ree ves RAD Adee ees strbawhstsasars ne “eee va Wtgh ches erdebedese . sth ate Ses = Ahad dealer ade atgorackdecaeeinee fe ee WS ficerkyny Rreecbeys At er te brs otro mrket . 5 > = eats i ENN EE nr ee Hey poleaeth: rbrdn Ocerineeuet ~~ ee “4 ee io ae cram at i rearey ee “fits. _ Steels ok bre ste > 4 hese 44 eet te tome Metco tires we = ee ty 9 ge vere See ree ee @ weer oe Rptighet tare’ e r me A tet > SA verre vat PR re ta fone coer mis ret pat be Ter ne 6 * . mi te hte my Sa 70 * * oy ~ i Aare ~ SF tee wer wey ee er es b+. b. 0.00.2. 2. 0 = in CC asnaye — bese he i) Th ij ps At ‘ : t i ae : he (ie Bah i ox ais 4 4 Be ITANIC A 2 . ‘4 — GARD PN FLORA MALESIANA ee BT a x Salo ANDS PLAN ENTUIN, BOTANIC GARDENS BUITENZORG, JAVA, INDONESTA meORA MALESIANA BEING AN ILLUSTRATED SYSTEMATIC ACCOUNT OF THE MALAYSIAN FLORA, INCLUDING KEYS FOR DETERMINATION: DIAGNOSTIC DESCRIPTIONS: REFERENCES TO THE LITERATURE: SYNONYMY, AND DISTRIBUTION, AND NOTES ON THE ECOLOGY OF ITS WILD AND COMMONLY CULTIVATED PLANTS PUBLISHED UNDER THE AUSPICES OF THE BOTANIC GARDENS, BUITENZORG, AND THE AUTHORITY OF THE CENTRAL GOVERNMENT OF INDONESIA, DEPARTMENT OF AGRICULTURE & FISHERIES PREPARED ON AN INTERNATIONAL CO-OPERATIVE BASIS UNDER THE SUPERVISION OF SEVERAL DIRECTORS OF BOTANIC GARDENS, KEEPERS OF HERBARIA AND VARIOUS PROMINENT BOTANISTS FOR THE PROMOTION OF BOTANICAL SCIENCE AND THE CULTURAL ADVANCEMENT OF THE PEOPLES OF SOUTH-EASTERN ASIA TO THE SOUTHWEST PACIFIC REGION SERIES I SPERMATOPHYTA VOLUME 4 PART 1 GENERAL EDITOR: Dre C. GG) J. VAN STEENIS SENIOR BOTANIST ’ BOTANIC GARDENS / BUITENZORG FOR SALE ONLY - NOT FOR EXCHANGE PUBLISHED BY NOORDHOFF-KOLFF N.Y.» BATAVIA 1948 JITAN GARD NO ELEGE Flora Malesiana is designed to represent a concise flora of the Malaysian region. © The present part is the first published. It contains the Ist instalment of vol. 4 of series I. The following independent series are planned: Series I. Spermatophyta (flowering plants) . ; . ca 15 volumes Series II. Pteridophyta (ferns and fern allies) : ; . ca 3 volumes Series III. Bryophyta (mosses and hepatics) . ; : . ca 5 volumes Series IV. Fungi & Lichenes (fungi and lichens) . . 2! 4 "Sa elomntics Series V. Algae (algae) Pavey fe ca 3 volumes At the present moment preparations have been made necessary for the start of series I only. As soon as possible the other series will be commenced and directed by special general editors.—The area covered by Flora Malesiana is indicated on the accompanying map by the hatched area. RPEAN P.OR.THE PUBLICATION OF S# REE SvEsB\K MeA T © P HY TA Volume 1. Cyclopaedia of Malaysian botanical collectors and collections, by Mrs M. J. VAN STEENIS-KRUSEMAN. This is a cyclopaedia of Malaysian collections of Phanerogams and Pterido- phytes. It contains over 3000 names of collectors, with short biographies, carefully . excerpted itineraries, and literature pertaining to the collections. GARDEN Indispensable when localizing herbarium sheets of Malaysian plants and inter- preting the scant notes on the labels of older collections. Contains also chapters on methods of collecting in the tropics, hints for travellers, notes on erroneously localized Malaysian collections, &c. — Ready for the press, probably available by the end of 1949. Appr. 600 printed pages. Sample treatment at the end of this copy. Volume 2. Malaysian plant life, by Dr C. G. G. J. VAN STEENIS. This is a second edition, much enlarged, of ‘Maleische Vegetatieschetsen’ (1935) by the same author. It deals with all vegetation types known in Malaysia, as far as described in literature and reports, or known to the author by personally acquired field knowledge, their status and interpretation, their relations, origin, distribution within Malaysia, and importance to mankind. Biological phenomena, both ex- plained or yet unexplained will be briefly touched on. This book will be copiously illustrated. The MS. can be expected to be finished in 1950. Appr. 500 printed pages. Sample treatment at the end of this copy. Volume 3. Malaysian plant geography, by Dr C. G. G. J. VAN STEENIS. This volume consists of two parts. Part one deals with floristic plant geography and contains tables showing the distribution of the ca 2200 recognized indigenous genera of Malaysian phanerogams, compiled from literature and from the Herbarium. Further there are chapters on the history of phytogeographical theories and discussions, a provisional list of the genera with their synonymy, an attempt to divide the Archipelago into phytogeographical districts based on the hitherto known facts of generic distribution, and a discussion of the phytogeographical character of the islands or island groups separately. The MS. is far advanced but not yet ready for the press. Sample treatment at the end of this copy. Part two will deal with the historical plant geography of the Malaysian region. This is only in the initial stage. Volume 4. Flora Malesiana proper. Onwards of vol. 4 the revisions of the families will be printed in the sequence in which they are finished, irrespective of the alliance of the groups concerned. Vol. 4 will be opened by an introductory essay containing chapters on the importance of variability in Malaysian plants, special aberrations with which the Malaysian botanist is confronted in the field and with which he must be acquainted to judge their importance in the often scanty specimens available in the herbaria. A chapter is added on the history of Malaysian phyto- graphy. An annotated list of existing revisions concludes the introduction. PROPOSED CONTENTS OF VOLU ME Ly Preface. 2. Introduction. 3. General considerations. m4. History of descriptive Malaysian botany (by Dr H. C. D. DE WIT). “5. General plan of revisions and hints to collaborators. 6. Keys for identifying Malaysian plants. ©7. Annotated list of former revisions. ~ 8. Photographs of principal botanical contributors to Malaysian botany. O9. Systematic revisions of families of Phanerogams, incl. Bignoniaceae, Podo- stemonaceae, Droseraceae, Umbelliferae, Convolvulaceae, Dioscoreaceae, Plumbaginaceae, etc. etc. ; MANAGEMENT OF FLORA MALESIANA Flora Malesiana will not be available for purposes of exchange; it is for sale only. Co-operating and collaborating institutions can obtain this flora at a reduced price. For subscribers to a complete series the price will be reduced. For substantial collaborators a special reduction will be fixed individually. General volumes 1—3 will also be sold separately to a limited extent. APPELCATUONs iO DE DIRE CTE Dame N.V. Erven P. Noordhoff, N.V. Noordhoff-Kolff, P.O, Box. 39, P20: Box103; Groningen, Holland. Batavia-C., Java. and in the Americas to: The Chronica Botanica Co., Book Department Waltham, 54, Massachusetts, U.S.A. SCIENTIFTEC COMMUNICA ERIONS concerning Flora Malesiana should be addressed to Dr C.G. G. J. van Steenis, c/o Riyksherbarium, Nonnensteeg 1, Leiden, Holland. INS EX page page Aceraceae . , ; i a SNVSSACeEde s |e 29 FACUIMIGIACEAG SSE. : 2) : 3/7’ Philydraceaeé ; 1 5 Ancistrocladaceae. . . . 6 " Sarcospermaceae .. > ee Aponogetonaceae. _.. ei: 11 Sphenocleaceae . . oe ee Bummaumiaceae . . | =. 13’ Stackhousiaceae . ~ "ees PRINTED IN THE NETHERLANDS BY JOH. ENSCHEDE EN ZONEN » HAARLEM MEMORIAE EORUM QUI SE FLORAE MALESIANAE PERSCRUTANDAE DEDERUNT ET NOBIS DUCES ET EXEMPLA FUERUNT GRATO ANIMO HOC OPUS DEDICANT AUCTORES FED 10 194d GARDE PREFACE There are only a few things left in common to the displaced and disjointed inhabit- ants of this Earth; they are the things spiritual. Among those treasures of the mind natural science has come to the fore only in the last three centuries, as a lofty and impartial principle that tends to join people instead of disrupting them. Through war, famine and pestilence the undying fire of science has remained a steady beacon. At the inception of a great work, which shall be the demonstration of the united effort of many workers, it seems meet to remember the function of Science, apart from its benificial or detrimental applications. In these days the adjectives ‘pure’ and ‘applied’ have lost much of their meaning, and the Masters amongst us were the least concerned with this classification. Nevertheless, as long as sentiment, politics, greed and bigotry rule this world, a purely scientific endeavour may become a binding force between individual groups, and maybe, even between nations. At the beginning of this great project we therefore see already a dark shadow cast by this unfortunate era, but rather than to dwell on darker thought let us invoke the light that is cast by those that lived and worked in these regions before us, and let us consider this work as an apotheosis of the ideals formulated by Melchior Treub who, half a century ago, became the initiator of co-ordinated scientific effort in the tropics. His ideals are still ours. To the General Editor, Dr C. G. G. J. van Steenis, we all want to express our gratitude for his initiative, for his boundless energy and, especially, for his faith in this project. To discuss the project of the Flora Malesiana from the technical side would be simply borrowing materials from the scientific collaborators. Rather than to plagiarize it seems fit to leave these matters to the specialist. But as Director of the Scientific Institutes, called “’s-Lands Plantentuin’ I may express the satisfaction that our Government, almost a century after the foundation of our Herbarium, after 130 years of effort of these institutes, after the publication of innumerable, chiefly disjointed contributions, has decided to further a unified effort. Amongst the many accusations that our workers have suffered in the last years, some at least have some foundation. We have plowed too deep, we were too few, we did not seek enough contact. But the greatest blight of tropical science has been the lack of continuity. Now we may plow deep, but with many to help. Now we may proceed together and, let us hope, with continuity guaranteed. The great ‘Horn of Plenty’, the cornucopia of our Malaysian Flora, which was opened by van Rheede and by Rumphius shall still flow for a long time. I wish great joy to those that shall have the privilege to examine its contents. (L.G.M.BAAS BECKING) Director of ’s-Lands Plantentuin 10. It. . Systematice plantas suas disponit verus Botanicus; Nec absque ordine easdem enumerat. . Fructificationis principium in theoretica dispositione agnoscit; Nec dispositionem secundum Herbam immutat. . Genera naturalia assumit; Nec Erronea ob speciei notam aberrantem conficit. . Species distinctas tradit; Nec e Varietatibus falsas fingit. Varietates ad species reducit; Nec eas, pari passu, cum speciebus obambulare sinit. Synonyma praestantissima indagat et seligit; Nec acquiescit in quacunque obvia nomenclatura. . Differentias characteristicas inquirit; Nec inania nomina specifica praeponit veris. Plantas vagas ad Genera amandare studet; Nec rariores obvias fugitivis oculis adspicit. Descriptiones complectentes differentias essentiales, compendiose sistit; Nec naturalissimam structuram oratorio sermone ebuccinat. Minimas partes attente scrutatur; Nec ea, quae maxime illustrant, flocci facit. Observationibus ubique plantas illustrat; Nec in vago nomine acquiescit. . Oculis propris quae singularia sunt observat; Nec sua solum, ex Auctoribus, compilat. LINNAEUS, Philosophia botanica INT RODY GEFON After the appearance of Rumpurius’s Herbarium Amboinense, the result of lifelong research into the botanical treasures of the Malaysian Archipela- go, the first comprehensive work on the flora of these islands was begun by C. L. BLuME, the second Director of the Botanic Gardens at Buitenzorg. His Bijdragen tot de Flora van Nederlandsch Indié (Contributions to the Flora of the Netherlands Indies) consisted of numerous brief botanical diag- noses mostly, however, of Javan species. Shortly after followed his Flora Javae and later Rumphia. None of these books represent a ‘flora’; neither completeness was aimed at nor keys were given. The first design for a flora of the whole of Malay- sia seems to have been drafted by the Swiss bota- nists H. ZOLLINGER and his teacher, A. Morrirzi.! I have not succeeded in tracing any further results of their plans. Since the publication? of the Flora van Neder- landsch Indié or Flora Indiae Batavae by F. A. W. Mique. (5 vols, 1854—°60)—which was no ‘flora’ in the present meaning of the word, keys being almost absent—no work has been conceived with the object of covering the Malaysian region. MIQUEL’s work? may be considered as a more or less critical compilation of descriptions, mostly copied or extracted.4 MIQUEL must have realized that by his Flora the proper work was only started. This may be concluded from the series of revisions which MIQUEL, together with some specialists, published in 4 volumes Annales musei botanici lugduno batavi (1863-"69),> Choix des plantes rares ou nouvelles (1863), to which was added his posthumous J//u- strations de la Flore del’ Archipel Indien (1870—71) by his successor at Leyden University, W. F. R. SURINGAR. Unfortunately, MriqueL had few pupils® which caused a serious shortage of well-trained systema- tists during half a century of botanical endeavour in the East Indies. The only Dutch scientists study- ing the Malaysian flora were: P. DE BoErR, who wrote his doctor’s thesis on the subject De Coniferis archipelagi indici (1866), and later became a professor of Pharmacology at Gro- ningen University, and R. H. C. C. SCHEFFER, an extremely able bota- (1) ZOLLINGER, Observationes phytographicae etc. Natuur- & Geneesk. Arch. 1 (1845) 375; cf. also J. K. HASsKARL, Flora 30 (1847) 299. (2) Made possible by a grant of the Ministry for the Colonies. (3) Dates of publication of the several parts in Bull. Jard. Bot. Btzg III, 13 (1934) 284. (4) Compare ZOLLINGER, Natuurk. Tijdschr. Ned. Ind. 13 (1857) 292-322; id. (in German), Vierteljahrschr. Naturf. Ges. Ziirich 2 (1857) 318-349. (5) Dates of publication of the several parts cf. NAKAI, Journ. Arn. Arbor. 6 (1925) 211-213. (6) Cf. the article in honour of Dr A. A. PULLE, who resumed MIQUEL’s work at Utrecht Uni- versity, Bull. Jard. Bot. Btzg III, 16 (1939) 103-105. nist whose thesis was entitled De Myrsinaceis archipelagi indici (1867). SCHEFFER was subsequently appointed as the (fourth) Director of the Botanic Gardens, Buiten- zorg, and ardently promoted the study of the Malaysian Flora, notwithstanding his feeble health. In his term of office he published several important papers, most on Annonaceae’ and Palmae.® De Boer had one pupil in systematic botany, TH. VALETON, who obtained his doctor’s degree on a monographic study of the Olacineae.? He eventu- ally was employed as a bacteriologist in the Sugar Experiment Station in Java but, soon after, joined the staff of the Botanic Gardens, Buitenzorg (1892). After the appointment of Dr M. Treus as the fifth Director of the Gardens in 1880, interest in the promotion of knowledge of the Malaysian flora revived, but TREUB was badly handicapped by the absence of trained Dutch systematists. TREUB—a contemporary of HooKer, EICHLER, BENTHAM, and HARVEY & SONDER, the editors of respectively the Flora of British India, the Flora Brasiliensis, the Flora Australiensis, and the Flora of tropical Africa—was well aware that systematic botany in the Netherlands Empire was on the verge of falling behind that in other tropical countries. He judged the advancement of systematics of pre- eminent importance. He engaged W. Burck, a pupil of SURINGAR’s at Leyden, later a teacher of botany at Buitenzorg, as a subdirector of the Gardens (1883) and charged him with critical research into Sapotaceae (getah- pertja family),'!° Mucuna,'! the Erythroxylaceae (cocafamily),!? and Dipterocarpaceae,'* mostly fa- milies of economic importance. TREUB, who tried continuously to raise a world- wide interest in the Gardens and its botanical in- stitutes, considered the compilation of a new Ma- laysian Flora to be premature. Collections were inadequate and of the vegetation of the surround- ing regions little was known. He advanced, therefore, the idea of composing a local flora of the surroundings of Buitenzorg, covering the region from the mangrove of Tand- jong Priok to the summit of Mt Gedeh at 3000 m. All altitudinal zones would thus be represented. This Flore de Buitenzorg would serve as a guide to botanically interested visitors of the Gardens and be equally acceptable to residents of Java. Dr J. G. BOERLAGE, then conservator of Leyden Her- barium, during a visit to Buitenzorg as a stipen- diate of the Dutch Buitenzorg Fund, had already made collections for the new flora (1889) and published an article on the grasses.!* (7) Ann. Jard. Bot. Btzg 2 (1885) 1-31. (8) Ibid. vol. 1 (1876) 103-164; O. BECcCARI, Reliq. ScHEFF. ibid. 2 (1885) 77-171. (9) Critisch overzicht der Olacineae (1886). (10) Ann. Jard. Bot. Btzg 5 (1886) 1-85. (11) Ibid. 11 (1893) 183-190. (12) Ibid. 11 (1893) 190-194. (13) Ibid. 6 (1887) 145-249. (14) Ann. Jard. Bot. Btzg 8 (1890) 47-78. VI FLORA MALESIANA [ser. I, vol. 4! Treus, however, found it difficult to rally workers to this local flora and so most of it was assigned to foreign visitors who sometimes were temporarily employed at the Gardens. Six volumes appeared viz the Myxomycetes by O. PENZIG (1898), Ferns and Fern Allies by M. RActBoRSKI (1898), Hepatics by V. SCHIFFNER (1900), Algae by E. DE WILDEMAN (1900), and Mosses by M. FLEISCHER (1900-22, 4 vols). The 6th and only volume on Phanerogams was written by J. J. SmirH (Orchida- ceae 1905, atlas 1908-14). None of the volumes of the Flore de Buitenzorg bears the character of a local flora; the majority deal with the whole of Java. FLEISCHER’s Musci even expanded to a standard work on the world’s mosses. Of the flowering plants apart from the Orchida- ceae, much material was collected by BurRcKk and H. HALLER who planned to elaborate a 7th volume of the Flore de Buitenzorg. A list of the species to be included is kept at Buitenzorg, but nothing ever appeared in print. During this period important revisions of fami- lies were published abroad by O. BeEccari in his 3-volume Malesia. Several monographs appeared in the 4° tomes of the Annals of the Royal Botanic Gardens, Calcutta, on the genus Ficus, the oaks and chestnuts, the bamboos, etc. Local floras of other parts of Malaysia were the 3rd edition of BLANCO’s Flora de Filipinas (1877-— °83)! by Naves & F.-VILLAR, SCHUMANN & HOLLRUNG’s Flora von Kaiser Wilhelmsland (1889), and SCHUMANN & LAUTERBACH’S Flora der Deut- schen Schutzgebiete in der Siidsee (1901) with the Nachtrdge (1905). These eastern floras resembled enumerations and were mainly indices of materials collected on expeditions. In 1890 BoERLAGE previously having published two critical studies of Malaysian plants, viz the genus Achyranthes? and the genera of Araliaceae,* started a work of quite another nature in the com- pilation Handleiding tot de kennis der flora van Nederlandsch Indié.* This comprised a description of the families and genera of Malaysian phanero- gams. The species were—especially in the last parts—only briefly enumerated. He added to a few families keys to the genera. The generic de- scriptions were mostly critically copied from BENTHAM & HOOKER’s Genera Plantarum, and occasionally emended. Phytographically Boer- LAGE’S Handleiding brought hardly anything new, but now a comprehensive review in the Dutch language of families and genera came within reach of interested persons in the colonies. However, as will be demonstrated later, this interest was and is still more directed towards species than genera. (1) On the dates of publication see MERRILL, Philip. J. Sc. 12 (1917) Bot. 113-117. (2) Ned. Kruidk. Archief II, 5 (1889) 420-430. (3) Ann. Jard. Bot. Btzg 6 (1887) 97. (4) In total 5 parts appeared, the last posthu- mously (1890-1903, 3 vols). The publication was made possible by a grant of the Ministry for the- Colonies. BOERLAGE’S work was more intended as a prelude to a general flora than as a final work. He accepted (1896) the post of subdirector of the Botanic Gardens and Head of its first Division (Herbarium and Botanical Museum), as a suc- cessor to BURCK and began a monograph of the Annonaceae>. Unfortunately he soon (1900) fell a victim to a tropical disease while on a tour in the Moluccas attempting to re-collect the plants mentioned by Rumpuius in his Herbarium Am- boinense. Another flora was started, at TREUB’s instigation, of trees growing in the island of Java. This was to be based mainly on the collections made by Forest officer S. H. Koorpers who gathered in the field notes on each species (occurrence, value, uses, etc.). Scientific descriptions and keys were by Tu. VALETON. This work is Bijdragen tot de kennis der Boomsoorten van Java (Additamenta ad cognitionem Florae Javanicae, pars 1, .Arbores). Thirteen vol- mes compose this standard work, the 12th volume is by J. J. Smiru, the concluding 13th by SmitH and VALETON. The work was begun in 1894, and finished in 1913. Later illustrations were edited by Koorpbers in his unfinished Atlas der Baumarten von Java (4 vols, 1913-18). The Bijdragen is an excellent work with critical descriptions and notes, and still very useful though, of course, now anti- quated. The descriptions of the species and genera are both in Dutch and Latin. During Treus’s directorate many collections, specially of the Outer Provinces,® were brought together. HALLIER made an important one in West Borneo, KoorDERS in Java and North Celebes, the SARASINS collected in Celebes, ForBEs and Koor- DERS in Sumatra, ForsBes in Timor, while WArR- BURG’S, SCHLECHTER’S, and BECCARI’s great col- lections equalled those of TEYSMANN’s and extended over the whole archipelago. These collections were partly inaccessible though together they could have served to a large measure as a reliable basis for a Flora Malesiana. Lack of trained taxonomists induced TREUB to engage J. J. SmitH, formerly an assistant curator of the Gardens, for taxonomic work. His revisions of Javan Euphorbiaceae, Ulmaceae, Urticaceae, and Orchidaceae proved his ability, and SmirH spent his life in describing Malaysian Orchids, Ericaceae, and Epacridaceae. Unfortunately, he did hardly any monographical work. For the same reasons TREUuB selected C. A. BACKER, a teacher in a primary school at Batavia who possessed already a thorough and critical knowledge of the local flora. BACKER intended to fill the still existing /Jacunae in the phanerogamic part of the Flore de Buitenzorg, which resulted in the publication of one volume of a Flora van Batavia (1907). This was followed by a preliminary schoolfiora’? and later by the Schoolflora (1911). (5) Icon. Bogor. 1 (1899) 79-208, t. 26—75. (6) That was: Netherlands Indian territory out- side the islands of Java and Madoera. (7) Voorlooper eener Schoolflora van Java (Pre- cursory Schoolflora of Java). Batavia (1908). Dec. 1948] Introduction VII The latter excellent work contained only + 25 °% of the Javan flora (Choripetalae). He later devoted all his time to the Javan flora, wrote (together with VAN SLOOTEN) a weed flora of tea plantations (1924), 3 instalments of a Handboek voor de Flora van Java (192428), a weed flora of sugar planta- tions (1928—34; vol. II (atlas) not yet completed), and is now engagedin completing the Flora of Java. ! An ill-advised enterprise was a flora of Java by S. H. KoorbDers who, when charged by the N.I. Government to write a flora of the Javan moun- tains, abandoned this concept and hurriedly com- piled an Exkursionsflora von Java (Jena, 1911—12, 3 vols) which did more harm than good and is scarcely of any value to a student of the Javan flora. The flora of the Malay Peninsula was originally included in the Flora of British India, but as the account remained very incomplete KING & GAMBLE, and RIDLEY, started to work on it, pub- lishing a true model of a critical local flora.* This was later followed by RIDLeEy’s decidedly uncritical Flora of the Malay Peninsula (5 vols, 1922-’25). On the Flora of Borneo a most helpful Biblio- graphic enumeration of Bornean plants was prepared by Dr E. D. MERRILL.? In the Philippines MERRILL, after 1902, energetic- ally undertook the research of the Philippine flora, this first resulting in an excellent local Flora of Manila (1912), in a large number of papers dealing with several aspects of the Philippine flora, and crowned by his Enumeration of Philippine flowering plants (1923-26). The results of frequent expeditions into the Dutch and German territories of New Guinea were published by Dr A. A. PULLE and others in the serial Nova Guinea (vols, 8, 12, 14, and 18), and by C. LAUTERBACH and others,* and in recent years those of Dutch and British parts by MERRILL and other collaborators.° The undesirability of compiling, at this stage, local floras in Malaysia. The studies of the mate- tials of various separate regions persuaded some leading Dutch botanists in the first quarter of our century—for some reasons they doubted the feasi- bility of a Malaysian flora as a whole—to propose several local floras e.g. one of Java, of Borneo, Sumatra, Celebes, etc. This caused the appoint- ment of HALLIER at Leyden to write a Flora of Borneo resulting in a small preliminary paper.® It is clear that this was a wrong policy, born from (1) Seven parts of a mimeographed emergency edition were issued up till now through the care of the Rijksherbarium, Leiden (1940-48), 9 vols. (2) The contributions of the former appeared under the title Materials towards a Flora of the Malay Peninsula in various numbers of the Journ. Asiat. Soc. Bengal, vol. 58 onwards (18891—915). (3) Journ. Str. Br. Roy. Asiat. Soc. Special number (1921). (4) Under the title Beitrdge zur Flora Papuasiens in many volumes of the Botanische Jahrbiicher (1912 onwards). (5) Journ. Arn. Arb. 9 (1928) et seq. (6) Beih. Bot. Centralbl. 2. Abt. 34 (1916) 19-53. either ignorance of the taxonomic position and the technique of writing revisions, or from the wish for dodging obstacles; the difficulties should be faced directly. Only temporary profit may be gained from making local floras, and both valuable time and money are wasted by the enormous duplication which is unavoidable when the goal of a flora of a plant-geographical unit is to be reached along this tortuous road. The natural sequence is to start with the large flora, eventually followed later by local floras, a procedure followed in the great floras of South America, tropical Africa, India, and Australia. The unnatural sequence of starting with the local flora has led, both in North America and Europe, to a most regrettable state of affairs. The absence of a general flora is also one of the causes that the flora of Java which BACKER has studied close on forty years is only now more or less to be completed. It contains several families which cannot be critically treated (Lauraceae, Araceae, Zingiberaceae, etc.) lacking revisions of these families in the whole Malaysian region. General Flora. A general flora was and is needed and prospects at the end of the first World War seemed favourable. The Forest Research Institute and the Museum for Economic Botany’ at Bui- tenzorg requested much service and urged the Herbarium of the Botanic Gardens to produce speedy results. This induced the Goverment to add to the staff of the Herbarium R. C. BAKHUIZEN VAN DEN BRINK (1917)—he was originally a plantation assistant—Dr D. F. vAN SLOOTEN and Dr H. J. La, the first pupils of PULLE at Utrecht (1919). In 1921 Dr H. C. CAMMERLOHER, a German biolo- gist, was appointed, and a professional collector engaged, H. A. B. BUNNEMEIJER. At the same time a scheme was made for critical revisions. These were to be published in the Bul- letin du Jardin Botanique, Buitenzorg* under the heading: Contributions a l’étude de la Flore des Indes Néerlandaises. Economically important fami- lies had priority. The method of treatment stood below that of KiNG & GAMBLE’s Materials in so far that descriptions were only admitted if species were new or critical. This was believed to save time. On the other hand extensive lists of herbarium numbers had to be compiled. If the latter had been left out and instead a concise characteristic of the occurrence of the species given, besides a good diagnostic description of each species, the Con- tributions would have made a most satisfactory foundation. Though the later Contributions are far more complete than the earlier, the manner of treatment and publication is so laborious and slow that at this rate the Flora Malesiana will never be completed. Till the present 34 Contributions have appeared, comprising 2000 species. Due to the post-war economic depression of 1921—22 the Staff of the Buitenzorg Herbarium (7) Head of this Museum was the late K. HEYNE, author of the standard work on useful plants of Indonesia (1927). (8) Bull. Jard. Bot. Btzg III, 5 (1923) 294 seq. VIII FLORA MALESIANA [ser. I, vol. 4! were reduced, and though towards 1930 there were a few constructive moments, a protracted slump set in after that year and the Staff at Buitenzorg were reduced to the barest minimum. Shortly before the Pacific War the Staff again increased but the circumstances limited advancement of the Flora to planning. I have always felt it as a shortcoming, and not in accordance with the standing of the great work at hand, that the contributions appeared in a peri- odical as scattered articles and not as a separate publication. The work was undertaken on full official au- thority but being printed in an irregularly inter- rupted series of articles in many volumes of a tech- nical journal, it was practically inaccessible to a wider non-professional public. A standard work of this scope and weight meant to be used by future generations and worthy of the wonders of nature in this great land ought to have commanded con- siderable interest in and beyond the tropics, speci- ally so in neighbouring countries. It would not have made a difference in expenditure to issue this work as a separate publication thus materially augmenting its practical importance, its intrinsic value remaining, of course, the same. This seemingly trivial technical-editorial point had very undesirable consequences. If the Govern- ment had once for all decided to order a standard work on the Malaysian flora to be written with all possible expediency and to be used many years afterwards, the halting and haphazard progress in the decade preceding the Pacific War would never have occurred. It is a gratifying thought that the turbulent times of the present could not prevent the Government now to put the Flora Malesiana in an advantageous and satisfactory position both as regards effective publication, and national and international colla- boration of systematists. Co-operation with foreign colleagues, whose help is invited and whose help is needed in order to finish the work within a reasonable time, will now, presumably, more easily be obtained. Evidently, it is far more attractive and stimulating to be entrusted with an individual part of a standard work than with writing an article in a journal. Prospect and scope of the Flora Malesiana. A general flora of Malaysia must result from a careful study of all previous publications, blending them into a harmonious whole, and so founding Malay- sian botany on a secure base of historical fact, ob- servation, and accurate description. This is, how- ever, the labour of a lifetime, and although I may be privileged in witnessing the laying of the founda- tions and the issue ofa number of volumes, I cannot hope to bring it to a conclusion; progress, more- over, will depend entirely upon circumstances at present beyond control. I have no doubt that when I will be called to abandon this endeavour the historical necessity for the completion of this work will compel someone to continue this task and, eventually, to finish it. It would, however, be wrong were I to convey the impression that this arduous undertaking had entirely originated with myself: on the contrary during many years the conviction has grown among plant taxonomists that the ample collections ac- cumulating in this country warranted the prepa- ration and publication of a Flora Malesiana. The collections are undeniably extensive having been gathered over a wide extent of country.! As I am anxious to render each portion of the work in itself as complete as possible, and desirous of enlisting those of our fellow-botanists as may be willing to take care of those families or groups they are most familiar with, the Flora Malesiana, when terminated will probably consist of a series of local-monographs. For these reasons it seems inadvisable and most inconvenient to arrange the families in the mode of sequence usually adopted in systematic works. I consider it important that the Flora Malesiana should embrace as wide an area as possible, being firmly convinced that no species can be properly defined, until it has been examined in all variations induced by the differences in climate, locality, and soil, which an extensive area affords. Also, the flora of an area cannot be worked out thoroughly with- out a knowledge of the botany of the surrounding countries (these have many plants in common), and so the greater the area encompassed, the better it will illustrate habits, forms, and variations of the species comprised within it. For this reason we have extended the limits of our Flora from Sumatra to New Guinea and from Luzon to Christmas Island, Timor and New Guinea. The use of the Flora Malesiana. In the preceding pages I have mentioned several times the public and the government. Both have a right to a clear understanding of the use of a flora of the scope and character of that now contemplated. Although it is difficult to explain theoretically the ‘use’, i.e. the material benefit of purely scien- tific standard works, many anecdotes and instances concerning scientists entirely possessed by their inventions, instruments, and desire for research, told in biographies and popular literature, exem- plify the eminently practical results based on seemingly impractical and abstract study. The same can be said about this Flora. Botany is not a cherished source of pleasure and interest to naturalists only; and I have but vague ideas of (1) Collections have increased enormously. From 1917 on, the Forest Research Station at Buitenzorg accumulated materials of arboreous plants from the islands outside Java (more than 30.000 numbers): The Museum for Economic Bo- tany furnished by its own collectors another 6000 numbers of those islands. The collectors of the Buitenzorg Herbarium in the past 30 years added to the collections more than 125.000 numbers. A similar increase of Malaysian collections in these last decades is due to the activities at Manila and Singapore; besides, private collectors substantially augmented the collections of New Guinea. A conservative estimate of the collections at Buiten- zorg alone runs to about 400.000 numbers of Malaysian plants. Dec. 1948] possible advantage and ultimate gain for the com- munity and practice by means of this registration of the Malaysian flora. I could refer, of course, to the fact that all other civilized nations have already made considerable progress in the task of making common knowledge of their vegetable resources. Actually the disentangling of confused species, the description of new or the rehabilitation of obsolete genera, the dissection of dried flowers and, in general, the establishment of law and order in ‘the hay loft’, and the publication of the results have less appeal to the lay public than the segregation of a new promising variety of rice or sugarcane, or devising a method to suppress a pest of coffee or of coconut plantations. The Flora of Malaysia contains besides highly interesting and even unique plant forms, instruc- tive vegetation types, and peculiar ecological and phytogeographical problems, numerous important industrial plants and economic products which, in their manifold kinds, add to human comfort and social prosperity, while, in their ranks, many treas- ures still await discovery, the latest accessions being pectin and mannan producing plants. Their value has come as a surprise both to taxonomists and economists. Nearly a century ago, one of the foremost of British botanists, Sir JosEpH DALTON HOOKER! wrote an introductory essay to the Flora of British India, one of the most instructive general essays ever written on tropical botany. This nearly one century old exposition of facts and thought meets the present state of knowledge of the Malaysian flora admirably. Its excellence induced me to copy the following from it:— ““With regard to economic botany, it is obviously impossible to do more than briefly enumerate, under their respective species, the various products which have been used in the arts: for detailed ac- counts of their value, we must refer our readers to the many excellent works on those subjects, which have been published by Indian botanists.” “‘Our work is intended to facilitate the progress of economists, by supplying their great desidera- tum, a critical description of the plants which yield the products they seek. We have had a considerable experience both in medical and economic botany and we announce boldly our conviction, that, so far as India is concerned, these departments are at a standstill, for want of an accurate scientific guide to the flora of that country. Hundreds of valuable products are quite unknown to science, while of most of the others the plants are known only to the professed botanists. The mass must indeed always remain so: just as the refinements of the laboratory and the calculations of the mathema- tician must ever be mysteries to the majority of manufacturers and navigators, whose operations are based on the sciences in question. It is a mistake to suppose that it can be otherwise; or that those who are engaged in forwarding a science so exten- (1) Hooker & THomson, Flora Indica (1855) 1-280, specially p. 3 et seq. Introduction IX sive and abstruse as philosophical botany, can command the time to become so familiar with the details of the commercial value of vegetable prod- ucts, as to be safe referees on these subjects. On the other hand, it is equally a mistake to suppose that those who devote themselves to the collection of economic products, can possess the experience and botanical knowledge necessary to render their identifications of tropical plants trustworthy in the eyes of men of science. It is therefore as a strictly scientific work that we offer this commencement of the Flora Indica to the public, but though the advancement of abstract science is indeed its pri- mary object, yet as we yield to none in our estimate of the value of economic botany, we confidently trust that... our labours will be found of material service.” “Had it been possible to take up the economic plants of India by themselves, and to present a history of them to the English reader, we should at once have devoted ourselves to the task, with the certainty of obtaining an amount of encourage- ment which a so-called paying work is sure to command, but which one ofa more scientific nature is not thought worthy of receiving. We should, however, only be deceiving the public, were we to propose a scheme which, in the present deplorably backward state of scientific Indian botany on the one hand, and the confusion of Indian economic botany on the other, is literally impracticable: the difficulties have increased fourfold, from scientific botany not having advanced pari passu with the economic branch; and so long as plants themselves remain undescribed, it is obviously impossible to recognize what are useful, or so to define them that they shall be known by characters that con- trast with those of the useless. Our principal aim, however, being purely botanical, the most insig- nificant and useless weed is as much the object of our attention as the Teak, Sal, and tea: in the vegetable kingdom, and in the great scheme of nature, all have equal claims on our notice, and no one can predicate of any, its uselessness in an economic point of view.” “Every one who has studied Indian plants, whether for economic purposes or for those of abstract science, must have felt the want of a gener- al work which should include the labours of all Indian botanists, to be a very serious inconvenience. Our own experience in India has convinced us of this; for we found it often impossible to determine the names of many of the most ordinary, and, in an economic point of view, often most valuable forms; and every day’s additional experience in the preparation of this volume has served to show more and more clearly, that whilst such a work is wanting satisfactory progress is impossible. At present the student has to search in general system- atic works, for the descriptions of species; and as all of these are imperfect, a multitude of scattered papers must be consulted for the additions which have from time to time been made. These too have unfortunately so often been published without reference to preceding works of a similar nature, that the same plant has been described as new by aK FLORA MALESIANA [ser. I, vol. 4! many successive botanists, ignorant or neglectful of the labours of their predecessors.’’ So far HOOKER. To emphasize our inability to foresee practical results of taxonomic work I intend to mention a few recent instances in Malaysia showing that plants which seem useless at the present may stand in the focus of attention at a future date. Twenty years ago it would have seemed the whim of a botanist to work on the species of a genus of foetid aroids, scientifically known as Amorphophallus. Few years later, however, the tubers of some species of this genus were found to be important commercially and industrially. The basic work on the distinction of the species, the notes on their distribution, their habit and structure proved to be most useful for agricultural purposes. The same holds for a genus of leguminous plants, Derris. The roots were found to contain a very valuable resin-like substance, rotenon, poison to fish and numerous insects but harmless to larger animals, also to man. As soon as its commercial value was recognized a sudden large demand for Derris rose. It soon appeared that not every species was valuable and so the original studies of Derris offered hold for a first segregation of promising material whereas the systematist was questioned about the characters by which the species could be recognized. The absence of any reliable taxonomic infor- mation of the genus Metroxy/on prevents at present well-founded research on the economic possi- bilities of the sago-producing species which supply a basic food to the whole population of East Malaysia and Melanesia. Invariably it is the duty of the taxonomic bota- nist to supply basic data to research in directed (= applied) botany. In all cases the name of the species, and eventu- ally its varieties, is the alpha of knowledge, as it represents the key to existing literature embodying earlier work on habits, life-history, on distribution geographical and altitudinal, ecology and growth habit, current native names if any, etc. and Flora Malesiana must serve for this purpose. In the past e.g. tropical plant-breeding in some cases followed a wrong direction and might have achieved better results more rapidly when the aid of taxonomists had been available or requested. From the discussion of some selected topics above it will be clear that the taxonomic botanist in composing the Flora Malesiana will be able to offer critical knowledge of numerous forest prod- ucts, plants containing vegetable oils, fats, and resins, rattan, timber, gums, fruits, spices, insec- ticides, fibres, dyes, and medicines, or species which may serve for afforestation, for ornamental use, as new green manures, fodder plants, or possibly, species withstanding drought or being resistant to fire or inundation, suitable for combating erosion, and other economic aspects. In addition to taxonomical information, the Flora Malesiana will contain ecological data. In anthropogenic areas and eroded lands biological control of necessity will seek guidance in its com- prehensive survey of facts. Large amounts of money and energy have been wasted in the absence of professional planning, through negligence of funda- mentals. I remember attempts, as expensive as they were fruitless, of planting mangroves to protect the coastal area of a tropical harbour, a waste which would have been avoided when the ecological potentialities of mangrove forest had been duly considered. ! In (re-)afforestation, the choice of trees has to rely partly on previous experience, but directions can be given by field-taxonomists and by means of general rules of tolerance capacities. Native trees occupy in our forest-types fitting ecological niches, but it should not be assumed that they grow always under optimal conditions. An example is probably found in swamp forest trees which have roots tolerant of a very low aeration of the soil, a virtue not practically utilized, as far as I know, when planting on very poorly aerated soils. The ecological misunderstanding that all plants grow in nature under optimal conditions for their growth led to ‘forest plantations’ of quinine by ~ JUNGHUHN. The Cinchona-crop was saved thanks to TEYSMANN who maintained that the plant should be grown in the open. Much trouble and still much more money could have been saved if this ecolo- gical principle had been better known. The Flora Malesiana is, therefore, of first in- terest to practice and may direct new research: it must give data as to where the plant occurs, in what quantity, under what life-conditions, and with what life-cycle. It ought to contain ecological and biological data, and a critical extract of the notes made by the collectors. None of us can pre- dict the industrial future of a neglected plant spe- cies, but we should be prepared for any coming _ rush on the botanical wealth of this vast archi- pelago, linking the Asiatic and Australian con- tinents. The aim of the Flora Malesiana is to compile a critical knowledge and a botanical standardization of the Malaysian flora of basic importance both to pure and to economic botany. How much of the flora is known? Often it is assumed—the majority of botanists being ac- quainted with the state of knowledge in Europe or North America—that the flora of these islands is sufficiently known, and the actual facts cause astonishment. For instance, not even the number of species is known otherwise than by very approximate cal- culation; 25.000 to 30.000 species of flowering plants is a conservative estimate. The Orchidaceae alone claim about 5000 species. Java possesses more than 500 species of ferns. The number of different species of trees in Malaysia is about 3000. The total number of genera is near 2400. The largest genera are found among the Orchids, Dendro- bium with ca 1110 and Bulbophyllum with about 933 recognized species. This is indeed astonishing if compared with the flora of Holland where the whole native flora (1) Kustaanwas en mangrove (Natuurwet. Tijdschr. Ned. Ind. 101 (1941) 82-85). Dec. 1948] Introduction XI amounts to little more than 1000 flowering plants. Counting all trustworthy and up to date revi- sions together, about 5000 out of a total of 25.000— 30.000 species are now more or less critically known. It appears that the bulk of the work re- mains still to be done. The area covered by the Flora Malesiana will besides Indonesia also include the Malay Pen- insula, Sarawak, Brunei & British North Borneo, the Philippines, Christmas Island, Portuguese Timor, and the whole of New Guinea (fig. 1). Fig. 1. Delimitation and main divisions of the flora of Malaysia. It may be asked whether this is not an unneces- sary extension of the task to include foreign border countries. To explain this it ought to be realized that the demarcation lines of natural units seldom coincide with political boundaries. As much as possible, however, the demarcation of a Flora should be based on scientific, that is, plant geographical limits. Plant geographically the natural demarcation lines of the Malaysian flora pass through the isth- mus of Kra, between the Philippines and Formosa, and through Torres Straits, and include the Louisi- ades and the Bismarck Archipelago. An extensive geographical survey of the distribution of the Malaysian flora will be published in the 3rd volume of this work. The outcome! is wholly in confir- mation with the suggested demarcation lines which were drawn first, as I have mentioned, about a century ago by ZOLLINGER.* In the NW quite a number of typical Malaysian genera of forest plants fail to occur any further in the Indochinese Peninsula, e.g. Rafflesia, Rhizan- thes, camphorwood (Dryobalanops), benzoin (Sty- rax benzoin), kauri or copal (Agathis), true ironwood (Eusideroxylon), menggaris (Koompassia), etc. The Philippines possess an essentially Malaysian flora, in contrast to Formosa’s Japano-Chinese (1) Tijdschr. Kon. Ned. Aardr. Gen. 65 (1948) 193-207, 7 fig. (2) Natuurk. Tijdschr. Ned. Ind. 13 (1857) 293-322. floral character which was definitely demonstrated by MERRILL.? The flora of New Guinea was formerly assumed to be essentially Australian in character. This in- terpretation was mostly based on zoological argu- ments and on the occurrence of few but very striking examples of plants which later appeared to be also spread westwards in the Moluccas and Celebes. O. WARBURG, in 1891,4 on account of important Statistics, already showed the essential Malaysian character of the Papuan flora. Technically the botanist must in each case —whether the Flora Malesiana is limited to a political or to a natural demarcation—study and compare critically all species of the natural phyto- geographical unit. Plants described hitherto only from East New Guinea almost certainly occur also in West New Guinea, numerous species originally described from the Philippines occur in Celebes, the Moluccas and New Guinea, and the same holds for the Malay Peninsula, where the flora is inti- mately allied to that of Sumatra and Borneo. In identifying plants of Malaysia in the narrow sense, that is, limited to the Netherlands Indian boun- daries. the botanist is always obliged to revise or critically to take into consideration the species described from the border areas. This will cost him about the same time and labour as when admitting them into the final work. If these species are omitted, the Flora Malesiana will doubtless be out of date early and unneces- sarily. Bibliographic advantage of the Flora Malesiana. The absence of any definitely indicated centre of publication for Malaysian plants has led to a rather chaotic taxonomic literature. At the present moment revisions of Malaysian plants are pub- lished more or less frequently in about 10 impor- tant periodicals scattered all over the world, and occasional publications are found in some 50 others. An annotated list of former revisions will be presented in this volume to facilitate future study. No single individual can be supposed to own these journals and it is thus more or less private knowledge to those, who have access to a well-stocked library. In Malaysia there are only two libraries where they are nearly all represented, viz at Buitenzorg and Singapore. This is of course a rather unsatisfactory situation to naturalists, foresters, agriculturists, phytoche- mists, veterinarians, pharmacologists, and inter- ested private persons desirous tostudy the flora according to the best available data. The Flora Malesiana will put students of systematic botany generally in possession of the essence of literature. Sequence of publication. It is commonly under- stood that in a flora the sequence of publication ought to be in agreement with the ‘natural system’. This has been—I feel sure—a serious obstacle mentally and practically to all those who, pre- viously, have considered the project of this flora. Arguments against this sequence are in the first (3) Bot. Jahrb. 58 (1923) 599-604. (4) Bot. Jahrb. 13 (1891) 230-455. XII FLORA MALESIANA [ser. I, vol. 4', Dec. 1948] place the existence of several ‘natural systems’; it is tacitly agreed that the last word in ‘the natural system’ will probably never be spoken. A system now adopted may be obsolete when this flora is finished. A choice seems, therefore, difficult, as most of the systems are advanced by leading botanists who among themselves, may claim little priority of preference. It would be possible that the editors of the Flora Malesiana advance a system of their own. However, this falls beyond the scope of this Flora which is solely intended as a practical work. This technical difficulty, which was already mentioned on p. viil, in connection with the adop- tion of a system is a serious obstacle to the progress of the work. Clearly not at every moment a specialist is available for every family of flowering plants. This is more or less a matter of chance. Rapid and regular publication is most desirable and so every opportunity should be made use of. A ‘natural system’ consequently involves the ‘waiting’ of some - manuscripts for many years because it is not yet their turn to be printed, and several volumes will be set up in one part but can be continued only at a remote period because for the ‘following’ family no specialist was available. The real disadvantage can be observed in works like the Flora of North America, in course of publication, of which, in 1941, were published 2 complete volumes and 55 loose parts belonging to 17 of the remaining 32 planned volumes. The same has been the case with the Flore Générale de Il’Indo-Chine where most volumes ranged over a period of about 30 years before they were completed and could be bound. In the meantime consultation was very difficult because the indexes appeared naturally in the final instalment. The handling of the loose parts is undesirable both from a bibliographical and a practical standpoint. In the newly started Flore de Madagascar the families are numbered according to the natural system and are separately published and paged. The idea is that after completion the subscribers can arrange them into sequence and bind them accordingly. We must be aware, however, that this will hardly bring any advantage as the number of families in the Malaysian flora is 211, and that among them 70 families are represented by less than about 10 species, so that also in this case one has to handle a large amount of small unbound fascicles. A long time is needed to complete the Flora Malesiana, about 25 years at least.! This is cer- tainly not overestimated if compared with floras of similar magnitude as Flora Brasiliensis (1840- 1906), Flora of Tropical Africa (1868—hodie), Flora Capensis (1894-1933), Flora of British India (1855- 1897), Flore générale de I’ Indo-Chine (1907-hodie), Flora Australiensis (1863-1878). The exact duration cannot be calculated, this (1) Under the most favourable conditions as regards funds, and co-operation. depends largely on opportunity and facilities, and the joining of forces. The editors are fortunate in having received the promise of much co-operation, and they hope to be able to extend their resources still more. Moreover a considerable amount of | recent publications exists which may easily be adapted to the flora. The here adopted scheme of ‘opportunity se- quence’ in the production of family revisions will remove any delay caused by the ‘natural system’. The addition of an up to date index to the contents of prior parts on the cover of each new appearing instalment will serve to verify in a moment if a desired group has already been revised. The size of the families is of course widely differ- - ent ranging from 1—5000 species. At least one figure illustrating characteristics will be added to each family and large genus. The volumes will not exceed 500-600 printed pages. They must be easy in the hand, agreeable to work with, and bound in covers which may not be attacked by tropical insects, as we hope that numerous subscribers will be found in the Old World tropics outside the official institutions. Completeness of the Flora. No perfection can ever be attained in any tropical flora. Always novelties and new localities will have to be recorded. No squadron of botanists can ever comb a tropical area engirdling !/7 of the equator. Although completeness is a first aim set for this work, its future value will depend mainly on the amount of critical original study which it contains. The Floras of British India? and Australia are now definitely incomplete, but they remain first class sources of information. BACKER’s Schoolflora voor Java, of 1911, still meets present demands nearly as well as at the time of its appearance. If we can keep our flora to so high a standard it will become the keystone to future Malaysian systematic botany. The Flora Malesiana will be started with the flowering plants (Series J). Series II will comprise the ferns and fern allies and is estimated to occupy 3 volumes. Series III will be devoted to mosses and hepatics. These will take about 5 volumes. Series IV will treat the fungi and lichens. The number of volumes can as yet not be estimated. Series V is intended for the algae and other groups of unicellular cryptogams. For the series II-V special editors will be ap- pointed. The general method of treatment may possibly deviate somewhat from the first and largest series but the needs of these can hardly be estimated at the moment. C. G. G. J. VAN STEENIS Buitenzorg / The Hague, Sept. °44/July °47. (2) Dr K. Biswas calculated that to the ‘Flora of British India’ consisting of ca 14000 species, ca 2000 have been added since its publication, a sur- prisingly low number in relation to its vast surface and variety of vegetation types (Proc. 30th I.S.C. pt II, sect. V, Bot., Pres. addr. p. 109). GENERAL CONSIDERATIONS We should endeavour to determine how few, not how many species are comprised in the Malaysian flora. In writing the following chapters I have kept in mind the exemplary ‘Introductory Essay’ of J. D. Hooker in his ‘Flora Indica’ (1855), the precursor of the ‘Flora of British India’. For the same reasons that moved Hooker, I felt obliged to introduce the Flora Malesiana prop er by some general considerations especially in- tended for co-operators less fortunate than I have been in acquiring an experience of long standing in the field. I may add that field experience often is invaluable when studying dried, always frag- mentary, materials in the Herbarium. Some of the subjects HOOKER treated are now too large to be included in one essay and, there- fore, the survey of the Malaysian collections, the physiognomy of the vegetation, and the genetic and floristic plant geography occupy the (introductory) volumes 1-3 of this work. The present essay will be entirely devoted to topics directly bearing on the study of systematic botany. Some of them I have previously discussed, or touched on, in my study of the origin of the Malaysian mountain flora.! As my intention is to further the study of Malaysian botany, I shall discuss only points of which a clear understanding is essential to the Malaysian naturalist. I will try to illustrate each case by reference to plants of this region. These points are: individual variation and racial Segregation, variation caused by the environment, the problem of speciation and specific centres, hybri- dization, views on the status of the species and subspecies, migration and adaptation, and the way to interpret these concepts. These theoretical points are inseparable from a philosophical study of plants, and I believe it to be essential for systematists to explain the prin- ciples which have guided them in the execution and design of their work. HOOKER’S general instructions have guided me in my work, and I am convinced that in the flux of botanical conceptions in general aspects the words of the Master still hold their own. I desire to express here my admiration for this classic work by quoting /iteratim some passages of his essay. “Tt may seem almost chimerical to look forward to a time when all the species of the vegetable world shall have been classified upon philosophical princi- ples, and accurately defined; and it must be con- fessed that the present state of descriptive botany does not hold out much prospect of the realization of so very desirable an object. This, we think, is in a great measure due, not to any want of students willing and anxious to take up the subject, but rather to a gradually increasing misapprehension of the true aim and paramount importance of systematic botany, and of the proper mode of pur- suing the study of the laws that govern the af- finities of plants. We are therefore desirous, at the outset of a work which is devoted to these subjects, (1) Bull. Jard. Bot. Btzg III, 13 (1935) 358-407. of explaining our views on them; and as we trust that our work will fall into the hands of many beginners who are anxious to devote themselves usefully to the furtherance of botanical science, but who have not an opportunity of acquiring in any other way its fundamental principles, we shall make no excuse for dwelling at some length on the subject. We are also anxious to refute the too com- mon opinion (which has been productive of much injury to the progress of botany) that the study of systems presents no difficulties, and that descriptive botany may be undertaken by any one who has ac- quired a tolerable familiarity with the use of terms.” “There can be no doubt that any observant per- son may readily acquire such a knowledge of ex- ternal characters, as will in a short time enable him to refer a considerable number of plants to their natural orders; though even for this first step more knowledge of principles is required, than to make an equal advance in the animal kingdom: but to go beyond this,—to develop the principles of classification, to refer new and obscure forms to their proper places in the system, to define natural groups and even species on philosophical grounds, and to express their relations by characters of real value and with a proper degree of precision, de- mands a knowledge of morphology and anatomy and often of physiology, which must be completely at command, so as to be brought to bear, when necessary, upon each individual organ of every species in the group under consideration. To follow the laws that regulate the growth of all parts of the plant, especially the structure of stems, the func- tions of leaves, the development and arrest of floral organs, and the form, position, and minute ana- tomy of the pollen and ovule, and to trace the whole progress of the ovule and its integuments to their perfect state in the seed, ought all to be fa- miliar processes to the systematic botanist who proceeds upon safe principles; but no progress can be made by him who confines his attention chiefly to the modification of these organs in indi- vidual or natural orders.’-—So far HOOKER. Variability in characters of minor importance and description of extreme forms have led to a rather confused state of affairs. I believe that among the scores of species described many microspecies should be reduced to a much smaller number of true species, with a normal area of distribution and a normal variability in characters typical for Lin- nean populations which are intermediate between the species of extreme ‘splitters’ and extreme ‘Jum- pers’. Much ‘splitting’ has been caused by de- scribing single extreme forms not exactly agreeing with the type or type-description; for practical purposes it is sometimes required to describe such forms as new species and to recognize them pro- visionally as new ‘entities’; the author’s conscience and eagerness to finish his task are thus tempo- rarily satisfied. This method has proved a failure and a serious handicap to the progress of tropical plant knowledge. XIV FLORA MALESIANA There are three methods of handling new col- lections, all being equally unsatisfactory. Firstly, provisional rapid identification of the material as to genus, or to species as far as is possible, and its insertion in the herbarium; collectors in general do not favour this method as only few final names can be provided onacursory examination. Secondly, a collection may be worked through by rough com- parisons to named specimens and with standard literature. This second method is rapid but all ex- treme forms and forms belonging to large genera or to difficult families which cannot be identified rom the available literature, are described as new (specimen description). By this method collectors get immediate results but science is burdened with a host of ‘endemic’ species which, as experience has shown, disappear by the score when a thorough monograph is made. Thirdly, a collection may be thoroughly studied, delaying results, as the iden- tification of extremes means in nearly every genus a preliminary revision. HooKER continues (/.c.):—‘‘A knowledge of the relative importance of characters can only be ac-. quired by long study; and without a due appreciation of their value, no natural group can be defined. Hence many of the new genera which are daily added to our lists rest upon trivial characters, and have no equality with those already in existence. A pro- neness to imitation leads to a gradual increase in their numbers, without a corresponding increase of sectional groups. Indeed, even when the sectional groups are well defined, and the genera in them- selves natural, a too great increase in the number of genera is detrimental, by keeping out of view those higher divisions which are of greater impor- tance. The modern system of elevating every minor group, however trifling the characters by which it is distinguished, to the rank of a genus, evinces, we think, a want of appreciation of the true value of classification. The genus is the group which, in consequence of our system of nomenclature, is kept most prominently before the mind, and which has therefore most importance attached to it.’’! (1) ‘““We may make our meaning more clear by a few examples. The genus Ficus is surely more natu- ral than the subgenera Pogonotrophe, Covellia, Urostigma, &c, into which it has been subdivided. So with the genera Anemone, Hedyotis, Erica, Andromeda, and others which have been split into many by modern systematists.””> R. BROWN, G. BENTHAM, J. D. HooKer and others, in all their works, laboured to keep this important principle in view, and to impress it upon others; they have, however, failed to check the prevalent tendency to the multiplication of genera. I add here other examples of genera occurring in Malaysia which are separated by trifling charac- ters: Voandzeia differs from Vigna only in fruit biology, viz its globular pods ripening subterrane- ously. In Urena and Pavonia now only one fruit character remains the decisive distinction, Dillenia and Wormia are distinguished only in their fruit biology, Berberis and Mahonia are distinct solely in the foliage, Kibessia and Pternandra differ only “The rashness of some botanists is productive | of still more detrimental effects to the science in the case of species; for though a beginner may pause before venturing to institute a genus, it rarely enters into his head to hesitate before proposing | a new species. Hence the difficulty of determining (ser. I, vol. 41 | synonymy is now the greatest obstacle to the pro- | gress of systematic botany; and this incubus un- | fortunately increases from day to day, threatening at no very distant period so to encumber the science,” that a violent effort will be necessary on the part of those who have its interests at heart, to relieve it of a load which materially retards its advancement. The number of species described is now so very great, and the descriptions are scattered through such a multitude of books, that even after long research it is difficult to avoid overlooking much that is already known; and when botanists with limited libraries and herbaria institute new species, it is almost certain that the latter will be found to have been already characterized. To such an extent is this carried, that we could indicate several works, in which one half and even more of the species are proposed in ignorance of the labours of other botanists. Indian Botany unfortunately, far from forming an honourable exception in this particular, presents a perfect chaos of new names for well-known plants, and inaccurate or incom- plete descriptions of new ones’’. “It must be remembered too that the Linnean canon, by which twelve words were allowed for a specific character, is now becoming quite inade- quate to the requirements of the science; and that the brief descriptions, which are now so generally substituted for definitions, unless prepared with the greatest skill, as well as care, and after an inspection of very numerous specitmnens, seldom express accurately the essential characters of a plant. Itis indeed becoming more and more evident, that in the great majority of instances no definition is sufficient to enable inexperienced botanists to determine with accuracy the species of a plant, even when the whole genus is well known; much more is this the case in genera, many of whose species are yet undiscovered; and most of all, in those where the forms, though sufficiently well known, are liable to much variation. In the last case their determination becomes a special study; in unimportant characters of the calyx tube, etc. The separating characters are far less important than those which, in other genera, serve to divide sections or subgenera. The more species are described the more differ- ences originally accepted as of generic rank tend to disappear. A redefinition of the generic charac- ters is often delayed, and the attitude in ‘local- monographs’ is mostly to keep at all costs the old delimitation in order to avoid laborious mono- graphic work. Suggestive casual remarks are often made in local works, but decisions deferred. (2) In entomology this has led to an intolerable chaos (cf. The New Systematics 1940, p. 475-491). The same holds for several large groups of the Fungi. Dec. 1948] and when attempted without access to authentic specimens, leads to inextricable confusion, and its evil effects are not confined to specific botany, but extend to all departments.” “The pages of our Indian Flora will supply numerous illustrations of these remarks, and we would direct the attention of those commencing the study to the lesson to be derived from these instructive errors; for where the first botanists of the day have failed, beginners cannot be expected to succeed. It cannot be too strongly impressed upon all students of botany, that it is only after much preliminary study, and with the aids of a complete library, and an herbarium containing authentic specimens of a very large proportion of known species, that descriptive botany can be effectively carried out; and it would be well for science if this were fully understood and acted upon.” “The prevailing tendency on the part of students of all branches of natural history, to exaggerate the number of species, and to separate accidental forms by trifling characters, is, we think, clearly traceable to the want of early training in accurate obser- vation, and of proper instruction in the objects and aim of natural science. Students are not taught to systematize on broad grounds and sound principles, though this is one of the most difficult processes, requiring great judgement and caution; or, what is worse, they are led by the example if not by the precepts of their teachers, to regard generic and specific distinctions as things of little importance, to be fixed by arbitrary characters, or according to accidental circumstances. As a consequence, the study of systematic botany is gradually taking a lower and lower place in our schools; and, being abandoned by many of those who are best qualified to do it justice, it falls into the hands of a class of naturalists, whose ideas seldom rise above species, and who, by what has well been called hair-splitting, tend to bring the study of these into disrepute.” “We therefore earnestly recommend to the Indian botanist the detailed study of individuals and their organs with the view of determining their limits of variation.” WiGcuHt and Arnott! formulated their warning to beginners as follows :— ““‘We shall perhaps be severely censured for cut- ting down species. We have all along considered it as trifling with nature to separate species on slight or variable grounds, nor could we ever under- stand the ‘cui bono’ for which so much ingenuity in splitting hairs has been wasted. Before we deter- mined what was a species, we examined with care numerous specimens from the same and different localities; and so far we have had an advantage over many other of the European botanists who have described Indian plants, they having only seen one or two isolated specimens. Numerous obser- vations too were made in the plants in their natural situation, the result of which went to prove, what we have frequently endeavoured to enforce by (1) Prod. Fl. Pen. Ind. Or. 1 (1834) p. xxxi. General considerations XV examples throughout the present volume, that no precise shape of leaf or quantity of pubescence is of any value, although both of these seem in each species to be limited within certain variations. With regard to varieties, we have seldom distinguished any unless well marked and tolerably constant; we are aware, indeed, that these correspond to what some naturalists call species, but our own obser- vations have convinced us, that varieties and forms, as well as species, may be constant in similar situ- ations, and even in widely different situations, for many years, if raised from seeds either obtained from the original locality or from cultivated plants; the cultivated cerealia and garden vegetables ought to lead to such an hypothesis without any addition- al proof.” So far Wicutr and ARNOTT. HOOKER continues :— “In relative size especially, the observer will find immense variation; for, unlike the animal creation, proportional dimensions are of small moment in the vegetable kingdom. This fact, so familiar to the botanist of experience, is always a puzzle to the zoologist, who fancies he perceives a vagueness and want of exactness in all botanical writings (except in those of the too numerous class that make a parade of measuring to lines organs that vary inches), that contrasts unfavourably with des- criptive zoology. Symmetry is again only a relative term amongst plants, for even such leaves as grow in pairs are never alike, and often differ much in form, texture, and colour; whilst the various sepals, petals, etc. of an individual flower, never so exactly correspond as the relative members of an animal do; and there are still greater differences between these organs, when taken from different flowers.” “Tt is hardly necessary to allude to the desira- bility of studying the various forms induced by artificial causes: the browsing of cattle on shrubs, for instance, which is almost invariably followed, by an abnormal state of foliage on the subsequently developed shoots, has been a prolific source of bad species; while there is scarcely an operation of man that does not tend to produce change in the vege- tation surrounding him.” “It will generally be found that botanists who confine their attention to the vegetation of a cir- cumscribed area, take a much more contracted view of the limits of species, than those who extend their investigations over the whole surface of the globe. This is partly, no doubt, owing to the force of bad example; and partly to the fact that the student who takes up the study of the flora of his native country, finds that the species are all toler- ably well known, and that no novelty is to be discovered. There is therefore a natural tendency to make use of trifling differences, from the scope which they afford for minute observation and critic- al disquisition; whilst the more close comparison of the few species which come under his investi- gation, leads the local botanist to attach undue importance to differences which the experienced observer knows may be safely attributed to local circumstances. To this tendéncy there can be no limit, when the philosophy of system is not under- stood; the distinctions which appeared trifling to XVI FLORA MALESIANA [ser. I, vol. 4! botanists a quarter of a century ago, are at the present day so magnified by this class of observers, that they constantly discover novelties in regions which have been thoroughly well explored; con- sidering as such, forms with which our predecessors were well acquainted, and which they rightly re- garded as varieties. ! ‘Another result of the depreciated state of systematic botany is, that intelligent students, being repelled by the puerilities which they everywhere encounter, and which impede their progress, turn their attention to physiology before they have ac- quired even the rudiments of classification, or an elementary practical acquaintance with the charac- ters of the natural orders of plants. Unfortunately, in botany, as in every other branch of natural science, no progress can be made in the study of the vital phenomena except the observer have a previous accurate acquaintance with the various modifications under which the individual organs of plants appear in the different natural orders, and such an appreciation of the comparative value, structural and morphological of these modi- fications, as can only be obtained by a careful study © of the affinities of their genera and species. Igno- rance of these general laws leads to misinterpre- tation of the phenomena investigated by the phy- siologist, and to that confusion of ideas which is so conspicuous in the writing of some of the astute physiological observers of the day.” “The modern system of botanical instruction attempts far too much in a very limited space of time, and sends the student forth so insufficiently grounded in any branch of the science, that he is unprepared for the difficulties which he encounters, let his desire to progress be ever so great. The history of botanical discovery, and the philosophy of its advance, form instructive chapters for the student in any department of natural science.” “We owe to LINNAEUS the establishment of the doctrine of the sexuality of plants; and we find by the writings of the same great naturalist, that be- sides foreseeing many physiological discoveries, he preceded GOETHE in the discovery of morpho- logy, a doctrine which, more than any other, has tended to advance scientific botany. A third great discovery, that of the nature of the ovule, and the relation of the pollentube to the ovary, received its principal illustration at the hands of BRown, our chief English systematist, and of BRONGNIART, also a practised botanist.’ “Tt should not be forgotten, that the relative importance of physiology is very different in the animal and vegetable kingdoms. In the former, structure and function operate so directly upon one another, that the great groups are, to a certain extent, defined by well-marked external characters, which are at once recognizable by the student, and are familiar, or at least intelligible, to those even (1) ‘‘Many of the species which have been revived in modern times were indicated by HALLER, RAy, TOURNEFORT, and other ancient botanists, but were reduced to the rank of varieties, when the science was reformed by LINNAEUS.” who have paid no attention to natural history. In the vegetable kingdom this is by no means the case: the processes of assimilation and secretion present but little of that complication which ren- ders the study of animal physiology so important; they are, on the contrary, uniform almost through- out its whole extent, and moreover so simple in their modus operandi, that this very simplicity pre- vents their being rightly understood. In conse- quence, even the two great classes of Monocotyle- dons and Dicotyledons are not distinguishable without considerable practice and study; and were we dependent upon actual inspection of the organs whence the essential characters of these groups are drawn, for the means of recognizing, Systematic Botany would be an impracticable study.” ‘Herein lies one great obstacle which meets the beginner on the very threshold of his botanical studies: he sees the great divisions of the animal kingdom to be recognizable by mere inspection, and that familiar characters are also natural, and available for purposes of classification: the very names of the groups convey definite information, and to a great extent give exact ideas. Birds, fishes, reptiles, etc. are all as natural as they are popular divisions; but what have we in the vegetable king- dom to guide the student through the two hundred and fifty natural orders of flowering-plants? As with a new language, he must begin from the very beginning, and also avail himself of artificial means to procure as much superficial knowledge of struc- ture and affinity as shall enable him to see that there is a way through the maze. Hence the obvious necessity of an artificial system of some sort to the beginner, who has, at the same time, to master a terminology, which, if not so complex as that of zoology, is more difficult at the outset, from the want of standards of comparison between the or- gans of plants and those he is familiar with in himself as a member of the sister kingdom. Ap- plying these remarks to practice, the botanical student finds that he has much to unlearn at the very outset; in many cases he has misapplied the terms root, stem, leaf, efc., and contracted most erroneous ideas of their structure and functions; while he is startled to find that the popular di- visions of plants into trees, shrubs, and herbs, —leafy and leafless, water and land, erect, climb- ing, or creeping,—are valueless even as guides to the elements of the science.” “Tt is not however to be supposed, because pure physiology is of secondary importance to the right understanding of the affinities of plants, that botany is therefore a less noble or philosophical study than zoology; since we find anatomy, development, and morphology, occupying a very far higher rank in proportion. Being deprived, as he is in most cases, of all technical aids to the determination even of the commoner exotic natural families, the systematist is compelled to commence with the knife and microscope, and can never relinquish these implements. Systematic Botany is indeed based upon development; and no one can peruse, however carelessly, the most terse diagnosis of a natural order or genus of plants, without being ~ Dec. 1948] General considerations XVII struck with the variety and extent of knowledge embodied as essential to its definition and recogni- tion. Not only are the situation and form, division or multiplication, relative arrest or growth, of the individual organs exactly defined, in strictly scien- tific and scrupulously accurate language, but the development of each is recorded from an early stage: the vernation and stipulation of the leaves; the aestivation of the young calyx and corolla, and their duration relatively to other organs; the de- velopment and cohesion of the stamens; the po- sition and insertion of the anther; its pollen; the cohesion or separation of the carpels, and the stages of their development from the bud to the mature fruit, and from the ovule to the ripe seed, are all essential points; all however minute, must in many cases be actually inspected before the position ofa doubtful genus can be ascertained in the Natural System; and this is not the exception, but the rule.” “The necessity for acquiring so extensive and detailed a knowledge indicates a power of variation in those organs from which the natural characters are drawn, that defeats any attempt to render one, or a few of them only, available for the purposes of classification; and hence it is that the study of morphology or the homologies of the organs, be- comes indispensable to the systematist; by this he reduces all anomalies to a common type, tests the value of characters, and develops new affinities. The number, form, and relative positions of organs may supply technical characters, by which ob- servers of experience recognize those natural orders under which a great number of plants arrange themselves; but a knowledge of structure and ana- tomy alone enable the botanist to progress beyond this, and to define rigidly: whilst the study of development affords him safe principles upon which to systematize and detect affinities, and mor- phology supplies the means of testing the value of the results, and reveals the harmony that reigns throughout the whole vegetable world.” **Physiology, again, is a branch of botany very much apart from these: its aim is the noblest of all, being the elucidation of the laws that regulate the vital functions of plants. The botanical student of the present day, however, is too often taught to think that getting up the obscure and disputed speculative details of physiology, is the most useful elementary information he can obtain during the short period that is given him to devote to botany; and that, if to this he adds the scrutiny of a few of the points under a microscope, he has made real progress as an observer. This, we maintain, is no more botany, than performing chemical experi- ments is chemistry, or star-gazing astronomy. A sound elementary knowledge of vegetable physi- ology is essential to the naturalist, and should indeed be a branch of general education, as it requires nothing but fair powers of observation and an ordinary memory to acquire it. For the student to confine his attention to this knowledge of the vegetable world, and to try and improve upon it by crude experiments of his own, under- taken in ignorance of the branches of pure botany we have enumerated, is a very rational amusement, but nothing more.” “The students are indeed, in too many cases, perfectly ignorant of the elements of natural science, and require some practical acquaintance with plants and their organs, before they can ap- preciate the relations of the different branches of botany to one another, or discriminate between what.is essential to understand first, and what is better acquired afterwards. Were the elements of science taught at schools, this would not be so: we should then have the student presenting himself at the botanical lectures fully prepared for the more difficult branches of science, and for making that progress in them for which the professor’s aid is indispensable. A sound practical knowledge of system we hold to be an essential preliminary to the study of the physiology of plants—a study which requires also a practical acquaintance with organic chemistry, consummate skill in handling the dissecting knife, and command over the micro- scope, a good eye, a steady hand, untiring per- severance, and aboveall, a discriminating judgment to check both eye, hand and instrument. A combi- nation of these rare qualities makes the accom- plished vegetable physiologist, and their indispensa- bility gives physiology its pre-eminence in practice.”’ “Tt has been with no desire of obtruding our views upon our readers that we have ventured to discuss these obscure subjects with relation to Indian plants, but from a conviction, that in the present unsatisfactory state of systematic botany it is the duty of each systematist to explain the principles upon which he proceeds; and we do it not so much with the intention of arguing the sub- ject, as of pointing out to students the many funda- mental questions it involves, and the means of elucidating them.” “To every one who looks at all beneath the surface of descriptive botany, it cannot but be evident that the word species must have a totally different signification in the opinion of different naturalists; but what that signification is, seldom appears except inferentially. After having devoted much labour in attempting to unravel the so-called species of some descriptive botanist, we have some- times been told that the author considers all species as arbitrary creations, that he has limited the forms he has called species by arbitrary characters, and that he considers it of no moment how many or how few he makes. So long as this opinion is founded on conviction, we can urge no reasonable objection against its adoption; but it is absolutely necessary that the principle should be avowed, and that those who think the contrary should not have to waste time in seeking for nature’s laws in the works of naturalists who seek to bind nature by arbitrary laws. So again with regard to specific centres; except we are agreed with an author as to whether the same species has been created in one or more localities, and at one or more times, we shall be at cross purposes when discussing points and principles relating to identity of species and geographical distribution.” “Great differences of opinion have from the XVIII FLORA MALESIANA [ser. I, vol. 4! earliest days of science always existed on the nature of species. The prevalent opinion has undoubtedly at all times been, that a species is a distinct creation, distinguishable from all others by certain perma- nent characters. Many eminent philosophers, how- ever, have taken a contrary view; of these the best known have been LAMARCK, and more recently the anonymous author of the ‘Vestiges of Creation’.”’ —So far HOOKER. Modern biological science has progressed rapidly in the last decades through the results of experi- mental genetics. Though it is far from easy to weld the often contradictory opinions into a satisfactory whole, views relating to matters of variation have much gained. In the following pages I will try to discuss on this new basis the value to be attributed to characters of less than specific importance and a number of considerations which may lead to increased accuracy in judging specific delimitation. Trifling characters, such as peloric and cleistoga- mous flowers, have led to the creation of worthless new genera; galls, insect bites, and parasitic fungi have been mis-interpreted and caused the publi- cation of new species of Phanerogams. Jndividual variations, either intrinsic or extrinsic have, in a similar way, induced systematists working on tropical plants to distinguish more species than Nature intended. It is not our intention to limit phytography to a merely administrative function in the study of botany, but to treat it as an essential of natural philosophy. The systematist ought to keep pace with cytogenetics, physiology and morphology,! phytochemistry, phytogeography, ecology, genetics, i. e. experimental taxonomy. Inadequate material? and information are the chief causes which prevent the phyto-systematist from applying the results obtained by these branches of botanical science. The systematist is seldom favourably regarded by the layman or student of directed botany. They are opposed to changes in nomenclature, being unable to gauge the force of the arguments for a ‘new’ name for a familiar plant and so rarely accept the judgment of taxonomists. When, on the other hand taxonomic problems are tackled by applied scientists nomenclature and specific distinction become chaotic; entomology, mycology, forestry, agriculture and horticulture supply many examples. A wish for simplification, impatience, or even per- sonal vanity or the desire for pecuniary gain have caused hosts of ‘species’ to be added to our lists (1) For the value of wood anatomy in taxonomy, see DEN BERGER, in Handel. 4e N.I. Natuurwet. Congres (1926) 397. (2) Cf. WiGuT, in a letter to GriFFITH, dated April 15, 1842:—‘“‘How people can work on dry plants I cannot imagine. I am daily convinced of the poverty of the study from such materials, unless a man has seen much of living structure.” MIQUEL ignored this remark, and on sterile and inadequate material based a host of species from Sumatra which even at present are not wholly elucidated. by applied workers. Not long ago a forestry officer made a study of Agathis? in Malaysia in which 13 species and 2 doubtful ones, that is 15 entities, were distinguished. In the same material the late Dr DANSER, whose judgment and experience cannot be doubted, distinguished only 3 divergent species with a number of local geographic variations. He found it very difficult to define the latter. Additional material showed that the keys and distinctions presented for the 13 species did not hold to the satisfaction of the Forest Research Station, from which this work emanated. In plant families of economic importance particularly in Gramineae, Rutaceae, and Leguminosae, similar work has re- sulted in multitudes of microspecies provided with binomials; by such a proceeding nothing is gained and much lost. An example of the difficulties arising between taxonomy and an applied science when a good revision is absent, is the following:—a Clausena of unknown origin was cultivated for economic purposes at Buitenzorg. I referred it to Clausena _ anisum-olens (BLCO)MERR. but the phytochemist was dissatisfied, the properties of the oil did not tally with data recorded from the same species in the Philippines. I then sent ample material with full notes to Dr TANAKA, Dr SWINGLE, and to the Kew and Paris Herbaria, for identification. The answers were all different and the phytochemist was, of course, disgusted with the practical results of taxonomy, because now he had the choice among 5 names for his plant. By way of comfort I expressed the hope that a systematist would some day make a satisfactory monograph of the genus.* In order not to raise his hopes too high I remarked that even then some research from him would be needed to establish the assumed constancy of the oil properties as a specific character. I also inform- ed him that taxonomy has sometimes scored by predicting phytochemical facts, e.g. when HALLIER supposed the presence of valerianic acid in Vibur- num> on phylogenetic grounds only. In the following two chapters general infor- mation on variation as a source of superfluous binomials is collected for the benefit of those with no field experience of the Malaysian flora. I distinguish variations induced by the environment from those belonging to the genetic composition of populations, and I have tried to illustrate them by examples in Malaysian phytography. Often the number of examples is too small, and chapters overlap, but in the course of time every student of Malaysian botany will meet with other equally telling cases. May they stimulate the wish to avoid lapses of this character by conscientious treatment of the revisions in Flora Malesiana. (3) Bull. Jard. Bot. Btzg III, 16 (1938) 455-474. (4) Compare R. WiGur in a letter to GRIFFITH, dated March 30, 1841:—... ‘‘as you say Botany is difficult, and increasingly so, but Botanists are to blame for this. No remedy will be so effectual as the publication of Monographs.” (5) Med. Rijksherb. Leiden no 14 (1912) 36; ibid. 37 (1918) 92. Cf. also V. valerianoides ELM. Dec. 1948] General considerations XIX VARIATIONS MOSTLY INDUCED BY THE ENVIRONMENT (Phenotypic modifications) Phenotypic modification is the response to environ- mental conditions, such as climate, soil, exposure, altitude, temperature, wind, fire and living organ- isms. The genetic qualities govern the character of the plant, but the environment in which the plant develops determines the actual and final appear- ance of the individual. The changes or differences from the ‘normal plant’ are called modifications. Such changed characters are not themselves in- herited, however, though the manner in which a plant reacts to environmental conditions is. In some cases an external change may be reversed by a change in the environment during the develop- ment of the individual but in other cases, when factors act in the seedling stage only, the effects in the individual are irreversible. It is necessary to agree about the concept ‘normal plant’. This is far from easy, as each speci- men grows under a different combination of CEB- factors (Climatic, Edaphic, Biotic). We might ap- proach the idea by saying that “‘the normal plant results froma genetically average individual under average natural environmental conditions”, average to be understood in the sense of optimal. This ‘normal’ individual is never a reality but remains an abstraction. Though the difference between phenotypic and genotypic variation is clear, the field botanist—and still more the herbarium botanist—is not always able to recognize it. Only experiments may furnish proof. For instance a dwarf shrub in an area sub- ject to fire or browsing animals may assume this stunted form through these CEB-factors but it is also possible that the stunted form is a specialized race adapted to these conditions and thus selected by nature itself from the specific population. Ex- perimental breeding must decide its constancy. I have arranged the phenotypic modifications under several headings—which partly overlap and interlock—in the following sequence: 1. Juvenile forms. , , ADs xix 2. Precocious flowering ( pacdocenest s). 2 ; ; ‘ XXI Grioecnesmorphesis 3. Dimorphous foliage : XXil 4. Dimorphous seeds and fruits XXiV Intrinsic 5. Dimorphous flowers XXV 6. Cleistogamous flowers XXV ine ee 7. Teratological forms XXV ee cca, ‘ 8. Phenotypic effect of altitude XXVi . { 9. Epiphytes XXVili Climatic. J OH DHROMD AIS) 110. Shade forms . XXix Hygro-morphosis . . 11. Influence of drought XXIX [ orsmorohos ; . 12. Seasonal variation XX1X Anemo-morphosis . 13. Wind forms XXX ( 14. Fumarole plants XXX |S senlesaiologk 15. Rock plants; calcareous and silicious soils XXXi Edaphic / 16. Solfatara plants XXXIii ( Hydro-morphosis . . 17. Water and swamp plants XXXIV Phyto-morphosis . . 18. Fungus and bacterial diseases, and symbiosis XXXV \ ( 19. Ant plants (myrmeco-morphosis ) XXXV Bioti Zoo-morphosis 20. Galls deceptive to phytographers ( cecilia: -morphosis) . XXXVI iotic 11. Infl fb | : 21. Influence of browsing animals (pascuo-morphosis) XXXVI ] ( 22. Influence of fire (pyro- nei XXXVii _ Anthropo-morphosis 23. Pioneer plants . XXXVili ( 24. Savannah trees . XXXVill Ontogeno-morphosis 1. Juvenile forms Juvenile forms often differ widely from the mature plant. Seedlings of many Leguminosae differ greatly from the adult in foliage and other charac- ters. The youth form of Cassia javanica L. pos- sesses large metamorphosed twigs acting as thorns (1). Thorny juvenile specimens are also found in Alangium. In general, flowering twigs have smaller leaves than sterile branches; this often gives rise to diffi- culty in identifying non-flowering material and s one of the pitfalls if new species are based on sterile material. An example is Campnosperma acutiauris BoeRL. & KoorD. (Anacardiaceae) described on sterile juvenile material from Sumatra. The leaves are large and conspicuously auriculate-amplexi- caulous. A similar juvenile form was later found in West Java, together with mature trees. These possessed much smaller non-auriculate leaves (fig. 2). The plant appeared to represent a species of Tristania (Myrtaceae) (2); its specific identity will probably remain obscure, however, as several species produce similar juvenile forms. XX FLORA MALESIANA [ser. I, vol. 4! Youth forms of Myrica longifolia T. & B. differ strongly from mature specimens in possessing distinct stipules and incised larger leaves. Incised leaves of seedlings occur in a score of arboreous plants, e.g. many Bignoniaceae, Pro- teaceae, Gmelina, Lonicera, Alangium, Vitex, &c. Leaves of young trees of Pangium edule REINW. Fig. 2. Large leaf of a juvenile specimen, small leaves of a mature tree of Tristania sp. (Myrt.) in W. Java, x 1/3. are 3-lobed whereas the leaves of mature trees are entire. The growth of different parts of the plant is often very disproportionate. In some Symplocos species I found the leaf teeth were mature and large in young leaves but inconspicuous in mature foliage: they possibly have some (?excretive) function during youth only. BACKER (3) found the leaf tip earlier developed than the blade in some species of Dioscorea; it disappears also sooner. A peculiar development occurs in the growing leaves of some Meliaceae, e.g. Chisocheton (fig. 3). Very peculiar juvenile forms greatly differing from the later normal foliage, have been describ- ed in various climbing plants such as some spp. of Adenia, Medinilla, Macrozanonia, Piper, Araceae, Ficus, &c. Juvenile specimens of these trunk climbers are always sterile. Their foliage is mostly broader than that of mature plants, and is appressed to rocks or tree trunks. The similarity in their appearance may cause considerable con- fusion as e.g. is shown by the type specimen of Ficus peltata BL. which was recently proved to represent a juvenile specimen of some climbing species of Piper. In several Malvaceae, Leguminosae, Sterculi- aceae, Tiliaceae, juvenile leaves are often different from mature ones (fig. 4). Sterculia polyphylla R. Br. is a juvenile stage of St. foetida L.; young trees often possess leaves having 10-15 narrow leaflets, mature trees have mostly 5—9-foliolate leaves with broader segments. Young Lasia spinosa THwW. is very different from the mature plant. Ficus basi- dentula Mia. is merely the juvenile form of F. cal- losa WILLD.; it is quite common in the hedges at Buitenzorg. The polymorphy in the habit and foli- age of Ficus quercifolia Roxs. and F. heterophylla L.f. is unbelievable. In juvenile forms of Nepenthes the shape of the pitchers may considerably differ from that in mature plants; asa result juvenile Nepenthes cannot with certainty be identified. In Carallia lucida Roxs. leaves of mature trees are oblong to obovate with very shallowly serrate to entire margins; saplings, however, have oblong to lanceolate leaves distinctly serrate (4). Other cases of old mature plants differing from young ones are found among lianas in which the shape of the stem may change considerably: Cissus . tuberculata Bu. has terete tuberculate stems but they later become flat and, in older stems, up to 60cm broad looking like gigantic ribbons! The latter were described as a separate species, Vitis lanceolaria WALL., but the two forms are merely two stages of one species. The stems of lianas generally change greatly with age, through the de- velopment of corky warts and wings, together with secondary wood not present in young flowering twigs. Spines sometimes disappear in lianas and trees with age; in some cases, on the other hand, they enlarge considerably. A peculiar case is repre- sented in two undescribed Cucurbitaceous lianas from the Lesser Sunda Islands, both having a Fig. 3. Growing leaf tip of Chisocheton sp. (Meliac.) (ole), PPM), SX She Dec. 1948] swollen base; in Gynostemma sp. this ‘podagric’ ‘base is smooth, in A/somitra sp. it is spiny (5). Habit also sometimes changes with age: Anci- strocladus is sometimes a shrub in youth whereas later it becomes distinctly scandent. Climbers which have no support may sometimes grow into semi-erect shrubs; this I once observed in a plant Fig. 4. Hibiscus sagittifolius KURZ (Malvac.), leaves from one specimen (Indramajoe, W. Java), * 1/2. of Smilax modesta DC. in a grass field on Mt Diéng. Spotted leaves often occur only or predomi- nantly in juvenile specimens. Strobilanthes picta KoorbD. was a new species proposed on account of silvery spots on the leaves. However, it is a juvenile stage of Str. cernuus BL. Similar cases are known in Begonia, Cissus, and other genera where these spots may disappear with age. In greenhouses these juvenile forms are preferred for ornamental purposes. Juvenile forms of plants with pinnate leaves sometimes have a much larger number of pinnae e.g.Campsis pandorana (ANDR.)STEEN. c.n. (6). An example of a new species based on a juvenile plant is found in Dacrydium: the type specimens of Dacrydium junghuhnianum Mig. from Sumatra consist of juvenile specimens of D. elatum WALL. with long loosely set needles. In the herbarium flowers sometimes open during drying and create the impression of being mature. This is specially the case with flowers having val- vate terete corollas e.g. Symplocos § Cordyloblaste, Styrax, Polyosma, Proteaceae, Loranthaceae, &c. MIQUEL described a new species of Lonicera from Sumatra L. sumatrana Mia. In his description short corollas are mentioned; owing to this mistake the species was subsequently placed in the wrong sec- tion and described twice again, once from Burma General considerations XxI and once from Sumatra (7). The examination of MIQUEL’S type specimen revealed that he described immature flowers, in fact buds which had opened in the herbarium. Immature woody capsules or strobili of Myrtaceae, Theaceae, Coniferae, Casua- rina, &c. also tend to open after drying. There is often a great similarity in the leaves of watersprouts of mature trees with those of saplings: large size, deeper incised teeth, thinner texture, e.g. in Symplocos, Ficus, Sapotaceae, Dipterocarpa- ceae, etc. A still unsolved case is that of Evonymus japo- nicus THUNB. of which a sterile slender climbing and rooting form is frequently found in the Javan mountain forests. I originally took it for a juvenile shade form (8). Not until 1941 did I succeed in finding it flowering and fruiting on the open sum- mit of Mt Jang. It is unknown whether the shade conditions in the juvenile stage determine the later morphology. Cited literature: (1) A. J. KoENS, De Trop. Na- tuur 2 (1913) 174; see also Koorpers, Bull. Jard. Bot. Btzg. III, 1 (1919) 168. (2) Tectona 22 (1929) 1336-1340. (3) Handboek Flora Java pt 3 (1924) 109. (4) Schoolflora voor Java (1911) 486. (5) Fig- ured in De Trop. Natuur 29 (1940) 6. (6) Bignonia pandorana ANDR. (7) Journ. Arnold Arbor. 27 (1946) 441, 445. (8) De Trop. Natuur 22 (1933) . 175-176. 2. Precocious flowering (paedogenesis) In several Malaysian plants pre- cocious flowering is observed. CosteRus (1) recorded flowering seedlings in Melia arguata DC. (fig. 5). BACKER found them in Melia azedarach L. and J. J. SMITH described (2) the same pheno- menon in Murraya paniculata L. In Cocos nucifera L. precocious flowering is often seen. The late Dr A. RANT observed flowering seedlings in Cinnamomum zeyla- nicum THw. (oral comm.). Other plants in Malaysia in which pre- cocious flowering has been observ- ed are Swietenia mahogani JACQ., Coffearobusta, Citrus decumana L., Nicotiana tabacum L., Sesbania sericea DC., Vigna sinensis ENDL., Teramnus labialis SPRENG., Tectona grandis L.f., Kalanchoe pinnata Pers., and Ailanthus sp. In plants which flower strictly periodically precocious flowering is sometimes controlled by the Fig. 5. date of sowing. If sown too late Precocious they flower together with full- flowering grown plants sown earlier. This (paedogenesis) is a fact well-known to agricul- in Melia turists (in Java e.g. in Hibiscus arguta DC. Spp.). (Meliac.). Precocious flowering may also (after be caused by poor soil or some COSTERUS) XXII FLORA MALESIANA [ser. I, vol. 4! methods of pruning. An example of the former cause is Osbeckia pusilla ZOLL. which is a flowering dwarf of O. chinensis L. on poor soils. Sometimes dwarfed plants flower when very small and represent distinct varieties or strains, e.g. the dwarf of Canangium odoratum BAIL. f. pumila (3) grown in pots in Malaysia (introduced Fig. 6. Monophyllaea horsfieldii R.Br. (Gesn.), adult plant, one cotyl large and leafy, the other (in front) bract-like, soon disappearing (W. Java, Kalapa Noenggal), x 2/s. from China), Aglaia odorata Lour. var. microphyl- lina DC., and a dwarf of Punica granatum L. The skill of Chinese and Japanese horticulturists in raising dwarfs is due partly to the selection of pygmy varieties but more important is their skill in impoverishing the plant without starving it (4). Many dwarfs are found near solfatara, on rocks, and on silicious soils (cf. § 15-16). Flowering juvenile forms are comparable to the neoteny found in the animal kingdom. Pteridophytes generally are apparently more plastic with regard to precocity than Phanerogams, and several species are known to form spores in dwarf or juvenile specimens which have some- times been described as distinct species. It has been assumed that pygmy species in Antrophyum may possibly represent neotenous stages of other species. COPELAND described in 1939 (5) a dwarf fern from Borneo as Holttumia, but it is Dr DONK’s conten- tion that this fern is a precocious stage of a Tae- nites. In the genera Teratophyllum, Stenochlaena and Lomariopsis, HOLTTUM (6) was able to demon- strate that a great deal of confusion is caused by the description of juvenile stages; being familiar with the living plants in the field he clarified the true status and affinities of a number of obscure species. DrELs (7) compiled an instructive book on juve- nile forms, giving instances where the juvenile foliage persists in the mature flowering plant, a course of development comparable to neotenous forms in zoology. Australian and New Zealand botanists have written a great deal about this phenomenon of heteroblasty which in those floras -has apparently an important bearing on speciation (8). I cannot remember a Malaysian plant sus- pected to represent such a case. Yet such strange plants as Monophyllaea (fig. 6) and ailied genera of the Gesneraceae living on the enlarged cotyle- dons might be examples. Cited literature: (1) Rec. Trav. Bot. Néerl. 1 (1904) 128. (2) De Trop. Natuur 1936, Jub. uitg. p. 73. (3) Now described as a separate species Canangium fruticosum CratB (Kew Bull. 1922, p. 166) being cultivated in Siam. (4) Compare F. A. McCLure, in Lingn. Sci. Journ. 12 (1933) Suppl. p. 119-149. (5) Philip. Journ. Sci. 74 (1941) 153-156. (6) Gard. Bull. Str. Settlem. 