the first author undertook this study of Naiadaceae of Illinois. GEOGRAPHICAL CONSIDERATIONS Clausen (1936), summarizing the geographical distribu- tions of Naias, listed five species from the northern United States, two of which were known from Illinois. Likewise, Winterringer and Evers (1960) and Jones (1963) recorded only the occurrence of the same two species, N. flexilis (Willd.) Rostk. & Schmidt and N. guadalupensis (Spreng.) Magnus. The results of this investigation demonstrated that four species of Naias inhabit Illinois waters. Lawrence (1955) stated that a total of seven species were native to the continental United States out of a world-wide total of approximately forty species. Two Illinois species have a general distribution in eastern North America from Newfoundland to Minnesota, south to Missouri and Virginia. Included here are Navas flexilis and N. gracillima (A. Br.) Magnus. The remaining two species exhibit quite dissimilar ranges. Naias guadalupensis is known from Massachusetts to South Dakota, south to Texas and Florida, and Tropical America. Naias minor All., an introduced Eurasian species, is locally established in New York and West Virginia, according to Fernald (1950) ; this species is also reported by Muenscher (1944) to occur in Alabama and Tennessee. In addition to the above localities, Naias minor is now recorded from Illinois. In Illinois, Naias flexilis is widely distributed, although not abundant, throughout the state; N. guadalupensis is found mainly in the western and southern parts of the state. Naias gracillima, mostly distributed in southern Illinois, was recently rediscovered in three counties. The first recent specimen was collected in Williamson County in 1952 (although heretofore unreported), while the other two were found during 1961. Fernald (1945) reported that a speci- men of Naias gracillima from Wabash County was listed in the Patterson Catalogue in 1876. Apparently, more than 218 Rhodora [Vol. 68 76 years have elapsed since this species was originally collected in Illinois. Possibly an adventive, Naias minor was discovered in June, 1963, in shallow alkaline waters of Lake Murphysboro in Jackson County. This naiad was growing in close asso- ciation with Naias flexilis, N. guadalupensis, Chara spp., and Jussiaea diffusa in an aquatic community commonly dominated by Potamogeton americanus. Naias minor, which is known only from this station, represents a new record for Illinois. Vascular plants such as Naias species are naturally oc- curring aquatics that were considered originally to have a valuable role in inland waters. Today, it is recognized that profuse submerged vegetation in ponds and lakes creates serious nuisance problems, particularly around aquatic recreational facilities, fish hatcheries, and drainage canals. Also, abundant clumps of Naias offer excessive escape cover to small fishes which may lead to over-population and stunting. For these reasons, dense stands of Naias are now often controlled chemically (Benson and Conner, 1956). TAXONOMY OF NAIAS MINOR Because of the lack of a detailed description of Naias minor in current manuals in the United States, an account of this species is presented here. Naias minor All. Fl. Pedem. 2:221. 1875. Slender to terminally compact annuals; stems slender, but slightly stouter than in Naias gracillima, moderately branched, the terminal nodes crowded, with lime-green internodes; leaves filiform, colored the same as the internodes, 2.0-3.5 em long, 0.2-0.3 mm wide, the distal end spinescent, recurved, the sheath abruptly distended, truncate, finely toothed, each leaf-margin with 8-16 barely macroscopic spinules more than 0.5 mm long; style and 2 stigmas 1.0-1.4 mm long; epicarp purplish; achene falcate, slender, oblique, 2.6-3.6 mm long, !4 as wide, marked with 10-18 regular vertical ribs of broad, rectangular reticu- lations. Although both Naias minor and N. gracillima typically have long filiform leaves with suddenly abrupt bases and spiny margins, N. minor is unique with respect to the 1966] Illinois Naiadaceae — Fore and Mohlenbrock 219 regular, rung-like pattern on the achenes. These markings usually consist of 12 parallel longitudinal rows of wider than high, rectangular areolae. In the field, N. minor is quite distinctive due to its branched, erect, bushy appearance, long narrow leaves, and pale, lime-green coloration which easily distinguishes this plant from its darker green, wider-leaved associates. Al- though growing to heights of 1-2 feet, N. minor is neverthe- less a rather fragile plant with stiffish leaves and stems so brittle that they fragment readily at the nodes. DISTRIBUTIONAL RECORDS OF ILLINOIS NAIADACEAE County distributional data for the four species of Naias in Illinois are given below. Naias flexilis (Willd.) Rostk. & Schmidt. Specimens examined. — COLES: Fox Ridge State Park, Anderson 11386 (ISM). CooK: Cedar Lake, Roush 814 (MO). JACKSON: Lake Murphysboro, Weber s.n. (SIU). JERSEY: Lake of Grafton, Ewers 38870 (ILLS). LAKE: Lake Marie, Winterringer 2671 (ISM). LEE: Amboy, Long 624 (ILL). MCHENRY: Crystal Lake, Pepoon 11425 (ILLS). MASON: Sand Lake near Havana, Richardsos s.m. (ILLS). MENARD: Without locality, Hall 2 (ism). PERRY: 4 miles W. of Du Quoin, Bell s.n. (SIU). RANDOLPH: Sinkhole N. of Chester, Evers 32786 (ILLS). ST. CLAIR: East St. Louis, Eggert 4258 (ILL). TAZE- WELL: Spring Lake, Chase 15843 (ILL). WABASH: Mt. Carmel, Schneck 91 (ILL). WILLIAMSON: Strip mine area near Cambria, Bell 4581 (SIU). Naias guadalupensis (Spreng.) Magnus. Specimens examined. — JACKSON: Along shore in shallow water of Lake Murphysboro, Fore & Stookey 105 (SIU). JEFFERSON: Mt. Ver- non Reservoir, Fore & Stookey 301 (SIU). LAKE: Honey Lake, 4 miles N. of Barrington, Steyermark 63897 (ILL). MACOUPIN: Riniker Lake near Carlinville, Adams 11429 (ISM). PEORIA: Douglas Lake, with- out collector s.n. (ILLS). PERRY: Du Quoin City Reservoir, along the shore, Fore & Stookey 304 (SIU). POPE: Lake Glendale, 2 miles N. of Dixon Springs State Park, Fore & Andrews 302 (SIU), RANDOLPH: Coulterville Reservoir, Fore & Stookey 303 (SIU). ST. CLAIR: Farm N. of Little Rolling Mill, Eggert s.n. (MO). TAZEWELL: Spring Lake, Evers 39969 (ILLS). WILLIAMSON: Shallows of Crab Orchard Lake, Voight 441 (SIU). Navas gracillima (A. Br.) Magnus. Specimens examined.— FORD: Hoppler Pond, Hilterbran s.n. 220 Rhodora [Vol. 68 (ILLS). JACKSON: Fountain Bluff Fish Farm near Gorham, Lewis s.n. (SIU). WILLIAMSON: Crab Orchard Lake, Evers 34356 (ILLS). Naias minor All. Specimens examined. — JACKSON. Shallow water of Lake Murphys- boro, Fore & Stookey 104 (stv), 108 (stv). The authors express their appreciation to the curators of the following herbaria for the congenial cooperation and assistance shown them during this study: University of Illinois (ILL), Illinois Natural History Survey (ILLS), Illinois State Musem (ISM), Missouri Botanical Garden (MO), and Southern Illinois University (SIU). LITERATURE CITED BENSON, N. G. and J. T. CoNNER. 1956. Use of CMU to control Naias. Prog. Fish-Cult. 18:(2) : 78-80. CLAUSEN, R. T. 1936. Studies in the genus Najas in the northern United States. Rhodora 38: 333-345. FERNALD, M. L. 1945. An incomplete flora of Illinois. Rhodora 47: 204-219. 1950. Gray's Manual of Botany. 8th ed. The Ameri- can Book Co., New York. 1632 pp. JONES, G. N. 1963. Flora of Illinois. 3rd ed. Amer. Midl. Nat. Monogr. 7: 1-401. LAWRENCE, G. H. M. 1955. Taxonomy of vascular plants. Mac- millan Co., New York. 823 pp. MUENSCHER, W. C. 1944. Aquatic plants of the United States. Comstock Publishing Co., Ithaca, N. Y. 374 pp. WINTERRINGER, G. S. and R. A. Evers. 1960. New records for Illinois vascular plants. Ill. State Mus., Sci. Pap. Ser. 11: 1-135. AQUATIC VASCULAR PLANTS NEW FOR ILLINOIS A project involving collection and identification of sub- mersed and floating aquatic plants was undertaken jointly by the Illinois State Museum and the Illinois Department of Conservation in 1964-65. Conservation biologists of the Division of Fisheries collected aquatic plants from hundreds of lakes, ponds and rivers throughout the state. In October 1965, at the close of the project, 1175 collections had been made. A uniform data sheet was included with each col- lection to insure accurate locality records for all specimens. The plants, sent by mail in heavy envelopes to the Illinois State Museum for study and processing, were kept moist in thin plastic bags. Notes on the data sheet enclosed in each bag, or with each collection, were made in pencil to prevent smudges. These data sheets were dried, numbered and filed for reference. Alvin Lopinot of the Department of Conservation and the writer are preparing a field manual to assist biologists and students in the identification of submersed and floating aquatic plants of Illinois. This manual will be particularly intended to help conservation biologists in the study and control of aquatic plants wherever their abundance interferes with fishing. One result of the project described above was the identi- fication of two submersed aquatic vascular plants new for Illinois. These plants are Najas marina L. and Ruppia maritima L. Specimens have been filed in the Illinois State Museum Herbarium. Collection data are as follows: LAKE CO., Najas marina L., 2Y2 mi. NE. of the village of Grayslake. Submersed in 3 feet of water in Druce Lake. The plants were abundant and established over a large area. The clear water of this glacial lake had a pH of 8.4. Oct. 5, 1964, G. S. Winterringer 22499 and 22503. In Wooster Lake, 1 mi. SW. of the village of Long Lake, Aug. 4, 1965, Ben L. Dolbeare 311 and 312. Originally collected in the Druce Lake locality, but not identified, by Gregg Tichacek, Sept. 9, 1964. VERMILION CO., Ruppia maritima L., 3 mi. NE. of Oakwood, Kickapoo State Park, in a strip mine pond called “Inland Sea”. Abundant along the shallow margin of the pond and apparently established in water with a pH of 8.5. Sept. 18, 1964, G. S. Winterringer 22449 and 22451. Originally col- lected in the same locality, but not identified, by Rudy Steinauer, 221 222 Rhodora [Vol. 68 Sept. 9, 1964. LAKE CO., R, maritima L., growing in Lake Zurich near Lake Zurich village. Sept. 28, 1965, Alvin Lopinot. A third aquatic plant found recently in Illinois, Najas minor All., was reported in 1964 by Stookey, Fore and Mohlenbrock in Castanea: 29, pages 151 and 153. This naiad was first brought to the writer's attention in October 1963 by Alvin Lopinot who had collected it in Wayne and Greene counties, The museum herbarium now has speci- mens of N. minor from 31 additional counties in the south- ern three quarters of Illinois, extending southward from Henderson and Iroquois counties. Procedure of handling specimens in the laboratory may be of interest. Each colleetion, with its numbered data sheet, was placed in plastic ice cream containers or floated in the sink during examination. Identification was made easier by working with fresh and mature specimens. In preparing material for mounting we used a technique of floating delicate species, e.g., Zannichellia palustris L., on letter size bond paper. The film of water on the paper and specimen permitted final arrangements of the material by moving it with a small brush. The sheet of paper with the specimen was lifted carefully to remove most of the water, or the sink was slowly drained before lifting the sheet. The bond paper bearing its specimen was numbered from the corresponding data sheet, placed on blotting paper or drying sheets and finally inserted in a folder of newsprint. Thus, à press of wet specimens was ready for the dryer. When specimens on bond paper were thoroughly dry they were pasted on standard herbarium sheets to avoid additional handling of the dried specimen. Drops of Duco cement or very narrow strips of gummed cloth were used to secure loose parts wherever needed. The cement also permitted fastening loose achenes of Potamogeton and fruiting parts of various other species. We have added approximately 2000 sheets of aquatics to the museum herbarium as a final result of the work described above. GLEN S. WINTERRINGER ILLINOIS STATE MUSEUM, SPRINGFIELD STUDIES IN THE FLORA OF BOLIVIA — IV. ROBERT C. FOSTER Continued from Vol. 68, No. 773 11. Briza L. See Parodi in Revist. Fac. Agron. Vet. Buenos Aires, 3: 113-158 (1920). Perennials, the erect culms 2-8 dm. high. Leaves chiefly basal, much shorter than the inflorescence, glabrous, often scabridulous, 1-2 mm. wide. Inflorescence a loose to dense racemose panicle. Spike- lets oblong-elliptic, longer than broad, but the shorter ones verging on the suborbicular, usually many-flowered, the crowded florets erect to nearly horizontal, the naked rachilla disarticulating above the glumes; glumes subequal, the margins scarious, awnless; lemmas awnless, more or less wing-margined and closely imbricate. Lemmas, and often the glumes, tuberculate-scabrid; lemmas about 2 mm. long; palea glabrous. .......4 1. B. Mandoniana. Lemmas and glumes not tuberculate-scabrid; lemmas to 5 mm. long; palea covered with gland-tipped hairs. ........ 2. B. paleapilif era. 1. Briza Mandoniana (Griseb.) Henrard in Meded. Rijks Herb. Leiden, no. 40: 70 (1921). B. Mandoniana var. tuberculata Henrard in Meded. Rijks Herb. Leiden, no. 40: 71 (1921). B. Mandoniana var. Herzogiana Henrard in Meded. Rijks Herb. Leiden, no. 40: 71 (1921). B. Mandoniana var. vallegrandensis Henrard in Meded. Rijks Herb. Leiden, no. 40: 71 (1921). Spikelets 3-5 mm. long, oblong-elliptic, with up to 8 florets. Glumes 2-2.5 mm. long, broadly ovate, often scabrid to tuberculate, the scabrid apex somewhat erose, the basal margins shortly ciliate. Palea shorter than the lemma, bicarinate, the keels ciliolate. LA PAZ: LARECAJA: vic. Sorata; Lacatia and Lancha de Cochipata, 3200-3600 m., Mandon 1356 (GH; type-number) ; Cerro del Iminapi, 2650 m., Mandon 1355 (GH); Sorata, 4300 m., Rusby 238 (GH); NOR YUNGAS: Pongo, 3640 m., Hitchcock 22773 (GH). 2. B. paleapilifera Parodi in Revist. Fac. Agron. Vet. Buenos Aires, 3: 124, fig. 2, no. 5 (1920). B. stricta of the Catalogue.* Plants to *The Catalogue refers to A Catalogue of Ferns and Flowering Plants of Bolivia, in Contrib. Gray Herb. no 184 (1958). 6 dm. high. Leaf-blades to 2 mm. wide, glabrous to slightly scabri- dulous above and beneath. Panicle 7-12 cm. long, the rachis, branches and pedicels scabrid, at least in part. Spikelets oblong-elliptic, to 7 mm. long, to 8-12-flowered. Glumes to 4.5 mm, long, broadly ovate, 5-7-nerved, subacute to obtuse. Lemmas to 3-5 mm. long, subquadrate, the apex coarsely 3-lobed, the lateral lobes broad, obtusely rounded, 223 224 Rhodora [Vol. 68 the central lobe triangular, acute, keel prominent, thickened, the lateral nerves with a basal vertical line of short, thick, unicellular hairs. Palea oval, subacute to obtuse, about 2 mm. long, the dorsal surface with numerous short gland-tipped hairs. bicarinate, the keels narrowly winged, short-ciliate. COCHABAMBA: CERCADO: Cochabamba, Holway 383 (us). The Holway collection is the basis on which Hitchcock ascribed Briza stricta to Bolivia in his treatment of the Andean grasses. No Bolivian material of B. Stricta has been seen. Briza spicigera (Presl) Steud. Nomencl. (ed. 2) 1: 225 (1840). No material has been seen from Bolivia, but it has been ascribed to that country by Pilger in Notizbl. 11: 780 (1933). The basis for this consists of two Troll collections, no. 632 from CHUQUISACA: TOMINA: Alto de Escalón, and no. 834, from the Alto de Cuchilla, which I have been unable to locate. If the species should be found, it can be distinguished by the laterally compressed lemmas which are finely and densely papillate, emarginate at the apex, with the markedly thickened and prominent keel excurrent as a short thick mucro or cusp. The species is apparently identical with Briza uniolae (Nees) Nees ex Steud. Syn. Pl. Glum. 1: 283 (1854). If this is the case, the latter name should be applied to it, for its basionym, Eragrostis uniolae Nees, was published a year earlier than the basionym of Briza spicigera, Chascolytrum spicigerum Presl. It has been illustrated by Parodi, l. c., under the name Briza uniolae. 12. Eragrostis Beauv. Annuals or perennials, from very low mat-forming plants to erect plants up to 2 m. high. Leaves flat or involute, seldom equaling or exceeding the inflorescence at maturity. Inflorescence a terminal race- mose panicle, from dense, with erect compact branching, to very open, with freely and widely spreading branches, the spikelets 2-36-flowered disarticulating above the awnless glumes, the rachilla usually glabrous. Lemmas awnless, occasionally the lowermost sterile, but sterile and reduced florets usually borne at the tip, the paleas usually persistent on the rachilla after the fall of the lemmas and caryopses. a. Palea long-ciliate, the conspicuous hairs up to 1 mm. long. ................ mE l. E. ciliaris. a. Palea short-ciliate at most, the hairs inconsicuous. b. Largest spikelets at least 1 cm. long. c. Axils of inflorescence, at least the lowermost, pilose. d. Glumes equal or subequal. e. Glumes gland-dotted on keel, especially near the apex. .... 5M 3. E. cilianensis. e. Glumes not gland-dotted on keel. ............ 12. E. bahiensis. 1966] Flora of Bolivia — Foster 225 d. Glumes distinctly unequal. .............. 5. E. maypurensis. c. Axils of inflorescence glabrous. ............ 2. E. hypnoides. b. Largest spikelets less, usually much less, than 1 cm. long. f. Axils of inflorescence glabrous. g. Plants to 2 m. high, the leaf-blades to 6 mm. wide, scabridu- lous on both surfaces. ........ HMM 4. E. glomerata. g. Plants less than 1 m. high, the leaf-blades 2-3 mm. wide. h. Glumes equal or subequal. i. Spikelets 1 mm. wide, 2-6-flowered. ........ 7. E. nigricans. i. Spikelets 2 mm. wide, with 7 or more Ore. 561 EIE dedinfwernioeeii ARTE RE 9. E. mexicana. h. Glumes definitely unequal. ........... 13. E. expamsa. f. Axils of inflorescence, at least the lowermost, pilose to some extent. j. Lemmas 1.5 mm. or less in length. k. Largest spikelets 7 mm. long, with up to 12 florets. ........ MR o or Undighunikacecuhna bla E NE 6. E. articulata. k. Largest spikelets 5 mm. long, with up to 9 florets. .........--- MEME HER UE Mes qa ELE TEE ES 11. E. montufari. j. Lemmas 2 mm. or more in length. l. Rachilla puberulent. ........eA C eem 10. E. lurida. l. Rachilla glabrous, or with 1-2 short hairs at most below a floret. m. Leaf-blades glabrous beneath, scabridulous above. ........ EN Lus EE E E ETT DNE 8. E. pilosa. m. Leaf-blades at least sparsely pilose above. n. Panicle to 20 cm. long, leaf-blades sparsely pilose. 14. E. lugens. CedewesquciesesauesqueceinaqeAFéqaqapnanecetutesitanuadunsasti too p ales pilose. eene nennen 15. E. polytricha. 1. Eragrostis ciliaris (L.) Link, Hort. Berol. 1: 192 (1827). Annual, to 4 dm. high. Leaf-sheaths long-pilose at apex, blades flat, to 4 mm. wide, sparsely long-pilose at base, scabridulous above. Panicle dense, interrupted below and often above, the rachis glabrous, branches and pedicels scabridulous. Spikelets about 3 mm. long, with up to 8 florets, the rachilla glabrous. First glume about 1 mm. long, 1-nerved, the keel scabridulous; second glume about 1.5 mm. long, 1-nerved, the keel scabridulous. Lowest lemmas 1.5 mm. long, 3-nerved, the lateral nerves submarginal, minutely scabridulous; width (folded) 0.5 mm. Palea shorter than lemma, the 2 keels long-ciliate, the hairs up to 1 mm. long. SANTA CRUZ: SARA: Dolores, Steinbach 1831 (GH). 2. E. hypnoides (Lam.) BSP. Prelim. Cat. N. Y. 69 (1888). Annual, often forming prostrate mats, culms to 1 dm. high. Leaf-sheaths ci- liate ard sparsely pilose at apex, the short blades to 2 cm. long, flat or somewhat involute, sparsely pilose above, glabrous beneath. Ra- chis and pedicels glabrous, or the pedicels with a few short hairs at 226 Rhodora [Vol. 68 base or apex. Largest spikelets to about 1.5 em. long, commonly about 1 cm. long, with up to 25 florets, the rachilla glabrous. First glume 1.5 mm. long, 1-nerved, the keel scabridulous; second glume to 2.5 mm. long, 1-nerved, the keel scabridulous. Lowest lemmas to 2.5 mm. long, 3-nerved, the keel scabridulous, width (folded) about 0.6 mm, Palea about 1 mm. long, the 2 keels distinctly ciliolate. BENI: CERCADO MOJOS: Trinidad, 200 m., Cárdenas 3536 (GH). LA PAZ: LA- RECAJA: Guanai, 650 m., Rusby 230 (GH). SANTA CRUZ: SARA: Palo- metilla, Steinbach 2914 (GH). 8. E. cilianensis (All.) ex Vign. Lut. in Malpighia, 18: 386 (1904). Annual, to 5 dm, high, but usually shorter, the leaf-sheaths ciliate, pilose around the apex, the blades flat, to 4 mm. wide, scabrid-ciliate, otherwise glabrous. Panicle to 1 dm. long, fairly compact, the rachis glabrous, the pedicels usually scabridulous. Spikelets to 1 cm. long, with up to 20 florets, the rachilla glabrous. Glumes equal, to 1.5 mm. long, 1-nerved, the nerve with 1-2 glandular dots and scabridulous at the apex. Lowest lemmas about 2 mm. long, 3-nerved, the keel gland- dotted and scabridulous at the apex, scabridulous between lateral nerves and margin, width (folded) 1 mm. Palea about 1.5 mm. long, the 2 keels ciliate. Bolivian Plateau: Bang 1078 (GH). COCHABAMBA: CERCADO: Cochabamba, 2700 m., Eyerdam 24882 (GH). SANTA CRUZ: SARA: Buenavista, Steinbach 1944 (GH). 4. E. glomerata (Walt.) L. H. Dewey in Contrib. U. S. Nat. Herb. 2: 543 (1894). Annual, to 2 m. high. Leaf-sheaths glabrous, the long- attenuate flat blades to 30 cm. long and 6 mm. wide, scabridulous on both surfaces. Panicle to 4 dm. long, narrow, the branches erect, appressed, rachis and pedicels scabridulous. Spikelets about 3 mm. long, or less, with up to 7 florets, the rachilla glabrous. Glumes equal, 1 mm. long or less, 1-nerved, the keel sparsely scabridulous. Lowest lemmas about 1 mm. long, 3-nerved, the keel scabridulous, width (folded) less than 0.5 mm. Palea about 0.5 mm. long, the keels short- ciliate. LA PAZ: SUR YUNGAS: Covendo, White (M. E.)* 915, 1000 (Hitchcock). 5. E. maypurensis (HBK.) Steud. Syn. Pl. Glum. 1: 276 (1854). Annual, to 3 dm. high, leaves mostly basal, short, the sheaths pilose, the narrow blades pilose, at least when young. Panicle to 15 em. long, the rachis and pedicels glabrous to pilose. Spikelets very vari- able, the largest 1-2.5 cm. long, with up to 36 florets, the rachilla glabrous. First glume to 2.5 mm. long, 1l-nerved; second glume to 3 mm. long, 3-nerved, broader than the first. Lowest lemmas to 3 mm. long, long-acute or acuminate, 3-nerved, the keel scabridulous, width (folded) 1 mm. Palea nearly 2 mm. long, the keels short-ciliate. *The initials M. E. refer to collections made on the Mulford Bio- logical Exploration. The number-series of these collections duplicates other number-series of the same collectors. 1966] Flora of Bolivia — Foster 227 SANTA CRUZ: SARA: Dolores, near Buenavista, 500 m., Steinbach 6138 (GH). 6. E. articulata Nees, Agrost. Bras. 502 (1829). Annual, to 4 dm. high. Leaf-sheaths sparsely pilose, the flat, short blades 2-4 mm. wide, densely papillose-hispid, especially beneath. Panicle usually about half the total length of culms, open, the branches distant, the rachis glabrous, branches and pedicels sparsely scabridulous, Largest spikelets to 7 mm. long, 1 mm. wide, with up to 12 florets, the rachilla glabrous. First glume 1 mm. long, 1-nerved, the keel scabridulous; second glume about 1.5 mm. long, obscurely 3-nerved, the keel scabri- dulous. Lowest lemmas about 1.5 mm. long, 3-nerved, width (folded) 0.5 mm. Palea shorter than lemma, bicarinate, the keels glabrous, the margins finely ciliolate. COCHABAMBA: MIZQUE: near Vilavila, 2700 m., Eyerdam 24965 (GH). SANTA CRUZ: SARA: Buenavista 500 m., Steinbach 5182, 6142, 6991 (all GH). Eragrostis articulata var. glabrescens Henrard in Meded. Rijks Herb. Leiden, no. 40: 69 (1921) was distinguished as having glabrous leaves, According to Hitchcock (Contrib. U. S. Nat. Herb. 24: 338), the “spikelet is more slender and has more florets than those of E. articulata, and appears to belong to a distinct species." 7. E. nigricans (HBK.) Steud. Nomencl. (ed. 2) 1: 563 (1840). E. subatra Jedw. in Mez, Bot. Arch. 5: 202 (1924). Annual, the decumbent culms to 5 dm. long. Leaf-sheaths glabrous but pilose at the base of the blade, the blades flat, to 3 mm. wide. Panicle dark, to 2 dm. long, open, the branches spreading, rachis, branches and pedi- cels glabrous. Spikelets 4-6 mm. long, with 2-6 florets, the rachilla glabrous. Glumes equal, about 2 mm. long, the second broader than the first, the keels scabridulous. Lowest lemmas 2.5 mm. long, ob- scurely nerved, the keel slightly scabridulous, width (folded) 0.7 mm. Palea 2 mm. long, keels ciliolate. LA PAZ: MURILLO: La Paz, 3800 m., Buchtiem 42 (GH), 3965 (GH), 3300 m., Bang 21 (GH), 80 (GH; type-number of E. subatra), Pennell 14216 (GH); Obrajes, 3300 m., Buchtien 562 (GH). POTOSÍ: SUR CHICHAS: Oploca, Hitchcock 22928 (GH). 8. E. pilosa (L.) Beauv. Ess. Agrost. 162 (1812). Annual, to 3 dm. high. Leaf-sheaths long-ciliate, pilose at apex, the flat blades about 2 mm. wide, scabridulous above, glabrous beneath. Panicle open at maturity, to 20 cm. long, rachis glabrous, the branches and pedicels often scabridulous. Spikelets to 5 mm. long, about 1 mm. wide, with up to 8 florets, the rachilla glabrous. First glume about 0.75 mm. long, 1-nerved, the nerve faintly scabridulous; second glume slightly over 1 mm., 1-nerved. Lowest lemmas about 2 mm. long, obscurely 3- nerved, the keel scabridulous, width (folded) about 0.5 mm. Palea somewhat shorter than the lemma, bicarinate. LA PAZ: LARECAJA: vic. Sorata; near Munaypata, 2650-2900 m., Mandon 1329 (GH); MURILLO: Obrajes, 3400 m., Buchtien 820 (GH). COCHABAMBA: CERCADO: Cocha- 228 Rhodora [Vol. 68 bamba, 2560 m., Cárdenas 2471 (GH). SANTA CRUZ: SARA: El Limén, Steinbach 1786 (GH). TARIJA: ARCE: Padcaya, 2100 m., Fiebrig 2528 (GH). 9. E. mexicana (Lag.) Link, Hort. Berol. 1: 190 (1827). Annual, to 4 dm. high. Leaf-sheaths ciliate, pilose at the apex, the blades flat, glabrous, 2-3 mm. wide. Panicle fairly open, the branches distant, to 30 cm. long, rachis glabrous, the branches and pedicels scabridulous, Spikelets to 7-8 mm. long, with 7 or more florets, the rachilla glabrous. Glumes equal or subequal, to 2.5 mm. long, the first 1-nerved, the keel scabridulous, the second obscurely 3-nerved, the keel scabridulous. Lowest lemmas to 2.5 mm. long, the keel scabri- dulous, apex obtuse to subacute, width (folded) 0.7 mm. Palea to 2 mm. long, the keels ciliate. LA PAZ: SUR YUNGAS: Sirupaya, Buchtien 428 in part (Hitchcock). 10. E. lurida Presl, Rel. Haenk. 1: 276 (1830). Perennial, very variable in height, up to 1 m. tall. Leaf-sheaths ciliate and pilose at apex, the flat blades to 2 mm. wide, sometimes very sparsely pilose. Panicle to 3 dm. long, the primary branches distant, the rachis glab- rous, pedicels finely scabridulous. Spikelets to 5 mm. long, 7-8-flowered, the rachilla puberulent. First glume 1.5 mm. long, keel scabridulous, the second glume 2 mm. long, the keel scabridulous. Lowest lemmas to 2.5 mm. long, 3-nerved, the scabridulous keel very prominent, emarginate at the obtuse apex, width (folded) 0.8 mm. Palea as long as lemma, the keels ciliate. La Paz: LARECAJA: vic. Sorata; Ullontiji, 2900 m., Mandon 1332 (GH; type-number of E. bahiensis var. boli- viensis); MURILLO: near La Paz, 4000 m., Rusby 49 (GH), 3800 m., Buchtien (GH). 11. E. montufari (HBK.) Steud. Nomencl. (ed. 2) 1: 563 (1840). E. Buchtienii Hack. in Fedde, Rep. Sp. Nov. 6: 157 (1908). E. boli- viensis Jedw. in Mez, Bot. Arch. 5: 205 (1924). Perennial, to 6 dm. high. Leaf-sheaths pilose, at least at apex, the flat blades to 4 mm. wide, pilose above and beneath. Panicle 10-30 cm. long, the primary branches distant, rachis, branches and pedicels scabridulous. Spike- lets to 5 mm. long, up to 9 florets, about 1 mm. wide, the rachilla glabrous except for an occasional short hair at the apex of a section. Glumes about equal, 1 mm. long, or a little more, l-nerved, the keels scabridulous. Lowest lemmas 1.5 mm. long, rounded, obscurely nerved, the keel scabridulous at apex, width (folded) 0.5 mm. Palea about 1 mm. long, bicarinate, the keels glabrous, the margins ciliolate. LA PAZ: LARECAJA: vic. Sorata; Cerro del Iminapi, 2650 m., Mandon 1330 (GH; type-number of E. boliviensis) ; MURILLO: La Paz, Bang 201 (GH); Cotana, 2450 m., Buchtien (GH). COCHABAMBA: CERCADO: Cochabamba, Hitchcock 22798 (GH). 12. E. bahiensis Schrad. ex Schult. Mant. 2: 318 (1824). Perennial, to 8 dm. high, Leaf-sheaths pilose at apex, the blades 2 mm. wide, flat or somewhat involute, sparsely pilose. Inflorescence a lax to 1966] Flora of Bolivia — Foster 229 compressed panicle, to 15 cm. long, the branches distant, the rachis glabrous, branches and pedicels scabridulous. Largest spikelets over 1 em. long, 3-4 mm. wide, with up to 15 florets, the rachilla glabrous. Glumes subequal, about 2 mm. long, obscurely nerved, the keels scabridulous at apex. Lowest lemmas to 3 mm. long, 3-nerved, the keel very prominent and scabridulous at apex, width (folded) 1 mm. Palea 2 mm. long, bicarinate, the keels glabrous, the margins ciliate. La Paz: NOR YUNGAS: Coripata, Hitchcock 22681 (GH). SANTA CRUZ: SARA: Buenavista, 500 m., Steinbach 5184 (GH), 6881 (GH); Rio Perdix, 400 m., Steinbach. 6889 (GH). 13. E. expansa Link, Hort. Berol. 1: 190 (1827). Perennial, to 8 dm. high. Leaf-sheaths pilose at apex, the blades flat to involute, 2 mm. wide, scabridulous above. Panicle open, to 15 cm. long, primary branches distant, rachis glabrous, branches and pedicels scabridulous. Largest spikelets about 5 mm. long (in some non-Bolivian material to 1.5 em.), 1 mm. wide, with up to 1l florets, the rachilla usually glabrous but occasionally with a few hairs. First glume 1 mm., the keel scabridulous; second glume 1.5 mm., the keel scabridulous. Lowest lemmas 2 mm. long, 3-nerved, the keel scabridulous at the apex, width (folded) 0.5 mm. Palea 1.5 mm. long, keels glabrous, margins ciliolate. SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 6853 (GH); Río Perdix, 400 m., Steinbach 6899 (GH). 14. E. lugens Nees, Agrost. Bras. 505 (1829). E. soratensis Jedw. in Mez, Bot, Arch. 5: 213 (1924). Perennial, to 5 dm. high. Leaf- sheaths pilose to sparsely pilose and ciliate, to more or less glabrate, the blades flat to involute, usually sparsely pilose. Panicle open, to 20 cm. long, the primary branches distant, the rachis glabrous, branches and pedicels somewhat scabridulous. Spikelets 5 mm. long, or less, about 1 mm. wide, 5-6-flowered, the rachilla glabrous. First glume 1 mm., the second 1.5 mm. long, the keels scabridulous. Lowest lemmas 2 mm. long, obscurely nerved, the keel and apex slightly scabridulous, width (folded) 0.8 mm. Palea somewhat shorter than the lemma, keels glabrous, margins ciliolate. Yungas: Bang 307 (GH). LA PAZ: LARECAJA: Sorata, 3300 m., Rusby 239 (GH); Cerro del Iminapi, 2650 m., Mandon 1283 (GH); Sicasica, 2650-2900 m., Mandon 1331 (GH; type-number of E. soratensis). COCHABAMBA: MIZQUE: near Vilavila, 2500 m., Eyerdam 25021 (GH). SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 7091 bis (GH). 15. E. polytricha Nees, Agrost. Bras. 507 (1829). Perennial, up to 1 m. high. Leaf-sheaths densely long-pilose, the blades flat to in- volute, densely long-pilose. Panicle to 3 dm. long, very open, the rachis, branches and pedicels scabridulous. Spikelets 5 mm. or less, 3-5-flowered, 1.3 mm. wide, the rachilla glabrous with occasionally a few short hairs below the florets. First glume 1 mm. long, the second, 1.5 mm., keels strongly scabridulous. Lowest lemmas to 2 mm. long, keel scabridulous, width (folded) 0.5 mm. Palea 1.5-2 mm. 230 Rhodora [Vol. 68 long, margins ciliolate. SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 6980 (GH). 13. Distichlis Raf. Perennial, plants dioecious, rhizomatous, culms erect, from 3 to 30 cm. high. Leaves glabrous or glaucous or softly puberulent or even pilose, particularly on the sheaths, obviously distichous, shorter to longer than the inflorescence, blades usually folded or involute, 1 mm. wide or less. Inflorescence a terminal racemose panicle, the panicle somewhat condensed. Spikelets few- to many-flowered, the staminate spikelets of a rather softer texture than the pistillate, glumes present, the naked rachilla disarticulating above the glumes, the lemmas rather obscurely nerved. a. Dwarf plants, less than 10 cm. high. b. Leaf-sheaths pilose, at least in part, blades usually glabrous; panicles of 1-3 spikelets, the panicle about 1 cm. long. ................ SN 2. D. spicata. l. Distichlis humilis Fhil. in Anal. Mus. Nac. Chile, 8: 86 (1891). Leaf-sheaths closely imbricate, the blades spreading, to 1 cm. long, or less. Spikelets barely or not at all exserted from leaves, to 7 mm. long, 3-6-flowered. Glumes equal to subequal, the second to 4 mm. long. Lemmas to 4.5 mm. long. Palea shorter than the lemma, bicarinate, the keels narrowly winged and minutely ciliolate. Potosi: sur CHICHAS: Atocha, Hitchcock 22876 (am). 2. D. spicata (L.) Greene in Bull. Calif. Acad. Sci. 2: 415 (1887). Plants generally much taller than the preceding, the leaf-blades much longer and often exceeding the inflorescence, the sheaths not closely imbricate. Panicle exserted from the leaves, to 7 em. long. Spikelets to 1.8 em. long and up to 20-flowered. Glumes unequal, the first to 1.5 mm. long, the second to 2.5 mm. Lemmas about 3.5 mm. long. Palea as long as the lemma, bicarinate. LA Paz: MURILLO: La Paz, 3300 m., Bang 108 (au). COCHABAMBA: CERCADO: Valle de Cocha- bamba, 2600 m., Steinbach. 8745 (GH). POTOSÍ: SUR CHICHAS: Oploca, Hitchcock 22883 (GH), 22884 (GH). 2a D. spicata var. andina Beetle in Revist. Argent. Agron. 22: 93 (1955). Differs in smaller size, short leaf-blades (to 2 em.), shorter panicles, smaller and fewer-flowered spikelets. LA PAZ: MURILLO: San Jorge, 3550 m., Buchtien 8537 (GH). Distichlis spicata gamma marginata (Phil.) O. Ktze. Rev. Gen. 3 (2) : 350 (1898) was recorded for Bolivia by Kuntze, 1966] Flora of Bolivia — Foster 281 on the basis of a collection made by him. In view of some of Kuntze's notorious misdeterminations, I hesitate to accept it on such a basis alone. 14. Arundo L. Arundo Donax L. Sp. Pl. 81 (1753). Rhizomatous perennial, the culms to 6 m. high, occasionally branched. Leaves very long, 3-7 cm. wide, the margins scabrid. Inflorescence a large, dense, terminal, drooping, racemose panicle to 6 dm. long. Spikelets to 1.2 cm. long, 3-4-flowered, the florets perfect, successively smaller upwards, the naked rachilla disarticulating above the glumes; glumes somewhat unequal, the first about 1 cm. long, the second 1.4 cm., as long as the spikelet; lemmas about 1 cm. long, hyaline, strongly 3-nerved, apex 2-toothed with a central awn to 3 mm. long, the rounded back and sides densely long-villous; palea to 5 mm. long, obtuse, with 2 promi- nent marginal nerves, the margins densely ciliate. LA PAZ: NOR YUNGAS: Coroico, Hitchcock 22724 (US). 15. Phragmites Trin. Phragmites communis Trin. Fund. Agrost. 134 (1820). Rhizomatous stoloniferous perennial, with culms to 4 m. high. Leaves long, to 5 cm. wide. Inflorescence a terminal, plumose, racemose, tawny panicle, the branches ascending, to 4 dm. long. Spikelets about 1.5 cm. long, 2-7-flowered, the long-villous rachilla disarticulating above the glumes; glumes unequal, the first nearly 5 mm, long, 3-nerved, the second, 6.5 mm, long, 5-nerved, much shorter than the spikelet; lowest floret staminate or sterile, the awnless lemma about 1.2 em. long; upper florets perfect, the glabrous lemmas long-attenuate into a 6 mm. pseudo-awn; palea about 5 mm. long, acute, slightly bidentate, the margins ciliolate. CHUQUISACA: SUR CINTI: Camataquí, about 2500 m., Fiebrig 2954 (US). 16. Cortaderia Stapf Large perennials, to 1.5 m. high. Leaves nearly as long as the culms, the blades to 1.5 em. wide, the margins and midrib scabrous beneath, old leaf-bases coiled at base of plant. Inflorescence a large, terminal, plumose, racemose panicle. Spikelets unisexual or perfect, if unisexual the plants dioecious, 1-several-flowered, the glumes often as long as the spikelet, hyaline, the villous rachilla disarticulating above the glumes; lemmas hyaline, tapering to a long awnlike pro- longation or bearing 2 such structures flanking a true awn. Plants dioecious; lemma not bidentate, with 1 long pseudo-awn. ........ 1. C. rudiuscula. Diodes laistaqisusvesoquédésansesevesaueoetéeesabaspéaenvendsseusatgoran aere e Esa ee 232 Rhodora [Vol. 68 1. Cortaderia rudiuscula Stapf in Gard. Chron. (ser. 3) 22: 396 (1897). Panicle yellowish or purplish, to 6 dm. long, the branches dense and drooping. Staminate spikelets about 1 em. long, 2-3-flowered (occasionally 1-flowered). Glumes 6-8 mm. long, 1-nerved. Lemmas 5 mm. long, with a 6 mm. pseudo-awn, villous on the back and sides. Palea as long as the lemma, Pistillate spikelets about 1.2 em. long, about 4-flowered. Glumes nearly equal, 8 and 9 mm. long, 1-nerved. Lemmas about 4 mm. long, the pseudo-awn about 6 mm. long, densely long-villous on back and sides. Palea as long as lemma, bicarinate, the keels ciliolate. LA PAZ: MURILLO: La Paz, 3750 m., Buchtien 507 (GH), Shepard 167 (GH), 3300 m., Bang 26 (GH); SUR YUNGAS: Chulu- mani, 1600 m., Hitchcock 22701 (GH). COCHABAMBA: CHAPARE: Inca- chaca, 2250 m., Steinbach 9493 (GH). PoTosí: CHICHAS: Quechisla, Cardenas 98 (GH). SANTA CRUZ: SARA: quebrada del Rio Blanco, Cerro Hosana, 1300 m., Steinbach 3372 (GH). TARIJA: CERCADO: Tarija, 2850 m., Fiebrig 2656 (GH); Paicho, 2900 m., Fiebrig 3043 (GH). 2. C. bifida Pilger in Engler, Bot. Jahrb. 37: 374 (1906). C. bifida var. grandiflora Henrard in Meded. Rijks Herb. Leiden, no. 40: 67 (1921). Panicle yellowish to purple, about 2 dm. long, laxer than in C. rudiuscula, Spikelets of perfect florets, 3-4 in number, about 8-10 mm. long (excluding the awns). Glumes unequal, 7 and 9 mm. long, hyaline, 1-nerved, long-aeute. Lemmas about 5 mm. long, the 2 teeth prolonged into pseudo-awns 6-18 mm. long, true awn 1.1-2.2 em. long, lemma ]long-villous, primarily near the base. La Paz: MURILLO: Unduavi, 3200 m., Buchtien 2575 (GH). SANTA CRUZ: SARA: Cerro La Negra, 1800 m., Steinbach 8171 (GH). 17. Gynerium Humb. & Bonpl. Gynerium sagittatum (Aubl.) Beauv. Ess. Agrost. 138 (1812). Large dioecious perennial, to 10 m. high. Leaf-blades to 2 m. long and 4-6 cm. wide, the margins scabrous-serrulate. Inflorescence a terminal, dense, plumose, racemose panicle to 1 m. long, the branches drooping. Spikelets unisexual, usually several-flowered, the naked rachilla dis- articulating above the glumes, glumes and lemmas quite different in staminate and pistillate spikelets. Staminate spikelets: 1-3-flowered, 4 mm. long; glumes somewhat unequal, 2.5 and 3 mm. long, hyaline, acute but not produced; lemmas to 3.5 mm. long, glabrous, acute but not produced. Pistillate spikelets: 2-3-flowered, 6 mm. long; glumes very unequal, the first 3-4 mm. long, acute, the second 9-10 mm. long, long-attenuate and produced into a pseudo-awn much exceeding the spikelet; lemmas to 5 mm. long, long-attenuate into a pseudo-awn about 4-5 mm. long, densely long-villous; stigmas black-purple. LA PAZ: NOR YUNGAS: Coripata, Hitchcock 22683 (Hitchcock); SUR YUNGAS: Covendo, White (M. E.) 901 (Us), 984 (Us). 1966] Flora of Bolivia — Foster 233 18. Melica L. See Papp in Fedde, Rep. Sp. Nov. 25: 97-160 (1928). Straggling perennials, culms up to 1 m. high, usually shorter. Leaf- sheaths scabrous, the blades flat or involute, occasionally equaling or even exceeding the inflorescence. Inflorescence a rather narrow, termi- nal, racemose panicle, the branches distant, spreading. Spikelets 1-4- flowered, the lowest 1-2 perfect, the upper 2-3 sterile and aggregated into a clavate mass, the rachilla glabrous or minutely puberulent, disarticulating above the glumes, the spikelet falling as a unit; lemmas densely puberulent or tuberculate-scabridulous. a. Lowest lemma with large thick hairs on lateral and submarginal naush DOURINCUURU D CN R E LEE 1. M. violacea. a. Lowest lemma without large thick hairs. b. Spikelets to 1 cm. long, perfect florets 1-2; leaf-blades flat, 2-6 mm. wide. eese Leian eene nennt nnne nennt nnne entente nennen 2. M. scabra. b. Spikelets to 7 mm. long, 1 perfect floret; leaf-blades narrow, TI DIL RUNS ARR 3. M. adhaerens. 1. Melica violacea Cav. Ic. 5: 47, t. 472 (1799). Spikelet about 7 mm. long, with 1 perfect floret. First glume suborbicular, about 7 mm. long, prominently nerved on the lower portion, the hyaline margins broad; second glume much narrower, about as long, densely scabridulous-puberulent. Fertile lemmas 5-6 mm. long, many-nerved, densely puberulent. Palea as long as lemma, bicarinate, the keels ciliolate, the inter-carinal area scabridulous-puberulent. SANTA CRUZ: VALLEGRANDE: Comarapa, 3000 m., Steinbach 8461 (GH). 2. M. scabra HBK. Nov. Gen. & Spec. 1: 164 (1816). M. scabra var. glabra Papp in Fedde, Rep. Sp. Nov. 25: 145 (1928). Spikelets to 6-10 mm. long, with 1-2 perfect florets. Glumes unequal, more or less hyaline, especially on the margin, 1-nerved, the first 8 mm. long, the second to 10 mm. Fertile lemmas to 8 mm. long, the upper portion hyaline, the lower part many-nerved, the nerves and intercostal areas densely tuberculate-scabridulous. Palea to 5 mm. long, bicarinate, the keels ciliate, the back usually puberulent. LA PAZ: LARECAJA: Sorata, Mandon 1343 (US). 3. M. adhaerens Hack. in Fedde, Rep. Sp. Nov. 6: 158 (1908). M. adhaerens var. tenuis (Papp) Papp in Notizbl. 10: 412 (1928). M. Mandonii Papp in Notizbl. 10: 356 (1928). Spikelet about 7 mm. long, with 1 perfect floret. First glume 7 mm. long, broadest above the middle, abruptly short-acute, l-nerved; second glume as long, much narrower, the nerves obscure, lower half puberulent-scabrid. Fertile lemma about 6 mm. long, apex hyaline, lower half many-nerved, densely tuberculate-scabrid. Palea about 4 mm. long, bicarinate, the keels ciliolate, the back densely puberulent. LA PAZ: MURILLO: Poto- poto, Río Chuquiaguillo, 3700 m., Mandon 1357 (GH; type-number of M. Mandonii) ; La Paz, 3300 m., Bang 30 (Gu), 3500 m., Buchtien 505 234 Rhodora [Vol. 68 (GH), 3700 m., Hitchcock 22565 (GH). COCHABAMBA: MIZQUE: Vila- vila, 2500 m., Eyerdam 24983 (GH). 19. Anthochloa Nees See Matthei in Gayana, Bot. no. 8: 11-15 (1963). Anthochloa lepidula Nees & Mey. in Meyen, Reise, 2: 14 (1835). A. rupestris Remy in Ann. Sci. Nat. (Ser. 3) 6: 347 (1846). Densely caespitose dwarf perennial, less than 10 cm. high. Leaf-blades 0.5- 1.5 mm. wide, somewhat scabridulous, often exceeding the inflorescence. Inflorescence a greatly reduced racemose panicle, more or less capitate in appearance, sometimes only 1 spikelet to a panicle-branch. Spike- lets about 8 mm. long, with 4-6 perfect florets, the naked rachilla disarticulating above the glumes. Glumes hyaline, the apex sometimes erose, 2 and 2.5 mm. long. Lemmas hyaline, scarious, 4 mm. long and 5 mm. wide, deeply bifid. Palea shorter than lemma, hyaline, deeply 2-3-cleft almost to the base. LA Paz: LARECAJA: vic. Sorata; Lacatia ad La Apacheta, 4500-5000 m., Mandon 1372 (Gu). 20. Orthoclada Beauv. Orthoclada laxa (Rich.) Beauv. ex Nees, Agrost. Bras. 522 (1829). Perennial, stoloniferous, the culms up to 1 m. high. Leaf-sheaths puberulous, the lance-ovate blades to 15 cm. long, and 2.5 em. wide, scabridulous above, the base long-petiolately attenuate, glabrous. In- florescence a terminal racemose panicle, the slender branches ulti- mately divergent, erect-appressed before maturity. Spikelets usually solitary, disarticulating below the glumes, usually with 1 perfect floret but often 2-flowered, fertile floret below sterile, 6-10 mm. long, the rachilla pubescent. Glumes unequal, to 4 and 5 mm. long, the first 3- nerved, main nerve scabrid, the second 5-nerved, all nerves scabrid on upper portion at least. Lemma to 7 mm. long, obscurely 5-nerved, scabrid-puberulent on upper portion. Palea about 4.5 mm., bicarinate, the keels ciliolate. SANTA CRUZ: ICHILO: Río Vibora, 350 m., Steinbach 7576 (GH); SARA: Rio Surutü, 400 m., Steinbach 7294 (GH). 21. Zeugites Schreb. Zeugites mexicana (Kunth) Trin. ex Steud. Nomencl. (ed. 2) 2: 798 (1841). Z. mexicana var. glandulosa Hack. in Fedde, Rep. Sp. Nov. 6: 758 (1908). A slender much-branched perennial, the wiry culms usually less than 30 em. high. Leaf-sheaths sparsely pilose, especially at the apex; leaf-blades long-petiolate, the petiole pubescent at base and apex, the blades broadly lance-ovate, acute, up to 3 em. long and 1.5 em. wide. Inflorescence a few-branched, few-flowered, open, race- mose panicle up to 5 cm. long. Spikelets few-flowered (2-3), the lowest floret perfect, the upper florets staminate and decidedly distant from the lowermost, the naked rachilla disarticulating above the glumes, the spikelet falling as a unit; spikelet to 6 mm. long. Glumes 1966] Flora of Bolivia — Foster 235 somewhat unequal, 1.5 and 2 mm. long, herbaceous, often purple- tipped, the second 5-nerved, the 3 central nerves connected by con- spicuous transverse veinlets, both glumes with 2-3 very short teeth at apex, and with slightly cilielate margins. Lemmas 3 mm. long, T-nerved, with a very short central apical tooth. Palea as long as the lemma, strongly bicarinate in the upper portion. LA PAZ: NOR YUN- GAS: Coripata, Bang 2131 (GH); Bella Vista, Hitchcock 22751 (GH). COCHABAMBA: CHAPARE: Incachaca, 2600 m., Steinbach 9872 (GH). 22. Triodia R. Br. Triodia avenacea HBK. Nov. Gen. & Sp. 1: 156 (1816). Caespitose dwarf perennial, from 3-10 (-20) cm. high. Leaf-sheaths sparsely pilose, the flat blades to 3 cm. long, 2 mm. wide, falcate, pilose above and beneath (at least when young), obtuse, shortly mucronate. Inflo- rescence a short (1-4 cm.), dense, terminal, racemose panicle, the rachis villous. Spikelets about 6 mm. long, with up to 5 perfect florets, the naked rachilla disarticulating above the glumes. Glumes unequal, the first 5 mm. long, with a ciliolate keel, the second, 7-8 mm. long, exceeding the spikelet, the keel scabridulous. Lowest lemma about 4 mm. long, the lemmas decreasing in size upwards on the rachilla, deeply bifid, with a scabridulous 4 mm. awn arising from the base of the sinus, the 2 lobes shortly 2-toothed, the rounded back densely villous at base, ciliate, the hairs increasing in length toward the apex. Palea much shorter than the lemma, bicarinate, the keels ciliate, long- villous between the keels and the margin. PoTosí: CERCADO: Potosí, 4000 m., Cárdenas 204 (GH). 23. Pappophorum Schreb. Caespitose perennials, the culms 2-5 dm. high. Leaves pubescent or pilose on sheaths and blades, the blades narrow, 1-2 mm. wide, not equaling the inflorescence or sometimes exceeding it. Inflorescence a terminal, narrow, racemose panicle, spicate in appearance. Spikelets 1-3-flowered, the lowest floret perfect, the rest sterile, the puberulent rachilla disarticulating above the thin-textured glumes; lemmas firn or indurate, the apex dissected into numerous awns. Panicle lead-colored; awns plumose, 3.5-5 mm. long. .... 1. P. Wrightii. Panicle light to purplish; awns scabrid, 0.5-1 cm. oaea A nore rere 2. P. mucronulatum. Q65000440000Q0400000400000090 52000090 009099000 09000004600 052400 000009009 904699508000 CADO: Tarija, Fries 1221 (US). 2. P, mucronulatum Nees, Agrost. Bras. 412 (1829). Panicle 5-10 236 Rhodora [Vol. 68 cm. long. Spikelets about 1.4 em. long (including the awns). Glumes glabrous, short-awned (awns 1-1.5 mm.), 5 and 6 mm. long. Lemmas about 3.5 mm. long, the awns about 1 cm. long, villous at base exter- nally, internally pubescent, at least at the apex, the nerves not visible on the exterior. Palea as long as the lemma, bicarinate, the keels ciliolate. (COCHABAMBA: CERCADO: Cochabamba, Holway 367 (vs). TARIJA: CERCADO: Tarija, Fries 1099 (vs). 24. Agropyron Gaertn. Perennial, more or less erect, the culms to 1-2 m. high, basally decumbent, sometimes stoloniferous. Leaf-sheaths usually glabrous but occasionally puberulent; leaf-blades to 12 cm. long and 4 mm. wide, much shorter than the inflorescence, usually glabrous, some- times puberulent, long-acute. Inflorescence a single spike or raceme, or slightly paniculate (rarely so), the spikelets sessile, subsessile or with a 1 mm. pedicel, sometimes rather unilateral in arrangement. Spikelets solitary, occasionally paired, the rachilla disarticulating above the glumes, few- to several-flowered; glumes awnless, or very short-awned; lemmas awnless or short-awned, Glumes at least 1 cm. long, with 4-6 prominent nerves; lemmas awn- less or shortly mucronulate (muero 0.75 mm.). .... 1. A. attenuatum. Glumes less than 1 cm. long, nerves inconspicuous; lemmas with an AWN to 7-8 mm. long. eanan. 2. A. breviaristatum. l. Agropyron attenuatum (HBK.) Roem. & Schult. Syst. Veg. 2: 751 (1817). Spike 7-17 em. long. Spikelet about 2 cm. long, several- flowered (up to 6), the uppermost a sterile lemma, the 1 below it apparently staminate. Glumes subequal, 1-1.2 em. long, obtuse. Low- est lemmas 1-1.2 cm. long, lowest shortly tridentate, 5-nerved, its palea 9 mm. long, 2-nerved, the nerves markedly ciliolate, Upper lemmas equal to or slightly shorter than the lowest, the uppermost much shorter. LA PAZ: MURILLO: La Paz, 3700 m., Buchtien 8532 (au); NOR YUNGAS: Pongo, 3640 m., Hitchcock 22766 (GH). 2. A. breviaristatum Hitchc. in Contrib. U.S. Nat. Herb. 24: 353 (1927). Spikes 10-20 em. long. Spikelets 1-1.5 em. long, 3-5-flowered. Glumes subequal, 7-10 mm. long, acuminate, scabridulous, especially on the nerves, Lemmas 8-9 mm. long, scabrid, awned. LA Paz: OMA- SUYOS: Isla del Sol, Asplund 6461 (Hitchcock). 25. Elymus L. Elymus angulatus Presl, Rel. Haenk. 1: 264 (1830). Perennial, culms spreading at base but erect, to 1 m. high, the plants glabrous. Leaf- blades much shorter than the inflorescence, to 6 mm. wide, acute, scabridulous beneath. Inflorescence a terminal spike, with 2 sessile spikelets at a node (rarely solitary ?), the spike, to 15 cm. long. Spikelets about 1.3 cm. long (excluding the awns), up to 6-flowered, always with more than 1 perfect floret, disarticulation above the 1966] Flora of Bolivia — Foster roi yd glumes; glumes equal or subequal, about 1 cm. long, awn 3 mm. long, apex ciliolate, mid-nerve and awn scabridulous to scabrid, at least on the upper portion of the glume; lemmas distinctly rounded, subterete, about 1 cm. long, awns 4-5 mm. long, scabridulous near the apex, palea nearly as long as the lemma, bicarinate, the keels densely scabrid-ciliate on the upper half; uppermost floret a rudi- ment, the one below it neuter or possibly sometimes staminate. Porosí: SUR CHICHAS: Oploca, Hitchcock 22892 (GH). 26. Hordeum L. Perennials, erect or spreading, or decumbent, sometimes caespitose, usually 2-4 dm. high. Leaves glabrous, the short flat blades to 3 mm. wide, acute, scabridulous beneath, never equaling nor exceeding the inflorescence. Inflorescence a short terminal spike, less than 10 cm. long, with 3 spikelets at a node, the rachis jointed and disarticulating. Spikelets 1-flowered, glumes present, the rachilla disarticulating above the glumes, the lemmas acuminate or shortly or long-awned, the palea bicarinate; central spikelet sessile, the 2 laterals somewhat pedicellate and reduced to slender setose glumes. Lemmas awnless or very shortly awned; glumes of fertile floret with awns 4 mm. long; rachilla obviously prolonged behind the palea. .... NUDO UE Se A LU ebrei EE E USE ER PEE 1. H. muticum. Lemma with a 6 mm. awn; glumes of fertile floret with awns 1 cm. long; rachilla only slightly or not at all prolonged behind the palea. EP AUIoorusdis dumis Min E daa meetin 2. H. halophilum. 1.Hordeum muticum Presl, Rel. Haenk. 1: 327 (1830). Spikelet, ex- cluding awns, about 6 mm. long. Glumes reduced virtually to scabridu- lous awns 4 mm. long. Lemma about 5 mm. long, seldom short-awned. Palea less than 5 mm. long, bicarinate. Sterile glumes reduced to awns 4-5 mm. long. LA Paz: OMASUYOS: vic. Achacache, 3950 m., Mandon 1379 (GH); MURILLO: La Paz, 3750 m., Buchtien 55 (GH), Bang 179 (GH). PoTosi: CERCADO: Potosi, 4000 m., Cardenas 207 (GH). TaRIJA: AVILES: Puna Patanca, 3700 m., Fiebrig 3193 (GH). Hordeum nodosum L. Sp. Pl. (ed. 2) 126 (1762) has been ascribed to Bolivia by Hitchcock on the basis of a collection made by Pearce. I have seen no South American material of this species. It is apparently like H. muticum but with longer awns. 2. H. halophilum Griseb. in Goett. Abh. 19: 249 (1874). Spikelet, excluding awns, about 5 mm. long. Glumes of fertile floret reduced to awns 1 cm. long. Lemma 5 mm. long, with a 6 mm. awn. Palea about 6 mm. long, bicarinate, the apex bidentate. Sterile glumes re- duced to awns over 1 cm. long. LA PAZ: MURILLO: La Paz, 8700: m. Buchtien 506 (GH). 238 Rhodora [Vol. 68 27. Lolium L. Annual or perennial, 2-8 dm. to 1 m. high. Leaf-blades long, often nearly equaling the inflorescence, to 1 em. wide, scabridulous beneath. Inflorescence a terminal spike, 2-25 cm. long, the sessile spikelets solitary at the nodes and placed edgeways to the rachis. Spikelets several-flowered, first glume absent from all but the uppermost spike- let, rachilla disarticulating above the glumes, lemmas dorsally round- ed, awned or awnless. a. Glume to 2 cm. long, usually as long as, often exceeding, the SPikelet. wo. ccc ccsecsscsssssceeseaesescsscssscssssavsecsestecsecseseses 1. L. temulentum. a. Glume less than 1 cm. long, usually much shorter than the spikelet. b. Lemmas awnless, esee 2. L. perenne. b. Lemmas awned. pra aan. 3. L. multiflorum. 1. Lolium temulentum L. Sp. Pl. 83 (1753). Annual. Spikelet 1.5-2 cm. long, with up to 7-8 florets. Second glume to 2 cm. long, almost always longer than the rest of the spikelet, very firm-textured. Lem- mas very firm, about 8 mm. long, obscurely nerved, the apex with 2 blunt hyaline teeth and a very short scabrid awn rising from between the bases of the teeth, or the uppermost lemmas sometimes very long- awned. Palea firm-textured, nearly as long as the lemma, the upper margins ciliolate. LA PAZ: LARECAJA: vic. Sorata; Cicirca, Munaypata, Mandon 1376 (au). la. L. temulentum var. arvense (With.) Bab. Brit. Bot. 377 (1843). Differs by awnless lemmas. No material has been seen. 2. L. perenne L. Sp. Pl. 83 (1753). Perennial. Spikelet about 1 cm. long, occasionally to 1.5 em., with up to 10 florets. Second glume about 5 mm. long, more or less obtuse. Lowermost lemmas about 5 mm. long, decreasing in length upward on the spikelet. Palea as long as the lemma, the margins densely and shortly scabrid-ciliate. LA Paz: MURILLO: La Paz, 3700 m., Buchtien 4487 (GH). 3. L. multiflorum Lam. Fl. Franc. 3: 621 (1778). Perennial. Spike- let (excluding awns) to 1.5 em. long, with up to 10 florets. Second glume about 1 cm, long, rather obtuse. Lemmas to 7 mm. long, the apex bidentate, awns to 6 mm. long. Palea as long as the lemma, the margins ciliate. COCHABAMBA: CERCADO: Valle de Cercado, 2700 m., Steinbach 9549 (GH). 28. Pariana Aubl. See Tutin in Journ. Linn. Soc. Bot. 50: 337-362 (1936). Perennial, from a creeping scaly rootstock, Leaves few, reduced to sheaths on fertile culms (except in no. 1), leaf-sheaths puberulent to glabrous, the orifice often fimbriate; blades broad, lanceolate to ovate, pubescent to glabrous, often scabrid beneath. Inflorescence condensed, spicate, male spikelets 1-flowered, pedicellate, in whorls of 4-6, usually 5, pedicels often fused marginally, a solitary 1-flowered 1966] Flora of Bolivia — Foster 239 female spikelet sessile within the whorl; rachis disarticulating be- tween the whorls. Glumes lateral, flanking the lemma, 1-3 nerved. Lemma ovate to oblong, coriaceous, usually 3-nerved. Palea shorter and thinner than the lemma. a. Leaf-sheaths without fimbriae at apex; lemmas less than 6 mm. long. b. Glumes 2-3-nerved; terminal spikelet pistillate. ........ 1. P. bicolor. b. Glumes 1-nerved; terminal spikelet sterile. ............ 2. P. gracilis. a. Leaf-sheaths fimbriate at apex; lemmas about 7 mm. long. ............ IRURE ce eT is ccdiripisanqaehteri n E 3. P. Swallenii. 1. Pariana bicolor Tutin in Journ. Linn. Soc. Bot. 50: 355 (1936). Sterile culms to 8 dm. high. Leaf-sheaths imbricate, glabrous, the efimbriate blades to 25 cm. long and 5 cm. wide, oblong to ovate, acu- minate, glaucous and scabridulous beneath, glabrous above. Fertile culms to 4.5 dm. high, leaves reduced to sheaths or the uppermost sheath sometimes with a rudimentary blade, sheaths puberulent, the orifice ciliate with short, stiff, white hairs. Inflorescence 5-9 cm. long. Male spikelets in whorls of five, pedicels 2-3 mm. long, puberu- lous, basally heavily bearded. Glumes 2-2.5 mm. long, acute to acumi- nate, glabrous or ciliolate, 2-3-nerved, the nerves confluent near the apex. Lemma 3.5-4.5 mm. long, lance-elliptic, acute, midrib prominent, glabrous. Female spikelet: glumes obtuse, 6 mm. long, lemma and palea 6 mm. long. LA Paz: LARECAJA: Mapiri, 1300 m., Rusby 232 (Us); San Carlos, Mapiri, 850 m., Buchtien 458 (GH; type-number) ; CAUPOLICÁN: San Buena Ventura, 500 m., Cárdenas (M. E.) 1889 (Us). 2. P. gracilis Doell in Mart. Fl. Bras. 2(2): 337 (1877). Sterile culms leafy, sheaths imbricate, fimbriae lacking. Leaf-blades oblong- lanceolate, to 15 cm. long and 2.5 em. wide, acuminate, Fertile culms leafless, the inflorescence slender. Male spikelets hardly imbricate, the pedicels to 3 mm. long, pubescent. Glumes to 3 mm. long, acute, 1-nerved. Lemma to 4.5 mm. long, oblong-ovate, acute, glabrous, sometimes red and purplish, the nerves obscure. LA PAZ: CAUPOLICAN: Mojos, d'Orbigny 147 (Tutin). No material has been available to me and this description has been derived from that of Tutin. 3. P. Swallenii R. C. Foster, spec. nov. Culmi steriles ad 6.5 dm., foliati, nodi dense breveque hispidi, vaginae puberulae vel glabrae, ad orem fimbriatae. Laminae (2) inaequales, late ovatae, acutae, ad 30 cm. longae et 10 cm. latae, supra glabrae, subtus glaucae et plus minusve scabridulae. Culmi fertiles ad 2.5 dm., efoliati, pubescentes, vaginis imbricatis glabris vel subpuberulis obtecti. Inflorescentia subcerassa, ad 13 cm. longa. Spiculae masculinae 5 in verticillo, pe- dicelli ad 3 mm. longi, puberuli, basi barbati, plus minusve coaliti. Glumae ad 4-5 mm. longae, 3-nervatae, acutae vel acuminatae, vel subsetaceae, ciliatae, minute papillatae. Lemma 7 mm. longum, ob- 240 Rhodora [Vol. 68 longum, obtusum vel subacutum, glabrum, 5-nervatum, nervis paucis connectentibus prope apicem. Spicula feminea; gluma prima ca. 7-8 mm. longa, 1-nervata, ciliata, puberula; gluma altera breviora, ciliata, glabra, 1-nervata; lemma ca. 6 mm. longum. COCHABAMBA: CHAPARE: Puerto Polonia, Río Coni, 14 km. east of San Antonio, 395 m., Oct., Cárdenas & Cutler 1358 (Typus, in Herb. Gray.). Since Mr. J. R. Swallen has described a number of new South American species in this genus, it is fitting that this apparent novelty should be named for him. Pariana lunata Nees, Agrost. Bras. 295 (1829), was ascribed to Bolivia by Hitchcock in his treatment of Andean grasses. The only specimen he cited, however, was sterile. The later monographer, Tutin, saw no Bolivian material, so that its presence in Bolivia is still problematical. Should fertile material be found, it can easily be distinguished from the three other Bolivian species by the fact that the fertile culms bear well-developed leaves rather than, at most, rudimentary leaf-blades. Pariana zingiberina Doell in Mart. Fl. Bras. 2(2) : 337 (1877) was also ascribed to Bolivia by Hitchcock. All the material cited by him was later cited by Tutin under the new species, P. bicolor Tutin. No Bolivian material was cited by Tutin for P. zingiberina, and it seems almost certain that it is not a member of the Bolivian flora. To be continued. | g we 29910 | 6 06 Volume 68, No. 774 including pages 121-240, was issued June 29, 1966. FARLOW REFERENCE LIBRARY Dodota JOURNAL OF THE | NEW ENGLAND BOTANICAL CLUB Conducted and published for the Club, by ALBION REED HODGDON, Editor-in-Chief ALBERT FREDERICK HILL D STUART KIMBALL HARRIS RALPH CARLETON BEAN ROBERT CRICHTON FOSTER , Associate Editors ROLLA MILTON TRYON | RADCLIFFE BARNES PIKE | LORIN IVES NEVLING, JR. Vol. 68 July-September, 1966 No. 775 CONTENTS: Studies in the Flora of Bolivia — IV. Robert C. Foster ccccsscosscovssevescossesengenscccsegcaseessscsscnecasccccecscccessoccee 241 Native North American Species of Lespedeza (Leguminosae). A'ndre F. Clewell 359 Environmental Control of a Taxonomic Character Separating Heterotheca S. Str. from Chrysopsis. C. John Burk and Chung-Hie Nah 406 The Nem England Botanical Club, Ine. Botanical Museum, Oxford St., Cambridge 38, Mass. SANE ETS EE AID PLO EN L ELE IE DEA TELE IES OE ETE TEE EE LEE leaded QTRbooora JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB Vol. 68 July-September No. 775 STUDIES IN THE FLORA OF BOLIVIA — IV ROBERT C. FOSTER Continued from Vol. 68, No. 774 29. Dissanthelium Trin. Annual or perennial, the plants seldom over 10 cm. high, caespitose. Leaves glabrous, the blades flat or involute. Inflorescence a short, terminal, racemose panicle, spicate in appearance. Spikelets mostly 2-flowered, occasionally 3-flowered or 1-flowered, the naked rachilla disarticulating above the glumes and prolonged beyond the uppermost floret; glumes equal, longer than the lemmas; glumes and lemmas awnless. a. Glumes 3 (-3.5) mm. long. b. Lemma puberulent, obtuse, with a central muero, ................... uc CREER RR UE e 1. D. macusamiense. b. Lemma scabridulous, obtuse, but not toothed. ...... 2. D. calycinum. a. Glumes 5 mm. long. c. Inflorescence 5-7 mm. broad, compact; spikelets numerous. ........ EGG o TEC EHE PRACT IE E OE indie 3. D. peruvianum. c. Inflorescence less than 5 mm. broad, lax, spikelets few. ................ ARE RNR T E P PRU E rero nen 4. D. Trollit. 1. Dissanthelium macusaniense (Krause) Foster and Smith in Phyto- logia, 12: 249 (1965). D. minimum Pilger in Engler, Bot. Jahrb. 56, Beibl. 123: 28 (1920). Annual, the leaf-blades flat. Inflorescence 1-2 cm. long, more or less oblong, the rachis, branches and pedicels only faintly scabridulous. Spikelets mostly 2-flowered. Lemma about 2.5 mm. long. Potosi: cERCADO: Potosí, 4000 m., Cárdenas 198 (GH). 2. D. calycinum (Presl) Hitche. in Journ. Wash. Acad. Sci. 13: 224 (1923). Perennial. Inflorescence to 3 em. long, oval, rachis, branches and pedicels puberulent-scabridulous. Spikelets 2-flowered. Lemma about 2.5 mm. long. Without locality: Bang 1873 (GH). 3. D. peruvianum (Nees & Meyen) Pilger in Engler, Bot. Jahrb. 37: 241 242 Rhodora [Vol. 68 878 (1906). Perennial. Apparently differs from D. calycinum pri- marily in having longer glumes and in spikelets frequently 1-flowered. LA PAZ: LARECAJA: Sorata, Mandon 1345 (GH). 4. D. Trollii Pilger in Notizbl. 11: 778 (1933). Perennial, about 5 cm, tall. Leaves with flat pungent blades to 4.5 em. long (usually shorter) and 2 mm. wide. Inflorescence narrow, about 2 em. long (2.5 cm. on the type), the rachis glabrous to scabridulous, pedicels scabri- dulous, sometimes almost glabrous, or scabrid on the margins and angles. Spikelet to 5 mm, long, 2-flowered. Glumes as long as the spikelet, acute to very acute. Lemmas 4 mm. long, hyaline. Palea hya- line, nearly as long as the lemma, bicarinate, slightly bidentate, the teeth obtuse (when flattened). L4 PAZ: MURILLO: Chacaltaya, 4800 m., Buchtien 1197 (us); La Cumbre, 5000 m., Tate 10 (us), Parodi 10042a (US); La Union, 4800 m., Troll 1966 (US; type-number). 30. Koeleria Pers. Koeleria cristata (L.) Pers. Synops. Pl. 1: 97 (1805). K. gracilis subsp. boliviensis (Domin) Domin in Biblioth. Bot, 65: 237 (1907). K. pseudocristata var. andicola Domin in Fedde, Rep. Sp. Nov. 2: 94 (1906). Perennial, to 5 dm. high, the culms pubescent. Leaves mostly basal, the sheaths from nearly glabrous to pubescent, the flat blades pubescent, at least when young. Inflorescence a dense, terminal, race- mose panicle, to 10 cm. long, interrupted at the base, the rachis, branches and pedicels pubescent, Spikelets 2-4-flowered, about 6 mm. long, the rachilla villous, disarticulating above the glumes. First glume 4.5 mm. long, 1-nerved, scabridulous on the nerve; second glume 5 mm. long, 3-nerved, the keel scabridulous. Lowest lemmas 5 mm. long, bidentate, with a 4 mm. awn arising from the base of the sinus, the keel and awn scabridulous, Palea about 4 mm. long, biden- tate, the keels ciliolate. L4 PAZ: NOR YUNGAS: Pongo, 3640 m., Hitch- cock 22767 (GH). With the form described here, there is also to be found one in which leaves and culms are more densely pubescent and the lemmas are awnless or, at most, shortly mucronate. Both forms are found on the Gray Herbarium sheet of Hitchcock 22767. 31. Trisetum Pers. Perennials, 1-5 dm. high, rarely to 1 m., the leaves and culms glab- rous, the leaf-blades involute or flat and sometimes scabridulous. In- florescence a terminal, dense, racemose panicle, spicate in appearance, interrupted at the base, 2-15 cm. long. Spikelets 2-3-flowered, uni- lateral, fertile florets below the sterile, disarticulating above the unequal glumes; lemmas awned. 1966] Flora of Bolivia — Foster 243 Glumes to 1 cm. long, acuminate, lemmas to 1.5 cm. long, rachilla PIADEOUIS 51 E A ee aaae de 1. T. floribundum. Glumes to 5 mm. long, acute, lemmas about 5 mm. long or less, MACH la villõüs. e aa a a e e iae 2. T. spicatum. 1. Trisetum floribundum Pilger in Engler, Bot. Jahrb. 37: 505 (1906). Dielsiochloa floribunda of the Catalogue. Bromus Mandonianus of the Catalogue. To 3 dm. high, leaf-blades involute. Panicle to 6 cm. long, more or less oval, branches and pedicels scabridulous. Spikelets (excluding awns) about 1.3 em. long. First glume to 7 mm. long, second to 1 cm. Lemmas deeply bidentate, the narrow teeth resembling awns, the true awn to 1.5 em. long. Palea to 1 cm. long, bidentate, the keels ciliolate. LA PAZ: LARECAJA: vic. Sorata; ad lacum Turi- guana, 4500 m., Mandon 1371 (GH; type-number of Bromus Mandoni- anus). 2. T. spicatum (L.) Richt. Pl. Eur. 1: 59 (1890). T. oreophilum Louis-Marie in Rhodora, 30: 221 (1928). To 3 dm. high, rarely to T m., leaf-blades flat, leaves chiefly basal. Panicle narrow, to 15 em. long, rachis, branches and pedicels pubescent. Glumes 3.5 and 4 mm. long, the keels scabridulous, Lemmas pubescent to villous, bidentate, the teeth not produced as pseudo-awns, the true awn about 3 mm. long. LA PAZ: MURILLO: La Paz, 3700 m., Buchtien 8540 (GH). 32. Deschampsia Beauv. See Parodi in Darwiniana, 8: 415-475 (1949). Perennial. Leaf-blades flat or folded. Inflorescence a dense to open racemose panicle, sometimes interrupted below. Spikelets 2-flowered, the villous rachilla prolonged beyond the upper floret, disarticulation above the glumes. Glumes equal or subequal, Lemmas thin to hyaline, the apex obtuse, 2-4-toothed, bearing a dorsal awn at or below the middle. Panicle dense, the short branches appressed, interrupted below; awn much exceeding the lemma. .........., HH 1. D. conferta. Panicle lax, the branches long, not appressed; awn equaling or slightly exceeding the lemma. ....... ee 2. D. caespitosa. 1. Deschampsia conferta (Pilger) Valencia in Revist. Argent. Agron. 8: 127 (1941). Aira conferta of the Catalogue. Plant to 10 dm. tall, usually shorter. Leaf-blades linear, flat, scabrid above and beneath, to 3 mm. wide. Panicle dense, to 20 cm. long, rachis glabrous or slightly scabridulous on the upper portion, branches and pedicels scabridulous. Spikelets to 5 mm. long (excluding the awns). Glumes to 5 mm. long, acute, the margins erosulous near the apex, the keel scabridulous. Lemmas about 3 mm. long, deeply bidentate, the teeth more or less acute; awn twisted for 1.5-2 mm., genieulate, scabrid, the whole to 5 mm. long. COCHABAMBA: CERCADO: north of Cocha- bamba, Hitchcock 22814 (US). 244 Rhodora [Vol. 68 2. D. caespitosa (L.) Beauv. Ess. Agrost. 91, 149, 160 (1812). Plant to 1.2 m. tall, densely caespitose, Leaf-sheaths glabrous; blades linear, to 4 mm. wide, flat or folded, scabrid above. Inflorescence an open panicle, to 3 dm. long, the branches rather long, branches and pedicels scabridulous. Spikelets to 4 mm. long, the rachilla sometimes bearing a reduced floret at the apex. Glumes subequal, about 4 mm. long, obtuse and erose to acute, often purple-tinged, thin, obscurely nerved. Lemma 3 mm. long, hyaline, the apex usually several-toothed, the awn equaling or somewhat exceeding the lemma, but not markedly so; palea narrow, slightly shorter than the lemma. No Bolivian material has been seen. The species is in- cluded on the authority of Parodi, l. c., who cites a specimen (collector unknown) from the vicinity of the city of Santa Cruz de la Sierra. 33. Avena L. Annuals up to 7.5 dm. high, the leaf-blades flat, to 4-8 mm. wide, scabrid, occasionally pilose. Panicle lax, the branches distant, race- mose, to 25 em. long, rachis and branches glabrous, the pedicels some- times puberulous-scabridulous. Spikelets 2-3-flowered, the rachilla villous, disarticulating above the nearly equal, many-nerved, papery glumes; lemmas bidentate, villous on the lower portion at least, awned, the awn arising from mid-lemma or below. Pedicels spreading, glumes to 2.5 cm. long, lemmas about 2 em. long, ACUTE. RRRRRRRRRRRRNENEMMMMMMM 1. A. fatua. Pedicels flexuous, glumes to 2 cm. long, lemmas about 1.5 cm. long, the teeth setaceous. wcccccccccccccccsccccsesecssscssscesseceseseeesees 2. A. barbata. 1. Avena fatua L. Sp. Pl. 80 (1753). Spikelets (excluding awns) about 2.5 cm. long. Awn to 3.5 em. long. COCHABAMBA: CERCADO: Cochabamba, Hitchcock 22836 (Hitchcock). 2. A. barbata Brot. Fl. Lusit. 1: 108 (1804). Spikelets (excluding awns) about 1.5-2 em. long. Awns to 3 em, long. LA PAZ: MURILLO: Palea, Pflanz 2541 (Hitchcock). 34. Holcus L. Holcus lanatus L. Sp. Pl. 1048 (1753). Perennial, to 1 m. high. Leaf-sheaths, blades and culms pubescent, the blades flat, about equal to the base of the mature inflorescence, to 6 mm, wide. Inflorescence a terminal, racemose, dense panicle to 10 em. long, the rachis, branches and pedicels pubescent. Spikelets 2-flowered, the lower floret perfect, the upper staminate, 1 spikelet sessile, 1 pedicellate, the rachilla glab- rous, disarticulation above the equal glumes, these to 3.5 mm. long, exceeding the florets, the first obscurely 3-nerved, the keel and upper margins ciliate, the second 3-nerved, broader, keel and margins ciliate, 1966] Flora of Bolivia — Foster 245 both glumes mucronulate. Pistillate lemmas 2 mm. long, nerveless, awnless, obtuse, shining, the palea as long as its lemma, the keels ciliolate. Staminate lemmas 2 mm. long, nerveless, with a 1.5 mm. stout hooked awn at apex, the palea shorter than its lemma, the keels ciliolate. LA PAZ:MURILLO: La Paz, 3300 m., Bang 155 (GH). COCHA- RAMBA: TOTORA: Pocona, 2500 m., Steinbach 8671 (GH). 35. Danthonia Lam. & DC. Danthonia cirrhata Hack. & Arech. in Arech. Gram. Uruguay, 307 (1894). Perennial, to 6 dm. tall, nodes glabrous. Leaves sparsely papillate-hirsute on sheaths and blades, the narrow blades involute, seldom equaling the inflorescence. Inflorescence a narrow panicle usually less than 10 cm. long, few-flowered. Spikelets several-flowered, the uppermost floret sterile; disarticulation above the glumes; axis glabrous to puberulent, the pedicels puberulent and often pilose at the base. First glume to 1.5 em. long, long-acute, glabrous, 5-nerved, the central nerve prominent, margins thin to hyaline; second glume similar, about 2 mm. shorter. Rachilla long-plumose at the base, hairs to 5 mm. long, white. Fertile lemmas to 6 mm. long, pilose near margin below the middle, deeply bidentate, the teeth produced as pseudo-awns 5 mm. long, a true awn, twisted, to 1 cm. long, arising at base of sinus between teeth. Palea thin, to 4 mm. long, the 2 keels briefly ciliolate. SANTA CRUZ: VALLEGRANDE: Samaipata, Herzog 1706, 3012 (Henrard). 36. Lamprothyrsus Pilger Lamprothyrsus Hieronymi (O. Ktze.) Pilger in Engler, Bot. Jahrb. 37:58 (1906). L. Hieronymi var. tinctus Pilger in Engler, Bot. Jahrb. 37: 59 (1906). Perennial, to 2 m. high, leaf-sheaths glabrous, the blades long, involute, scabrid on the upper surface, often nearly equal- ing the mature inflorescence, the old sheaths ultimately coiled at base of plant. Inflorescence a terminal, racemose panicle to 30 em. long, fairly open at maturity, rachis glabrous, branches and pedicels often scabridulous. Spikelets (excluding awns and pseudo-awns) about 7 mm. long, with up to 6 florets, the uppermost reduced, disarti- culation above the glumes, rachilla villous. Glumes equal or subequal, to 6 mm. long, transparent, without apparent nerves. Body of lemma 3 mm. long, villous, bidentate, the teeth produced as pseudo-awns to 1.7 cm. long, the true awn from the base of the sinus, geniculate shortly above its base, to 2 cm, long. Palea 3 mm. long, keels ciliate, villous between the keels and the margins. La PAZ: LARECAJA: vie. Sorata; San Pedro, 2650 m., Mandon 1360 (GH). SANTA CRUZ: VALLE- GRANDE: Cerro La Negra, 1700 m., Steinbach 8171 (GH); SARA: Cerro Hosana, 1000 m., Steinbach 3453 (GH). TARIJA: ARCE: Bermejo, 1400 m., Fiebrig 2099 (GH; type-number of L. Hieronymi var. tinctus). 246 Rhodora [Vol. 68 27. Calamagrostis Adans. See Tovar in Mem. Mus. Hist. Nat. Javier Prado, no. 11: 3-88 (1960). Perennials, Leaves flat or involute. Inflorescence a panicle, often reduced and condensed. Spikelets disarticulating above the glumes, l-flowered, with the rachilla more or less prolonged as a bristle behind the palea. Glumes usually more or less equal. Lemma usually shorter than the glumes, thinner-textured, with a dorsal awn (1 species awn- less), callus more or less pilose or glabrous, the hairs of varying lengths. a. Lemmas awnless. a, 3. C. calderillensis. a. Lemmas awned. b. Lemmas stipitate below callus. c. Panicle very open, not Spicate. noscere 2. C. eminens. c. Panicle dense, spicate. d. Panicle elongate, slender; awn geniculate. .... 4. C. amoena. d. Panicle short, thick; awn straight. e. Awn shorter than the lemma. ................ 7. C. chrysantha. e. Awn exceeding the lemma, mostly by 1 mm. .......... eee DM 8. C. ovata b. Lemmas sessile below the callus. f. Rachilla glabrous or with a tuft of hairs at the apex only. £&. Rachilla glabrous. ............ ess 6. C. rigescens, g. Rachilla with an apical tuft of hairs. h. Rachilla 2 mm. long; awn 9 mm. long. .......eeeeseeeeerereneen REN 27. C, longiaristata. h. Rachilla 1 mm, long; awn 6 mm. long. ......... esee RR 30. C. viridiflavescens. f. Rachilla villous. i. Awn 2-2.5 mm. long. j- Spikelet 3 mm. long. .......... ee 29. C. polygama. j. Spikelet 5-8 mm. long. k. Spikelet to 5 mm, long. .......... 13. C. curvula. k. Spikelet to 8 mm. long. .......... 31. C. spicigera. i, Awn 4 mm. or more in length. l. Awn less than, equaling, or only slightly exceeding the lemma, m. Inflorescence narrow, dense. ............ 10. C, eryptolopha. m. Inflorescence lax. n. Rachilla 1.5 mm. long, the longest hairs to 1.5 mm. LONG. RM 19. C. tarijensis. n. Rachilla 2-2.5 mm. long, the longest hairs to 4 mm. long... treten 23. C. intermedia. l. Awn distinctly exceeding the lemma. o. Inflorescence lax, sometimes spreading. 1966] Flora of Bolivia — Foster 247 p. Panicle about 1 dm. long. ..........-.- 18. C. tarmensis. p. Panicle 2 dm. or more in length. q. Lowest branches of panicle naked for 2-5 cm. r. Spikelets to 7-8 mm. long; rachilla-hairs to 2 mm. long. ..........e eere rne a tn aanonunuo 1. C. leiophylla. r. Spikelets to 5-6 mm. long; rachilla-hairs to 4 mm. longe e bocce tsicsusentevseccessasteamtuecs 22. C, Orbignyana. q. Lowest branches of panicle spikelet-bearing from nearly the base. s. Rachilla-hairs to 1.5 mm long. ........ 26. C. recta. s, Rachilla-hairs to 4 mm. long. .... 21. C. antoniana, o. Inflorescence dense, sometimes interrupted, but not spreading. t. Panicle over 10 cm. long. u. Spikelets 4 (-4.5) mm. long. ... 5. C. heterophylla. u. Spikelets 6-8 mm. long. v. Spikelets to 6 mm. long; callus glabrous or nearly so, if a few hairs present, the longest to 0.4 mm. long. ....... reete ene tnn nonne ennt 28. C. densiflora. v. Spikelets 8 mm. long; callus somewhat villous, the longest hairs to 1 mm. long. ....... m OST eel ea PEe REFERRE AVES ANN EEERSR E 11. C. glacialis. t. Panicle less than 10 cm. long. w. Panicle less than 2 cm. long. x. Inflorescence narrow, spicate. .... 24. C. minima. x. Inflorescence dense, ovoid. .............. 15, C. curta. w. Panicle 2 cm. or more in length. y. Panicle not interrupted, or only slightly so at the base. z. Pedicels villous. ...................--- 12. C. Jamesonii. z. Pedicels scabridulous to pubescent, or nearly glabrous. aa. Callus densely villous, the longest hairs 1 mm. long. seeen 16. C. brevifolia. aa. Callus sparsely short-villous, the longest hairs 0.5 mm. long. ........ 9. C. vicunarum. y. Fanicle obviously much interrupted, bb. Panicle to 2.5 cm. long. .......... HA dansesyunnawssesrdhshscenans puscntesensesqersane 14. C. cephalopora. bb. Panicle 4-9 cm. long. ec. Awn to 7 mm. long. ......... 20. C. Fiebrigü. cc. Awn to 4 mm. long. dd. Rachis, branches and pedicels glabrous. br Eben qme 17. C. wiolacea. dd. Rachis, branches and pedicels sparsely géabrid, oae 25. C. trichophylla. 248 Rhodora [Vol. 68 1. Calamagrostis leiophylla (Wedd.) Hitche. in Contrib. U. S. Nat. Herb. 24: 367 (1927). C. toluccensis var. laxiflora O, Ktze. Rev. Gen. 3(2): 345 (1898). Caespitose, erect, 0.5-1 m. high. Leaf-sheaths glabrous to scabridulous, ligule less than 1 mm. long; leaf-blades flat to subinvolute, to 2.5-3 mm. wide, Inflorescence to 3 dm. long, narrow but lax, at least the lowermost branches whorled, axis glabrous below to sparsely scabridulous above, the branches, branchlets and pedicels scabrid. Spikelets to 5-8 mm. long. Glumes to 5-8 mm. long, acute, obscurely nerved, the keel prominent, scabrid. Lemma sessile below the callus, about 2 mm. shorter than the glumes, bidentate, scabri- dulous; callus glabrous to sparsely short-villous, rachilla about 2 mm. long, the longest hairs about 2 mm. long, not quite equaling the lemma; awn geniculate, to 5-6 mm. long, inserted at the middle of the lemma and exceeding it. COCHABAMBA: CERCADO: Cochabamba, Holwa 379 (us). 2. C. eminens (Presl) Steud. Nomencl. (ed. 2) 1: 250 (1840). C. elegans (Wedd.) Henrard in Meded. Rijks Herb. Leiden, no. 40: 61 (1921). Deyeuxia polystachya Wedd. in Bull. Soc. Bot. France, 22: 177, 178 (1875). C. eminens var. tunarieasis O. Ktze. Rev, Gen. 3(2): 344 (1898). C. eminens var. sordida O. Ktze. Rev. Gen. 3(2): 344 (1898). Caespitose, somewhat rhizomatous, stout, to 1 m. high, in damp or wet places. Leaf-sheaths glabrous, ligule to 1.8 cm.; leaf- blades involute, finely scabridulous to glabrous, Inflorescence open, to 3 dm. long, the spikelets glomerulate or subglomerulate at the ends of branches and branchlets, the rachis, whorled branches and pedicels glabrous to scabridulous. Spikelets to 4-5.5 mm. long. Glumes to 4-5.5 mm. long, acute and erosulous to subobtuse, the keel faintly prominent. Lemma distinctly stipitate below the callus, about 1 mm. shorter than the glumes, obtuse or truncate and erosulous at the apex; callus densely villous, with hairs up to 3 mm. long; rachilla about 1.5 mm. long, long-villous, the hairs chiefly basal; awn delicate, straight, equaling or shorter than the lemma, inserted below the middle of the lemma. La PAZ: LARECAJA: vic. Sorata; Chicani, 3600-3800 m., Mandon 1309 (GH), 1310 (GH; co-types of Deyeuxia elegans). COCHA- BAMBA: ARANI: pass, Cordillera de Cochabamba, 4500 m., Eyerdam 24850 (GH). 3. C. calderillensis Pilger in Engler, Bot. Jahrb. 42: 72 (1908). Plant to 13 dm. tall. Leaf-sheaths glabrous, ligule about 3 mm. long; leaf-blades flat to somewhat inrolled, to 7 mm, wide, the margins scabrid, otherwise glabrous above and beneath. Inflorescence to 22 cm. long, the rachis, whorled branches and pedicels glabrous. Spikelets to 4 mm. long. Glumes to 4 mm. long, the second glume longer than the first, acute to subobtuse. Lemma as long as the first glume, sessile below the callus, obtuse or truncate; callus glabrous, with an apical ring of short, sparse, fine hairs; rachilla 0.5 mm. long, sparsely villous, the longest hairs less than 1 mm. long, not reaching the middle 1966] Flora of Bolivia — Foster 249 of the lemma; awn lacking. TARIJA: CERCADO: Calderillo, 3300 m., Fiebrig 3172 (US; type-number). 4. C. amoena (Pilger) Pilger in Engler, Bot. Jahrb. 42: 60 (1908). C. filifolia (Wedd.) Pilger in Engler, Bot. Jahrb. 42: 67 (1908). To 6 dm. high. Leaf-sheaths glabrous, ligule to 3 mm. long; leaf-blades involute, filiform, minutely scabridulous. Inflorescence 6-15 cm. long, somewhat lax, rachis glabrous below, scabrid above, the whorled branches, branchlets and pedicels scabridulous to scabrid. Spikelets to 5-5.5 mm. long. Glumes to 5-5.5 mm. long, long-acute, entire, the faintly prominent midrib keeled and scabridulous above, Lemma shortly stipitate below the callus, as long as the first glume, scabridulous, with 3-4 very slender, short, apical teeth; callus villous at the apex, the longest hairs 1 mm. long; rachilla to 2 mm. long, the upper portion bristle-like, moderately villous, the longest hairs to 1.5 mm. long, reaching 2/3 the length of the lemma; awn geniculate, to 4.5 mm. long, much exceeding the lemma, the base twisted, the upper portion straight, inserted well below the middle of the lemma. LA PAZ: LARECAJA: vic. Sorata; Marcamocreani, 3500-3600 m., Mandon 1306 (GH); NOR YUNGAS: Milluguaya, 2700-3900 m., Mandon 1301 (GH); MURILLO: Talca Chuquiaguilla, Bang 805 (GH; type-number). 5. C. heterophylla (Wedd.) Pilger in Engler, Bot. Jahrb. 42: 64 (1908). C. boliviensis Hack. in Fedde, Rep. Sp. Nov. 6: 156 (1908). Plants to 8 dm. tall. Leaf-sheaths usually glabrous, the ligule to 5 mm. long; blades linear, flat, about 2-4 mm. wide, usually shorter than the inflorescence, glabrous to scabridulous or somewhat short-pilose above and beneath, at least when young. Panicles to 2 dm. long, narrow, dense, interrupted (mostly on the lower half), rachis and branches glabrous, pedicels scabrid, the branches short. Glumes to 4 (-4.5) mm. long, acute, the narrow keel scabridulous. Lemma ses- sile below the callus, about 0.5-1 mm. shorter than the longer glume, with 3 slender apical teeth, the lateral teeth shorter than the central tooth; callus sparsely short-villous, the hairs not over 0.5 mm. long; rachilla 0.5-1 mm. long, sparsely villous, the longest hairs not over 1 mm. long, about 1/3 the lemma-length; awn geniculate, to 5.5 mm. long, loosely twisted at the base, inserted at or below the middle of the lemma. LA Paz: MURILLO: La Paz, 3700 m., Buchtien 8834 (GH). 6. C. rigescens (Presl) Scribner in Rept. Missouri Bot. Gard. 10: 37 (1899). Deyeuxia imberbis Wedd. in Bull. Soc. Bot. France, 22: 177, 180 (1875). Caespitose, rhizomatous, to 3 dm. tall. Leaf-sheaths glabrous, ligule to 1 mm. long; blades involute, shorter than inflores- cence, glabrous to somewhat scabridulous, Inflorescence to 1 dm. long, narrow, the branches short, interrupted below, rachis usually glabrous below, puberulent-scabridulous above, branches and pedicels puberu- lent-scabridulous. Glumes to 5 mm. long, acute, the green midrib keeled and scabridulous above. Lemma about 0.5 mm, shorter than glumes, sessile below the callus, with 3-4 fine, short, apical teeth; 250 Rhodora [Vol. 68 callus sparsely short-villous, the hairs less than 0.5 mm. long; rachilla glabrous, to nearly 2 mm. long; awn straight, to 4-4.5 mm. long, inserted at the middle of the lemma. LA Paz: MURILLO: near La Paz, 3300 m., Bang 104 (GH). C. chrysantha (Presl) Steud. Nomencl. (ed. 2) 1: 250 (1840). Deyeuxia phalaroides Wedd. in Bull. Soc. Bot. France, 22: 177, 180 (1875). D. leiopoda Wedd. in Bull. Soc. Bot. France, 22: 177, 180 (1875). Stylagrostis leiopoda of the Catalogue. More or less rhizom- atous, to 5 dm. tall. Leaf-sheaths glabrous, the ligule to 1.8 cm. long; blades involute, the apex pungent, much shorter than the inflo- rescence, glabrous, or scabridulous near the apex. Inflorescence golden- brown, oblong, to 6 em. long, dense, the very short branches ascending, rachis, branches and pedicels mostly glabrous. Glumes to 7 (rarely 8) mm. long, acute or shortly bidentate at the apex, the midrib not promi- nent. Lemma stipitate below the callus, about 1-1.5 mm, shorter than the longer glume, 2-4-toothed; callus densely villous, the longest hairs 4-5 mm. long; rachilla to 1.5 mm. long, densely villous, the longest hairs to 5 mm. long; awn straight, 2-3 mm. long, shorter than the lemma, quickly deciduous, inserted about 1 mm. above the base of the lemma. LA Paz: MURILLO: near La Paz, on the road to Coroico, 5000 m., Mandon 1319 (GH), 8. C. ovata (Presl) Steud. Nomencl. (ed. 2) 1: 254 (1840). Dey- euxia nivalis Wedd. in Bull. Soc. Bot. France, 22: 176, 180 (1875). D. anthoxanthum Wedd. in Bull. Soc. Bot. France, 22: 176, 180 (1875). D. capitata Wedd. in Bull. Soc. Bot. France, 22: 176, 180 (1875). C. Pflanzii Pilger in Engler, Bot. Jahrb. 49: 184 (1912). Caespitose, to 3 dm. high, Leaf-sheaths glabrous, the ligule to 1.5 em. long; blades flat to subinvolute, much shorter than the inflorescence. Panicle golden- brown, to 6-7 em. long, dense, ovate, the branches short, ascending, rachis and branches mostly glabrous, pedicels sparsely puberulous. Glumes to 1 cm. long (rarely longer), long-acute, the midrib mod- erately prominent, often scabridulous. Lemma about 4 mm. long, stipitate below the callus, rather deeply bidentate; callus densely villous, the longest hairs 3-4 mm. long; rachilla to nearly 1.5 mm. long, glabrous below, the apical hairs to 3 mm. long; awn straight, mostly exceeding the lemma by about 1 mm., inserted somewhat above the base of the lemma. LA PAZ: LARECAJA: vic. Sorata; Anilaya, 3800-4500 m., Mandon 1320 (GH); MURILLO: near La Paz, on the road to Coroico, 5000 m., Mandon 1318 (US; type-number of Deyeuxia nivalis); NOR YUNGAS: at top of La Cumbre, 4500 m., Eyerdam 25112 (GH). 9. C. vieunarum (Wedd.) Pilger in Engler, Bot. Jahrb. 42: 62 (1908). C. pentapogonoides O. Ktze. Rev. Gen, 3(2): 344 (1898). Plants 5-25 cm. tall, caespitose. Leaf-sheaths glabrous, the ligule to 2 mm. long; blades pungent, filiform, involute, curled or flexuose, shorter (usually) than the inflorescence, glabrous above, the margins 1966] Flora of Bolivia — Foster 251 scabrid, puberulous to scabridulous beneath, ultimately glabrescent below. Inflorescence oblong, spicate in appearance, occasionally inter- rupted below, 2-3 cm. long, the very short branches ascending, ap- pressed, rachis, branches and pedicels rather sparsely scabridulous. Spikelets to 7 mm. long. Glumes as long as the spikelet, long-acute, scabridulous, especially near the upper portion of the scabridulous keel. Lemma sessile below the callus, 2-3 mm. shorter than the glumes, with 3-4 slender apical teeth; callus sparsely-villous, the longest hairs about 0.5 mm. long; rachilla 0.5-0.75 mm. long, slightly villous, the longest hairs about 1 mm. long; awn geniculate, basally twisted, about 5-6 mm. long, inserted 1 mm. above the base of the lemma, LA Paz: MURILLO: La Paz, 4100 m., Buchtien (GH). COCHABAMBA: CERCADO: Putuyuni, 4000 m., Steinbach 4086 (GH). 10. C. eryptolopha (Wedd.) Hitchc. in Contrib. U. S. Nat. Herb. 24: 374 (1927). To 8 dm. tall. Leaf-sheaths glabrous, the ligule 2-4 mm. long; blades mostly shorter than the inflorescence, nearly filiform, involute, scabridulous, Inflorescence narrow, dense, to 15 cm. long, sometimes interrupted below, the short (to 5 cm.) ascending branches appressed, rachis, branches and pedicels scabridulous. Glumes to about 6 mm. long, long-acute, scabridulous on and near the subprominent keel. Lemma sessile below the callus, about 1-1.5 mm, shorter than the glumes, shortly bidentate; callus rather sparsely villous, the longest hairs about 1 mm. long; rachilla about 1 mm. long, densely villous, the longest hairs about 2 mm, long; awn geniculate, the lower half loosely twisted, 4-5 mm. long, inserted about 1.5 mm. above the base of the lemma. La Paz: LARECAJA: Mt. Illampu, 3800 m., Mandon 1313 (GH; type-number). 11. C. glacialis (Wedd.) Hitche. in Contrib. U. S, Nat. Herb. 24: 375 (1927). Deyeuxia picta Wedd. in Bull. Soc. Bot. France, 22: 177, 180 (1875). Height 3-5 dm. Leaf-sheaths scabridulous, ligule to 6 mm. long; blades shorter than or equaling the inflorescence, involute, stiff, scabridulous to glabrous. Inflorescence tawny-purple, to 20 cm. long, spicate in appearance, the rachis mostly glabrous, branches and pedicels sparingly scabrid cn the angles. Glumes to 8 mm. long, acute to long-acute, glabrous to scabridulous, especially on the subpromi- nent keel. Lemma to about 6 mm. long, sessile below the callus, deeply bidentate; callus rather sparsely villous, the longest hairs about 1 mm. long; rachilla about 2 mm. long, densely villous, the longest hairs about 4 mm. long, equaling the lemma; awn about 4 mm. long, straight, exceeding the lemma, inserted at or slightly below the middle of the lemma. LA PAZ: LARECAJA: vic. Sorata; Millipaya, 5000 m., Mandon 1312 (GH; type-number) ; MURILLO: Cerro de Huaina Potosí. Germann 1 (US); Illimani, 4850 m., Hitchcock 22590 (GH). 12. C. Jamesonii Steud. Syn, Pl. Glum, 1: 191 (1854). Plant 7-35 em. high, somewhat rhizomatous. Leaf-sheaths glabrous, the ligule to 2 mm. long; blades involute, shorter than or equaling the in- 252 Rhodora [Vol. 68 florescence, obtuse to subacute, glabrous to puberulous. Inflorescence more or less ovoid, dense, 2-4 cm. long, occasionally somewhat inter- rupted below, the rachis, very short branches and pedicels villous to pubescent. Glumes to 6 mm. long, acute to subacute, glabrous to puberulent, the rather sharp keel pubescent-ciliate. Lemma sessile below the callus, about 4 mm. long, deeply bidentate, the teeth entire or toothed; callus very sparsely short-villous, the longest hairs less than 1 mm. long; rachilla 1.5 (-2) mm. long, densely long-villous, the longest hairs to 4 mm. long, equaling the lemma; awn straight, to 4 mm. long, much exceeding the lemma, inserted well below the middle of the lemma. La PAZ: MURILLO: Chacaltaya, 4800 m., Buchtien 1193 (vs). This description is based on four topotypes, one of which is a probable isotype. 13. C. curvula (Wedd.) Pilger in Engler, Bot. Jahrb, 42: 60 (1908). C. variegata O. Ktze. Rev. Gen. 3 (2): 354 (1898). Densely caespitose rhizomatous plants, to 3 dm. high. Leaf-sheaths glabrous, the ligule to 2 mm. long; blades involute, pungent, filiform, arcuate, scabridulous. Inflorescence spicate in appearance, oblong, to 4.5 cm, long (rarely to 7 cm.), the short branches ascending, rachis and branches glabrous, the pedicels glabrous to sparsely puberulent. Glumes to 5 mm. long, acute, glabrous, the keels scabridulous, Lemma sessile below the callus, about 3.5 mm. long, bidentate, the teeth broad and erosulous; callus villous, the longest hairs about 1 mm. long; rachilla about 2 mm. long or slightly longer, the longest hairs equaling or exceeding the lemma; awn straight to flexuose, but not geniculate, about 2 mm. long, equaling the lemma, inserted at the middle of the lemma or lower. POTOSÍ: SUR CHICHAS: Atocha, Hitchcock 22873 (GH). 