Classification of the Outlying Species of New World Panicum (Poaceae: Paniceae)

Home , Echinolaena, Homolepis, Ibirataia, Ichnanthus, Leptocoryphium, Mesosetum

SMITHSONIAN COKTRIBUTIOSS TO BOTANY NUMBER 59

Fernando 0. Zuloaga and Thomas R. Soderstrom

SMITHSONIAN INSTITUTION PRESS City of Washington 1985 ABSTRACT Zuloaga, Fernando O., and Thomas R. Soderstrom. Classification of the Outlying Species of New World Panicum (Poaceae: Paniceae). Smithsonian Contributions to Botany, number 59, 63 pages, 25 figures, 2 tables, 1985.- In the present study a number of species originally assigned to Panicum but not fitting tvell ivithin its boundaries have been examined. Some of the characters that have been used in the past to distinguish genera in the tribe have been re-examined, including type of hilum, spikelet compression, relative length of the glumes, consistency of the upper anthecium and whether crested or not, and carriage of the spikelet relative to the axis. Further characters of taxonomic importance have been revealed by studies made lvith the scanning electron niicroscope (SEM) at higher magnifications than previously possible. The results of these studies indicate that several species should be removed from Panicum and transferred to other genera. Accord- ingly the following changes have been made: P. megastachyum Nees ex Trinius and P. albicomum Sr%.allen& Garcia Barriga have been transferred to Bra- chiaria, P. glutinosum Swartz and P. villaricense Mez to Homolepis, P. grandifolium Doell to Ichnanthus, P. killipii Hitchcock to Paspalurn, and P. macranthum Trinius to Streptostachjs. Panicum arnacites Trinius forms the basis of a net\ genus, Tatianjx. U'ith these changes in content the descriptions of Homolepis and Streptostachjs are emended, two new species-Brachiaria tatianae and Streptostachjs ramosa-are described, and the unusual species P. aristellum Doell is characterized and a possible relationship discussed. A chart is presented that conipares the characters of the genera covered in the present investigation, P. aristellum and the ttvo new species are illustrated, and SEX1 photomicrographs of all taxa are included.

OFFICIALPUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIESCOVER DESIGN: Leaf clearing from the katsurd tree Cerczdzphyllum japonicum Siebold & Zuccarini.

Library of Congress Cataloging in Publication Data Zuloaga, Fernando. Classification of the outlying species of New W'orld Panicum (Poaceae: Paniceae). (Smithsonian contributions to botany ; no. 59) Bibliography: p. Sum. of Docs. no.: SI 1.29:59 1. Panicurn-Classification. 2. Grasses-America-Classification. 3. Grasses-Classifica- tion. 4. Botany-America-Classification. I. Soderstrom, Thomas R. 11. Title. 111. Series QKl.S2747 no. 59 [QK49.5.G54] 581s [584'.92] 84-600221 Contents

Page Introduction ...... 1 Methods and Materials ...... 3 Acknowledgments ...... 3 Key to the Genera of Paniceae ...... 3 Brachiaria ...... 4 Brachiaria megastachya. new combination ...... 5 Brachiaria tatianae. new species ...... 7 Brachiaria albicoma. new combination ...... 11 (Comparison of Hornolepis and Panicum) ...... 11 Hornolepis. emended ...... 14 Key to the Species of Homolepis ...... 14 Hornolepis aturensis ...... 15 Hornolepis glutinosa. new combination ...... 19 Hornolepis isocalycia ...... 24 Hornolepis longispicula ...... 27 Homolepis villaricensis. new combination ...... 29 Ichnanthus ...... 31 Ichnanthus grandifolius. new combination ...... 31 (Taxonomic Position of Panicum aristellum) ...... 33 Panicurn ...... 36 Panicum aristellum ...... 36 (Taxonomic Position of Panicum killipii) ...... 40 Paspalurn ...... 40 Paspalum killipii. new combination ...... 40 (Taxonomic Position of Streptostachys) ...... 42 Streptostachys Desvaux. emended ...... 46 Key to the Species of Streptostachys ...... 46 Streptostachys asperifolia ...... 47 Streptostachys macrantha. new combination ...... 50 Streptostachys ramosa. new species ...... 52 (Taxonomic Position of Tatianyx) ...... 53 Tatianyx Zuloaga & Soderstrom. new genus ...... 56 Tatianyx arnacites. new Combination ...... 56 Literature Cited ...... 61

... 111

Classification of the Outlying Species of New World Panicum (Poaceae: Paniceae)

Fernando 0. Zuloaga and Thomas R. Soderstrom

Introduction genera is that of Pilger (1 93 l), who divided the The genus Panicum, described by Linnaeus in genus into eight subgenera and 11 sections, and 1753, is one of the largest genera of grasses, with of Hsu (1965), who made a complete analysis of approximately 600 species of worldwide distri- the tribe-Panicum in particular-reorganizing bution; the majority of these are tropical or sub- it into five subgenera and 25 sections. Hsu uti- tropical. Because of its size and widespread oc- lized newly recognized characters pertaining to currence the genus has been subjected to the such structures as the ligule, midnerve of the scrutiny of many investigators but none has stud- blade, lodicules, and epidermis of the leaf and ied it in its entirety. The African members re- upper lemma. ceived a rather thorough review in the early part Since the time of Chase, especially during the of this century by Otto Stapf of Kew, who allo- past decade, herbarium holdings of tropical cated them to a number of sections. On this side American Panicum have expanded dramatically of the Atlantic the American agrostologists due to an increase in collecting activity. As new Alfred S. Hitchcock and Agnes Chase made de- species have been described and known ones tailed studies of the North American species and restudied, the most interesting problems have grouped most of them according to natural affin- continued to be the delimitation of the genus ity. Their chief studies were published in “The itself and its position within the tribe. North American species of Panicum” in 19 10 and In the present study we have examined a num- “‘Tropical North American species of Panicum” ber of species that were originally described in in 1915. Chase also wrote a series of papers Panicum but did not seem to be well accommo- (1906, 1908, 191 1) giving decailed analyses of dated in it. This review has prompted us to re- New World genera in the tribe Paniceae. Among examine some of the characters that have been the most recent works on Panicum and allied used in the past to distinguish genera in the tribe, such as type of hilum, spikelet compression, rel- Fernando 0. Zuloaga, Instituto de Botanica Danuinion, Casilla de Correo 22, Sun Isidro, Argentina. Becario del Consejo Na- ative length Of the glumesj consistency Of the cional de Investigaciones Cientijicas y Tecnicas (CONKET) de upper anthecium and whether crested or not, la Republica Argentina. Thomas R. Soderstrom, Department of Botany, A’ational Museum of Natural History, Smithsonian In- and carriage Of the ’pikelet relative to the axis. stitution, Washington, D. C. 20560. Some of the most important characters that we

1 2 SMITHSONIAN CONTRIBUTIONS TO BOTANY

I I I I

1 I I 33

\ IL

n(d NUMBER 59 3 have found to be of generic value are illustrated cium” instead of “anthoecium” and “androe- in Figure 1. ci u m .” Information derived from studies made with Data in “Additional Specimens Seen”and “Par- the scanning electron microscope (SEM) on the atypes” sections have been quoted from specimen surface of the lemmas and paleas of the upper labels without editing, except for abbreviations floret has been especially useful in the present of distance and direction. study. Characters revealed at the higher magni- A synopsis of New World Paniceae is reflected fications often corroborate decisions based on in the Contents, with background notes and com- gross morphology. parisons set off with angle brackets. We have concluded that several species de- METHODS AND MATERIALS.-ClaSSiCal tax@ scribed in Panirum should be transferred to other nomic methods have been used in the present genera, such as Brachiaria, Ichnanthus, and Pas- study, utilizing a Wild M5 dissecting microscope palum. One of the species does not fit within the and Leitz Ortholux compound microscope. Ob- boundaries of any described genus and we are servations at higher magnification were made thus elevating it to that rank with the name of using the scanning electron microscope. For Ta tia njx. these studies, anthecia were removed from dried A study of gross morphological characters as herbarium specimens, secured on stubs, carbon well as those revealed by the SEM should be coated in a vacuum evaporator, and then coated extended to all species of Panicurn. When these with a gold-palladium alloy. The specimens so data are collected for the entire genus we will be treated were then viewed in a Cambridge S4- 10 able to compare them more meaningfully and or Cambridge Stereoscan 2.50 Mk 2 scanning arrive at a fuller understanding of the genus and electron microscope operating at 1 0-20 kV. relationships of allied taxa. In addition to the extensive herbarium collec- Rather than refer to the two florets (anthecia) tions available at US, we relied on specimens lent of the typical panicoid spikelet as fertile and by the following herbaria: B, BAA, BAF, CTES, sterile, we designate them by their position in the F, GH, LIL, LP, M, MO, NY, S, and SI. spikelet. The fertile antheciuni is thus the “upper ACKNOWLEDGMENTS.-The senior author is anthecium” and rvhat is usually called the sterile grateful to the Consejo Nacional de Investiga- floret is referred to as the “lower anthecium.” ciones Cientificas y Tecnicas de la Replublica This avoids the biased use of the word sterile, Argentina (CONICET) for a grant that allowed which is really incorrect, because this floret or him to spend two years (1982-1 983) at the anthecium usually contains stamens. Our use of Smithsonian Institution as a post-doctoral fellow. the term anthecium is for the unit that includes The junior author appreciates the opportunity the lemma and palea and when we refer to these afforded him by the New York Botanical Garden structures we again use their position: upper to participate in an expedition to the Brazilian lemma, upper palea, lower lemma, lower palea. cerrados in 1964, at which time he collected As the earlier spelling of “oecology” has given material of the new genus Tatianyx. To Ms. Alice way to “ecology”and ”gynoecium”to “gynecium” Ruth Tangerini we are both indebted for her in modern American usage, we have adopted skillful rendering of the illustrations of panicoid these more evolved forms. Following the same grasses in this paper and the chart of genera and rules we are also using “anthecium” and “andre- characters depicted in Figure 1.

Key to the Genera of Paniceae

(Acroceras and Hymenachne are not treated in this paper) 1. Hiluni of caryopsis linear ...... 2 2. Spikelets densely villous and positioned obliquely on the falcate pedicel, with a callus on the 4 SMITHSONIAN CONTRIBUTIONS TO BOTANY

dorsal face ...... Tatianyx, new genus 2. Spikelets glabrous and positioned evenly on the truncate pedicel, without a callus ...... 3 3. Lower and upper glumes equal in length, shape, and nervation; upper anthecium chartaceous with the margins of the lemma flat, the lemma not crested at the apex and rachilla not thickened between the lower glume and upper anthecium ...... Homolepis, emended 3. Lower glunie shorter than the upper, differing in shape and nervation; upper anthecium indurate with the margins of the lemma inrolled, the lemma crested at the apex or with the rachilla thickened between the lower glume and the upper anthecium ...... ,4 4. Upper anthecium crested at the apex; rachilla not thickened between the lower glume and upper anthecium ...... Acroceras 4. Upper anthecium not crested at the apex; rachilla thickened between the lower glume and upper anthecium ...... Streptostachys, emended 1. Hilum of caryopsis punctiform, rarely oblong...... 5 5. Upper anthecium membranous, hyaline, with the palea free at the apex, not covered by the lemma; lower palea absent; panicles spiciform to contracted, dense ...... Hymenachne 5. Upper anthecium indurate, when membranous the palea completely covered by the lemma at its apex; lower palea sometimes present; panicles lax to contracted, rarely spiciform . , .6 6. Spikelets with the back of the upper lemma turned away from the rachis, the lower glume facing it ...... Brachiaria 6. Spikelets with the back of the upper lemma facing the rachis, the lower glume turned awayfromit ...... 7 7. Upper anthecium with appendages or “excavations” at the base; spikelets keeled and somewhat compressed laterally ...... Zchnanthus 7. Upper anthecium without appendages or “excavations” at the base; spikelets not keeled nor compressed laterally ...... 8 8. Spikelets plano-convex, shortly pedicellate on unilateral racemes; lower glume cotnmonly absent, small when present ...... Paspalum 8. Spikelets biconvex, commonly arranged in panicles, less frequently in unilateral racemes, variously pedicellate; lower glume always present, variable in size ...... Panicum

Brachiaria upper anthecium is frequently papillose-rugose, a character that Stapf (19 19) also used to delimit In 1853 Grisebach elevated to the level of the genus. genus the section Brachiaria of Panicum, which Hitchcock (1935, 1951), Gardner and Hub- Trinius had established in 1826. As the type- bard (1938), Pilger (1940, 1954), and Jacques- species of his new genus, Grisebach cited B. eru- F6lix (1962), among others, considered spikelet cijormis (Panicum erucqorme Smith). Chase (1 9 1 1) position as an important character to differen- accepted the genus and underscored its distinc- tiate the genus. Ascertaining the position of the tive type of panicle and spikelets, the former lower glume in species with long pedicels, how- consisting of spike-like racemes and the latter ever, is difficult, but Gardner and Hubbard inverted in position, with the lower glume adja- pointed out a way to overcome this. They sug- cent to the rachis. She also pointed out that the gested that the spikelet just below the terminal NUMBER 59 5 one on the axis or on a branch of the panicle be the Paniceae; and while the texture of the upper examined to determine the position of the lower anthecium may be rugose or papillose in most glume, because the spikelet is generally short- Brachiarias, it is smooth in the type species! Pan- pedicellate and appressed to the rachis. They icum maximum, P. millegrana, P. sellowii, and P. commented that the orientation is still occasion- trichoides are examples of species in which the ally unclear due to suppression or rudimentary upper anthecium is rugose. development of the lower spikelet of the pair, in In the present study spikelet position is consid- which case the terminal spikelet appears to be ered to be of generic value, in concordance with solitary and in an abaxial position. most agrostologists, and is the basis on which we In 1969 Parodi, detailing and confirming the are transferring the following species of Panicum findings of Gardner and Hubbard, utilized the to Brachiaria: P. megastachyum Nees ex Trinius, character of spikelet position in transferring to 1826; P. megastachyum Nees, 1829, not Nees ex Brachiaria five species that had previously been Trinius, 1826; and P. albicomum Swallen 8c Gar- included in the Fasciculata group of Panicum. In cia-Barriga. some of these-B. echinulata, B. lorentziana, and B. mollis-the spikelets are borne on rather long Brachiaria megastachya (Nees ex Trinius) pedicels and their position is difficult to ascertain, Zuloaga 8c Soderstrom, new combination a situation reminiscent of some African species such as Brachiaria ramosa (L.) Stapf and B. deflexa FIGURE2 (Schumacher) Hubbard. Although she noted dif- Panicum megastachyum Nees ex Trinius, 1826:205. [Type: ficulty in differentiating the Brazilian species of “V. spp. Brasil. Langsdorff.” Holotype, LE, not seen; Brachiaria on the basis of spikelet position, Sen- fragment of the type, US sheet no. 974713. “In saxosis dulsky (1 978) accepted the character as valid and umbrosis Serra da Lapa,” cited under Panicum megastach- did not provide further ones. yum Nees, 1829:155, refers to this specimen.] Unlike previous authors, Blake (1958) did not Plants perennial (?), the base not seen. Culms consider spikelet position a definitive enough cylindrical, hollow, glabrous. Nodes obscure, gla- character to separate Brachiaria from Panicum, brous. Sheaths pale, glabrous. Ligule -1.5 mm offering other characters instead: the texture of long, membranous, long-ciliate above; collar gla- the upper anthecium, presence or absence of a brous, inconspicuous. Blades lanceolate, acumi- rupture line, and type of areole on the dorsal nate, flat, approximately 12 cm long, 0.8-1 cm face of the upper lemma toward its base. Accord- wide, subcordate with the adaxial surface pubes- ing to him, cent, the abaxial glabrous and margins scabrous. In one, characteristic of Panicum subgen. Panicum (the Panicles lax, diffuse, many-flowered, -25 cm Panicum type), the fertile floret is more or less biconvex long, 10 cm wide, the branches alternate to and muticous with the lemma smooth over the whole of subopposite and diverging from the rachis, the the back and the rupture line and areole resemble the spikelets diffused; axis angular, scabrous, with rest of the back in surface in curvature. In the trend represented by Urochloa (Urochloa-type), the fertile floret thick evenly distributed papillose-pilose hairs; is plano-convex or nearly so, sometimes apiculate or branches hirsute with hairs similar to those pres- aristulate, with the lemma more or less strongly trans- ent on the axis, the axils densely pubescent, the versely rugose, the rupture line is prominent and the spikelets 2 or 3 on each branch, one subsessile, areole more or less flattened or depressed, less rugulose the other long-pedicellate, the pedicels 1-1 2 mm than elsewhere. long, triquetrous, hirsute. Spikelets long-ellipsoid, Unfortunately, these characters are not invaria- 3.8-3.9 mm long, 1.4-1.5 mm wide, stramineous ble. Pohl (1980) discounted the presence or ab- to brown, with the lower glume adjacent to the sence of a rupture line as a generic character in rachis. Lower glume ovate, acute, 1.8-2.2 mm 6 SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE2.-L?rachiaria megastachya, new combination: a, upper anthecium, palea side (X 100); b, apex of lemma, detail of a showing flat macrohairs (X 400); c, upper anthecium, lemma side (X 100); d, detail of c showing simple papillae evenly distributed over the surface, making it rugose (X 500). All photomicrographs based on Riedel96 (US). NULIRER 59 7

long, glabi.ous, 3-.?I-nerved, the nerves anasto- Brachiaria tatianae Zuloaga & Soderstrom, inosed to\vard the glabrous to pubescent summit. new species 1,Pper glume obtuse to subacute, 3.3-3.3 mm long, usually not covering the upper anthecium, FIGURES?I, 4 5-7-nerved, the nerves anastomosed to\vard the TYPESPECIMEN.-BR~Z%I L, )TINAS (;ERAIS: sl1oi.t -lariat e sii iii 111 it . Louler lemma subacute, " Habit ;at i 11 ii I ;i rg i n i b 11s s y 1 va r ii I 11 D i st r i ct 11 s Ada- glume-like, lanate on the margins, 3.6-3.7 iiim mantum ad Tejuco etc. provinciae 5linaruni ge- long, .i-nei-ved, the nerves anastomosed. Lower neraliiim." (Holotype, M;fi-agment of the Iiolo- pa lea iiai.imi.\.Iv el liptic, conspicuous, 3.2-3.4 nim type, CS sheet no. 9747.52.) Basis of Panicum long. 1 - 1.2 ii1111 it.ide, menibratious, glabrous megastachyum Nees, 1829:134, not Nees ex Ti+ mcept foi- flat Iiairs to\t.ard the apex; niale flower iiiais, 1826. The specimeii rvxs collected bv Xlar- piwent jvitli 2 lodicules and 3 anthers, 2.2 mni long. [$per antherium nai-roivlv ellipsoid, acute, tius. 3.2-3.4 111111 long, 1.1-1.3 inn1 it.ide, plano-con- Culmi erecti, simplices. Ligula membranaceo- \.ex, st i.;im ineoiis to broiln, glabrous, covered ciliata. Laminae lanceolatae, subcordatae. Pani- \\.it11 evenl\. distributed simple or compound pa- culae lame, diffusae, pyramidales, 30-35 cm lon- pillae, the apes of the upper lenlnia \%.it11iiiimer- gae, 15-30 crn latae, raceinis spiciformibus spi- oils hooks. (,'arjopsis not seen. culas binatas ferentibus. Spiculae ellipsoidae, 6- ~!I)DI-IION.AL SPECIMENSSEEN.-BRAZIL. M'ith- 6.8 mm longae. Gluma inferna ovata longitudine out localit\-, Kieclel 96 (US), 1143 (SY). lh spiculae aequans, 3-7-nervata. Gluma supera I)ISCL&N.-III 1829 Trinius added the fol- obtusa, 9-nervata. Lemma infernum 5-7-nerva- loiviiig d;ita iii his description of the species (as t ii 111 . Pa lea i 11feima lo tigi-el 1i psoi dea , floi-em mis- tl~anslated): culum gereiis. ;Znthoeciuni supei-iim longi-ellip- ii i , 1. ti i I 1 , I ii ii ii rt iii t 11 (;ulins 1 .X-2.4 111 tall, cylindrical, glabrous, branching, soi deu g 1ab on i o co fe i e aeq ;a 1 - shortly geniculate. Sheaths glabrous. Blades linear-lan- iter papillosum; rhactiilla super- ;arithoecium sii- ceolate, flat. Panicle rniich-branched, with the branches pei.um ext ell sa. spreading from the rachis, more than 30 cm long. Perennial (?). the base not seen. (;ulms erect, IHis illusti.atioii depicts a much-branched panicle c\,lindrical, sparsely pubescent near the nodes to tvith spikelets occurring mostly in pairs on the glabrous, niany-noded, simple. ,Yodes brown, pu- i';iceiiies, one s~ibsessile and the other long-pedi- bescent. Sheaths longer. than the internodes, (ella t e. -1 3-1.5 ciii long, pubescent, especially toiLxrd On a sheet ivith a fragment of the type speci- the upper part, pale. Ligule menibraiious, ciliate, iiieii ;it US, Cliase made the following annota- - 1.5 iiini long; collar villous. Blades lanceolate, t ion: acuminate, flat or ivith involute niargins, -23 ciii

Culms 4-6 ft., blades 1 ft. or more, Ih inch wide, gla- long, 1 ciii ivide, rigid, subcordate at the base, brous, sheath wholly glabrous or silky on the collar, pubescent 011 both surfaces, especially toward the panicle about 1 ft., erect, contracted, rachis 8c branches base, the mai.gins smooth. Panicles lax, diffuse, mid pedicels pilose, spikelets about 4 mm, fruit slightly pyi-amidal, many-floivered, 30-3.5 ciii long, 1 .?I- roughened, not shiny. Type of Nees ex Trin. 1826, not 30 ciii \tide, spikelets in pairs on spike-like ra- Nees ipse. cemes, one subsessile, the other short-pedicellate; Even tlioiigli the terminal spikelet in this spe- axis angled, hispid, the branches alternate or cies is boi-ne 011 a long pedicel, the inherent subopposite, the lokver verticillate, diverging position of the spikelet is obvious in a subsessile from the axis, naked at the base, hispid above, meiiiber, ivhich al\vays occurs kvitti the lower flexuous; pedicels 0.8-4 iiiiii long, shorter than glume adjacent to the rachis. the spikelets, triquetrous, \\'it11 long thick hairs FIGUREJ.--Brachiaria tatianae, new species: a, habit of culm (X 1); b, panicle (X 1); c, spikelet, upper glume side (X 12); d, spikelet, lower glume side (X 12); e, upper anthecium, palea side (X 12);J; upper anthecium, palea side showing prolongation of rachilla (X 12); g, lower glume (X 12); h, upper glume (x 12); i, lower lemma (X 12);j, lower palea (X 12); k, spikelet, lateral view (X 12); I, branch of panicle (X 9); m,developed lemma of third anthecium (X 12); n, upper palea with gynecium and lodicules (X 12); 0, upper palea with stamens and lodicules (X 12). All draivings based 011 the holotype (M). NUMBER 59 9