5 (1932) 245 seq.; ibid. 9 (1937) 139 seq. (7) Jugendformen und Bliitenreife im Pflanzenreich 1905. (8) cf. COCKAYNE, 13th Meeting Australas. Ass. Adv. Sci. (1912) 217 seq. 3. Dimorphous foliage It was observed by F. W. WENT (1) that in trees generally the foliage of the lower branches is larger than that of the upper twigs. He ascribes this to the amount of water available to different shoots (internal water-conducting capacity) ; so, in mature trees the upper foliage would be insufficiently pro- vided with water. The leaves of water sprouts, on the other hand, are mostly exceedingly large as their water supply is abundant. Leaves of these shoots are mostly hardly recognizable in the her- barium, as they may reach disproportionate di- mensions. Foliage for description in the herbarium ought therefore to be comparable and preferably that of flowering twigs. The dimorphy of the foliage is mostly linked up with a difference between flowering and non- flowering parts of the plant, similar to that found in Hedera. It is conspicuous in several climbing Ficus. Piper, Araceae, and in some Conifers. A strik- ing example of plagiotropically flowering twigs is that of Abroma angusta L.f. A good illustration is also Luvunga sarmentosa (BL.) Kurz (Rutaceae). The stem shoots of this Dec. 1948] liana possess large straight axillary thorns and 1-foliolate leaves. The climbing shoots, however, possess conspicuously curved thorns and 3-foliolate leaves and the flowering parts of these are often unarmed. L. eleutherandra DALZ. was based on a type different from BLUMe’s but is actually the same species, as was found by Kurz (2). Fig. 7. Heterophylly in Ficus deltoidea JACK (= F. diversifolia Bu.) (Morac.), Mt Gedeh, W. Java, x 2/3. Putting into practice what he had read of Hedera helix in a botanical manual, Mr BoLt made a remarkable application of the dimorphy of Piper cubeba L. Near Semarang, instead of cultivating it as a climber he took cuttings of the flowering twigs, and got shrublets which, though small, pro- duced abundantly ‘tail pepper’. Plants with dimorphous foliage are very numer- ous in Malaysia and species are frequently named after this peculiarity. Ficus deltoidea JACK (= Ficus diversifolia BL., fig. 7) is one of them; L. VAN DER Pu (3) could not find any regularity in its hetero- phylly. In Faradaya dimorpha PULLE from New Guinea there are two kinds of twigs, with decussate and with 3-whorled leaves of different shape. Phy- tocrene macrophylla BL. has both entire and 3-lobed leaves on one individual, as have Broussonetia sumatrana Miq., Knema heterophylla WARB., sever- al species of Gmelina and Sterculiaceae, Tiliaceae, Artocarpus varians MiqQ. A good case is also Uraria picta Desv. (fig. 8). Heterophylly is common in ferns. Polymorphy in leaf shape among different indi- viduals of a population is a subject which ought to have a separate heading. It is of universal oc- currence in the Malaysian flora, and has (e.g. in Cucurbitaceae) given rise to a multiplication of names. In Coccinea, MIQUEL (4) distinguished two species, one with incised leaves and one with angu- lar entire leaves: according to BACKER they are identical, the incised leaves mostly belong to juve- General considerations XXII Fig. 8. Uvaria picta Desv. (Legum.), with heterophyllous foliage, Kangean Island, moist Imperata fields at low alt., x 1/2. XXIV FLORA MALESIANA nile specimens. In Gymnopetalum cochinchinense (Lour.) Kurz there is even more confusion: speci- mens with incised leaves have been described as G. septemlobum Mia., G. qguinquelobum Mia. and G. quinquelobatum CoGN., those with angular or entire leaves as G. piperifolia Miq. and G. hors- fieldii MQ. There is probably a host of other names X ‘A SF V7 4 Fig. 9. Macrobiocarpy in Callistemon speciosus DC. (Myrt.) with 3 sets of fruits below the terminal bud, originating from 3 previous flowering periods, x 1/4. for this species which is very variable in foliage. In Trichosanthes a similar polymorphy of the foliage caused superfiuous description of species. Tr. gran- diflora BL. is, according to BACKER, a form of Tr. globosa BL. with lobed leaves. In Gynostemma CLARKE and BACKER assume a variability in foliage (3-foliolate to pedately lobed leaves) which will cause a considerable reduction of the number of species. Similarly scores of superfluous names are found in polymorphic species such as Urena lobata [ser. I, vol. 4! L. etc. In the Oleaceae, Nyctanthes dentata BL. is only a dentate-leaved form of N. arbor-tristis L. In some Pteridophytes heterophyllous leaves are well known. The most striking examples occur in the genera Teratophyllum, Stenochlaena and Loma- riopsis where according to Hortrum (5) hetero- phylly has caused much taxonomic confusion. An other striking case is that of the plant which is mostly cited as Lindsaya repens (BORY) BEDD. as demonstrated by W. TROLL (6). Cited literature: (1) Handel. Se Ned. Ind. Nat. Wet. Congres (1928) 385-392 (1929). (2) Journ. As. Soc. Beng. 39 (1870) 69. (3) De Trop. Natuur 27 (1938) 89. (4) Flora Ind. Batavae 1, 1 (1855) 673. (5) Gard. Bull. Str. Settlem. 5 (1932) 245; ibid. 9 (1937) 139. (6) Flora 126 (1932) 408. 4. Dimorphous seeds and fruits Of heterocarpy (1) only few examples are known in the Malaysian flora. In some Compositae the marginal fruits are sometimes strikingly different - from those produced by the central tubular flowers, as was described for Synedrella nodiflora GAERTN. by A. Ernst (2). It is also known that in Tragia volubilis L. normal and 2-hooked one-seeded fruits may occur together. In Umbelliferae normal fruits and fruits with one half reduced may sometimes be observed. In Leguminosae also different types of fruit are sometimes found on one plant. In Desmodium heterocarpum DC. the lower pods are 1-seeded, the upper 5—7-seeded. Dimorphous fruits and seeds are known in Aeschynomene spp. and in the genus Jussieua. One of the most curious cases of dimorphous fruits is that detected by BACKER (3) in the common Acalypha indica L. in Java where the tip of the male spikelets is crowned by a single female flower de- veloping into a T-shaped fruit with a central fertile and 2 lateral sterile cells; the central cell seems to be sunken in the tip of the axis of the rachis. The normal capsule consists of 3 equal cocci. A special case is that of macrobiocarpy (4) when not all fruits dehisce at the end of the season but a number remain closed on the plant and grow for years larger and woody. Sometimes fruits of 3-4 seasons are found on one twig, which thus keeps a reserve of seeds. Macrobiocarpy seems to be mainly restricted to the semi-arid climates and is of definite advantage in fire-swept areas. It is very common in some genera of capsular Myrtaceae (fig. 9), viz Leptospermum, Eucalyptus, Melaleuca, Agonis, Metrosideros, Syncarpia. It possibly also occurs in some Proteaceae, Coniferae, Casuarina, and some Rubiaceae. The woody structure, large size and modified shape of the fruits formed in previous seasons must be allowed for in identifying the species. Inadequate material may cause considerable confusion. Cited literature: (1) DELPINO, Mem. R. Ac. Sc. Inst. Bologna V, 4 (1894). (2) Ber. Deutsch. Bot. Ges. 24 (1906) 450-459. (3) Onkruidfiora Jay. Sui- ker. (1930) 406-407. (4) WINKLER, Ann. Jard. Bot. Btzg 20 (1905) 37-41. Dec. 1948] General considerations XXV 5. Dimorphous flowers A most peculiar case of flower dimorphy occurs in some Orchidaceae viz in Renanthera lowii Rcup. f. (1) and Grammatophyllum speciosum BL. (2). The shape and colour of the lower flowers in a raceme are very much different from those of the upper ones. In these Orchids the occurrence of aberrant lower flowers is a normal phenomenon. It seems also to occur in some species of Arachnis and less obviously in some species of Bulbophyllum. In Oberonia imbricata LINDL. the upper flowers of the spike are abnormal and their gynaecium is reduced. Dimorphous flowers are also frequently found in dioecious and polygamous plants. Male and female flowers are sometimes very different in shape and size, e.g. in Mangifera, Brucea, He- vea, &c. In thyrsoid inflorescences the marginal flowers are often different from the central ones, or some- times the central ones are reduced or deformed. A conspicuous instance is Mussaenda where some flowers of the inflorescences have one calyx lobe large and leafy. Other cases are found in Hy- drangea, Sambucus javanica REINW., some Umbel- liferae, some Araliaceae, e.g. Boerlagiodendron, and some Mimosaceae. Cited literature: (1) WINKLER, Ann. Jard. Bot. Btzg 20 (1906) 1. (2) CosTERUs, Dodonaea 6 (1894) 24. 6. Cleistogamous flowers Cleistogamous flowers occur frequently in the Ma- laysian flora. A general survey has hitherto not been compiled. They were described in Clitoria by HArms (1) and RANT (2) where they are sometimes more fre- quent than normal flowers. The description of specimens with cleistogamous flowers has led here to phytographical confusion: the American genus Martia LEAND. SACR. was based onacleistogamous leguminous plant which is, actually, according to BENTHAM and Harms (/.c.) nothing but the cleis- togamous state of Clitoria. Cleistogamy also occurs in Malaysian species of Viola. It is stated by BECcARI (3) to occur in several Bornean Annonaceae. A very typical example is described in Commelina benghalensis L. by J. VAN WELSEM (4): cleistoga- mous flowers are present on subterranean shoots. Another well known case in a common plant is Ruellia tuberosa L. mentioned by VAN WELSEM (5) and A. F. G. Kerr (6). Cleistogamous (better: cleistopetalous) aie ers are common in Orchidaceae as J. J. SmitH and R. SCHLECHTER both frequently mentioned. The for- mer gives a list of cases known to him in connection with his experience.on autogamy (8); the latter studied the occurrence of cleistogamy especially in New Guinea (7) and found it in several genera, and both in the lowland and in the mountains. Some- times in several specimens all the flowers are cleis- togamous, e.g. in Eria rugosa LINDL. and Dendro- bium gemellum LINDL. SMiTH even found species which are only known in the cleistogamous state (8, p. 138), or of which normal flowers have only occasionally been found. SMITH suggests that cleistogamy is more common in the rainy season, and he mentions that R. SCHLECHTER also got the impression that cleisto- gamy was common in very wet places in Sumatra and in the mossy forests of New Guinea more frequent in the rainy season than in the dry period. The same phenomenon has been observed by C. A. BACKER (9) for cleistogamy in Dicliptera canescens Nees (Acanth.) in Java; in moist countries or dur- ing wet periods in the dry season this plant pro- duces minute white cleistogamous flowers the corollas of which drop in the early morning. It is certainly noteworthy that a single trivial character like cleistogamy can so upset taxono- mical judgment that a new genus has been based on this abnormal state of a plant; this character changes the whole floral development, and sup- presses the manifestation of numerous genom tendencies in the mature plant. Physiologically this can only be explained by some break in the physiological chain reactions in an early stage of the development of the flower. The field observa- tions mentioned above may show how this problem may be studied experimentally. Cited literature: (1) Ber. Deutsch. Bot. Ges. 25 (1907) 165-176. (2) Ann. Jard. Bot. Btzg 44 (1935) 239-242; Bull. Jard. Bot. Btzg III, 4 (1922) 241. (3) Wanderings in the great forests of Borneo (1904) 402. (4) De Trop. Natuur4 (1915) 142; see also BACKER, Handb. Flora Java pt 3 (1924) 25. (5) De Trop. Natuur 2 (1913) 53-58, 68. (6) Journ. Siam Soc. Nat. Hist. Suppl. 10 (1935) 66-67. (7) Die Orchid. Deutsch Neu Guinea, FEDDE, Re- pert. Beih. 1 (1914) p. Li. (8) Natuurk. Tijdschr. Ned. Ind. 88 (1928) 122-140, Orch. Rev. 37 (1929) 75, Nova Guinea 14 (1929) 359. (9) Onkruidfiora Jay. Suiker (1931) 676, in nota. Teratologo-morphosis 7. Teratological forms Malaysia can boast of a series of good articles by J. J. SmitH & J. C. Costerus (1) dealing with teratological phenomena in plants. Though several of these teratological forms are due to some hereditary factor, others are apparently caused by external factors. Some are possibly the result of a fungus’s attack though no fungus has been found. Pometia pinnata Forst. almost always has pe- culair large brown structures like witches’ brooms by which the tree can easily be recognized in the riverine forest: they suggest inflorescences. Invirescentia are quite a common phenomenon in several Compositae (fig. 10); the fact that they are often found together in colonies in several different species suggests that they may be due to some virus (?). Monstrous flowers occur rather frequently in Orchidaceae and have often confused systematists. XXVI FLORA MALESIANA [ser. I, vol. 41 J. J. SmiruH (2) has given an interesting account of them. The absence of a rostellum is closely con- nected with autogamy. As a result the flowers often hardly open, do not develop well, and their colour is paler than normal e.g. in Phajus tankervilliae BL. a={f ESS } LA Fig. 10. Invirescence of Emilia sonchifolia DC. (Comp.) from Mt Abang, Bali, sandy riverbed, ca 1000 m, xX 3/2. Sumatran specimens are apparently more normal than Javan. Of quite a number of these abnormal Orchidaceae no normal specimen is as yet known. Another abnormality is a variation in the num- ber of anthers, which, in Dilochia pentandra RcusB. f., is five; this ‘species’ is, however, a mere form of D. wallichii LINDL. In other cases the third stigma- tic lobe is changed into a rostellum and the ros- tellum has become a stigmatic lobe. J. J. SmirH remarks that the phenomenon of peloria occurs in different degrees. Mostly the peculiarities of the labellum disappear, sometimes the tepals show some characteristics of the labellum: As peloria is for the most part inherited these forms are treated in more detail in the following chapter, paragraph 2. Teratological aberrations frequently cause such large changes in the structure of flowers that they strongly suggest some taxonomic novelty. An ad- ditional example is: an interesting 3-seeded coco- nut (3). DE Wit & PostHuMus collected at Buiten- zorg, Sept. 1944, a specimen of Cassia mimosoides L. of which each flower possessed 2 ovaries. This character is considered to be primitive or ancient in the Leguminosae; it has been reported to occur in several Caesalpiniaceae, e.g. in African Schwart- zia (6) and Indian Caesalpinia (7). In Archidendron, a genus of Mimosaceae centred in New Guinea, it is a generic character. Monstrous forms occur frequently in ferns where the plasticity seems greater than in Spermatophyta. Forked, lobed, and crisped leaves occur in many genera. Sometimes these monstrosities seem to be inherited and of racial rank (4). Even precocious spore formation may be partly inherited. Teratological aberrations merge gradually into individual variations. It is questionable whether an individual of Cassia mimosoides L. with two ovaries is to be classed as a teratological or individual variation. I will mention only a few examples of individual variation. MELCHIOR found (5) some flowers in Aphania masakapu MeELCH. with free anthers. BACKER found individuals of Alysicarpus rugosus DC. with 2-3-foliolate leaves. The leaves of Cis- sampelos pareira L. are sometimes both peltate and non-peltate in one plant. Some specimens of Ama- ranthus spinosus L. are unarmed. There is no end to this kind of individual vari- ation which sometimes affects typically structural characters. Experiments are needed to ascertain whether these aberrant plants are sports of the genom and hereditary or not. Cited literature: (1) Ann. Jard. Bot. Btzg vols 13, 19; 23; 24,28, 29, 32) 33, 39) 42.1895 =S1Dsie (2) Natuurk. Tijdschr. Ned. Ind. 88 (1928) 122-140. (3) Natuurwet. Tijdschr. Ned. Ind. 101 (1941) 144. (4) O. PostHumMus, De Trop. Natuur 25 (1936) 177-178. (5) Notizbl. Berl.-Dahl. 10 (1928) 277. (6) JACQUES-FELIX, Bull. Soc. Bot. Fr. 92 (1945) 158. (7) WicutT & Arnott, Prod. Fl. Pen. Ind. Or. (1834) 281. Hypselo-morphosis 8. Phenotypic effect of altitude G. BONNIER, and later F. E. CLEMENTS, experi- mented on the effect of altitude on plants. BONNIER even assumed that species might change under pro- longed exposure to different conditions into other species but it seems that his experiments are un- trustworthy (1). In the Malaysian mountains where collectors are often compelled to follow ridges, plants from exposed situations are frequently brought home. Their foliage is often reduced, the leaves roundish, margins recurved, texture coriaceous, venation prominent, petioles reduced, habit compact. It is Dec. 1948] not always certain that these characters are a ‘nor- mal feature’ of the species. It is, therefore, of the greatest importance to try to collect such species from less exposed habitats (light, wind, poor soil), i.e. from the more fertile, sheltered, though less Fig. 12. Histiopteris alte-alpinav. A.v.R.(Polypod.), an altitudinal form of H. incisa J.SM., in its habitat between ‘sterile’ rocks on the summit of Mt Kerintji, W.Sumatra, ca 3750 m alt. (FREY WIJSSLING) accessible slopes. Extensive notes and large col- lections may show that such variability exists and serve to define the position of transitional speci- mens. The same species may be a crooked gnarled shrub when growing on a ridge and a moderately tall tree 50 m lower on the slope. The dwarfing of trees towards the summits of mountains and ridges is chiefly due to the gradual disappearance of the bole with increasing altitude. This is partly a consequence of the development of the young plants under a gradually increasing light intensity which stimulates branching close to the base. I observed a striking example in the field of dwarfing in Casuarina junghuhniana Mia. on Mt Soeket, Idjen volcano, E. Java. Herbs too are generally dwarfed at high altitude, e.g. Erigeron linifolius WILLD. I studied an instructive case of variation induced by altitude combined with poor rocky soil in the grass Isachne pangerangensis Z.M. (fig. 11). A large series of transitions from tall to dwarfed speci- mens were represented. An example of a ‘hypselo-morphosis’ which has been described as a local-endemic species is that of the fern Histiopteris alte-alpina v. A. v. R. (fig. 12) from the summit of Mt Kerintji, West-Central Sumatra, which is found at ca 3700m alt., on a barren rocky ridge. This is certainly only a form of the common volcanophile H. incisa J. SM. It is difficult, however, to single out the various factors associated with increasing altitude viz more wind, sudden and large changes of temperature, strong insolation, poorer soils, lower atmospheric General considerations XXVII | | | | Fig. 11. Variable habit of Jsachne pangerangensis Z.M. (Gram.) in N. Sumatra, Mt Losir, a. on burnt ridge in thick humus, 1500 m, c. on ridge with ericoid scrub, half-shade, thinner soil, 2000 m, d-e. open sandy flats on poor soil, 3000 m, f. on-rocky windswept summit, soil nearly absent, 3440 m, x 1/4. XXVIII FLORA MALESIANA [ser. I, vol. 4! pressure, different fluctuations of atmospheric hu- midity, greater difference between day and night temperatures, &c. In the absence of experiments one can only make some suggestion, in many cases based on observation in the field only. I know of only few species which are hairier in the mountains than in the lowland, e.g. Hydrocotyle sibthorpioides LAMK, of which BLUME described the hairy form as H. hirsuta Bu. nonal. However, glabrous forms of this species also occur on the mountains! An other example is that of Dodonaea viscosa (L.) JACQ. Kurz (2) in his ‘Sketch of the Vegetation of the Nicobar Islands’ has remarked on the apparent absence of any general relation between hairiness and environment. There is no general rule that flowers are brighter coloured in the mountains. Ageratum houstonia- num MILL. has larger capitules and brighter blue flowers in the mountains than at low altitude buton Mt Pakiwang, S. Sumatra, Ifound the reverse (3), Scutellaria javanica JUNGH. var. sumatrana BACKER having here blue flowers at the base of the peak but white ones towards the summit. Of Dendrobium jacobsonii J.J. 8S. (§ Pedilonum) from the Casuarina forests 2400-2900 m alt. in East Java, J. J. SMITH says that at Bandoeng at 700 m alt. cultivated specimens had smaller and paler coloured flowers with a slightly different flower shape: mentum not bent and differences in the labellum; the inflorescences were, moreover, sometimes 2-flowered (4). Fruiting and flowering are also strongly influ- enced by altitude, as I demonstrated elsewhere (5). Experiments on the influence of altitude, the morphological and physiological behaviour of Malaysian plants have been scarcely made. TEys- MANN made some observations in his pioneer work on Mt Gedeh in West Java but did not comment; COSTER (6) wrote a note on the beech specimen planted by the former. In the Malay Peninsula RmLey (7) made some notes on the acclimatization of plants and the ways in which they can be accomodated at low altitude. Cited literature: (1) The New Systematics 1940, p. 55 seg. (2) Journ. Asiat. Soc. Beng. new ser. pt II, 45 (1876) 126. (3) Bull. Jard. Bot. Btzg III, 13 (1933) 16. (4) Bull. Jard. Bot. Btzg II, no 26 (1918) 41. (5) Bull. Jard. Bot. Btzg III, 13 (1935) 331-343. (6) Ann. Jard. Bot. Btzg 35 (1926) 105. (7) Agric. Bull. Str. & Fed. Mal. St. volumes 6-7 (1907-08). Photo-morphosis 9. Epiphytes It is sometimes wrongly assumed that epiphytism is confined to specific plants which are restricted to this mode of life. The amount of light appears to be the main factor. On the floor of closed forest the shade prevents epiphytes from making use of patches of bare soil, which in primary forest are always present. Exposed places, such as rocks, lava streams, landslides, poor silicious soils, mud streams and solfatara, however, offer conditions suitable for their growth, and are indeed often the places where many epiphytes are assembled, i.e. selected from the neighbouring forest. Though epiphytes may withstand dry conditions well, they mostly need a rather high atmospheric humidity which, in these exposed places, becomes a limiting factor. Most astonishingly rich communities of epiphytes I found on the often misty slopes of Mt Telong in N.Sumatra which from 1800 m upwards is like a rock garden carpeted with normally epi- phytic orchids amidst luxuriant dripping cushions of hepatics and mosses with some isolated dwarf Rhododendrons. It is sometimes contended that these terrestrial epiphytes are epilithes but I have CC * es 74/ \X Fig. 13. Vaccinium laurifolium Mia. (Eric.) as a hemi-epiphyte, height ca 5 m, along a road above Trétés, 1500 m, N.slope of Mt Ardjoeno, E. Jaya. Dec. 1948] also found them in deep humic soil between the rocks. I did not succeed in detecting any essential differences in habit between terrestrial and epi- phytic specimens. In other species, though, the terrestrial speci- @ Gy es ee Sess Fig. 14. Habitat variations of Gentiana quadrifaria BL. (Gent.) in Java. The condensed pin-cushion shape is found on open dry windswept habitats, the loose habit on marshy or slightly shady soil, x 1/2. mens may differ considerably in habit from epi- phytic; they become more rigid and condensed, often fastigiate. Vaccinium lucidum (BL.) MIQ., as an epiphyte is a loosely and irregularly branched shrublet with a tuberous woody base. Terrestrial specimens on ridges are mostly cupressus-shaped miniature trees without the woody tuberous base. Similar differences are found in Ficus deltoidea JACK of which the epiphytic and terrestrial speci- mens may differ considerably in habit. It goes without saying that a proposal by NAKAI (2) to distinguish the Ericaceous Agapetes and Vac- cinium by a terrestrial habit in the latter and an epiphytic habit in the former did not meet with the approval of SLEUMER. Some species begin their life as epiphytes but, when their roots subsequently reach the soil, they may grow into trees and sometimes show no sign of their early history. Such is found e.g. in Ficus, Fagraea, Schefflera, Wightia (1), and I even found it once in Vaccinium laurifolium Mig. (fig. 13). Many, however, are equally able to germinate ter- restrially and grow normally to trees. This is, in Wightia, even more common than the hemi-epi- phytic habit. As a small tree it is gregarious on the sunbaked lava streams of Mt Idjen in East Java, but on the forested outer slopes of the same moun- tain it is a hemi-epiphyte. Cited literature: (1) Revision of Wightia, Bull. Jard. Bot. Btzg III, 18 (1948) in the press. (2) Japan. Journ. Bot. 12 (1936) 37-38. 10. Shade forms Shade forms are found both in the lowland and the mountains. In general they possess larger, thin- ner leaves, longer internodes, &c. Shade and nor- mal leaves may occur in one individual. A very good example is Gentiana laxicaulis Z.M. de- scribed from Java, which appears to be a shade form of G. quadrifaria BL. Sometimes compact tussocks of the latter bear on one side shoots of ‘lJaxicaulis’ in one individual plant (fig. 14). Slender modifications of herbs can be observed in tall grass fields, comparable with those in temper- ate corn fields. These weeds growing in the damp dark micro-climate between the closely set culms of Saccharum spontaneum L., Andropogon amboi- nicus (L.) MERR., efc. strive for light. They show reduced leaves and inflorescences in relation to their lank habit. All herbs unable to emerge from the tops of the grasses show a similar habit, a kind of etiolated growth combined with some degree of nanism. For the effect of light on the habit of forest trees see the paragraph on savannah trees. Hygro-morphosis 11. Influence of drought Hardly anything is known of the influence of drought, and the changes induced by it in the mor- phology and physiology of Malaysian plants. In Gerbera jamesonii Bo.us I observed in the dry year 1945 at Buitenzorg an astonishing reduction in length of the peduncles in relation to leaf length. The size of the leaves was very much reduced dur- ing the same period in Turnera subulata Sm. (T. trioniflora AIT.). Similar behaviour is mentioned by BACKER (1) in Jatropha gossypifolia L. var. elegans M.A., a plant which is thoroughly naturalized in the dry regions of Java and the Lesser Sunda Islands; dur- ing the driest period of the dry season only minute, short-petiolate dark-brown leaves are produced. Flowering of some trees, e.g. Dipterocarpaceae, and probably bamboos coincides with unusually dry years. Higher fungi fructify after a dry spell. Cited literature: (1) Onkruidflora Jay. Suiker. (1930) 411. Hora-morphosis 12. Seasonal variation Seasonal variation as described in Europe (1) I have not found recorded from Malaysia. In the cultivated Hibiscus sabdariffa L. I have seen fruiting specimens flowering a second time; these flowers, however, were only half the normal size and, also, paler.in colour. Field botanists should search for ‘autumn forms’ in periodically dry regions. XXX FLORA MALESIANA [ser. I, vol. 4! Seeds of seasonal plants germinating in the wrong season may sometimes grow into dwarfs. I ob- served such forms also in Hibiscus sabdariffa L. at Buitenzorg. These dwarfs were 10-15 cm high and Fig. 15. Oblique, wind-trimmed Tamarindus indica L. (Leg.) on a ridge at ca 600 m on Noesa Penida, SE of Bali Island (DE VooGp) had 2-3 flowers producing good seeds; the flowers were mostly much smaller than those of specimens flowering in the optimal season. Of leaf-shedding trees flowers are often collected with immature foliage which may deviate consider- ably from mature leaves. A peculiar case is that j Dichrocephata Hyptis brevipes Ne Fig. 16. Compact dwarfing of plants near the fumaroles on the summit of Mt Kembar, Ardjoeno, E. Java, 3100 m alt. Normal specimens left, dwarfed ones right, x !/3. chrysonthemifolia’ « of some leguminous trees which produce leaves in flushes; the latter consist of pale or white or even pink-coloured limply hanging leaves which only slowly get their normal texture (Maniltoa). The distinction between annuals, biennials and perennials causes many difficulties in species growing both inside and outside the tropics, spe- cially when the duration of life is used as a charac- ter to establish taxonomic limits. I assume e.g. Centrolepis to be annual in N. Sumatra, though its perenniality in S. temperate regions is used as a distinctive generic character against allied genera. In some Gramineae species may be similarly vari- able, specially in tropical localities, and thus deviate from temperate representatives of the same species in a character which is, in grasses, generally assumed to be of importance for the delimitation of species if it runs parallel with other morpholo- gical differential characters. It is puzzling me how it is possible to interprete from herbarium speci- mens the duration of life of perennials flowering during their first year and collected in that state. -The use of the duration of life as a character in keying out species must be limited to very clear cases based on wide experience. Cited literature: (1) R. Vv. WETTSTEIN, Unt. ii. d. Saison-Dimorphismus im Pflanzenreiche. Wien 1900. 42 pp. Anemo-morphosis 13. Windforms A peculiar aberrant habit in shrubs and trees can be caused by constant winds. I have described this from Noesa Penida and Bali (1) in Terminalia catappa L., Barringtonia asiatica (L.) KURZ, Calo- phyllum inophyllum L., Bischofia javanica BL., Ficus sp., and Tamarindus indica L. (fig. 15). Other more recent examples are Dodonaea viscosa JACQ. near the Wijnkoops Bay, S. Java, and plants from Pa- dang Bolak in N. Sumatra described by M. VAN DER VoortT (2). These plants possess sometimes a peculiar oblique condensed one-sided habit and always show a decreased leaf size apparently owing to desiccation of the buds. They are found both on seashores and inland. Cited literature: (1) De Trop. Natuur 26 (1937) 69-78, 14 fig. (2) De Trop. Natuur 28 (1939) 201-209. Edapho-morphosis 14. Fumarole plants I have described (1) very aberrant modifications from some mountain summits viz Mt Ardjoeno in East Java(2) and Mt Agoeng in Bali(3) at 2900-3000 m alt. Some common lowland weeds, have through chance dispersal by wandering pilgrims and/or by deer established themselves in the immediate neighbourhood of fumaroles. Owing to the heat and moisture emitted by the fumaroles they are able to grow at these high altitudes. They are very much reduced in size and in habit very condensed, and their leaves are very small (fig. 16). Without Dec. 1948] flowers their identification would be difficult. They live in what may be called ‘open air hothouses’ in the subalpine zone, and the altitude, insolation, &c. are doubtless the factors which have induced their WA Fig. 17. Dwarf of Pemphis acidula Forst. (Lythr.) in flower and fruit, seashore of Oedjoeng Koelon, W. Java, xX 2/s. aberrant mode of growth and resulted in what seems to be an ‘alpine habit’. The species concerned were: Hyptis brevipes Porr., Dichrocephala chrysanthemifolia (BL.) DC., Lycopodium cernuum L., Emilia sonchifolia DC., Bidens pilosus L., Oldenlandia herbacea Roxe., Fimbristylis capillaris A. GRAY, Lindernia crustacea F. v. M. Fig. 18. Full-grown specimen of Pemphis acidula Forst., NE. coast of P. Tioman, Mal. Peninsula. (CORNER) General considerations XXXI Cited literature: (1) The Gard. Bull. Str. Settlem. 9 (1935) 63-69. (2) De Trop. Natuur 23 (1934) 119-120. (3) De Trop. Natuur 25 (1936) 158-159. 15. Rock plants; calcareous and silicious soils Both rocks and silicious soils may bring about rather conspicuous changes of habit in some plants, apparently owing to the small amount of nutrients available. These modifications can occur either at low or high altitude. VN =S FS Fig. 19. Flowering and fruiting dwarf of Lepto- spermum flavescens J.SM. (Myrt.) on dry sterile sands of Toba highlands, Central Sumatra, in a heath-like vegetation, * !/2. Mr C. N. A. DE Voocp collected dwarf speci- mens of Pemphis acidula Forst. (fig. 17) on the rocky coast of SW. Java resembling subalpine ‘Spa- lierstraucher’; normally this littoral species is a bush or small tree (fig. 18). On the so-called ‘padangs’, the gravelly or sandy flats of various geological history which sometimes occupy large areas in Sumatra and Borneo, many species are dwarfed: Leptospermum flavescens SM. when growing under optimal conditions is a me- dium sized cedar-like tree (fig. 20); here it is a dwarf, 10-20 cm high, which flowers and fruits abundantly (fig. 19). Many other species behave similarly. If herbarium specimens are not provided with good field notes, a botanist who has never visited the tropics is of course confronted with a puzzle. He may even find some other slight charac- XXXII FLORA MALESIANA [ser. I, vol. 4! ter not known to occur in the normal population and may think that they represent a different species: in this way another ‘paper species’ is created. Residents in the tropics ought to experi- ment with seeds gathered from dwarf individuals. Abandoned mining grounds in Banka, Billiton and Borneo are rich in dwarf forms of the most diverse species, which flower precociously as very small individuals (1). flowers, an unusual character in the species (2). Though no experiments have been made it is likely to be an edaphical form only. Scores of dwarfed species, mostly of shrubs or small trees but also of herbs (e.g. Dianella nemorosa LAMK. f. nana SCHLITTL. from Camarines and f. monophylla SCHLITTL. from New Guinea) occur in the Philippine Islands, and especially in New Guinea on ridges in the mossy forest and the sub- Fig. 20. Full-grown specimens of Leptospermum flavescens J.SM. (Myrt.) on the slopes of Mt Bonthain, SW. Celebes. (L. VAN DER PIJL) On poor unweathered volcanic ash on the slopes of mountains the vegetation as a whole is dwarfed, e.g. on the slopes of the easily accessible Mt La- mongan, E. Java. Here the black gravel and sand is continually rejuvenated and gradually runs down. On these ash slopes all the common Javan mid- mountain trees and shrubs are dwarfed but flower and fruit profusely e.g. Radermachera gigantea (BL.) MiQ., Parasponia parviflora MiQ., and Wein- mannia blumei (BL.) PLANCH., &c. flower and fruit on 1-2 m high shrubs. This observation induces me to suspect that Radermachera brachybotrys Me_rr. from Leyte merely represents a dwarf speci- men of some other species; KORTHALS found a similar specimen in the padangs of Borneo. I have also found dwarfs on wooded limestone cliffs in NW. Bali at 100-200 m mostly of her- baceous species. One of them was so aberrant that I described it as a new variety, Anisomeles indica (L.) O.K. var. biflora STEEN.; this had solitary alpine zone. No experiments have been done and the ‘normal’ habit of these plants is thus unknown. On rock cones, e.g. Mt Idjen, E. Java (fig. 21) and Mt Agoeng, Bali, I found Casuarina (fig. 22), Vaccinium, Rhododendronas extremely small shrubs and ascribed this to the very poor soil, though on these cones the influence of climate and soil are not readily separable. W. TROLL found precocious spore formation in Gleichenia vulcanica BL. on Mt Gedeh. This was certainly not caused by altitude but by the locally poor rocky soil. In cracks of rock on the summit Argapoera, of Mt Jang, E.Java, I have collected microphyllous specimens of a Polygonum which I originally took for P. chinense L. but which DANSER afterwards identified as an aberrant form of P. runcinatum Down (fig 23). On Mt Kerintji were found minute fruiting specimens of Aralia ferox BL. which I have distinguished as f. nana (3). Dec. 1948] Cited literature: (1) TEYSMANN, Nat. Tijdschr. Ned. Indié 32 (1873) 84; DUNSELMAN, De Trop. Natuur 27 (1938) 97-104. (2) Bull. Jard. Bot. Btzg III, 17 (1948) 389. (3) Bull. Jard. Bot. Btzg III, 17 (1948) 394. 