CONTRIBUTIONS FROM THE UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 17 27 April 1990 Pooh ae bx gti ee eee sig? Cay ie ide J sy +2 SOE TS Ry te ee i a le CwWws N Arbor Wright/Ann Larry E TEWART ~ re RALPH R. CONTRIBUTIONS FROM THE UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 17 University of Michigan Herbarium Ann Arbor, Michigan 27 April 1990 CONTRIBUTIONS FROM THE UNIVERSITY OF MICHIGAN HERBARIUM ISSN 0091-1860 Copyright 1990 University of Michigan Herbarium All rights reserved Printed in the United States of America Volume 17 Editor: Christiane Anderson For information about the availability and prices of previous volumes of the Contributions, address the Director, University of Michigan Herbarium, North University Building, Ann Arbor, Michigan 48109-1057, U.S.A. 27 April 1990 CONTENTS Dedication to Ralph R. Stewart WILLIAM R. ANDERSON Ralph R. Stewart MICHAEL G. PRICE Seven New Species of Stigmaphyllon (Malpighiaceae) from Brazil HRISTIANE ANDERSON The Taxonomy of Jubelina (Malpighiaceac) WILLIAM R. ANDERSON Ill WILLIAM R. ANDERSON Notes on Neotropical Malpighiaceae An Evaluation of Familial Limits among the Genera Traditionally Aligned with the Thuidiaceae and Leskeaceae WILLIAM R. Buck & HowarD CRUM Comments on Sphagnum sect. Sphagnum in South America OWARD CRUM A New Look at Sphagnum sect. Acutifolia in South America OWARD CRUM Preliminary Notes on Sphagnum sect. Subsecunda in South America HowarpD CRUM Systematics of Henrya (Acanthaceae) THOMAS F. DANIEL New and Reconsidered Acanthaceae. III. Justicia THOMAS F. DANIEL New, Reconsidered, and Little-Known Mexican Species of Ruellia (Acanthaceae) THOMAS F. DANIEL New Species of Malvaceae from South America PAUL A. FRYXELL The Malvaceae Published by Turczaninow PauL A. FRYXELL & ANTONIO KRAPOVICKAS Botanical Results of the Sessé & Mocino Expedition (1787-1803). IV. The Library and the Herbarium of the Expedition RoGers McVAUGH The Taxonomy of Carex bromoides (Cyperaceac) Rosert F. C. Naczi Systematics of the Scutellaria angustifolia Complex ene ARD G. OLMSTEAD Philippine Fern Notes MICHAEL G. PRICE Four New Species of Carex (Cyperaceae) from Mexico, with Notes on the Mexican Carex Flora A. A. REZNICEK Notes on the Archaeinae and other Russulas RoBERT L. SHAFFER A Mexican Pine Promoted to Specific Status: Pinus praetermissa B. T. Styces & RoGeERs MCVAUGH Moonworts (Botrychium subg. ee Ophioglossaceae) of the pper Great Lakes Region, U.S.A. and Canada, with Descriptions ot Two New Species W. HL WAGNER, JR., & F. S. WAGNER Observations on Haraldia and Calloseris, Two Rare Genera of Delesseriaceae (Rhodophyta) from the Western Atlantic MICHAEL J. WYNNE Records and Notes on Alaskan Marine Algae. III MICHAEL J. WYNNE nie 295 307 Sto O27 335 Contr. Univ. Mich. Herb. 17:1—2. 1990. DEDICATION TO RALPH R. STEWART Taxonomy is an historical discipline. One of the goals of our research is to deduce the evolutionary history of the organisms we describe. In the more recent sense of history, we are dependent upon and constrained by the work of our predecessors of the last three centuries, and while one might claim the same for any branch of science, taxonomists are probably more conscious of that fact than any other group of natural scientists. A good taxonomist builds on the work of those who went before, beginning with their publications and trying to improve on them. Because our application of names is governed by the principle of priority, we must have access to and understanding of the publications and specimens of our predeces- sors, and in the process of trying to understand their work we inevitably become interested in them as people too. Our research forces us to study carefully the itineraries and other records of collectors, so that we can correctly identify the specimens they collected and the names they published, and it is no wonder that while studying their histories we marvel at the achievements of those intrepid travellers. For all these reasons, taxonomists collect, conserve, and consult the literature and specimens of our predecessors, and many of us take great interest in the lives of those who labored before us to describe and interpret the earth’s biological diversity. Another peculiarity of taxonomists is that our most substantial contributions often come late in life, especially in the form of synoptic works like monographs and floras. Taxonomic research is information-dense and involves many skills and traditions that are learned only gradually and only by those with the patience and foresight to undertake ambitious projects at the expense of immediate recognition. One has only to look at the publication records of many plant taxonomists to see the frequent repetition of a pattern of long series of relatively modest publications followed by acceleration toward major works in the fifth, sixth, and later decades of life. One of the joys of working at major herbaria is the opportunity to know, observe, and emulate some of those senior colleagues as they produce their great works. Such acquaintances have greatly enriched my own professional and personal life. They have served as models of solid scholarship and persistence toward goals that they knew to be worthwhile, regardless of whether or not they happened to be currently fashionable or in favor with administrators and bureaucrats dispensing money and other rewards. Those colleagues have forged a strong link with the past for me, giving life to our history and showing how I and my students can join their ranks and make our own contributions to taxonomy, the queen of the sciences. One of the most memorable plant taxonomists I have known is Dr. Ralph R. Stewart. When I came to Michigan as a graduate student in 1964, Dr. Stewart was already here, working on his monumental Annotated Catalogue of the Vascular Plants of West Pakistan and Kashmir. Throughout the years of my graduate study I came to know and admire him, and I was delighted to find him still here and active when I returned as a staff member in 1974. Until his second retirement in 1981, Dr. Stewart was one of our most regular and diligent researchers. Aside from his botanical achievements, which are described in the following sketch by Michael Price, Dr. Stewart was and is a good companion, with a sharp mind and a quick wit that suffers neither fools nor foolishness. His has been a life of productive service. 2 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 17 He embodies much that is good in plant taxonomy, and he also, as of this year, embodies a full century of our history. It therefore gives me great pride and plea- sure, as director of the University of Michigan Herbarium, to dedicate this volume of our Contributions to Dr. Ralph R. Stewart in celebration of his one-hundredth birthday on 15 April 1990. : William R. Anderson Contr. Univ. Mich. Herb. 17:3-5. 1990. RALPH R. STEWART Dr. Ralph R. Stewart worked at the University of Michigan Herbarium with the title of Research Associate from 1960 to 1981, and during those years produced the two volumes for which he is now most heralded, An Annotated Catalogue of the Vascular Plants of West Pakistan and Kashmir, which treated 5783 taxa, and History and Exploration of Plants in Pakistan and Adjoining Areas, as well as a dozen other substantial botanical papers, and identified an enormous number of unnamed bo- tanical specimens from the Indian subcontinent. He arrived in Ann Arbor at the age of 70, in 1960, at the invitation of Dr. Rogers McVaugh, at which time the backlog of undetermined Himalayan specimens exceeded 30,000. The vast majority were collected by T. R. Chand and W. Koelz with the support of Prof. H. H. Bartlett of the University of Michigan. Dr. Stewart’s Michigan years were especially significant, because he was able to concentrate on botany more than at any other prolonged period. After graduating with honors from Columbia University in 1911, he accepted a position teaching biology at Gordon College in Rawalpindi, now in Pakistan, but then the major northern fortified frontier city of British India. Soon he began serious plant collecting, securing specimens from most parts of the western Hima- layas and all districts of what is now Pakistan over the next fifty years. The 60,000 mounted specimens he amassed, mainly his own collections, but also those of his students, and from exchanges and gifts, now form the nucleus of the National Herbarium of Pakistan in Islamabad, which is named the Stewart Collection. Out- side of Pakistan the most substantial holdings of Dr. Stewart’s plants are at the New York Botanical Garden, the Smithsonian Institution, and the Royal Botanical Gar- dens at Kew. Until the present Dr. Stewart continues to collect and study plants on vacations and visits, and in the vicinity of his current retirement home in California. n 1914, Dr. Stewart returned to Columbia University as