FIGURE4.--Brachzaria tatzanae, new species: a, upper anthecium, palea side, showing compound papillae (X 1000); b, upper anthecium, palea side, showing apiculate summit of lemma and flat macrohairs (X 100); c, upper anthecium, palea side at base showing prolongation of rachilla (X 50); d, detail of summit of lemma as shown in b (X 1000). All photomicrographs based on the I1olot)pe (M). 10 SMITHSONIAN CONTRIBUTIONS TO BOTANY

toivard the apex and base. Spikelets ellipsoid, 6- Sellou~ 1276" (fragments, BAA, US sheet no. 6.8 iiiiii long, 2.2-2.4 niiii \t.ide, lanate on the 9 7 47 .i2). apex of the gliinies and loiver lemnia, otherwise DrscvssrON.-In his De Graminibus Paniceis of glabrous, tlie gl tiiiies and loi\.er lemma ivith 1826, Trinius described niany species recognized pi-oiiiinent iien'es, the upper glunie, loiver bv C.G. Nees voii Esenbeck, indicating after each lemma, and upper antliecium subequal. Lower iiaiiie: "S.ah Es! iii Slai-t. FI. Bras. ined." Three glume o\'ate i\.itli ;I rounded to acute apex, 4-4.3 years later Nees in his .Igrostologia Brasiliensis nil11 long, half ;IS long ;is the spikelet or a little published these taxi ;is nelv, presumably unaivare longer, 5-7-iierved, the nerves aiiastoiiiosed to- that they had already appeared in print. In most i\.ai.d the apex, tlie summit and upper margins of tlie species described-/'. chloroticum, P. ele- short-lanate. l'pper glume obtuse, j.3-6.3 iiim phantipes, P. decipiens, P. milioides, P. cyanescens, long. 9-nerved, Ianate, not covering the apex of and P. ligulare, for example-the descriptions of the tipper antlieciuni. Lozler lemma glunie-like, both aiit1ioi.s refer to the same species and usually aciite, 5.5-6.3 iiiiii long, .i-7-nerved, the nerves cite the saiiie type specimen. Hoivever, in at least aiiastoniosed. Lorcer palea long-elliptical, 4.9-5.9 tivo ciises the tvpe indicated bv both authors is miii long, 1.6-2.1 iiim [vide, membranous, short- different. One of these, P. campestre Sees ex pilose or glabi.oiis in the middle part and on the Trinius. 1826, is a valid species of the Rudgeana riiargins: niale flol\.er present ivith 2 lodicules gi-oup and distinct froni P. campestre Nees, 1829, and 3 ;intliei-s. Lpper anthecium long-ellipsoid, ivhich is ;I valid species of the section Panicum. 4.7-.5.9 iiiiii long, 1.6-2.1 iiiiii \vide, brown, Henrard described the latter as new in 1940, densel!. papillose throughout, the papillae simple giving it the 1i;iiiie of Panicum peladoense. or coiiipouiid, tlie niargins of the lerii~iiacover- The otlier case in \t.hich tivo species have mas- ing the plea, [vitli iiuiiiei-ous hooks on the apex queraded under the saiiie name is Panicum me- of the leriinia and flattened hairs on its upper gastachpm. The trvo are characterized in Table iiiargins; i-acliilla pi~olongedinto a short acute 1. iiiiicro above the upper antliecium, conspicuous Nees in fact referred to both specimens in his in txi-).ing degrees 011 different spikelets; lodi- original desci-iption, the one a collection of Afar- ciiles 2, truncate; anthers 3, violet. Caryopsis not tiiis and the other of Langsdorff. Hoivever, the seen. description of Nees agrees closely ivith the Mar- PARATYPE.-BRA%IL. "Panicum megastach- ti 11s specimen but not lvith that of Langsdorff. yum. FI. Bras. \,arpeduizculis basi et apice barbatis, Ti-inius cited onlv the Langsdorff gathering.

TABLE1 .-Comparison of P. megastachyum Nees ex Trinius and P. megastachyum Nees.

P. megastachjum Nees ex Trinius, 1826 P. megastachyum Nees, 1829 spikelets 3.8.-3.9 mtn long spikelets 6-6.8 mm long spikelets long-pedicellate spikelets short-pedicellate rachilta not prolonged beyond the upper an- rachilla prolonged as a conspicuous mucro t heci uni beyond the upper anthecium axis and branches hirsute with thick papil- axis and branches hispid with papillose-pilose lose-pilose hairs; pedicels hirsute with thin, hairs only at the base: pedicels papilloze- evenly distributed hairs pilose at the apex and base lower glume 3-5-nerved lower glume 5-7-nerved upper glume 5-7-nerved upper glurne 9-nerved NUMBER 59 11

Iloell (in llartius, 1877) reiterated what Nees mm \\ide, hyaline, glabrous; male floiver present, had i.\.i-itten but noted that spikelet size and pu- rvith 3 stamelis. Upper anthecium ovoid, 3.3-3.7 bescence of the pedicels differed in the two spec- nini long, 1.6-1.8 inin rvide, glabrous, strami- imens. Steudel (183.5) had also observed the dif- neous to brorvn, transversely rugose, \\.ith round ference in pedicel pubescence. papillae distributed evenly; lemma apiculate, 5- nerved Ivith a scabrous apex: lodicules 2, truncate, plicate, enclosing the basal margins of the Brachiaria albicoma (Swallen & Garcia palea: Lvith 3 stamens. Caryopsis not seen. Barriga) Zuloaga & Soderstrom, So far knoivn only from the Department of new combination Cauca, Colombia, at elevations of 1600 to 1900 FIGURE5 111. ADDITIONALSPECIMENS SEEN.--COLOILI- Panicum albicomum Swallen k Garcia Barriga, 1943:303, fig. BIA. 11Tithout state and locality, 'Llutis 5522 (US). c. [Type: (hlombia: "Dept. del Cauca. El Ramal to Rio Sucio, r\'est of Popayan, Cauca valley, Pennell 3 Killip CAUCA:El Tanibo, Garcia krriga 4426 (US): 8129." Holotype, US;isotypes, GH, NY.] Chisquia, Finca Los Derrumbos, Asplund 10778 (US). Annuals 1.6-2 in tall. Culms cylindrical, erect, DIscussIoN.-This species is apparently most geniculate ;it the base, simple or branched, hol- closely related to Brachiaria mollis (Srvartz) Par- Ion., hispid. Sodes short- to long-pubescent r\.ith odi, lvhich differs in its smaller spikelets, 3.5-4 u.Iiitish hairs. Sheaths usually longer than the nini long, and \\.idel-pubescent glunies. intei-nodes, 12- 19 cni long, densely to sparsely hispid, short-ciliate on the n~argins.Ligule mem- Comparison of Homolepis and Panicum branous, ciliate, 1 - 1 .6 iiini long: collar lanate. H 1a d es 1 i I i ea r-1 an ce ol a t e , ac u ni i n a te , 18 - 2 4 cm Homolepis \\.as established in 19 1 1 by Agnes long, 1.4- 1.8 cni \vide, flat, subcordate, densely Chase, rvho based the genus on Panicum aturense Iiispid, especially toivard the base on the adaxial H.B.K., deriving its name from the Greek homo surfiice, tlie niai-gins scabrous lvith short, ap- (similar) plus lepis (scale), in reference to the pressed hairs, tlie niidnerve prominent. Panicles subequal glumes. In addition to the type species, oblong, lax, 22-30 cni long, 8-14 cni rvide, the H. aturensis (H.B.K.) Chase, trvo further species bra n c h es xcen d i n g , t tie s pi k el e t s short- pedi ce 1- were included: H. isocalyia (Meyer) Chase and late in pairs or in threes, \$.ith the lower glume H. longispicula (Doell) Chase, both transferred f,icing : the axis: branches and pedicels angular, from Panicum, densely hirsute to villous, lvith long lvhitish hairs Diagnostic characters given by Chase Lvere the up to .5 nini long, thick toward the apex and Mith subequal glunies that are 7-9-nerved and enclose shorter hairs intermixed: the branches alternate, both anthecia; enclosure of the upper antheciuni \\.idelv sp;iced and bearing spikelets to the base. by the leninia of the lotvei. aiitheciun~,Mhich is Spikelets long-ellipsoid, 4.5-5.1 mm long, 1.6- subequal to the glunies: and presence of a palea I .8 nini \vide, glabrous, pale to dark purple, the in the loiver antheciuni. Furtherinore, the upper glumes and lo\~.erlenirna Lvith thin nerves. Lower antheciuni is not indurate but rather delicate and glume o\.ate, acute to obtuse, 1.9-2.5 nini long, fragile and the margins of its leinnia are flat ;I third to tialf as long as the spikelet, 3-nerved, rather than inrolled, enclosing most of the palea. the apex scabrous. ["pper glume acute, 7-nerved, Several vegetative characters were also given for the nerves anastotnosed ton-ard the scabrous the genus: plants perennial, stoloniferous or apex. Lower lemma glume-like, 7-nerved, the creeping, decumbent and rooting on the loiver nerves anastoniosed toward the scabrous apex. nodes. These latter characters, houever, do not L,ouler palea lanceolate, 3.7-3.8 nini long, 1.3 apply to plants of H. longispicula, u,hich are ces- 12 SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE.i.-Brachiaria albicoma, new combination: a, upper antheciuni, lemma side, showing apiculate summit of lemma (X 1000); b, detail of lemma surface, shoiving rugose surface and simple papillae (X 200): c, upper antheciuni, lemma side at base, showing rupture line (X 70); d, upper anthecium, palea side, shoiving bicellular microhairs and prickle hairs (X 200). All photomicrographs based on Garcia Barrzga 4426 (US). NUMBER 59 13 pitose, erect, and iii vegetative and habitat char- longipedicellaturn, and in species of the Pat-vifolia ;icters totally distinct from the remainder of the group, for example. M’ith feiv exceptions, the species. Its spikelets, nevertheless, are those of upper antheciuni is indurate in Panicurn and the Hornolepis. margins of the ~ippei-lemnia are inrolled over Regional floras \vith references to Hornolepsis the palea (Figure 12). In Hornolepis the upper include Hitchcock (19 13, 1922, 1927, 1930a,b), antheciuni is ch;ii.taceous and the m;irgins of the S~vallen (1 943, 19.55), Amshoff and Henrard lemma ai-e flat and not ini.olled over the palea (1 94H), and Pohl (1980). In all of these Hornolepis (Figure 9). The lo\z.er lemina of Hornolepis noi’- is distinguisl~edfrom Panicurn by the long loiver mallv exhibits differing degrees and types of glume, noti-inctui.ate ~ipperantheciuni, and thin, pubescence, ivliile tlie glunies are glabi.ous (pilose flat niargins of’ the upper lenima. These criteria up to the margins in H. aturensis and H. longis- i$.ei.ealso used 11:. Pilger in 19.34, but Hsu in his picula). C:on\.ersely, \%.lienthe spikelet is pubes- 1965 studv of‘ the Paniceae made no mention of cent in Panicurn, the hairs occur on both glumes tlie genus. and inore sparsely over the loiver lemnia. 0111- st utlies Iiave shown that two additional Carjopsis: The hiluni of Hornolepis is linear, a species of Panicurn, P. glutinosurn and P. villari- chai.acter that clearly distinguishes it froni Pani- cense, shai-e the generic characters of Hornolepis cum, in \$.Iiicli this structure is punctifoim or and should be transfei-red to it, bringing to five rarely oblong. il’hile linear in H. glutinosa and tlie nunibei- of species in this American genus. H. viilaricensis it is sonieF\.hat broadei- and occu- Tlie forniei. beconies tlie most \$.idel?distributed pies half or ;I little inore tlie length of the car- menibei-, occurring from Mexico and the M’est yopsis. Chase niade no mention of the caryopsis I ndies to Argeiitiiia. Hornolepis isocalyia is found in her desct-iptioii of the genus. in liexico (iv1ie1.e only the type of one of its Silica Cells: The presence of a particular type s~~non~~nis,Panicurn langei, has been collected), of silica cell in the epideimis of the upper anthe- Rinaina, atid South America, ivhile H. aturensis ciuin is ;I peviously unrecorded charactel- that ranges fi-oni hiexico to Bolivia and Brazil. T~vo further differentiates the t\vo genera. Such silica species are know^ otil>. fiorn South America: H. cells in Hornolepis are dumbbell-shaped and alike longispiczclu is endeniic to the cerrados of Brazil in all species. Silica cells also occur toivard the and H. uillaricensis is restricted to Paraguay, Ar- tip of the upper antheciuni in certain species of gentina, and southern BI the Imi-ea and Lisa groups of Panicurn but are New features, as revealed by SEM studies, rare, randonil>,disposed, and niai.kedly different ~OTTO~OTJt e the previous separation of Pa nicurn from those in Hornolepis. and Hornolepis that ivas based on characters of The 1iteratui.e contains relativel!. few studies the spikelet, flo\\.er, and fruit. The major differ- concerned \zit11 the surface of glunies and anthe- ences bet\\.een these genera ai-e summarized in ciurn, among these Baum (1 97 l), Baum and Had- the follo\ving paragmphs. land (1 973), Clark and Gould (197.i), Thonias- Spikelet: The lo~,erand upper glumes in son (197Xa,b), Shaw and Seims (1 gig), SIiaFv and Hornolepis are similar, equal in length and not Webster (1 98 l), and Tei-rell and M’ergin (1979, differing iii their morphology, and the margins 198 1). Clark and Gould (1973) utilized differ- of all bracts are o\,erlapping. The length of the en tial chai-acters of the ant heci ii ni s LI r face to Io~.erglunie in Panicurn varies from ]/r, to Y4 that separate Ilichantheliurn from Panicurn, pointing of the spikelet, not equalling it nor the upper out that the upper palea in the former is covered glunie. In 21 feiv species, where the lower glume evenly \$.it11 simple papillae, rvliile in the latter is more de\.eloped, it differs from the upper in they do not occur or are cro.tvded together and such features as type of apex, nei distributed at randoni. Terrell and \Vergin pubescence. This occurs in Panicurn obtusurn, P. (198 1) studied silica cells present on the surface 14 SMITHSONIAN CONTRIBUTIONS TO BOTANY of the pistillate lemmas in Zizania, showing that void, blunt or ivith an acute or acuminate apex, the four species of the genus could be separated greenish or brown, compressed dorsiventrally, by the different types of silica cells present in articulated below the glumes; glumes subequal each. or the lolvei. somewhat larger than the upper, Chromosome ‘Yumbers: Gould and Soderstrom completely covering the lower lemma and upper (1970) reported the first counts for the genus anthecium or occasionally leaving exposed the \\.it11 211 = 40 for H. isocalyia and 211 = 22 for summit of both anthecia. Lower glume 5-9- H. aturensis, although Pohl and Davidse (1971) nerved, glabrous, its margins enclosing the up- and Davidse and Pohl (1978) found the latter to per. Llpper glume 7-9-nerved, commonly gla- be 211 = 20, concluding that the genus has a basic brous, its margins enclosing the lower lemnia. number of x = 10. These authors have also Lower lemma glume-like, 5-7-nerved, pilose, its reported counts for H. glutinosa (as Panicurn glu- margins broadly enclosing the upper antheciurn. tinosurn) in their papers of 1971, 1972, 1974, Louier palea membranous, often equal in size to and 1978. The most common basic number in its lemma and conspicuous; r-nale flower sonie- Panicum is x = 9, although x = 10 occurs in times present. Lpper anthecium ellipsoid, charta- species of the Laxa and Mertensia groups. ceous, compressed dorsiventrally, acute or acu- Leafblade Anatomy H. glutinosa is a non- minate, glabrous (sometimes with bicellular rni- Kranz or G4 species according to the data given crohairs or hooks toward the apex), pale, covered by Bro\zn (1977) under Panicum glutinosum. rvith suborbicular silica cells evenly distributed Broivn also listed Homolepsis aturensis and H. over the lenima and palea; lemma with flat mar- isocalyia ;is C:3 s?ecies. gins, not inrolled over the palea and enclosing the greater part of it; palea of similar texture; lodicules 2, conduplicate; anthers 3; stigmas 2, Hornolepis Chase, 1911, emended Zuloaga 8c plumose. Caryopsis obovoid to oblong, broivn, Soderstrom with a linear hilum and embryo more or less half its length. Hornolepis Ch‘ise, 1 9 1 1 : 146. An exclusiirely American genus kvith species Plants perennial, stoloniferous or creeping, de- occupying huiiiid habitats and shady places in cumbent and rooting at the loizer nodes, then forests or woods from Mexico to Bolivia, Brazil, becoming erect, rarel) cespitose. Spikelets short- Paraguay, and Argentina, with one species en- to long-pedicellate, arranged in lax panicles. demic to the cerrados of Brazil. Spikelets lanceolate to long-ellipsoid or obo- TYPE-SPECIES.-panicum aturense H.B.K.

Key to the Species of Hornolepis

Spikelets 9-1 1 mm long, 2-2.9 mm wide; glumes and lower lemma pubescent, lower lemma with long submarginal hairs; lower palea conspicuous, as long as its lemma; male flower present; plants cespitose with the culms few-noded, erect, not branching; ligule ciliate; sheaths keeled, rigid; blades linear-lanceolate, rigid, involute, 0.2-0.6 cm wide. Cerrados in the north of Brazil ...... H.longispicula Spikelets 2.7-8 inn1 long, 1-1.8 mm \\side; upper glume usually glabrous, lower lemma sparsely pube3cent; louer palea small, never equal in size to the lemma; male flower rarely present; plants stoloniferous to decumbent, rooting and branching at the lower nodes; ligule membranous to membi-anous-ciliate; sheaths rounded, herbaceous; blades oblong-lanceolate to ovate-lanceolate, flat, 0.6-2.3 cm wide. Inhabiting forests or woods, never in cerrados ...... 2 NUMBER 59 15

2. Spikelets 2.7-3.7 mni long; lower lemma without thick hairs in the central part: caryopsis Mith a broadened linear hilum half or a little more than half as long as the carvopsis; upper anthecium scabrous at the apex ...... 3 3. Panicle 12-25 cm long, 7-22 cm wide, with the lower branching verticillate; plants robust, 0.6- 1.8 111 tall; blades 10-43 cm long, 0.6-2.3 cm wide ...... H. glutinosa, new combination 3. Panicle 5-1 1 cm long, 1.5-5 cm wide, with opposite or alternate branches, not verticillate; plants small, 0.2-0.4 m tall: blades 4-12 cm long, 0.6-0.9 cm wide ...... H. villaricensis, new combination 2. Spikelets 4.6-8 mni long: lower lemma with thick hairs in the central part; caryopsis with a narro\<,linear hilum as long as the caryopsis; upper anthecium glabrous...... 4 4. Spikelets ellipsoid, acute, 4.6-7.5 mm long; lower lemma sparsely pubescent between the lateral nerves arid with thick hairs on the middle part or glabrescent; upper anthecium acute, chartaceous ...... H. isocaljcia 4. Spikelets lanceolate, acuminate, 6.2-8 mm long; lower lemma densely pubescent between the lateral nerves and with thick papillose-pilose hairs up to the apex of the middle and toward the sides; upper anthecium acuminate, thin, of weak consistency ...... H.aturensis

Hornolepis aturensis (H.B.K.) Chase the upper part, 0.3-0.6 mm long. Blades oblong- lanceolate, acuminate, 6-1 2 cm long, 0.7-1.5 cni FIGURE6 \vide, flat, i<.ith a short pseudopetiole, the base