16. Solfatara plants Specimens collected in craters are often of a surpris- ingly dwarfed habit even when old. SCHROTER (1) figured a dwarf plant of Vaccinium varingifolium Mia. of East Java which was probably 50 years old and had the appearance of some alpine ‘Spalier- strauch’. At a short distance from these strongly insolated, edaphically dry and often wind-swept barren rocky places on slopes or summits, the same species occurs in hollows or other sheltered places as well-developed shrubs or small trees. The dwarf shrubs of craters are often wholly appressed to the soil (with rooting branches!), with a matted and prostrate habit. Owing to the poisonous gases emitted by the solfatara or effect of the wind on ridges, their surfaces are flat and look as if clipped Fig. 21. Vaccinium varingifolium Mia. (Eric.) as poor prostrate shrubs (‘Spalierstraucher’) near Kawah Idjen, E. Java, ca 2000 m alt. Exceedingly poor, eroded, very young volcanic soil. This species also grows in the mountain forest on the ridge behind in ca 3-6 m tall trees. General considerations XXXII or pruned (2) (fig. 21, 24). The solfatara may shift its outlet and so release these plants from its influence: I found some partly grown into a fresh bush, proving that the plant had recently escaped from the reach of the gases, the prostrate section being the oldest part. The reverse may also occur; erect shrubs may be affected later by crater gases (3) which makes them one-sided (fig. 25). b oA A GEA Ee” (LLL A Fig. 22. Casuarina junghuhniana MiqQ. (Casuar.). Old dwarf from the summit of Mt Agoeng, Bali Island, 3100 m, on a rocky windswept cone, x 1/2. Solfatara plants thus represent forms different in habit, and herbarium botanists must handle these materials cautiously. In a general sense the ‘poor soil’-conditions cause nanism. In Malaysia adaptability to habitat and varia- bility in habit under extreme conditions is far greater than the average herbarium botanist sus- pects. It is difficult to interpret aberrant specimens from remote regions without a thorough field knowledge. Unfortunately this has led to the des- cription of many ‘paper species’ which may seem specifically distinct but, when studied under various natural environments appear gradually to merge in the range of modifications existing in many Linnean populations. XXXIV FLORA MALESIANA [ser. I, vol. 4! Cited literature: (1) Vierteljahrschr. Naturf. Ges. Ziirich 73 (1928) 584. (2) De Trop. Natuur 24 (1935) 142-144, fig. 2-5. (3) SO2, H2S, Cl2, &c. Hydro-morphosis 17. Water- and swamp plants Phenotypic variations comparable to those known in Europe are also known in the Malaysian flora. Jussieua repens L., when growing on muddy soil Fig. 23. Polygonum runcinatum Don. (Polygon.). Below: apex of a normal plant. Above: a very uncommon form of Mt Argapoera (Jang massif, E. Java) from clefts in rocks (St. 10960), x 2/s. | hh 4 x AYA NK\Y| y IW ‘ bi i x sNRianinte SS AV AINA AD ii = NUN aie! uD A, = Ss through the lowering of the water level, changes into a conspicuously different land form with minute hairy leaves and very condensed habit; it takes some time to recognize this land form (1). Land forms are also known in Potamogeton, e.g. from Sumatra, and in Utricularia. Of Neptunia plena Bru. a land form is known. BACKER suggests that the endemic WN. javanica Miq., a terrestrial endemic species in Java, is merely a land form of the common N. oleracea Lour. (2). Limnophila sessiliflora Bui. and L. indica (L.) Druce, in shallow water, have deeply divided leaves below the surface with gradual transitions to lobed and toothed upper leaves above the water level. In very deep water pinnatifid leaves predomi- nate, but in marshy grass fields only pinnatisect to dentate leaves are present. The amount of aerenchyma is closely related to the depth of the water. Many swamp plants fail to flower in deep water, but come rapidly into bloom when the water recedes (Lemna, Blyxa, Pistia, Azolla, Salvinia, Marsilea, &c.). Swamp forest trees are also affected by the amount of water in the soil. A conical base to the trunk, so well known in Taxodium, is found fre- quently in other swamp species e.g. in Gluta renghas L., Alstonia spathulata BuL., &c. but, in non-inun- dated soils, the swollen base of the trunk is not or scarcely developed. The same is true of aerial roots at the base of the trunk. In deep swamps they may resemble the stilt roots of mangrove. Such roots may not de- velop in the same species when it is growing on dry land e.g. Acmena (Eugenia) operculata (ROXB.) MerrR. & PERRY. Root production in these cases is doubtless a direct response to the habitat. Cited literature: (1) De Trop. Natuur 2 (1913) 83, fig. 3. (2) Schoolflora voor Java (1911) 428. Fig. 24. Sketches of prostrate clipped habit of shrubs in the crater of Mt Papandajan, W. Java, ca 2000 m, through the combined action of wind and sulphurous vapours from solfatara. Above: Vaccinium varingifolium MiQ., below: Rhododen- dron retusum (BL.) BENN. (Eric.) (drawn after photographs). Fig. 25. Oblique growth of Vaccinium varingifolium Mia. caused by sulphurous gases of the crater of Mt Papandajan, W. Java, ca 2000 m alt., ca 1 m high. Dec. 1948] Phyto-morphosis 18. Fungus and bacterial diseases, and symbiosis Malformations caused by fungi have led to some errors in identifying Malaysian plants. Loranthus maculatus BL. is, according to DANsgrR, the com- mon Dendrophthoe pentandra Mia. with a fungus on the leaves causing black spots (1). In specimens of Cassytha filiformis L. from New Guinea Dr Hatusima found some tetramerous Fig. 26. Pseudo-flowering of bamboo; galls caused by Epichloe treubii (Fungi). Bot. Gardens, Buitenzorg, Java. flowers with a central column marked by little pits. The slender inflorescence was glabrous and the rest of the plant hairy. It was evidently a maifor- mation caused by a Peziza-like Ascomycete. Root deformities caused by Cyanophyceae are found in Cycas, Gunnera, &c. Structures like witches’ broom are often found in bamboos, and often regarded as immature flow- ering parts. These pseudo-flowers are galls caused by a fungus (fig. 26). A curious malformation in Pilea trinervia WIGHT consisting of conspicuous swellings of the inter- nodes was described by Mrs WEBER VAN Bosse (2) and is caused by a parasitic alga: Phytophysa treubii W. v. B. In Pavetta, bacteria cause dark often thickened spots in the leaves. According to BREMEKAMP the symbiosis is mostly restricted to particular species. Similar bacteria are found in species of Psychotria, General considerations XXXV Ardisia, the tips of the leaves of Smilax, &c. The presence of absence of bacterial nodules is used in the identification of Rubiaceae, a rather singular method. Cited literature: (1) Compare BoEDIJN, Bull. Jard. Bot. Btzg III, 13 (1935) 497-SOl1, fig. 1. (2) Ann. Jard. Bot. Btzg 8 (1890) 165-186. Zoo-morphosis 19. Ant plants (myrmeco-morphosis ) Several Malaysian plants are inhabited by ants. TREUB (1) proved that the cavities in which the ants live in the tubers of Myrmecodia and Hydno- phytum are also formed in the absence of ants. In other instances, however, ants presumably bite their way into internodes and remove and carry away the pith. This was shown to occur in Endospermum moluccanum T. & B. (E. formicarum Becc.) by DocTEeRS VAN LEEUWEN (2); I am able to confirm this. DOCTERS VAN LEEUWEN also found some specimens uninhabited by ants. In Endosper- mum, therefore, whether the internodes are hollow or not is certainly not a good specific distinction, hough used by PAx in his key to the species of Fig. 27. Above: Kibessia sessilis BL. (Melast.) being based on a galled swollen fruit of K. azurea BL. (W. Java). Below: peculiar galls of Styrax benzoin DRYAND. (Styrac.) from Sumatra, x 2/3. XXXVI FLORA MALESIANA [ser. I, vol. 4! Endospermum subg. Capellenia (3). Moreover, as the other character used by PAX, viz the number of cocci in the fruit, varies from 3—5, through igno- rance of data on points the collector ought to have noted on the label, the whole key breaks down. In Wightia borneensis Hook. f. some individuals are attacked by ants which remove the pith from the upper internodes. The withdrawal of the inner tissue causes the hollow internodes to assume a cigar-like shape. Cited literature: (1) Ann. Jard. Bot. Btzg 3 (1883) 129-153. (2) Treubia 10 (1929) 1-7. (3) Pfl. Reich Heft 52 (1912) 34. 20. Galls deceptive to phytographers (cecidio-morphosis ) Cecidia caused by animals have sometimes de- ceived botanists when describing plants. An ex- ample is Ceratostachys arborea BL., a genus based on a galled fruit of Nyssa javanica (BL.) WANG. Kibessia sessilis BL. is merely the galled and enlarged fruit of K. azurea BL. (fig. 27). According to RmpLey (1) Apteron lanceolatum Kurz, described as a distinct genus, is identical with Ventilago kurzii RIDL.; KURZ mistook some insect galls for the ripe fruit. MiQuEL described (2) an abnormal tree from Sumatra which was actually a species of Styrax, a genus in which most peculiar galls (fig. 27) are very common. Otopetalum micranthum Mia. is an Apocynacea described from Java. According to BOERLAGE (3) the plant was referred to the wrong tribe because MIQUEL erroneously took galled flowers for 1- seeded berries; the former author suspects that it is related to Micrechites. Insects (mostly cicads and larvae of Hemiptera) cause a singular malformation of the flowers in some species of the genus Sterculia. H. C. CAM- MERLOHER (5) Observed that they are attracted to so-called ‘sugar hairs’ which occur on the inside of the perianth. The insects injure both the hairs and the outer tissue in an early stage of develop- ment of the flower. The calyx becomes enlarged, thicker and tough, and opens hardly in anthesis; its lobes remain short and triangular, and the tube is relatively large. These flowers are conspicuously different from the normal ‘uninhabited’ flowers and, according to ADELBERT (6), ought not to be used when describing or identifying plants. BACKER (7) described in Hibiscus schizopetalus (Mast.) Hook. f. malformations of the vegetative parts and of the flowers caused by plant lice. The Philippine species Euphoria malaanonan was described by BLANCO and by him referred to Sapin- daceae but MERRILL stated (8) that it is merely based on specimens of the echinate galls of Shorea guiso BL. of the Dipterocarpaceae. W. M. DocTERS VAN LEEUWEN has published (4) an illustrated book on zoocecidia of Indonesia. Cited literature: (1) Flora of the Malay Peninsula 5 (1925) 300. (2) Linnaea 26 (1853) 285. (3) Hand- leiding Fl. Ned. Ind. 2? (1899) 380. (4) The Zooce- cidia of the Netherlands Indies, Batavia 1926; Supplement, Ned. Kruidk. Arch. 51 (1941) 122- 251. (5) De Trop. Natuur 22 (1923) 147. (6) In BAcKER, Flora van Java, Nooduitg. [VB (1944) fam. 107, p. 18. (7) Flora van Java, Nooduitg. [Vc (1943) fam. 109, p. 27. (8) Spec. Blanc. (1918) 33. 21. Influence of browsing animals (pascuo-morphosis) In some parts of Malaysia browsing cattle and deer (1) can induce changes in the morphology of plants which might be termed pascuo-morphosis. In Fig. 28. Casuarina junghuhniana MiqQ. (Casuar.) on Mt Jang, E. Java; crown trimmed below by deer. the deerpark of the Buitenzorg Palace, deer regu- larly feed on the pendent air roots of Ficus and prevent them from reaching the soil. The trees therefore remain single-stemmed and do not form thickets of pillar-like roots. A similar effect is caused by deer on Mt Jang, in East Java, where deer eat the hanging branches of Casuarina junghuhniana Mia. as high as they can reach (fig. 28). The trees look as if clipped (2) at the underside of the crown like those on the lawns at Buitenzorg. Much the same was observed on Mt Rindjani, Lombok Island (3). The broom- like appearance of the grass Pogonatherum pani- ceum HACK. on Mt Diéng was ascribed to grazing cattle by the late Mr LooGEN (4), an excellent ama- teur field botanist. Browsing of animals has in general the same effect on the vegetation as frequent burning: the plants acquire a low habit and flower at an early age (pseudo-nanism). This occurs very commonly on the closely cropped fields of fine grass of Mt Dec. 1948] General considerations XXXVII Jang. It is only in hedges, on steep slopes or some- where out of reach of deer that plants grow to normal dimensions. In Central and East Java, and the Lesser Sunda Islands pascuo-morphosis is due to browsing of cattle in the dry season; it is known in Zizyphus jujuba L., Streblus asper L. and other shrubs. These assume a fastigiate habit as high as the browsing animals can reach: above about 2 m the twigs are again spreading and form a globular crown. A transition to anthropo-morphosis is the clip- ping and pruning of plants which can sometimes produce an aberrant habit. According to BACKER (5) repeated cutting of plants along roadsides pro- duced a peculiar table-shaped densely branched tile. aye ee FOO Y \ y op (us ZGEy aie Z ERE Ro AEC CSE SES PR, ENS (YA Na esate ne Ting ge as BONA ARO PSST ASS AKAs HARA) YS KHOR EAD EAA AS NIDA Rr ah ZIG IS SO Ce TL V4 EIR a XX postelleags fe : plane eS: EA LA aOR Fig. 29. Seedling of Butea monosperma TAUB. (Leg.) sprouting in its 3rd year in the fire-swept savannahs of Indramajoe, W.Java, and developing a lignotuber, x 1/3. form in Sida retusa L. near Batavia. In the Lesser Sunda Islands cattle are sometimes fed in the dry season with leaves of trees, as is done in Africa. For this purpose the people lop the lower branches of trees in order that cattle may reach the foliage. This causes a tendency to umbrella-shaped trees. Deer may cause the same change of habit. Cited literature: (1) Mentioned for India by Hooker & THOMSON, Flora Indica (1855) 29. (2) De Trop. Natuur 21 (1932) 27. (3) De Trop. Natuur 30 (1941) 123. (4) De Trop. Natuur 30 (1941) 70. (5) Flora van Batavia (1907) 102. Anthropo-morphosis 22. Influence of fire (pyro-morphosis) The changes in habit and structural characters induced by fire were named pyro-morphosis by PERRIER DE LA BATHIE who made observations in the island of Madagascar. In Malaysia there are few reliable data. One of the changes induced by regular burning of the vegetation is that plants are stunted, and flower when small. Owing to the damage done to the surface part of the plant the underground parts thicken, and the upper portion of the rootsystem and lower por- tion of the stem form gradually a thickened half- subterranean ‘lignotuber’, which sprouts after fires have swept the plains. I made some observations in the Indramajoe plains (W. Java), where species of Grewia, Butea, Dillenia, Morinda, Phyllanthus emblica, Zizyphus, &c. sprouted from _ these thickened bases (1) (fig. 29). Some instances of phytographical importance have come to my knowledge. RANT found (2, 3) that Psidium cujavillus BurM. f. can originate spon- Fig. 30. Park-like savannah in SW. Soembawa; trees with short boles and rounded crowns. (DE VooGD) XXXVIII FLORA MALESIANA [ser. I, vol. 4! taneously from root shoots of Ps. guajava L. It is distinguished from Ps. guajava L. mostly by differ- ences of size. Ps. cujavillus BURM. f. must, therefore, be reduced to a sport of Ps. guajava. Fire-resistant trees are often crippled beyond recognition. Plants described from semi-arid (4) regions where fires occur annually are sometimes known only in this crippled state, e.g. Fordia fruti- cosa CRAtB, from N. Siam, described (5) as a shrublet 40 cm tall. The late A. F. G. KERR, a most able and experienced field botanist, stressed in a note made in the field that the plant was growing in an area subject to fire; this may explain such a habit in this otherwise arboreal genus. The normal plant will, in all probability, prove to be a tree. Cited literature: (1) De Trop. Natuur 25 (1936) Jub. nummer, p. 117-118. (2) Ann. Jard. Bot. Btzg 41 (1930) 27-32. (3) Natuurk. Tijdschr. Ned. Ind. 94 (1934) 112. (4) That is: regions which are periodically wet and dry, and show two distinct seasons coinciding with the monsoons. (5) Kew Bull. 1927, 60; Fl. Siam. En. I (1928) 395. 23. Pioneer plants In the preceding paragraphs 9, 14-16, and 22, several examples have already been given of pioneer plants. I am decidedly of the opinion that this term should not be restricted to plants peculiar to land- slides and other bare soils. Trees settling in savan- nahs or devastated areas, epiphytes settling on rocks, &c. are just as well ‘pioneer plants’. They constitute seral vegetation types. In the initial stages of revegetation forest trees may appear as pioneer shrubs, flowering and fruiting early. It is rather baffling to find Schima noronhae REINW. flowering and fruiting as a lax shrub 2m tall, when one is familiar with the gigantic full grown tree in the forest (height sometimes over 50 m, columnar bole over 1 m diam.). BACKER found (1) near Batavia flowering specimens 15 cm tall of Grewia microcos L., usually a tree up to 17 m high. The same phenomenon can occur in Adinandra, various Urticaceae, Leguminosae, Ulmaceae, &c. In New Guinea some endemic species were originally described as tall trees but Mr Brass has recently found them as gregarious pioneer shrubs in differ- ent seral vegetation types. A promising shade plant, A/bizzia sumatrana STEEN., described from the Westcoast of Sumatra, was found to be a forest tree. In the plantations it is a weed tree flowering and fruiting at a very early age. In the forest, its native habitat, flowering is apparently suppressed by the deep shade, the seedlings grow into pole trees, slender and tall with a minute crown and real growth only starts when the crownlet emerges from the canopy. The behaviour and appearance of forest trees when growing in the open cannot be predicted. The adaptive capabilities of most Malaysian forest trees are unknown, though data on these points would be of great importance for practical forestry. See also chapt. 2 on precocious flowering, a phenom- enon frequently observed in pioneer plants. Cited literature: (1) Flora van Batavia (1907) 196. 24. Savannah trees Trees of the savannah generally differ in habit from trees of the closed forest in their short bole and spherical crown (fig. 30). If young forest trees Fig. 31. Altingia excelsa Nor. (Hamam.) at Tjibodas, W. Java, ca 1450 m. Left: forest-grown tree, clear bole ca 25 m. Right: planted on the lawns of the mountain garden, at ca 200 m distance. Dec. 1948] in deforested areas become exposed, or when they are planted as roadside trees, they acquire this shape: a striking example is A/tingia excelsa Nor., a forest giant of the West Java midmountain forest between 600-1600 m. The clear bole is usually a characteristic feature of the tree; it is columnar, up to 14m in diameter, and up to 20-30 m from the ground unbranched (fig. 31). Specimens 50 years old, however, planted on the lawns at Tji- bodas mountain garden, have grown into low spreading trees with hardly any bole at all (fig. 31). Descriptions of the habit of trees and shrubs taken from specimens grown in private and botanic gar- General considerations XXXIX dens, will therefore generally not agree with those taken from specimens in the forest. THORENAAR made similar observations in Javan oaks, and other trees such as Podocarpus imbricata BL., Quercus, etc. The habit of trees grown in open gardens resembles the shape of trees of the savannahs where the rounded crowns on a short bole often charac- terize the physiognomy of the open savannah for- est. Physiologically this tree form is in all proba- bility determined by the high amount of light pres- ent during the juvenile stages of growth. The crowns of mature forest giants exposed after de- forestation also tend to become rounded. VARIATIONS BOUND TO THE GENOTYPE (Genotypic variation) . General remarks ; XXXIX . Peloric flowers . ; ; xii . Other teratological or deviating forms xlii . Distribution of the sexes. xlili . Geographical segregation of Malaysian AkWNe specific populations . xliv (a) Ecological xlvi (b) Regional xlvii (c) Topographical : 4 xlix 6. The problem of speciation . : : li 7. The effect of isolation . lii 8. Centres of speciation in the Malaysian flora . : ; ; ‘ ; j F lii Variation bound to the genotype is intimately related to the species concept and to geographical distribution. The ideas advanced by J. D. HOOKER in his introductory essay to the ‘Flora Indica’ (1855) have gained in ‘philosophical’ importance through the progress of basic research in experi- mental taxonomy in the last decades in Europe and the United States, in particular by E. Baur, F. v. WETTSTEIN, N. H. NILsson, G. TURESSON, A. Muintzinc, O. WinGe, G. D. KARPECHENKO, B. H. DANsER, W. B. TURRILL, J. CLAUSEN, J. P. Lotsy, H. DE Vries, N. I. VAviLov, and many others. This work is summarized in several useful symposia and textbooks such as ‘The New System- atics’ edited by J. HUxLEyY, CAIN’s ‘Foundations of Plant Geography’ and CLAUSEN’s c.s. ‘Experi- mental Studies on the Nature of Species’ (1945). It lies outside the scope of this essay to consider the various view points on these subjects. Some of them I have already touched on in a study of Malaysian mountain plants. ! Now I will try to explain briefly some current views, including my own, and will illustrate them. by examples taken from Malaysian botany. 1. General remarks Scientific names of species rest on the ‘International Rules of Nomenclature’ and depend on the identi- ty of the ‘type specimen’. This need not imply that only studies in which all type specimens have been (1) Bull. Jard. Bot. Btzg III, 13 (1935) 358-391. 9. Centres of generic development in Malaysia . : : lv 10. Local-endemic species and genera ; Ivi 11. Parallel or homologous variation ; Ivii 12. Reticulate affinities. Iviii 13. Vicariism in the Malaysian flora . : lix 14. Adaptation and migration . f Ix 15. Hybrids in the Malaysian flora . P Ixii 16. Polyembryony, parthenogenesis and apogamy . ‘ 2 Ixiv 7) late origin of native aliens d ‘ F Ixv 18. The origin of Malaysian cultigens Ixvi 19. Extinct plant groups in Malaysia. Ixvili examined, full synonymy is given, in which nomen- clature is in accordance with the Rules, and in which new species or other taxa have been de- scribed by careful Latin descriptions, are sound and durable. I know some excellent works in which nomenclature is neglected and synonymy is obso- lete but in which botanical distinction and de- scription are superb, and specific delimination is carefully drawn. Such books give the impression that the author is master of his art. I become more and more convinced that in the past two decades the care for outward appearance has come to take a too predominant share of the attention of some botanists who wrongly assumed that the examination of type specimens is the last word in real taxonomical research. It is sometimes not realized that type specimens often are only deficient, poor and miserably dried single plants chosen at random from billions of specimens growing in Nature in the past, the present and the future, which, together, according to the Linnean principle, compose the specific population. Very often these type specimens by no means represent the ‘average’ or ‘most common type’ of the population. The ‘z-typica’-distinction, therefore, has only nomen- clatural, i.e. administrative but no botanical value. It needs no comment that an up to date nomen- clature is a conditio sine qua non for any taxonomic work, and it will be tried to reach a high standard in this Flora. At the same time, however, is is hoped that the contributors will not be satisfied when writing formally correct revisions, but also carefully consider the status of both genera and XL FLORA MALESIANA [ser. I, vol. 4! species and the structural differences distinguishing them, not merely limiting themselves to the dis- tinction of taxa for reasons of convenience. Inadequacy of material and lack of field knowl- edge are both sources for the provisional dis- tinction of ‘species of convenience’, which mark, in tropical floras, as a rule the initial stage of ex- ploration. Whilst the inadequacy of material is an immo- vable obstacle, and not every revisor will have the privilege to acquire field knowledge, a large fund of experience has been collected on the subject of variation in tropical plants. The following pages contain a discussion, valu- ation, and illustration of variation in Malaysian plants. In general the new systematics, based on modern experimental taxonomy holds that a narrow spe- cies concept is not in accordance with the structure of nature. Modern insight offers no support to so-called ‘splitters’, one of whom declared to me as his ‘prin- ciple’ that he felt obliged to distinguish the smallest - distinguishable entities and to assign to these a binomium. On the average the standard of the specific concept proposed by LINNAEUS in his works and rules cannot be questioned and for binomiums the Linnean canon has priority of conception. I fail to understand how that conscientious splitter who in matters of nomenclature adheres strictly to priority and applies the binomial system of LIN- NAEUS can simultaneously call wide-spread poly- morphic populations ‘collective species’. To do so is intentionally to depreciate the time-honoured and scientifically sound Linnean standards. The difficulties confronting the systematist are manifold; no clue exists to the causes of polymor- phy. Systematists are still far from being able to explain why some species are polymorphic and variable and why others show a narrow amplitude of divergency. This is a fundamental barrier to the methods of ‘weighing and measuring’, a common basis in the natural sciences generally. Polymorphy is apparently not related to speciation, as mono- specific units such as Homo sapiens, Cocos nucifera, &c. are very variable and species of large genera are sometimes not very polymorphic, though it will be observed that in most genera at least one species is widely distributed and rather polymor- phic. Ignorant of the laws underlying his taxonomic distinctions, the systematist should be aware that he deals with unequal entities, though we may try to shape them as consistently as possible. A basic research in this connection is the work of the late E. BAurR, the geneticist, who made un- surpassed long-range efforts in the combined fields of taxonomy, field-work and experiments, to disen- tangle the genus Antirrhinum sect. Antirrhinastrum (1). This section had some dozens of local species described from the West Mediterranean distin- guished by characters which were the despair of taxonomists. BAUR proved that the population of the section falls apart in numerous ‘colonies” or ‘partial populations’ which are isolated and cross mutually (convivia, sensu DANSER). Each colony has its own type, the larger the colony the wider the local diversity (fig. 33c). All these local types can be freely intercrossed with fertile offspring, and there is no doubt that, if not isolated in nature, they would together merge into a still more diverse popu- lation with transitions and intermediates. BAuR, moreover, obtained experimentally many forms not realized in nature (/.c. p. 289), showing that the potential variability (2) or polymorphy (3), i.e. the total number of possible forms (genetic capabilities) is not exhaustively represented. There is a great reserve of possible combinations, and the genus manifests itself to us at present under a limited number of combinations. Hardly any plant species is evenly spread within its area, and many occur in aggregates or colonies. BAurR’s findings are thus of the utmost value for the Malaysian archipelago where isolation is a normal factor in specific populations, in the low- land owing to the insular discontinuous nature of this region and in the mountains still more accen- tuated by the often long distances between the summits. We cannot expect that the whole plant world will be subjected to accurate and thorough experimental research like BAuR’s Antirrhinum- studies, but, judging from his results, it appears that geographical distribution is an important argu- ment when determining the status of taxa. Well- defined allied species possess in general overlap- ping areas of distribution proving the independence of the populations. If, however, several allied species exclude each other geographically one must be on the alert, and check the differential charac- ters again because the specific population may well be differentiated into a number of races, subspecies, or ecotypes. Especially along the frontier—horizontal and altitudinal—of the area, a species population has a different facies from that at its centre. KERNER (4) showed that in Cytisus sect. Tubocystis aberrant forms occurred along the border of the area (fig. 33e). Migrating plant individuals (seeds, spores, fruits, root-stocks, &c.) carry only a part of the potential polymorphy of the genus, and their off- spring will possess a special facies. Hence, along the frontiers, combinations can be expected which are not realized within the centre of the population. According to VAVILOV such pioneer aggregates are found to be recessive homozygotous; this is of great practical importance. It is worthy of note, as was pointed out by E. C. ANpReEws (5) that BENTHAM realized long ago that the geographical station of a waif or colonist im- poses variations upon it almost from the moment of its arrival. ANDREWS adds, that Eucalypts plant- ed in New Zealand, California, etc. present marked differences in general appearance from the same species in Australia. Stimulated apparently by HOOKER (6), HUGO DE Vries (7), the Master of experimental taxono- mic botany, remarked that the initial stages of new species will be found most easily in luxuriant alien vegetations. His classic example, Oenothera, show- ed these ‘mutations’ — which they indeed are if the TAXONOMIC REVISIONS ACERACEAE! (S. Bloembergen, Buitenzorg) 1. ACER LinnE, Sp.Pl. (1753) 1054; Pax, Pfl.R. 8 (1901) 1; K. & V. Bijdr. 9 (1903) 252. Trees or shrubs, buds with many perules. Leaves decussate, petiolate, entire, palmate or pinnate, appearing simultaneously with the flowers or later, exstipulate. Inflorescence racemose, corymbose or spicate, terminal with 2-4 leaves, or rarely terminal or axillary without leaves. Monoecious or dioecious, flowers actinomor- phic, d and 9, ovary in the d fls more reduced than stamens in 9 fis. Calyx and corolla 4—5-merous. Stamens 4-10, mostly 8, hypogynous or perigynous. Disc extra- or intrastaminal. Ovary superior, 2-celled, laterally flattened, each cell with 2 ovules. Fruit a samara, splitting into 2, rarely 3, winged usually 1-seeded parts. Seed without endosperm, radicle elongate, cotyledons foliaceous, or thickened, plicate, involute or flat. Distr. Ca 200 spp. in the N. hemisphere, only in Malaysia crossing the equator. Notes. By BLUME, BENTHAM & HOOKER, MIQUEL, &c. this genus was included in the Sapindaceae. In Malaysia only one species. 1. Acer niveum BL. Rumphia 3 (1847) 193; Pax, Bot. Jahrb. 6 (1885) 293; ibid. 7 (1886) 207, cum var. cassiaefolium; WESMAEL, Bull. Soc. Bot. Belg. 3/4 mm (in Q slightly smaller and not dehiscent). Disc fiat, glabrous to woolly. Ovary 2 mm broad, densely woolly, in od (with the styles) usually 29 (1890) 41, cum var. praec.; SCHWERIN, Gartenfl. 42 (1893) 228, cum var. laurinum & praec.; PAX, in E. & P. 3, 5 (1896) 267, 271; Pfi.R. 8 (1901) 4, 31; K. &. V. Bijdr. 9 (1903) 254; Backer, Schoolfli. (1911) 272; HEYNE, Nutt. Pl. (1927) 987; Koorp. Fl. Tjib. 2 (1923) 153; Merr. En. Philip. 2 (1923) 493; STEEN. Bull. J.B.B. m1, 13 (1936) 148.—A. java- nicum (non BuroM. f., 1768) JUNGH. Monatsber. Berl. Geogr. Ges. 1842; JUNGH. & DE Vr. Tijd. Nat. Gesch. & Phys. 10 (1843) 138.—A. laurinum HAssk. ib. nomen; Mia. FI. Ind. Bat. 1, 2 (1859) 582, Suppl. (1860) 200, 511; Boert. Handl. 1 (1890) 281.—Laurus alba BL. Rumphia 3 (1847) 193, in syn.—A. cassiaefolium BU. l.c.—A. philip- pinum MerR. Goy. Lab. Publ. 35 (1906) 36.—A. curranii MeERR. Philip. J. Sc. 4 (1909) Bot. 285.— Fig. 1. Tree up to 48 m, clear bole up to 28 m by 70 cm, buttresses to 2m high. At the start of the dry season foliate twigs sprout 2-4 together simultane- ously from last year’s buds; inflor. appearing in the axils of fallen leaves. Both flower and shoot- buds ca 4mm long with 4-9 pairs of decussate ca 2mm long caducous perules. Leaves simple entire glabrous, glossy dark green above, glaucous, whitish or light blue-grey below; petiole 1!/4-10 cm; blade elliptic to lanceolate, 3- to slightly 5-plinerv- ous at the base, apex acuminate to tailed, tip subacute. Inflor. corymbose, either d or Q, rarely with few fis of the other sex, glabrous, 2!/2-10 (in fruit to 19) cm long; peduncle 1/2~-3!/2 cm, pedicels 4-17 mm. Flowers pale yellowish. Sepals and petals (3-)5, free, resp. 21/2-3 and 1!/2-2!/2 mm long. Stamens (4-)6(-8) in 1 whorl, sometimes isomerous and then alternating with the petals, attached on the disc in pits; filament in d 5 mm, in9 2.2 mm; anther Fig. 1. Acer niveum BL. x 1/3, flower enlarged. strongly reduced. Styles 2, 11/2 mm long. Wings of fruit 4-7 by 1-2!/2 cm, asymmetric and obovate, inside narrowed or straight, hairy; mericarp proper 8-13 mm long, ovate. : (1) In Malaysia only one genus. 4 FLORA MALESIANA [ser. I, vol. 4', Dec. 1948] Distr. Cf. fig. 2; in the Malay Peninsula re- cently collected in the hills near Cameron High- lands (CF. 27181, 27344, 36281, 37745, 45489), in W. Borneo once near Simpang at 27 m (bb 13518) and once in Sarawak (HAVILAND 2092), in the other islands many localities. Fig. 2. Distribution of Acer niveum BL.; in Borneo and the Malay Peninsula it is very rare. Ecol. In primary, rarely in secondary or devas- tated, forests, often common but scattered, 900- 2550 m, in Flores descending to 750 m, in Sumatra to 630 m, in Celebes to 450 m, and at only 27 m near Simpang, Borneo. Fls in April—Aug., fr. July—Nov. In Casuarina forest saplings have been found. When flowering the leafless crown swarms with Hymenoptera collecting honey. Trees are easily located in the forest by the fallen leaves which are glaucous and fine-reticulate-veined underneath. Vern. Some dozens of native names have been recorded, none of these fixed (HEYNE, /.c.). Uses. Timber unimportant, no distinct heart- wood is present; used for building purposes, fit for boxes. Notes. Pax inserted A. niveum in his sect. Integrifolia in which the inflor. is terminal on short leafy twigs. I could examine this character in A. oblongum WALL. (WALL. 1222 A), A. laevigatum WALL. (THOMSON s.n.) and ‘A. niveum’ (HELFER s.n.) from India. A. niveum BL. with its axillary leafless inflorescences is, however, possibly related to PAx’s sect. Lithocarpa and does not belong in sect. Integrifolia. Excluded Acer javanicum BurM. f. Fl. Ind. (1768) 221 = Actinophora fragrans R.Br. sec. BACKER in Herb. Bog.; according to BURRET it is Colona javanica, both Tiliaceae. PHILYDRACEAE (C. Skottsberg, Goteborg) Erect herbs with a short rhizome. Leaves linear radical or crowded at the stem base, distich, equitant, parallel-nerved. Flowers zygomorphic, bisexual, solitary in the axil of spathaceous bracts. Perianth corolline, segments 4, 2-seriate. Stamen 1, inserted at the base of the abaxial segments. Filament flattened; anther 2-celled; cells straight or twisted, opening lengthwise by slits. Ovary superior, 3-celled with axile placentas, or l-celled with parietal placentas. Style simple. Ovules ~, ana- tropous. Capsule with 3 valves. Seeds ~. Distr. Centering in Australia, comprises 4 genera with 5 species. KEY TO THE GENERA 1. Outer tepals free. Anthers spirally twisted. Ovary 1-locular F 1. Philydrum 1. Outer tepals united at the base. Anthers straight. Ovary 3-locular 2. Helmholtzia 1. PHILYDRUM BANKS & SOL. ex GAERTN. Fruct. 1 (1788) 62; Mia. FI. Ind. Bat. 3 (1855) 250; HAssk. Bull. Soc. Bot. Fr. 16 (1869) xxiv; Rmpv. Fl. Mal. Pen. 4 (1924) 347; Seorrss. Bull. J: B. B: Il, 13 (1933) 111. Outer tepals free, inner ones more or less united at the base with the filament. Anther spirally twisted. Pollengrains in tetrads. Ovary 1-locular. Capsule loculicid. Testa spirally striate. Distr. Monotypic, E.-SE. Asia, and Australia, rare in Malaysia. 1. Philydrum lanuginosum BANKS & SOL. ex GAERTN. /.c.; Mia. /.c.; HAssk. /.c.; BANKS & SOL. Bot. Cooxk’s Voy. 3 (1905) t. 310; MeErR. Philip. J. Sc. 10 (1915) Bot. 88; Rip. /.c.; Skotrss. /.c.; YAMAMOTO, J. Soc. Trop. Agric. 10 (1938) 119; STEEN. J. Arn. Arb. 28 (1947) 420.—Fig. 1. Perennial caespitose herb, caudex short. Leaves densely rosulate, isolateral monofacial, glabrous, thick and of soft texture, 40-80 cm long incl. the sheath; sheath 14-30 by 1-1'/2cm and 24mm thick. Scape 1 m high or more, slender, terete, gla- brate below, villous towards the woolly inflior., with few cauline leaves gradually passing into the alternate bracts. Inflor. a terminal spike, simple or paniculate. Bracts ovate, clasping, abruptly acu- minate and subulate, 2—7 by 3/4-1 cm enclosing the buds, reflexed in anthesis, again embracing the fruit. Flowers sessile, yellow. Perianth thin, outer tepals 12-15 by up to 10 mm, acute, co-nerved, long-villous outside, margins inflexed, the posterior with 2 stronger veins and bidentate; inner petals united below 1—2 mm with the filaments, 8 by 2mm 3-nerved, spathulate, base hairy outside. Stamen 8-9 mm, glabrous; anther + spherical, 1'/2 mm across. Ovary 6-7 by 2-3 mm, densely long-wool- ly; style3-4 mm, glabrous; stigma broad-triangular long-papillose. Capsule triangular-oblong, 9-10 by 4-5mm. Seeds o, dark-reddish, bulb-shaped, 0.80.9 by 0.3-0.4 mm. Distr. E. to SE. Asia (Riu Kiu Isl., Formosa, Kwantung, Hongkong, Indo-China, Siam, Burma, Andaman Isl.) and NE. Australia, in Malaysia: only in the Malay Peninsula, and in SE. New Guinea, to be expected locally elsewhere. Ecol. In ponds, marshes, and rice-fields at low altitude, in New Guinea in sedge swamps and moist savannahs. Notes. According to MERRILL (1915) the Cu- MING specimen credited to the Philippines came from the Malay Peninsula; the HILLEBRAND speci- men is certainly erroneously believed to occur wild in Java. 2. HELMHOLTZIA F. v. M. Fragm. 5 (1866) 202; Skottss. Bot. Jahrb. 65 (1932) 260; Bull. J. B. B. mb 135(1933) 112. Tepals united to form a short cupular tube, the inner connate to half their length with the filament. Anther straight. Pollen grains single. Ovary 3-locular. Berry leathery, (apparently) indehiscent. Seed with long funicle, outer testa lengthwise striate and not spirally so. Distr. 2 species, one in Australia, the other in E. Malaysia. FLORA MALESIANA [ser. I, vol. 4! aa | Fig. 1. Philydrum lanuginosum BANKS, x 2/5 (after BANKS & SOLANDER). Dec. 1948] 1. Helmholtzia novoguineensis (KRAUSE) SKOTTSB. Il.cc.; STEEN. J. Arn. Arb. 28 (1947) 419.— ‘?Xerotidae sp.’ TEysM. Nat. Tijd. N.I. 37 (1877) 132-133.