14. C. cephalantha Filger in Engler, Bot. Jahrb. 42: 61 (1908). Deyeuxia Lagurus Wedd. in Bull. Soc. Bot. France, 22: 176, 180 (1875). Dwarf caespitose plants, usually to 4 em. high, occasionally to 15 em. high. Leaf-sheaths glabrous, inflated, the ligule to 2.5 mm. long; blades involute, 8 mm. to 2 em. long, pubescent. Inflorescence ovate, to 2.5 em. long, the very short branches ascending, rachis, branches and pedicels puberulent-scabridulous. Glumes to about 6 mm. long, acute, glabrous to puberulous above, Lemma sessile below the callus, to 4.5 mm. long, the apex several-toothed; callus densely villous, the longest hairs to 1 mm. long; rachilla to 3 mm. long, densely villous, the longest hairs to 4 mm. long, much exceeding the lemma; awn geniculate, the lower half twisted, to 6.5 mm. long, inserted some- what below the middle of the lemma. La PAZ: MURILLO: Chacaltaya, 4800 m., Buchtien 1195 (us). 15. €. curta (Wedd.) Hitche. in Contrib. U. S. Nat. Herb. 24: 376 (1927). Perennial, probably less than 10 cm. tall. Leaf-sheaths glabrous; blades glabrous, to 2.5 em. long and less than 1 mm. wide, 1966] Flora of Bolivia — Foster 253 flat or semi-involute, obtuse. Inflorescence a short, condensed, ovate panicle, less than 2 cm. long, the branchlets and pedicels puberulent. Glumes 6 mm. long, the midrib carinate and ciliate, upper portion of glumes .scabridulous. Lemma nearly 4 mm. long, the apex with 4 short fine teeth; callus sparsely short-villous, the hairs about 0.5 mm. long; rachilla to nearly 2 mm. long, villous, the hairs about 1 mm. long; awn 4 mm. long, geniculate, inserted below the middle of the lemma and exceeding it. LA PAZ: LARECAJA: vic. Sorata, 4200 m., Mandon 1316 (US; type-number). 16. C. brevifolia (Presl) Steud. Nomencl. (ed. 2) 1: 240 (1840). Caespitose cushion-plants, the culms 2-4 dm. tall. Leaf-sheaths gla- brous, the ligule to 2 mm, long; blades involute, filiform, arcuate or flexuose, pungent, scabridulous above, glabrous beneath, 2-10 cm. long. Inflorescence somewhat dense, oblong to oblong-ellipsoid, 3-7 cm, long, the short branches ascending-appressed or somewhat spreading at the base, the rachis usually glabrous, or sometimes sparsely puberulous or scabridulous, branches and pedicels glabrous to pubescent, Glumes to 7 mm. long, acute to subobtuse, the keel subprominent and scabri- dulous above. Lemma sessile below the callus, to 5 mm. long, the apex 3-4-toothed, the teeth short, delicate; callus densely villous, the longest hairs about 1 mm. long; rachilla to 3 mm. long, densely villous, the longest hairs to 4 mm. long, equaling or slightly exceeding the lemma; awn straight or flexuose, to 4 mm. long, slightly exceeding the lemma, inserted about 1.5-2 mm. above the base of the lemma. LA Paz: MURILLO: La Paz, Asplund 6519 (Hitchcock). 17. €. violacea (Wedd.) Hack. ex Buchtien, Contrib. Fl. Bolivia, 1: 75 (1910). Caespitose, to 3 dm. tall. Leaf-sheaths puberulent- scabridulous, at least when young, the ligule acute, to 4 mm. long; blades filiform, involute, pungent, arcuate, scabridulous-puberulent, shorter than the inflorescence. Inflorescence rather open, to 6 cm. long, the branches 1.5-2.5 cm. long, rachis, branches and pedicels glabrous. Glumes to 6 mm. long, acute to shortly setaceous, purplish dorsally, the keel prominent and scabridulous above. Lemma about 0.5-1 mm. shorter than the glumes, sessile below the callus, with 3-4 short setaceous teeth at the apex; callus densely villous, the longest hairs to 2 mm. long; rachilla densely villous, the longest hairs 3 mm. long, equaling the lemma; awn straight or flexuose, about 4 mm, long, somewhat exceeding the lemma, inserted above the middle of the lemma. LA PAZ: OMASUYOS: near Achacache, 3950 m., Mandon 1307 (GH), 18. C. tarmensis Filger in Engler, Bot. Jahrb. 42: 70 (1908). C. rosea var. macrochaeta Hack, in Fries in Ark. Bot. 8: 40 (1908). Plants caespitose in dense tufts or fans, to 8 dm. tall. Leaf-sheaths glabrous, the truncate ligule 2-3 mm. long, erose; blades involute, filiform, flexuose to arcuate, stiffly coriaceous, scabridulous, shorter than to equaling the inflorescence. Panicle lax, 4-11 em. long, narrow, 254 Rhodora [Vol. 68 the branches ascending and appressed to somewhat spreading, rachis, branches and pedicels scabridulous to scabrid. Glumes 5-6 mm. long, acute, the keel prominent and scabridulous above. Lemma sessile below the calllus, 4 (-5) mm. long, with 3-4 fine, short, apical teeth, sometimes scabridulous above; callus sparsely short-villous, the longest hairs about 0.8 mm. long; rachilla about 2 mm. long, rather sparsely villous, the longest hairs 1.5-2 mm. long; awn geniculate, the tightly twisted basal half not equaling nor exceeding the glumes, inserted about 1 mm. above the base of the lemma, at least 6 mm. long. COCHABAMBA: CERCADO: south side of Mt. Tunari, 3600 m., Steinbach 9783, 9785, 9788 (all GH), 19. C. tarijensis Pilger in Engler, Bot. Jahrb. 42: 71 (1908). Plants caespitose, to 9 dm. tall. Leaf-sheaths glabrous to scabridulous, the ligule to 2 mm. long, acute or subacute; blades involute, scabri- dulous. Panicle to 20 cm. long, lax, the branches to 5-6 cm. long, rachis, branches and pedicels scabridulous, Glumes about 5 mm. long, acute, the keel prominent and scabridulous above, Lemma sessile below the callus, about 4 mm. long, somewhat scabridulous above, with 4 short, slender, apical teeth; callus villous, the hairs to 1 mm. long; rachilla 1.5 mm. long, longest hairs to 1.5 mm.; awn geniculate, basal half twisted, equaling or exceeding the glumes, 6 mm. or more in length, inserted 1.5 mm. above the base of the lemma. COCHABAMBA: CERCADO: Mt. Tunari, 3600 m., Steinbach 9873 (GH). TARIJA: AVILES: Pinos, Fiebrig 3119 (GH). 20. C. Fiebrigii Pilger in Engler, Bot. Jahrb. 42: 68 (1908). Caespitose, to 3 dm. tall. Leaf-sheaths glabrous, the ligule to 4 mm. long, acute; blades involute, erect, finely scabridulous beneath. In- florescence 4-6 cm. long, rather dense, the short branches ascending, appressed, rachis and branches glabrous to scabridulous, the pedicels hispidulous. Glumes to 6 mm. long, acute, the keel scabridulous. Lemma sessile below the callus, to 5 mm. long, acute, shortly bidentate; callus villous, longest hairs 1 mm. long; rachilla 2.5 mm. long, villous, the longest hairs 3 mm. long, nearly equaling the lemma; awn genicu- late, to 7 mm. long, basal half tightly twisted, inserted about 1.5 mm. above the base of the lemma and much exceeding it. TARIJA: CERCADO: Calderillo, 3300 m., Fiebrig 3191 (US; type-number). 21. C. Antoniana (Griseb.) Steud. ex Hitche. in Contrib. U. S. Nat. Herb. 24: 378 (1927). Caespitose, somewhat rhizomatous bunch-grass, to 6-8 dm. high. Leaf-sheaths usually glabrous, sometimes scabridulous, the ligule to 12 mm. long, acute; blades nearly as long as the inflo- rescence, involute, stiff, pungent, more or less scabridulous, Inflo- rescence narrow but rather lax, to 2 dm. long (but usually shorter ?), the branches to 3-4 em. long, ascending-appressed to somewhat spread- ing, rachis and branches mostly glabrous, the pedicels sparsely scabrid- puberulent, especially near the apex. Glumes to 7 mm. long, acute to subacuminate, the keel seabridulous. Lemma sessile below the callus, 1966] Flora of Bolivia — Foster 255 about 5 mm. long, scabridulous, bidentate, the thin teeth shortly bidentate; callus villous, the longest hairs to 1 mm. long; rachilla 1-1.5 mm. long, villous, the longest hairs to 4 mm. long, distinctly exceeding the lemma; awn geniculate to flexuose, the basal portion loosely twisted, to 4-6 mm. long, much exceeding the lemma, inserted at or somewhat below its middle. LA PAZ: MURILLO: La Cumbre, Asplund 6514 (vus). 29. C. Orbignyana (Wedd.) Hitche. in Contrib. U. S. Nat. Herb. 24: 378 (1927). Caespitose, to 8 dm. tall. Leaf-sheaths glabrous, the ligule to 3 mm. long, acute to obtuse, emarginate to bidentate; blades filiform, involute, flexuose, much shorter than the inflorescence, scabri- dulous, the margins scabrid-ciliate. Inflorescence to 2 dm. long, narrow but lax, the branches to 4-5 cm, long, ascending to somewhat spreading, rachis and branches glabrous, the pedicels very sparsely scabrid to glabrous. Glumes about 5-6 mm. long, the keel scabridulous. Lemma sessile below the callus, 4-5 mm. long, very shortly 3-toothed ; callus villous, the longest hairs 1.5 mm. long; rachilla 2 mm. long, villous, the longest hairs to 4 mm. long, equaling or slightly exceeding the lemma; awn about 4 mm. long, somewhat geniculate, the lower half loosely twisted, inserted at or below the middle of the lemma and somewhat exceeding it. LA PAZ: MURILLO: La Paz, 3750 m., Buchtien 180 (GH). 23. C. intermedia (Presl) Steud. Nomencl. (ed. 2) 1: 250 (1840). C. Humboldtiana of the Catalogue. Deyeuxia gracilis Wedd. in Bull. Soc. Bot. France, 22: 179 (1875). D. Mandoniana Wedd. in Bull. Soc. Bot. France, 22: 179, 180 (1875). Caespitose, rhizomatous, 3-10 dm. high. Leaf-sheaths finely scabridulous to glabrous, the ligule to 1 em. long, apically narrowed and toothed; blades involute, stiff, coriaceous, usually about as long as the inflorescence, or at least reaching its base, scabridulous, pungent. Inflorescence to 30 cm. long, lax, the lowest branches sometimes to 8-10 em. long, ascending-spread- ing, somewhat interrupted, the rachis mostly glabrous, branches and pedicels quite sparsely scabrid. Glumes to 7 mm. long, acute to acuminate, the keels apically scabridulous. Lemma sessile below the callus, to 6 mm. long, usually densely scabridulous-puberulous, biden- tate, the slender teeth bidentate; callus villous, the longest hairs nearly 1 mm. long, rarely a little longer; rachilla 2-2.5 mm. long, villous, the longest hairs about 4 mm. long, equaling the lemma; awn geniculate, twisted, inserted well below the middle of the lemma and much exceeding it. LA PAZ: MURILLO: La Paz, 4100 m., Buchtien 123 (GH); Palca, 3800 m., Buch tien 122 (GH); LARECAJA: vic. Sorata; Velatulini, 3700 m., Mandon 1308 (GH; type-number of Deyeuxia Mandoniana). 24. C. minima (Pilger) Tovar in Mem. Hist. Nat. Javier Prado, no. 11: 52 (1960). Dwarf, pulvinate, to 6 cm. tall. Leaf-sheaths glabrous, the ligule less than 0.5 mm. long, apically truncate and 256 Rhodora [Vol. 68 ciliate, blades to 1 em. long, involute, the apex obtuse, margins scabrid- ciliate. Inflorescence to 1.5 em. long, narrow, spicate in appearance, few-flowered, the rachis, branches and pedicels glabrous to sparsely scabridulous. Glumes to 5.5 mm. long, acute, the upper Margins some- times erosulous, the keel scabridulous above. Lemma sessile below the callus, about 4 mm. long, scabridulous above, with several very fine apical teeth; callus villous, the longest hairs less than 1 mm. long; rachilla about 0.5 mm. long, sparsely villous, the longest hairs to 1 mm. long; awn nearly 5 mm, long, genieulate, inserted about 0.5 mm. above the base of the lemma and much exceeding it. La Paz: MURILLO: La Paz, 4200 m., Asplund 6518 (US); La Cumbre, 4650 m., Asplund 6511 (us); Chacaltaya, 4800 m., Buchtien 1202 (us). 25. C. trichophylla Pilger in Engler, Bot. Jahrb. 42: 67 (1908). Caespitose, rhizomatous, to 5 dm. high. Leaf-sheaths glabrous, the ligule to 2.5 mm. long, subobtuse; blades involute, filiform, flexuose to straight, margins scabridulous, subacute to obtuse, shorter than the inflorescence. Inflorescence lax, open, narrow, to 9 em, long, rachis, branches and pedicels sparsely scabrid. Glumes to 5 mm. long, acute, the keel scabridulous. Lemma sessile below the callus, about 3.5 mm. long, bidentate, the slender teeth bidentate; callus villous, the longest hairs about 1 mm. long; rachilla to 1.5 mm. long, sparsely villous, the longest hairs somewhat less than 1 mm. long; awn 4 mm. long, geni- culate, inserted less than 1 mm. above the base of the lemma and exceeding it. COCHABAMBA: CHAPARE: 50 km. northeast of Cocha- bamba, on road to Chimoré, 1700 m., Eyerdam 25038 (GH). 26. C. recta (HBK.) Trin. ex Steud. Nomencl. (ed. 2) 1: 251 (1840). Y Deyeuzia sulcata Wedd. in Bull. Soc. Bot. France, 22: 178, 180 (1875). Bunch-grass, to 9 dm. high. Leaf-sheaths glabrous or scabridulous near the apex, ligule to 5 mm. long, truncate, the apex erose; blades involute, stiff, obtuse, nearly equaling or exceeding the inflorescence, Panicle to 25 em. long, lax, interrupted, the lower branches to 7-8 em. long, rachis, branches and pedicels scabrid to hispidulous. Glumes to 7 mm. long, acute, the prominent keel scabridulous. Lemma sessile below the callus, about 5 mm. long, bidentate; callus villous, the longest hairs somewhat less than 1 mm. long; rachilla to 2 mm. long, the longest hairs about 1.5 mm. long; awn geniculate, to 6-7 mm. long, inserted 1.5-2 mm. above the base of the lemma and much exceeding it. LA PAZ: LARECAJA: vic. Sorata; Puerta del Inca, 3700 m., Mandon 1308 bis (GH; type-number of Deyeuxia sulcata). COCHA- BAMBA: CERCADO: Cuesta Tabacruz, Tunari, 4500 m., Steinbach 9789 (GH). 27. C. longiaristata (Wedd.) Hack. in Sodiro in Anal. Univ. Quito, 1889: 8 (1998). C. Beyrichiana of the Catalogue. C. montevidensis var. linearis Hack, in Fedde, Rep. Sp. Nov. 6: 156 (1908). Plant to 8 dm. tall. Leaf-sheaths glabrous, the ligule to 4 mm, long, truncate, laciniate-dentate; blades flat, to 20 em. long and 7 mm. wide, scabrid * pg(EO XI 1966] Flora of Bolivia — Foster 201 above and beneath. Inflorescence lax, open, to 2.5 dm. long, the lowest branches up to 15 cm. long, ascending-spreading to arcuate, rachis, branches and pedicels scabridulous, Glumes to 6 mm. long, very narrow, acute, the inconspicuous keel finely scabridulous. Lemma sessile below the callus, to 4.5 mm. long, acute, bidentate; callus densely villous, longest hairs as long as the lemma; rachilla nearly 2 mm. long, the lower portion glabrous, the apex sparsely villous with hairs 2.5-3 mm. long; awn to 9 mm. long, straight, divergent from the very base, inserted well above the middle of the lemma and far exceeding it. LA PAZ: NOR YUNGAS: Unduavi, Buchtien 8910 (US); SUR YUNGAS: Sirupaya, Buchtien 430 (us). 28. C. densiflora (Presl) Steud. Nomencl. (ed. 2) 1: 250 (1840). Caespitose, to 6 dm. tall. Leaf-sheaths glabrous, the ligule to 7 mm. long, subtruncate, erose-dentate; blades involute, acute to pungent, to 15 em. long, scabridulous. Inflorescence to 18 cm. long, dense, the short ascending branches appressed, rachis, branches and pedicels scabridulous-puberulous. Glumes about 6 mm. long, acute, scabri- dulous on the upper portion and keel. Lemma sessile below the callus, almost as long as the glumes, somewhat scabridulous, bidentate; callus glabrous or with a very few hairs to 0.4 mm. long; rachilla about 2 mm. long, villous, the longest hairs 3 mm. long; awn straight, to 5 mm. long, inserted at or below the middle of the lemma and exceeding it. LA PAZ: MURILLO: Palca, Pflanz 296 (us). The Pflanz specimen is without data except for the number. However, I have records for his nos. 295 and 297, which were collected at Palca. Thus, it seems probable that this is the locality for no. 296. 29. C. polygama (Griseb). Parodi in Physis, 9: 18, 14 (1928). C. Lilloi of the Catalogue. Caespitose, to 1-1.8 m. high. Leaf-sheaths glabrous, the ligule to 2.5 mm. long, acute to subobtuse; blades flat to folded, shorter than the inflorescence, scabridulous. Inflorescence a lax open panicle to 4 dm. long and 15 cm. wide, the spreading branches distant, rachis, branches and pedicels scabridulous. Glumes 3 mm. long, acute, scabridulous on the keel. Lemma sessile below the callus, as long as the glumes or a little longer, scabridulous, bidentate; callus villous, the longest hairs 0.8 mm. long; rachilla 1.5 mm. long, the longest hairs 1.5 mm. long, equaling the lemma; awn flexuose, about 2 mm. long, inserted shortly below the middle of the lemma and about equaling it. COCHABAMBA: CERCADO: west of Cochabamba, Hitchcock 22838 (GH); Vinto, 2500 m., Parodi 10234 (Parodi). 30. C. viridiflavescens (Poir.) Steud. Nomencl. (ed. 2) 1: 251 (1840). C. montevidensis of the Catalogue. Plant to 1 m. tall. Leaf-sheaths glabrous, ligule about 1 mm. long, deeply dissected-fimbriate; blades flat, to 7 mm. wide, usually much shorter than the inflorescence, scabridulous above and beneath. Inflorescence to 3 dm. long, Joose, rather narrow, somewhat nutant, the lower branches sometimes to 10 258 Rhodora [Vol. 68 em. long, rachis and branches mostly glabrous, the pedicels scabri- dulous. Glumes to 4 mm. long, acute to acuminate, the keel scabri- dulous. Lemma sessile below the callus, about 3 mm. long, bidentate, the 2 teeth shortly fimbriate; callus densely villous, the longest hairs exceeding the lemma; rachilla about 1 mm. long, glabrous except for a very few hairs at the apex, hairs to 2.5 mm. long; awn divergent from the base, straight, about 6 mm. long, inserted above the middle of the lemma and much exceeding it. LA PAZ: SUR YUNGAS: San Felipe, 2800 m., Hitchcock 22601 (GH). 31. C. spicigera (Presl) steud. Nomencl. (ed. 2) 1: 251 (1840). Deyeuxia obtusata Wedd. in Bull. Soc. Bot. France, 22: 177, 180 (1875). D. subsimilis Wedd. in Bull. Soc. Bot. France, 22: 178, 180 (1875). Stoloniferous, to 5 dm, tall. Leaf-sheaths glabrous, the ligule to 2 mm. long; blades involute, arcuate to flexuose, usually glabrous, occasionally puberulous, mostly shorter than the inflorescence. In- florescence to 6 cm. long, dense, spicate in appearance, the short branches ascending, appressed, rachis glabrous to puberulent, branches and pedicels puberulent to hispidulous. Glumes to 8 mm. long, acute, the keel scabridulous. Lemma sessile below the callus, to 6.5 mm. long, bidentate, the 2 teeth bidentate; callus villous, the longest hairs to 2 mm. long; rachilla to 2.5 mm. long, villous, the longest hairs to 5 mm. long, equaling or slightly exceeding the lemma; awn geniculate, the basal portion twisted, inserted about 2 mm. above the base of the lemma and equaling or slightly exceeding it. LA PAZ: LARECAJA: vic. Sorata, Mandon 1311 (Tovar). 38. Agrostis L. Perennials. Leaf-blades flat to involute. Inflorescence a terminal racemose panicle, from dense and spicate in appearance to lax and diffuse. Spikelets 1-flowered, disarticulation above the glumes. Glumes subequal, or, if unequal, the first longer than the second. Lemma hyaline, awned or awnless, much shorter than the glumes; palea present and obvious, or very minute, or absent, a. Panicle sometimes narrow, but open to diffuse, not Spicate. b. Panicle narrow, spikelets many. c. Palea present, obvious, d. Lemmas awned. ....ccccccccccccsecscessecesccessecesscecsscesseess 1. A. gelida. d. Lemmas awnless. ...cccccccccccccesssessssseeceessecceseece 2. A. stolonifera. c. Palea absent or very minute. nesses 8. A. exasperata. b. Panicle open to diffuse, spikelets few. e. Lemmas awnless. |... e D. A. perennans. e. Lemmas awned. f. Spikelets all long-pedicellate. ................ 4. A. montevidensis. f. Spikelets partly short-pedicellate, partly long-pedicellate. dusuusenssunseessssssvyvenyseqesectssesessiuessoyoveeuseccvessesssvccecscsess 6. A. boliviana. a. Panicle narrow, but not open nor diffuse, sometimes spicate. 1966] Flora of Bolivia — Foster 259 g. Palea present. ........ eene 3. A. verticillata, g. Palea absent. h. Panicle spicate; lemma 1.5 mm. long. i. Panicle short (to 7 em.), not interrupted, .......eeenenmmmmm 7, A. Haenkeana. PPPTTETTTT TT h. Panicle narrow, loose; lemma 2 mm, long. .... 9. A. araucana, 1. Agrostis gelida Trin. in Mém. Acad. St. Pétersb. (ser. 6) Sci. Nat. 4(1): 343 (1841). Laxly caespitose, to 1 m. tall. Leaf-sheaths glabrous to scabridulous, ligule to 8 mm. long, abaxially pubescent; blades often long, flat, to 7-8 mm. wide, usually narrower, scabri- dulous beneath, Panicle narrow but lax, to 3 dm. long, the primary branches verticillate, bearing spikelets only from well above the middle, rachis mostly glabrous, branches and pedicels sparsely to densely scabridulous to scabrid. Glumes 3-4 mm, long, acuminate to subaristate, more or less sparsely scabridulous. Lemma to 2 mm. long, more or less truncately obtuse at the apex, the straight awn about 3 mm. long, inserted below the middle of the lemma and equaling or exceeding the glumes; palea nearly as long as the lemma, about 1.5 mm. La Paz: MURILLO: Pongo, Hiteheock 22781 (us). 2. A.stolonifera L. Sp. Pl. 62 (1753). Basally spreading, the culms to 5 dm. long. Leaf-sheaths glabrous, the ligule to 6 mm. long, glabrous; blades flat, to 3 mm. wide, scabridulous above and beneath. Panicle to 20 cm. long, rather open but narrow, the primary branches verticillate, rachis glabrous, the branches bearing spikelets nearly to the base, the branches and pedicels scabridulous to nearly glabrous. Glumes to 2-2.5 mm. long, acute, the keel scabridulous on the upper half. Lemma about 1.5 mm. long, acute to subobtuse, awnless (rarely awned); palea half as long as the lemma, obtuse. LA PAZ: MURILLO: Pongo, Hitchcock 22784 (US). 3. A. verticillata Vill. Prosp. Pl. Dauph. 16 (1779). Decumbent to spreading, sometimes stoloniferous, the culms to 5 dm. long (rarely up to 1 m.). Leaf-sheaths glabrous, the ligule 4-5 mm. long, abaxially short-scabrid, apically ciliate-fimbriate; blades mostly short, flat, to 7 mm. wide, scabrid above and beneath. Panicle dense, interrupted, to 15 cm. long, rachis glabrous, branches and pedicels mostly glabrous, occasionally sparsely scabridulous, the verticillate primary branches short, bearing spikelets to the base. Glumes about 2 mm. long, acute, scabridulous, the keel scabrid. Lemma awnless, about 1-1.5 mm. long, obtuse and toothed at the apex; palea as long, or nearly so. COCHA- BAMBA: CERCADO: Valle de Cochabamba, 2600 m., Steinbach 8773 (GH). 4. A. montevidensis Spreng. ex Nees, Agrost. Bras. 403 (1829). Slender, erect, culms to 2-3 dm. high. Leaf-sheaths glabrous, ligule to 4 mm. long, apically rounded, glabrous; blades short, flat, 1(-3) 260 Rhodora [Vol. 68 mm. wide, scabridulous above and beneath. Panicle rather diffuse, rachis, branches and elongated pedicels variously scabridulous, primary branches verticillate and bearing spikelets only near the apex. Glumes 2-3 mm. long, acute, the keels scabrid. Lemma about 1.5 mm. long, the awn to 3.5 mm. long, geniculate, inserted nearly at the base of the lemma; palea absent. COCHABAMBA: CHAPARE: Llanta-Aduana, 2800-3200 m., Steinbach 8830 (GH, US); Cerro San Benito, 3400 m., Steinbach 9649 bis (GH). 9. A. perennans (Walt.) Tuckerm. in Amer, Journ. Sci. 45: 44 (1843), Decumbent to erect, the culms to 5-10 dm. high. Leaf-sheaths glabrous, ligule to 5 mm. long, glabrous, narrowed above, the apex obtuse and sparsely erose-dentate; blades relatively short, flat, to 3 mm. wide, scabridulous above and beneath. Panicle open, to 2 dm. long, rachis, branches and pedicels glabrous, the primary branches verticillate, rebranched, the spikelets borne at or near the apices on long pedicels. Glumes 2-3 mm. long, acute to acuminate, the keels sometimes scabridulous near the apex. Lemma 1.5-2 mm, long, biden- tate, awnless or with a short awn (to 1.25 mm.) inserted at base of sinus between the teeth; palea absent or extremely minute, L4 Paz: MURILLO: Unduavi, 3200 m., Buchtien 2585 (GH); NOR YUNGAS: Bella Vista, Hitchcock 22757 (GH). COCHABAMBA: CHAPARE: Incachaca, 2250 m., Steinbach 9497 (GH); La Aduana, 2850-3000 m., Steinbach 9695 (GH). 6. A. boliviana Mez in Fedde, Rep. Sp. Nov, 18: 1 (1922), Some- what lax, culms to nearly 6 dm. tall, Leaf-sheaths glabrous, the ligule to 6 mm. long, glabrous, apex bluntly rounded to deeply fim- briate; blades about 3 mm. wide, scabridulous above and beneath. Panicle open, to 18 em. long, the rachis and branches glabrous, the pedicels sometimes sparsely scabridulous, the primary branches re- branched verticillately, the long-pedicellate spikelets borne toward the ends of the secondary branches, Glumes to 3.5 mm. long, acute, the keel finely scabridulous above. Lemma 2 mm. long, deeply biden- tate, the flexuose to geniculate awn to 4.5 mm. long, inserted below the middle of the lemma; palea absent. TARIJA: CERCADO: Calderillo, 3200 m., Fiebrig 2905 (GH). 7. A. Haenkeana Hitche. in Contrib. U. S. Nat. Herb. 24: 881 (1927). Caespitose, spreading to erect, the culms to 5 dm. tall, but usually much shorter. Leaf-sheaths glabrous, the ligule to 4 mm. long, rounded or fimbriate-dentate at the apex, glabrous; blades usually short, to 3 mm. wide, flat to involute, glabrous above and beneath. Panicle dease, spicate in appearance, to 7 cm. long, rachis, branches and pedicels glabrous, or the ped'cels scabridulous, the short branches erect, appressed, bearing spikelets to the base (the lowest spikelets often aborted and reduced to pedicellate rudiments), Glumes to 3.5 mm. long, acute, somewhat scabridulous, the keel scabridulous above. Lemma 1.5 mm. long, the apex truncate and finely denticulate, 1966] Flora of Bolivia — Foster 261 awned, the awn straight, flexuose or geniculate, to 2.5 mm. long, in- serted above the middle of the lemma; palea absent or extremely minute. LA PAZ: LARECAJA: Puerto del Inca, 3800 m., Mandon 1292 (GH). 8. A. exasperata Trin. in Mém. Acad. St. Pétersb. (ser. 6) Sci. Nat. 4(1): 352 (1841). Erect to decumbent, the culms to 1 m. tall, usually shorter. Leaf-sheaths glabrous, the ligule to 5 mm. long, abaxially scabrid or scabridulous, the apex subobtuse to truncately acute, erose-dentate; blades flat, to 3-4 mm. wide, scabridulous above and beneath. Panicle rather dense, subpyramidal, to 15 cm. long, the rachis glabrous below, scabridulous above, the whorled primary branches and pedicels scabridulous to scabrid. Glumes mostly 3 mm. long, acute to long-acuminate and subaristate, densely scabridulous to scabrid, the keel markedly scabrid. Lemma to 1.5 mm. long, rather deeply bidentate, the straight awn to 2.5 mm. long, inserted at the middle of the lemma and equaling or very slightly exceeding the glumes; palea about 0.4 mm. long. LA Paz: MURILLO: Palca, Buchtien 2531 (US). 9. A.araucana Phil. in Anal. Univ. Chile, 94: 14 (1896). Caespitose, to 5 dm. tall. Leaf-sheaths glabrous, the ligule to 4 mm. long, abaxially scabridulous, the apex truncate and dentate; blades flat to loosely involute, to 3 mm. wide, scabridulous above and beneath. Panicle narrow, to 15 em. long, the rachis, ascending-appressed branches and pedicels scabridulous to scabrid, the long-pedicellate spikelets borne nearly to the base of the branches. Glumes to 3.5 mm. long, acute, glabrous or sometimes faintly scabridulous, the keel finely scabridulous. Lemma to nearly 2 mm. long, bidentate, the geniculate awn to 4 mm. long, inserted below the middle of the lemma and much exceeding the glumes; palea absent or very minute. LA PAZ: MURILLO: Unduavi, Buchtien 2581 (Hitchcock). 10. A. tolucensis HBK. Nov. Gen. & Sp. 1: 135 (1816). Erect or spreading, to 6 dm. tall. Leaf-sheaths glabrous, the ligule to 5 mm. long, acute, glabrous; blades flat to involute, to 2 mm. wide, scabri- dulous above and beneath. Panicle narrow, to 15 cm. long, dense, interrupted below, more or less spicate in appearance, rachis and branches glabrous to scabridulous, the pedicels scabridulous. Glumes 2.5-3 mm. long, acute, glabrous to scantily scabridulous, the keel scabridulous. Lemma 1.5 mm. long, bidentate, the straight awn about 2.5 mm. long, inserted at or above the middle of the lemma and much exceeding the glumes (awns sometimes absent); palea absent. LA PAZ: SUR YUNGAS: San Felipe, 2800 m., Hitchcock 22600 (GH). 39. Alopecurus L. See Parodi in Revist. Fac. Agron. Vet. Buenos Aires, 7: 845-369 (1931). 262 Rhodora [Vol. 68 Erect or decumbent perennials or annuals, 50 cm. to 1 m. high. Leaf-blades flat, to 6 mm. wide, glabrous beneath, scabrid above, the uppermost sometimes equaling or slightly exceeding the inflorescence, Inflorescence a condensed panicle, spicate in appearance. Spikelets l-flowered, strongly flattened, disarticulation below the equal glumes. Lemmas about as long as the glumes, awned; palea lacking. Awns nearly 5 mm. long; inflorescence to 3 em. long, nearly 1 cm, WIE, RR 1. A. bracteatus. Awns 1 mm. long; inflorescence about 5 cm. long, about 4 mm. wide. RE 2. A. aequalis. 1. Alopecurus bracteatus Phil. in Anal. Univ. Chile, 94: 6 (1896). Perennial. Spikelets 3-4 mm. long. Glumes 3-4 mm. long, the keel long-ciliate, the hairs up to 1.5 mm. long, the 2 lateral veins silky- villous. Lemmas 3-3.5 mm. long, the upper margins ciliolate, other- wise glabrous, the awn inserted well below the middle of the lemma. LA PAZ: LARECAJA: vic. Sorata; Alto de Ticonguaya, 4100 m., Mandon 1244 (GH); MURILLO: La Paz, 3750 m., Buchtien 431 (GH), 502 (Gu), 861 (GH); Pelaya, 4000 m., Hitchcock 22585 (GH). 2. A. aequalis Sobol. Fl. Petrop. 16 (1799). Annual. Spikelets about 2.5 mm. long. Glumes about 2 mm. long, the keel ciliolate but not densely long-silky. Lemmas 2.5 mm. long, the awn about 1 mm. long, hardly or not at all exserted from the glumes, inserted near the apex of the lemma. LA PAZ: MURILLO: vic. La Paz, 5000 m., Mandon 1243 (Gu). Alopecurus Hitchcockii L. R. Parodi in Revist. Fac. Agron, Vet. Buenos Aires, 7: 366 (1931). Described as having the inflorescence 4 cm. long, 5-6 mm, wide. Spikelets 2.5 mm. long, Lemma-awn 1.5 mm. long, inserted at 1/3 the length of the lemma, not, or only very slightly, exceeding the glumes. From the description, this seems unduly close to A. aequalis. It probably should not be maintained as distinct. 40. Polypogon Desf. Perennial, in wet or damp habitats, to 1 m. high, leaf-blades flat. Inflorescence a narrow, dense, interrupted panicle, the 1-flowered spikelets disarticulating below the equal, awned glumes; lemma shorter than the glumes, thin-textured, usually awned, the awn in- serted below the apex. Glumes gradually attenuate into the awns, about 3 mm, long. ............ sassesendessasacessecnscecesnaveesestensroscssevenssassesssesssuvenssesesasasseeresiaseseces 1. P. elongatus. Glumes abruptly obtuse, not narrowed into the awns, about 2 mm. long. sesesereeeensessesoeaseseecoseresosserereossrsvoesoesorsesessresrrrrsenroanan 2, P. interruptus. 1. Polypogon elongatus HBK. Nov. Gen. & Sp. 1: 134 (1816). Awns 1-1.5 mm. leng, glumes scabrid-puberulent, Lemma 2 mm. long, the apex 2-4-toothed, the awn about 2 mm. long. LA PAZ: LARECAJA: 1966] Flora of Bolivia — Foster 263 vic. Sorata, Mandon 1294 (GH); SUR YUNGAS: San Felipe, 2800 m., Hitchcock 22599 (GH). 2. P. interruptus HBK. No. Gen. & Sp. 1: 134 (1816). P. lutosus of the Catalogue. Awns 2.5-3 mm, long, glumes, especially the first, densely scabrid-puberulent. Lemma about 1 mm. long, apex toothed, awn about 2.5-3 mm. long. LA PAZ: MURILLO: canon of Río de La Paz, 3355 m., Shepard 160 (GH); La Paz, 3300 m., Rusby 40 (GH), Bang 47 (GH), 3650 m., Buchtien 169 (GH), 238 (GH). COCHABAMBA: CERCADO: foothills of Mt. Tunari, 3400 m., Steinbach 9768 (GH); Valle de Cochabamba, Steinbach 8773 (GH), 8789 (GH); CHAPARE: Sacaba, 2900 m., Steinbach s.n. (GH); MIZQUE: Vilavila, 2600 m., Eyerdam 25321 (GH). Potosi: CERCADO: Potosí, 4000 m., Cárdenas 206 (GH). 41. Lycurus HBK. Lycurus phleoides HBK. Nov. Gen. & Sp. 1: 142 (1816). Perennial, somewhat caespitose, to 3 dm. high. Leaves mostly basal, crowded, the sheaths puberulent to scabrid. the short flat blades scabridulous above and beneath, scabrid-ciliate. Inflorescence a terminal dense panicle, spicate in appearance, to 5 cm. long, the branchlets and pedicels sca- bridulous. Spikelets 1-flowered, paired, 1 spikelet perfect, the other staminate or sterile, the pair falling as a unit, disarticulation below the glumes. Glumes about equal, or the second longer than the first, 1.5-2 mm. long, 2-awned, the awns to 6 mm, long, the keels scabrid- ciliolate. Lemmas to 4 mm. long with a 3 mm. straight awn, 3-nerved, the margins villous-ciliate on the lower half. Palea as long as the lemma, villous on the keels. L4 Paz: MURILLO: Capi, Bang 762 (GH). COCHABAMBA: CERCADO: west of Cochabamba, Hitchcock 22822 (GH). Potosi: CERCADO: Potosi, 4000 m., Cárdenas 204a (GH). 42. Muhlenbergia Schreb. See Parodi in Fhysis, 9: 205-222 (1928). Annual or perennial, from dwarf plants a few centimeters high to about 1 m. high. Leaf-blades flat to involute. Inflorescence a race- mose panicle, from dense and spicate to diffuse. Spikelets 1-flowered (2-3-flowered in one species), disarticulation above the awnless glumes. Lemma firm-textured, awned, the awn terminal, or very shortly cuspidate; palea similar to lemma and about as long, awnless. a. Second glume deeply tridentate. menn 1. M. peruviana. a. Second glume entire. b. Panicle spicate in appearance. «eet b. Panicle not spicate. c. Panicle open to diffuse. d. Lemma to 1.5 mm. long, awned, the awn to 5 mm. lotis. 3. M. tenuissima. 7. M. angustata. UE Micro essoesacotescaccoseetbo ses bi so eniro $e PP ERI HMM 264 Rhodora [Vol. 68 d. Lemma to 2 mm, long, awnless, but shortly cuspidate. ........ vosssneacascecceseeesssncssssssstecececssessssssesecsecsensnsessssseseeseees 4. M. asperifolia. c. Panicle narrow. e. Lemma awnless. f. Leaf-blades flat; rhizome absent. ............ 5. M. ligularis. f. Leaf-blades involute; slender scaly rhizome present. ........ (REN 6. M. fastigiata. e. Lemma awned. g. Awn over 1 em. long. h. Lemma 2.5-3 mm. long. .......... sss 2. M. quitensis. h. Lemma to 6-7 mm. long. c. 9. M. rigida. g. Awn 2-4 mm. long. ......... cess 8. M. Holwayorum. 1. Muhlenbergia peruviana (Beauv.) Steud. Nomencl. (ed. 2) 1: 41 (1840). M. Herzogiana Henrard in Meded. Rijks Herb. Leiden, no. 40: 58 (1921). Caespitose annual, 2-20 cm. high. Leaf-sheaths glabrous, ligule to 2 mm. long, acute to subobtuse; blades involute, very short, glabrous. Panicle to 2 cm. long, narrow, rather dense, rachis, branches and pedicels scabridulous. First glume entire, acute, to 1.5 mm. long; second glume to 2.25 mm. long, deeply tridentate. Lemma to 2.25 mm. long, scabridulous to pubescent, slightly bidentate, the straight or flexuose awn to 4 mm. long (rarely to 1.5 cm.) ; palea nearly as long as the lemma, sparsely pubescent. LA Paz: MURILLO: La Paz, 4100 m., Buchtien 72 (GH). Potosi: CERCADO: Potosi, 4000 m., Cardenas 196 (GH); LINARES: Lagunillas, 3800 m., Cárdenas 483 in part (GH; originally entangled in a Stevia, no. 483). COCHABAMBA: CERCADO: Mt. Tunari, 3800 m., Steinbach 9849 (GH). TARIJA: CERCADO (?): Cuesta San Anaton, 3000 m., Fiebrig 3277 (Gn). 2. M. quitensis (HBK.) Hitche. in Contrib, U. S. Nat. Herb. 17: 292 (1913). Annual, 3-9 dm. high. Leaf-sheaths slightly scabridulous, the ligule to 4 mm. long; blades flat or involute, scabridulous, short (to 5 cm.), 2 mm. wide. Inflorescenca narrow but loose, interrupted, to 2 dm. long, the primary branches bearing spikelets almost to the base. First glume to 2 mm. long, acute; second glume to 2.5-3 mm. long, entire, subulate-aristate. Lemma about 2.5-3 mm. long, sub- bidentate, the awn to 2 cm. long; palea pubescent at the base. With- cut locality: Bang 947 (Hitchcock). 3. M. tenuissima (Presl) Kunth, Rév. Gram. 1: suppl. XVI (1830). Delicate annual, to 2 dm. tall, occasionally taller. Leaf-sheaths glabrous to puberulous, the ligule to 2 mm. long, the apex rounded to subacute; blades short, flat, about 1 mm. wide, sparsely pubescent above and beneath, or almost glabrous. Panicle 10 em. long, open, the primary branches apparently secund on occasion, rachis, branches and pedicels mostly glabrous. First glume to 1 mm. long, acute; second glume to 1.5 mm. long, long-acute or acuminate. Lemma (ex- cluding awn) 1.5 mm. long, scabridulous, subbidentate, the straight awn to 5 mm. long; palea as long as lemma, somewhat puberulent at 1966] Flora of Bolivia — Foster 265 base. LA Paz: SUR YUNGAS: La Florida, 1840 m., Hitcheock 22622 (GH). COCHABAMBA: TARATA: Anzaldo, Cárdenas 2477 (GH). 4. M. asperifolia (Nees & Meyen) Parodi in Physis, 9: 210 (1928). Perennial, decumbent, to 4 dm. tall, with creeping rhizomes. Leaf- sheaths finely scabridulous to glabrous, the ligule 1-2 mm. long, apex rounded, ciliate or fimbriate-denticulate; blades short, flat, to 1.5 mm. wide, not falcate, scabridulous above and beneath. Panicle open, very diffuse, the primary branches rebranched, the branchlets bearing solitary long-pedicellate spikelets at the ends, branchlets and pedicels sparsely to rather densely scabridulous. Spikelets often 2-flowered, rarely 3-flowered, the florets distant on a glabrous rachilla. First glume to 1.75 mm. long, acute, somewhat scabridulous, especially on the keel; second glume to 2.25 mm. long, scabrid on the keel, the apex narrowly subtruncate, the midrib excurrent as a short scabridulous cusp. Lemma dark purple, at least in part, to 2 mm. long, slightly bidentate, the cusp less than 0.5 mm. long; palea as long as the lemma, bidentate, each tooth cuspidate, PoTosí: SUR CHICHAS: Oploca, Hitch- cock 22891 (GH). 5. M. ligularis (Hack.) Hitche. in Contrib. U. S. Nat. Herb. 24: 388 (1927). Decumbent tufted perennial, not rhizomatous, culms to 10 em. high. Leaf-sheaths glabrous to finely papillate-scabridulous, the ligule to 4 mm, long, more or less acute; blades flat, to 2 mm. wide, rarely over 2 cm. long, finely papillate-scabridulous above and beneath. Panicle short, hardly 4 cm. long, narrow, few-branched, rachis, branches and short pedicels finely scabridulous. Glumes subequal, about 1 mm. long, the apex usually rounded, occasionally truncately subacute, the keel not prominent. Lemma to 2 mm. long, callus glabrous, finely scabridulous above, the midrib slightly excurrent as a cusp 0.2 mm. long; palea as long as the lemma, LA PAZ: MURILLO: vie. La Paz, 3300 m., Bang 109 (GH); LARECAJA: Millipaya, 3700 m., Mandon 1286 (GH). 6. M. fastigiata (Presl) Henrard in Meded. Rijks Herb. Leiden, no. 40: 59 (1921). Tufted creeping perennial, a few centimeters high, with spreading, slender, scaly rhizomes. Leaf-sheaths finely papillate, ligule about 1-1.5 mm. long, acute; blades involute, falcate, seldom over 1 cm. long, finely papillate. Panicle to 1 cm. long, narrow, few- flowered, rachis, branches and short pedicels finely papillate to scabri- dulous. Glumes subequal, slightly over 1 mm. long, acute, keel prominent, scabridulous. Lemma to 2.5 mm. long, acute, the apex with a cusp about 0.2-0.4 mm. long; palea about as long as the lemma, hardly cuspidate. LA Paz: MURILLO: La Paz, 4100 m., Buchtien (GH). TARIJA: AVILES: Puna Patanca, 3700 m., Fiebrig 2633 (GH). 7. M. angustata (Presl) Kunth, Rév. Gram. 1: suppl. XVI (1830). Caespitose perennial, to 8 dm. tall. Leaf-sheaths scabridulous, the ligule to 1.1 em. long, rounded to acute at the apex; blades long, folded to involute, densely scabridulous. Panicle dense, spicate in 266 Rhodora [Vol. 68 appearance, to 15 em. long, rachis, branches and short pedicels densely scabrid to densely short-hispid. Glumes equal or subequal, to 6-7 mm. long, acute, scabridulous. Lemma as long as the glumes, scabridulous, the straight awn to 3 mm. long; palea slightly shorter than the lemma, scabridulous, acute to subbidentate, LA PAZ: LARECAJA: near Milli- paya, 3600 m., Mandon 1279 (GH). TARIJA: CERCADO: Calderillo, Fiebrig 3173 (GH). 8. M. Holwayorum Hitche. in Contrib. U. S. Nat. Herb. 24: 389 (1927). Caespitose perennial to 6 dm. high. Leaf-sheaths glabrous, the ligule to 5 mm. long; blades flat to involute, to 3 mm. wide, sca- bridulous. Panicle narrow, to 2 dm. long, the rachis, appressed branches and pedicels scabridulous. Glumes subequal, about 1.5 mm. long, acute. Lemma to 3-4 mm. long, the awn to 3-4 mm. long; palea as long as the lemma. LA PAZ: LARECAJA: Sorata, Holway & Holway 530 (US; type). 9. M. rigida (HBK.) Kunth, Rév. Gram. 1: 63 (1829). M. phragmi- toides of the Catalogue. Caespitose perennial, to 8 dm. tall. Leaf- sheaths finely puberulent to scabridulous, the ligule to 4 mm. long, apically dentate; blades firm, involute, scabridulous (at least above), acute, Panicle narrow, lax, not spicate in appearance, to 3 dm. long, the rachis, ascending branches and long pedicels scabridulous. Glumes subequal, to 2 mm. long, acute. Lemma to 6-7 mm. long, scabridulous, the awn to 1.5 em. long; palea slightly shorter than the lemma. La Paz: MURILLO: Obrajes, 3350 m., Buchtien 819 (GH); LARECAJA: San Pedro, 2600-2700 m., Mandon 1280 (GH). 43. Triniochloa Hitchc. Triniochloa stipoides (HBK.) Hitche. in Contrib. U. S. Nat. Herb. 17: 303 (1913). ‘Perennial, to 1 m, high. Leaf-sheaths and blades puberulent, the blades very narrow and sometimes involute. Inflo- rescence a lax, few-flowered panicle, to 2 dm. long, the branches scabridulous. Spikelets 1-flowered, disarticulation above the glumes, to 1.2 em. long (excluding the awn). Glumes equal, thin-textured, 2-3 mm. long, acute, 1-nerved, the keels scabridulous. Lemma nearly terete, rather indurate, to 1.2 em. long, scabrid-puberulent on the upper portion, bidentate, the dorsal geniculate awn to nearly 2 cm. long, the lower half loosely twisted, the callus densely villous with hairs to 1.5 mm. long. Palea as long as the lemma and as firm- textured, deeply bidentate, the keels scabridulous. LA PAZ: NOR YUNGAS: Unduavi, 3800 m., Buchtien 4172 (GH). 44. Aciachne Benth. See Chase in Journ. Wash. Acad. Sci. 14: 364-366 (1924). Aciachne pulvinata Benth. in Hook. Ic. Pl. 4: 44, t. 1362 (1881). Low-growing, less than 10 em, high, densely caespitose perennial, in 1966] Flora of Bolivia — Foster 267 wide-spread clumps. Leaf-blades involute, glabrous, falcate, pungent, about 1 em. long. Inflorescence a single 1-flowered perfect spikelet, terminal on a solitary peduncle, hardly, or not at all, raised above the leaves, disarticulation above the glumes. Glumes equal, about 2 mm. long, more or less indurate, produced into a long, sharp awn- like point. Palea much shorter than the lemma and more or less enclosed by it, the rounded apex bifid. LA PAZ: LARECAJA: vic. Sorata; Chilcani, 3600-4200 m., Mandon 1287 (GH). COCHABAMBA: CHAPARE: Cumbre de San Benito, 3600 m., Steinbach 9888 (GH). Locality uncertain: Fiebrig 3571 (GH), Bang 1843 (GH). 45. Sporobolus R. Br. See Parodi in Revist. Fac. Agron. Vet. Buenos Aires, 6: 115-168 (1928). Annual or perennial. Ligule obsolete, reduced to a semi-lunate line of hairs. Leaf-blades involute or flat. Inflorescence a panicle, open or spicate in appearance, often subpyramidal. Spikelets 1-flowered, disarticulation above the unequal glumes, Lemma and palea similar in appearance, the palea frequently splitting. Caryopsis turbinate- pyriform to oblong-ovoid, readily shed. a. Primary branches not in whorls. b. Spikelets hardly 1 mm. long; panicle very open and diffuse. .... S EE cc oosixar DEOR UC ODD Huet Cea 1. S. tenuissimus. b. Spikelets 2-3 mm. long. c. Panicle narrow, spicate in appearance; spikelets to 3 mm. long GEAR he EL imesh a cae IIR cdi UR 4. S. Poiretü. c. Panicle more open, not spicate; spikelets 2 mm. long, or less. a. Primary panicle-branches in whorls. d. Spikelets about 2 mm. long. .................. 2. S. argutus. d. Spikelets 3-4 mm. long. ................... n 3. S. aeneus. 1. Sporobolus tenuissimus (Mart. ex Schrank) O. Ktze. Rev. Gen. 3(2): 869 (1898). Annual, delicate, to 7 dm. high, glabrous, the culms compressed. Leaf-blades linear, to 3 mm. wide, sometimes equaling the inflorescence. Panicle to 3.5 dm. long, narrow, very open, the primary branches not whorled, branches ascending to spreading, pedicels deli- cate, long. Spikelets 1 mm. long, or less. First glume to 0.3 mm. long, narrowly linear; second glume 0.6 mm. long, narrowly deltoid. Lemma ovate to obovate, 1 mm. long or less, acute; palea as long, broader, the apex truncate to obtusely rounded. Caryopsis turbinate- pyriform, broadest at the apex or very near it. SANTA CRUZ: SARA: between the Río Cuchi and Río Piray, Herzog 1446 (Hitchcock). 2. S. argutus (Nees) Kunth, Enum. Pl. 1: 215 (1833). Spreading to suberect perennial, to 5 dm. tall. Leaf-sheaths glabrous; blades glabrous, the margins scabrid, linear, to 7-8 em. long and 5 mm, wide, 268 Rhodora [Vol. 68 flat to folded or subinvolute. Panicle to 10 cm. long, narrow, nar- rowly pyramidal at maturity, the primary branches in whorls, rachis, branches and pedicels glabrous. Spikelets about 2 mm. long. First glume 0.5 mm. long, oblong-ovate, subobtuse; second glume 1.75 mm. long, acute. Lemma 1.5 mm. long, acute; palea shorter and broader, Caryopsis oblong-ovoid, broadest at the middle. LA PAz: MURILLO: La Paz, 3300 m., Bang 79 (GH). 3. S. aeneus (Trin.) Kunth, Rév. Gram. 1: suppl. XVII (1830). Densely caespitose perennial, to 6 dm. tall. Leaf-sheaths glabrous, the margins ciliate; blades long, narrow, folded or involute, pilose at the base on the adaxial surface, the margins ciliate. Panicle subpyra- midal, narrow, to 3 dm. long, primary branches in distant whorls, floriferous nearly to the base, rachis, branches and pedicels glabrous. Spikelets 3-4 mm. long. First glume about 1.5 mm. long, lance-linear; second glume as long as the spikelet, 1-nerved. Lemma to 3-4 mm. long, lanceolate, acute, 3-nerved; palea as long as lemma. Caryopsis oblong-ovoid, broadest at the middle. LA PAz: LARECAJA: Hacienda Casana, Buchtien 7147 (Hitchcock). 4. S. Poiretii (Roem. & Schult.) Hitchc, in Bartonia, 14: 32 (1932). S. Berteroanus of the Catalogue. Perennial, to 1 m. tall. Leaves glabrous, the blades sometimes equaling the inflorescence, flat below to involute above, to 5 mm, wide at the base. Inflorescence to 4 dm. long, usually much shorter, spicate in appearance, the primary branches not in whorls, rachis glabrous, branches and pedicels usually glabrous, sometimes sparsely scabridulous. Spikelets to 3 mm. long. First glume 0.5-1 mm. long, oblong-ovate, the apex truncate and erosulous; second glume 1.5 mm. long, acute, the upper margins erose. Lemma nearly 3 mm. long, acute to subobtuse; palea similar, but slightly shorter. Caryopsis turbinate-pyriform, broadest near or at the apex. LA PAZ: LARECAJA: Sorata, Mandon 1285 (GH); MURILLO: La Paz, 3700 m., Buchtien 179 (GH); NOR YUNGAS: Millu- guaya, 1300 m., Buchtien 4187 (GH). COCHABAMBA: CHAPARE: Sacaba, 2900 m., Steinbach 8808 (GH); CERCADO: Mollemolle, 2800 m., Steinbach 4054 (GH). SANTA CRUZ: SARA: Buenavista, 500 m., Steinbach 5148 (GH). 5. S. indicus (L.) R. Br. Prodr. Fl. Nov. Holl. 1: 170 (1810). Perennial, to 8 dm. tall. Leaves glabrous, the blades chiefly involute, to 2 mm, wide at the base, shorter than the inflorescence. Panicle interrupted at the base, branches not whorled, moderately long and ascending to spreading, rachis, branches and pedicels glabrous to sparsely scabridulous. Spikelets 2 mm. long or less. First glume less than 0.5 mm. long, obtuse to truncate and erosulous; second glume somewhat less than 1 mm. long, acute or subacute. Lemma about 2 mm. long, acute; palea similar, usually somewhat shorter. SANTA CRUZ: ICHILO: Rio Palometillas, 400 m., Steinbach 7933 (GH); SARA: 1966] Flora of Bolivia — Foster 269 Buenavista, 450-500 m., Steinbach 6701 (GH), 6843 (GH), 6844 (GH), 6852 (GH), 6854 (Gu). Sporobolus eximius (Nees) Ekm. was included among the Bolivian species by Hitchcock, on the basis of a Fiebrig collection from Toldos. This locality is in Argentina. I have seen no Bolivian material of the species. 46. Nassella (Trin.) E. Desv. See Parodi in Darwiniana, 7: 369-395 (1947). Perennials, often caespitose. Leaf-blades flat or convolute. Inflo- rescence a panicle, the 1-flowered spikelets disarticulating above the glumes. Glumes equal or unequal, rather thin, mostly hyaline. Lemma obliquely obovoid, gibbous toward the apex, often somewhat apiculate or mucronulate, enclosing the short membranous palea. Awn ex- centrieally attached, caducous. a. Glumes equal, or if unequal, not markedly so; lemma villous, callus villous. b. Lower portion of awn villous; callus acute. .... 1. N. Asplundii. b. Lower portion of awn scabrous; callus obtuse. .... 2. N. pubiflora. a. Glumes markedly unequal; lemma glabrous or nearly so; callus glabrous or with a few short hairs. .................... 3. N. Meyeniana. 1. Nassella Asplundii Hitchc. in Contrib. U. S. Nat. Herb. 24: 394 (1927). Caespitose, to 6 dm. tall. Leaf-sheaths glabrous or sparsely villous at the apex; blades involute, to 5 em. long (occasionally longer). Panicle to 1.5 dm. long, open, the capillary branches naked below, axis and branches glabrous, pedicels scabrous-hirsutulous. Glumes to 4 mm. long, 3-nerved, the first glume ciliate on the keel. Lemma densely villous, 2.5 mm. long, the oblique apex unilaterally apiculate. Awn to 1.5 em. long. Potosi: Porco: Uyuni, 3500 m., Asplund 6548 (vs; type). 2. N. pubiflora (Trin. & Rupr.) Desv. in Gay, Fl. Chile, 6: 264 (1853). N. flaccidula Hack. and N. flaccidula var. humilior Hack. in Fedde, Rep. Sp. Nov. 6: 155 (1908). N. deltoidea Hack. in Fedde, Rep. Sp. Nov. 11: 23 (1912). ? Stipa tiraquensis O. Ktze. Rev. Gen. 3(2): 313 (1898). Stipa Pflanzii Mez in Fedde, Rep. Sp. Nov. 17: 206 (1921). Caespitose, 5-9 dm. tall. Leaf-sheaths glabrous to scabrous; blades flat or subconvolute, glabrous, to 20 cm. long. Panicle to 2 dm. long, the lower branches naked on the basal third, axis scabridulous, branches and pedicels puberulent to shortly pubescent. Glumes 3.5-5 mm. long, 3-nerved, the first glume sparsely ciliate on the keel. Lemma villous, 2-2.4 mm. long, the obtuse callus villous; awn geniculate, to 1.5 cm. long, the base scabrous. LA PAZ: LARECAJA: vic. Sorata; Puquesani, 2700-3000 m., Mandon 1270 (GH); Catarguata, 3000 m., Mandon 1270 bis (GH; in part); MURILLO: La Paz, Buchtien 8541 270 thodora [Vol. 68 (GH), 9309 (GH); Cerro del Calvario, 3800 m., Buchtien 145 (Hitch- cock); Palca, 3700 m., Hitchcock 22579 (GH). 8. N. Meyeniana (Trin. & Rupr.) Parodi in Darwiniana, 7: 379 ((1947). N. corniculata Hack. in Fedde, Rep. Sp. Nov. 5: 155 (1908). Caespitose, to 6 dm. tall. Leaf-sheaths glabrous or somewhat pubes- cent at the apex; blades flat or subconvolute, to 20 cm, long, glabrous or dorsally pubescent. Panicle narrow, somewhat interrupted, to 25 «m. long, branchlets covered by spikelets to the base, axis and branch- lets glabrous, the pedicels hispidulous on the angles. Glumes glabrous, markedly unequal, the first glume 4-4.5 mm. long, the second glume 3.5-4 mm. long. Lemma 1.8-2.2 mm. long, glabrous, shining, the obtuse callus glabrous or with a few short hairs, awn 1-1.5 em. long, weakly or not at all geniculate. LA PAZ: MURILLO: La Paz, Bang 40 (GH); La Paz, 3700 m., Buchtien 366 (GH). 47. Piptochaetium Pres] See Parodi in Revist. Mus. La Plata (n. s.) Bot. 6: 213-310 (1944). Piptochaetium panicoides (Lam.) E. Desv. in Gay, Fl. Chile, 6: 270 (1853). P. setifolium and P. tuberculatum of the Catalogue, Caespi- tose perennial to 4 dm. tall. Leaves mostly basal, shorter than the inflorescence, the glabrous involute blades to 15 cm. long and 0.5 mm. wide. Inflorescence a terminal racemose panicle, to 1 dm. long, few- flowered, the rachis, branches and pedicels puberulent-scabridulous. Spikelets 1-flowered, disarticulation above the glumes. Glumes sub- equal, 3.5-4 mm. long, acuminate, several-nerved, glabrous, Lemma more or less D-shaped, to 2.5 mm. long, thick, firm-textured, sur- rounding the caryopsis and almost concealing the palea, finely striate, with a short apical collar strongly oblique at the summit, bearing a geniculate, loosely twisted excentric awn to 1 cm. long. Palea cori- aceous, with 2 keels. (Department and province unknown): Alto de Cuchilla, 2600 m., Troll 839 (Parodi). P. panicoides f. subpapillosum (Hack.) Parodi in Revist. Mus. La Plata (n. s.) Bot. 6: 302 (1944). A form with the lemma more or less papillose-tuberculate. LA PAZ: LARECAJA: vic. Sorata; Munay- pata, 2650-2900 m., Mandon 1274 (GH). 48. Stipa L. See Hitchcock in Contrib. U. S. Nat. Herb. 24: 263-289 (1925). Perennial. Leaf-blades mostly involute, sometimes flat. Inflores- cence a panicle. Spikelets 1-flowered, disarticulation above the glumes; callus sharply pointed. Glumes hyaline-membranous. Lemma indurate, mostly fusiform, with a terminal twisted awn. a. Glumes unequal, the outer much longer than the inner, b. Longest glumes 1 cm. or more in length, c. Lemma with an apical crown. 1966] Flora of Bolivia — Foster 271 d. Lemma about 8 mm. long. .......... oie. 9. S. Neesiana, d. Lemma about 3 mm. long. .................... 11. S. depauperata. c. Lemma without an apical crown. .................... 18. S. Holwayi. b. Longest glumes to 7 mm. long. ................ 3. S. pampagrandensis. a. Glumes equal or subequal, but the outer not much longer than the inner. e. Longest glumes about 1 cm. or more in length. f. Lemma with an apical crown. g. Lemma 5-6 mm. long, villous below and nearly to apex; crown conspicuous, ciliate with short, thick, hispid hairs. 8. S. mucronata. déttsssssskshssssssssosssosssosesesssosshosesessssasocoshsesosesososostooee g. Lemma 4 mm. long, appressed-villous, crown short. ............ 12. S. brachyphylla. Messsssssosoossssossesesossoseseoseesosevesesecsssosoonvoeseeececscseos f. Lemma without an apical crown. h. Awn plumose on one segment. i. Lemma to 9 mm. long; awn plumose on basal segment Only e a e e a FERUNT rp] ak ee 1. S. speciosa. i. Lemma to 5 mm, long; awn plumose on terminal segment Onlyi. OEE pees sien Reman riya eee LUE 2. S. plumosa. h. Awn not plumose, but sometimes pubescent or subvillous. j. Lemma to 4 mm. long, villous, the apical hairs to 3 mm. Jong n Eo Meese sieves crate TIRES 6. S. nardoides. j. Lemma to 8 mm. long, sparsely villous, or glabrous, the apicalhairs, If any, short. UA 14. S. curviseta. e. Lengest glumes less than 1 cm. long; lemma lacking an apical crown. k. Awn straight or flexuose, but not geniculate. l. Apex of lemma with a dense ring of hairs 3-4 mm. long; lemma fusiform or subfusiform, m. Lemma glabrous or sparsely villous at base or apex. wur" MM 7. S. Hans-meyeri. m. Lemma sparsely pubescent or short-villous. n. Lemma to 3 mm. long. ........."... ener 4. S. Ichu. n. Lemma to 4-5 mm. long. ........1..... 5. S. capilliseta. l. Apex of lemma without a dense ring of hairs. o. Lemma-apex broad, truncate; awn slightly excentric. ...... NNMERO UNE E 19. S. florulenta. o. Lemma-apex attenuate, not truncate; awn not excentric. p. Glumes acute; lemma glabrous above, sparsely pubes- cent below. 5 oic OPE 16. S. polyclada. p. Glumes more or less obtuse; lemma glabrous, save for a few short hairs near apex. escai 15. S. obtusa. k. Awn geniculate. q. Lemma obovoid, apex truncate. ............ 10. S. inconspicua. q. Lemma narrowly ellipsoid to fusiform, apex attenuate. r. Lemma 3 mm. long, sparsely villous. .... 18. S. dasycarpa. 212 Rhodora [Vol. 68 r. Lemma at least 4 mm. long. s. Plants to 2.5 dm. tall; lemma short-pilose. ...................- ——— HHÉÁÉÁRPRRRRRRRRRRRREN 17. S. illimanica. s. Plants to 4-6 dm. tall; lemma sparsely short-pilose. —— ÁÉÁÉEERRRRRREEÜ 20. S. Trollii. l. Stipa speciosa Trin. & Rupr. in Mém. Acad. St. Pétersb. (ser. 6) Sci. Nat. 5(1): 45 (1843). To 6 dm. tall, rigid, erect, the brown bases glabrous to pubescent. Leaves involute, acute to pungent, glabrous, or the margins loosely long-ciliate. Panicle rather dense, to 1 dm. long, the axis glabrous to scabrid to pubescent, the pedicels scabrid to pubescent and villous at the apex. Glumes thin, subequal or the outer shorter, long-acuminate, to 2.2 cm. long, 1-3-nerved, glabrous. Lemma to 9 mm. long, densely pubescent to villous. Awn to 6 cm. long, once-geniculate, twisted and plumose below the geniculation, glabrous above it. ORURO: PooPó: Pasña, Buchtien 1184 (US) ; PORCO: Uyuni, Asplund 6552 (us). 2. S. plumosa Trin. in Mém. Acad. St. Pétersb. (ser. 6) Sci. Nat. 4(2): 87 (1836). Erect to straggling, somewhat woody, to 2 m. tall, the base leafless or covered with old, glabrous to pubescent leaf- sheaths. Leaf-blades involute or subinvolute, long-ciliate, acute. Panicle somewhat lax, narrow, to 2 dm, long, the axis and branchlets mostly glabrous, pedicels glabrous to seabrid and short-pubescent on the angles. Glumes thin, papery, long-acuminate, 3-nerved, sparsely villous, usually on the midrib, to 1.2 cm. long. Lemma to 5 mm. long, pubescent to short-villous, glabrous near the apex and with an apical ring of sparse stiff hairs. Awn strongly geniculate or curved, to 4 cm. long, the basal section twisted and pubescent to short-villous, the upper section plumose. SANTA CRUZ: VALLEGRANDE: Comarapa, Herzog 1885 (US). PoTosí: PORCO: Uyuni, Asplund 6563 (Us). 3. S. pampagrandensis Speg. in Anal. Mus. Nac. Montevideo, 4: 158 (1910). To 10 dm. tall, erect, glabrous. Leaf-blades involute. Panicle narrow, lax to 2 dm. long, somewhat exceeding or nearly equaled by the leaf-blades, axis glabrous, pedicels scabrid. Glumes unequal, the outer much longer than the inner, to 7 mm. long, long-attenuate to almost aristate, thin, 2-nerved. Lemma more or less obovoid, glabrous, irregularly rugulose or papillate-striate, to 3 mm. long. Awn to 5 cm. long, geniculate or strongly curved, the lower portion loosely twisted and scabridulous, the upper portion straight and glabrous to minutely scabridulous. TARIJA: CERCADO: Junaca, Fries 1308 (US). 4. S. Ichu (Ruíz & Fav.) Kunth, Rév. Gram. 1: 60 (1829). Caespi- tose, erect, glabrous, to 1.5 m. high. Leaf-blades involute, pungent. Panicle to 4 dm. long, lax, rather narrow, often silvery in color, the axis glabrous, branchlets and pedicels finely scabridulous. Glumes subequal or equal, thin, papery, long-acute, to 8 mm. long. Lemma fusiform, to 3 mm. leng, sparsely pubescent, with a dense apical ring of hairs to 3-4 mm. long. Awn to 1.7 em. long, scabridulous, not 1966] Flora of Bolivia — Foster 273 geniculate but somewhat flexuose, the basal portion loosely twisted. LA PAZ: LARECAJA: Río Challasuyos between Lojena and Illampü, 2650-3800 m., Mandon 1273 (GH); MURILLO: La Paz, 3750 m., Buch- tien 185 (GH); Palca, Hitchcock 22581 (GH). PoTosí: CERCADO: Potosí, 4000 m., Cárdenas 199 (GH), 205 (GH). COCHABAMBA: CHAPARE: Cuchicanehi, 3300 m., Steinbach 9632 (GH). 5. S. capilliseta Hitche. in Contrib. U. S. Nat. Herb. 24: 271 (1925). Caespitose, glabrous, to 5 dm. tall. Leaf-blades about equaling the base of the inflorescence, pungent, involute. Panicle narrow, to 15 cm. long, axis, branches and pedicels scabrid. Glumes subequal, to 7 mm. long, hyaline, papery, obscurely 3-nerved, scabridulous near base and along midrib, long-acute. Lemma 4-5 mm. long, fusiform, sparsely pubescent to short-villous, especially below, the apex with denser hairs to 4 mm. long. Awn to 2 cm. long, scabridulous, not geniculate but the lower portion loosely twisted. Potosi: NOR LIPEZ: Chiguana, 3700 m., Asplund 6556 (US; type). 6. S. nardoides (R. A. Phil.) Hack. ex Hitche. in Contrib. U. S. Nat. Herb. 24: 271 (1925). Densely caespitose, to 4 dm. high. Leaf- sheaths short-pubescent, short-villous at the apex; blades short, involute, obtuse to subpungent. Panicle narrow, to 1 dm. long, axis, branches and pedicels puberulent to scabridulous. Glumes subequal, glabrous, to 1 em. long, long-acute, obscurely 3-nerved, purplish, the margins and apex white-hyaline. Lemma to 4 mm. long, terete or subfusiform, densely short-villous, the longer apical hairs to 3 mm. long. Awn to 2 cm. long, geniculate, basal portion tightly twisted, somewhat puberulous, upper portion shortly puberulous. ORURO: PORCO: Uyuni, Asplund 6539 (us). 7. S. Hans-meyeri Pilcer in Engler, Bot Jahrb. 56, Beibl. 123: 24 (1920). Caespitose, glabrous, to 5 dm. tall. Leaf-blades involute, pungent. Panicle dense, to 10 (-20) cm. long, purple or brownish- purple, axis, branches and pedicels scabridulous. Glumes equal or subequal, to 7 mm. long, long-acute, glabrous. Lemma subfusiform, to 3.5 mm. long, sparsely villous at base and apex, the apical ring of hairs dense and to 3 mm. long. Awn about 1 cm. long, curved but not strongly geniculate, the base puberulent, the upper portion finely scabridulous. LA PAZ: LARECAJA: inter Millipaya et Velatutini, 3700 m., Mandon 1272 bis (GH). 8. S. mucronata HBK. Nov. Gen. & Sp. 1: 225 (1816). Caespitose, to 10 dm. tall, erect or geniculate, mostly glabrous. Leaf-blades often about equaling the inflorescence, flat or somewhat folded, sparsely scabrid beneath, the margins coarsely ciliate. Panicle to 3 dm. long, lax, the axis glabrous, branches and pedicels somewhat puberulent- scabridulous. Glumes subequal, or the outer slightly longer, about 1 cm. long, long-acute, the midrib and margins often coarsely and sparsely hispid-ciliate. Lemma to 5-6 mm. long, subfusiform, villous below and nearly to the summit, the apex developed into a crown 274 Rhodora [Vol. 68 terminated by short stiff hairs. Awn to 3 em. long, twice bent, the lowest portion tightly twisted and pubescent, the central portion loosely twisted and scabridulous, the terminal portion straight and only minutely scabridulous. LA PAZ: MURILLO: La Paz, 3750 m., Buchtien 3692 (GH), 430 (GH); LARECAJA: vic. Sorata, 2600-2700 m., Mandon 1276 (GH). COCHABAMBA: CERCADO: Mollemolle, 2800 m., Steinbach 4056 (GH). 9. S. Neesiana Trin. & Rupr. in Mém. Acad. St. Pétersb. (ser. 6) Sci. Nat. 5(1):27 (1843). To 10 dm. tall, caespitose, mostly glabrous, the nodes usually pubescent to villous. Leaf-blades flat to folded, scabridulous beneath, the margins hispid-ciliate. Panicle to 2 dm. long, axis glabrous, branches and pedicels scabridulous to hispidulous. Glumes somewhat unequal, the outer longer, to 1.5 em. long, long- acute, the midribs often hispid. Lemma to 8 mm. long, pubescent to villous below, the upper portion glabrous, smooth to papillate-rugose, the apical crown often long. Awn twice-bent or twice-geniculate, to 1.5 cm. long, the lowest portion tightly twisted and pubescent, the central portion loosely twisted and puberulent, the terminal portion straight and puberulent to scabridulous. LA Paz: LARECAJA: Sorata, Rusby 237 (us). COCHABAMBA: CERCADO: Cochabamba, Holway 328 (Us). Extreme forms of S. Neesiana and S. mucronata appear quite distinct, but enough intergradation occurs to make their specific distinctness appear somewhat questionable. 10. S. inconspicua Presl, Rel. Haenk. 1: 277 (1830). Nassella flac- cidula Hack. in Fedde, Rep. Sp. Nov. 6: 154 (1908). Caespitose, to 4.5 dm. tall, glabrous. Leaf-blades flat or involute, the uppermost equaling or exceeding the inflorescence. Panicle to 10 cm. (occasion- ally to 18 cm.), narrow, the branches quite distant, axis and branches glabrous, branchlets and pedicels puberulent to scabridulous. Glumes subequal or the outer slightly longer, acute, several-veined, about 3.5 mm. long. Lemma more or less obovoid, 2.5-3 mm. long, sparsely pubescent. Awn to 1.5 em. long, once-geniculate, scabridulous, the lower portion twisted, the upper portion straight. L4 PAZ: MURILLO: La Paz, Holoway 600 (vs). ll. S. depauperata Pilger in Engler, Bot. Jahrb. 56, Beibl. 123: 23 (1920). Caespitose, spreading, to 5 dm. tall, mostly glabrous. Leaves short, the involute blades rarely equaling the base of the inflorescence. Panicle narrow, to 15 cm. long, brownish to purplish, axis glabrous, branchlets and pedicels from glabrous to pubescent, Glumes unequal, to 1 em. long, long-acute, the midribs minutely scabridulous, otherwise glabrous. Lemma fusiform, to 3 mm. long, pubescent (sometimes sparsely so), the apical crown quite short. Awn to 1.5 cm. long, geniculate, the lower portion tightly twisted and short-pubescent, the upper portion straight and puberulent to scabridulous. La Paz: MURILLO: La Paz, Holway 492 (vs). 1966] Flora of Bolivia — Foster 275 12. S. brachyphylla Hitche. in Contrib. U. S. Nat. Herb. 24: 275 (1925). Caespitose, to 4 dm. tall, glabrous. Leaves short, the blades involute, pungent, upper surface scabrid. Panicle lax, narrow, to 1 dm. long, axis and branches glabrous, branchlets and pedicels puberulent to short-hispid. Glumes somewhat unequal or subequal, to 1 em. long, acute or acuminate, the midrib of outer glume often sparsely appressed-hispid. Lemma fusiform, about 4 mm. long, or a little longer, appressed-villous, the apical crown short. Awn to 2.5 em. long, twice-geniculate, lowest portion tightly twisted and pubes- cent, central portion more loosely twisted and puberulent, upper portion straight, minutely puberulent to scabridulous. LA Paz: MURILLO: La Paz, Holway 433 (US); oMASUYOS: Yumari, 3850 m., Asplund 6557 (US). 13. S. dasycarpa Hitchc. in Contrib, U. S. Nat. Herb. 24: 281 (1925). To 3-4 dm. tall. Leaf-blades involute, glabrous on the lower surface, puberulous on the upper surface. Panicle rather lax, to 1.5 dm. long. Glumes equal, 5 mm. long, acuminate. Lemma about 3 mm. long, fusiform, sparsely villous, apical crown lacking, callus 0.5 mm. long. Awn 2 cm. long, twice-geniculate, the lower 2 segments appressed- villous. PoTosí: NOR LIPEZ: Chiguana, Asplund 6562 (Hitchcock). 14. S. curviseta Hitchc. in Contrib. U. S. Nat. Herb. 24: 282 (1925). Caespitose, to 5 dm. tall. Leaf-blades involute, glabrous or scabridu- lous beneath, pubescent above. Panicle lax, to 2 dm. long, axis glabrous, branches and pedicels pubescent to scabrid. Glumes sub- equal, to 1.5 em. long, long-acuminate. Lemma terete, to 8 mm. long, sparsely villous to glabrate, apical crown obscure, ciliate, callus to 3.5 mm. long. Awn to 7 em. long, twice-geniculate, the lowest portion twisted and villous, the terminal segment flexuose, decreasingly villous toward the end. Potosi: porco: Uyuni, 3700 m., Asplund 6551 (US; type); SUR CHICHAS: Atocha, 3700 m., Asplund 6550 (vs), 6547 (US); NOR LIPEZ: Chiguana, 3700 m., Asplund 6554 (US). 15. S. obtusa (Nees & Mey.) Hitche. in Contrib. U. S. Nat. Herb. 24: 284 (1925). S. boliviensis Pilger in Engler, Bot. Jahrb. 56: 26 (1920). Caespitose, to 4 dm. high, glabrous. Leaf-blades involute, pungent, scabridulous, some equaling or exceeding the inflorescence- base. Panicle narrow, to 15 cm. long, axis, branches and pedicels scabridulous. Glumes equal or subequal, about 3 mm. long, broadly ovate, abruptly short-acute or subobtuse, the second glume often sub- tridentate. Lemma narrowly elliptic-fusiform, to 3 mm. long, glabrous save for a few short sparse hairs at or near the apex. Awn to 1 cm. long, not geniculate, not twisted, scabrid to scabridulous. LA PAZ: MURILLO: La Paz, 4100 m., Buchtien 184 (GH). COCHABAMBA: CERCADO: south side of Mt. Tunari, 3600 m., Steinbach 9786 (GH). 16. S. polyclada Hack. ex Stuck. in Anal. Mus. Nac. Buenos Aires, 21: 80 (1911). Straggling, the branches fascicled at the nodes. Leaf- blades involute to flat, pungent, short, Panicle lax, to 1 dm. long, 276 Rhodora [Vol. 68 axis and branches glabrous (sometimes scabridulous), pedicels puberu- lent to scabridulous. Glumes hyaline, about 3 mm. long, acute, sub- equal. Lemma to 2.5 mm. long, narrowly ellipsoid, glabrous above, sparsely appressed-pubescent below, the apex attenuate. Awn to 2.5 cm. long, variously bent but not geniculate, scabridulous. I have seen no Bolivian material on this species. In his treatment of the Andean grasses (1927) and his synopsis of South American Stipa (1925), Hitchcock cited Asplund 6563 as the basis for the inclu- sion of Bolivia in the range. The sheet of this number at Washington seems to me to be Stipa plumosa. Consequently, the occurrence of Stipa polyclada in Bolivia is problematical. 17. S. illimanica Hack. in Fedde, Rep. Sp. Nov. 11: 22 (1912). Caespitose, to 2.5 dm. tall. Leaf-blades flat or loosely involute, the margins scabrid. Panicle narrow, to 1.4 dm. long, axis glabrous. Glumes subequal, to 6 mm long, acute, 3-nerved. Lemma to 4-4.5 mm. long, short-pilose, the apex scabrid, apical crown lacking, callus short (0.5 mm.), pubescent. Awn to 3 cm. long, geniculate, the base loosely twisted and pubescent, the terminal portion puberulent to sca- brid. LA PAZ: MURILLO: Cotana, 2450 m., Buchtien 3134 (US; type- number). 18. S. Holwayi Hitchc. in Contrib. U. S. Nat. Herb. 24: 287 (1925). To 1 m. tall. Leaf-blades flat or involute, narrow, glabrous beneath, short-pubescent on upper surface. Inflorescence a narrow, lax panicle to 2.5 dm. long, axis glabrous, branches and pedicels shortly puberu- lous to hispid, especially near the apex of panicle. Glumes to 1.1 cm. long, unequal, the outer 1.5 mm. longer than the inner, 3-nerved, acuminate. Lemma to 5-6 mm, long, the callus to 1 or 1.5 mm. long and densely pubescent, fusiform, villous, the apex with an obscure crown. Awn to 4.5 em. long, twice-geniculate, the 2 lower portions twisted and scabrid-pubescent, the terminal portion to 2-3 cm. long, flexuose, glabrous to scabridulous. ORURO: ABAROA: Challapata, 2900 m., Asplund 6546 (US). COCHABAMBA: CERCADO: Cochabamba, Holway 380 (US; type). 19. S. florulenta (Pilger) Parodi in Rev. Mus. La Flata (n. s.) Bot. 6: 307 (1944). Oryzopsis florulenta of the Catalogue. To 1 m. tall. Leaf-blades flat or somewhat involute, scabridulous. Panicle lax, to 2.5 dm. long (sometimes to 4 dm. ?), axis more or less glabrous, branches and pedicels hispid-scabridulous. Glumes subequal, to 4 mm. long, acute to acuminate, 3-nerved. Lemma about 3-4 mm. long, obovoid, apex broad and truncate without a crown or ciliation, sparsely pilose to almost glabrous. Awn to 2 em. long, not geniculate, the scabridulous base somewhat loosely twisted, the remainder somewhat flexuose or straight. La PAZ: LARECAJA: Catarguata, 3000 m., Man- don 1270 bis (GH; mixed with Nassella pubiflora). 20. S. Trollii Pilger in Notizbl. 11: 777 (1933). To 4-6 dm. tall. Leaf-blades somewhat involute, the upper surface scabridulous. 1966] Flora of Bolivia — Foster 277 Panicle to 1.2 dm. long, the axis glabrous. Glumes 5-6 mm. long, lanceolate, the midrib somewhat excurrent. Lemma 4 mm. long, sparsely short-pilose. Awn to 1.5 cm. long, once-geniculate, glabrous. Potosi; porco: Salasala-Tasnapalco, 3400 m., Troll 3374. No material of this species has been seen. This descrip- tion is adapted from the original description. 49, Aristida L. See Caro in Kurtziana, 1: 123-206 (1961). Annual or perennial, Leaf-blades flat or involute. Inflorescence a compact to lax panicle. Spikelets 1-flowered, disarticulation above the glumes, the callus sharply pointed. Glumes unequal to equal or sub- equal. Lemma terete to fusiform, with 3 apical awns, these free to the base or nearly so, or basally united in a short or long twisted column. a. Callus bidentate at tip. |... 4. A, riparia. a. Callus entire. b. Glumes subequal or equal, or the outer longer than the inner. c. Glumes very unequal, the outer about twice the length of the MMC uL CARA a afeiecuis nttandtt za exis D AES 5. A. mendocina. c. Glumes subequal or equal. d. Leaf-blades involute. .............. e 6. A. enodis. d. Leaf-blades flat (at least at base). «se 7. A. circinalis. b. Glumes unequal, the outer shorter than the inner, e. Lemma to 3 mm. long. ........." 1. A. capillacea. e. Lemma to 8-10 mm. long. f. Lemma-awns united in a column 2-3 em. long. ...... cr a so adoauanenensndenunnenoresnncirariantaneas: OE Loa AIMO. f. Lemma-awns not united in a column. g. Longest glumes to 7 mm. long. ...........- 2. A. adscensionis. g. Longest glumes to 1.5 em. long. seeesessreeee 8. A. laevis. 1. Aristida capillacea Lam. Tabl. Encycl. 1: 156 (1791). A. Man- doniana Henr. in Meded. Rijks Herb. Leiden, no. 40: 55 (1921). Annual to 2 dm. high. Leaf-blades narrow, somewhat involute, scabri- dulous on upper surface, with a few scattered long hairs near the base. Panicle narrow but rather lax, to 8-10 cm. long, axis and branches glabrous, pedicels glabrous to puberulent. Glumes unequal, the outer shorter, to 4 mm. long, acute, glabrous, the prominent midrib scabridulous. Lemma to 3 mm. long. Awns united basally in a pu- berulent column, the free portions to 4-5 mm. long, puberulent to scabridulous. LA PAZ: SUR YUNGAS: Guanai, 1600 m., Rusby 208 (GH). SANTA CRUZ: SARA: Buenavista, 500 m., Steinbach 6140 (GH). 2. A. adscensionis L. Sp. Pl. 82 (1753). Caespitose annual, glabrous, the culms branching from the lower nodes, to 3 dm. tall. Leaf-blades involute. Panicle to 1.5 dm. long, compact to spreading, axis, branches 278 Rhodora [Vol. 68 and pedicels usually scabridulous. Glumes unequal, the outer shorter, to 7 mm. long, acute, the midrib scabrous. Lemma to 8 mm. long, the apex long-attenuate, the upper portion puberulent-scabrid. Awns not united basally into a column, to 2 cm. long, the central awn exceeding the laterals, markedly scabrous. COCHABAMBA: CERCADO: Cocha- bamba, Hitchcock 22817 (GH). Potosi: SUR CHICHAS: Villazón, 3500 m., West 6368 (GH). LA PAZ: MURILLO: Cotana, 2450 m., Buchtien 208 (GH); Obrajes, 3300 m., Buchtien 140 (GH). 3. A. implexa Trin. in Mém. Acad. St. Pétersb. VI, Sci. Nat. 2(1): 48 (1836). Perennial, to 6 dm. tall. Leaf-blades flat or involute, the upper surface densely pilose near the base. Panicle to 2 dm. long, axis, branches and pedicels scabrid. Glumes unequal, the outer much shorter, to 2 cm. long, scabridulous, long-acute. Lemma to 8 mm. long, glabrous, the apex attenuate. Awns united basally into a twisted column to 2-3 cm. long, the free portions to 3-5 em. long, the central awn longer than the laterals, scabridulous. SANTA CRUZ: SARA: Buenavista, Steinbach. 5352 (GH), 7029 (GH). 4. A. riparia Trin. in Mém. Acad. St. Pétersb. VI, Sci. Nat. 2(1): 48 (1836). Perennial, caespitose, to 8 dm. high. Leaf-blades flat or involute, rather thick. Panicle to 1.5 dm. long, axis, branches and pedicels scabridulous. Glumes unequal or subequal, the outer longer, awned, to 1.4 em. long (including the awn), scabridulous, the midrib of the outer glumes prominently so. Lemma to 6 mm. long, glabrous, the callus bidentate at the tip. Awns united basally in a twisted column to 1 em. long, the free portions equal, to 2 cm. long, scabri- dulous. L4 PAZ: SUR YUNGAS: Chulumani, 1600 m., Hitehcock 22702 (GH). 5. A. mendocina R. A. Phil. in Anal. Univ. Chile, 36: 205 (1870). A. inversa Hack. in Fries in Ark. Bot. 8(8): 37 (1908). Perennial, caespitose, to 6 dm. high. Leaf-blades somewhat involute, scabridulous on the upper surface. Panicle lax, narrow, to 2 dm. long, axis, branches and pedicels scabridulous. Glumes very unequal, the first to 11 mm. long, midrib scabrid, the second to 5 mm. long, scabridulous, both elumes often shortly mucronate, Lemma terete to fusiform, somewhat scabridulous on the upper portion, to 9 mm. long. Awns free to the base or nearly so, the central awn to 3 cm. long, the laterals slightly shorter, scabridulous. SANTA CRUZ: SARA: Buena- vista, Steinbach 1927 (GH). I have followed Hitchcock in uniting A. mendocina and A. inversa, but Caro has shown some differences which may warrant at least varietal segregation. 6. A. enodis Hack. in Fedde, Rep. Sp. Nov. 11: 21 (1912). A. Pflanzii Mez in Fedde, Rep. Sp. Nov. 17: 151 (1921). ? A. A splundii Henr. in Meded. Rijks Herb. Leiden, no 54: 42 (1926). Perennial, caespitose, to 4 (-5) dm. tall Leaf-blades involute, often somewhat scabridulous. Panicle rather dense, often purple, to 10 em. long. 1966] Flora of Bolivia — Foster 279 Glumes equal or subequal, to 1 cm. long. scabrid to nearly glabrous, the scabrid midrib excurrent as a mucro. Lemma narrowly fusiform, to 1 em. long, glabrous. Awns distinct to the base, to 3 cm. long, minutely scabridulous. LA PAZ: LARECAJA: prope Munaypata, 2700 m. Mandon 1277 (GH). SANTA CRUZ: SARA: Buenavista, 400 m., Steinbach 6807 (GH). 7, A. circinalis Lindm. in Sv. Vet. Akad. Handl. 34(6): 13 (1900). ? A. Friesii Henr. in Meded. Rijks Herb. Leiden, no. 54: 186 (1926). Perennial, caespitose, to 8 dm. tall. Leaf-blades long, flat near the base, becoming involute-filiform, glabrous. Panicle to 2 dm. long, axis, branches and pedicels usually glabrous. Glumes subequal or the outer longer, to 8 mm. long, glabrous, acute, or the midrib excurrent as a mucro. Lemma fusiform, to 7 mm. long, glabrous. Awns sub- equal, free to base, to 3 cm. long, minutely scabridulous. LA PAZ: SUR YUNGAS: Chulumani, Hitchcock 22711 (US). 8. A. laevis (Nees) Kunth, Enum. Pl. 1: 192 (1838). A. complanata of the Catalogue. A. longiramea var. boliviana, Henr. in Meded. Rijks Herb. Leiden, no. 40: 56 (1921). Perennial, caespitose, to 7 dm. high. Leaf-blades flat, somewhat scabridulous. Panicle narrow, to 2.5 dm. long; axis, branches and pedicels scabridulous to scabrid. Glumes equal or subequal, the outer shorter, to 1.5 em. long, aristate or mucro- nate, the midrib scabrid. Lemma tubular-fusiform, to 1 cm. long. Awns free to base, subequal, about 3 cm. long, scabridulous. SANTA CRUZ: SARA: Buenavista, 500 m., Steinbach 5355 (GH). 50. Tragus Hall. Tragus racemosus (L.) All. Fl. Pedem. 2: 241 (1785). Weedy annual. Stems somewhat geniculate, to 17 cm. high. Leaf-blades flat, lanceolate, to 2.5 cm. long, 6 mm. wide, acute, ciliate. Inflorescence a terminal spike, spikelets 1-flowered, paired, contiguous by flat faces, first glume absent, second glume about 3-4 mm. long, ovate, convex, the margin and three nerves bearing stout hooked spines, lemma and palea thin-textured. LA PAZ: LARECAJA: Sorata, Mandon 1268 (GH); NOR YUNGAS: Milluguaya, 1300 m., Buchtien 4176 (GH); SUR YUNGAS: Chulumani, 1600 m., Hitchcock 22649 (GH). 51. Aegopogon Humb. & Bonpl. Annuals. Culms to 20 cm. long (rarely to 30 cm.), spreading or decumbent. Leaf-blades short, linear, flat, 1-2 mm. wide. Spikelets 1-flowered, pectinately arranged, in groups of 2 or 3, 1 perfect and nearly sessile, the 1 or 2 laterals pedicellate and either staminate or neuter, often purple; glumes narrow, half the length of the lemmas or less, scabridulous, with a rather short delicate awn, lemma with a long central awn and 2 shorter lateral awns, palea with 2 short awns. 280 Rhodora [Vol. 68 Spikelets in groups of 3. sss 1. Ae. cenchroides. Spikelets in groups of 2. sss 2. Ae. bryophilus. 1. Aegopogon cenchroides Humb. & Bonpl. ex Willd. Sp. Pl. 4: 899 (1806). Leaf-blades scabridulous. Fertile spikelets (excluding the awns) about 3 mm. long. Central awn of lemma to 8 mm. long. La PAZ: NOR YUNGAS: Unduavi, Buchtien 2578 (Hitchcock). CocHa- BAMBA: CERCADO: Cochabamba, Hitchcock 22820 (Hitchcock). 2. Ae. bryophilus Doell in Mart. Fl. Bras. 2(3): 239 (1880). Ae. geminiflorus var. muticus Pilger in Engler, Bot. Jahrb. 27: 25 (1899). Ae. Fiebrigii Mez in Fedde, Rep. Sp. Nov. 17: 145 (1921). Leaf- sheaths glabrous to scabridulous; blades scabridulous. Fertile spike- lets (excluding the awns) about 3 mm. long. Central awn of lemma to 4-6 mm. long. LA Paz: LARECAJA: Cerro del Iminapi, 2650-3100 m., Mandon 1296 (GH); Sorata, Bang 1307 (GH; type-number of Ae. geminiflorus var. muticus). COCHABAMBA: TOTORA: Vacas, 3500 m., Steinbach 3965 (GH). TARIJA: ARCE: Camacho, Fiebrig 2865 (Hitch- cock). 52. Leptochloa Beauv. Annual or perennial, the culms decumbent or erect, 3-8 dm. or even to 1 m. high, leaf-sheaths glabrous, glaucous, scabridulous or even sparsely pilose, blades flat, glabrous to sparsely pilose, rarely equaling or exceeding the mature inflorescence. Inflorescence a terminal, nar- row to spreading panicle composed of few to many racemes, Spikelets usually many, few- to many-flowered, unilateral on the rachis, the rachilla disarticulating above the glumes, shortly pedicellate, ap- pressed and somewhat imbricate; glumes unequal to nearly equal, l-nerved; lemmas broad and notched at apex or acute, sometimes awned or mucronate, 3-nerved, glabrous, or pubescent between the lateral nerves and the margin, at least at the base, the uppermost lemma reduced. a. Spikelets at least 4 mm. long, usually to 7-9 mm. long. b. Lemmas much exceeding the minutely ciliolate palea. ................ RM 1. L. uninervia. b. Lemmas about equal to, sometimes exceeded by, the distinctly ciliate palea. coeccccccccsccsssscsssssscesssscessecscsssessversestesteceeccs 2. L. dubia. a. Spikelets less than 4 mm, long, never to 7-9 mm. long. c. Leaf-sheaths and blades glabrous, glaucous or rarely scabri- dulous, not pilose. vices 3 L. virgata. c. Leaf-sheaths and blades not glaucous, sparsely long-pilose. ........ RM 4. L. domingensis. l. Leptochloa uninervia (Presl) Hitche. & Chase in Contrib. U. S. Nat. Herb. 18: 383 (1917). Annual. Leaf-sheaths glabrous, blades scabrid. Spikelets (4-) 7-8 mm. long. First glume to 2.5 mm. long, the central nerve carinate and scabrid, acute; second glume 3 mm. long, scabrid on the central nerve, acute, sometimes mucronulate. 1966] Flora of Bolivia — Foster 281 Fertile lemmas about 4 mm. long, much exceeding the palea, palea minutely ciliolate. COCHABAMBA: CERCADO: Cochabamba, 2560 m., Steinbach 8746 (GH). 2. L. dubia (HBK). Nees in Syllog. Pl. Ratisb. 1: 4 (1824). Peren- nial. Leaf-blades scabrid. Spikelets to 8-9 mm. long. First glume about 2.5 mm. long; second glume very broad, about 3 mm. long. Fertile lemma to 4-5 mm, long, obtuse, emarginate, with a short central mucro, often pubescent basally between the lateral nerves and the margin. Palea nearly as long as the lemma, occasionaliy exceeding it, densely ciliate. LA PAZ: MURILLO: Cotana, 2450 m., Buchtien 3136 (GH). 3. L. virgata (L.) Beauv. Ess. Agrost. 166 (1812). Perennial. Leaf- sheaths and blades glabrous, glaucous, or rarely both scabridulous. Spikelets 3-3.5 mm. long. First glume about 1.5 mm. long; second glume 2 mm. long, both glumes acute, seabridulous on the central nerve. Lowest fertile lemma about 2.5 mm. long, occasionally the lowest 1 or 2 with short awns, usually the lowest puberulent along the lateral nerves. Palea nearly as long as the lemma, ciliolate. SANTA CRUZ: SARA: Río Surutt, 400 m., Steinbach 6834 (GH). 4. L. domingensis (Jacq.) Trin. Fund. Agrost. 133 (1820). Peren- nial. Leaf-sheaths and blades sparsely pilose, not glaucous. Spikelets 2-3.5 mm. long. First glume about 1.5 mm. long; second glume about 2 mm. long. Fertile lemma about 2.5 mm. long, the lowest 1 or 2 awned, the awns to 2 mm. long, margins long-ciliate. Palea some- what shorter than the lemma, not ciliolate or only faintly so at the apex. LA PAZ: NOR YUNGAS: Coripata, Hitchcock 22692 (GH). SANTA Cruz: SARA: Dolores, 450 m., Steinbach 1891 (GH); Buenavista, 450- 500 m., Steinbach 1179 (GH), 5222 (GH), 6730 bis (GH). 53. Gouinia Fourn. See Swallen in Amer. Journ. Bot. 22: 31-41 (1935). Tufted perennials, the culms erect, to 1-1.5 m. high, leaf-sheaths glabrous, leaf-blades scabrid beneath, to 8-14 mm. wide, not equaling the inflorescence. Inflorescence an open racemose panicle. Spikelets few-flowered, the uppermost reduced, in racemes to 15 cm. long, pedi- cellate; rachilla glabrous; glumes subequal or the second longer, 3-nerved; lemmas longer than glumes, densely villous on the nerves, awned, the awns scabridulous. Second glume rather obtuse, long-mucronate; panicle-branches bearing spikelets from the base; awn to 1.6 cm. long. ................... eene smi ENEMIES I 1. G. brasiliensis. Second glume acute, not mucronate; panicle-branches bearing spikelets from the middle upward; awn about 5 mm. long. ........... pc MURDER NN ERE cnet ceee 2. G. latifolia. 1. Gouinia brasiliensis (S. Moore) Swallen in Amer. Journ. Bot. 22: 36 (1935). First glume 4 mm. long; second glume 6 mm. long. Lower 282 Rhodora [Vol. 68 lemmas 7-8 mm. long, the awns to 1.6 em. long. SANTA CRUZ: CORDIL- LERA: Puerto Suarez, Chase 11151 (GH). 2. G. latifolia (Griseb.) Vasey in Contrib. U. S. Nat. Herb. 1: 365 (1895). Glumes subequal, about 5 mm. long. Lower lemmas 7 mm. long, the awns about 5 mm. long. LA PAz: NOR YUNGAS: Chulumani, 1600 m., Hitchcock 22650 (GH). 54. Tripogon Roem. & Schult. Tripogon spicatus (Nees) Ekman in Ark. Bot. 11(4): 36 (1912). Densely tufted low perennial. Leaves filiform, glabrous, chiefly basal, much shorter than the culms. Inflorescence a single terminal spike, the entire culm and inflorescence usually less than 25 em. long (60 cm. in one instance). Spikelets 5-9 mm. long, several- to many- flowered, with more than 1 perfect floret, sessile in 2 rows on the axis but usually appearing unilateral, the rachilla disarticulating above the glumes and between the florets; first glume 2 mm. long or less, subacute, second glume 2.5 mm. long, obtuse; lemmas about 3 mm. long, 3-nerved, the central nerve somewhat carinate and pro- longed as a slender awn about 1 mm. long from between 2 small apical teeth, at least the lowermost lemmas with a small basal tuft of hairs; palea hyaline, much shorter than the lemma and enclosed by it. SANTA CRUZ: [province uncertain]: between Río Piray and Río Cuchi, Herzog 1449 (Hitchcock). 55. Eleusine Gaertn. Eleusine indica (L.) Gaertn. Fruct. & Sem. 1: 8 (1788). Weedy annual, the culms ascending to spreading, rarely to 1 m. high, but usually less than 5 dm. Leaves nearly glabrous, the sheaths hirsute near the apex and with a ring of hairs at the apex; leaf-blades flat or folded, to 8 mm. wide, some equaling or exceeding the inflorescence, glabrous. Inflorescence terminal, with 2-5 digitate or subdigitate spikes to 10 em. long, the spikelets in 2 rows but unilateral on the axis. Spikelets few- to several-flowered, with more than 1 perfect floret below sterile florets, sessile, 3-4 mm. long, the rachilla dis- articulating above the glumes and between the florets; glumes awn- less, strongly keeled, the keel scabrid, acute, first glume about 2 mm. long, the second glume about 3 mm. long; lemmas strongly keeled, the keels scabridulous, about 3 mm. long; palea bicarinate, the nerves green; caryopsis subglobose to subtrigonous, brown, finely and trans- versely striate. LA PAZ: LARECAJA: Hacienda Simaco, 1400 m., Buchtien 5332 (GH); NOR YUNGAS: Polo-Polo, near Coroico, 1100 m., Buchtien (in 1912) 238 (GH); Milluguaya, 1300 m., Buehtien (in 1917) 238 (GH). SANTA CRUZ: SARA: Río Surutü, 400 m., Steinbach 6837 (GH); Buenavista, 500 m., Steinbach 5149 (GH), 1871 (GH). 1966] Flora of Bolivia — Foster 283 56. Dactyloctenium Willd. Dactyloctenium aegyptium (L.) Willd. Enum. Hort. Berol. 1029 (1809) [as aegyptiacum]. Weedy, rather pilose annual, spreading by stolons and forming mats, culms erect, to 6 dm. high, leaf-sheaths glabrous or nearly so, the flat leaf-blades pilose and ciliate, shorter than the inflorescence. Inflorescence terminal, of 2-5 digitately ar- ranged spikes 2-5 em. long, the rachis exceeding the spikelets as a short subulate point, spikelets in 2 rows, unilateral on the axis. Spikelets several-flowered, sessile, imbricate, about 9 mm. long, the rachilla disarticulating above the glumes and between the florets; first glume about 3 mm. long, 1l-nerved, acute, the second glume longer, with a short scabridulous awn arising from below the apex; lemma about 6 mm. long, 3-nerved, the central nerve keeled, scabri- dulous, the apex acuminate to short-awned; palea a little shorter than the lemma; caryopsis nearly globose, wrinkled or ridged. SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 3132 (GH). 57. Cynodon L. Rich. Cynodon Daetylon (L.) Pers. Synops. Pl. 1: 85 (1805). Low peren- nial, the stolons spreading extensively and rooting at the nodes, the culms produced at the nodes, with a few basal leaves, the sheaths glabrous, the short (2-3 cm.) blades scabridulous, the culms clothed in leaf-sheaths with greatly reduced free tips; fertile culms 10-40 em. high. Inflorescence terminal, digitate, 3-6 spikes to 5 cm. long, the sessile spikelets in 2 rows along the axis, unilateral, imbricate. Spikelets 1-flowered, the rachilla disarticulating above the glumes and produced as a slender bristle behind the palea; glumes almost equal, to 3 mm. long, or the first glume shorter than the second, narrow, strongly keeled, the keel scabrid; lemma awnless, to 3 mm. long or a little longer, usually somewhat exceeding the second glume, navieulate, brown-hyaline, the central keel prominent and densely pubescent, the margins long-ciliate; palea somewhat shorter than the lemma. SANTA CRUZ: CERCADO: playa del Río Piray, 450 m., Stein- bach 3246 (GH). 58. Microchloa R. Br. Microchloa indica (L. f.) Beauv. Ess. Agrost. 13 (1812). Tufted annual, usually low in stature but sometimes the erect culms to 25 cm. high, leaves glabrous, the very narrow blades flat or folded, not equaling the inflorescence. Inflorescence a solitary terminal spike to 10 em. long, the axis bifariously puberulent on the side toward the spikelets, the spikelets unilateral on the axis. Spikelets 1-flowered, about 3 mm. long, sessile or subsessile; disarticulation above the glumes; glumes subequal, 2.5 mm. long, 1-nerved, acute; lemma and 284 Rhodora [Vol. 68 palea subequal, about 2 mm. long, acute, hyaline, the upper portion more or less densely covered by a long silky pubescence; caryopsis brown, subtrigonous, finely reticulate-striate. LA PAZ: MURILLO: La Paz, 3600 m., Buchtien 3130 (GH); Obrajes, 3450 m., Buchtien 9132 (GH). 59. Gymnopogon Beauv. Lcosely tufted perennials, the slender culms erect or spreading, 3-8 dm. high, leaves glabrous, the blades 2-5 cm. long and 2-5 mm. wide, reduced upward on the culm, never equaling nor exceeding the inflorescence. Inflorescence a terminal panicle of slender racemes 5-25 cm. long. Spikelets many, pedicellate, with 1-2 perfect florets below 1-3 sterile florets, rachilla disarticulating above the glumes; glumes subequal; fertile lemma long-awned, more or less terete. Racemes to 5 em. long, collected near the apex of the culm; spikelet 2-flowered. .......... seen thee terne trt 1. G. jubiflorus. Racemes 10-25 cm. long, distant along the main axis; spikelet 1- flowered, with 1-2 much reduced florets. .................... 2. G. spicatus. 1. Gymnopogon jubiflorus Hitchc. in Contrib. U. S. Nat. Herb. 24: 412 (1927). Glumes subequal, the second slightly longer than the first, 3.5-4 mm. long, lance-acuminate. Fertile lemma about 2 mm. long with an apical tuft of hairs to 1.5 mm. long, the awn about 8 mm. long. Palea about as long as the lemma. LA PAZ: LARECAJA: Guanay, Rusby 215 (us; type). 2. G. spicatus (Spreng.) O. Ktze, Rev. Gen. 3(2): 354 (1898). First glume 3.5 mm. long; second glume 4 mm. long. Fertile lemma very narrow, 2.5 mm. long, apical hairs very short, awn 8-10 mm. long. SANTA CRUZ: SARA: Buenavista, 450-500 m., Steinbach 1932 (am), 2357 (GH), 5386 (GH), 6977 (GH). 60. Chloris Swartz Annual or perennial, with decumbent to erect culms 2 dm. to 1 m. high, leaves glabrous or scabrid or sparsely pilose, the short blades to 1 cm. wide, not equaling the inflorescence. Inflorescence terminal, composed of digitately arranged spikes in 1 or 2 whorls, the sessile spikelets unilaterally arranged. Spikelets with 1 perfect floret below 1-2 (-8) reduced and rudimentary florets, rachilla disarticulating above the glumes; glumes unequal, lemmas mucronulate or awned (awnless in no. 1). a. Lemmas awnless. ..........eeeee s 1. C. distichophylla. a. Lemmas awned or mucronulate. b. Fertile lemmas not villous on keel (sometimes sparsely villous above), nor ciliate; sterile floret narrow, acute at apex. c. Spikelets about 5 mm. long; fertile lemma 3-5 mm. long. d. Spikes 6-10, 5-8 em. long. ....... s 4. C. Beyrichiana. 1966] Flora of Bolivia — Foster 285 d. Spikes 6-15, 7-15 em. long. ........ 5. C. halophila. c. Spikelets about 3 mm. long; fertile lemma about 2 mm. long. 6. C. radiata. ae0099909990090900900000002000009000099900900909009090005000099909099009090000090009 00090000999 b. Fertile lemmas villous on keel, at least on lower half (rarely glabrous), margins ciliate; sterile floret broad and truncate at the apex. e. Lemma long-ciliate from base to apex, the hairs 3 mm. long; perennial. .............. eee enn enne 2. C. polydactyla. e. Lemma only shortly ciliate, or glabrous at base, the hairs increasing in length upward, the keel rarely glabrous; annual. 