Hornolepis aturensis (H.B.K.)Chase, 191 1:146. cordate, the njargins ciliate or scabrous, \vith Panicurn aturense Huniboldt, Bonpland, and Kunth, strigose pubescence on both surfaces or glabrous, 1816: 103, pl. 33. [Type: "Crescit ad cataractas Aturen- tessellate. Panicles lax, ellipsoid, 3.5-9 cni long, sis." Holotype, P, not seen; fragment of the type, US sheet 1-4 cm \vide, the branches shortly divergent 110. 80.535.1 from the axis lvith the spikelets congested along Panicurn nzridzjlorurn Nees, 1829: 135. [Type: "Habitat in the branches; axis glabrous, smooth, undulate, herbario nostro occurrit, nesco nunde illatum." Type not Ioca ted. ] the branches alternating, infrequently opposite, Panicurn blepharoporum J.S. Presl in K.B. Presl, 1830:312. glabrous, the axils glabrous. Spikelets fusoid, ac- [Type: "Habitat in Mexico." Holotype, PR, not seen; uminate, 6.2-8 nim long, 1.3- 1.8 tnni !vide, fragmenr of the type3 US sheet no. 80486.1 greenish to pale, herbaceous, strongly coni- Panicurn turnescens Tr.inius, 1834:s 16. [Type: "V. spp. Bras." pressed dorsiventrally, glunles and loiver- leninia Holotype, LE, not seen; isotype, NY; fragment of the faintly nerved. Lower glume acuminate, as long as type, US sheet no. 81 158.1 the spikelet, enclosing part of the the upper Stoloniferous perennials, 20-60 ciii tall. Culms glume, 5-9-net-ved, glabi-ow. Cyper glume 7-9- compressed, glabrous, erect or geniculate-as- nerved, frequently tessellate, short- to long-pu- cending and rooting at the lokver nodes, the bescent betLveen the lateral nerves in the zone internodes hello\<., 2- 1 1 cni long. lYodes brown, covered by the loi$.ei.glunie, the hairs delicate, con i pressed, g I a b 1-ous or 1%. it h 1on g 1%. hit i s h hairs . the rest of the surface glabrous. Louler lemma Sheaths ?-7 ciii long, the margins pilose with thin glume-like, 7-nerved, densely pubescent betlveen r\.hitish hairs, glabrous over the rest of the sur- the lateral nerves, the hairs purplish and whitish, f'xe 01- I~;ipillose-pilose:collar broLvn, pubescent the central nerves inconspicuous, ivith thick tub- to glabrous. Ligule niembranous, short-ciliate on erculate hairs in the middle and oblong bicellular 16 SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGUREri.--Homolepis aturensis: a, upper anthecium, lemma side (X 500); b, detail of upper palea shoi\.ing a suborbicular silica cell (X 5000); c, lower lemma showing base of a macrohair (X .iOO); d, lower lemma showing macrohairs (X 100). Photomicrographs a and b based on Arrher 1426 (CS), c and d on Prance 2860 (US). NUMBER 59 17 microhairs. Lower palea lanceolate, 2-3.2 mm Moura, Negro river, Baldwin 3365 (US); Reserva long, 0.5-0.9 nini wide, whitish, hyaline, gla- Fiorestdl Ducke, km 26, Prance et al. 2627 (US); brous; male flower rarely present. Upper anthe- Santa Isabel, Negro river, Baldwin 3457 (US). cium ellipsoid, acuminate, 4.6-5.3 mm long, 1.1- GOIAS: Rio Doce, 65 km W of Rio Verde, Chase 1.4 mm wide, glabrous, pale to whitish, herba- 11727 (US); W of Rio Verde, Chase 12072 (NY, ceous, with silica cells distributed evenly, the US). MARANHAO:Barra do Corda to Grajau, lemma 5-nerved, its wide margins enclosing the Swallen 3670 (US); kni 427, margens da Rod. palea. Caryopsis broadly obovoid, 2.5 mm long, BR-316, Rosa €3 Vilar 3136 (NY). PARA:Belem, 1.3 mm wide, brown, with a linear hilum, as long Kuhlmann s.n., 13 Aug 1923 (US); Braganga as the caryopsis; embryo less than half the length railroad, Goeldi 43 (NY, US); Castanha, Fazenda of the caryopsis. Setenta-dois, Mexia 5909 (US); Lagoa Agua DIsTRrBuTIoN.-~idely distributed from Preta, Drouet 2028 (NY, US); Mosqueira, Swallen Mexico to Bolivia and northern Brazil. Found 4891 €3 1/2 (US); Santarem, Spruce 30 (NY, US), commonly in margins of forests or woods or in 765 (US); 36 kni SE of Vigia, Dauidse et al. 17591 open inundated or swampy places and along the (NY). PERNAMBUCO:Dois Irmaos, vicinity of Re- margins of rivers in wet and sandy soils. This cife, Chase 7725 (US); Recife, Pickel 3608 (US). species, which occurs from sea level to 1700 COLOMBIA. Angelopolis, vicinity of Medel- meters elevation, has been reported as a weed of lin, Taro 270 (NY, US); Bnenaventura, Hitchcock plantations in Brazil. 19905 (US); vicinity of MedelIiA, Toro 474 (NY, ADDITIONALSPECIMENS SEEN.-BELIZE. EL US); Santa Marta, Smith 170 (NY, US). AMA- CAYO: Central Farm, Reyes 4 (US); Cohune ZONAS: about 6 kin N of Leticia at Santa Isabel, Ridge, Lundell 6425 (US); Augustine, Mountain Gillett €3 Dickenson 16520 (US); Loretoyacu river, Pine Ridge, Hunt 101 (US); Cay0 Road, Gentle Schultes 6015 (US). ANTIOQUIA:Puerto Berrio, 9704 (US). STANNCREEK: near. Stann Creek Juzepcruk 4862 (US); Santa Catalina, Bosse 3670 town, Gentle 9097 (US); Stann Creek, Schipp 896 (US); Venecia, granja de las Mercedes, Gutierrer (NY, US). TOLEDO:Punta Gorda-Cero Road, CY Barkley s.n. (US 2184160). CAUCA:Chisquio, Gentle 6835 (US); edge of pineland, condemned Finca Los Derrumbos, Asplund 10558 (US); C6r- Branch Pine Ridge, Gentle 7752 (US); in sacatal, doba, Dagua valley, Pittier 560 (US); El Tambo, near Machaca, Gentle 6891 (US); between cattle Garcia Barriga 4416 (US); La Esmeralda, near landing and Rio Grande, Gentle 6879 (US); Gra- Jamundi, Cauca valley, Pittier 936 (US); Rio Su- cie Rock, Sibun River, Gentle 1629 (NY, US); cio, W of Popayan, Pennell €3 Killip 8163 (US). Temash river, Schipp 1373 (NY, -US); Masons CHOCO:Atrato river, above Quibd6, Sabuidero, Lease, Smart 54 (US); Little Cocquericot, Belize Cuatrecasas €3 Llano 24062 (US); Andagoya, Kil- river, Lundell 3878 (US). lip 35497 (US); Atrato river, near Quibd6, Mol- BOLIVIA. BENI: Guayamerin, Krapouickas €3 irta €3 Barkley s.n. (US 2206657); between La Schinini 35109 (US). SANTACRUZ: Buena Vista, Oveja and Quibd6, Archer 1690 (US); Quibd6, Ste3nnbach 1844 (NY), 5120 (US). quebrada la Platina, Idrobo €3 Cuatrecasas 2705 BRAZIL. ACRE: Cruzeiro do Sul, road Cruz- (US); San Juan river, Cuatrecasas 21372 (US); eiro do Sul-Japini, Prance et al. 2860 (US). AMA- Serrano river, Forero et al. 1371 (NY). EL VALLE: ZONAS: Barcelos, Negro river, Baldwin 3330 Rio Calima, Aguaclara, Cuatrecasas 21 167 (US); (US); Icana river, Santa Ana, Froes 28472 (US); Alto del Dinde, between Cartago and Alcali, Japura river, Rodrigues 620 (US); 30-50 km from Cuatrecasas 22939 (US); Buenaventura, AsplCnd Manaus, along the Manaus-Itacoatiara road, Ei- 4984 (US); Buenaventura, Cbrdoba, Idrobo 6250 ten €3 Eiten 5267 (US); Manaus, km 8 da BR 17, (US); Cajambre river, Pacific coast, Cuatrecasas Rodrigues €3 Chagas 3083 (US); Manaus, rua Du- 16955 (US); Calima river, Pacific coast, Cuatre- que de Caxias, Rodrigues €3 Coelho 3076 (US); casas €3 Willard 26071 (US); Pacific coast, Yu- 18 SMITHSONIAN CONTRIBUTIONS TO BOTANY runiangui river, i’eneral, Cuatrecasas 15895 8.5, M’ of road, Contreras 2955 (US); Puerto (US); Isla del Guayabal, in the mouth of Ca- Barrios, Brenckle 47-184 (KY),Hitchcock 9156-11 jambre ri\,er, Cuatrecasas 16222A (US); Naya 2 (US); vicinitv of Puerto Barrios, Standley 24725, river, Puerto \lei-izalde, Pacific coast, Cuatreca- 24978 (US). sas 14001 (US): Calima river, bettveen La Esper- FRENCH GCIANA. CAYENNE:vicinity of anm y Bellavista, Cuatrecasas 16796 (US). META: Cayenne, Broadway 703 (NY, US). along Rio Ocoa, SE of Villavicencio, Kill$ 34371 HONDURAS. CORTES:Yojoa Lake, Garcia 93 (US); i’illavicencio, Cuatrecasas 1973 (US), Pen- (NY): Agua ilzul, on shores of Lake Yojoa, ,!lor- nell 1514 (US), Pe‘rez 5336 (US). SANTANDER: ton 7699 (US): Puerto Cortess,Carleton 623 (US); Granja Expel-iniental Agua Caliente, N of between Agua Azul y Pito Solo, ,Vlolina 5584 Plaron, Robinson Jr. 3010 (US); La Gomez, 40 (US). MORAZAN: nearEl Hombre river, NW of kni E of Puerto M‘ilches, Fosberg L3 Fassett 21 767 La Venta, Molina 13420 (US). GRACIASA DIOS: (US); Puerto Wilches and vicinity, Killip 3 Smith along Platano river, Pohl €3 Gabel 13342 (NY). 14813 (NY, US). VAUPE SIS~S:Rio GuainIa, Puerto MEXICO. CHIAPAS:near the junction of the Colombia and vicinity, Schultes et al. 18149 (US). Rio Perlas and Rio Jatate at San Quintin and COSTA R I (;A. CARTAGO:I nstit uto Intera- near Laguna Miramar, Sohns 1633 (US). OAX- iiieric;ino de Ciencias Agricolas , Turrialba, Dav- ACA: Chiltepec, ,Zlartiner Calderon 721 (US), 981 idse 935 (US); betr\.een Pavones y Charia, Tur- (NY); Comaltepeque, Galeotti 5859 (US); Santa rialba, Jimenez 751 (KY). LIMONN: Hamburg Maria de Lovani, Vera Santos 3440 (NY, US); 1 Finca, on the Rio Keventazon below Cairo, Stan- km S of i’alle Sacional, on the road to Sari dlej 3 Valerio 48682 (US); 3 mi S of Puerto hlateo, Rredowski 33792 (NY). TABASCO:Mayito, Limon, Lathrop 5542 (US); betlveen Rio Linion Rouirosa 425 (KY, US); Ocuapan, near Huirnan- v Rio Banano,Jimeenez 2830 (NY); Rio Cimarron, guiilo, Archer 3973 (US). i’ERACRUZ: along the crossing of highivay, E of Siquirres, Pohl 13185 Trans-Isthmian Highivay, 4 Km NE of Minati- (KY):v;ille).s of Sixaloa river, 2 km Mi of Barnbu, tah, King 1070 (US); El Palniar, Zongolica, Vera Pohl 3 Pinette 13198 (NY).PUNTARENAS: 0.5 krn Santos 2304 (NY, US); Sabana Palenque, iblatuda \Y of Buenos :!ires, Pohl 3 Pinette 13262 (NY); 3736 (US); Jicaltepec, Liebmann 391 (US). Buenos Aires, Tondur 4876 (US). NICARAGUA. State unknown: La Tron- ECUADOR. ESMERALDES: Playa Rica, Parro- quera, ,!lolina 14956 (NY); region of Braggnian’s quia de Concepcion, .Llexia 8946 (US); in clear- Bluff, Englesing 295 (US); El Recreo, Semple 110 ings near Borbon, Jativa €3 Epling 2141 (NY). (US). ZELAYA:field S of airport, Seill6485 (NY). LOS RIos: Hacienda Sta.Luci a, canton Vinces, PANAMA. BOCASDEL TORO:Su’ of Bocas, iblexia 6584 (US); Rio Palenque Biological Sta- Mlacatv Hills, von Il’edel 529 (US); Isla Colon, von tion, Dodson 5813 (US). KAPO-PASTAZA:Tena, Il’edel 2801 (US). CHIRIQUI::Chiriqui CU~: Viejo river, Asplund 8954 (US). PICHINCHA:between Santo .ZlcCorkle 690 (NY); vicinity of El Boquete, Hitch- Domingo and Quininde, Acosta Solis 13694 (US). cock 8352 (US). COLON: a roundEmpire, Canal (; U A7’Ehl12 LIZ. ALTA VERAPAZ:Chama, Zone, Pittier 3727 (US); Barro Colorado Island, Johnson 456 (US); Coban, von Tuerckheim 1255 Fairchild Point, Ebinger 278 (US); Fort Sher- (SY,US), 3772 (US): Cubilquitz, uon Tuerckheim ni;iiin hlilitary Keservation, bettveen Chagres 7698 (YY, US): E’inca Socuyo, SE of Carcha, Batteries and Fort San Lorenzo, Canal Zone, Standlej 70285 (US); S of Cubilquitz, Steyermark .\.laxon 3 Valentine 7031 (US); bet\\.een France 44560 (US); vicinity of Secanquini, Pittier 248 Field and Catival, Standley 3039 (US); bettveen (US). IZARAI.: betM.een Bananera and La Presa, Corozal and Ancon, Canal Zone, Pittier 2635 in Xlontania del >lice, Steyermark 38074 (US); (US): bettveen Gorgona and Tabernilla, Canal Cristina, Blake 7598 (US); vicinity of Quirigua, Zone, Hitchcock 7963 (.US); between hlindi and Standlej 24017 (NY, US). PETEN:Dolores, km Colom, Canal Zone, Hitchcock 7940 (US); Culebra, NCMRER 59 19

Canal Zone, Hitchcock 7893 (US);Darien Station, Panicurn obtu5zjlorum Richard, 1853:305. [Type: “Crescit ad Canal Zone, Standley 31558 (US); Ft. Kobbe, marginem rivuli Cauta in provincia Santiago de Cuba (Linden 11. 2143).” Holotype, P, not seen; fragment of the near beach, Duke 4181 (US); Juan Mina, Canal type, US sheet no. 80887.1 Zone, Piper 5268 (US): Gatun Lake, Goodyear, Panicum lindenii Grisebach, 1866:233. [Based on Panicurn Seibert 1527 (US);Toro Point, Canal Zone, Hitch- obtuszTorum Richard.] cock 8051 (US): jicinity of Fort Shermann, Canal

Zone, Standley 30997’(US);vicinity of Summit, Robust perennials 0.6-1.2 111 tall. Culms de- Chnal Zone, Standley 30071 (US); Canal Zone, cumbent, rooting and branching at the lower Popenoe 14 (NY). DARIEN:Tuyra river, near the nodes, rarely stoloniferous, geniculate, cylindri- mouth of Rio Yape, Allen 332 (US). PANAMA: .., cal to compressed, pubescent (more so toward near ,2rraijam, IV’oodsonJr. 1379 (NY, US); Chor- the upper part) to glabrous; internodes hollow, rera, Hitchcock 8129 (US): E of Orange river, 5-20 cm long. ,Yodes purplish, compressed, gla- (iormann 2572 (US):Tecumen, Duyer 2558 (US); brous. Sheaths .5.5-18.4 cm long, generally ‘I’ecumen river, Standley 26513 (US). VERAGUAS: shorter than the internodes, papillose-pilose to road between Santiago & David, Stern et al. 1004 glabrescent. Ligule meinbranous, truncate, 0.1- (US); :Inchon Hill, Killip 4196 (US). 0.3 mni long, lvith short hairs on the adaxial PERU. CORONELPORTILLO: environs of Pu- surface toivard the base of the blade; collar dark calipa, 011 Ucayali river, Fosberg 28887 (NY). brown, densely pubescent kvith ivhitish hairs. 1.oRrro: Caballo-Cocha, on the Amazon river, Blades oblong-linear or lanceolate, 10-43 cm Il‘illiams 2205 (US): lquitos, Anderson 1086 (US), long, 0.6-2.3 cm rvide, flat, glaucous, distally ..isplund 13922 (US), IV’illiarns 2964 (US); May- attenuate, papillose-pilose 011 both surfaces to nas, Nanay river, Picuruyacu road below Bellav- glabrescent, the margins scaberulous to papillose- ista, Rimachi 4816 (NY): lower Huallaga river, pilose belolv, the base narroLved, the midnerve It‘illiams 4210 (US): lorver Nanay river, IVilliams prominent. Panicles lax, diffuse, many-flowered, 262, 31 1 (US): Santa Maria, about 25 km below exserted (included or not ivhen young in the Yurimaguas 011 Rio Hiiallaga, Allard 22452 (US); upper leaves), 12-25 cm long, 7-22 ciii wide, \’aguas\.;icii river, Plowman et al. 6798 (US); Yu- the branches spreading, the spikelets separated riniagua, outside of town, Anderson 254 (US), on the branches: axis MY~VYand scaberulous, the Ferrejra 10177 (US). loiver branghes in verticils, the upper alternate \‘EWF.ZUElA. AMAZONAS:Alto Orinoco, Es- and opposite, the axils of the branches densely nieralda savanna, Il‘illiams 15338 (US); El Tem- pubescent; pedicels scaberulous, 2-1 1 iiiiii long. blador, Orinoco river betrveen San Antonio and Spikelets obovoid to ellipsoid, 2.7-3.5 inn1 long, Kirare, Foldats 256‘4 (US); Guainia river, just E 1 - 1.6 miii \vide, greenish to olivaceous, viscous of Xlaroa, ‘Llaguire et al. 41703 (US).TACHIRA: at maturity, the gliimes rvhitisli along the mar- Gi-aiija del FAado, San Cristobal, Tamayo 2294 gins. Lozcer glume acute to obtuse, 2.4-3.1 mni (LTS). TRU-JILLO:]/4 mile from La Ceiba, Reed 932 long, 5-9-nerved, as long as or longer than the (US). upper glunie, glabrous. Upper glume obtuse, 2.4- ~:HROMOSOSIENUMRERS.-~~ = 20 (Pohl and 2.8 iiim long, 7-9-nerved, usually not enclosing Davidse, 197 1): 211 = 22 (Gould and Soderstrom, the upper antheciuni, glabrous. Lower lemma 1970): = 10 (Davidse and Pohl, 1978). n glume-like, acute, 2.6-3.3 miii long, .5-7-nerved, Homolepis glutinosa (Swartz) Zuloaga & slightly to markedly scabrous at the apex, cov- Soderstrom, new combination ered ivith oblong bicellular microhairs, lvith the FIGURE7 basal cell short and the upper cell elongated, 43- Panicum glutinmum Sxvart7, 1788:24. [Type: “Jamaica.” 80 yni long, 11-22 ym wide. Lower palea ovate- Holotype, S. not seen; fragnients of the type, BAA, US lanceolate, smaller than the lemina, 1.9-2.4 iiiiii sheet no. 80678.1 long, 0.4-0..3 miii wide, membranous, glabrous; 20 SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE7.-Homolepis glutinosa, new combination: a, upper anthecium, lemma side (X 100); b, detail of a shotving dense aggregation of prickle hairs and bicellular microhairs (X 200); c, upper anthecium, palea side, showing the same as in b (X 200 ); d, detail of upper palea showing suborbicular silica cells (X 1000). All photomicrographs based on Eiten 3183. NUMBER 59 21 inale flo~verabsent. C,-pper anthecium ellipsoid, (F, h40, NY, US); \.’icosa,Bailej 1011 (US),Kuhl- 2.5-3.3 nini long, 1-1.5 nini \vide, smooth, with mann s.n., 9 Xpr 193.5 (US). PARANA: without iiunierous hooks and bicellular microhairs at the locality, Dusen 3828 (US); Acungui, Mattos 3 apex, shining, pale and Lvhitish, covered through- Labouriau s.n., 1 Mar 1948 (US); Carijo, Hatsch- out by silica cells; anthers light broivn. Caryopsis bach 3040 (US); Curitiba, Swallen 8512 (US); rounded, 1 .7-2 iiiiii long, 1.3-1.8 mm \\ride, Doce Fino, Hatschbach 401 1 (US); Fazenda Re- pale; hiluni linear, someivhat broad, half as long serva, 85 km SW of Guarapuava, Lindeman et al. ;IS the car!.opsis or ;I little longer; embryo half 4692 (NY, US); Guarapuava, Swallen 8822 (US); the length of the caryopsis. Ponta Grossa, Swallen 8357 (US); Rio Espin- DISTRIRUTION.-~I~forests of Mexico, Central garda, Hatschbach 4401 (US); Jaguariahyva, Du- llnierica, the M’est Indies, and South Anierica to sen 1671 7 (MO, LS); Tres Barras, Dusen 17551 Pai-agii;iy and :1rgentina; from sea level to 2500 (NY); Vossoreoca, Kummrov 318 (NY); Rio La- ni elevation. geado, Hatschbach 13745 (US); Rio Passa Dois, r~DDITTONAL. SPECIXlENS SEEN.-ARGES- Hatschbach 15990 (US); Fazenda Lagoa, Hatsch- TIN,,l. (:oRRIt:Nms: arroyo Chimiray, Krapov- bach 14259 (US). Rro GRANDEDO SUL: Porto ickas et al. 25199 (CTES); arroyo Santa Isabel y Alegre, Facultade de Agronomia, Valls 2005 ruta 12, Schinini 3 Quarin 8495 (CTES);deseni- (US); Quinta near Rio Grande do Sul, Malme ImcaduI-a ctel arroyo Garape en el Rioo Parani, 1574 (US); Azevedo, near Cai, Rambo 41438 4.5 kni k: de Ituzaingo, Schinini et al. 11089 (US); Estacao Guianuba, Rambo 40548 (US); Ja- (CTES); Estancia Santa Teresa, Pedersen 2645 car;, Rambo 40950 (US); Morro Agudo, near (US); Kincon (>nibu Chico, Schinini et al. I1260 Porto Alegre, Rambo 41040 (US); Pareci, near ((TES); Santa Ana, Carneuali 2365 (CTES); ruta Montenegro, Rambo 52770 (US); SaoLeopoldo, Leopoldo, 40, 18 kin NE de Santo Tome, Krapovickas et al. Rambo 36987 (US). RIO DE JANEIRO:Alto Ma- 25306 (CTES). ~IISIONES:Bernai-do de Irigoyen, cah;e Glaziou 18647 (F, US); Alta Boa Vista, Burkart 28255 (SI); Canipo i’iera, ,2Jontes 15364 Chase 121?6 (US); Alto da Serra, S of Petropolis, (SI);Fracrh, Parodi 5674 (BAA); Loreto, iVJontes Chase 9791 (US); Petropolis, Chase 12178 (US); 1 I180 (LP); hlontecarlo, Montes 15397 (BAA), Tijuca, Brade s.n., 31 Mar 1929 (US), Smith 2 Porta 203 (BAlI), Schwindt 1329 (US); Parque Brade 2206 (US). SANTACATARINA: 33 kin SE: of IL’,‘ic :ional Iguasui, Cabrera, Gebhard J Corte 201 Cacador, Smith 3 Klein 12189 (NY, US); If kni (LP): Picada San \fartin, Schwarz 4486 (LIL); W of Canipo Alegre, Smith 2 Klein 12035 (NY, Puerto Ylani, Schwarz 10274 (LIL); San Pedro a US); Campo IIlegre, Morro do Iquererim, Reitz Tobuna, Schziindt 4080 (US); San Pedro, Parodi 3 Klein 6389 (US); Canela Gaucha, Sao Miguel 5614 (Bi411);Santa Ana, Rodriguez 77 (MO, US). do Oeste, Klein 5068 (US); Chapeco, Klein 5287 BOLIVIA. State unknown: Hacienda Siniaco, (US); Ilha de Santa Catarina, Morro Itacorubi, road to Tipuani, Buchtien 5316 (F, MO, SI, US); Reitz 4582 (US), Smith 3 Reitz 6166 (US); Itajai, Mapiri, Rusbj 244 (US); San Carlos, Buchtien 23 Praia Braba, Klein 374 (US); Serra dos Ilheus, (BAA. CS). LA PAX:Coroico, De la Sota 454 Smith 3 Klein 16032 (US); 14 kin N of Abelardo (€31114);%ongo valley, Renvoize 3 Cope 4251 Lu7, Smith 3 Klein 11437 (US); Papanduva, Pi- (\lo).SANTA (:RuZ: Buena i’ista, Steinbach 2015 cadas, km 18 1 da ERF, Reitz 3 Klein 12510 (US); (SI), 14952 (US). Pinheiral, 11 km M’ of Cacador, Smith 3 Klein BKAZI I,. M’ithout state and locality, bl’eddell 10911 (NY, US); Rio Cacador, Swallen 8246 1932 (NY). GOIAS:vicinity vicinity of Goiabeira, of Chase Goiabeira, (US). SAo PAULO: Chasei\.ithout locality, Esteri 9916, 11496 (US). MINAS GERAIS:rvithout locality, 9918 (US); Cambucy, Chase I I158 (US); Fazenda Chase 9536 (US), Regnell III. 1370 (US), 12‘idgren Canchim, 8 km NE of Sao Carlos, Eiten et al. 912, 1299, 1845 (US), Campinas, Novaes 1247 318? (MO, US); Estafao Florestal do Paraguaccu (US), Chase 9442 (US); Juiz de Fora, Chase 8570 Paulista, Clayton 4558 (NY, US); Parque do Es- 22 SMITHSONIAN CONTRIBUTIONS TO BOTANY