an Assistant in the Botany Department, and obtained a Ph.D. in Botany in 1916 under the professor- ship of N. L. Britton. His thesis on the Flora of Western Tibet was published in the Bulletin of the Torrey Botanical Club. Also in 1916, he married the botanist Isa- belle Darrow, sister of Dr. George M. Darrow, who later achieved fame as a pomologist for the U.S. Department of Agriculture. Isabelle was a fellow student at Columbia, earned an M.S. in plant breeding, and until her death in 1953, she was usually his companion even in the field, on collecting trips. From 1917 to 1960, Dr. Stewart taught botany and other subjects at Gordon College, as Instructor, Vice-principal, and Principal. His legacy includes the many pupils he inspired, several of whom have gone on to eminent botanical careers, notably Prof. E. Nasir, co-editor with Dr. S. I. Ali of the Flora of Pakistan. In 1955, he was obliged to retire as Principal due to age restrictions, but continued in a supporting role for another five years. Dr. Stewart married again, in 1954, to Hladia Porter, a poet, painter, and teacher, daughter of an earlier Principal of Gordon College, and sister to the plant taxonomist Cedric L. Porter, especially noted for his work on the flora of Wyoming. Hladia died in 1984. During furloughs, Dr. Stewart devoted his efforts to identifying his plant collec- tions at the New York Botanical Garden, where he spent one year (1932-33) working at the side of Elmer D. Merrill, who achieved for Philippine botany what 4 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 17 R. R. Stewart has for Pakistan. Prevented from returning to Rawalpindi from a home visit by the outbreak of World War II, Dr. Stewart accepted the post of Curator of the Oriental Herbarium at the New York Botanical Garden, and worked there continuously for 2% years. Dr. Stewart spent fourteen summers of his Michigan period at the Royal Botani- cal Gardens at Kew, England, with side trips to the British Museum and the Royal Botanic Garden at Edinburgh, and became the undisputed authority on the flora of Pakistan and the Western Himalayas; he is better known in Asia than in his home country. Two ferns have been named in his honor, Dryopteris stewartii Fraser- Jenkins and Lepisorus stewartii Ching, and he discovered many new species of flowering plants, some of which are named after him. Dr. Stewart attributes his longevity only partially to a good heritage; other factors include frequent outdoor activity, a consistent work schedule, and refraining from excess. He 1s in position to challenge the longevity records for botanists held by the Canadian mycologist John Dearness (1852-1954) and the British taxonomist famous for his study of the Malayan flora, Henry N. Ridley (1855-1956). Dr. Stewart continues to be active, lecturing and campaigning for environmental causes, and writing a botanical column for a monthly journal, Gleanings, in Duarte, California. He has recently authored a semi-technical booklet on the woody plants, including those in cultivation, of Westminster Gardens in Duarte, and a companion volume on herbaceous plants is near completion. Because of his diverse activities, his correspondence is very extensive, and he has received as many as 61 pieces of mail in a single day. All of us at the University of Michigan Herbarium wish him many more years of good health, good friendships, and good botany. Michael G. Price BOTANICAL PUBLICATIONS OF RALPH R. STEWART = Some observations on the Flora of the Northwest Himalaya. Torreya 15(12): 251-260. 1915. . The Flora of Ladak, Western Tibet. I. Discussion of the Flora. II. List of Ladak plants. Bull. Torrey Bot. Club 43: 571-590. 1916; 625-650. 1917. 3. Alpine plants of Kashmir. Torreya 24(3): 41-44. 1924. 4. Studies in the Ice Age in India. Carnegie Institute, 1939 (with H. de Terra & Patterson). The Ferns of Mussoorie and Dehra Dun. 150th Annual Volume of the Royal Botanic Gardens. Calcutta. Pt. 2: 159-172. 1942. 6. Alpine flowers of Kashmir. Bull. Amer. Rock Garden Soc. Jan.—Feb. 1943: 20- 24 io) wa 7. The Scrophulariaceae of the Western Himalayas. Philad. Acad. Sci. Monog. 5: 1-163. 1943 (with F. W. Pennell). 8. Vegetation of Kashmir Valley and Lakes. J. New York Bot. Gard. 45: 199-207. 1944. 9. The Grasses of Northwest India. Brittonia 5: 404-468. 1945. 10. Ferns of Kashmir. Bull. Torrey Bot. Club 72: 399-426. 1945. 11. Notes on the Flora of Kashmir. Punjab (Pakistan) For. Records 2: 1-13. 1950. 12. Grasses of Rawalpindi District. Punjab (Pakistan) For. Records 3: 1-24. 1951. 13. Ferns of Pahlgam, Kashmir. J. Indian Bot. Soc. 30: 137-142. 1951. 1990 PRICE: RALPH R. STEWART 5 b> — mn an ww es es) oe aw =) ja N 4, Catalog of the plants of Rawalpindi District. Pakistan Forestry Dept., 1-90. 1952: . Plant collecting in Kashmir. Pakistan J. Sci. 4: 89-93. 1952 (excerpted in The Women’s Missionary Magazine, June 1952: 601-602). . A bibliography of the flowering plants of West Pakistan and Kashmir. Biologia (Pakistan) 2: 221-230. 1956. . The wild flowers of Swat. Pakistan Quarterly 6: 36-40. 1956. . Materials for a revision of Parker’s Forest Flora. Pakistan J. Forest., Jan. 1957: 9 Seal . The ferns and fern allies of West Pakistan and Kashmir. Biologia (Pakistan) 3: 133-164. 1957 (reprint pagination 1-32). . The Flora of Rawalpindi District, West Pakistan. Pakistan J. Forest., Oct. 1957: 237-300; Jan. 1958: 13-111 (also reprinted by R. R. Stewart, i-x + 163 p). . Quetta and Baluchistan. Pakistan Quarterly 8: 20-26. 1958 (with Hladia P. Stewart). . The Flora of Baluchistan. Pakistan Quarterly 8: 27-30. 1958. . Grasses of West Pakistan. Biol. Soc. Pakistan, Monog. III, 388 p. 1958-59 (with Sultan Ahmad). . Additions and corrections to the Rawalpindi District Flora. Pakistan J. Forest. 11: 51-63. 1961 (reprint pagination 1-13). . The Flora of the Deosai Plains. Pakistan J. Forest. 11: 225-295. 1961. The Flora of the Kurram Valley (W. Pakistan). Biologia (Pakistan) 13: 47-124. 1967. . Plant collectors in West Pakistan and Kashmir. Pakistan J. Forest. 17: 337-363. . Check list of the plants of Swat State, Northwest Pakistan. Pakistan J. Forest. 17: 457-528. 1967. . The grasses of Kashmir. Bull. Bot. Surv. India 9: 114-133. (1967) 1968. . The Cyperaceae of Kashmir—A check list. Bull. Bot. Surv. India 9: 152-162. (1967) 1968. . The grasses of West Pakistan and Kashmir. Biologia (Pakistan) 16: 1-46. 1970 . An annotated catalogue of the vascular plants of West Pakistan and Kashmir. Flora of West Pakistan. Karachi, Dec. 1972. xviii + 1028 p. . Validation of some new names of Pakistan Plants. Kew Bull. 29: 443-444. 1974. . The first plant collectors in Kashmir and the Punjab. Taxon 28: 5-11. 1979. 5. Missionaries and clergymen as botanists in India and Pakistan. Taxon 31: 57— 64. 1982. History and exploration of plants in Pakistan and adjoining areas. Flora of Pakistan. Islamabad, March 1982. 186 p. . The Flora of Panjar and Karot Valleys, Kahuta Tehsil, Rawalpindi District. Pakistan J. Bot. 14: 131. 1982. Pteridophyte genera—The meaning of their names. Fiddlehead Forum, Bull. Amer. Fern Soc. 10: 21-36. 1983 (with David M. Johnson & John T. Mickel). . How did they die? Taxon 33: 48-52. 1984. . Remarks on North Western Himalayan ferns. Indian Fern J. 1: 41-45. 1984. . Pleistocene plant fossils of the Lower Karewa Beds of Kashmir. M. B. Raizada Commemorative Volume 1-4. 1985. _ Trees, shrubs and vines in Westminster Gardens. Duarte, CA, 1985. 59 p. ; - 7 Contr. Univ. Mich. Herb. 17:7—19. 1990. SEVEN NEW SPECIES OF STIGMAPHYLLON (MALPIGHIACEAE) FROM BRAZIL Christiane Anderson University of Michigan Herbarium North University Building Ann Arbor, Michigan 48109-1057 Since the publication of Niedenzu’s treatments of Stigmaphyllon (1899, 1900, 1928), one of the wing-fruited genera of the Malpighiaceae, many new collections from South America have shown this Neotropical genus of yellow-flowered vines to be larger and more diverse than previously thought. Seven new species from Brazil are here described. They all exhibit the characters shared by most species: large and long-petioled leaves, flowers aggregated into umbels or pseudoracemes, a hetero- morphic androecium with the stamens opposite the sepals modified, and styles bearing the apical foliole for which the genus is named. The samaras of the three species for which fruiting material is available also agree with the form typical for the genus, a nut bearing a large dorsal wing and often also lateral winglets. Stigmaphyllon bradei C. Anderson, sp. nov.