—‘Liliacea’ J. J. SmirH, Teysm. 12 (1902) 168, 329.—Astelia novoguineensis KRAUSE, Bot. Jahrb. 59 (1924) 559. Perennial herb. Rhizome stout, ascending to erect, woody, covered with leaf sheaths, up to 35 by 1-1'/2 cm; roots coarse, shoots flat, fan-shaped. Leaves densely rosulate, ensiform, 75—150 by 3—4!/2 cm; sheath 20-30 cm long, inside with scanty very long thin arachnoideous hairs, linear, acute, gla- brous, of firm texture, a bundle veins on each side forms aprominent costa dissolving to the apex, with short oblique transverse veins; blade monofacial arched or horizontal, with secondary upper and lower surface. Scape terminal 25—50 cm, erect, ob- tuse-angular, upwards covered with a dense light- grey wool, leafless in its lower half, thence carrying 5-10 reduced ensiform leaves or spathes passing into bifacial alternate bracts. Branches of Ist order of the panicle supported by a spathe, the largest 10-40 by 1'/2-3 cm, 2-3 lowermost with few branchlets of 2nd order 2-8 cm long. Bracts linear subulate 1—2 by 1/4—!/2 cm, 1—3-nerved, base woolly convolute enclosing the bud. Flowers sessile, white, PHILYDRACEAE (Skottsberg) 7. glabrous except the 2—2!/2 mm high tube. Outer tepals narrow-triangular, convolute with filiform apex, posterior one bicarinate-bicuspidate, with in- flexed margins, 9—12'/3 by 4—5 mm, anterior one 8-11 by 2!/2-3'/2 mm. Inner tepals and filament adnate to the tube, small, l-nerved, 4-5 mm long, irregularly 3-dentate, free portion 1'/2-2!/2 by 3/4-1!/2 mm. Free part of the stamen 3!/4—33/4 mm; anther 2—2!/2 by 1—1!/2 mm. Ovary 2 by 1 mm den- sely grey-woolly. Style 3-sulcate, 23/4—4'/2 mm long, stigma small triangular. Berry white, slightly 3-sul- cate, 7-8 by 6 mm, pericarp tough leathery. Seeds ~%, 2-2!/4 by 1/2 mm, _ cylindrical-flattened, often slightly curved, dark-brown with a transparent striate outer testa prolonged at both ends. Distr. Malaysia: Moluccas (Ambon, Boeroe, Ceram) and New Guinea, 600-1500 m. Ecol. In groups in muddy or moist, humic open spots in rain forests, and along ponds and margins of lakes. Fl. & fr. throughout the year. Notes. Closely allied to H. acorifolia F. v. M. from E. Australia, which has an almost glabrous scape, a more robust habit, a trifle smaller flowers, outer petals hairy on the back, style 5—6!/2 mm long, seeds mostly a little less than 2 mm. ANCISTROCLADACEAE (C. G. G. J. van Steenis, Buitenzorg) ANCISTROCLADUS (WALL. Cat. (1832) 1052) ex Arnott, Nov. Act. 18 (1836) 325; PLANCH. Ann. Sc. Nat. III, 13 (1849) 316; ScHerr. Nat. Tijd. N. I. 32 (1873) 407; BoerL. Handl. 1 (1890) p. XVH, XX; Kina, J. As. Soc. Beng. 42, II (1893) 137; MAssart, Ann. J: B. B. 12:(1895) 121; GiiG; in E & P; ed: 2, 21 (1925) 589, f. 269-70 Fs Reims Mal. Pen. 1 (1922) 250.—Bembix Lour. Fl. Coch. (1790) 282, nom. rej., cf .MOORE, J. Bot. 65 (1927) 279.—Wormia VAHL, Skrift. Nat. Selsk. Kjobenh. 6 (1810) 104, non ROTTB.—Bigamea KOEN. ex ENDL. Gen. PI. 1183 (1840). Scandent shrubs (often erect in youth), without resin; branches sympodial with a series of circinate woody hooks in one plane. Leaves spread, simple, entire, often rosette-crowded, cuneiform, penninervous, reticulate-veined, glabrous, both surfaces minutely pitted, each pit with a peltate small hair secreting a wax- like substance; petiole articulated, scar on the twigs often saddle-shaped; stipules absent. Flowers 3, actinomorphic small.-Inflor. few or several times dichotomous or spike-like, often provided with said hooks and single reduced bract-like leaves, branches often recurved. Pedicels articulated. Bracts with a glandular-thickened base, margin fimbriate-membranous. Calyx tube short, at length adnate to the base of the ovary; lobes 5 inequal imbricate, enlarged and wing-like in fruit. Petals 5, united at the base, slightly contorted in bud. Stamens mostly 10, rarely 5, the episepalous slightly longer. Filaments with broadened base; anthers basifixed, + introrse to + latrorse, 2-celled, opening lengthwise. Ovary for the greater part inferior, consisting of 3 carpels, 1-celled, protruding into a nipple- shaped elongation bearing 3 articulated erect styles with a punctiform or horse- shoe-shaped stigmatic apex; nipple enlarging in fruit. Ovule 1, basal, ascending, with 2 integuments. Nut not dehiscent, crowned by the enlarged calyx. Seed round- ish with testa intruding between the cerebral-like folds of the endosperm. Exocarp leathery. Embryo straight, erect, obliquely placed; cotyledons diverging; hypo- cotyl rather thick. Distr. Disjunct, ca 3 spp. in trop. W. Africa, and 9 in SE. Asia, from the Deccan to Burma, Indo- china, Hainan, S. China, the Malay Peninsula, Borneo and Sumatra (cf. fig. 2). Uses. Except for some local information nothing is known (cf. BURKILL). Ecol. In mixed rain forests, but most common on silicious soil in so-called ‘padang-scrub’, from the lowland to the hills. KERR noted of A. wallichii (his no 7006) that all specimens grew erect, and it is reported by GAGNEPAIN to be erect in youth. RmpDLey also found it on the ground as a bush, or ascending trees, and this is also observed in specimens from Sumatra and Borneo. In the open padang-scrub it is zither erect or trailing. Notes. This monogeneric family has been subsequently been referred to several families; it is now mostly placed next to the Dipterocarpaceae but differs by the 1-celled ovary, basal ovule, peculiar en- dosperm, climbing habit, sympodial structure, absence of stipules, and presence of hooks. HALLIER f. brought it to the Linaceae-Hugoniaceae, suggested already by MiqueL. The bark of the twigs shows a peculiar cracking viz lengthwise superficial splitting of the thin grey corky outer bark and further by deeper transverse cracks. In A. extensus I found peculiar rather large crateriform glands on the base of the bracts of the inflor. Similar glands I found on 2-3 or all 5 sepals, distinctly elevated, 1-3 together. I have not found any stipules, neither in A. extensus nor in abundant living material of A. hamatus (VAHL)GILG; there are rather large bracts leaving scars amidst the leaf-tufts but these belong apparently to the leaf-spiral. GAGNEPAIN (FI. Gén. I. C. 1 (1910) 393) mentions 3—5 styles, but I found only 3. HUTCHINSON (Fam. FI. Pl. 1 (1926) 178) apparently assumes the style to be represented by the nipple- shaped extension of the ovary above the calyx on tip of which 3 free stigmas are articulated, but the tip of the latter I found distinctly ‘stigmatic papillose’ so that I assume the styles to be articulated with the ovary. The stigmatic surface is punctiform or horse-shoe-shaped. The nipple enlarges in fruit and forms a distinct part of it. All authors assume the presence of a ruminate endosperm, but HUTCHINSON Dec. 1948] ANCISTROCLADACEAE (Vv. Steenis) 9 denies its presence and assumes the embryo to be constituted of remarkably ‘folded cotyledons’. I had no seedlings at my disposal but an examination of the seeds did not confirm HUTCHINSON’s statement. The embryo is lying loose in the endosperm. The flowers are mostly deficient or absent in our rather rich material and when drying shrink to poor and brittle remnants. However, in A. extensus I found laterally slit anthers and not introrse cells, con- trary to GAGNEPAIN’S statements. BOERLAGE mentions slits which are turned somewhat towards the inner surface. The size of the leaves varies much both in shape and dimensions in one specimen, specially between sterile and fertile twigs. In cultivated A. hamatus I found leaves of flowering twigs 6-9 by 2—2!/2 cm, and those of sterile twigs 35—40 by 4!/2-5'/2 cm. Notwithstanding the scanty flowering material I am perfectly satisfied that only one species occurs in Malaysia. 1. Ancistrocladus tectorius (LouR.) MerrR. Lingn. Sc. J. 6 (1930) 329; Comm. Lour. (1935) 275.— Bembix tectoria Lour. Fl. Coch. (1790) 282.—A. extensus (WALL. Cat. 1052, nomen) PLANCH. Ann. edie MN Ne Sat PASS staat rie - en ae TBS Ts a Siegen: DE Za ey Ee ae Fig. 1. Ancistrocladus tectorius (LOUR.) MERR., from Borneo, X 2/s. Sc. Nat. III, 13 (1849) 318; Kina, J. As. Soc. Beng. 42, I1 (1893) 137; BoerRL. Cat. pl. phan. Hort. Bot. Bog. pt 2 (1901) 114; Burk. Dict. (1935) 155.—A. pinangianus (WALL. Cat. 1054, nomen) PLANCH. l.c.; Mig. Fl. Ind. Bat. 1, 2 (1859) 587; ScHEFF. Nat. Tijd. N.I. 31 (1870) 348; 32 (1873) 407; Dyer, in Hook. f. Fl. Br. Ind. 1 (1874) 300; Ript. FI. Mal. Pen. 1 (1922) 251, f. 25.—A. extensus var. pinangianus KING, J. As. Soc. Beng. 42, II (1893) 137; GaAGN. l.c.; CratB, 1.c.—A. hainanensis HAyATA, Ic. Pl. Form. 3 (1913) 46.—Fig. 1—2. Liana, in the youth and in open scrub often a shrub, later often trailing; main shoots provided with scattered + erect small leaves, between and near which arise spreading non-foliate tendril-like shoots provided with 3-6 curved hooks, lower 2 rarely 3 hooks getting woody, hooks mostly uni- lateral, rarely 1—2 alternate; these ‘tendrils’ later woody, becoming branches, upper part vanishing. Leaves crowded mostly immediately above the 2nd hook, variable in size and shape, sessile, mostly obovate-oblong, tapering towards the base, apex obtuse, rounded, acute or even acuminate, blade 9-30 by 3-10 cm; nerves 4-8 on either side, spread- ing, connected by a slightly looped intramarginal vein and a 2nd feebler outer one, rather straight, numerous secondary veins often becoming as strong as the main nerves and parallel. Inflor. be- tween the crowded leaves, very rarely lateral in the place of a ‘tendril’ on the main shoot, repeatedly dichotomous, branches divaricate, 8-15 cm long. Flowers rather crowded at their tips. Calyx lobes inequal, oval, thin-margined, glabrous except the Fig. 2. Localities of Ancistrocladus tectorius MERR. short ciliate rounded apex, some or all lobes pro- vided with 1—3 conspicuous crateriform prominent glands, mostly shorter than the corolla, 13/s—2!/2mm long, soon enlarging. Petals oblique-oval, one mar- gin often involute, acute, 3—3'/2 by 13/4 mm. Styles erect, nearly as long as the nipple-shaped ovary- 10 FLORA MALESIANA _ [ser. I, vol. 4', Dec. 1948] top, both !/2 mm high, stigma punctiform. Stamens alternately inequal; filament broadened at the base; cells free, acute, more or less latrorse. Fruit with spreading calyx wings slightly decurrent on the obconical sub-5-angular smooth tube, oblong-cune- ate to spathulate, inequal, often oblique, apex blunt to rounded, with 3 larger nerves and numerous smaller densely reticulate ones, overlapping at the base, smallest mature ones measured 2!/2 by !/2cm, largest 5 by 13/4 cm; nipple broad-obcampanulate, + 3 mm high protruding, solid, not filled with part of the seed. Seed obconical with flat apex, ca 5 mm high, mostly consisting of a ruminate endosperm; germ ca 2—2!/2mm high, erect, straight, obliquely inserted. Distr. Burma, Siam, the Andamans, and Indo- china to S. China and Hainan, in Malaysia: Malay Peninsula, Riouw & Lingga Arch., Anambas Isl., W. Dutch Borneo, Karimata, Banka, Billiton, once collected in S. Sumatra (fig. 2). Ecol. Low altitude, often near the sea, some- times on the margin of the beach, mostly on sili- cious soils, both in mixed forest and padang scrub fr. fl. March-Aug. Vern. akar (be)boeloes, beloeloes, meloeloes (Banka), mendjoeloeng (Lepar), troeng boeloes (Billiton). Notes. I agree with BURKILL that no differences of importance can be found between A. extensus and A. pinangianus. I have tentatively accepted MERRILL’s name, though Moore stated that the type in the Br. Mus. is inadequate for specific iden- tification. It was collected in the classical locality but I am not satisfied that no other species grows there; in tropical regions the identification ‘by exclusion’ is a somewhat dangerous procedure. Excluded Ancistrocladus pentagynus Wars. Bot. Jahrb. 13 (1891) 385 = Durandea (Linac.) acc. to HALLIER f. (B.B.C. 39, IL (1921) 68-78). Ancistrocladus sagittatus WALL. = glabra Mia. (Theac.). Tetramerista APONOGETONACEAE (C. G. G. J. van Steenis, Buitenzorg) 1. APONOGETON LinnE f. Suppl. (1781) 32; ENGL. & Krause, Pfl. R. 24 (1906). Perennial lactiferous freshwater herbs, rhizome short tuberous with fibrous roots. Leaves radical, submerged or floating, base sheathing, oblong to linear, entire or crisped, often long-petiolate; nerves lengthwise parallel, connected by numerous oblique transverse veins. Spike emerging from the water, simple or 2-8-forked, without bracts, subtended by a mostly caducous basal sheath (spathe). Flowers bisexual (rarely by abortion unisexual), small, spicate-scapose, white, rose, purple, yellow or yellowish-green. Perianth segments 2 (1-3, or absent), equal or unequal, usually persistent. Stamens in 2 rows, 6 (or more), free, hypogynous, persistent; filament filiform; anthers extrorse, small, 2-celled. Pollen subglobose or ellipsoid. Gynaecium superior, apocarpous; carpels 3-6, sessile, each with a simple style. Ovules 1-8 (or more), anatropous. Mature carpels inflated, opening along the back. Seeds without endosperm; outer testa often loose; embryo straight, elongate. Distr. About 40 spp. described, Africa, Ma- dagascar, Ceylon, SE. Asia, through Malaysia (very rare) to N. Australia, centering in Africa and Madagascar. Ecol. The few Malaysian specimens were col- lected in lowland stony streams both on calcareous and other rock. The testa contains in some spp. air between the two coats and float on the water; it soon decays and the embryo sinks to the bottom. Uses. The starchy tuberous rootstock is said to be edible in some spp. Notes. Monogeneric family. Next to the single indigenous species, A. fenestralis with its unique fenestral-leaved foliage is cultivated in the Bot. Gard. Buitenzorg, and may be found in private gardens as a curiosity. SS . SKS SS 1. Aponogeton loriae MARTELLI, Nuovo Giorn. Bot. Ital. II, 3 (1897) 472, t. 8; ENGL. & KRAUSE, Pfl. R. 24 (1906) 12; Domin, Bibl. Bot. 20 (1915) 254; Camus, Bull. Soc. Bot. Fr. 70 (1923) 672-3; RENDLE, J. Bot. (1923) Suppl. 58; STEEN. Journ. Arn. Arb. 28 (1947) 419.—A. crispus (non THUNB.) F. v. M. Descr. Not. Pap. Pl. 8 (1886) 51; RIDL. J. Bot. 24 (1886) 359.—A. monostachyum (non L. f.) HEMSL. Kew Bull. (1899) 113.—Fig. 1. Submerged; rootstock roundish !/2-11/2 cm. Leaves green or brown, distinctly petiolate (2-15 cm), blade linear-spathulate, 10-35 by 14cm, mostly gradually tapering into the petiole, base narrow-cuneate, apex rather broadly cuneate and + blunt, primary nerves 2 on both sides and a marginal vein; paren- chyma opaque dotted brown-punctate; margin slightly undulate-crisped to + flat. Scape 5-40 cm. Spathe 1/2-11/2 cm long, ovate-acute, lengthwise nerved, persistent, decaying gradually from the apex towards the base, green, concave, subam- Fig. 1. Aponogeton loriae MARTELLI. Plant x '/2, plexicaulous, apex mucronulate. Flowers greenish- flower, tepal with stamen, stamen, and fruit yellow, the lower ones over 2—3 cm densely set and enlarged (after MARTELLI). 12 FLORA MALESIANA [ser. 1, vol. 4', Dec. 1948] | with developing fruits, stamens about equal- ling the tepals on 2 mm long filaments, the upper ones rather abruptly as it seems male with 3 mm long stamens, very small ovaries and flowers set laxly to remote. Spikes 11/2-7 elongating in flower up to 18-20 cm. Tepals obovate ca 1!/2-13/4 by 1-11/4 mm, concave, apex broadly rounded. Stamens 6; anthers roundish oval, no dehisced ones observed by me. Carpels 3, ca 2'/2 mm long, ovate, + bluntly trigonous, rather abruptly beaked by a distinct recurved rostrate style about !/2mm long. Seeds (in TEYSMANN 12792) 1-6 with a delicate loose outer coat 6-winged or -ribbed, transparently brown- reticulate-netted-celled, 2 by 2/3 mm. Inner testal coat oblong, opaque, darkbrown, smooth, 11/4 by 1/2 mm, closely enveloping the straight embryo, easily splitting on slight pressure, rounded at both sides. Distr. Queensland (DiELs 8397, n.yv.), in Ma- laysia: New Guinea, and SW. Celebes in the cal- careous Maros-Pangkadjene distr. (TEYSMANN 11901, 12792). Ecol. In shallow stony streams in forests and savannahs, 100-600 m. Notes. There is a remarkable yet unexplained dimorphy in the flowers of the spike, the lower ones setting fruit only and differing in length of anthers. BURMANNIACEAE (F. P. Jonker, Utrecht) Annual or perennial, saprophytic or autotrophic herbs; the saprophytic species often colourless. Leaves usually spread or alternate, entire, simple, without stipules; non-saprophytic species with a radical rosette of linear leaves; stem leaves often reduced to small scales; sometimes the basal part of the stem provided with many decurrent, grass-like leaves. Flowers ¢, usually actinomorphic, solitary or in capi- tate or cymose inflorescences. Perianth corolline; limb consisting of 2 whorls; tube sometimes 3-winged. Anthers 3, subsessile in the perianth throat and dehiscing laterally with horizontal slits,or 6, hanging down in the perianth tube and dehiscing with longitudinal slits. Connective large, often appendiculate. Style filiform or shortly cylindrical or conical. Stigmas 3, sometimes connate. Ovary inferior, 1-celled with parietal placentation, or 3-celled with axile placentation. Ovules «, anatropous, with 2 integuments; funicles often rather long. Fruit usually capsular, sometimes fleshy, crowned by the persistent perianth tube and the style, or by a thickened persistent basal ring of the perianth tube, dehiscing irregularly or with transverse slits at the top. Seeds ~, small, subglobose to linear, sometimes with loose, reticulate testa, with endosperm. Distr. About 125 species, widely distributed in the tropics of both hemispheres, also in subtropical America, Chicago area, Mocgambique, Southern China, Japan, Southern Australia, New Zealand and Tasmania. As many species are rare, it is possible that only a part of their areais known. Most of them are found in moist regions. Among the autotrophic Malaysian Burmanniaceae there are 3 rather common species which are widely spread, viz Burmannia coelestis, B. disticha and B. longifolia. The latter two are absent from Java and the Lesser Sunda Islands, the former occurs in Java proper only in its western part. Of the saprophytic Malaysian species only 3 have been often collected, viz Burmannia championii, B. lutescens, and Gymnosiphon affinis. Ecol. The autotrophic species provided with green leaves occur in grass-fields, along road sides and river-banks, among brush-wood and in forests or on moist swampy soil, up to about 3000 m alt. The saprophytic species usually occur in dense primary or secondary forests on soils rich in humic matter by decaying wood and leaves, up to ca 1500 m alt. They are also found sometimes in bamboo bushes and parks. Notes. Treatment mostly after JoNKER, A monograph of the Burmanniaceae, Thesis, Utrecht, 1938; also in: Meded. Bot. Mus. & Herb. Utrecht no 51; slightly revised. In collecting Burmanniaceae it is necessary to collect plants with complete flowers, as the limb with the stigmas and stamens is often caducous. The fruits are also important. The colour of the flowers, stems and leaves must be noted. Preservation of collections in 60 °%/o spirits is recommended. In the field the saprophytic species are often found in colonies together with other saprophytic plants belonging to the Orchidaceae, Triuridaceae, and Gentianaceae. From the extreme rarity of a number of species it may be assumed that by further collecting these tiny plants several novelties will be found. KEY TO THE GENERA 1. Perianth tube cylindrical or trigonous, persistent on the capsule. Style of equal length as the tube. Anthers 3, subsessile in the perianth throat. Thecae dehiscing laterally with transverse slits TRIBE Burmannieae MIERS 2. Ovary and capsule 3-celled with axile placentation. Perianth as a whole persistent on the capsule. Ovary and perianth often prominently 3-winged, sometimes 3-costate or wingless. Ovary without glands. Capsule mostly dehiscing irregularly . . nae 1. Burmannia 2. Ovary and capsule 1-celled with parietal placentation. Perianth limb with the stamens and stigmas deciduous. Ovary and perianth wingless. Both sides of the top of each placenta inside the ovary provided with a gland. Capsule reticulate-perforated . . . . ee 2. Gymnosiphon 1. Perianth tube urceolate, circumscissile, only a small basal ring persistent on the fruit. Style very short, cylindrical or conical. Anthers 6, ae down in the tube. Thecae dehiscing introrsely with longi- tudinal slits .. Oa TRIBE Thismieae MIERS 3. Inner perianth lobes free, ¢ or converging at their tops or connate to a mitre with 3 holes, the latter without appendages at the apex : . © ~3. Thismia 3. Inner perianth lobes connate to a mitre with 3 holes, crowned by l or 3 appendages. 4. Mitre crowned by 3 erect, thick, filiform appendages, clavately swollen at theirtops . 4. Geomitra 4. Mitre crowned by 1 erect thick column, bearing at its apex 3, more or less connate, glandular MODES a, Vs, os fold com Reh es Wenge act tee tee er aie eam oe eee 5. Scaphiophora 14 FLORA MALESIANA | [ser. I, vol. 4! - Fig. 1-8. Burmanniaceae. 1. Burmannia bifaria J. J. S., x 41, 2. Scaphiophora gigantea JONK., X 1/3, 3. Thismia aseroe BrEcc., 8/s, 4. Th. episcopalis (BECC.) F. v. M., x 2/3, 5. Burmannia coelestis Don, x 3/2, 6. Burmannia championii Taw., * 1/1, 7. Gymnosiphon aphyllus BL., < 2/3, 8. Burmannia . longifolia BEcc., X 2/s. WN Dec. 1948] BURMANNIACEAE (Jonker) l 1. BURMANNIA LINnNE, Sp. Pl. ed. 1 (1753) 287; JoNKER, Monogr. (1938) 18, 57. Annual or perennial, saprophytic and colourless or chlorophyllose. Flowers often 3-winged. Perianth limb usually consisting of 6 lobes; the outer ones being much larger; inner 3 often minute, sometimes lacking. Perianth tube cylindrical to trigonous. Anthers 3; connective sometimes with 2 apical crests and/or a hanging, median, basal spur. Style filiform, branching into 3 short branches, each bearing a stigma, or 3 sessile stigmas at the apex of the style. Ovary trigonous. Fruit cap- sular, mostly dehiscing irregularly. Seeds many, oblong or ellipsoid. Distr. 57 species, tropics of both hemispheres, also in the S. United States, S. part of S. America Mogambique, S. China, Japan and S. Australia. KEY TO THE SPECIES 1. Perennial, leafy green herbs. Greater part of the stem beset with grass-like, linear or ensiform, de- current, imbricate leaves. Inflorescence usually many-flowered. Flowers hanging, very narrowly 3-winged in the basal part . . . . (Sect. Foliosa Jonx.). 1. B. longifolia 1. Annual or perennial, saprophytic or green herbs. Stem leaves reduced to small scales. The non- saprophytic species with a radical rosette of linear leaves. Stem usually 1- to few-flowered. Flowers ercer =: - + (Sect. Euburmannia MALME) 2. Non- saprophytic, " chlorophyllose herbs with a Tosette of green leaves at the base; rosette often consisting of only 1-3 leaves. 3. Flower wings narrower than the perianth tube or reduced to ribs. 4. Basal rosulate leaves few. Stem bearing 1-2 flowers at its apex. Connective with 2 apical crests, basal spur lacking. Ovary as long as the perianth orlonger . . 2. B. geelvinkiana 4. Basal rosette well developed. Stem bearing at its apex a usually bifid inflorescence. Connective provided with 2 apical crests and a basal, hanging spur. Ovary shorter than the perianth 3. B. bancana 3. Flower wings as wide as the perianth tube or wider. 5. Margin of the perianth lobes double. Connective with a basal hanging spur and 2 apical crests. Thecae separated. 6. Robust herbs with a well developed rosette of ae up to 15 cm long leaves. Inflorescence usually a bifid, many-flowered cyme . . . . . 4, B. disticha 6. Slender herbs. Basal rosulate leaves about i cm. Stem bearing at its apex a single flower or a cluster of few flowers . . . . . 5. B. coelestis 5. Perianth lobes with single margin. ‘Connective with 2 apical crests; basal spur lacking. Thecae connate below the basal connective margin . eerie ee) SSG a 2. Saprophytic herbs without chlorophyll. Radical rosette absent. 7. Flowers wingless, 3- or 6-costate. 8. Stem scales many, imbricate in the lower part of the stem. Ovary as long as the perianth or longer”. =. > » « « « Js Bs sphacnoides 8. Stem scales not imbricate. Ovary shorter than the perianth. 9. Flowers 6-costate. Perianth limb thick, fleshy, more or less succulent ye 8. B. bifaria 9. Flowers 3-costate to narrowly 3-winged. Limb not fleshy. 10. Inflorescence usually capitate. Inner perianth lobes spathulate, sometimes slightly papillose. Connective mucronate at the apex, obtuse at the base . . . . 9. B. championii 10. Flowers usually pedicellate. Inner perianth lobes broadly obovate: distinctly papillose. Con- nective not mucronate, acute atthe base . . . ..... -. 10. B. micropetala 7. Flowers 3-winged. 11. Inner perianth lobes absent. 12. Perianth lobes simple. Connective with an apical, pepe crest and a basal, hanging, obtuse spur : «5, Os, je bis eS eeutats 12. Perianth lobes bind: Connective without crest and ‘spur = 35S es ee 12. B. oblonga 11 Inner perianth lobes not lacking, sometimes very small. 13. Very delicate plants. Connective with a basal, hanging spur . . . . . 13. B. steenisii 13. Plants not very delicate. Connective without basal, hanging spur. p 14. Stem rather robust, often many-flowered. Inner perianth lobes minute, orbicular. Flower wings variable, linear to half cuneate or quadrangular . . . . . 14, B. lutescens 14. Slender herbs, usually 1-flowered. Inner perianth lobes lanceolate. Flower wings elliptical 15. B. malasica 16 FLORA MALESIANA [ser. I, vol. 4! 1. Burmannia longifolia BEcc. Malesia 1 (1877) 244; JONKER, Monogr. (1938) 20, 59.—B. leucantha ScHLtr. Bot. Jahrb. 49 (1913) 107.—Fig. 8. Perennial, 12-50 cm. Stem usually simple, forked at the top into the bifid inflorescence or bearing a simple cincinnus. Leaves linear, sometimes keeled, decurrent, stem-clasping, acute, sometimes subul- ate, parallel-veined but midrib more prominent, growing smaller towards the top, 4-20cm by 2-9 mm. Upper part of stem beset with appressed, scattered, lanceolate, acute scales, 5-33 mm. Basal part of stem with brownish, dried leaves. Inflo- rescence 32-1-flowered, branches up to 4cm. Bracts. scale-like, linear-lanceolate, 5-10 mm. Flowers subsessile, hanging, white, often with pale- violet or bluish limb, 8-16 mm. Outer perianth lobes deltoid, acute, 2—4'/2 mm; margin fleshy at the base. Inner ones broad-obovate to orbiculate, entire and rounded, or retuse, or bilobate, 1!/2-2 mm. Perianth tube cylindrical, sometimes swollen in the upper part, 3-5 mm; lower part of tube and ovary very narrowly 3-winged. Stamens inserted just be- low the inner perianth lobes. Connective broad, oblong, crowned by two, rather wide crests. Fila- ments short, broad. Style thick, branching into 3 very short branches, each bearing a curved funnel- shaped stigma. Ovary obovoid, 4-7 mm. Capsule obovoid, dehiscing transversally, irregularly. Seeds oblong to scobiform, appendaged at both sides; testa loose, reticulate. Distr. All over Malaysia, except Java and the Lesser Sunda Islands. Ecol. In mountain forests, brush-wood, along mossy trails, often on ridges, scattered, ascending to 2800 m alt. 2. Burmannia geelvinkiana BEcc. Malesia 1 (1877) 244; JoONKER, Monogr. (1938) 111. Annual, 7-12 cm. Stem filiform, simple, bearing 1 or, sometimes, 2 flowers. Rosulate /eaves 2-5, linear, subulate, 3-nerved, 3-5 mm by 1 mm. Stem leaves scale-like, appressed, linear-lanceolate, acuminate to subulate, up to 3 mm. Bracts ovate- lanceolate, long-acuminate, 3-nerved, 1!/2 mm. Flowers bluish, very narrowly 3-winged, 7 mm. Outer perianth lobes triangular to broad-ovate, apiculate, about 11/2 mm. Inner ones linear, obtuse, 1/2-1 mm. Connective thick, triangular, obtuse at the base, bearing two divergent, slightly papillose crests at the apex. Style rather short and thick, bearing 3 sessile, funnel-shaped stigmas; style with stigmas about 11/2 mm. Ovary ellipsoid to narrowly obovoid, about 4 mm. Flower wings linear, about 51/2 mm by '/4 mm. Capsule obovoid, dehiscing with transverse slits. Seeds ovoid, bright yellow. Distr. Malaysia: West New Guinea (Wandam- men Peninsula, Geelvink Bay), once collected. 3. Burmannia bancana Mia. FI. Ind. Bat. Suppl. 1 (1860) 617; JoNKER, Monogr. (1938) 24, 113.—B. graminifolia WARB. in FEDDE, Rep. 18 (1922) 330. Annual, 20-37 cm. Stem simple, terete, forked into the inflorescence. Rosette distinct; eaves many, linear to lanceolate, subulate, parallel-veined, 2!/2-7 cm by 6 mm. Stem leaves few, scale-like, ap- pressed, lanceolate, subulate, 1-3 cm. One rosette sometimes bearing 2 or 3 stems. Inflorescence a double cincinnus, 3- to 5-flowered; branches up to 3 cm. Flowers blue or purplish, narrowly winged, 6— 13 mm. Outer perianth lobes lanceolate-triangular, acute, with 3 prominent, fleshy nerves inside, up to 3 mm. Inner ones linear-lanceolate, obtuse, up to 2'/2mm; midrib prominent, fleshy. Perianth tube cylindrical-trigonous, up to 4!/2 mm. Connect- ive oblong, provided with a basal, hanging, obtuse spur and 2 apical, divergent obtuse crests. Style filiform, branching into 3 short branches, each bearing a slightly curved, funnel-shaped stigma. Ovary truncate-ellipsoid, 3-7 mm. Flower wings linear, 11 by '/2-1 mm. Capsule ellipsoid to obovoid. Testa of the seeds elongate. Distr. Malaysia: Sumatra, Banka, Billiton, Borneo. Ecol. Wet places, along streams, &c. Vern. Roempoet taroem, oemboet oemboet (Bil- liton). . 4, Burmannia disticha LINNE, Spec. Plant. 1 (1753) 287; JonKEeR, Monogr. (1938) 115.—B. distachya R. Br. Prod. Fl. Nov. Holl. 1 (1810) 265.—B. suma- trana Mia. FI. Ind. Bat. Suppl. 1 (1860) 616.—B. disticha var. sumatrana Hook. f. Fl. Br. Ind. 5 (1888) 664. Robust annual, up to 75 cm. Stem usually simple, forked into the inflorescence. Rosette distinct; leaves linear or lanceolate, acute, up to 15 cm by 13 mm. Stem leaves reduced to appressed, lance- olate, acute or acuminate scales, up to 7cm by 7 mm, imbricate in the lower part of stem; upper part of stem often leafless. Inflorescence branches up to8cm. Bracts lanceolate, acute, about 5-12mm. Flowers sessile or shortly pedicellate, blue or pur- plish, often with yellow-tipped, greenish lobes, rarely yellow, 10-20 mm. Outer perianth lobes tri- angular, acute, 2!/2mm; margin thick, doublein the basal part. Inner lobes linear-lanceolate, fleshy, obtuse, 1—1'/2 mm. Perianth tube cylindrical-trigo- nous, 3—4!/2 mm. Connective broad, provided with 2 distinct, acute apical crests and a broad, obtuse to almost truncate, basal, hanging spur. Style thick- filiform, bearing 3 sessile, funnel-shaped stigmas; style with stigmas about 3 mm. Ovary ellipsoid to obovoid, truncate, attenuate towards the base, up to 1 cm. Flower wings elliptical, 10-18 by 11/2-2!/2 mm, continuing as crests on the back of the outer perianth lobes, decurrent along the short pedicel. Capsule obovoid, truncate, irregularly dehiscing with transverse slits. Distr. Widely distributed in the tropics of Asia and Australia: Ceylon, India, Siam, Indo-China, China, through Malaysia to Australia, in Malaysia hitherto not found in Java, the Lesser Sunda Is- lands, Moluccas, and Philippines. Ecol. A species with a large ecological ampli- tude. It has been collected in brush-wood, swamps and bogs among Sphagna, moist hollows, open grasslands, mountain meadows, marshy plateaus, on bare rocks, and has even been recorded as growing in water; ascending to ca 3500 m alt. Vern. Si goeroe goeroe (Sumatra). Dec. 1948] 5. Burmannia coelestis DON, Prod. Fl. Nep. (1825) 44; JONKER, Monogr. (1938) 120.—B. javanica BL. Enum. FI. Jav. 1 (1827) 28.—B. triflora Roxs. FI. Ind. 2 (1832) 117.—B. azurea GrirF. Not. 3 (1851) 326.—B. selebica Becc. Malesia 1 (1877) 243.—B. borneensis, B. chinensis, B. malaccensis & B. rigida GANDOG. Bull. Soc. Bot. Fr. 66 (1919) 290.—Fig. 5. Autotrophic annual, up to 30 cm. Stem simple or, sometimes, branched, bearing a single flower or a cluster of few flowers. Rosulate Jeaves linear or lanceolate, acute or acuminate, 3-nerved, about 1 cmby 1!/2-3 mm. Stem leaves appressed, imbricate in the basal part, linear-lanceolate, subulate, rather long, up to 2cm. Bracts lanceolate, acute, 4 by 1mm. Flowers prominently 3-winged, blue, pur- plish or white, often with yellow lobes, about 11!/2 mm. Outer perianth lobes ovate, apiculate, with double margin, about 1'/2 mm. Inner ones lanceolate, apiculate, with double margin, about 1/2 mm. Tube cylindrical-trigonous, about 5 mm. Connective provided with 2 apical, divergent, ob- tuse to truncate crests and a basal hanging, rather long, obtuse spur. Style thick-filiform, bearing 3 sessile, funnel-shaped stigmas with swollen margin. Style with stigmas about 4 mm. Ovary ellipsoid to obovoid, truncate, attenuate towards the base, about 5mm. Flower wings half elliptical to half obovate, about 10 by 2!/2 mm. Capsule obovoid, truncate, transversely dehiscing. Distr. Widely spread in tropical Asia: India, Siam, Indo-China, S. China, and the Caroline Is- lands, in Malaysia throughout the Archipelago, in Java only in the W. part, once collected in the S. part of Madoera Island, and once in Bali, other- wise absent from the Lesser Sunda Islands. Ecol. Grass-fields, among alang-alang (/mpe- rata), in mountain meadows, parks and plantations, ascending to ca 1700 m alt. 6. Burmannia connata JONKER, Monogr. (1938) 128. Autotrophic annual, 15-30 cm. Stem simple, bearing 1-3, shortly pedicellate flowers. Rosulate leaves few, linear, acute or acuminate, 1-nerved, 4-8 by 1 mm. Stem leaves scale-like, appressed, linear, acute, 2-5 mm. Bracts linear-lanceolate, acute, 1'/2 mm. Flowers prominently 3-winged, 6-8 mm. Outer perianth lobes acute, with involute margin, 1 mm. Inner ones broadly ovate, obtuse, nearly '/2mmlong. Perianth tube cylindrical, 3 mm. Connective rather broad, provided with 2 apical, divergent, obtuse crests. Thecae bright yellow, ap- pressed against the connective and connate below the basal connective margin. Basal hanging spur lacking. Style as long as the tube, branching into 3 short branches, each bearing a peltate, disk- shaped stigma. Ovary ellipsoid to obconical, 2'/24 mm. Flower wings half oblanceolate, decur- rent along the pedicel, 8 by 2mm. Capsule ellipsoid, dehiscing with transverse slits ,4—6 mm. Seeds scobi- form, yellow. Distr. Malaysia: Sumatra, Eastcoast Resi- dency (Kota Pinang, Soengei Kana). 7. Burmannia sphagnoides Becc. Malesia 1 (1877) 246; JONKER, Monogr. (1938) 135. BURMANNIACEAE (Jonker) 17 Saprophyte, 4!/2-12cm. Stem simple, thick, beset with many lanceolate, acute, in the lower part im- bricate, 3-6 mm long, scale-like /eaves and bearing 2—5 subsessile flowers at the apex. Bracts broad- lanceolate to ovate, acute, 3-6 mm. Flowers 6-cost- ate, white, about 8'/2 mm. Outer perianth lobes broadly triangular, with swollen margin, about 8'/2 mm. Inner ones fleshy, obtuse to rounded, papillose, almost 1 mm. Perianth tube cylindrical, about 2 mm. Connective oblong, acute at the base crowned by 2 divergent, obtuse crests. Style thick- filiform, bearing 3 sessile, obconical stigmas. Ovary large, broadly ellipsoid to subglobose, 4-5 mm. Distr. Malaysia: Malay Peninsula, Sumatra (Eastcoast Res.), and Borneo (Sarawak). Ecol. A rare species, occurring in decaying matter in forests. 8. Burmannia bifaria J.J.S. Icon. Bogor. 4 (1914) 379; JONKER, Monogr. (1938) 136.—B. engganensis JONKER, Blumea 3 (1938) 108; Monogr. (1938) 137. —Fig. 1. Saprophyte, 5-13 cm. Stem simple or branched, beset with scale-like, ovate to lanceolate, 1-nerved, sometimes distichous, up to 5 mm long /eaves and bearing 1 flower or branching into a bifid, up to 9-flowered cyme. Flowers shortly pedicellate, 6- costate, white or somewhat purplish, 9-13 mm. Limb fleshy. Outer perianth lobes triangular, ob- tuse, with involute, crenate margin, 1!/2-2 mm. Inner ones ovate to orbicular, !/4-1!/2 mm. Perianth tube cylindrical-trigonous, 2!/2-5 mm. Connective obtri- angular, crowned by 2 divergent, papillose crests. Style thick-filiform, branching into 3 short branch- es, each bearing a somewhat funnel-shaped stigma with a broad, rotundate, membranous, hanging appendage. Style with stigmas 4~4!/2 mm. Ovary ellipsoid, 3-6 mm. Seeds ovoid, brown. Flower wings reduced to narrow, linear ribs. Distr. Malaysia: West Java (vicinity of Buiten- zorg) and Enggano Island (off SW. Sumatra). Ecol. Among decaying leaves in forests, as- cending to 1000 m alt. 9. Burmannia championii THw. Enum. Pl. Zeyl. (1864) 325; JoNKER, Monogr. (1938) 138.—B. tuberosa Becc. Malesia 1 (1877) 245.—B. capitata (non Mart.) MAKINO, Bot. Mag. Tok. 4 (1890) 23. —B. japonica MAxIM. ex MAK. Ill. Fl. Jap. 1, no 7 (1891) 4.—B. dalzieli RENDLE, Journ. Bot. 40 (1902) 311.—B. chionantha SCHLTR. Bot. Jahrb. 49 (1913) 107. Fig. 6. Saprophyte, 2-18cm. Rhizome tuberous or elongate, covered with hair-like roots, producing small, adventitious tubers. Stem simple, beset with lanceolate, acute, appressed, scale-like, 1'/2-4 mm long Jeaves. Bracts lanceolate, acute, about 3 mm. Flowers subsessile in a capitate inflorescence at the top of the stem, 3-costate, white, 5-12 mm. Outer perianth lobes triangular, acute, with involute margin in the upper part, 1—2!