tado, 1 lim E of Salo Paulo, Fonseca 52 (MO, US); Kipe, ,Llorton 3 Acuna 3208 (US), Shafer 299? near Piedade, 124 km from S. Paulo, Smith 6324 (NY). PINARDEL RIO: Taco-Taco, Sierra de Los (US); Sao Paulo, 10 kni S of center of city, Organos, Morton 4389, 4484 (US). SANTA Sendufsky 339 (US). CLARA:banks of Yayabo river, Leon 3998 (US). COLOMBIA. W’ithout locality, ‘lloritz 759 ECUADOR. ARCHIPIELAGODE COLON: Isla (US). ANTIOQUIA:Hatillo, Daniel 417 (US); Ve- Santa Cruz, above Bella Vista, Wzggins C3 Porter necia, Graiija de las Mercedes, Barkley 3 Gutier- 644 (US); Mt. Crocker, Indefatigable Island, re: 1657 (NY, US). CALDAS:Salento, Killip et al. Howell 9224 (US). ORO:betM.een Portobelo and 876 1 (US). CAECA:Chisquio, Finca 10s Derrum- El Tambo, Hitchcock 21300 (US). SANTIAGO-ZA- bos, Asplund 10561 (US); El Ranial to Rio Sucio, MORA: Macas, Asplund 1920 (NY). M’ of Popavan, Penizef et al. 8090 (US); El Tambo, GUATEMALA. ALTAI’ERAPAan: Cobin,John- Kjell von Sneidern 21 16 (F, US); Popayan, near son 29 (US). BAJA L’ERAP.4Z: Sierra de las hlinas, Tunia, Arbelaez et al. 6326 (US). CHOCO:between 3 km SE of Purulha, IZ’illiams et al. 43168 (F, La Oveja and Quibdo, Archer 1680 (US). CUN- US). HUEHUETENANGO: bet\j.een Barillas and DINAMARCA: Fus>igdsuaa,Pennel 2697 (NY, Us). Cerro I’ictoria, Steyermark 49697 (US); Ixcan and EL I’ALLE:Cordillera Occidental, Pichinde, Cua- Finca San Rafael, Sierra de 10s Cuchumatanes, trecasas 18237 (US); Cordillera Occidental, a1 S Steyermark 49466 (F, US). QUEZALTENANGO:Re- de Las Brisas, Cuatrecasas 22644 (US); La talhuleu, Koninck 236 (US). Cumbre, Pennel 5165 (KY, US); La Manuelita, GUY.4NA. Kamana Falls, ,Ifaguire et al. Palniira, Pennel et al. 6172a (US); Las Nieves, W 45963~(NY); U’aru-Waru creek, Pomeroon, of Cali, Kiffip et al. 39209 (US); Monte Frio, Bartfett 8124 (US). Yanaconas, Killip et al. 33852 (LS). HCILA:San HAITI. Without locality, Ekman 623 (US); Agustin, Daniel 4096 (US). MAGDALENA:Sierra vicinity of Furcy, Leonard 4337 (NY, US); vicinity ISewda de Sta. Marta, Barklej 3 Gutierrez 1874 of Marinelade, Leonard 8212 (US); north\%.ard (US). TOLIMA:La Trinidad, Libano, Pennel3329 from hlorne Jeffrard, Bartlett 17557 (US); (XI’,US); \.icinity of Medellin, Toro 1020 (NY). Source-Rouille, hlassif de la Selle, Ekman 6827 (:OSTA RICA. Without province: Cucasacha, (US); Terre Froid, Holdridge 1760 (US). Lankester 21 1 (US); Helechales del General, Pit- HOh’DUR,4S. EL PARAISO:Yuscaran Yuscaran moun- tier 12002 (US); ALAJEELA:Piedades, near San tain, Molina 613 (US). Ramon, Brenes 4477 (US); Rioo Jeulis de San Ra- JAXlAIC.4. Without locality, Hart 792 (US); mon, Brenes 22890 (NY); La Palma de San Ra- Arntully, Orcutt 2643, 31 10 (US); Cinchona, Har- nion, Brenes 4477 (KY). CARTAGO:2 km NE of ris 11264 (US), 11440, 11508 (MO, US), Hitch- Sari Cristobal, Interseccion Norte, Pohl 13253 cock 9708 (US);Diablo Mt., Ridley 25, 38 (US), (NY); .ikm E of Bajo Pacuare, along road, Pohl Alaxon 3 Kilfip 443 (US); Elvarton to hloneague, 13148 (NY); Alto de la Estrella, Standley 39042 Hitchcock 945 1 (US); Go]-dontown, Hitchcock (US). PCNTARENAS:Canias Gordas, Pittier 11017 93216 (US); near Ipswich, Hitchcock 9627 (US); (US). SANJosE: Santa Maria de Dota, Standley Mason River Savanna, Tt‘ of Kellits, Fosberg 41582 (US). 42773 (US); Neivport, Britton 3193 (US); Red CUBA. U’ithout locality, Wright 757 (NY, US); Hills, St. Andre\\., Harris 11836 (MO, US); road Las Villas, Buenos Aires, Morton 4153 (Us). ORI- to Salt Hill, Harris 11395 (US); vicinity of St. ENTE:Grdn Piedra, Shafer 9014 (NY, Us); s of Helen’s Gap, St. Andrew, ,Llaxon 3 Killip 595 Guaro, Hitchcock 23416 (US); La Bayamesa, crest (US): St. Thomas, Cuna Cum trail, lZfaxon 3 of the Sierra Maestra, lLlorton 9282 (US); La Killip 185 (US); Troy, Harris 12607 (MO, US), Perla, Leon 3790 (NY, US); Rio Yao, W of Sierra Hitchcock 9785 (US), ;Ilaxon 2816 (US). hlaestra, Morton et al. 3489 (US); Sierra Maestra, MEXICO. Il’ithout state: Mirador, Mohr s.n., Alto de Iberia, Ekman 8298 (US); crest of Sierra in 1857 (US). HIDALGO:Zacual[ti]pan, Purpus NUMBER 59 23

2903 (US). CHIAPAS:Turubula, il'elson 3357 Alto de Choroni, Chardon 181 (US); road be- (US). OAXACA:Hiiantla de Jimenez, Schultes et tiveen Choroni and Maracay, Pittier 23923 (US), al. 185 (US); San Ildefonso de ITiIla Alta, Yetze- Soderstrom 976 (US), Tamayo 1636 (US); Parque lala, \'era Santos 3503 (MO, US). ORIZABA:San Nacional, Chase 12456 (US). BOLIVAR:Santa Cristobal, Bourgeau 3192 (US). PCERLA:30 kni Teresita de Kavanayen, Steyermark 60529 (F, N of %ac;ipoaxtla, Koch 7724 (US). VERACRUZ: US). DISTRITOFEDERAL: on the old road to Ca- near Jalapa, Hitchcock s. n. (US); Zacualpin and racas from La Guayra, Pittier 5910 (US); Cerros Imiciiiitv, Purpus 2156 (US). del Avila, Altos de Galipin, Pittier 8301, 9915 I',.ZK:ZGU:ZY. Lt'ithout state: Alto Parana, Na- (US); Cordillera Costanera, Junquito, Chase cunday, Alontes 10949 (US); Asuncion, Jardin 12432 (US): Hacienda Puerto La Cruz, Pittier Bota'inico, Rojas 14546 (BAA); Caacupe, Insfran 8060 (US). IARA: betrveen La Escalera y Cubiro, 1087 (SI);Carapegua,Rojas Rojas 3400 (US); Carobeni Burandt jr et al. 0032V (US);near Sanare, Parque Suevo, ixxid to Itape, Rosengurtt 5896 (LP); Isla Yacamb;i, Smith 132917 (US). MIRANDA:La Prov- ?'ei.esa, Bertoni 4294 (LIL); Lago Ipacaray, Has- idencia, Pittier 23762 (US); road froni Los Can- sler 22544 (F,110, US); Pedro Juan Caballero, ales to El Encanto, Lasser 660 (US); hills above Rojas 6690 (BAA); Pirareta, Schinini 12183 I,os Teques, Pittier 21247 (US). TACHIRA:La La (CTES): :Zpa Ki\.ei., Hassler 11449 (US); Sapu- hlulera, Tamayo 2302 (US). cay, Hassler 13027 (US); Tacuru Pucu, Fiebrig CHROMOSOMESUMRERS.--~I~ = 'LO (Davidse 6502 (BX;\); Puerto Bertoni, Rojas 8070 (BAA). and Pohl, 197'La,b, 1974, 1978), 211 = 40 (Pohl PERU. AMAZONAS:1-5 kni SE of Yambras- and Ilajidse, 197 1). bamba, ll'urdack 1033 (US). HUAUCO:Pampa Pampa DrscvssIoN.-This species u'as not placed in Hermosa, between Puente Durand 8c Exito, any group b). Hitchcock and Chase in their pub- ,\.lexia 824? (F, MO, US); Pampayacu, mouth of lication of 19 10. 111 an unpublished manuscript Rio Chinchao, ,blurbride 5101 (US). JUNIN:Pichis Pichis Chase later included it in the group Glutinosa, trail, Yapas, Killip & Smith 25389 (F, US). SAN along ivith P. millegrana and P. selloxlii, taxi to MARTIN:Juan Juan Guerra-Mayo River, Anderson ivhich ive do not believe it to be related. 1096 (US); road between Tarnpoto and Juan Homolepis glutinosa and H. villaricensis were Guerra, Anderson 1108 (US). included in section Glutinosa of Panicum (Zu- I'UEKTO KICO. Alta de Bandera, E of Ad- loaga, 1978) on the basis of glume length and by juntas. O'hase 6469 (US); Guavate, Liogier 9771 the type of pubescence in the spikelet. In both (US); Alaricao Insular forest, Schubert 337 (US); species the lolver and upper glunies reach about hlai-icao, wad to Indiei-a Fria, Chase s.n., 1913- the same sire and developnient and are indistin- Oct-23 (US); \.icinity of hlaricao, Chase 6199, guishable niorphologically; frequently the loiver 6250 (US); Mayaguei., Sintenis 357 (US); Sierra glurne exceeds the upper in length. In relation de Yabucos, Sintenis 2609 (US). to the spikelet the length of the glumes is vari- DO\ll~I~~,~ZSREPUBLIC. AZCA:Cordillera able, in certain cases enclosing the upper anthe- (:enti.al, Sail Juan, Ekman 13431 (US). LA \.r~~~:ciuni entirely, in others leaving the apex exposed vicinity of Constanza, Allard 16451, 16488 (US); (especially on the dorsal face of the spikelet, as Constanza, near Rio Grande, Jimenez 2129 (US); the upper glume is smaller). In young spikelets vicinity of Jarabacoa, Allard 14886, 14909 (US); both glumes frequently enclose the anthecia en- 1,a Cienaga, south bank of Ri o Los Guanos, tirely. In this state the spikelet is compressed Gastony et al. 220 (US); Lonia Canipana, Allard dorsiventrally and its form is ellipsoid. U'hen the 28216Bahia.“ Holotype, P, not seen; fragment of the the type, covered ivith evenly distributed silica cells; US sheet no. 2903503.1 lemma .?-nerved. Carjopsis obovoid, 2.5 mm NUMBER 59 25

FIGURE8.--Homolepis isocalycia: a, upper anthecium, lemma side (X 100); b, detail of upper palea showing suborbicular silica cells (X 500); c, detail of lower lemma showing bicellular microhairs and prickle hairs (X 430); d, detail of upper anthecium, palea side, showing bicellular niicrohair and silica cells (X 1000). All photomicrographs based on Duarte 6092 (US). 26 SMITHSONIAN CONTRIBUTIONS TO BOTANY

long, 1.3 iiim !vide, bi-olvn; Iiilum linear, as long plateau of Serra Tepequem, Prance et al. 4482 ;IS the caryopsis, embryo ‘h as long as the caryop- (410). sis. COLOMBIA. ANTIOQUIA: Cisneros, Toro DISTRIBUTION.--~I~moist and shaded wood- 1027 (NY, US). SANTANDER:Mesa 10s Santos, S lands or forests from Mexico and Panama to of Bucaramanga, Robinson Jr. 3181, 3195 (US). iiorthern Brazil. FRENCH GUIANA. Road Iracoubo-Organ- IZDDITIONAI. SPECIMENS SEEN.-BRAZIL. abo, kni :313, Black 3 *Muller 54-172393 (US); BAHIA: near Bahia, Salzmann s.n. (US); Bahia, route de L‘ilcarouany, Hoock s.n., 1 1 hov 1955 Dorsett 3 Popenoe 4446 (US); de Brejao a Mu- (NU). cuge, Lemos Fro es 19982 (US); Parafuso, 39 km GUYAS,2. Il‘ithout locality, Jenman 6000 NE: of Bahia. Chase 7976 (MO, US); Campo Santo (US): Kamakusa, upper Mazaruni river, De la to Bahia, Chase 8048 (MO, US); 40 kni S of Cruz 2251, 421 7 (110, NY, US); Wanalva river, Eunapolis, Porto Seguro, Arruda 1 (MO); 5 km De la Cruz 4057(NY,US); Bartica, Lengs.n. (NY, SE: of Feira de Santana, Davidse et al. I1702 US): hiazaruni river, Bartlett 8059 (US); Kaieteur (h1O); Serra do Rio de Contas, 11 km S of Rio Fall, forest along left bank of Potaro river, Cowan de Contas on side road to LV of the road to 3 Soderstrom 2002 (NY, US); vicinity of Penal I,i\wmento do Brunlado, Harley et al. 15077 settlement, on ryest side of Essequibo river, near (US), 12-14 km K of toivn of Rio de Contas on mouth of 1lazaruni river, Hitchcock 17112 (NY, the road to Slato Grosso, Harlej et al. 15179 US); Il’aranar\.a Ranch, -3 mi from Jaca\va, Har- (VS): Serra do Sincorai, 9 km SIZ’ of Mucuge. on rison 1097 (US): Kaiken, Paruima, Leakley 227 I-oadfrom Cascavel by the Rio Paraguacu, Harley (NY); Kaieteui- Plateau, .Iiaguire 3 Fanshawe et al. 15977 (US): 17.5 km N of Ubaitaba, near 23287 (NY): Xl:izai.uni river, Kalacoon, Archer Sao ;lntonio de Jesus, on the BA 101 highr\.ay to 2479 (US): Kartabo Point, at confluence of Cu- Feira de Santaiia, Harley et al. 17033 (US), 5.6 yuni and hlazaruni rivers, Sandwith 644 (US). Liii N of SI~icug&on road to Andarai, Harley et PilNrl1l~l.1.5 miles S of Goofy Lake, Dzyer al. I8880 (US), -10 km N of Barra de Estiva, on 8027 (S10); Campo Tres, 3 mi NE Altos de Ibicoara road, by the Rio Preto, Harley et al. Pacora, Liesner 528 (MO): Cerro Azul, Duyer s. n. 15862 (US), -6 km N of Barra da Estiva on (MO), Ebinger 405 (LIO), Godfrej 2047 (NY), Ibicoara road, Harley et al. 15543 (US); south Tjson 2132 (510): Cerro Jefe, Antonio 4701 part of peninsula, vicinity of Bahia, Chase 7858 (h1O), Davidse 101 I6 (MO),Hammel 4814 (h1O); (110, US): Marau , a 3 km do Porto, Mori et al. road to <;erro Jefe, 21 km N of Panamerican 11390 (hlO); hlarau-Ubai taba, 45-50 km E of Highway, Folsom 3 Hartman 4620 (MO); road Lbaitaba, iIlori et al. 11950 ($10); 34 krn E of to Cerro Jefe, shore of Goofy Lake, Tl’itherspoon Moi-ro do Chapeu,Davidse et al. e t al. 11870 (MO);8439 (MO); road to Ceri-o Jefe, beyond Goofy Porto Seguro, BR 5, km 18, Duarte 6092 (US); Lake along road, Correa 3 Dressler 464 (MO); 7’. de Fi-eitas, 12 kin r\; of Cachoeira, Chase 8076 top of Cei-roJefe, Tyon et al. 441 1 (MO). (US); Cruz das iZliiias, Pinto 124 (US); Serra da SURIKAM. Kayser airstrip, 45 km above con- Agua de Rega, 27 km N of Seabra, Irwin et al. fluence with Lucie River, Irwin et al. 55980 (310, 31010 (SY).CEARA: oCamp Grande, Swallen NY, US); Kaysei- airstrip, 25 km above conflu- 4518 (US). ~IINASGERAIS: 10-15 kni da cidade, ence Ivith Luck River, ,\laguzre et al. 53956 (MO, I’llagalhaies 14143 (US); 17 kni SW of the Minas hY, US); Sipalwini savanna, area on Brazilian Gel-ais-Bahia border, along highway, Davidse et frontier, Oldenburger et al. 355 (NY). al. 11615 (NY). PARA: Alto Quim ina, Sampaio \‘ENE%UELA. BOLIVAR:Hato Divina Pastora, 5784 (US); Igarape. das Borboledas, Sampaio Gran Sabana, a1 pie del cerro Ceita, Tamajo 2918 5580 (US); Rio Para, Sampaio 5516 (US). PER- (US); Icabaru-Santa Elena, 70 km NE of Icabaru, NAMBUCO: Olinda, Pickel 108 (US). RORAIMA: Croat 54228 (MO): Santa Elena, Tamayo 2780 NUMBER 59 27

(US). TACHIoadroad from from Palmira Palmira to Mulera, to Mulera,Cespitose perennials, shortly rhizoinatous, 40- ,Vfuller 889 (US). TRUJILLO:near Escuque, along 90 cm tall. Culms few-noded, cylindrical, gla- road to \’alera, Pittier 13141 (MO, NY, US). brous, erect, not branching, covered amply at DrscvssroN.-Although the species appears to the base by the sheaths; internodes hollow, 6-26 be iiiost common in northern South America, cm long. ‘Vodes obscure, compressed, glabrous. the type specimen of one of its synonyms, Pani- Sheaths 6-2 1 cm long, the basal ones abbreviated, cum langei, conies from Mexico. Curiously, we shorter than the lower internodes, the upper have no specimens from Central America, except ones longer, pale, keeled, glabrous or papillose- recent collections from Panama. Steudel cited pilose to\<.ard the upper margins. Ligule a short only “Paraguay. Bahia” for another synonym of arc, ciliate, formed by a line of \\.hitish hairs 0.3- this species, Panicum renggeri. Given the geo- 0.6 mm long; collar glabrous or densely pubes- graphical distribution of H. isocalyia, we may cent. Blades linear-lanceolate, subulate to acute, assiiiiie there is an error in the labeling of the 6-20 ciii long, 0.2-0.6 cni wide, rigid, narrow at type material and that it was collected in the state the base, not clearly differentiated from the of Bahia, Brazil. sheath, pubescent on both sides (more densely so 111 spikelet shape, pubescence, and consistency on the abaxial side) to glabrous, the margins of the upper- arithecium, this species appears to convolute, ciliate nith thin, fragile hairs, inid- be niost closely related to H. glutinosa and H. nerve prominent. Panicles contracted, exsei-ted, villaricensis. The upper anthecium in these three 6-20 cm long, 1-7 cm wide, the branches ap- species is sonielvhat more rigid than in H. longi- pressed to the axis, the spikelets borne on long spicula and H. aturensis. pedicels; axis cylindrical, undulate above, gla- The lilbel dmof Irwin 55980 indicate that the brous, smooth, purplish, the branches alternate culiiis of this species attain heights of 2.3 ni; data or subopposite, smooth, glabrous, purplish, the fi-om Chase 8076 indicate the plant has stolons pedicels claviform, 3-2.3 inn1 long, glabrous, reaching 3 iii 01-more in length. In this species smooth. Spikelets fusiform, 9- 1 1 inm long, 2-2.9 there is great wi-iation in the size of the plants, nim \t.ide, acuniinate, greenish-violet, herba- paiiicles, and spikelets. The spikelets, for exani- ceous, glunies and anthecia separated by distinct ple, range froin 4.6 to 7.4 mm in length, a internodes. Louler glume acute, 9- 10 iiiiii long, \,ariation that does not correlate with other char- greenish, .i-.l-nerved, glabrous or sparsely hir- acters. There are examples of small plants Lvith sute lzith caducous hairs. Cpper glume acuminate, spikelets 7 iiim or more long and robust plants 8.2- 10 iiim long, .?-nerved, pubescent toiz,ard \\.itli siiiallei. spikelets. For this reason kve con- the margins, the hairs \vhitish and violet, the sider it prudelit to maintain the material exam- margins hyaline, the middle part hirsute, it,ith ined under ;I single species, H. isocalyia. caducous tuberculate hairs. Lower lemma -8 inn1 long, 3-7-nerved, densely pubescent (especially HomoZepis longispicula (Doell) Chase tolvard the niargins), the long hairs whitish and purplish with thick tubei-culate hairs on the mid- FIGCRE9 dle part (more so to\zard the apex and margins) Hornolepis longispicula (Doell) Chase, 191 1 : 147. and oblong, hicellular microhairs. Lower palea Panicum longtflorum Trinius, 1834:317, not Ginelin, 1796. conspicuous, lanceolate, 6-7 niiii long, 1 - 1 ..i [Tvpe: “Habitat in Brasilia, loco accuratis non adnotato mm \\.ide, as long as the lemma, hyaline, trans- (herb. Acad. Petropolit.). Holotype, LE, not seen; frag- parent, glabrous or pilose toizard the upper mar- ment of the liolotype, US sheet no. 80743.1 gins; male flou.er present lvith 3 stamens. C’pper Panicum longispirulurn Doell in Martius, 1877:26 1. [Type: the same :is that of Panicum longzjlorurn Trinius.] anthecium fusifoim, 6.7-7.8 iiiiii long, 1.4-2 iiim Ichnanthus longij7orus (Trinius) Bentham, 188 1: 145. ivide, herbaceous, rvhitish, glabrous or pubescent Hornolepis longzyora (Trinius) Pilger, 1940:35. tolzard the upper- part, densely covered with 28 SMITHSONIAN CONTRIBUTIONS TO BOTANY