—TyPe: BraziL. Sao Paulo: Registro, 21 Jun 1963, Moura s.n. [SP123438| (holotype: SP!). Liana. Laminae 6.5—14.3 cm longae, 6.8-12.5 cm latae, cordatae vel suborbi- culares, supra glabrae, subtus glabrae vel nervis sparsim sericeae, margine sparsim glanduloso; stipulae glandulosae. Inflorescentia solitaria (vel dichasialis?) constata ex umbellis, floribus in quaque umbella ca. 15-25. Pedunculi rudimentales vel usque 1.5 mm longi; pedicelli 8—13.5 mm longi. Petala limbo orbiculari; limbi petalorum antico-lateralium marginibus erosis vel irregulariter denticulato-erosis; limbi petalorum postico-lateralium et petali postici marginibus fimbriatis. Stamina heteromorpha; antherae sepalis lateralibus oppositae loculis redactis instructae vel steriles, antherae ceterae fertiles; antherae glabrae. Stylus anticus 2.6-3.2 mm longus, sparsim pubescens, utroque foliolo 1.2—-1.4 mm longo, 1.1-1.2 mm lato, triangulari vel parabolico; styli postici 3.3-4 mm longi, lyrati, glabri vel sparsim pubescentes, foliolo 1.8-2 mm longo latoque, subquadrato vel suborbiculari. Sa- mara ignota. Vine. Laminas 6.5—14.3 cm long, 6.8-12.5 cm wide, cordate to suborbicular, apex mucronate in larger leaves to caudate in smaller ones, base auriculate in larger leaves to cordate in smaller leaves, glabrous above, glabrous or sparsely sericeous only on major veins below, margin with scattered stalked glands (0.2—-0.3 mm in diameter, 0.3-0.6 mm long) and/or filiform glands (up to ca. 2 mm long), with a pair of prominent but sessile glands borne on the petiole 1-2.5 mm below the base of the lamina, each gland 1.5—1.8 mm in diameter, up to 0.5 mm high; petioles 2.8—7 cm long, pubescent in younger leaves to glabrate in older and larger ones; stipules 1.3— 2.5 mm long, 0.9-1.6 mm wide, the proximal % a circular gland, the distal 7 herbaceous and narrowly triangular to linear. Flowers ca. 15-25 per umbel, the umbels borne solitary (and in dichasia?). Peduncles rudimentary to 1.5 mm long; pedicels 8—13.5 mm long; peduncles up to 0.1 times as long as the pedicels. Bracts a 8 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 17 ca. 1.5—2.5 mm long, ca. 1.5—2 mm wide, broadly triangular, bracteoles ca. 1.3—2 mm long, 1.1—1.3 mm wide, broadly triangular, eglandular. Sepals ca. 2.2-2.4 mm long, ca. 2.5-2.7 mm wide, glands ca. 2.5-2.8 mm long, 1.3—1.5 mm wide. All petals with the limbs orbicular, glabrous, those of the anterior-lateral petals with the margin erose or irregularly denticulate-erose, the teeth up to 0.3 mm long, those of the posterior-lateral and posterior petals with the margin fimbriate, the fimbriae up to 0.6 mm long; anterior-lateral petals: claw 3-3.2 mm long, limb ca. 11 mm long and wide, posterior-lateral petals: claw 2.5—-3 mm long, limb ca. 10 mm long and wide; posterior petal: claw 3.2—4 mm long, apex very slightly indented, limb ca. 7.5— 8 mm long and wide. Stamens unequal, those opposite the posterior styles the largest, anthers of those opposite the lateral sepals with the connective enlarged and the locules reduced or absent; anthers glabrous. Anterior style 2.6-3.2 mm long, shorter than the posterior two, erect, with scattered hairs in the proximal 4; each foliole 1.2—1.4 mm long, 1.1-1.2 mm wide, triangular to parabolic. Posterior styles 3.3-4 mm long, lyrate, glabrous or with scattered hairs in the proximal 4; foliole 1.8-2 mm long and wide, subsquare to suborbicular. Samara not seen. Fig. 1. ADDITIONAL SPECIMEN EXAMINED. BRAZIL. Sao Paulo: Iguape, Morro das Pedras, 1921, Brade 7987 ): Stigmaphyllon bradei, known only from two flowering collections from coastal Sao Paulo, is readily distinguished by its unusual stipules, each bearing a basal at) ey l Ly, yay ge Y FIG. 1. eieaply lon bradei. a. Flowering branch (x0.5) and enlargement of stipules (5). b. Large leaf (0.5). c. Detail of base of umbel (x5); the base of each pedicel is hidden by two bracteoles and one bract: the te and ans are in such close association because the peduncle is ru tary. d. Posterior petal (2.5). e. Androecium (x5); second stamen from left opposes the oe petal. f. Gynoecium (x5); ee styles bent slightly outward to show anterior style, which is in the middle. (a, c-e based on Moura s.n. [SP123438]; b based on Brade 7987.) 1990 C. ANDERSON: STIGMAPHYLLON 9 gland (Fig. 1a), and by its inflorescence structure. The flowers are aggregated into congested umbels, which are borne singly (and perhaps in dichasia) on long primary axes. The peduncles, often rudimentary, are at most 1.5 mm long, so that the bract and the pair of bracteoles (borne at the apex of the peduncle) are in close proximity. Stigmaphyllon bradei is named for the noted Brazilian botanist Alexandre Curt Brade, who first collected it. Stigmaphyllon hatschbachii C. Anderson, sp. nov.—Type: Brazi_. Minas Gerais: Realeza, orla da mata, 15 Oct 1983, Hatschbach 46863 (holotype: MBM!; isotype: MICH!). Liana. Laminae ca. 10 cm longae, 6—7 cm latae, anguste ovatae vel ellipticae, supra glabrae vel glabratae costa pilis T-formibus instructa, subtus densissime pu- bescentes pilos T-formes aureos ferentes, margine sparsim glanduloso. Inflores- centia thyrsiformis constata ex umbellis, floribus in quaque umbella ca. 20-30. Pedunculi 7-9 mm longi; pedicelli 7-10 mm longi. Petala limbo orbiculari, margine fimbriato. Stamina heteromorpha; antherae sepalis antico-lateralibus oppositae 1— 2 loculis redactis instructae, antherae sepalis postico-lateralibus oppositae steriles, antherae ceterae fertiles pubescentes. Stylus anticus ca. 3.5 mm longus, glaber, utroque foliolo ca. 1.8 mm longo, ca. 1.4 mm lato, parabolico; styli postici ca. 4mm longi, lyrati, glabri, foliolo ca. 2.7 mm longo latoque, suborbiculari. Samara ignota. Vine. Laminas ca. 10 cm long, 6-7 cm wide, narrowly ovate to elliptical, apex mucronate, base attentuate to truncate, glabrous to glabrate above but with T- shaped hairs on the midvein, very densely pubescent with golden T-shaped hairs below, irregularly spaced stalked glands (each ca. 0.3 mm in diameter, ca. 0.4—0.5 mm long) borne adjacent to the margin below, with a pair of prominent but sessile glands at the apex of the petiole, each gland 1.8—2 mm in diameter; petioles 2.5-3 cm long, densely pubescent with T-shaped hairs; stipules 0.5—-1 mm long and wide, triangular, eglandular. Flowers ca. 20-30 per umbel, the umbels borne in small thyrses. Peduncles 7-9 mm long; pedicels 7-10 mm long; peduncles 0.7—1 times as long as the pedicels. Bracts 2.8-3.2 mm long, 1—1.3 mm wide, narrowly triangular, bracteoles 1.8-2.2 mm long, I-1.2 mm wide, narrowly triangular, eglandular. Se- pals ca. 3 mm long, 2.3-2.5 mm wide, glands 1.8—2 mm long, ca. | mm wide. All petals with the limbs orbicular, glabrous, margin with fimbriae up to 0.5 (—0.6) mm long; anterior-lateral petals: claw ca. 1.5 mm long, limb 13-14 mm long and wide; posterior-lateral petals: claw 1-1.3 mm long, limb 11-12 mm long and wide; poste- rior petal: claw 3—3.5 mm long, apex not indented, limb ca. 10 mm long and wide. Stamens unequal, those opposite the posterior-lateral petals (and the posterior styles) the largest, anthers of those opposite the anterior-lateral sepals with the connective enlarged and bearing only 1 or sometimes 2 reduced locules, anthers of those opposite the posterior-lateral sepals sterile; fertile anthers pubescent. Ante- rior style ca. 3.5 mm long, shorter than the posterior two, glabrous; each foliole ca. 1.8 mm long, ca. 1.4 mm wide, parabolic. Posterior styles ca. 4 mm long, lyrate, glabrous; foliole ca. 2.7 mm long and wide, suborbicular. Samara unknown. Fig. 2. Stigmaphyllon hatschbachii, known only from the type, is easily recognized by the copious golden pubescence of most of the vegetative parts. The dense indumentum of the abaxial leaf surface obscures the epidermis and hides the briefly stalked marginal glands. The anthers of stamens opposite the anterior-lateral sepals bear only one (sometimes two) reduced locule and those of the stamens opposite the posterior-lateral sepals are sterile. 10 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 17 FIG. 2. Stigmaphyllon hatschbachii. a. Flowering branch and portion of large leaf (<0.5), and enlarged T-shaped hair (20). b. Detail of margin of abaxial leaf surface with some of the hairs ue to show position of gland (x10), se view of marginal gland (x10). c. Posterior petal (x Androecium (5); second stamen from left opposes the posterior petal. e. ee with anterior style to the right (5). (Based on pn en 46803.) This distinctive species is named in honor of the Brazilian botanist Gert Hatschbach, discerning collector of many novelties. Stigmaphyllon macedoanum C. Anderson, sp. nov.