3. C. virgata. auacecscq)o c[ t OUR 0000990000000 020400000020409000049909009000000000009099902000000009099200 1. Chloris distichophylla Lag. Gen. & Sp. Nov. 4 (1816). Perennial. Spikelets about 3 mm. long. First glume somewhat less than 2 mm. long, 1-nerved, acute; second glume about 2.5 mm. long, 1-nerved, often mucronulate. Fertile lemma about 3 mm. long, mucronulate, the margin ciliate with long hairs decreasing in length upward. Palea shorter than the lemma. Sterile lemma shorter than the fertile, glabrous, obtuse at apex but not broadest at apex. La Paz: Yungas, Bang 267 (GH); NOR YUNGAS: Polo-Polo, near Coroico, 1100 m., Buchtien 228 (GH); SUR YUNGAS: Chulumani, 1600 m., Hitchcock 22663 (GH). SANTA CRUZ: SARA: Buenavista, 450-500 m., Steinbach 5359 (GH), 6867 (GH). 2. C. polydactyla (L.) Swartz, Prodr. 26 (1788). Spikelets to 3.5 mm. long. First glume about 2.5 mm. long, the single nerve sca- bridulous; second glume about 3.5 mm. long, the central nerve scabrid. Fertile lemma about 2.5 mm. long, with a 3.5 mm. awn. LA PAZ: NOR YUNGAS: Milluguaya, 1300 m., Buchtien 739 (GH); Coripata, Hitchcock 22689 (GH). 8. C. virgata Swartz, Fl. Ind. Occ. 1: 203 (1797). Spikelets about 4 mm. long. First glume 2 mm. long; second glume 4 mm. long, mucronulate to short-awned. Fertile lemma about 3 mm. long, the awn about 1.1 cm. long. Palea about 3 mm. long, the 2 nerves lateral and ciliolate, COCHABAMBA: CERCADO: Valle de Cochabamba, 2600 m., Steinbach 8796 (GH). 4. C. Beyrichiana Kunth, Rév. Gram. 1: 289 (1830). Perennial. Spikelets about 5 mm. long. First glume 3 mm. long, long-acute, the nerve scabridulous; second glume about 4.5 mm. long, long-acute, the nerve scabridulous. Fertile lemma about 4.5 mm long, strongly biden- tate, the awn to 1.7 cm. long. Palea shorter than lemma, densely ciliolate. LA PAZ: MURILLO: La Paz, 3300 m., Bang 82 (GH). COCHA- BAMBA: MIZQUE: Vilavila, 2500 m., Eyerdam 24987 (GH). TARIJA: CERCADO: Tarija, 2000 m., Cárdenas 200 (GH). 5. C. halophila L. R. Parodi in Revist. Argent. Agron. 12: 45 (1945). Perennial. Spikelets 5-6 mm. long. First glume 2.5 mm. long; second glume 4-4.5 mm. long. Fertile lemma about 5-5.5 mm. 286 Rhodora [Vol. 68 long, bidentate, the awn 2 cm. long, the margins sparsely ciliate above. LA PAZ: MURILLO: Obrajes, 3400 m., Buchtien 573 (GH). The original description cited several collections from the depart- ments of Cochabamba and Tarija. I have not seen these. 6. C. radiata (L.) Swartz, Prodr. 26 (1788). Annual. Spikelets about 3 mm. long, but sometimes up to 5 mm. First glume about 1.5 mm. long; second glume about 3 mm. long, long-acute to very shortly awned, margins sparsely ciliate above. Palea as long as the lemma, bidentate. LA PAZ: NOR YUNGAS: near Coripata, 1700 m., Buchtien 8056 (GH), Bang 2173 (GH). 61. Trichloris Fourn. Tufted perennials, the erect culms to 1 m. high, usually shorter, leaves scabrid on sheaths and blades, blades 2-10 mm. wide, not ex- ceeding the inflorescence. Inflorescence a group of terminal spikes 5-15 em. long. Spikelets 2-5-flowered, fertile florets below the sterile, the rachilla disarticulating above the glumes, subsessile, in 2 rows, uni- lateral on the rachis; first glume shorter than the second, mucronate or shortly awned; lemmas with a tuft of hairs at the base, 1-3-awned, all awns scabridulous. Leaf-blades to 4 mm. wide; lemmas 3-awned, the awns all long; spikelets 2-flowered, the upper floret much reduced; inflorescence DEM 1. T. mendocina. Leaf-blades to 10 mm. wide; lemmas with lateral awns much reduced or obsolete; spikelets 3-5-flowered, the uppermost much reduced; inflorescence much more open. ....cccccsccccsseccecerseeceseesees 2. T. pluriflora. 1. Trichloris mendocina (Phil.) Kurtz in Mem. Fac. Cienc. Exact. Univ. Córdoba, 1897: 37 (1897). Spikelet about 3-4 mm, long (ex- cluding the awns). First glume about 1 mm. long, very narrow, the awn 1.5 mm. long; second glume 2 mm. long, the awn 3 mm. long. Lowest lemma to 4 mm. long, pubescent on the upper portion, awns to 1.2 em. long. SANTA CRUZ: CERCADO: Mataral, Herzog 1804 (Hitch- cock). 2. T. pluriflora Fourn. Mex. Pl. 2: 142 (1886). Spikelet about 6 mm. long (excluding the awns). First glume about 1.5 mm. long, the awn much shorter; second glume about 2.5 mm, long, the awn about 0.5 mm. long. Lowest lemma 4 mm. long, ciliate, awns about 6 mm. long. SANTA CRUZ: VALLEGRANDE: Comarapa, Herzog 1928 (Us). 62. Bouteloua Lag. See Griffiths in Contrib. U. S. Nat. Herb. 14: 343-428 (1912). Annual or perennial, the culms erect or decumbent, mostly 10-20 cm. high, occasionally to 1 m. high, leaves generally glabrous, some- times sparsely hirsute near the edge of the sheath and on the base 1966] Flora of Bolivia — Foster 287 of the blade, always shorter than the inflorescence, the blade narrow. Inflorescence terminal, a solitary spike or few to many short racemose spikes. Spikelets sessile, 1-flowered, with a rudimentary floret above the perfect one, unilaterally arranged; glumes present, unequal; fertile lemma 3-5-toothed, awned or awnless. a. Spike 1, persistent, the spikelets falling separately. ...... mmm 1. B. simplex. a. Spikes few to several, each spike falling entire. b. Awns 2-3 em. long. ......: HH 2. B. megapotamica. b. Awns much shorter. c. Spikes at maturity erect or spreading; rachis sharply pointed at base and retrorsely bearded. „sesser 3. B. aristidoides. c. Spikes at maturity deflexed, rachis-base neither pointed nor bearded. ........ eere nennen nnne nnne nnne nnne nnn 4. B. curtipendula. 1. Bouteloua simplex Lag. in Varied. Cienc. 29(4): 141 (1805). Annual First glume 3.5 mm. long, faintly 1-nerved; second glume to 5 mm. long, acute, strongly l-nerved. Fertile lemma with a tuft of hairs at base, to 3 mm. long, the central awn 3 mm. long, sparsely pilose at base and along midrib. LA Paz: MURILLO: La Paz, 3300 m., Bang 81 (GH); LARECAJA: vic. Sorata; Lorecasa, 2650-3200 m., Mandon 1325 (GH). Potosi: CERCADO: Potosí, 4000 m., Cárdenas 197 (GH). TARIJA: AVILES: Puna Patanca, 3700 m., Fiebrig 2629 (GH). 2. B. megapotamica (Spreng.) O. Ktze. Rev. 3(2): 341 (1898). Perennial. Spikelet with basal tufts of hairs. First glume 3 mm. long with a 3 mm. awn, 1-nerved; second glume 7 mm. long with a 3 mm. awn, very strongly 1-nerved. Fertile lemma 4 mm. long with the central awn 1 cm. long. Palea longer than the lemma, bidentate. Reduced florets with awns to 3 cm. long. TARIJA: ARCE: Padcaya, 2100 m., Fiebrig 2552 (GH). 8. B. aristidoides (HBK.) Griseb. Fl. Brit. W. Ind. 537 (1864). Annual. First glume 2.5 mm. long, acute, l-nerved; second glume 7 mm. long, puberulent at base, scabridulous above, awnless. Fertile lemma about 6 mm. long, glabrous, awnless. Palea as long as lemma, bidentate. Awns of sterile lemmas to 6 mm. long. TARIJA: Gran Chaco, Fries 1691 ( Hitchcock). 4. B. curtipendula (Michx.) Torr. in Marcy, Red River Rept. 300 (1853). Perennial, the culms up to 1 m. high. First glume 3 mm. long, attenuate into a 2 mm. awn, the prominent central rib hispi- dulous; second glume 7 mm. long, awnless, bidentate, with a short point between the teeth, scabridulous near the margin and on the prominent central nerve. Fertile lemma about 6 mm. long, glabrous below, scabridulous above and on the 8 long teeth. Palea as long as lemma, bidentate, the upper half scabridulous. LA PAZ: LARECAJA: Sorata, Mandon 1380 (GH). COCHABAMBA: CERCADO: west of Cocha- bamba, 2700 m., Eyerdam 24875 (GH). TARIJA: ARCE: Padcaya, Fiebrig 3281 (GH), 3282 (GH). 288 Rhodora [Vol. 68 63. Munroa Torr. See L. R. Parodi in Revist. Mus. La Plata, 34: 171-193 (1934). Low-growing, more or less tufted, spreading annuals, 2-15 cm. high, the leaves very short, the sheaths very broad, blades a few cm. long, the apex pungent. Inflorescence condensed into heads or glomerules, these more or less concealed by the sheaths of the fascicled sub- tending leaves. Spikelets aggregated in groups of 2 or 3, the lower 3-4-flowered, the upper 2-3-flowered, the first floret in the spikelet perfect, the others sterile, usually 3 groups in a cluster, the rachilla disarticulating above the glumes; glumes usually quite unequal; lemmas 3-nerved, the nerves much or little prolonged as awns or mucros. Caryopsis dorsally compressed. a. Lowest fertile lemma 4-lobed with 3 awns, the central awn longest; back of lemma with 2 tufts of hairs below the central sinus of the lobes; 2 central lobes broader than the 2 lateral, the former rather obtuse, the latter acute; margins long-ciliate for over half their length; palea with puberulent nerves, ................ 1. M. andina. a. Lowest fertile lemma 2-3-toothed, the central tooth narrower than the laterals, its apex subaristate, the laterals mucronulate; palea glabrous. b. Lowest lemma papery at base, 2-toothed with a central awn, the back with 2 tufts of hairs, the margin long-ciliate below or above the middle, the hairs reaching neither apex nor base; palea broadest at apex. a, 2. M. decumbens. b. Lowest fertile lemma coriaceous at base, 3-toothed; dorsal tufts of hairs absent, the margin long-ciliate above the middle; palea broadest at the base. sss 3. M. argentina. 1. Munroa andina Phil. in Anal. Mus. Nac. Chile, 8: 90 (1891). Leaf-blades about 1 cm. long, 1 mm. wide. Glumes unequal, the second to 4 mm. long, mucronulate. Lowest lemma about 3.5 mm. long. La PAZ: PACAJES: Calacota, 3800 m., Asplund 2653. la. M. andina var. breviseta Hack. ex Stuckert in Ann. Conserv. & Jard. Bot. Genéve, 17: 294 (1914). 'This variety is distinguished by the lowest lemma with very short awns, hardly, if at all, exceeding the lemma-lobes. LA Paz: PACAJES: Charana, 4050 m., Asplund 2691. 2. M. decumbens Phil. in Anal. Mus. Nac. Chile, 8: 90 (1891). Leaf-blades to 2.5 cm. long and 2 mm. wide. Glumes unequal, the second to 4.5 mm. long. Lowest lemma about 4 mm. long, with a 3 mm. central awn between the 2 lobes. Potosi: NOR LIPEZ: Chiguana, 3700 m., Asplund 6504 (Us); PORCO: Uyuni, 3700 m., Asplund 6487 (us). 3. M. argentina Griseb. in Goett. Abh. 24: 300 (1879). Leaf-blades 2 em. long, 1 mm. wide. Glumes subequal, about as long as the lowest lemmas. Lowest lemma to 5 mm. long. Potosi: SUR CHICHAS: Atocha, 1966] Flora of Bolivia — Foster 289 9700 m., Asplund 6485 (US). CHUQUISACA: SUR CINTI: Camataquí, 2500 m., Fiebrig 2801. With three exceptions, all specimen-citations have been taken from Parodi's treatment of the genus. 64. Phalaris L. Annuals, the culms erect. Leaf-blades linear, flat, acute, from much shorter than to exceeding the inflorescence, to 6 mm. wide. Inflo- rescence a terminal spicate panicle. Spikelets strongly flattened, densely imbricate, the perfect floret terminal with 2 sterile lemmas below it, all concealed by the glumes; disarticulation above the glumes; glumes papery white with green stripes, the green keel broadly winged, or narrowly winged near the acute apex; sterile lemmas one-half to one-third as long as the appressed-pubescent coriaceous fertile lemma, the bicarinate palea enclosed by the fertile lemma. Culms 3-6 dm. tall, inflorescence ovate, broad, to 5 cm. long or less; glumes broadly winged, 7-8 mm. long, sparsely long-hirsute, the hairs usually appressed: ei ario a 1. P. canariensis. Culms 1-1.5 m. tall, sometimes shorter; inflorescence narrowly cylin- dric, 6-15 cm. long; glumes not winged or only narrowly so, espe- cially near the apex, 3.5-4 mm. long, shortly ciliolate on keel and 2 chief nerves odie E ce a NUN 2. P. angusta. 1. Phalaris canariensis L. Sp. Fl. 54 (1753). LA PAZ: MURILLO: La Paz, Buchtien 239 (Hitchcock); NOR YUNGAS: Coroico, Buchtien 3641 (Hitchcock). 2. P. angusta Nees ex Trin. Gram. Icon. 1: t. 78 (1827). LA Paz: LARECAJA: Sorata, Mandon 1246 (Hitchcock). 65. Leersia Swartz Leersia hexandra Swartz, Prodr. 21 (1788). Perennial, often sub- aquatic, scabrous or scabridulous, many culms decumbent, rhizomes slender and producing leafy stolons, fertile culms erect, sometimes to 2 m. tall, usually shorter. Leaf-blades narrow, linear-lanceolate, to 15-20 em. long and 5 mm. wide, long-acute. Inflorescence a narrow panicle to 15 cm. long, the branches appressed. Spikelets solitary, 1-flowered, the floret perfect, disarticulating from the very short pedicel, strongly flattened, glumes lacking, lemma and palea similar in appearance, to 4-5 mm. long, often pinkish or purplish, subequal in length and width, oblong to narrowly ovate, acute, with 3 prominent ribs, these often hispidulous or the entire lemma hispidulous, the margins coarsely ciliate. SANTA CRUZ: SARA: playa del Río Perdix, 450 m., Steinbach 7453 (GH); Buenavista, 450 m., Steinbach 2180 (GH), 7116 (GH). 290 Rhodora [Vol. 68 66. Luziola Juss. Luziola peruviana Gmel. Syst. Nat. 1: 637 (1791). Perennial, usually of wet or muddy areas or even in water, the slender culms to 40 cm. long, but usually much shorter, often decumbent, branching, fertile culms erect. Leaf-blades more or less linear, long-acute, to 30 em. long, usually much shorter, 1-4 mm. wide, exceeding the inflo- rescences. Spikelets 1-flowered, unisexual, disarticulating from the filiform pedicels, pistillate and staminate spikelets in separate panicles on the same plant, the staminate panicles terminal, the pistillate terminal and axillary, both short, to 6 em. long; pistillate spikelets 2-3 mm. long, the stigmas long-plumose, the staminate spikelets 4-7 mm. long; glumes absent in both; lemma and palea similar in appear- ance and subequal, thin-textured, with several to many nerves, the nerves obten scabridulous; caryopsis globose, rather shiny, finely striate, about 1 mm. long or longer, sometimes shortly beaked with the persistent base of the style. SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 1946 (GH), 6811 (GH). TARIJA: ARCE, La Merced, near Bermejo, 1700 m., Fiebrig 2189 (GH). 67. Pharus L. Perennials, often in wet places, the culms to 1 m. high, solitary and erect, or creeping at the base and rooting at the basal nodes, forming mats. Leaf-blades petiolate, the petiole once-twisted, elliptic- oblong or lanceolate, the principal veins connected by numerous and obvious small parallel eross-veins. Inflorescence a reduced to expanded small panicle; spikelets 1-flowered, appressed, paired, the small stami- nate spikelet pedicellate, the pistillate subsessile; 2 glumes present, dark-brown; disartieulation above the glumes; lemma terete, much exceeding the glumes, somewhat indurate, beaked, entirely or partially pubescent, the hairs somewhat uncinate, the margins incurved and concealing the palea. a. Lemma entirely pubescent. b. Culms creeping at base and forming mats; inflorescence usually reduced and few-branched. .................. esses 3. P. parvifolius. b. Culms erect, solitary; inflorescence larger and more open. ...... ccveuceeesecesessececcuscceucercestseeeesseeeseeeeceeseeesseeeecesecenseeeeensecensesensas 1. P. glaber. a. Lemmas pubescent only near the beak. .................... 2. P. latifolius. 1. Pharus glaber HBK. Nov. Gen. & Sp. 1: 196 (1816). Spikelets about 1 cm. long, about twice the length of the glumes. Leaf-blades to 25 em. long and 5 em, wide, abruptly short-acuminate. LA Paz: NOR YUNGAS: Milluguaya, 900 m., Buchtien 4173 (GH). 2. P. latifolius L. Syst. Nat. (ed. 10) 2: 1269 (1759). Spikelets 1.5-2 em. long, exceeding, but seldom twice the length of, the glumes. Leaf-blades generally broader. No Bolivian material which is un- 1966] Flora of Bolivia — Foster 291 questionably this species has been seen, but it seems probable that the species will be found there. 3. P. parvifolius Nash in Bull. Torr. Bot. Club, 35: 301 (1908). Snikelets less than 1.5 cm. long. Leaf-blades generally broadest below the middle, sometimes long-acuminate. LA PAZ: SUR YUNGAS: Covendo, White (M. E.) 1043 (Hitchcock). 68. Arundinella Raddi Tufted perennials, the erect culms from 1-2.5 m. high. Leaf-sheaths glabrous; leaf-blades shorter than or equaling the inflorescence, to 5 mm. wide, hirsutulous to scabrid along the nerves, or glabrous. Inflorescence a rather dense panicle, the pedicellate spikelets solitary. Spikelets 2-flowered, the lower floret staminate or sterile, the upper pistillate; first glume much shorter than the second, the apices divergent and often somewhat acuminate, strongly 3-nerved, the second glume at least twice the length of the sterile and fertile lemmas, or equal to the sterile lemma; disarticulation above glumes; fertile lemma shortly villous on the basal callus, scabridulous, bearing a geniculate awn about 1 cm. long or less, completely enclosing the palea. Culms to 2.5 m. high; inflorescence-branches and pedicels densely and retrorsely scabridulous; glumes 2-2.5 mm. long; lemmas 2-2.5 mm. long; awn basally twisted, about 2 mm. loni on 1. A. confinis. Culms usually less than 1 m. high; inflorescence-branches and pedicels glabrous to faintly scabridulous; glumes 3-5 mm. long; lemmas 1.5 mm. long; awn about 1 cm. long, not twisted basally. ..............- 2. A. Berteroniana. Mos ea Roce ks va suAqA So RRAORERASURSARRIOSEPERÓKAN SaMQLGOERRARLISQAS AL E20SANSENTS ESAE bach 6888 bis (GH). 2. A. Berteroniana (Schult.) Hitche. & Chase in Contrib. U. S. Nat. Herb. 18: 290 (1917). LA PAZ: LARECAJA: Mapiri, Buchtien 1175; SUR YUNGAS: Covendo, White (M. E.) 995. BENI: YACUMA; Rurrenabaque, Cárdenas (M. E.) 1166. Citations from Hitchcock. 69. Trichopteryx Nees Trichopteryx flammida (Trin.) Benth. ex Jacks. & Hook. f. Index Kew. 4: 1109 (1895). Tufted perennial, the culms 1.5-2 m. high. Leaf-sheaths glabrous to hirsute; leaf-blades sometimes exceeding the inflorescence, flat, to 7 mm. wide. Inflorescence terminal, a rather dense panicle, to 60 cm. long, the spikelets solitary, pedicellate, the pedicels bearing a few long hairs at the apex. Spikelets 2-flowered, the lower floret staminate, glabrous and awnless, the upper floret pistillate; first glume 4-5 mm. long, strongly 3-nerved, the nerves ciliolate, with 3-5 erect hairs 3-4 mm. long near the apex, second 292 Rhodora [Vol. 68 glume 4-5 mm. long, less strongly nerved, glabrous; disarticulation above the glumes; lemma of staminate floret about 6 mm. long, glabrous, nearly twice the length of the hyaline palea; lemma of pistillate floret 3-4 mm. long, densely long-pubescent, bearing a scabridulous awn up to 2 cm. long, this geniculate shortly above its base. LA PAZ: LARECAJA: San Carlos, Mapiri, 850 m., Buchtien 9 (GH). SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 1884 (GH), 5358 (GH), 6958 (GH). The authorship of this combination is often given as Bentham in Journ. Linn. Soc. Bot. 19: 59 (1882), but ex- amination shows that the combination was not actually made there. It was merely implied. 70. Tristachya Nees Tristachya chrysothrix Nees, Agrost. Bras. 460 (1829). Tufted perennial, the culms to 1 m. high, the leaf-sheaths hirsute to hispi- dulous or glabrous. Leaf-blades flat to involute, hirsute to almost glabrous, not exceeding the inflorescence, 'to 4 mm. wide (when flat- tened). Inflorescence a lax, nearly simple panicle, the branches puberulent to glabrous below, hirsute above, the hairs golden-brown, spikelets sessile in groups of 8, terminal on the branches. Spikelets 2-flowered, the lower floret staminate and awnless, the upper floret pistillate; first glume about 1 cm. long, 3-nerved, each lateral nerve with a row of short, dark brown tubercles, each bearing a long golden-brown hair, second glume to 1.5 cm. long, pubescent along the margins in the upper portion; disarticulation above the glumes; lemmas of staminate floret glabrous, about 1.2 cm. long, the palea 2-nerved, ciliate; lemma of pistillate floret about 7 mm. long, densely pubescent, the base surrounded by longer, white, silky hairs, bearing a twisted awn 4-7 cm. long, rising between 2 slender teeth, the lower portion brown and hispidulous, the upper portion green and more nearly scabridulous. SANTA CRUZ: VALLEGRANDE: Samaipata, 2000 m., Steinbach 3772 (GH). 71. Leptocoryphium Nees Leptocoryphium lanatum (HBK.) Nees, Agrost. Bras. 84 (1829). Perennial, to 1 m. high. Leaves glabrous, the blades flat or often involute, to 8 mm. wide, linear, Inflorescence a rather lax, narrow, racemose, terminal panicle, the rachis, branches and pedicels glabrous. Spikelets 2-flowered, ihe lower floret a sterile lemma, the other perfect, disarticulation below the glumes. First glume absent; second glume to nearly 5 mm. long, strongly several-nerved, long-pilose, the hairs mostly confined to the nerves. Sterile lemma as long as the glume and similar in appearance and pilosity. Fertile lemma about as long 1966] Flora of Bolivia — Foster 293 as the sterile, thin-textured, the margins sparsely ciliate, the apex slightly erose-fimbriate. Palea nearly as long as the fertile lemma, thin, not carinate. LA PAZ: LARECAJA: Hacienda Casana, 1600 m., Buchtien 7146 (GH). SANTA CRUZ: SARA: Buenavista, 450 m., Stein- bach 3269 (GH), 6639 (GH), 6640 (GH). 72. Trichachne Nees Perennials, the bases somewhat swollen and felty-pubescent or sealy. Leaf-sheaths glabrous or papillate-hirsute; leaf-blades glabrous or scabridulous. Inflorescence a panicle of shortish racemes. Spikelets paired, pedicellate, 2-flowered, the lower floret a sterile lemma, the upper floret perfect, silky-villous; disarticulation below the glumes. Glumes very unequal; second glume and sterile lemma similar and subequal; fertile lemma firm-textured and acuminate; palea similar but narrower. a. Sterile lemma silky-villous dorsally. .........................- 1. T. insularis. a. Sterile lemma glabrous dorsally. b. Leaf-sheaths glabrous; blades to 7-8 mm. wide; fertile lemma about B mm. Jong. 01922505 de steep reni ds 2. T. sacchariflora. b. Leaf-sheaths sparsely papillate-hirsute (at least when young); blades to 5 min. wide, glabrous to puberulent; fertile lemma 2.b mm. LONG. ............-- ee ee eena eee hann nuoto enano toot aee en oa 3. T. californica. 1. Trichachne insularis (L). Nees, Agrost. Bras. 86 (1829). Plants to 1.5 m. tall Leaf-sheaths rather sparsely papillate-hirsute; leaf- blades to 1.5 cm. wide, scabridulous above. Panicle to 3 dm. long, the racemes to 15 em. long, tawny when mature. Spikelets to 4 mm. long. First glume over 0.5 mm. long, obtuse; second glume 3 mm. long, acute, silky-villous. Sterile lemma 4 mm. long, not glabrous dorsally. Fertile lemma 3.5 mm. long. LA PAZ: NOR YUNGAS: Millu- guaya, 1200 m., Buchtien 4188 (GH), COCHABAMBA: CHAPARE: Anta- huacana, Espíritu Santo, 750 m., Buchtien s. n. (GH). 2. T. sacchariflora (Raddi) Nees, Agrost. Bras. 87 (1829). T. affinis Swallen in Rhodora, 65: 355 (1963). Plants to 8 dm. tall. Leaf- sheaths glabrous; leaf-blades to 7-8 mm. wide, glabrous. Panicle to 20 cm. long, the racemes often more than 5 cm. long. Spikelets about 3-4 mm. long, silky-villous, the hairs white to purplish. First glume less than 0.5 mm. long, apex broadly truncate and usually erosulous, glabrous; second glume as long as the spikelet, acute, the hairs basal, marginal and on the nerves. Sterile lemma as long as the spikelet, acute, apically somewhat lanate-penicillate, the remaining hairs be- tween the margins and the marginal nerves, Fertile lemma about 3 mm. long, longitudinally striatulate. LA PAZ: LARECAJA: San Carlos, Mapiri, 850 m., Buchtien 31 (GH). SANTA CRUZ: SARA: Buena- vista, 450 m., Steinbach 6638 (GH), 6876 (GH); Dolores, 450 m., Stein- bach 1911 (GH). 3. T. californica (Benth.) Chase in Journ. Wash. Acad. Sci. 23: 294 Rhodora [Vol. 68 455 (1933). T. saccharata of the Catalogue. Plants 4-10 dm. tall. Leaf-sheaths glabrous to sparsely papillate-pilose; leaf-blades to 5 mm. wide, glabrous to puberulent. Panicle to 10 (-15) cm. long, racemes few, mostly to 5 em. long. Spikelets about 3-3.5 mm. long, mostly obscured by the long white hairs. First glume about 0.5 mm. long, truncately obtuse to subacute. Sterile lemma as long as the spikelet, 3-nerved, densely villous between the marginal nerves and the margin, glabrous dorsally. Fertile lemma 2.5 mm. long, glabrous, finely striatu- late. COCHABAMBA: CERCADO: Cochabamba, Holway 321 (Hitchcock), 368 (Hitchcock). 73. Digitaria Heister See Henrard, J. Th. Monograph of the Genus Digitaria. Leiden, 1950. Annuals or perennials, the culms 0.4-1 m. tall. Leaf-sheaths and blades often papillate-hirsute, the blades sparsely so, to glabrous or glabrate. Inflorescence a panicle of racemes, these digitate, subdigitate, solitary or irregularly arranged along the axis. Spikelets 2-flowered, the lower floret a sterile lemma, pedicellate in groups of 2 or 3, uni- lateral, in 2 rows. Glumes very unequal, the first glume minute or wanting; second glume and sterile lemma similar, Fertile lemma firm- textured and somewhat pitted-striatulate. a. First glume present, minute. b. Spikelets about 3 mm. long (2.5-3.5); rachis of inflorescence without hispid hairs. ..........seseeeee 1. D. adscendens. b. Spikelets 2 mm. long; rachis with sparse, long, hispid hairs, especially at the nodes. scce. 2. D. horizontalis. a. First glume absent. c. Spikelets 1.5 mm. long, or less. ......... s 3. D. violascens. c. Spikelets 2.5 mm. long. d. Second glume obtuse, shorter than the dark brown fruit. ........ TR 4. D. leiantha. d. Second glume acute, longer than the pale fruit. sss "REN 5. D. lanuginosa. 1. Digitaria adscendens (HBK.) Henr. in Blumea, 1: 92 (1934). Weedy annual, decumbent and spreading, the culms to 1 m. long but usually much shorter. Leaf-sheaths papillate-hirsute; leaf-blades to l em. wide, pubescent to glabrous. Racemes to 10 (-15) cm. long, the rachis 0.5 mm. wide, with margins as wide as the central rib, digitate or in several whorls. First glume less than 0.5 mm. long; second glume about 1.5 mm. long, ciliate. Sterile lemma acute, 3-nerved, pubescent along the nerves but mostly between the lateral nerves and the margins. Fertile lemma about 3 mm. long, light in color and finely pitted. LA PAZ: LARECAJA: Hacienda Simaco, 1400 m., Buchtien 5318 (GH); NOR YUNGAS: Polo-Polo, near Coroico, 1100 m., Buchtien (in 1912) 444 (cu); Milluguaya, 1300 m., Buchtien (in 1966] Flora of Bolivia — Foster 295 1917) 444 (GH). SANTA CRUZ: SARA: Río Surutá, 400 m., Steinbach 6839 (GH); Buenavista, 450 m., Steinbach 6848 (GH), 7057 (GH). 2. D. horizontalis Willd. Enum. Pl. 92 (1809). Annual. Leaf- sheaths usually papillate-hirsute; blades to 1 cm. wide, densely pu- bescent. Raceme-rachis almost without a margin, with scattered, long, hispid hairs, these prominent at the nodes. Spikelets 2 mm. long. First glume minute; second glume about 1 mm. long, subacute, ciliate on upper margin. Sterile lemma 2 mm. long, puberulent near the margin. Fertile lemma pale, acute, shorter than the sterile lemma. SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 6986 (GH). 3. D. violascens Link, Hort. Berol. 1: 229 (1927). Annual, decum- bent, to 8 dm. tall. Leaf-sheaths usually glabrous; leaf-blades to 6 mm. wide, glabrous or pubescent. Racemes 2-5, digitate or subdigitate, the rachis-margins very narrow. Spikelets to 1.5 mm. long, puberulent or glabrous. First glume wanting; second glume about 1 mm. long. Sterile lemma as long as the spikelet. Fertile lemma dark-brown, shining. LA PAZ: ZXARECAJA: Hacienda Simaco, 1400 m., Buchtien 5335 (GH); NOR YUNGAS: Coripata, Hitchcock 22674 (GH). 4. D. leiantha (Hack.) Parodi in Physis, 7: 57 (1923). Perennial, to 1 m. tall, glabrous. Racemes stiff, erect. Spikelets 3 mm. long, glabrous. First glume absent; second glume obtuse, shorter than the spikelet. Sterile lemma as long as the spikelet. Fertile lemma acute, shining, chestnut-brown. SANTA CRUZ: VALLEGRANDE: Coripata, Herzog 1763 (US). 5. D. lanuginosa (Nees) Henr. in Meded. Rijks Herb. Leiden, no. 61: 5 (1930). Perennial, to 1 m. tall. Leaf-sheaths and blades pu- bescent. Racemes to 10 cm. long. Spikelets 2.5 mm. long, pubescent on the nerves, acuminate. First glume absent; second glume and sterile lemma as long as the spikelet. Fertile glume shorter than the sterile lemma and glume, pale in color. SANTA CRUZ: SARA: Buena- vista, 450 m., Steinbach 6849 (GH), 6877 (GH). 74. Thrasya HBK. Thrasya campylostachya (Hack.) Chase in Proc. Biol. Soc. Wash. 24: 115 (1911). Perennial, to about 5 dm. high. Leaf-sheaths densely ciliate, the linear blades to 5 mm. wide, sparsely ciliate and densely puberulent. Inflorescence solitary, terminal or axillary racemes to 9 cm. long, the base often partly sheathed by a stem-leaf, the rachis rather broadly winged and partly concealing the bases of the paired, shortly pedicellate spikelets. Spikelets about 3 mm. long, unilateral, disarticulation below the glumes, 2-flowered, the lower a sterile lemma. First glume usually hyaline, 1 mm. long or less, the second glume about 2.5 mm. long, about 5-nerved, the back rounded, often puberu- lent along the nerves, the apex shortly penicillate. Lemma about 2.5 mm. long, usually slightly shorter than the second glume, indurate, finely striate, the apex shortly penicillate. Palea shorter than the 296 Rhodora [Vol. 68 lemma, indurate, striate, the 2 keels thick, rounded and shining, La PAZ: NOR YUNGAS: San José, 1500 m., Hitchcock 22733 (GH); Cori- pata, Hitchcock 22694 (GH). 75. Pseudechinolaena Stapf Pseudechinolaena polystachya (HBK.) Stapf in Prain, Fl. Trop. Afr. 9: 495 (1919). Sprawling annual, in moist habitats, rooting at the nodes, the fruiting culms to 5 dm. high. Leaf-sheaths densely pilose, the lance-ovate blades to G cm. long, sparsely pilose above and beneath. Inflorescence axillary or terminal, the racemes solitary or arranged in a lax panicle, the rachis, branches and pedicels scabri- dulous-puberulent, the individual racemes 2-5 cm. long. Spikelets 2-flowered, the lower floret sterile or staminate, the upper perfect, about 4 mm. long, disarticulation below the glumes, First glume about 3 mm. long, 5-nerved, rather sparsely short-hispid; second glume 4 mm. long, several-nerved, short-hispid when young, ventricose and uncinate-spiny at maturity. Sterile lemma firm, minutely papillate, slightly shorter than the second glume, its palea nearly as long, but much narrower. Fertile lemma about 2 mm. long, shining, indurate, the narrow indurate palea about as long. LA PAZ: LARECAJA: Ha- cienda Simaco, 1406 m., Buchtiem 53920 (GH); Polo-Polo, 1100 m., Buchtien 265 (GH); Milluguaya, 1300 m., Buchtien 265 (GH). COCHA- BAMBA: CHAPARE: Locotal, 1500 m., Steinbach 9054 (GH), 9070 bis (GH). 76. Rhynchelytrum Nees Rhynchelytrum repens (Willd.) C. E. Hubb. in Kew Bull. 1934: 110 (1934). Tricholaena rosea of the Catalogue. Annual (or sometimes perennial ?), often decumbent at the base, the culms to 1 m. tall. Leaf-sheaths glabrous to sparsely papillate-hirsute; leaf-blades flat, to 2-7 mm. wide, glabrous. Inflorescence an open terminal panicle, to 15 em. long, the axis glabrous below to puberulent above, branches puberulent around the nodes, pedicels glabrous. Spikelets 2-flowered, the lower floret a sterile or staminate lemma, about 4-5 mm. long (excluding hairs), disarticulation below the glumes. First glume to 1.5 mm. long, extremely obtuse or truncate, silky-villous; second glume to 4 mm. long, apically obtuse and bidentate, with a 1.5 mm. awn arising from the base of the sinus, long-ciliate, densely silky- villous (at least below), the hairs exceeding the spikelet by 4-5 mm. Sterile lemma as long as second glume, bidentate, shortly awned, silky-villous. Fertile lemma 2.5-3 mm. long, glabrous, rather shining, obtuse, palea similar and nearly as long, but narrower. COCHA- BAMBA: CERCADO: about 5 km. southeast of Cochabamba, 2800 m., Eyerdam 24918 (GH). 1966] Flora of Bolivia — Foster 297 The panicle of this ornamental grass varies in color from white and gray to pale or deep rosy-purple. 77. Mesosetum Steud. See Swallen in Brittonia, 2: 363-392 (1937). Mesosetum cayennense Steud. Syn. Gram. 118 (1854). Perennial, caespitose, 3-12 dm. high. Leaf-sheaths pilose, ciliate, the flat blades to 5 mm. wide, papillose-pilose above and beneath. Inflorescence a solitary terminal raceme to 20 cm. long, the spikelets solitary, uni- lateral in 2 rows on the rachis. Spikelets 4-5 mm. long, 2-flowered, the lower floret a sterile lemma, or sometimes staminate, the upper floret perfect, disarticulation below the glumes. Glumes unequal, the first to 3 mm. long, truncate, the apex somewhat erose, the margins appressed-pilose below with a tuft of long hairs on each side at the middle, the back toward the rachis; second glume to 5 mm. long, the margins appressed-pilose on the lower half, a tuft of hairs on each side at the middle, and a tuft of long hairs on each margin well below the erose or bidentate apex. Sterile lemma somewhat shorter than the second glume, a tuft of long hairs on each margin at the middle, a few hairs at the base. Fertile lemma smooth, shining, about 3 mm. long. LA PAZ: CAUPOLICÁN: Ixiamas, White (M. E.) 2318 (Swallen). 78. Eriochloa HBK. Erect perennials, 2 dm. to 1 m. tall. Inflorescence a terminal panicle of 2-many somewhat distant racemes. Spikelets short-pedicel- late, unilateral in 2 rows, 2-flowered, the lower floret a sterile lemma, the first glume a minute sheath adnate to the first rachilla-joint as a thickened callus; second glume and sterile lemma similar in appear- ance and size, the glume sometimes awned; fertile lemma indurate, finely papillate-rugulose, mucronate or awned or merely penicillate. Pedicels pilose, with an apical ring of long hairs; leaf-blades narrow (to 4 mm.) ; racemes 2 (-3). eee 1. E. distachya. Pedicels puberulent, lacking an apical ring of long hairs; leaf-blades to 1.5 cm. wide; racemes numerous. .......eeeennnm 2. E. punctata. 1. Eriochloa distachya HBK. Nov. Gen. & Sp. 1: 95 (1816). Plants 2.4 dm. tall, branched, the nodes pubescent, leaves glabrous, blades to 4 mm. wide. Inflorescence of 2, rarely 3, racemes 1-2 cm. long; main axis, raceme-axes and pedicels pilose to pilose-hispid, the pedicels wth an apical ring of long hairs. Spikelets about 4 mm. long (excluding the awn). Second glume and sterile lemma pilose, acute, the glume sometimes awned. Fertile lemma 2.5-3 mm. long, the apex penicillate. SANTA CRUZ: SARA: Río Dolores, Stein- bach 1940 (GH). 298 Rhodora [Vol. 68 2. E. punctata (L.) Desv. ex Hamilton, Prodr. Pl. Ind. Occ. 5 (1825). Plants to 1 m. or more in height, basally decumbent, branched, nodes puberulent, leaves glabrous, the blades flat, to 1.5 em. wide. Inflorescence of numerous racemes mostly over 2 cm. long; main axis and raceme-axes pilose, pedicels puberulent and without an apical ring of long hairs. Spikelets about 4 mm. long. Second glume and sterile lemma sparsely pilose, acute. Fertile lemma over 2 mm. long, the apex not penicillate but awned, the awn about 1 mm. long. LA PAZ: NOR YUNGAS: Coripata, Hacienda El Choro, 1700 m., Buchtien 8040 (GH); SUR YUNGAS: Chulumani, 1600 m., Hitchcock 22666 (GH). 79. Brachiaria (Trin.) Griseb. Brachiaria plantaginea (Link) Hitchc. in Contrib. U. S. Nat. Herb. 12: 212 (1909). Annual, decumbent, rooting at the nodes, culms to 5 dm. high. Leaves glabrous, the lance-ovate blades to 1.5 cm. wide. Inflorescence a series of terminal, unilateral racemes to 7 cm. long, the solitary spikelets in 2 rows along the rachis. Spikelets 2-flowered, the lower floret sterile or staminate, the upper floret perfect, to 5 mm. long, disarticulation below the glumes. Glumes subequal to unequal; first glume about 2 mm. long, but often longer, completely surround- ing the base of the second glume, rather thin-textured, about 7-nerved, its back toward the rachis, second glume to 5 mm. long, firmer- textured, about 7-nerved, the apex sometimes mucronulate. Sterile lemma as long as the second glume and similar in appearance. Fertile lemma about 3 mm. long, somewhat indurate, densely papillate-striate. Palea similar to fertile lemma, enfolded by the margins of the latter. LA PAZ: NOR YUNGAS: Polo-Polo, 1100 m., Buchtien 443 (GH); Millu- guaya, 1300 m., Buchtien 158 (GH); Coripata, 1700 m., Buchtien 8041 (GH). 80. Axonopus Beauv. See Black in Advancing Frontiers of Plant Science, vol. 5 (1963). Perennial (1 Bolivian species annual), mostly tall. Leaf-blades involute or flat or foided. Inflorescence a panicle or subpaniculate, of few to many racemes conjugate, digitate, or distant along a common axis. Spikelets solitary, unilateral in 2 rows along the rachis, dis- articulation below the glumes, 2-flowered, the lower floret a sterile lemma, the perfect floret above it. First glume absent; second glume and sterile lemma more or less alike in size and appearance; second glume and fertile lemma turned away from the rachis. Fertile lemma indurate, whitish or yellowish to dark brown, the margins curved over the edge of the similar palea but not enclosing it, a. Rachis hispid or pilose. b. Rachis-hairs not hispid nor papillate-based. ........ 10. A. pilosus. b. Rachis-hairs hispid, papillate-based. 1966] Flora of Bolivia — Foster 299 c. Hairs not yellow nor golden-brown. .........- 14. A. elegantulus. c. Hairs yellow to golden-brown. d. Spikelets in cupular cavities along the rachis. .e.e..e.seesee 17. A. chrysoblepharis. d. Spikelets not in cupular cavities. e. Rachis not obviously flexuose; glume with papillate-hispid hairs on the nerves. ........ m 16. A. Herzog. e. Rachis obviously flexuose; glume without papillate-hispid hairs. f. Glume and sterile lemma glabrous, occasionally very slightly puberulent. .......á 19. A. chrysites. f. Glume and sterile lemma shortly and finely pubescent. 18. A. exasperatus. syecsssssdettesssosacosesesiseccssnseerestossas o edes deos uaum ET a. Rachis neither hispid nor ciliate. g. Spikelets obviously hirsute, enn 2. A. marginatus. g. Spikelets not hirsute but glabrous to sparsely puberulent. h. Leaves papillate-pilose, at least when young. i. Leaf-blades to 4 mm. wide. ......m 4. A. hirsutus. i. Leaf-blades to 3 CM. wide. ...ccceeeeeeeer sree: 12. A. scoparius. h. Leaves essentially glabrous, at least on upper surface. j. Culm-nodes not glabrous. k. Racemes 2-4 in panicle; if 4, the upper 2 conjugate. C E NR 6a. A. compressus var. macropodius. k. Racemes much more numerous in panicle. |l Racemes up to 20; distant and divergent on the com- mon AXIS. .eenne me 13. A. iridifolius. 1. Racemes more than 20; approximate and ascending on the common axis. m. Spikelets about 2 mm. long; leaf-blades subpetiolate at base. .. eH 7. A. eminens var. bolivianus. m. Spikelets about 2.5 (-3) mm. long; blades not sub- petiolate. serene 8. A. barbigerus. j. Culm-nodes glabrous. n. Leaf-blades abruptly rounded at base. o. Spikelets to 1.6 mm. long. ..sessseessereeseees 15. A. capillaris. o. Spikelets to 2.4 mm. long. ......« 6. A. compressus. n. Leaf-blades not abruptly rounded at base. p. Blades tightly involute, appearing terete. ...... enn MN 9. A. siccus. p. Blades f'at or folded. q. Blades acuminate, acute. een 11. A. andinus. q. Blades not acuminate, obtuse. r. Culms mostly 2 dm. tall. seese. 5. A. fissifolius. r. Culms about 6 dm. tall. s. Glumes pilose on the nerves, especially at base and apex. «eene 1. A. Purpusii. 300 Rhodora [Vol. 68 s. Glumes sparsely puberulent to glabrous. ............ IRE 3. A. affinis. l. Axonopus Purpusii (Mez) Chase in Journ. Wash. Acad. Sci. 17: 144 (1927). Perennial, to 7 dm. tall, culm-nodes usually glabrous. Leaf-sheaths ciliate, at least near summit; blades flat or folded, to 4 mm. wide, shorter than inflorescence, glabrous or sparsely papillate- pubescent, ciliate at the base. Inflorescence of 2-7 erect racemes, to 7 em. long, the rachis scabridulous, flexuose, the pedicels sometimes densely hairy. Spikelets to 2.2 mm. long, ovoid to oblong, greenish or brownish-purple. Glume and sterile lemma equal, mostly 2-nerved, the mid-nerve absent, pilose on the nerves, at least at base and apex. Fertile lemma a little shorter than or equaling the spikelet, yellowish, glabrous or minutely penicillate at apex. SANTA CRUZ: SARA: Buena- vista, 450 m., Steinbach 7060 bis, in part (au). The description given is based in part on an isotype in the Gray Herbarium, Purpus 2450, 2. A. marginatus (Trin.) Chase in Contrib. U. S. Nat. Herb. 17: 226 (1913). Perennial, to 8.5 dm. high, sometimes stoloniferous or rhizomatous. Culm-nodes pubescent or barbate, sometimes glabrous. Leaf-sheaths glabrous, or pilose near the apex; blades 2-4 mm. wide, to 30 em. long, flat or folded, ciliate at base, glabrous above, pilose or sparsely pilose beneath, the obtusely acuminate apex naviculate, In- florescence of up to 13 ascending racemes to 15 cm. long, the rachis scabrid, pedicels pubescent, especially on the margins. Spikelets 2-3 mm. long, ellipsoid to oblong. Glume and sterile lemma equal, 2-4- nerved, mid-nerve absent, densely hairy along (but seldom on) the nerves, the hairs dark purple to silvery, becoming light to dark brown. Fertile lemma about equal to sterile lemma, white or nearly so, the apex penicilate. LA PAZ: LARECAJA: Hacienda Casana, 1600 m., Buchtien 7118 (US). 3. A. affinis Chase in Jourm. Wash. Acad. Sci. 28: 180 (1938). Caespitose perennial, to 6.5 dm. tall. Culms erect or geniculate, the nodes glabrous. Leaf-sheaths ciliate toward apex, otherwise glabrous; blades folded or flat, the base sometimes sparsely pilose, otherwise glabrous. Inflorescences terminal the panicle of 2-7 racemes to about 8 cm. long, rachis glabrous or slightly scabridulous, straight to flexu- ose. Spikelet to 2.1 mm. long, elliptic to ovate. Glume and sterile lemma equal, 2-4-nerved, midrib absent, lateral nerves sparsely pubes- cent to glabrous. Fertile lemma equal to or somewhat shorter than the sterile lemma, pale yellow, glabrous or the apex somewhat penicil- late, SANTA CRUZ: SARA: Buenavista, 450-500 m., Steinbach 5183 (GH), 6724 (GH), 6847 (GH). 4. A. hirsutus G. A. Black in Advanc. Front. Pl. Sci. 5: 55 (1963). Perennial, the culms to 2.5 dm. tall, nodes densely pilose. Leaf-sheaths carinate, densely ciliate, densely pilose at the apex; blades to 20 cm. long, 4 mm. wide, flat, linear, attenuate at the apex, densely papillate- 1966] Flora of Bolivia — Foster 301 pilose, ultimately glabrescent. Inflorescence terminal, solitary, of 4-5 subdigitate racemes to 8 cm. long, rachis scabrous, flexuose. Spikelet to 1.6 mm. long, ovate to oblong-elliptic. Glume and sterile lemma equal, 2-4-nerved, sparsely pubescent along the inner nerves. Fertile lemma slightly shorter, stramineous to brown, shining, glabrous. SANTA CRUZ: SARA: Matacú [Motacá] — Portachuelo, Steinbach 1979 (GH; probable isotype). 5. A. fissifolius (Raddi) Kuhlm. in Comm. Linh. Telegr. Estrat. Matto Grosso Amazonas, Publ. 67, Annexo 5, Bot. 11: 87 (1922). Perennial, to 4.5 dm. tall (usually about 2 dm.), nodes glabrous. Leaf-sheaths ciliate at the apex; blades to 4 mm. wide, usually gla- brous, ciliate at the base, obtuse. Inflorescences mostly terminal panicles of 1-3 racemes to 7 cm. long (occasionally to 10 cm.) ; rachis glabrous, straight to flexuose. Spikelet to 2.1 mm. long, ovate to elliptie. Glumes and sterile lemma equal, 2-4-nerved, sparsely pubes- cent on the nerves, stramineous. Fertile lemma as long as the sterile, light yellow to whitish, glabrous or minutely penicillate. LA PAZ: LARECAJA: Hacienda Simaco, 1400 m., Buchtien 5326 (GH); NOR yungas: Polo-Polo, 1100 m., Buchtien 267 (GH); Coroico, 1560 m., Hitchcock 22716 (GH); SUR YUNGAS: La Florida, 1840 m., Hitchcock 22625 (GH). SANTA CRUZ: SARA: Buenavista, Dolores, Steinbach 1908 (GH). 6. A. compressus (Swartz) Beauv. Ess. Agrost. 154 (1812). Peren- nial, stoloniferous. Culms to 8 dm. high, but usually to 4.5 dm., nodes glabrous to pubescent. Leaf-sheaths ciliate, pubescent near the apex or glabrous; blades flat, to 2 cm. wide, lance-linear to ovate, usually ciliate, sometimes the upper surface pubescent to pilose. In- florescences terminal or terminal and lateral, the panicle of 2-5 racemes to 13 em. long, the uppermost conjugate; rachis scabrid to pilose, flexuose or nearly straight. Spikelet to 3 mm. long on pedicels up to 1 mm. long. Glume and sterile lemma equal, 2-5-nerved, sparsely pubescent along the outermost nerves, or glabrous, stramineous to purple. Fertile lemma to 1.8 mm. long, glabrous or shortly penicillate. SANTA CRUZ: SARA: Buenavista, 500 m., Steinbach 5159 (GH); Dolores, Steinbach 1907 (GH). 6a. A. compressus var. macropodius (Steud.) G. A. Black in Advance. Front. Pl. Sci. 5: 82 (1963). Perennial, to 6 dm. high, culm-nodes pilose. Leaf-sheaths glabrous except along the margins and at the apex; blades to 2 dm. long and 12 mm. wide, elliptic, ciliate near the base; petiolately narrowed at the base, apex obtuse. Panicle of 2-4 racemes, these often conjugate, rachis and pedicels glabrous. Spikelet to 2.5 mm. long. Second glume and sterile lemma 2-4-nerved, very sparsely short-pilose along the nerves, especially at the base. Fertile lemma about 0.75 mm. shorter than the sterile lemma, whitish, more or less striate or papillate-striate. COCHABAMBA: CHAPARE: Anta- huacana, Espíritu Santo, 750 m., Buchtien 2505 (US). 302 Rhodora [Vol. 68 7. A. eminens var. bolivianus G. A. Black in Advanc. Front, Pl. Sci. 5: 93 (1963). Perennial, robust, to 2.5 m. tall (fide Steinbach), the culm-nodes densely barbate. Leaf-sheaths densely ciliate and pilose near the apex; blades involute, subpetiolate at the base, becoming flat, to 1 em. wide, upper surface pilose near the base, the apex long- acuminate to pungent. Panicle terminal, to 35 em. long, of numerous approximate racemes (usually at least 50) to 18 cm. long; rachis scabrous. Spikelet to 2 mm. long, ovoid; glume and sterile lemma equal, 5-nerved, glabrous or sparsely pubescent. Fertile lemma shorter than the sterile, white, glabrous to penicillate. SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 6976 (GH). 8. A. barbigerus (Kunth) Hitche. in Contrib. U. S. Nat. Herb, 24: 433 (1927). Stout perennial, the culms to 1-2 m. tall, nodes usually densely pilose but occasionally glabrous. Leaf-sheaths ciliate at the apex, the base sometimes pubescent; blades to 9 dm, long and 6 mm. wide, long-attenuate at the apex, involute or folded at the base. Inflorescences terminal, usually solitary, of 10-50 or more ascending racemes to 20-30 em. long, rachis scabrous with occasional long hairs. Spikelet about 2.5 mm. long (but occasionally to 3.1 mm.). Glume and sterile lemma equal, o-nerved, glabrous or sparsely pubescent between the nerves. Fertile lemma about equaling the sterile, or shorter, pale yellow to white, glabrous or penicillate. LA Paz: [prov- ince unknown]: Puente Villa, Cárdenas 3608 (GH). SANTA CRUZ: SARA: Buenavista, 450-500 m., Steinbach 1920 (GH), 5377 (GH). 9. A. siccus (Nees) Kuhlm. in Comm. Linh. Telegr. Estrat. Matto Grosso Amazonas, Publ. 67, Annexo 5, Bot. 11: 87 (1922). Slender perennial, the culms mostly to 9 dm. tall but occasionally much taller, the nodes usually glabrous. Leaf-sheaths ciliate, at least near the apex; blades tightly involute, appearing terete, the apex slender and filamentous. Panicles terminal, of 4-30 racemes to 15 em. long, rachis scabrid. Spikelets to 2.1 mm. long. Glumes and sterile lemma equal, 5-nerved, glabrous, or puberulent between the nerves. Fertile lemma equalling the sterile, white to stramineous, glabrous or penicillate. SANTA CRUZ: VALLEGRANDE: Samaipata, Herzog 1709 (Us). TARIJA: O'CONNOR: Narvaez, 2700 m., Cárdenas 4935 (US). 10. A. pilosus G. A. Black in Advanc. Front. Pl. Sci. 5: 100 (1968). Perennial, the culms to 5 dm. tall with pubescent nodes. Leaf-sheaths somewhat pilose, especially near the apex, the hairs tufted at the base of the blade; blades to 2.5 mm. wide, folded, ciliate at the base, the upper surface pilose, the apex very long-acuminate or filamentous. Inflorescence a solitary terminal panicle of many (to 25) racemes to 10 em. long; rachis flexuose, densely pilose, the hairs not hispid nor yellow-brown, mostly aggregated around the short pedicels. Spikelets to 2.7 mm. long, long-ellipsoid, often purplish. Glume and sterile lemma equal, 5-nerved, glabrous to sparsely pubescent. Fertile lemma 1966] Flora of Bolivia — Foster 303 greenish to white, penicillate. TARIJA: MENDEZ: Tucumilla, 2600 m., Fiebrig 2782 (GH; type-number). 11, A. andinus G. A. Black in Advanc. Front. Pl. Sci. 5: 111 (1963). Perennial, to 1 m. tall, culm-nodes glabrous. Leaf-sheaths somewhat short-pilose at the apex, otherwise glabrous; blades long, to 5 mm. wide, loosely folded or flat, glabrous or with a few hairs on the upper surface at the base. Panicles of 5-7 racemes to 10 em. long; rachis and pedicels scabrid on the angles. Spikelets about 2.5 mm. long, lanceolate. Second glume and sterile lemma equal, often purplish (at least on the type), 2-4-nerved, the nerves short-pilose. Fertile lemma shorter than the sterile, yellowish-white, obscurely striatulate, the acute apex penicillate. LA PAZ: LARECAJA: Hacienda Simaco, 1400 m., Buchtien 5328 (US; type). 19. A. scoparius (Flügge) Kuhlm. in Comm. Linh. Telegr. Estrat. Matto Grosso Amazonas, Publ. 67, Annexo 5, Bot. 11: 45 (1922). Paspalum tripinnatum Mez in Fedde, Rep. Sp. Nov. 15: 64 (1917). Stout perennial, the culms to 1.5 m. tall, nodes glabrous. Leaf-sheaths glabrous; blades to 5 dm. long and 3 em, wide, the apex obtuse, the upper surface papillate-pilose. Inflorescences terminal (sometimes axillary as well), of few to 100 racemes to 20 em. long; rachis scabrid and sometimes sparsely short-pilose. Spikelets to 2.4 mm. long. Glume and sterile lemma equal, usually 5-nerved, short-pubescent between the nerves. Fertile lemma mostly shorter than the sterile, pale yellow, glabrous or penicillate. La PAZ: LARECAJA: Hacienda Simaco, 1400 m., Buchtien 5324 (GH). COCHABAMBA: CHAPARE: Locotal, 1500 m., Steinbach 9093 in part (GH). The Steinbach specimen is only an inflorescence-fragment and the present attribution may be an error. 13. A. iridifolius (Poepp.) G. A. Black in Advance. Front. Pl. Sci. 5: 125 (1963). Perennial, to 1 m. tall, the culm-nodes densely pu- bescent. Leaf-sheaths scabrid, carinate, pubescent to long-pilose at the apex; blades flat, to 1.9 cm. wide, obtuse, upper surface scabrid or strigose at base. Inflorescence a terminal panicle of 6-20 distant racemes to 18 em. long; rachis somewhat flexuose, scabrid, Spikelets to 3 mm. long, greenish to purple. Glume and sterile lemma equal, 4-5-nerved, often puberulent between the nerves. Fertile lemma shorter than the sterile, stramineous, the apex penicillate. LA PAZ: NOR YUNGAS: Polo-Polo, 1100 m., Buchtien 448 (GH). 14. A. elegantulus (Presl) Hitche. in Contrib. U. S. Nat. Herb. 24: 433 (1927). Perennial, the culms to 7.5 dm. tall, the nodes glabrous. Leaf-sheaths papillate-pilose to glabrous, papillate-ciliate; blades folded at base, flat above, to 25 cm. long and 5 mm. wide, long- attenuate, papillate-pilose to glabrous. Inflorescence terminal, of up to 12 subdigitate racemes to 10 cm. long; rachis ciliate, with few to no hairs immediately below the spikelets, hairs papillate-based but neither hispid nor golden-brown. Spikelets to 2.5 mm. long, greenish 304 Rhodora [Vol. 68 to purple. Glume and sterile lemma equal, 2-3-nerved, glabrous. Fer- tile lemma sometimes slightly longer than the sterile, glabrous, shiny brown. LA Paz: LARECAJA: San Carlos, Mapiri, Buchtien s. m. (Black). SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 7060 (at least as to GH sheet). 15. A. capillaris (Lam.) Chase in Proc. Biol. Soc. Wash. 24: 133 (1911). Lax annual, to 7.5 dm. high, nodes glabrous, the lowest some- times rooting, culms geniculate. Leaf-sheaths ciliate; blades flat, to 8 mm. wide, lanceolate, acuminate, the edges undulate, abruptly rounded and more or less clasping at the base, pilose at the base, the main portion of the blade glabrous or pubescent. Inflorescence terminal, of 2-6 paired or subdigitate racemes to 8 cm. long; rachis and peduncles capilary, the rachis markedly flexuose and scabri- dulous. Spikelets to 1.6 mm. long, sessile, ellipsoid, obtuse to acute. Sterile lemma and glume equal, with 2-4 inconspicuous nerves, glabrous to sparsely pubescent. Fertile lemma shining, brown, glabrous. LA PAZ: LARECAJA: Mapiri, 1600 m., Rusby 241 (GH) ; Hacienda Simaco, 1400 m., Buchtien 5897 (GH); SUR YUNGAS: Chulumani, 1600 m., Hitchcock 22657 (GH). SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 6879 (GH). 16. A. Herzogii (Hack.) Hitche. in Contrib. U. S. Nat. Herb. 24: 431 (1927). Perennial, to 8 dm. tall, the culm-nodes glabrous. Leaf- sheaths glabrous; blades folded, to 15 cm. long and 3 mm. wide, papil- late-ciliate near the base. Panicle of 2-7 subdigitate racemes to 10 cm. long, axis pilose, rachis papillate-ciliate, the hairs yellowish- to golden-brown, long. Spikelet to 2.5 mm. long, lanceolate or lance-ovate, purplish, the pedicels minute. Second glume and sterile lemma equal, 5-nerved, the glume papillate-hispid on the nerves, the sterile lemma with shorter and finer white hairs between the nerves. Fertile lemma nearly as long as the sterile, rather dark brown, obscurely, if at all, striatulate, the acute apex penicillate. SANTA Cruz: CHIQUITOS: Cerro San Miserate, Santiago, Herzog 866 (US; type-number). 17. A. chrysoblepharis (Lag.) Chase in Proc. Biol. Soc. Wash. 24: 134 (1911). Annual or perennial, to 10 dm. tall, the culm-nodes gla- brous or pubescent. Leaf-sheaths glabrous to densely pubescent, often hispid-ciliate, especially near the apex; blades to 1.5 cm. wide, usually narrower, pilose to glabrous, acuminate. Inflorescence terminal or lateral, of 2-10 racemes to 4 cm. long, subdigitate; rachis not flexuose, the margins and center densely papillate-hispid, the golden or golden- brown hairs dense, to 3-4 mm. long, basally fused, apex often without spikelets. Spikelets to 1.5 mm. long, fitting into cavities in the rachis, ovoid, obtuse. Glume and sterile lemma equal, glabrous, 2-nerved, very thin. Fertile lemma purple or brown, glabrous, obtuse to sub- acute. BENI: YACUMA: Reyes, Rusby 216 (Black). LA PAz: NOR YUNGAS: Hacienda Anacuri, Holway 708 (Black). 18. A. exasperatus (Nees) G. A. Black in Advanc, Front. Pl. Sci. 1966] Flora of Bolivia — Foster 305 5: 168 (1963). Straggling perennial to 8 dm. tall, culms often branched at the base, nodes glabrous. Leaf-sheaths glabrous; blades mostly less than 1 dm. long, to 5 mm, wide, with a few stiff hairs near the base, the apex long-acuminate. Inflorescences terminal, 1-2, of 2-7 racemes to 9 em. long; rachis somewhat flexuose, papillate- hispid, the hairs golden-brown, with a dense tuft of hairs subtending each spikelet, mostly more than 10 hairs in a tuft. Spikelets to 1.5 mm. long, green to purple, ovoid to obovoid, obtuse to subacute. Glume and sterile lemma equal, the 2-4 nerves obscure, short-pubescent, usually sparsely so, or the hairs in 2 rows. Fertile lemma shining brown, glabrous. SANTA CRUZ: SARA: lomas arenosas, 450 m., [Buenavista ?], Steinbach 1815 (GH). 19. A. chrysites (Steud.) Kuhlm. in Comm. Linh. Telegr. Estrat. Matto Grosse Amazonas, Publ. 67, Annexo 5, Bot. 11: 88 (1922). A. aureus of the Catalogue, at least in part. Perennial, straggling to erect, to 7.5 dm. tall, culm-nodes glabrous. Leaves glabrous except for a few hispid hairs at base of upper surface of blade, the blades to 7 mm. wide, attenuate. Inflorescence terminal, 1-2 panicles, the racemes to 8 cm. long; rachis somewhat flexuose, long-hispid on the margins and mid-ridge, with a cluster of hairs to 2.5 mm. long below each spikelet, the hairs yellow- or golden-brown. Spikelets to 1.5 mm. long, obovoid, obtuse, purple to fulvous. Glume and sterile lemma equal, the 2-4 nerves obscure (or absent), glabrous or slightly pubes- cent. Fertile lemma shining brown, glabrous. SANTA CRUZ: SARA: Buenavista, 450-500 m., Steinbach 5426 (GH), 6948 (GH). 81. Paspalum L. See Chase in Contrib. U. S. Nat. Herb. 28: 1-310 (1929). Annual or perennial. Leaf-blades flat or folded at the base. Panicle of 1 to many racemes on a common axis. Rachis of racemes narrow to very broadly winged, sometimes prolonged beyond the ultimate spikelets. Spikelets glabrous or variably pubescent, sessile, sub-sessile or pedicellate, solitary or paired, unilateral on the rachis, often biconvex or planoconvex, disarticulation below the glumes, 2-flowered, the lower floret a sterile lemma, the upper perfect. First glume usually absent, present in some species and occasionally present in others. Second glume usually present, absent in a few species, similar to the sterile lemma in size and appearance. Fertile lemma usually somewhat indurate, often shining, usually somewhat striate or striate- papillate. a. Rachis prolonged beyond uppermost spikelet. b. Spikelets glabrous. c. Glumes absent. d. Spikelets over 2 mm. long. seeserecretenenn 4. P. candidum. d. Spikelets less than 2 mm. long. 306 Rhodora [Vol. 68 e. Culms dark; leaf-sheaths glabrous or sparsely pilose along marginsg. ....eeseesesseeeeeeeeerereeeene trenes 5. P. depauperatum. e. Culms light; leaf-sheaths densely long-pilose. ..................- 6. P. lineispatha. metttssssisstosesoieesssssesósseooososooososstesssstssceseeseseceeeeees c. Second glume present. f. Spikelets less than 2 mm. long; rachis less than 2 mm. wide. £. Plants over 10 cm. high; leaves sparsely pilose to gla- DYOUS. MM 1. P. penicillatum. g. Plants less than 10 em. high; leaves densely pilose. ........ RM 2. P. pygmaeum. f. Spikelets over 2 mm. long; rachis at least 2 mm. wide. 3. P. prostratum. Metettsotvostoscssssesocoseeeovoscesesoocceceeceeoseooeessepvososecteoscoceeooo b. Spikelets not glabrous. h. Rachis very broadly winged. i. Leaf-blades flat, 3-8 mm. wide; racemes usually more than Bs eewestinsntertbashssseraanvacsngeneasysanuenigeristiserosaredystessiseisoceess 7. P. Ceresia. i. Leaf-blades very narrow, involute, pilose; racemes rarely more than 1. usss 8. P. stellatum. h. Rachis very narrowly winged. eese.. 39. P. saccharoides. a. Rachis not prolonged beyond uppermost spikelet. j. Spikelets glabrous. k. Glumes absent. l. Rachis puberulent; spikelets mostly over 2 mm. long. .......... RR 38. P. boliviense. l. Rachis glabrous, with a few long cilia; spikelets mostly less than 2 mm. long. ............. es 37. P. malacophyllum. k. Second glume present. m. Rachis coarsely long-ciliate. n. Rachis sparsely ciliate, mainly puberulent-scabrid; spike- lets mostly over 2 mm. long. .................— 32. P. millegrana. n. Rachis densely ciliate; spikelets mostly less than 2 mm. o CRRNNNNNEENNENNMMMMMMMMN 33. P. densum. m. Rachis not coarsely long-ciliate. o. Racemes not conjugate. p. First glume present on 1 or both of the paired spikelets. q. Spikelets about 1.5 mm. long. ........ 14. P. decumbens. q. Spikelets about 2.5 mm. long. .............. 35. P. pilosum. p. First elume absent or rarely present on 1 spikelet of the pair. |... 21. P. inaequivalve. o. Racemes conjugate. r. Spikelets about 1 mm. long. ................. 20. P. pictum. r. Spikelets 2-3 mm. long. s. Spikelets 2.5-3 mm. long. voces 25. P. notatum. s. Spikelets 2-2.5 mm. long. ............. 26. P. minus. j. Spikelets not glabrous. t. Rachis coarsely seabrid-ciliate. ............. sss 30, P. virgatum, 1966] Flora of Bolivia — Foster 307 t. Rachis not coarsely scabrid-ciliate. u. Spikelet-hairs swollen-globose. .........-- 19. P. multicaule. u. Spikelet-hairs not swollen-globose. v. Spikelets with long silky hairs. w. Racemes neither solitary nor conjugate. x. Spikelets to 2.5 mm. long. seese 24. P. Urvillei. x. Spikelets 3.5 mm. long, or more. y. Sterile lemma glabrous. z. hacemes 2-5; leaf-blades stiffly pilose (at least on margins). .....HR 9. P. Humboldtianum. z. Racemes 8-16; leaf-blades glabrous or nearly so. 11. P. Buchtienii, coredétltasasossbaeertestossnesees Sue senos uut Ea Eo Ue y. Sterile lemma not glabrous. aa. Leaf-blades ciliate at base, otherwise glabrous. ROUTE E E utem one re rr 23. P. dilatatum. aa. Leaf-blades puberulent or pilose. bb. Blades puberulent on both surfaces. ...........«.-. SURE RENT meu s 12. P. polyphyllum. bb. Blades sparsely to densely pilose. ......... 10. P. erianthwm. eecossececsosssesco aet bosses sone ut ttt esae eeoeee w. Racemes solitary or conjugate. cc. Racemes solitary. ...... 36. P. Ekmanianum. cc. Racemes conjugate. ......... HM 22. P. conjugatum. v. Spikelets lacking long silky hairs. dd. Racemes conjugate (rarely a third raceme present below). eene 13. P. distichum. dd. Racemes not conjugate. ee. First glume present on 1 of the paired spikelets. ff. Plants to 8.5 dm. tall; second glume sometimes »uberulent on the margin only. ...... Hem NNNM 15. P. inconstans. ff. Plants to 4.5 dm. tall; second glume obscurely puberulent or pubescent at the summit. eesse. 16. P. lepidum. "———— A Edid ee. First glume lacking. gg. Spikelets 2 mm. or less in length. hh. Fruit dark brown. ....... 27 P. melanospermum. hh. Fruit pale. ii. Sterile lemma pubescent. .. 17. P. paniculatum. ii. Sterile lemma glabrous. ...... 18. P. Juergensti. gg. Spikelets over 2 mm. long. jj. Fruit pale. kk. Leaf-blades densely pubescent on both sur- FACES, cccececsssesscterncssocansesssesecssnce 16. P. lepidum. kk Leaf-blades glabrous, scabrous on margins. 34. P. remotum. D isis saacaseusaseqonosond sata ARDOR eR ESSERE 308 Rhodora [Vol. 68 jj. Fruit dark. ll. Sterile lemma transversely rugulose near base. RR 28. P. plicatulum. ll. Sterile lemma not rugulose. mm. Spikelets to 2.5 mm. long. eee mm. Spikelets 2.7-3 mm. long. ........ ee RR 31. P. conspersum. 1. Paspalum penicillatum Hook. f. in Trans. Linn. Soc. 20: 171 (1851). Annual, decumbent, to 4 dm. tall. Leaf-blades to 13 cm. long and 1.5 em. wide, glabrous beneath, sparsely pilose above, ulti- mately glabrate and somewhat pilose-ciliate near the base. Panicle of several racemes, usually paired, to 3 cm. long, usually nearer 2 cm., the rachis of each raceme extended well beyond the uppermost spikelet, less than 2 mm. wide. Spikelets to 1.7 mm. long, glabrous, second glume present. Sterile lemma as long as spikelet, thin, several- veined. Fertile lemma dull white. LA PAZ: LARECAJA: Sorata, 2650- 2800 m., Mandon. 1250 (GH); Sorata, Bang 1310 (GH); Sorata, 2800 m., Rusby 240 (GH); NOR YUNGAS: Coripata, 1700 m., Buchtien 8050 (GH); SUR YUNGAS: La Florida, 1840 m., Hitchcock 22643 (GH). 2. P. pygmaeum Hack. in Fedde, Rep. Sp. Nov. 11: 18 (1912). P. pygmaewm var. glabrescens Hack. in Fedde, Rep. Sp. Nov. 11: 18 (1912). Matted, branched annual, culms to 1 dm. long, usually shorter. Leaf-sheaths rather densely pilose; blades short, usually to 3 cm. long, more sparsely pilose above and beneath. Panicles small, with 2-6 racemes 0.5-2 cm. long, the rachis less than 2 mm. wide, extended beyond the uppermost spikelet. Spikelets less than 2 mm. long, glabrous. Second glume present, 3-nerved, sterile lemma similar, the 3 nerves in a central position. Fertile lemma smooth, not highly shining. LA PAZ: MURILLO: La Paz, 3900 m., Buchtien 2490 (GH; type-number of P. pygmaeum var. glabrescens) ; Talca Chuquiaguillo, Bang 808 (GH). COCHABAMBA: CHAPARE: La Aduana, 3100 m., Steinbach 9579 (au). 3. P. prostratum Scribn. & Merr, in U. S. Dept. Agric. Div. Agrost. Bull. no. 24: 9 (1900). Annual, decumbent or prostrate, the culms to 3 dm. long, often rooting at the nodes. Leaf-sheaths sparsely pilose-ciliate or glabrous to densely pilose; blades to 8 cm. long and 1.2 em. wide, papillate-pilose above and beneath. Panicle of 3-12 solitary to somewhat fasciculate racemes 1-3 cm. long, the rachis about 2.5 mm. wide, extended beyond the uppermost spikelet. Spikelet to 2.2 mm. long, glabrous, second glume present, sterile lemma similar, 3-nerved. Fertile lemma nearly as long as the Spikelet, smooth, shining. La PAZ: LARECAJA: Sorata, Holway 507 (us). COCHA- BAMBA: CERCADO: Morochata, 2900 m., Cárdenas 3414 (US). 4. P. candidum (Humb. & Bonpl. ex Flügge) Kunth in Mém. Mus. Hist. Nat. 2: 68 (1815). Annual, decumbent, straggling, the culms 1966] Flora of Bolivia — Foster 309 often to 1 m. or more in length, the dark nodes somewhat enlarged and strigose to villous. Leaf-sheaths ciliate near the apex, otherwise pilose or glabrous; blades to 12 cm. long and 1.2 cm. wide, finely pilose on both surfaces. Panicle lax, with 7-20 ascending solitary racemes to 4 cm. long, rachis to 2.5 mm. wide, prolonged beyond the uppermost spikelet. Spikelets to 2.5 mm. long, glabrous, both glumes absent. Sterile lemma 3-nerved. Fertile lemma shorter than the sterile, smooth, ivory-white, shining. LA PAZ: SUR YUNGAS: San Felipe, 2800 m., Hitchcock 22597 (GH); Sirupaya, 2100 m., Buchtien (in Baenitz Herb. Amer.) 1408 (GH). 5. P. depauperatum Presl, Rel. Haenk. 1: 215 (1830). P. vinosum Mez in Fedde, Rep. Sp. Nov. 15: 28 (1917). Decumbent annual, the dark, purplish culms to 1.8 dm. long. Leaf-sheaths glabrous, or, in youth, pilose near the apex, ciliate; blades to 7 em. long and 4 mm. wide, pilose or papiliate-pubescent on both surfaces. Panicle of up to 7 solitary racemes, 15-3 cm. long, the rachis prolonged beyond the uppermost spikelet. Spikelets glabrous, less than 2 mm. long, glumes absent. Sterile lemma 3-veined. Fertile lemma shining. COCHABAMBA: CHAPARE: Incachaca, 2250 m., Steinbach 9495 (GH). 6. P. lineispatha Mez in Fedde, Rep. Sp. Nov. 15: 27 (1917). Rather straggling annual, the light-colored culms to 30 em. tall. Leaf-sheaths densely papillate-pilose; blades to 6 cm. long and 9 mm. wide, densely papillate-pilose beneath, sparsely so above. Panicle of 2-12 solitary racemes to 1.5 cm. long, the rachis prolonged beyond the uppermost spikelet, less than 2 mm. wide. Spikelets glabrous, less than 2 mm. long, glumes absent. Sterile lemma 3-veined. Fertile lemma shining. LA PAZ: LARECAJA: Hacienda Casana near Tipuani, 1400 m., Buchtien 7112 (GH). 7. P. Ceresia (O. Ktze.) Chase in Niles in Contrib. U. S. Nat. Herb. 24: 153 (1925). P. membranaceum of the Catalogue. Perennial, to 6 dm. tall, villous at the base, shortly rhizomatous. Leaf-blades to 15 em. long and 8 mm. wide, pilose near the margin on the upper surface. Panicle of 1-4 distant racemes, the rachis-proper dark, with orange-brown broad margins, to 1 cm. wide overall, prolonged beyond the uppermost spikelet. Spikelets to 4 mm. long, with a dense silver- white pilosity, the longest hairs as long as the spikelet, hairs mostly papillate-based. Second glume present. Without locality: Bang 1080 (GH). LA Paz: LARECAJA: Cerro del Iminapi, 2600-2700 m., Mandon 1255 (GH). SANTA CRUZ: SARA: Buenavista, 500 m., Steinbach 5421 (GH). 8. P. stellatum Humb. & Bonpl. ex Flügge, Monogr. Pasp. 62 (1810). Caespitose perennial to 5 dm. tall. Leaf-sheaths densely long-pilose. Inflorescence of 1 arcuate raceme (rarely 2) to 10 cm. long, the broad yellow-brown rachis prolonged beyond the uppermost spikelet. Spikelets to 2.5 mm. long, second glume present; second glume and sterile lemma with very short white hairs and long shining hairs, 310 Rhodora [Vol. 68 these more or less stellate in arrangement. Fertile lemma shorter than the spikelet. SANTA CRUZ: SARA: Buenavista, 450 m., Stein- bach 1988 (GH), 7103 (GH). 9. P. Humboldtianum Flügge, Monogr, Pasp. 67 (1810). Caespitose perennial to 8 dm. tall, the base rather woody. Leaf-sheaths papillate- pilose, sometimes glabrate; blades flat, to 18 cm. long and 1.5 cm. wide, papillate-pilose or glabrate. Panicle of 2 to several racemes, neither solitary nor conjugate, the narrow rachis not coarsely scabrid- ciliate, not exceeding the uppermost Spikelet, usually dull purple. Spikelets mostly purple, to 3.5-4 mm. long. Second glume present, shortly pubescent with appressed hairs, the margins with numerous long papillate-based hairs. Sterile lemma shorter than second glume, glabrous. Fertile lemma dull white, thin-textured. LA PAZ: LARE- CAJA: Cerro del Iminapi, 2650 m., Mandon 1254 (GH). COCHABAMBA: MIZQUE: near Vilavila, 2500 m., Eyerdam 24977 (GH); CHAPARE: Locotal, 1900 m., Steinbach 9093 in part (GH). SANTA CRUZ: SARA: Cerro del Amboró, 6618 (GH); Buenavista, 450 m., Steinbach 6644 (GH); VALLEGRANDE: Cuesta de La Negra, Steinbach 3712 (GH). 10. P. erianthum Nees in Trin. Gram. Pan. 121 (1826). Perennial, densely caespitose, to 1 m. tall Leaf-sheaths variably pilose and ciliate; blades sparsely to densely pilose. Panicle of few to several erect racemes, raceme-rachis narrow, not prolonged beyond the upper- most spikelet, not coarsely scabrid-ciliate. Spikelets to 5 mm. long, ovate, acute. Second glume sparsely long-villous, with a marginal fringe of long silky hairs. Sterile lemma more nearly glabrous except at apex. Fertile lemma pale, shining, finely reticulate-striate, SANTA CRUZ: SARA: Buenavista, Steinbach 3543 (GH). 11. P. Buchtienii Hack. in Fedde, Rep. Sp. Nov. 6: 153 (1908). Cae- spitose perennial, somewhat straggling, to 10 dm. tall. Leaf-sheaths usually densely long-ciliate, otherwise glabrous; blades to 15 em. long and 1.5 em. wide, usually glabrous or glabrate on the upper surface. Panicle of 8-16 purplish racemes to 10 em. long, neither solitary nor conjugate, the very narrow rachis not extended beyond the uppermost spikelet, not coarsely scabrid-ciliate. Spikelets to 3.5-4 mm. long, second glume present, shortly acuminate, ciliate with a long fringe of stiff white hairs. Sterile lemma glabrous, Fertile lemma shorter than the spikelet. YUNGAS: Bang 265 (GH); Puente Villa, 1400 m., Cárdenas 3597 (GH). LA PAZ: NOR YUNGAS: Polo-Polo, 1100 m., Buchtien 266 (GH), 447 (GH); SUR YUNGAS: Chulumani, 1600 m., Hitchcock 22664 (GH); La Florida, 1840 m., Hitchcock 22648 (GH). 12. P. polvphylla Nees ex Trin. Gram. Pan. 114 (1810). Perennial, to 6 dm. tall. Leaf-sheaths somewhat pilose at the apex, otherwise glabrous; blades to 5 em. long, about 3 mm. wide, flat to subinvolute, puberulent on both surfaces. Panicle of 1-3 racemes to 7 em. long, light in color, the narrow rachis not coarsely scabrid-ciliate, not prolonged beyond the uppermost spikelet. Spikelets to 3.5 mm. long, 1966] Flora of Bolivia — Foster 311 second glume present, 5-nerved, densely ciliate with long, stiff, white hairs. Sterile lemma 5-nerved, ciliate on at least the upper margin. Fertile lemma as long as the sterile, thin-textured. SANTA CRUZ: VALLEGRANDE: Cuesta de La Negra, 1500 m., Steinbach 3718 (GH); SARA: Buenavista, 500 m., Steinbach 5385 (GH), 5762 (GH), 6808 (GH). 13. P. distichum L. Syst. Nat. (ed. 10) 2: 855 (1759). Stoloni- ferous perennial to 6 dm. tall, the nodes pubescent. Leaf-sheaths sparsely hispid-ciliate at the apex, otherwise glabrous; blades subin- volute to flat, to 6 mm. wide, the base sparsely hispid-ciliate, other- wise glabrous. Panicle of 2 conjugate racemes (rarely 3) to 7 cm. long, the rachis not coarsely scabrid-cilate, not prolonged beyond the uppermost spikelets. Spikelets 2.5-3.5 mm. long. First glume often present, but not over 0.5 mm. long; second glume abruptly acute, pubescent to puberulent, the midrib prominent. Sterile lemma similar but glabrous, the midrib prominent. Fertile lemma shorter than spikelet, greenish-white, somewhat shining, often shortly penicillate at the abruptly acute apex. LA PAZ: MURILLO: La Paz, Buchtien 4518 (Us). COCHABAMBA: CERCADO: Cochabamba, Hitchcock 22793 (us). SANTA CRUZ: SARA: Rio Surutü, 400 m., Steinbach 6835 (GH). 14. P. decumbens Swartz, Prodr. 22 (1788). Branched, decumbent perennial, the culms to 6 dm. tall. Leaf-sheaths finely pubescent and densely short-ciliate; blades flat, to 10 em. long and 8 mm. wide, pubescent on both surfaces, the margins ciliate, at least near the base. Inflorescence a solitary raceme 2-3 cm. long, the narrow rachis not prolonged beyond the uppermost spikelet, not coarsely long-ciliate. Spikelets about 1.5 mm. long, obovate to suborbicular, paired, glabrous. First glume present on one or both spikelets in a pair, very short ; second glume rounded, half the length of the spikelet. Sterile lemma similar but usually twice as long. Fertile lemma shining, longi- tudinally regularly striate, the ridges finely and regularly papillate. La Paz: LARECAJA: Hacienda Simaco, 1400 m., Buchtien 5314 (GH). COCHABAMBA: CHAPARE: Antahuacana, Espíritu Santo, 750 m., Buchtien s. n. (GH). SANTA CRUZ: SARA: Buenavista, 450 m., Stein- bach 7016 (GH). YUNGAS: Bang 266 in part (at least as to GH sheet). 15. P. inconstans Chase in Contrib. U. S. Nat. Herb. 24: 446 (1927). Perennial, to 8.5 dm. tall, the culms branched above. Leaf-sheaths short-ciliate, pubescent at the apex; blades flat, to 15 cm. long and 1.2 cm. wide, rather densely pubescent on both surfaces. Panicle of 1-8 (usually 2-5) non-conjugate racemes, the narrow rachis not pro- longed beyond the uppermost spikelet and not coarsely scabrid-ciliate. Spikelets paired, about 2.5 mm. long. First glume usually present, at least as a rudiment, on the lower of the paired spikelets; second glume shorter than the spikelet, 3 (-5) -nerved, the margin puberulent. Sterile lemma somewhat longer, sometimes slightly puberulent on the margin. Fertile lemma somewhat shining, very finely papillate-striate. LA 312 Rhodora [Vol. 68 PAZ: NOR YUNGAS: San José, 1500 m., Hitehcock 22734 (GH); SUR YUNGAS: La Florida, 1840 m., Hitchcock 22609 (GH). 16. P. lepidum Chase in Contrib. U. S. Nat. Herb. 27: 447 (1927). Densely caespitose perennial, to 4.5 dm. tall, the culms simple. Leaf- sheaths pubescent at the apex, otherwise glabrous to pubescent; blades to 11 em. long and 1 em. wide, densely pubescent on both surfaces. Panicle of 3-4 non-conjugate racemes, the narrow rachis not coarsely scabrid-ciliate, not prolonged beyond the uppermost spikelet. Spikelets paired, to 2.4 mm. long. First glume seldom present; second glume 5-nerved, somewhat puberulent or pubescent at the summit. Sterile lemma slightly longer, 5-nerved, glabrous. Fertile lemma very finely papillate-striate. LA PAZ: SUR YUNGAS: San Felipe, 2800 m., Hitch- cock 22605 (GH). 17. P. paniculatum L. Syst. Nat. (ed. 10) 2: 855 (1759). Perennial, branching, to 1 m. or more in height, the culm-nodes pilose to pubes- cent. Leaf-sheaths glabrous, ciliate; blades flat, to 30 cm. long and 1.5 em. wide. Panicle of numerous, crowded, non-conjugate racemes, the narrow rachis not coarsely scabrid-ciliate, not prolonged beyond the uppermost spikelet. Spikelets paired, suborbicular, about 1.2-1.4 mm. long. Second glume present. Sterile lemma pubescent. Fertile lemma minutely papillate-striate. La Paz: LARECAJA: Hacienda Simaco, 1400 m., Buchtien 5323 (GH); NOR YUNGAS: Polo-Polo, 1100 m., Buchtien 269 (GH); Milluguaya, 1300 m., Buchtien 269 (GH); SUR YUNGAS: La Florida, 1840 m., Hitchcock 22624 (GH). 18. P. Juergensii Hack. in Fedde, Rep. Sp. Nov. 7: 312 (1909). Perennial, to 1 m. high, somewhat straggling or decumbent, the nodes dark, often with long hispid hairs. Leaf-sheaths villous-ciliate, often with papillate-based hairs; blades flat, to 20 cm. long and 1.5 cm. wide, long-pubescent above and beneath with papillate-based hairs, or glabrate. Panicle of several lax non-conjugate racemes, rachis of racemes not coarsely scabrid-ciliate, not prolonged beyond the upper- most spikelet. Spikelets paired, to 1.7 mm. long, second glume present, pubescent, rather blunt. Sterile lemma glabrous. Fertile lemma some- what shining, very finely papillate-striate. YUNGAS: Bang 308 (GH). 19. P. multicaule Poir. Encycl. Suppl. 4: 309 (1816). Annual, much- branched from the base, about 1 dm. tall. Leaf-sheaths hispid-ciliate, often pilose; blades narrow, linear, pilose, the margins hispid-ciliate. Panicle of 2 conjugate divergent racemes, rarely 1, or still more rarely 3; raceme-rachis not prolonged beyond the uppermost spikelet, not coarsely scabrid-ciliate. Spikelet suborbicular, 1-1.25 mm. long, second glume present. Glume and sterile lemma with short, clavate to faleate or globose, swollen, transparent, unicellular hairs. Fertile lemma shining, minutely papillate-striate. LA Paz: LARECAJA: San Carlos, Mapiri, 850 m., Buchtien. 16 (GH). 20. P. pictum Ekman in Ark. Bot. 10 (17): 11 (1911). Annual, somewhat tufted, to 3.5 (-5) dm. tall, the nodes glabrous. Leaf-sheaths 1966] Flora of Bolivia — Foster 313 glabrous; blades very narrow, linear, flat to conduplicate, scabridulous above, glabrous beneath or with a few papillate-based hispid hairs. Racemes usually 2 (occasionally 1 or 3), somewhat conjugate; raceme- rachis not prolonged beyond the uppermost spikelet, not coarsely long- ciliate. Spikelets obovoid, to 1 mm. long, glabrous, second glume present. Fertile lemma broader than second glume and sterile lemma, markedly papillate-striate at maturity. SANTA CRUZ: SARA: Buena- vista, Steinbach 1894 (GH). 21. P. inaequivalve Raddi, Agrost. Bras. 28 (1823). Perennial, the culms broadly spreading and rooting at the lower nodes, Leaf-sheaths finely pubescent to glabrous, the hairs papillate-based; blades flat, to 25 em. long and 1.5 em. wide, cordate-clasping at the base, sparsely pilose above and beneath with papillate-based hairs. Panicle of few to many spreading non-conjugate racemes, the narrow raceme-rachis not coarsely long-ciliate, not prolonged beyond the uppermost spikelet. Spikelets paired, elliptic, to 2 mm. long. First glume absent or some- times present on 1 spikelet of the pair, very short; second glume half as long as the spikelet, finely pubescent to glabrous. Sterile lemma finely pubescent or glabrous. Fertile lemma shining, pale green-brown. La Paz: NOR YUNGAS: Coripata, Hitchcock 22691 (GH). 22. P. conjugatum Berg. in Act. Helv. Phys. Math. 7: 129 (1762). Perennial, creeping, the flattened culms to 1 m. high, but usually shorter. Leaf-sheaths densely short-ciliate, with an apical ring of long soft hairs; blades flat, to 20 cm. long and 8 mm. wide, the mar- gins shortly scabrid-ciliate, otherwise usually glabrous. Panicle usually of 2 long (10-12 cm.) arcuate, divaricate, conjugate racemes, rarely a third raceme below; raceme-rachis narrow, not coarsely scabrid-ciliate, not extending beyond the uppermost spikelet. Spikelets to 1.5 mm. long, strongly flattened, suborbicular. First glume absent, second glume thin, with a marginal fringe of long silky hairs. Sterile lemma thin, glabrous. Fertile lemma pale, rather shining. LA PAZ: LARE- CAJA: Hacienda Simaco, 1400 m., Buchtien 6325 (GH); NOR YUNGAS: Polo-Polo, 1100 m., Buchtien 268 (GH). SANTA CRUZ: SARA: Buena- vista, 450 m., Steinbach 1862 (GH), 500 m., Steinbach 5160 (GH); Dolores, 450 m., Steinbach 6836 (GH). 23. P. dilatatum Poir. Encycl. 5: 35 (1804). Perennial, to 1.5 m. tall, from a short rhizomatous base. Leaf-sheaths sometimes pilose at the base, otherwise glabrous, ciliate near the apex; blades flat, to 2.5 dm. long and 1.2 em. wide, the base somewhat ciliate. Panicle usually of 3-5 non-conjugate racemes, the narrow rachis not prolonged beyond the uppermost spikelet, the margin not coarsely scabrid-ciliate. Spikelets to 3.8 mm. long, ovoid, acute. First glume absent, second glume covered with long silky hairs, especially dense along the margin. Sterile lemma slightly shorter than the second glume, the margin not densely silky. Fertile lemma pale, finely papillate-striate. TARIJA: CERCADO: Tarija, 2000 m., Cárdenas 202 (GH). 314 Rhodora [Vol. 68 24. F. Urvillei Steud. Syn. Pl. Glum. 1: 24 (1854). Stout peren- nial, to 2.5 m. tall. Leaf-sheaths hirsute, ciliate, or the upper sheaths glabrous; blades to 4.5 dm. long, pilose at the base, otherwise glabrous. Panicle of numerous, ascending, non-conjugate racemes, the raceme- rachis narrow, margin scabrid but not coarsely scabrid-ciliate, not prolonged beyond the uppermost spikelet. Spikelets to 2.5 mm. long, ovoid, obtuse. Second glume and sterile lemma with many long silky hairs on the margin, the glume sparsely hairy on the back, the lemma glabrous on the back. Fertile lemma pale, smooth or nearly so. LA PAZ: LARECAJA: Sorata, Gunther (us). 25. P. notatum Flügge, Monogr, Pasp. 106 (1810). Perennial, from a short rhizome, to 5 dm. tall. Leaf-sheaths ciliate at the apex, other- wise glabrous; blades flat, to 3 dm. long and 1 cm. wide, ciliate at the base, otherwise glabrous. Panicle of 2 divergent racemes, conjugate or nearly so (rarely a third raceme present); raceme-rachis narrow, glabrous, not extended beyond the uppermost spikelet. Spikelets un- paired, glabrous, obovoid to ovoid, about 3 mm. long. Second glume present, as long as the sterile lemma, firm-textured. Fertile lemma rather indurate, finely papillate-striate. SANTA CRUZ: SARA: Dolores, 450 m., Steinbach 1833 (GH); Buenavista, 450-500 m., Steinbach 5273 (GH), 7020 (GH). TARIJA: O'CONNOR: near Entre Rios, 1300 m., West 8249 (GH). 26. P. minus Fourn. Mex. Pl. 2: 6 (1886). Densely matted peren- nial, to 3 dm. high. Leaf-sheaths mostly glabrous, long-ciliate. Panicle of 2 conjugate or nearly conjugate racemes (occasionally 3) ; raceme- rachis narrow, not prolonged beyond the uppermost spikelet, not coarsely scabrid-ciliate. Spikelets solitary, glabrous, to 2.5 mm. long, broadly ovoid, subobtuse. Second glume present, sterile lemma similar but slightly shorter, both with prominent green midribs. Fertile lemma shorter, pale, somewhat shining, finely striate. LA PAZ: LARECAJA: San Carlos, Buchtien 79 (us). 27. P. melanospermum Desv. in Encycl. Suppl. 4: 315 (1816). Spreading annual, eulms to 7 dm. long, the bases decumbent. Leaf- sheaths glabrous; blades flat or folded, to 15 cm. long and 1 em. wide, pilose at the base on the upper surface, otherwise glabrous. Panicle of 2-7 non-conjugate racemes; raceme-rachis very narrow, not coarsely scabrid-ciliate, not extended beyond the uppermost spikelet. Spikelets usually paired, to 2 mm. long, obovoid. Second glume present, thin, somewhat strigose; sterile lemma thin, the margin wrinkled. Fertile lemma dark-brown and shining, obscurely striate. SANTA CRUZ: SARA: Buenavista, Steinbach 5459 (GH). 28. P. plicatulum Michx. Fl. Bor.-Am. 1: 45 (1803). Perennial, 0.5-1 m. tall. Leaf-sheaths usually glabrous but papillate-hispid on the margins; blades to 5 dm. long and 1 cm. wide, papillate-pilose at the base of the upper surface. Panicle of several to many non-conju- gate racemes; raceime-rachis narrow, not coarsely scabrid-ciliate, not 1966] Flora of Bolivia — Foster 815 extended beyond the uppermost spikelet. Spikelets paired, to 3 mm. long, obovoid. Second glume present, thin, sparsely appressed-pubes- cent (occasionally glabrous). Sterile lemma thin, glabrous, trans- versely rugulose near the base. Fertile lemma dark chestnut-brown, shining, striate. LA Paz: NOR YUNGAS: Coripata, Hitchcock 22675 (GH). SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 1829 (GH), 1869 (GH), 6873 (GH), 6887 bis (GH), 7036 (GH). 29. P. collinum Chase in Contrib. U. S. Nat. Herb. 27: 451 (1927). Perennial, to 6.5 dm. tall. Leaf-sheaths glabrous, with a few marginal hairs near the apex; blades flat, to 2 dm. long and 9 mm. wide, rather sparsely long-ciliate. Panicle of 4 distant ascending racemes 2-6 cm. long, the raceme-rachis about 0.7 mm. wide, not coarsely scabrid- ciliate, not prolonged beyond the uppermost spikelet. Spikelets paired, pedicellate, the pedicels glabrous, to 2.5 (-2.7) mm. long. Second glume shortly appressed-pubescent. Sterile lemma more sparsely pubescent than the glume, 5-nerved. Fertile lemma dark brown, papillate-striate, shining. LA PAz: NOR YUNGAS: Coroico, 1560 m., Hitchcock 22723 (US; type). 30. P. virgatum L. Syst. Nat. (ed. 10) 2: 855 (1759). Stout peren- nial, to 1-2 m. tall. Leaf-sheaths papillate-hirsute on the margins and at the apex; blades flat, to 7.5 dm. long and 2.5 em. wide, upper surface papillate-hirsute at the base. Panicle of several to many racemes; raceme-rachis narrow, not prolonged beyond the uppermost spikelet, margin seabrid-ciliate, with scattered, long, hispid hairs. Spikelets paired, to 2.5 mm. long, obovoid. Second glume with long silky hairs around margin. Sterile lemma puberulent to glabrous. Fertile lemma shining chestnut-brown, papillate-striate. La PAZ: LARECAJA: San Carlos, Mapiri, 850 m., Buchtien 52 (GH); NOR YUNGAS: Hacienda El Choro, Coripata, 1700 m., Buchtien 8044 (GH). 31. P. conspersum Schrad. ex Schult. Mant. 2: 174 (1817). Peren- nial, to 1.2 m. tall. Leaf-sheaths glabrous or papillate-pilose; blades sometimes subcordate at the base, flat, sometimes papillate-pubescent on the upper surface. Panicle of several (to 16) non-conjugate racemes; raceme-rachis narrow, the margin scabridulous to scabrid, but without long hispid hairs, not prolonged beyond the uppermost spikelet. Spikelets paired, to 3 mm. long, ellipsoid, acute. Second glume and sterile lemma thin, 5-nerved, one or both shortly pubescent. Fertile lemma chestnut-brown, papillate-striate. LA PAZ: NOR YUNGAS: Coripata, Hitchcock 22696 (US). 32. P. millegrana Schrad. ex Schult. Mant. 2: 175 (1824). Stout perennial, 1-2 m. tall. Leaf-sheaths glabrous or with a few hairs near the apex; blades folded, to 7.5 dm. long and 1.5 cm. wide, the upper surface often papillate-pubescent. Panicle of several to many non- conjugate racemes to 1.5 dm. long; raceme-rachis coarsely scabrid- serrate and sparsely hispid-ciliate, not prolonged beyond the uppermost spikelet. Spikelets paired, obovoid to suborbicular, glabrous, to 316 Rhodora [Vol. 68 2.5 mm. long. Second glume and sterile lemma firm, 3-nerved. Fertile lemma pale, finely papillate-striate. BENI: YACUMA: Reyes, Cárdenas (M. E.) 1656 (Hitchcock). 33. P. densum Poir. Encycl. 5: 32 (1804). Stout perennial, to 1.2 m. tall. Leaf-sheaths glabrous, or the margin ciliate, or somewhat pubescent to hirsute at the apex; blades folded, often to 1 m. long and 2 em. wide, but generally shorter and narrower. Panicle of many non-conjugate racemes; raceme-rachis narrow, not prolonged beyond the uppermost spikelet, coarsely scabrid and densely hispid-ciliate. Spikelets paired, suborbicular, to 2.2 mm. long, glabrous. Second glume and sterile lemma 3-nerved, thin. Fertile lemma pale, shining, finely papillate-striate. SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 3264 (GH), 6898 (GH). 34. P. remotum Remy in Ann. Sci. Nat. (ser. 8) Bot. 6: 349 (1846). Perennial, decumbens to 10 dm. tall, glabrous. Leaf-blades to 2 dm. long and 1 cm. wide, the margins strongly scabrous. Panicle of 2-5 non-conjugate racemes; raceme-rachis narrow, not coarsely scabrid- ciliate, not prolonged beyond the uppermost spikelet. Spikelets paired, to 5.2 mm. long, ovoid. Second glume and sterile lemma green, pubes- cent, the glume 5-nerved, the lemma 3-nerved, Fertile lemma pale, shining, finely papillate-striate. LA PAZ: LARECAJA: Sorata, Mandon 1252 (GH), Bang 1312 (GH). COCHABAMBA: CERCADO: Cochabamba, Hitchcock 22807 (GH); Mollemolle, 2800 m., Steinbach 4055 (GH). 35. P. pilosum Lam. Tabl. Encycl. 1: 175 (1791). Perennial, some- what tufted, to 11.5 dm. tall, usually shorter. Leaf-sheaths papillate- pubescent to glabrous, with ciliate margins; blades flat, to 4 dm. long and 8 mm. wide, pubescent above and beneath, margins and base often with papillate-hispid hairs. Racemes solitary, to 15 cm. long, the narrow rachis with few to many papillate-based hispid hairs, not prolonged beyond the uppermost spiketlet. Spikelets paired, glabrous, ovoid, to 3 mm. long. First glume small to minute; second glume 5-nerved, firm, shorter than mature fruit. Sterile lemma 5-nerved, firm. Fertile lemma pale, markedly papillate-striate. LA PAZ: LARE- CAJA: San Carlos, Mapiri, 850 m., Buchtien 15 (GH). 36. P. Ekmanianum Henr. in Meded. Rijks Herb. Leiden, no. 40: 49 (1921). Perennial, to 6 dm. tall. Leaf-sheaths ciliate; blades to 1 dm. long, the upper surface papillate-pilose. Racemes solitary (very rarely 2 racemes on a culm), the narrow rachis not prolonged beyond the uppermost spikelet, not coarsely scabrid-ciliate. Spikelets paired, oblong-ellipsoid, to 2 mm. long. Second glume and sterile lemma covered with white hairs of varying lengths, the longest more than twice the length of the spikelet. Fertile lemma pale, rather shining, smooth. SANTA CRUZ: SARA: Buenavista, 450-500 m., Steinbach 1818 (GH), 5320 (Gu), 6923 (GH). 37. P. malacophyllum Trin. Gram. Pan. Icon. 3: t. 271 (1831). Perennial, to 1-2 m. tall. Leaf-sheaths sparsely papillate-pilose; 1966] Flora of Bolivia — Foster 317 blades to 25 cm. long and 2.5 cm. wide, densely to sparsely pubescent beneath. Panicle of numerous non-conjugate racemes; raceme-rachis narrow, sparsely ciliate, not prolonged beyond the uppermost spikelet. Spikelets paired, mostly less than 2 mm. long, often purplish, glabrous. Glumes absent, sterile lemma firm, midrib prominent. Fertile lemma often purplish, strongly ribbed, papillate-striate. LA PAZ: LARECAJA: Hacienda Casana, Buchtien 7110 (US). 38. P. boliviense Chase in Contrib. U. S. Nat. Herb. 24: 454 (1927). Perennial, 1-1.5 m. high. Leaf-sheaths pubescent or pilose at the base, otherwise glabrous; blades flat, to 2 dm. long and 2 cm. wide, margins scabrid, occasionally a few long hairs near the base, otherwise glabrous. Panicle of numerous non-conjugate racemes; raceme- rachis puberulent, narrow, not prolonged beyond the uppermost spikelet. Spikelets paired, usually purplish, mostly over 2 mm. long, glabrous. Glumes absent. Sterile lemma with a prominent midrib. Fertile lemma strongly 7-nerved, strongly papillate-striate. LA Paz: LARECAJA: Sorata, Bang 1306 (GH; type-number) ; prope Munaypata, Cerro del Iminapi, 2650 m., Mandon 1253 (GH). COCHABAMBA: CERCADO: Cochabamba, Hitchcock 22866 (US). 89. P. saccharoides Nees in Trin. Gram. Icon. 1: t. 107 (1828). Perennial, stoloniferous, to 1.2 m. high. Leaf-sheaths usually glabrous, sometimes papillate-pilose or papillate, densely ciliate; blades flat, to 3 dm. long and 1.5 em. wide, the upper surface papillate-pilose, the lower surface glabrous. Panicle of many slender drooping racemes, more or less flabellately arranged; raceme-rachis very narrow, hardly winged, prolonged beyond the uppermost spikelet. Spikelets solitary, distant, lanceolate, acute or acuminate, to 3 mm. long. Second glume often puberulent on the back, marginal fringe of long white hairs, mixed with shorter hairs. Sterile lemma thin, glabrous, shorter than the glume. Fertile lemma thin, not indurate. LA PAZ: LARECAJA: San Carlos, Sarampiuni (Mapiri), 500 m., Buchtien 65 (GH); NOR YUNGAS: Coripata, Hitchcock 22686 (GH). 82. Panicum L. Annual or perennial; leaf-blades not or barely reaching the inflo- rescence, rarely exceeding it, linear to lanceolate to elliptic-oblong, acute, glabrous or variously pubescent. Inflorescence a simple to complex panicle, with spikelets from densely racemose on the branches to remote and solitary at the ends of the branchlets. Spikelets pedi- cellate, disarticulating below the 2 glumes; 2-flowered, the lower floret a sterile lemma (2 sterile lemmas in P. quadriglume), or some- times a staminate floret, the upper floret perfect; fruit indurate or chartaceous-indurate, the lemma-margins inrolled and partly conceal- ing the similar palea. a. Inflorescence simple, a central axis with the primary branches densely flowered one-sided racemes. 318 Rhodora [Vol. 68 b. Primary branches unbranched, short, very densely flowered. c. Spikelets hispidulous to hispid. d. Sterile lemma with 2 crateriform glands; fruit not rugose. — ÁREA 16. P. pulchellum. d. Sterile lemma glandless; fruit rugose. .... 1. P. echinulatum. c. Spikelets glabrous. e. Longest leaf-blades not over 5 cm, long and 2 mm. wide. stsvonscconsasecccesesonssssssvecassecessssocnssssenensenscstseseca sens 14. P. stoloniferum. e. Longest leaf-blades 9-15 cm. long and 1.5-2 em. wide. f. Inflorescence-axis pilose with long hairs; second glume 1.5 mm. long. .......... eee 11. P. pilosum. f. Inflorescence-axis puberulous to short-pubescent, but not pilose; second glume over 2 mm. long. ........... ces /— —————— ÁREA 15. P. frondescens. b. Primary branches branched, usually from the base, less densely flowered. g. Stem-nodes pubescent to hirsute. ............ 