FICCREF).-Homolepis longispicula: a, upper anthecium, palea side, showing flat margins of lemnia over palea and macrohairs on the lemma (X 100); b, lower lemma showing oblong bicellular niicrohair with collapsed upper cell (X 1400); c, upper palea showing suborbicular silica cells (X 1000); d, lower lemma with thick and thin macrohairs (X 80). Photomicrograph a based on Chase 10349 (US), b and d on Hatschbach 27965 (CS), and c on a fragment of the holotype (US). NUMBER 59 29 silica cells throughout, the hyaline margins of the Stoloniferous perennials 15-40 cm tall. Culms lemma enclosing almost entirely the palea. Car- cylindrical or compressed, geniculate, branching yopsis oblong, 2.9 mni long, l .5 mm wide, brown; at the lower nodes, glabrous; internodes hollow, hilum linear, as long as the caryopsis; embryo 1.5-1 1 cm long. Nodes glabrous, obscure. Sheaths less than half as long as the caryopsis. shorter than the internodes, 2-8 cm long, the DISTRIBUTION.-~n sandy soils of dry and margins papillose-pilose to glabrescent. Lzgule open areas of the Brazilian cerrado in the states membranous, laciniate, 0.4-0.8 mm long; collar of GoiaS, Distrito Federal, and Minas Gerais, dark brown, glabrous to pilose. Blades oblong- between 1000 and 1500 m elevation. Plants of lanceolate, acuminate, 4-1 2 cm long, 0.6-0.9 cm this species are exposed to frequent fires as evi- wide, flat, narrowed at the base, the margins denced bj, the large number of herbarium spec- scaberulous, the lower with caducous tuberculate imens with burnt bases. hairs, strigose on both surfaces, the midnerve ADDITIONALSPECIMENS SEEN.-BRAZIL. prominent. Panicles lax, diffuse, 5-1 1 cm long, DISTRITOFEDERAL: 10 km S of Brasilia on road 1.5-5.5 cm wide, the axis scabrous, the lower to Belo Horizonte, Irwin et al. 8591 (MO, NY, branches alternate or opposite, the remaining US). GorAs: 2 kni NW of Veadeiros, road to ones alternate, the axils finely pubescent, pedi- Cavalcante, Irwin et al. 9489, 9491 (NY, US); cels scabrous, 3- 18 mm long. Spikelets ellipsoid vallev of Rio Corumba, Glaziou 22470 (US). to obovoid, 3.2-3.7 mm long, 1.2-1.5 mm wide, MINAS GERAIS:Diamantina, summit of Serra de glabrous, dark green to olivaceous, viscous at Sao Antonio, Chase 10349 (MO, NY, US); 5-10 maturity, the glumes usually enclosing the lower km W of Diairrantina, serra de Sao Antonio, lemma and upper anthecium. Lower glume acute, Chase 10426 (US); Serra do Espinhaco, Hatsch- 3.1-3.4 mm long, as long as or longer than the bach 27809 (NY, US); Serra do Cipo, along the upper glume, 5-9-nerved, glabrous. Upper glume road at km 134-135, Eden & Eiten 6750 (MO, acute, 2.9-3.4 mm long, 7-9-nerved, glabrous. US), 6754 (ISY, US); road BR 259, Hatschbach Lower lemma glume-like, acute, 2.8-3.3 mm long, 27965 (US); Serra do Cipo, 110 km NE of Belo 5-7-nerved, scabrous at the apex, the remainder Horizonte, Chase 921 I (MO, US). of the surface densely covered with bi-, tri-, or DrscussroN.-This species is clearly separable (rarely) quadricellular microhairs, globose, obo- from others in the genus by its habit and habitat. void, 40-70 pm long, 19-30 pm wide. Lower The plants are cespitose with erect culms and palea ovate-lanceolate, 1.9-2.4 mm long, 0.4- densely covered at the base by hard, keeled 0.7 mm wide, membranous, hyaline, glabrous; sheaths that are resistant to fires. The blades are male flower absent. Upper antheczum ellipsoid, lanceolate, involute, rigid, and without a sharp 2.8-3.2 mm long, 1.1-1.2 mm wide, smooth, differentiation between the sheath and the blade, shining, pale, covered with evenly distributed and the ligule is ciliate, unlike that of the remain- dumbbell-shaped silica cells, the apex of the ing species. The spikelet differs most notably in lemma with numerous hooks and dense bicellular its conspicuous lower palea, which frequently microhairs. Caryopsis obovoid, 1.8-2 nim long, encloses a well-developed male flower. In the 1.4-1.8 mm wide; hilum linear, half as long as remaining characters the spikelet conforms to the caryopsis or a little longer; embryo half the the type common in the genus. length of the caryopsis. DISTRIBUTION.-southern Brazil, Paraguay, Hornolepis villaricensis (Mez) Zuloaga 8c and northwestern Argentina, in shaded and hu- Soderstrom, new combination mid habitats where it forms dense clumps be- FIGURE10 neath trees. Panicurn uillaricense Mez, 192 1 : 1. [Type: “Paraguay, Balansa ADDITIONAL SPECIMENS SEEN.-ARGEN- n. 2.” Isotype, BAF; fragments of the type, BAA, US sheet TINA. CORRIENTES:Garape s tream,mouth of no. 2808940.1 Parana River, 45 km E of Ituzaingo, Schinini et 30 SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE10.-Hornolepis uillaricensis, new combination: a, upper anthecium, lemma side (X 100); b, detail of a showing prickle hairs, bicellular microhairs, and suborbicular silica cells (X 300); c, upper anthecium, palea side, showing macrohairs, bicellular microhairs, and suborbicular silica cells (X 200); d, lower lemma, showing glandular microhairs, mostly collapsed (X 140). Photomicrographs a-c based on Rojas 3410 (US), and d on Schwindt 3266 (US). NUMBER 59 31 al. 11096 (CTES), Quarin et al. 3375 (CTES); The size of the scars varies among the species Estancia Santa Teresa, Pedersen 35 (US); Loreto, in Foveolatus, from prominent as in Zchnanthus Ahumada @ Giberti 67 (SI, US). MISIONES:Lor- pallens to hardly apparent in I. camporum, I. eto, Gruner 764 (BAA), Novat 78 (BAB); San minarum, and 1. procurrens. Stieber pointed out Pedro, Parodi 5609 (BAA, US); Yabebyri, Montes that the species in this section are further char- 747 (LP); San Ignacio, Giambiagio s.n., 1914 (SI acterized by a dense aggregation of silica cells at 13499), Quiroga s.n., 3 Apr 1914 (NY); San the apex of the upper lemma. Antonio, Cabrera, Gebhard €3 Corte 108 (BAA); Our study of Panicum grandifolium has shown Campo Grande, Montes I5357 (SI, US); Salto that a pair of diminutive scars is present at the Encantado, Schmidt 4555 (LIL); ruta 14, km 304, base of the upper lemma, suggesting an alliance Schwindt 3266 (LIL, US); Teyucuare, Montes with species of Foveolatus. An SEM analysis of 2110 (US). the upper anthecium has also revealed the pres- BRAZIL. PARANA: Guarapuava, Swallen 8909 ence of numerous silica cells at the summit of the (US). upper palea and lemma, densely aggregated on PARAGUAY. Villa Elisa, Pedersen 5902 (Ped- the palea, sparser and more evenly arranged on ersen, US); Villarica, Jorgensen 4094 (BAA, F, the lemma. Additional characters that indicate a US), Ramirez 828 (BAA); Carobeni Nuevo, Ro- relationship to Zchnanthus include the keeled con- sengurtt 85895 (F, US); Carobeni Nuevo-Itape, dition of the glumes and lateral compression of Ramirez 676 (BAA); Ipacaray Lake, Hassler the spikelets. Because this species is so unlike any 12469 (US), 12537 (MO); Alto Parana, niacun- other within Panicum, instead sharing many char- day, ,Llontes 10906 (US); Santiago, Estancia La acters with Zchnanthus, it seems appropriate to Soledad, Pedersen 5948 (Pedersen, US); Alto Par- include it within this latter genus. Interestingly, aguay, Priniavera, Woolston 69 (SI); Asuncoh, Stieber (1982) removed Zchnanthus nervosus from Jardin Boakico, Rojas 12565 (BAA); Colonia the genus and made it synonymous with Panicum Independencia, Rosengurtt 59?9 (BAA); Carape- grandifolium Doell. gua, Rojas 3410 (BL4A,US); Caaguazu, Hassler 8919 (BAA, NY); Puerto Bertoni, Alto Parana, Zchnanthus grandifolius (Doell) Zuloaga 8c Rojas 7951 (BAA),Bertoni 3644 (US), 4701 (US); Soderstrom, new combination Tobati, Cerro Tobati, Arenas I145 (CTES), Has- sler 500 (NY). FIGURE11 DIscussroN.-Parodi (1925) considered this Panicum grandijohm Doell in Martius, 1877:195. [Type: species to be conspecific with Panicum glutinosum, “Habitat in sylvis ad Itahypie fluviurn et Carnacorum vicum believing that the smaller size of the plants was S. Pedro d’Alcantara provinciae Bahiensis (Martius).” the result of logging and of trampling by cattle. Holotype, M, not seen; fragment of the type, US.] Ichnanthus neruosus Swallen, 1964:147. [Type: “Brazil, Per- narnbuco, Tapera, in a woods, 24 Apr. 1935,Pickel 3855.” Zchnanthus Holotype, US sheet no. 164517 12.1 Stieber, in his revision of Zchnanthus (1982), Cespitose perennials, 60-200 cni tall. Culms divided the genus into two sections, Zchnanthus ascending, cylindrical to compressed, simple, and Foveolatus, including in the first those species short-pubescent below the nodes, otherwise gla- M ith prominent appendages attached to the base brous; internodes solid, short at the base, becom- of the upper lemma and in the second those with ing longer above, 7- 18 cm long. iVodes obscure, scars at the base of the upper lemma. He noted glabrous to short-pubescent. Sheaths 11-13 cm that other authors (Zuloaga, 1981; Shaw and long, the lower ones longer than the internodes, Webster, 1981) had demonstrated a common the upper ones shorter, villous at the base and origin for appendages and scars. around the collar, the remaining sparsely pubes- 32 SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE 1 1 .-Ichnanthus grandijolius, new combination: a, upper anthecium, lemma side (X 100); b, detail of a showing silica cells and stoma (X 2000); c, upper anthecium, lemma side (X 500); d, detail of upper palea (X 500). All photomicrographs based on Pickel 1629 (US). NUMBER 59 33 cent toglabrous. Ligule nienibranous-ciliate, 0.7- sides of the leniiiia. (;arjopsis not seen. 1.1 iiim long, \\-it11 loiig hairs adaxially toward DIsTRIRvTIo~.-~CcUl~rillgfrolii sea level to tlie base of the blade: collar dark broivii, glabrous 300 111 elevatioii in eastern Brazil-from Per- to pubescent. Blades lanceolate, acuminate, 22- mnibuco to Espii-ito Santo-in riioist places 1 ciii long, 3..i-6 ciii \vide, flat, pseudopetiolate, ivitliin or at tlie edges of 12.oodlands or forests. \\.it11 ;in asyniiiietric apes and base, the lower A D D ITION A I. SPECIMENSS I.: EN. -B K A Z I L . 111 argi ii s g la t1i.o 11s to Ion g-p u bescen t , with thin , BAHIA:2323 km [El of Ibirataia, on side road iion-tuhet.culate liaii-s, otherwise glabrous, the connecting Ivith BK-101, Fanenda Macedonia, iiiidnewe proniiiient; pseudopetiole 0.3-2 cni Calderdn et al. 2362 (US); Fazenda Uruguayaiia long, short-pilose to glabrous. Panicles lax, and vicinitv, 8 kiii S of Saiita Cru%da i'itoria, essei-red, diffuse, pyi-aniidal, many-floivered, Soderstrom et al. 2109 (US): 6 kni E of Ubaira 1.5-.50 ciii loiig, 5-1 4 cm \\.ide; axis angular, city: Rio Jequirica, Calderon 3 Pinheiro 2249 scabrous to liii-sute, the branches alternate to (US); rodo\.ia Itabuna-Urucuca,Belem 3 Aguiar subopposite, diverging, triquetrous, the axils 1248 (US). EsPIRI~roSANTO: Keserva Biologica long-pilose, the spikelets in pairs and appressed de Sooretania, Soderstrom 3 Sucre 1894 (US). along both sides of the branches, one subsessile, PERNAMRUCO:Saziii-e. da llata, .Lloraes 1103 the other shortly pedicellate; pedicels scabrous (US): Tapera, Pickel 1629, 1928, 2347, ?543, to hirsute, claviform. Spikelets long-ellipsoid, 4- 3545 (US). 4..i inm long, 1-1.4 mni wide, greenish, shining, Drscvssrow.-The sheaths of one specimen, laterally compressed, pubescent only at the apex Soderstrom 3 Sucre 1894, are distinctive in their- of the lorver glume, otherwise glabrous, glumes purplish mottling. and IoM-ei- lemma rigid and with prominent nerves; upper glume and lower lemma subequal Taxonomic Position of Panicum aristellum in length. Lowr glume ovate-lanceolate, acute, In an uriedited key to South Aiiierican species 2.6-3.4 mm long, 2/i to V4 as long as the spikelet, of Panicum, Chase considered P. aristellum to its margins broadly enclosing the upper glume, have little I-elationship to any other species of the 3-nerved, the midnerve scabrous toward the geniis, because of the presence of ;i\viis on both apex and M.ith thick long whitish hairs, scabrous glumes. She placed it next to the only other at the apex of the inner surface. Upper glume species Ivith aivned glumes, Panicum najadum acute, 3.7-4.4 mni long, 5-nerved, both mid- Hackel, Lvliich w.;is later transferred to Oplismen- nerve and inner surface scabrous. Lower lemma opsis najada bv Parodi. In fact, Chase later glume-like, acute, 3.6-4.3 mm long, 5-nerved, agreed Fvith other authors in regarding Panicum scabrous on the inner surface, completely em- as a genus i\.itli unarvned glumes. :Ii\.ned glunies bracing the upper anthecium at the base. Lower are, ho\vever,characteristic of several other pan- palea lanceolate, 3.1-3.6 mm long, 0.6-1 mm icoid genera, such as Chaetium, Echinochloa, and wide, membranous, pubescent along the edges, Oplismenus. otherwise glabrous; male flower usually present The thin and membranous uppei- antheciuin with 3 stamens, the anthers 1.6 mm long. Upper of P. aristellum resembles that of Hqmenachne anthecium fusiform, 3.3-3.7 nim long, 1 mm (Figure 13), lvliicti rvas described by Beauvois in wide, laterally compressed, smooth, shining, 18 12 in the follo\ving iiiaiiiier- (as translated from membranous, the lemma enclosing the palea with the Latin): its i\.ide margins, the lemma and palea with numerous silica cells and stomata at the apex; silica Panicle simple, spiciform: branches dense. - Glunies cells rounded, lvith 2-4 sinuses, more or less unequal, herbaceous, acute: the lower much shorter than the upper. Lower anthecium sterile: Lower lemma acute, e\.enly distributed over tlie upper lemma and the palea very short, membranous, hyaline. Upper anthe- densely so oii tlie apex of the palea; small scars cium hermaphrodite: the lemma and palea membranous, present ;it tlie base of the antheciurn on both acute. 34 SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE12.-Panicum species: a, upper anthecium of P. vaseyanum, palea side (X 100);b, upper anthecium of P. quadriglume, lemma side (X 200);c, base of upper anthecium of P. hispidifoliurn (X 100); d, upper anthecium of P. difffusum, palea side (X 200). Photomicrograph a based on Pringle 1415 (US), b on Chase 8897 (US), con Valerio Rodriguez 1981, and d on Eggers 76 (CS). NUMBER 59 35

FIGVRElJ.-Hymenachne species: a, upper anthecium of H. hemitoma showing the thin lemma and palea (X 200); b, detail of upper anthecium, palea side, of a showing bicellular microhairs and silica cells (X 1000); c, detail of upper anthecium of H. amplexicaulis showing prickle hairs (X 1000); d, upper antheciuni of H. donacijolia (X 100). Photomicrographs a and b based on Tracy 8847 (US), c on Rosengurtt 6175 (US), and d on Ule 88? (US). SMITHSONIAN CONTRIBUTIONS TO BOTANY

Chase (1908) and later Pilger (1940, 1954) are shared by Hjmenachne, P. aristellum, and the considered the distinguishing character of Hj- above species of Panicum. Panicum pernambucense menachne to be the nieinbranous consistency of occ~irsfrom Bahia to the north of Argentina and the upper leninia and palea, which is the basis of P. grumosum from Santa Catarina to the province the generic iimie (hymen, membranous, plus of Buenos Ail-es, Argentina. ,411 of these species achne, scale). In Panicum the upper antheciuni is are chaixcteristic of ri\.er or lake margins and comnionly indurate Mith the margins of the habitats periodically covered by rvater. lemma inrolled over the palea (Figure 12). Chase Panicum, Panicum aristellum, and Hq'menachne included iii Hymenachne, H. amplexicaulis, a spe- ma\' be distinguished by the characters listed in cies !\.it11 spike-like panicles, and H. donacfolia, a Talle 2. species Ivith long and narrow panicles whose If the consistency of the upper antheciuni is branches sometimes diverge from the axis. accepted as a character of generic importance to Panicum aristellum differs from Hq'menachne differentiate Hjmenachne froni Panicum, then mainl?. iii the laxness of the panicle, the aristate Panicurn aristellum and species of the Laxa group condition of the glumes, and the presence of a ~vouldbe congeneric \lit11 Hymenachne. In order plea and inale florver in the loiver- anthecium. to test this hypothesis, a study should be made of Sonie species of the Laxa group of Panicum (P. all of the species iii Hq'menachne as \\.ell as those pernambucense and P. grumosum) share ivith HJ- of the Laxa group of Panicum. Because r3.e have menachne and P. aristellum a niembranous upper not made such a study l+.eare not transferring P. antheciuiii. In these species the spikelets are aristellum to another genus at this time. shoi-t-pec1icell;iteand foim dense racemes, which Panicum are 11ioi.e01- less divergent from the axis and not much diffei-ent from H. donacfolia or H. conden- Panicum aristellum Doell sata. In Hjmenachne and P. aristellum the glumes FIGURES14, I3 are acuiiiiiiate or aristate, whereas in the other "Panicurn .-\riste//a" Doell in Rlartius, 1877:22 1. [Type: "In species of Panicum cited above, the glumes and provincia hlinaruni a cl. M'idgren lectum (herb. Holm.)." lo~verlernnia are acute. The palea of the lower Type, S, not seen; isotype, US sheet no. 702302; fragment of the tvpe, US sheet no. 80468.1 floret is usually absent in P. pernambucense but is present in P. grumosum. Cespitose or shortly rhizoinatous perennials, A similar habitat and geographical distribution 1-2 111 tall. (;ulms erect, many-noded, simple,

TARLL:2.--Comparison of Panicurn, Panzcum arzstellum, and Hyrnenachne.

Panicum Character I Panicum aristellum Hymenachne Upper anthecium indurate or membra- membranous, lemma membranous, lemma nous, lemma cover- covering the palea not covering the pa- ing the palea at the at the apex lea at the apex apex Lower anthecium palea present or ab- palea and male flower palea and male flower sent and sometimes present absent a male flower Glumes I and I1 obtuse, acute or acu- awned, the awns of acuminate, not aivned minate. not awned equal length to the body of the glumes Inflorescence panicle lax to con- panicle lax, diffuse panicle spike-like to tracted contracted NUMBER 59 37

FIGURE14.--Panicum aristellum: a, habit (X 1); b, panicle (X 1); c, branch of the panicle (X 12); d, ligule (X 3); e, scheme of sheath in cross section;f; lower glurne (X 25);g, upper glume (X 25); h, upper anthecium, leninia side (X 25); i, upper anthecium, lateral view (X 25);j, upper antheciuni, palea side (X 25);k, stamens (X 25);1, upper palea with gyneciurn and lodicules (X 2.5); m, lodicules of the upper antheciurn (X 40); n, lower anthecium, lateral view (X 25); 0, lower anthecium, palea side (X 25). Drawing a based on Hatschbach 28413 (US), b-o on Hatschbach 28413 (US), Kuhlmann 2774 (US), and Widgren 924 (US). 38 SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE15.-Panzcurn arktellurn: a, upper anthecium, lateral view showing thin lemma and palea (X 1 j0); b, detail of palea in a showing prickle hairs, silica cells, and stomata (X 500); c, detail of upper antheciuni, lemma side, showing bicellular microhair, prickle hair, and sinuous silica cells (X 1000); d, detail of upper palea showing stoma (X 1500). Photomicrographs a and d based on M’idgren 924 (US), b and c on Hatschbach I4888 (US). NUMBER 59 39 cylindrical, glabrous to hirsute; internodes solid lemma .i-nerved: lodicules 2, flat, covering the or with a small lunien, 15-40 ciii long. iyodes base of the gyneciuiii: stamens 3, stigmas 2 with obscure, compressed, glabrous. Sheaths shorter the styles united at the base. Caryopsis dark than the internodes solid or with a small lumen, browm ivith aii oblong hiluni; embryo Ih the 10-2.3 ciii long, keeled, rigid, overlapping at the length of the caryopsis base, glabimus or papillose-pilose along the mar- Kote: In some specimens the rachilla is pro- gins. Lzgule iiieriibranous-ciliate, 0.5-0.7 iiim longed in the form of a small point above the long; collar inconspicuous. Blades linear-lanceo- upper antheciuiii. late, acuniinate, 4.3-7.3 cm long, 1-1.8 cni \vide, COMMON ~AILlE.-”CaIlar~ll~.” narrow and involute torvard the base, becoming SPECIFIC EPITHET.-This species was de- flattened and Fvider, sometimes tvith the margins scribed by Doell under the name of Panicum involute, rigid, pubescent toivard the base on the Aristella instead of aristellum, rvhich would be the adaxial sui-face, otherivise glabrous, the margins correct neuter agreement. Comnienting on this, scabrous, the midnerve prominent. Panicles lax, Smith and Lt‘assliausen (1 978) felt that the epi- diffuse to contracted, pyramidal to oblong, thet Aristella should be retained and argued that iiianv-floIvered, 20-40 cm long, 7-20 cni wide, the specific epithet had been taken from a genus the branches diverging from the axis with the named Aristella 21s it appeared in this form in the spikelets congested along secondary racemes; index to the Flora Brasiliensis of hfar-tius axis cylindrical, scabrous, the branches alternate, (1 877:347-348). However, that index contains a triqueti-ous, markedly scabrous, densely to number of specific epithets that are capitalized, sparsely hirsute, the axils glabrous to pubescent; none of \vhich corresponds to the iiaiiie of a pedicels short, scabrous or occasionally hirsute. genus; Arnacites, ,.iturense, and Longispicula are Spikelets fusoid, 3.8-5.7 nirii long, 0.7-1 mm examples. The index is therefore not helpful and \$,ide (3.1-3.9 iiiiii long without the awns), \ve feel there is no strong argument to retain bror$m, soiiie\vhat compressed laterally, glumes Aristella. and lotvei. leni ma hei-baceous, weak, prom in en tl y DIsTRIRu.I.ION.-SoutheTn Brazil, from the scabrous on the nerves. Lower glume ovate-lan- state of Rio de Janeiro to Rio Grande do Sul. A ceolate, 2.3-3.6 iniii long M’ith an a1t.n 0.9-1.8 hygroplivtic species inhabiting marshes or sea- iitiii long, about as long as the lower lemma, 1- sonally inuiid;ited lot%.landsat elevations up to 3-nerved, the inidnerve and awn as well as the 1000 111. According to Smith and U‘asshausen iiia rgi iis marked I y scabrous, the inner surface (1 982) it is a coiiiiiioii species in fields of the scabrous toi\.ard the apex. Upper glume 3.7-5.7 central planalto of Santa Catarina, \$.here it is iiiiii long, niai-kedly scabrous at the apex on the associated m*ith Hjpogjnium virgatum and Andro- i ii 11e i- s u r face , .3 -11e r ved , the 11er ves scabrous , the pogon lateralis. midnerve extending into an awn 0.7-2 mm long, ADDITIONALSPECIMENS SEEN.-BRL4ZIL. this tlvisted or not. LozLler lemma oblong, acute, PARANA:Foz do Rio Taquaral, Hatschbach ??37 2.8-3.6 iiiiii long, aIvnless, 3-5-nerved, the mid- (US): Guarapuava, Kio das Pedras, Pereira 3 nerve scabrous. Lower palea conspicuous, lanceo- Hatschbach 8004 (US); Prainha, Hatschbach late, 2.3-3.1 inn1 long, 0.j-0.7 mm \$.ide, hya- 28413 (US); hlallet, Hatschbach 15?71 (US): Rio line, glabrous, ivith scabrous, denticulate niar- das Cinzas, Barra do Perdizes, Hatschbach 8491 gins: niale floiver present Lvith 3 stamens. Upper (US): road to Porto \’itoria, Hatschbach 14888 anthecium fusiform, 2.3-3.1 iiiiii long, 0.3-0.8 (US); Piraquara, -22 km E of Curitiba, Tessmann iiiiii \vide, inembranous, smooth, pale to brown, s.n., 1 Jan 1949 (US): Rio das Pedras, Hatschbach M.ith prominent Iiooks toryard the apex of the 10615 (us).RIO DE JANEIRO:Lvithout locality, lemnia and palea and on the margins of the Glaziou 20576 (US). RIO GRANDEDO SUL:Tu- lemma, also \vith silica cells, bicellular microhairs paceretan, Araujo 338 (US). SANTACATARINA: 6 and stomata ;it the apex of both lemrna and palea; km E of Porto Uniao, Smith 3 Klezn 15279 (US); 40 SMITHSONIAN CONTRIBUTIONS TO BOTANY