—Type: BRazi_. Minas Gerais: Capinopolis, Fazenda Santa Terezinha, as margens do Rio Paranaiba divisa de Goias e Minas Gerais, 27 Jan 1989, A. Macedo 5486 (holotype: RB!:; isotypes: G! K! MICH! NY! R! UB! US!). Liana. Laminae 9—18.5 cm longae, 6.3-19 cm latae ambitu, majores palmatim —5(—7)-lobatae vel late ovatae vel late ellipticae, minores 2—3-lobatae vel late ovatae, supra sparsim pubescentes vel glabratae, subtus sparsim sericeae, margine sparsim glanduloso. Inflorescentia dichasialis constata ex pseudoracemis, floribus in quoque pseudoracemo ca. 15-25. Pedunculi 2.5—4 mm longi; pedicelli 5.5—6 mm longi. Petala limbo orbiculari, margine fimbriato. Stamina heteromorpha; fila- menta sepalis lateralibus opposita longissima; antherae omnes fertiles pubescentes. Stylus anticus ca. 2.5 mm longus, utroque foliolo ca. 0.8 mm longo, 0.6—-0.7 mm lato, triangulari vel late triangulari; styli postici ca. 3 mm longi, parum lyrati, foliolo ca. 1.2 mm longo, ca. | mm lato, triangulari. Samara ala dorsali 1.8-2.1 cm longa, ca. 1.2 cm lata; alulae laterales absentes; nux ca. 7 mm alta. Vine. Laminas 9—18.5 cm long, 6.3—19 cm wide in outline, the larger palmately —5(—7)-lobed to broadly ovate or broadly elliptical, the smaller 2—3-lobed to broadly ovate, apex cuspidate, base deeply cordate to auriculate or in the smaller 1990 C. ANDERSON: STIGMAPH YLLON 1] sometimes sagittate, sparsely pubescent to glabrate above, sparsely sericeous be- low, margin with filiform glands (up to 1.5 mm long), these terminating the major veins, with a pair of shallowly cupulate glands at the apex of the petiole, each gland (.9-2 mm in diameter, ca. 0.5 mm high; petioles 4-15 cm long, sericeous, confluent across the node and forming a corky ridge bearing the stipules; stipules 0.2—0.5 mm long, 0.5-0.6 mm wide, triangular, eglandular. Flowers ca. 15-25 per pseudo- raceme, the pseudoracemes borne in dichasia or compound dichasia. Peduncles 2.5-4 mm long; pedicels 5.5-6 mm long, somewhat inflated; peduncles 0.4—0.7 times as long as the pedicels. Bracts 1.1-1.5 mm long, 0.6—0.9 mm wide, triangular; bracteoles 0.8—1 mm long, 0.5-0.8 mm wide, triangular, eglandular. Sepals ca. 2 mm long and wide, glands 1.8—2 mm long, ca. | mm wide. All petals with the limbs orbicular, those of the lateral petals with the margin fimbriate, fimbriae up to 0.4 mm long; anterior-lateral petals: claw 1.6-1.7 mm long, limb ca. 9.5 mm long and wide; posterior-lateral petals: claw ca. 1 mm long, limb ca. 7.5 mm long and wide; posterior petal: claw ca. 3 mm long, apex not indented, limb ca. 6.5 mm long and wide, margin with fimbriae up to | mm long. Stamens equal in shape but unequal in size, those opposite the lateral sepals with the longest filaments; anthers all fertile, glabrous. Anterior style ca. 2.5 mm long, shorter than the posterior two, glabrous; each foliole ca. 0.8 mm long, 0.6—0.7 mm wide, triangular or broadly triangular. Posterior styles ca. 3 mm long, slightly lyrate, glabrous; apex 1.5 mm long and incurved, foliole ca. 1.2 mm long, ca. 1 mm wide, triangular. Dorsal wing of samara 1.8-2.1 cm long, 1.2 cm wide, upper margin with a blunt tooth; lateral winglets absent; nut ca. 7 mm high, ca. 3.5 mm in diameter, areole plane, carpophore up to 2.3 mm long. Seed ca. 7.8 mm long, embryo ovoid, outer cotyledon ca. 6.6 mm long, ca. 4.2 mm wide, distally curved but not folded over the inner cotyledon, inner cotyledon ca. 6 mm long, ca. 3.8 mm wide, straight. Fig. 3. ADDITIONAL SPECIMEN EXAMINED. BRAziL. Minas Gerais: Ituiutaba, Capindépolis, Fazenda Santa Terezinha, ramosa de mata em Pedreiras, 22 Nov 1971, A. Macedo 5057 (MBM). Stigmaphyllon macedoanum, known only from one locality in Minas Gerais, is easily recognized by its polymorphic, sparsely pubescent leaves, whose petioles are fused across the node into a corky ridge bearing the tiny caducous stipules (Fig. 3b), a condition not found in any other species. All laminas are cuspidate, and the major veins terminate in filiform marginal glands. The petiole glands are shallowly cupulate. The flowers are arranged in pseudoracemes. The androecium is unusual in that the stamens opposite the lateral sepals have the longest filaments and unreduced or only slightly reduced locules. The small samara lacks lateral winglets. This unusual species is named for Engenheiro Agr6nomo Cassio Macedo at the request of his brother Amaro Macedo, who collected the type and paratype. Stigmaphyllon matogrossense C. Anderson, sp. nov.—TyPe: BRazi_. Mato Grosso: Mpio. Caceres, 9 km ENE de Porto Esperidiao, BR-174, 150 m, 18 May 1985, Krapovickas et al. 40114 (holotype: MICH!; isotype: CTES!). Liana. Laminae 5.5—9.6 cm longae, 3—5.8 cm latae, ovatae vel anguste ovatae, supra glabrae, subtus tomentosae, margine eglanduloso. Inflorescentia solitaria vel dichasialis constata ex umbellis, floribus in quaque umbella ca. 15. Pedunculi 3.5—4.5 mm longi; pedicelli 5—8.5 mm longi. Petala limbo orbiculari, margine glanduloso- fimbriato; limbus petali postici supra basin fimbriis crassis glandulosis instructus. VOLUME 17 : rs a CONTR. UNIVERSITY OF MICHIGAN HERBARIUM aGee ans) a Rent tal (<3). e. FIG. 3. Stigmaphyllon macedoanum. a. Flowering branch (0.5). b. Detail of node showing corky ridge formed by confluent bases of petioles (1.5); note stipule scars. ¢. Detail of abaxial surface of leaf wing sparse appressed pubescence and basal glands (2.5). d. Posterio second stamen from left opposes the posterior petal. f ee ote ae anterior style to the left (<5). g. Samara (<1). h. Embryo (X2.5). (Based on A. Macedo 5480. ) base show Androecium (x5) 1990 C. ANDERSON: STIGMAPHYLLON 13 Stamina heteromorpha; antherae sepalis postico-lateralibus oppositae steriles, an- therae ceterae fertiles glabrae. Stylus anticus ca. 3.5 mm longus, glaber, utroque foliolo ca. 1.7 mm longo, ca. 2.2 mm lato, subrectangulari; styli postici ca. 4.3 mm longi, lyrati, adaxaliter pubescentes, foliolo ca. 2.2 mm longo, ca. 2.7 lato, suborbiculari. Samara ignota. Vine. Laminas 5.5—9.6 cm long, 3-5.8 cm wide, ovate or narrowly so, apex mucronate or emarginate-mucronate, base cordate or shallowly cordate, glabrous above, tomentose below, margin eglandular, with a pair of prominent but sessile glands at the apex of the petiole, each gland 1.4—2 mm in diameter; petioles 1.3-4.5 cm long, sericeous; stipules 1-1.4 mm long, 0.8—1.3 mm wide, triangular, egland- ular. Flowers ca. 15 per umbel, the umbels borne solitary or in dichasia or com- pound dichasia. Peduncles 3.5—4.5 mm long; pedicels 5—8.5 mm long; peduncles 0.5—0.8 times as long as the pedicels. Bracts 1.5—2.2 mm long, 1.2-1.6 mm wide, triangular; bracteoles 1.6-2 mm long, 1.2-1.5 mm wide, triangular, eglandular or each bracteole with a pair of inconspicuous glands (each 0.1—0.2 mm in diameter). Sepals 2.8-3 mm long and wide, glands ca. 2.2 mm long, 1-1. mm wide. All petals with the limbs orbicular, glabrous, margin with gland-tipped fimbriae up to 0.5 mm long; anterior-lateral petals: claw 2-2.3 mm long, limb ca. 13 mm long and wide; posterior-lateral petals: claw ca. 1.5 mm long, limb ca. 10 mm long and wide; posterior petal: claw 3—3.2 mm long, apex indented, limb 9—9.5 mm long and wide, the margin at the base with 1-3 stout glandular fimbriae, these 0.6—0.7 mm long, ca. 0.2 mm wide. Stamens unequal, those opposite the posterior-lateral petals (and the posterior styles) the largest, anthers of those opposite the lateral sepals sterile; anthers glabrous. Anterior style ca. 3.5 mm long, shorter than the posterior two, glabrous; each foliole ca. 1.7 mm long, ca. 2.2 mm wide, subrectangular. Posterior styles ca. 4.3 mm long, lyrate, adaxially with a row of hairs in the proximal 4-4; foliole ca. 2.2 mm long, ca. 2.7 mm wide, suborbicular. Samara not seen. Fig. 4. Stigmaphyllon matogrossense, known only from the type, has the leaves tomentose below and flowers with several distinctive features. The limbs of the petals have glandular-fimbriate margins, but the limb of the posterior petal also bears 1-3 stout gland-tipped fimbriae on each side near the base. The stamens opposite the lateral sepals are sterile. The posterior styles only bear an adaxial row of scattered hairs in the proximal %4—¥ of their length. Stigmaphyllon paraense C. Anderson, sp. nov.—TypPe: BraziL. Para: Rio Pa- ranapebas, control point at entrance to Serra Norte, ca. 39 km E of AMZA camp N-5, 6°04’S, 49°55'W, ca. 150 m, Sperling 6322 (holotype: INPA!; isotypes: MICH! NY!). Liana. Laminae 9.2-17.5 cm longae, 5.5—11.5 cm latae, anguste triangulares vel triangulares vel anguste ovatae vel ovatae, supra sparsim sericeae vel glabrae, subtus sericeae, margine sparsim glanduloso. Inflorescentia dichasialis constata ex umbellis, floribus in quaque umbella ca. 12-20. Pedunculi 4—13.3 mm longi; pedicelli 4.5—7 mm longi. Petala limbo orbiculari, margine fimbriato. Stamina heteromorpha; antherae sepalis lateralibus oppositae 1-2 loculis redactis instructae; antherae glabrae. Stylus anticus 3.8-4.5 mm longus, glaber vel sparsim pubescens, utroque foliolo 1.8-2.3 mm longo, 2—2.4 mm lato, subquadrato; styli postici 4.5—5 mm longi, lyrati, adaxa- liter pubescentes, foliolo 2.5-3.2 mm longo, 2.2—2.5 mm lato, subquadrato vel inter- dum subrectangulari. Samara ala dorsalis ca. 4 cm longa, ca. 1.6 cm lata; nux ca. 8.5 mm alta, pari alularum lateralium instructa; loculus cavernulis aériis circumcinctus. 