10. P. polygonatum. g. Stem-nodes glabrous. h. Longest leaf-blades to 20 em. long and 1.4 em. wide, basally cordate and subpetiolate. .............. eee 13. P. boliviense. h. Longest leaf-blades to 10 cm. long and 1 em. wide, basally not cordate nor subpetiolate. .................. ess 12. P. laxwm. a. Inflorescence complexly branched, not composed of one-sided racemes; if racemose, the racemes not one-sided and the spikelets not densely crowded. i. Spikelets not solitary at ends of branchlets. j. Spikelets hirsutulous to long-hirsute. k. Spikelets 5.5-7 mm. long; fruit TUQOSE., ....esssssssseeee nee wenaspsssesscsssesgnsavapagsassaeeanusssessssssscsssesesvasevonevsse 29. P. paucispicatum. k. Spikelets 4 mm. long or less; fruit not rugose. l. Spikelets long-hirsute, 3.5-4 mm. long. .. 32. P. procurrens. l. Spikelets hirsutulous, about 2 mm. long. neseser. breresesssessaeseteecctesesoooserosnearreesesseosoesssseosocoreneenase 26. P. pantrichum. j. Spikelets glabrous. m. Primary panicle-branches (except at uppermost nodes) in vertieils, ......sesseseseeeeeee eere eerte 30. P. megiston. m. Primary panicle-branches not in verticils (except some- times at the lowest node only). n. First glume 1 mm. or less. ........ 2. P. aquaticum. n. First glume 2-3 mm. o. Leaf-blades to 10 em. long and 1 em. wide. ........ — ÉD 27. P. cordovense. o. Leaf-blades to 20 em. long and 4 mm. wide. o. Ssaasaoesnasunpdeansesessacesbssusseotersneesssarenessseoons 9. P. tricholaenoides. i. Spikelets remote, solitary at ends of branches, or appearing so. p. Spikelets minutely puberulent to sparsely hispidulous. 1966] Flora of Bolivia — Foster 319 q. Inflorescence-axis glabrous; first glume less than 0.5 mm. lup ER ME E 17. P. trichanthum. q. Inflorescence-axis not glabrous; first glume 1-2 mm. long. r. Spikelets 2 mm. long; axis and branches pilose below. IA aid eI pdt teas Ste cuc cc ish) Ras RENE VES IRIS SEM 25. P. sciurotis. r. Spikelets 3.5 mm. long; axis and branches hirsute-hispid TN We NN e er mRNA 23. P. Rudgei. p. Spikelets glabrous. s. Spikelets 6-8 mm. long. ....... 31. P. olyroides. s. Spikelets less than 5 mm. long. t. Spikelets with 2 sterile lemmas. ........ 7. P. quadriglume. t. Spikelets with 1 sterile lemma. u. Fruit rugose or rugulose. v. Second glume to 3.5 mm. long; fertile lemma 2.5 mm. long, rugose. ......nennenn een 8. P. maximum. v. Second glume 2 mm. long; fertile lemma 2 mm, long, PUgaOlone. asiento A e HA 24. P. Sellowii. u. Fruit not rugose nor rugulose. w. Leaf-sheaths papillate-pilose. x. Spikelets 3-4 mm. long. y. Spikelets 3.5-4 mm. long, acute; first glume 1 mm. long. .............. eene eene onnnne 5. P. stramineum. y. Spikelets 3 mm. long, acute to acuminate; first glume 1.5-2 mm. long. ..............- 6. P. peladoénse. x. Spikelets not over 2.5 mm. z. First glume 1.5 mm. long; leaf-blade to 1 cm. wide; fertile lemma about 1.5 mm. A. Leaf-blades pilose above and beneath. ............ Hox c cU NUMES S 3. P. cayennense. A. Leaf-blade glabrous above, pubescent to glab- rate beneath. odi denn 4. P. hirticaule. z. First glume 1 mm. long; leaf-blade to 1.5 cm. wide; fertile lemma about 2 mm. long. «+ "E ansa cm PM MM E A 24. P. hebotes. w. Leaf-sheaths glabrous; or, if somewhat pubescent, not papillate-pilose. B. Glumes distinctly unequal. C. First glume 1 mm. long; leaf-blades 4 mm. wide, not cordate at base. seiirt 19. P. cyanescens. C. First glume 0.5 mm. long; leaf-blades to 1 cm. wide, cordate-clasping at base. «scenes MENTRE D Ir er 20. P. nervosum. B. Glumes subequal or nearly so. D. Spikelets viscid-glutinous. .... 18. P. glutinosum. D. Spikelets not viscid-glutinous. E. Leaf-blades about 2 cm. long. ............ em acce E See. - 22. P. parvifolium. 320 Rhodora [Vol. 68 REN 28. P. versicolor. l. Panicum echinuülatum Mez in Notizbl. 7: 62 (1917). P. echinu- latum. var. boliviense Henr. in Meded. Rijks Herb. Leiden, no. 40: 50 (1921). Annual, spreading, culms to 3.5 dm. high. Leaf-sheaths pubescent along the margins, otherwise glabrous; blades lanceolate, acute, to 11 em. long and 1.4 cm. wide, firm-textured, the thickened margins undulate and ciliolate, glabrous above and beneath. Panicles less than 10 em. long, axis, branches and pedicels glabrous, scabri- dulous on the margins. Spikelets to 4 mm. long, usually about 3.5 mm., acute, hispid. Glumes very unequal; first glume broadly ovate, abruptly acute, to 1.5 mm. long, 3-nerved; second glume 3.5 mm. long, acute, 5-nerved. Sterile lemma as long as second glume and similar in appearance. Fertile lemma shorter than the sterile lemma, sub- obtuse, indurate, transversely rugose. TARIJA: ENTRE RIOS: Villa Montes, Pflanz 607 (Hitchcock). SANTA CRUZ: CERCADO: Santa Cruz, 450 m., Steinbach 5259 (GH). 2. P. aquaticum Poir. Encycl. Suppl 4: 281 (1816). Perennial, often decumbent and spreading, in wet places or actually in water, the culms to 10 dm, tall. Leaf-sheaths glabrous, the lance-linear blades to 25 cm. long and 1 em. wide, acute, glabrous. Panicle to 25 cm. long at maturity, the branches somewhat erect but spreading, axis, branches and pedicels glabrous, their margins sometimes scabridulous. Spikelets glabrous, acute, to 3 mm. long. Glumes quite unequal; first glume 1 mm. long, or less, truncately acute, broadly clasping at the base, l-nerved; second glume 3 mm. long, acute, about 9-nerved. Sterile lemma similar to second glume, slightly shorter and fewer- nerved. Fertile lemma about 2 mm. long, acute, shining, longitudinally Striatulate. SANTA CRUZ: SARA: Buenavista, Steinbach 3180 (GH). 2a. P. aquaticum var. chloroticum (Nees ex Trin.) R. C. Foster, comb. nov. P. chloroticum Nees ex Trin, Gram. Pan. 236 (1826). Sympatric with and similar in all respects to P. aquaticum except for smaller spikelets, 2-2.5 mm. long. BENI: YACUMA: Reyes, Rusby (M. E.) 1636 (vs). 3. P. cayennense Lam. Encycl. Tabl. 1: 173 (1791). Annual, erect, the culms to 5 dm. tall. Leaf-sheaths papillate-pilose, the linear blades to 16 cm. long and 1 cm. wide, pilose above and beneath. Panicles to 20 em. long, rather open, the axis hispid-pilose, branches and pedicels glabrous, the margins scabridulous. Spikelets to 2.5 mm. long, usually a little less, glabrous, somewhat acute. Glumes unequal; first glume nearly 1.5 mm. long, acute, 5-nerved; second glume a little over 2 mm. long, acute, 7-nerved. Sterile lemma as long as second glume, 5-nerved. Fertile lemma 1.5 mm. long, obtuse, firm, longitudinally striatulate. SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 6935 (GH). 4. P. hirticaule Presl, Rel. Haenk. 1: 308 (1830). Annual, spread- 1966] Flora of Bolivia — Foster 321 ing, the culms usuaily less than 5 dm. tall. Leaf-sheaths variably hispid, but mostly papillate-pilose, the blades lance-linear, to 15 cm. long and 8 mm. wide, variably hispid to pubescent or glabrate be- neath. Panicle open, to 15 cm. long, usually shorter, axis and main branches sparsely long-hispid, branchlets and pedicels scabridulous on the margins. Spikelets about 2.5 mm. long, acute, glabrous. Glumes unequal; first glume 1.5 mm. long, acute, 3-5-nerved, the nerves not anastomosing; second glume about 2.5 mm. long or a little shorter, 7-nerved. Sterile lemma about 2.5 mm. long. Fertile lemma about 1.5 mm. long, shining, warm brown, longitudinally striatulate. BENI: CERCADO MOJOS: Trinidad, 200 m., Cárdenas 3534 (GH). SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 7086 bis (GH). 5. P. stramineum Hitchc. & Chase in Contrib. U. S. Nat. Herb. 15: 67 (1910). Annual, erect or spreading, the culms to 5 dm. tall but usually shorter. Leaf-sheaths usually papillate-pilose, but some- times almost glabrous, the lance-linear blades to 20 cm. long and 8 mm. wide, glabrous to sparsely pilose, at least beneath, the margins sparsely long-ciliate to glabrous. Panicle open, about 15 cm. long, axis, branches and pedicels glabrous but scabridulous on the margins. Spikelets 3.5-4 mm, long, glabrous, acute. Glumes very unequal; first glume about 1 mm. long, obtuse or abruptly acute, 3-5-nerved, the nerves anastomosing; second glume 3 mm. long, acute, many-nerved. Sterile lemma as long as or slightly longer than the second glume, subobtuse, many-nerved. Fertile lemma about 2.5 mm. long, shining, firm-textured. BENI: YACUMA: Reyes, Cárdenas (M. E.) 1647 (US). 6. P. peladoénse Henr. in Blumea, 4(3): 504 (1941). Perennial, to 4 dm. tall. Leaf-sheaths papillate-pilose; blades scabridulous-puberu- lent above and beneath, sparsely papillate-hirsute above and beneath, the long hairs hispid, blade to 4 mm. wide. Panicle to 12 cm. long, axis and branches glabrous, branchlets and pedicels scabridulous. Spikelets glabrous, acute to acuminate, 3 mm. long. Glumes unequal; first glume ovate, 1.5-2 mm. long, subacute, 5-nerved; second glume 2.5-3 mm. long, 5-nerved, acute. Sterile lemma about 3 mm. long, or slightly longer, acute to acuminate. Fertile lemma about 2 mm. long, dark-brown, shining, obtuse. SANTA CRUZ: VALLEGRANDE: Cuesta de los Monos, 1400 m., Herzog 1896j (cited in original descrip- tion; not seen). The only material of this species which has been seen is an isotype, Balansa 4357, in the United States National Herbarium. The description is based entirely on this. Since no Bolivian material has been seen in two large herbaria, it is possible that the attribution of the species to Bolivia rests on a misdetermination. 7. P. quadriglume (Doell) Hitchc. in Contrib. U. S. Nat. Herb. 24: 460 (1927). Perennial, the culms to 4 dm. tall, the nodes densely 322 Rhodora [Vol. 68 pilose. Leaf-sheaths papillate-pilose, blades to 20 cm. long and 5 mm. wide, linear, pilose above and beneath. Panicle rather open at ma- turity, to 15 em. long, axis, branches and pedicels glabrous, the branches and pedicels scabridulous on the margins, Spikelets to 3 mm. long, glabrous, acute, with 2 sterile lemmas. Glumes unequal; first glume 1.5 mm. long, broad, acute, 5-nerved, the lateral nerves more or less anastomosing; second glume 2.5 mm. long, 7-nerved. Sterile lemmas 3 mm. long, about 7-nerved. Fertile lemmas about 2 mm. long, firm-textured, shining, dark brown (paler in immaturity). LA PAZ: NOR YUNGAS: Milluguaya, 1100 m., Buchtien 4174 (GH). SANTA CRUZ: SARA: Buenavista, 450-500 m., Steinbach 5299 (GH), 7076 bis (GH), 6979 (at least as to GH sheet). 8. P. maximum Jacq. Coll. Bot. 1: 76 (1786). Perennial, to 2 m. tall, culms with pubescent to somewhat hirsute nodes. Leaf-sheaths glabrous to sparsely papillate-hirsute, blades to 30 em. long and 7 mm. wide, linear, glabrous above and beneath. Panicle open, to 3.5 dm. long, axis, branches and pedicels glabrous, the branches often shortly pubescent at the very base. Spikelets to 3.5 mm. long, glabrous, acute. Glumes unequal; first glume 1.5 mm. long, rather obtuse, obscurely nerved; second glume to 3.5 mm. long, about 7-nerved, the nerves rather obscure. Sterile lemma about as long as the second glume, 7-nerved. Fertile lemma 2.5 mm. long, strongly rugose, the apex often minutely puberulent. LA PAz: NOR YUNGAS: Milluguaya, 1200 m., Buchtien 442 (in 1917) (GH); Polo-Polo, near Coroico, 1100 m., Buchtien 442 (in 1912) (am). 9. P. tricholaenoides Steud. Syn. Pl. Glum. 1: 68 (1854). ? P. giganteum O. Ktze. Rev. Gen. 3(2): 360 (1898). Perennial, slender, erect, rhizomatous, the culms to 1 m. tall. Leaf-sheaths glabrous, the glabrous linear blades to 20 em. long and 4 mm. wide, often inrolled. Panicle rather narrow, to 20 em. long, the branches erect to rather spreading, axis glabrous, branches scabrid on the margins, the ulti- mate pedicels often ciliate with long stiff hairs, a group of such hairs often subtending the spikelets. Spikelets to 3 mm, long, glabrous, acute. Glumes unequal; first elume to 2 mm. long, rather narrowly acute, 3-nerved; second glume as long as the spikelet, 5-7-nerved. Sterile lemma shorter than second glume, obscurely nerved. Fertile lemma less than 2 mm. long, somewhat obtuse, shining, firm. SANTA CRUZ: SARA: Buenavista, campos de Dolores, 450 m., Steinbach 3220 (GH). 10. P. polygonatum Schrad. ex Schult. Mant. 2: 256 (1824). Peren- nial, somewhat sprawling, the culms 2-6 dm. tall, the nodes pubescent to hirsute. Leaf-sheaths ciliate, the lanceolate blades to 20 cm. long and 2 cm. wide, pilose above and beneath, at least when young, long- acute, the base subcordate and often subpetiolate. Panicle spreading, to 2.5 dm. long, the spikelets unilateral on the branches, the axis and branches scabridulous and sparsely hispid, the pedicels scabri- 1966] Flora of Bolivia — Foster 828 dulous. Spikelets about 1.5 mm. long, acute, glabrous. Glumes unequal; first glume i mm. long, 1-nerved, the nerve apically scabri- dulous, acute; second glume as long as the spikelet, 5-nerved. Sterile lemma shorter than second glume, about 3-nerved, Fertile lemma 1 mm. long or slightly longer, acute, shining. LA PAZ: LARECAJA: Hacienda Simaco, 1400 m., Buchtien 5334 (GH); San Carlos, Mapiri, 850 m., Buchtien 74 (GH). SANTA CRUZ: SARA: Buenavista, 400 m., Steinbach 5231 (GH); Rio Surutá, 500 m., Steinbach 6840 (GH). 11. P. pilosum Swartz, Prodr. 22 (1788). Spreading to ascending perennial, the culms to 6 dm. tall. Leaf-sheaths long-ciliate on the margins, otherwise glabrous to slightly pilose, the broadly linear blades to 15 em. long and 1.5 em. wide, usually glabrous beneath, scattered- pilose above, acute, abruptly rounded at the base. Panicle to 15 cm. long, the main axis glabrous or somewhat puberulent, the primary branches with long stiff hairs much exceeding the spikelets, pedicels glabrous to subpuberulous, the component racemes short (seldom more than 3 em. long), densely flowered, ascending to somewhat spreading, usually arranged in verticils. Spikelets about 1.5 mm. long, sub- acute. Glumes unequal; first glume 1 mm. long, 3-nerved; second glume as long as the spikelet, 5-nerved. Sterile lemma like second glume, but sometimes slightly shorter, rather blunt, obscurely 5-nerved. Fertile lemma shorter than sterile, indurate, shining. LA PAZ: LARE- CAJA: San Carlos, Mapiri, 750 m., Buchtien 78 (GH). 12. P. laxum Swartz, Prodr. 23 (1788). Perennial, basally decum- bent or spreading, culms 2-6 dm. tall. Leaf-sheaths densely ciliate, blades to 10 cm. long and 1 cm. wide, lanceolate, acute, the base abruptly contracted, glabrous above and beneath. Panicle fairly open, to 2 dm. long, the branches spreading and branched, the numerous spikelets unilaterally arranged, axis, branches and pedicels glabrous but the margins seabridulous. Spikelets to 1.5 mm. long, acute, glabrous. Glumes unequal; first glume less than 1 mm. long, 1-nerved, acute; second glume as long as the spikelet, 5-nerved. Sterile lemma as long as second glume, several-nerved, rather obtuse. Fertile lemma about 1 mm. long, shining. YUNGAS: Bang 308a (GH). LA PAZ: LARECAJA: Charopampa, 570 m., Buchtien 1168 (GH); San Carlos, 750 m., Buchtien 1179 (GH); Hacienda Simaco, 1400 m., Buchtien 5315 (GH). SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 2064 (GH), 5161 (GH), 5338 (GH), 6850 (GH); VALLEGRANDE: Coma- rapa, Yungas de San Mateo, 2500 m., Steinbach 8508 (GH). 13. P. boliviense Hack. in Fedde, Rep. Sp. Nov. 11: 19 (1912). Perennial, decumbent to spreading, the culms to 3-5 dm. tall. Leaf- sheaths ciliate, blades lanceolate to lance-oblong or occasionally lance- linear, to 20 cm. long and 1.4 cm. wide, rather long-acute, the base cordate to subcordate and subpetiolate, glabrous above and beneath. Panicle quite open at maturity, to 20 cm. long, axis, branches and pedicels often scabridulous on the margins, the lower primary branches 324 Rhodora [Vol. 68 sometimes sparsely hispid in the axils, the spikelets more loosely arranged than in P. laxum. Spikelets 1.5 mm. long, acute or sub- obtuse, glabrous. Glumes unequal; first glume about 0.75 mm. long, l-nerved or rather obscurely 3-nerved; second glume as long as the spikelet, 5-nerved. Sterile lemma as long as second glume, 5-nerved, subobtuse. Fertile lemma slightly longer than 1 mm., subobtuse, not markedly shining but rather dull, obscurely 3-nerved, longitudinally striatulate. SANTA CRUZ: SARA: Buenavista, 400-450 m., Steinbach 6857 (US, GH), 6944 bis (GH). COCHABAMBA: CHAPARE: Antahuacana, Buchtien 2501 (US; type-number). 14. P. stoloniferum Poir. Encycl. Suppl. 4: 274 (1816). Perennial, creeping and spreading, the flowering culms to 3 dm. tall Leaf- sheaths pubescent, at least sparsely so, blades lanceolate, to 3.5-4 cm. long and 5 mm. wide, puberulent above and beneath, at least when young, acute, abruptly narrowed at the base and usually subpetiolate. Panicle small, occasionally to 5 cm. long, the component racemes seldom over 1 cm. long, axis glabrous or nearly so, branches and pedicels glabrous to somewhat puberulous. Spikelets 2.5-3 mm. long, acute, glabrous. Glumes very unequal; first glume 1 mm. long, or slightly longer, 3-nerved, the central nerve prominent; second glume 2.5 mm. long, 5-nerved. Sterile lemma about 3 mm. long, 5-nerved. Fertile lemma 1.5 mm. long, indurate, shining, acute, obscurely nerved, not rugose. LA PAZ: LARECAJA: Mapiri, Buchtien 1167 (Us). 15. P. frondescens Meyer, Primit. Fl. Esseq. 56 (1818). Annual (?), decumbent, rooting at the nodes, the flowering culms to 8 or even 5 dm. tall. Leaf-sheaths ciliate, blades to 9-10 em. long and 2 cm. wide, lance-ovate, acuminate, the wide base abruptly narrowed and shortly petiolate, scabridulous above and beneath. Panicles to 13 cm. long, main axis puberulent to pubescent, branches and pedicels glabrous. Spikelets to 3 mm. long, usually shorter, acute, glabrous. Glumes very unequal; first glume 1 mm, long, broadly ovate, abruptly acute, rather obscurely 3-nerved, the midrib scabridulous above; second glume to 2.25 mm. long, shorter than the spikelet, acute, 5-nerved. Sterile lemma similar to second glume but 2.5-3 mm. long. Fertile lemma about 1.5 mm. long, narrow, acute, indurate, shining, smooth. LA PAZ: NOR YUNGAS: Polo-Polo, near Coroico, 1100 m., Buchtien 264 (GH), 1173 (GH). SANTA CRUZ: SARA: Buenavista, 400 m., Steinbach 5130 (GH), 6855 (GH). 16. P. pulchellum Raddi, Agrost. Bras. 42 (1823). Slender spreading perennial, the flowering culms to 2-3 dm. tall. Leaf-sheaths shortly ciliate, the blades lance-ovate, to 6 em. long and 1.7 em. wide, basally subcordate, glabrous above, sometimes sparsely puberulent beneath. Panicle to 10-11 em. long, the branches remote, axis and branches sparsely pubescent to hispidulous, the pedicels mostly glabrous. Spikelets to 2.5 mm. long, from nearly glabrous to hispidulous or hispid, acute. Glumes unequal; first glume 1 mm. or slightly less, 1966] Flora of Bolivia — Foster 825 ovate, acute, 3-nerved; second glume 2 mm. long, ovate, acute, 5- nerved. Sterile lemma about 2 mm. long, long-acute, 5-nerved. Fertile lemma 1.5 mm. long, chartaceous, shining, not rugose. LA PAZ: SUR YUNGAS: Guanay, Rusby 217 (US). 17. P. trichanthum Nees, Agrost. Bras. 210 (1829). Straggling perennial, the culms to 1 m. tall. Leaf-sheaths long-ciliate, otherwise glabrous, blades to 15 cm. long and 2 cm. wide (rarely wider), sparsely to densely pubescent above and beneath, at least when young, lanceo- late, acute, basally abruptly rounded and somewhat clasping. Panicle large, diffuse, to 20 cm. long, axis, branches and pedicels glabrous. Spikelets minutely puberulous to glabrous, solitary, to 1.5 mm. long. Glumes very unequal; first glume less than 0.5 mm. long, white- papery, not obviously nerved; second glume as long as the spikelet, 5-nerved. Sterile lemma as long as second glume and similar in appearance. Fertile lemma less than 1.5 mm, long, indurate, not rugose, shining. LA PAZ: NOR YUNGAS: Coroico, Buchtien 445 (GH), 446 (US); SUR YUNGAS: Chulumani, Hitchcock 22660 (us). SANTA CRUZ: CORDILLERA: Puerto Suarez, Agnes Chase 11155 (GH). 18. P. glutinosum Swartz, Prodr. 24 (1788). Perennial, to 1-2 m. tall, decumbent at the base. Leaf-sheaths glabrous with a hispid collar at the apex, the blades to 30 cm. long and 2.5 em. wide, the margins hispid-ciliate, especially at the base, otherwise glabrous or glabrescent. Panicle open, to 3 dm. long, axis, branches and pedicels glabrous, or the margins finely scabridulous. Spikelets glutinous, subobtuse, to 3-3.5 mm. long. Glumes approximately equal; first glume 2.25 mm. long, broad, 5-nerved, obtuse; second glume about 2.5 mm. long, 7-nerved, obtuse. Sterile lemma 3 mm. long, 5-nerved. Fertile lemma 2.5 mm. long, firm-textured, minutely puberulent at the apex. LA Paz: LARECAJA: Mapiri, 830 m., Rusby 244 (GH); Hacienda Simaco, near Tipuani, 1400 m., Buchtien 5316 (GH). SANTA CRUZ: SARA: Monte del Fraile, Buenavista, 450 m., Steinbach s. n. (GH). 19. P. cyanescens Nees, Agrost. Bras. 220 (1829). Slender peren- nial, the culms to 6 dm. tall. Leaf-sheaths glabrous, sparsely hispid- pilose at the apex, the linear blades mostly less than 10 em. long, 4 mm. wide, glabrous above and beneath. Panicle diffuse, less than 10 cm. lone, axis, branches and pedicels glabrous. Spikelets to 2 mm. long, ovoid, glabrous. Glumes unequal; first glume 1 mm. long, sub- obtuse, 1-nerved, with 2 obscure lateral nerves; second glume 1.5 mm. long, obtuse, 5-nerved. Sterile lemma slightly over 1 mm. long, obscurely nerved. Fertile lemma 1.5 mm. long, shining, longitudinally striatulate. SANTA CRUZ: SARA: Buenavista, 500 m., Steinbach 5175 (GH), 6940 in part (GH). 20. P. nervosum Lam. Encycl. 4: 747 (1798). Perennial, the culms 3.6 dm. tall. Leaf-sheaths somewhat hispid-ciliate, otherwise glabrous, the blades to 8-9 em. long and 1 cm. wide, glabrous above and beneath, acute, cordate-clasping. Panicle diffuse, to 12-15 cm. long, axis, 326 Rhodora [Vol. 68 branches and pedicels glabrous. Spikelets about 1.5 mm. long, glabrous, rather narrowly ovoid, subobtuse. Glumes markedly un- equal; first glume 0.5 mm. long, obtuse to subacute, obscurely nerved; second glume 1.5 mm. long, obscurely 7-nerved. Sterile lemma slightly shorter than second glume, or equal to it. Fertile lemma slightly over 1 mm. long, subacute, shining, longitudinally striatulate. SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 1213 (GH); CERCADO: banado de Piray, 450 m., Steinbach 1980 (GH). 21. F. Sellowii Nees, Agrost. Bras. 158 (1829). Perennial, spreading, the culms to 1 m, tall. Leaf-sheaths ciliate, pubescent to pilose, often glabrate with age, the lance-linear blades to 20 cm. long and 2 em. wide, pilose above and beneath, or glabrate with age, acute, the base often abruptly rounded and subpetiolate. Panicle very open at ma- turity, to 15 em. long, the branches remote and with the relatively few spikelets along the upper half on slender pedicels, axis and branches sparsely pilose below, pedicels and upper branches scabri- dulous. Spikelets 2-2.5 mm. long, glabrous. Glumes unequal; first glume to 1.5 mm. long, acute, l-nerved; second glume 2 mm. long, 5-nerved, subacute. Sterile lemma to 2.5 mm. long, obscurely 5-nerved. Fertile lemma about 2 mm. long, shining, transversely rugulose. LA PAZ: LARECAJA: San Carlos, near Mapiri, 850 m., Buchtien 53 (GH); Hacienda Simaco, 1400 m., Buchtien 5317 (GH); NOR YUNGAS: Bella Vista, Hitchcock 22752 (GH); SUR YUNGAS: La Florida, 1840 m., Hitchcock 22638 (GH). COCHABAMBA: CHAPARE: Locotal, 1600 m., Steinbach 9095 (GH). 22. P. parvifolium Lam. Encycl. Tabl. 1: 173 (1791). Straggling perennial, the culms to 3 dm. tall. Leaf-sheaths glabrous, the blades about 2 cm. long and 5 mm. wide, pilose above and beneath, occa- sionally glabrous, lanceolate, acute, cordate or subcordate at the base. Panicle open, to 5 cm. long, axis, branches and pedicels glabrous. Spikelets about 1.5 mm. long, glabrous, more or less elliptic-ovoid, obtuse. Glumes somewhat unequal; first glume a little over 1 mm. long, 3-nerved; second glume 1.5 mm. long, 5-nerved, subacute. Sterile lemma as long as second glume, 5-nerved, obtuse. Fertile lemma slightly over 1 mm. long, longitudinally Striatulate, shining, SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 6646 (GH), 6865 (GH), 7011 (GH), 6940 in part (GH). 23. P. Rudgei Roem. & Schult. Syst. Veg. 2: 444 (1817). Tufted perennial, the often flexuose culms to 1 m. tall. Leaf-sheaths densely papillate-pilose, blades more or less linear, acute, to 30 em. long and 8 mm. wide, equaling the inflorescence, puberulent to pubescent above, pubescent to short-pilose beneath. Panicle large, to 3-4 dm. long, the branches erectly spreading, the axis hirsute-hispid below, glabrous above, branches scabridulous on margins, spikelets remote and solitary at ends of branches. Spikelets to 3.5 mm. long, at least the first glume somewhat hispid. Glumes unequal; first glume 2 mm. long, 1966] Flora of Bolivia — Foster 327 5-nerved, very acute, hispid on upper portion; second glume 3 mm. long, 5-nerved, mostly glabrous. Sterile lemma 3 mm. long, glabrous, T-nerved. Fertile lemma to 2.5 mm. long, obtuse, indurate, shining, longitudinally striatulate. LA PAZ: LARECAJA: San Carlos, 950 m., Buchtien 32 (GH). 24. P. hebotes Trin. in Mém. Acad. St. Pétersb. (ser. 6) Sci. Nat. 1: 301 (1834). Perennial, the culms to 6.5 dm. tall. Leaf-sheaths papillate-pilose, the lanceolate blades to 15 em. long and 1.5 em. wide, pilose above and beneath, acute, basally cordate. Panicle rather dif- fuse, to 15 cm. long, axis densely pilose, primary branches more sparsely pilose below, glabrous above, branchlets and pedicels gla- brous. Spikelets 2 ram. long, acute, glabrous. Glumes unequal; first glume 1 mm. long, acute, 3-nerved, sometimes obscurely so; second glume 2 mm. long, acute, 9-nerved. Sterile lemma slightly longer than the second glume, about 7-nerved. Fertile lemma slightly less than 2 mm. long, subacute. YUNGAS: Bang 493 in part (at least as to GH sheet). 25. P. sciurotis Trin. Gram. Pan. 228 (1826). Slender perennial, caespitose, culms to 4 dm. tall, rarely to 6 dm. Leaf-sheaths pubes- cent, blades lanceolate, to 7 cm. long and 1.2 cm. wide, puberulent above and beneath, acute, the base abruptly cordate-clasping. Panicle much-branched, usually less than 10 cm. long, the branching fine, axis and branches pilose below, glabrous above, pedicels glabrous. Spikelets solitary, to 2 mm. long, minutely puberulous to glabrous. Glumes unequal; first glume 1 mm. long, acute, ]-nerved; second glume 1.5 mm. long, subobtuse, 5-nerved. Sterile lemma nearly 2 mm, long, T-nerved, subobtuse. Fertile lemma 1.5 mm. long, acute, shining, more or less indurate. LA Paz: SUR YUNGAS: Chulumani, 1600 m., Hiteh- cock 22669 (GH), 22709 (GH). 26. P. pantrichum Hack. in Verh. Zool. Bot. Ges. Wien, 1915: 72 (1915). Spreading perennial, lower nodes rooting, culms 2-3 dm. tall, but usually shorter. Leaf-sheaths somewhat papillate-pilose, blades lanceolate, to 7-8 cm. long and 8 mm. wide, acute, puberulent above and beneath. Mature panicle spreading, to 15 em. long, axis and branches scabridulous, the long pedicels puberulent to scabridulous. Spikelets 2 mm. long, or a little longer, hirsutulous to nearly glabrous, ovoid, obtuse. Glumes unequal; first glume 1.5 mm. long, 3-nerved; second glume as long as the spikelet, 5-nerved. Sterile lemma as long as the spikelet, 5-nerved. Fertile lemma indurate, smooth, shining, sometimes obscurely striatulate, LA PAZ: LARECAJA: Mapiri, Buchtien 1171 (Us). 27. P. cordovense Fourn. Mex. Pl. 2: 26 (1886). Perennial, strag- gling, culms to 20 cm. tall. Leaf-sheaths papillate-pilose, densely ciliate, the blades to 10 cm. long and 1 em. wide, lanceolate, acute, glabrous. Panicle to 15 cm. long, axis, branches and pedicels glabrous. Spikelets to 3-3.5 mm. long, acute, glabrous. Glumes somewhat un- 328 . Rhodora [Vol. 68 equal; first glume about 3 mm. long, 3-nerved; second glume as long as the spikelet, abruptly acute, 7-nerved. Sterile lemma slightly shorter than second glume, 7-nerved, obtuse. Fertile lemma about as long as the sterile, obtuse, rather indurate, shining. LA Paz: SUR YUNGAS: La Florida, 1840 m., Hitchcock 22647 (GH). 28. P. versicolor Doell in Mart. FI. Bras. 2(2): 254 (1877). Peren- nial, shortly rhizomatous, erect, the culms to 6 dm. tall. Leaf-sheaths hirsute-ciliate, otherwise sparsely pubescent to glabrous, the linear blades to 20 cm. long and 8 mm. wide, pubescent to glabrous or glabrate. Panicle open, to 10-15 cm. long, axis, branches and pedicels glabrous. Spikelets 2.5 to nearly 3 mm. long, glabrous, asymmetric, racemose on remote slender pedicels along the upper portion of branches, appearing solitary at ends of branchlets. Glumes nearly equal; first glume 1 mm. long, subacute to obtuse, obscurely 3-nerved; second glume slightly over 1 mm. long, about 5-nerved. Sterile lemma about 2.5 mm. long, containing a staminate floret, winged, splitting at maturity. Fertile lemma 2 mm. long, dark brown, shining, with a marked dorsal ridge. SANTA CRUZ: SARA: Río Dolores, Steinbach 3542 (GH); Buenavista, 450 m., Steinbach 6643 (GH), 6656 (GH). 29. P. paucispicatum Morong in Ann. N. Y. Acad. 7: 262 (1893). Coarse perennial, to 1 m. high, usually less, the culms pubescent. Leaves oblong to lanceolate, acute, to 6.5 cm. long and 1.5 em. wide, densely short-pubescert and ciliate, blades flat. Panicle about 10 em. long, few-branched, axis, branches and pedicels densely puberulent, the pedicels with an apical collar of long stiff hairs subtending the spikelet. Spikelets fusiform, acuminate, hirsutulous, 5.5-7 mm. long. Glumes unequal, hirsutulous; first glume about 3 mm. long, strongly 5-nerved; second glume as long as the spikelet, strongly 7-nerved. Sterile lemma 5-7 mm. long, hirsutulous above, Sparsely so below, about 5-nerved. Fertile lemma indurate, rugose, abruptly acute and subrostrate, SANTA CRUZ: CERCADO: Palmar, Herzog 1524 (vs). 30. P. megiston Schult. Mant. 2: 248 (1824). Stout perennial, creeping at the base, the culms to 1-2 m. tall. Leaf-sheaths sparsely papillate-pilose, the blades lanceolate, to 40 cm. long and 3 em. wide, long-acute, glabrous above and beneath. Panicle to 6 dm. long, spread- ing, the primary branches in distant verticils (usually opposite or alternate at the uppermost node), axis hispid at the nodes, otherwise glabrous, branches and pedicels scabridulous on the margins, Spike- lets glabrous, to 4 mm. long, broadly ovoid, abruptly acute, usually solitary or in groups of 2 along the branches. Glumes very unequal; first glume about 1.5 mm. long, abruptly subacute, strongly 3-nerved, broader than long and basally completely clasping; second glume about 3.5 mm. long, abruptly acute, 9-11-nerved. Sterile lemma about 3 mm. long, subobtuse, 7-nerved. Fertile lemma about 3 mm. long, abruptly acute, indurate, shining. LA PAZ: CAUPOLICÁN: Ixiamas, White (M. E.) 1135 (Hitekcock). 1966] Flora of Bolivia — Foster 329 31. P. olyroides HBK. Nov. Gen. & Sp. 1: 102 (1816). Erect peren- nial, the culms to 10 dm. tall. Leaf-sheaths markedly imbricate, glabrous, the linear firm blades to 30 cm. long and 8 mm. wide, glabrous above and beneath. Panicle open, to 3 dm. long, the axis, branches and pedicels densely scabrid-ciliate, the few spikelets on long slender pedicels near the branch-ends. Spikelets 6-8 mm. long, glabrous, fusiform, long-acute. Glumes unequal; first glume to 6 mm. long, long-acute, 5-7-nerved; second glume to 8 mm. long, long-acute, about 9-nerved, with 2 internal basal tufts of subhispid hairs. Sterile lemma to 3.5 mm. long, acute, the nerves prominent near the apex only. Fertile lemma to 3.5 mm. long, obtuse, shining, the base with 2 lateral patches of short, white, crispate hairs. SANTA CRUZ: SARA: Buenavista, 450-500 m., Steinbach 5378 (GH), 6978 (GH). 32. P. procurrens Nees ex Trin. Gram. Pan. 183 (1826). Decumbent perennial, spreading, the culms to 6 dm, tall, usually shorter. Leaf- sheaths glabrous with an adaxial apical collar of hairs, blades lance- linear, to 5-6 cm. long and 4-5 mm. wide, acute, white-margined, glabrous to puberulent, often scabridulous beneath. Panicle to 15 cm. long, axis pubescent, the erect branches usually mostly glabrous, the spikelets often somewhat approximate. Spikelets about 3.5-4 mm. long, acute to subacuminate, long-hirsute. Glumes unequal; first glume 2.5 mm. long, strongly 3-nerved, long-acute; second glume 3-3.5 mm. long, 5-nerved, subacute to subobtuse. Sterile lemma as long as second glume, 5-nerved, acute. Fertile lemma shorter than sterile lemma, smooth, shining, chartaceous. SANTA CRUZ: SARA: Dolores, 450 m., Steinbach 1801 (GH); Buenavista, 450-500 m., Steinbach 5339 (GH), 6940 bis in part (GH), 7012 (GH). 83. Ichnanthus Beauv. Perennials, erect, decumbent or creeping. Leaves pubescent or glabrous. Inflorescence a panicle. Spikelets pedicellate, disarticu- lating below the glumes, 2-flowered, the lower floret a sterile lemma or staminate, the upper floret perfect. Glumes 2, the lemma shortly stipitate, the stipe with a pair of membranous appendages basally adnate to the base of the fertile lemma, free above, or the appendages wanting and represented by a pair of small pits or scars. a. Lemma-appendages present. b. Lemma-appendages suborbicular, 0.4 mm. long; spikelets 3 mm. long, obtuse. ....... eene 1. I. calvescens. b. Lemma-appendages narrowly deltoid, 0.6 mm. long; spikelets to 5 mm. long, acute. ........... nennen enn . 2. I. peruviana. a. Lemma-appendages wanting. c. Leaf-sheath auricles well-developed, to 4 mm. long. «seers Eu Ot a sisi Lcawaks lobes iuba vo cET Pr E 3. I. breviscrobs. c. Leaf-sheath auricles not developed. d. First glume shorter than second. 330 Rhodora [Vol. 68 e. Inflorescences terminal and axillary, usually less than 10 em. long. f. Spikelets glabrous, or if with a few hairs, these not papillate-based. l. 4. I. pallens. f. Spikelets sparsely pilose, the hairs papillate-based. ........ IDE 5. I. candicans. e. Inflorescences terminal only, to 20 cm. long. sesser ln RJ 6. I. Ruprechtii. d. First glume longer than second. cecccccccccccss--..... 7. I. minarum. 1. Ichnanthus calvescens (Nees) Doell in Mart. Fl. Bras. 2(2): 285 (1877). Decumbent, the culms to 1 m. tall. Leaf-sheaths pubescent to villous, or glabrous; blades long, lance-elliptic, long-acute, 1-2 cm. wide, densely pubescent above and beneath. Inflorescence to 2 dm. long, narrow, the branches ascending, rachis, branches and pedicels scabridulous, the rachis and branches villous-hispid at the nodes. Spikelets about 3 mm. long, obtuse. First glume about 2.25 mm. long, acute, 3-nerved, the keel scabridulous; second glume about 3 mm. long, 5-nerved. Sterile lemma 3 mm. long. Fertile lemma about 2 mm. long, the appendages suborbicular, 0.4 mm. long. LA PAZ: NOR YUN- GAS: Milluguaya, Buchtien 737 (Hitchcock); SUR YUNGAS: Chulu- mani, Hitchcock 22653 (Hitchcock). 2. I. peruviana Mez in Fedde, Rep. Sp. Nov. 15: 129 (1918). Spread- ing, to 1 m. tall. Leaf-sheaths pubescent to villous; blades lanceolate, to 10 cm long and 1.5 cm. wide, pubescent above and beneath, or scabridulous above. Panicle to 2 dm. long, open, the few branches short, distant, few-flowered, rachis, branches and pedicels scabridulous on the margins. Spikelets to 5 mm. long, acute. First glume to 5 mm. long, acute, puberulent at the apex, 3-nerved, the keel scabridulous; second glume 4 mm. long, obtuse, puberulous at the apex, 5-nerved, the keel scabridulous above. Sterile lemma about 4 mm. long, obtuse, puberulent, somewhat shortly penicillate, at the apex, rather obscurely 5-nerved. Fertile lemma about 2.5-2.75 mm, long, shining, the apex incurved as a short thick beak, the appendages narrowly deltoid, subobtuse, 0.6 mm. long. LA PAZ: NOR YUNGAS: Coroico, 1560 m, Hitchcock 22715 (GĦ). 3. I. breviscrobs Doell in Mart. Fl. Bras. 2(2): 294 (1877). Erect, stout, to 3-5 m. tall. Leaf-sheaths glabrous, with a pair of well- developed auricles to 4 mm. long at the apex; blades long, lanceolate, to 4 em. wide, scabridulous above and beneath. Panicle open, to 3 dm. long, the branches ascending, rachis, branches and pedicels puberulent. Spikelets 4 mm. long, obtuse. First glume 2.5 mm. long, obtuse, pu- berulent, 3-nerved, the nerves scabridulous-ciliate; second glume 4 mm. long, obtuse, puberulent, 7-nerved. Sterile lemma 4 mm. long, puberu- lent above. Fertile lemma 3 mm. long, the apex incurved as a short, thick, blunt beak, appendages wanting, the pits small. LA Paz: LARECAJA: San Carlos, Mapiri, 750 m., Buchtien 60 (GH). 1966] Flora of Bolivia — Foster 331 4. I. pallens (Swartz) Munro ex Benth. Fl. Hongk. 414 (1861). Creeping, the culms to 3 dm. long. Leaf-sheaths papillate-pilose, auricles not developed; blades ovate to lanceolate, to 10 em. long, 1-2 cm. wide, asymmetric, glabrous to papillate-pilose above and beneath. Inflorescences terminal and axillary, to 6 cm. long, the branches ascending-spreading, rachis and branches somewhat pilose, the pedicels scabridulous-puberulent. Spikelets to 4 mm. long, acute, glabrous or often with a few long hairs near the nerves and margin, the hairs not papillate-based. First glume 3 mm. long, long-acuminate, 3-nerved; second glume to 4 mm. long, acute, 5-nerved. Sterile lemma 3 mm. long, acute, 5-nerved. Fertile lemma about 2 mm. long, shining, obscurely keeled, the subacute apex only slightly incurved, appendages wanting. LA PAZ: LARECAJA: Hacienda Simaco, 1400 m., Buchtien 5322 (GH); SUR YUNGAS: Sirupaya, 1650 m., Buchtien (in Baenitz) 1401 (GH). COCHABAMBA: CHAPARE: Antahuacana, Espíritu Santo, 750 m., Buchtien 153 (GH). SANTA CRUZ: SARA: bosques del Fraile, 450 m., Steinbach 2258 (GH). 5. I. candicans (Nees) Doell in Mart. Fl. Bras. 2(2): 291 (1877). Creeping plant, much-branched. Leaf-sheaths papillate-villous, the auricles not developed; blades papillate-villous above and beneath, ovate, to 5 em. long and 1.5 em. wide. Inflorescences terminal and axillary, to 7-8 cm. long, branches ascending, rachis, branches and pedicels pubescent to scabridulous-puberulent. Spikelets to 4 mm. long, acute, the glumes sparsely papillate-pilose. First glume about 2.5 mm. long, long-acuminate, 3-nerved; second glume about 4 mm. long, shortly acuminate. Sterile lemma about 3 mm. long, obtuse, 5-nerved. Fertile lemma about 2 mm. long, shining, obscurely 3-nerved, very obtuse, the apex not incurved as a beak, appendages wanting. LA PAZ: LARECAJA: San Carlos, Mapiri, 850 m., Buchtien 6 (GH). The line of demarcation between this species and I. pallens seems tenuous and rather difficult to draw. 6. I. Ruprechtii Doell in Mart. Fl. Bras. 2(2): 293 (1877). Spread- ing, to 1-2 m. tall Leaf-sheaths mostly glabrous, auricles not de- veloped; blades lanceolate, to 15 cm. long and 1 cm. wide, glabrous to sparsely pubescent above and beneath. Inflorescences terminal, to 2 dm. long, rachis, branches and pedicels pubescent to pilose, especially at and just below the nodes. Spikelets about 3-4 mm. long. First glume to 3 mm. long, subobtuse, 3-nerved, the keel scabrid-ciliate; second glume 3.5 mm. long, subobtuse. Sterile lemma 3 mm. long, subobtuse. Fertile lemma about 2.5 mm. long, shining, slightly in- curved at the subacute apex, appendages wanting. LA PAZ: LARECAJA: Hacienda Simaco, 1400 m., Buchtien 5336 (GH). 7. I. minarum (Nees) Doell in Mart. Fl. Bras. 2(2): 294 (1877). Straggling, sprawling plant. Leaf-sheaths glabrous, auricles not de- veloped; blades lanceolate, to 12 cm. long and 2 em. wide, glabrous or sparsely pubescent above and beneath, the hairs often papillate- 332 Rhodora [Vol. 68 based. Inflorescence a terminal panicle of a few unbranched racemes, the panicle to 10 em. long, rachis, branches and pedicels puberulent- scabridulous. Spikelets dense on the raceme-axes, to 4.5 mm. long. First elume 4.5 mm. long, long-acute, scabridulous below, the keel scabridulous, a few long hairs near the apical margins; second glume 3.5 mm. long, similar to the first glume in pilosity. Sterile lemma 3 mm. long, glabrous, 5-nerved, subobtuse. Fertile lemma shining, about 2.5 mm. long, subacute, appendages wanting. LA PAZ: LARE- CAJA: Cerro del Iminapi, 2650 m., Mandon 1256 (GH); SUR YUNGAS: La Florida, 1840 m., Hitchcock 22619 (GH), 84. Lasiacis (Griseb.) Hitchc. Perennials, sprawling or clambering, to 10 m., much-branched, woody-stemmed. Leaves flat, rather firm, lanceolate to more or less elliptie, subpetiolate. Inflorescence a rather open panicle; spikelets obliquely placed on the pedicels, 2-flowered, the sterile lemma below the fertile, glumes markedly unequal, lanate at the apex; second glume and sterile lemma similar in appearance; fertile lemma and palea shining, indurate to bony, their apices briefly tufted with short woolly hairs in a slight crateriform depression; disarticulation below the glumes. Ligule prominent, brown, to 2 mm. long; leaf-blades glabrous above. ieteeeeaeeeeeceeaeseessessecesaeeeessesescsseesseeessstsesststteaessatetsusceeccssecs 2. L. sorghoidea. l. Lasiacis ligulata Hitche. & Chase in Contrib. U. S. Nat. Herb. 18: 337 (1917). Plants to 5-10 m. long. Leaf-blades narrowly lanceo- late, to 12 cm. long, somewhat pubescent to puberulous beneath. Panicle to 10 em. long, rachis, branches and pedicels puberulent to scabridulous. Spikelets ovoid to obovoid, to 5 mm. long. First glume to 2.5 mm. long, many-nerved; second glume as long as the Spikelet, many-nerved, the central nerve more prominent than the others. Sterile lemma as long as second glume, more shining. Fertile lemma slightly shorter than sterile lemma, longitudinally striatulate but not rugose. Glumes and sterile lemma black at maturity. La PAZ: SUR YUNGAS: Colaya, 1760 m., Mexia 4307 (GH). COCHABAMBA: CHAPARE: Locotal, 1600 m., Steinbach 9091 (GH); Incachaca, 2400 m., Stein- bach 9091 (GH). 2. L. sorghoidea (Desv.) Hitche. & Chase in Contrib. U. S. Nat. Herb. 18: 338 (1917). Plants to 5-7 m. long. Leaf-blades lanceolate to somewhat elliptic, to 2.5 em. long; sheaths sparsely to densely pubes- cent or villous, blades pubescent to papillate-hispid above and beneath. Panicle to 20 em. long and nearly as broad at maturity, axis pubescent to sparsely hispid, branches and pedicels puberulent to scabridulous. Spikelets obovoid to subglobose, to 3.5-4 mm. long. First glume 1.5-2 1966] Flora of Bolivia — Foster 333 mm. long, not conspicuously nerved, the central nerve prominent; second glume as long as the spikelet, many-nerved. Sterile lemma as long as second glume. Fertile lemma slightly shorter than sterile lemma, obtuse, prominently striatulate. LA PAZ: NOR YUNGAS: Milluguaya, 1300 m., Buchtien 157 (GH); SUR YUNGAS: Guanay, 630 m., Rusby 191 (GH). COCHABAMBA: CHAPARE: Antahuacana, Espíritu Santo, 750 m., Buchtien (in 1909) 161 (GH); CERCADO: Cochabamba, Bang 1289 (GH). SANTA CRUZ: SARA: 400-450 m., Steinbach 1931 (GH), 5551 (GH), 7111 (GH). 85. Sacciolepis Nash Sacciolepis myuros (Lam.) Chase in Proc. Biol. Soc. Wash. 21: 7 (1908). Annual, to 1 m. high, in wet or marshy habitats. Leaf- blades flat or folded, long, linear, to 5 mm. wide, usually glabrous, but occasionally sparsely pilose above and beneath. Inflorescence a terminal, racemose, compact, narrow panicle, spicate in appearance, generally 10 cm. or less in length, occasionally to 20 cm., the rachis, branchlets and pedicels more or less densely puberulent. Spikelets about 2 mm. long, pointed, disarticulation below the glumes, 2-flowered, the lower floret a sterile lemma, occasionally a staminate floret, the upper floret perfect. Glumes unequal, the first to 1.5 mm. long, 3-nerved, the second inflated or somewhat saccate, to 2 mm. long, 7-9-nerved, often puberulous near the apex. Sterile lemma as long as the second glume and similar in appearance; fertile lemma shorter than the sterile and concealed by it, hard-textured, shining, some- times sparsely and minutely puberulent, the margins not inrolled but revealing the hard shining palea, this shorter than but similar to the fertile lemma, the fruit minutely stipitate. SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 7030 (GH), 7101 (GH). 86. Hymenachne Beauv. Hymenachne donacifolia (Raddi) Chase in Journ. Wash. Acad. Sci. 13: 177 (1923). Perennial, to 1 m. high, in wet or swampy habitats. Leaf-sheaths coarsely ciliate, the glabrous blades broadly lanceolate, to 20 em. long and 3 cm. wide, the bases cordate-clasping or even somewhat auriculate. Inflorescence a terminal, racemose panicle to 3 dm. long, somewhat open, the branches mostly ascending, the rachis, branches and pedicels finely and densely puberulent. Spikelets clus- tered, acuminate, to 3 mm. long, 2-flowered, the lower floret a sterile lemma, the upper floret perfect; disarticulation below the glumes. Glumes very unequal, the first about 1 mm. long, acute, faintly 3-nerved, the keel ciliolate on the upper portion; second glume to 2 mm. long, acuminate, strongly 3-nerved, neither saccate nor inflated. Sterile lemma about 2.5 mm. long, acuminate, similar to the second glume; fertile lemma shorter than the sterile, without apparent 334 Rhodora [Vol. 68 nerves, smooth, rather shiny, minutely puberulent near the apex, not firm and indurate. Palea as long as the fertile lemma, similar in appearance and texture. SANTA CRUZ: SARA: Curichis, monte de Palometillas, 400 m., Steinbach 2150 (GH). 87. Homolepis Chase Homolepis aturensis (HBK). Chase in Proc. Biol. Soc. Wash. 24: 146 (1911). A decumbent to sprawling perennial, culms to 5 dm. high. Leaf-sheaths densely ciliate and sparsely pilose, the narrowly lanceo- late blades flat, to 1.5 em. wide, often sparsely pilose when young and glabrescent with age, the margins scabridulous. Inflorescence a terminal, racemose, rather narrow panicle to 10 em. long, the rachis, branches and pedicels glabrous. Spikelets to 7 mm. long. somewhat fusiform, 2-flowered, the upper floret perfect, the lower sterile or sometimes staminate, disarticulation below the glumes. Glumes equal or subequal, to 7 mm. long, the first glume 9-nerved, glabrous; second glume slightly shorter, fewer-nerved, the margins appressed-villous. Sterile lemma shorter than the glumes, enclosing the fertile lemma, densely appressed-villous along the margins; fertile lemma shorter than the sterile, glabrous, shining, firm-textured but not indurate, partially enclosing the similar but shorter palea. SANTA CRUZ: SARA: Buenavista, 450-500 m., Steinbach 1844 (GH), 5120 (GH), 7013 (GH). 88. Isachne R. Br. Isachne arundinacea (Swartz) Griseb. Fl. Brit. W. Ind. 553 (1864). Climbing perennial, the branches up to 6 m. long. Leaf-blades flat, lanceolate, to 20 cm. long and 1.5-2 em. wide, scabrid above and beneath. Inflorescence a terminal racemose panicle to 12 cm. long, the branches ultimately spreading, the rachis and branches puberulent- scabridulous, the pedicels usually glabrous. Spikelets subglobose, about 2 m. long, disarticulation below the glumes, 2-flowered, the upper floret perfect, the lower floret staminate or perfect. Glumes nearly equal, often suborbieular, about 1.5 mm. long, with a tuft of a few short hairs below the apex. Staminate lemma and palea equal in length, about 2 mm. long, indurate, shining; fertile floret slightly smaller; both lemmas with a few short basal hairs. YUNGAS: Bang 297 (GH). 89. Oplismenus Beauv. Oplismenus hirtellus (L.) Beauv. Ess. Agrost. 54, 168 (1812). Spreading, much-branched perennial, the culms 3-5 dm. high. Leaf- sheaths glabrous to densely hirsute, the lanceolate or lance-ovate blades to 10 cm. long, glabrous above, sometimes scabridulous beneath, the margins undulate. Inflorescence a terminal few-branched panicle 1966] Flora of Bolivia — Foster 335 of 3-6 short (1-3 em.) densely flowered racemes, the spikelets paired or solitary, unilateral, very shortly pedicellate, the rachis scabridulous to pubescent or coarsely ciliate. Spikelets 2-flowered, the lower floret a sterile lemma, the upper floret perfect, disarticulation below the glumes. Glumes subequal, about 2.5-3 mm. long, 3-5-nerved, variably pubescent to nearly glabrous, awn of first glume to 6 mm. long, awn of second glume about 2 mm. long. Sterile lemma about 3 mm. long, about 7-nerved, variably pubescent, especially on the nerves, shortly awned or mucronate. Fertile lemma to 2.5 mm. long, indurate, shining, very faintly nerved, the flat margins clasping the palea. LA Paz: LARECAJA: Hacienda Simaco, 1400 m., Buchtien 5331 (GH); NOR YUN- GAs: Milluguaya, 1300 m., Buchtien 738 (GH); Coripata, Bang 2079 (GH). SANTA CRUZ: SARA: Buenavista, 450-500 m., Steinbach 1067 (GH), 1859 (GH), 5681 (GH), 1010 (GH); Dolores, 450 m., Steinbach 2036 (GH). 90. Echinochloa Beauv. Echinochloa crus-pavonis (HBK.) Schult. Mant. 2: 269 (1824). Annual, erect to decumbent at base, the culms to 1 m. tall. Leaf- sheaths glabrous; blades linear, to 1.2 cm. wide, glabrous, the margins scabridulous. Inflorescence a terminal panicle of racemes, the spikelets solitary or unilaterally clustered on the racemes, the rachis hispid at the nodes, branches and pedicels scabrid. Spikelets about 3 mm. long, plano-convex, 2-flowered, the lower floret sterile or staminate, the upper floret perfect. First glume about 1.5 mm. long, broadly ovate, acute, few-nerved scabrid to hispidulous; second glume 3 mm. long, awned, few-nerved, the nerves strongly short-hispid. Sterile lemma 3 mm. long, few-nerved, the nerves short-hispid, the awn to 4 mm. long. Fertile lemma indurate, about 2.5 mm. long, with a terminal awn or mucro 0.5 mm. long, shining; palea similar but narrower and slightly shorter. LA PAZ: LARECAJA: vic. Sorata, Mandon 1258 (GH); Hacienda Simaco, 1400 m., Buchtien 5313 (GH); NOR YUNGAS: Cori- pata, Bang 2108 (GH); Polo-Polo, near Coroico, 1100 m., Buchtien 445 (GH). 91. Setaria Beauv. See Rominger, J. M. Taxonomy of Setaria (Gramineae) in North America. Illinois Biological Monographs, no. 29. 1962. Annvals or perennials, erect to decumbent and spreading. Leaf- sheaths and blades glabrous to varyingly pubescent. Inflorescence a condensed panicle, the branches seldom obvious, mostly spicate in appearance. Spikelets mostly plano-convex (very obscurely so in 1 species), subtended by 1 to several antrorsely or retrorsely scabrous bristles, 2-flowered, the lower floret a sterile lemma (or occasionally a staminate floret), the upper floret perfect; disarticulation below the 336 Rhodora [Vol. 68 glumes. Glumes unequal; fertile lemma coriaceous to indurate, smooth in 2 species, or rugose or rugulose. a. Bristles antrorsely scabrous. b. Fertile lemma neither rugose nor rugulose. c. Leaves plicate; spikelet 3-4 mm. long; bristles 1-2 cm. long. LM 1. S. Poiretiana. c. Leaves not plicate, spikelet 2-2.5 mm. long; bristles 5-6 mm. [oT RM 6. S. oblongata. b. Fertile lemma transversely rugose or rugulose. d. Nodes pubescent to villous, v.ccccccccccscssscccosececesses. 9. S. barbinodis. d. Nodes glabrous. e. Bristles below spikelet 5 or more. ............ 2. S. geniculata. e. Bristles below spikelet 1, or 2.3 by abortion of spikelets. f. Leaf-base rounded, not petiolate nor subpetiolate, ........... IM 8. S. argentina. f. Leaf-base gradually tapered, petiolate or subpetiolate. £. Main panicle about 10 cm. long; bristles to about 1 em. long. |t 10. S. trichorhachis. g. Main panicle about 20-30 em. long; bristles to about 2-8 em. long. ... sss T. S. vulpiseta. a. Bristles retrorsely scabrous, at least on upper half. h. Spikelets 2.5-3 mm. long. a 3. S. tenax. h. Spikelets 1.5 mm. long. i. Bristles 2-5 mm. long. «se 4. S. scandens. i. Bristles 1-1.5 em. long. «Lese 5. S. tenacissima. l. Setaria Poiretiana (Schult. in Roem. & Schult.) Kunth, Rév. Gram. 1: 47 (1829). Ferennial, the culms to 1-2 m. tall, sometimes villous near the nodes, but the nodes glabrous. Leaf-sheaths papillate- hirsute; leaf-blades plicate, to 5 cm. wide, often pubescent or pu- berulous below, scabridulcus above and beneath, the base tapering and at least subpetiolate. Panicle to 6 dm. long, the branches ascend- ing, the rachis, branches and pedicels puberulent. Spikelets acute, 3-4 mm. long, secund on the branches, slightly plano-convex, subtended by 1 bristle 1-2 cm. long. First glume 1.5 mm. long, obtuse, 3-nerved; second glume about 2 mm. long, obtuse, 5-nerved. Sterile lemma as long as the spikelet, acute, 2-nerved. Fertile lemma slightly shorter than the sterile, acute to somewhat acuminate, not rugose but promi- nently longitudinally striatulate. La Paz: LARECAJA: San Carlos, Mapiri, 850 m., Buchtien 18 (GH); NOR YUNGAS: Coroico, 1560 m., Hitchcock 22719 (GH). COCHABAMBA: CHAPARE: Locotal, 1500 m., Steinbach 9019 (GH). SANTA CRUZ: SARA: Cerro del Amboró, 1000 m., Steinbach 2998 (Gn). 2. S. geniculata (Lam.) Beauv. Ess. Agrost. 51, 178 (1812). S. gracilis, S. gracilis f. brevispica and S. gracilis f. penicillata of the Catalogue. Perennial, to 12 dm. tall, the culms geniculate, nodes glabrous. Leaf-sheaths glabrous; blades glabrous beneath, scabridu- 1966] Flora of Bolivia — Foster 337 lous above, to 8 mm. wide. Panicle to 10 em. long but usually shorter, the branches extremely short, rachis, branches and pedicels densely short-pubescent or hispid. Spikelets 2-2.5 mm. long, strongly plano- convex when mature, subtended by 5 or more bristles, these to 1 cm. long on early-season plants or 2-3 mm. long on late-season plants. First glume about 1 mm. long, obtuse, 3-nerved; second glume about 1.5 mm. long, abruptly acute at apex, about 5-nerved. Sterile lemma as long as the spikelet, about 7-nerved, subacute, enclosing a well- developed palea and often a staminate floret. Fertile lemma as long as the sterile, markedly transversely rugose, the palea-apex free at maturity. YUNGAS: Bang 218 (GH), 273 (GH). LA PAZ: LARECAJA: vic. Sorata, Mandon 1259 (GH); SUR YUNGAS: La Florida, 1840 m., Hitchcock 22634 (GH); Sirupaya, 2200 m., Buchtien 7029 (GH). COCHAMBAMBA: CERCADO: Cochabamba, 3300 m., Shepard 259 (GH), 2500 m., Eyerdam 24666 (GH); Mollemolle, 2800 m., Steinbach 4057 (GH). SANTA CRUZ: SARA: Buenavista, 500 m., Steinbach 6199 (GH). 3. S. tenax (L. Rich.) Desv. Opusc. 78 (1831). Perennial, to 1-2 m. tall, the nodes glabrous. Leaf-sheaths glabrous to puberulent, the margins papillate-ciliate; blades to 2 cm. wide, abruptly rounded at the base but not petiolate, scabridulous to pubescent. Panicle to 3 dm. long, the rachis short-villous, the short branches and pedicels pu- berulous to scabridulous. Spikelets 2.5-3 mm. long, plano-convex to suborbicular, each subtended by a single bristle 1-2 cm. long, this antrorsely scabrous at the base and retrorsely scabrid above. First glume about 1 mm. long, broadly ovate, subacute, 5-nerved; second glume 1.5-2 mm. loag, subobtuse, about 7-nerved. Sterile lemma as long as the spikelet, usually with a well-developed palea. Fertile lemma as long as the sterile, smoothish at the apex, strongly trans- versely rugose below. LA PAZ: NOR YUNGAS: Coripata, Hitchcock 22685 (GH). 4. S. scandens Schrad. ex Schult. in Roem. & Schult. Syst. Veg. Mant. 2: 279 (1824). Annual, to 8 dm. tall, the culms geniculate at the base, the nodes glabrous. Leaf-sheaths glabrous, the margins ciliate; blades to 1 em. wide, abruptly narrowed at the base, scabrous- pubescent above and beneath. Panicle to 10 cm. long, the branches very short, rachis short-pubescent with scattered, long, hispid hairs, branches and pedicels puberulous-scabridulous. Spikelets about 1.5 mm. long, plano-convex, subtended by 1-3 bristles retrorsely barbed on the upper portion, the bristles 2-5 mm. long. First glume about 0.75 mm. long, broadly ovate, subacute, 3-nerved; second glume nearly 1.5 mm. long, 5-nerved, subobtuse. Sterile lemma 5-nerved, as long as the spikelet with a palea half as long. Fertile lemma transversely rugose or rugulose, as long as the sterile lemma. LA PAZ: LARECAJA: Hacienda Simaco, 1400 m., Buchtien 5330 (GH); NOR YUNGAS: Polo- Polo, near Coroico, 1100 m., Buchtien 449 (GH). 5. S. tenacissima Schrad. ex Schult. in Roem. & Schult. Syst. Veg. 338 Rhodora [Vol. 68 Mant. 2: 279 (1824). Annual, to 2 m. tall, nodes glabrous. Leaf- sheaths glabrous below, scabrous above; blades to 1 em. wide, pubes- cent and scabrous above and beneath. Panicle to 2 dm. long, the branches short (less than 1 cm.), rachis pubescent to hispid, branches and pedicels pubescent, Spikelets plano-convex, 1.5 mm. long, each subtended by a retrorsely scabrous bristle 1-1.5 em. long. First glume about 1 mm. long, 3-nerved, subobtuse; second glume nearly 1.5 mm. long, 5-nerved. Sterile lemma 1.5 mm. long, 3-5-nerved. Fertile lemma as long as the sterile or slightly shorter, rugulose on the lower portion, LA PAZ: LARECAJA: Hacienda Simaco, 1400 m., Buchtien 5329 (GH); NOR YUNGAS: Coroico, 1560 m., Hitchcock 22717 (GH). 6. S. oblongata (Griseb). Parodi in Physis, 9: 88 (1928), Perennial, to 1.5 m. tall, nodes glabrous. Leaf-sheaths glabrous below, scabrid above; blades long, to 1 em. wide, scabrous above and beneath. Panicle to 1 dm. long, the branches very short, rather lax, rachis short-villous, branchlets and pedicels puberulent. Spikelets 2-2.5 mm. long, plano-convex, each subtended by a single antrorsely scabrous bristle 5-6 mm. long. First glume 1.5 mm. long, subobtuse, 5-nerved; second glume about 2 mm. long, 7-nerved. Sterile lemma as long as the spikelet, 5-nerved. Fertile lemma as long as the sterile, smooth, shining. LA PAZ: LARECAJA: vic. Sorata, Mandon 1260 (Hitchcock). T. S. vulpiseta (Lam.) Roem. & Schult. Syst. Veg. 2: 495 (1817). Perennial, culms to 2 m. tall, nodes glabrous. Leaf-sheaths some- what carinate, glabrous below, scabrous to pubescent above; blades basally petiolate or subpetiolate, to 3 cm. wide, scabrous above and beneath. Panicle to 3 dm. long, the ascending branches to 3-4 (occa- sionally nearly 5) em. long, rachis and branches villous, pedicels puberulent. Spikelets to 2.5 mm. long, plano-convex, each subtended by 1-2 antrorsely scabrous bristles 9.3 em. long. First glume about 1 mm. long, 3-5-nerved; second glume about 2 mm. long, 7-nerved. Sterile lemma as long as the spikelet, 5-nerved, with a well-developed palea. Fertile lemma 2.5 mm. long, smooth near the apex, markedly rugose below. SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 1178 (GH); Rio Surutt, 400 m., Steinbach 6842 (GH). 8. S. argentina Herrm. in Beitr. Biol. Pflanz. 10: 54 (1910). Peren- nial, to 10 dm. tall, the nodes glabrous. Leaf-sheaths glabrous below, sparsely papillate-hirsute above, long-ciliate; blades abruptly rounded at base, to 1 cm. wide, papillate-pubescent above and beneath. Panicle to 12 em. long, the branches very short, the rachis, branches and pedicels puberulent. Spikelets plano-convex, to 2.5 mm. long, each subtended by 1 antrorsely scabrous bristle (occasionally 2) 8-10 mm. long. First glume nearly 1.5 mm. long, subacute, 3-nerved; second glume about 2 mm. long, acute, 5-nerved. Sterile lemma 2.5 mm. long, acute. Fertile lemma nearly as long as the sterile, smooth above, rugulose below. LA PAZ: MURILLO: Cotana, 2450 m., Buchtien 286 (GH); LARECAJA: vic. Sorata, Mandon 1262 (GH). COCHABAMBA: 1966] Flora of Bolivia — Foster 339 CERCADO: near Cochabamba, Hitchcock 22803 (GH); Valle de Cocha- bamba, 2700 m., Steinbach 9714 (GH). 9. &. barbinodis Herrm. in Beitr. Biol. Pflanz. 10: 60 (1910). Peren- nial, to 10 dm. high, the nodes pubescent to villous. Leaf-blades linear, long, to 7 mm. wide, scabrid above and beneath, with occasional hairs but hardly pubescent. Panicle to 15 cm. long, interrupted below, the branches short, rachis short-pubescent with few to many long hairs intermixed, the branches and pedicels short-pubescent. Spikelets sub- globose, to 3 mm. long, each subtended by 1 antrorsely scabrous bristle to 1 em. long. First glume about 2 mm. long, acute, 3-nerved; second glume 2.5 mm. long, broad, shortly apiculate, 5-7-nerved. Sterile lemma 3 mm. long, shortly apiculate, 5-nerved. Fertile lemma slightly shorter than the sterile lemma, rugulose to rugose. LA PAZ: LARE- CAJA: vic. Sorata, Mandon 1261 (US; type-number). 10. S. trichorhachis (Hack.) R. C. Foster, comb. nov. S. Liebmannii f. trichorhachis Hack. in Fedde, Rep. Nov. Spec. 8: 46 (1910). Peren- nial, to 8 dm. tall, the nodes glabrous. Leaf-sheaths glabrous below, sparsely pubescent above, ciliate; blades gradually tapered and sub- petiolate at base, to 1.5 cm. wide, scabrous above and beneath. Panicle to 8 em. long, the branches very short, rachis short-pubescent with numerous longer hairs intermixed, branches and pedicels pu- berulent. Spikelets plano-convex, 3 mm. long, each subtended by 1 antrorsely scabrous bristle about 1 em. long. First glume about 1 mm. long, subacute, 5-nerved; second glume 2 mm. long, obtuse, 7-nerved. Sterile lemma about 2.5 mm. long, 5-nerved, acute. Fertile lemma 2.5 mm. long, rugose below, rugulose to smooth above. Without locality: Bang 2160 (GH). 92. Pennisetum L. Rich. Perennials, to 5 m. tall. Leaf-blades flat, sparsely pubescent to glabrous and scabridulous, petiolate to subpetiolate. Inflorescence more or less spicate in appearance, the spikelets solitary to few in a fascicle, sessile or subsessile, subtended by a ring of equal to unequal, basally plumose or antrorsely puberulent bristles, 2-flowered, the lower floret a sterile lemma or sometimes a staminate floret, the upper floret perfect; disarticulation below the glumes. Glumes unequal. Fertile lemma chartaceous to herbaceous, mostly smooth. a. Culm -nodes villous. ........... ee 4. P. latifolium. a. Nodes glabrous. b. Bristles plumose. c. Longest bristles 1.5 cm. long; spikelets 7 mm. longo c ees NNNM -c2£. 1. P. setosum. c. Longest bristles 4 cm. long; spikelets to 12 em long s 2. P. villosum. a9604000060000040606000000050000000000000090000000000000090000 0009008 8 980 0 PR b. Bristles not plumose. 340 Rhodora [Vol. 68 d. Longest bristles about as long as the spikelet (4-5 mm.) ........ ————— RR 5. P. mutilatum. d. Longest bristles much longer than the spikelet (about 1.2 cm. long). e. Second glume 2-3 mm. long; spikelet to 7 mm. long. ....... ——— HÁÉRRRRRRRRRRRRRERRRRRRREERRREA 6. P. chilense. e. Second glume less than 1 mm. long; spikelet to 5 mm. long. /———— ÉERERRRERRRRRREMM 3. P. tristachyum. 1. Pennisetum setosum (Swartz) L. Rich. in Pers. Synops. Pl. 1: 72 (1805). Culms 1-2 m. tall, nodes glabrous. Leaf-sheaths glabrous; blades to 4 dm. long and 1.8 em. wide, pubescent or scabridulous above and beneath. Inflorescence to 25 cm. long, purplish. Spikelets about 4 mm. long. Bristles in 2 series; outer series shorter to slightly longer than the spikelet, antrorsely puberulent; inner series about twice the length of the spikelet (1 bristle much longer than the rest, to 1.5 em. long), densely silky-plumose basally. First glume about 0.5 mm. long, sometimes absent; second glume 3.5 mm. long, longer than the sterile lemma, acute, puberulous on the upper portion. Sterile lemma about 3 mm. iong, obtuse, puberulous, the apex finely ciliolate. Fertile lemma about 2.5 mm. long, the apex penicillate. LA Paz: NOR YUNGAS: Coripata, Bang 2168 (GH). 2. P. villosum R. Br. in Fresen. in Mus. Sencken. 2: 134 (1837). Culms to 0.5 m. tall, nodes glabrous. Leaf-sheaths glabrous; blades 3-5 mm. wide, scabridulous beneath, occasionally sparsely pilose above. Inflorescence to 8 em. long, sometimes pale-tawny. Spikelets to 1.2 cm. long, 1-4 in a shortly pedunculate fascicle, the peduncle densely short-villous at the base. Bristles to 4 (-5) em. long, the outer shorter and sparsely or not at all plumose, the inner long and plumose. First glume about 0.5 mm. long; second glume about 3.5 mm. long, long- acute. Sterile lemma about 1 cm. long, long-acute, scabridulous on the nerves above the middle. Fertile lemma 1 mm. shorter than the sterile, acuminate. CHUQUISACA: OROPEZA: outskirts of Sucre, 2700 m., Cardenas 4116 (GH). 3. P. chilense (Desv.) Jacks. ex R, E. Fries in Nov. Act. Soc. Sci. Upsal. (ser. 4) 1: 172 (1905). Culms 1-2 m. tall, from a distinct rhizome, the nodes glabrous. Leaf-sheaths glabrous; blades scabrous beneath, sometimes scabridulous above, to 1 em. wide. Inflorescence to 2 dm. long, grayish to purplish. Spikelets to 7 mm. long. Bristles of varying lengths, from the length of the spikelet to 1.5 cm. long, antrorsely puberulent, none plumose. First glume nearly 1 mm. long, obtuse; second glume about 2-3 mm. long, subacute, sometimes ciliolate. Sterile lemma 6 mm. long, scabridulous on the upper portion, long-acute. Fertile lenma 6 mm. long, long-acute, L4 PAZ: MURILLO: La Paz, 3700 m., Buchtien 81 (GH), 161 (GH), Bang 55 (GH), Mandon 1265 (GH); Palca, 3700 m., Hitchcock 22583 (GH). COCHABAMBA: CHAPARE: Sacaba, 2900 m., Steinbach 8809 (GH). Potosi: SUR 1966] Flora of Bolivia — Foster 341 CHICHAS: Oploca, Hitchcock 22898 (GH). TARIJA: CERCADO: Con- cepción, 1900 m., Fiebrig 2595 (GH). 4. P. latifolium Spreng. Syst. Veg. 1: 302 (1825). Culms to 1-3.5 dm. tall, the nodes villous. Leaf-sheaths glabrous to pilose on the upper portion, or papillate-hirsute, the margins hispid-ciliate; blades to 35 em. long and 5 em. wide, usually narrower, scabridulous above and beneath. Inflorescence to 10 cm. long, greenish. Spikelets about 4.5 mm. long, acute. Bristles few, not plumose, shorter than or as long as the spikelet, 1 bristle to 1 cm. long. First glume 0.5 mm. long, subacute; second glume about 1 mm. long, ovate, subacute, ciliolate at the apex. Sterile glume as long as the spikelet, acute, scabrid- puberulous. Fertile lemma slightly shorter than the sterile lemma, subacuminate, slightly scabridulous at the apex. LA PAZ: LARECAJA: vie. Sorata; Luemapampa, Iminapi, 2600 m., Mandon 1263 (GH). COCHABAMBA: CERCADO: west of Cochabamba, Hitchcock 22847 (GH); CHAPARE: Locotal, 1500 m., Steinbach 9014 (GH). 5. P. mutilatum (O. Ktze.) Hack. ex Index Kew. Suppl. 2: 137 (1904). P. sagittatum Henr. in Blumea, Suppl. 1: 229 (1937). Culms 1-2 m. tall, nodes glabrous. Leaf-sheaths sparsely papillate-hirsute to glabrous; blades glabrous or puberulent at base when young, to 1.5 cm. wide. Inflorescence to 10 cm. long. Spikelets about 4 mm. long. Bristles few, short, equaling or slightly exceeding the spikelet, not plumose. First glume between 0.5-1 mm. long, obtuse; second glume 2.3 mm. long, acute, puberulous above. Sterile lemma nearly as long as the spikelet, acute, antrorsely puberulent on entire dorsal surface. Fertile lemma to 4 mm. long, occasionally scabridulous near the acute apex. La PAZ: LARECAJA: vic. Sorata; between Munaypata and Challasuyo, 2650 m., Mandon 1264 (GH). 6. P. tristachyum (HBK.) Spreng. Syst. Veg. 1: 302 (1825). P. tristachyum subsp. boliviense Chase in Contrib. U. S. Nat. Herb. 24: 486 (1927). Culms 2-6 m. tall, branched, nodes glabrous. Leaf-sheaths glabrous; blades to 4 dm. long and 3 or more em. wide, petiolate, the petioles villous and the blades sparsely villous above and beneath at the base. Inflorescence to 10 cm. long. Spikelets about 5 mm. long, acute. Bristles unequal in length, the longest to 1.5 em. long, not plumose. First glume about 0.25 mm. long; second glume less than 1 mm. long. Sterile lemma as long as the spikelet, acuminate, scabridu- lous toward the apex. Fertile lemma slightly shorter than the sterile lemma, acuminate, glabrous or nearly so. LA PAZ: NOR YUNGAS: Coroico, 1100 m., Buchtien 451 (Gu); Colapampa, Bang 2340 (GH). SANTA CRUZ: SARA: Cerro Hosana, 1300 m., Steinbach 3379 (GH); Cerro La Negra, 1809 m., Steinbach 8172 (GH). 93. Cenchrus L. Annual or perennial, often to 1 m. tall. Inflorescence a terminal raceme. Spikelets 1-4 in a group, surrounded by an involucre (modi- 342 Rhodora [Vol. 68 fied sterile branchlets) forming spiny burs, the spines usually retrorsely barbed; disarticulation below the burs. Spikelets 2-flowered, the lower floret a sterile lemma, the upper floret perfect. Glumes unequal, the first glume sometimes obsolete. a. Burs with 1 spikelet; involucral lobes united at the base only. MM 1. C. myosuroides. a. Burs with more than 1 spikelet; involucral lobes united to about the middle. b. Burs with a basal ring of bristles. co.cc... 4. C. pauciflorus. c. Involucral lobes interlocking; burs about 4 mm. wide. ............ uM 2. C. viridis. c. Involucral lobes not, or seldom, interlocking; burs 5-7 mm. Wide. MM 9. C. echinatus. b. Burs without a basal ring of bristles. ................ 4. C. pauciflorus. 1. Cenchrus myosuroides HBK. Nov. Gen. & Sp. 1: 115 (1816). Perennial, rather woody, to 1.5 m. tall, glaucous, glabrous. Leaf-blades 5-12 mm. wide. Raceme to 25 em. long, dense; burs 1-flowered, to 5 mm. wide, the lobes united at the base only, free above, the base hispidulous, the outer bristles of the involucre shorter than the inner, both series erect, the inner about equaling the spikelet. Spikelet about 5 mm. long, acute. First glume about 2 mm. long, broadly ovate, acute, l-nerved; second glume 3.5-4 mm. long, 5-nerved, the midrib prominent. Sterile lemma 4.5 mm. long, similar to second glume. Fertile lemma 5 mm. long, acute; palea almost as long, nar- rower, COCHABAMBA: TARATA: Anzaldo, 3000 m., Cárdenas 2425 (GH); CERCADO: Cochabamba, Hitchcock 22806 (GH); Cochabamba Railway Station, 2560 m., Cárdenas 2391 (GH); Taquifia, 2700 m., Steinbach 9801 (an). 2. C. viridis Spreng. Syst. Veg. 1: 301 (1825). Annual, up to 1 m. tall, glabrous. Leaf-blades to 12 mm, wide. Raceme to 10 cm. long, rather dense; burs usually 3-flowered, subglobose, 4 mm. wide (ex- cluding the bristles), lobes interlocked at maturity, the base pubescent. Spikelet about 7 mm. long, acute. First glume narrowly ovate, 3 mm. long, 1-nerved; second glume 5 mm. long, acute, 5-nerved. Sterile lemma to 6 mm. long, acute, several-nerved. Fertile lemma slightly longer. LA PAZ: SUR YUNGAS: Guanay, Rusby (M. E.) 190 (Hitch- cock). 3. C. echinatus L. Sp. Pl. 1050 (1753). Annual, to 6 dm. high. Leaf-blades to 8 mm. wide, sparsely pilose above near the base. Raceme to 10 cm. long; burs usually 4-flowered, lobes united to the middle, seldom interlocking at maturity, the tips spinose, 5-7 mm. wide (excluding the bristles), pubescent. Spikelet about 7 mm. long. First glume ovate, 2 mm. long, 1-nerved, acute; second glume 4 mm. long, 5-nerved. Sterile lemma 5 mm. long. Fertile lemma 5 mm. long, long-acute. LA PAZ: NOR YUNGAS: Milluguaya, 1300 m., Buchtien 1966] Flora of Bolivia — Foster 343 4171 (GH). SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 6980 bis (GH): Río Surutu, 400 m., Steinbach 5576 (GH), 6838 (GH). 4. C. pauciflorus Benth. Bot. Voy. Sulphur, 56 (1840). Annual, often in mats, to 9 dm. tall. Leaf-blades to 7 mm. wide. Raceme to 8 cm. long; burs usually 2-flowered, 4-6 mm. wide, pubescent, the lobes united to the middle. Spikelet about 6 mm. long. First glume about 2 mm. long, acute, 1-nerved; second glume 4 mm. long, 3-nerved. Sterile and fertile lemmas as long as the spikelet, several-nerved. LA PAZ: LARECAJA: vic. Sorata; San Pedro, 2650 m., Mandon 1267 (GH). 94. Olyra L. Perennials, often very tall, often rather woody, plants monoecious. Leaf-blades variable in size, often very large, mostly glabrous, petio- late or subpetiolate. Inflorescence a compact to lax panicle; pistillate spikelets at ends of branches, the staminate spikelets below, these pedicellate, smaller, or the upper branches all with pistillate spike- lets and the lower branches all staminate. Pistillate spikelets: rather large, 2-flowered, the lower floret a sterile lemma, the first glume absent, the fruit indurate or bony; disarticulation below the glumes. Staminate spikelets: both glumes and the sterile lemma absent, the fertile lemma and palea membranaceous. a. Leaf-blades oblong- or triangular-deltoid. b. Panicle short, 2-2 cm. long; glume 2-3 mm. long, not caudate; fruit glabrous. ........... eene nennen 1. O. lateralis. b. Panicle to 10 cm. long; glume to 2 cm. long, caudate; fruit pubescent. .....eeeseeeeeeeeeennnnn nen enne nennen ennt 5. O. ciliatifolia. a. Leaf-blades not deltoid. c. Fruit pitted. d. Glume to 1 cm. long, densely pubescent externally: sete Jr idee du oe A ca iie eeddeetueasese mee LENIN 6. O. micrantha. d. Glume to 3 cm. long; externally glabrous. .... 3. O. heliconia. c. Fruit not pitted. e. Glume 2-4 em. long, caudate. f. Glume about 2 cm. long. ............. mA 7. O. latifolia. f. Glume about 4 em. long. ..............:... sas inis 4. O. Buchtienit. e. Glume 1 cm. long, not caudate. ............ 2. O. ecaudata. 1. Olyra lateralis (Presl) Chase in Proc. Biol. Soc. Wash. 21: 179 (1908). Straggling. Leaf-blades oblong-deltoid, to 4 cm. long and 1.2 em. wide, the base truncate to subcordate, glaucous beneath, pube- rulent above when young. Panicle 2-3 em. long, few-branched, the branches usually whorled. Pistillate spikelet: ovoid, 2-3 mm. long, glabrous; glume with 3 prominent nerves, sterile lemma similar; fruit shining, not pitted, glabrous. Staminate spikelet: linear, acute, 2.5-3 mm. long, puberulent at base. LA PAZ: LARECAJA: San Carlos, Mapiri, 850 m., Buchtien 25 (GH) ; Hacienda Simaco, 1400 m., Buchtien 5333 (GH). 344 Rhodora [Vol. 68 2. O. ecaudata Doell in Mart. Fl. Bras. 2(2): 326 (1877). Erect. Leaf-blades oblong, to 20 em. long and 2.5 cm. wide, scabrid above. Pistillate spikelet: 6-8 mm. long, oblong, acuminate, not caudate. Staminate spikelet: subulate, the lemma cuspidate. PANDO: COBIJA: Ule 9146 (Hitchcock). 3. O. heliconia Lindm. in Sv. Vet. Akad. Handl. 34(6): 11 (1900). Stout, little-branched, to 2 m. tall. Leaf-blades oblong, to 35 em. long and 8 em. wide, the subtruncate base asymmetric, sparsely puberu- lent when young. Panicle to 30 cm. long, the few branches in 2-3 distant whorls. Pistillate spikelet: glume to 3 em. long, long-caudate, sterile lemma to 2 cm. long, long-acute to subcaudate, both puberulent within and glabrous externally; fruit to 1 cm. long, glabrous but conspicuously and regularly pitted. Staminate spikelet: about 1 cm. long, narrowly subulate, often purple. LA PAz: NOR YUNGAS: Coroico, 1560 m., Hitchcock 22720 (GH); Polo-Polo, 1100 m., Buchtien 450 (GH). SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 2010 (GH), 4. O. Buchtienii Hack. in Fedde, Rep. Sp. Nov. 11: 20 (1912). To 1.5 m. tall. Leaf-blades oblong-lanceolate, to 30 em. long and 7 em. wide, the base subtruncate, somewhat asymmetric. Panicle to 15 cm. long, few-branched. Pistillate spikelet: glume and sterile lemma long-caudate, glume to 4 cm. long; glume and lemma puberulent within, scabridulous-puberulent externally; fruit 1 cm. long, glabrous. Staminate spikelet: narrowly ellipsoid, to 4 mm. long, sparsely pu- berulent. L4 PAZ: LARECAJA: San Carlos, Mapiri, 800 m., Buchtien. 40 (GH). 5. O. ciliatifolia Raddi, Agrost. Bras. 19 (1823). Seldom 1 m. tall. Leaf-blades triangular-deltoid, to 10 em. long and 4 em. wide, the truncate base asymmetric, glabrous, or pubescent at base when young. Panicle to 10 cm. long, at least the lower branches in whorls. Pistillate spikelet: glume 2 em. long, very slender-caudate, the sterile lemma similar but shorter, glume and lemma glabrous internally and ex- ternally; fruit to 7 mm. long, pubescent. Staminate spikelet: to 7 mm. long, very slender, glabrous. BENI: YACUMA: Lago Rogagua, Rusby (M. E.) 1669 (Us). 6. O. micrantha HBK. Nov. Gen. & Sp. 1: 199 (1816). Very large, to 5 m. tall. Leaf-blades oblong-lanceolate, to 30 em. long and 10 cm. wide, the base rounded but not cordate, glabrous. Panicle to 30 cm. long. Pistillate spikelet: glume to 1 em. long, long-caudate, densely pubescent; sterile lemma similar but much shorter; glume and lemma minutely puberulent internally; fruit 3 mm. long, glabrous but regularly and deeply pitted. Staminate spikelet: very narrow, 7 mm. long, the lemma somewhat caudate, glabrous. SANTA CRUZ: ICHILO: Río Vibora, 350 m., Steinbach 7572 (GH). LA PAZ: LARE- CAJA: San Carlos, Mapiri, 750 m., Buchtien 76 (GH). 7. O. latifolia L. Syst. Nat. (ed. 10) 2: 1261 (1759). To 5 m. tall. 1966] Flora of Bolivia — Foster 345 Leaf-blades lanceolate-oblong, to 30 em. long and 8 cm. wide, glabrous, acuminate, the base asymmetric. Panicle to 15 cm. long. Pistillate spikelet: glume to 2 cm. long, long-caudate, minutely puberulent externally, hispid-ciliate near the base, internally densely puberulent above the middle; sterile lemma similar but shorter; fruit 5 mm. long, shining, apparently somewhat rugose above the middle, glabrous. Staminate spikelet: slender, to 7 mm. long, the lemma short-caudate. La PAZ: LARECAJA: Mapiri, Buchtien 1155 (Hitchcock). 95. Imperata Cyrillo Caespitose, rhizomatous perennials, the rhizome scaly, the culms unbranched, to 1-2 m. tall. Leaf-sheaths often ciliate, otherwise glabrous; leaf-blades flat or involute, glabrous beneath, pubescent to hirsute above (especially near the base) when young. Inflorescence a panicle of racemes. Spikelets 2-flowered, the lower floret a sterile lemma, the upper floret perfect, pedicellate, awnless; disarticulation below the glumes. Glumes mostly subequal, membranaceous to hyaline, more or less densely villous, the hairs much exceeding the spikelet. Sterile lemma and fertile lemma and palea hyaline, thin. a. Leaf-blades flat. b. Spikeiet 4 MM: Jong. ura cence ctae ira 1. I. brasiliensis. b. Spikelet 2 mm. Jong: cere inima 3. I. minutiflora. a. Leaf-blades involute (the uppermost sometimes flat). ..................... EERS P E D BR EPI II Em 2. I. tenuis. 1. Imperata brasiliensis Trin. in Mém. Acad. St. Pétersb. (ser. 6) Math. Phys. Nat. 2: 331 (1832). To 1 m. tall. Leaf-blades flat, to 1.8 cm. wide, mostly glabrous, the margins scabrid. Panicle about 10 em. long. Spikelets 4 mm. long, glumes acute, the first glume somewhat shorter than the second. BENI: YACUMA: Reyes, White (M. E.) 1535 (Hitchcock). 2. I. tenuis Hack. in DC. Monogr. Phan. 6: 689 (1889). Plants 0.8-2 m. tall. Leaves mostly involute, the uppermost occasionally flat. Panicle to 2 dm. long. Spikelets 4 mm. long, glumes long-acute, the first glume shorter than the second. SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 1824 (GH), 6928 (GH); Río Dolores, 450 m., Stein- bach 1933 (GH). 3. I. minutiflora Hack. in DC. Monogr. Phan. 6: 100 (1889). Plant 0.4-1 m. tall. Leaf-blades to 4 mm. wide. Panicle to 2-3 dm. long. Spikelets 2 mm. long, glumes acute, hairs mostly basal. SANTA CRUZ: SARA: Curichi, 400 m., Steinbach 2410 (GH); Río Guendá, 400 m., Steinbach 6894 (GH). 96. Saccharum L. Saccharum officinarum L. Sp. Pl. 54 (1753). Very large perennial, to 5 mm.. tall, the culms 2-3 em. thick. Leaf-sheaths closely imbricate, 346 Rhodora [Vol. 68 the lowermost falling from the culms; blades to 6 cm, wide, midrib very prominent. Inflorescence a plumose panicle to 6 dm. long, the racemes drooping, the rachis disarticulating. Spikelets about 3-4 mm. long, awnless, paired, both with a perfect floret, one sessile, a ring of long hairs at base of each spikelet, 2-flowered. Glumes subequal, acute, subindurate. COCHABAMBA: CHAPARE: Espíritu Santo, Buch- tien 2514 (Hitchcock). 97. Eriochrysis Beauv. Erect perennials to 1-2 m. tall. Leaf-blades flat to involute, glab- rous to sparsely or densely pubescent. Inflorescence a terminal continuous or interrupted panicle of racemes, the raceme-rachis ulti- mately disarticulating below the spikelets. Spikelets paired, perfect spikelets sessile, pistillate spikelets pedicellate, both long-villous at the base. Glumes subequal, rather indurate; lemma somewhat hyaline; palea hyaline. Leaf-blades flat, densely velvety-pubescent; panicle dense, almost un- interrupted, deep golden-brown. ............ esee l. E. cayennensis. Leaf-blades usually involute, glabrous or sparsely pubescent; panicle obviously interrupted, pale to medium-brown, ............ e MMMNNMMMMMMMMMMMNM 2. E. Warmingiana. 1. Eriochrysis cayennensis Beauv. Ess. Agrost. 8 (1812). Un- branched plant, 1-2 m. tall. Panicle to 12 cm. long, the individual racemes very short. Spikelets about 3 mm. long. First glume obtuse, long-villous on upper portion and along margins; second glume sub- acute, convex, sparsely long-villous on upper keel. SANTA CRUZ: SARA: Portachuelo, Steinbach 1974 (Gn); Buenavista, 450-500 m., Steinbach 5174 (GH), 6915 (au). Occasionally, an intermediate with a more interrupted inflorescence, but almost the typical coloring, will be found, such as Steinbach 6975 bis (GH), from Buenavista. 2. E. Warmingiana (Hack.) Kuhlm. in Comm. Linhas Telegr. Estrat. Matto Grosso, 67: 29 (1922). Caespitose, about 1 m. tall, the nodes pubescent. Panicle about 15 cm. long, the racemes 2-4 cm. long. Spikelets to 5 mm. long. First glume long-acute, the upper portion short-puberulent, with a few long hairs, otherwise glabrous; second glume convex, puberulent, with a few long lax hairs on the upper margin. SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 1916 (GH), 7032 (GH). 98. Erianthus Beauv. Perennials, 1-3 m. tall, the nodes pubescent to long-villous, Leaf- sheaths sparsely to densely pubescent; leaf-blades flat, pubescent or scabrous. Inflorescence a terminal, rather dense, panicle of racemes. Spikelets perfect, paired, 1 sessile, 1 pedicellate, the pedicel villous 1966] Flora of Bolivia — Foster 347 in part, the raceme-rachis disarticulating below the spikelets. Glumes equal or subequal, silky-villous, at least at the base, sterile lemma hyaline, fertile lemma hyaline and long-awned; palea small and hyaline. Leaf-blades to 1.5 em. wide; panicle light-colored, 3-4 dm. long. ........ Waa Sete aR ound ta Ta tales E AAA DERE ENSIS 1. E. Trini. Leaf-blades less than 1 cm. wide; panicle usually purplish, 1.5-2.5 dm. IDEE al Sie ee huis nS I CIUS 2. E. angustifolius. 1. Erianthus Trinii (Hack.) in DC. Monogr. Phan. 6: 185 (1889). Plants 2-3 m. tall, leaf-blades to 1.5 cm. wide. Panicle to 3-4 dm. long, the base usually enclosed in the inflated upper leaf-sheath, the culms pubescent below the panicle. Spikelets to 7 mm. long (excluding the awn). Glumes acute to slightly bidentate, the upper portion soft- pubescent. Awn of fertile lemma about 1 cm. long, not geniculate. La Paz: NOR YUNGAS: Coripata, Hitchcock 22682 (Hitchcock). 2. E. angustifolius Nees, Agrost. Bras. 316 (1829). Plants about 1 m. tall, leaf-blades 4-8 mm. wide. Panicle 1.5-2.5 dm. long, the base mostly free from the leaf-sheath, culm densely pubescent below panicle. Spikelets to 8 mm. long (excluding the awn). Glumes biden- tate, short-pubescent on upper portion. Awn of fertile lemma about 1 em. long, not geniculate. LA PAZ: NOR YUNGAS: San José, 1500 m., Hitchcock 22737 (GH). 99. Andropogon L. Perennials, to 1-2 m. tall. Inflorescence a panicle of racemes, soli- tary or 2 or more at apices of branches, the common peduncle usually enclosed by a sheathing spathe. Spikelets 2-flowered, with the lower floret a sterile hyaline lemma, paired on a jointed rachis, 1 spikelet ses- sile and perfect, the other pedicellate, staminate or neuter, or greatly reduced; disarticulation below the glumes. Glumes of perfect spikelet subequal or somewhat unequal, coriaceous, the first glume bicarinate, the margins inflexed. Fertile lemma hyaline, awned or awnless. Pedicellate spikelet sometimes with the first glumes aristate, some- times reduced to a pedicel, the pedicels often conspicuously villous. a. Spikelets awnless. b. Rachis-joints clavate, with a cupular hollow at the summit. ........ AUR osse avi TE N ced ts cant neha tanisaad E L E dag 2. A. virgatus. b. Rachis-joints not clavate, the summit without a cupular hollow. c. First glume of sessile spikelet glabrous. d. Culms 1-2 m. tall; leaf-blades long-attenuate, not boat- shaped at the tip. eem 9. A. bicornis. d. Culms 1 m. tall or less; leaf-blades not long-attenuate, boat- shaped at the tip. .......... ee 11. A. Selloanus. c. First glume of sessile spikelet not glabrous. e. Glume pilose on and near keels. ...............- 6. A. Hassleri. e. Glume merely ciliolate on keels. ........ 10. A. leucostachyus. 348 Rhodora [Vol. 68 a. Spikelets awned, f. Rachis-joints clavate, with a cupular hollow at the summit. g. First glume of sessile spikelet glabrous. h. Racemes spreading, rachis flexuose. ........ 1. A. condensatus. h. Racemes appressed, rachis straight. i. Sessile spikelet 4(-5) mm. long. ................ 4. A. tener. i. Sessile spikelet 5-6 mm. long. .............. 5. A. cirratus. g. First glume of sessile spikelet not glabrous. j. Sessile spikelet 9-10 mm. long; glume of sterile spikelet long-aristate. .........eseseeeeeeee eene 3. A. hirtiflorus. j. Sessile spikelet 5 mm. long; glume of sterile spikelet not arlstate. ........seseeeeeeeeeeee nennen nennen 4. A. tener hirtiglumis. f. Rachis-joints not clavate, not with a cupular hollow at the summit. k. First glume of sessile spikelet pilose on keels. ............... ne m——— HÁÉK T. A. saccharoides. k. First glume of sessile spikelet glabrous. l. Glume with a dorsal pit. seen 8. A. altus. l. Glume not pitted. m. Leaf-blades glabrous. n. Blades less than 10 mm. wide at base, base not cordate. sasbsdeheenuass tessensnesvorsassnntssonserensceanecenrersenutsvaltieeere 12. A. lateralis. n. Blades 15-20 mm. wide at base, base cordate. ................ Sosvsveseessdapsevuusssaseesssessuvuvenesveceeiessepisesserssaeeseyers 13. A. cordatus. 1. Andropogon condensatus HBK. Nov. Gen. & Sp. 1: 188 (1816). Culms to 1 m. or more high. Leaves glabrous, the blades flat, linear, acute to subobtuse. Inflorescence much-branched, corymbose, the ra- chises flexuose, the rachis-joints clavate, cupular at the apex, uni- laterally long-pilose. Sessile spikelets 4.5-5 mm. long, the glumes subequal, the first glume strongly bicarinate, the keels scabrid on upper portion; awn slender, flexuose, to 1 cm. long. Pedicellate spike- lets reduced, the pedicels curved and unilaterally long-pilose. YUNGAS: Bang 276 (GH). LA Paz: NOR YUNGAS: Milluguaya, 1300 m., Buch- tien 38 (GH); Polo-Polo, near Coroico, 1100 m., Buchtien 205 (GH); SUR YUNGAS: La Florida, 1840 m., Hitchcock 22639 (GH). SANTA CRUZ: SARA: Buenavista, 400-500 m., Steinbach 3302 (GH), 5165 (GH), 6809 (GH), 6951 (GH), 6952 (GH), 6953 (GH). 2. A. virgatus Desv. ex Hamilt. Prodr. Pl. Ind. Occ. 9 (1825). Culms to 2 m. high. Leaf-blades long, narrowly linear, glabrous above, pubescent beneath (glabrate in old leaves ?). Inflorescence elongate, much-branched, the rachis straight, spikelets densely appressed and ascending, rachis-joints shortly ciliate or ciliolate on the 2 margins, clavate with a cupular apex, racemes 1-2 cm. long, somewhat enclosed in the subtending red-brown or purplish spathes. Sessile spikelets 1966] Flora of Bolivia — Foster 349 about 2.5-3 mm. long, the first glume strongly bicarinate, the keels scabrid-ciliolate, otherwise glabrous; lemma awnless. Pedicellate spikelets not reduced, about 2.5 mm. long, the glumes ciliolate on the keels, pedicels puberulent. SANTA CRUZ: SARA: Buenavista, 450-500 m., Steinbach 5344 (GH), 6945 (GH). 3. A. hirtiflorus (Nees) Kunth, Rév. Gram. 2: 569 (1832). A. Riedelii of the Catalogue. Culms to 1 m. tall, reddish. Leaves flat, linear, glabrous. Inflorescence much-branched, the rachis straight, racemes and spikelets ascending, rachis-joints pubescent, clavate, with a cupular apex, racemes 6-10 cm. long. Sessile spikelets to 9-10 mm. long, the first glume villous, aristate; lemma awned, the awn twisted, about 1-1.5 em. long. Fedicellate spikes reduced, aristate, the pedicels pubescent. La PAZ: LARECAJA: vic. Sorata; between Toquirani and San Pedro, 2700 m., Mandon 1385 (GH); between Choquecoa and Mt. Illampu, 3600 m., Mandon 1275 (GH); SUR YUNGAS: Chulumani, 1600 m., Hitchcock 22658 (GH). 4. A. tener (Nees) Kunth, Rév. Gram. 2: 565 (1832). Densely tufted, the culms to 1 m. tall. Leaf-blades flat or involute, sometimes sparsely long-pilose above. Inflorescence very slender, the rachis straight, rachis-joints clavate, ciliate around the cupular apex and on 1 side, or nearly glabrous. Sessile spikelets 4-5 mm. long, the first glume glabrous or ciliolate on the keels; awn to 1 cm. long. Pedicellate spikelet reduced. shortly aristate. LA PAZ: LARECAJA: Sorata, Mandon 1383 (GH); Catarguata, 2700 m., Mandon 1381 (GH). TARIJA: ARCE: Camacho, 2500 m., Fiebrig 2859 (GH). 4a. A. tener subvar. hirtiglumis Henr. in Meded. Rijks Herb. Leiden, no. 40: 42 (1921). Resembling A. tener in most respects, but with the first glume of the sessile spikelet densely pilose. TARIJA: AVILES: Pinos, 2300 m., Fiebrig 3154 (GH; type-number). 5. A. cirratus Hack. in Flora, 68: 119 (1885). Culms to 7 dm. tall, tufted. Leaf-blades flat, linear, scabrid. Inflorescence narrow, the rachis straight, racemes appressed-ascending, rachis-joints uni- laterally long-ciliate, clavate, with a cupular apex. Sessile spikelets to 5-6 mm. long, the first glume glabrous; awn to 1 cm. long. Pedi- cellate spikelets somewhat reduced, the first glume long-aristate, pedi- cels long-ciliate. TARIJA: CERCADO: Tucumilla, 2200 m., Fiebrig 2787 (GH). 6. A. Hassleri Hack. in Bull. Herb. Boiss. (ser. 2) 4: 266 (1904). Culms about 1 m. tall, the nodes glabrous. Linear leaf-blades scabridu- lous beneath. Inflorescence oblong, to 12 em. long, pale, silky-villous, racemes appressed-ascending, rachis-joints long-silky-villous, especially on the margins. Sessile spikelets about 3 mm. long, pilose, especially on and near the rather rounded keels; awnless. Pedicellate spikelets greatly reduced, the aborted glume short-pubescent near the apex, pedicels densely long-villous. LA PAZ: SUR YUNGAS: Chulumani, 1600 m., Hitchcock 22659 (GH). 350 Rhodora [Vol. 68 7. A. saccharoides Swartz, Prodr. 26 (1788). Culms tufted, to 1 m. tall, nodes hispid. Leaf-blades glabrous (occasionally some pubes- cence present), glaucous. Inflorescence to 15 cm. long, exserted, mostly silvery-white, silky-villous, the rachis-joints long-villous. Ses- sile spikelet about 5 mm. long, the first glume pilose on the keels; awn twisted below, 1(-1.5) em. long. Pedicellate spikelets somewhat reduced, the pedicels long-villous. La Paz: MURILLO: Obrajes, 3400 m., Buchtien 574 (Git). COCHABAMBA: CERCADO: Cochabamba, Hitch- cock 22794 (GH); Quillacolla: Liriuni, 3000 m., Steinbach 9847 (GH). Ta. A. saccharoides subsp. laguroides (DC.) Hack. in DC. Monogr. Phan. 6: 495 (1889). A. ternatus var. macrothrix of the Catalogue. Culms shorter, nodes glabrous, and panicle smaller than in A. sac- charoides, COCHABAMBA: CERCADO: Cochabamba, Hitchcock 22823 (Hitchcock). Tb. A. saccharoides subsp. parvispiculus Hitchc. in Contrib. U. S. Nat. Herb. 24: 497 (1927). A. saccharoides var. barbinodis of the Catalogue. Panicle longer, gray or purplish, sessile spikelets 3-4 mm. long. Leaf-blades often pubescent. LA PAZ: LARECAJA: Sorata, 2600- 2700 m., Mandon 1389 (GH); MURILLO: La Paz, 3300 m., Bang 102 (GH); La Paz, 3500 m., Buchtien 8846 (GH); SUR YUNGAS: La Florida, 1840 m., Hitehcock 22615 (GH). COCHABAMBA: MIZQUE: Vila- vila, 2500 m., Eyerdam 24975 (au). "c. A. saccharoides subvar. perforatus (Trin.) Hack. in DC. Monogr. Phan. 6: 496 (1889). First glume of sessile spikelet with a suprame- dial dorsal pit. Otherwise like A. saccharoides. COCHABAMBA: CER- CADO: Cochabamba, Holway 325 (Hitchcock). 8. A. altus Hitchc. in Contrib. U. S. Nat. Herb. 17: 208 (1913). Culms mostly solitary, the nodes bearded, to 1-2 m. high. Leaf-blades flat, linear, scabrid, with a few long hairs at base of upper surface. Panicle oblong, to 2 dm. long, greenish-white, rachis-joints densely long-villous. Sessile spikelet to 5 mm. long, the glabrous first glume with a supramedial dorsal pit; awn to 2 em. long. Pedicellate spikelet greatly reduced, the pedicel long-villous. LA PAZ: NOR YUNGAS: Milluguaya, Buchtien 4258 (Hitchcock). 