/2 mm. Inner ones spathulate, rounded, slightly papillose at the mar- gin, about 3/4 mm. Connective broadly oblong, ob- tuse at the base, crowned by 2 indistinct, divergent, obtuse crests and provided with a median small 18 FLORA MALESIANA [ser. I, vol. 4! point at the apex, usually directed inwards and then hardly perceptible. Style thick-filiform, bearing 3 subsessile funnel-shaped stigmas; style with stig- mas 3 mm. Ovary ellipsoid to obovoid, 2-3 mm. Fig. 9. Burmannia lutescens Brcc., with broad wings (Mt Gedeh, W. Java), < !/1. Distr. Ceylon, S. China, Japan and Malaysia: Batoe Islands (off W. Sumatra), Banka, W. Java, Borneo, and New Guinea. Ecol. A species with a large ecological ampli- tude, occurring in humus of moist forests and also in parks, plantations, bamboo bush, on rocks in streams. 10. Burmannia micropetala RIDL. Trans. Linn. Soc. II, 9 (1916) 228; JoNKER, Monogr. (1938) 140. Saprophyte, 7!/2-15 cm. Stem simple, beset with acute or acuminate, 1-nerved, often keeled, 2-5 mm long, scale-like /eaves. Bracts linear-lance- olate, acute, 1-nerved, about 4 mm. Pedicels up to 5mm. Flowers shortly pedicellate, in contracted 3-8-flowered bifid or, sometimes, pseudo-umbel- late cymes, very narrowly 3-winged to 3-costate, 7-9 mm. Outer perianth lobes triangular, acute, about 2 mm, in the upper part provided with small, rounded, crenate lateral lobes. Inner ones broadly obovate, rounded, papillose at the margin, !/2 mm. Perianth tube cylindrical, about 3 mm. Connective oblong, acute at the base, crowned by 2 divergent, acute crests. Style thick-filiform, branching at the apex into 3 very short branches, each bearing a funnel-shaped stigma with 2 small, apical points. Style with stigmas about 3 mm. Ovary ellipsoid, truncate, about 2!/2 mm. Distr. Malaysia: New Guinea only. 11. Burmannia tridentata Becc. Malesia 1 (1877) 246; JONKER, Monogr. (1938) 141. Saprophyte, 6-14cm. Stem simple or branched, beset with appressed, lanceolate, acute, 1-veined, slighly keeled, scale-like, 1!/2-2 mm long leaves. Bracts ovate-lanceolate, acuminate, 1-veined, about 1!/2mm. Stem or branches bearing 1-3, pro- minently winged, 2-7 mm long flowers. Outer peri- anth lobes triangular to ovate, obtuse, swollen at the margin, 1—1!/2 mm. Inner lobes absent. Perianth tube cylindrical, about 2mm. Connective quad- rangular with a broad, swollen, obtuse, hanging, basal spur, and an apical, erect, papillose, obtuse crest. Style as long as the tube, bearing 3 subsessile, funnel-shaped stigmas. Ovary subglobose, about 2mm. Flower wings half elliptical to half-orbi- culate, about 4 by 2mm. Capsule subglobose, about 2!/2 mm. Seeds scobiform. Distr. Malaysia: Borneo, Sarawak (Mt Mat- tang), once collected. 12. Burmannia oblonga RIDL. J. Str. Br. R. As. Soc. 41 (1904) 33; JoNKER, Monogr. (1938) 25, 142.— B. bifida GAGNEP. Bull. Soc. Bot. Fr. 54 (1907) 462. Saprophyte, 7-15 cm. Stem simple or branched, bearing 1-2 flowers, beset with appressed, ovate to lanceolate, obtuse, scale-like, about 1!/2mm long leaves. Below the flower 2 lanceolate, scale-like bracts, 2!/2 mm. Flowers white, sometimes with yellow limb, 8-10 mm. Outer perianth lobes bifid, obtuse, about 1!/2 mm, papillose in the upper half at the margin, in the lower half with 2 yellow bags inside, provided with 2 involute, narrow triangular lateral lobes. Inner ones absent. Tube conical, 4-4!/2mm long. Connective oblong, yellow, without crests or spur. Style thick-filiform, bearing at the apex 3 sessile, funnel-shaped curved stigmas. Style with stigmas about 4!/2 mm. Ovary subglobose, 2'/2-4 mm. Flower wings obovate, truncate, white, 5-7!/2 by 3-4 mm. Distr. Hainan, Indo-China and Malaysia: Ma- lay Peninsula, N. Sumatra (Atjeh and Eastcoast Res.). Ecol. On forested rocks or loamy soil in dense jungle or forest, ascending to 1300 m. 13. Burmannia steenisii JONKER, Monogr. (1938) 158. Delicate saprophyte, 2-6 cm. Stem simple or branched, bearing 1-2 flowers, beset with lance- olate, acute, scale-like, !/2-11/2 mm long eaves. Below each flower 1 or 2 lanceolate, 1-veined, acute bracts, about 1'/2mm. Flowers pure white with yellow Dec. 1948] limb, prominently 3-winged, 3-7 mm. Outer peri- anth lobes triangular, subobtuse, with swollen mar- gin, about 1 mm. Inner ones orbiculate, minute. Perianth tube cylindrical-trigonous to conical-trig- onous, about 2!/2 mm. Connective quadrangular, with a broad, obtuse, basal hanging spur and crowned by 2 short, thick, straight, obtuse, di- vergent crests. Style thick-filiform, bearing 3 ses- sile, slightly curved, bilabiate, funnel-shaped stig- mas. Ovary subglobose, about 2 mm. Flower wings half elliptical to half-quadrangular, pure white, about 4!/2 by 1!/2mm. Capsule subglobose, dehiscing with transverse slits. Seeds scobiform. Distr. Malaysia: E. Java, Pasoeroean Resi- dency (Mt Lamongan). Ecol. Collected on the SW. slope of Mt La- mongan on coarse, volcanic sandy soil in brush- wood, ca 600 m alt. It is the only species of the family hitherto reported from East Java. 14. Burmannia lutescens BEcc. Malesia 1 (1877) 246; JONKER, Monogr. (1938) 24, 148.—Gonianthes candida BLUME, Cat. Gew. Buitenzorg (1823) 20.— Gonyanthes candida BLUME, Flora 8 (1825) 123.— B. candida (Bu.) ENGL. Nat. Pfl. Fam. 2,6 (1889) 50, not B. candida GriFF. ex HooK. f.—B. gracilis Rw. J. Str. Br. R. As. Soc. 22 (1890) 335.—B. papillosa STAPF, Trans. Linn. Soc. II, 4 (1894) 232. —B. novae-hiberniae SCHLTR. in K. Scu. & LAUT. Nachtr. Fl. D. Sch. Geb. (1905) 73.—B. gjellerupii J.J.S. in FEDpE, Rep. 10(1912) 487.—B. gonyantha Hocure. Candollea 2 (1925) 325.—Fig. 9-10. Saprophyte, up to 23 cm. Stem thickly filiform to robust, simple or branched, 1-flowered or forked into the inflorescence. Leaves lanceolate to ovate, acute, 1-3!/2 mm. Bracts lanceolate to ovate, often keeled and carinate. Cincinni bifid, up to 11-flow- ered; branches up to 3cm. Flowers pedicellate, white, sometimes with yellow limb, seldom bluish, about 8!/2mm. Outer perianth lobes ovate or tri- angular, apiculate, about 1!/2 mm; margin fleshy. Inner lobes minute, fleshy, orbiculate. Perianth tube trigonous, 21/2-5 mm. Connective truncate, rounded at the base, slightly 2-lobed at the apex into 2 very short, papillose crests. Basal spur ab- sent. Style thick, bearing 3 subsessile, funnel- shaped to bowl-shaped stigmas. Style with stigmas as long as the tube. Ovary subglobose to truncate- globose, 3-5 mm. Flower wings various, linear, or elliptical, or rather broad, half-cuneate or qua- drangular, running from the base of the limb to the middle or the base of the ovary. Capsule subglo- bose, dehiscing with large horizontal slits. Distr. Malaysia: Sumatra, Malay Peninsula, Borneo, Java, New Guinea, and New Ireland. Ecol. Usually in the humus of shady moist forests, up to 1500 m. Notes. Specimens with narrow perianth wings have been described as B. lutescens, B. novae-hiber- niae, B. gjellerupii, with elliptical ones as Goni- anthes candida, and rather broad-winged speci- BURMANNIACEAE (Jonker) 19 mens as B. gracilis. They all belong to one species variable in that respect. In Java the species was often called B. candida (BL.) ENGL. but this is a later homonym; B. candida GrirF. ex HooK. f. is Fig. 10. Burmannia lutescens BEcc., Mt Salak, W. Java. Form with narrow perianth wings, x 2/3. an allied species, occurring in Burma, Siam and the Langkawi Islands. 15. Burmannia malasica JONKER, Monogr. (1938) 152.—Burmannia lutescens (non BECC.) WINKLER, Bot. Jahrb. 48 (1913) 96. Saprophyte, 5!/2-8 cm. Stem simple, 1-flowered, seldom 2-flowered, beset with few appressed, lance- olate, subacute, 1-veined, slightly keeled, 11/2-2mm long, scale-like /Jeaves. Bracts elliptical, acuminate, l-veined, 1!/2 mm. Flowers purple or white with yellow jlimb, prominently winged, about 5 mm. Outer perianth lobes triangular with swollen mar- gin, acuminate to apiculate, about 1 mm. Inner lobes erect, lanceolate-ovate, obtuse, about !/2 mm. Perianth tube cylindrical, 11/2 mm. Connective obtriangular, obtuse at the base, provided with 2 short, divergent crests at the apex. Style cylindrical, bearing 3 sessile, funnel-shaped stigmas. Ovary subglobose to ellipsoid, 2!/2by 2 mm. Flower wings half-orbiculate to half-elliptical, about 4 by 2 mm. Capsule ellipsoid to obovoid, dehiscing with a transverse slit. Seeds scobiform to fusiform. Distr. S. Siam and Malaysia: SE. Borneo. 20 FLORA MALESIANA [ser. I, vol. 4! 2. GYMNOSIPHON BiumeE, Enum. Pl. Jav. I (1827) 29; JONKER, Monogr. (1938) 27, 168. Saprophytic annuals, without chlorophyll. Leaves scale-like. Perianth limb con- sisting of 6 lobes, the 3 outer being much larger and slightly 3-lobed. Anthers 3, sessile in the throat. Thecae bursting horizontally. Ovary ovoid to globose, 1-celled with 3 parietal placentas, each placenta provided with a large, globose gland at both sides of the top. Style filiform, branching into 3 short branches, each bearing a stigma. Perianth limb, stamens and the upper part of style with the stig- mas caducous after flowering. Capsule crowned by the persistent perianth tube. Seeds ovoid to globose, reticulate. Distr. 29 species, tropics of both hemispheres, not in Australia. Notes. In Asia, and Malaysia, this genus is represented by the section Eugymnosiphon URBAN only, characterized by the reticulate-perforated capsule dehiscing at the top. KEY TO THE SPECIES 1. Anthers inserted above the middle of the perianth. 2. Flowers pedicellate, in loose, many-flowered cincinni or bifid cincinni 1. G. aphyllus 2. Flowers - sessile in a 1- or sparsely flowered inflorescence. 3. Capsule + globose. Margin lobes of the outer perianth lobes crenate . 3. Capsule conical-ovoid. Margin lobes of the outer perianth lobes entire 2. G. oliganthus 3. G. minahassae 1. Anthers inserted in or below the middle of the perianth. 4. Anthers inserted below the middle of the perianth. Connective apiculate at the top. Ovary elongate- conical, tapering to the pedicel 4. G. affinis 4. Anthers inserted in the middle of the perianth. Connective not apiculate. Ovary marked from the pedicel. 5. Flowers sessile or subsessile in loose cincinni or bifid cincinni . 5, G. papuanus 5. Stem 1-2-flowered, or many sessile flowers in a capitate inflorescence at the top of the stem. 6. Outer perianth lobes acuminate, a third of the length of the whole perianth. Connective narrow. Stigmas with dorsal crest. Inflorescence 2- to many-flowered 6. G. neglectus 6. Outer perianth lobes deltoid, short, a fifth of the whole Bonen Stigmas without crest. Inflo- rescence 1—3-flowered 1. Gymnosiphon aphyllus BLUME, Enum. PI. Jav. 1 (1827) 29; JonKER, Monogr. (1938) 30, 170.—G. borneense BEcc. Malesia 1 (1877) 241.—G. pedicel- latum SCHLTR. Bot. Jahrb. 49 (1913) 105.—Fig. 7. Stem up to 17 cm, forked into a bifid cincinnus or bearing a simple cincinnus. Leaves acute, often keeled, appressed, 1—2!/2 mm. Bracts ovate, obtuse, scale-like. Pedicels 1-5 mm. Inflorescence often loose and many-flowered. Perianth white or lilac; tubular part up to 4mm; limb about 2!/2 mm. Outer perianth lobes ovate, obtuse, provided with a narrow, crenate lateral lobe at both sides. Inner ones linear-lanceolate, minute. Stigmas curved, funnel-shaped, inappendiculate. Capsule about 3 mm. Distr. S. Siam, throughout Malaysia. Ecol. On humus or decaying wood or leaves in the shade of moist forests, below 1500 m alt. 2. Gymnosiphon oliganthus SCHLTR. Bot. Jahrb. 49 (1913) 101; JoNKER, Monogr. (1938) 172. Stem tender, simple or branched, 4—9!/2 cm, 1- or 2-flowered. Leaves and bracts minute, to 1 mm, keeled and appressed. Pedicels up to 3 mm. Flowers white to bluish lilac, up to 5 mm. Outer perianth lobes triangular, subobtuse, provided with crenate lateral lobes. Inner ones small, cuneate, obtuse to truncate. Stigmas auriculate, soup-plate-shaped. 7. G. pauciflorus Distr. Malaysia: NE. Brit. New Guinea, once collected. Ecol. In forests, 450 m alt. 3. Gymnosiphon minahassae SCHLTR. Bot. Jahrb. 49 (1913) 104; JonKER, Monogr. (1938) 172. Stem usually simple, 2-5-flowered, 7-12 cm. Leaves acute, appressed, up to 1 mm. Bracts more or less obtuse, keeled. Pedicels 1-3 mm. Perianth white with bluish limb. Outer perianth lobes ob- tuse; lateral lobes entire. Inner ones lanceolate. Perianth limb deciduous above the anthers. Stigmas quadrangular, truncate at the apex, apiculate at the base. Distr. Malaysia: N.Celebes (Minahasa), once collected. Ecol. In humic soil, 800 m alt. 4. Gymnosiphon affinis J.J.S. Nova Guinea 8 (1909) 194; Jonker, Monogr. (1938) 31.—G. tor- ricellense SCHLTR. Bot. Jahrb. 49 (1913) 101. Stem 7-13 cm, simple or branched, white, lilac or rose-coloured, forked into a bifid cincinnus or bearing a simple cincinnus of 1-3 flowers. Leaves ovate, acuminate, 1-2 mm. Bracts to 3 mm. Pedi- cels 14mm. Perianth white, limb 2!/2 mm, tube 1!/2mm. Outer perianth lobes ovate, obtuse; lateral lobes crenate. Inner ones small, rather broad, Dec. 1948] obovate, obtuse. Tube swollen at the insertion of the stamens. Connective quadrangular, acute-api- culate at the apex. Stigmas rather large, soup-plate- shaped. Ovary obconical, swollen in the upper part, about 1!/2 mm. Capsule thick-ellipsoid, about 3-3'/2 mm, crowned by the short, cylindrical to conical, 2mm long, persistent part of the tube. Seeds brownish, fusiform, reticulate. Distr. Malaysia: New Guinea. Ecol. In forests, in humic soil, ascending to ca 700 m alt. 5. Gymnosiphon papuanus Becc. Malesia 1 (1877) 241; JonKEeR, Monogr. (1938) 174.—G. celebicum ScHLTrR. Bot. Jahrb. 49 (1913) 104. Stem simple or branched, 4-14 cm, colourless, bearing rather loose simple or bifid cymes of 3-many subsessile flowers. Leaves acuminate, up to 2!/2 mm. Bracts up to 3!/2 mm. Perianth whitish- purplish; limb about 1'!/2mm; tube up to 4!/2 mm. Outer perianth lobes ovate, triangular, obtuse; lateral lobes entire, involute. Inner ones small, linear. Connective deltoid, at the top 3-lobed, pro- vided with a forked thickening. Stigmas rather large, soup-plate-shaped, obtuse, cordate, auricu- late at the base. Ovary ovoid, about 1!/2 mm. Cap- sule obovoid to truncate-subglobose, 2!/2-5 mm long; crowned by the 2-2!/2 mm long, cylindrical persistent part of the tube. Distr. Micronesia (Palau Islands), in Malaysia: Celebes and New Guinea. Ecol. Moist forests, in humic soil, ascending to ca 500 m alt. BURMANNIACEAE (Jonker) 21 6. Gymnosiphon neglectus JoNKER, Monogr. (1938) Wis: Stem simple or branched, 7!/2-11 cm, bearing 1 or few flowers or a capitate inflorescence, consisting of contracted cymes. Leaves lanceolate-ovate, acu- minate, keeled, 1—2!/2 mm. Bracts to 5 mm. Flow- ers subsessile. Perianth dirty white; tube avout 2mm; limb about 2mm. Outer perianth lobes ovate, acuminate; lateral lobes crenate. Inner ones small, linear, acute. Stigmas funnel-shaped, curved, dorsally cristate. Ovary ovoid, about 2 mm. Cap- sule nearly globose, crowned by the 2 mm long persistent part of the tube. Distr. Malaysia: Java (Preanger Regencies and Semarang). Ecol. Moist forests, ascending to 1000 m alt. 7. Gymnosiphon pauciflorus SCHLTR. Bot. Jahrb. 49 (1913) 102; JonkER, Monogr. (1938) 176. Stem simple, colourless, 2!/2—-9!/2 cm, bearing 1-3 sessile or subsessile flowers. Leaves ovate, acumi- nate, 1-1'/2 mm. Bracts to 3 mm. Perianth whitish to purplish; tube about 4 mm; limb very short. Outer perianth lobes ovate, acute, about 1—-1!/2 mm; lateral lobes entire. Inner ones minute. Connective not apiculate, forked at the top. Style branches rather long, each bearing an ovoid, in transverse section somewhat triangular, stigma. Ovary obo- void, about 2 mm. Capsule ovoid, to 31/2 mm; per- sistent part of the tube 1!/2-2!/2 mm. Distr. Malaysia: New Guinea (Kani Mts), once collected. 3. THISMIA GRIFFITH, Proc. Linn. Soc. 1 (1844) 221; JONKER, Monogr. (1938) 42, 227. Saprophytic, fleshy herbs. Underground part in the Malaysian species coralliform or vermiform and creeping. Stems usually short, seldom branched. Leaves small, scale-like. Below the flowers 1 or more bracts, sometimes forming an involucre. Flowers actinomorphic or, sometimes, zygomorphic, urceolate to campanulate. Perianth lobes 6, occasionally free and of equal length and size, or inner ones larger, sometimes inner lobes connivent at the apex or connate in the apical part, then forming an erect mitre with 3 holes, in that case outer lobes very small. Stamens 3, free or, usually, stuck together to an anther tube, hanging at an annulus in the perianth throat. Filaments short, ribbon-shaped. Style thick, short, cylin- drical or conical, persistent, bearing at its apex 3 simple or bilabiate stigmas. Ovary obconical or obovoid; the 3 placentas inserted at the bottom or parietally in the basal part of the ovary. Fruit fleshy, cup-shaped, crowned by the persistent, fleshy, basal ring of the perianth tube and the style with the stigmas. Distr. 24 species, in tropical America (Sect. Myostoma and Ophiomeris), tropical Asia (Sect. Euthis- mia and Sarcosiphon), Chicago area, New Zealand and Tasmania (Sect. Rodwaya). 22 FLORA MALESIANA [ser. I, vol. 4! KEY TO THE SPECIES 1. Inner perianth lobes free, spreading or erect. Underground part vermiform, creeping 2. Perianth lobes equal in length and size 3. Flowers zygomorphic, geniculate 3. Flowers actinomorphic. 4. Stems several, flowers 4-6 inaraceme . . 4. Stem simple; flowers usually 1—3, terminal. 5. Leaves and bracts beset with distinct, prominent, blunt processes . 5. Leaves and bracts without processes. 6. Perianth lobes lanceolate, acute to acuminate, flat é 6. Perianth lobes triangular at the base, tapering into long, filiform tentacles. (Sect. Euthismia SCHLTR.) .(Sussect. Odoardoa SCHLTR.) Ses . 1. T. chrysops 2. T. racemosa 3. T. grandiflora 4. T. fumida 7. Perianth tube with horizontal bars inside. Stigmas lanceolate. 8. Anthers provided with 3 thick-filiform appendages at the free apical margin. Perianth yellow- ish in the basal part, bright orange-yellow in the upper part and limb. Tentacles bright orange-red at the base. Perianth lobes with tentacles up to 10 mm 5. T. aseroe 8. Anthers with 1 thick-filiform, median appendage and 2 Jateral, short teeth at the free apical margin. Perianth white with 6 ochre-brown streaks; lobes with tentacles c. 20 mm 6. T. alba 7. Perianth tube without bars. Apical margin of the anthers with 2 teeth, each bearing a globose body at the top. Stigmas funnel-shaped with prominent margin 2. Inner perianth lobes larger 7. T. ophiuris (Sussect. Brunonithismia JONK.) 9. Perianth very zygomorphic, bilabiate. Upper lip ‘fleshy, bent over the opening of the tube 9. Flowers actinomorphic. 8. T. labiata 10. Inner perianth lobes simple. Tube with prominent horizontal bars inside. 11. Anthers with 3 distinct teeth at the free apical margin, each tooth pesca a stiff hair. Outer perianth lobes broadly ovate, obtuse, erect . 11. Anthers slightly dentate apically. Outer perianth lobes short, ear- -shaped . . 9. T. javanica . 10. T. arachnites 10. Inner perianth lobes consisting of 3 parts. Basal part erect, short, bearing the transverse part, hamate at the base and broadened at the apex. Third part awl-shaped, inserted on the broad apex of the second part. Perianth tube without bars 11. T. neptunis 1. Inner perianth lobes connected at the apex to an erect mitre with 3 holes. Underground part coral- liform . (Sect. Sarcosiphon (Bu.)JONK.) 12. Inner perianth lobes linear, connate at the tips, “forming a mitre ‘with large holes. Anthers ciliate in the basal part, toothed at the apex. 13. Apical margin of the anthers provided with 2 constricted in the middle, below the thecae teeth, each bearing a stiff hair. Anthers slightly 12. T. clandestina 13. Apical margin of the anthers provided with 3 teeth, each bearing a stiff hair. Anthers constricted at the base, just above the thecae . 13. T. episcopalis 12. Inner perianth lobes spathulate, connate to a | mitre “with rather small holes. Anthers not ciliate, truncate at the apex . 1. Thismia chrysops RipL. Ann. Bot. 9 (1895) 323; JONKER, Monogr. (1938) 237. Stem usually simple and 1-flowered, about 15 cm. Leaves and bracts linear-lanceolate, acute more or less imbricate, up to 4 mm. Perianth tube geniculate; part below knee c. 3 mm, pink with longitudinal striae; the c.5 mm long, upper part and limb chocolate-brown; perianth mouth yellow. Perianth lobes lanceolate, about 7 mm, tapering to filiform tentacles. Annulus prominent, slightly 6-lobed. Anthers quadrangular, provided with a thick hair on both sides of the free, apical margin and a broad, wing-like appendage, inserted on the midline of the connective. Stigmas oblong, bifid. Fruit stalk elongate. Distr. Malaysia: Malay Peninsula (Malacca, Mt Ophir), once collected. 2. Thismia racemosa RIDL. J. Str. Br. R. As. Soc. 69 (1915) 13; JoNKER, Monogr. (1938) 238. Stems several, occasionally branched. Leaves linear, acuminate. Flowers 4—6in a raceme; pedicels 14. T. crocea 1—1!/2 cm long. Perianth lobes short, triangular- ovate, blunt. Annulus prominent. Tube cylindrical, about 6 mm long. Distr. Malaysia: Malay Peninsula (Pahang), once collected. 3. Thismia grandiflora RimpL_. Ann. Bot. 9 (1895) 324; JoNKER, Monogr. (1938) 239. Stem simple, 1-flowered, 3 cm, provided with 2 basal, opposite, scale-like lanceolate Jeaves, about 5 mm, and 2 apical bracts, of the same shape and size as the basal leaves; both leaves and bracts beset with stiff, terete, blunt processes on the back. Perianth urceolate; tube pink with longitudinal striae, 8 mm. Lobes patent, ovate-triangular in the basal part, tapering at the apex to filiform tenta- cles. Annulus prominent. Anthers not or scarcely stuck together, provided with 2 lateral teeth at the free apical margin and a wing-like appendage in- serted at the middle of the connective. Stigmas lanceolate, bifid, acute, papillose. Ovary about 4mm, obovoid, truncate. Dec. 1948] Distr. Malaysia: Malay Peninsula (Johore), once collected. 4. Thismia fumida Ripv. J. Str. Br. R. As. Soc. 22 (1890) 338; JoNKEerR, Monogr. (1938) 240. Stem slender, conspicuous, unbranched, about 10 cm, bearing 1-2 flowers. Leaves very small, ap- pressed, lanceolate, acute. Flowers up to 1 cm. Perianth lobes lanceolate, acute, greenish-grey, constricted above the ovary and broadened below the limb, white with pink stripes. Annulus pro- minent. Ovary obconical. Capsule cup-shaped, rib- bed and scabrid, crowned by the crenulate, basal ring of the perianth. Distr. Malaysia: Malay Peninsula (Singapore and Selangor). 5. Thismia aseroe Becc. Malesia 1 (1877) 252; JONKER, Monogr. (1938) 240.—Fig. 3. Stem simple or, sometimes, branched, 1- or 2- flowered, up to 8!/2cm high. Leaves few, lanceolate, obtuse, to 4mm long. At the base of the flowers an involucre, consisting of lanceolate bracts. Peri- anth obconic-campanulate, dirty-yellow in the basal part, bright orange-yellow in the upper part and in the limb. Perianth tube about 11 mm; the basal 5 mm inside with transverse bars. Lobes tri- angular, 3 mm long, ending in bright orange tentacles, red at the base, 6mm long. Annulus prominent. Anthers with 3 short thick-filiform ap- pendages at the free apical margin; in the lower part of the anther, inserted at the middle of the connective, a broad, dorsal quadrangular wing, wider than the anther. Thecae oblong, in the basal part of the anther; in the apical part 2 nectaries on the line of junction of one connective with the next. Ovary obovoid, 3 mm. Stigmas narrow-lan- ceolate, rather long, acute. Capsule ribbed, about 5 mm. Fruit stalk lengthening about 5-7 mm above the involucre. Seeds ellipsoid, ribbed. Distr. Malaysia: Malay Peninsula (Singapore and Perak). Ecol. In humic forests. 6. Thismia alba HoLTTUM, mis. A Th. aseroe differt antheris singulis munitis una tantum appendice mediana crassi-filiformi, porro margine apicali libera instructa dentibus 2 late- ralibus brevibus. Perianthium album signatumstriis 6 longitudinalibus ochraceo-brunneis; perianthii lobis tentaculis + 2 cm longis praeditis. Stem simple, 1- to 3-flowered, up to 10cm. Leaves few, 3—4!/2 mm long, lanceolate, acute or acuminate. At the base of the flowers an involucre, consisting of lanceolate, acute bracts. Perianth ob- conic-campanulate, white with 6 thin ochraceous- brown streaks, leading down from each perianth lobe, alternating with 6 thin yellow lines. Perianth tube about 10 mm, the basal part with transverse bars inside. Lobes triangular, 3-4 mm long, pale- yellow at the base, terminated by white, tentacles about 15 mm long. Annulus prominent, bright yellow. Anthers with 1 thick-filiform, median ap- pendage and 2 lateral short teeth at the free apical margin; in the lower part of the anther, inserted BURMANNIACEAE (Jonker) 23 at the middle of the connective, a broad, dorsal, quadrangular wing, wider than the anther. The- cae oblong, in the basal part of the anther; in the apical part 2 nectaries on the line of junction of one connective with the next. Ovary semi-globose, about 2 mm. Stigmas lanceolate, retuse, papillose. Capsule obconical, about 6 mm. Distr. Malaysia: Malay Peninsula (Pahang). 7. Thismia ophiuris Becc. Malesia I (1877) 252; JONKER, Monogr. (1938) 242. Stem 2—6mm, simple or branched, 1- or 2-flow- ered. Leaves lanceolate, obtuse, to 4mm long. Below the flowers an involucre of several, lance- olate, 3-4 mm long and 1 mm wide bracts. Perianth urceolate, yellowish brown. Tube about 9 mm; lobes triangular, terminated by long, filiform ten- tacles, about 13 mm. Annulus broad and thick. Insertion of the stamens broad, then narrowed into a ribbon-shaped filament and again broadened into the quadrangular anthers. Apical free margin of the anthers provided with 2 teeth, each with a globose body at the top. Ovary ovoid, about 5 mm. Style bearing 3 sessile, funnel-shaped, circumval- lated stigmas. Capsule ribbed; seeds oblong with longitudinal ribs; funicles about the same length as the seeds. Distr. Malaysia: Borneo (Sarawak & Br. N. Borneo). 8. Thismia labiata J.J.S. Bull. Jard. Bot. Btzg III, 9 (1927) 220; JoNKER, Monogr. (1938) 44, 243. Stem simple, 22 mm long. Leaves ovate, acute, appressed, 1!/2mm. Flowers with an involucre of 3 ovate-lanceolate, acute, 5'/2 mm long bracts. Pe- rianth urceolate in the basal part, bilabiate-zygo- morphous in the upper part. Outer perianth lobes 2!'/2 mm, broad-ovate at the base, rounded, witha subulate appendage inserted below the top. Inner lobes linear to filiform, subulate, 5mm. A thick, fleshy upper lip bent over the perianth mouth; on the back of the upper lip | inner and 2 outer peri- anth lobes. The other 2 inner lobes between the 2 lips. The third outer lobe inserted on the middle of the lower lip. Stamens rounded and ciliate at the free, apical margin; thecae elongate; outer side of the stamen provided with scattered hairs. The 3 stigmas connate to a capitate, 3-lobed stigma. Ovary obconical. Distr. Malaysia: Sumatra (Eastcoast Res.) once collected. 9. Thismia javanica J.J.S. Ann. Jard. Bot. Btzg 23 (1910) 32; Jonker, Monogr. (1938) 245.—Fig. 11. Stem simple or branched, up to 12 cm, 1- to 5- flowered. Leaves ovate or lanceolate-ovate, obtuse, 3 mm. At the base of a flower an involucre of 3 bracts. Perianth tube 7 mm, urceolate, whitish with 12 longitudinal, orange stripes, inside with longi- tudinal bars connected by many transverse bars. Outer perianth lobes obtuse, ovate; inner ones triangular, terminated in up to 3 cm long, filiform tentacles. Anthers 3-toothed at the free apical mar- gin; each tooth terminating in a hair. On the outer side of the anther, inserted in the middle, a qua- 24 FLORA MALESIANA [ser. I, vol. 4! drangular appendage, wider than the stamen. Margin of the appendage strigose. Style orange- coloured; stigmas sessile, ovate, truncate. Ovary obovoid, 3 mm. Capsule orange-coloured, about 6 mm. Seeds ellipsoid. Fig. 11. Thismia javanica J. J. S. Doengoes Iwoel, x 3/2. (LIEFTINCK) Distr. Malaysia: Sumatra and W. Java. Ecol. Shade of forests, on humus, 1000 m alt. Vern. Angkrek rambut (Java). Notes. Perhaps conspecific with the following species. below 10. Thismia arachnites RipL. Journ. Str. Br. Roy. As. Soc. 44(1905) 197; JONKER, Monogr. (1938) 247. Stem simple, 1—71/2 cm, bearing 1—3 flowers. Leaves few, lanceolate, acute, about 5 mm. At the base of the flowers an involucre, consisting of ovate-lance- olate, acute, 5—7 mm long bracts. Perianth urceolate to obconical, about 8 mm long, inside with longi- tudinal bars, connected by many transverse bars; tube transparent, white with 6, vertical, red streaks in the apical part. Perianth lobes pale red. Outer lobes very short, about 4 mm, ear-shaped; inner ones triangular, about 1 mm, terminating in up to 3 cm long, thin, filiform tentacles. Annulus prominent, yellow. Anthers slightly 3-toothed at the apical free margin, the lateral teeth somewhat larger than the median one, each tooth terminating in an indistinct, very thin, fragile hair. On the outer side of the anther, inserted in the middle, a quad- rangular strigose appendage, wider than thestamen. Style thick, conical; stigmas lanceolate, obtuse. Fruit stalk lengthening above the involucre. Distr. Malaysia: Malay Peninsula (Perak & Pahang). Note. Closely related to the preceding species, perhaps conspecific. 11. Thismia neptunis BeEcc. Malesia I (1877) 251; JONKER, Monogr. (1938) 43, 243. Stem 4-25 mm, simple, 1-flowered, beset with few, lanceolate, acute, about 3 mm long Jeaves. At the base of the flower an involucre of 3 lanceolate, acute, 4-6mm long bracts. Perianth tube urceolate, with 6 longitudinal stripes, 6 mm. Outer perianth lobes simple, recurved, filiform with triangular base, 4!/2 mm. Inner ones erect, about 15mm, con- sisting of an erect, short, basal part; a transverse part with hamate base and a broadened, rounded apex and, inserted on the apex of the transverse part, anerect, long, awl-shaped part. Annulus prom- inent. Anther quadrangular, 3-toothed at the free apical margin. Stigmas lanceolate, acute. Ovary obovoid, truncate, 1!/2 mm. Distr. Malaysia: Borneo (Sarawak, Mt Mat- tang), once collected. 12. Thismia clandestina (BL.) Mia. Fl. Ind. Bat. 3 (1855) 616; JoNKER, Monogr. (1938) 252.—Sarco- siphon clandestinus But. Mus. Bot. Lugd. Bat. 1 (1849) 65. Stem up to 5cm, 1—2-flowered. Leaves appressed, acuminate, to 5mm. At the base of the flower 3 bracts. Perianth tube urceolate, about 21/2 mm, greenish-grey, with 12 longitudinal, brownish- black stripes. Outer perianth lobes almost absent, inner ones connate to a 2!/2mm long, acuminate mitre. Annulus prominent, 6-lobed. Margin of the filaments and upper part of the anthers with short hairs. Thecae oblong, inserted on the margins of the anthers. Free apical margin of the anthers with 2 teeth, tapering to stiff hairs. Inserted on the midline a large, wing-like appendage, provided with bundles of hairs on the angles. Stigmas ovate, bilobate, papillose, whitish. Ovary obovoid, about 3 mm. Funicle about the same length as the ovules. Capsule papillose, about 5 mm. Distr. Malaysia: W. Java. Ecol. In humus of forests, ascending to ca 1000 m alt. 13. Thismia episcopalis (BEcCc.) F. v. MUELL. Pap. & Proc. R. Soc. Tasm. for 1890 (1891) 235; JoNKER, Monogr. (1938) 46, 253.—Geomitra episcopalis Becc. Malesia 1 (1877) 250.—Bagnisia episcopalis ENGL. Pfl. Fam. 2, 6 (1889) 48.—Sarcosiphon episco- palis Scutr. Notizbl. 8 (1921) 38.—Fig. 4. Stem simple or branched, 1—8 flowered, up to 19 cm. Leaves appressed, ovate, acute, 2-5 mm. Perianth tube urceolate, yellow with black stripes, 6-9 mm. Outer perianth lobes almost lacking; inner connate to a slightly acuminate, about 5 mm long mitre. Filaments constricted. Thecae diver- gent. Margin of the anthers and the winglike ap- pendage hairy, apical part of the anther, below the insertion of the appendage, darker coloured; free apical margin 3-toothed, each tooth terminated by a stiff hair. Stigmas bilobate. Ovary obovoid, 3 mm. Funicles about as longas the ovules. Capsule ribbed, about 3 mm. Fruit stalk lengthened. Dec. 1948] BURMANNIACEAE (Jonker) 25 Distr. Malaysia: Borneo (Sarawak, Br. N. Borneo). Ecol. In humus of forests, ascending to ca 1700 m alt. 14. Thismia crocea (BeEcc.) J.J.S. Nova Guinea 8,1(1909) 193; JoNKER, Monogr. (1938) 44, 251. —Bagnisia crocea Becc. Malesia 1 (1877) 249.— Thismia versteegii J.J.S. Nova Guinea 8,1 (1909) -193.—Sarcosiphon croceus SCHLTR. Notizbl. 8 (1921) 38.—Sarcosiphon versteegii SCHLTR. Notizbl. 8 (1921) 38. Stem simple, 1—3-flowered, about 6 cm. Leaves appressed in the basal part, lanceolate, acute, to 6 mm. At the base of the flowers 3 ovate lanceolate, acute bracts. Perianth tube urceolate, ribbed, red- dish-brown in the upper part, yellowish-orange in the middle and white at the base, about 6 mm. Outer perianth lobes broad, short, rounded; inner ones connate to a thick, 2mm long mitre with 3 narrow holes and 3 prominent midribs. Annulus slightly 12-lobed. Anthers quadrangular, not hairy; thecae oblong, parallel; inserted in the apical part of the anther a broad appendage with curled mar- gins. Stigmas ovate, obtuse, papillose. Ovary light reddish-brown, about 2mm. Funicles as long as the ovules. Fruit ribbed, obovoid. Fruit stalk thickened and lengthened after flowering to 16 mm above the bracts. Distr. Malaysia: West New Guinea. Notes. In Perak (Malay Peninsula), RIDLEY observed a Thismia, described by him in Mat. Fl. Mal. Pen. 2 (1907) 75, as Bagnisia crocea var. brunnea. This specimen was apparently not preserved; it is highly improbable that it belongs to T. crocea. 4. GEOMITRA BECCARI, Malesia 1 (1877) 250; JONKER, Monogr. (1938) 46, 254. Underground part unknown. Stem beset with scale-like Jeaves. Flowers rather large, with an involucre at the base. Tubular part of the perianth urceolate. Outer perianth lobes free, very small. Inner ones connate at the top to an erect mitre with 3 holes, crowned by 3 apical, long, thick-filiform, erect, clavately swollen appendages. Basal ring of the perianth tube thickened, persistent on the fruit. Throat margin of the perianth thickened to a 6-lobed annulus. Stamens 6, hanging at the annulus; anthers stuck together to a tube. Style short, cylindrical, fleshy, bearing 3 erect stigmas. Ovary with 3 stalked placentas; funicles short. Capsule cup-shaped, crowned by the persistent, basal perianth ring and the style. Distr. One species, known only from Borneo (Sarawak). 1. Geomitra clavigera BEcc. Malesia 1 (1877) 251; JONKER, Monogr. (1938) 46, 255.—Thismia clavi- gera F. v. MUELL. Vict. Nat. (1890) 235.—Sarco- siphon clavigerus SCHLTR. Notizbl. 8 (1921) 39. Stem simple, up to 12 cm, bearing about 3 flow- ers. Leaves lanceolate, acuminate or acute, 2-6 mm. Bracts lanceolate, acuminate, 6-7 mm. Perianth tube about 9mm. Outer perianth lobes erect, 3-5 mm, hooked at the apex. Filiform appendages 8-12 mm long. Anthers quadrangular; free apical margin with 3 teeth, each bearing a stiff, trans- parent hair. Anther tube about 4mm. Stigmas lanceolate, bilobate; lobes acute. Ovary obovoid, truncate, about 3 mm. Distr. Malaysia: Borneo (Sarawak), once col- lected. broadly triangular, about 1mm. Mitre about 5. SCAPHIOPHORA SCHLTR. Notizbl. 8 (1921) 39; JoNKER, Monogr. (1938) 46, 256. Roots coralliform. Stem provided with scale-like /eaves; at the base of the flower an involucre. Perianth tube urceolate. Outer perianth lobes small; inner ones narrow in the basal part, broadened at the apical part, connate to an erect mitre with 3 holes in the basal part. Mitre crowned by a long, stiff column, bearing at the top 3 lobes. Stamens 6, hanging, inserted at an annulus in the perianth throat. Filaments ribbon-shaped. Anthers stuck together to an anther tube; each anther provided with a wing-like appendage, inserted in the middle and broader than the anther. Placentas stalked; stalks inserted peripherically at the bottom of the ovary. Basal perianth ring and style persistent on the fruit. Distr. Two species, one in New Guinea, the other in the Philippines. 26 FLORA MALESIANA [ser. I, vol. 4', Dec. 1948] KEY TO THE SPECIES 1. Flowers 3-6!/2 cm long (without column). Column 1!/2-6 cm long, at the apex broadened to 3 fleshy, connate lobes . bodies . 1. S. gigantea 1. Flowers 1 cm long Guithout column). Column 5 mm long, bearing at the apex 3 cup-shaped 1. Scaphiophora gigantea JONKER, Monogr. (1938) 257.—Fig. 2. Stem 4-10!/2 cm, partly subterranean. Leaves lanceolate, acute, 2-4 mm, the lower ones keeled. Bracts ovate, lanceolate, acute, about 18 mm. Peri- anth tube 15-21 mm, pale rose-coloured with yellow veins, reticulate below the inner perianth lobes. Outer perianth lobes ear-shaped. Mitre 5-9 mm long, orange to yolk-yellow. Stamens about 7mm. Anthers prominently nerved; free apical margin provided with 3 median and 2 lateral teeth; each bearing a stiff, transparent hair. Appendix of the anther greenish-blue; lateral margins bearing 3 bundles of short hairs; apical margin pilose. Thecae divergent, ovate. Style truncate-conical. Stigmas sessile, obovate, 2-lobed, papillose outside and in the upper part inside. Fruit cup-shaped. Placentas connate at the apex, stalked; stalks about the same length as the placentas. Distr. Malaysia: Philippines (Luzon), twice collected. 