14 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 17 yta uy FIG. 4. Stigmaphyllon matogrossense. a. Flowering branch and portion of large leaf (0.5). b. Detail of abaxial leaf surface (x15). c. Posterior petal (<2). d. Androecium (x5); second stamen from left opposes the posterior petal. e. Gynoecium (x5); posterior styles bent slightly outward to show anterior style (in center). (Based on Krapovickas et al. 40114.) Vine. Laminas 9.2-17.5 cm long, 5.5-11.5 em wide, narrowly triangular to triangular to narrowly ovate to ovate, apex acuminate to sometimes almost caudate, base truncate to cordate, sparsely sericeous to glabrous above, sericeous below, margin very shallowly crenate to subentire and with irregularly spaced glands (0.5—0.8 mm in diameter) at the sinuses, with a pair of prominent but sessile glands at the apex of the petiole, each gland 1.5-2.2 mm in diameter; petioles 1.8- 8.5 cm long, sericeous; stipules 0.6-1 mm long, 0.8-1 mm wide, triangular, eglandular. Flowers ca. 12-20 per umbel, the umbels borne in dichasia or com- pound dichasia. Peduncles 4—13.3 mm long; pedicels 4.5—-7 mm long; peduncles 0.9—2.3 times as long as the pedicels. Bracts 1.4-3 mm long, 1.2-1.8 mm wide, triangular; bracteoles 1.2—2.3 mm long, 1-1.6 mm wide, triangular, each bracteole usually with a pair of inconspicuous glands 0.2—0.5 mm in diameter, sometimes one or both glands absent. Sepals 2.5-3.2 mm long, 2.8-3.5 mm wide, glands 2.3-3 mm long, 1.2-1.4 mm wide. Limbs of the lateral petals orbicular, the margin with fimbriae up to 0.8 mm long; anterior-lateral petals: claw 2.8-3.4 mm long, limb 14— 15 mm long and wide; posterior-lateral petals: claw 1.5-2 mm long, limb 12-13.5 mm long and wide; posterior petal: claw 3—3.3 mm long, apex indented, limb 10-11 mm long and wide, orbicular or broadly obovate, margin fimbriate or fimbriate- lacerate, the fimbriae/teeth up to 0.8 (—1) mm long. Stamens unequal, those oppo- site the posterior-lateral petals (and the posterior styles) the largest, anthers of those opposite the anterior-lateral sepals with the connective enlarged and the locules reduced, anthers of those opposite the posterior-lateral sepals commonly with only | reduced unopened locule and thus essentially sterile or sometimes with 1990 C. ANDERSON: STIGMAPHYLLON ibs) 2 tiny locules; anthers glabrous. Anterior style 3.8—4.5 mm long, shorter than the posterior two, glabrous or with a few scattered hairs in the proximal %; each foliole 1.8-2.3 mm long, 2-2.4 mm wide, subsquare. Posterior styles 4.5-5 mm long, lyrate, with a row of hairs adaxially in the proximal 4—¥4; foliole 2.5-3.2 mm long, 2.2-2.5 mm wide, subsquare to sometimes subrectangular. Dorsal wing of samara ca. 4cm long, ca. 1.6 cm wide, upper margin with a blunt tooth; nut bearing a pair of lateral winglets, each 8-11 mm long, 3-4.5 mm wide, semicircular to lunate, sometimes also with a spur ca. 0.1 mm long; nut ca. 8.5 mm high, ca. 6 mm in diameter, inflated, the thick-walled locule flanked by incompletely partitioned nar- row air chambers, areole concave, carpophore up to 4.5 mm long. Seed ca. 8 mm long, embryo narrowly ovoid, ca. three times as long as wide, outer cotyledon ca. 7.1 mm long, ca. 2.6 mm wide, straight, inner cotyledon ca. 6.7 mm long, ca. 2.2 mm wide, straight. Fig. 5. Phenology. Collected in flower from April through June, in fruit in August. Distribution. Collected in savanna and varzea in Para, Brazil. ADDITIONAL SPECIMENS EXAMINED. BraAZziL. Parad: Serra do ee Mpio. Itaituba, estrada Santarém-Cuiaba, BR-163, Km 794, 9°22’S, 54°54'W, Amaral et al. 961 (MICH, NY); Mpio. Itaituba, estrada Santarém-Cuiaba, BR-163, Km 1011, margem direita do Rio eae 7°40'S, SS°1S'W, Amaral et al. 1230 (MICH, NY); Boa Vista, 1931, Carr s.n. (F); Rio Tapajos, Fordlandia, M. Silva 1641 (MG). The leaves of S. paraense are silvery-sericeous below and have sessile glands in the sinuses of the very shallowly crenate leaf margin. The peduncles commonly FIG. 5. Stigmaphyllon paraense. a. Flowering branch (0.5). b. Detail of abaxial leaf surface with marginal gland (x15). c. eae (<5); second stamen from left opposes the posterior petal. d. Gynoecium, with anterior style in center (5). e. Samara (x0. 75). f. Longitudinal section through nut (<1.5); the thick-walled locule is es by aay) partitioned air chambers. g. Embryo (2.5). (a-d based on Sperling 6322, e-g based on M. Silva 1641. 16 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 17 exceed the pedicels. The anthers are glabrous, and the stout posterior styles bear an adaxial row of scattered hairs. The samara is unusual in that the thick-walled locule is surrounded by an array of incompletely partitioned air chambers. Stigmaphyllon saxicola C. Anderson, sp. nov.—Tyre: Brazit. Espirito Santo: Mpio. Ibatiba, Corrego S. Jodo, sobre bloco de rochas, 14 Jun 1985, Hatschbach 49397 (holotype: MBM!; isotype: MICH!). Liana. Laminae 10-17 cm longae, 8-11 cm latae, ellipticae vel ovatae vel suborbiculares, supra tomentulosae vel glabratae, subtus dense tomentosae, mar- gine sparsim glanduloso. Inflorescentia dichasialis vel thyrsiformis constata ex um- bellis, floribus in quaque umbella ca. 15, axibus secondariis distalibus interdum alternantibus divaricatis. Pedunculi 6—9.5 mm longi; pedicelli S-7.5 mm longi. Petala limbo orbiculari, margine fimbriato. Stamina heteromorpha; antherae sepalis antico-lateralibus oppositae 2 loculis redactis instructae, antherae sepalis postico- lateralibus oppositae | loculo redacto instructae; antherae sparsim pubescentes. Stylus anticus ca. 2.4 mm longus, utroque foliolo ca. 1.2 mm longo, ca. 1.3 mm lato, subquadrato; styli postici ca. 2.8 mm longi, lyrati, foliolo ca. 1.5 mm longo latoque, subquadrato vel suborbiculari. Samara ala dorsali 5.8—6.2 cm longa, ca. 1.8 cm lata; nux 9.5—10 mm alta, alulis lateralibus vel calcaribus vel cristis instructa. Vine. Laminas 10-17 cm long, 8-11 cm wide, elliptical to ovate to subor- bicular, apex emarginate-mucronate, base cordate, tomentulose to glabrate above, densely tomentose below, irregularly spaced prominent glands (each 0.2—0.3 mm in diameter) borne adjacent to the margin below, with a pair of flush glands at the apex of the petiole or borne halfway on the lamina, each gland 1.5-2 mm in diameter; petioles 2.5—4.5 cm long, densely pubescent with stout T-shaped hairs; stipules 1—1.2 mm long, 0.6—0.8 mm wide, narrowly triangular, eglandular. Flowers ca. 15 per umbel, the umbels arranged in dichasia or compound dichasia or small thyrses, the distal secondary axes sometimes alternate and divaricate and the inflorescence then appearing racemiform. Peduncles 6—9.5 mm long; pedicels 5—7.5 mm long; peduncles 1—1.4 times as long as the pedicels. Bracts 1-1.5 mm long, 0.8- I mm wide, triangular; bracteoles 1.3-2 mm long, 0.8-1 mm wide, triangular, eglandular. Sepals 2—2.2 mm long, 2.2—2.4 mm wide, glands 2—2.2 mm long, 1 mm wide. All petals with the limbs orbicular, margin with fimbriae up to 0.3 mm long: anterior-lateral petals: claw ca. 1.8 mm long, limb ca. 8 mm long and wide; posterior-lateral petals: claw ca. | mm long, limb ca. 7 mm long and wide; posterior petal: claw ca. 2 mm long, apex slightly indented, limb ca. 6 mm long and wide. Stamens unequal, those opposite the posterior-lateral petals (and the posterior styles) with the longest filaments, anthers of those opposite the anterior-lateral sepals with the connective enlarged and the locules reduced, anthers of those oppo- site the posterior-lateral sepals with only | reduced locule; anthers very sparsely pubescent. Anterior style ca. 2.4 mm long, shorter than the posterior two, glabrous or with a few scattered hairs in the proximal 3; apex ca. 1.5 mm long, each foliole ca. 1.2 mm long, 1.3 mm wide, subsquare. Posterior styles ca. 2.8 mm long, lyrate, pubescent in the proximal “%2-%; foliole ca. 1.4 mm long, ca. 1.5 mm wide, subsquare to suborbicular. Dorsal wing of samara 5.8—6.2 cm long, ca. 1.8 cm wide, upper margin with a slight blunt tooth; nut bearing a grossly dentate lateral winglet on each side, these up to 1.2 mm long, and/or spurs and crests, these 1.5—6 mm long, 1.5-3.3 mm wide; nut 9.5-10 mm high, 5.5—-6 mm in diameter, areole con- cave, carpophore up to 2.5 mm long. Mature seed not seen. Fig. 6. 1990 C. ANDERSON: STIGMAPHYLLON 17 FIG. 6. Stigmaphyllon saxicola. a. Flowering branch (0.5) and enlargements of stem hairs (20). b. Detail of abaxial leaf surface with submarginal gland (25) and e nlargements of T-shaped hairs (x20). c. Androecium (<7.5); second stamen from right opposes the posterior petal. d. Gynoecium, with anterior style in center (<5). e. Samara (0.5). (a-d based on eee 49397, e based on Hatschbach 49399. ) NAL SPECIMEN EXAMINED. BRAZIL. Espirito Santo: Mpio. Ibatiba, Corrego S. Joao, 14 Jun 1985, pe ee 49399 (MBM, MICH Stigmaphyllon saxicola, known only from two collections from Espirito Santo, has relatively small flowers borne in umbels arranged in sometimes eccentrically branched inflorescences. Although the branching is basically in the dichasial pat- tern, the distal branches may be alternate and somewhat divaricate, so that the inflorescence appears racemiform (Fig. 6a). The limbs of the petals are less than 1 cm in diameter, and the styles are less than 3 mm long. The anthers are very sparsely pubescent with a few apical hairs; those of the stamens opposite the posterior-lateral petals bear only one reduced locule. The broad leaves are so densely tomentose below that the prominent submarginal glands are hidden. The samaras are rather large; the dorsal wing is ca. 6 cm long and the nut ca. 1 cm high. Seats ce stylopogon C. Anderson, sp. nov.