9. A. bicornis L. Sp. Pl. 1046 (1753). Culms to 1-2 m. tall, the nodes glabrous. Leaf-blades flat, linear, long-attenuate, scabrid. In- florescence a corymbose, feathery, villous panicle, rachis-joints long- villous. Sessile spikelets about 3 mm. long, the first glume glabrous. Pedicellate spikelets greatly reduced, the pedicels long-villous, LA PAZ: LARECAJA: Hacienda Simaco, 1400 m., Buchtien 5312 (GH); NOR YUNGAS: Polo-Polo, near Coroico, 1100 m., Buchtien (in 1912) 204 (GH); Milluguaya, 1300 m., Buchtien (in 1917) 204 (GH). SANTA CRUZ: SARA: Buenavista, 450-500 m., Steinbach 5274 (GH), 6949 (GH). 10. A. leucostachyus HBK. Nov. Gen. & Sp. 1: 187 (1816). Densely tufted, the culms usually less than 1 m. tall, the nodes glabrous. 1966] Flora of Bolivia — Foster 351 Leaf-blades linear, flat, glabrous. Inflorescence a rather short villous panicle, the rachis-joints long-villous. Sessile spikelet awnless, about 3 mm. long, the first glume ciliolate on the keels and sometimes puberulent on the upper portion. Pedicellate spikelets greatly reduced, the pedicels long-villous. LA PAZ: NOR YUNGAS: Milluguaya, 1200 m., Buchtien 735 (GH). SANTA CRUZ: SARA: Dolores, 450 m., Stein- bach 1888 (GH); Buenavista, 450 m., Steinbach 6517 (GH), 6845 (GH), 6846 (GH), 6848 (GH). 11. A. Selloanus (Hack.) Hack. in Bull. Herb. Boiss. (ser. 2) 4: 266 (1904). Culms mostly less than 1 m. tall, nodes glabrous. In- florescence a densely villous panicle, the rachis joints densely long- villous on the upper portion. Sessile spikelets awnless, to 4 mm. long, the first glume glabrous. Pedicellate spikelets greatly reduced, the pedicels densely long-villous. SANTA CRUZ: SARA: Buenavista, 450- 500 m., Steinbach 1928 (GH), 5143 (GH), 6515 (GH), 6516 (GH), 6723 (GH), 6874 (GH), 6971 bis (GH). 12. A. lateralis Nees, Agrost. Bras. 329 (1829). Culms to 1 m. tall, often purple, the nodes glabrous. Leaf-blades flat, linear, the base not cordate, to 6 mm. wide, glabrous. Inflorescence of 2-4 racemes to 4 em. long, the rachis-joints long-villous. Sessile spikelet to 4-5 mm. long, the first glume glabrous; awn to 1 cm. long. Pedicellate spikelet not reduced, about 4 mm. long, the pedicel long-villous. SANTA CRUZ: SARA: Rio Quimory, 400 m., Steinbach 7328 (GH); Buenavista, 450- 500 m., Steinbach 5427 (GH), 6851 (GH). 13. A. cordatus Swallen in Contrib. U. S. Nat. Herb. 29: 275 (1948). Culms to nearly 2 m. tall. Leaf-blades glabrous, linear- attenuate, the base 15-20 mm. wide and cordate, Inflorescence to 5 dm. long, racemes to 4.5 cm. long, the rachis-joints villous. Sessile spikelet to 4 mm. long, the first glume glabrous; awn to 1.5 em. long. Pedicel- late spikelets to 6.5 mm. long, the pedicel long-villous. COCHABAMBA: TOTORA: Chimoré, 1000 m., Cárdenas 2083 (US; type). 14. A. leptocladus Hack. in Flora, 68: 122 (1885). Culms to 4 dm. high, the nodes short-hispid. Leaf-blades bristly-pilose at base, pu- bescent on both surfaces (at least when young), linear, long-attenuate. Inflorescence exserted, of few racemes, the rachis-joints long-hispid. Sessile spikelet about 6 mm. long, the first glume glabrous or with a few short hairs near the base; awn strongly geniculate, to 4 cm. long. Pedicellate spikelet to 5-6 mm, long, the short pedicel bilaterally ciliate. SANTA CRUZ: SARA: Rio Guenda, 400 m., Steinbach 7939 (GH). 100. Hyparrhenia Anderss. Hyparrhenia bracteata (Humb. & Bonpl. ex Willd.) Stapf in Prain, Fl. Trop. Afr. 9: 360 (1918). Perennial, to 1 m. high, or more. Leaf- blades long, linear, narrow (to 4 mm.), appressed-pilose beneath. Inflorescence terminal, to 3 dm. or more, a complex panicle of paired 352 Rhodora [Vol. 68 racemes, these subtended by narrow pilose bracts, 1-2.5 em. long, the ultimate racemes few-flowered. Spikelets 1-flowered, paired, 1 sessile, fertile, and awned, the other (sometimes 2) pedicellate, the pedicel long-villous, sterile and awnless; disartieulation below the glumes. Fertile spikelet: about 6 mm. long (excluding the awn), the glumes somewhat indurate; first glume with involute margins, bidentate, a few longish hairs near the apex, second glume similar but not biden- tate; fertile lemma thin, delicate, shorter than the glumes, the sub- plumose awn to 2.5 em. long, loosely twisted, Sterile spikelet: shorter than the fertile, not bidentate, the apices pubescent and subaristate. LA PAZ: SUR YUNGAS: Chulumani, Hitchcock 22704 (Hitchcock). 101. Sorghum Moench Sorghum vulgare Pers. Synops. Pl. 1: 101 (1805). Robust annual, the culms to 1.5 m. high. Leaf-blades long, 2-6 cm. wide, glabrous. Inflorescence a terminal panicle, often condensed, of several few- jointed racemes, these tardily disarticulating. Spikelets paired, the perfect spikelet sessile, the staminate or sterile spikelet pedicellate. Perfect spikelet: glumes subequal, about 5 mm. long, broadly ovate, the first glume few-nerved, more or less shortly silky-villous, the second glume indurate, shining; lemma shorter than the glumes, awned, the awn geniculate, basally twisted, to 1 cm. long, quickly deciduous. Sterile spikelet: pedicel villous, glumes 3-4 mm. long, few-nerved, very sparsely villous. LA PAZ: NOR YUNGAS: Coroico, Buchtien 3615 (Hitchcock). The perennial S. halepense (L.) Pers. Synops. Pl. 1: 101 (1805) may also be found. It has an extensive, creeping, scaly rhizome and, in general, is somewhat more robust than is S. vulgare. 102. Sorghastrum Nash Perennials, to 1 m. high, the culm-nodes pubescent to shortly villous, the leaf-sheaths rather prominently auriculate, the blades glabrous to scabrid. Inflorescence a panicle of short racemes, the rachis dis- articulating. Spikelets paired, the perfect spikelet sessile or sub- sessile, the sterile spikelet reduced to a pubescent or villous pedicel. Glumes coriaceous, the fertile lemma hyaline, awned. a. Awn to 8 em. long. wi.ccccccecccssscsssessscesseeceescecsecsecscessecess 1. S. minarum. a. Awn much less than 8 cm. long. b. Awn less than 1 cm. long, straight or once geniculate, not tightly twisted at base. sesser 2. S. parviflorum. b. Awn to nearly 2 cm. long, twice geniculate, tightly twisted at DD 3. S. stipoides. 1. Sorghastrum minarum (Nees) Hitche. in Contrib. U. S. Nat. 1966] Flora of Bolivia — Foster 853 Herb. 24: 501 (1927). Leaf-blades long, mostly flat. Panicles rather dense, somewhat spicate in appearance. Perfect spikelet: glumes equal or subequal, obtuse, to 1 cm. long, the first glume glabrous to sparsely villous on the upper portion, the apex shortly ciliate; second glume glabrous, the apex shortly ciliate. Awn once or twice geniculate, the basal portion tightly twisted, plumosely ciliate, dark brown, the upper portion light brown, straight, shortly scabrid- pubescent. SANTA CRUZ: SARA: Buenavista, 450-500 m., Steinbach 5430 bis (GH), 7028 (GH). 2. S. parviflorum (Desv.) Hitchc. & Chase in Contrib. U. S. Nat. Herb. 18: 287 (1917). Tufted, the leaf-blades flat to involute. Panicle more open, the pedicels and spikelet-bases villous. Perfect spikelet: glumes subequal, 4-5 mm. long, subacute, glabrous or the first glume sparsely villous. Awn glabrous to faintly scabridulous. SANTA CRUZ: SARA: Buenavista, 450-500 m., Steinbach 1819 (GH), 5428 (GH), 6984 (GH), 7050 bis (GH). 3. S. stipoides (HBK.) Nash, N. Amer. Fl. 17: 129 (1912). Slender, leaf-blades narrow, convolute. Perfect spikelet: glumes equal, 5-6 mm. long, subobtuse, the first glume sparsely villous. Awn light brown, the tightly twisted basal portion somewhat plumose-ciliate. LA PAZ: LARECAJA: vic. Sorata, Mandon 1382 (Hitchcock). SANTA CRUZ: VALLEGRANDE: Vallegrande, Herzog 1789 (Hitchcock). 103. Heteropogon Pers. Perennial or annual. Racemes solitary, digitate, or subdigitate. Staminate spikelets pedicellate, awnless, paired (2-5 pairs) on lowest portion of rachis, this remaining entire. Fertile spikelets sessile, 2-flowered, awned, the lower floret a sterile lemma, each paired with a pedicellate staminate spikelet above the paired staminate spikelets, the rachis disarticulating obliquely, leaving a sharp villous callus below each fertile spikelet. a. Racemes 2-7, digitately arranged (occasionally solitary on axillary branches); awns to 5 cm. long; pedicels villous. .... 1. lH. villosus. a. Racemes solitary; awns 12-15 cm. long; pedicels glabrous. be Glumes all elandless. 4... ereeeenteeeseeern soto Don 2. H. contortus. b. First glume of staminate spikelets with a dorsal row of punc- late Plandg. .eeasciiteetrrinncn iia sot arai eo ERIT IET NI 3. H. melanocarpus. 1. Heteropogon villosus Nees, Agrost. Bras. 362 (1829). Andro- pogon villosus and Agenium villosum of the Catalogue. Perennial, erect or spreading, the culms to 1 m. tall, nodes pubescent to villous. Leaf-sheaths pubescert to glabrate, ciliate, the linear acute blades to 10 cm. long and 4 mm. wide, pubescent above and beneath. Inflo- rescence a terminal panicle of 2-7 digitately or subdigitately arranged racemes on villous peduncles to 1 cm. long, the racemes to 3 cm. long (excluding awns). Staminate spikelets: pedicels villous; first glume to 6 mm. long, acute, hispid to papillate-hispid on upper portion, 354 Rhodora [Vol. 68 many-nerved; second glume somewhat shorter, glabrous, thin-textured, several-nerved; lemma shorter, hyaline, ciliate, the hairs soft and curling. Fertile spikelets: 5 mm. long, excluding awns; first glume 4.5 mm. long, obtuse, villous, only slightly papillate; second glume slightly shorter, acute, sparsely pubescent; lemma about 3 mm. long, thin, terminated by an awn 2.5-5 em. long, basally twisted, geniculate, ultimately straight, pubescent to apically puberulent. SANTA CRUZ: CERCADO: Santa Cruz, Herzog 1308 (Hitchcock). 2. H. contortus (L.) Beauv. ex Roem. & Schult. Syst. Veg. 2: 836 (1817). Perennial, to 8 dm. or 1 m. tall. the culms compressed, nodes glabrous. Leaf-sheaths glabrous, carinate, the linear, acute blades to 15 em. long and 7 mm. wide, scabridulous above but scarcely so be- neath. Inflorescence a terminal, 1-sided raceme about 5 cm, long (ex- cluding awns). Staminate spikelets imbricate, to 1 cm. long, pedicels glabrous; first glume to 1 cm. long, acute, finely short-ciliate, sparsely papillate-hispid above, many-nerved; second glume nearly as long, thin, 1-nerved, glabrous; lemma shorter, hyaline. Fertile spikelets to 7 mm. long; glumes equai, the first brown, densely pubescent, enclosing the second; lemma short, bearing a geniculate, basally twisted awn to 12 (-15) em. long, the base pubescent to plumuse, the apex finely vuberulent. LA PAZ: NOR YUNGAS: Coripata, Hitchcock 22678 (GH). CHUQUISACA: SUR CINTI: Camataqui, 2500 m., Fiebrig 3074 (GH). 3. H. melanocarpus (Ell) Benth. in Journ. Linn. Soc. Bot. 19: 71 (1882). Annual, to 2 m. tall. Leaf-sheaths glabrous, carinate, the keel with a row of small glands, especially on sheaths of upper leaves, the lance-linear blades to 30 em. long and 1 cm. wide, scabrid above and beneath. Inflorescences often densely crowded in upper leaves, the racemes to 6 em. long (excluding the awns); inflorescence to 2-3 dm. long. Staminate spikelets to 1.5-2.5 em. long, glabrous, the pedicels glabrous; first glume acute, many-nerved, the margins hyaline, with a dorsal line of flat punctate glands in the middle of the glume. Fertile spikelets about 7-10 mm. long; glumes equal, firm, subacute to obtuse, dark brown, pubescent; lemma shorter, thin, bearing a 10-12 em. geniculate awn, the base twisted, pubescent to subplumose, the upper half straight, puberulent. LA PAZ: NOR YUNGAS: Coripata, Hitchcock 22678 (GH). 104. Trachypogon Nees Perennials. Racemes terminal, 1-3 in an inflorescence, the rachis disarticulating. Spikelets paired, the staminate spikelet awnless, ses- sile or nearly so, persistent; perfect spikelet awned, pedicellate, the pedicel disarticulating, leaving a sharp villous callus below the spikelet. a. Leaf-blades glabrous to scabrid. b. Racemes usually solitary; awn pubescent; glume of staminate spikelet pubescent. ............... esee 1. T. montufari. 1966] Flora of Bolivia — Foster 355 b. Racemes 2-3; awn plumose; glume of staminate spikelet glab- MUCH EU E ME iU EMEN ET 2. T. plumosus. "YES VES TOS pad Eoo EID CPU SRI NDS M e ea aer aaas 3. T. canescens. 1. Trachypogon montufari (HBK.) Nees, Agrost. Bras. 342 (1829). T. polymorphus var. bolivianus Pilger in Engler, Bot. Jahrb. 27: 22 (1899). Culms to 1 m. tall, the nodes hispid to villous. Leaf-sheaths glabrous, auricles erect, to 1 cm. long, the flat or involute blades to 8 mm. wide, shorter than the inflorescence, glabrous but scabridulous. Inflorescence usually a solitary raceme to 15 em. long. Staminate spikelet: first glume to 7 mm. long, subacute, dorsally rounded, many-nerved, sparsely pubescent and shortly hispid-ciliate; second glume to 8 mm. long, acute, few-nerved, glabrous or softly and sparsely pubescent along the margins; lemma shorter, hyaline, long-ciliate, the hairs soft. Perfect spikelet: glumes nearly equal, to 7-8 mm. long, few-nerved, firm-textured, pubescent; lemma shorter, hyaline, the geniculate awn to 6 cm. long, densely long-pubescent, the hairs shorter toward the apex. Bolivian Plateau: Bang 1079 (GH; type- number of T. polymorphus var. bolivianus). SANTA CRUZ: SARA: Buenavista, 500 m., Steinbach 5379 (Gu). TARIJA: ARCE: Camacho, Fiebrig 2864 (GH). 2. T. plumosus (Humb. & Bonpl. ex Willd.) Nees, Agrost. Bras. 344 (1829). Taller than T. montufari; racemes 2-3 (rarely 1); awn plumose; first glume of staminate spikelet glabrous. LA PAZ: LARE- CAJA: Hacienda Casana, Buchtien 7145 (Hitchcock). 3. T. canescens Nees, Agrost. Bras. 943 (1829). Very similar to T. montufari, but the leaf-sheaths sparsely hispid to papillate-hispid, the involute blades villous. Awn plumose, at least below. LA PAZ: NOR YUNGAS: Puente Villa, 1700 m., Cárdenas 3613 (GH). SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 6947 (GH). 105. Elyonurus Humb. & Bonpl. ex Willd. Perennial, up to 1 m. tall, the terminal racemes usually solitary, to 10-15 em. long, often much shorter, the rachis ultimately disarticu- lating. Spikelets paired, awnless; staminate spikelet pedicellate; per- fect spikelet sessile, appressed against the rachis; spikelet-pairs fall- ing attached to a rachis-joint. Glumes entire or bidentate, slightly glandular on the marginal nerves; lemmas hyaline, thin-textured; palea obsolete. First glume of staminate spikelet entire. ......... ee 1. E. adustus. First glume of staminate spikelet bidentate (sometimes only slightly ^9 BERT 2. E. tripsacoides. 1. Elyonurus adustus (Trin.) Ekman in Ark. Bot. 13(10): 6 (1913). Caespitose, the bases slightly bulbous. Leaf-sheaths glabrous, the linear, flat or inrolled, scabridulous leaf-blades to 4 mm. wide, shorter than the inflorescence. Raceme-rachis and pedicels pubescent to villous. Staminate spikelets: glumes subequal, about 5 mm. long, entire, acute, 356 Rhodora [Vol. 68 the first elume more or less densely hirsute and hispid-ciliate on the submarginal keels, the margins enclosing the second glume, the second glume slightly shorter, pubescent, few-nerved, the upper margin sparsely ciliate with soft curling hairs. Perfect spikelets: glumes subequal, the first glume about 6 mm. long, acute, densely hispid- villous, the second glume slightly shorter, pubescent. LA PAZ: LARE- CAJA: Hacienda Casana, 1600 m., Buchtien 7138 (GH); NOR YUNGAS: Polo-Polo, near Coroico, 1100 m., Buchtien s. n. (GH). SANTA CRUZ: SARA; Buenavista, 450-500 m., Steinbach 5244 (GH), 5404 (GH), 6620 bis (GH), 6641 (GH), 6642 (GH), 6875 (GH); near Portachuelo, 400 m., Steinbach 3136 (GH). 2. E. tripsacoides Humb. & Bonpl. ex Willd. Sp. Pl. 4: 941 (1806). E. tripsacoides var. ciliaris (HBK.) Hack. in DC. Monogr. Phan. 6: 333 (1889). E. tripsacoides var. brevidentatus Hack. in DC. Monogr. Phan. 6: 334 (1889). Very similar to E. adustus, but the bases often more bulbous and shortly rhizomatous. Leaf-blades somewhat pilose above, near the base. First glume of staminate spikelet bidentate, sometimes only shortly so. COCHABAMBA: CERCADO: 5 km. southeast of Cochabamba, 2800 m., Eyerdam 24914 (GH); Arani, 2700 m., Cárdenas 2382 (GH). SANTA CRUZ: SARA: Buenavista, 400-450 m., Steinbach 3270 (GH), 6514 (GH), 6895 (GH), 6896 (GH). TARIJA: ARCE: Camacho, 2500 m., Fiebrig 2577 (GH). Three forms are known in Bolivia. The first glume of the staminate spikelet may be glabrous and strongly ciliate (typical), pubescent (var. brevidentatus) or villous (var. ciliaris). The generic name is often spelled Elionurus, but I have retained the original spelling, Elyonurus, 106. Manisuris L. Perennials, often tall, in moist or wet places. Leaves glabrous, culms glabrous, Inflorescences racemes, these sometimes aggregated in the upper leaf-axils. Spikelets awnless, paired on a disarticulating rachis, the perfect spikelet sessile, appressed against or adnate to the rachis, the sterile spikelet pedicellate or reduced to a pedicel only; glumes firm, coriaceous, lemma and palea hyaline. First glume of perfect spikelet bidentate, foveolate-pitted on lower nIY MEMMERNEMENRRERENMMMMMMMMMEMKFNISSWKEMMN 2. M. aurita. First glume of perfect spikelet not bidentate, not foveolate-pitted. munseasfessesseenaessessanusbesessaysensssssgusvevedercossssiee;ssusesssssassecsarseneseeses 1. M. altissima. 1. Manisuris altissima (Poir.) Hitehe. in Journ. Wash, Acad. Sci. 24: 292 (1934). M. fasciculata (Lam.) Hitch. in Amer. Journ. Bot. 2: 299 (1915). Culms to 1 m. tall. Leaf-blades flat or folded, to 5 mm. wide; apex of leaf-sheaths densely long-ciliate, base of blades sparsely 1966] Flora of Bolivia — Foster 357 short-ciliate. Racemes to 10 em. long, usually shorter, the rachis rather tardily disarticulating. Perfect spikelet: glumes unequal, first glume 5-7 mm. long, abruptly acute or subobtuse, 9-11-nerved, the nerves rather prominent, the keel narrowly winged on the upper portion, second glume about 1 mm. longer, long-acute, few-nerved, the margins inflolded, with the glume thus somewhat cucullate. Sterile spikelet: glumes short and equal, otherwise resembling the perfect spikelet. LA PAz: SUR YUNGAS: Espía, White (M. E.) 627 (Hitch- cock). 2. M. aurita (Steud.) Hitche & Chase in Contrib. U. S. Nat. Herb. 18: 276 (1917). Culms to 1-1.5 m. tall. Leaf-blades flat or folded, to 8 mm. wide, usually narrower, glabrous. Racemes to 10 cm. long, usually less, the base often partially enclosed by a leaf-sheath, the rachis quickly disarticulating. Perfect spikelet: glumes unequal, the first glume 3-4 mm. long, wing-margined above, bidentate, the teeth broad and overlapping, usually foveolate-pitted on the lower half (some glumes occasionally not pitted), nerves not conspicuous, second glume much shorter, the keel winged or at least pronounced, other nerves absent or inconspicuous. SANTA CRUZ: SARA: Buenavista, 450 m., Steinbach 6973 bis (GH). 107. Hackelochloa O. Ktze. Hackelochloa granularis (L.) O. Ktze. Rev. Gen. 2: 776 (1891). A much-branched annual, 3-10 dm. high, coarsely hirsute on leaf-sheaths and blades, the sheaths papillate, the blades ciliate. Inflorescences axillary and terminal racemes (often fascicled) to 2.5 cm. long. Spikelets paired, the lower sessile and perfect, clasping the pedicel of the upper, this sterile or staminate; disarticulation in the rachis. Fertile spikelet: first glume about 1 mm. long, globose and globosely pitted or alveolate, finely and minutely puberulent, indurate, conceal- ing the second glume and lemma. Sterile spikelet: glumes equal, about 2 mm. long, broadly lanceolate, several-nerved, not indurate, bicarinate, the keels ciliolate, the margins of the first glume partially enclosing the second glume. SANTA CRUZ: SARA: Monte de Palo- metillas, 400 m., Steinbach 2363 (Gu); Buenavista, 800 m., Steinbach 5301 (GH). LA PAZ: LARECAJA: vic. Sorata, Bang 1310 in part (GH); NOR YUNGAS: Milluguaya, 1300 m., Buchtien 4189 (GH). 108. Tripsacum L. Tripsacum australe Cutler & Anderson in Ann. Missouri Bot. Gard. 28: 259 (1941). T. dactyloides of the Catalogue. Tall perennial, the culms to 2 m. high. Leaf-sheaths pilose at the margins and at the auricled apex, tomentose within; leaf-blades long, flat, to 4 cm. wide, subpetiolate at the base. Inflorescence terminal, of 1-4 spikes, stami- nate spikelets above the pistillate, but on the same rachis. Pistillate 358 Rhodora [Vol. 68 spikelets solitary at the nodes, embedded in the axis, with 1 fertile floret and 1 sterile lemma; first glume coriaceous, about 5-10 mm. long, the margin ciliate at base, covering the spikelet, the second glume similar but smaller; fertile lemma shorter than the glumes, hyaline. Staminate spikelets paired at the nodes of rachis, both sessile or 1 pedicellate; glumes firm, chartaceous, about as long as the pistillate glumes. L4 PAZ: NOR YUNGAS: Coroico, 1560 m., Hitchcock 22721 (GH). 109. Coix L. Coix lachryma-jobi L. Sp. Pl. 972 (1753). Tall perennials (or some- times annual ?), branched, the culms to 1 m. high. Leaf-blades much exceeding the inflorescences, to 4 cm. wide, lance-ovate, the base cordate-clasping. Inflorescence long-pedunculate, of 2-3 short pistil- late spikelets at the base, enclosed in a hard bony involucre to 1 em. long, only the spikelet-tips exserted; staminate spikelets usually 3, on a slender axis ultimately exserted from the involucre, the whole raceme about 1 cm. long. COCHABAMBA: CHAPARE: Espíritu Santo, Buchtien 2521 (Hitchcock). GRAY HERBARIUM, HARVARD UNIVERSITY NATIVE NORTH AMERICAN SPECIES OF LESPEDEZA (LEGUMINOSAE)* ANDRE F. CLEWELL The genus Lespedeza occurs in eastern North America, eastern Asia, and Australia. The native American species are perennial herbs, mostly occurring in dry, open sites, such as old fields, roadsides, prairies, and savannahs, The Asian species include shrubs and annuals, the latter some- times being separated into Kummerowia Schindl. (Hutch- inson 1964). Two annuals, L. striata (Thunb.) H. & A. and L. stipulacea Maxim., and one herbaceous perennial, L. cuneata (Dumont) G. Don, have been introduced widely in eastern North America from Asia as forage for cattle and wildlife, for preventing soil erosion, and for green manure. All three species have become well-established weeds along roadsides and in similar sites. An Asian shrub, L. bicolor Turez., has become much less widely established after its introduction for purposes of wildlife management, and several other Asian introductions either have not es- caped or have remained local. None of the Asian species are treated in this paper; Wilbur (1963) has described the common Asian introductions in detail. I have examined *This work was supported by a National Science Foundation grant, GB-1365, and by funds from the Florida State University Research Council. My appreciation is extended to the curators of the following herbaria who allowed me to examine their specimens. Abbreviations for herbaria follow Lanjouw & Stafleu (1964), Al specimens from CINC, DAO, DUKE, FLAS, FSU, GA, IND, ISC, LAF, LL, MISSA, MO, NCU, NEBC, NY, OC, PUL, SMU, TENN, TEX, UMO; regional collections from CM, ILLS, IA, KSC, KY, MICH, MIN, NEB, NYS, OKL, OS, RBGH, SDU, TRT, WVA; selected species from F, GH, KANU, US; types from PH, I thank R. K. Godfrey for his encouragement and for critically reading the manuscript. Many people have helped make this work possible; some are men- tioned in the text, others include H. E. Ahles, J. R. Coleman, B. N. Culbertson, D. Demaree, J. Ewan, C.R. Gunn, C. B. Heiser, Jr., N. E. Hill, D. Isely, T. Kakac, F. C. MacKeever, T. E. Melchert, Mrs. J. T. Mickel, the late E. J. Palmer, W. W. Payne, L. H. Shinners, A. M. Torres, and D. B. Ward. 360 Rhodora [Vol. 68 specimens of about 36 Asian species; more have been de- scribed. Van Steenis (1957) pointed out that certain Asian species intergrade and are in need of revision. Hiroyoshi Ohashi (pers. com.) at the University of Tokyo recently began studying the Japanese species. Until the Asian species are satisfactorily circumscribed, attempts to divide the genus into subgenera or sections will be premature; Isely (1955) has summarized the previous attempts. Lespedeza has long been considered a member of the Hedysareae which, according to Isely (1955) and au- thors cited by him, is an arbitrarily defined tribe of poly- typic origin. Hutchinson (1964) has redefined the tribes of the Leguminosae and has described a new tribe, the Lespe- dezae, to which Lespedeza and several other genera belong (but not Desmodium). LITERATURE Important contributions from the older literature to a taxonomic understanding of the American species include those by Maximowicz (1873) who prepared the first syn- opsis on a world-wide basis after most of the species had been discovered; Britton (1893) who first revised the North American species ; and Schindler (1913) who wrote the only comprehensive world-wide monograph of Lespedeza. More recent critical treatments of the North American taxa are by Blake (1924) on selected species, Fassett (1939) on those occurring in Wisconsin, Fernald (1941) the white-flowered species, Gambill (1953) the species from Illinois, Isely (1955), a detailed account of those occurring in the north- central United States, Turner (1959) the Texas species, and Wilbur (1963), accurate, detailed descriptions of the species in North Carolina. Papers concerned with popula- tion studies, breeding systems, cytology, ecology, hybridiza- tion, and evolution are Clewell (1964b, 1966). Many of the data in those papers and in this one came from a disserta- tion, Clewell (1963). Many references to other papers on Lespedeza are cited in the Botany Subject Index (compiled by the U.S.D.A. Library, 1958) vols. 5: 4467-69 and 7: 7060-62. 1966] Lespedeza — Clewell 361 AN EVALUATION OF SOME TAXONOMIC CHARACTERS To characterize and identify species, it is important to distinguish cleistogamously-formed pods from chasmoga- mous pods. With rare exceptions cleistogamous and chasmogamous flowers occur on separate racemes. In con- trast to racemes with chasmogamous flowers, cleistogamous- flowered racemes generally are shorter, are borne farther down the stem, and bear a few more flowers. Hanson (1943) and Hanson & Cope (1955) have described cleistogamous Fig. 1. Chasmogamous and cleistogamous pods. la: Lespedeza intermedia (chasmogamous) ; 1b: L. intermedia (cleistogamous) ; 1c: L. violacea (chasmogamous, style broken); 1d: L. violacea (cleistog- amous); le: L. repens (cleistogamous). 362 Rhodora [Vol. 68 flowers of Lespedeza in detail. They pointed out that in bud the style recurves until the stigma rests upon one or more anthers, the filaments of which do not elongate. Pollen grains germinate within the anthers upon which the stigma rests and the tubes grow through the indehiscent anther wall into the stigma (cleistantheric pollination). The style is persistent on mature pods and reflects its mode of develop- ment by being sharply recurved. Sometimes an anther adheres to the stigma of a mature pod, evidently held there by the pollen tubes. Styles persistent on chasmogamous pods are longer and either erect or variously curved but not sharply recurved. Cleistogamous pods can be distinguished from chasmogamous ones primarily by the shape of the style and secondarily by the slightly shorter length of the calyxes (Fig. 1). Cleistogamous pods also tend to be shorter and more orbicular, rather than elliptic. Pollen grains, taken from dried plants and stained with cotton blue in lactophenol, revealed only nominal variation both within and between species. The grains have 3 pores and no ornamentation. Most are 27-33 microns in diameter ; however, some plants have slightly larger grains, particu- larly certain plants of Lespedeza capitata and L. violacea. Gross pollen morphology is not taxonomically significant. Chromosome numbers cannot be employed in distinguish- ing the American taxa, because all plants examined had 10 pairs of chromosomes (Clewell 1966). Pierce (1939) found the chromosomes to be small and rather uniform with no significant variation between species. My own observations confirm his conclusions, thereby negating karyotypic varia- tion as a useful character. D. A. Levin (pers. com.) has prepared chromatograms from pigments extracted from the leaves of several lespe- dezas that I sent to him. Included were a specimen of a hybrid and specimens of its parental species and of several other species, including one native to Asia. He concluded that, "the patterns of the different species are quite similar and further study of Lespedeza using the present techniques does not appear warranted." Levin discovered a close simi- 1966] Lespedeza — Clewell 363 larity between some of the chromatograms of the lespedezas and a chromatogram of a plant of Desmodium ciliare (Muhl.) DC. This suggests that a comparative chromato- graphic study of representatives of the Hedysareae might be rewarding in assessing relationships between genera. CONCEPT OF SPECIES I have observed many populations of Lespedeza over a period of 8 years on the Atlantic Coastal Plain and adjacent physiographic provinces from the Carolinas to the Missis- sippi River, in most of Indiana, and in parts of Texas, Iowa, and the Ozark Plateau. Field observations and analysis led me to my recognition of 11 species. Subsequently progeny tests yielded data supporting my concepts of these species, as did observations on wild plants transplanted and grown in a greenhouse. Each progeny test consisted of removing seeds from one plant, which was collected in a natural popu- lation, and planting them under uniform conditions in a garden. The mature offspring were then compared with each other and with their parental plant. With the exception of 6 obvious hybrids, all 378 sibs of the 24 progeny tests clearly resembled their respective parental plants. The species tested were L. angustifolia, L. capitata, L. hirta (both subspp.), L. intermedia, L. procumbens, L. stuevet, L. violacea, and L. virginica (Clewell 1966). Examination of herbarium specimens from throughout the geographic ranges of all species has contributed further to the recog- nition of each species and has yielded information which has allowed not only the precise delimitation of the species but also the assessment of the geographic variation existing within each species. Usually a few plants in each population are difficult to identify. Some of these have proved to be shade forms which one can learn to correlate with the variation in the “good” species (see notes on L. intermedia in the taxonomic treat- ment). Other plants appear to be hybrids. Virtually every population of about 50 plants or more consisted of 2-7 species and contained at least one hybrid. Not infrequently 364 Rhodora [Vol. 68 hybrids composed 156 or more of the plants present. The parentage of nearly all hybrids was determined easily in the field, even though several hybrid combinations were present. This was possible because most hybrids appeared to be F,s, and hybrid swarms with many notomorphs were rare. The parentages of hybrids that have been deposited in herbaria are less easily postulated than of hybrids seen in the field with their parental species. My identifications of Lespedeza angustifolia X hirta and L. procumbens X stuevei have gained credence through comparisons with artificially synthesized hybrids of these combinations. The identifications of 24 plants of hybrid origin representing 9 hybrid combinations have been verified through progeny tests yielding a total of 175 mature sibs. These sibs com- prised a series of segregants, often with forms characteristic of one parent or the other and including various forms be- tween them (Clewell 1966). My success in having field identification of both hybrids and species verified by progeny tests has convinced me that with a few exceptions hybrids can be distinguished easily from species and that the par- entages of hybrids can be postulated accurately. Only one of the 48 plants upon which progeny tests were run proved to be misidentified in the field. I have used two criteria for cireumscribing species: First, plants of any species must be morphologically distinct from plants of other species in a population. Exceptions are shade forms and hybrids. Second, the morphological variation within a species throughout its geographic range has precise limits, and no species seemingly merges into another species in any region. If taxa merge, then they are assigned sub- specific status. This species concept applies to Lespedeza as it presently occurs and not necessarily to other genera or to Lespedeza at other times in its evolutionary history. In the taxonomic treatment the descriptions of species are prepared so that almost any character of one species can be compared with the same character as it occurs in every other species. Only the more important diagnostic characters are described, thereby emphasizing specific de- 1966] Lespedeza — Clewell 365 limitation. I have omitted lists of specimens examined. My annotations on the great majority of the specimens I examined serve as documentation of this study. These speci- mens are deposited in the herbaria listed in the acknowl- edgements. Mimeographed lists of specimens serving as vouchers for the distribution maps (Figs. 2-13) are avail- able from me on request; a copy will be deposited in the library of Florida State University. No index to exsiccatae was prepared. Identification of duplicate specimens in other herbaria by consulting such an index would be hazardous, because in many instances duplicates under one collection- number represent more than one species and, more com- monly, hybrids. TAXONOMIC TREATMENT* Lespedeza Michx. Fl. Bor.-Amer. 2: 70 (1803). (Type species: L. procumbens). Hedysarum L. Sp. Pl. 2: 745 (1753). (in part). Medicago L. Sp. Pl. 2: 778 (1753). (in part). Lespedezia Hornem, Hort. Reg. Bot. Hafn. 2: 699 (1815). Perennial herbs from woody rhizomes bearing 1-45 aerial stems, usually branched, and occasionally a short, underground stolon termi- nating in a new plant; INDUMENT of appressed trichomes (appressed- pubescent, sometimes silvery or sericeous) or soft, erect or spreading trichomes (pilose if short, villous if long and undulating); LEAVES trifoliolate (rarely some with 4-5 leaflets); LEAFLETS entire, nearly equal in shape, often mucro-tipped, the terminal one often slightly longer than the lateral ones; STIPULES inconspicuous, persistent, the longer ones linear or setaceous, the shorter ones subulate; STIPELS minute, usually absent; RACEMES axillary, sometimes spicate or capi- tate, commonly some or all reduced to axillary fascicles, rarely panic- ulate; FLOWERS borne in pairs, each pair subtended by 3 bracteoles on the peduncle, each flower pedicellate to nearly sessile and subtended by 2 bracteoles borne near or at the apex of the pedicel opposite the flat surfaces of the pod; CHASMOGAMOUS FLOWERS showy, CALYX campanulate, persistent in fruit, the lobes subequal, the upper 2 being more connate than the others, COROLLA violet, roseate, cream-colored, or white with a purplish spot near the base of the standard; STANDARD suborbicular to oblong, clawed; WINGS falcate-oblong, about equaling the keel; STAMENS diadelphous; ANTHERS uniform; STYLE long, fili- *Synonyms of Lespedeza are limited to those used for native Ameri- can plants. The generic description also applies only to native American plants. 366 Rhodora [Vol. 68 form, often persistent in fruit; PODS ovate to orbicular, sometimes somewhat asymmetrical, lenticular or laterally compressed, reticulate, mostly indehiscent; CLEISTOGAMOUS FLOWERS often present, sometimes exclusively, about 0.1 the size of chasmogamous flowers, nearly always in separate racemes from chasmogamous flowers, often axillary and averaging a few more flowers per raceme than racemes of chasmoga- mous flowers; CALYX as in chasmogamous flowers though often slightly shorter; COROLLA greatly reduced; STYLE short, usually persistent in fruit, the distal half sharply recurved and (nearly) touching the proximal half; Pops as in chasmogamous flowers, sometimes tending to be more orbicular and slightly shorter than in chasmogamous flowers; Pops of both chasmogamous and cleistogamous flowers l-seeded; SEEDS straw-colored to black, sometimes mottled, oval and somewhat laterally compressed. Ricker (1934) explained that Michaux commemorated the governor of East Florida, 1784-1790, in the name, Lespedeza. The governor’s name was not D. Lespedez, as stated by Michaux (1803) but Vincente Manuel de Cespedes. Ricker suggested that the change in spelling may have stemmed from an inability of Michaux's son or of L. C. M. Richard to interpret Michaux's notes which were damaged in a ship- wreck. The spelling of Lespedeza must be retained under Article 73 of the International Code of Botanical Nomer- clature (Lanjouw 1961). Through the efforts of Hochreu- tiner (1934), Lespedeza now serves as an illustration for the application of this Article. I was unable to examine the types of several names ap- plied to the lespedezas, because these types are deposited in European herbaria and were unavailable to me, I have justified the identification of some of these names with certain species in the commentaries on these species. I have associated the remainder of these names with various species both on the basis of the original descriptions and from the comments of Schindler (1913). Schindler examined the type specimens in the European herbaria, and he identified the names in nearly all previous treatments of the genus with the corresponding taxa in his monograph. His lucid discussions made it possible for me to compare my species concepts with his and to gain insight into the concepts of prior authors. 1966] Lespedeza — Clewell 367 KEY TO NATIVE AMERICAN SPECIES OF LESPEDEZA 1. Stems procumbent to weakly ascending. 2. Stems procumbent, pilose throughout (uncommonly mostly pilose in b terana eae RR E TEES L. procumbens 2. Stems appressed-pubescent. 3. Stems erect up to 1.5 dm, then trailing; the longer stipules 4-8 mm long; leaflets firm-textured, gray-green beneath; plants ofi central Texase aon e E aE L. texana 3. Stems procumbent to weakly ascending; the longer stipules 1.5-6 mm long; leaflets membranous, green beneath. 4. Stems procumbent; keel about equaling wings; calyx of the cleistogamous pods 0.25-0.5 times the length of the pod; stipules mostly subulate, 1.5-4 mm long. ..............-. L. repens 4. Stems weakly ascending; keel usually longer than wings; calyx of cleistogamous pods up to 0.25 mm long, mostly shorter; stipules mostly filiform, 2.5-6 mm long. ..................- PR IE M ELM d UE M E L. violacea 1. Stems erect or strongly ascending. 5. Calyx nearly equaling or exceeding the mature pod; flowers white with a purple throat (L. leptostachya sometimes has light purple flowers) ; racemes with 10-44 flowers. 6. The larger terminal leaflets more than half as wide as long. [RON Chee E PRRRPRPRRRIRURC INI: T (A. SENI: L. hirta 6. The longer terminal leaflets less than half as wide as long. 7. Rachises longer than their petioles; racemes included within or exserted up to 1.5 times beyond their subtending leaves; calyx mostly 8-18 mm long. ............... een L, capitata 7. Rachises of the longer terminal leaflets equaling or gen- erally shorter than their petioles; racemes 1-4 times longer than their subtending leaves; calyx 4-9 mm long. 8. Leaflets glabrous above and stems appressed-pubescent or leaflets pubescent and stems densely pilose; racemes dense, the flowers limited usually to the terminal third of the peduncle; plants of the Atlantic Coastal Plain. ........ puc T A A E L. angustifolia 8. Leaflets above and stems appressed-pubescent; racemes open, the flowers scattered on the terminal half of the peduncle; plants of the upper Mississippi Valley. ............ L. leptostachya 5. Calyx half as long as the mature pod or shorter; flowers purple; racemes with 4-14 flowers. 9. Keel often noticeably longer than wings; some racemes usually exserted 2-5 times beyond their subtending leaves; calyx of the cleistogamous pods up to 0.2 (rarely 0.25) times the length of the mature pod; stems sparsely appressed-pubescent ; 368 Rhodora [Vol. 68 Stipules 2.5-6 mm long. sees L. violacea, 9. Keel equaling or shorter than wings; racemes included within or slightly exserted beyond their subtending leaves; calyx of the cleistogamous pods 0.2-0.4 times the length of the mature pod; stems appressed-pubescent or pilose; stipules rarely ex- ceeding 4 mm long. 10. Leaflets glabrous above (rarely appressed-pubescent along the midrib), the longer ones more than 0.3 times as wide as long; stems moderately appressed-pubescent. .................... IM L. intermedia 10. Leaflets appressed-pubescent, at least along the midrib, to pilose, the longer ones 0.1-0.5 times as wide as long; stems appressed-pubescent to pilose. ll. Terminal leaflets 0.1-0.3 times as wide as long, sparsely appressed-pubescent above, sometimes only along the midrib. alll r raa, L. virginica 11. Terminal leaflets 0.3-0.5 times as wide as long, appressed- pubescent to pilose above. a. L. stuevei Lespedeza angustifolia (Pursh) Ell. Sketch Bot. S. C. 2: 206 (1824). (T.: Clayton 934 [BM]). Lespedeza capitata var. angustifolia Pursh Fl. Amer. Sept. 480 (1814). Lespedeza hirta var. angustifolia Maxim. Acta Horti Petr. 2: 379 (1873). Lespedeza angustifolia var. brevifolia Britt. Trans. N.Y. Acad. Sci. 12: 68 (1893). (T.: A. W. Chapman s. n., Florida [NY!]). Lespedeza angustifolia f. subvelutina Fern. Rhodora 42: 457 (1940). (T.: M. L. Fernald & Bayard Long 9077, sphagnous bog about 1 mile n, w. of Dahlia, Greenville Co., Va., 20 August 1938 [GH!]). Lespedeza hirta var. intercursa Fern. Rhodora 43: 572 (1941). (T.: M. L. Fernald, Ludlow Griscom, & Bayard Long 6622, argil- laceous & siliceous swales & swaley thickets, south of Zuni, Isle of Wight Co., Va., 20, 22 Aug. 1936 [GH!]). STEMS erect or ascending to about 1.8 m tall, short-appressed-pubes- cent or sometimes short-pilose; STIPULEs filiform to narrowly subulate, 2-4 mm long; PETIOLES short, the terminal leaflets of the larger leaves mostly being at least 2 times longer than the combined lengths of their petiole and rachis and at least 3 times longer on leaves among the racemes; RACHISES of terminal leaflets of the petiolate leaves equaling or shorter than their petioles; TERMINAL LEAFLETS linear or narrowly oblong, the longer ones up to 6 cm long and .07-.25 times as wide, glabrous or occasionally appressed-pubescent above, if ap- pressed-pubescent above, then stems Short-pilose; STIPELS often present on some leaves, up to 1 mm long; RACEMES 1-3.5 times longer 1966] Lespedeza — Clewell 269 than their subtending leaves and mostly bearing 10-26 flowers on the terminal 0.3-0.25 of the peduncle; BRACTEOLES mostly exceeding the base of the lobes of the calyx; PEDICELS 1-2 mm long; FLOWERS 5-7 mm long; CALYX 5-9 mm long, nearly chorisepalous, about equaling to much exceeding the mature pod which is easily visible between the lobes; COROLLA white or cream-colored; WINGS equaling or slightly exceeding the keel; STANDARD equaling or slightly exceeding the wings; PODS oval or elliptic, 3.5-5 mm long, moderately to densely appressed-pubescent ; CLEISTOGAMOUS PODS commonly present, general- ly oval, 3-4 mm long, appressed-pubescent, exceeded by the calyx; SEEDS greenish-yellow to black. Commonly on moist sand in open pine flatwoods, wet savannahs, along railroad tracks, and in open flood plains; occasionally on drier long leaf pine-scrub oak-wire grass sand ridges, particularly where disturbed; uncommonly on loam soils of pinelands, old fields, and roadsides; from Cape Cod along the Atlantic coast to southeastern Virginia and then both coastal and inland on the Atlantic Coastal Plain to northern Florida; disjunctions in the Georgia Piedmont (Walton Co.), western North Carolina (Henderson Co.), central Tennessee (Coffee, Warren Cos.) central Florida (Hillsborough Co.), and coastal Mississippi (Jackson Co.) (Fig. 2). Flowers from early August through mid-Novem- ber. N. K. B. Robson (pers. com.) has examined the holotype and said that it “agrees well" with the key characters. Whether or not Britton (1893) intended to publish the name, Lespedeza angustifolia var. brevifolia, is uncertain ; his only mention of that name is the following: “I have specimens of a peculiarly short-leaved form of the plant from Florida, communicated by Dr. Chapman, which may claim recognition as var. BREVIFOLIA.” The type is a frag- ment of typical L. angustifolia, which Britton received from Chapman in April, 1891. Most plants of Lespedeza angustifolia have leaflets gla- brous above and scattered, appressed pubescence on their stems; however, some have both stems and leaflets densely short-pilose. Both types occur sympatrically throughout the range of the species. The latter, pilose type has been rec- ognized as L. angustifolia f. subvelutina. If this form is 370 Rhodora [Vol. 68 recognized, then it is reasonable that many other forms of equal insignificance in the genus should be named, causing a cumbersome and impractical classification. Eight specimens annotated as Lespedeza hirta var. inter- cursa by Fernald, all of which were collected by Fernald in southeastern Virginia and deposited in GH and US, are typical L. angustifolia. I have searched without success for Lespedeza angusti- folia from the Choctawhatchee River in western Florida into eastern Louisiana. It occurs sympatrically with L. capitata in Florida. The occurrence of L. capitata westward near the Gulf coast into eastern Texas is in the same kinds of habitats occupied by both species in Florida while L. angustifolia occurs in old fields and roadsides and in fre- quently burned-over pineland in the Tallahassee Red Hills, an area resembling the Piedmont in soil and physiography. The disjunct populations in central Georgia and western North Carolina might owe their appearance to migrations inland along the flood plains of the major river systems. I hesitate to explain the occurrence of the populations in Tennessee. A. J. Sharp (in verb.) stated that several coastal plain species of other genera occur there. Relatively little morphological variation exists in the species, and no varia- tion is associated with any particular geographic region or habitat. Closely allied morphologically to Lespedeza angustifolia is L. leptostachya, an upland prairie endemic of the north- central states. The latter differs from L. angustifolia by having a uniformly appressed pubescence on both stems and leaves and densely so on the cleistogamous pods; flowers less densely arranged on the peduncles; narrower calyx lobes subtending the cleistogamous pods; and corolla some- times light purple. Lespedeza capitata Michx. Fl. Bor.-Amer. 2: 71 (1803). (T.: Michaux [P]). Hedysarum umbellulatum Walt. Fl. Car. 184 (1788), non L. 1753. Hedysarum frutescens Muhl. in Willd, Sp. Pl. 3: 1193 (1803), non L. 1753. 1966] Lespedeza — Clewell 371 Hedysarum conglomeratum Poir. in Lam. Encycl. 6: 416 (1804). Lespedeza frutescens Ell. Sketch Bot. S. C. 2: 206 (1824). Lespedeza stuevei DC. Prodr. 2: 349 (1825), non Nutt. 1818. Lespedeza capitata var. sericea Hooker Comp. Bot. Mag. 1: 23 (1835). (T.: Drummond 227, Jacksonville, herb. not indicated). Lespedeza capitata var. vulgaris T. & G. Fl. N. Amer. 1: 368 (1840). Lespedeza velutina Bickn. Torreya 1: 102 (1901). (T.: E. P. Bick- nell s. n., Woodlawn, N. Y., 28 Aug. 1898 [NY!]). Lespedeza bicknellii House Torreya 5: 167 (1905). (T.: E. P. Bick- nell s. n., Woodlawn, N. Y., 28 Aug. 1898 [NY!]). Lespedeza capitata var. velutina Fern. Rhodora 10: 51 (1908), Lespedeza capitata var. stenophylla Bissell & Fern. Rhodora 14: 92 (1912). (T.: F. E. McDonald s. n., open, dry ground, rare, Peoria, Peoria Co., Ill, Sept., 1904 [GH!]). Lespedeza capitata var. typica Fern. Rhodora 43: 576 (1941). Lespedeza capitata var. stenophylla f. argentea Fern. Rhodora 43: 579 (1941). (T.: H. A. Gleason s. n., sanddunes, Havana, Mason Co., Ill, Aug. 1903 [GH!]). STEMS erect or ascending, to about 2 m tall, short appressed-pubescent to long-villous with scattered to dense hairs; STIPULES filiform to nar- rowly subulate, to about 7 mm long; PETIOLES short, the terminal leaflets of the larger leaves being at least 2 times longer than the combined lengths of their rachises and petioles and 3-9 times longer on leaves among the racemes; RACHISES of terminal leaflets rarely equaling and mostly longer than their petioles; TERMINAL LEAFLETS linear, oblong, narrowly rhombate, or elliptie, the longer ones up to 5 cm long and 0.1-0.45 times as wide, glabrous, glabrate, appressed- pubescent, or sericeous above; STIPELS rarely present; RACEMES capi- tate, included or exserted by a stout peduncle up to 1.5 times beyond their subtending leaves and generally bearing 14-30 flowers; PEDICELS 1-3 mm long; BRACTEOLES mostly exceeding the base of the lobes of the calyx; FLOWERS mostly 7-9 mm long; CALYX (7-)8-13 mm long, deeply lobed and nearly chorisepalous, the sinuses often very narrow, partially hiding and greatly exceeding the pod; COROLLA white or cream-colored; STANDARD longer than the keel and longer than to nearly equaling the wings; PODS narrowly elliptic, often somewhat asymmetrical, mostly 5-7 mm long; moderately pubescent; CLEISTOG- AMOUS PODS common, elliptie, often somewhat asymmetrical, mostly 4-5 mm long, moderately pubescent; SEEDS greenish-yellow to black. On upland prairies, sand dunes, old fields, dry roadsides, and well-drained, disturbed habitats throughout most of its range; in wet savannahs, open flood plains, and moist ditches along roads and railroads on the Atlantic Coastal Plain and distributed from Maine to northern Minnesota, central Kan- 372 Rhodora [Vol. 68 sas, and the Texas Panhandle south to southeastern Georgia, western Florida, and southeastern Texas. Common in west- ern prairies and in glaciated regions, less common on the Atlantic Coastal Plain, uncommon on the Appalachian Plateau (Fig. 3). Flowers from early July until mid October, mostly in August and September. The type specimen is mounted on the same sheet with plants of Lespedeza angustifolia and intermediates, prob- ably hybrids, between L. angustifolia and either L. capitata or L. hirta (see photograph of this sheet in Fassett, 1939). Fernald (1941) designated the shoot to the extreme right as the holotype of L. capitata. As Lespedeza capitata is highly polymorphic, all of the synonyms published since 1835 reflect attempts to delimit various segregates of the species. The indument, the shape of the leaflet, and the length of the peduncle in relationship to the length of the subtending leaf are three of the more conspicuously variable characters. Fassett (1939, p. 102) planted seeds from different varieties of L. capitata and obtained offspring which segregated widely in pubescence and leaflet shape from both their parental plants and their sibs. He found that there would be eight morphological races (varieties) of L. capitata in Wisconsin by using the same characters of pubescence and leaflet-shape that were used in describing the four varieties recognized by Fernald. Pierce (1939) planted seeds from cleistogamously formed pods from two widely divergent specimens of L. capitata and obtained considerable variation among each of their progenies. Little sense can be made of this variation until several hundred specimens of Lespedeza capitata from all parts of its geographic range are amassed for comparisons. After making such comparisons, two conclusions become evident: that populations of L. capitata near or above the southern limit of the Wisconsin glaciation are highly polymorphic and populations below this region are relatively uniform; and that certain forms of L. capitata are much more char- 1966] Lespedeza — Clewell 373 acteristic of particular geographic regions than they are of other regions. This geographic variation is as follows: 1. Lespedeza capitata with nearly linear leaflets, taper- ing gradually from the middle to the base and apex, glabrous above, and stems with appressed trichomes (L. c. var. stenophylla Bissell & Fern.). This form occurs primarily from northeastern Indiana to eastern Iowa, south to south- western Illinois; however, individuals have been collected in Rhode Island, Virginia and elsewhere. 2. Lespedeza capitata with narrowly elliptic or oblong leaflets, sericeous above, and with stems densely pubescent with very short, nearly straight trichomes, giving a silvery or cinnamon-colored appearance. (L. c. var. capitata sensu Gl. ; var. sericea Hook., var. typica Fern.). This is the most common form and occurs frequently in all parts of the range of the species with the possible exception of the Appalachian Plateau. It is by far the most common type in western Iowa, South Dakota, Nebraska, Kansas, Oklahoma, and along the Atlantic Coastal Plain from North Carolina to Florida and westward to Louisiana. The populations from the South- east are strikingly uniform. There is a slight tendency for southeastern plants to have oblong leaflets, while midwest- ern plants of this form have narrowly elliptic leaflets. In Iowa and Illinois many plants have been collected that re- semble this form except that they have leaflets glabrous above. I have also observed a few populations in the South- east with glabrous leaflets. Many plants from Ontario have exceptionally long, appressed trichomes on the leaflets, and these plants are not quite as silvery as others of this form and appear velutinous. These plants from Ontario and others like them, especially from coastal New England, have been named L. velutina Bickn. (L. c. var. velutina (Bickn.) Fern.). Prostrate forms of what might otherwise be called var. velutina have been observed by F. C. MacKeever (pers. com.) on Nantucket Island, Massachusetts. Attempts to grow living plants of this prostrate form which were sent to me by MacKeever were unsuccessful. Intergrades be- tween plants that have been called var. capitata and var. 374 Rhodora [Vol. 68 stenophylla are named L. c. var. stenophylla f. argentea Fern. 9. Lespedeza capitata with (narrowly) elliptic leaflets, glabrous above, stems with variable pubescence, usually (short-) villous (L. c. var. vulgaris T. & G.). This is the most common form in the Appalachians; however, I have examined only a Zew specimens from the southern Appala- chians, and some of these have been anomalous forms with leaflets glabrous above. Intergradations between the various forms, the occur- rence of many anomalous forms, and the heterogeneity en- countered by Fassett (1939) and Pierce (1939) in progeny tests preclude the practicability of naming each form. De- limitation of forms would have to be arbitrary, a “waste basket" taxon would be necessary for anomalous forms, and identification would remain difficult for many plants. An excellent character separating L. capitata from all other American species has been overlooked by previous authors: the distinctly greater length of the rachis as com- pared to the length of its petiole, especially of the larger leaves. From the distribution of L. capitata (Fig. 3) it seems logical that the plants occurring in the northern part of the lower peninsula of Michigan arrived there recently via a post-glacial migration through Wisconsin and the upper peninsula of Michigan, not northward through southern Michigan. Heard (1962) reported a similar migration route for certain sphaeriid clams. Range extensions of L. capitata in Michigan in the future will be noteworthy. Lespedeza hirta (L.) Hornem. Hort. Reg. Bot. Hafn. 2: 699 (1815). STEMS erect or ascending, to 1.8 m tall, short-appressed-pubescent to long-villous; STIPULES filiform to subulate, 3-6 mm long: PETIOLES long, the terminal leaflets of the longer leaves being 1-2.5 times longer than the combined lengths of the petiole and rachis and mostly 2-4 times longer on leaves among the racemes; RACHISES of the longer terminal leaflets equaling or shorter than their petioles; TERMINAL LEAFLETS elliptic or (cb-)ovate, the longer ones up to 4 em long and (0.45-)0.5-0.8 times as wide, glabrous, appressed-pubescent, or pilose above; STIPELS rarely present; RACEMES open, mostly 1.5-4 times 1966] Lespedeza — Clewell 375 i S } LESPEDEZA LEPTOSTACHYA ) T » LESPEDEZA INTERMEDIA Figs. 2-7. Distribution by county of 6 taxa. Fig. 2, Lespedeza angustifolia; Fig. 3, L. capitata; Fig. 4, L. hirta subsp. hirta; Fig. 5, L. hirta subsp. curtissii; Fig. 6, L. intermedia; Fig. 7, L. leptostachya. 376 Rhodora [Vol. 68 longer than their subtending leaves and bearing up to 16-44 flowers on the terminal half of the peduncle; BRACTEOLES extending to, or slightly bevond, the sinuses of the calyx; PEDICELS 1-2 mm long; FLOWERS 5-8 mm long; CALYX 6-12 mm long, lobed about 0.75 of its length or more, about equaling or slightly exceeding the pod which is easily visible between the lobes; COROLLA white or cream-colored; WINGS about equaling the standard and longer than the keel; PODS ovate or elliptic, 5-8 mm long, moderately to densely appressed- pubescent or appressed-pilose; CLEISTOGAMOUS PODS ovate or elliptic, mostly 5-7 mm long, moderately to densely appressed-pubescent or appressed-pilose, their calyces about equaling the pods; SEEDS mostly black, sometimes tan. KEY TO SUBSPECIES Leaflets glabrous, strigose, or pilose above, not silvery; stems villous OV pilose RREEEMMMMMMMM subsp. hirta Leaflets silvery with fine, dense, appressed hairs; stems appressed- pubescent or densely short-pilose .......c.cc:ccccseceesseesseeeseees subsp. curtissii Lespedeza hirta (L.) Hornem. subsp. hirta Hedysarum hirtum L. Sp. Pl. 748 (1753). (T.: Clayton 510, Va. [LINN, microfiche!]). Lespedeza polystachya Michx. Fl. Bor.-Amer. 2: 71 (1803). Lespedeza hirta var. sparsiflora T. & G. Fl. N. Amer. 1: 368 (1840). Lespedeza hirta var. typica Schindl. Bot, Jahrb. 49: 623 (1913). Lespedeza hirta var. calycina Schindl. Bot. Jahrb. 49: 624 (1913). (T.: J. Reverchon s. 'n., Dallas Co., Tex., Aug., 1874 [Gu!]). Lespedeza capitata var. calycina Fern. Rhodora 43: 572 (1941). In open, well-drained old fields, roadsides, sand dunes, and borders of woods from southern Maine, southern Ontario, southern Michigan, northeastern and southern Illinois, southern Missouri, and central Oklahoma south to central Florida and eastern Texas; rare on sandy soils of the Atlantie Coastal Plain (Fig. 4). Flowers from (June) mid- August through October. Michaux cited Hedysarum hirtum L. as a synonym of L. polystachya, and his description and plate of L. poly- stachya are identifiable as L. hirta (L.) Hornem. Linnaeus (1753) cited Gronovius (1739, p. 173, Trifolium fruticosum hirsutum ete.) who, in turn, cited Clayton 510 which thus serves as the lectotype. Hornemann published Lespedezia hirta, incorrectly spelling the name of the genus but validly 1966] Lespedeza — Clewell 377 publishing the combination. Hornemann described the species and cited the description by Muhlenberg (in Will- denow 1803) who cited Hedysarum hirtum L. I have been unable to locate the type of Lespedeza hirta var. sparsiflora but have examined a photograph (GH) of it. This plant resembles a shade form of L. hirta or possibly the hybrid, L. hirta X violacea. Cleistogamous pods occur much less frequently in this subspecies than in Lespedeza hirta subsp. curtissii. Com- ments on variation within subsp. hirta are included in the discussion under subsp. curtissv. Lespedeza hirta subsp. curtissii Clewell Brittonia 16: 75 (1964). (T.: Clewell 532, sandy field along State hwy. 24, 4.5 mi. e. of Ocean, Carteret Co., N. C. 12 Sep. 1961 [IND!]). Lespedeza hirta sensu Ell. Sketch Bot. S. C. 2: 207 (1824); non (L.) Hornem. (1815). In wire grass — long leaf pine — turkey oak sand ridges, among sand pine of the Florida scrub, along roadsides and old fields in dry, sandy soils, and occasionally on loam in frequently burned-over pinelands from southeastern Vir- ginia south along the Atlantic Coastal Plain to central Florida and west to southeastern Alabama and adjacent Florida (Fig. 5). Flowers from mid-August through October. Plants of this coastal subspecies have been identified as Lespedeza hirta var. appressipilis Blake. The type of this variety has narrow leaflets, glabrous above, and is identi- fiable as L. hirta X angustifolia, and therefore the silvery plants of L. hirta from the coastal plain cannot be ascribed to it. Since the coastal plain plants of L. hirta are distinct morphologically, geographically, and ecologically from other plants of L. hirta, they were recognized as a subspecies (Clewell 1964a). Much of the apparent geographic overlap of these two subspecies (Figs. 4 & 5) reflects the irregular boundary between the Piedmont and the Atlantic Coastal Plain, as well as piedmont-like regions, such as the Talla- hassee Red Hills, surrounded by coastal plain sands. I have studied populations of L. hirta on the coastal plain from 378 Rhodora [Vol. 68 North Carolina to Louisiana and have discovered a few populations which include plants of both subspecies (Long Co., Ga., Clewell 923, 937, 939; Allendale Co., S. C., Clewell 863, 867, 868 (FSU)). These plants might owe their occur- rence to dispersal by man, or they may represent local genetic variation in a subspecies that is generally much more uniform on the coastal plain. Intergradation between the two subspecies may occur locally where their ranges meet ; however, they often remain distinct. On deposit at FSU is a folder with plants from many populations of Lespedeza hirta from near the common boundary of the ranges of these subspecies, mostly collected in North Carolina. The nature of the variation encountered near this boundary may be examined from these specimens. Some plants from the pine barrens of New Jersey appear to be intergrades between the subspecies. If this intergra- dation were more extensive or if the silvery, coastal plain plants occurred with frequency within the range of subsp. hirta, then the variation would parallel that existing in L. capitata, If this situation occurred, then the identification of too many plants would be arbitrary, and to recognize distinct taxa would be impractical. I have searched without success for plants of this sub- species near the Gulf Coast from Pensacola to New Orleans. I can distinguish seven geographie races of Lespedeza hirta, which are not distinct enough to be recognized nomen- claturally but which may indicate the directions of evolution within the species. The first two races comprise subsp. curtissii, the others subsp. hirta. l. Plants of subsp. curtissii from the Carolinas and north- ern Georgia have stems with erect trichomes, often with the tips of the trichomes bent upwards, and leaflets with trichomes longer than 0.25 mm above. 2. Plants from western Florida have stems appressed- pubescent and leaflets with trichomes mostly shorter than 0.25 mm above. Intergrades between this and the first race occur in southern Georgia and in northeastern and peninsular Florida. 1966] Lespedeza — Clewell 379 3. Plants roughly from within the Gray's Manual range (New England south to Virginia and west to northern Arkansas) usually have stems densely villous and leaf- lets moderately appressed-pubescent above. 4. Plants from the same range as race 3 less commonly have stems moderately villous and leaflets glabrous above. Seeds from a plant of race 4 yielded offspring representing both forms (Clewell 1964a). 5. Off the coast of Massachusetts on Nantucket and Martha's Vineyard Islands plants of L. hirta resemble those of race 3 but with denser, longer trichomes. On these islands other species of Lespedeza tend to have more pronounced induments than elsewhere. 6. In Texas, plants have induments approaching that of subsp. curtissii but are not as silvery. In addition the calyces tend to be longer than in other areas, and the calyx lobes are slightly broader, tending to have a cuspi- date, rather than an acute, apex. These plants have been recognized as L. hirta (or L. capitata) var. calycina. Fernald (1941, p. 579) discussed the typification of this name. 7. On the Piedmont of the Carolinas and Georgia and extending westward into nearly all of Alabama, Missis- sippi and southeastern Arkansas are plants with leaf- lets which are 10% narrower, on the average, (0.6 times as wide as long) and with shorter and denser trichomes on the stems than plants of races 3 and 4. This variation within Lespedeza hirta is correlated with geological provinces and is not clinal. Most of this varia- tion exists in the southern half of its range, while northern populations are uniform. This contrasts with L. capitata which is polymorphic in the North and uniform in the South. It seems plausible that geographic differentiation has oecurred to a noticeable degree in L. hirta, and the activities of man in dispersing seeds and in modifying habitats have caused minor amounts of intergradation to occur between these races. In contrast, L. capitata, which is just as poly- morphic as L. hirta, appears to be à species in which much 380 Rhodora [Vol. 68 less racial differentiation has occurred, or if considerable differentiation had occurred previously, then it was pri- marily in the North and was mostly obliterated by glacia- tion. Lespedeza intermedia (S. Wats.) Britt. Trans. N. Y. Acad. Sci. 12: 63 (1893). (T.: M. A. Howe s. n., Brattleboro, Windham Co., Vt., Sept., 1891 (GH!)). Hedysarum violaceum Muhl. in Willd. Sp, Pl. 3: 1195 (1803). Lespedeza sessiliflora Nutt. Gen. Amer. Pl. 2: 107 (1818). Lespedeza violacea var. sessiliflora G. Don, Gen. Syst. 2: 807 (1832), Lespedeza reticulata var. sessiliflora Maxim. Acta Horti Petrop. 2: 365 (1873). Lespedeza reticulata S. Wats. Bibliog. Index 233 (1878), non Pers. 1807. Lespedeza stuevei var. intermedia S. Wats. in Gray, Man, Ed. 6 141 (1890). Lespedeza frutescens Britt. Mem. Torrey Bot. Club 5: 205 (1894). Lespedeza virginica var. sessiliflora Schindl. Bot. Jahrb. 49: 614 (1913). Lespedeza frutescens var, acutifructa Farwell, Rep. Mich. Acad. Sci. 22: 184 (1921). STEMS erect or ascending, to about 1 m tall, appressed-pubescent ; STIPULES filiform to narrowly subulate, mostly 2-4 mm long; PETIOLES long, the terminal leaflets of the larger leaves being 1-2.5 times longer than the combined lengths of the petiole and rachis and mostly 1-5 times longer on leaves among the racemes; RACHISES of longer termi- nal leaflets shorter than their petioles, often by 2 times; TERMINAL LEAFLETS oblong, elliptic, or narrowly ovate, the longer up to 4 cm. long and 0.3-0.55 times as wide, glabrous above or, rarely, glabrate along the midvein; RACEMES included or exserted up to 1.5 times beyond their subtending leaves and bearing 4-10 flowers; BRACTEOLES extending up to the sinuses of the calyx; PEDICELS 2-3 mm long; FLOWERS 5-7 mm long; CALYX 3-4.5 mm long and 0.3-0.5 times the length of the pod, lobed about half of its length; COROLLA purple; WINGS at anthesis (nearly) equaling the standard and distinctly longer than the keel; Pops ovate or elliptic, 5-7 mm long, short- appressed-pubescent; CLEISTOGAMOUS PODS commonly present, some- what orbicular, 2.5-5(-6) mm long, the calyx being 1-2.5 mm long, mostly 0.2-0.33 times longer than the pod. Dry roadsides, old fields, sand dunes, and borders of woods from soutnern Maine, southern Ontario, northern Indiana, central Missouri, and eastern Oklahoma south to 1966] Lespedeza — Clewell 381 Georgia, western Florida, and eastern Louisiana with dis- junctions in southeastern Minnesota, northwestern Illinois, and central Texas (Fig. 6). Flowers from late J uly through early October. Lespedeza intermedia has long been called L. frutescens (L.) Britt., based on Hedysarum frutescens L. (T.: Clayton 174). Schindler (1913) identified Clayton 174 as what is now known as L. violacea on the basis of its long keel. Blake (1924) recognized the name, L. intermedia, and pursued its nomenclatural history. I agree with his position and will not be able to elaborate further on the typification of this species until such time as I am able to examine Clayton 174. Both from the description and a photograph of the type of Lespedeza frutescens var. acutifructa I am unable to detect any differences between it and typical L. intermedia. Lespedeza intermedia is remarkable for its lack of geo- graphical variation. A study of phenotypic plasticity in L. intermedia by means of reciprocal transplants showed that the size of the leaflets and the length of the internodes vary considerably according to the degree of shading but that pubescence and the floral characters are stable (Clewell 1964b). Leaflet shape may vary nominally but not to the point that it be- comes significant in separating species. An understanding of this plasticity is important for accurate identification of species. From observations in the field and from compari- sons of field- and greenhouse-grown shoots from the same rhizome, other species exhibit similar plasticity. I have examined a number of specimens which had been mis- identified, evidently because they were shade forms of L. virginica, that superficially resembled plants of L. inter- media, or because they were shade forms of L. intermedia that resembled fruiting plants of L. violacea. Chasmogamous and cleistogamous pods of L. intermedia can be compared in Fig. 1, a and b. I have yet to see a specimen of, or visit a population of, L. intermedia in sandy pinelands in the Southeast. It is 382 Rhodora [Vol. 68 uncommon in the Southeast except in the Blue Ridge and on the adjacent Piedmont. Lespedeza leptostachya Engelm. in A. Gray, Proc. Amer. Acad. 12: 57 (1876). (T.: C. E. Bessey s. m., Emmet Co., Iowa, 1871 [Mo 146617!]). sTEMS erect or ascending, to about 1 m tall, appressed- pubescent; STIPULES filiform to narrowly subulate, up to 5 mm long; PETIOLES short, the terminal leaflets of the larger leaves mostly 1-4 times longer than the combined lengths of the petiole and rachis and at least 4 times longer on leaves among the racemes; RACHISES of the longer leaves shorter than their petioles, often 4 times shorter; TERMINAL LEAFLETS linear or narrowly oblong, up to 3.5 em long and 0.15-0.25 times as wide, appressed-pubescent above; RACEMES open, mostly 1.5-4 times longer than their subtending leaves, bearing up to about 16-30 flowers; FLOWERS 5-6 mm long; BRACTEOLES equaling or exceeding the bases of the lobes of the calyx; CALYX 4.5-5 mm long, nearly chorisepalous; COROLLA white to light purple; WINGS at anthesis equaling or slightly shorter than the standard and longer than the keel; CLEISTOGAMOUS PODS commonly present in open racemes, nearly orbicular, 3-4 mm long, densely pubescent, exceeded by the calyx, the calyx 4-6 mm long, nearly chorisepalous, the lobes very narrow, bracteoles greatly exceeding the base of the lobes of the calyx; SEEDS greenish-yellow to dark tan. In upland prairies on gravelly moraines from central Minnesota south to northeastern Illinois and northwestern Iowa (Fig. 7). Flowers from late July through mid Septem- ber. Extant specimens of Lespedeza leptostachya mostly were collected before 1900, thereby reflecting the intolerance of this species to plowing and grazing. In 1964 I searched for it for two days in the northern tier of counties in lowa along railroad tracks, in unplowed strips between fields, and in other prairie remnants, including where it had been collected previously. I discovered only one population, that being on a moraine of black, gravelly soil in Cayler Prairie, a preserved virgin prairie about 3 miles west of Lake Okoboji, Dickinson Co., Iowa. Roughly 500 plants grew in a three acre area, and a few plants were scattered else- where. Lespedeza capitata was also present. Recent plowing of remnant virgin tracts by farmers interested in capital- izing on the Federal Soil Bank Program probably has con- tributed to the pending extinction of L. leptostachya. 1966] Lespedeza — Clewell 383 Gleason and Cronquist (1964, p. 197) reported L. lepto- stachya near Hawarden, Sioux Co., Iowa. In addition to the two collections from Wisconsin (Fig. 7) Fassett (1939) reported seeing specimens collected in the 1860's from Grant and La Crosse counties. He searched all four localities without discovering plants of L. leptostachya. He also saw specimens which I have not seen from northwestern Illinois and from southeastern Minnesota. Steyermark and Swink (1955) reported a recent collection from a dry, gravelly prairie in Cook Co., Illinois. Most collections have cleistogamous pods. This degree of self-pollination helps explain the lack of variation between individuals. See the comments under Lespedeza angustifolia for a comparison with that species. Lespedeza procumbens Michx. Fl. Bor.-Amer. 2: 70 (1808). (T: A. Michaux [P]). STEMS procumbent, to about 1.8 m long, short-pilose to nearly villous, usually dense; STIPULES filiform to narrowly subulate, 2-4(-5) mm long; PETIOLES long, the terminal leaflets of the larger leaves mostly 0.5-1.5 times longer than the combined lengths of their petioles and rachises and mostly 1.5-5 times longer on leaves among the racemes; RACHIS of the longer terminal leaflets equaling or shorter than their petioles; TERMINAL LEAFLETS ovate, sometimes elliptic or oblong, the longer ones up to 2.5 cm long and 0.45-0.7 times as wide, sparsely appressed-pubescent, pilose, or rarely densely pilose above; RACEMES mostly 2-7 times longer than their subtending leaves, occasionally 2 per node, bearing up to 8-12(-14) flowers scattered along the terminal half of the peduncle; PEDUNCLES pilose; PEDICELS 0.5-1.5 (-2) mm long; FLOWERS 5-9 mm long; BRACTEOLES extending about to the base of the lobes of the calyx; CALYX 2-4 mm long and extending (nearly) half the length of the pod, lobed 0.5-0.7 of its length; COROLLA purple; WINGS at anthesis about as long as the keel and shorter than the. standard; STANDARD equaling or slightly longer than the keel; PODS elliptic, about 5 mm long, short appressed-pubescent to nearly glabrous; CLEISTOGAMOUS PODS commonly present, in axillary clusters or clustered near the apexes of peduncles, elliptic to orbicular, 3.5-5 mm long, mostly sessile, their calyces 1-2 mm long and 0.2-0.3 times the length of the pod; SEEDS greenish-yellow to dark tan. Dry roadsides, old fields, and open woods from southern New England, central Illinois, and central Oklahoma south to Georgia, western Florida, and eastern Texas (Fig. 8). [Vol. 68 Rhodora 2 * 1, LI MN H $. > * .. c $$. e "s A B * ul Lu $.. 2d HA © o wee ,ee * v . Q A et^ . { d . o sa . eo. i . e? * LRL Se v 3 See ^. fe . e ° et. fee ° Ye Y .* “wee Hist o’ j ; . 8 N ] v jV À | ex / . b J ^ me . . es J ESPEDEZA REPENS In "e 1 eo, .f ( d ..*5 < . s. * . . . " l . BA Ss. $ n "o ee ^ egdeee J {e ° PM ' $2" . E vl han LAO . ee Kj * * . e QUEE. lá E e e 9. ; . eve fo Aest NO 4 «gees A O LEA ej 9.9 t. ee ae 9 | e "e P oe LEELEE T *9 on "n à ene ve i . n te /. e° M ^, ei EM * V, da aè «ser, we st ^" "v. . N lind x ~ 10 v Figs. 8-10. Distribution by county of 3 taxa. Fig. 8, Lespedeza procumbens; Fig. 9, L. repens; Fig. 10, L. stuevei. 1966] Lespedeza — Clewell 385 Flowers from (mid-June) late August through September. Mlle. A. Lourteig kindly compared a typical plant of L. procumbens from Brown Co., Indiana, with the type in the Michaux Herbarium (P) and wrote me that, “it matches perfectly." She said it did not match a plant of typical L. repens which I had also included. On the Atlantic Coastal Plain I have seen L. procumbens only on clays and loams, not on soils of pure sand. I have not detected any geo- graphie variation in L. procumbens. There is close resemblance between Lespedeza procumbens and L. repens. While L. procumbens has stems and peduncles pilose, typically 8-12 flowers per raceme, and cleistog- amous pods either pedunculate or axillary, L. repens has stems and peduncles sparsely short-appressed-pubescent, typically 4-8 flowers per raceme, and cleistogamous pods commonly pedunculate. Pubescence alone is quite sufficient in distinguishing them. Lespedeza repens (L.) Bart. Prod. Fl. Phil. 2: 77 (1817) in part. (T.: Clayton 85, Va. [BM]). Hedysarum repens L, Sp. Pl. 749 (1753). Hedysarum prostratum Muhl. in Willld. Sp. Pl. 3: 1200 (1803). Lespedeza prostrata Nutt. Gen. Amer. PI. 2: 107 (1818). STEMS procumbent or sometimes ascending when young, to about 1 m long, sparsely short-appressed-pubescent; STIPULES subulate, 1.5-3 (-4) mm long; LEAVES generally of the same size on a stem, gradually becoming smaller towards the tip with an occasional axillary bud expanding into one very small leaf; PETIOLES long, the terminal leaflets of the larger leaves being mostly 1-4 times longer than the combined lengths of their petioles and rachises and mostly 2-5 times longer on leaves among the racemes; RACHISES of the longer terminal leaflets equaling or shorter than their petioles; TERMINAL LEAFLETS mem- branous, elliptic to (ob-)ovate, the longer ones up to 2.5 em long and 0.33-0.8 (mostly 0.45-0.7) times as wide, glabrous or sparsely ap- pressed-pubescent above, green in color below; RACEMES mostly 2-7 times longer than their subtending leaves, sometimes 2 per node, bearing 4-8(-12) flowers; PEDUNCLES filiform, appressed-pubescent ; PEDICELS 0.5-3(-4) mm long; BRACTEOLES extending up to the base of the lobes of the calyx; FLOWERS 5-7(-8) mm long; CALYX 2-4 mm long, lobed about half its length and extending (nearly) half the length of the mature pod; COROLLA purple; WINGS about equaling or shorter than the keel; STANDARD equaling or slightly exceeding the keel; PODS ovate or orbicular, sometimes slightly asymmetrical, 4-5 mm long, 386 Rhodora [Vol. 68 sparsely short-appressed-pubescent; CLEISTOGAMOUS PODS commonly present, orbicular or occasionally somewhat elliptical, rarely asym- metrical, 3-4(-5) mm long, mostly pedunculate, sometimes a few pods in axillary clusters, their calyces being up to 0.25-0.5 times the length of the mature pod; SEEDS greenish-yellow. Dry roadsides, old fields, borders of woods, and sandy pinelands from Long Island, Ohio, central Missouri and south-central Kansas south to northern Florida and eastern Texas, also along the Mississippi River to southwestern Wisconsin (Fig. 9). Flowers from (mid-April) late May through October. The lectotype of Lespedeza repens, Clayton 85 (BM), was cited by Gronovius (1739, p. 86) as Hedysarum caulibus procumbentibus, etc. Gronovius also cited a plant from Ceylon under this name by referring to a plate in Dillenius' Hortus Elthamensis. The plant from Ceylon is not a Lespedeza but might be an Alysicarpus (Robson, pers. com.) or a Desmodium (Britton 1893). Robson, who kindly examined Clayton 85 for me, questioned the name, Lespe- deza (Hedysarum) repens, because Linnaeus' description matches more closely the plant from Ceylon than Clayton 85 which lacks obcordate leaflets described by Linnaeus. Two possibilities may explain this discrepancy; over the years the lower leaflets, which are often the only obcordate ones in L. repens, were lost from Clayton 85, or Linnaeus based his description on additional plants that he did not cite, such as 921-42 in the Linnaean Herbarium. Since Linnaeus re- ferred to Clayton 85 by citing Gronovius and since he described H. repens as occurring only in Virginia, then I recognize the Linnaean name as legitimate. Britton (1893) and Schindler (1913, p. 593) both examined Clayton 85 and recognized it as the lectotype. Plants of Lespedeza repens in the southeastern states often bloom in the late spring. These plants may have at least some of their stems ascending while in flower. Later in the year continued growth of these same stems will pro- vide sufficient weight to make them lie procumbent. Other- wise, I have detected no geographic variation in L. repens; however, I have seen an occasional specimen from the lower 1966] Lespedeza — Clewell 387 Mississippi Valley with fruits resembling those of L. vio- lacea in size and shape. Plants of Lespedeza repens are easily distinguished from L. violacea in the field; however, pressed specimens of these species in fruit are often misidentified. The most reliable character in separating such specimens is the ratio between the length of the mature, cleistogamous pod and its sub- tending calyx. In L. repens (Fig. 1e) the calyx extends at least a quarter of the way up the pod (not including the style). In L. violacea (Fig. 1d) the calyx reaches up to two tenths of the way up the pod, usually much less. Other char- acters are: length of stipules — 1.5-3 mm in L. repens, up to 2.5.6 mm long in L. violacea; cleistogamous pods — mostly pedunculate, 3-4 mm long, and symmetrical in L. repens, mostly axillary, 3-7 mm long, and often asymmetrical in L. violacea; leaves generally of one size, though gradually be- coming somewhat shorter towards the apex of the stem in L. repens, leaves of two or more distinct sizes, attributable to one or few smaller leaves of different sizes occurring in the axils of the larger leaves in L. violacea. Lespedeza stuevei Nutt. Gen. Amer. Pl. 2: 107 (1818). (T: W. Stüve [PH.]). STEMS erect or ascending, to about 1.8 m tall, densely appressed- pubescent or pilose; STIPULES filiform to narrowly subulate, up to 5 mm long; PETIOLES long, the terminal leaflets of the larger leaves being 0.5-2 times longer than the combined lengths of the petiole and rachis and mostly 1-5 times longer on leaves among the racemes; RACHISES of the longer terminal leaflets shorter than their petioles; TERMINAL LEAFLETS elliptic or oblong, the longer ones up to 4 cm long and 0.3-.55 times as wide, uniformly appressed-pubescent, occasionally pilose, rarely sparsely appressed-pubescent along the midvein above; RACEMES included or exserted up to 1.5 times beyond their subtending leaves and bearing up to (6-)8-14 flowers; PEDICELS up to 2 mm long; BRACTEOLES extending about to the base of the lobes of the calyx; FLOWERS 4.5-6.5 mm long; CALYX 3-5 mm long, extending (nearly) half the length of the pod; lobed about 0.5-0.6 of its length, COROLLA purple; WINGS at anthesis (nearly) equaling the standard and distinctly longer than the keel; PODS ovate or elliptic, mostly 5-6 mm long, sparsely short-appressed-pubescent to densely appressed-pilose; CLEIS- TOGAMOUS PODS commonly present, ovate or elliptic, mostly 4-6 mm long, sparsely short-appressed-pubescent to densely appressed-pilose, the calyx being 0.3-0.4 times the length of the pod. 388 Rhodora [Vol. 68 Dry roadsides and old fields, borders of dry woods, and open sandy pinelands along the Atlantic Coast from Massa- chusetts to North Carolina, then west to southern Indiana and eastern Kansas and south to northern Florida and central Texas (Fig. 10). Flowers from (June) August through October. Lespedeza stuevei was named for W. Stiive and was spelled L. stüvei by Nuttall (1818) and some subsequent authors. Article 73 of the International Code of Botanical Nomen- clature (Lanjouw 1961) necessitates the orthographic change to stuevei. N. K. B. Robson (pers. com.) stated, “Medicago caule erecto vix ramoso, racemo dense spicato terminatrice (Gronov., Fl. Virgin.: 86) appears to belong to this species, according to the Clayton specimen in our Herbarium (BM). Linnaeus did not take it up in Sp. Pl." Geographie variation within Lespedeza stuevei is pro- nounced and recognition of two subspecies is perhaps war- ranted. Plants from the Ozark Plateau and from along the Atlantie coast above North Carolina have long trichomes. The stems of these plants are often pilose, and the other- wise brown pods at maturity are often hoary with appressed trichomes. Plants from much of Texas and from the south- eastern states, particularly in dry pinelands, have excep- tionally fine, short, appressed trichomes which, though dense, may barely be discerned without a lens. Specimens of these southeastern plants sometimes have been mistaken for L. intermedia, I have seen two nearly glabrous plants of L. stuevei: Kral 8159 (LAF) and Cocks in 1900 (NO) from Jackson and Rapides Parishes, respectively, Louisiana. Plants of Lespedeza stuevei or L. virginica with the larger terminal leaflets about 0.3 times as wide as broad must be identified arbitrarily. Most collections of such plants are probably hybrids. I have examined many populations in which both species were present, and in nearly all popula- tions the great majority of plants can be identified easily to one or the other species. Undoubtedly these species are closely related, both phenetically and genetically, but there is little reason to consider lumping them. 1966] Lespedeza — Clewell 389 Lespedeza texana Britt. in Small, Fl. S. E U. & 64t (t908): (T.: A. Arthur Heller 1914, Kerrville, Kerr Co., Tex., 19-26 June 1894 [NY!]). STEMS erect up to about 1.5 dm, the more distal portions procumbent, up to 1.5 m long, appressed-pubescent to short-pilose; STIPULES fili- form, the longer ones (3-)4-8 mm long; PETIOLES long, the terminal leaflets of the larger leaves being 1-3 times longer than the combined lengths of their petioles and rachises and mostly 2-5 times longer on leaves among the racemes; RACHISES of the longer terminal leaflets mostly shorter than, or about equaling, their petioles; TERMINAL LEAFLETS firm-textured, elliptic, the longer ones up to 3.5 em long and 0.35-0.55 times as wide, glabrous, glabrate, or sometimes short-pilose, above, gray-green to white below; RACEMES mostly 2-7 times longer than their subtending leaves, sometimes 2 or 3 per node, bearing 4-8(-10) flowers; PEDUNCLES appressed-pubescent or short-pilose; PEDICELS (1-)3-5 mm long; BRACTEOLES extending to the base of the lobes of the calyx; FLOWERS 5.5-8.5 mm long; CALYX 2-4 mm long, and extending (nearly) half the length of the pod, lobed about half its length; COROLLA purple; WINGS shorter than or about equaling the keel; STANDARD equaling the keel; Pops ovate, 6-7 mm long; sparsely short-appressed-pubescent; CLEISTOGAMOUS PODS commonly present, 4-6.5 mm long, ovate, sometimes asymmetrical, axillary or peduncu- late, their calyces being 0.1-0.3 times the length of the pod. In thin, rocky soil of scarp woodlands on or near the Edwards Plateau in central Texas; disjunction in eastern Coahuila, Mexico (Fig. 11). Flowers from mid-June through mid-October. I did not recognize Lespedeza texana as a species until I saw living plants of it in natural populations. Its most distinctive characters are not preserved in dried specimens. Its long stipules, up to 4-8 mm, often break off when dry. The somewhat coriaceous texture to the leaflets is easily overlooked on mounted specimens. The greenish-gray to white undersurface of the leaflets contrasts with the green undersurfaces in related species; this color escapes attention or is disregarded until seen in the field. The habit, which is entirely unlike that of any other American species, is the most striking character in the field, yet it cannot be recog- nized from pressed specimens. The stems are erect up to about 15 em high, the next 10 cm are arching, and the rest of each stem, which may extend over a meter in length, is trailing. Other distinctive characters are the long, cleistog- 390 Rhodora [Vol. 68 amous pods, 4-6.5 mm long (as compared to 3-4 mm long in L. repens), and the keel and wing petals which are about of equal lengths (in contrast to L. violacea). I have seen only one plant in flower, and it was greenhouse-grown. Its flowers were uniformly dark purple and were more deeply pigmented than the flowers of plants of the other species. Although I do not know Lespedeza texana in the field as well as I know the other species, I believe that L. texana tends to be less weedy than the other species and does not invade roadsides. It is essentially allopatric; I have collected L. procumbens with it in Robertson Co., Texas, and know of no other sympatric populations. The collection from Coahuila is Palmer 249 (PH) from Mt. Caracol, 21 miles southeast of Monclova. Lespedeza violacea (L.) Pers. Syn. 2: 318 (1807). (T.: Kalm [LINN 921-41]). Hedysarum frutescens L. Sp. Pl. 748 (1753), non Muhl. in Willd, 1803. Hedysarum violaceum L. Sp. Pl. 749 (1753). Hedysarum divergens Muhl. in Willd, Sp. Pl. 3: 1196 (1803). Lespedeza divergens Pursh, Fl. Amer. Sept. 2: 481 (1814). Lespedeza violacea var. divergens G. Don, Gen. Syst. 2: 307 (1832). Lespedeza violacea var. prairea Mackenz. & Bush, Trans, Acad. Sci. St. Louis 12: 14 (1902). (T.: B. F. Bush 93, common, Lee's Sum- mit, Jackson Co., Mo., 2 Sept. 1895 [Mo!, isotype Nv!]). Lespedeza prairea Britt. in Small, Fl, S. E. U. S. 641 (1903). STEMS erect to weakly ascending, to about 0.7 m tall, sparsely ap- pressed-pubescent; STIPULES filiform, up to 2.5-6 mm long; LEAVES gen- erally of several different sizes on a stem with axillary buds commonly expanding into single, smaller leaves of various sizes; PETIOLES long, terminal leaflets of the larger leaves mostly 0.5-1.5 times longer than the combined lengths of their petiole and rachis and mostly 1-4 times longer on leaves among the racemes; RACHISES of the longer leaflets shorter than their petioles; TERMINAL LEAFLETS membranous, elliptic or ovate, the longer ones up to 4.5 cm long and 0.35-0.7 times as wide, glabrous or sometimes glabrate above, light green or green below. RACEMES exserted up to 2-5 times beyond their subtending leaves or, uncommonly, all included within their subtending leaves, often 2 per node, bearing 4-6(-8) flowers; PEDUNCLES filiform, appressed-pubes- cent; PEDICELS 0.5-3 mm. long; BRACTEOLES extending about to the base of the lobes of the calyx; FLOWERS 5.5-9 mm long; CALYX 3-6 mm long, about half the length of the mature pod; lobed about 0.6 of its length, COROLLA purple; WINGS and STANDARD distinctly shorter than, 391 1966] Lespedeza — Clewell veg ^. i e Iz utet ee à ee e. . H oe i f SN . Siete ) ~ \ ` i ee , | TN MET AE b ved e. 4 (— 3 d Sa Srt y p LESPEDEZA TEXANA \ \ \ . 