Born Retiro, Reitz 3 Klein 5500 (US); 10 kni E cylindrical, glabi.ous; internodes hollow, 12-28 of Pop Preto, Smith 3 Klein 1573'6 (US); Porto cni long. Sheaths 12-1 7 ciii long, long-ciliate on L'niao, Reitz 3 Klein 13671, 13680 (US); \'allos, tlie margins, otherivise glabrous. Ligule membra- Reit- 3 Klein 3706, 13571 (US); Faxenda Frei nous, laciiiiate, ivitli long i\.hitisli hairs toivard Kogerio, Poi-to Uiiiao, Reitz 3 Klein 1?6l? (US); the base of the adaxial surface of the blade, llorro do Caiiipo ;\legre, Sao Francisco do Sul, brotvn. Blades Ianceolate, acuminate, 20-33 cm Reitt 3 Klein 10062 (US); Campo do Areao, long. 2-3.8 ciii ivide, flat, tlie upper ones cor- Santa Cecilia, Reitz U Klein 14150 (US). SAO date, the loi\,er ones M.itli >I pseudopetiole up to PAULO:Boraceia, margem do Rio Claro, Kuhl- 3 ciii long, the abaxial surface glabrous, adaxial mann 2774 (US). glabrous to sparsely pubescent, the margins scabrous to short-ciliate, the basal ones long-ciliate; Taxonomic Position of Panicum killipii upper blades small. Panicles lax, diffuse, pyrami- This species displays no apparent affinity to dal, 15-33 cni long, 10-1 7 ciii Ivide, the an!. other species of Panicum, as Hitchcock hini- branclies divergent, alternate, \%.idel?separated; self pointed out in tlie original description. Ho~v- axis angular , sca br o LI s , tlie branches flattened, ever, its affinities to Paspalurn, to Lvhich lve are scabrous, the axils sliort-pubescent, the spikelets transferring it, become clear in tlie following in pairs and ari-anged more or less unilaterally enunieration of characters: on the raceiiiose branches, one subsessile, the 1 . The spikelets are arranged on unilateral other sliort-pedicellate; pedicels shorter than the 1-aceiiies,ivitli one subsessile and the other short- spikelet, scabrous. Spikelets ovoid, plano-convex, pedicellate. .~Iltliougliboth spikelets are usually 2-2.4 niiii long, 0.8-1 iiim ]vide, purplish, gla- subsessile and appressed to the rachis in Pas- brous or occasionally with some hairs toivard the palzim, soiiie meiiibers, such as Paspalurn jlavum sunitnit of tlie upper glume and IoLver lemma. Presl and P. corcovadense Raddi, have pedicellate Loxler glume o\xe, acute, 0.2-1 iiini long, nerve- spikelets like Panicum killipii. less, short-pubescent. Cpper glume and lower 2. The spikelets are plano-convex and obtuse, lemma subequal, obtuse, herbaceous, .%nerved, characters that are particularly noticeable in the glabrous or pubescent. Lower palea elliptical, upper anthecium (the presence of a lower male 1.8-2.2 mni long, O..i-0.8 inn1 \vide, hyaline, flolver obscures this in certain cases). Indeed, the glabrous, the upper margins short-ciliate; male spikelets of Panicum killipii and those of Paspalum floiver present, Ivith 2 lodicules and 3 stamens, jlazum show ;I great deal of similarity. the anthers 1.3 iiini long. Cpper anthecium ovoid, 2-2.3 iiini long, 0.8- 1 tiitii Fvide, plano-convex, Paspalurn pale, shining, papillate throughout, scabrous at the apex of the lemma. Caryopsis 1.4 tiirii long, Paspalurn killipii (Hitchcock) Zuloaga 8c 0.7 iiini i\,ide; hilum punctiform, the embryo Soderstrom. new combination about half as long as the caryopsis. Drs-rRIf$uTronr.-This species is knowm only FIGURE16 from the Peruvian departtnents of Ayacucho, Panicum killipii Hitchcock, 1930b:383. ["Type in the L. S. Cuxco, and Huancavelica, at 750 to 1200 meters National Herbarium no. 1357803, collected in thickets, el eva t iot1. Aina, betrveen Huanta and Rio Apurimac, Dept. Ayacu- ADDITIONAL SPECIMENS SEEN.-PERU. cho, Peru, alt. 750 to 1000 meters, May 1929, by E. P. CVZCO:tvitliout locality, Bues s.n., Jun 1930 (F, Killip and .4. C. Smith (no. 22804)." Holotype, US.] US). AYACUCHO:Aina, betJveen Huanta and Rio Perennials 80- 160 cm tall Fvith thick adventi- Apurimac, Killip 3 Smith 22803 (US); Ocarrapa, tious roots ;it the base. Culms erect, rarely decum- betrveen Huanta and Rio Apuriniac, Killip 3 bent and rooting on the loiver nodes, branching, Smith 22491 (US). HUANCAVELICA:Vuelo-Pata, NUMBER 59 41

FIGURE16.--Paspalurn killipii, new combination: a, upper anthecium, lemma side (X 100); b, detail of a showing simple papillae (X 500); c, detail of immature upper anthecium, lemma side (X 500); d, upper antheciuni, Iemrna side, showing bicellular microhairs (X 500). All photomicrographs based on Touar 4615 (US). 42 SMITHSONIAN CONTRIBUTIONS TO BOTANY

arriba de L‘irgen-Pampa, a1 SE de Tintay, Tovar rather than asperous. In the following year Des- 4615 (US). vaux (1 8 13) redefined the genus and followed DrscvssroN.-The species exhibits great vari- the modifications made by Beauvois. Trinius ation in the length of the lower glume, from (1820) accepted the genus and also used the spikelets in tvhich it is almost absent to those in name of S. hirsuta. tvhich it attains about half the length of the In 182.3 Sprengel transferred the species to spikelet. Panicum and called it P. streptostachys, and Tri- The species is endemic to Peru, a distribution nius (1826), Kunth (1829), Steudel (1855), and that fits ivell Ivith Paspalum, in which genus more Doell (in Martius, 1877) followed his lead. Doell than 2.5 species occur exclusively in the Andean addition a 1 1 y 1i s t ed severa 1 s y n on y ins: P. ba la n ites region of Colombia, Ecuador, Peru, and Bolivia. Trinius, P. perfoliatum Nees, P. vaginaejlorum On the other hand, the only species of Panicum Steudel, P. rohri Nees ex Steudel, S. asperfolia that are found here are those of wide distribu- Desvaux, and S. hirsuta Beauvois, and listed P. tion. macrophjllum Raddi as doubtful. Benthani (1 883) One specimen, Bues s. n., differs from the tvas the first to indicate the position of the species others in its smaller panicles with densely hirsute in the genus by assigning it to section Grandzjlora axis and branches and densely hispid sheaths and rvith P. macranthum and P. zizanioides. Hitchcock foliage. The specimen consists only of the upper, (1922) accepted the species as valid in Panicum densel!. branched, part of a culm. but effected the combination P. asperzjolium (Des- va ux) Hi tchcoc k. It ivas not until 1931 that the original genus Taxonomic Position of Streptostachys was re-established by Pilger. For the first time A.N. Desvaux established Streptostachys in since its inception 12 1 years earlier, Streptostachys 1810 i\.ith ;I single species, S. asperfolia, lvhich Ivas described on the basis of several natural he described ;IS iiionoecious and with spikelets of characters: the oblong shape of the lower glunie, both sexes intermixed. He further described the lvhich reaches 3/4-4L5 the length of the spikelet, female spikelets as having 18-20 glumes. The abscence of a sterile palea, and linear hilum. name \vas based on the Greek streptos (twisted) Pilger further pointed out that the base of the plus stachp (spike), in reference to the tH.isted spikelet is flat and strongly thickened with re- disposition of the racemes. Unfortunately, the curved margins. In 1940 and 1934 the same type material ~vasa sterile monstrosity and nei- author ernphasized the generic characters of long ther the description nor generic nanie reflected lo\\.er glume, thickened and flattened base, and the true nature of the genus. In a footnote to the the prominently thickened margins. description, Desvaux noted that the disposition Butzin (197Oa) grouped together a whole se- of the flowers \vas quite unusual and served to ries of genera of Paniceae on the basis of leaves separate the genus from any other known grass. Ivith tessellate venation, adding that all of these After examining specimens that lvere part fer- possessed a s)ndronie of characters that he con- tile and part sterile, Palisot de Beauvois (1812) sidered priniitive: the “bambusoid habit”; diffuse emended the original description of the genus. panicles; presence of a loiver, usually many- Although he described the sterile racemes in the nerved glume; elongation of the rachilla between same iiianner 21s had Desvaux, he added a de- the lo\\.er glume and the upper antheciuni; pres- scription of the fertile panicle and normal spike- ence of a developed lower floLver; and indurate lets and illusti-ated both types of inflorescence upper anthecium. He cited genera that “prove” togethei. Ivitli details of the spikelet, lodicules, that the Paniceae kvere derived from ancestors and ovai-!. of ;I iiornial panicle. At the same time ~vi t h 111an y - fl o rv ered s pi ke 1e t s , i 11 c1 u di 11g among he changed the iiaiiie of the species to S. hirsuta, these Streptostachy, for bvhich he cited teratolog- pointing out that the leaves \vere pubescent ical panicles in lvliich the spikelets denionstrate a NUMBER 59 43

multiplication of glumes and absence of fertile macrantha and S. ramosa it is erect and promi- floivers. Butzin regarded this as an extreme case nently thickened. of spikelet dinlorpliism that could be interpreted Caryopsis: The hilum is linear and as long as as a link betlveen putative many-flowered ances- the caryopsis. Streptostachys shares this particular tors and the genera that presently comprise the character Lvith fe\v genera of Paniceae, including tribe. Acroceras, Echinochloa, Homolepis, Mesosetum, Bu t 7, i ii related St rep tosta c hys p h y logen e tical 1y Oplismenus, and Tatianyx. to Acroceras, enumerating the following shared Upper Anthecium: Round papillae are evenly characters: amplexicaul leaves, panicles with spi- distributed over the lemnia and palea. The an- kelets ainnged in pairs in racemose branches, thecium is generally densely pubescent at the and caryopses with a linear hilum. He distin- apex and sparsely so or glabrous toward the base, guished .4croceras by its laterally compressed spi- varying in the three species. The hairs are uni- kelets and presence of a rudimentary lower cellular, long and wavy in S. macrantha and S. flo\\rer. (Although the spikelets of Streptostachys ramosa, Lvhereas in S. asperfolia they are bicel- are indeed dorsally compressed, a lower flower lular and shorter with the distal cell elongated. is frequently present. Along with Oplismenus, Lower Lemma: In different specimens varia- these t\vo genera are further distinguished by a tion in the consistency of the lower lemma may linear hiluiii. Homolepis belongs here as well, occur; in sonie spikelets it is niembranous, although Butyin included it with genera having lvhereas in others the median portion is hard and a punc ti form hi lu 111 .) the edges menibranous. There are also spikelets In another paper Butzin (1970b) published a Lvith the louver lemma indurate throughout and systematic treatment of the Paniceae and divided Lvith an ornamentation similar to that of the it into 11 subtribes, giving to the group already upper antheciuni, i.e., with abundant hairs and mentioned the rank of subtribe, which he called evenly distributed papillae. hlicrocalaniinae, characterized by leaves with tes- Ligule: This may be ciliate-formed by a line sellate venation. Curiously, he placed Panicum in of whitish hairs-to absent, with all variations another subtribe even though many of its species between these extremes occurring within a single share the character assigned to his Microcalami- specimen. nae. We do not consider the tkvo follorving charac- 111 spite of iiionstrous material and his niisin- ters, M.hich r\'ei'e enumerated by Pilger, to be of terpretations of it, Desvaux's recognition of a taxonomic value: new genus, Streptostachys, has been validated by Lower Glume: Pilger pointed out that the the years. To the type-species bve are adding a lorver glume is generally about :Y4 to 4/5 the length iier$r one and transferring yet another from Pan- of the spikelet and utilized the character to sep- icum. The observations that follow refer to the arate Streptostachys from the remaining genera of genus of three species as tve now recognize it. Paniceae. It'hen S. macrantha and S. ramosa are Kachilla: This is manifest with the internodes included in the genus, the length of the lower prominent Iietrz.een the loi\.er glume and upper glume varies from ]/:3 to 4/5 the length of the antlieciuni. At the same time it is thickened and spikelet. In Panirum itself the length of the loiver gives the spikelet a particular aspect at its base. glume varies considerably; for exaiiiple, in the As ;I consequelice of this peculiar type of rachilla, groups Parviglumia and Dichotoniiflor-a, it tlie gliriiies arid lo~.erand upper anthecia are reaches to 'h the length of tlie spikelet, to sti-oiigl!, adnate to it and do not break apart Y4 in Panicum, Rudgeana, Sarmentosa, Laxa, easil),, for \i7hicIi reason the spikelet falls at nia- Penicillata, and approxiniately V4 in Dura and tirrit), \vitli all of its parts together. In S. asperi- Parvifolia. In species such as P. obtusum and P. folia the rachilla is recurved at the base and longipedicellatum, the lower glunie is about as covei's tlie upper part of the pedicel, but in S. long as the upper glunie and lower lemnia. '41- 44 SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE1 'i.--Acroceras zzzanioides: a, upper antheciurn, lateral view, showing thin and crested suniniit~of leninia and palea (X 100): b, upper anthecium, lemma side, showing the crested summit (X 100): c, detail of apes of upper lemma, showing prickle hairs and rounded silica cells (X .i00); d, same as c but at higher magnification (X 1000). All photomicrographs based 011 Gillett I6486 (US). NULIHER 59 4 5

FIC~L-RI.:1 ti.-,$croceras paucispicatum: a, upper anthecium, lemma side, showing the crested sutiitiiit and rugose surface (X 50); b, upper anthecium, palea side, showing crested sunimit and rugose turbce (X 50); c, detail of summit of lemma shoi\,ing bicellular microhairs. rounded papillae, and prickle hairs (X 500); d, detail of apex of upper anthecium, palea side, showing S~IIICstructure as in c (X 200). All photomicrographs based on Hassler 12914 (US). 46 SMITHSONIAN CONTRIBUTIONS TO BOTANY

though this character generally remains constant sar tic u la t in g be low the g1 u mes, cad uco us, blunt , ivithin each group, it is not considered to be of glabrous to shortly hirsute; rachilla thickened generic value. between the lower glunie and upper anthecium Lower Palea: Contrary to published accounts, and with conspicuous internodes. Glunies 2, her- a lower palea is developed in sonie specimens of baceous, pale to purplish, unequal (the lower S. asperqolia. Although rudimentary or reduced glume rarely equalling the upper). Lower glume in sonie cases, it reaches half the length of the oblong to ovate, Ih to 4/5 as long as the spikelet, loiirer lemma in others. In S. macrantha and S. 1-7-nerved. Upper glume 5-7-nerved, obtuse to ramosa the lolver palea is conspicuous, attaining acute, frequently not covering the upper anthe- the same size ;is its lemma and sometimes enclos- cium. Lower lemma 3-3-nerved, acute. Lower pa- ing ;I male flower. In Panicum the lower palea is lea conspicuous, lanceolate, equal in size to the absent in the groups Parviglumia and Cordov- lemma or reduced to absent; male floFver some- ensia and in some species of the groups Panicuni, times present. Upper anthecium fusoid to nar- Lam, and Penicillata. roivly-ellipsoid, pale, shining, indurate, papillose- pilose over the entire surface (more so toward Streptostachys Desvaux, 18 10, emended the summit), and ~vithround papillae distributed Zuloaga & Soderstrom evenly, margins of the upper lemma enclosing Streptostachys Desvaux, 18 10: 190. the palea; lodicules 2, conduplicate; stamens 3, Plants perennial, cespitose or with short rhi- styles 2, stigmas plumose. Caryopsis fusoid, pale 7oiiies. Culms decumbent or ascending, simple or to light bro\vti: hilum linear: embryo less than branched: internodes hollow. Ligule ciliate, half the length of the caryopsis. formed by a line of whitish hairs, or lacking. A hew W'orld genus, occurring in Trinidad Blades ovate-lanceolate, flat, subcordate to cor- and South Anierica and favoring open places, date, amplexicaul. Panicles lax, diffuse to con- savaiiiias, or cerrados r\itIi sandy soils; less fre- tracted, the spikelets short-pedicellate, aggre- quent in margins of forests or woods. gated in racemes diverging from the rachis. TYPE-SPECrEs.-Streptostachjs asperi$olia Des- Spikelets oblong, compressed dorsiventrally, di- vaus.

Key to the Species of Streptostachys

1. Spikelets 3.9-5.1 iiim long; lower glume obtuse, 3/4-4-% as long as the spikelet; rachilla forming a thick, recurved disc around the pedicel; lower palea small, ovate, '13 as long as the lemma or absent; iipper antheciuni with bicellular hairs toward the apex, the basal cell short, the distal cell elongated; papillae encircled by a raised process; axillary panicles present, usually reaching the length of the terminal panicle; leaves amplexicaul, 2-4.3 cni wide ...... S. asperifolia 1. Spikelets .5.9-9.3 iiim long; lower glume acute, ]/3-'/2 as long as the spikelet: rachilla thickened, not recurved at the base; lower palea conspicuous, lanceolate, equal in size to the lemma; upper aiitheciiim Lvitli long unicellular hairs throughout; papillae not encircled by a raised process; axillary panicles absent: leaves subcordate, not amplexicaul, 0.7-1.3 cm wide ...... 2 2. Panicles 5-10 cm long, 2-4 cm wide; axils of the racemes glabrous or occasionally sparsely pubescent; plants 50-70 cni tall; blades 7-12 cm long, 0.7-1.3 cm wide ...... S. macrantha, new combination 2. Panicles 30-45 cm long, 15-30 cm wide; axils of the racemes densely pubescent with long whitish hairs; plants 1.5-2 m tall; blades 35-55 cm long, 1-1.2 cm wide...... S. ramosa, new species NUMBER 59 47