—TyPe: BraziL. Rond6énia: Mpio. Pimenta Bueno, rodovia Cuiaba-Pérto Velho, BR-364, Km 188, 11°12'S, 61°62 W, Cid et al. 4648 (holotype: INPA!; isotypes: MICH! NY!). Liana. Laminae 7.5-10.5 cm longae, 5-9 cm latae, ovatae, supra sparsim tomentulosae vel glabratae, subtus dense tomentosae, margine parum crenato vel subintegro sinubus glanduliferis. Inflorescentia dichasialis vel thyrsiformis constata ex umbellis, floribus in quaque umbella ca. 10-15. Pedunculi 5.5—8.5 mm longi; 18 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 17 pedicelli 3-9.5 mm longi. Petala limbo orbiculari, margine fimbriato; limbus petali postici supra basin fimbriis crassis glandulosis instructus. Stamina heteromorpha; antherae sepalis antico-lateralibus oppositae loculis redactis instructae, antherae sepalis postico-lateralibus oppositae | loculo redacto instructae; antherae glabrace. Stylus anticus 3.7—4.5 mm longus, adaxaliter pubescens, utroque foliolo 2-2.3 mm longo, ca. 2 mm lato, subquadrato; styli postici ca. 5 mm longi, lyrati, adaxaliter pubescentes, foliolo 3-3.7 mm longo, 3—3.2 mm lato, subquadrato. Samara ignota. ine. Laminas 7.5-10.5 cm long, 5-9 cm wide, ovate, apex mucronate or emarginate-mucronate, base deeply cordate to sometimes auriculate, sparsely tomentulose to glabrate above, densely tomentose below, margin very shallowly crenate to subentire and with irregularly spaced glands (0.4—0.6 mm in diameter) at the sinuses, with a pair of prominent but sessile glands at the apex of the petiole, each gland 1.5-2.5 mm in diameter; petioles 2.2-4 cm long, densely sericeous to glabrate; stipules 1-1.7 mm long, 1.2-2 mm wide, triangular, eglandular. Flowers ca. 10-15 per umbel, the umbels borne in dichasia or compound dichasia or small thyrses. Peduncles 5.5—8.5 mm long; pedicels 3-9.5 mm long; peduncles 0.7—2.2 times as long as the pedicels. Bracts 1.6-2.2 mm long, |.2—1.7 mm wide, triangular; bracteoles 1.5—2 mm long, 1-1.5 mm wide, triangular, eglandular or each bracteole with a narrow band of glandular tissue along the margin. Sepals 2.7-3 mm long, 2.5-3 mm wide, glands 2.6—3 mm long, 1.3—1.5 mm wide. All petals with the limbs orbicular, those of the lateral petals with the margin with fimbriae up to 0.5 mm long; anterior-lateral petals: claw 3-3.5 mm long, limb 12-13 mm long and wide; posterior-lateral petals: claw 2.2-2.5 mm long, limb 11-12 mm long and wide; FIG. 7. Stigmaphyllon stylopogon. a. Flowering branch and portion of large leaf (0.5). b. Detail of abaxial leaf surface with marginal gland (x 10). c. Posterior petal (2.5). d. Androecium (<5); second stamen from left opposes the posterior petal. e. Gynoecium (x5); posterior styles bent slightly outward to show anterior style (in center). (Based on Cid et al. 4648.) 1990 C. ANDERSON: STIGMAPHYLLON 19 posterior petal: claw 3.5—4 mm long, apex not indented or only slightly so, limb 10- 11 mm long and wide, margin with fimbriae up to 0.7 mm long and at the base on each side with 2-3 stout gland-tipped fimbriae (0.6-1 mm long, ca. 0.2 mm wide). Stamens unequal, those opposite the posterior-lateral petals (and the posterior styles) the largest, anthers of those opposite the lateral sepals with the connective enlarged and the locules reduced, anthers of those opposite the posterior-lateral sepals with only 1 reduced locule; anthers glabrous. Anterior style 3.7—4.5 mm long, shorter than the posterior two, adaxially with a row of hairs in the proximal V;—¥s; each foliole 2-2.3 mm long, ca. 2 mm wide, subsquare. Posterior styles ca. 5 mm long, ee adaxially with a row of hairs from the base nearly to the stigma; foliole 3-3.7 mm long, 3—3.2 mm wide, subsquare. Samara not seen. Fig. 7. ADDITIONAL SPECIMEN EXAMINED. BraziL. Mato Grosso: Rio Juruena, Aripuana, Fontanilha, RADAM SC-21-YD, descampado da beira do rio, 25 Jun 1977, M. G. Silva 3192 (MG, NY). Stigmaphyllon stylopogon, a species of western Brazil, is named for the bearded styles, all bearing an adaxial row of hairs. The anthers are glabrous, and those of stamens opposite the posterior-lateral sepals bear only one reduced locule. This species, like S. matogrossense, bears several marginal, stout, gland-tipped fimbriae near the base of the limb of the posterior petal. The leaves are densely tomentose below and bear sessile glands in very shallow sinuses along the margin. ACKNOWLEDGMENTS I thank William R. Anderson for his comments and suggestions, and Stephen F. Smith and Amaro Macedo for prov iene material of S. macedoanum. | am grateful to the curators of the following herbaria for permitting me to examine their specimens: CTES, F, INPA, MBM, MG, MICH, NY, R, SP, US Karin Douthit drew the beautiful ieee LITERATURE CITED Niedenzu, F. 1899. De genere Stigmatophyllo (pars prior). Ind. Lect. Lyc. Brunsberg. p. hiem. 1899— 1900: 1-33. . 1900. De genere Stigmatophyllo (pars posterior). Ind. Lect. Lyc. Brunsberg. p. aest. 1900: 1-32. . 1928. Malpighiaceae. In Das Pflanzenreich, ed. A. Engler, IV. 141: 1- 870. Leipzig: Wilhelm Encelnenct - i a OO oo _ a es oe niche aril Contr. Univ. Mich. Herb. 17:21-37. 1990. THE TAXONOMY OF JUBELINA (MALPIGHIACEAE) William R. Anderson University of Michigan Herbarium North University Building Ann Arbor, Michigan 48109-1057 The Malpighiaceae are not as numerous or diverse in the Amazonian lowlands as in drier, more open habitats like the Planalto of central Brazil, but some genera are mostly or entirely Amazonian. One such is Jubelina, four of whose six species occur in the Amazonian drainage system, the other two being found in nearby areas whose floras have a strong Amazonian component (Fig. 1). My purpose in this little monograph is to bring together what is known about the genus, and to offer some suggestions about its evolutionary history. Jubelina Adr. Juss. in Delessert, Icon. Sel. 3: 19, pl. 32. 1837 [1838]. Sprucina Nied., Arbeiten Bot. Inst. K6nigl. Lyceums Hosianum Braunsberg 3: Diplopterys subgenus Jubelina (Adr. Juss.) Nied., Arbeiten Bot. Inst. Konigl. Lyceums Hosianum Braunsberg 4: 16. 1912. Woody vines. Leaves opposite, the petiole eglandular, the lamina flat or very slightly revolute at margin, bearing impressed glands or rarely eglandular, the lat- eral veins prominent below and interconnected by + parallel “scalariform” tertiary veins; stipules small or minute, triangular, borne on base of petiole. Inflorescences axillary and terminal, decompound, thyrsiform, containing much-reduced bractlike leaves below the floriferous bracts, the flowers ultimately borne in umbels of 4 or corymbs of 6; bracts and bracteoles large, pubescent on both sides, persistent; peduncle shorter than pedicel. Sepals 5, nearly distinct narrowly ovate, obovate, or oblong, spreading to expose outermost petal in enlarging bud, the anterior sepal eglandular, the lateral 4 usually bearing 1 large gland each, formed by + complete fusion of 2 (except in J. uleana, with 6-8 distinct glands), occasionally all sepals eglandular. Petals pink or yellow, 5, at least the anterior-lateral 2 abaxially seri- ceous, the lateral 4 spreading, the posterior erect. Receptacle glabrous. Stamens 10, glabrous; filaments very stout opposite posterior-lateral petals. Ovary of 3 carpels, 1 anterior and 2 posterior, all fertile, adaxially adnate to a common axis; styles 3, subterminal, the apex with a large internal stigma and dorsally truncate or short- hooked. Fruit breaking apart into 3 1-seeded samaras on a high pyramidal torus; samara with a narrowly elliptical or linear ventral areole 1-3 mm wide, a semicircu- lar, entire or repand, central dorsal wing often extended forward at apex between lateral wings, and 2 large lateral wings usually confluent at base, each lateral wing with a complex structure comprising at least an outer membranous wing and a sterile cavity developed in its base, parallel to the fertile locule, during maturation of the fruit, and frequently bearing additional wings, winglets, crests, or irregular out- growths between outer wing and central dorsal wing. Embryo with thick flat sub- equal cotyledons, | slightly longer than the other and bent back over it at the apex. Type. Jubelina riparia Adr. Juss. pe CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 17 Jubelina @ grisebachiana FIG. 1. Distribution of all species of