4 E Trisetum spicatum var. molle © NOVA SCOTIA seare 4 - ji B^ LAL; h At Al yO L 4 D X 22 k.. ^o Carex capillaris ó © ANN 3 Carex capillaris var. major (f Draba norvegica [n] Oxyria digyna ) Saxifraga Aizoon var. neogaea €) Saxifraga aizoides Fig. 4 C y. d C d T AT Tu ELO LU [- E Luzula spicata @ CT as a 3 | Phyllodoce n) Solidago multiradiata (D Diapensia lapponica Betula borealis €) Pinguicula vulgaris (9 Betula glandulosa Å Fig. 5 v a es eet xc a Kj a ee Aw igj p” y NOVA SCOTIA D ae m o hr E oe 3 9G OP a 4 à- "i + t X k.. P Vy Vaccinium uliginosum var. alpinum (f) OEN n Draba pycnosperma @ Arnica chionopappa © ori dn w Festuca prolifera var. lasiolepis (3 Fig. 6 412 Cystopteris fragilis var. laurentiana (f) Dryopteris fragrans var. remotiuscula Qe fw NOVA SCOTIA seare Polystichum Lonchitis o Cryptogramna Stelleri«Q Asplenium Trichomanes © 6 Fig. 7 Ls det hs "- M. P, PFa Ta NOVA SCOTIA Schizachne purpurascrns (f Festuca prolifera (3 Phleum alpinum @ Seirpus cespitosus var. delicatulus © ie seat Fig. 8 Zo arex atratiformis © Draba arabisans @ Arabis hirsuta var. pycnocarpa @ Arabis Drummondi © 413 Epilobium Hornemanni (f) Primula mistassinica Q Erigeron hyssopifolius o Achillea borealis © Betula Michauxii @ Pig. 10 NOVA SCOTIA 0 sene YH Senecio pauperculus o Luzula parviflora @ Luzula parviflora var. melanocarpa @ Betula pumila @ Juncus alpinus var. rariflorus © Tofieldia glutinosa Parnassia parviflora d Fig: 11 Oxytropio jJohannensis ® Vaccinium cespítosum [-] Vaccinium ovalifolium o Vaccinium boreale © vid a NOVA SCOTIA seare 414 1966] Flora of Nova Scotia — Hounsell and Smith 415 region of Kings county where they occupy exposed head- lands and such similar sites. The accompanying distribution maps (Figs. 1-12) show the bicentric distribution quite clearly. Each symbol, in most cases, represents several small stations, each of which is separated from the others of its group by distances ranging from a few hundred yards up to five miles; for example, a number of stations found along the cliffs of a river valley are represented here by a single distribution dot. The counties of Nova Scotia plus other pertinent geo- graphical features are shown in Fig. 1 along with the distribution pattern for five Arctic-alpine species. Of these five species all are found in the northern Cape Breton region (henceforth NCB) whereas three are known from the Minas Basin-Bay of Fundy region (henceforth MB-BF). None are found outside these two areas. When the overall Arctic-alpine picture is assessed one sees that of the 26 species dealt with in this survey (see Table 1, part A) 25 are found in NCB, nine in MB-BF and only 2 beyond these two centers. The two found outside these areas (as well as within them) are: Woodsia ilvensis (L.) R.Br., from Halifax and Lunenburg counties and Sedum Rosea (L.) Scop., which has been collected from Louis Head, Shelburne county. These two widespread Arctic-alpine species plus a third, of perhaps even wider distribution, Lycopodiwm Selago L., are shown in Fig. 2. Of these 26 species only one, Carex capillaris L., has been found in MB-BF but not in NCB. Most of these Arctic-alpines inhabit river gorge cliffs or exposed outcroppings and headlands but three are plants of the high barrens and bogs: Betula glandulosa Michx., B. borealis Spach and Vaccinium uliginosum L. var. alpi- num Bigel. An examination of the distributions of the 32 species classified as Boreal disjuncts and Gulf endemics (Figs. 7-12 and Table 1, part B) reveals that 31 of these are found in NCB, 14 in MB-BF and two only occur in other regions as wel. Asplenium Trichomanes L. has been reported from 416 Rhodora [Vol. 68 Guysborough Co. by Roland (1947) but since the exact location of the station is not known to the present authors à solid black dot has been placed in the centre of the county (Fig. 7). Betula Michauzii Spach grows in Guysborough Co. and on Brier Island, Digby Co. but not in NCB (Fig. 10). Brier Island represents the southwestern extremity of the elevated section of land that lies between the Annap- TABLE 1 Numbers of Arctic-alpine, Boreal disjunct and Gulf endemic stations in Nova Scotia (NCB — Northern Cape Breton, MB-BF — Minas Basin and Bay of Fundy) A. Arctic-alpine Species NCB MB-BF others total Lycopodium Selago L. 18 9 — 27 Woodsia ilvensis (L.) R. Br. 19 17 5 41 Woodsia alpina (Bolton) S. F. Gray 5 — — 5 Woodsia glabella R. Br. 6 1 — 7 Asplenium viride Huds. 6 2 — 8 Poa alpina L. ] — — 1 Poa glaucantha Gaudin 5 4 — 9 Trisetum spicatum (L.) Richter var. molle (Michx.) Beal 6 — — 6 Trisetum spicatum (L.) Richter var. pilosiglume Fern. 7 3 — 10 Carex scirpoidea Michx. 8. — — 8 Carex capillaris L. — 1 — 1 Carex capillaris L. var. major Blytt 6 — — 6 Juncus trifidus L. 8 — — 3 Luzula spicata (L.) DC. 1 — — 1 Betula borealis Spach 4 — — 4 Betula glandulosa Michx. 1 — — 1 Oxyria digyna (L.) Hill 1 — — 1 Draba norvegica Gunn 2 — — 2 Sedum Rosea (L.) Scop. 7 7 5 19 Saxifraga aizoides L. 1 —. — 1 Saxifraga Aizoon Jacq. var. neogaea Butters 6 3 — 9 Phyllodoce caerulea (L.) Bab. 1 — — 1 Vaccinium uliginosum L. var. alpinum Bigel. 10 — — 10 Diapensia lapponica L. 1 — — 1 Pinguicula vulgaris L. 8 — — 8 Solidago multiradiata Ait. 4 — — 4 25/26 9/26 2/26 ——————————— 1966] Flora of Nova Scotia — Hounsell and Smith 41T B. Boreal Disjuncts, Gulf Endemics(*). Species NCB MB-BF others total Cystopteris fragilis (L.) Bernh. var. laurentiana Weath. 4 — — 4 Dryopteris fragrans (L.) Schott var. remotiuscula Komarov 5 5 — 10 Polystichum Lonchitis (L.) Roth 13 — — 13 Asplenium Trichomanes L. T5 3 1 19 Cryptogramma Stelleri (Gmel.) Prantl 2 — — 2 Schizachne purpurascens (Torr.) Swallen 11 i — 18 Festuca prolifera (Piper) Fern. 3 — — 3 Festuca prolifera (Piper) Fern. var. lasiolepsis Fern.* 2 — — 2 Phleum alpinum L. € — — 3 Scirpus cespitosus L. var. delicatulus Fern. 5 Z — — 5 Carex atratiformis Britt. 10 1 — 11 Juncus alpinus Vill. var. rariflorus Hartm. 1 — — 1 Luzula parviflora (Ehrh.) Desv. 12 4 — 16 Luzula parviflora (Ehrh.) Desv. var. melanocarpa (Michx.) Buchenau 1 — — 1 Tofieldia glutinosa (Michx.) Pers. 1 — — 1 Betula pumila L. 8 — — 8 Betula Michauxii Spach — 1 1 2 Draba pycnosperma Fern. & Knowlt.* 1 — — 1 Draba arabisans Michx. 5 5 — 10 Arabis hirsuta (L.) Scop. var. pyenocarpa (M. Hopkins) Rollins 2 3 — 5 Arabis Drummondi Gray 6 2 — 8 Parnassia parviflora DC. 2 — — 2 Oxytropis johannensis Fern. it 1 im 2 Epilobium Hornemanni Reichenb. 12 — — 12 Vaccinium cespitosum Michx. 3 1 — 4 Vaccinium ovalifolium Sm. 2 — — 2 Vaccinium boreale Hall & Aalders 3 — — 3 Primula mistassinica Michx. T 2 — 9 Erigeron hyssopifolius Michx. 10 5 — 15 Achillea borealis Bong. 11 4 — 15 Arnica chionopappa Fern.* 2 — — 2 Senecio pauperculus Michx. 2 — -— 2 | 31/32 14/32 2/3 Grand totals 56/58 23/58 4/58 418 Rhodora [Vol. 68 olis Valley and the Bay of Fundy. This elevated area, known as North Mountain, is about 500 feet high at Cape Blomidon at the Minas Basin end but becomes lower to the southwest so that at Brier Island the land is low, flat and boggy. Combining the Arctic-alpine and Boreal disjunct figures one sees that of the 58 species 56 have been collected in NCB but only 23 in MB-BF. This fact can be correlated with the greater number of suitable habitats in NCB, particularly damp, cool cliffs along the many river valleys and gorges. A following paper will present descriptions of these cliff- face stations, discussing the ecological implications. The following collections, used in this study, have not been reported previously : Poa alpina L. Cliff top, Ciboux Island, Victoria Co. Smith et al. 10958. First collection for province, Luzula spicata (L.) DC. Headlands at 1000 feet near Bay St. Law- rence, Victoria Co., H. Harries, 804. First collection for province. Pinguicula vulgaris L. Rare on moist ledges in sphagnum, cliff near mouth of Cheticamp River, Inverness Co., D. H. Webster, 626. Two plants in small crevices at river edge, Big Southwest Brook, Inver- ness Co., Smith, Pick and Hounsell, 22134. This species had pre- viously been collected only from St. Paul's Island by Perry and Roscoe, 352, in 1929. Arnica chionopappa Fern. The collection from cliff ledges, Big South- west Brook, Smith, et al. 6903, represents a second station for the province. The distribution maps were compiled from collections deposited in the Herbarium of Acadia University. Additional distributional in- formation was obtained from Roland (1947). PERRY BIOLOGICAL LABORATORIES, ACADIA UNIVERSITY, WOLFVILLE, NOVA SCOTIA. LITERATURE CITED ERSKINE, D. S. 1951. Species newly or rarely reported from Nova Scotia and Cape Breton Island. Rhodora 53: 264-270. FERNALD, M. L. 1950. Gray's Manual of Botany, American Book Co., Boston 1966] Flora of Nova Scotia — Hounsell and Smith 419 LawsoN, G. 1884. Notice of new and rare plants. Trans. N. S. Inst. Sci. 6: 68-75. Linpsay, A. W. H. 1887. A catalogue of the flora of Nova Scotia. Proc. N. S. Inst. Sci. 4: 184-222. RoLAND, A. E. 1947. Flora of Nova Scotia. Trans. N. S. Inst. Sci. 21: 97-642. ScHOFIELD, W. B. V. Results of exploration in Cumberland county. 301-310. Scoccan, H. J. 1950. The flora of Bic and the Gaspe Peninsula, Quebec. National Museum of Canada Bull. 115. SMITH, E. C. & W. B. SCHOFIELD. 1952. Contributions to the flora of Nova Scotia I and II. Rhodora 54: 220-228. & D. S. ERSKINE. 1954. Contributions to the flora of Nova Seotia IV. Rhodora 56: 242-252. 1955. Contributions to the flora of Nova Scotia: Rhodora 57: SOME ADDITIONS AND CORRECTIONS TO THE FLORA OF TEXAS — III DONOVAN S. CORRELL Additional data regarding the flora of Texas contained in this paper have been uncovered in the course of my work on a Manual of the vascular plants of Texas with Marshall C. Johnston. This project is being supported, in part, by a grant from National Science Foundation (GB-2409). All of the specimens cited, unless otherwise noted, are in the Lundell Herbarium (LL) of Texas Research Foundation. I wish to acknowledge the help of my wife and colleague, Dr. Helen B. Correll, in the preparation of this paper. Erythronium americanum Ker, Bot. Mag., Pl. 1113. 1808, TEXAS : Sabine County, abundant in the woods at Patroon Creek, campground about 10 miles north of Milam, flowers yellow, outer petals red-brown on the lower surface, [no date], Betty E. Lemmon 830. San Augustine County, rich hardwoods, 7 miles east of San Augustine, east of Apolo-gacho stream, Cousins’ farm, black ankle community, March 17, 1962, D. S. Correll & Helen, B. Correll 24829. As far as we know, this species is new to Texas. It has previously been reported to occur only as far southwest as Arkansas and Oklahoma. It differs from A. albidum Nutt. var. mesochorewm (Knerr) Rickett, east- and north-cen- tral Texas, in having yellowish instead of pinkish or bluish white flowers. Triphora trianthophora (Sw.) Rydb. var. Schaffneri Camp, Rhodora 42: 55-56. 1940. TEXAS: Smith County, low woods about 1 mile east of Hopewell Baptist Church at Swan, in clumps of 100 or more plants scattered, plants up to 3 dm. high, stems reddish purple, capsule erect, flowers pink, readily deciduous, October 20, 1965, D. S. Correll & Helen B. Correll 32034. This distinctive variant, distinguished from var, trian- thophora by its smaller flowers and erect, not pendent, fruits, is exceedingly abundant in a large tract of virgin forest in the above-cited locality in Texas. It is new to the flora of the state. 420 1966] Donovan S. Correll 421 Emex spinosa (L.) Campd., Mon. Rum. 58, tab. 1, fig. 1. 1819. Texas: Kleberg County, in sandy loam, old school ground, Riviera, flowers white, March 10, 1966, John R. Crutchfield 1090. This Mediterranean species has become introduced in various parts of the world. As far as we know, this is the first collection from Texas. Sesuvium erectum Correll, sp. nov. Herba perennis, typica erecta vel recto-patens, cum globulis crys- tallinis plus minusve velata, fusca vel fumea in siccitate; caules ad 5 dm. vel magis longi; folia oblanceolata vel spathulata vel aliquando lineari-oblonga, ad apicem obtusa, infra in basem amplectentem de- crescentia; flores plurimi, e foliorum et ramorum axillis orti, sessiles vel pedicellis ad 5 mm. longis; lobi calycis late elliptiei vel ovato- lanceolati, ad apicem obtusi vel subacuti, intra roseo-violacei, appen- dice subapicali dorsali apicem sepali aequanti vel superanti; styli 3 ad 5, comspicui, typice atri ubi exsiccati; stamina plurima; capsula ovoidea, ad apicem obtusa et truncata; semina turgida, atra, circa 1 mm. longa. Plants perennial, typically erect to erect-spreading or sometimes decumbent, more or less adorned throughout with erystalline globules, when dry brownish black to dark gray; stems up to 5 dm. or more long; leaves oblanceolate to spatulate or sometimes linear-oblong, ob- tuse at apex, tapered below to a clasping base, up to 4 cm. long and 8 mm. wide; flowers numerous, in the axils of leaves and branches, sessile or with pedicels rarely up to 5 mm. long; calyx-lobes broadly elliptic to ovate-lanceolate, obtuse to subacute at apex, rose-purple within, 4-8 mm. long, with a subapical dorsal appendage about equal to or greatly exceeding the sepal-apex; styles 3 to 5, conspicuous, typically black when dry; stamens numerous; capsule ovoid, blunt and truncate at apex, about 5 mm. long and 3 mm. in diameter; seeds black, plump, about 1 mm. long. TEXAS: Kenedy County, edge of pond, 25 miles south of Sarita, along U. S. rte. 77, July 12, 1957, D. S. Correll & I. M. Johnston 17868 (holotype, LL). Brooks County, in depression among sand hills near gypsum quarry, several miles southeast of Falfurrias, plant erect (only one plant seen), July 10, 1957, D. S. Correll & I. M. Johnston 17825 (paratype, LL). According to our interpretation of the complex genus Sesuvium, there are six species in Texas. One of these, S. verrucosum Raf., is found in the interior of west and north- west Texas, and, of special interest, when dry the plants are usually a light brown or tan-color. The other five taxa are essentially coastal with the exception of S. erectum which occurs northwestward along the Rio Grande into Trans- 422 Rhodora [Vol. 68 Pecos, Texas. Interestingly, all of these coastal plants, ex- cept S. trianthemoides, are dark gray or nearly black when dry. Of these taxa, S. trianthemoides and S. maritimum (Walt.) B.S.P., with their 5-staminate flowers, and S. Por- tulacastrum L., with its large, long-pediceled flowers, are quite distinctive. A small intricate plant, typically of coastal clay dunes and flats, which has sessile, narrowly linear-ob- long leaves and is densely covered with crystalline globules, is referred to S. sessile Pers. The remaining robust, erect to erect-spreading, typically more inland, plants are re- ferred to S. erectum. Sesuvium trianthemoides Correll, sp. nov. Herba annua, succulenta, fulva in siccitate, basiramifera, cum globulis dispersis crystallinis grandibus; caules ad 3.5 dm. vel magis longi, internodiis 4-5 cm. longis; folia opposita, oblanceolata vel spathulata, ad apicem cbtusa, minimum 3 em. longa (petiolo incluso) et 1 em. lata supra medium, infra in petiolum conspicuum attenuata; petiolus basi late scarioso-alatus et amplectens; flores solitarii, ses- siles, e foliorum et ramorum axillis orti; lobi calycis triangulo-ovati, ad apicem subacuti, valde nervati, circa 3.5 mm. longi, margine hyalino, appendice apicali dorsali parva; stamina 5, filamentis graci- libus cirea 1 mm. longis; ovarium ovoideum bicellulare; styli 2, circa 0.5 mm. longi; capsula ovoideo-ellipsoidea ad apicem aeuta, 4-5 mm. longa, circumscissilis; semina circa 10 in quaque capsula, cirea 1.5 mm. longa, conspicue rugosa cristis irregularibus ferrugineis gran- ularibus, plagis gilvis in ambitu vagis e hilo extensis. Plant annual, fleshy, brown when dry, branched from the base, the herbage with scattered large crystalline globules; stems up to 3.5 dm. long or more, the internodes 4-5 cm. long; leaves opposite, oblance- olate to spatulate, obtuse at apex, at least 3 cm. long (including the petiole) and 1 em. wide above the middle, tapered below into a con- spicuous petiole; petiole broadly scarious-winged at base and clasp- ing; flowers solitary in the axils of leaves and branches, sessile; calyx-lobes triangular-ovate, subacute at apex, strongly nerved, about 3.5 mm. long, with hyaline margins, the dorsal apical appendage small; stamens 5, with slender filaments about 1 mm. long; ovary ovoid, 2-celled; styles 2, about 0.5 mm. long; capsule ovoid-ellipsoid, pointed at apex, 4-5 mm. long, cireumscissile; seeds about 10 in each capsule, about 1.5 mm. long, conspieuously rugose with brownish granular irregular ridges, with additional light patches extended in irregular lines from the hilum. TEXAS: Kenedy County, dunes, July 11, 1947, B. C. Tharp 47431 (holotype, TEX). Similarly to S. maritimum, the flowers of this species 1966] Donovan S. Correll 423 have only 5 stamens. However, the conspicuously rugose, not smooth, seeds readily set it apart from that species. Also, as its name implies, it resembles very closely in habit and appearance the rather common Trianthema Portulacas- trum L. Oddly enough, its rugose seeds also resemble those of Trianthema. Paronychia Jamesii T. & G. var. praelongifolia Correll, var. nov. Varietate Jamesii similis, sed foliis calycem superantibus. Similar to var. Jamesii but with leaves exceeding the calyx. TEXAS: Brewster County, North Sunny Glenn, July 17, 1938, O. E. Sperry T683 (TAES); top of Old Baldy Mt., June 18, 1941, B. H. Warnock 20355. Childress County, Childress, May 1930, Biol. Class 18 (TEX). Crockett County, infrequent perennial, limestone soil, 15 miles north of Juno, alt. 1500 ft., June 7, 1957, B. H. Warnock & W. D. McBryde 15191. Culberson County, off highway U. S. 62, lime- stone foothills of Guadalupe Mountains, entrance to Guadalupe, per- ennial herb, August 8, 1945, C. L. Lundell & Amelia A. Lundell 14374 (holotype, LL); Beach Mts., 8 miles north of Van Horn, frequent on slopes and summit, June 3, 1938, H. C. Cutler 29644 (TAES). El Paso County, crevices of igneous rock, west slope of Franklin Mts., east of Canutillo, July 4, 1958, D. S. Correll & I. M. Johnston 19245. Howard County, Big Spring, May 24, 1945, V. L. Cory 49279. Potter County, along a small stream, 10 miles north of the Canadian River on Hy. #87, flowers yellow, June 2, 1957, D. S. Correll 16616. Tay- lor Ccunty, 18 miles southwest of View, May 27, 1943, W. L. Tolstead 7302 (TEX); San Angelo, common on gravelly arid soil, May 19, 1899, W. L. Bray 359 (TEX). Wilbarger County, 3-3/4 miles south of Vernon, May 12, 1934, H. D. Parks & V. L. Cory 13179 (TAES). The calyx of var. Jamesii usually exceeds or is equal to the subtending leaves, but the calyx of var. praelongif olia is exceeded by the leaves, often conspicuously so, and the plants are found mainly in west and northwest Texas. Cuphea carthagenensis (Jacq.) Maebr., Field Mus. Pub., Bot. 8: 124. 1930. Texas: Hardin County, 4.5 miles north of Sour Lake, forming a large colony on edge of low wet woods, flowers pink, October 10, 1965, D. S. Correll 31987. This species, which has been known from North Carolina and Louisiana, is new to Texas. It is quite different from C. petiolata (L.) Koehne that occurs from Oklahoma north- eastward in its sparsely hispid stems and branches, flowers half the size of that species and its much-abbreviated, gla- 424 Rhodora [Vol. 68 brous style in contrast to the long, pubescent style of C. petiolata, Frankenia Johnstonii Correll, sp. nov. Frutex ad 3 dm. altus, basi lignea a radicibus fuscis filo metallico similibus sustinenti et caules plures vel multos elongatos arcuatos vel valde recurvatos emittenti; planta omnis cana vel glaucescens; caules subherbacei, circa 1 mm. lati, sparsim pilosi, pilis brevibus albidis appressis vel incurvatis; internodia vulgo 1-1.5 em. longa; folia op- posita, certe petiolata, oblanceolata vel oblongo-elliptica, ad basim rotundata, ad apicem rotundatum minute apiculata, ad 13 mm. longa et 4 mm. lata, margine revoluto, supra pilosis appressis sparsim brevipilosa, infra sparsim farinosa et incana; petioli flavovirentes, 1-2 mm. longi, leviter alati, alis pubescentibus membranaceis circum nodum conjunctis; flores sessiles, solitarii, ad apicem ramulorum axil- larium; calycis tubus valde costatus, circa 4 mm. longus, 5 lobis triangularibus circa 2 mm. longis; petala 5, unguiculata, alba ellip- tico-oblanceolata, ad apicem erosa, calyce duplo longiora; stamina 6, inclusa longitudine varia; styli 3-fissi. Shrub up to 3 dm. high, the woody base supported by dark brown wiry reots and giving rise to several or many elongated arching or strongly recurved willowy stems, the entire plant grayish or bluish green; stems subherbaceous, about 1 mm. thick, sparsely pilose with short whitish appressed or incurved hairs; internodes mostly 1-1.5 cm. long; leaves opposite, distinctly petioled, oblanceolate to oblong- elliptie, rounded at base, minutely apieulate at the rounded apex, up to 13 mm, long and 4 mm. wide, the margins somewhat revolute, the upper surface sparsely short-pilose with appressed hairs, the lower surface farinaceous and hoary; petioles brownish-green, 1-2 mm. long, slightly winged with the membranous pubescent wings united around the node; flowers sessile and solitary at the apex of short axillary branchlets; calyx-tube strongly ribbed, about 4 mm. long, the 5 tri- angular lobes about 2 mm. long; petals 5, clawed, white, elliptic-ob- lanceolate, erose at the apex, about twice as long as the calyx; stamens 6, of various lengths, included; styles 3-cleft. TEXAS: Zapata County, rocky hillside 24.5 miles northeast of San Ygnacio on road to Aguilares, small shrub up to 1 foot high, flowers white, March 16, 1966, D. S. Correll 32254 (holotype, LL; isotypes, UC, GH, TEX, US). Starr County, on salt flats along Arroyo Los Olmos just east of El Sauz, spreading shrub 1 foot high, flowers white, March 17, 1966, D. S. Correll 22308 (paratypes, Ds, K, LL, S). This species is characterized by its elongated, strongly recurved stems with long internodes, and distinctly petioled leaves that are farinaceous on their lower surface. These characteristics readily distinguish it from the other four Species found in the United States and northern Mexico. 1966] Donovan S. Correll 425 It is à pleasure to name this distinctive species for my friend and colleague, Marshall C. Johnston, who kindly di- rected my attention to its uniqueness. Myriophyllum verticillatum L., Sp. Pl. 992. 1753. Texas: Bowie County, attached and floating in Club Lake about 3 miles west of New Boston, May 12, 1962, D. S. Correll & E. C. Ogden 25243. Harrison County, Crip's Camp, east of Uncertain, floating on Caddo Lake, May 10, 1962, D. S. Correll & E. C. Ogden 25213. Henderson County, in water at head of Big Lake (Preserve), Koon Kreek Klub, off rte. 19, 12 miles south of Athens, May 5, 1965, D. S. Correll, Helen B. Correll & John R. Crutchfield 30938. Rusk County, in water of Lake Cherokee, near Monroe off rte. 322 from Longview to Henderson, June 3, 1965, D. S. Correll & Helen B. Cor- rell 31094. Indefinite: “Texas,” E. Dapprich 7577 (SMU). This species is new to Texas. It is readily distinguished from M. heterophyllum Michx., which it superficially re- sembles, by having its floral bracts pinnately dissected or lobed instead of being entire or only merely toothed. Limonium limbatum Small var. glabrescens Correll, var. nov. Ramuli ultimi inflorescentiae elongati et aliquantum laxi; calycis costae glabrae vel pilis non nisi paucis sparsis. Ultimate branchlets of the inflorescence elongated and rather laxly flowered; calyx-ribs glabrous or with only a few scattered hairs. TEXAS: Reeves County, in saline soils just north of Pecos, flowers nearly white, July 27, 1957, D. S. Correll & I. M. Johnston 18564 (holotype, LL). The ultimate branchlets of the inflorescence in var. gla- brescens are somewhat elongated and more laxly flowered than in var. limbatum whose flowers are borne in dense ab- breviated clusters. The calyx-ribs of var. glabrescens are also glabrous or are provided with only a few scattered hairs instead of being densely pubescent as in var. limba- tum. Scrophularia marilandica L., Sp. Pl. 619 1753. Texas: Red River County, down Red River from Hy. 39 [37], first collection in Texas, August 25, 1948, H. B. Parks s. n.; On river terraces in open woods along the Red River above bridge, just west of Rte. 37, north of Negley, plants up to 7 ft. high, August 29, 1966, D. S. Correll 33564. This species has apparently never been reported from Texas. It was first collected by Parks too late to include in his and Cory's Catalogue of the Flora of Texas (Texas Agr. Exp. Sta. Bull. 550: 1937). Also, for some unknown reason, 426 Rhodora [Vol. 68 it was not included in Cory’s unpublished revision copies of the Catalogue. It was also omitted from Gould’s Texas Plants, published in 1962, Proboscidea sabulosa Correll, sp. nov. (Plate 1335) Herba annua, procumbens et ramosissima, ad circa 12 dm. lata et 4 dm. alta, omnino viscido-pubescens; folia petiolis incrassatis ad 10 cm. vel magis longis, triangulo-ovata vel late subreniformia, late cordata, apice rotundata vel obtusa, margine undulato-repando, ad ‘circa 12 em. longa et lata; flores plures, in racemo congesto super pedunculo abbreviato prodientes; inflorescentia folia brevicz; pedi- celli per anthesin graciles circa 1 cm. longi, maturitate crassi circa 2 cm. longi; bracteae 2, oblongo-ellipticae, 7-8 mm. longae; calyx 1-1.8 em. longus, supra medium irregulatim 5-lobatus, ad basim ven- traliter fissus; corolla tubulari-cylindracea, non nisi leviter ventri- cosa; corollae tubus extra cremeus et sparsim glandulosus, circa 2 cm. longus, orificio 1 em. lato; faux glabra, cremea, maculis parvis rubiginosis et linea crocea pro longitudine tubi facie inferiore ex- tensa; corollae lobi 5, ovati vel aliquantum quadrati, violacei, 4-5 mm. longi, recurvati, lobo infimo plicato; capsulae corpus oblongo- ellipsoideum, aliquantum laterale complanatum, endocarpio grosse sculpto, pagina dorsali carinatum, circa 7 cm. longum et 1.5-2 cm. latum, in cornu gracile recurvatum, longitudine corporis duplo longius protractum, Plant annual, sprawling and much-branched, up to about 12 dm. across and 4 dm. high, the herbage viscid-pubescent throughout; leaves with thickened petioles up to 10 em. long or more, triangular- ovate to broadly subreniform, broadly cordate, rounded to obtuse at apex, the margins undulate-repand, up to about 12 cm. long and broad; flowers several, borne in a congested raceme on an abbreviated peduncle, greatly exceeded by the foliage; pedicels slender and about l em. long in flower, becoming stout and about 2 cm. long in fruit; bracts 2, oblong-elliptic, 7-8 mm. long; calyx 1-1.3 em. long, irregularly 5-lobed above the middle, ventrally split to the base; corolla tubular- cylindrie, only slightly ventricose; corolla-tube cream-color and sparsely glandular without, about 2 cm. long and 1 cm. across the orifice, the glabrous throat cream-color with small reddish spots and à deep yellow line running the length of the tube on the lower side; corolla-lobes 5, ovate to somewhat quadrate, purplish red, 4-5 mm. long, recurved, the lower lobe pleated; fruit-body oblong-ellipsoid, somewhat laterally compressed, the endocarp coarsely sculptured, keeled on the dorsal surface, about 7 cm. long and 1.5-2 cm. thick, tapered into a slender recurved horn that is about twice as long as the body. TEXAS: Crane County, on shinnery oak dunes along Ranch Road #1233 about 1.5 miles west of U. S. Rte. 385, north of Crane, August 6, 1966, D. S. Correll 33328 (holotype, LL, isotypes, GH, TEX, UC, US); 1966] Donovan S. Correll 42'1 Plate 1335 PROBOSCIDEA SABULOSA Correll: Upper, fruits, about X 1/3; lower, portion of plant, note tiny flower in center of picture, about X 2, from Crane Co., Texas, Correll, no. 33328. 428 Rhodora [Vol. 68 in deep sand of dunes 11 miles northwest of Crane on Ranch Road #1601 to Penwell, August 6, 1966, D. S. Correll 33332 (paratype, LL). This is apparently the smallest flowered species in this genus, The abbreviated inflorescences are concealed by the foliage and it is necessary to search the plant in order to locate flowers. Because of its exceeding abundance in the dunes where it occurs in Crane County it is most unusual that this species has not come to light before now. Utricularia juncea Vahl, Enum. Pl. 1: 202, 1804. TEXAS: Angelina County, along marshy edge of brook, in pine land, off U. S. 63, near Beans Place, northwest of Jasper, September 8, 1942, C. L. Lundell 11797; boggy stream, 3 miles west of Bouton Lake, Angelina National Forest, June 6, 1963, D. S. Correll & D. C. Wasshausen 27522, Henderson County, on mud, boggy area above Withers Lake, Koon Kreek Klub, off Rte. 19, 12 miles south of Athens, May 5, 1965, D. S. Correll, Helen B. Correll & John R. Crutch- field 30945. This species is apparently new to Texas. Is resembles U. cornuta, Michx., but the flowers are about half the size of that species. Plantago insularis Eastw. var, fastigiata (Morris) Jepson, Man. FI. Pl. California 956. 1925. P. minima Cunn., Proc. Ind. Acad. Sci. 1896: 202. 1897. P. scariosa Morris, Bull. Torrey Bot. Club 27: 117. 1900. TEXAS: Brewster County, beds near Terlingua, Big Bend National Park, April 18, 1961, D. S. Correll & R. C. Rollins 23631; mouth of Tornillo Creek at Hot Springs on gravel bars, Big Bend National Park, April 22, 1966, D. S. Correll & R. C. Rollins 32642. Presidio County, on red and brownish black cobbly slope of small open can- yon, about 24.5 miles southeast of Redford, February 7, 1965, D. S. Correll 30682. This species, which heretofore has been known to occur from California to Utah and Arizona, is new to Texas. Its cottony leaves that grow in a dense clump and its broad scarious bracts are distinctive. My wife, who has been studying the genus Plantago, called my attention to this addition to the flora of Texas. TEXAS RESEARCH FOUNDATION RENNER, TEXAS CANDOLLEODENDRON, A NEW GENUS OF THE LEGUMINOSAE (CAESALPINIOIDEAE) RICHARD S. COWAN When I studied the type of Swartzia brachystachya A. P. DC. in the DeCandolle Herbarium in 1953, I doubted that the species could be left in the genus to which it had been assigned by DeCandolle, but I was reluctant to make that decision with no other material available for study. Then, two Froes collections were received in materials borrowed from the Instituto Agronómico do Norte at Belém, Brazil; there is no doubt whatever that DeCandolle’s type and these Brazilian collections are in all respects con- specific. I have chosen the name for the new genus to commemo- rate one of the most perceptive pioneers in the study of tropical legumes, Aug. Pyrame DeCandolle. Candolleodendron Cowan, gen, nov. A Swartzia antheris basifixis uniformibus, stigma uni- laterale stylo saepe curvato adapicem differt; a Zollernia alabastris globosis vel late ovalibus, petalo uno, foliis pinnatis, stigma unilaterale differt. TYPE SPECIES: Candolleodendron brachystachyum (A. P. DC.) Cowan. Candolleodendron brachystachyum (A. P. DeCandolle) Cowan, comb. nov. Swartzia brachystachya A. P. DeCandolle, Prod. 2: 423. 1825: Mem. Leg. 408. May 1826. Tree 3-7 m. tall, the branchlets glabrous and nitid, the stipules minute, triangular, ca. 0.4 mm. long and wide; leaves glabrous, pinnate, the petioles 4-6.5 em. long, terete, the rachis terete, slender, 7.5-10 cm. long; leaflets 2-jugate. strongly discolorous, much paler beneath, elliptic, the base acute, the apex acute to subacuminate, the venation plane above but salient beneath, the primary vein-branches strongly ascending; inflorescence 8-9 em. long, ramuliger- ous, the axis sparingly strigulose, the bracts persistent, triangular, ca. 0.4 mm. long and wide, minutely strigulose and ciliolate sparingly, the bracteoles lacking; pedicels 8-11 429 Figure 1.— a-e, Candolleodendron brachystachyum (a--d from Froes 32364, e from Froes 32975): a, leaf, X lo; b %, c, inner and outer faces of stamens, X 4; fruit, X 1. f, stamens of Bocoa Sp., X 4. X 3; h, stamens, X 4; i, gynoecium, X 4. , inflorescence, X d, gynoecium, X 4; e, g--i, Zollernia sp., g, bud, 1966] Candolleodendron — Cowan 431 mm. long, slender, sparingly strigulose, the buds globose to broadly oval in outline, sparingly strigulose, 9-11 mm. long and wide; calyx entire in the bud but opening in about four more or less equal segments, glabrous within, sparingly strigulose and irregularly striate externally, the striae brownish-red and longitudinally oriented ; petal one, yellow, glabrous, the claw ca. 1.5 mm. long, the blade oval, concave, ca. 15 mm. long and 10 mm. wide; stamens numerous, uni- form, glabrous, the filaments 1-2 mm. long, ligulate, the anthers basifixed, linear-lanceolate, acute to subacuminate, 5-6 mm. long, 0.6-0.8 mm. wide; gynoecium excepting the style sericeous, the stigma unilateral, obliquely oriented relative to the style, truncate, the style slightly uncinate or merely curved apically, 3.5-4.5 mm. long, glabrous, the ovary narrowly oblong, 3.5.4.5 mm. long, 1-1.5 mm. wide, the gynophore 0.4-0.8 mm. long; fruit (mature) oblong to oblong-oval, elliptic in cross-section, ca. 5.5 em. long, 2.5-3 cm. wide, glabrous but finely tesselate (drying?) and dis- tinctly verruculose (sublente), the carpophore ca. 2.5-3 mm. long, minutely strigulose; seeds not seen. TYPE: J. B. Patris s. n. (holotype G-DC), “Cayenne.” Distribution: Lowland soils above level of seasonal in- undation in French Guiana and along the lower reaches of tributaries of the Amazon River near its mouth. Additional specimens examined : BRAZIL: Pará: Municip. Porto do Móz: Left bank of Rio Xingü at Souzel, Froes 32354 (IAN) ; Portel, Rio Tapact, vicinity of the Rio Anapü, Froes 32975 (IAN). Earlier, in the course of my review of Swartzia and re- lated genera, I had questioned the supposed near relation- ship of Zollernia, as well as that of several other presumed relatives. The new genus combines characteristics of Swart- zia, Zollernia, and Bocoa; the latter shares with the new genus a common aspect of the leaves but there are obviously more characters in common with the other two relatives. The buds and petals of Candolleodendron are much like those of Swartzia but the basifixed, linear-lanceolate anthers and the form of the gynoecium are much more suggestive of Zollernia. Unlike either relative, Candolleodendron has 432 Rhodora [Vol. 68 the style apex abruptly hooked or curved, with the stigmatic surface unilaterally disposed. SMITHSONIAN INSTITUTION WASHINGTON, D. C. OBSERVATIONS REGARDING THE OCCURRENCE AND RELATIONSHIPS OF RUELLIA LORENTZIANA (ACANTHACEAE) IN SOUTHERN FLORIDA: For a number of years taxonomists interested in the flora of southern Florida have been aware of a Ruellia not de- scribed in Small's manual of the Flora of Southeastern United States. Thus far the plant is known only as a naturalized weed on the grounds of the University of Florida Subtropical Experiment Station and immediate environs in Homestead, but it appears to be spreading aggressively into neighboring fields, particularly those disturbed by agricul- ture. Identification of these plants proved difficult, and transplants were grown in the experimental garden of the University of South Florida in order that individuals could be observed over a period of time. Vigorous plants yielding abundant fertile seeds were produced (fig. 1). They were compared with transplants of other species of Ruellia and proved to be distinctly unlike any other taxon from eastern United States, but similar to transplants of R. occidentalis (A. Gray) Tharp & Barkley (R. nudiflora Urban var. occi- dentalis Leonard) from Texas. Preliminary crossing at- tempts indicated that the unknown plants were intersterile with eastern United States taxa such as R. caroliniensis (J. F. Gmel.) Steud. and R. succulenta Small, but cross compatible with R. occidentalis. "Contribution No. 23 from the Botanical Laboratories, University of South Florida. Aided by grant GB-1026 from the National Science Foundation. Plate 1336. Garden culture of Ruellia lorentziana. The following brief description based on garden cultures may serve to characterize the plants : Erect, simple or divergently branching near the base, up to 0.9 m. high, stems quadrangular and grooved, covered with pale cystoliths, glabrescent below to cglandular-strigose above especially at the nodes and on young leaves and branches, larger internodes 11.6-13.7 cm. 434 Rhodora [Vol. 68 long, larger leaves oblong-ovate to slightly cordate, blades 9.8-10.2 em. long, 5.8-6.1 cm. wide, with conspicuously darker veins, margins obscurely to irregularly serrate, petioles 2.5-2.7 em, long; inflorescence a narrow terminal panicle, densely glandular pubescent, sepals not keeled, corolla pale blue-violet, pubescent, capsule 2.2-2.4 cm. long, glandular pubescent, stalked, with 17-18 seeds. REPRESENTATIVE SPECIMEN: FLORIDA, DADE Co., Homestead, spreading as weed on grounds of Univ. of Florida Subtropical Experi- ment Station. June 8, 1962, R. W. Long 1130 (USF). Although this plant clearly belongs to the R. nudiflora-R. tuberosa L. complex, it is not easily assigned to any of the Species or varieties given by Leonard (Jour. Wash. Acad. Sci. 17:509-520, 1927). It is referred here provisionally to the Argentine species R. lorentziana Griseb., but it differs in having eglandular stems, sepals not keeled, and pubescent corollas, Lindau (In: Die natürlichen Pflanzenfamilien 4(3b): 274-354, 1895) places R. lorentziana in sect. Dipteracanthus along with numerous other South American taxa, but places R. tuberosa in sect. Euruellia. This illustrates the highly unsatisfactory state of our understanding of this complex group of species that includes R. tuberosa, R. nudiflora, R. intermedia Leonard, and R. lorentziana. ROBERT W. LONG DEPARTMENT OF BOTANY AND BACTERIOLOGY UNIVERSITY OF SOUTH FLORIDA, TAMPA, FLORIDA NOMENCLATURAL NOTES ON MONOCOTS EDWARD G. VOSS In the course of preparing a flora of the vascular plants of Michigan, I have accumulated considerable data relating to the names of the native and introduced species of (chiefly) the Great Lakes region. It seems well to present these data in advance of the flora and with more documen- tation than will be appropriate therein. Rather than publish a diverse series of short notes, cluttering the literature with a paragraph or two each time a discovery arose, I have preferred to present such notes at one time for a major group of plants (in this case, the Monocotyledones). This report may then cause less pain to bibliographers and may even be more useful as a reference list for other students of the flora. With the latter aim in mind, brief notes have been in- cluded on certain names for which good arguments have been published by others in the past — but too often and unjustifiably ignored. Thus, the present listing includes, in addition to some new combinations, most if not all of those instances where I consider the name of a local monocot in current manuals or monographs to require some clarifica- tion or replacement for largely nomenclatural (as distin- guished from taxonomic) reasons. Having previously discussed at some length (Taxon 14: 154-160. 1965) my interpretation of Article 46 of the Inter- national Code of Botanical Nomenclature and its plain requirement that the author publishing a name (1. e., sup- plying the description) be invariably cited (whether or not one takes the option of citing in addition a non-publishing author), I do not list below those instances in which only a change in author citation seems required, unless there are special reasons for discussing the case. The following list is organized under names, in boldface type, which I feel are correct, although a few of them are ones which for taxonomic reasons I choose not to employ at present. Comments will be welcomed on overlooked publica- 435 436 Rhodora [Vol. 68 tions, names, or interpretations which would alter the conclusions here set forth. ACKNOWLEDGMENTS Publications not available in Ann Arbor have been sought during several visits to the Missouri Botanical Garden, whose librarian, Dr. George Van Schaack, has been ever helpful. Every reference cited below has been examined by me, except for two specifically mentioned; not all refer- ences are cited, especially when a particular case has been thoroughly discussed and documented by others to whose work attention is called. For help on certain specific points, as acknowledged in the text or footnotes, I am indebted to F. J. Hermann, B. Peterson, A. E. Schuyler, and W. T. Stearn. Over the years Dr. Rogers McVaugh has patiently saved me from many (doubtless not all!) nomenclatural blunders and guided me in application of the Code; he has kindly read the manuscript of the present paper but is, of course, not to be blamed for any errors or rashness which it may yet contain. SPARGANIACEAE Sparganium chlorocarpum Rydb. f. acaule (Beeby) E. G. Voss, comb. nov, S. simplex var. acaule Beeby in Macoun, Cat. Canad. PI. 2(5): 367. 1890. The extreme state of this Species deserves, at most, recognition in formal rank. JUNCAGINACEAE Triglochin maritimum L. Triglochin palustre L. These are the only two species described by Linnaeus (Sp. PL, pp. 338, 339. 1753). Both epithets were given neuter form by him, the gender in accord with most Greek usage of the substantive adopted as the generic name. (Glochis, in contrast to triglochin, is chiefly feminine.) Since Linnaeus consistently employed the classical Greek gender for his genus, Triglochin, there seems to be no rea- 1966] Monocots — Voss 437 son at all, under Recommendation 73A(1) to alter it to feminine, as is often done in American publications (but not, e.g. in the treatment by Lóve & Lóve, Nat. Canad. 85: 156-165. 1958). HYDROCHARITACEAE Elodea Michx. As has been carefully pointed out by Shinners (Rhodora 58: 162. 1956), Elodea is correct without the conservation once proposed for it. (Cf. Art. 75 examples, where Elodea is cited as a different name not likely to be confused with Elodes Adans. Not only was the latter misspelled Elodea by Jussieu (Gen. Pl, p. 255. 1789) but it was also given by him in the synonymy of Hypericum, which would give it no nomenclatural standing even if it were different.) Continued use of Anacharis instead of Elodea is incorrect. GRAMINEAE 'Triticeae Dumort. As has been fully documented by the synonymy of Pilger (Bot. Jahrb. 76: 312. 1954), briefly summarized by Bowden (Canad. Jour. Bot. 37: 659, 1959), the tribal name Triticeae was validly published by Dumortier in 1823 (Obs. Gram. Belg., p. 82). Thus it has priority over Hordeae Benth. (Jour. Linn. Soc. Bot. 19: 31. 1881). Agropyron dasystachyum (Hook.) Scribn. var. psammophilum (Gil- lett & Senn) E. G. Voss, comb. nov. A. psammophilum Gillett & Senn. Canad. Jour. Bot. 39: 1170. 1961. Typical var. dasystachyum of the plains tends to have the foliage less glaucous, the lemmas less villous, and the glumes less attenuate than does this endemic variety of the Great Lakes shores (cf. Guire & Voss, Mich. Bot. 2: 107- 108. 1963). However, the differences do not seem to be as cleareut as implied by Gillett and Senn in describing the Great Lakes plant as a new species, and I prefer to consider the latter as a local variety in many ways paralleling Calamovilfa longifolia var. magna, Scribn. & Merrill. Agrostis stolonifera L. var. palustris (Huds.) Farw. This combination by Farwell (Rep. Mich. Acad. 21: 351. 438 Rhodora [Vol. 68 1920) should have been employed by Gleason (Ill. Fl. 1: 166. 1952) rather than var. compacta Hartman (Skand. FI. Handb., ed. 4, p. 24. 1843), since the former is based on the oldest epithet in varietal rank (A. polymorpha [var.] . palustris (Huds.) Huds., Fl. Angl., ed. 2, p. 32. 1778). Since Agrostis alba L. has been shown by Philipson (Jour. Linn. Soc. Bot. 51: 91. 1937) to be based on a species of Poa, the familiar “redtop” grass may be called A. sto/oni- fera var. major (Gaud.) Farw., as Gleason does in a con- servative treatment, or A. gigantea Roth, as does Fhilipson (op. cit., p. 90). (The name A. palustris has sometimes been misapplied to this species.) Fernald's treatment of this group in Man., ed. 8, is unique and cannot be correlated with other works. Agrostis hyemalis (Walt.) BSP. var. tenuis (Tuck.) Gleason f. seti- gera (Fern.) E. G. Voss, comb. nov. A. scabra var. septentrionalis Fern. f. setigera Fern. Rhodora 35: 210. 1933. A. scabra [var. scabra] f. tuckermani Fern. Ibid. 207. 1933. Errors and omissions in the synonymy of Agrostis scabra in Hitchcock’s Manual of Grasses (ed. 2, pp. 807, 808. 1951) help to confuse the nomenclatural picture, especially if one chooses to follow Gleason in considering this plant to repre- sent only a variety of A. hyemalis. From the synonymy in Man. Gr., one might conclude that Gleason did not transfer the oldest available epithet in varietal rank, for “Agrostis michauxii var. laxiflora A. Gray, N. Amer. Gram. and Cyp. 1: 17. 1834. Based on Trichodium laxiflorum Michx.” antedates Tuckerman’s epithet by nine years. However, Michaux's binomial (Fl. Bor.-Am. 1: 42. 1803) is illegitimate, as he lists Cornucopiae hyemalis (the basionym of A. hyemalis) as a synonym. Michaux's superfluous name (Art. 63) is thus a synonym of A. hyemalis and not (if two species are maintained) of A. scabra, as listed in Man. Gr. and in Ind. Gr, Sp. (Chase & Niles, 1962). Under Art. 72, Gray's varietal name is to be treated “as new," and not as a recombination of Michaux's illegitimate 1966] Monocots — Voss 439 epithet. Gray' provided no original description but cited the Michaux reference and figure, which is probably A. scabra. One might be able to interpret Gray's name as applying to the A. scabra concept; on the other hand, in the absence of a clear precedent in the Code, I consider it to fall in the same synonymy as Michaux's name and thus avoid the necessity of making yet another new combination. (Michaux's description is not clearly referable to either species, and because of his synonymy his type is the same as the type of A. hyemalis according to the controversial but often useful Art. 7, Note 4. Since the application of Michaux's name is governed by its type, and since Gray's name is validated only by reference to Michaux, Gray's type can be considered the same.) If all of “A. scabra" is treated as a variety of A. hyemalis (as var. tenuis), then a single epithet is in order for the awned form. Of Fernald's two available epithets of the same date, I choose f. setigera as being more descriptive. Neither Fernald's var. septentrionalis nor f. setigera is listed in the synonymy of any taxon in Man. Gr. (although both are in Ind. Gr. Sp., 1962). Andropogon gerardii Vitman Article 73, Note 3, of the Code states that “wrong use of the terminations i, ii, . . . is treated as an orthographic error,” but this does not completely solve the problem of what constitutes “wrong use.” Rec. 73C states that if a personal name “is already Latin or Greek, the appropriate Latin genitive should be used.” Andropogon gerardi has been defended on the grounds that there is a Latin form gerardus. But there is a Latin form for a great many names, and Rec. 73C does not refer to names which have a Latin form (i. e., have ever been Latinized by any- one) but to names which are “already Latin." I believe its intent is merely to avoid such barbarisms as linnaeusti. If one is basing a genitive epithet on the already-Latin T am indebted to Dr. A. E. Schuyler, of the Academy of Natural Sciences of Philadelphia, for examining for me a label from Gray’s exsiccata. (For the significance of Gray’s Gram. & Cyp., see Rickett & Gilly, Bull. Torrey Bot. Club. 69: 461-466. 1942.) 440 Rhodora [Vol. 68 gerardus, then of course it must be gerardi, or at least not gerardusii; but if one bases it simply on the French name of Gérard, whom this plant honors, then it seems to me that Art. 73 prefers, for consistency, gerardii. Aristida necopina Shinners According to Shinners (Rhodora 56: 30. 1954) this is the plant which, in the Midwest, has been referred to A. intermedia Scribn. & Ball; true A. intermedia is A. longe- spica Poir. var. geniculata (Raf.) Fern. Bromus pubescens Willd. f. glabriflorus (Wieg.) E. G. Voss, comb. nov. B. purgans L, f. glabriflorus Wiegand. Rhodora 24: 92. 1922. Wagnon (Rhodora 52: 213-215. 1950; Brittonia 7: 455. 1952) has indicated that the type of B. purgans L. belongs to the species long called B. latiglumis (Shear) Hitchcock, so that the older name supplants the latter. The plant long known ( erroneously) as B. purgans then becomes B. pubes- cens Muhl. ex Willd. The above transfer is therefore neces- sary to make available a name for the rare form with the lemmas and paleas all completely glabrous. Deschampsia cespitosa (L.) Beauv. The epithet was originally spelled cespitosa by Linnaeus (Sp. PL, p. 64. 1753). While this may be less preferable Latin than caespitosa, it was clearly not a typographical error (cf. Philos. Bot., No. 274, and other usage by Lin- naeus), and alteration of the original spelling is unjustified. Echinochloa muricata (Beauv.) Fern. Echinochloa crusgalli (L.) Beauv. var. frumentacea (Link) W. F. Wight The case for the name and author citations of E. muricata was adequately presented by Fairbrothers and Reeder (Rhodora 58: 48 & 331-332, respectively. 1956.) Since Pani- cum muricatum Michx. is illegitimate, the basionym (under Art. 72) becomes Setaria muricata Beauv. In making the combination in Echinochloa (Rhodora 17: 106. 1915), Fer- nald cited the Michaux reference, not Beauvois. Acceptance of Fernald as author of the combination seems to be a good illustration of Art. 33, Note 2 (“Bibliographic errors of citation do not invalidate the publication of a new combina- tion.” ) — even though the “error” in this case is citing the 1966] Monocots — Voss 441 wrong author and hence place of publication of the ba- sionym. A similar situation involves the epithet frumentacea. Panicum frumentaceum Roxb. (Fl. Ind. 1: 307. 1820), usually cited as the basionym, is an illegitimate homonym of P. frumentaceum Salisb. (Prod. Stirp. Hort. Chap. All., p. 18. 1796), which was itself an illegitimate substitute name for Holcus sorghum L. (cited as “Linn. Sp. Pl. ed. 2. p. 1404"). Therefore, Echinochloa frumentacea Link (Hort. Reg. Bot. Berol. 1: 204. 1827) is, under Art. 72, treated “as new,” and becomes the basionym of E. erusgalli var. frumentacea. Eragrostis cilianensis (All.) E. Mosher Shinners (Rhodora 56: 26, 27. 1954) discussed the no- menclature of this species, tentatively attributing the com- bination to Hitchcock (whose 1919 usage is not a “nomen nudum" as termed by Shinners, for no description is re- quired for a new combination and even citation of the basionym was not required prior to 1953 [Art. 83]). Shinners cautiously pointed out that it was possible that some author published the combination prior to Hitchcock. The name was used in Mosher's Grasses of Illinois (lll. Agr. Exp. Sta. Bull. 205: 381. 1918) — but Shinners' warn- ing still stands: there may be yet a prior use. Festuca pratensis Huds. This name has been used for some time in Europe (the native home of the species) for what many manuals con- tinue to call F. elatior L. (Sp. Pl, p. 75. 1753.), a name which may be rejected under Art. 69 as a nomen confusum; otherwise, it supplants F. arundinacea Schreb. (cf. Dandy, List Brit. Vasc. Pl., p. 157. 1958). Koeleria macrantha (Ledeb.) Spreng. Shinners (Rhodora 58: 94. 1956) has pointed out not only that K. cristata Pers. is an illegitimate name but also that it applies to a European species, so that our American plant must apparently be identified as K. macrantha. Panicum commutatum var. ashei Fern. Not P. ashei Pearson in Ashe. Jour, Elisha Mitchell Sci. Soe. 15: 35. 1898. 442 Rhodora [Vol. 68 Panicum ashei Pearson was originally published as a nomen novum for P. commutatum Schult. var. minor Vasey. The reference cited is Contr. U. S. Nat. Herb. 3: 32 (1892) and not the original place of publication of var. minor (U. S. D. A. Bot. Div. Bull. 8: 34. 1889). How- ever, Vasey's description is identical in both places, and bibliographie errors of citation do not invalidate the publi- cation of a new combination (Art. 33, Note 2) — or, pre- sumably, of a nomen novum. Panicum ashei and P. com- mutatum var. minus (as ‘minor’) are synonymous. Since the latter is apparently not taxonomically distinct from typical P. commutatum (cf. synonymy of Hitchcock and Chase, and implication in Fernald, Rhodora 36: 83. 1934), P. ashei also belongs in the synonymy of P. commutatum. Fernald cited P. ashei Pearson as basionym for his new combination (loc. cit.), but definitely stated "excluding synonym." Fernald's varietal name must be treated as new, dating from 1934, validated by reference to the de- scription and citations by Pearson in Ashe but specifically excluding the cited synonym which unfortunately makes the binomial P. ashei Pearson apply to a different plant from the varietal name P. commutatum var. ashei Fernald. Those who believe that the weakly distinguished var. ashei should be accorded specific rank will have to provide a new name for it. (The type of P. ashei, as a nom. nov., must be the type of P. commutatum var. minus. Hitchcock & Chase’s designation of Ashe’s 1898 collection from Ithaca as a type is irrelevant.) "Hitchcock and Chase (N. Am. Sp. Panicum, Contr. U. S. Nat. Herb. 15: 301. 1919) argue that the synonym is cited in error and that P. ashei, as a “sp. nov.” was not really intended as a nom. nov. However, the phrase following the listing of Vasey’s name, “Not P. capillare var. minus Muhl. (1817)" indicates that Pearson (or Ashe) intentionally avoided creating a homonym of P. minus (Muhl.) Nash [Bull. Torrey Bot. Club 22: 421. 1895]. Since Vasey's varietal epithet could thus not be raised to specific rank, a new name was required. Note, too, that no type was designated, whereas for truly new species in the same paper Ashe usually designated a type when more than one of his collections was cited. 9 1966] Monocots — Voss 443 Panicum depauperatum Muhl. var. involutum (Torr.) Wood P. depauperatum var. psilophyllum Fern. It is not clear why Fernald (Rhodora 23: 193. 1921) found it necessary to provide a new name for “the common plant with sheaths sparsely pilose or quite glabrous” since Hitchcock and Chase (N. Am. Sp. Panicum, Contr. U. 5 Nat. Herb. 15: 151. 1910) had long previously noted that the type of P. involutum Torrey (Fl. No. & Mid. States, p. 144. 1824) had “sparsely pilose sheaths.” Hence, Fer- nald's name is considerably antedated by f involutum (Torr.) Wood (Class-book, ed. 1860, p. 786). Panicum rigidulum Nees P. agrostoides Spreng. As was pointed out long ago by Hitchcock and Chase (N. Am. Sp. Panicum, Contr. U. S. Nat. Herb. 15: 100. 1910) Sprengel (Pl. Pugill. 2: 4. 1815) cited P. agrostidi- forme Lam. as a synonym of his proposed new P. agro- stoides. Hence, under Art. 63, Sprengels name is illegitimate as superfluous when published, for by citing Lamarck’s name he apparently intended to circumscribe his species so as to include Lamarck’s plant — even though Sprengel did in fact have something new. As was also pointed out by Hitchcock and Chase (loc. cit.), P. rigidulum Bosc. ex Spreng. (Syst. Veg. 135520. 1825) is a name merely published in synonymy (of P. an- ceps Michx.). Thus the 1825 name is invalid and has no status under the Code (Art. 12); it is not illegitimate. Hence, the later described P. rigidulum Bosc. ex Nees (Agrost. Brasil. [= Mart., Fl. Brasil. 2(1) ], p. 163. 1829) is evidently the earliest correct name for this plant. (It is of interest to note that as long ago as 1897 (U. S. D. A. Div. Agrost. Bull. 4: 21), Holm reported that the specimen of P. agrostoides in Willdenow's herbarium has “P. rigidulum Bosc.” added by Willdenow on the label.) Paspalum muhlenbergii Nash Although described by Nash (Britton, Man., p. 75. 1901) as a species distinct from “Paspalum pubescens Muhl.," this was not maintained as a distinct entity by Mrs. Chase (N. Am. Sp. Paspalum, Contr. U. S. Nat. Herb. 28: 83. 1929) 444 Rhodora [Vol. 68 or by Fernald (Rhodora 36: 20. 1934). Mrs. Chase used the name P. pubescens Muhl. both in her monograph and in the revised edition of the Manual of Grasses (as had Hitch- cock in the original edition), in spite of the fact that in the monograph (loc. cit.) she herself stated: “Willdenow cites P, ciliatifolium Michx. as a synonym." Thus, Paspalum pubescens Muhl. ex Willd. (Enum. Plant. Hort. Reg. Bot. Berol., p. 89. 1809) is an illegitimate name, superfluous when published (Art. 63). Not only does Will- denow quote Michaux's description fully (after his own), bu: "Habitat in Carolina" is also apparently taken from Michaux. Although the type sheet of P. ciliatifolium Michx. (Fl. Bor.-Am. 1: 44. 1803) is mixed (Chase, op. cit., p. 86), Willdenow gives no evidence of intentionally basing his species on one of the included elements. Whatever speci- mens Muhlenberg or Willdenow may have had before them, they are irrelevant to the nomenclatural situation of pro- posing what is, in effect, a substitute name. If one does not follow Fernald in treating these plants as P. ciliatifolium var. muhlenbergii (Nash) Fern., they must be called P. muhlenbergii Nash. P. pubescens Muhl. ex Willd. must be cited in the synonymy of P. ciliatifolium Michx. Phalaris arundinacea L., f. variegata (Parnell) Druce P. arundinacea f. picta (L.) Asch. & Graebn. The combination by Druce (FI. Berks., p. 556. 1897) was clearly based on P. arundinacea var. variegata Parnell (Gr. Brit., p. 188. 1845). The latter was evidently intended as a new taxon (no prior authors are cited in the manner employed by Parnell for most other taxa), although the variety is undoubtedly the same as Calamagrostis variegata Withering (Bot. Arr. Brit. Pl., ed. 3, 2: 124. 1796). Since Withering cited P. arundinacea L., his binomial is super- fluous anyway, and could not serve as basionym for Par- nel's name, which in any event, therefore, is treated as new — although it was taxonomically unnecessary in view of P. arundinacea var. picta. L. (Sp. PL, p. 55. 1753). The later formal combination by Ascherson and Graebner (Syn. Mitt.-Eur. Fl. 2: 24. 1898), based on P. arundinacea 1966] Monocots — Voss 445 [var.] picta L. (the page number they cite [80] is that of Sp. Pl, ed. 2, although said to be ed. 1), is taken up by Anderson in his recent monograph of Phalaris (Iowa State Jour. Sci. 36: 38. 1961). Neither Anderson nor Chase and Niles (Ind. Gr. Sp., 1962) nor Man. Gr. list f. variegata, and the latter two do not list f. picta. Hence, it seems valu- able to summarize the nomenclature. The attractive “Rib- bon Grass" may be called P. arundinacea var. picta L. or f. variegata (Parnell) Druce — but not f. picta nor var. variegata (unless, of course, one makes the taxonomic judgment that there exist two different taxa of ribbon grass). Fuccinellia fernaldii (Hitche.) E. G. Voss, comb. nov. Glyceria pallida (Torr.) Trin. var. fernaldii Hitchcock, Rhodora 8: 211. 1906. Glyceria fernaldii (Hitche.) St. John, Rhodora 19: 76. 1917. Torreyochloa fernaldii (Hitche.) Church, Am. Jour. Bot. 36: 164. 1949. Torreyochloa pallida (Torr.) Church ssp. pallida var. fernaldii (Hitchc). Dore, Canad. Jour. Bot. 42: 874. 1964. This combination is proposed with some reluctance, but it is necessary if one follows the suggestion of Clausen (Rhodora 54: 42-45. 1952) that Torreyochloa Church (the Glyceria pallida group) be treated as a section of Puccinel- lia. In defending Torreyochloa, Church (Rhodora 54: 187- 200. 1952) questioned the recognition of fernaldii at specific rank, and Clausen evidently had doubts on this point also. Since in Michigan it is morphologically and geographically distinct from P. pallida (Torr.) Clausen, I recognize it, at least at present. Setaria glauca (L.) Beauv. Reeder (Rhodora 53: 27-30. 1951) discussed this name under the title: “Setaria lutescens an Untenable Name." Rominger in his recent monograph of Setaria (Ill. Biol. Monogr. No. 29, pp. 97-98. 1962) rehashes the old argu- ments against S. glauca, rejects Reeder's conclusion with a backhanded compliment (referring to it as a strong argu- ment and nowhere directly refuting it), and ignores com- pletely Reeder's assertion that the assumed basionym, Panicum lutescens Weigel, cannot in any event be accepted 446 Rhodora [Vol. 68 since it is invalid under Art. 33, which requires for valid publication that an author definitely indicate that an epithet is to be used in a particular combination. Weigel's publieation was first questioned by Weatherby, et al. (Rhodora 31: 109. 1929), who stated that he “no- where directly makes the combination Panicum lutescens.” Exactly the same assertion ("nowhere") is made by Reeder. Unfortunately for this argument, “nowhere” is not true. In the explanation of his plates, Weigel (Obs. Bot., 1772) does indicate that on Plate II, Fig. 1 is “of Panicum lutes- cens" [“Panici lutescentis"] and Fig. 2 is of Panicum virescens. With this evidence of his intent in proposing (p. 20) the epithets, I think we must accept them as validly published binomials. Otherwise, for those who are con- vinced that S. glauca is incorrect, the basionym must pre- sumably become, under Art. 72, Chaetochloa lutescens Stuntz (U. S. D. A. Bur. Pl. Ind. Inv. Seeds & Pl. Imp. 31: 86. 1914 — incorrectly cited as p. 84 by Weatherby and as p. 83 by Reeder). Sorghum bicolor (L.) Moench S. vulgare Persoon A clear summary of this rather clearcut but often ignored case has been given by Shinners (Baileya 4: 141-142. 1956). Sorghum bicolor was specified as the type of the genus by Clayton in his proposal for conservation of Sorghum (Taxon 10: 242, 243. 1961). To the usual synonymy of this species (e.g., Snowden, Cultivated Races of Sorgnum, p. 166. 1936) should be added Panicum frumentaceum Salisb. (cf. under Echinochloa, above). Spartina pectinata Link var. suttiei (Farwell) Fernald It is extremely doubtful whether this variety has any taxonomic significance whatsoever, and it is mentioned here only to call attention to a slight correction necessary in the indication of type locality in Man. Gr. (rev. ed., p. 959. 1951) and in the Ind. Gr. Sp. (Chase & Niles, 1962). These sources give “Orchard Lake, Mich., Suttie” in designating the type. However, in describing S. michauxiana var. suttiei, Farwell (Rep. Mich. Acad. 21: 352. 1920) said: "Island Lake, July 16, 1905, No. 1487a. Also Dr. George 1966] Monocots — Voss 447 Suttie, Orchard Lake, July 29, 1892, and Waterford, August 27, 1893." Priority in this listing is given to Island Lake, and further confirmation is Farwell’s specimen No. 1487a, marked *Type" by him (BLH), as pointed out by MeVaugh, Cain, & Hagenah (Farwelliana, Cranbrook Inst. Sci. Bull. 34: 71. 1953). The type locality is Island Lake [Livingston Co.], not Orchard Lake [Oakland Co.], Michigan, the type being Farwell 14872. Sporobolus vaginiflorus (Torr.) Wood var. neglectus (Nash) Scribn. S. vaginiflorus var. neglectus (Nash) Shinners Scribner's combination (Am. Gr. 2, U. S. D. A. Div. Agrost. Bull. 17, rev. ed., p. 170. 1901) considerably ante- dates the same combination by Shinners (Rhodora 56: 29. 1954), but what started as a routine check on an apparently obvious duplication revealed a tangle of confused citations, upon which some light has already been shed by Rickett & Gilly (Bull. Torrey Bot. Club 69: 463. 1942). Scribner used the spelling S. vaginaeflorus, and the first source which might lead one astray (and suggest that Shinner's combination was indeed required) is the Gray Index card for “Sporobolus vaginaeflorus Vasey, not Wood.” First of all, the reference on this card to Vasey in Watson & Coulter (Man., ed. 6, p. 645. 1889) is later than Vasey, Cat. Gr. U. S., p. 45 (1885). Secondly, there is no justification for the assertion “Vasey, not Wood.” Vasey (1885) cited “(Vilfa, Torr.)" — which is more of a basionym than one might have expected, or than was re- quired as of that date. I see no reason to question that Vasey’s Sporobolus vaginaeflorus is based on Vilfa vagini- flora Torr. (in Gray, N. Am. Gram. & Cyp. 1: 3. 1834). Rickett & Gilly (loc. cit.) have pointed out that Torrey later used the spelling vaginaeflora (Fl. N. Y. 2: 438. 1843). These are merely orthographic variants, and the present Code (Rec. 73Gd) favors the first spelling as the correct form. The combination by Wood (Class-book, ed. 1860, p. 775) antedates Vasey's and has the same basionym (*'va- ginaeflorus Torr"). If S. vaginaeflorus Wood and S. vaginaeflorus Vasey are indeed the same, Scribner's 1901 var. neglectus is correct with the former specific combina- 448 Rhodora [Vol. 68 tion, for those who do not consider S. neglectus a distinct species. Sporobolus vaginiflorus (Torr.) Wood This combination, as noted above, is based on Vilfa va- giniflora Torr. (in Gray, Gram. & Cyp. 1: 3. 1834). This is often cited “Torr. ex Gray," but a study of the situation sug- gests that this is not the case and, moreover, that the type locality is generally given erroneously. The printed label with Gram, & Cyp. 1: 3 reads as follows: 3. Vilfa vaginiflora. Torrey, Synop. Flora, ined. Agrostis Virginica. Muhl. Gram. p. 14, Torrey, Flora, v. 1, p. 89, non Elliott et Auct. HAB. — Dry barren fields, New-Jersey. What is being proposed is a nomen novum for Agrostis virginica sensu Muhlenberg (and, later, Torrey). The true A. virginica L. [— Sporobolus virginicus (L.) Kunth] oc- curs (or occurred) from southeastern Virginia and North Carolina along the coast to Texas (and southward). The more northern plant, previously confused with it, required a new name. This was proposed by Torrey in manuscript and published by Gray, the publication validated by refer- ence to a previously published description [Muhlenberg (1817) antedates Torrey (1824) and is cited first]. If a description had been required, and had been provided by Gray for Torrey's name, the citation would be “Torrey ex Gray." Since no description was required, Torrey's recognition of the need for a new name presumably re- quired no further work on the part of Gray, and “Torrey in Gray" is an acceptable citation (as it would be assumed for a new combination, for which a description is also not required; cf. Voss, Taxon 14: 154-160, esp. p. 158. 1965). Since the new name was validated by reference to a pre- vious description, the New Jersey collection of Gray's Gram. & Cyp. has no relevance to the selection of a type, which would be the same as that for the plant previously described under the mistaken identity (Art. 7, Note 4). Muhlenberg (p. 75) stated: *Habitat in aridis, floret Sept. Penns. Caro- lina, Georgia." Since Pennsylvania plants would be per- sonally familiar to him and would undoubtedly be vagini- 1966] Monocots — Voss 449 flora rather than true virginica, the type locality would more likely be Pennsylvania than the Carolinas or Georgia. It would in any event not be New Jersey, a state not in- cluded in the description.? Trisetum spicatum (L.) Richt. var. molle (Kunth) Beal Trisetum spicatum var. maidenii (Gandoger) Fernald As is clear from the recent treatment of the complex by Hultén (Sv. Bot. Tidskr. 53: 203-228. 1959), the wide-rang- ing Trisetum spicatum is taxonomically complicated. In Gray’s Manual, ed. 8, Fernald adopts the varietal treatment proposed by him earlier (Rhodora 18: 195-198. 1916). I do not at this time intend to imply that this treatment (in- cluding the varieties named above) is necessarily the most helpful, but want merely to call attention to necessary cor- rections in citation for two of Fernald's three varieties (the third, var. pilosiglume, was described as new by him). The Rhodora treatment of T. spicatum var. molle (Michx.) Piper was corrected to var. molle (Michx.) Beal in the Manual (p. 145). However, the basionym is incor- rectly attributed to Michaux, for Michaux's Avena mollis was (as is clear, e.g., from the synonymy in Man. Gr., rev. ed., p. 977. 1951; in Hultén, op. cit.; and in Chase & Niles, Ind. Gr. Sp., 1962) illegitimate when published, being a homonym of two earlier names. Trisetum molle of Kunth (Rév. Gram. 1: 101. 1829), based on A. mollis Michx., is therefore, under Art. 72, treated as new. Since Beal (Gr. N. A. 2: 377, 1896) cited Kunth as well as Michaux, the authorship of the variety should be cited as var. molle (Kunth) Beal. Similarly, those who follow Hultén's treatment would have ssp. molle (Kunth) Hultén. A further discussion of this species complex has recently been published by Löve and Löve (Univ. Colo. Stud. Ser. Biol. 17: 608. 1965), who recognize the illegitimacy of Michaux’s name but neglect to observe that Kunth's name "I am indebted to A. E. Schuyler and C. W. Laskowski, who have examined the Muhlenberg collections in the Academy of Natural Sciences of Philadelphia in an unsuccessful search for a specimen with data which might definitely determine the type locality. 450 Rhodora [Vol. 68 (which they date from 1833 rather than 1829) is legitimate (by Art. 72) although it may be a taxonomic synonym of Melica triflora Bigel. and therefore may serve as a legiti- mate basionym for combinations in other ranks, e.g., T. spicatum ssp. molle (Kunth) Hultén or T. triflorum ssp. molle (Kunth) Löve & Löve (the latter combination, as published with Hultén as author of basionym, an example of bibliographic error of citation). The variety T. spicatum var. maidenii (Gandoger) Fern. is based, according to the citations in Rhodora (op. cit.), in the Gray Index, in Hultén (op. cit.), and in Ind. Gr. Sp., on T. subspicatum f. maidenii Gandoger (Bull. Soc. Bot. France 49: 182. 1902). Whether this incorrect citation is repeatedly given as a result of independent error or un- critical copying I do not know. But the original publication of T. subspicatum f. maidenii Gandoger was three years earlier (Bull. Soc. Bot. France 46: 393. 1899). The 1902 note was merely a defense of the validity of the taxon, it having been challenged by Maiden (quoted, op. cit. 49: 72. 1902). The use of boldface type in the indices of these vol- umes to indicate new taxa clearly supports the intent of the 1899 publication, which was complete with Latin diagnosis, to describe a new form. CYPERACEAE Carex One of the most brutal products of Fernald's acid (and usually accurate!) pen was his review (Rhodora 49: 49- 52. 1947) of Beetle's conspectus of sections in the genus Scirpus. Under the title, *Unverified Bibliography of Scir- pus,” Fernald called attention to a number of Supposed sec- tional names which were not published as sections by the authors to whom Beetle attributed them. It is, therefore, not a little surprising to find in Fernald’s own edition of Gray's Manual (1950) 58 sections recognized in the genus Carex, most of which are subject to the same criticisms pre- viously leveled at Beetle's Scirpus sections. I will not take the space here to itemize the several sup- posed sectional names in Carex which I have checked and 1966] Monocots — Voss 451 found not to have been validly published in that rank or, if so published, not by the author to whom ascribed in the Manual. Since such names are not in the usual indexes, and only partial synonymies appear in monographs, the work of verifying all of them is a task of greater magnitude than I have the time or competence to undertake when it is not required. The simple solution is to refer to the traditional names in Carex as merely names of convenient groups, without nomenclatural standing and not designated as sec- tions. Even so, it would be in the spirit of the Code to change the designation of the group long known as Acutae to Carex, since it is the group which includes C. acuta L., the type species of the genus. (This change would be required, under Art. 22, in the rank of section.) Carex bebbii (Bailey) Fern. This name is usually attributed to Olney (Car. Bor.-Am. 2:12. 1871), but as is clear from Mackenzie (N. Am. Fl. 18: 147. 1931), the first valid publication of the epithet was as C. tribuloides var. bebbii Bailey (Mem. Torrey Bot. Club i 55. 1889). The combination in specific rank was made (ap- parently more or less inadvertently) by Fernald (Proc. Am. Acad. 37 [= Contr. Gray Herb. 22]: 478. 1902,) who also pointed out that Olney's name was a nomen nudum. Mac- kenzie (op. cit., p. 148) observed that the few words by Bailey earlier (Bot. Gaz. 10: 379. 1885) "can scarcely be regarded as a publication of the name"; regardless of the number of words, Bailey at that point considered C. bebbii to be a synonym of either C. lagopodioides or C. scoparia — in any event, the name is there only in synonymy and hence without nomenclatural standing. Carex communis Bailey f. gynandra (Farwell) E. G. Voss, comb, nov. C. communis var. gynandra Farwell, Am. Midl. Nat. 12: 52. 1930. It is doubtful whether taxa such as this merit recognition even in the rank of form, but the above combination is made for consistency with, e.g., C. pensylvanica f. androgyna F. J. Hermann. Carex houghtoniana Dewey C. houghtonii Torrey As was pointed out by Butters and Abbe (Rhodora 55: 452 Rhodora [Vol. 68 134. 1953) and Rittenhouse and Voss (Mich. Bot. 1: 66. 1962), C. houghtoniana Torrey ex Dewey (Am. Jour. Sci. 30: 63. 1836) is validly published and there is no sanction in the Code for Torrey's emendation to C. houghtonii ( Ann. Lyc. Nat. Hist. N. Y. 3: 413. 1836). Carex hystericina Willd. The original spelling was C. hystericina Muhl. ex Willd. (Sp. Pl., ed. 4, 4(1) : 282. 1805) and in the absence of any evidence that Willdenow committed an unintentional ortho- graphic error, emendation to C. hystricina is unjustified — as in the previous instance. Carex lasiocarpa Ehrh. var. latifolia (Boeckl.) Gilly C. lasiocarpa var. latifolia (Boeckl.) Gleason C. lanuginosa Michx. Although I prefer to recognize C. lanuginosa as a good species, it is perhaps worth noting that the combination by Gleason (Phytologia 4: 22. 1952) used in the Ill. Fl. (1: 349. 1952) is quite unnecessary, having been made six years previously by Gilly (Iowa St. Coll. Jour. Sci. 21: 125. 1946), with good bibliography. Gleason's combination is therefore properly not included in the Gray Index. Carex leptalea Wahl. var. harperi (Fern.) Weathb. & Grisc. C. leptalea ssp. harperi (Fern.) Stone The varietal combination is usually attributed to Stone (Pl. So. N. J., p. 305. 1912), whose work clearly indicates that his trinomials were intended to represent subspecies (cf., e.g., pp. 35 & 119). The first use of the epithet in varietal rank was apparently by Weatherby & Griscom (Rhodora 36: 39. 1934). Stone's subspecific combination is given as an undesignated trinomial in the Gray Index. The unnecessary “comb. nov.” as subspecies by Calder and Tay- lor (Canad. Jour. Bot. 43: 1391. 1965), based on “var.” harperi, presumably resulted from failure to examine Stone's publication carefully. Carex livida (Wahl.) Willd. var. radicaulis Paine C. livida var. grayana (Dew.) Fern. In publishing var. grayana, Fernald (Rhodora 28: 8. 1926) stated that to take up Paine’s name for “ a very ex- ceptional departure” would “lead only to confusion.” Un- fortunately, the Code makes no allowance for exclusion of a 1966] Monocots — Voss 453 name based on an unusual specimen if it is included in a taxon as cireumscribed, and hence the varietal name of Paine (Rep. N. Y. St. Cab. 18: 159. 1865) has priority over Fernald's superfluous name. Carex rugosperma Mack. var. tonsa (Fernald) E. G. Voss, comb. nov. C. umbellata Willd. var. tonsa Fern. Proc. Am. Acad. 37 [— Contr. Gray Herb. 22]: 507. 1902. Sufficient intermediacy exists between tonsa and typical C. rugosperma that varietal status for the former seems desirable. Those who follow such Carex authorities as Mac- kenzie and Hermann in applying the name C. umbellata Willd. to what Fernald calls C. abdita Bickn. will require this combination with C. rugosperma Mack. (C. umbellata sensu Fern. et al., not Schk. ex Willd.). Eleocharis erythropoda Steudel E. calva Torrey Svenson (N. Am. Fl. 18: 525. 1957) clarified the nomen- clature of this species. It is discouraging to contemplate for how many years the plant was called E. calva Torr. (Fl. N. Y. 2: 346. 1843), when Torrey merely mentioned certain specimens at the end of a paragraph under E. palustris and stated: “It may be regarded as a provisional species, under the name of E. calva." No more straightforward example of a “provisional name," invalid under Art. 34, can be im- agined. Since a name not validly published has no status under the Code (Art. 12) — i.e., is not even illegitimate — later valid publication of E. calva would not be considered a homonym. Such publication evidently did not occur, how- ever, prior to FE. erythropoda Steud. (Syn. Pl. Glum. II, Cyp., p. 76. 1855). Rhynchospora glomerata (L.) Vahl var. minor Britton R. glomerata var. capitellata (Michx.) Kuek. The combination by Kükenthal (Bot. Jahrb. 15: 98. 1950) is illegitimate; he cites R. “capitellata” [error for glo- merata] var. minor Britt. (Trans. N. Y. Acad. 11: 89. 1892) in synonymy. Those who wish to follow his tax- onomic disposition of R. capitellata (Michx.) Vahl should use the latter name. The spelling Rhynchospora is con- 454 Rhodora [Vol. 68 served, a point apparently overlooked by some recent au- thors who have used Rynchospora. JUNCACEAE Luzula multiflora (Retz.) Lejeune Being reluctant to propose a new name for this weil known species, I believe the Code can be interpreted to al- low retention of the familiar name. The nomenclatural! problems have been discussed only in part by Fernald (Rho- dora 47: 268-271. 1945) and by Hylander (Uppsala Univ. Arsskr. 7: 109-110. 1945). The chief synonyms and their status may be summarized as follows: Juncus multiflorus Retz. (Fl. Scand. Prodr., ed. 2, p. 82. 1795) [Not J. multiflorus Desf. (1798) (fide Hylander).] Juncus intermedius Thuill. (Fl. Env. Paris, ed. 2, p. 178. 1799) [Not J. intermedius Host (1805).] Juncus multiflorus Ehrh. ex Hoffm. (Deutschl. Fl., rev. ed., 1: 169. 1800) [Illeg. as homonym of Retz. (1795) if based on different type.] Juncus erectus Pers. (Syn., p. 386. 1805) [Illeg. as superfluous under Art. 63.] Luzula multiflora Lej. (Fl. Env. Spa, p. 169. 1811) [Based on J. multiflorus Hoffm., with J. intermedius Thuill. & J. erectus Pers. as synonyms (fide F, J. Hermann, in litt.).‘] Luzula intermedia Spenner (Fl. Friburg. 1: 177. 1825) [J. multiflorus Hoffm. cited as synonym, and also cited under f multiflora as is L. multiflora Hagenb.] Since Hagenbach (Tent. Fl. Basil., p. 336. 1821) included J. intermedius Thuill. in the synonymy of his L. “multiflora Hoffm.,” we can interpret this as the assumed basionym of Spenner's L. intermedia, Under e congesta, Spenner's only other var., J. congestus Thuill. is cited and is obviously the basionym. In any case, Spenner's binomial is illegitimate as a homonym of L. intermedia Baumg. (1816) and of Nocca & Balbis (1816), both of which were (fide Hermann) based on J. intermedius Host. The 1816 names should be treated as new, however, under Art. 72, since the name of Host (Icon. Descr. Gram. Austr. 3: 65. 1805) is superfluous under Art. 63, for he cited J. glabratus Hoppe (ex Rostk., Monogr. Gen. Junc., p. 27. 1801). (Host cited no publica- tion, and was perhaps unaware that Hoppe's unpublished name had been validated by Rostkovius.) 1966] Monocots — Voss 455 Fernald apparently assumed that L. multiflora Lejeune was based on J. multiflorus Retzius. Unfortunately Lejeune* did not cite Retzius, but Hoffman (which was based on a presumed nomen nudum of Ehrhart, as described by Fer- nald,® and on J. campestris 8 of the Flora Lapponica). Hoff- man's name, if based on a different type, is a homonym of that of Retzius; but Lejeune's name cannot be treated as new under Art. 72 since it would be nomenclaturally super- fluous under Art. 63 (as he included the prior J. intermedius Thuill. in his synonymy). Furthermore, it would be tax- onomically superfluous if not based on J. multiflorus Retz., if we assume the latter to be the same species as J. multi- florus Ehrh. ex Hoffm. (as is maintained, e.g., by Buchenau, Pflanzenr. IV (36) : 94. 1906). Juncus multiflorus Hoffm. is a homonym of J. multiflorus Retz. only if it is based on a different type (Art. 64). There is, of course, the remote possibility that the two names are not based on different types and that Retzius also had a sheet No. 127 from Ehrhart's exsiccata as cited by Hoff- man, Unfortunately, Retzius cited no such specimen, and I am informed by B. Peterson of Lund (in litt., 1958) that no specimen named J. multiflorus can be found in Retzius' herbarium (nor in the Riksmuseum nor in Bergius' her- barium in Stockholm). Although no help is thus available ‘I am indebted to Dr. F. J. Hermann for checking the Lejeune and some other references for me and for considerable helpful corre- spondence (1957-1959) on this problem — although he is not to be held responsible if I have reached any erroneous conclusions. "Fernald expressed some doubt as to the status of a 13th decade of Ehrhart's exsiccata, and appears not to have observed that on the Tth (unnumbered) page of his Introduction, Hoffman cites *Ehrhart's, Calamariae, Gramina et Tripetaloideae Linn. Dec. 1-14. 1789-1793. Fol" Additional evidence is that Buchenau, in his monograph (Bot. Jahrb. 12: 162. 1890), cites “Collect. EHRHART, Calam., 127 (1) under L. campestris var. multiflora, although the highest number cited in the "Index collectionum" (p. 473) is 126. Ehrhart's specimens seem to have been intended to illustrate authoritatively the Linnaean species, and one cannct help noting the striking coincidence (if indeed it is that) that No. 127 is the same number as Juncus campestris B in the Flora Lapponica, the only other reference cited by Hoffman under J. multiflorus. 456 Rhodora [Vol. 68 to make clear what Retzius had, there is the additional possibility that Hoffman was aware of the earlier name and used it without citing its author. In view of the uncertainties here suggested, the fact that J. multiflorus Retz. and J. multiflorus Hoffm. apparently refer to the same species whether or not based on the same type, and the desirability of a stable nomenclature, the best solution seems to me to invoke Art. 33, Note 2: “Biblio- graphic errors of citation do not invalidate the publication of a new combination." (Cf. comment under Echinochloa, above.) We can accept Luzula multiflora (Retz.) Lej. if we treat Lejeune’s citation of Hoffman rather than of Retzius as a bibliographic error. This course would also explain Lejeune’s failure to base his binomial on the earlier J. inter- medius Thuill., which he cites. It is not clear that there is any fully satisfactory alter- native other than a new name. Without laboring the point further here, suffice it to say that several other epithets which (from Index Kewensis or other sources) might be thought applicable have been checked by Hermann or me and found to be taxonomically or nomenclaturally inadmis- sible. If L. multiflora and L. congesta are considered con- specific, one could take up L. congesta (Thuill) Lej. (op. cit., p. 168) for the species and make multiflora a variety or subspecies of it. [Spenner (op. cit., p. 178) was evidently the first to treat multiflora in varietal rank, under his L. intermedia; if we are, in this line of thought, rejecting Hoff- man’s name as a homonym of Retzius’s name, then it is il- legitimate and Spenner’s varietal use of the epithet would be treated as new. There may even be some earlier epithet available in varietal rank.] This is not the place for a mono- graph of Luzula, and I leave the subject with attention called to the unusual nomenclatural complexities. LILIACEAE Hemerocallis lilio-asphodelus L. H. flava (L.) L. Continued use by some authors of the illegitimate H. flava suggests that it is not inappropriate to call attention to dis- 1966] Monocots — Voss 457 cussions of the case by Farwell (Am. Midl. Nat. 11: 51. 1928), Hylander (Uppsala Univ. Arsskr. 7: 112. 1945), and Dress (Baileya 3: 107-108. 1955). The original H. lilio- asphodelus L. must be retained for one of the two elements into which it was later divided — even if the later divider was the revered Linnaeus himself. Muscari atlanticum Boissier & Reuter M. racemosum Auct., non Miller, non DeCandolle This common grape-hyacinth, which occasionally escapes from cultivation, was originally called Hyacinthus race- mosus L. (Sp. Pl, p. 318. 1753). In 1768, Miller (Gard. Dict., ed. 8, no. 3) described a Muscari racemosus, which too many authors (e.g., Fernald) have assumed to be based on H. racemosus L. However, it is clear, from the Clusian synonym which both cite and from the common names given, that Miller was instead providing a new name for the musk-hyacinth, H. muscari L. (p. 317). In order to avoid the tautonym Muscari muscari when transferring these species into Muscari, Miller was perfectly correct (ac- cording to present rules) in providing a new epithet. He was free to use any epithet he wanted (provided it was not otherwise in use in Muscari), and it is unfortunate that he happened to select a common epithet, already in use by Linnaeus for another species which ought to have been transferred to Muscari. Therefore Muscari racemosum Mill. (loc. cit.)* applies to the species later called M. moschatum Willd. (Enum. Pl. Hort. Reg. Bot. Berol., p. 378. 1809), the name by which it usually appears in works on cultivated plants. Even if one should now reject M. racemosum Mill. as a nomen am- "When making the transfers into Muscari under the binomial system (Gard. Dict., ed. 8. 1768), Miller treated the generic name as mascu- line; Hylander (Uppsala Univ. Arsskr. 7: 88. 1945) has therefore termed incorrect the usual usage as neuter. However, valid publica- tion of the genus dates from the 4th abridged edition of Miller's Gardeners Dictionary (1754), wherein the genus was consistently treated as neuter. The fact that Miller changed his mind 14 years later does not alter the original gender, which followed that of Tournefort, and which has been followed by Medicus, DeCandolle, and Turrill (in the works cited here), as well as others. 458 Rhodora [Vol. 68 biguum under Art. 69, the plant cannot be called M. mos- chatum, since the latter name is distinetly antedated by M. muscarimi Medicus (Ann. Bot. Usteri 2: 15. 1791), a name clearly proposed as a substitute for Hyacinthus muscari L. in a paper on the Linnaean genus Hyacinthus. Medicus cites not only the Linnaean name (transfer of which would have created a tautonym) but also the Clusian synonym. (Cf. Turrill, Bot. Mag. 167, sub t. 124. 1950). [Another alternative, for those who prefer small genera, is Mus- carimia, muscari (L.) A. Losink., Fl. URSS 4: 411. 1935.] The combination *Muscari racemosum (L.) DC.,” which is often seen, is of course a homonym of M. racemosum Mill, and therefore illegitimate even if it were based on the Linnaean name. However, DeCandolle (in Lam. & DC., Fl. Franç., ed. 3, 3: 208. 1805) cited first, M. racemosum Mill., and second, H. racemosus L. “Spec. 455” [a reference to the 2nd (or 3rd) ed. of Sp. Pl.]. Thus, DeCandolle was evidently confused and thought that Miller's name was based on H. racemosus L.; that he did not equate Miller's name with H. muscari (as he should have) is clear from his citation (op. cit., p. 207) of the latter under his M. am- brosiaceum Moench, the odoriferous species (our “musk- hyacinth’’) .7 Since the name Muscari racemosum is unavailable for the common grape-hyacinth, a taxonomic judgment must be made as to what other name, if any, in the literature is ap- plicable to this species. Pending the monographie work which this genus needs, the appropriate name seems to be M. atlanticum Boiss. & Reut. (Pugill. Pl. Nov., p. 114. 1852). This name was taken up by Dandy (List Brit. Vasc. Pl., p. 599. 1958), who has been followed by Clapham, Tutin, & Warburg (Fl. Brit. Isles, ed. 2, p. 975. 1962). Polygonatum commutatum (Schult. f.) A. Dietr. P. canaliculatum Auct. non Convallaria canaliculata Muhl. ex Willd. This synonymy was worked out by R. Ownbey (Ann. Missouri Bot. Gard. 31: 393 & 403. 1944) and I am unaware ‘I am greatly indebted to W. T. Stearn of the British Museum for his help in clarifying this problem (in litt., 1961). 1966] Monocots — Voss 459 of any reason not to accept her conclusions, although Fer- nald did not do so in Man., ed. 8. Smilax ecirrata (Kunth) Watson The usual spelling S. ecirrhata is given as a "bad ex- ample" of word construction by R. W. Brown (Composition of Scientific Words, p. 245 [& cf. p. 4]. 1954). It happens, however, that this is a bad example of a bad example. The original spelling was correct [Copromanthus herbaceus B ecirratus Engelm. ex Kunth, Enumerat. 5: 266. 1850] and there was no reason for Watson in assuming specific rank (Man., ed. 6, p. 520. 1889) or anyone else to adopt an in- ferior spelling. IRIDACEAE Sisyrinchium angustifolium Mill. The nomenclature of Sisyrinchium, especially revolving around this name, is no less complicated than its taxonomy. Both have been carefully investigated by Dr. Daniel B. Ward, whose results are not yet published, so it is prema- ture here to concur in his work, which has seemed sensible insofar as I have been aware of it. Thus postponing this large problem, I confine myself to a comment concerning a Michigan species: Sisyrinchium farwellii Bicknell This is noted by Fernald (Man., ed. 8, p. 458. 1950) as “scarcely separable” from S. arenicola Bickn. (Bull. Torrey Bot. Club 26: 496. 1899). One fears that the reason for Fernald's reluctance to state directly that they are the same is that S. farwellii (Bull. Torrey Bot. Club 26: 298. 1899) has a few months' priority and hence the familiar name for the eastern coastal species would have to be abandoned. In any event, Michigan plants (only the distinctive type col- lection is known) must be called S. farwellii, whether or not the coastal ones are different. Sisyrinchium farwellii differs in several important characters from S. atlanticum Bickn., of which for some reason Alexander (in Gleason, Ill. Fl., 1: 452. 1952) considered it only a form. 460 Rhodora [Vol. 68 ORCHIDACEAE Calopogon tuberosus (L). BSP. C. pulchellus (Salisb.) R. Brown Manuals continue to use Brown's illegitimate name in spite of a clear elaboration of the case by Mackenzie (Rho- dora 27: 193-196. 1925), whose opinion is apparently ac- cepted by Rickett & Stafleu (Taxon 8: 257. 1959). Corallorhiza trifida Chatelain In 1907, Rendle and Britten (Jour. Bot. 45: 442) called attention to the fact that the earliest valid publication of the generic name Corallorhiza was by J. J. Chatelain (Speci- men Inaug. Corall., 1760). This fact was reiterated by Fer- nald in 1946 (Rhodora 48: 193). In 1909, Ames (Rhodora 11: 106) made no mention of the correct citation of the generic name but did state (as had Rendle & Britten, whom he did not credit with the discovery) that since an inadmis- sible tautonym would result from the transfer of Ophrys corallorhiza L. to this genus, the valid name for the plant previously known as C. innata R. Br. (1813) must be C. trifida Chat. (1760). In the meantime, apparently no one has questioned taking up C. trifida. The generic name may, of course, validly date from Chatelain's work even if his specific names are for any reason to be rejected. In his little 15-page paper, Chatelain accepted two species of Corallorhiza, “Species Prima" is “Corallorhiza nectarii labio trifido" with the marginal epithet “Trifida,” as re- printed by Ames (loc. cit.). "Species Secunda," which has not been mentioned by those who have called attention to Chatelain's previously neglected paper, is *Corallorhiza nec- tarii labio integro," with no trivial epithet indicated in the margin. C. ruppii Hall. is cited as a [pre-Linnaean] syno- nym. Among the synonyms cited for C. nectarii labio trifido is the Linnaean polynomial (but not the Linnaean epithet, under Ophrys, of Corallorhiza). From this evidence, and knowing the reluctance of Haller (for whom Chatelain's publication was a doctoral disserta- tion) to accept binomial nomenclature wholeheartedly, we might seriously wonder whether this may not be a work 1966] Monocots — Voss 461 *in which the Linnaean system of binary nomenclature for species was not consistently employed" (Art. 23). One spe- cies has a binomial; the other does not. The consequences of such a conclusion can probably be avoided. Either we can consider that Chatelain in his “Spe- cies Secunda" was validating C. ruppit of Haller, or we can assume that a trivial epithet integra was accidentally omitted from the margin. The latter suggestion is sup- ported by the first sentence of his concluding paragraph: “Ex descriptione data Corallorhizae integrae, à trifida dis- crepare videtur, in eo, quod nectarii labium integrum sit." Apparently he did intend that his second species be known as Corallorhiza integra. This second species is presumably not taxonomically dis- tinct from C. trifida as now understood and generally con- sidered to be the only European representative of the genus. So whatever Chatelain intended to call it, the rule (Art. 57) of the “first reviser" (Rendle & Britten?) would cause us to accept C. trifida over the other name of the same date. No name C. integra, nor any other epithet of Chatelain’s except trifida, has been listed in the Index Kewensis, which finally caught up with trifida in the Fifth Supplement, and which still (11th suppl.) credits the genus (spelled ‘“Coral- lorrhiza" to Robert Brown. I do not propose to abandon the now familiar name C. trifida. But for consistency, if we assume that Chatelain did use binary nomenclature (thus validating trifida), his other species, even though undoubtedly a taxonomic syno- nym, should be properly indexed and accepted as validly published. Cypripedium X andrewsii Fuller Garay (Science 148: 65. 