Streptostachys asperifolia Desvaux 15-35 cni long. Spikelets fusoid, 3.9-5.1 mni long, 1.5-1.9 niiii wide, glabrous to hirsute with FIGURE19 a truncate and prominently thickened base. Streptostachy asperifolia Desvaux, 1810: 190. [Type: “Habitat Lower glume oblong, obtuse, 3.2-4.7 nim long, in America calidiore.” Holotype, P, not seen; fragment of %-4/5 as long as the spikelet, 5-7-nerved, the the type, US.] nerves anastomosing toward the apex. Upper Streptostachy hirsuta Beauvois, 18 1250, pl. 10: fig. 1 1. [Type: The same as that of Streptostachys asperifoh Des- glume 3.5-4.7 mm long, 3-7-nerved, the nerves vau x . ] anastomosing tolvard the obtuse apex. Lower Panicum streptostachys Sprengel, 1825:316. lemma acute, 3.4-4.6 nini long, 3-nerved, gla- Panicum perfoliatum Nees, 1829: 142. [Type: “Habitat in brous to sparsely pubescent at the base and cen- sylvaticis umbrosis ad flumen Amazonum ejusque con- tral zone, hirsute at the apex, membranous to fluentes provinciae Paraensis et Fluminis Nigri passum.” indurate (in ivhich case the midnerve and adja- Holotype, M,iiot seen.] Panicum balanites J.B. Trinius, 1836:297. [Type: “Colarea cent nerves not manifest), its margins enclosing ad ost. fl. Ama7 (America calidiores, Poeppig).” Holotype, the upper anthecium. Lower palea lanceolate, LE, not seen: isotype, US sheet no. 1126919.1 1.3-1.9 iiiiii long, 0.4-0.9 mm wide, brown or Panicum vaginaeflorum Steudel, 1835:80. [Type: “Guiana.” whitish, hyaline, glabrous, frequently absent; Holotype, P, not seen: fragment of the type, US.] male floIver absent. Upper anthecium oblong to Panicum streptostachys f. paleacea Doell in Martius, 1877:231. [No type cited.] narroivly ellipsoid, acute, 3.1-4.1 mm long, 1.3- Panicum asperifolium (Deskaux) Hitchcock, 1922:489. 1.6 mm tvide, pale, shining, finely papillose throughout with the pa pi 1lae distributed evenly , Perennials, 30- 100 cm tall, shortly rhizonia- pubescent toward the apex; lodicules 2, condu- tous. Culms geniculate, decumbent and some- plicate. Carq’opsis oblong, 2.4-2.8 nini long, 1.2- times rooting at the lou.er nodes, then becoming 1.3 mm wide, pale or broLvn; hiluni linear; eni- erect, simple or branching, cylindrical to com- bryo less than half the length of the caryopsis. pressed, sparsely hirsute to glabrous, many- DIsTRIBrrTIoN.-Inhabitiiig forests in shaded noded; internodes Iiollo~,4-20 cni long, pale to spots or more frequently in savannas or cerrados, purplish. ‘Yodes purplish, compressed, glabrous in sandy soils and among shrubs, from Trinidad to pubescent. Sheaths 5-10 cni long, densely hir- to l’enezuela, the Guianas, and northern Brazil. sute to glabrous, Lvitli long tuberculate hairs to- ADDITIONALSPECIMENS SEEN.-BRAZIL. Mard the mai.gins. Ligule in an arc, shortly ciliate Without locality, Burchell 9291 (US), Gardner or lacking [\.it11 long Lvhitish hairs toward the 6108 (US); Gratijeiro, Luetzelburg 25974 (F, US). base of the blade; collar hirsute to glabrous. AMAPA:Matapi , Pires & Silva 4799 (US); Porto Blades ovate-lanceolate, acuniinate, 14-28 cni Platon, ,Matto.c 10120 (US); hlapati, kni 116, Black long, 2-4.3 cin rvide. flat, cordate, amplexicaul, c” Lobato 50-9676 (US); 2 kni from Rio Araguari, coninionly tessellate, densely hirsute on both sur- along road from Porto Platon to hlacapa, Pires faces to glabrous, the niargins ciliate with long et al. 51084 (US); road to Amapa, kin 108, be- tuberculate hairs, the midnerve prominent on ginning of Rio Pedreira, Pires 3 Cavalcante the abaxial surface. Panicles lax, oblong to pyra- 521 79 (F, NY); Calcoene, frequent at roadside, niidal. 6- 16 ciii long, 1-9 cm wide, the branches road to gold mines, Pires & Cavalcante 52520 often diverging from the axis, spikelets short- (NY). BAHIA:wLvithout locality, Zehntner 3758 pedicellate and aggregated in pairs on the (US); Cruz das Alnias, Pinto 139 (US); Serra do branches: axis angular, scabrous to hirsute, the Apora, hluritiba, Pinto 256 (US); road Maracas- tira n c 11 es ;I 1 t ei-ii ati n g , flat ten ed , g la bro u s to Contendas do Sincora (BA 026), kin 2, Mattos sparsely pubescent, the axils pubescent; pedicels Silva et al. 213 (NY); Road BR 330, trecho Je- scabrous. angular, sometimes bearing whitish quie-Ipiau, 4 kin de Jequie, ,Uori c“ King 12200 hairs; axillary panicles numerous, similar to the (KY):on road to Abaira, -8 km N of the town terniinal one and borne on long peduncles, these of Rio de Contas, Harlej et al. 15235 (NY); 31 48 SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGCRF1 F).--Streptostachys asperfolia: a, upper anthecium, lemma side (X 100); b, detail of a showing ~nacrohairs,bicellular microhairs, and rounded papillae (X 200); c, detail of upper palea showing bicellular microhairs and simple papillae (X 200); d, detail of simple, rounded papillae of upper palea (X 2000). All photomicrographs based on Swallen 4898 (US). NUMBER 59 49 km S of Olindina, along highway BR-116, Dav- Martius s.n. (US); 32.8 km along road S of i\mar- idse et al. 11750 (NY); 16 km NW of Lagoinha, ante, 25 km S of Piaui river, Eiten 3 Eiten 10371 on side road to Minas do Mimoso, Harley et al. (US). RIO GRANDEDO KORTE: Pedro Velho, 16642 (NY). CEARA:Serra de Baturite, Eugenio Emygdio 1655 (US); Natal, Swallen 4675 (US); 287 (US); Campo Salles to Crato, Swallen 4?17 Nova Cruz to Montanhas, Swallen 481 I (US). (US); Campo Grande, Swallen 4517 (US); Cam- FRENCH GUIANA. Without locality, Sagob ocim to Granja, Swallen 46?1 (US); Baturite to 1395 (US), Leprieur s.n. (US). CAYENNE:St. Ma- Guaramirango, Swallen 4431 (US). GOIAS: 2 km deleine road, vicinity of Cayenne, Broadway 745 N of Araguaina, Irwin et al. 211?? (F, NY, US). (US). MARANHAO:S5o Sao Luis to Sao Jose, Swallen ?46? GU Y A NA. Without 1oca 1it y , Sc ho m b u rgk 7 74 (US); Caxias to Barra do Corda, Swallen ?485 (F, US). Waranawa Ranch, Harrison 1047 (US). (US); 39 km NE of city of Grajau, on old road to SURINAM. Kayser airstrip, 25 km above con- Barra do Corda, Eiten €3 Eiten 10266 (US); Car- fluence with Lucie River, Maguire et al. 5?95? olina to Sao Antonio de Balsas, Swallen 4144 (US); Zuid River, Kayser airstrip, Irwin et al. (US); about 25 km S of Loreto, between the Rios 5599 I (F); Kaysergebergte airstrip, Freund R-34- Balsas and Parnaiba, Eiten €3 Eiten 414? (US); B (US). Ilha de Balsas, region between the Balsas and TRINIDAD. Chaguasamas, Broadway s.n., 12 Parnaiba rivers, Santa Barbara, 55 km SSE of Jul 1927 (US). Loreto, Eiten €3 Eiten 10704 (US). PARA:without VENEZUELA. BOLIVAR:between Los Castil- locality, Spruce 95 (US); Marapanim, estrada para 10s de Guayana and Piacoa, Steyermark 86270 Jgarape Grande, Silva 496 (US); Marajo Island: (US); Morichal, Santa Isabel, Cardona 655 (US). Caracara river, in plantations, Goeldi 99 (F, US), MONAGAS:17 km NNW of San Felix and Rio near Soure, Goeldi 299 (F, US); Ilha de Mos- Orinoco along hwy 10, Davidse 4604 (NY). queiro, near Para, Killip & Smith ?0482 (US); Drscuss1oN.-The pubescence in this species Ilha de Marajo, Salvaterra, Kauffmann 4 (F, US); is quite variable, both on the vegetative parts and Soure, Black 50-9071 (US), Swallen 4975 (US); in the inflorescence, with hirsute to totally gla- Fazenda Camburupy, near Soure, Swallen 4956 brous specimens encountered. The panicle of (US); Rio Moju, Fabrica, Black 54-16242 (US); Swallen 44?1, for example, contains both hirsute estrada Joao Coelho-Vigia, km 40, Black et al. 52- and glabrous spikelets. 14200 (US); Santarkm, Kuhlmann 1740 (US), The lower lemma is frequently indurate with Swallen 3267, ?29? (US); near Santarkm, Spruce a consistency similar to that of the upper; in such 15, 606, 697 (US); Boim, Tapajos, Kuhlmann examples the midnerve and adjacent nerves are 1922 (US); Mosqueira, Swallen 4898 (US); Breu not manifest. In other cases the lower lemma is Branco, kni 40 S of Represa Tucurui along high- membranous and similar to the glumes. Even way BR 422, Plowman et al. 9659 (NY); 36 km though the lower palea is generally lacking or SE of Vigia, along highway PA-140 to Belem, poorly developed, it occasionally reaches half the Davidse et al. 17597 (NY); Campo de Matapi- length of its lemma, as in Pires 4799. Axillary quara, -1 km E of Matapiquara, 73 km NNE of panicles are frequent and numerous in this spe- Castanhal by road, Davidse et al. 17896 (NY); cies and reach the same length as the terminal -73 airline km NE of Castanhal; savanna adjoin- one. ing the village of Martins Pinheiro, Davidse et al. The upper anthecium is sparsely pubescent 17960 (NY). PARAIBA:Areia, Coelho de Moraes toward the apex with bicellular hairs on the 854, I156 (US). PERNAMBUCO:Dois Irmaos, vi- lemma and palea. Sparse long unicellular hairs cinity of Recife, Chase 7728 (F, US); Olinda, and numerous hooks also occur on the the apex Pickel 2273 (US); Joao Pessoa, Pickel 3852 (US). of the lemma, as revealed by SEM. Except toward PIAU~:without locality, Lutzelburg 5544, 5547 the margins of the lemma the entire surface of (US), Ule 7425 (US), Gardner 2?41 (US); Oeiras, the anthecium is evenly covered with rounded 50 SMITHSONIAN CONTRIBUTIONS TO BOTANY papillae, each of these centered on a swelling of with manifest internodes. Louwr glume ovate, the anthecium. acute, 2.3-3.5 iiiiii long, lh-lh the length of the A ligule is generally absent. When it does occur spikelet, 1 -3-nerved, the midnerve scabrous or it consists of a line of short hairs. not to\\xrd the apex. Gpper glume 5-3.8 mm Bi-okvn (1 977) reported the species to be Cs, long, not completely enclosing the upper anthe- or non-Kran7, rvith a "C/"C index of -27.0. cium, sparsel!. hispid to glabrous at the obtuse apex, .5-7-nerved, the nerves anastomosed to- Streptostachy macrantha (Trinius) \Yard the apex. LozLler lemma 5.4-6 inn1 long, Zuloaga & Soderstrom, new combination acute, 3-.5-nerved, the nerves anastomosed to- FIGURE20 Mard the apex, densely hispid to glabrous, membranous to indurate. Lower palea lanceolate, con- Panicum macranthum Trinius, 1826:209. [Type: "V. spp. Brdsii. (Langsd.)." Holotype, LE, not seen; fragment of spicuous, 4.3-3.1 nniii long, 1 - 1.4 mni r+.ide, the type, US sheet no. 974708.1 hvaline, glabrous; nnale florver sometimes pres- Panicum imginatum var. alpha et beta Nees, 1829: 156. [T\pe: ent, Ivitli 2 lodicules, and 3 stamens. Upper anthe- "Habitat in Brasilia meridionali (Sellow). \'idi in Herb. cium ovoid, 5.1 -.5.9 mi11 long, 1.8-2.1 mm !vide, Reg. Berol." Holotype, B, not seen; fi-agnient of the type, pale, shiiiing, pubescent, covered lvith long, thin us s.n.1 u 11ice 11 u 1 a I- I i 1xi-oha i rs , g la bresce 11 t to ward the Short-rhizomatous perennials, 30-70 cm tall, base, finely papillose throughout, the papillae covered at the base bj. densely villous cataphylls. rounded and distributed evenly; lemma 5- Culms compressed, simple, glabrous; internodes nerved. Carppsis fiisiforni, 3.6 iiiiii long, 1.7 mm hollou., 5- 1 .5 ciii long. 'Yodes pilose to glabrous, uide; hiluni linear; embryo less than half the co 11113 ressed , dark ; basa I 1eaves penis te n t , c u r 1- length of the cai-yopsis. ing. Sheaths 6- 14 ciii long, pale to purplish, gla- DIsTRIRvTroN.-~ccurriiig in the Brazilian brous to short-pubescent, the basal ones short, ceri-ados of Goak and \lato Gr-osso betiveen 400 overlapping, the upper elongated. Ligule ciliate, and 1200 111 elewtion and reported from the 0..5-1 iinni long, lacking in the lower leaves; collar Amaiiiba). range of Paragua),. pale to purplish, sliort-pubescent to glabrous. AIIDITIONAL SPECIMENS SEEN.-BRAZIL. Blades ovate-laiiceolate, acunlinate, 7- 14 cm iVithout state: Rio de Saii Francisco, Pohl 329 long, 0.7-1 .3 ciin M.ide, flat, occasionally rvith (US); Lagoa Santa, Warming s. n. (IVY). DISTRITO involute margins, cordate and sometimes am- FEDERAL:20 km S of Brasilia on road to Goiania, plexicaul, densely hirsute to glabrous, the mar- near Rio Melchior, Irwin et al. 8658 (US); Ro- gins ciliate, the midnerve not differentiated; up- dovia DF6-Brasilia, Allem U Vieira 1087 (MO). per blades coninnoiily small or absent. Panicles GOIAS:without locality, without Glaziou 22526 locality, (US); 33 Glaziou 2 2526 (US); 33 lax, open, .5-10 ciii long, 1 .3-4 cm wide, long- kni S of Caiaponia on road to Jatai, Irwin 8 exserted, peduncles 16-33 cni long, the spikelets Soderstrom 7131 (MO, US); Serra do Caiapo, 30 more or less spread along the branches; axis kni h of Jatai on road to Caiaponia, Irwin 8 angular, glabrous, the loiver branches verticil- Soderstrom 7307 (US); -20 km S of Brasilia on late, opposite or alternate above, flexuous, gla- road to Goiania, near Rio Melchior, Irwin et al. brous, smooth, uithout spikelets at the base, di- 8658 (NY). MINAS GERAIS:Serra do Cip6, 110 verging fi-on) the axis, axils of the Iorver branches km NE of Belo Horizonte, Chase 9297 (US); glabrous to sparsely pubescent; pedicels scaber- Moi-ro do Cruzeiro, Lagoa Santa, 42 km N of ulous, claviforin. Spikelets fusiform, 5.9-6.8 mm Belo Horizonte, Chase YO18 (NY, US); 38 km de long, 2.1-2.5 miin wide, glabrous to hispid on Paraopeba, Ferreira 1463 (MO). the apes of the upper glume and lo\ver lemma, PARAGUAY. Sierra de Amambay, Estrella, greenish to purplish, the rachilla thickened be- Hassler 10115 (BAA), Rojas 6298 (BAA). tlveen the loiver glume and upper anthecium and DISCCSSION.--ASin S. asperifolia this species is NUMBER 59

FIGUREZO.--Streptostachys macrantha, new combination: a, upper anthecium, lemma side (X 100);b, detail ofa showing macrohairs and simple rounded papillae (X ZOO); c, upper antheciutn, palea side (X 100); d, upper anthecium, palea side (X 50). All photomicrographs based on Glazzou 22526 (US). 52 SMITHSONIAN CONTRIBUTIONS TO BOTANY

variable in the consistency of the lower lemma, Short-rhizomatous perennials, 1.5-2 m tall which may be membranous to indurate through- with densely villous cataphylls. Culms erect, few- out. In Irwin et al. 8658, for example, there are noded, simple, cylindrical, glabrous; internodes spikelets with the lower lemma totally membra- hollow, 8-20 cm long, Nodes purplish, glabrous nous and similar in aspect to both glumes; in to short-pubescent. Sheaths 15-40 cm long, others the central part of the lower lemma is longer than the internodes, overlapping at the indurate, smooth, shining, pale, and densely pu- base and villous, prominently distichous, keeled, bescent toward the apex, bearing rounded pa- sericeous to glabrous above, the margins short- pillae as well and not distinguishable from the ciliate. Ligule ciliate, formed by a line of whitish upper anthecium. In this latter case the midnerve hairs 0.8-2.5 mm long or lacking; collar dark and lateral nerves are not manifest. The lower brown to purplish, glabrous to short-pubescent. lemma remains membranous and purplish to- Blades linear-lanceolate, acuminate, 35-62 cm ward the margins and here the nerves are visible. long, 1-1.4 cm wide, flat or with the margins Both Kunth (1829) and Doell (in Martius, involute, rigid, cordate, the margins short- to 1877) made note of the thickened base of the long-ciliate, scabrous, hispid toward the base of spikelet and also mentioned that the base of the the adaxial surface, the remainder glabrous or panicle was either pubescent or glabrous. The densely pubescent on both surfaces with long illustration in the former work agrees exactly appressed hairs. Panicles lax, diffuse, pyramidal, with what we here consider to be the species. 30-45 cm long, 15-30 cm wide, spikelets solitary The lower sheaths are persistent and com- along the branches, subsessile; axis cylindrical, pletely envelop the culm, giving protection to undulate, glabrous, the branches verticillate or the plants that are affected by periodic fires. opposite, naked at the base, glabrous, wavy, axils densely pubescent, the hairs long, whitish, the Streptostachys ramosa Zuloaga 8c Soderstrom, pulvini dark brown, pedicels short, angular, new species smooth, glabrous, thickened at the apex. Spikelets FIGURES2 1, 22 fusoid, 6.3-9.3 mm long, 2.1-2.5 mm wide, glabrous, greenish to brown or purplish, glumes Type Specimen.-BRAZIL, BAHIA:22 22 km S and lower lemma with prominent nerves, pubes- of the Rio Galheirao along highway BR-020, 8 cent or scabrous toward the apex of the inner Apr 1976, 860 m, G. Davidse, T.P. Ramamoorthy surface. Lower glume acute, 2.2-4.4 mm long, €+ D.M. Vital 12216 (Holotype, MO; isotype, %-l/z the length of the spikelet, 3-nerved, short- NY). pilose at the apex. Upper glume obtuse or acute, Gramina perennia rhizomatibus brevibus, 1.5- 5.3-8.1 mm long, commonly leaving exposed the 2 m alta. Culmi erecti, simplices. Foliorum va- apex of the upper anthecium, 5-nerved. Lower ginae carinatae, manifeste distichae. Ligula cil- lemma subacute, 6.2-8.5 mm long, 3-5-nerved. iata. Laminae lineari-lanceolatae, 35-62 cm lon- Lower palea conspicuous, lanceolate, 5.3-7.9 mm gae. Paniculae laxae, diffusae, 30-45 cm longae, long, 1-2.2 mm wide, hyaline, glabrous; male axillis raniorum dense pubescentibus. Spiculae flower sometimes present, with 2 lodicules and 3 oblongoideae, 6.3-9.3 mm longae, 2.1-2.5 mm stamens, the anthers 4.3 mm long. Upper anthe- latae, glabrae. Gluma inferna acuta, longitudine cium ovoid to narrowly-ellipsoid, 5.4-7.9 mm '/3-'/2 spiculae aequans, 3-nervata. Gluma supera long, pale to brown, 1.9-2.4 mm wide, shining, obtusa vel acuta, 5-nervata. Lemma infernum pubescent, with thin whitish hairs, more abun- subacutum, 3-5-nervatum. Palea inferna lanceo- dant toward the apex, finely papillose through- lata, 5.3-7.9 mm longa. Anthoecium superum out; lodicules 2, the stigmas purplish. Caryopsis oblongum ad anguste ellipsoideum, 5.4-7.9 mm ellipsoid, 3.8 mm long, 1.9 mm wide, olivaceous; longum, pubescens, omnino subtiliter papil- hilum linear; embryo a little less than half the losum. Caryopsis ellipsoidea, hilo lineari. length of the caryopsis. FIGURE‘LI.--Streptostachys ramosa, new species: a, habit (X 1); b, panicle (X 1); c, spikelet, lateral view (x 12); d, scheme of longitudinal section through base of spikelet (X 12); e, car)opsis, sho~inglinear hilum (X 12);J lower glume (X 12); g, upper glume (X 12); h, lower anthecium (X 12); i, lower lemma (x 12);j, lower palea (X 12); k, stamen (X 12); 1, upper anthecium, leninia side (X 12); m, upper anthecium, palea side (X 12); n, upper palea with gyneciuni and lodicules (X 12); 0, lodicules of upper anthecium (X 25); p, upper anthecium, lateral view (X 12). Didwing a based on Davidse et al. 12216 (MO), b on Hatschbach et al. 36145 (MO),and C-o 011 both. 54 SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE22.--Streptostachys ramosa, new species: a, detail of upper anthecium, lemma side (X ZOO): b, upper anthecium, palea side (X 40); c, base of spikelet showing thickened internodes (X 40); d, detail of upper palea showing macrohairs and simple rounded papillae distributed evenly over the surface (X 200). Photomicrographs a, b, and d based on Harley 16876 (US), c 011 Heringer 9590 (NY). NCMRER 59 55

PARATYPES.-BRAZIL.BAI~A: 15 kni S of the 1849). In his publication Mez gave very brief intersection of highl\’ay BR-020 and the Rio descriptions of his new species, making their Roda \‘elha, G. Davidse et al. 12085 (MO, NY, identification difficult. The author’s reference to US); Sen-a do Curral Feio, 22 km NW of La- the large size of the plant and striking similarity goinha, on side road to Minas do Minioso, 6 Mar of spikelet characters betiveen P. mirabile and S. 1974, R..Zl. Harley et al. 16876 (MO, NY, US); ramosa suggest that ive may be dealing ~viththe BR 242, km 216, 12 k~iii% of Lencois, S. Mori same species. A fi.agment of the type, rvhich 13335 (NY); Espigao Mestre, -100 kni WSW of consists of a racemose branch of the panicle, is B.‘11 ._I eiras, It’. R. ‘Anderson et al. 36657 (MO, NY). at US and is labeled “Panicummirabile Mez. Brasil GOIAS:region of Chapadaregion dos i’eadeiros, 38 km occ. Tambei-lik.”The holotype from the Berlin N of Sao Joao da Alianfa, at W long. 47”30’, S Museum consists only of a fragment of the pani- kit. 14”30’, fi;. Y. Dazllson 14554 (US). M.4TO cle and a leaf and even though the aspect of the <;ROSSO: Chapado dos Guimaraes, G. Hatschbach panicle matches that described for S. ramosa, the et al. ?5145 (hIO, NY); eiitre Queiniada e Rio absence of fuller material does not allow us to \‘el.de, \vitliout collector (MO sheet no. arrive at a definitive conclusion regarding the 23 12029). MINAS GERAIS:Tres Marias, 26 Feb identity of P. mirabile. The point is not critical, 1964, E.P. Heringer 9590 (NY); -20 kni S of ho\vever, because the name of this species is Alto do Paraiso (formerly Veadeiros), H.S. Irwin invalidated by tiz~oearlier epithets. et al. 24716 (NY); -33 kni YE of Francisco Sa, CHROMOSOMENUMRERS.--n = 27 (Davidse road to Saliiias, H.S. Irwin et al. 23126 (SY). and Polil, 1978, as Panicum macranthum Trinius). DIsTRIRuTIoN.-Species of the Brazilian cer- Taxonomic Position of Tatianjx rados, occurring betlveen 800 and 1 100 111 elevation. Among the niost beautiful of paiiicoid grasses DrscussroN.--‘I’Iiis species is related to S. ma- in tropical Anierica is a species that was named crantha, from i$.hich it is distinguished by the size Panicum arnacites by Trinius for a grass that of the plants, leaves, and panicles, the branches inhabits the “campo rupestre” of Brazil in the of the latter. iiiiicli longer and the spikelets borne states of Bahia, Para, Goais, llinas Gerais, and iiiore distaiit froni one another than in S. ma- Mato Grosso. The epithet arnacites, Ivhich conies crantha. l’he densely pubescent axils of the pan- from the Greek ivord for “~.oolly,”aptly de- icle branches furthei- distinguish it. scribes the long-hairy spikelets of this species. The only basis for assigning Glariou 20123~ Our comparison of this Panicum Lvith other (US) to this species is a fragiiient of the panicle members of the tribe confirms its unique posi- Lvith the follo\ving annotation by A. Chase: tion. A linear hilum immediately distinguishes it Specimen in Paris lib. labeled Panicum grandijlorum n. from other species in the genus as does the sp. agrees with Glaziou 22526 and Chase collections but position of the pedicel relative to the spikelet. is 70 cm tall, the panicle 25 cm long, the ascending- The pedicel is falcate at the apex and the spikelet spreading branches to 9-1 0 cni long. held in an oblique position, behind the upper l’here is also a fragnient of the same specimen glunie and upper lenima Lvher-e it form a pubes- from \’ienna. The type-locality must be erro- cent callus. In Panicum the hilum is round and neous ;is given on the label: “Brazil, Rio. Glaziou basal and the pedicel is truncate at the apex with 20123a.” Obviously the species does not occur in the spikelet held erect. A linear hilum is found Rio de Jaiieiro and the \,eracity of this label- in other panicoid genera such as Streptostachjs, like so niany of Glaziou-is open to question. and a callus is found in Chaetium and Digitariopsis. It is possible that this species was first described Trinius did not refer to the caryopsis when he bv Xlez under the miiie Panicum mirabile Mez described the species nor \\.as the hilum men- (1 92 1) not ;I.Braun (1 846) nor M’alpers (1848- tioned in the latter references of Trinius (1834), 56 SMITHSONIAN CONTRIBUTIONS TO BOTANY