Jubelina. Flora Neotropica base map no. 1, copyright the University of Utrecht. NOTEWORTHY GENERIC CHARACTERS Jubelina is one of the vining genera that have the principal wings of the samara lateral (see further discussion below under Extra- and Intrageneric Relationships). It is set off as a genus by the following combination of character-states: scalariform tertiary veins, large decompound terminal inflorescences with the flowers in um- bels, large bracts and bracteoles, long narrow sepals, at least the two anterior- lateral petals abaxially sericeous, and the samaras with a high narrow ventral areole and complex lateral wings, each of which contains a sterile cavity parallel to the fertile locule and usually bears at least one crest or wing between the outer lateral wing and the central dorsal wing. In addition, five of the six species are notable for having the paired calyx glands (if present) connate to form one large gland in the center of the sepa The sterile cavities in the fruit are especially interesting, because they are unique in this genus. They seem likely to be an adaptation for dispersal by water, and suggest that even in species like Jubelina uleana and J. grisebachiana, which have among the largest samaras in the family, water may be an important factor in their dispersal. In J. magnifica there seems to have been a shift toward more dependence on water, reflected in reduced fruit wings, and in J. riparia this trend has culminated in samaras with minimal wings but very large inflated bladders. Another significant trend in the genus is for the reduced leaves, bracts, bracteoles, and sepals in the large inflorescence to become colored. These characteristics com- bine to produce a picture of plants well adapted to life in wet lowland forests. They are strong vines that can reach the tops of large trees. Their large inflorescences full of colored parts presumably attract pollinators from some distance, an important adaptation since individual plants are often far apart in such forests. And their fruits are well adapted for primary dispersal by wind followed by secondary dis- persal by water. 1990 W. R. ANDERSON: JUBELINA 23 EXTRA- AND INTRAGENERIC RELATIONSHIPS Most neotropical genera of Malpighiaceae can be placed in one of two subfami- lies. The one that has most of the less specialized character-states is the Byrsonim- oideae, in which the pollen is tricolporate and the plants are trees or shrubs and bear unwinged fruits (Anderson 1978). The other large subfamily must be called the Malpighioideae, because it includes the genus Malpighia, which is atypical in bearing fleshy fruits but is descended from the wing-fruited genus Mascagnia. Most species in the Malpighioideae are vines and bear winged fruits, and the pollen is of several derived types. Within the Malpighioideae Mascagnia is central to a group of genera in which the principal wings of the samara are lateral, not dorsal. When one looks beyond the samaras, one finds the mascagnioid genera to be rather diverse. Hiraea and Mascagnia were separated long ago, and only recently Johnson (1986) has segre- gated Callaeum from Mascagnia, which still remains a probably unnatural assem- blage. Jubelina is mascagnioid, but it is not very similar in most characters to other mascagnioid genera. Niedenzu (1928) included Jubelina with Mezia as a subgenus of his genus Diplopterys. Gates (1982) has shown that Diplopterys sensu stricto is a segregate from Banisteriopsis, not at all closely related to Jubelina and Mezia. The latter two certainly deserve to be recognized as separate genera, as was done by Cuatrecasas in 1958, but I agree with Niedenzu that Mezia is closely related to Jubelina. It has a similar inflorescence, long narrow sepals, and samaras whose lateral wings are basally confluent and often bear intermediate elaborations. How- ever, Mezia is advanced in several characters (Anderson 1981) and can only be regarded as a sister genus to Jubelina; neither could reasonably be considered di- rectly ancestral to the other. Moving further afield, one naturally looks to Mascagnia and its segregates for other outgroups with which to compare Jubelina. The best of these is probably Callaeum (Johnson 1986), which often has umbellate inflorescences and sericeous petals, and one species of which (C. nicaraguense) bears a complex structure between the lateral and dorsal wings of the samara. Within Mascagnia itself, the best group to which I can compare Jubelina is Niedenzu’s section Pleuropterys (1928), which has hairy petals and sometimes bears intermediate elabo- rations on the lateral wings of the samara. However, the lateral wings are usually free at the base and the inflorescence is racemose, so the similarity is less than compelling. No help is to be obtained in this case from consideration of chromosome numbers or pollen. The only chromosome number known for Jubelina is my count of n = 10 in J. magnifica. Most wing-fruited Malpighiaceae I have counted have ten pairs of chromosomes or a multiple of that number, so that count simply confirms that Jubelina is correctly placed in the Malpighioideae. The pollen, as described by Lowrie (1982), is of a moderately specialized rugose type common among mas- cagnioid Malpighiaceae. A believable phylogenetic analysis of Jubelina is made difficult by the lack of an obvious outgroup to guide the assessment of the polarity of characters. Neverthe- less, I am including a simple analysis, based on rather few characters, in order to summarize some clear relationships between the species and present my best esti- mate of phyletic trends. The principal outgroup employed was Mezia, with secon- dary consideration given to Callaeum and Mascagnia section Pleuropterys. The characters and character-states used in the analysis are given below, with my rea- sons for assigning polarity as I did: a. Leaf hairs shifting from sessile (0) to stalked (1). Sessile hairs are common in all three outgroups and stalked hairs are rare, so it seems best to consider stalked hairs derived within Jubelina. 24 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 17 OR . Leaf glands shifting from uniseriate (0) to scattered (1). Having the leaf glands scattered is an unusual condition, apparently not found in any of the outgroups, so it is surely derived within Jubelina. . Sepals uninflated (Q) to distally inflated with aerenchyma (1). This condition is unknown in any of the outgroups, so it is probably derived within Jubelina. . Calyx glands distinct (0) to connate (1). Almost all outgroup species (and indeed almost all neotropical Malpighiaceae) have distinct calyx glands, so connate glands are surely a derived condition. . Lateral petals shifting from yellow (0) to pink (1). All species of all three outgroups have yellow petals, so that seems likely to be the ancestral color in Jubelina. . Inner wing on lateral wing of samara well developed (0) to reduced or absent (1). Intermediate wings are present in Mezia, and were probably present in the common ancestor of the two genera; a residual crest or winglet 1s often formed in the two species of Jubelina that have experienced this loss. . Transverse outgrowths of winglets on the lateral wings of the samara absent (0) to present (1). Elaborations like these have evolved repeatedly in mascagnioid lines, including Mezia. I consider them an adaptation for dispersal by water, because they increase the surface area of the fruit and probably retard wetting. This adaptive significance, added to the complete absence of even rudiments of such structures in half the species of Jubelina, leads me to consider them a derived feature. This decision is based primarily on intrageneric considerations, not on outgroup comparison. . Lateral wings of the samara well developed (0) to partially reduced (1) to rudimentary (2). This step, too, seems likely to be part of a shift from dispersal by wind to dispersal by water, so polarity is assigned partly on the basis of intrageneric considerations. However, it is also true that almost all species in the outgroups have large, well-developed lateral wings on their samaras. Q jae — oO = ga — —_— When the six species of Jubelina are scored for these eight characters, the result is the matrix shown in Table 1. From this I derive the diagram shown in Figure 2, which has ten steps, including a parallel loss of the inner wing on the lateral wing of the samara in J. grisebachiana and J. wilburii. One could construct a tree of the same length by postulating that that inner wing was lost in a step between u/eana and the rest of the genus, then regained in the common ancestor of rosea, riparia, and magnifica. That would not change the branching pattern of the tree. In several ways I| find the tree in Figure 2 intuitively satisfying. Jubelina uleana and J. wilburtt have many characteristics in common and surely belong close to- gether