1965) has criticized the use of C. X andrewsii in Case's *Orchids of the Western Great Lakes Region" (Cranbrook Inst. Sei. Bull. 48, col. pl. 2. 1964). It should be borne in mind that when our yellow lady-slippers are treated as a single species (usually with two varieties, as in Case), the hybrids, no matter of which variety, with C. candidum would under Art. H. 1 receive the same name: “Where binary ‘specific? names of Latin 462 Rhodora [Vol. 68 form are used for hybrids, all offspring of crosses between individuals of the same parent species receive the same binary name. ... When polymorphic parental species are involved and if infraspecific taxa are recognized in them, greater precision may be achieved by the use of formulae than by giving the hybrids ‘specific’ names.” Without admitting that the Code adequately or clearly treats all problems in the nomenclature of hybrids, there would seem to be a good case for accepting C. X andrewsii Fuller (Rhodora 34: 100. June 3, 1932) over the slightly later C. X favillianum Curtis (Rhodora 34: 242. Dec. 14, 1932). Into the same synonymy would fall the backcross C. X landonii Garay (Canad. Jour. Bot. 31: 660. 1953). (Cf. Boivin, Nat. Canad, 87: 32. 1960.) Orchis rotundifolia Pursh f. lineata (Mousley) E. G. Voss, comb. nov. O. rotundifolia var. lineata Mousley, Canad, Field-Nat. 55: 65. 1941, Only formal rank seems appropriate for this striking variant, which was originally described from Alberta and later discovered in Ontario; The albino form of this spe- cles was named O. rotundifolia f. beckettii Boivin (Nat. Canad. 87: 42. 1960); since it is named for the industrious and amiable Mrs. Eva Beckett (formerly of Churchill, now of Fort William), the spelling should be corrected to beckettae (Art. 73, Note 3 & Rec. 73C). Spiranthes lacera (Raf.) Raf. In 1946 Fernald pointed out (Rhodora 48: 6 et Sqq.) that S. lacera (Raf.) Raf. (based on Neottia lacera Raf., 1818) would have priority over S. gracilis (Bigel.) Beck (based on Neottia gracilis Bigel., 1824). However, Fernald found "Elliott (Rhodora 62: 174. 1960) reported this form from Sibley Provincial Park, on the north side of Lake Superior. Unaware of his discovery, on July 22, 1961, Robert L. Jeanne and I discovered this form to be local in a black spruce — sphagnum bog at the head of the north arm cf Fork Bay, at the southeast end of the Sibley Pen- insula. The lin was white, streaked longitudinally with bold purplish lines becoming confluent. The normal form with spotted lip was also in the area. Specimens of f. lineata (Voss 10203) have been distrib- uted to the herbaria of the University of Michigan, University of Toronto, the author, and Sibley Provincial Park. 1966] Monocots — Voss 463 it unnecessary completely to supplant S. gracilis, determin- ing that it applies to a predominantly southern species whereas S. lacera applies to a more northern species not previously distinguished by most authors. Most subsequent authors have also not been able to dis- tinguish the two (e.g., Correll, Nat. Orch. N. A n JL. 1950; Gleason, Ill. Fl. 1: 470. 1952). However, they have neglected Fernald's original point that S. lacera has priority over S. gracilis. Our northern plant, in any event, is to be called S. lacera (Raf.) Raf. Those who find the southern population different may call it S. gracilis (Bigel.) Beck; otherwise, they must call it, too, S. lacera. HERBARIUM, THE UNIVERSITY OF MICHIGAN ANN ARBOR, MICHIGAN NOTES ON LESSER ANTILLEAN FERNS. II GEORGE R. PROCTOR Continuing studies on the Pteridophyte flora of the Lesser Antilles have revealed further taxonomic problems to be solved, and a few more undescribed populations of ferns which require names, The present paper is a sequel to one published in Rhodora in 1961 (vol, 63, pp. 31-35), and arises largely from studies made at the Harvard University herbaria during September and October, 1965, supported by a Mercer Research Grant from the Arnold Arboretum which is gratefully acknowledged. Special thanks are due to Dr. Robert C. Foster for his preparation of the Latin descriptions of new taxa. LYCOPODIACEAE Lycopodium reflexum Lam. var. rigidum (J. F. Gmel.) Proctor, comb. nov., based on Lycopodium rigidum J. F. Gmel. in L., Syst. Nat. ed. 13, 2: 1289, 1791. This variety typically differs from var. reflecum in its more rigidly ascending leaves with less strongly denticulate margins (sometimes nearly entire), and in its slightly larger spor- angia. It has usually been treated as a species distinct from reflexum, but considerable intergradation occurs, and I suspect that some of the differences may be due to the influence of varying growth-conditions; var. rigidum usually grows at relatively higher elevations in more ex- posed situations. HYMENOPHYLLACEAE Hymenophyllum hirtellum Swartz was not credited to the Lesser Antilles by Morton (Contr. U.S. Nat. Herb. 29: 164-165, 1947) in his treatment of the group to which this species belongs. Subsequently, Kramer (in Stoffers, FI. Neth. Ant. 1: 19-20. 1962) attributed H. hirtellum to several Lesser Antillean islands. The material on which Kramer’s records were based is apparently equivalent to Hymenophyllum gratum Fée, more recently (Brit. Fern 464 1966] Antillean Ferns — Proctor 465 Gaz. 9: 73. 1962) reduced to varietal status under H. hirsu- tum by the present writer. It must be conceded that Hymenophyllum hirtellum in its typical form is confined to the Greater Antilles. A closely related population occurring in Guatemala was assigned specific rank (H. maxonii) by Morton, and the Lesser Antilles material somewhat resembles that taxon, even to the occurrence of two rather well-marked varieties similar to those of maxonii, However, the Lesser Antilles plants differ from their Guatemalan counterparts in subtle dif- ferences of texture, particularly in the somewhat coarser hairs on the fronds, and the wide geographic disjunction suggests that the two populations evolved separately, per- haps from the same stock as the present Greater Antillean hirtellum. In assigning names to the Lesser Antilles material, several alternative courses are possible. One of these is to recognize the species H. gratum Fée, and then to subdivide this into two varieties. Objections to this course are the apparent intergradation toward typical H. hirtellum shown by certain specimens, and the difficulty of clearly distin- guishing gratum from maxonii on a morphological basis. Another course is to relate gratum directly with maxonii, but this is objectionable on phyletic grounds. The course adopted is to regard the Lesser Antilles populations as geographic varieties of H. hirtellum. Hymenophyllum hirtellum Swartz, var. gratum (Fée) Proctor, comb. nov., based on H. gratum Fée, Mém. Foug. 11: 118, t.80, Tl, 1866. H. hirsutum var. gratum (Fée) Proctor in Brit. Fern Gaz. 9: 13. 1962. The assignment of this taxon as a variety under H. hir- sutum was due to the close resemblance of some forms to that spe- cies, but the very much larger size and convergence with H. hirtellum now suggests that the latter species probably has a closer relation- ship. Hymenophyllum hirtellum var. vincentinum Proctor, a varietate typica frondibus laxe pendentibus, laminis linearibus vel lanceo- linearibus, ad 27 cm. longis, plerumque 3-4.5 cm. latis (raro ad 6 cm.); stipitibus peranguste marginatis, non alatis, soris 1.2 mm. latis non excedentibus differt. TYPE from southeast slopes of the Soufriére volcano, St. Vincent, 466 Rhodora [Vol. 68 in mossy woodland (Proctor 25987, IJ, isotypes A, U, USF), collected 26 Feb. 1965. Numerous other specimens of this entity have been seen from St. Vincent, where the population is quite uniform and distinct. It is connected with var. gratum, however, by intermediate forms occurring especially in St. Lucia, but also from several other islands. POLYPODIACEAE Diplazium legalloi Proctor, sp. nov. Asplenium callipteris sensu W. Hook. & Bak., Syn. Fil. 231. 1867, in part; Duss, Foug. & Lye. Ant. Fr. 85. 1903, not Diplazium callipteris Fée, 1852. Rhizoma erectum, apice squamis pallide brunneis, ovatis, lacerato- fimbriatis vestitum. Frondes ad 1.7 m. longae, vel ultra, crassae, coriaceae. Stipites ad 60 cm. longi, alte sulcati, basin versus squamati, sparse appresseque arachnoidei. Laminae ovato-oblongae saepius plus quam 1 m. longae, ad 60 cm. latae; rhachis alte sulcata in partem superiorem. Pinnae 9-11 pares, lanceo-oblongae, 12-30 cm. longae, 5-10 em. latae, inferiores breve stipitatae, apicibus acutis vel subfal- catis, marginibus subintegris vel sinuato-crenatis (raro late truncato- lobatis) ; atro-virides, supra glabrae, infra sparse appresso-pilosae. Venae prominulae, in fasciculis, pinnato-ramosae, curvato-conniventes ad margines subcartilaginei. Sori elongati, ad 3 cm. longi; indusium augustum, atrobrunneum, lacerato-ciliolatum. TYPE from the vicinity of Ravine Chaude, above Matouba, Guade- loupe (Proctor 20161, A, isotype IJ), collected Nov. 26, 1959, in the company of Pére C. LeGallo, for whom the species is named. Pére LeGallo, for many years stationed at the Roman Catholic church of Vieux Fort, Guadeloupe, has a wide botanical knowledge of this and other islands, has collected extensively (especially mosses), and is a worthy member of the long and distinguished line of French priest-botanists who (beginning with Charles Plumier) have contributed much to our knowledge of Antillean plants, The species now described as Diplazium legalloi has long been known as D. callipteris Fée. The latter name, how- ever, was based on Linden 233 (ascribed by Fée to Cuba in error), which originated in Venezuela and represents the plant known by the earlier name D. celtidifolium Kunze. Fée himself misidentified the Guadeloupe plant in his trea- tise on Antillean ferns (1866), as his illustration (1.10, f.2) makes clear. D. legalloi differs from D. celtidifolium in its 1966] Antillean Ferns — Proctor 467 usually wider pinnae of a different shape and texture, in the longer sori with lacerate-ciliolate (instead of entire) indusium, and in other details. Diplazium limbatum (Willd.) Proctor, comb. nov., based on Asple- nium limbatum Willd. in L., Sp. Pl. 5: 810. 1810, Asplenium margi- natum L., Sp. Pl. 2: 1082. 1753; Hemidictyum marginatum (L.) K. B. Presl; Diplazium marginatum (L.) Diels, not Blume, 1828. There are no convincing characters by which Hemi- dictyum can be maintained as distinct from Diplazium, but under the latter genus a different specific epithet is neces- sary. Elaphoglossum petiolatum (Swartz) Urban, var. dussii (Underw. ex Maxcn) Proctor, comb. nov., based on Elaphoglossum dussii Underw. ex Maxon, Pter. Porto Rico 398. 1926. Grammitis phlegmaria (J. Sm.) Proctor, comb. nov., based on Polypodium phlegmaria J. Sm. in London Jour. Bot. 1: 194. 1842. Grammitis phlegmaria var. antillensis Proctor, var. nov. Poly- podium flabelliforme sensu Griseb., Fl. Br. W.I. 700. 1864, not Poir. in Lam., 1804. A varietate typica in segmentis angustioribus oblongibusque, marginibus subintegris (non pergibbosis et sinuato-crenulatis) , rhizomatis squamis latioribus clathratioribus, griseo-brunneis (non luteo-bruneis) differt. TYPE from along Trace Victor Hugues near Grande Découverte, Guadeloupe (Proctor 20307, 1J), collected Dec. 8, 1959. Known also from Dominica, Martinique, St. Vincent, and Grenada. Typical G. phlegmaria is chiefly a plant of northern South America. True Polypodium flabelliforme is an older name for P. rigescens Bory ex Willd., and originated from the island of Réunion. Grammitis stipitata Proctor, sp. nov. Subg. Melanoloma. Rhizoma breve, decumbens vel erectum; squamae attenuati-deltateae, ad 3 mm. longae, ochraceo-brunneae, subclathratae. Frondes fasciculatae, in textura firmae, longe stipi- tatae; stipites 4-6 cm. longi, sursum anguste marginati, hirtelli in toto, pilis brevibus, patentibus, pluricellularibus. Laminae anguste lancec-lineares vel lanceatae, 8.5-14 em. longae, ad 1.3 cm. latae infra medium, ad apicem anguste acutae, ad basim cuneatae, nervo primario infra et marginibus atro-scleroticis sparse hirtellis sicut in stipite. Venae plus minusve 1-2-furcatae, liberae vel paucae ad apices con- junctae, margines non attingentes; sporangia glabra. TYPE from Guadeloupe without definite locality (E. A. Marie 6, P, isotypes BM, IJ, NY). Although known only from the type collection, this spe- cies appears to be absolutely distinct. Copeland (in Phil. 468 Rhodora [Vol. 68 Jour. Sci. 80: 258. 1952) mentioned it merely “as an abnor- mal form of G. limbata", but it clearly differs from the latter in its larger, long-stipitate fronds, forked and occasionally anastomosing veins, and hispidulous margins. It seems much more nearly related to G. marginella, but aside from the elongate stipes, the fertile veinlets are not prolonged beyond the sori as they are in that species. Thelypteris consimilis (Fée ex Bak.) Proctor, comb. nov., based on Gymnogramma gracilis “B, G. consimilis" Fée ex Bak. in W. Hook. & Bak., Syn. Fil. 377. 1868. Dryopteris consimilis (Fée ex Bak.) C. Chr., 1907, as to name. D. mollicella Maxon in Proc. Biol. Soc. Wash. 36: 49. 1923. TYPE from Guadeloupe, L'Herminier. It has been customary to place the name consimilis in the synonymy of Thelypteris gracilis, but in the present writer's opinion these are distinctly different species, as shown par- ticularly by the structure of the hairs. No materia] of true T. gracilis has been seen from the Lesser Antilles. It should be noted that in Morton's recent treatment of West Indian Thelypteris with “red” glands (in Amer. Fern Jour. 53: 57-70. 1963), this species was omitted. In his key, it would come out to T. piedrensis, from which it differs in shape of pinnae and especially in having the lower surfaces rather densely puberulous with hamate hairs; in T. piedrensis, underside hairs are practically con- fined to the vascular parts, and are coarser and always acicular. The latter species is apparently known only from Cuba and Puerto Rico, according to available records. Thelypteris cooleyi Proctor, sp. nov. Subg. Thelypteris. Rhizoma erectum, 0.4-1 em. crassum, ad apicem squamis numerosis, lucidis, brunneis, lanceo-attenuatis, minute glandu- losis vestitum. Frondes fasciculatae, in textura elasticae. Stipites comparate perbreves, 1.5-5 cm. longi, minute stipitato-glandulosi, cetera glabri. Laminae oblanceatae, 20-75 cm. longae, 7-15 cm. latae, acuminatae, deorsum longi-attenuatae, pinnis infimis distantibus, au- riculiformibus; rhachis straminea, pars superior bifariam minuteque cincurvo-puberula, cetera glabra. Pinnae plerumque 25-35 pares, oppositae, anguste lanceo-lineares, subaequilaterales, ad basin sessiles 5-15 mm. latae (pinna aerophora parvo, conico subtenta), ad apicem attenuato-faleatae, alte pinnatifidae, glabrae, Segmenta subobliqua, deltato-oblonga, plerumque ad basin 2-3 mm. lata, acuta; segmentum infimum acroscopicum pinnae plerumque dilatatum, ovatum, rhachi- dem excessum, venis furcatis et marginibus serratis; venae segmen- 1966] Antillean Ferns — Proctor 469 torum plerumque in paribus 4-6, simplices; margines segmentorum fertilium plerumque revolutae, soros in parte obscurantes. Sori sub- marginales; indusium rotundato-reniforme, ca. 0.8 mm. latum, rufo- brunneum, minute stipitato-glandulosum, persistens; sporangia glabra. TYPE from outer slopes and rim of the Soufriére crater, St. Vincent (Proctor 26008, 1J, isotypes A, U, US, USF), collected 26 Feb. 1965 in company with Mr. George R. Cooley, for whom the species is named in recognition of his contributions to West Indian botany. Paratypes (all from St. Vincent) Cooley 8214, 8417, 8445, and 8591; also Proctor 25985. Thelypteris cooleyi is closely related to T. limbata (Sw.) Proctor, but differs from the latter in its narrower fronds always of clearly definite growth, in its glandular rhizome- scales whose cells are smaller and less transparently cla- thrate, in its entire ultimate segments, with the sori not located on teeth, and in its much paler indusium. Further, the tissues of T. limbata are characteristically beset with reddish or yellowish, sessile, resinous glands on the under- side, while such glands are entirely absent from T. cooleyi. INSTITUTE OF JAMAICA, KINGSTON, JAMAICA A NEW SPECIES OF ASTERELLA FROM MEXICO R. M. DEL ROSARIO! Asterella evansii del Rosario sp. nov. Monoica et tenuis frequenter viridis omnino sed interdum tincta purpureo, praesertim in super- ficie ventrali. Thalli usque ad 15 mm. longi, 4 mm. lati, ramificantes furcalis, interdum simplicibus ex apice innovatis aliquando praesenti- bus. Thalli plani vel parvo concavi cum margine crispato et parvo sinuato. Cellulae epidermidis cum tenuibus parietibus sed interdum cum aliquantum distinctis trigonis. Stomata 80-90 u longa et 65-80 u lata, cireumventa 6-7 seriebus cellularum, cum 3 cellulis in quibusque seriebus et cum radialibus parietibus parvo crassatis aliquantum dis- tinctis minutisque trigonis. Squamae ventrales aliquantum purpureae et interdum hyalinis, subulatis et inconstricto basi, acutis usque ad acuminatis, totis vel aliquantum dentatis. Pedunculus cum sparsis paleis tenuibus et parvo confertiore fasciculo in apice, viridosus sed interdum purpurarius usque ad fuscasus basi, circa 1 cm. altitudine. Carpocephalum lanum vel fere planum cum parva media parte parvo elatiore, cum inferis et salebrosis tuberculis; involucra dubie sinuata- crenata, vix attingentia margines loborum, Pseudoperianthia alba vel sufflava et interdum tincta purpureo et directa in obliquum desuper. Sporae fulvasae, 60-80 & diametro cum tota superficie tecta subtile et incomposito reticulo cum retiaculis 1-8 wu latis. Sporae minute punctulata. Elateres fulvosi, 200-500 u et 8-10 x lati, etiam plurime bispirales omnino. Thalli 0.5-1.5 cm. long, 1.5-4 mm. wide, thin, usually green through- out but sometimes more or less tinged with purple, the pigmentation usually more extensive in the ventral surface, branching by forking, sometimes simple, apical innovations sometimes present, plane or slightly concave, with slightly sinuate crispate margins, not incurved when dry, keel broad and rounded. Epidermal cells averaging 40-60 qu, with thin walls but sometimes with more or less distinct trigones, cells containing oil bodies 10 or fewer, scattered. Pores slightly ele- vated, 80-90 „u in length, and 65-80 & in width, surrounded by 6-7 ‘Philippine National Herbarium, National Museum, Manila, I wish to express my appreciation to Dr. Ronald L. McGregor, head of the Botany Department, University of Kansas for his suggestions and criticisms. To the curators of the herbaria of New York Botanical Garden and U. S. National Museum, I am indebted for the loan of specimens. I must also extend my thanks to the National Museum of the Philippines and the University of Kansas whose financial aid made possible this study. To Prof. Austin Lashbrook, Classics De- partment, University of Kansas, my sincere thanks for editing the Latin diagnosis. 470) 1966] Asterella — del Rosario 471 THALLUS IN VENTRAL (x8) VIEW (x8) THALLUS WITH FEMALE RECEPTACLE Asterella evansii del Rosario, n. sp. series of cells with 3 cells in each series, radial walls slightly thickened with more or less distinct trigones. Green tissue loose, air chambers sparingly subdivided by supplementary partitions. Ventral tissue composed of cells with thin unpitted walls. Ventral scales more or less purple, sometimes with hyaline margins and appendages, with 10 or fewer oil bodies, appendages 1-2 subulate from an unconstricted base, 0.25-0.4 mm. long and 0.12-0.18 mm. wide, acute to acuminate, entire or somewhat dentate with occasional oil bodies in the ventral portion. Paroicous; the antheridia forming a small and irregular median group near the peduncle of the female receptacle, the ostioles low. Peduncle with scattered slender paleae and a slightly denser cluster at the apex, greenish but sometimes purplish to brownish at base, about 1 cm. high. Disk of receptacle, 3-4 mm. wide, flat or nearly so, with a small slightly elevated center, with low but coarse tubercles, rugose when dry, normally 4-lobed, the lobes spreading almost horizontally, the involucre vaguely sinuate-crenate, barely reaching the margins of the lobes. Pseudoperianth white or yellowish, sometimes pigmented with purple, directed obliquely downward, 8-10- cleft, the divisions narrowly lanceolate, connate at apex, operculum remaining intact at dehiscence. Spores yellowish-brown, 60-80 & in diameter, with wavy crenulate wings, 8-14 u wide along the edges, the entire surface covered over with a fine and irregular reticulum, the meshes 1-8 & wide, the surface, in addition, minutely punctulate. Elaters yellowish-brown, 200-250 & and 8-10 & wide, variously coiled, mostly bispiral throughout. Growing on banks. Distribution: Mexico. Type locality: Canada Santa Magdalena, Mexico. SPECIMENS CITED — Cañada Santa Magdalena, 1908, Barnes and Land 441 (NY, holotype, mixed with Targionia hypophylla L.); 472 Rhodora [Vol. 68 Asterella evansii del Rosario. n. Sp. © (x-190) VENTRAL SCALES (x-200) Amecameca, 1908, Barnes and Land 339 in part (NY, mixed with Targionia hypophylla L.); banks along Avenida Hidalgo and path to Teziutlan, Puebla, Barnes and Land 555 (NY). Since 1920, the specimens of this new species have been identified as A. venosa, in spite of the fact that the charac- teristics of their epidermal pores, ventral scales, and spore markings differ from those of the latter species. Evans (1920) was unable to study the type of A. venosa. Its dis- tinguishing characteristics which were listed in an annota- tion made in 1955 are as follows: The radiating series of cells around the pores are 4-6 and the radial walls are not thickened. The appendages of the ventral scales are constricted in the base and the tip is blunt or obtuse with one or two apical cells. In general, the cells of the append- ages are globose and do not have oil bodies. The meshes of the spores are larger, 10-24 in diameter. On the other hand, in the new species, A. evansii, the radiating series of cells around the pores are 6-8 and the radial walls are thickened. The appendages of the ventral scales are apicu- 1966] Asterella — del Rosario 473 È &, ARS E SPORES INNER FACE QUTER FACE (x 810) (x 914) Asterella evansii del Rosario, n.sp. lated, unconstricted at the base and have oil bodies. The meshes of spores are much smaller, being 1-8 p wide. There is another specimen identified also as A. venosa which was collected in Venezuela by Chardon in 1932. I determined this specimen as A. macropoda, which is a South American species. Its distinguishing characteristics are a larger thallus with lateral intercalary branches, larger epidermal pores, autoicous inflorescence, larger disk of female receptacle, and yellowish-brown pseudoperianth, and 12-16 cleft. The spores are immature but the charac- teristic brown color is well shown. A. macropoda has coarsely reticulate spores 20-30 , in diameter. Thus, A. venosa is again restricted to Brazil with possible occurrence in other parts of South America. The extension of the range of this species to Mexico was mainly based on the specimens described above which, as said earlier, were incorrectly determined. A. evansii is named after the late Alexander W. Evans who was the leading authority on the genus in North America. PHILIPPINE NATIONAL HERBARIUM NATIONAL MUSEUM, MANILA LITERATURE CITED Evans, A. W. 1920. The North American Species of Asterella, Contr. U. S. National Herbarium 20: 286. RUBUS SUBGENUS EUBATUS IN NEW ENGLAND A CONSPECTUS! ? A. R. HOGDON AND FREDERIC STEELE This paper is an attempt to define the species of black- berries in New England, to produce a workable key, and to assess the previous work on the group in our area. It is not intended to be a monographic treatment, as no attempt has been made to survey the subgenus throughout its entire geo- graphic range. It is rather a provisional treatment for the New England area. Rubus is an inherently difficult genus, as the different species intergrade in a perplexing manner. More cytotaxonomie and genecological information is needed before a definitive treatment can be made. Never- theless, 15 years of field study have convinced the authors that there are evolutionary populations that can be treated as species by competent botanists in the field, or in the her- barium, providing there is adequate data. Although we feel that our species are “good,” further research, especially on hybridization and introgression, might reduce the number slightly. The first major step in taxonomy certainly must be to establish a reasonable assemblage of taxa that are well enough defined to permit botanists to communicate facts and ideas about them. This premise appears so reasonable that it should not be necessary to mention it at this time. However, our blackberries have become essentially unwork- able taxonomically largely through the reliance of authors solely on morphological criteria for the interpretation of species. As a result the circumscriptions of new taxa have been related to single specimens rather than populations. There has resulted a multiplicity of "species" a large pro- "Published with the approval of the Director of the New Hampshire Agricultural Experiment Station as Scientific Contribution No. 406. "This research has been supported by the New Hampshire Agricul- tural Experiment Station and by grants from the Central University Research Fund of the Graduate School of the University of New Hampshire and from the Northern New England Academy of Science. 474 1966] Rubus in New England — Hodgdon & Steele — 475 portion of which have no validity in evolution as such and actually are expressions of intraspecifie variation or else are interspecific hybrids of many kinds. Hodgdon and Steele (1962) showed that glandularity in Rubus alleghen- iensis is a very variable character which makes it difficult to separate other “species” from R. allegheniensis on grounds of glandularity. More recently Steele and Hodgdon (1963) demonstrated that a number of so-called species could be referred to hybrids of Rubus hispidus and R. setosus. A large number of field observations and labora- tory measurements support our belief that nearly all of our species of blackberry hybridize in nature. Also it is quite apparent that there is much intraspecific variability throughout most of the group involving such characters as the type of inflorescence, the quality and number of prickles and bristles, the stature and growth habit, as well as gland- ularity. Apomixis is a further complicating factor in the taxonomy of blackberries. In Europe Gustafsson (1946, 1947a, b) showed that Rubus fruticosus L., considered as a collective species, contained both amphimictic and apom- ictic lesser taxa, the apomixis often being facultative. Com- menting on the implications of apomixis in taxonomy Valentine & Lóve (1958) wrote, “In many apomictic groups, very large numbers of species have been described taxo- nomically: the fact that each individual may be true-breed- ing means that discrimination can be much finer than in the amphimictic groups. As demonstrated by Gustafsson (l.c.), the resulting classifications vary a great deal in their usefulness and validity. Once a taxonomist dealing with a particular group has decided on his criteria he must main- tain them; and in apomictic groups, this may lead to the description and naming of very many species which are very local or even ephemeral.” Einset (1951) worked with 24 “different forms of Rubus” determined by L. H. Bailey and grown at the New York State Agricultural Experiment Station at Geneva, New York. The chromosome numbers of parent plants were de- termined and various pollinations were tried. Einset then grew seedlings and made chromosome counts of these. He 476 Rhodora [Vol. 68 found that from 80-100% of the progeny resulted from the parthenogenetic development of unreduced eggs. He con- cluded that the principal type of apomixis occurring was pseudogamy with the unreduced eggs developing partheno- genetically. He noted also that some hybrid offspring were produced, indicating facultative apomixis. Vegetative spreading also may take place in blackberries with great rapidity. Considerable areas thus can be occu- pied by populations, depending on the length of time in- volved and barriers to spreading, With the fragmentation of such colonies a single clone may come to occupy a suffi- ciently extensive area in such a way as to suggest that it be recognized as a distinct “species” in the flora. It is evident to us that the “species” that have been de- scribed, variously fit all the categories discussed above; some are “good species”, some are better treated as falling within the range of variations of these “species”, others are recognizable hybrids between these “good species" while still others are more obscure populations whose status is yet quite uncertain. Until the biological nature of these popula- tions can be ascertained we are not ready to accord the lesser ones taxonomic status. The fact that many of these populations will be shown to be of hybrid origin and are thus distinct and that they tend to produce essentially sim- ilar offspring by apomixis or vegetative spreading, does not mean to us that they should be regarded as species, How- ever distinct the hybrid population may have been when it first appeared, we don't consider it to be a species unless it has undergone subsequently a recognizable amount of evolu- tionary change. Such change is expressed in the develop- ment of evident and distinct morphological characteristics and those adaptive traits which permit the population to occupy a definite geographic range. Our work on Rubus started with an attempt to compile a list of species for New Hampshire. At one time there was much interest in creating hybrids of horticultural value in Rubus at the University of New Hampshire under the di- rection of the late Dr. Albert F. Yeager. Several phases of research were completed which are of interest to taxono- 1966] Rubus in New England — Hodgdon & Steele — 477 mists. One by Dr. Donald Craig (1959) related to our pres- ent study. It demonstrated that R. canadensis, a triploid, could be crossed with various blackberry cultivars. At the beginning of our study we used the eighth edition of Gray’s Manual expecting confidently to locate each spe- cies that Fernald attributed to New Hampshire. This soon led us to try out Bailey’s elaborately presented studies in Gentes Herbarum (1932-49). Both treatments helped us make a start beyond which they became frustrating and, particularly true of Bailey, essentially unworkable. Because of the great prestige and massive productivity of both Fer- nald and Bailey we were too willing to accept their “species” without reservation but very soon we perceived that by relying solely on morphological criteria for “setting up” new species the end was by no means in sight, for if we wanted to add to the plethora of species, we too could pro- ceed on the same basis and describe still more. Our field studies and collecting have taken us throughout most of New England and also led us into parts of New Brunswick, Nova Scotia, and New York. The most signifi- cant omissions have been upper Maine, where there is rea- son to believe the number of species is relatively small, western Vermont and western Massachusetts. We have studied in the field and made collections of all indigenous species that we are recognizing in this treatment except Rubus cuneifolius. We have taken pains to make sure that all our collections of blackberries have careful descriptive notes on the growth-habit and include both primocanes and floricanes. To provide evidence of our research we have ac- cumulated large collections of Rubus at the University of New Hampshire (NHA) and in Tamworth, N. H. (private herbarium of Steele). Necessarily a considerable part of our work has been de- voted to the study of herbarium specimens and particularly in recent years, to a careful evaluation of nomenclatural types. We wish to express our appreciation to the curators and staffs of the following herbaria which we have con- sulted, the Gray, Arnold Arboretum, and New England 478 Rhodora [Vol. 68 Botanical Club herbaria at Harvard University, the Pringle Herbarium of the University of Vermont, the Jesup Her- barium of Dartmouth College, and the Bailey Hortorium collection of Cornell University. Bailey and Fernald set a high standard of collecting which often has been main- tained by those who have come after them, but a very large proportion of the specimens in most herbaria are not sub- ject to critical determination, since they are usually without habit notes and very often consist only of primocanes or floricanes, not both. Thus the value of these large collec- tions in elucidating the taxonomic problems in the group is somewhat less than one might expect. Our principal ap- proach has been the study of populations in nature and the collecting of much appropriate material followed by inten- sive herbarium study consisting of rather exact measure- ments of parts, the tabulation of results and the subsequent analysis of the population. One test of the validity of scien- tific work is predictability. Now with our background of experience in observing populations and analyzing them, when we enter a characteristic type of environment and find certain kinds of blackberries present, we can often antici- pate finding the putative parents or hybrid offspring in the same general area. Taxonomic work on Rubus subgenus Eubatus in New England in this century has been carried on by an interest- ing group of botanists. William Henry Blanchard (1850- 1922) a farmer and country schoolmaster of southern Vermont described many species, amongst which we are recognizing the following in this paper: R. recurvicaulis, R. arenicola, R. semisetosus, R. vermontanus and R. elegan- tulus. Fernald (1950) accepted 19 additional Blanchard species. Several of these we interpret as rather commonly occurring hybrids having distinctive morphological features by which they can be recognized, R. jacens and R. tardatus for example. Blanchard had a keen collector’s eye but most of his speci- mens lack habit-notes. Also his method of collecting in- volved the grouping of several separate collections under 1966] Rubus in New England — Hodgdon & Steele 479 one collection number. This applies to types as well as to other specimens and make his specimens confusing to study. Bailey has selected lectotypes from Blanchard’s “types” for most of the critical material. Ezra Brainerd (1844-1924), also from Vermont, was at one time the President of Middlebury College and he had established a botanical reputation for his work on the genus Viola. His interest in blackberries largely was related to their breeding behavior and in fact his chief contribution was his recognition of the importance of hybridization in the group. With A. K. Peiterson (1920) he published a very interesting and suggestive paper on the subject. Essentially they recognized 12 basie species in New England which were postulated to have crossed and given rise to all the myriad of forms to be found in the area. Our chief criticism of their work centers around the inadequate evidence that was advanced to support their conclusions. A large part of the statements made about the parentage of their hybrids seem to be hypothetical and not proven. Also the taxonomic basis for the paper was faulty ; for example they recognized R. Baileyanus Britt. which we have been unable to confirm and they failed to distinguish R. recurvicaulis Blanch. (R. plicatifolius) which we find to be fully distinct as a species. A. K. Peiterson (1921) published a study of variation and hybridization in the blackberries. He performed transplant experiments to study the effect of environment on growth habit and armature. The indication was that plants grown in full shade were more erect than those in sun, the latter showing a marked tendency to trail. Primordial prickles and hairs were present on all blackberries. Their develop- ment is controlled by both genetic and environmental fac- tors. In one experiment, suckers from the root of a plant grown in full sunlight produced many more prickles and glandular hairs than suckers from the same plant grown in shade. Infertility of blackberries, due to defective pollen, was found to be common. The two extremes, R. alleghenien- sis and R. Baileyanus, showed less infertility than any others possibly indicating that they were primitive species. 180 Rhodora [Vol. 68 Many others may have been of hybrid origin. Eleven spe- cies were used in hybridization experiments and all crosses attempted were successful even though some species were remotely related. As a result of these experiments, Peiter- son concluded, and the authors of this article concur, that hybridization is common in the Eubati and that a large number of so-called species are hybrids. Although Peiterson's work was significant, he did not preserve voucher specimens of his crosses and it is difficult to know what the actual results were in terms of the species as we interpret them. Observations of herbarium material in the Pringle Herbarium at the University of Vermont in- dicate that several of Peiterson's species represent a varied assemblage of plants. For example any low blackberry of the Flagellares section may be designated as R. Baileyanus and any of three species may be designated as R. frondosus. It would be highly desirable to repeat the experiments pre- serving voucher specimens at all stages. Nonetheless had Bailey and Fernald accepted Brainerd's and Peitersen's ex- planation for the great diversity of blackberries as attribu- table to crossing, the study of Rubus subgenus Eubatus would probably have been brought into some order before now, In Gray's Manual (1950) are to be found two species of Brainerd’s, R. severus and R. sceleratus both of which are highly localized aberrant populations presumably of hybrid origin which we are not recognizing as species. The name of Eugene P. Bicknell (1859-1925) must be introduced in relation to Blanchard and Brainerd. Bicknell (1910) published an article entitled *Have we enough New England Blackberries." He recognized 11 species including Rubus flagellaris Willd. which up to that time had not been interpreted properly. He accepted none of Blanchard's out- pouring of species. Of the then recently published species of Blanchard and a few other newly named species proposed by other authors he made the following comment, “It would scarcely yet be the part of wisdom to accept any one of these new names as denoting a valid species nor, on the 1966] Rubus in New England — Hodgdon & Steele 481 other hand, is there sufficient warrant in our present knowl- edge for separating all of them as being without standing. I should suppose however, that some sixty per cent of the number might be allowed to pass into the category of sym- onyms; the remainder, possibly with a few exceptions, ap- pear to disclose themselves as scarcely doubtful hybrids". ]t is ironie that Bailey (1941) named a species for Bicknell — a very localized blackberry of Nantucket which most certainly is of hybrid origin. Bailey concluded his remarks following the description of R. Bicknellii with the state- ment *. ... ; his publication had much influence on the Rubus work of Rydberg and Brainerd”. Yet Bicknell in the afore-mentioned discussion of New England blackberries had acknowledged Brainerd’s influence in interpreting the great diversity of blackberries in York County, Maine as the result of hybridization. Thus there were in the early years of this century two schools of thought concerning Rubus which have continued to the present. Per Axel Rydberg (1913) tended to follow the conserva- tive approach of Bicknell and Brainerd. However, for New England, he recognized 25 species, 12 of which we still maintain. He was more discerning than Bicknell in evalu- ating Blanchard’s species: we find him accepting Rubus ver- montanus, R. arenicola and R. elegantulus. He recognized R. plicatifolius of Blanchard which was published at the same time as was R. recurvicaulis, also Blanchard's per- gratus which now is considered to be synonymous with Poiret’s pensilvanicus and finally Blanchard's recurvans which we consider a state of frondosus. He followed the practice of Bicknell in listing the possible hybrids for each species. The diffieulty with this approach to the problem of hybridization lies in its speculative quality; no evidence was provided and no specimens cited. Liberty Hyde Bailey (1858-1954) did a prodigious amount of work on Rubus. He improved the terminology, devising such terms as primocane, floricane, parcifrond and novirame. He brought order and method to the study of the 182 Rhodora [Vol. 68 group and he established an herbarium of the finest quality at Cornell University. Bailey proposed five of the seven native Sections of Rubus subgenus Eubatus that are often recognized in New England. These are Flagellares, Setosi, Cuneifolii, Canadenses and Alleghenienses. He provided a scholarly basis for further taxonomic work by clarifying types and designating them where necessary. His major treatments in Gentes Herbarum are clearly prepared and carefully illustrated. Because of the orderliness and com- pleteness of his work it is easy to see his shortcomings. As pointed out by Steele & Hodgdon (1963) pp. 262, 263, Bailey refused to recognize hybridization as a taxonomic problem in blackberries. Also he paid scant attention to the variabil- ity within the species. Since there are no evident limits to variability in the group L. H. Bailey described hundreds of new species. Some of the folders at the Bailey Hortorium bear evidence that Dr. Bailey became confused at times ; too often the other specimens in the folder fail to resemble the types. Bailey never accepted Blanchard's Rubus elegantulus although this is without doubt one of the most abundant and distinctive species throughout central and northern New England and eastern Canada. The only one of Bailey's spe- cies that we accept is Rubus Jaysmithii which is readily sep- arable from its closest relative R. flagellaris by its pubescent leaves. Unless other distinctions can be found it may be bet- ter eventually to treat R. Jaysmithii as a lesser taxon. M. L. Fernald (1873-1950) also made substantial contrib- utions to our knowledge of Rubus. Although in a footnote (p. 822 in Gray’s Manual 8th Ed. 1950) he recognized that hybridization was rife and accounted for many of the mor- phologically distinctive trends, in only one case (R. ab- brevians) did he designate a species as a probable hybrid. This is in marked contrast to his treatment of Salix, for ex- ample, where he described a number of hybrids, and indicated specifically a great many crosses. Fernald estab- lished the Section Tholiformes and described Many new species in it, several of which were confined to central and northern New Hampshire, others more widely dispersed in 1966] Rubus in New England — Hodgdon & Steele 483 the Northeast and with some local species elsewhere. Fer- nald also described new species in several of the other Sec- tions. He was somewhat more discriminating than Bailey in the sense that his “species” were more likely to be well defined morphologically. In an earlier paper on hybrids (1963) we indicated that we considered some of Fernald's species to be hybrids of Rubus hispidus and R. setosus. We were able to locate R. aculiferus Fern. which belongs in this category of hybrids at North Woodstock, New Hampshire. On Cape Cod we succeeded in finding Rubus paludivagus Fern. which we consider to belong to R. argutus. ln Fer- nald's treatment (1950) the New England species fall into eight sections. The Tholiformes seems to be a collection of hybrids and aberrant forms without any common unifying factor. The other sections represent reasonably natural di- visions with some overlapping at times but, because in New England several would contain only one or two species, there has seemed little point in our recognizing them. We see no merit in regarding the sections as superspecies, which seems to be an artificial concept of doubtful convenience. Fernald's major contribution in Rubus seems to have been his often sensible evaluation of the work of others such as Blanchard, Brainerd, and Bailey. He seemed to know blackberries in New England better than Bailey did. He was fully aware of the distinctiveness of Blanchard's R. elegan- tulus for example. Fernald screened out a vast array of Bailey's minor "species" though it must be admitted that he might better have omitted many others along with a number of his own making. Through experience we have found that Fernald's treat- ment in Gray's Manual with all its prolixity, provides the best starting place for taxonomic work on the group in New England. The recent handling of our blackberries by Gleason and Cronquist (1963) reflects the commendable decision of the authors to reduce the group to workable size. However the treatment, so far as New England, is concerned, is not sat- isfactory. The following species are omitted, Rubus elegan- 484 Rhodora [Vol. 68 tulus, R. vermontanus and R. semisetosus while R. Enslenii is stated to extend north only to Pennsylvania. Two other species which we do not recognize are given for New Eng- land, Rubus orarius and R. ostryifolius. Longley (1924) and Einset (1945) have reported chro- mosome numbers of many species of Rubus. Many of Long- ley's determinations must remain in considerable doubt since there is much question as to the identity of his species and the chromosome numbers are often at variance with the better documented work of Einset whose plants were identified by L. H. Bailey. It seems clear from Einset's findings that Rubus allegheniensis and R. setosus (R. Boot- tianus) have 2N—14 chromosomes, that R. canadensis is a triploid with 3N—21 while both R. flagellaris and R. recurv- tcaulis have 63 chromosomes. Other species reported by Einset were all polypoids, some of them with variation in chromosome number. Plants of R. bellobatus for example, reported to have been collected from two different localities near Ithaca, had chromosome numbers of 21 and 28. Einset reported Rubus abactus with 35 chromosomes and R. avipes to have 21. Both in our opinion belong in the species R. pen- silvanicus. We have been able to identify most mature Rubi that we encounter in the field by means of the present treatment, (providing the primocanes and floricanes are both present) except for young plants that have been excessively stunted by mowing. Many plants encountered would be classified as hybrids in which ease it is nearly always possible to specify one or both putative parents. It is hoped in a future article to describe the more common hybrids so that they can be recognized. It is not possible to identify all herbarium specimens. Early collectors, notably Blanchard and Brainerd, did not include growth-habit notes on the label. Thus a guess as to growth-habit, from the appearance of the plant on the sheet, must be made. At best this is an unsatisfactory procedure and usually is quite impossible. Many collections contain either primocanes or floricanes but not both. Little can be 1966] Rubus in New England — Hodgdon & Steele 485 done with these. The most characteristic part of the primo- cane in the middle part of the cane with two or three at- tached leaves. When only the tip is collected, identification is less certain, and in the case of sheets consisting of mere fragments it is impossible. The characteristics used in the description of Rubus re- quire special terms. Growth-habit is the most important single characteristic for major division of species. The pri- mary distinction is between upright and trailing plants. It should be noted, however, that normally erect plants will sometimes become bent over at the edge of a colony and may reach the ground. Species that are normally trailing may arise to a considerable distance off the ground if supported on vegetation. Young primocanes are often erect and do not attain their trailing habit until late in the season. The terms used by us to describe growth habit are erect, arch- ing, doming, trailing, and prostrate. Erect plants are nearly vertical. Doming plants are 1’ to 3’ high and curve so that their tips reach the ground. Prostrate plants are in contact with the ground for their full length, whereas trailing ones start off as low-domers. The term reclining is used for flori- canes that are close to the ground, apparently having been lodged by snow. The nature of the prickles is distinctive. At one extreme are the stout broad-based prickles usually 5 mm or more long. At the other extreme are acicular prickles with slight- ly dilated bases, but needle-shaped. Thin prickles are inter- mediate between the two. Bristles are no thinner than acicular prickles but are soft to the touch. Prickles if di- rected backwards are hooked if curved, retrorse if straight. In the treatment to follow two introduced species are in- cluded in the key but otherwise the study is confined to na- tive Rubi. It must be recognized that cultivars may escape to the wild and persist. At present introduced blackberries do not constitute a major taxonomic problem in New Eng- land. KEY ro RUBUS SUBGENUS EuBATUS IN NEW ENGLAND Note, hybrids can not be identified with the key although it may provide a clue to their origin. 486 Rhodora [Vol. 68 Inflorescence mostly a raceme or resembling a cyme or corymb, if paniculate, lacking abundant prickles on pedicels; indigenous species. Mature primocane doming, trailing or prostrate. Primocane low-doming, long-trailing or prostrate. Primocane bristly without strong prickles; leaflets usually 3, coriaceous and lustrous. .....ccceccccscessssssessessscerseesees 6. R. hispidus Primocane with few to many strong and hooked or straight and short prickles, rarely setose; leaflets 3 or 5, not coriaceous. Cane very slender, 1-2 mm in diameter; inflorescence 1-2- flowered; plant of oak-or oak-hickory-woods. .. 3. R. Enslenii Cane slender to medium, 1.5-4 mm in diameter; well-devel- oped inflorescences 1-12 flowered. Pedicels erect, up to 6 (averaging 4) cm long; inflores- cences 1-3 (5)-flowered. Leaves glabrous or subglabrous beneath ....... ees seesssseeaasecececeeeesesessenessssesseceesssseesesasessececesens 1. R. flagellaris Leaves pubescent beneath eec. 2. R. Jaysmithii Some pedicels ascending or divergent, central ones short (.5 cm); inflorescences 1-12 flowered. Leaves glabrous or nearly so beneath .................... —— RR 4. R. recurvicaulis Leaves soft-pubescent or velvety beneath ............. /— — ERR 5. R. arenicola Primocanes low-or high-doming, tips often reaching ground. Canes slender, 1-4 mm in diameter, armed with thin prickles or bristles. Canes armed with hooked or thin and retrorse sharp prickles .....................ss. 4. R. recurvicaulis or 5. R. arenicola Canes armed with bristles or straight acicular prickles 100- 200 per decimeter ............ 7. R. setosus or prickly forms of 8. R. vermontanus 9. R. semisetosus and 12. R. elegantulus Canes stout 3-9 mm in diameter, armed with strong straight prickles .. 15. R. frondosus or extremes of 13. R. alleghensis Mature primocanes arching or erect. Well developed plants low, 0.3-1 m high, armed with bristles, acicular prickles or hooked prickles; leaves medium-sized averag- ing 8-18 em broad. Leaves glabrous beneath. Armature of numerous soft bristles ................... 7. R. setosus Armature of acicular prickles resistant to touch. Central petiolule 1.4-2 em long. .......... 8. R. vermontanus Central petiolule 2.5-4 cm long. .......... 12. R. elegantulus Leaves pubescent beneath. Leaves pilose to velvety beneath, armature of acicular priekles. ..........ssssseeeeeeeeeeeee retenue 9. R, semisetosus 1966] Rubus in New England — Hodgdon & Steele 487 Leaves densely tomentose beneath, armature of strong, stout hooked prickles, ........... ene 10. R. cuneifolius Well-developed plants 1-2 m high; leaves large, averaging 12-24 cm broad. Leaves glabrous beneath. Armature of 0-10 thin prickles per decimeter. .... DOE ATP CUM, cas E A E 11. R. canadensis Armature of 10-100 prickles per decimeter. ....... nm 12. R. elegantulus Leaves pubescent or velvety beneath. Axis of inflorescence pubescent and with numerous stipitate glands. EE 13. R. allegheniensis Axis of inflorescence pubescent, glandless or rarely with glands. Leaflets lanceolate, ratio of width to length 0.5 or less, base rounded to cuneate. ........eee HH 14. R. argutus Leaflets ovate, ratio greater than 0.5, base cordate to rounded. Leaves chartaceous or subchartaceous; prickles thin to stout, 4-6 mm long, often numerous, 4-60 per dej. c.m Riu OAM TUNE 15. R. frondosus Leaves subchartaceous to membranaceous; prickles thin to medium, 2-4 mm long, usually few 0-20 per dE esee 16. R. pensilvanicus 4écanadaéasaedenoseseteecasasqabsenacsovesqevonadseeeqeovortédeeceuón Inflorescence a large terminal panicle, its branches and pedicels with abundant prickles; introduced species of southern New England. Leaflets deeply cut, green beneath. ........ HA R. laciniatus Leaflets merely toothed, white-velutinous beneath. ........ R. bifrons 1. Rubus flagellaris Willd. PRIMOCANE trailing to prostrate (some- times climbing over low vegetation), commonly tip-rooting; FLORI- CANE trailing to prostrate; ARMATURE of hooked prickles 1-3(4) mm long, 5-50 per dm, occasionally more; DIAMETER OF CANES 1.5-3 mm; PRIMOCANE LEAFLETS mostly 3, sometimes partly divided or 5 in number, mostly chartaceous, ovate or rhombic to suborbicular, the ratio of width to length 0.6-0.8, abruptly acuminate to apex, rounded to base, glabrous above, glabrous or very lightly pubescent on veins beneath; DIMENSIONS OF COMPOUND LEAF 66-1313 cm; INFLORES- CENCE a narrow slender raceme 6-11 cm long with 1-4 (5) flowers on nearly erect long pedicels, lowest and longest pedicels 3-4 (5) em long, the uppermost 1-2 cm; axis of inflorescence and pedicels usually somewhat pubescent and often prickly, not bearing any glands; early flowering; FRUIT globular or subglobular, ripening early, often of good quality; HABITAT, fields, sandy areas, roadsides and ledges; RANGE, central Maine south and west, common at low elevations. 488 Rhodora [Vol. 68 Five species in this treatment: R. flagellaris, R. Jay- smithii, R. Enslenii, R. recurvicaulis and R. arenicola belong to the section Flagellares of Gray’s Manual. Of these five, the first three appear to be closely related, intergrade fre- quently, and could, with some justification be considered to form one superspecies. When they are stunted because of mowing, or from an unfavorable site, or when they are not fruiting well, they are difficult to separate. However, when well developed with good floricanes, they can usually be readily identified in the field. The basic division is be- tween the flagellaris group and the recurvicaulis group. Further study may indicate that R. Jaysmithii and R. areni- cola would better be considered as varieties of R. flagellaris and R. recurvicaulis respectively. They are not well repre- sented in herbaria, but are often encountered in the field, where they can usually be recognized. It may be assumed that all of these species hybridize with each other on occa- sion, but it would be difficult to establish with certainty the nature of the hybrids. In identifying members of the group, it should be noted that new primocanes start as arching shoots and do not develop trailing tendencies until the middle of the season. In favorable sites, such as new roadside soil where there is no competition, the primocanes are so vigorous that they develop mounding tendencies, thus falling in the category of the section Tholiformes so far as habit is concerned. On bare gravel or ledge, the primocanes may become excessively long trailing. This seems to be a direct response to en- vironment and is of no diagnostic significance. By mid- August it is common to find vigorous primocane colonies in open areas with masses of canes and few or no fruits. These cannot be identified as to species. All of these species have small sharp prickles, usually hooked, and are unpleasant to touch. They usually can be recognized as belonging to the flagellares group by this character alone. Under favorable conditions they have fruit of good quality, large and sweet. Because of the prostrate habit and numerous prickles, the fruit is not as easily har- 1966] Rubus in New England — Hodgdon & Steele 489 vested as that of the erect species, but it is often of better quality. As indicated in the key, Rubus flagellaris is distinguished from R. recurvicaulis primarily by the inflorescence. In addition it can be noted that R. flagellaris tends to be lower than R. recurvicaulis and trailing or prostrate, while the latter may be high trailing or doming. Rubus flagellaris may trail off the ground on vegetation but will not, under these conditions, get as high above the ground as R. recurvi- caulis. There also may be a difference in leaves with those of flagellaris being smaller and duller. As indicated above, hybridization is undoubtedly frequent within the flagellares group. These are indications that Rubus flagellaris hybridizes with several species in other sections but these hybrids are uncommon. REPRESENTATIVE SPECIMENS: MAINE: SOMERSET CO., Fair- field, Fernald & Long 13901 (NEBC); ANDROSCOGGIN CO., Auburn, July 21, 1924, Bean (NEBC) ; CUMBERLAND CO., Standish, Fernald & Long 13908 (NEBC); YORK CO., Arundel, August 3, 1905, Blanchard (NEBC, as R. geophilus Blanchard) ; Alfred, Fernald & Long 13897 (NEBC). NEW HAMPSHIRE: COÓS CO., Whitefield, Pease 37861 (NEBC) ; CARROLL CO., Bartlett, Steele 3496 (STEELE) ; Tamworth, Steele, 2780 (STEELE); STRAFFORD CO., Milton, Hodgdon & Steele 10908 (NHA); Durham, Hodgdon 5159 (NHA); BELKNAP CO., Alton, iIHodgdon & Steele 12257 (NHA); MERRIMACK CO., Hooksett, Hodgdon 10912 (NHA); SULLIVAN CO., Charlestown, Hodgdon & Steele 11365 (NHA); ROCKINGHAM CO., Seabrook, Hodgdon & Steele 10911 (NHA). VER- MONT: WINDSOR CO. Hartland, Wheeler 26770 (NEBC); RUTLAND CO., north of Rutland, July 1, 1914, Brainerd 11 (NEBC) ; WINDSOR CO., Newfane, Wheeler 27000 (NEBC). MASSACHUSETTS: ESSEX CO., Salis- bury, Steele & Hodgdon 3501 (STEELE); PLYMOUTH CO., Plymouth, Fernald & Hunnewell 15232 (NEBC) ; BRISTOL CO., Dartmouth, August 12, 1910, Hervey (NEBC); BARNSTABLE CO. Barnstable, Fernald. 18589 (NEBC); NANTUCKET CO. Nantucket Island, Bicknell 11907 (NEBC); WORCESTER CO., June 26, 1921, Weatherby (NEBC); BERK- SHIRE CO., Sheffield, May 30, 1919, Bean & Fernald (NEBC); CON- NECTICUT: TOLLAND CO., Stafford Springs, Steele & Hodgdom 3893 (STEELE); HARTFORD CO., Southington, Andrews 5051 (NEBC); NEW LONDON CO., Stonington, Hodgdon & Steele 14828 (NHA); NEW HAVEN co., Oxford, Harger 729 (NEBC). 2. Rubus Jaysmithii Bailey. Differs from R. flagellaris in having leaflets soft-pubescent or velvety beneath to the touch; in similar 490 Rhodora [Vol. 68 habitats and probably of about the same range as R. flagellaris, but less common. This species is closely related to R. flagellaris. REPRESENTATIVE SPECIMENS: NEW HAMPSHIRE: CARROLL co., Sandwich, Hodgdon & Steele 14829 (NHA); GRAFTON CO., Ply- mouth, Fernald, 11174 (NEBC); Rummey, Steele 2869 (STEELE) ; STRAFFORD CO., Rochester, Hodgdon 10915 (NHA); MERRIMACK CO., Franklin, ‘Hodgdon 7270 (NHA); ROCKINGHAM co., Nottingham, Hodgdon 14830 (NHA); Salem, Steele 2281 (STEELE). VERMONT: RUTLAND CO., Rutland, September 15, 1912, Kirk (NEBC) ; WINDHAM co., West Wardsboro, July 25, 1913, Brainerd 10 (NEBC) ; Westmin- ister, Steele 3498 (STEELE). MASSACHUSETTS: ESSEX co., Danvers, Steele 768 (STEELE); NORFOLK CO., Walpole, Hodgdon & Steele 18431 (NHA); BARNSTABLE CO., Harwich, Fernald 16930 (NEBC) ; Chatham, Fernald 16932 (NEBC); DUKES CO., Marthas Vineyard, July 1, 1913, Bicknell (NEBC) ; NANTUCKET CO., Nantucket Island, Bicknell 11894 (NEBC); FRANKLIN CO., East Charlmont, July 4, 1921, Churchill (NEBC). RHODE ISLAND: NEWPORT CO. Block Island, Jansson 19 (NEBC); PROVIDENCE CO. East Providence, Wiegand 1009 (NEBC). 3. Rubus Enslenti Tratt. PRIMOCANE mostly trailing, sometimes low-arching (0.3 mm); FLORICANE of similar habit; ARMATURE of thin sharp and hooked prickles, 1-2 mm long, 2-60 per dm; DIAMETER of cane 0.5-2 (2.5) mm; PRIMOCANE LEAFLETS mostly 3, sometimes 5 in number, membranaceous to subchartaceous, ovate, the ratio of width to length 0.6-0.7, subacute to acuminate at apex, rounded to subeuneate at base, glabrous above, essentially glabrous beneath (sometimes finely pubescent on veins); DIMENSIONS OF COMPOUND LEAF 77-1212 em, AVERAGING SMALLER than R. flagellaris; IN- FLORESCENCE a narrow raceme 2-6 (8) cm long, with 1-2 (3) flowers on nearly erect pedicels, solitary or longest pedicels 2-6 (8) em long; axis of inflorescence and pedicels mostly pubescent and with or without prickles, lacking glands; early flowering; FRUIT sparse, often of good quality; HABITAT, dryish woods mostly of oak-hickory and on ledges; RANGE, southern Maine mostly near coast, central and southern New Hampshire southward and westward, less common to the north. Although this blackberry is at times close to reduced forms of R. flagellaris it is the most distinctive of the Flagel- lares and clearly a good species. In addition to morphologi- cal differences it has a distinct range and characteristic environment. Its range in New England is somewhat re- stricted. It reaches its northern limits in central N. H. and southern Maine. The most characteristic habitat of Rubus Enslenii is open oak-hickory woods, but it is also found in 1966] Rubus in New England — Hodgdon & Steele — 491 mixed woods and small rocky openings. A small trailing prickly blackberry in hickory-woods is certainly this species. It is one- or two-flowered with thin stems and small dull glabrous leaves, usually three in number. In shady sites the floricanes may not produce any flowers at all. Rubus flagel- laris is often one- or two-flowered and may have slender canes, but usually inflorescences can be found with more flowers while the leaves average larger and are often 5 fo- liate. REPRESENTATIVE SPECIMENS: MAINE: CUMBERLAND CO., Brunswick, June 28, 1913, Furbish (NEBC) ; YORK COUNTY, Biddeford, Hodgdon & Steele 14832 (NHA); York, Hodgdon & Steele 14833 (NHA). NEW HAMPSHIRE: CARROLL CO., Tuftonboro, Hodgdon & Steele 12243 (NHA); STRAFFORD CO., Durham, Hodgdon 3032 (NEBC) ; Durham, Hodgdon & Steele 14834 (NHA) ; MERRIMACK co., Boscawen, Pease 19150 (NEBC); ROCKINGHAM CO., Nottingham, Hodgdon & Steele 10916 (NHA); Seabrook, Hodgdon & Steele 10917 (NHA). MAS- SACHUSETTS: MIDDLESEX CO., Pepperel, Steele 2777 (STEELE); Con- cord, June 1, 1960, Eaton (NEBC); Lincoln, June 30, 1956, Eaton (NEBC); SUFFOLK CO. Arnold Arboretum, E. J. Palmer 44596 (NEBC); BARNSTABLE CO., Harwich, Fernald & Long 16934 (NEBC); NANTUCKET CO., Nantucket Island. Bicknell 12044 (NEBC) ; WORCES- TER CO., Leominster, Fernald & Bean 14058 (NEBC). RHODE ISLAND: WASHINGTON CO., Charlestown, Fernald & Phelps 16936 (NEBC). CON- NECTICUT: WINDHAM CO., Putnam, June 11, 1922, Bill & Grigg (NEBC); NEW LONDON CO., Franklin, May 25, 1906, Woodward (NEBC); Stonington, Steele & Hodgdon 3890 (STEELE). 4. Rubus recurvicaulis Blanch. PRIMOCANE trailing to low doming or low arching to 0.3-0.6 m high, in dense vegetation often appear- ing suberect early in growth; FLORICANE trailing or reclining to low arching; ARMATURE of stiff hooked or straight retrorse prickles 2-4 mm long, (2-) 10-50 (-100) in number per dm; DIAMETER OF PRIMO- CANE 2-4 mm: PRIMOCANE LEAFLETS 3-5 in number, chartaceous, ovate (rhombic), ratio of width to length 0.6-0.8, acuminate at apex, rounded to subcordate at base, glabrous above and glabrous beneath or with slight pubescence on veins beneath; DIMENSIONS OF COM- POUND LEAF 88-1414 cm; INFLORESCENCE à compact to diffuse raceme (corymb of many authors) with 2-8 (1-12) flowers, 3-9 (2- 10) em long, 2-5 (1-6.5) cm wide, pedicels spreading to ascending, the longest 1.5-4, the upper 0.5-1 em long; axis of inflorescence and pedicels mostly with slight to medium pubescence, prickles often present, glands absent; early flowering; FRUIT, poor quality to good; HABITAT, pastures, fields, railroad banks and roadsides; range throughout New England, at low elevations. 492 Rhodora [Vol. 68 This species occupies the same habitats as Rubus flagel- laris and often resembles it. When properly developed, the inflorescences are distinctive with some flowers on short pedicels, some of which are ascending or diverging as op- posed to the erect pedicels of R. flagellaris. At times, espe- cially in poor soils, the primocanes do not assume the trailing habit but remain low-arching, not close to the ground. At other times, when there is little competition, the primocanes may form large tangled doming and trailing colonies with few well-developed inflorescences. The colonies may be difficult to classify, but growth-habit alone will sug- gest an affinity to R. recurvicaulis rather than R. flagellaris. The fresh leaves of this species vary from dull to lustrous and the margins may be plicate when the leaves are young. This tendency has led to the “Manual” species R. plicati- folius not here recognized. Most blackberries grow in disturbed locations and thus the stations tend to be temporary. An apparently natural location for the flagellares-group is pockets of soil on ledges and cliffs. Most New England cliffs will contain one of these species presumably growing in natural and virgin sites, as such habitats have been little disturbed by agriculture and lumbering. REPRESENTATIVE SPECIMENS: MaINE: AROOSTOOK CO., Fort Kent, July 6, 1906, Fernald (GH); Ashland, August 8, 1901, Fernald (NEBC); PENOBSCOT CO., Oldtown, Fernald & Long 13883 (NEBC); PISCATAQUIS CO., Abbot, Fernald & Long 13875 (NEBC); OXFORD CO., Rumford, Fernald & Pease 2516 (NEBC); WASHINGTUw co., Roque Bluff, July 25, 1913, Knowlton, (NEBC) ; KNOX CO., Matini- cus, July 19, 1920, C. A. E. Long (NEBC); LINCOLN CO., Monhegan Island, Hodgdon & Hodgdon 12941 (NHA); Southport Island, Hodg- don 7228 (NHA); KENNEBEC CO., Vassalboro, Fernald 13904 ( NEBC) ; CUMBERLAND CO. Harpswell, Hodgdon & Steele 12169 (NHA); YORK CO., Alfred, Fernald & Long 13898 (NEBC); Kittery, Hodgdon & Steele 12171 (NHA). NEW HAMPSHIRE: COÓS co., Shelburne, Pease 11162 (NEBC) ; CARROLL CO., North Conway, Hodgdon & Steele 12189 (NHA); Sandwich, Hodgdon & Steele 14835 (NHA); GRAFTON CO., Lineoln, Fernald 15796 (NEBC) ; Campton, Hodgdon & Steele 11354 (NHA); STRAFFORD CO. Durham, Hodgdon 12261 (NHA); Milton, Hodgdon & Steele 10920 (NHA); ROCKINGHAM CO., Northwood, Hodg- don & Steele 10930 (NHA): Seabrook, Hodgdon & Steele 10928 (NHA); HILLSBORO CO., Litchfield, Steele & Hodgdon. 3842 (STEELE). 1966] Rubus in New England — Hodgdon & Steele 493 VERMONT: WINDHAM CO., Westminster, Steele 3479 (STEELE); BEN- NINGTON CO., Searsburg, June 20, 1925, Carpenter, Churchill & Knowlton (NEBC). MASSACHUSETTS: MIDDLESEX CO., Concord, June 19, 1958, Eaton (NEBC); BARNSTABLE CO., Barnstable, Woodward & Fernald 1523 (NEBC); WORCESTER CO., Royalston, Steele & Hodgdon 3819 (STEELE); HAMPSHIRE CO., Middlefield, Fernald & Long 9678 (NEBC); HAMPTON CO., Springfield, June 17, 1914, Andrews (NEBC). Specimens from Rhode Island and Connecticut in the herbarium of the New England Botanical Club are hardly adequate for purposes of citation. 5. Rubus arenicola Blanch. Differs from R. recurvicaulis chiefly in having leaves abundantly pilose to soft-velvety beneath and axis of inflorescence and pedicels with medium to strong pubescence; HABITAT, dry fields, roadsides, alluvial plains, and ledges: RANGE, southern and western Maine, northern New Hampshire and south- ward at low elevations. This blackberry is closely related to R. recurvicaulis, but at times seems markedly distinct. In its best marked states it occurs in large doming and trailing colonies on sandy soil, with fruit of superior quality. REPRESENTATIVE SPECIMENS: MAINE: YORK CO. Kenne- bunkport, July 6, 1905, Blanchard (NEBC). NEW HAMPSHIRE: COÓS co., Shelburne, August 9, 1921, Deane (NEBC); CARROLL CO., Tam- worth, Hodgdon & Steele 7782 (NHA); Ossipee, July 14, 1951, Steele (NHA); Sandwich, Steele & Hodgdon 764 (STEELE); GRAFTON CO., Holderness, Hodgdon, Steele & Lincoln 11317 (NHA); Plymouth, Hodgdon & Steele 10918 (NHA); STRAFFORD CO., Milton, Steele & Hodgdon 2932 (STEELE); Durham, Hodgdon, Pike & Schreiber 7260 (NHA); ROCKINGHAM CO., Deerfield, Steele & Hodgdon 3769 (STEELE); Seabrook, Hodgdon & Steele 10919 (NHA); HILLSBORO CO., Pelham, Hodgdon & Eldredge 7256 (NHA). MASSACHUSETTS: ESSEX co., Newburyport, Steele 3834 (STEELE); PLYMOUTH CO., Marshfield, July 4, 1914, Knowlton (NEBC); BARNSTABLE CO., Chatham, Fernald & Long 18593 (NEBC); Harwich, Steele & Hodgdon 3924 (STEELE) ; Sandwich, Steele & Hodgdon 3895 (STEELE); DUKES CO., West Tis- bury, Fernald & Fogg 920 (NEBC); NANTUCKET CO., Nantucket Island, Bicknell 11952 (NEBC); WORCESTER CO., Sterling, Steele & Hodgdon 3897 (STEELE). RHODE ISLAND: NEWPORT CO., Portsmouth, July 28, 1924, Sanford (NEBC, Cotype of R. rhodinsulans Bailey). 6. Rubus hispidus L. PRIMOCANE prostrate and often tip-rooting in late summer or sometimes trailing over vegetation; FLORICANE prostrate or trailing; ARMATURE of thin hooked prickles resistant to touch or of soft green to purple bristles 2-3 mm long, 0-2000 per dm: DIAMETER OF CANES 1-2(3) mm; PRIMOCANE LEAFLETS 8 in num- ber, occasionally 5, lustrous, coriaceous (sometimes dull), tending 494 Rhodora [Vol. 68 to be evergreen, ratio of width to length 0.65 or more, obovate (broad- est above the middle), apex obtuse, abruptly pointed or acute (never acuminate), the base rounded or subeuneate, 3.5-5(7) em long, glab- rous above and beneath, teeth somewhat rounded or abruptly pointed; DIMENSIONS OF COMPOUND LEAF 66-99 cm; INFLORES- CENCE a compact raceme 2-4.5 em long with 4-10 flowers on ascend- ing to divergent pedicels, the axis pilose, setae often present, glands lacking or few and of uniform length; late flowering, FRUIT usually not developing or, if maturing, small and Sour; HABITAT, fields, open woods especially pine, gravel pits, peaty soil and peat bogs; RANGE, common throughout New England at low elevations. This is one of the oldest and best-marked species. Except when reclining on vegetation, it is low-trailing or prostrate and tip-rooting, the lowest of the blackberries. The arma- ture is of bristles or weak prickles not painful to touch. The leaves are small, obovate, coriaceous and usually shiny. These features serve to easily identify this blackberry in most cases. Like some of the others, it grows in a variety of situations, but is the only one to invade acid peat bogs. It flowers later than most species and the fruit, if it de- velops at all, is sour and of inferior quality. The authors have investigated in detail its hybridization with R. setosus (Rhodora 65: 262-270). These hybrids are frequent and several have been designated as species. The hybrids frequently backcross with one of the parents, and the introgression of R. setosus into the plant gives rise to forms of R. hispidus which are not typical and may suggest new species. There is fairly good evidence of hybridization with R. allegheniensis and vermontanus but the details need to be worked out. A large-leafed form of Rubus hispidus occurs with leaves twice as large as the typical but similar in other respects. Although at times this may appear to be distinctive, there is insufficient evidence to consider it a separate species. REPRESENTATIVE SPECIMENS: Maine: PENOBSCOT CO., Oronc, July 9, 1888, Fernald (NEBC) ; SOMERSET CO., Lexington Plan- tation, Pease 34502 (NEBC); WASHINGTON CO., Machias, Aucust 13, 1942, (NEBC); Hancock Co., Mt. Desert, July 14, 1897, Williams (NEBC); WALDO CO. Burnham, July 24, 1940, Bean (NEBC); KNOX co., Friendship, Pease 747 (NEBC); LINCOLN CO., Southport, August 1, 1894, Fernald (NBEC); Boothbay Harbor, Hodgdon 12254 (NHA); 1966] Rubus in New England — Hodgdon & Steele 495 SAGADAHOC CO., Georgetown (Five Islands), Rehder 1045 (GH); ANDROSCOGGIN CO., East Livermore, Furbish (NEBC); YORK CO., Al- fred, Fernald & Long 13893 (NEBC). NEW HAMPSHIRE: coos CO., Colebrook, Pease 29760 (NEBC); Jefferson, Pease 33126 (NEBC) ; CARROLL CO., Madison, June 28, 1951, Steele (NEBC); Tamworth, Steele 2859 (STEELE); GRAFTON CO., Waterville, Steele 2921 (STEELE) ; STRAFFORD CO., Milton, Hodgdon & Steele 11202 (NHA); Durham, Hodgdon 12168 (NHA); BELKNAP CO., Alton, Hodgdon & Steele 12191 (NHA); MERRIMACK CO., Bradford, Hodgdon, Steele, Lincoln & Har- rington 7443 (NHA); SULLIVAN CO., Charlestown, Hodgdon & Steele 11363 (NHA); ROCKINGHAM CO., Northwood, Hodgdon & Steele 11195 (NHA); HILLSBORO CO., Peterboro, October 3, 1907, Batchelder (NEBC); CHESHIRE CO., Rindge, August 19, 1916, BATCHELDER (NEBC). VERMONT: ORLEANS CO., East Charleston, Pease 34755 (NEBC) ; CHIT- TENDEN CO., June 27, 1931, Kittredge (NEBC); WINDSOR CO., North Woodstock, July 15, 1914, Forbes (NEBC); RUTLAND CO., Castleton, August 20, 1940, Knowlton (NEBC); WINDSOR CO., Stratton, July 15, 1912, Wheeler (NEBC). MASSACHUSETTS: ESSEX CO., Rockport, Forbes 1902 (NEBC); MIDDLESEX CO., Billerica, August 20, Manning (NEBC) ; BRISTOL CO., Westport, September 10, 1916, Sanford (NEBC); NAN- TUCKET CO., Nantucket, June 24, 1910, Bicknell (NEBC); WORCESTER co. Holden, July 7, 1946, Gates & Gates (NEBC); FRANKLIN CO. Rowe, May 15, 1915, Harger & Schweinfurth (NEBC); BERKSHIRE co., Sandisfield, July 19, 1912, Hoffman (NEBC). RHODE ISLAND: PROVIDENCE C0., Providence, June 1878, W. W. Bailey (NEBC) ; WASH- INGTON CO., South Kingston, June 10, 1927, Bill, Eaton & Gilbert (NEBC) ; NEWPORT CO., Block Island, August 13, 1919, Graves (NEBC). CONNECTICUT: WINDHAM CO. Thompson, July 6, 1921, Weatherby (NEBC); HARTFORD CO. East Hartford, Clark & Weatherby 120 (NEBC); NEW LONDON CO., Stonington, Hodgdon & Steele 14837 (NHA); NEW HAVEN CO, Oxford, August 4, 1901, Harger (NEBC). 7. Rubus setosus Bigel. PRIMOCANE arching to erect (sometimes doming) of medium height; FLORICANE reclining or trailing on ground; ARMATURE of abundant soft bristles and glands (600) 2000- 5000 per dm; CANE DIAMETER 2-5 mm; PRIMOCANE LEAFLETS 5, chartaceous with impressed veins, elliptic to obovate, usually broadest at middle, ratio of width to length 0.5 —0.7, acute or acuminate, 6-11 cm. long, glabrous above and beneath, teeth acute, leaf-base rounded to cuneate; dimensions OF COMPOUND LEAF 66-1414; INFLORESCENCE a compact to diffuse raceme 7-11 cm long; late- flowering, number of flowers 5-11; floral axis pubescent, bristles often present, glands frequent to abundant on stalks of uneven lengths; FRUIT failing to develop or of poor quality, small; HABITAT, alluvial plains, poorly drained soil, grassy swamps, open land; RANGE, common in northern New England, becoming less frequent southward to central Connecticut, at low elevations. 496 Rhodora [Vol. 68 This is a long-recognized and distinctive species best de- veloped in poorly drained soil and alluvial meadows. It oftens grows in swales or bushy swamps but hardly ever in peat bogs. It is essentially an erect plant of medium height with rather slender canes. The abundant soft bristles are characteristic. These are not unpleasant to touch as compared with the sharp thin prickles of R. vermontanus. In deep vegetation the plant grows tall, the bristles may be much less numerous, and the floricanes poorly developed and diffieult to locate. The fruit usually does not develop and when it does it is small and of inferior quality. REPRESENTATIVE SPECIMENS: MAINE: PENOBSCOT CO., Winn, Fernald & Long 13889 (NEBC) ; PISCATAQUIS CO., Brownville, Parlin 1886 (GH); FRANKLIN co., Farmington, July 24, 1915, Knowl- ton (NEBC); OXFORD CO., Bethel, August 7, 1929, Wheeler (NEBC) ; WASHINGTON CO., Princeton, July 22, 1909, Fernald (GH); HANCOCK co., Mt. Desert, July 2, 1896, Rand (NEBC) ; ANDROSCOGGIN CO., South Poland, 1895, Furbish (NEBC); CUMBERLAND CO., Baldwin, Fernald & Long 18885 (NEBC) ; YORK CO., South Berwick, September 26, 1897, Parlin (NEBC). NEW HAMPSHIRE: COÓS co., Berlin, Pease 17060 (NEBC) ; CARROLL CO., Ossipee, August 4, 1948, Knowlton (NEBC) ; North Conway, Hodgdon & Steele 11204 (NHA); GRAFTON CO., Easton, Fernald 15826 (NEBC); Plymouth, Hodgdon & Steele 11173 (NHA); STRAFFORD CO., Rochester, Hodgden 3461 (NEBC); BELKNAP CO., La- conia, Fernald 15776 (NEBC); SULLIVAN CO., Langdon, August 23, 1963, Richardson & Seymour (NEBC) ; ROCKINGHAM CO., Nottingham, Hodgdon 10199 (NEBC) ; HILLSBORO CO., Peterboro, July 10, 1888, Deane (NEBC). VERMONT: ESSEX co., Brighton, Pease 33683 (NEBC); FRANKLIN CO., Berkshire, July 1916, Knowlton (NEBC); ADDISON CO., Ripton, July 19, 1900, Brainerd & Eggleston (NEBC); RUTLAND CO., Wallingford, Brainerd 1225 (NEBC) ; WINDHAM CO., Stratton, Under- wood 3038 (NEBC). MASSACHUSETTS: MIDDLESEX CO., Lexington, August 19, 1908, Kennedy (NEBC) ; WORCESTER CO., Ashland, Gates 31415 (NEBC); Sterling, Steele & Hodgdon 3947 (NHA, STEELE). CONNECTICUT: WINDHAM CO., Hampton, Hunnewell 12620 (NEBC) ; NEW LONDON CO. Franklin, June 26, 1905, Graves (GH); TOLLAND co., Bolton, Weatherby 3166 (NEBC); Strafford, August 29, 1903, Graves (GH): HARTFORD CO., Southington, Andrews 5014 (NEBC) ; LITCHFIELD CO., Colebrook, Weatherby 2668 (NEBC): FAIRFIELD CO, Botsford, June 27, 1897, Eames (NEBC). 8. Rubus vermontanus Blanch. PRIMOCANE erect or arching, of medium height (0.3-0.9), less often doming or high trailing; FLORI- CANE arching or reclining; ARMATURE usually of acicular straightish prickles 2-4 mm long, stiff and unpleasant to touch, less often hooked 1966] Rubus in New England — Hodgdon & Steele 497 50-500 (10-250) per dm; DIAMETER OF CANE 2-4 mm; PRIMOCANE LEAFLETS 5, chartaceous or subchartaceous, veins impressed, rhombic, elliptic or obovate, broadest at middle or above middle, ratio of width to length 0.5-0.7, apex acute or abruptly acuminate, 6-12 cm long, glabrous on upper and lower surfaces usually with slight pubescence on main veins evident under magnification, teeth acute, base rounded to cuneate, central petiolules 1.4-2.0 (0.8-0.3) cm. long, longest lateral petiolules 0-1 (1.7) em long; DIMENSIONS OF COM- POUND LEAF 9X9-1818 cm; INFLORESCENCE similar to R. setosus but thin prickles usually present as well as glands; early flowering; FRUIT small, often underdeveloped, of poor to fair quality; HABITAT, lumbered areas, sterile fields, thickets and bases of cliffs; RANGE, northern New England common, becoming less abundant southward to Connecticut, medium and low elevations. This is a distinctive species in northern New England, common in open lumbered areas and run out fields. It most closely resembles R. elegantulus and at times is difficult to distinguish from that species, perhaps because of hybridiza- tion, The following characters serve to separate it; height averaging low, 1-3’ (R. elegantulus, 2-4’) ; length of petio- lules, central usually less than 2 cm, (R. elegantulus more than 1 cm); floral axis frequently with glands, (R. elegan- tulus never with glands). In growth-habit and appearance this plant resembles R. setosus, but the armature is distinc- tively different. Although the leaves apparently are glab- rous, some hairs can usually be seen under magnification on the main veins beneath. However these two species frequent- ly hybridize forming various intermediate states, many of which have been designated as R. Groutianus. Rubus ver- montanus also occurs in doming, excessively prickly forms which have been designated as R. severus. This possibly represents introgression with some of the flagellares. Rubus vermontanus appears to hybridize with several other spe- cies producing puzzling mongrel populations. In southern New England it tends to be replaced by R. semisetosus and to hybridize with it. At high elevations above 300 feet in boggy places a black- berry occurs which we have assigned to R. elegantulus but it has some of the characteristics of R. vermontanus. It does not seem to be a hybrid of them, however, as neither of the putative parents is present. 498 Rhodora [Vol. 68 REPRESENTATIVE SPECIMENS: MAINE: AROOSTOOK CO., Fort Fairfield, August 10, 1909, Fernald & ‘Wiegand (GH); PENOBSCOT co., Stacyville, July 3, 1900, Churchill (NEBC); PISCATAQUIS CO., Dover, September 2, 1896, Fernald (NEBC); FRANKLIN CO., Chain of Ponds, Pease & Bean 33605 (NEBC) ; OXFORD CO., Canton, Parlin 2002 (NEBC); Mason, Hodgdon & Steele 11313 (NHA); WASHINGTON CO., Cutler, July 30, 1902, Furbish (NEBC); Pembroke, August 18, 1909, Fernald & Wiegand (GH); HANCOCK CO., Mt. Desert, July 1, 1896, Hand (NEBC); KNOX co., Thomaston, Bissell, Fernald & Chamberlain 7697 (NEBC); LINCOLN CO., Boothbay (Ocean Point), Hodgdon 12269 (NHA); YORK CO., Kennebunk, July 3, 1905, Blanchard (NEBC) ; Bidde- ford, Hodgdon & Steele 12255 (NHA). NEW HAMPSHIRE: COÓS CO., Shelburne, Fernald & Pease 15746 (NEBC) ; CARROLL CO., North Con- way, Hodgdon & Steele 11208 (NHA); Conway, Pease 28776 (NEBC) ; GRAFTON CO., Woodstock, Fernald 11743 (NEBC); Plymouth, Hodgdoia & Steele 11211 (NHA); STRAFFORD CO., New Durham, Steele & Hodg- don 3703 (STEELE) ; Rollinsford, Hodgdon 11367 (NHA); BELKNAP CO., Laconia, Fernald 15777 (NEBC, as var. viridiflorus); Alton, Hodgdon & Steele 12195 (NHA); SULLIVAN CO., Grantham, Hodgdon, Steele, Page & Stearns 7781 (NHA); ROCKINGHAM CO., Seabrook, Hodgdon & Steele 1180 (NHA); HILLSBORO CO., Nashua, Hodgdon & Steele 11299 (NHA). VERMONT: ESSEX co., Ferdinand, Pease 25694 (NEBC) ; ORLEANS CO. Westmore, Floyd 2264 (NEBC); ADDISON CO., Ripton, July 27, 1914, Brainerd (NEBC); RUTLAND CO. East Wallingford, June 27, 1909, Kent (NEBC) ; WINDHAM CO., West Dover, August 13, 1928, Eaton (NEBC) ; Windham, June 20, 1904, Blanchard (Gu type; BENNINGTON C0., Woodford, June 29, 1939, Knowlton (NEBC). MAssa- CHUSETTS: ESSEX CO. Plum Island, Steele & Hodgdon 3833 ( STEELE) ; BERKSHIRE CO., Becket, July 9, 1920, Hoffmann (NEBC). RHODE ISLAND: PROVIDENCE co., Diamond Hill, E. J. Palmer 45604 (GH as R. significans Bailey). CONNECTICUT: WINDHAM CO. Thompson, Weatherby 5051 (NEBC); HARTFORD CO., Berlin, Hodgdon & Steele 14838 (NHA); East Hartford, Ashe & Weatherby 5356 (NEBC). 9. Rubus semisetosus Blanch. PRIMOCANE erect or arching, of medium height (0.3-0.9 m); FLORICANE erect or reclining; ARMATURE of acicular to thin prickles 2-4 mm long, 25-100 per dm; DIAMETER OF CANE 25 mm; PRIMOCANE LEAFLETS 5 (occasionally 3), membrana- ceous or chartaceous, ovate or rhombic, 6-9 em long, ratio of width to length averages 0.6-0.7, glabrous or occasionally minutely pilose above, strongly pilose to velvety beneath, base of leaflet round to cuneate rarely cordate, central petiolule 1-1.8 cm long, longest lateral petiolules 0.2-0.7 mm long; DIMENSIONS OF COMPOUND LEAF 8X8- 14X14 (17X17) cm; INFLORESCENCE similar to R. vermontanus, floral axis always pubescent, thin prickles often present, glands rarely present; FRUIT undeveloped or poor; HABITAT, swales, thickets 1966] Rubus in New England — Hodgdon & Steele 499 and open fields; RANGE, coast of Maine, southern New Hampshire occasional, moderately common from Massachusetts southward. This blackberry is most closely related to Rubus ver- montanus and replaces it in southern New England. The two differ in pubescence of leaves; Ff. vermontanus has leaves glabrous on the underneath side with a minute pubes- cence on the veins whereas R. semisetosus has leaves soft- velvety to the touch. In southern New Hampshire some plants with only a slight pubescence are found along with good R. semisetosus. Farther south R. vermontanus is rare with R. semisetosus replacing it. Thus there appear to be two distinct species that have a transition zone of inter- gradation. Rubus semisetosus is also close to the Arguti, especially to R. argutus. It is a smaller plant, however, and the leaf- shape is usually distinctive. Rubus semisetosus typically has rather broad obovate or rhombic leaves, whereas R. argutus has narrow lanceolate leaves. It definitely hybrid- izes with R. setosus and R. vermontanus and probably with other species. REPRESENTATIVE SPECIMENS: MAINE: HANCOCK CO., Mt. Desert Island, July 8, 1894, Rand (NEBC) ; Mt. Desert Island, Septem- ber 7, 1896, Rand (NEBC). NEW HAMPSHIRE: ROCKINGHAM CO., Not- tingham, Hodgdon 14839 (NHA); HILLSBORO CO., Merrimack. Steele & Hodgdon 3830 (STEELE). MASSACHUSETTS: MIDDLESEX CO., Pep- perel, Hodgdon & Steele 14880 (NHA); Concord, June 16, 1962, Eaton 5205 (NEBC); Concord, July 16, 1961, Eaton (NHA); NOR- FOLK CO., Millis, Hodgdon & Steele 14841 (NHA); Walpole, Hodg- don & Steele 14842 (NHA); Medway, Hodgdon & Steele 14843 (NHA); PLYMOUTH CO., Carver, Fernald, Hunnewell & Long 9667 (NEBC) ; BARNSTABLE CO., Brewster, Steele & Hodgdon 3878 (STEELE) ; WOR- CESTER CO., Sterling, Hodgdon & Steele 14844 (NHA); HAMPDEN CO., Springfield, Weatherby 5918 (NEBC). RHODE ISLAND: WASHINGTON co., Kingston, July 30, 1906, Blanchard (GH type); Kingston, Steele & Hodgdon 3877 (STEELE); Kenyon, Fernald & Weatherby 843 (GH, NHA). CONNECTICUT: HARTFORD co., East Hartford, Weatherby 4905 (NEBC) ; New Canaan, Steele 3948 (STEELE); NEW HAVEN CO., North Branford, August 17, 1940, Harger (NEBC); FAIRFIELD CO., Green- wich, June 20, 1913, Bissell & Weatherby (NEBC). 10. Rubus cune*folius Pursh PRIMOCANES stiff-erect to arching or occasionally trailing, of medium height (0.3-0.9 m); FLORICANES sim- ilar, cften branched; ARMATURE of numerous (20-50 per dm) thin 500 Rhodora [Vol. 68 to stout stiff straight or hooked prickles; PRIMOCANES usually PUBES- CENT, DIAMETER OF CANE 2-4 mm; PRIMOCANE LEAFLETS 3 or 5, char- taceous or subcoriaceous, glabrous above, closely white-or gray-tomen- tose beneath, obovate, base cuneate, broadest above the middle, ratio of width to length averaging 0.6-0.7, apex rounded or abruptly pointed, length 2-5 cm; DIMENSIONS OF COMPOUND LEAF average 6X6- 8X8 cm, INFLORESCENCE prickly-pubescent, of few-flowered small racemes or cymes; early flowering; FRUIT dry and seedy or sweet and of good quality; HABITAT, sandy or rocky and open dry lands; RANGE, southern Connecticut, where uncommon and isolated in Hills- boro, N. H. This is a very distinctive southern species of the Coastal Plain and Lower Piedmont which enters New England only in southern Connecticut except for a highly disjunct station in Hillsboro County, New Hampshire. REPRESENTATIVE SPECIMENS: New HAMPSHIRE: HILLS- BORO CO., Hillsboro, October 22, 1966, William Cottrell (NEBC, NHA). CONNECTICUT: HARTFORD co., Southington, June 13, 1900, Bissell (NEBC); MIDDLESEX CO., Portland, June 24, 1897, Starmer (NEBC) ; NEW HAVEN CO. North Branford, June 18, 1927, Harger (NEBC); LITCHFIELD CO., Woodbury, August 12, 1926, Harger (NEBC); FAIR- FIELD CO., Bridgeport, July 4, 1890, Johnson (NEBC). ll. Rubus canadensis L. PRIMOCANE and FLORICANE 1-2 (2.5) m high, erect to somewhat arching; armature of acicular to weak prickles or none, 0.5-3 mm long, 0-7 (9) per dm; DIAMETER OF PRIMO- CANE 2-7 mm; PRIMOCANE LEAFLETS 5, membranaceous, ovate, ratio of width to length 0.5-0.7, acuminate or occasionally abruptly pointed, 7-14 em long, glabrous on both surfaces, cordate to rounded at base, central petiolules 1.5-5 em long, longest lateral petiolules 0.5-2 cm; dimensions of compound leaf 1010-24 24 cm; INFLORESCENCE vary- ing from long racemes (11 em long by 5 em) to short racemes (7 em long by 6 em) to extremes of small diffuse racemes with thin pedi- cels, the flowers 6-14 in number; axis of inflorescence and pedicels pubescent to subglabrous, lacking glands and prickles; flowering in midseason; FRUIT sour, often poorly developed, occasionally of fair quality; HABITAT, moist thickets, openings in northern hardwood forest, bases of cliffs, occasionally ascending mountains to 1100 m; RANGE, northern New England south to mountainous areas of Mas- sachusetts and Connecticut, common in north, less common southward. This species, along with R. elegantulus, and R. vermont- anus appear to be the three northernmost blackberries, all occurring in Newfoundland. In New England R. canaden- sis is characteristic of mountainous areas and ascends higher than any other blackberry. It is often found in open 1966] Rubus in New England — Hodgdon & Steele 501 hardwoods or near ledges at 2000 feet and has been collected on Mt. Washington at 3600 feet which is the highest re- corded altitude for any blackberry in New England. It occurs in a variety of lowland habitats but its most char- acteristic environment seems to be open hardwoods and moist thickets. It is usually a distinctive well-marked plant, a high black- berry with glabrous leaves and a stem nearly devoid of prickles, the few present being small and weak. Close ex- amination of the leaves under magnification indicates that they are either completely glabrous or have a few hairs on the main nerves. Any pubescence that could be detected without a lens would indicate there was introgression with some other species, presumably R. pensilvanicus. REPRESENTATIVE SPECIMENS: MAINE: AROOSTOOK CO., Van Buren, July 17, 1914, Knowlton (NEBC); PENOBSCOT CO., Bangor, June 27, 1909, Blanchard (GH); PISCATAQUIS CO., Abbot, Fernald & Long 13879 (NEBC); FRANKLIN CO., Industry July 9, 1896, Fernald (NEBC) ; OXFORD CO., Greenwood, June 13, 1931, Bill, Eaton, Fernald, Griscom & Hunnewell (NEBC); WASHINGTON CO., Pembroke, July 7, 1909, Fernald & Wiegand (GH); HANCOCK CO., Dedham, Fernald & Long 13877 (NEBC); WALDO CO., Islesboro, Woodward, Bissell & Fer- nald 9672 (NEBC); LINCOLN CO., Monhegan Island, Hodgdon & Hodg- don 12808 (NHA); ANDROSCOGGIN CO., South Poland, 1893, Furbish (NEBC); YORK CO. Kittery, Fernald & Long 13880 (NEBC). NEW HAMPSHIRE: COÓSs co., Dartmouth College Grant, Pease & Harris, 33915 (NEBC); Dummer, Hodgdon & Steele 11269 (NHA, STEELE) ; CARROLL CO., Sandwich, Steele 3547 (STEELE); GRAFTON Co., Thornton, Hodgdon & Steele 12373 (NHA); Franconia, August 7, 1919, Deane (NEBC); STRAFFORD CO., Barrington, Hodgdon & Steele 7279 (NHA); BELKNAP CO., Gilford, Fernald 15654 (NEBC); HILLSBORO CO., Peter- boro, Ware 4127 (NEBC); Hollis, Steele & Hodgdon 4008 (STEELE) ; CHESHIRE CO., Gilsum, Fernald 321 (NEBC) ; Alstead, June 14, 1903 — August 16, 1903, Blanchard (GH all marked type). VERMONT: ORLEANS co., Lowell, August 9, 1915, Winslow (NEBC); ADDISON CO., Ripton, July 5, 1914, Brainerd (NEBC); WINDSOR CO., North Woodstock, July 15, 1914, Forbes (NEBC); RUTLAND CO. Pittsfield, July 27, 1933, Knowlton (NEBC); WINDHAM CO., Marlboro, June 28, 1939, Knowlton (NEBC). MASSACHUSETTS: WORCESTER CO., Princeton, July 22, 1913, Weatherby (NEBC); FRANKLIN CO., Ashfield, June 19, 1921, Churchill, Hunnewell, Knowlton & Svenson (NEBC) ; BERKSHIRE CO., New Marl- boro, July 12, 1912, R. Hoffmann (NEBC). CONNECTICUT: TOLLAND co., Union, August 31, 1903, Graves (GH); HARTFORD CO., Hartland, 502 Rhodora [Vol. 68 Weatherby 3093. (NEBC) ; LITCHFIELD CO., Colebrook, September 5, 1909, Fernald (GH); FAIRFIELD CO., Danbury, Blewitt 2047 (NEBC). 