Steudel (1 855), or Doell (in Martius, 1877). The Cespitose perennials 40- 100 cm tall with unusual placement of the pedicel and spikelet densely villous cataphylls. Culms erect, cylindrical had apparently also never been considered as an to compressed, glabrous or sparsely pubescent important character to separate this species from near the nodes, simple; internodes hollow, 4-20 others in Panicum. cm long. Sodes purplish, long to shortly pubes- In comparison with other panicoid grasses, cent bvith whitish hairs. Sheaths 5-26 cm long, Panicum arnacites is superficially similar to An- longer than the internodes, the lower ones en- thaenantiopsis and Leptocoryphium, in which the veloping the culni, densely to sparsely pubescent, spikelets are lanate or pilose. In the latter the the upper ones glabrous. Ligule ciliate, formed anthecium is delicate, not rigid, and the margins by an arc of whitish hairs 0.3-0.4 mm long; collar of the upper lemma are hyaline and do not cover glabrous or short-pubescent. Blades linear-lan- the apex of the palea. The anthecium is open at ceolate, acuniinate, 10-36 cm long, 0.2-0.4 cm the summit and the lower glume is diminutive. wide with the margins involute, rarely flat, nar- In the former genus the upper anthecium is also rowed at the base, ciliate along the lower mar- open at its summit and the spikelets are arranged gins, the adaxial surface sericeous, the abaxial on unilateral racemes rather than in a panicle. pubescent; upper blades small, inconspicuous. Because Panicum arnacites cannot be accom- Panicles lax, diffuse, pyramidal, many-flowered, modated in any known genus of the Paniceae, 6-1 3 cm long, 4-9 cm wide, exserted, peduncles we are establishing a new genus for it, which we 15-24 cm long, almost completely included in are calling Tatianyx, in honor of Sra. Tatiana the upper leaves when young, the spikelets borne Skvortzov Sendulsky (born 1922, Harbin, Man- on long flexuous pedicels, axillary panicles some- churia), of the Instituto de Botanica inSaoo Paulo, times present; axis cylindrical or angular, an authority on the grasses of Brazil. smooth, glabrous, usually purplish, the branches alternate, opposite or verticillate, smooth, thin, Tatianyx Zuloaga & Soderstrom, new genus glabrous, wavy, the axils densely pubescent to (Poaceae: Panicoideae) glabrous; pedicels long, glabrous, smooth, flex- Gramen perennis. Culmi erecti, simplices. Lig- uous, falcate at the apex, positioned obliquely on ula ciliata. Laminae lineari-lanceolatae. Paniculae the dorsal side of the spikelet. Spikelets ellipsoid, laxae, pyramidales; pedicelli longi, flexuosi, apice 4-4.5 mni long, 1.2-1.4 mni wide, silvery-villous falcati, facie dorsali spiculae oblique affixi. Spi- with long rigid rvhitish hairs, exceeding the apex culae ellipsoideae, argenteae. Gluma inferna lon- of the glumes and lower lemma in length; glumes gitudine ‘/~-~/4 spiculae aequans. Gluma supera and lower lemma herbaceous, glumes and anthe- 5-7-nervata. Lemma infernum 5-7-nervatum. cia separated by prominent internodes. Lower Palea inferna lanceolata, interdum florem mas- glume acute, 2.3-3 mm long, ‘h to 3/4 as long as culum continens. Anthoecium superum ellipso- the spikelet, 3-5-nerved, glabrous on the inner ideum, induratum, papillas compositas superfi- surface. Upper glume acute, 2.8-3.5 mm long, ciares gerens. Caryopsis ellipsoidea, hilo lineari. leaving exposed the apex of the upper anthecium, 5-7-nerved. Lower lemma acute, 3.2-4 mm TYPE-SPECrEs.-Tatianyx arnacites (Trinius) Zuloaga 8c Soderstrom, new combination. long, glume-like, 3-7-nerved. Lower palea lanceolate, 3.2-4.3 nim long, 0.7-1 mm wide, long to Tatianyjc arnacites (Trinius) Zuloaga & short-pilose on the upper margins, otherwise gla- Soderstrom, new combination brous, hyaline; male flower sometimes present with 3 stamens, the anthers -3.3 mm long. Upper FIGURES23, 24, 25 anthecium ellipsoid, 2.9-3.7 mm long, 1-1.4 mm Panicum arnacites Trinius, 1832:117. [Type: “Figura ad wide, glabrous, indurate, shining, pale, with long specimen Brasilianuni.” Holotype, LE, not seen; fragment prickle hairs at the apex of the lemma, this and of the type, US sheet no. 974682.1 the palea covered by compound papillae distrib- FIGUR~:P:J.--Tatianyx arnacites, new combination: a, habit (X 1); b, stamens (X 18); c, lower pale~i,inner surface (X 18); d, lower palea, outer surface (X 18); e, caryopsis, embryo side (X 18);f; IoM.~~lenima, inner surface (X 18); g, caryopsis, showing linear hilum (X 18); h, lower gl~iiiie(X 18); i, upper glunie (X 18);j, lower lemma (X 18); k, upper anthecium, lemma side (x 18); I, pail. of spikelets, showing oblique attachment of-pedicels (X 12); m, spikelet, upper gluiiie side, showing oblique callus (X 12); n, upper anthecium, lateral view (X 18); o, upper antheciuni. plea side (X 18);p, lodicules (X 35);9, upper palea showing gynecium and lodicules (X 18); r, upper lemma (X 18). All drawings based on IrzLlin 2 Soderstrom 6959 (US) except e aiid g based on Pires 2947 (US). 58 SMITHSONIAN CONTRIBUTIONS TO BOTANY

FIGURE24.--Tatianjx arnacites, new combination: a, base of spikelet showing oblique callus (X 200); b, base of spikelet, upper glurne side, showing oblique callus (X 150); c, falcate pedicel, lateral view (X 200); d, pedicel, locus of attachment with spikelet (X 200). All photomicrographs based on E'iten 6752 (US). NUMBER 59 59

FIGURE 25.--Tatianyx arnacites, new combination: a, upper anthecium, palea side (X 200); b, upper antheciuni, lateral view, showing prickle hairs on apex of lemma and stomata below (X 200); c, upper anthecium, lemma side (X 500); d, detail of upper palea showing compound papillae (X 2000). All photomicrographs based on Eiten 6752 (US). 60 SMITHSONIAN CONTRIBUTIONS TO BOTANY uted evenly over the surface. Caryopsis ellipsoid, do Cipo, along the road at km 134-1 33, Eiten 2.2 nini long, 1.1 nim \vide, brown; hilum linear, 6752 (US). PARA:A riramba, Ducke 8015 (US). ;IS long ;is the caryopsis, embryo -’h as long as DrscussroN.-Although it is most common in the caryopsis. Panicum for the spikelet to be even on the pedi- ~rsTRI~L~~rIoN.-~ndeniicto the “campos ru- cel, it is sometinies twisted as in the case of P. pestres” of Brazil in the states of Para, Goias, hirtum, P. cervicatum, and P. rudgei. In these Minas Geixis, and Mato Grosso, in rocky or sandy species, however, the pedicel is claviform and soils at elevations of 800-1 400 m: occasionally truncate at the apex and there is no callus. burned over. In the genus Plagiantha Renvoize the pedicel ADDITIONALSPECIMENS SEEN.-BRAZIL. is also truncate and the spikelet lacks a callus at BAH~A:near 6 km N of the town of Rio de Contas its base, hvitli the spikelets disposed in oblique on road to Abaira, Harley et al. 15085 (MO, KY, form in the panicle. The morphology of this US). GOIAS:near 33 km S of Caiaponia on road genus is therefore similar to that of the species to Jatai, Irwin 3 Soderstrom 6959 (US). MATO of Panicum mentioned above. GROSSO:Fazenda Capa0 da Taquara, Hatschbach Tt4.o specimens from Mato Grosso (Hatschbach 32469 (RIO); Faxenda Urubu Branco, Pires @ 32469 and 33145) are about 1 111 tall, exception- Silva 11318 (US); 7 Quedas, Hatschbach 33145 ally robust, possess broad leaves that are flat ($10, NY, US). MINAS GERAIS:Estrada Concei- throughout, and have lax and diffuse panicles. $50 do Cerro, Occhioni s/n., Dec 1940 (US): Serra W’e consider these to represent extremes in the do Cipo, perto do Hotel Palacio, Black 3 Magal- variation of the species, as in smaller specimens hies 51-12039 (NY, US); Chapeo de Sol, Serra (Irmin 3 Soderstrom 6959, for example) small and do Cipo, Chase 9191 (US), 9207 (NY, US): Serra involute blades occur along with larger flat ones. Literature Cited

Amshoff, G.J.H., and J.Th. Henrard tral American Grasses. Canadian Journal ofBotany, 1948. Gramineae. In A. Pulle, Flora of Suriname, 501273-283. 1( 1):273-442. 1972b. Chromosome Numbers, Meiotic Behavior and Baum, B.R. Notes on Some Grasses from Central America and 197 1. Additional Taxonomic Studies on Avena Fatuoids: the West Indies. Canadian Journal of Botany, Some Morphological Attributes Seen Using the 50:1441-1452. Scanning Electron Microscope. Canadian Journal 1974. Chromosome Numbers, Meiotic Behavior and of Botany, 49:647-649. Notes on Tropical American Grasses (Gramineae). Baum, B.R., and V.E. Hadland Canadian Journal of Botany, 52:317-328. 1973. A Scanning Electron Microscope Study of Epicu- 1978. Chromosome Numbers of Tropical American ticular Waxes of Glumes in Avena magna, A. mur- Grasses (Gramineae). Annals of the ,Missouri Botan- phyi and A. sterilis. Canadian Journal of Botany, ical Garden, 65(2):637-649. 51:2381-2383. Desvaux, A.N. Bentham, G. 1810. Extrait d'un memoire sur quelques nouveaux 1881. Kotes on Gramineae.Journal of the Linnean Society, genres de la famille des Graminees. Nouveau Bul- Botany, 19: 14- 1 34. letin des Sciences par la Societe Philomatique de Paris, Bentham, G., and J.D. Hooker 2:187-190. 1883. Genera Plantarum . . . . Volume 3. London: L. 1813. Memoire sur les Graminees, et sur quelques Reeve 8c Company. genres et especes nouvelles de cette famille. Jour- Beurling, P.J. nal de Botanique, 1:65-77. 1854. Bidrag till Portobellos Flora. Kongliga Svenska Ve- Fournier, E. tenskapsakademiens Handlingar, 1854:11 2-1 75. 1886. Gramineae. In Mexicanas Plantas. . . , 2: 160 Blake, S.T. pages, 7 plates. Paris: Imprimerie hationale. 1958. New Criteria for Distinguishing Genera Allied to Gardner, C.A., and C.E. Hubbard Panicum. Proceedings of the Royal Society of Queens- 1938. Brachiaria occidentalis Gardner et C.E. Hubbard. land, 70(3)'15-19. In Hooker's Icones Plantarum. . . , plate 3363. Lon- Brown, W.V. don. 1977. The Krani Syndrome and Its Subtypes in Grass Could, F.W., and T.R. Soderstrom Systematics. Memoirs of the Torrey Botanical Club, 1970. Chromosome Numbers of Some Mexican and Co- 23(3): ii + 97 pages. lombian Grasses. Canadian Journal of Botany, Butzin, F. 48: 1633-1639.

1970a. Die Blattnervatur der Paniceae in ihrer Bedeu- Grisebach, A.H.R. ' tung fur die systematische Gliederung dieser Tri- 1853. Gramineae. In F.K. Ledebour, Flora Rossica, sive bus. Willdenowia, 6( 1): 1.67- 178. enumeratio plantarum in totius imperii rossici prov- 1970b. Die systematische Gliederung der Paniceae. Willd- inciis europaeis, asiaticis et americanis hucusque ob- enowia, 6(1):179-192. servatarum. Volume 4. Stuttgart. Chase, A. 1864. Catalogus plantarum Cubensium, exhibens collecti- 1908. Notes on Genera of Paniceae, 111. Proceedings of onem Wrightianam aliasque minores ex Insula Cuba the Biological Society of Washington, 21:175-188. Missas. 301 pages. Leipzig: W. Engelmann. 19 1 1. Notes on Genera of Paniceae, IV. Proceedings of Hitchcock, A.S. the Biological Society of Washington, 24: 103-1 60. 1913. Mexican Grasses in the United States National Clark, C.A., and F.W. Could Herbarium. Contributions from the Cnited States 1975. Some Epidermal Characteristics of Paleas of Di- ivational Herbarium, 17(3): 389 pages. chanthelium, Panicum and Echinolaena. American 1922. Grasses of British Guiana. Contributions from the Journal of Botany, 62(7):743-748. United States 6ational Herbarium, 22(6): Davidse, G.W., and R.W. Pohl 439-514. 1972a. Chromosome Numbers and Notes on Some Cen- 1927. The Grasses of Ecuador, Peru and Bolivia. Contri-

61 62 SMITHSONIAN CONTRIBUTIONS TO BOTANY

butzonsfrom the C.nited States R’ational Herbarium, h‘ees \‘on henbeck, C.G.D 24(8): v + 291-356 + xx. 1829. Agrostologia Brasiliensis . . . . 608 pages. Stuttgart 1930a. The Grasses of Central America. Contributions and l‘iibingeii. [This represents volume 2, part 1,

(7 .. from the C’nited States ‘Yational Herbarium, 71 ariiineae, of Flora Brasiliensis seu Enumeratio 2 4(9): .5 5 7 - 7 6 2, . . . , edited by C.F.D. von Martius.] 1930b. Four Neil. Grasses. Journal ofthe lthshington Acad- Palisot de Beauvois, ,,I.M,F,J, emj ofSciences, ‘20(1.5):381-384. 18 12. Essai d’une nouuelle agrostographie; ou nouveaux 193,j. Manual of the Grasses of the United States. L‘nited genres des Graminees; avec figures representant les States Department of.4gricultui-e .%liscellaneous Pub- caracteres de tous les genres. 184 pages [text], 25 lication, 200: 1040 pages. [ Revised 195 1 by Agnes plate5 [atlas]. Paris: Fain. Cliase, second edition, 105 1 pages.] Parodi, I..K. Hitchcock, A.S., and A. Chase 192.5. Notas sobre Gramineas de la flora Argentina. 191 0. The North Aniericari Species of Panicum. Contri- Phq’sis, 8:6 1-7 1, butions from the L‘nited States ‘Vational Herbarium, 1969. Estudios sistematicos sobre las Graniineae-Pani- 1 .i:1-396. cuc argentirias ? uruguayas. Darwiniana, 15( 1- I9 1.5. Tropical North Anierican Species of Panicum. 2):6.5- 1 1 1, (,‘ontributions from the tinited States ,l’ational Her- Pilger, R. barium, 1 7(6):4.59-539. 199 1, Bemerkungen ZLI Panicurn und verwandten Gat- HSU,C.-C. tungeri. Sotizblatt des Botanischen Gartens und ,Wu- 196.5. The Classification of Panicum (Gramineae) and Its seum2s zu Berlin-Dahlem, 104(1 1):237-247. Allies, with Special Reference to the Characters 1940. C;ramineae 111: Unterfani. Panicoideae. In Engler of l.odicule, Stylebase and Leninia. Journal of the and Prantl, editors, Die A’aturlichen P’anzenfamz- Facultj of Science, L‘nzversitj of Tokyo, section 3 lien, second edition, volume 14 [ 1 volume], 208 (Botall)), 9(3):4?J-150. pages, illustrated. Leipzig. Huniboltit, F.H.A. von, A.J. Bonpland, and C.S. Kunth 19.54. Das System der Graniineae unter .Ausschlus der 18 16. ,\‘ova genera et species plantarum . . . . \‘olume 1, Banibusoideae. Botanische Jahrbucher fur Systema- 302 pages. Paris. [This ivork consists of 7 volumes, tik, I1flan:engeschichte und Pflanzengeographie, published bet\veen 18 16 and 1825, which coni- 76(3):281-384. prise part 6 (botany), section 3, of Humboldt and Pohl, K.\V. Bonpland, editors, L‘ojage aux regions eyuinoctiales 1980. Graiiiineae, In LV. Burger, Flom Costaricensis. du noz’eau continent,fait en 1799-1804.j Fieldiana: Botany, 4: 608 pages. Jacq ues-Felix, H. Pohl, R.\V. aiid (;. Davidse 196‘2. l.es Graniinees (Poaceae) d’Afrique tropicale, I: 197 1. Chromosome Numbers of Costa Rican Grasses. (ekelittes, classification, description des genres. Brittonia, 23(3):293-324. Iristitut de Recherches Agronomiques Tropicales et des Presl, K.B., editor (:ultures I.’ivrie.res, Bulletin Scientijque (Paris), 8: xi 1890. Graniineae. In Reliquiae Haenkeanae . . . , l(4- + 34.5 pges. .5): ‘2 07 - 3 .iI,. Prague: J .G . Gal ve . liunth, <:.s, Richard, I\. 1829. Re\ision des Graniinees . . . . In Huniboldt and 185.1. \’ascular Plants. In R. de la Sagra, Flora Cubana, Bonpland, editorb, Voyage aux regions equinoctiales o descripcion botanica, usos y aplicaciones de las du noueau continent, fait en 1799-1804, part 6 plantas reunidas en la isla de O’uba. Volume 3. (Botany), section 6, pages. Paris: Librairie- P;ii.is: Libreria de la Viuda de Arthus Bertrand. <;ide. SKlldU~S~\,T. hlartius, <:.F.P. voii, editor 1978. Brachiaria: Taxonomy of Cultivated and Kative 1877. Tribe 3, P;iniceae. In Flora Brasiliensis, 2(2):33- Species in BIa/il. Hoehnea, 7:99-139, 16 figures. 342, plates 12-49 (fascicle 72). Munich and \’i- Scholi., H. enna. [ 1 .i\oIunies (in 40) published in 130 fasci- 1978. Flaschenformige Slikrohaare in der Gattung Pan- cles betWKe11 1840 alld 1906.1 icum (G~aniineae).Il’ildenowia, 8:5 1 1-5 15. hleyer, G.F.\V. Sliaw, K.B.. and F.E. Seiins 18 18. Primitiae florae essequeboensis adjectis descriptioni- 1979. Epidermal Chai-acteristics of the Callus in Erio- bus centum circiter stirpium novarum, observationi- rhloa (P~xEIK). American Journal of Botany, busyue criticis. 3 I6 pages, 2 plates. Gottingen. 66(8):907-913. Me/, <:. SIia\\, K.B., and K.D. \Vebster 192 1. Neue Gr;iiiiineen. In H.G.A. Engler, Botanzsche I98 1, \‘ariation aiid the Taxonomic Significance of .An- Jahrbucher. . . , 56(4)(supplenient 125):1-12. theciuni Characteristics in Ichnanthus (Poaceae: NUMBER 59 63

Pariiceae). Botanical Societj of America, dtliscella- 1.52 + 6 [index]pages. Stockholm: M. Swederi. neous Series, 160:78. Terrell, E.E., and M'.P. M'ergin Svith. L.B.. and D.C. M'asshausen 1979. Scanning Electron Microscopy and Energy Dis- 1978. Notas sobre Gramineas brasileirds, I. Bradea, persive X-ray Analysis of Leaf Epidermis in Ziza- 2 (3.5):2 4 9- 2 .i0. nia (Gr~~tiiiiieae).Scanning Electron .\.licroscopy, Sprengel, I\. 1979(3):81-88. 1825. (iaroli Linnaei , , , , sytema vegetabilium, editio de- 198 1. Epidernial Features and Silica Deposition in Lern- rima sexta. \'olume 2, 939 pages. Gottingen. [The mas and :\\\.tis of Zizania (Gramineae). American coniplete ~vorkconsists of 3 volumes, published Journal of Botanj, 68(.i):697-707. bettveer 1825 and 1828.1 Thomasson, J.R. Sqf,0. 1978a. Epidermal Patterns of the Lemma in Some Fossil 191 7-1 930. (;ramineae. In D. Prain, editor, Flora of atid Living Grasses and Their Phylogenetic Sig- Tropical ..?frica, 9: 1 132 pages. London: L. Reeve. nificance. Science, 199:975-977. Stant, k1.Y. 1978b. Observations on the Characteristics of the Lemma 1973. Scanning E:lectron Xlicroscopy of Silica Bodies and and Palea of the Late Ceno7oic Grass Panicum Other Epidermal Features in Gibasis (Tradescan- elegans. .4merican Journal OfBotanj, 65(1):34-39. tza) l.eaf. Botanical Journal of the Linnaean Societ)), Trinius, C.B. 6 6(:3): 2 3 3 - 2 44. 1820. Fundamenta agrostographiae siue . , . . 208 pages, Steudel, E.(;. 18 plates. L'ienna: J.G. Heubner. S)nopsis plantarum Graminearurn. In Synopsis 1826. De Graminibus Paniceis: Dissertatio botanica altera. plantarum Glumacearum, 1 : 475 pages. Stuttgart: 29 1 pages. St. Petersburg: Impensis Academiae J.B. hIet/ler. [This book appeared between 1853 I mperialis Scientiarum. and 18.ii: pages 1-80 in 1853, 81-400 in 1854, 182.5-1 836. Species Graminurn iconibus et descriptionibus and 40 1-475 in 1855.1 illustrauit. 3 volumes, 360 plates. Petropoli. Stieber. 11.l'. 1834. Panicearum genera retractauzt speciebusque complu- 1982. Revi\iori of Ichnanthus sect. Irhnanthus (Grami- ribus illustravit. 267 pages. [The copy in the nex, I'anicoideae). Systematic Botany, 7(1): Smithsonian Libraries, dated June 1834, in pencil, 85- I 15. is paginated from 1 through 267, and the title Swallen. J.K. page indicates neither cit) nor publisher. It is a 1943. Graniineae. In R.E. Woodson, Jr., and R.Lt'. prept-int of the paper jvith the same title that was Sc.lier>, Flora of Panama. Annals of the 'Missouri published in 183.5 in 'Li'emoires de l',lcademie Im- Botanical Garden, 30:104-280. periale des Sciences de Saint-Petersbourg, sixth series 19.5.5. Flora of Guatemala, Part 11: Grasses of Guateniala. (Sciences Saturelles), 1 :89-335.] FiPldiana: Botany, 24(2): 390 pages. Trinius,J.B. 1964. New South American Ichnanthus. Phytologia, 1836. Gratiiinuni in America Calidiore. Linnaea, 1 1 (:+): 14.;- 1 .i1. 10:291-308. Swllen. J.R., and H. Garcia Barrigd Zuloaga, F.O. 194:1. Fi\e New Crrasses from Colombia. Caldasia, 1978. Una nueva seccion del genero Panicum (Granii- 2(8):301-306. neae). Hickenia, l(29):1 .56- 159. s\vart/, 0.P. 1981. Las especies argentinas del genero lchnanthus 1788. lYova genera L+ species plantarum x+ (Gramineae). Daminiana, 23( 1): 189-22 1.