in any phylogenetic analysis. Similarly, J. rosea, J. riparia, and J. magnifica TABLE |. Species/character matrix for Jubelina. See text for description and discussion of characters. a b c d € ii 2g h grisebachiana 1 0 i l | 1 0 0 gnifica 1 | | ] 0 ] | riparia ] 0 1 | 1 0 l 2 rosea 1 0 I | I 0 I 0 ilean 0 0 0 0 0 0) 0 0 wilburil 0 0 0 l 0 | 0 0 1990 W. R. ANDERSON: JUBELINA 25 uleana f { wilburii d f Beryl { grisebachiana a Cc € i t T TOSe€a & a h ‘ riparia b magnifica FIG. 2. Phylogeny of Jubelina. Crossbars indicate shifts from ancestral to derived character-states; letters above crossbars denote characters. See text for description of characters and Table 1| for distribu- tion of character-states. form a complex of obviously related species. And J. grisebachiana, while generally more like the rosea group than the two yellow-flowered species, is intermediate in its stem hairs and hooked styles, and probably deserves a somewhat intermediate place on the tree. On the other hand, I am not so confident about the orientation of the tree. Parallelism and reversal have been common in the evolution of the Malpighiaceae, and just a few changes in my analysis of characters would root the tree differently. For example, the distinct calyx glands of J. uleana could be a reversal, and the bright yellow petal pigments of J. uleana and J. wilburii may not be homologous with the darker pigments in Mezia, in which case the ancestral condi- tion in Jubelina might actually be pink petals, which are common in Mascagnia section Mascagnia. Therefore, I urge the reader not to regard the tree in Figure 2 as a definitive phylogeny of Jubelina, but as a first attempt based on the information now at Nand. GEOGRAPHICAL DISTRIBUTION The distribution of all six species of Jubelina is mapped in Figure 1. Several features of that distribution merit comment. Only J. grisebachiana and J. magnifica have been found in the same vicinity, near San Carlos de Rio Negro, and I do not know that they are actually sympatric there. Two species are extra-Amazonian, but in both cases there are strong ties between their floras and the Amazonian flora. Jubelina rosea occurs only in the Atlantic drainage of French Guiana and Suriname, where the lowland forest is continuous with that of adjacent Amazonia. Jubelina wilburii is known mostly from Central America, but it has also been collected at Anori in Colombia, home of disjuncts from western Amazonia like Dicella julianit (Anderson 1975) and Ectopopterys soejartoi (Anderson 1980). One related pair, J. uleana and J. wilburii, share the western part of the genus’s range, and another close pair, J. rosea and J. riparia, share the east. 26 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 17 KEY TO THE SPECIES OF JUBELINA I. Leaves sparsely sericeous to nearly glabrate, the hairs sessile, + straight, strongly appressed; lateral 4 petals yellow, the posterior yellow or pink and white; all 5 petals abaxially sericeous; hairs on samara 0.2—0.4 mm long. 2. Petiole of larger leaves 20-40 mm long; oui bracts 7.5-10.5 mm long, 5-6 mm wide, with white vesture; sepals tomentose on both . the anterior eglandular, ih lateral 4 each bearing 2 glands or | or both of the glands pee to the anterior sepal much reduced or absent; apex of styles with an obvious dorsal hook 0.2—0.5 mm long; ae wing of samara bearing a well- een inner wing parallel to central dorsal wing, 7-14 mm wide; Amazonian Ecuador and P 1. J. uleana. pop eae of larger leaves 11 a ie 25) mm long; floriferous bracts 2—4 (-6) mm long, 1-2 (-: mm wide, with yellow or brown vesture; sepals abaxially tomentellous, adaxially glabrous or sparsely pilose near apex, the anterior eglandular, the lateral 4 all eglandular or all bearing | large central gland; apex of styles dorsally truncate or with a rounde up to (0.1 mm long; lateral wing of samara bearing an inner crest or winglet parallel to dorsal wing, up to 5 mm wide, this occasionally absent; Costa Rica, Panama, and northern Colombia. 2. J. wilburii. Leaves persistently velutinous or tomentose, the hairs stalked, their arms varying from suberect to parallel to the lamina and straight to serpentine; lateral 4 petals pink, the posterior pink or pink and white; lateral 4 petals see sericeous or the posterior-lateral 2 glabrous, the posterior glabrous; longest hairs on samara |—2.7 mm long 3. Samara without wings or winglets Peed oe and lateral wings, at most only acrest | mm wide parallel to dorsal wing; stems subsericeous or appressed-tomentose, the limb of hairs at right angles to the stalk; calyx glands revolute at apex; posterior-lateral 2 petals eglandular- dentate or -fimbriate; northwestern Brazil and Amazonian Colombia and Venezuela. o — — Bod: nee 3. Samara with well- sal eee wings and winglets between central dorsal and outer later wings; stems velutinous, the hairs ral erect; calyx glands attached at apex; Bie lateral 2 petals often biaganiae: fimbria 4. Lamina of larger leaves 10-21 cm one 5-14 cm wide, rounded or abruptly short- acuminate at apex with the acumen up to 5 (-10) mm long, bearing below on each sic 0-3 glands near base and up to 4 nen zt a single row distally; central dorsal wing af samara as wide as to (usually) wider than pe inner wings borne on lateral wings. 5. Lateral wing of samara with its outer membranous wing 22—36 mm wide; bracts and bracteoles |.4—2.5 (~3) mm wide; lamina ete without g aaa near base, occasion- ally 1; Suriname and French Guiana. 4. J. rosea, 5. Lateral wing of samara with its outer membranous wing 5—7 mm wide; bracts and bracteoles 4—5 mm wide; lamina bearing (0— A —2 (—3) glands near base on each side of midrib; northeastern Brazil and French Guiana. 5. J. riparia. 4. Lamina of larger leaves 18-28 cm long, cece cm wide, acuminate at apex with the acumen often up to 20 (~25) mm long, bearing (4—) 5-15 scattered glands below on each side; central dorsal wing of samara narrower than and hidden by parallel inner wings borne on lateral wings; Amazonian Venezuela. 6. J. magnifica. The terminology used here follows that of my 1981 treatment of the Mal- pighiaceae of the Guayana Highland. See pp. 24-26 of that paper for definitions of vesture types, bract, peduncle, bracteole, and pedicel, and for an explanation of the floral symmetry in the family. Here, as there, I have described all wings of the samara such that width is measured at right angles to the ventral areole and height is measured parallel to it. 1. Jubelina uleana (Nied.) Cuatr., Webbia 13: 445. 1958. Diplopterys uleana Nied., Arbeiten Bot. Inst. Konigl. Lyceums Hosianum Braunsberg 4: 18. 1912. Stems sericeous to glabrate, the hairs white or pale yellowish, sessile, straight and mostly strongly appressed, very short. Lamina of larger leaves 14-24 cm long, 1990 W. R. ANDERSON: JUBELINA De 8-17 cm wide, broadly elliptical or somewhat ovate or obovate, obtuse or rounded at base, usually abruptly acuminate at apex with the acumen 5—20 mm long, bearing 2-5 glands below on each side in a row set in from margin, sparsely sericeous to nearly glabrate on both sides, the hairs sessile, straight, strongly appressed, 0.2—-0.4 mm long; petiole 20-40 mm long, sericeous to glabrate. Inflorescence proximally sericeous, distally tomentose or velutinous; bracts and bracteoles reddish, densely white-tomentellous or velutinous, 7.5—10.5 mm long, 5-6 mm wide, broadly ovate, obtuse, the bracteoles somewhat falcate; peduncle 2.5-4.5 mm long; pedicel 6—9.5 mm long, short-velutinous. Sepals reddish, 4.5-7 mm long, 1.5-2 mm wide, nar- rowly oblong, acute or obtuse at apex, reflexed, apparently not inflated with aerenchyma, tomentose on both sides, the anterior eglandular, the lateral 4 each bearing 2 elliptical or ovate glands 1.6-2 mm long and 0.8-1.5 mm wide, or | or both of the glands adjacent to anterior sepal much reduced or absent. Lateral 4 petals yellow, orange in age, abaxially sericeous except glabrous near margin, the claw 2-3.5 mm long, the limb 3.5—6.3 mm long, 3.5—6 mm wide, rotund, concave, subentire or erose, the anterior-lateral 2 notably larger than the posterior-lateral 2; posterior petal white with red veins, abaxially sericeous only in center, the claw 2.8-4.3 mm long, the limb 2.6-3.5 mm long and wide, flat, ovate or nearly square, bearing small sessile or short-stalked glands all around margin. Filament opposite posterior petal notably shorter than others, 1.5—2 mm long, the rest 2.3-4 mm long, longest opposite anterior sepal, connate up to 1 mm; anthers 0.8—1.3 mm long, the connectives flat or somewhat enlarged opposite sepals. Ovary velutinous; styles glabrous or piliferous, 3-3.7 mm long, subequal or the anterior shorter, the anterior nearly straight or somewhat sigmoid or arcuate, the posterior 2 arcuate- or sigmoid- ascending, the apex of all 3 with a protuberant internal stigma and an obvious dorsal hook 0.2-0.5 mm long. Samara elliptical, 80-100