12. Rubus elegantulus Blanch. PRIMOCANE and FLORICANE arching or less often doming 0.3-1 m high; armature of thin to acicular prickles 1-4 mm long, 10-60 (100) per dm; DIAMETER OF CANE 3-6 mm; PRIMOCANE LEAFLETS 5, membranaceous (occasionally charta- ceous), lanceolate, ovate or elliptic, 7-14 em long, apex acuminate or sometimes abrupt-acuminate, base of central leaflet cordate, lateral leaflet rounded, glabrous above and below: petiolules of central leaflet 2.5-4 (1.5-5) em long, longest lateral petiolules 1-2 (0.3-2.5) em long; DIMENSIONS OF COMPOUND LEAF 1111-2222 em: INFLORESCENCE in shape similar to R. vermontanus, 6-17 em long; floral axis pubes- cent often with small prickles but no glands; flowering in midseason, the number of flowers 3-15 (21); FRUIT undeveloped or small (rarely large and of good quality) ; HABITAT, old fields, moist or dry thickets, woods, bases of ledges, ascending to 1000 m in moist peaty soil; RANGE, northern New England where common, becoming less abun- dant southward in New Hampshire and Vermont and occasional in areas of high elevation in Massachusetts. Although this species was not recognized by Bailey, it seems to be a distinct and wide-ranging blackberry of north- ern New England. It would be difficult to understand the blackberry-flora without recognizing this as a species. It has been considered as a variety of R. canadensis, from which it is separated principally by the number of prickles on the primocane, and there are occasional states of R. can- adensis that are difficult to assign to either species. In most cases, however, R. elegantulus is closer to R. vermontanus, and can be distinguished only by careful examination as noted in the description of the latter species. Some of the mountain-forms are difficult to assign to either species and possibly are stable hybrid colonies of considerable age. Thus R. elegantulus represents a transition between R. vermont- anus and R. canadensis, but has a range of sufficient extent and is constant enough to be maintained as a species in its own right. It appears to cross more frequently with R. setosus than any other. Shorter hybrids are impossible to distinguish from hybrids of R. setosus and R. vermontanus, except by taking note of the occurrence of the putative par- ents in the area. The leaves are typically glabrous beneath as in R. cana- 1966] Rubus in New England — Hodgdon & Steele 503 densis. Good R. elegantulus may have very light pubescence on the main veins not noticeable without a lens. More ob- vious pubescence indicates introgression with R. pensilvani- cus, REPRESENTATIVE SPECIMENS: MAINE: OXFORD CO., Mason, Steele & Hodgdon 2773 (STEELE); WASHINGTON CO., Pembroke, July 6, 1909, Fernald & Wiegand (GH); HANCOCK CO., Mt. Desert Island, July 18, 1896, Rand (NEBC); LINCOLN CO., Boothbay Harbor, Hodg- don 12199 (NHA); South Bristol, Hodgdon 12174 (NHA); YORK CO., Kennebunkport, Steele & Hodgdon 3701 (STEELE); York, Steele & Hodgdon 3744 (STEELE). NEW HAMPSHIRE: coós co, Northumber- land, Fernald & Pease 15832 (NEBC); Dummer, Fernald & Pease 15792 (NEBC); CARROLL CO., Sandwich, Steele 3932 (STEELE) ; Mad- ison, Steele 3543 (STEELE); GRAFTON CO., Bath, Fernald 15668 (NEBC); Franconia, Hodgdon & Steele 11375 (NHA, STEELE) ; Rum- ney, Hodgdon & Steele 11271 (NHA, STEELE); STRAFFORD CO, New Durham, Hodgdon 9426 (NHA); Milton, '‘Hodgdon & Steele 11270 (NHA); ROCKINGHAM CO., Seabrook, Hodgdon & Steele 11261 (NHA); HILLSBORO CO., Sharon, Hodgdon 7216 (NHA). VERMONT: ESSEX CO., Granby, Pease 26804 (NEBC); WINDHAM CO., Saxtons River, Steele 3548 (STEELE); BENNINGTON CO., Woodford, June 29, 1939, Knowlton (NEBC). MASSACHUSETTS: BERKSHIRE CO., Pittsfield, July 14, 1919, Hoffmann (NEBC). CONNECTICUT: TOLLAND CO., Willington, Weather- by 5322 (NEBC). 13. Rubus alleghenicnsis Porter. PRIMOCANE and FLORICANES erect to arching 1-2 m high, sometimes low-doming in late summer; AR- MATURE of broad-based stout or occasionally thin priekles 2-7 mm long, 1-20 per dm.; PRIMOCANES glabrous or pubescent, mostly glandu- lar particularly on upper parts and early in the season: FLORICANES usually glabrous and glandless; DIAMETER OF PRIMOCANE 4-8 (3-9) mm; PRIMOCANE-LEAFLETS 5, membranaceous ovate, ratio of width to length 0.6-0.75, acuminate, the terminal leaflets cordate to sub- cordate, the lateral leaflets rounded to subcordate at base, 8-15 mm long, glabrous to pubescent above, velvety beneath, central petiolules 2.6 cm long and longest laterals 1.5-5 m long; DIMENSIONS OF COM- POUND LEAF 1212-23238 cm; INFLORESCENCE when best developed and most typical forming cylindric racemes 8-20 cm long but often reduced to short racemes or corymbs to 3 em long, flowers 9-22 in number but at times reduced to as few as 3 in the shorter clusters, axis of inflorescence pubescent and with abundant glands, with or without prickles; pedicels divergent to ascending 1.5-5 cm long, some- times with weak scattered prickles; early flowering; FRUIT elongate to thimble-shaped or less often globular, of good quality; HABITAT, roadsides, old fields and cut over woods, RANGE, throughout New England at low elevations, common. 504 Rhodora [Vol. 68 In spite of considerable variation, this is one of the most distinctive and earliest known species of blackberry. It is the common high blackberry, when well developed, produc- ing long cylindrical racemes of good fruit. It is widespread, occurring throughout New England in fields and roadsides and quickly invading new lumbered areas. Chromosome counts indicate that it is one of the few diploid species. In good sites the globular, or more commonly thimble-shaped, fruits occur in abundance and are easy to pick. This species and some of the Arguti occasionally have flowering primocanes, forming inflorescences that would be difficult to classify, unless their true nature is appreciated. An inflorescence with each single flower subtended by a leaf belongs in this category. Primocanes are fresh-green with buds at the end; floricanes are red or purple often with dead ends. Although R. allegheniensis is normally arching and 4 to 6’ tall, stunted specimens only 1' or 2' high are frequent. These might be mistaken for another species, but the glan- dular floral axis and the soft pubescence on the underneath side of the leaves give clues as to the true nature of the plant. Colonies of this species have been found in which some of the outer plants dome and reach the ground. This may represent an extreme state of the plant modified by environmental factors or it may represent introgression with another species. Rubus allegheniensis, on occasion, hybridizes with most other species. These hybrids are infrequent to rare, the most frequent being a cross with R. setosus which gives rise to various forms intermediate between the two parents, sev- eral of which have been designated as species. REPRESENTATIVE SPECIMENS: This species is well enough understood and clearly enough marked to require a minimum of specimen citations. MAINE: PENOBSCOT co., Milford, Fernald 13867 (NEBC) ; PISCATAQUIS CO., Dover, September 1, 1896, Fernald (NEBC) ; SOMERSET CO., Fairfield, Fernald & Long 13862 (NEBC) ; WASHINGTON co., Lubec, Hodgdon & Pike 7223 (NHA); HANCOCK CO., Dedham, Fernald & Long 13861 (NEBC) ; KNOX CO., Isle au Haut, September 3, 1918, Kidder (NEBC); LINCOLN CO., Southport, August 2, 1894, Fer- nald (NEBC); ANDROSCOGGIN CO. Auburn, August 29, 1932, Bean 1966] Rubus in New England — Hodgdon & Steele 505 (NEBC); YORK CO. Wells, Fernald & Long 13860 (NEBC). NEW HAMPSHIRE: C00s CO., Dummer, Fernald & Pease 15793 (NEBC) ; CAR- ROLL C0., Conway, Hodgdon & Steele, 11277 (NHA); GRAFTON CO., Wocdstock, Fernald 11752 (NEBC); STRAFFORD CO., Durham, Hodg- don 3031 (NEBC); MERRIMACK CO. Hooksett, August 24, 1925, Batchelder (NEBC) ; SULLIVAN CO., Plainfield, August 9, 1962, Cowden, Eaton, Hodgdon, Poole & Wilson (NHA); ROCKINGHAM CO., Rye, Pease 16937 (NEBC); CHESHIRE CO. Hinsdale, August 28, 1919, Batchelder (NEBC). VERMONT: ESSEX CO., Concord, Pease 33722 (NEBC); ORLEANS CO., Newport, June 7, 1913, Knowlton (NEBC) ; CHITTENDEN CO., Hinesburg, July 17, 1940, Knowlton (NEBC) ; WIND- SOR CO., Hartford, June 12, 1920, Eaton & St. John (NEBC); RUTLAND co., Wallingford, June 21, 1909, Kent (NEBC) ; WINDHAM CO., Vernon, June 19, 1925, Churchill & Knowlton (NEBC). MASSACHUSETTS: ESSEX CO., Manchester, June 7, 1913, Hubbard (NEBC); MIDDLESEX co., Concord, June 20, 1958, Eaton (NEBC); PLYMOUTH CO., Marsh- field, June 26, 1926, Knowlton (NEBC); BARNSTABLE CO., Falmouth, August 9, 1927, Fernald (NEBC); DUKES CO. Marthas Vineyard, September 23, 1913, Bicknell (NEBC); WORCESTER CO., Douglas, Fer- nald 15226 (NEBC): FRANKLIN CO., Colerain, June 18, 1921, Churchill, Knowlton & Schweinfurth (NEBC) ; HAMPSHIRE CO., Ware, September 2. 1916, Forbes (NEBC); BERKSHIRE CO., Sheffield, August 27, 1902, Hoffmann (NEBC). RHODE ISLAND: PROVIDENCE CO., East Providence, May 30, 1911, Wiegand (NEBC): BRISTOL CO., Barrington, May 30, 1911, Winslow (NEBC). CONNECTICUT: WINDHAM CO., Thompson, Weatherby 5034 (NEBC); NEW LONDON CO., Franklin, July 5, 1912, Woodward (NEBC); TOLLAND CO., Bolton, Weatherby 5318 (NEBC) ; HARTFORD CO., Hartland, Weatherby 4343 (NEBC); MIDDLESEX CO., Cornwall, Weatherby 4302 (NEBC); NEW HAVEN CO, Waterbury, Blewitt 1314 (NEBC); LITCHFIELD CO., Norfolk, June 10, 1919, Evans (NEBC) ; FAIRFIELD CO., Danbury, July 19, 1912, Harger (NEBC). 14. Rubus argutus Link. PRIMOCANE and FLORICANE erect or high- arching (occasionally low-arching) 0.9-1.8 m high; ARMATURE of thin to stout prickles, 4-6 mm long, 10-30 per dm; DIAMETER OF CANE 3-5 (2-7) mm; PRIMOCANE-LEAFLETS 5, membranaceous, lanceolate, narrow, the ratio of width to length. 0.4-0.45, acuminate at apex, 8-11 cm long, finely pilose to glabrous above, sparsely pilose to velvety beneath, rounded to cuneate at base, the central base cuneate or subcuneate, the lateral rounded; DIMENSIONS OF COMPOUND LEAF 1318-17 x 17.em; INFLORESCENCE a short diffuse raceme (corymb) 3-7 em long and 3-5 em across, the flowers 5-10 (4-12) in number, the axis and pedicels often with prickles, glands absent; medium-to late-flowering; HABITAT, thickets, edges of woods and bog-margins; RANGE, central and south- ern Massachusetts and southward at low elevations, uncommon. The three species; R. argutus, R. frondosus and R. pen- silvanicus are often classed as Arguti. Rubus argutus ap- 506 Rhodora [Vol. 68 pears to be a distinctive blackberry, although of limited range. The leaflets are narrow, less than one half as broad as long, and the base of the central one is wedge-shaped. In other related species, the bases are rounded, or, more usually heart-shaped. The species appears to be uncommon in New England but more frequent southward. There a number of collections from Virginia. In Massachusetts the northernmost station is south of Boston, and most of the other stations are on Cape Cod. It appears to be commonest at or near the coast and absent from upland or mountainous regions. No doubt with the development of the coast much suitable environ- ment for it was destroyed and it may have been more com- mon in the past. REPRESENTATIVE SPECIMENS: MASSACHUSETTS: NORFOLK co., Stoughton, Blake 12127 (GH, Type of R. Blakei Bailey) ; BRISTOL co., New Bedford, June 9, 1910, Hervey (NEBC); BARNSTABLE CO., Harwich, Fernald & Long 16924 (GH, type of R. paludivagus Fer- nald); Bourne, Fernald & Clark 16898 (NEBC) ; Barnstable, St. John & Hubbard 11726 (NEBC) ; Falmouth, August 9, 1927, Fernald 608 (NEBC); Brewster, Steele & Hodgdon 3915 (NHA, STEELE). CON- NECTICUT: NEW LONDON Co. Franklin, July 8, 1915, Woodward (NEBC). 15. Rubus frondosus Bigel. PRIMOCANE and FLORICANE erect to arching or even doming to 0.9-1.9 m high; ARMATURE of thin to stout straight prickles 4-6 mm long, 4-60 in number per dm; DIAMETER OF CANE 4-6 (3-7) mm; PRIMOCANE LEAFLETS 5, subchartaceous to chartaceous (occasionally membranaceous), ovate to obovate-rhombic, leaflets often short and broad, ratio of width to length 0.6-0.8 (0.55), acuminate, 6-12 cm long, subglabrous to glabrous above, velvety be- low, terminal leaflet cordate to subcordate or rounded at base, lateral leaflets with bases rounded to subcuneate: petiolules of central leaf- let from 1.2-3 (4.5) em long, of middle lateral leaflets 0.1-1 (2.3) cm long; DIMENSIONS OF COMPOUND LEAF 1111-1919 cm; IN- FLORESCENCE a compact raceme (corymb) 4-9 em long by 2-4 em broad, flowers 3-10 (2-12) in number, axis pilose with few to 0 prickles, leafy bracts 1-7, sometimes lacking, often conspicuous; FRUIT globular to subglobular, large, juicy and often of exceptional quality; HABITAT, thickets, old fields and roadsides; RANGE, coast of Maine, southern New Hampshire and southern Vermont southward, common at low elevations, This species is essentially southern in distribution, reach- ing its northern limit along the coast of Maine, and is more 1966] Rubus in New England — Hodgdon & Steele 507 distinctive in the southern part of its range where it is as common as R. allegheniensis and occupies similar environ- ments. It often has good fruits and presumably was the source of some cultivated varieties. It is closest in appearance to Rubus pensilvanicus and no doubt frequently intergrades with it. There is a difference in the texture of the leaves, those of R. frondosus being darker and subchartaceous, which gives the plant a distinc- tive appearance even at a distance. The prickles tend to be stouter and more numerous than those of R. pensilvanicus and the fruit is of better quality. The primocanes frequently have lateral branches which elongate and may crawl over bushes or reach the ground, thus giving the plant a doming appearance. We include the state described as R. multispinus in our concept of R. fron- dosus. Because there is so much variation in the degree of doming in R. frondosus itself it would not seem to be a re- liable character on which to separate two species. In some instances the doming habit is caused no doubt by intro- gression with some of the trailing species. Although there seems to be a tendency in Rubus fron- dosus to have pronounced leafy bracts in the inflorescence this is by no means a constant or reliable character; often the inflorescences in perfectly good material are no more leafy than R. pensilvanicus. REPRESENTATIVE SPECIMENS: MAINE: KNOX CO. Matini- cus, July 9, 1920, C. A. E. Long (NEBC); Isle Au Haut, August 2 1919, Kidder (NEBC); LINCOLN CO., Boothbay Harbor, August 19, 1955, Steele & Hodgdon 2154 (STEELE); Southport, Hodgdon 7230 (NHA). NEW HAMPSHIRE: STRAFFORD CO., Durham, Hodgdon & Steele 12188 (NHA); HILLSBORO CO., Hollis, Steele & Hodgdon 4009 (STEELE). MASSACHUSETTS: SUFFOLK CO., Boston, June 25, 1908, Rich (NEBC); BARNSTABLE CO., Harwich, Fernald 16895 (NEBC) ; Harwich, Fernald 16917 (NEBC) ; Harwich, Steele & Hodgdon 3925 (NHA, STEELE) ; Chatham, Fernald & Long 18578 (NEBC) ; Brewster, Hodgdon & Steele 14845 (NHA); DUKES CO., Chilmark, Steele 4011 (STEELE); NANTUCKET CO., Nantucket, Bicknell 11932 (NEBC); WOR- CESTER CO., Sterling, Hodgdon & Steele 14846 (NHA); Berlin, Hodg- don & Steele 14847 (NHA); HAMPDEN CO., Springfield, June 30, 1914, Andrews (NEBC). RHODE ISLAND: WASHINGTON co., Charlestown, Hodgdon & Steele 14848 (NHA). CONNECTICUT: NEW LONDON CO., 508 Rhodora [Vol. 68 Mystic, Hodgdon & Steele 14849 (NHA); Salem, Steele & Hodgdon 3906 (STEELE); TOLLAND CO., Stafford Springs, Hodgdon & Steele 14850 (NHA); HARTFORD CO., Hartford, Wright (NEBC); Berlin, Steele & Hodgdon 3931 (NHA, Steele); Southington, Steele & Hodg- don 3910 (STEELE, NHA); MIDDLESEX CO., Haddam, Hodgdon & Steele 148501 (NHA); NEW HAVEN CO., Orford, Harger 5552 (NEBC). 16. Rubus pensilvanicus Poir. PRIMOCANE and FLORICANE erect to somewhat arching 0.6-2.2 m high; ARMATURE of thin to medium prickles 2-4 (5) mm long, tending to be rather few, 0-20 (27) per dm; DIAMETER OF CANE 4-9 (2-10) mm; PRIMOCANE LEAFLETS 5, mem- branaceous to subchartaceous, ovate, ratio of width to length 0.55- 0.75 (0.5), apex acuminate, 8-13(16) em long; glabrous or nearly so above, moderately pilose to velvety beneath, leaf base cordate or subcordate to rounded; petiolules of central leaflet 2-5(6) cm long, of medium lateral leaflets 0-2.5(4.5) cm, the larger lateral petiolules frequently more than 1 cm long; DIMENSIONS OF COMPOUND LEAF 11X 11-22 22(3030), often large; INFLORESCENCE varying, from long to short and somewhat compact racemes (corymbs), flowers 7-12 (4-20) in number, axis pilose with occasional prickles and no glands, leafy bracts 1-2(0-6); FRUIT fair to good, not often produced in abundance; HABITAT, thickets, roadsides, swamps and woods; RANGE, throughout New England, at low elevations, scarcer north- ward, a scattered and not very common species. Although this Species has a wide distribution extending from northern New Hampshire southward throughout New England, it has certain puzzling aspects and is not always clearly defined. A plant from the northernmost known New Hampshire station, in Franconia Notch, resembles R. cana- densis except for a thin pubescence on the underneath side of the leaves. Specimens from the few scattered stations immediately southward also resemble R. canadensis or a hybrid of R. allegheniensis and R. canadensis. Thus there is a question about the specific identy of the northern speci- mens, which might account for some of the diversity. In southern New Hampshire the plant becomes more com- mon and better marked. The prickles instead of being in- frequent and minute as in R. canadensis are of medium-size and more abundant. In southern New England the prickles tend toward those of R. frondosus. Folders in herbaria contain some plants with slightly glandular inflorescences, which must represent some in- fluence of R. allegheniensis. Chromosome counts indicate 1966] Rubus in New England — Hodgdon & Steele 509 that this plant may contain both triploid and tetraploid ele- ments, which might account for some of the diversity. This is another of the species of Rubus in which flowering primocanes are not uncommon and represent a possible source of confusion. REPRESENTATIVE SPECIMENS: MaINE: PENOBSCOT CO., Mil- ford, Fernald 13866 (NEBC) ; OXFORD CO., Bethel, Pease 37595 (NEBC) ; LINCOLN CO., Monhegan Island, September 2, 1921, Churchill (NEBC) ; Boothbay Harbor, Hodgdon 2719 (NHA); KENNEBEC CO., Clinton, July 17, 1915, Bean (NEBC); ANDROSCOGGIN CO., Auburn, August TT, 1916, Bean (NEBC); YORK CO., Kennebunkport (Parson Beach), July 8, 1905, Blanchard (NEBC) ; Kennebunkport, Hodgdon & Steele 12485 (NHA). NEW HAMPSHIRE: Cos C0., Dummer, Harris 26588 (NEBC) ; Jefferson, Pease 16899 (NEBC); Northumberland, Fernald & Pease 15675 (NEBC); Colebrook, Fernald & Pease 15794 (NEBC); CARROLL co., Tuftonboro, Hodgdon & Steele 12187 (NHA); Sandwich, Steele 4001 (STEELE); GRAFTON CO., Haverhill, Fernald 15769 (NEBC) ; Bath, Fernald 15593 (NEBC); Thornton, Hodgdon, Steele, & Lincoln 11265 (NHA, STEELE); Rumney, Hodgdon & Steele 11297 (NHA, STEELE) ; STRAFFORD CO., New Durham, Hodgdon & Steele 12259 (NHA); Dur- ham, Hodgdon & Steele 11298 (NHA); MERRIMACK CO., Hooksett, August 16, 1925, Batchelder (NEBC); Newbury, Hodgdon 11377 (NHA); SULLIVAN CO., Plainfield, August 9, 1962, Cowden, Eaton, Hodgdon, Poole & Wilson (NHA); HILLSBORO CO., Peterboro, Batch- elder 693 (NEBC); CHESHIRE CO., Nelson, Batchelder 2812 (NEBC). VERMONT: RUTLAND CO. Wallingford, September 1, 1912, Kent (NEBC); BENNINGTON CO. Searsburg, June 20, 1925, Carpenter, Churchill & Knowlton, (NEBC). MASSACHUSETTS: MIDDLESEX CO., Arlington, August 15, 1885, Batchelder (NEBC); Concord, July 13, 1935, Eaton (NEBC); BRISTOL CO., New Bedford, June 15, 1900, Her- vey (NEBC); BARNSTABLE CO. Sandwich, Fernald & Long 18575 (NEBC) ; Falmouth, August 9, 1927, Fernald (NEBC); WORCESTER CO., Harvard, August 6, 1916, Forbes (NEBC); FRANKLIN CO., Shelburne Falls, July 4, 1921, Churchill (NEBC) ; BERKSHIRE CO., New Marlboro, July 9, 1912, Hoffmann (NEBC). CONNECTICUT: WINDHAM CO., Plain- field, Harger 6419 (NEBC); NEW LONDON CO., Groton, Hodgdon & Steele 14852 (NHA); HARTFORD CO, New Canaan, Steele 3934 (STEELE) ; LITCHFIELD CO., Colebrook, Weatherby 2680 (NEBC) ; FAIR- FIELD CO., Stratford, Hames 2120 (NEBC). REJECTED SPECIES The following list of rejected New England species pro- vides a suggested interpretation of each. No attempt is made to deal here with subspecific taxa. The generic name ma 510 Rhodora [Vol. 68 Rubus is understood and the name of Bailey is also con- sidered to be superfluous as the author of most of the names. abactus = pensilvanicus abbrevians Blanch. = setosus X allegheniensis aculiferus Fern. = allegheniensis adjacens Fern. = setosus alius = arenicola alter = hispidus X setosus alumnus = allegheniensis X pensilvanicus amicalis Blanch. = elegantulus amnicola Blanch. — pensilvani- cus Andrewsianus silvanicus apparatus = vermontanus aptatus = recurvicaulis arcuans Fern. & St. John — re- curvicaulis X setosus arundelanus Blanch. = recurvi- caulis ascendens Blanch. — semisetosus auroralis = allegheniensis avipes = pensilvanicus Baileyanus Britt. (type quate = Enslenii barbarus — pensilvanicus bellobatus = frondosus Bicknellii = setosus X recurvi- caulis biformispinus Blanch. = hispi- dus X allegheniensis Bigelovianus = semisetosus Blakei — argutus Blanchardianus = setosus Boottianus = setosus Brainerdii Rydb. = arenicola brevipedalis = recurvicaulis coloniatus = recurvicaulis conanicutensis = pensilvanicus cubitans Blanch. — hispidus setosus hispidus Blanch. = pen- inade- hispidus X curtipes — arenicola Deaneanus = vermontanus eflagellaris — hybrid involving one of the Flagellares electus = hispidus X alleghe- niensis facetus — pensilvanicus felix — flagellaris Fernaldianus = allegheniensis frondisentis Blanch. — setosus geophilus = flagellaris glandicaulis Blanch. — setosus X allegheniensis gnarus = identity unknown Gravesii (Fern.) Bailey = al- legheniensis Groutianus Blanch. = x vermontanus harmonicus = hispidus X seto- sus heterogeneous = frondosus heterophyllus Willd. — identity unknown hispidoides — semisetosus honorus — pensilvanicus insons = pensilvanicus insulanus = frondosus invisus (Bailey) Britt. — not in New England according to Bailey setosus jacens Blanch. = hispidus X setosus jactus = hispidus X allegheni- ensis Janssonii — arenicola Jeckylanus Blanch. — recurvi- caulis jugosus = argutus junceus = vermontanus junior = setosus Kennedyanus Fern. = elegantu- lus laevior (Bailey) Fern. = his- 1966] pidus X allegheniensis latens — hybrid involving alle- gheniensis Lawrencei = setosus licitus — allegheniensis X pen- silvanicus longissimus = allegheniensis mainensis — hispidus X flagel- laris maniseesinsis — flagellaris Millspaughii Britt. — canaden- sis miscix —— vermontanus X pen- silvanicus montpelierensis Blanch. = seto- sus X allegheniensis multiformis Blanch. = elegan- tulus multilicius — vermontanus >d elegantulus multispinus Blanch. = frondo- sus nigricans Rydb. — setosus nigrobaccus — allegheniensis notatus = setosus novanglicus — identity unknown obsessus =— arenicola orarius Blanch. = pensilvanicus ortivus =— semisetosus ostryifolius Rydb. = pensilvani- cus paludivigus Fern. = argutus Parlinii — setosus X vermonta- nus paulus = allegheniensis X pen- silvanicus pauper = arenicola peculiaris — vermontanus X pensilvanicus pergratus Blanch. = pensilvani- cus perinvisus — semisetosus permixtus Blanch. = hispidus X allegheniensis perpauper = arenicola pervarius = hispidus Rubus in New England — Hodgdon & Steele 511 philadelphicus Blanch. = silvanicus plicatifolius Blanch. = recurvi- caulis positivus = recurvicaulis procumbens Muhl. = flagellaris prosper Bailey = arenicola pudens = hispidus X setosus pugnax = allegheniensis Randii Rydb. — canadensis (de- pauperate) ravus = vermontanus X alle- gheniensis recurvans Blanch. = frondosus rhodinsulanus = recurvicaulis rixosus — hispidus X setosus roribaccus (Bailey) Rydb. = doubtfully present in New England, an escape from cul- tivation in the North pen- Rosa = allegheniensis X pen- silvanicus Rossbergianus Blanch. = fron- dosus saltuensis — allegheniensis Sanfordii == hispidus X alle- gheniensis sativus Brain. — allegheniensis scambens — Jaysmithii (speci- men called R. scambens from New England is flagellaris sceleratus Brain. — setosus X allegheniensis semierectus Blanch. = recurvi- caulis sempervirens Bigel. = Enslenii severus Brain. = recurvicaulis X vermontanus significans == setosus singulus = vermontanus spiculosis Fern. = hispidus tardatus Blanch. — extreme of vermontanus tetricus — Jaysmithii tholiformis Fern. — hispidus X setosus 512 Rhodora [Vol. 68 trifrons Blanch. — hispidus X wnivocus — setosus X vermont- setosus anus udus = setosus vigoratus — hispidus X setosus usus = recurvicaulis viridifrons — hispidus X ver- MONTANUS DEPARTMENT OF BOTANY, UNIVERSITY OF NEW HAMPSHIRE DURHAM ST. MARY'S-IN-THE-MOUNTAINS, LITTLETON, NEW HAMPSHIRE LITERATURE CITED BAILEY, L. H. 1932. Gent. Herb. Il: 271-423, —————— 1984. Gent. Herb. III: 247-271. — — —— 1941-1945. Gent. Herb. V: 1-932. — — ——— 1947. Gent. Herb. VII: 181-349. L—— . 1949. Gent. Herb. VII: 481-526. BICKNELL, E. P. 1910. Have we enough New England Blackberries. Bull. Torr. Bot. Club. 37: 393-403. BRAINERD, E. & A. K. PEITERSEN. 1920. Blackberries of New Eng- land. Vt. Ag. Exp. Sta. Bull. 217: Burlington, Vermont. CRAIG, D. L. 1959. The Cytology and Breeding Behaviour of Rubus canadensis L. Thesis, Univ. of New Hampshire, Durham, New Hampshire. EINSET, J. 1945. Chromosome Studies in Rubus. Gent. Herb. 7: 181-192. —————. 1951. Apomixis in American Polyploid Blackberries. Am. Jour. Bot. 38: 768-772. FERNALD, M. L. 1950. Gray’s New Manual of Botany 8th Ed.: 822. GLEASON, H. A. and ARTHUR CRONQUIST. 1963. Manual of Vascular Plants: 375-379. GUSTAFSSON, A. 1946. Apomixis in higher plants. Part I. The mechanism of Apomixis Acta Univ. Lund. II 42: 1-67. 1947a. Apomixis in higher plants. Part II. The casual aspect of Apomixis. Acta Univ. Lund. II 43: 71-179. ————————————. 194b. Apomixis in higher plants. Part LII. Bio- type and species formation Acta Univ. Lund. II 43: 183-370. HopGDON, A. R. and F. L. STEELE. 1962. Glandularity in Rubus AI- legheniensis Porter. Rhodora 64: 161-168. LoNGLEY, A. E. 1924. Cytological Studies in the Genus Rubus, Am. Jour. Bot. 11: 249-280. 1966] Sisymbrium pauciflorum — Rollins 513 PEITERSEN, A. K. 1921. Blackberries of New England — Genetic Studies. Vt. Ag. Exp. Sta. Bull: 218 Burlington, Vermont. RYDBERG, P. A. 1913. North American Flora. 22: Part 5, 428-480. STEELE, F. L. and A. R. HODGDON. 1963. Hybridization of Rubus hispidus and R. setosus. Rhodora 65: 262-270. VALENTINE, D. H. and ASKELL LOVE. 1958. Taxonomic and Biosys- tematic categories. Brittonia 10: 153-166. THE IDENTITY OF SISYMBRIUM PAUCIFLORUM (CRUCIFERAE). — In the Flora of North America, Tor- rey and Gray (1838) published as Sisymbrium pauciflorum, a name from Nuttall’s manuscript that has since been largely ignored. Robinson (1895) put it succinctly when he stated, “S. PAUCIFLORUM, Nutt. .. .. has always been obscure, and appears to have been founded upon immature specimens of Arabis canescens.” Payson (1922) in his treatment of Sisymbrium failed to mention S. pauciflorum, even though he attempted to account for the North Ameri- can species of the genus, north of Mexico. Also, Floras covering the Rocky Mountain Region, where the type ma- terial of S. pauciflorum was collected by Nuttall, have not taken the name into account. My own earlier attempt to deal with the name (Rollins, 1941) was prompted by Robinson's suggestion that the plant Nuttall described belonged to the genus Arabis. On the basis of the original description and some notes about the type provided by a colleague, it was clear to me then that the type specimen was indeed an Arabis. Since the specific epithet pauciflora was preoccupied by Arabis pauci- flora (Grimm) Garcke, the exact positioning of the Nuttall name in Arabis as to species had no particular nomencla- tural consequences and the most abundant species of Arabis to which the description and notes might apply was A. holboellii, var. retrofracta, this is where I listed Sisymbrium pauciflorum as a synonym. A few months ago, I was privileged to examine the type of Sisymbrium pauciflorum in the British Museum. The identity is unmistakable. It is Arabis fendleri var. spati- 514 Rhodora [Vol. 68 folia (Rydberg) Rollins. This is rather unexpected because the taxon is not commonly found in Wyoming on Nuttall's collecting route. However, the original description is accu- rate and now that the identity is known, there is no diffi- culty in following it. REED C. ROLLINS GRAY HERBARIUM, HARVARD UNIVERSITY LITERATURE CITED PavsoN, E. B. 1922. Species of Sisymbrium Native to North America North of Mexico. Univ. Wyo. Publ. 1: 1-27. ROBINSON, B. L. 1895. Synoptical Flora of North America 1: 138. RoLLINS, R. C. 1941. A Monographie Study of Arabis in Western North America, Rhodora 43: 441. TORREY, J. and A. GRAY, 1838. Flora of North America 1: 93. BOTANICAL NOTES FROM THE PRINGLE HERBARIUM I. In the course of compiling data at the Pringle Herbarium of the University of Vermont for my forthcoming book on the Flora of New England, a number of items have come to my attention which seem worth publication. Most of these items consist of species new to the State of Vermont, meaning that they are not listed in the latest edition of the Flora of Vermont by E. J. Dole (1937). A new edition of that Flora is in preparation by this author. In some cases, the occurrence of these taxa may have been published else- where, but not to my knowledge. Even so, it is desirable for them to be listed together under the heading of Ver- mont. The voucher specimens unless otherwise indicated are deposited in the Pringle Herbarium. l. JUNIPERUS COMMUNIS L. var. COMMUNIS. Vermont: Dorset Hollow, 6 Aug. 1965, Alfred H. Gilbert. Mr. Gilbert writes, 6 Aug. 1965: “This is distinctly an upright form." 1966] Botanical Notes — Seymour 515 2. BROMUS COMMUTATUS Schrader. Occasional, several lo- calities; the earliest coll. C. G. Pringle as early as 1880, without definite locality. This species has been much con- fused with B. secalinus L. from which it differs in having the margins of the lemmas concealing the rachis, not in- rolled as in B. secalinus. 3. BROMUS COMMUTATUS Schrader forma perglaber forma nova. B. commutatus f. commutatus has the sheaths, even the upper ones, with abundant long spreading hairs. In some specimens of this species, all the sheaths are glabrous in striking contrast to the typical form. Therefore, the specimens with glabrous sheaths deserve recognition by name. I designate them as forma perglaber forma nova. Vaginis glabris. Type: Maine, North Berwick, July 1893, J. C. Parlin (NEBC). Among other specimens of this new form are: New Hampshire, Dover, 18 June 1937, A. R. Hodgdon (NEBC) ; Mass., Swampscott, 13 June 1891 J. KR. Churchill. (NEBC) ; Dedham, 27 June 1897, F. F. Forbes (NEBC). An example of an intermediate is: Connecticut: Fairfield, 12 June 1895, E. H. Eames (NEBC), in which the lowest sheaths are slightly hairy. 4. ERAGROSTIS PILOSA (L.) Beauv. Vermont: Woodstock, Miss E. Billings, 1905. This species is easily confused with E. pectinacea (Mx.) Nees. In the specimen cited, the spike- lets are not over 1 mm wide and the branches of the panicle are thread-like and flowerless near the main axis. The Manual of Grasses, ed. 2, by Agnes Chase, figure 201, indi- cates by a dot that it has been found in Vermont, but this specimen by Miss Billings is the only one I have seen from Vermont. 5. AIRA CARYOPHYLLEA L. A considerable colony in a drive- way in front of a tent-site in the State Camping Grounds in Stillwater, Groton, near the shore of Lake Groton, 27 June 1963, Seymour 20, 650. 6. ELEUSINE INDICA (L.) Gaertner. Hartford Center, 9 516 Rhodora [Vol. 68 Sept. 1938, L. A. Charette 858. This species appears to bo spreading rapidly but has not been reported previously from Vermont. 7. PHLEUM PRATENSE L. var. NODOSUM (L.) Hudson. Burl- ington and Middlebury. This variety is doubtless abundant, but it has not previously been reported from Vermont. 8. LEPTOLOMA COGNATUM (Schultes) Chase. Burlington, 10 Aug. 1938, E. J. Dole 642. Misidentified as Eragrostis spectabilis (Pursh) Steudel var. sparsihirsuta Farw. which it strongly resembles in general habit. This new station for an uncommon grass is noteworthy because it is far from the other known stations in Vermont, all of which are in the valley of the Connecticut River. FRANK C. SEYMOUR, PRINGLE HERBARIUM, UNIVERSITY OF VERMONT, BURLINGTON A NEW METHOD FOR MAINTAINING THE CONES OF ABIES AND CEDRUS INTACT FOR STUDY AND STORAGE In the cones of Abies and Cedrus, both the scales and bracts together become deciduous at maturity. Unless some means of preservation is accomplished, these cones disinte- grate leaving a somewhat bare, spike-like persistent axis. In the past those wishing to retain cones of Abies or Cedrus in good condition for herbaria, display, or as teaching aids were forced to resort to binding each cone with wire or string. This method is often unsuccessful and results in un- attractive specimens which must be handled with much care. We have found that the clear plastic described by Archer (1950) and now used by many herbaria for mounting speci- mens can be used for treating cones of Abies and Cedrus (Figure 1). The plastic is prepared according to Archer’s formula as modified by Rollins (1955), and placed in a 1966] Cones — MacDonald and MacDonald 517 Plate 1337. Cones which have been treated with clear plastic; twelve months after treatment. Left to right: Abies magnifica, Abies concolor, Cedrus atlantica. small, easy to handle, glass container (a 16-ounce size works well). Specimens to be treated should have 14- to 14-inch of the stem attached. After tying a 6-inch piece of string to the stem at the base of the cone, hold the cone by the base over a disposable tray and slowly pour plastic over it. Rotate the cone to insure a thin, even coating. It is imporant that the entire cone be coated and that no holes are left. When the cone is covered, it should be hung by the string to dry. As it dries excess plastic should be trimmed from the tip of the cone. R. D. MACDONALD AND M. E. MACDONALD UNIVERSITY OF TENNESSEE ARBORETUM, OAK RIDGE BIBLIOGRAPH Y ARCHER, W. ANDREW. 1950. New plastic aid in mounting herbarium specimens. Rhodora 52: 298-299. RoLLINs, REED C. 1955. The Archer method for mounting her- barium specimens. Rhodora 57: 294-299. SOME PLANTS NEW TO TENNESSEE! Plant collections made during the summer of 1965: in- cluded representatives of six genera and fifteen species and varieties which are, to the best of our knowledge, previously unreported from Tennessee. Specimens have been deposited in the State Herbarium at The University of Tennessee. Nomenclature and systematic arrangement of the list is according to Fernald (1950), and Hitchcock (1951). The annotations include the county in which collected, the col- lection number or numbers, the general range of the taxon, and comments on the present collection if it represents an extension in range. POA PALUSTRIS L. Carter County. 34665. The general range is from Newfoundland to Alaska, south to Virginia, western North Caro- lina, West Virginia, Missouri, Nebraska, New Mexico, Colorado and California. Our collection extends its range to include Tennessee. ERAGROSTIS INTERMEDIA Hitche. Shelby County. 33510. This species is reported from Georgia to Arizona and New Mexico, north to Missouri and Kansas. The collection in west Tennessee appears to represent a northern extension of its known range. TRIPLASIS PURPUREA (Walt.) Chapm. Shelby County. 34465, 33525. Ontario and Maine to Minnesota and Nebraska, south to Florida and Texas; Colorado. SPOROBOLUS CRYPTANDRUS (Torr.) A. Gray. Shelby, Lake Counties. 34771, 34467, 33257, 33532. Maine to Washington, south to North Carolina, West Virginia, Indiana, Illinois, Louisiana, Texas, south- ern California, and Mexico. MUHLENBERGIA SOBOLIFERA var. SETIGERA Scribn. Sevier County. 34291. This variety has been reported only from Arkansas and Texas. Our collection is an eastward extension in range. CHLORIS VERTICILLATA Nutt. Hardin County. 33603. Nebraska to Texas, and southern California; Maine and Massachusetts; Ohio, Indiana, and North Carolina to Florida; Louisiana and Missouri. The collection from near the Tennessee River in Tennessee was 'Contributions from the Botanical Laboratory, The University of Tennessee, N. S. 256. "Field collecting was supported primarily by a contract between the Tennessee Game and Fish Commission and The University of Ten- nessee, and was part of a continuing study for the preparation of a manual of grasses, their identification, habitat, distribution, and use by game species in Tennessee, 518 1966] Tennessee Plants — Rogers and Underwood 519 from an area sown to cultivated grass. This is a southward ex- tension in its recorded range. LEPTOCHLOA UNINERVIA (Presl) Hitchc. and Chase. Shelby County. 34062, 33567, 33505, 33649. North Carolina; Mississippi to Colo- rado and California, south to Argentina; introduced in Maine, Massachusetts, and New Jersey. Our collections appear indigenous and thus represent a northward extension in its known natural range from the Coastal Plain. ERIOCHLOA GRACILIS (Fourn.) Hitche. Lake County. 33672. Oklahoma io southern California, south to Mexico. This collection is from a county that is 80-90% under cultivation, and it very probably represents an introduction. ERIOCHLOA CONTRACTA Hitche. Lake County. 33671. Nebraska to Colorado, Louisiana and Arizona; adventive in Missouri and Vir- ginia. This species, as with the preceding one, is primarily a western grass, and its occurrence in Tennessee represents an east- ward extension of range. PANICUM SPHAEROCARPON var. INFLATUM (Scribn. & Sm.) Hitchc. Shelby, Fayette, and Dyer Counties. 33511, 335 46, 33518, 33696, 33515. Coastal Plain, Delaware to Florida and Texas, north to Oklahoma and Missouri. JUNCUS VALIDUS Coville. Hardeman County. 34907. Florida to Texas, north to Georgia, Missouri, and Oklahoma. CYCLOLOMA ATRIPLICIFOLIUM (Spreng.) Coult. Shelby County. 33569, 34773. Mississipi to Texas, Arizona, Manitoba, and Massachusetts. AMARANTHUS TAMARISCINUS Nutt. Shelby County. 33521. A plant of hybrid origin for which distribution information is not available. AEGOPODIUM PODAGRARIA L. Cocke County. 34005. Newfoundland to Michigan, south to Nova Scotia, New England and North Carolina. The specimen was collected at an abandoned homesite, where it may have been planted. HETEROTHECA SUBAXILLARIS (Lam.) Britt. and Rusby. Shelby County. 34409. Florida to Arizona and Mexico, north to Delaware, New Jersey, Illinois, Missouri, and Kansas. KEN E. ROGERS and J. K. UNDERWOOD DEPARTMENT OF BOTANY UNIVERSITY OF TENNESSEE, KNOXVILLE LITERATURE CITED FERNALD, MERRITT LYNDON. 1950. Gray's Manual of Botany. 8th ed. American Book Company, New York. Hitcucock, A. S. 1951. Manual of the Grasses of the United States. 2nd. ed. revised. U. S. Dept. of Agric. Misc. Publ. No. 200. SPERGULA MORISONII IN AMERICA In 1944, Fernald first reported the occurrence of Spergula pentandra L. (Caryophyllaceae) in America, from a col- lection made by O. H. Brown at the Cold Spring Cemetery, Cape May, New Jersey, on April 8, 1942. A description and illustration (based on European specimens at NY) of S. pentandra were included by Maguire (1952) in the New Britton and Brown Illustrated Flora. In the spring of 1965, I collected a Spergula at Lake- hurst, New Jersey. The plants had seeds with conspicuous wings, and were tentatively identified as S. pentandra. They belong, rather, to the closely related S. morisonii Boreau (1847), as determined by Dr. Arthur Cronquist of the New York Botanical Garden. Spergula morisonii is reported here for the first time in America. Spergula morisonii differs from S. pentandra chiefly in the obtuse, slightly overlapping petals, and the brownish, striate wing of the seed. The petals of S. pentandra are acute, not overlapping, and the seed is surrounded by a striking, white wing. Examination of the NY specimens of both species indicates considerable blurring of these distinctions in European material. If, however, the two are considered distinct, the Lakehurst plants are certainly S. morisonii. The Brown specimen (GH) has immature seeds, but another collection, also at GH, from North Caro- lina (H. E. Ahles 52617) shows typical, mature seeds of S. pentandra. Collection data for the Lakehurst plants are as follows: In ballast along railroad tracks at Lakehurst, Ocean County, New Jersey. G. L. Smith 2377. April 30, 1965 (NY). GARY L. SMITH NEW YORK BOTANICAL GARDEN BRONX PARK BRONX, NEW YORK BIBLIOGRAPH Y BoREAU, M. A., 1847. Note sur deux espèces d'Alsinées confondues sous le nom de Spergula pentandra. Rev. Bot. (Duchartre) 2: 421-424. 520 1966] Book Review — Hodgdon 521 FERNALD, M. L., 1944. Spergula pentandra in America. Rhodora 46: 88. MAGUIRE, B., 1952. Caryophyllaceae. In GLEASON, H. A., The New Britton and Brown Illustrated Flora of the Northeastern United States and Adjacent Canada. 2:118-145. The New York Botani- cal Garden, New York. REVISED CHECK-LIST OF THE VASCULAR PLANTS OF MAINE' This new up-to-date amplified and corrected check-list of the vascular flora of Maine is a welcome contribution. It is a pleasure to note that the volume is dedicated to Ralph C. Bean whose scholarly contributions to New England botany and particularly that of Maine are too numerous to summarize here. Indeed the preparation of the revised check-list is primarily the result of Mr. Bean's painstaking study of specimens and recording of records from Maine in the herbarium of the New England Botanical Club. Many of the obvious deficiencies of the earlier check-list’, which serves as an exact model for the present volume, have been corrected; the list is much more complete; introduced species appear now with an asterisk before the specific epi- thet and some of the arboreal species that were listed in the earlier volume such as Ginkgo biloba and Sciadopitys ver- ticillata are thankfully omitted. In the years intervening since the first list, certainly most of the extant herbarium material must have been studied. We can happily be con- fident that this is not a rush job; it is a dependable list which will serve as a secure base for further collections. It is to be noted that the late Dr. Anne Perkins of South Ber- wick, Maine who was particularly dissatisfied with the earlier list because of the many omissions, sent to the pres- 'Ralph C. Bean, Richards, Charles D. and Hyland, Fay. Revised Check-list of the Vascular Plants of Maine. Josselyn Bot. Soc. Me. Bull. No. 8 71 pp. Orono, Me. July 1966. *E. C. Ogden, Steinmetz, F. H. and Hyland, F. Check-list of the: Vascular Plants of Maine. Josselyn Bot. Soc. Me. Bull. No. 8. 1948. 522 Rhodora [Vol. 68 ent reviewer about the year 1949, a "partial list of York Co. plants omitted from check-list". There were twenty-six taxa excluding Carices so specified. Of these all except eight are reported from York Co. in the new list. It is quite pos- sible that specimens of those missing from the present list may be found at Cornell University where Dr. Perkins de- posited her collections. In this reviewer's opinion one aspect of the present vol- ume should provoke some discussion. Professor Fernald reviewed the earlier check-list at some length’. Much of the burden of Fernald's critique was directed to the prob- lem of properly indicating for each species in a flora its correct native or introduced status, In fact he went so far (p. 8) as to suggest some criteria to indicate the degree of establishment of introduced species and relative abundance of native species. Admittedly this would be difficult to do but some such distinctions, if not too finely drawn, would greatly enhance the value of any floristic work. Most floras as well as Gray's Manual itself too often are misleading or else lacking in any sort of information about known rela- tive abundance of the various taxa under consideration. The Maine check-list still hasn't resolved this problem; in fact a certain degree of inconsistency seems to prevail. quoting from the Foreword to the first edition which is reprinted in the revised edition, “Those plants growing on wool-waste, ballast, abandoned-gardens and other habitats where they may not yet be part of our flora but show indi- cations of becoming so are indicated by minus (— ) signs." A large number of both herbaceous and woody taxa were indicated as occurring in the appropriate counties by (—) signs in the earlier list but all or nearly all of the herba- ceous taxa in the more recent list have been changed to (+) signifying that they now are “considered to be a part of our flora", whereas the woody taxa have been left es- sentially as they were in the earlier treatment except for the few that have been excluded. Fernald (loc. cit.) cited “Fernald, M. L. An abbreviated Flora of Maine. RHODORA 51: 5-8, 1949 1966] Book Review — Hodgdon 523 Deschampsia elongata, Chenopodium graveolens and Collo- mia linearis as species whose permanent status in the Maine flora might be questioned and therefore merit the (—) sign; yet in the revised list these still are indicated by the (+) sign as they were earlier. Presumably it has not been possible to demonstrate that these no longer occur in the wild. It is hoped that a future edition would correct a few errors in the woody plants and add some indications of relative abundance. The authors are to be commended for leaving out Ginkgo biloba and Sciadopitys verticillata from the revised edition but what shall we say of Abies concolor, (about which there is no mention in Gray's Manual 8th edi- tion) of Juglans nigra, Magnolia acuminata, Liriodendron tulipifera and Gymnocladus dioica (in Gray's Manual not stated to spread from cultivation in New England beyond their natural ranges) or of Cladrastis lutea (“spreads from cultivation north to Massachusetts") ? In New Hampshire we have no evidence that these produce new individuals in nature and the same probably holds true in Maine. How- ever Mr. Richard Eaton states that Juglans nigra spreads naturally in eastern Massachusetts and he has found it in thin woods (old pastures) in southern Vermont. On the other hand it is quite evident that Berberis vul- garis and Berberis Thun bergii are part of the flora in some areas of Maine. The latter has become decidedly weedy in Bayville (part of Boothbay Harbor, Maine). In fact it has to be pulled up constantly from beneath a heavy growth of Abies balsamea, Picea glauca and P. rubens to prevent it from overrunning the woods and appears abundantly throughout this community at the head of Linekin Bay. On a recent visit to Monhegan Island, plants in varying stages of growth were observed along trails and in the woods at some distance from plantings. The capacity of Syringa vul- garis to persist for generations after the original planting and to spread by vegetative means over extensive areas must certainly cause it to be as much a part of the flora as are certain of the wool-waste species that reproduce only- locally by seeds. 524 Rhodora [Vol. 68 The reviewer may be pardoned for dwelling on the de- ficiencies. It is his ambition to do a similar check-list for New Hampshire in the near future and if he can succeed as well and produce as complete and accurate a list as this one is in most particulars he will be well pleased. ALBION R. HODGDON, UNIVERSITY OF NEW HAMPSHIRE, DURHAM Errata for RHODORA « p. 3 line 15 for altissiumum, read altissimum , P. 256 line 27 for Deyeuzia, read Deyeuxia Volume 68, No. 776 including pages 409-543, was issued December 28, 1966. Corax pp. 120, + 524 v.GT v, og" INDEX TO VOLUME 68 New scientific names and combinations are printed in bold face type Abies and Cedrus, A New Method for Maintaining the Cones of Intact for Study and Storage 516; balsamea 523; concolor 517 PL, 523; magnifica 517 Pl. Acer pensylvanicum 186; spica- tum 186 Achillea borealis 414 fig., 417 Aciachne pulvinata 266 Acorus brachystachys 28 Aegopodium podagraria 519 Aegopogon bryophilus 280; cenchroides 280; Fiebrigii 280; geminiflorus, var. muticus 280 Agaricus coloreus 176; lilacinus 181; luteoolivaceus 176; rubes- centifolius 176 Agenium villosum 353 Agropyron attenuatum 236; breviaristatum 236; dasystach- yum, var. psammophilum 437; psammophilum 437 Agrostis alba 438; araucana 259, 261; boliviana 258, 260; exas- perata 258, 261; gelida 258, 259; gigantea 438; Haenkeana 259, 260; hyemalis 439; var. tenuis, f. setigera 438, 439; michauxii, var. laxiflora 438; montevidensis 258, 259; palus- tris 438; perennans 258, 260; polymorpha. var. palustris 438; scabra 438, 439; var. scabra, f. tuckermanii 438; var. septen- trionalis, f. setigera 438, 439; stolonifera 258, 259; var. com- pacta 438; var. major 438; var. palustris 437; tolucensis 259, 261; verticillata 259; virginica 448 Aira caryophyllea 515; conferta 243 Alnus tenuifolia 186 Alopecurus aequalis 262; brac- teatus 262; Hitchcockii 262 Amaranthus tamariscinus 519 Amphibromus scabrivalvis 115 Andresen, John W. A Multi- variate Analysis of the Pinus chiapensis-monticola- strobus Phylad 1 Andropogon altus 348, 350; bi- cornis 347, 350; cirratus 348, 349; condensatus 348; cordatus 348, 351; gerardii 439; Hassleri 347, 849; hirtiflorus 348, 349; lateralis 348, 351; leptocladus 348, 351; leucostachyus 347, 350; Riedelii 849; saccharoides 348, 350; var. barbinodis 350; ssp. laguroides 350; ssp. parvis- piculus 350; subvar. perfor- atus 350; Selloanus 347, 351; tener 348, 349; subvar. hirti- glumis 348, 349; villosus 353; virgatus 347, 348 Anthochloa lepidula 234; rupes- tris 234 Aphanelytrum procumbens 115 Aquatie Vascular Plants New for Illinois 221 Arabis canescens 513; Drum- mondii 413 fig., 417; fendleri, var. spatifolia 513; hirsuta, var. pyenocarpa 413 fig., 417; holboellii, var. retrofracta 513; pauciflora 513 Arctic-Alpine and Boreal Dis- juncts, Distribution of; Con- tributions of the Flora of Nova Scotia VIII 409 Aristida adscensionis 277; As- plundii 278; capillacea 277; circinalis 277, 279; complanata 527 Cc: = r^ : > Jr 525-4] iy T. P. 528 279; enodis 277, 278; Friesii 279; intermedia 440; inversa 278; implexa 277, 278; laevis 277, 279; longiramea. var. bo- liviana 279; longispica, var. geniculata 440; Mandoniana 277; mendocina 277, 278; necopina 440; Pflanzii 278; riparia 277, 278 Arnica chionopappa 412 fig., 417, 418 . Arundinaria Herzogiana 109 Arundinella Berteroniana 291; confinis 291 Arundo Donax 231 Asplenium callipteris 466; limba- tum 467; marginatum 467; Trichomanes 418 fig., 415, 417; viride 410, 411 fig., 416 Asterella evansii 470, 471 fig., 472 fig., 473 fig.; from Mexico, A New Species of 470; macro- poda 473; venosa 472, 473 Athrostylidium racemiflorum 108 Athyrium filix-femina 127 Avena barbata 244; fatua 244; mollis 449 Axonopus affinis 300; andinus 299, 303; aureus 305; barbi- gerus 299, 302; capillaris 299, 304; chrysites 299, 305; chry- soblepharis 299, 304; compres- sus 299, 301; var. macropodius 299, 301; elegantulus 299, 303; eminens, var. bolivianus 299, 302; exasperatus 299, 304; fissifolius 299, 301; Her- zogii 299, 304; hirsutus 299, 300; iridifolius 299, 303; mar- ginatus 299, 300; pilosus 298, 302; Purpusii 299, 300; sco- parius 299, 303; siecus 299, 302 Ballota alba 94; foetida 93; nigra 93, 94; in the Eastern Rhodora [Vol. 68 United States, Varieties of 93; ssp. foetida 93, 94; var. alba 94; var. foetida 93; ssp. nigra 93; var. ruderalis 93 Banks, D. J., Paspalum minus (Gramineae) in Louisiana and Mississippi 94 Barr, Margaret E. and H. E. Bigelow, Contribution to the Fungus Flora of Northeastern North America, IV 175 Beaman, J. H. and C. E. Seiden- schnur, Cuchumatanea, A New Genus of the Compositae (Heliantheae) 139 Betula borealis 412 fig., 415, 416; glandulosa 412 fig., 415, 416; Michauxii 414 fig., 416, 417; pumila 414 fig., 417 Berberis Thunbergii 523; vulgaris 523 Bigelow, Howard E. and Mar- garet E. Barr, Contribution to the Fungus Flora of North- eastern North America, IV 175 Bolivia, Studies in the Flora of, IV., Gramineae 97, 223, 241 Boreal Disjuncts, Distribution of Arctic-alpine and, Contribu- tions to the Flora of Nova Scotia 409 Bouteloua aristidoides 287; curti- pendula 287; megapotamica 287; simplex 287 Brachiaria plantaginea 298 Brachypodium mexicanum 112 Britton, D. M. and Rolla Tryon, A Study of Variation in the Cytotypes of Dryopteris spin- ulosa 59 Briza Mandoniana 223; var. Herzogiana 223; var, tubercul- ata 223; var. vallegrandensis 223; paleapilifera 223; spici- gera 224; stricta 228, 224; uniolae 224 1966] Bromus angustatus 111; arena- rius 168, 169; arvensis 170, 173; brizaeformis 170, 173; Buchtienii 111; catharticus 110, 111; commutatus 168, 169, 171, 178, 515; f. commutatus 171, 173, 515; f. perglaber 515; f. pubens 171, 173; var. pubens 168, 171, 173; lanatus 111; lati- glumis 440; lepidus 169, 170, 172; f. lasiolepis 170, 172; f. lepidus 170, 172; f. micromollis 172; Mandonianus 243; mollis 168, 169, 171; and Allies in New England 168; f. leiostach- ya 169, 171; f. mollis 169, 171; mutabilis, var. hirtus 172; Pflanzii 111; pitensis 111; pratensis 173; pubescens, f. glaberiflorus 440; purgans, f. glabriflorus 440; racemosus 168, 170, 173; segetum 110, 111; secalinus 169, 170, 172, 515; var. hirsutus 170, 172, 173; var. secalinus 170, 172; f. hirtus 170, 172; f. secalinus 170, 172; f. submuticus 170, 172; Thom- inii 168, 170, 171; f. hirsutus 170, 171; f. Thominii 170, 171; Trinii 110, 111; unioloides 111 Buchloé dactyloides 36 Burch, Derek, The Application of the Linnaean Names of Some New World Species of Euphorbia subgenus Cham- aesyce 155 Burk, John C. and Chung-Hie Nah, Environmental Control of a Taxonomic Character Separating Heterotheca s. str. from Chrysopsis 406 Calamagrostis amoena 246, 249; Antoniana 247, 254; Beyrich- iana 256; boliviensis 249; brevifolia 247, 253; calderil- Index to Volume 68 529 lensis 246, 248; cephalantha 247, 252; chrysantha 246, 250; cryptolopha 246, 251; curta 247, 252; curvula 246, 252; densiflora 247, 257; ele- gans 248; eminens 246, 248; var. sordida 248; var. tunarien- sis 248; Fiebrigii 247, 254; fili- folia 249; glacialis 247, 251; heterophylla 247, 249; Hum- boldtiana 255; intermedia 246, 255; Jamesonii 247, 251; leio- phylla 247, 248; Lilloi 257; longiaristata 246, 256; minima 247, 255; montevidensis 257; var. linearis 256; Orbignyana 247, 255; ovata 246, 250; Pflanzii 250; pentapogonoides 250; polygama 246, 257; recta 247, 256; rigescens 246, 249; rosea, var. macrochaeta 253; spicigera 246, 258; tarijensis 246, 254; tarmensis 247, 253; toluccensis, var. laxiflora 248; trichophylla 247, 256; varie- gata 252, 444; vicunarum 247, 250; violacea 247, 253; viridi- flavescens 246, 257 Calamovilfa longifolia, var. magna 437 Callistosporium luteoolivaceum 176, 178; psilocybe 176 Calopogon pulchellus 460; tuber- osus 460 Candolleodendron brachystachum 429, 430 fig.; A New Genus of the Leguminosae (Cesalpini- oideae) 429 Capparidaceae, Studies in the, VIII Polanisia dodecandra 9) DC 4 Carex abdita 453; acuta 451; atratiformis 413 fig., 417; bebbii 451; capillaris 412 fig., 415, 416; var. major 412 fig., 416; communis, f. gynandra 530 451; var. gynandra 451; hough- toniana 451; houghtonii 451, 452; hystericina 452; hystricina 452; lagopodioides 451; lanu- ginosa 452; lasiocarpa, var. latifolia 452; leptalea, ssp. harperi 452; var. harperi 452; livida, var. grayana 452; var. radicaulis 452; pensylvanica, f. androgyna 451; rugosperma, var. tonsa 453; scirpoidea 411 fig., 416; scoparia 451; tribu- loides, var. bebbii 451; umbel- lata, var. tonsa 453 Cedrus and Abies, A New Method for Maintaining the Cones of Intact for Study and Storage 516; atlantica 517 PI. Cenchrus echinatus 342; myosu- roides 342; pauciflorus 842, 343; viridis 342 Chaetochloa lutescens 446 Chaetosphaeria phaeostroma, var. phaeostromoides 190; phaeostromoides 190 Chamaesyce jenningsii 160; niru- rioides 160 Chascolytrium spicigerum 224 Chenopodium graveolens 523 Chloris Beyrichiana 284, 285; dis- tichophylla 284, 285; halophila 285; polydactyla 285; radiata 285, 286; virgata 285; verticil- lata 518 Chromosome numbers in nonia (Compositae) 49 Chromosome numbers in Zinnia and Sanvitalia, New 215 Chrysopsis berlandieri 406; En- vironmental Control of a Tax- onomie Character Separating Heterotheca s. str. from 406 Chusquea delicatula 109, 110; longipendula 110; peruviana 109, 110; quitensis, var. paten- tissima 110; scandens 110; ser- Ver- Rhodora [Vol. 68 rulata 109, 110; spicata 110; uniflora 109, 110 Cirsium altissimum 3; discolor 3 Cladrastis lutea 523 Cleome aspera 44; dodecandra 41, 42, 43, 44, 45, 46, 47; gynandra 43; icosandra 43; monophylla 43; polygama 46; serrata 46; viscosa 43, 44, 46 Clewell, Andre F., Native North American Species of Les- pedeza (Leguminosae) 359 Clitocybe albimontana 178, 179; asterospora 179 Coix lachryma-jobi 358 Collomia linearis 523 Collybia colorea 176; var. rubes- centifolia 176; luteoolivacea 176; rubescentifolia 176 Convallaria canaliculata 458 Copromanthus herbaceus, 9 ecir- ratus 459 Corallorhiza innata 460; integra 461; ruppii 460, 461; trifida 460, 461 Cornucopiae hyemalis 438 Correll, Donovan S., Some Addi- tions and Corrections to the Flora of Texas III 420 Cortaderia bifida 231, 232; var. grandiflora 232; rudiuscula 231, 232 Cowan, Richard S., Candolleoden- dron, A New Genus of Legumi- nosae (Cesalpinioideae) 429 Cryptogramma Stelleri 413 fig., 417 Cuchumatanea A New Genus of Compositae (Heliantheae) 139; steyermarkii 139, 140 fig., 143, 144 Cuphea carthagenensis 423; peti- olata 423, 424 Cycloloma atriplicifolium 519 Cynodon Dactylon 283 Cypripedium X Andrewsii 53, 1966] 56, 57, 461, 462; Calceolus, var. parviflorum 53, 54, 55, 56, 57; candidum 53, 54, 55, 56, 57, 461; X favilianum 462; X landonii 462; in Manitoba, Nat- ural Hybrids of Lady's Slip- pers 53 Cystopteris fragilis, var. lauren- tiana 413 fig., 417 Dactyloctenium aegyptium 283 Danthonia cirrhata 245 Del Rosario, R. M., A New Spe- cies of Asterella from Mexico 470 Deschampsia caespitosa 243, 244, 440; conferta 243; elong- ata 523 Desmodium ciliare 363 Deyeuxia anthoxanthum 250; capitata 250; gracilis 255; im- berbis 249; Lagurus 252; leio- poda 250; Mandoniana 255; nivalis 250; obtusata 258; phalaroides 250; picta 251; polystachya 248; subsimilis 258; sulcata 256 Diapensia lapponica 412 fig., 416 Dielsiochloa floribunda 243 Digitaria adscendens 294; hori- zontalis 294, 295; leiantha 294, 295; lanuginosa 294, 295; violascens 294, 295 Diplazium callipteris 466; celtidi- folium 466; legalloi 466; limb- atum 467; marginatum 467 Dissanthelium calycinum 241, 242; macusaniense 241; mini- mum 241; peruvianum 241; Trollii 241, 242 Distichlis humilis 230; spicata 230; var. andina 280; y mar- ginata 230 Draba arabisans 413 fig., 417; norvegica 410, 412 fig., 416; pycnosperma 412 fig., 417 Index to Volume 68 531 Dryopteris X Boottii 90; con- similis 468; cristata, var. Clin- toniana X spinulosa, var. intermedia 91; cristata X spin- ulosa 90; X spinulosa, var. intermedia 90, 91; dilatata 127; fragrans, var. remotiuscula 413 fig., 417; Goldiana X margin- alis 90; linnaeana, f. glandulosa 133, 134; marginalis X spinu- losa, var. intermedia 90; mol- licella 468; spinulosa 59, 60, 67, 69. Th 1T Tih Te ik., 70 fg, MI M D. ot; A Study of Variation in the Cytotypes of 59; var. inter- media 59, 60, 61 fig., 63, 64, 65, 66, 67, 68 fig., 69, 70, 71, 72 fig., 78, 74, 75, 76, 77, 78 fig., 79 fig., 80, 81, 82, 83, 84, 85, 86, 87, 88, 89, 90, 91, 92; X var. spinulosa 62 fig., 64, 67, 68 fig. 69, 70, 71, 73 fig., 75, 76, 11, 718 fig, 79 fig., 80, 81, 82, 84, 85, 86, 87, 88, 90, 91, 92; var. spinulosa 59, 60, 63 fig., 64, 65, 66, 67, 68 fig., 69, 70, 71, 78, 74 fig., 75, 76, 77, 78 fig., 79 fig., 80, 81, 82, 83, 84, 85, 86, 87, 88, 89, 90, 91, 92 Duncan, W. H. and S. B. Jones, Jr, Chromosome Numbers in Vernonia (Compositae) 49 Echinochloa crus-galli, var. fru- mentacea 440, 441; crus-pavo- nis 335; frumentacea 441; muricata 440 Elaphoglossum dussii 467; petio- latum, var. dussii 467 Elocharis calva 453; erythropoda 453; palustris 453 Eleusine indica 282, 515 Elymus angulatus 236 Elyonurus adustus 855, 356; 532 tripsacoides 355, 356; var. brevidentatus 356; var. ciliaris 356 Emex spinosa 421 Epilobium Hornemanni 414 fig., 417 Eragrostis articulata 225, 227; var. glabrescens 227; bahien- sis 224, 228; boliviensis 228; Buchtienii 228; cilianensis 224, 226, 441; ciliaris 224, 225; ex- pansa 225, 229; glomerata 225, 226; hypnoides 225; intermedia 518; lugens 225, 229; lurida 225, 228; maypurensis 225, 226; mexicana 225, 228; montufari 225, 228; nigricans 225, 227; pectinacea 515; pilosa 225, 227, 515; polytricha 225, 229; sora- tensis 229; spectabilis, var. sparsihirsuta 516; subatra 227; uniolae 224 Erianthus angustifolius 347; Trinii 347 Erigeron hyssopifolius 414 fig., 417 Eriochloa contracta 518; dis- tachya 297; gracilis 518; punc- tata 297, 298 Eriochrysis cayennensis Warmingiana 346 Erythronium americanum 420 Euphorbia brasiliensis 160; chamaesyce 155, 163, 164; glomerifera 160; hypericifolia 155, 159, 160, 161, 162, 163; hyssopifolia 160, 161, 163; lasiocarpa 160, 161, 163; macu- lata 155, 156, 157, 158, 159, 160, 161, 164, 165; nutans 160, 163; pilulifera 160; preslii 160, 163; prostrata 155, 158, 163, 164; subgenus Chamaesyce, The Application of Linnaean Names of Some New World 346; Rhodora [Vol. 68 Species of 155; thymifolia 155, 158, 164 Ferns, Gymnocarpium, New Data on North American Oak 121; Notes on Lesser Antillean II 464 Festuca arundinacea 441; Buch- tienii 114; dolichophylla 113, 114; elatior 441; Fiebrigii 113; laeteviridis 114; megalura 112, 118; orthophylla 113, 114; var. boliviana 114; Pflanzii 114; pratensis 441; procera 112, 113; prolifera 413 fig., 417; var. lasiolepis 412 fig., 417; riges- cens 113, 114; sublimis 112, 113; ulochaeta 112, 113 Fore, Paul L. and R. H. Mohlen- brock, Two New Naiads From Illinois and Distributional Rec- ords of the Naiadaceae 216 Foster, R. C., Studies in Flora of Bolivia, IV. ineae 97, 223, 241 Frankenia Johnstonii 424 Fungus Flora of Northeastern North America, IV., Contribu- tion to the 175 the Gram- Gentiana X billingtonii 213; X curtisii 213; gracilis 212; lin- earis 211; puberula 209, 210 PL, 211, 212, 213, 214; puber- ulenta 210 PL, 211, 214; A Known But Unnamed Species of the North American Prai- ries 209; X Gentiana alba 213; X Gentiana andrewsii 213; rigida 212; rubricaulis 211, 212; saponaria 211, 212; B linearis 211, 212; shortiana 212; torreyana 212 Ginkgo biloba 521, 523 Glyceria fernaldii 445; pallida, var. fernaldii 445 Gouinia brasiliensis 281; latifolia 1966] 281, 282 Gramineae, Studies in the Flora of Bolivia, IV 97, 223, 241 Grammitis limbata 468; margin- ella 468; phlegmaria 467; var. antillensis 467; stipitata 467 Grear, John W., Jr., Cytogeog- raphy of Orontium aquaticum (Araceae) 25 Gross, A. T. H., G. A. Stevenson and H. H. Marshall, Natural Hybrids of Lady's Slippers (Cypripedium) in Manitoba 53 Gymnocarpium continentale 128 129, 135; dryopteris 121, 124 fig., 125, 126, 127 fig. 128, 131, 132, 133, 134, 135, 136, 137; var. glandulosa 134; var. disjunctum 122 Pl, 128, 129, 130 Pl, 135; X var. dryopteris 131; dryopteris, var. pumilum 134; var. dryopteris 128, 129, 136; X robertianum 133; heterosporum 123 Pl., 124 fig., 127 fig., 130 fig., 131, 132, 134, 135, 136, 137; New Data on North American Oak Ferns 121; robertianum 121, 125, 126, HT fig, LS, 19], 133, 10A, 135, 136 Gymnocladus dioica 523 Gymnogramma gracilis 468 Gymnopogon jubiflorus 284; spicatus 284 Gynerium sagittatum 232 Hackelochloa granularis 357 Hedysarum conglomeratus 371; divergens 390; frutescens 370, 881, 390, 392; hirtum 376, 377; prostratum 385; repens 385, 386; reticulatum 393; sessili- florum 393; umbellatum 370; violaceum 880, 390, 392 Hehre, Edward J., Polanisia dodecandra in New Hampshire Index to Volume 68 533 48 Helianthemum tripetalum 208 Hemerocallis flava 456; lilio-as- phodelus 456, 457 Hemidictyum marginatum 467 Herbarium, Botanical Notes from the Pringle, I 514 Heteropogon contortus 353, 354; melanocarpus 353, 354; vil- losus 353 Heterotheca latifolia 406; var. macgregoris 406; s. str., En- vironmental Control of a Tax- onomic Character Separating Chrysopsis from 406; subaxil- laris 406, 407, 519; var. petio- laris 406 Hodgdon, A. R., Revised Check- list of the Vascular Plants of Maine (Review) 521; and Frederic Steele, Rubus Sub- genus Eubatus in New Eng- land 474 Holcus lanatus 441 Homolepis aturensis 334 Hordeum halophilum 237; muti- cum 237; nodosum 237 Hounsell, R. W. and E. C. Smith, Contributions to the Flora of Nova Scotia VIII. Distribution of Arctic-alpine and Boreal Disjuncts 409 Hyacinthus muscari 457, racemosus 457, 458 Hygrophorus lilacinus 179, 181; monticola 180 Pl. 181, 182; nitiosus 182; tahquamenonen- sis 182; violeipes 181 Hymenachne donacifolia 333 Hymenophyllum gratum 464, 465; hirsutum 465, var. gratum 465; hirtellum 464, 465; var. gratum 465, 466; var. vincentinum 465; maxonii 465 244; sorghum 458; | Hyparrhenia bracteata 351 534 Ichnanthes breviscrobs 329, 330; calvescens 329, 330; candicans 330, 331; minarum 380, 331; pallens 330, 381; peruviana 329, 330; Ruprechtii 330, 331 Illinois, Aquatic Vascular Plants New for 221 Iltis, Hugh H., Studies in the Capparidaceae VIII. Polanisia dodecandra (L.) DC. 41 Imperata brasiliensis 345; min- utiflora 345; tenuis 345 International Plant Index, Au- thors of Plant Genera and the (Review) 35 Isachne arundinacea 334 Jaegeria discoidea 145; gracilis 145; hirta 140 fig., 148, 144, 145 Jones, S. B., Jr. and W. H. Dun- can, Chromosome Numbers in Vernonia (Compositae) 49 Juglans nigra 523 Juncus alpinus, var. rariflorus 414 fig., 417; campestris 8 455; congesta 454; erectus 454; glabratus 454; intermedius 454, 455, 456; multiflorus 454, 455, 456; trifidus 411 fig., 416; vali- dus 519 Juniperus communis, var. com- munis 514 Jussiaea diffusa 218 Keissleriella caudata 185, 187 fig., cladophila 185 Koeleria cristata 242, 441; gra- cilis, ssp boliviensis 242; mac- rantha 441; pseudocristata, var. andicola 242 Lactuca muralis 138; in New England 138 Lamprothyrsus Hieronymi 245; var. tinctus 245 Rhodora [Vol. 68 Lasiacis ligulata 332; sorghoidea 332 Lasiosphaeria chrysentera 185; ovina 185 Lechea brevifolia 205; cernua 203; cinerea 194; (Cistaceae), Notes on Rafinesque's Species of 192; corymbosa 200; divari- cata 203; deckertii 203; flori- dana 203, 204; furfuracea 195; glomerata 200; heterophylla 199, 200; var. major 200; var. minor 200; var. parviflora 200; intermedia 193, 195, 198; juncifolia 196, 197; laxiflora 195, 196, 197; var. brevifolia 196; leggettii 193, 194, 195, 197, 198, 201, 203, 207; linifolia 208; major 197; maritima 193, 198, 205; mexicana 208; minor 194, 198, 199, 200, 201, 202, 203, 204, 205, 206, 207; mucro- nata 197, 198, 200, 201, 202, 203, 204; var. ramosa 199; var. sessiliflora 199; var. sim- plex 199; var. typica 199; paucifolia 208; pauciflora 205; patula 194, 203, 204, 207; pulchella 193, 194, 196, 197, 203; racemulosa 193, 195, 197, 198, 201, 202, 203, 204; re- curvata 197; revoluta 202; secundiflora 204; sessiliflora 207, 208; stellata 206, 207; stricta 195; surculosa 201, 202; tenuifolia 196, 197, 198, 199, 200, 202, 203, 204, 205; terni- folia 194, 195; torreyi 194, 203, 207; thymifolia 194, 205, 206; uniflora 206; verna 196, 197; villosa 197, 198, 199, 200, 202, 203, 204, 207; var. macrotheca 199; var. schaffneri 199; vir- gata 202, 203; var. bracteata 203; var. breviflora 203 Ledum groenlandicum 189 1966] Index to Volume 68 535 Leiosphaerella falcata 186, 187 fig., phoenicis 186; praeclara 186 Leptochloa domingensis 280, 281; dubia 280, 281; uninervia 280, 519; virgata 280, 281 Leptocoryphium lanatum 292 Leptoloma cognatum 516 Lespedeza acuticarpa 403; an- gustifolia 363, 367, 369, 3870, 372, 375 fig., 383, 393, 394, 395, 399; X capitata 396, 400 fig.; X hirta 364, 377, 396, 398, 399, 401 fig.; X repens 396, 400 fig.; X stuevei 396, 400 fig.; xX virginica 396; var. brevi- folia 368, 369; f. subvelutina 368, 369; bicknellii 371; bi- color 359; brittonii 402; capi- tata 362, 363, 367, 370, 372, 378, 374, 375 fig., 379, 382, 393, 394, 398, 399; X hirta 396, 398, 399, 401 fig.; X inter- media 396, 397, 400 fig.; X leptostachya 396, 401 fig.; X repens 396, 401 fig.; X stuevei 396, 400 fig.; violacea 396, 399, 400 fig.; virginica 396, 397, 399, 401 fig.; var. angustifolia 368; var. calycina 376, 399; var. capitata 373; var. hirti- formis 398, 399; var. longifolia 399; var. sericea 371, 373; var. stenophylla 371, 373, 374; f. argentea 371, 374; var. typica 371, 373; var. velutina 371, 373; var. vulgaris 371, 374; cuneata 359; divergens 390, 393; frutescens 371, 380, 381, 392, 393; var. acutifructa 380, 381; hirta 363, 367, 368, 372, 374, 377, 378, 379, 399; X in- termedia 396, 397, 399, 400 fig., 402; X procumbens 396, 401 fig.; X repens 396, 403; X stuevei 396, 397, 401 fig.; X violacea 377, 396, 400 fig.; X virginica 896, 397, 403; ssp. curtissii 375 fig., 876, 377, 378, 379, 398; ssp hirta 375 fig., 376, 377, 378; var. angusti- folia 368; var. appressipilis 377, 398; var. calycina 376, 379; var. dissimulans 398, 399; var. intercursa 368, 370; var. longifolia 398; var. oblongi- folia 398; var. sparsiflora 376, 377; var. typica 376; inter- media 361 fig., 363, 368, 375 fig., 380, 381, 388, 392, 393, 397, 402; X procumbens 396, 401 fig.; X repens 396, 400 fig.; X stuevei 396, 403 fig.; X violacea 396, 400 fig.; X vir- ginica 396, 400 fig.; f. hahnii 402; var hahnii 401; (Legumi- nosae), Native North Amer- ican Species of 359; lepto- stachya 367, 370, 375 fig., 382, 383, 395; longifolia 398, 399; manniana 399; neglecta 402, 403; X nuttallii 397; nuttallii 399; var. manniana 399; var. nuttallii 402; oblongifolia 398; polystachya 376; prairea 390, 393; procumbens 363, 364, 367, 383, 384 fig., 385, 390, 392; X repens 396, 400 fig.; X stuevei 364, 396, 403 fig.; X violacea 396, 400 fig.; X virginica 396, 401 fig., 402; var. elliptica 402; prostrata 385; reticulata 380, 393; var. angustifolia 393; var. sessiliflora 380; repens 361 fig., 367, 384 fig., 385, 386, 387, 390, 392, 393; X stuevei 396, 401 fig.; X violacea 396, 401 fig.; virginica 396, 403 fig.; sessiliflora 380, 393; sim- ulata 397, 399; stuevei 363, 368, 371, 384 fig., 387, 388, 395, 402; X violacea 396, 401 fig.; 536 X virginica 396, 402, 403 fig.; f. angustifolia 402, 403; var. angustifolia 402, 403; var. in- termedia 380; var. neglecta 402; stipulacea 359; striata 359; texana 367, 389, 390, 391 fig, 395; velutina 371, 373; virgata 401; virginica 363, 368, 381, 388, 391 fig. 893, 394, 395; f. deamii 402, 403; var. deamii 395; var. sessiliflora 380; var. typica 394; violacea 361 fig, 362, 363, 368, 381, 387, 390, 391 fig., 392, 393; X virginica 396, 400 fig., 403; var, angustifolia 393, 394; var. divergens 890; var. prairea 390; var, reticulata 393; var. sessiliflora 380 Limonium limbatum, var. gla- brescens 425; var. limbatum 425 Linnaea borealis, var. americana 187, 188 Linnaean Names of Some New World Species of Euphorbia subgenus Chamaesyce, The Application of 155 Liriodendron tulipifera 523 Lolium multiflorum 238; perenne 238; temulentum 238; var. arvense 238 Long, Robert W., Observations Regarding the Occurrence and Relationships of Ruellia Lor- entziana (Acanthaceae) in Southern Florida 432 Louisiana and Mississippi, Pas- palum minus in 94 Luziola peruviana 290 Luzula congesta 456; intermedia 454, 456; multiflora 454, 455, 456; parviflora 414 fig., 417; var. melanocarpa 414 fig., 417; spicata 412 fig., 416, 418 Lycopodium reflexum, var. re- Rhodora [Vol. 68 flexum 464; var. rigidum 464; rigidum 464; Selago 411 fig., 415, 416 Lycurus phleoides 263 Lysichitum americanum 28; camtschatcense 27 MacDonald, R. D. and M. E. Mac- Donald, A New Method for Maintaining the Cones of Abies and Cedrus Intact for Study and Storage 516 Magnolia acuminata 523 Maine, Revised Check-list of the Vascular Plants of (Review) 521 Manisuris altissima 356; aurita 356, 357; fasciculata 856 Manitoba, Natural Hybrids of Lady's Slippers (Cypripedium) in 53 Marshall, H. H., A. T. H. Gross and G, A. Stevenson, Natural Hybrids of Lady's Slippers (Cypripedium) in Manitoba 53 Medicago virginica 393 Melica adhaerens 233; var. tenuis 233; Mandonii 233; scabra 233; var. glabra 233; triflora 450; violacea 233 Mesosetum cayennense 297 Mexico, A New Species of As- terella from 470 Microchloa indica 283 Mississippi, Paspalum minus in Louisiana and 94 Mohlenbrock, R. H. and Paul L. Fore, Two New Naiads from Illinois and Distributional Records of the Naiadaceae 216 Monocots, Nomenclatural Notes on 435 Morton, C. V., Varieties of Bal- lota nigra in the Eastern United States 93 Muhlenbergia ^ angustata 263, 1966] 265; asperifolia 264, 265; fastigiata 264, 265; Herzo- giana 264; Holwayorum 264, 266; ligularis 264, 265; peru- viana 263, 264; phragmitoides 266; quitensis 264; rigida 264, 266; sobolifera, var. setigera 518; tenuissima 263, 264 Munroa andina 288; var. brevi- seta 288; argentina 288; de- cumbens 288 Muscari ambrosiaceum 458; at- lantieum 457, 458; moschatum 457, 458; muscari 458; mus- carimi 458; racemosum 457, 458 Mycena atkinsoniana 182 Mycosphaerella crepidophora 188; ebulina 188; linnaeae 187, 188, 189; minor 188 Myriophyllum heterophyllum 425; verticillatum 425 Nah, Chung-Hie and John C. Burk, Environmental Control of a Taxonomic Character Separating Heterotheca s. str. from Chrysopsis 406 Naiads from Illinois and Dis- tributional Records of the Naiadaceae, Two New 216 Naias flexilis 217, 218, 219; gracillima 217, 218, 219; guadalupensis 217, 218, 219; minor 217, 218, 219, 220 Najas marina 221; minor 222 Nassella Asplundii 269; corni- culata 270; deltoidea 269; flac- cidula 269; var. humilior 269; Meyeniana 269, 270; pubiflora 269 Naucoria kauffmanii 183 Neottia gracilis 462; lacera 462 New England, Lactuca muralis in 138; Rubus Subgenus Euba- tus in 474 New Hampshire, Polanisia dode- Index to Volume 68 537 candra in 48 Nova Scotia, Contributions to the Flora of VIII. Distribution of Arctic-alpine and Boreal Dis- juncts 409 Olyra Buchtienii 343, 344; ciliati- folia 348, 344; ecaudata 343, 344; heliconia 343, 344; lati- folia 343, 344; lateralis 343; micrantha 343, 344 Omphalia luteolilacina 181 Omphalina oreades 178, 179 Ophrys corallorhiza 460 Oplismenus hirtellus 334 Orchis rotundifolia, f. beckettae 462; f. beckettii 462; f. lineata 462; var. lineata 462 Orontium aquaticum 25; (Ara- ceae), Cytogeography of 25; fossile 28 Orthoclada laxa 234 Oryzopsis florulenta 276 Oxyria digyna 410, 412 fig., 416 Oxytropis johannensis 414 fig., 417 Panicum agrostidiforme 443; ag- rostoides 443; anceps 443; aquaticum 318, 320; var. chlor- oticum 320; ashei 441, 442; bo- liviense 318, 323; capillare, var. minus 442; cayennense 319, 320; chloroticum 320; commu- tatum, var. ashei 441, 442; var. minor 442; cordovense 318, 327; cyanescens 319, 325; depauperatum, var. involutum 443; var. psilophyllum 443; echinulatum 318, 320; var. bo- liviense 320; frondescens 318, 824; frumentaceum 441, 446; giganteum 322; glutinosum 319, 325; hebotes 319, 327; hirticaule 319, 320; involutum 443; laxum 318, 323, 824; 538 lutescens 445, 446; maximum 319, 322; megiston 318, 328; muricatum 440; nervosum 319, 325; olyroides 319, 329; pan- trichum 318, 327; parvifolium 319, 326; paucispicatum 318, 328; peladoénse 319, 321; pilo- sum 318, 3823; polygonatum 318, 322; procurrens 318, 329; pulehellum 318, 324; quadrig- lume 317, 319, 321; rigidulum 443; Rudgei 319, 326; sciurotis 319, 327; Sellowii 319, 326; sphaerocarpon, var. inflatum 519; stoloniferum 318, 324; stramineum 319, 321; trichan- thum 319, 325; tricholaenoides 318, 322; versicolor 320, 328; virescens 446 Pappophorum mucronulatum 235; Wrightii 235 Pariana bicolor 239, 240; gracilis 239; lunata 240; Swallenii 239; zingiberina 240 Parnassia parviflora 414 fig., 417 Paronychia Jamesii, var. Jamesii 423; var. praelongifolia 423 Paspalum boliviense 306, 317; Buchtienii 307, 310; candidum 305, 308; Ceresia 306, 309; ciliatifolium 444; var. muhlen- bergii 444; collinum 308, 315; conjugatum 307, 313; consper- sum 308, 315; decumbens 306, 311; densum 306, 316; depaup- eratum 306, 309; dilatatum 307, 313; distichum 307, 311; Ek- manianum 307, 316; erianthum 307, 310; Humboldtianum 307, 310; inaequivalve 306, 313; in- constans 307, 311; Juergensii 307, 312; lepidum 307, 312; line- ispatha 306, 309; malacophyl- lum 306, 316; melanospermum 307, 314; membranaceum 309; millegrana 306, 315; minus 94, Rhodora [Vol. 68 95, 96, 306, 314; (Gramineae) in Louisiana and Mississippi 94; muhlenbergii 443; multi- caule 307, 312; notatum 95, 306, 314; paniculatum 307, 312; penicillatum 306, 308; pictum 306, 312; pilosum 306, 316; plicatulum 308, 314; poly- phyllum 307, 310; prostratum 306, 308; pubescens 443, 444; pygmaeum 306, 308; var. glab- rescens 308; remotum 307, 316; saccharoides 306, 317; stella- tum 306, 309; tripinnatum 303; Urvillei 307, 314; virgatum 306, 315 Paxillus curtisii 183 Pennisetum chilense 340; lati- folium 339, 341; mutilatum 340, 3841; sagittatum 341; setosum 339, 340; tristachyum 340, 341; ssp. boliviense 341; villosum 339, 340 Phaeocollybia kauffmanii 183, 184 Fhalaris angusta 289; arundi- nacea 444; f. picta 444, 445; f. variegata 444, 445; canarien- sis 289 Pharus glaber 290; latifolius 290; parvifolius 290, 291 Phleum alpinum 413 fig. 417; pratense, var. nodosum 516 Phragmites communis 231 Phyllodoce caerulea 412 fig., 416 Picea glauca 523; rubens 523 Pinguicula vulgaris 412 fig., 416 418 Pinus ayacahuite 16; chiapensis 1, 2, 4, 5, 8 Pl, 10 PL, 11, 14, 15, 16, 20, 21, 23; -monticola- strobus Phylad, A Multivariate Analysis of the 1; contorta 3; flexilis 13; griffithii 16; monti- cola 4, 8 PL, 10 Pl., 12, 14, 15, 16, 20, 21, 23; X strobus 23; 1966] oocarpa 14; parviflora 16; peuce 16; strobiformis 13; strobus 1, 2, 4, 5, 7, 8 PL, 10 Pioi 12 aA T5, 18 25 4b 23; var. chiapensis 1, 2, 4, 5, 14 Piptochaetium panicoides 2/05 E subpapillosum 270; setifolium 270; tuberculatum 270 Plagiophiale petrakii 189 Plagiostigma petrakii 189 Plant Genera and the Interna- tional Plant Index (review) 35 Plantago insularis, var. fastigiata 428; minima 428; scariosa 428 Plectosphaeria salicis 189 Pleurotus elongatipes 184, 185 Poa alpina 411 fig., 416, 418; androgyna 119; annua 116, 119; asperiflora 116, 118; bo- liviensis 118; Buchtienii 116; var. subacuminata 116; can- damoana 116, 120; denticulata 116, 119; dumetorum 119; var. unduavensis 119, 120; glaucan- tha 411 fig, 416; gym- nantha 116, 117; var. aperta 117; Hieronymi 116, 118; hor- ridula 116, 119; humillima 116, 117; Lilloi 116, 118; myriantha 118; palustris 518; perligulata 117; Pflanzii 118; pratensis 116, 118; scaberula 116, 117 Podosphaera clandestina, var. clandestina 189; oxyacanthae, var. oxyacanthae 190 Polanisia dodecandra 41, 42, 46, 47, 48; in New Hampshire 48; (L.) DC., Studies in the Cap- paridaceae VIII 41; ssp. trachysperma 47; ssp. dode- candra 47; graveolens 41, 42, 45, 46; trachysperma 47; uni- glandulosa 47 Polygonatum canaliculatum 458; commutatum 458 Polypodium flabelliforme — 467; Index to Volume 68 539 phlegmaria 467; rigescens 467 Polygonum viviparum 189 Polypogon elongatus 262; inter- ruptus 262, 263 Polystichum Lonchitis 413 fig., 417 Poole, James P., Lactuca muralis in New England 138 Potamogeton americanus 218 Primula mistassinica 414 fig., 417 Pringle, James S., Gentiana puberulenta, sp. nov., a Known but Unnamed Species of the North American Prairies 209 Proboscidea sabulosa 426, 427 PI. Proctor, George R., Notes on Lesser Antillean Ferns II 464 Pseudechinolaena polystachya 296 Psoralea melitoides 392; psora- lioides 392 Puccinellia fernaldii 445; gena 115; parvula 115 oresi- Rafinesque's Species of Lechea (Cistaceae), Notes on 192 Rhoades, Mary H., Seed Ger- mination of Shortia galacifolia T & G Under Controlled Con- ditions 147 Rhododendron canadense 36 Rhynchelytrum repens 296 Rhynchospora capitellata 453; glomerata, var. capitellata 453; var. minor 453 Rogers, Ken E. and J. K. Under- wood, Some Plants New to Tennessee 518 Rollins, Reed C., Authors of Plant Genera and the Interna- tional Plant Index (Review) 35; The Identity of Sisymbrium pauciflorum (Cruciferae) 513 Rubus abactus 484, 510; abbre- vians 482, 510; aculiferus 483, 510; adjacens 510; alius 510; 540 allegheniensis 475. 484, 486, 487, 494, 508, 504, 507, 508, 510, 511; X pensilvanicus 510, 511; alter 510; alumnus 510; ami- calis 510; amnicola 510; An- drewsianus 510; apparatus 510; aptatus 510; arcuans 510; arenicola 478, 481, 486, 488, 493, 510. 511; argutus 483, 487, 499, 505, 510, 511; arundelanus 510; ascendens 510; auroralis 510; avipes 510; Baileyanus 479, 480, 510; barbarus 510; bellobatus 484, 510; Bicknellii 481, 510; biformispinus 510; bi- frons 487; Bigelovianus 510; Blakei 510; Blanchardianus 510; Bootianus 484, 510; Brain- erdii 510; brevipedalis 510; canadensis 477, 484, 486, 500, 502, 508, 511; coloniatus 510; conanicutensis 510; cubitans 510; cuneifolius 477, 487, 499; curtipes 510; Deaneanus 510; eflagellaris 510; electus 510; elegantulus 478, 481, 482, 483, 486, 487, 497, 500, 502, 503, 510, 511; Enslenii 486, 488, 490, 510, 511; facetus 510; felix 510; Fernaldianus 510; flagel- laris 480, 482, 484, 486, 487, 488. 489, 490, 491, 492, 510, 511; frondisentis 510; fron- dosus 480, 481, 486, 487, 505, 506, 507, 508, 510, 511; frutico- sus 475; geophilus 510; glandi- caulis 510; gnarus 510; Gravesii 510; Groutianus 497, 510; harmonius 510; hetero- geneous 510; heterophyllus 510; hispidoides 510; hispidus 475, 488, 486, 493, 494, 510, 511; X allegheniensis 510, 511; X flagellaris 511; X setosus 510, 511, 512; X vermontanus 512; honorus 510; insons 510; in- Rhodora [Vol. 68 sulus 510; invisus 510; jacens 478, 510; jactus 510; Janssonii 510; Jaysmithii 482, 486, 488, 489, 511; Jeckylanus 510; jugosus 510; junceus 510; junior 510; Kennedyanus 510; laciniatus 487; ]laevior 510; latens 511; Lawrencii 511; licitus 511; longissimus 511; mainensis 511; maniseesinsis 511; Millspaughii 511; mis- cix 511; montpelierensis 511; multiformis 511; multilicius 511; multispinus 507, 511; nigricans 511; nigrobaccus 511; notatus 511; novangli- cus 511; obsessus 511; orarius 484, 511; ortivus 511; ostryi- folius 484, 511; paludiyagus 488. 511; Parlinii 511; paulus 511; pauper 511; peculiaris 511; pensilvanicus 481, 484, 487, 501, 508, 505, 507, 508, 510, 511; pergratus 481, 511; per- invisus 511; permixtus 511; perpauper 511; pervarius 511; philadelphicus 511; plicatifolius 479, 481, 492, 511; positivus 511; procumbens 511; prosper 511; pudens 511; pugnax 511; Randii 511; ravus 511; recur- vans 481, 511; recurvicaulis 478, 479, 481, 484, 486, 488, 489, 491, 492, 498, 510, 511, 512; X setosus 510; X ver- montanus 511; rhodinsulanus 511; rixosus 511; roribaccus 511; Rosa 511; Rossbergianus 511; saltuensis 511; Sanfordii 511; sativus 511; scambens 511; sceleratus 480, 511; semierectus 511; semisetosus 478, 483, 486, 497, 498, 499, 510, 511; sem- pervirens 511; setosus 475, 483, 484, 486, 494, 495, 497, 499, 502, 504, 510, 511, 512; X al- 1966] legheniensis 510, 511; X ver- montanus 510, 511, 512; severus 480, 497, 511; signifi- cans 511; singulans 511; spicu- losus 511; Subgenus Eubatus in New England 474; tardatus 478. 511; tetricus 511; tholi- formis 511; trifrons 512; udus 512; usus 512; univocus 512; vermontanus 478, 481, 484, 486, 494, 496, 497, 498, 499, 500, 502, 510, 511; X allegheniensis 511; X elegantulus 511; X pensil- vanicus 511; vigoratus 512; viridifrons 512 Ruellia caroliniensis 432; inter- media 434; lorentziana 438 P1.. 434; (Acanthaceae) in South- ern Florida, Observations Re- garding the Occurrence and Relationships of 432; nudiflora 434; var. occidentalis 432; oc- cidentalis 432; succulenta 432; tuberosa 434 Ruppia maritima 221 Saccharum officinarum 345 Sacciolepis myuros 333 Sanvitalia abertii 215; fruticosa 216; New Chromosome Num- bers in Zinnia and 215; tenuis 215, 216; versicolor 216 Saxifraga aizoides 410, 412 fig., 416; Aizoon, var. neogaea 409, 412 fig., 416 Schizachne purpurascens 413 fig., 417 Sciadopitys verticillata 521, 523 Scirpus caespitosus, var. deli- catulus 413 fig., 417 Scrophularia marilandica 425 Sedum Rosea 411 fig., 415, 416 Seidenschnur, C. E. and J. H. Beaman, Cuchumatanea, A New Genus of the Compositae (Heliantheae) 139 Index to Volume 68 541 S:lloa plantaginea 144 Senecio pauperculus 414 fig.. 417 Sesuvium erectum 421, 422; mari- timum 422; Portulacastrum 422, 423; sessile 422; trianthe- moides 422; verrucosum 421 Setaria argentina 336, 338; bar- binodis 336, 339; geniculata 336; glauca 445, 446; gracilis 336; f. brevispica 336; f. peni- cillata 336; Liebmannii, f. trichorachis 839; muricata 440; oblongata 336, 338; Poiretiana 336; scandens 336, 337; tena- cissima 336, 337; tenax 336, 337; trichorachis 336, 339; vul- piseta 336, 338 Seymour, Frank C., Botanical Notes from the Pringle Her- barium, I. 514; Bromus mol- lis and Allies in New England 168 Shortia galacifolia 147, 151, 152, 154; T & G Under Con- trolled Conditions, Seed Ger- mination in 147 Siegesbeckia repens 144 Sisymbrium pauciflorum 513; (Cruciferae), the Identity of 513 Sisyrinchium angustifolium 459; arenicola 459; atlanticum 459; farwellii 459 Smilax ecirrata 459 Smith, C. E., Jr., Identity of Witchweed in the Southeastern United States 167 Smith, E. C. and R. W. Hounsell, Contributions to the Flora of Nova Scotia VIII. Distribution of Arctic-alpine and Boreal Disjuncts 409 Smith, Gary L., Spergula moris- onii in America 520 Solidago multiradiata 412 fig., 416 542 Sorghastrum minarum 352; par- viflorum 352, 353; stipoides 352, 353 Sorghum bicolor 446; halepense 352; vulgare 352, 446 Sparganium chlorocarpum, f. acaule 436; simplex, var. acaule 436 Spartina michauxiana, var. sut- tiei 446; pectinata, var. suttiei 446 Spergula morisonii 520; in Amer- ica 520; pentandra 520 Sphaerella eucarpa 189; tonii 188 Sphaeria phaeostromoides 190 Spiranthes gracilis 462, 463; lacera 462, 463 Sporobolus aeneus 267, 268; ar- gutus 267; Berteroanus 268; cryptandrus 518; eximius 269; indicus 267, 268; neglectus 448; Poiretii 267, 268; tenuissimus 267; vaginiflorus 448; var. neg- lectus 447; virginicus 448 Steele, Frederic, and A. R. Hodg- don, Rubus Subgenus Eubatus in New England 474 Stipa boliviensis 275; brachy- phylla 271, 275; capilliseta 271, 273; curviseta 271, 275; dasy- carpa 271, 275; depauperata 271, 274; florulenta 271, 276; Hans-meyeri 271, 273; Hol- wayi 271, 276; Ichu, 271, 272; illimanica 272, 276; incon- spicua 271, 274; mucronata 271, 273, 274; nardoides 271, 273; Neesiana 271, 274; obtusa 271, 275; pampagrandensis 271, 272; Pflanzii 269; plumosa 271, 272, 276; polyclada 271, leigh- 275, 276; speciosa 271, 272; tiraquensis 269; Trollii 272, 276 Stevenson, G. A., H. H. Marshall, Rhodora [Vol. 68 and A. T. H. Gross, Natural Hybrids of Lady’s Slippers (Cypripedium) in Manitoba 53 Striga asiatica 167; lutea 167 Stylagrostis leiopoda 250 Swartzia brachystachya 429 Symplocarpus foetidus 27 Syringa vulgaris 523 Targionia hypophylla 471, 472 Tennessee, Some Plants New to 518 Texas, Some Additions and Cor- rections to the Flora of, III 420 Thaxteria fusca 190 Thelypteris gracilis 468; con- similis 468; cooleyi 468, 469; limbata 469; noveboracensis 125; palustris 124; piedrensis 468 Thrasya campylostachya 295 Tofieldia glutinosa 414 fig., 417 Torres, Andrew M., New Chro- mosome Numbers in Zinnia and Sanvitalia 215 Torreyochloa fernaldii 445; pal- lida, ssp. pallida, var. fernaldii 445 Trachypogon canescens 355; montufari 354, 355; plumosus 355; polymorphus, var. bolivi- anus 355 Tragus racemosus 279 Trichachne affinis 293; cali- fornica 293; insularis 293; sac- charata 294; sacchariflora 293 Trichloris mendocina 286; pluri- flora 286 Trichodium laxiflorum 438 Tricholaena rosea 296 Tricholoma rubescentifolium 176 Trichopteryx flammida 291 Triglochin maritimum 436; pal- ustre 436 Triniochloa stipoides 266 3 1753 00341 3900 1966] Triodia avenacea 235 Triphora trianthophora, var. Schaffneri 420; var. triantho- phora 420 Triplasis purpurea 518 Tripogon spicatus 282 Tripsacum australe 357; dacty- loides 357 Trisetum floribundum 243; oreo- philum 243; spicatum 243, 449; var. maidenii 449, 450; var. molle 411 fig., 416, 449; var. pilosiglume 411 fig., 416, 449; ssp molle 450; subspicatum, f. maidenii 450; triflorum. ssp. molle 450 Tristachya chrysothrix 292 Tryon, Rolla and Donald M. Britton, A Study of Variation in the Cytotypes of Dryopteris spinulosa 59 Underwood, J. K. and Ken E. Rogers, Some Plants New to Tennessee 518 Utricularia cornuta 428; juncea 428 Vaccinium angustifolium 189; boreale 414 fig., 417; caespito- sum 414 fig., 417; ovalifolium 414 fig., 417; uliginosum 189; var. alpinum 412 fig.. 415, 416 Vernonia acaulis 50; altissima 51; angustifolia 51; var. an- gustifolia 51; var. mohrii 51; var. scaberrima 5l; anthel- mintica 51; baldwinii 49, 51; blodgettii 51; cinerea 49, 51; (Compositae), Chromosome Index to Volume 68 543 Numbers in 49; colorata 51; conferta 51; X concinna 51; xXdissimilis 51; glabra 51; greggii 51; guadaloupensis 51; guineensis 51; interior 51; karvinskiana 51; kotschiana 51; lindheimeri 51; marginata 51; noveboracensis 52; nudi- flora 52; ovalifolia 52; perot- tetii 52; platensis 52; primu- lina 52; pulchella 52; X recurva 52; texana 52 Vilfa vaginiflora 447, 448 Voss, Edw. G., Nomenclatural Notes on Monocots 435 Wagner, W. H., Jr., New Data on North American Oak Ferns, Gymnocarpium 121 Wilbur, Robert L., Notes on Rafinesque's Species of Lechea (Cistaceae) 192 Winterringer, Glen S., Aquatic Vascular Plants New for Il- linois 221 Witchweed in the Southeastern United States, Identity of 167 Woodsia alpina 411 fig., 416; glabella 411 fig., 416; ilvensis 411 fig., 415, 416 Zannichellia palustris 222 Zea mays 167 Zeugites mexicana 234; var. glandulosa 234 Zinnia maritima 215; purpusii 215; and Sanvitalia, New Chromosome Numbers in 215