2. S. appendiculata 2. Scaphiophora appendiculata (SCHLTR.) SCHLTR. Notizbl. 8 (1921) 39; JoNKER, Monogr. (1938) 259. —Thismia appendiculata SCHLTR. Bot. Jahrb. 55 (1918) 202. Stem 15-20 mm high, partly subterranean, usu- ally 1-flowered. Leaves ovate to lanceolate, 2-3 mm. Bracts lanceolate, acute, about 5mm. Perianth tube 6mm, yellowish white in the lower part. Outer perianth lobes small, ear-shaped. Mitre 3-6 mm, orange-coloured. At the base of each peri- anth lobe, on the inner side, a glandular, bowl-shaped body. Column + broadened towards the apex, bearing 3 thick, fleshy, cup-shaped bodies. Stamens about 3 mm. Appendage of the anther crenulate at the apical margin. Thecae divergent. Style trun- cate-conical, 1!/2 mm. Stigmas sessile, obovate, 2- lobed, 1 mm. Ovary 3!/2 mm. Placentas stalked, above the fertile part suddenly narrowed again into a filiform, apical appendage. Placentas attach- ed to the bottom of the ovary by the stalks and to the roof by the apical appendages. Distr. Malaysia: Northeast New Guinea, once collected. SPHENOCLEACEAE (H. K. Airy Shaw, Kew) Mart. ex LINDL. Nat. Syst. ed. 2 (1836) 238; DC. Prod. 8 (1939) 548; WiGur, III. Ind. bot. 2 (1850) 115; Mia. Fl. Ind. Bat. 2 (1857) 569; Boiss. Fl. Or. 3 (1875) 963. Annual (?)laticiferous herbs, with the habit of Phytolacca. Stem erect, somewhat succulent. Leaves spirally arranged, simple, entire, exstipulate. Inflorescences termi- nal, densely spicate, acropetal. Flowers subtended by a bract and two bracteoles, bisexual, actinomorphic. Calyx tube adnate to the ovary; segments 5, united below, imbricate, connivent, persistent. Corolla campanulate-urceolate, perigynous; lobes 5, imbricate. Stamens 5, epipetalous, alternating with the corolla lobes; filaments short; anthers rounded, 2-locular, dehiscing longitudinally. Ovary semi-inferior, 2-locular; style short, stigma capitate; ovules ~, attached to large spongy stipitate axile placentas. Capsule cuneate-obconic, 2-locular, membranous, circumscissile; seeds ~, minute, oblong, rugose-costate, albumen very scanty or none (?); embryo axile, straight, subterete. Distr. Mono-generic, almost pantropical. Ecol., Uses, Vern., see below under S. zeylanica. Notes. The maintenance of Sphenocleaceae as a separate family is abundantly justified; there is no evidence of affinity with Campanulaceae, with which it has hitherto been associated. The habit resembles that of Phytolacca, and the anatomy shows several significant features occurring in members of the Phytolaccaceae and related families. Other characters suggest Primulaceae, and provisionally it is sug- gested that the family represents a ‘half-way house’ between the families mentioned. From the Centro- Spermae it deviates in the semi-inferior ovary, gamopetalous corolla and straight embryo, and from the Primulaceae principally in the alternipetalous stamens. A separate note on the classification will be published in the Kew Bulletin. 1. SPHENOCLEA GAERTN. Fruct. 1 (1788) 113, t. 24, f. 5; Mig. Fl. Ind. Bat. 2 (1857) 569; B. & H. Gen. Pl. 2 (1876) 560; BAILL. Hist. Pl. 8 (1886) 327, 362, f. 158-161; SCHONLAND, in E. & P. 4, 5 (1889) 60; BoeRL. Handl. 2, 1 (1891) 257. For characters see family description. Distr. Two species, one pantropical, one endemic in W. Africa. 1. S. zeylanica GAERTN. Fruct. /.c.; Bl. Bijdr. 16 nivent. Corolla whitish, 2!/24 mm long, caducous, (1826) 1138; Morirzi, Syst. Verz. (1845-6) 66; segments ovate-triangular, obtuse or acute, united BLANCO, FI. Filip. ed. 2(1845) 62, ed. 3, 1(1877) 117, t. 143; Mia. l.c.; F.-VILL. Nov. App. (1880) 121; K. & G. Mat. Fl. Mal. Pen. no 16 (1905) 57; Koorp. Exk. Fl. 3 (1912) 301; Merr. Fl. Man. (1912) 462; Sp. Blanc. (1918) 374; En. Philip. 3(1923) 588; RIDL. Fl. Mal. Pen. 2 (1923) 204; BAcKer, Onkruidfi. Jav. Suik. (1931) 742; OcHsE & BAKH.V.D. BR. Veg. D. E. I. (1931) 93, f. 55, 349._—Pongatium spongi- osum BLANCO, FI. Filip. (1837) 86.—Reichelia palustris BLANCO, I.c. 220; ed. 2 (1845) 155; ed. 3, 1 (1877) 277, t. 143.—Fig. 1. Roots long, cord-like. Stem hollow, 7-150 cm. Leaves oblong to lanceolate-oblong, attenuate at both ends, acute or obtuse, glabrous, 2!/2-12'/2 by 1/2-5 cm; petiole 3-30 mm. Spikes 3/4—7!/2 cm long, cylindric; peduncle 1-8 cm. Bracts and bracteoles + spatulate, the green apices arched over the calyx before and after anthesis. Flowers crowded, thomboid or hexagonal by compression, sessile, wedge-shaped below, attached longitudinally to the tachis by a linear base. Calyx segments deltoid- semicircular, obtuse, ultimately accrescent and con- slightly more than half-way, connivent. Stamens inserted half-way up tube of corolla, filaments slightly dilated at base. Ovary obovoid, 2!/2 mm long, apex broad, free, truncate. Capsule 4-5 mm in diam., dehiscing below the calyx segments which fall with the lid, leaving the scarious persistent base. Seeds yellowish-brown, + '/2 mm long. Distr. Trop. America (introduced), trop. Africa (incl. Madagascar) (indigenous; cf. BENTH. in Journ. Linn. Soc. Bot. 15 (1875) 13), SW. Persia to Turkestan, India and Formosa (prob. intro- duced), in Malaysia (prob. introduced): Malay Peninsula (scarce, mainly in the prov. Kedah and Wellesley), 7?Sumatra, Philippines (Luzon, Biliran, Negros), Java, Bali, SW. Celebes and Timor. Ecol. A weedy annual occurring in almost any kind of damp ground at low alt. up to 350 m: river banks and dry riverbeds, damp marshy or periodic- ally inundated depressions, seasonal swamps, sides of ponds, ditches, and stagnant water generally, especially rice-fields, both in continuously rainy and in seasonal climates. Almost every flower on 28 FLORA MALESIANA every inflor. sets fruit; only one or two flowers are open at once on any one head. In Malaysia never gregarious, nor growing on mud of tidal creeks, as in Africa. Uses. In Java young plants and tips of older plants are steamed and eaten with rice; they have a slightly bitter taste; leaves are sold under the name goenda padi. Vern. Java: goenda, M, J, Sd, g. rawah, g. lalab, g. padi, g. sapi, Sd, goendha, Md, gondo, J; Bali: gonda; Celebes: gangang karaéng, Mk., gonra, Mk, Bg; Philippines: mais-mais (Panay, Bisaya), silisi- [ser. I, vol. 4', Dec. 1948] lihan (Tagalog); the Javanese names are also ap- plied to the superficially similar Hydrophyllaceous Hydrolea zeylanica (L.) VAHL. Notes. The plant is described as laticiferous but METCALFE reports that ‘typical laticiferous ca- nals are absent from the phloem, although occa- sional elongated cells have been observed in this tissue with granular contents which may represent coagulated latex’. Miss M. C. VREEDE, Anatomist in the Treub Lab., Buitenzorg, Java, reported, July 6, 1948, that in fresh material she could find neither milky juice nor laticiferous elements. a\4 >” Fig. 1. Sphenoclea zeylanica GAERTN. X 1/4. A rich-flowering individual. NYSSACEAE! (J. Wasscher, Groningen) 1. NYSSA LINNE, Sp. Pl. (1753) 1058; Gen. Pl. ed. 5 (1754) 478; WasscHeErR, Blumea 1 (1935) 343.—Agathisanthes & Ceratostachys Bl. Bijdr. (1825) 644; Mia. Fl. Ind. Bat. 1, 1 (1856) 838.—Agathidanthes Hassk. Cat. Hort. Bog. (1844) 254.—Daphniphyllopsis Kurz, J. As. Soc. Beng. 44, II (1875) 201. Dioecious trees or shrubs. Leaves simple, scattered. Stipules 0. Flowers uni- sexual, often in heads, in the axils of a bract and with 2 bracteoles. d: in axillary heads or short racemes; calyx entire or 5-toothed; petals 5, imbricate, often small, alternate with the calyx; stamens 8-16 in 2 alternating whorls; anthers small, dorsifixed with lateral lengthwise slits; disk pulvinate; style rudimentary. 9: soli- tary, axillary or in 2-10-flowered heads; ovary inferior, l-locular, connate with the 5-toothed or entire calyx; petals 5-8 often minute; stamens of inner whorl partly sterile, both petals and anthers soon dropping; style with 2 appressed later diver- gent often torulose branches stigmatose on their inside, brittle, often deficient in the herbarium. Ovule 1, hanging from the apex of the cell, anatropous with 2 integu- ments. Fruit drupaceous ovoid to oblong. Distr. Ca 6 spp., 4 in Atlantic N. America, 1 in China, 1 from India to W. Malaysia. Ecol. The American spp. mostly in swamp forests, the Asiatic one not so. Notes. The flowers are often deficient in the herbaria. The polymorphy of N. javanica suggests that perhaps more than one species is present in Malaysia. 1. Nyssa javanica (BL.) WANG. Pfl. R. 41 (1909) 15; WASSCHER, Blumea 1 (1935) 344.—Ceratostachys arborea BL. Bijdr. (1825) 644; Mia. FI. Ind. Bat. 1, 1 (1856) 839.—Agathisanthes javanica BL. Bijdr. (1825) 645; Mia. FI. Ind. Bat. 1, 1 (1856) 839.— Agathidanthes javanica HAssK. Cat. Hort. Bog. (1844) 254.—WNyssa sessiliflora Hook. f. & Th. Gen. Pl. 1 (1867) 952.—Ilex daphniphylloides Kurz, J. As. Soc. Beng. 39, II (1870) 72.—Daphniphyllopsis capitata KuRzZ, l.c. 44, II (1875) 201; For. Fl. Burm. 1 (1877) 240.—Nyssa arborea Koorp. Exk. FI. Jav. 2 (1912) 731.—Nyssa bifida Crap, Kew Bull. (1913) 69.—Fig. 1. Dioecious tree up to 40m, 30-100 cm diam., clear bole 13-23 m, buttresses mostly absent. Twigs tomentose, glabrescent. Leaves rather den- sely set, oblong-lanceolate to obovate, rarely sub- ovate, base acute, apex abruptly acuminate, cori- aceous, entire, sparsely hairy to tomentose on midrib and nerves beneath, further glabrous, 5—23 by 2!/2-8 cm; in seedlings the Ist pair of leaves is opposite; nerves 8-11 pairs; petiole 1—-3!/2 cm long, flat or slightly sulcate, hairy or glabrous. Flowers pallid, in pedunculate nearly globose axillary heads 12-18mm diameter; peduncles flat- tened towards the apex 3/4s—-5 cm long, their apex 2-5 mm broad, glabrous or hairy, ca halfway with 1-2 sessile small acute bracts 3-4 by 1 mm. Re- ceptacle globose to ellipsoid, flattened, 2-3 and 4-5 mm. Flowers enveloped by 1 bract and 2 half- way connate bracteoles, all broad-ovate, sericeous- ciliate, 2-2!/2 by 1!/2-3 mm, in Q persistent.—d: Flowers 20-40 capitate, !/2-4 mm pedicellate; calyx teeth 4-5 rounded, ‘/2—-/4 by 1-1'/2 mm, outside appressed-hairy, ciliate; petals 4-5 free, ovate with broad base, curled back, 3—5 by 1'/2-3 mm, both sides very short spreading hairy; stamens 8-10, those of outer whorl 3—5 of inner 24 mm long; anthers elliptic 1!/2 by 1 mm, outer loculi often larger than inner ones; disk !/2-1 by 1-2 mm, 8-10 lobed.—9o: Flowers usually 3-8 rarely up to 18, sessile; calyx campanulate 2-3 by 1!/2 mm, densely appressed-sericeous; lobes 4—5 irregular, rounded, 1/2-] by 2'/2 mm or almost absent; petals 4-5 as in 6 but smaller 3-4 by 2!/2-3 mm; stamens 8-10, probably of inner whorl at least sterile, smaller than in d; style 11/2-2 by 1/2-1 mm, with 2 divergent (in bud one branch longer and incurved over the other), or curled branches 1—2 mm long. Fruit ellipsoidal, little flattened, 11/2-2!/s by 4-1!/2 cm, 3/4-1'/2 cm thick, crowned by the disk & calyx limb 1 by 2 mm, immature yellow, ripe purple. Exocarp coriaceous glabrescent, mesocarp spongious-fleshy. Stone flat- tened obovate, acute 1-2 by 4/2-1'/4 cm, 2-6 mm thick, on one side with 5 length-grooves, the other side with few tubercles above the middle and a length-keel. Distr. SE. Asia and Malaysia: Sumatra, Mal. Peninsula, Borneo, Java, 600-1600 m alt. (in Siam and Sumatra once at 100 m, in the Himalaya ascending to 2400 m acc. to Hook. f). Ecol. Common or scarce, never gregarious, in mixed evergreen mountain forests both in ever-wet and periodically dry regions (E. Java), not in sec- (1) The family consists of 3 genera, 2 of which are endemic in Central Asia. Nyssa occurs from Malaysia to Asia and N. America. Formerly the family was included in the Cornaceae sens. ampl. 30 FLORA MALESIANA [ser. I, vol. 4! Fig. 1. Nyssa javanica (BL.) WANG. Twig, x 2/3, a. male flower, < 6, b-d. female flowers, x 6, e. galled fruits, x 2/3, f. fruit, x 2/3, g. seed, Xx 2/3. Dec. 1948] NyYSSACEAE (Wasscher) 31 ondary forest, mostly fl. Jan.—May, fr. July—Dec. The thick end-bud produces young pale brown-red foliage and flowers in the rainy season. Fruits often deformed into worm-shaped galls. Vern. Malay names in Sumatra mostly ‘me- dang’ with some additional name (also common for Laur.), in Java hiroeng, or kiroeng, Sd; Javanese names very variable. Uses: Rather heavy dense wood not highly esti- mated. Bark grey, smooth, dingy yellow in section. Fruits are said to be edible and have a sweet odour, but a bitter acid taste. Notes. The fruits are often deformed to a gall on which BLumeE based his Ceratostachys arborea. The variability in the flowering parts, their early dropping, and the brittleness of the flowers in the herbarium have caused many discrepancies in literature. Excluded Nyssa hollrungii K. Scx. Nachtr. Fl. D. Schutzgeb Siids. (1905) 334 = Alangium javanicum (BI.) WANG. var. papuanum (MANsF. & MELCH.) BLoemsB. Blumea 1 (1935) 284. SARCOSPERMACEAE (H. J. Lam, Leyden) 1. SARCOSPERMA Hook. f. in B. & H. Gen. Pl. 2 (1876) 655; RipL. Fl. Mal. Pen. 2 (1923) 260; H. J. Lam, Bull. J. B. B. III, 7 (1925) 248; Ic. 8 (1926) 18; Philip. J. Sc. 49 (1932) 143; Blumea 3 (1938) 183; /.c. 3 (1939) 261; I.c. 4 (1941) 322.—Bracea KING, J. As. Soc. Beng. 64, II (1896) 101.—Apoia Merk. Philip. J. Sc. 17 (1920) 605. Trees or shrubs, at least two spp. laticiferous. Leaves simple, entire, subopposite or opposite, rarely subverticillate; often with some alternate ones between, penni- nerved; petiole sometimes with auricles at the top; blade often with glandular pits in the axils of the secondary nerves or scattered on the undersurface; tertiary nerves slender but conspicuous, transverse and usually crowded, more or less perpen- dicular to the midrib. Stipules small, caducous. Flowers bisexual, in small fascicles or solitary, placed along racemose or more or less broadly paniculate axillary shoots; bracts minute deltoid. Sepals 5, quincuncially imbricate, two inner ones with scarious margins. Corolla infundibuliform, tube short, slightly thickened; lobes spreading, imbricate in bud. Staminodes 5, alternipetalous, inserted in the throat. Stamens 5, epipetalous; filaments short, connate with the base of the petals; anthers basifix, slightly extrorse, 2-celled, longitudinally dehiscent. Ovary su- perior, 1—2-celled, glabrous, contracted into a short stout style; cells with 1 apotro- pous, ascending ovule, attached to the basis of the central axis; stigma truncate, capitate or faintly 2-lobed. Fruit drupaceous, 1-(2)-seeded, ovoid to oblong; peri- carp thin. Seeds with a thin-crustaceous pale dull testa. Hilum small, round, basal; albumen absent; cotyledons thick; radicle inferior. Distr. 6 spp. of this mono-generic family occur in SE. Asia and Malaysia. Ecol. Scattered in mixed forests from the lowland up to + 1100 m. Uses. The wood of the moderate-sized S. paniculatum is rather soft and not durable; in Sumatra it is used as timber (HEYNE, Nutt. Pl. (1927)1245); Marco described it in detail anatomically (Trop. Woods 5 (1933) 1). Notes. The genus is closely allied to the Sapotaceae; its wood anatomy is only slightly different. Herbarium specimens are often not recognized and confused with other families. KEY TO THE SPECIES 1. Apex of the petiole with distinct auricles. Leaves oblong to elliptic, 6-28, by 3!/2-10 cm, dark brown s.s; glandular pits scattered on undersurface; secondary nerves 6-11. Ovary 2-celled 1. S. paniculatam 1. Auricles absent. Leaves rather broad, 11-36 by 3-13 cm, slightly Eater below, light brown s.s.; glandular pits absent; secondary nerves 7-16. Ovary 1-celled oe : 2. S. uittienii 1. Sarcosperma paniculatum (KING) STAPF & KING, _ thickish, waxy yellow to pale greenish white, fascicl- Ic. Pl. 7 (1901) t. 2690; Lam, //.cc.—Bracea panicu- lata KING, l.c.—Discocalyx macrocarpa ELMER, Leafl. 8 (1915) 2781.—Apoia macrocarpa MERR. 1.c.—Sarcosperma breviracemosum H. J. LAM, Bull. J. B. B. III, 8 (1926) 21. Laticiferous tree, 12-27 m, unbranched bole 6-17 m, 20-50 cm diam., crown spreading. Leaves glabrous, opposite or scattered, oblong to elliptic, base acute to ++ acuminate, apex gradually bluntly acuminate, 6-28 by 3!/2-10 cm, auricles acute 1/22 mm long; petiole 1—2!/2 cm. Stipules subulate, glabrous, ca 4 mm long. Inflor. glabrous or slightly tomentose, either paniculate, 1-14 cm long with branches 2-9 cm, or hardly branched 8-16 mm long; bracts glabrous, acute, 1 mm long. Flowers ed or solitary; pedicels 1—-1!/2 mm. Sepals round- ish or broadly acute 2 mm through. Corolla tube 1 mm, lobes ovate 3 by 2 mm. Staminodes acute 1 mm. Stamens ovoid 1 mm through. Ovary 2 by 1 mm; style 1 mm. Fruit ovoid, 17—20 by 17-15 mm, 1-(rarely 2-) seeded, 3 mm stalked; hilum 3 by 4mm. Distr. Malaysia: Mal. Peninsula (Perak), N. Sumatra, Br. N. Borneo, Mindanao, S. Celebes, Ternate, Flores, and E. New Guinea. Ecol. Scattered in dense or open mixed rain- forests, rarely in forest borders, among bamboos, or in secondary jungle, in the Malay Peninsula below 250 m, elsewhere ascending to 1100 m. FI. and fr. irregularly. 33 SARCOSPERMACEAE (Lam) Dec. 1948 a ae Suse Cae IN SAWN \ \ b. bud, c. flower diagram, d. part of ering branch, x 1/2, erma uittienii H. J. L. a. flow Fig. 1. Sarcosp d cross-section. e. ovary in longitudinal an the corolla and calyx within, 34 FLORA MALESIANA [ser. I, vol. 41, Dec. 1948] Vern. Not:constant, few noted. Notes. It is probable that more specimens are hidden among indeterminates in several families. 2. Sarcosperma uittienii H. J. LAm, Bull. J. B. B. III, 8 (1926) 19, f. 1, &c.—S. sumatranum UITT. ex Lam, /.c.—Fig. 1. Tree. Leaves fairly opposite, oblong-elliptic to ovate or obovate, both base and apex acuminate, glabrous above glabrescent below; petiole 12—20 mm. Inflor. densely minutely tomentose, generally broadly and laxly paniculate, sometimes almost unbranched, 33/4s—-131/4 cm long, branches 1!/2—63/4 cm long; bracts tomentose, deltoid 1-2 mm long. Flowers fascicled or solitary only known in bud; pedicals 2-4 mm. Calyx densely tomentose, 21/2 by 2mm. Corolla tube !/2 mm long, lobes obovate, 2-21/2 mm. Staminodes deltoid !/2 by '/3 mm. Sta- mens ovoid, | mm through. Ovary glabrous, 2!/2 by 1'/2 mm. Style 1 mm. Fr. unknown. Distr. Malaysia: only known from Sumatra (Eastcoast Res.). Ecol. Forests, ca 500 m. Fl. June—July. Vern. Only once noted. Notes. Inadequately known. Closely related to S. kachinense (KING & PRAIN) EXELL from Burma & China, and to S. arboreum Hook. f. from India to China. Excluded Sarcospermum petasites REINW. eX DE VRIESE, Rein- wardt’s reize (1858) 576= Gunnera macrophylla Bu. (Halorrh.). STACKHOUSIACEAE (F. I. Brouwer, Groningen) STACKHOUSIA J. SMITH, Trans. Linn. Soc. Lond. 4 (1798) 218; PAmp. Bull. Herb. Boiss. II, 5 (1905) 912; Brouwer, Blumea 3 (1938) 173; MatTrTr., in E. & P. ed. 2, 20b (1942) 240. Annual, or perennial herbs with a rhizome. Leaves scattered, entire. Stipules 0 or very small. Racemes terminal. Flowers bisexual, regular, 5-merous, in groups in the axils of bracts. Sepals usually more or less connate, rarely free. Corolla perigynous or almost hypogynous, petals long-clawed, rarely entirely free, usually free at the base, connate in the upper portion of the claws, lobes imbricate spread- ing. Stamens 5, inserted on the margin of the calyx tube, free, usually unequal (2 shortest), included in the corolla tube. Ovary (2-)3(-5) celled, lobed, each cell with | erect ovule. Style with (2-)3(-5) stigmatic lobes, partly sunk in the ovary. Fruit with (2-)3(-5) one-seeded cocci and a columella. Distr. Ca 19 spp. in Australia, 4 in Tasmania, 1 in New Zealand and 1 in Malaysia, Australia and Micronesia (Palau, Jap). Notes. The family consists next to the genus Stackhousia, the single one by which it is represented in Malaysia, of 2 other monotypic genera, and is practically confined to Australia. It is not directly allied to any other family and has been compared with e.g. Euphorbiaceae, Celastraceae, Sapindales, &c. 1. Stackhousia intermedia F. M. BAILEy, Q. Agric. J. 3, 4 (1898) 174; Q. Fl. (1899) 264; Pamp. l.c. 1149, cum f. philippinensis; BROUWER, Blumea 3 (1938) 174; STEEN. J. Arn. Arb. 28 (1947) 422. —S. muricata (non LINDL.) auct. plur. quoad Philip. —St. viminea (non J. SM.) VOLKENS, Bot.Jahrb. 31 (1902) 467; id. var. micrantha LautsB. Nachtr. FI. Deut. Sch. Geb. Siids. (1905) 305.—St. tenuissima var. ramosa STEEN. Nova Guinea 14 (1927) 307.— Fig. 1—2. Erect, glabrous annual, 6—50 cm long. Root fusi- form, up to 5 cm long, 1!/2 mm diam. at the base, attenuate, with fibrous ramifications. Stem gradu- ally attenuate to the almost filiform angular apex, little branched and leafy below, terete, striate, inter- nodes !/2-3 cm long. Leaves linear, sessile, base attenuate, 7!/2—20 by !/s—2 mm, lower obtuse, upper acute to mucronate, nervation absent or midrib visible. Racemes 1—20cm long. Flowers minute yellow, upper groups |-3 fls and 2 bracteoles, low- er groups with more bracteoles and up to 5 fils. Bracts roundish ovate, strongly acuminate, fim- briate, dentate, 3/4s-1 by !/2 mm, membranous ex- cept the midrib. Bracteoles like the bracts but more dentate and less acuminate. Pedicels 3/4—11/4 mm. Calyx-tube '/2mm high, lobes ovate-acuminate, 1/2 mm long, irregularly fimbriate-dentate, margin membranous. Corolla inserted on the margin of the calyx-tube, sympetalous, hypocraterimorphous, tube cylindrical, 2 by !/2 mm, divided into 5 petals in the lower portion over !/4 mm, lobes ovate-ob- long, strongly acuminate, ca 3/4 mm long. Fila- ments filiform, 2 shorter ones reaching the middle, 3 longer ones the margin of the corolla-tube; anthers oblong, very obtuse and emarginate at base and apex, 0.6 by 0.3 mm, introrse, dithecic, 4-lo- cular. Ovary subglobose, 0.3-0.4 mm diam. 3-lo- bate, 3-celled. Style straight, 0.4 mm long, with 3 linear stigmas. Cocci 3, roundish ovate, 1'/2 by 1 mm, reticulate. Distr. Australia, Micronesia, and Malaysia: Sumatra (Toba-Batak Lands), N. Celebes, Philip- pines (Luzon, Culion, Guimaras), Moluccas (Boe- roe, Ambon, Saparoea), New Guinea, 10—100—300— 600-1500 m alt.—Fig. 1. Fig. 1. Localities of Stackhousia intermedia BAILEY in Malaysia. Ecol. Lank herb mostly in grassfields, savan- nahs and abandoned fields, in both wet and dry spots, in Sumatra at 600-1400, but in E. Malaysia & Micronesia below 300m, in the Philippines ascending to 1500m. F/. mostly in April-May together with the grasses. Notes. St. tenuissima, virgata, aphylla and mi- crantha Pamp. l.c. are most probably all identical with this species. 36 FLORA MALESIANA (ser. I, vol. 4!, Dec. 1948] ‘ \ Fig. 2. Stackhousia intermedia BAILEY, X 1/2, fruits and flowers enlarged. ACTINIDIACEAE (C. G. G. J. van Steenis, Buitenzorg) 1. ACTINIDIA LINDL. Nat. Syst. ed. 2 (1836) 439; B. & H. Gen. Pl. 1 (1862) 177; BENTH. FI. Hongk. (1861) 26; KiNG, Ann. R. Bot. Gard. Calc. 5, 2 (1896) 145, t. 176; E. & P. ed. 2, 21 (1925) 36. Trailing shrubs or lianas without special organs for climbing, branches rarely flexuose; stem with wide vessels, raphides in the flowering parts: bark often with short linear lengthwise lenticels. Growth in flushes from terminal and axillary buds. Indumentum of stellate or simple hairs. Stipules minute, obsolete, or absent. Leaves simple, scattered, petiolate, serrate or callous-dentate, penninervous, midrib sulcate, veins in cross-bars, veinlets reticulate. Inflor. lateral, often on a common peduncle forked at the apex, cymose, often pseudo-umbellate; bracts 2, at the apex of the peduncle. Flowers mostly white, dioecious (or polygamous), 5(—4)— merous. Sepals distinctly imbricate (rarely valvate), free or subconnate at the base, persistent. Stamens (10—)«, in 9 fls with short filaments and small sterile anthers; filaments thin, anthers versatile, base divaricate, attached in the middle, reflexed in bud, dehiscing lengthwise. Disc absent. Ovary free, superior, tomentose (or glabrous), (5—) ~-celled; ovules attached on the central axis. Styles free, (5-)~, persistent, elongating after flowering in 9, +clavate, spreading, in d ovary small, with minute styles. Berry glabrous (or hairy), often spotted by lenticels, oblong. Seeds ~, small, biconvex, oblong, immersed in pulp; testa cartilagineous, reticu- late-pitted, dark when dry; albumen copious; integuments 1; embryo cylindrical straight, cotyledons short. Distr. Ca 30 spp. from W. Malaysia & Himalaya to Sachalin, Japan and Formosa, centering in China and Japan. Ecol. Forests and forest borders, in the montane zone mostly. Notes. Both Malaysian species appear to be strictly dioecious; the number of dg and 9 sheets in A. callosa is about equally large; on Mt Kinabalu only 9 have been found of A. /atifolia. The total number of specimens examined is inconsiderable; the species are either rare or little collected being inconspicuous. The genus Actinidia is often included in Theaceae, Dilleniaceae, or even Ericaceae, and it is closely related to Saurauiaceae from which it differs in its trailing or climbing habit, absence of scale-like emer- gences (except in A. strigosa), mostly dioecious fis, ebracteate pedicels, lengthwise dehiscing anthers, numerous styles, and a multilocular ovary. I wish to express my sincere thanks to Mr H. K. Atry SHAW and to Mr M. R. HENDERSON for verifying the MS. of this contribution with the materials preserved at London and Singapore respectively. KEY TO THE SPECIES 1. Leaves either glabrous or subglabrous, or provided with simple pluri-celled hairs. Petals glabrous. Inflorescences short 1. A. callosa 2. Leaves glabrous or subglabrous var. callosa 2. Leaves rather distinctly subtomentose beneath var. pubescens 1. Leaves stellate-tomentose beneath. Petals pubescent on the back. Inflorescences often well-developed. Peduncle 1!/2-8 cm 1. Actinidia callosa LINDL. Nat. Syst. ed. 2 (1835) 439, s.l.; K. & V. Bijdr. 3 (1896) 280; BACKER, Schoolfi. (1911) 102; Dunn, J. Linn. Soc. 39 (1911) 405; Koorp. Exk. Fl. 2 (1912) 602; Fl. Tjib. 2 (1923) 179; BaKeEr, J. Bot. (1924) Suppl. 9; STEEN. Bull. J.B.B. III, 13 (1934) 174.—See further under var. pubescens. Rambling or trailing shrub or liana up to 30 m, twig-lenticels distinct, wood and inner bark orange. Petiole red s.v.,1-4cm, blade rather variable in shape ovate-elliptic or obovate, acuminate, midrib 2. A. latifolia red s.v., 5—10!/2 by 2!/2-6 cm,sidenerves ca 5-6 pairs rather steeply ascending and substraight, insertion decurrent, margin distinctly serrate or dentate, teeth erect at the end of a vein, apex acuminate, base rounded to subcuneate. Indumentum meagre or absent, consisting of short often somewhat crisped pluri-celled simple hairs. Peduncle, pedicels and calyx thin-tomentose. Peduncle '/4-1'/2 cm, pedicels !/2-1!/s cm, all thin. Dioecious, flowers white, anthers yellow. Sepals ovate-orbicular, ca 6 by 5 mm. Petals oblique-broad-spathulate, sub- 38 FLORA MALESIANA [ser. Diveks Fig. 1. Actinidia latifolia (GARDN. & CHAMP.) MERR., habit x 1/2 (after KING). 4! Dec. 1948] ACTINIDIACEAE (v. Steenis) 39 fleshy, margin + irregular, ca 10 by 7 mm. Stamens © in ca 2 rows, filaments subequal, ca 6 mm (in 9 very short); anthers 17/3 by 1 mm, apex subapiculate (in Qsterile, hardly dehiscing); 9 fls unknown to me. Ovary stout cylindric, styles ca 2mm (in 6 very small, reduced). Berry grey-green, spotted grey or brown, entirely syncarp, oboval to broad-ellip- tic, often oblique, apex often concave, 17-27 by 14-18 mm, base rounded, sepals recurved. Seeds elliptic, 3 by 1'/2 mm. Distr. SE. Asia, China, Formosa, in Malaysia: Sumatra, Java. Ecol. Mountain forests, forest borders, 1000— 2040 m, rather rare. Notes. Young shoots edible. Leaves sometimes with -raspberry-coloured zoocecidia consisting of crowded-hairy portions. In Java a juvenile shoot was collected with subcordate subglabrous leaves resembling in shape those of A. Jatifolia. A rather variable species; some of the forms dis- tinguished by DUNN are now taken up as species, wrongly itseems. A. indochinensis MERR. apparently belongs here. var. pubescens DuNN, /.c. 406.—Saurauia tomen- tosa KORTH. nomen ex K. &. VY. Bijdr. 3 (1896) 280.—Actinidia pubescens RID. J. Fed. Mal. Stat. Mus. 8, 4 (1917) 18.—Leaves 6!/2-11!/2 by 4!/2-61/2 cm, thinly tomentose beneath. Distr. Assam, in Malaysia: Malay Peninsula (HENDERSON 23436), Sumatra (KORTHALS, FORBES). Notes. Apparently rare, may be confused with A. latifolia. The Sumatra specimen has glabrous twigs, the others hairy ones. The indumentum seems partly caducous. I assume KORTHALS’s specimens came from Sumatra, not from Java. 2. Actinidia latifolia (GARDN. & CHAMP.) MERR. J. Str. Br. R. As. Soc. 86 (1922) 330.— Heptaca latifolia GARDN. & CHAMP. in Hook. J. Bot. & Kew Gard. Misc. 1 (1849) 243.—Kadsura pubescens Mia. Fl. Ind. Bat. Suppl. (1860) 620; Kurz, J. As. Soc. Beng. 45, Il (1876) 119, non A. pubescens RIDL. 1917.—A. championi BENTH. Fl. Hongk. (1861) 26; FINET & GAGN. FI. Gén. I. C. 1 (1907) 28; Rip-. Fl. Mal. Pen. 1 (1922) 206.—A. miquelii Kina, J. As. Soc. Beng. 59, Il (1890) 196, nomen illeg.; Ann. R. Bot. Gard. Calc. 5 (1896) 145, t. 176.— Fig. 1. Rambling shrub or liana to 20m long, twigs dark-coloured s.s., innovations, inflor. and under- surface of the /eaves thinly cinnamon- (s.y. rusty- red-)stellate-tomentose. Petiole 2-4 cm; blade broad-ovate, obovate to suborbicular, 5!/2-11 by 3-9 cm, base reniform-cordate to rounded or cun- eate, apex acuminate, margin subentire with small callous teeth, veins rusty in distinct cross-bars, reticulations below hidden by a pale closed indu- mentum, upper surface puberulous. Peduncle rather stout, -- remote from the petiole, 1!/2-8 cm long, apex forked, + pseudo-umbellate, rich-flow- ered, pedicels in fr. apparently enlarging. Flowers velvety, light-brown, yellow inside, stamens yellow (ex coll.). Only seen Q buds, these depressed-glo- bose. Sepals tomentose outside. Petals pubescent outside, apex imbricating, basal parts free, blunt, rather roundish, pale green in bud apparently smaller than in A. callosa. Anthers numerous + 1 mm long, on !/2-4/4 mm long filaments, sterile hardly dehiscing. Ovary depressed-globose, densely pilose, 1'/2 mm high. Styles co, + 2 mm long, slen- der-clavate, overtopping flatly the anther clump. Berry acorn-shaped, 3-4 by 2 cm, brown, spotted pale. Seeds broad-elliptic, + 13/4-2 by more than | mm. Distr. China, Indochinese Peninsula, Hong- kong, ?Formosa, Hainan, in Malaysia: Malay Peninsula, Sumatra, Borneo. Ecol. Hill forests, rather rare, ca 900-1500 m, fi. April—July. Vern. Once noted, S. Sumatra, wait boerah. Notes. There is some variability in the size of the inflor. 4. formosana HAYATA probably belongs here. Expected to occur in the Philippines. PLOR A SmALESIANA 4 \ « CERATOPHYLLACEAE (C. G. G. J. van Steenis, Buitenzorg) lL. CERATOPHYLLUM LInng, Sp.Pl. (1753) 992; Mia. Fl. Ind. Bat. 1, 1 (1856) 799; BogrRL. Handl. 3, 1 (1900) 399; STEEN. Bull. J.B.B. III, 13 (1933) 102; Backer, Bekn. FI. Java 3 (1941) no 31. Submerged, rootless, monoecious freshwater plants. Leaves verticillate, 2-4 times forked, segments linear dentate. Flowers actinomorphic, solitary, axillary, uni- sexual. Perianth valvate, segments 9-12, persistent, narrow. d: stamens 8-24; anthers nearly sessile rather broad, connective pointed, the 2 cells mostly crowned by a minute bristle; ovary rudiment absent. 9: ovary superior, sessile, 1-celled with | ovule; style persistent, subulate, sulcate towards the apex; stamen rudiments absent. Fruit oblong, compressed, warty, not dehiscent, near the base with 2 straight or curved soft spines, or unarmed. Distr. Ca 2 spp., both ubiquitous. Ecol. In stagnant pools, small slow streams, shallow parts of lakes and ditches, in ponds, often gregarious. The specimens are frequently rough by incrustations of lime. They ‘collect’ mud on their surface and contribute to the clearing of water as do Hydrilla, Blyxa, &c. The stems die at their distal end and grow at the top. They propagate easily vegetatively but remain often sterile; near Batavia they are locally rather frequently found flowering in shallow water. The leaves are slack though in C. demersum slightly stiffer than in C. submersum. On extracting material from the water the leaves join brush-like. KEY TO THE SPECIES 1. Leaves mostly 2 times forked, segments 4, rarely 2, 3, or 5. Fruit with 1 apical and 2 basal spines 1. C. demersum 1. Leaves mostly 3—4 times forked, segments 7-12. Fruit with 1 apical spine only 2. C. submersum 1. Ceratophyllum demersum LINNE, Sp.Pl. (1753) 992; K. Sco. & Laut. Nachtr. (1905) 261; VAL. Bull. Dép. Agr. Ind. Néerl. 10 (1907) 11; BACKER, Teysm. 22 (1911) 503; Merr. Fl. Manila (1912) 202; SYBRANDI, De Trop. Natuur 5 (1916) 126; Merr. En. Born. (1921) 247; MeErR. En. Philip. 2 (1923) 141; HEYNeE, Nutt. Pl. (1927) 616; STEEN. Arch. Hydr. Suppl. 11 (1932) 305; Bull. J.B.B. III, 13 (1933) 103; McCann, J. Bomb. Nat. Hist. Soc. 37 (1934) 681; BACKER, Bekn. Fl. Java 3 (1941) no 31.—C. tuberculatum CHAM. Linnaea 4 (1829) 504; Mia. Fl. Ind. Bat. 1, 1 (1856) 799.—Fig. 1. Plant often much branched up to 3 m long, stems mostly red, internodes 1—3cm. Leaves in whorls of 7-10, dark green when adult, 1-4 cm long, base gradually or abruptly thickened above the middle. Pedicel3/4—-1!/2 mm. Perianth segments 9-12, linear, transparent greenish-white with numerous short brown lines, near the apex with few bristles, 1'/2-2 mm long. d: stamens 8 or more. 9: ripe fruit black, 4-5 mm long, with 3 spines, apical spine (style) 11-12 mm long, soft, 2 basal ones straight (in Mal. material), patent to slightly recurved, ~ 9-11 mm long. Distr. Throughout Malaysia, not yet known from the Malay Peninsula, ascending to over 1500 m. Ecol. See above. The ‘internodes’ of the leaves are sometimes inflated and imitate an articulated structure. Vern. Kantjil, M, ganggeng, Sd., ganggang, J, hoornblad (Dutch); the Indonesian names are also used for other waterplants. 2. Ceratophyllum submersum LINNE, Sp.Pl. ed. 2,2 (1763) 1409; Zo. Syst. Verz. 2 (1854) 86; STEEN. Bull. J.B.B. III, 13 (1933) 103; BAcKErR, Bekn. FI. Java 3 (1941) no 31; MERR. & PERRY, J. Arn. Arb. 23 (1942) 390. Fig. 1. Ceratophyllum demersum L. Habit, detached leaves, x 3/4, fruit enlarged. 42 FLORA MALESIANA [ser. I, vol. 42, Dec. 1949] Much like the preceding sp., up to 1m long, leaves in whorls of 6-8, adult ones 13/4—-2 cm long, bright green. Flowers not yet found in Malaysia. Q: ripe fruit black, + 4 mm long, only the style thickened into an apical soft spine, basal spines absent. Distr. In Malaysia rather rare: N. Sumatra, Java, Celebes, and New Guinea, at low altitudes. Ecol. In places as the prec. sp., the New Gui- nean specimens were found massed in open water in Melaleuca leucadendron swamp forest, plant brown (BRASS). ‘-HYDROCARYACEAE (C. G. G. J. van Steenis, Buitenzorg) IS cRAPA LINNE, Sp.Pl. 1 (1753) 120; Mia. Fl. Ind. Bat. I, 1 (1855) 635; ANON. Tijdschr. Nijv. & Landb. 30 (1877) 97-100; Boer. Handl. 1, 2 (1890) 561; VoRDERMAN, Teysm. 6 (1895) 313-321; ANON. Tropenpfl. (1905) 703; WIGMAN, Teysm. 22 (1911) 547; WEBSTER, Philip. Agr. Rev. 6 (1913) 138, t. 12; FLerorr, Bull. Jard. Bot. Rep. Russ. 24 (1925) 13; Gams, Pfl. Areale I, 3 (1927) 39; Ocnsg, Ind. Vrucht. (1927) 98; HEYNE, Nutt. Pl. (1927) 1206; Burk. Dict. Ec. Prod. Mal. Pen. 2 (1935) 2173; BACKER, Bekn. Fl. Java 4a (1942) no 75, p. 11. Floating aquatic herbs with dimorphic /eaves, submerged ones opposite pinna- tifid rootlike, apical ones ina rosette, rhomboid, dentate, with spongy often inflated petiole, arranged in leaf-mosaic; stipules 4-8, minute. Flowers bisexual, small, solitary, axillary, short-pedicelled, 4-merous, white or lilac. Petals imbricate. Disk present. Ovary half-inferior with | style and 2-4 persistent sepals turning often to thorns or horns. Fruit mostly 1-celled, 1-seeded, shell bone-hard; thorns after withering often set with barbs at the apex. Seed often producing 2-5 free germ-stalks. Distr. Several species in the Old World, but not known from Australia. Ecol. Trapa is very rare and was for the first time reported in 1877 in Java. It is surely a component of the old lowland euthrophous swamp flora such as is preserved in the Danoe swamp (Bantam). Use. The fat-containing kernels of T. bicornis are highly nutritious and are eaten in West Java but not on a scale comparable to that of several parts of Asia, where it is a staple food. Notes. Sometimes included in the Onagraceae or Halorrhagaceae. The names are those given by H. GLtckx in sched. Herb. Bog. There is little agreement about the specific distinctions in the genus Trapa. KEY TO THE SPECIES 1. Fruit triangular to 4 cm high, 7-8 cm broad with 2 large horns. Leaves green . 1. T. bicornis 1. Fruit tetrahedral, 1—1!/2 cm wee thorns 4, acca Cae often unequal. Leaves spotted black at the base Sess hat eg ; Ant ee 2. T. maximoviczii 1. Trapa bicornis OsBeck var. cochinchinensis (Lour.) GLuUcx.—Ospeck, Dagb. Ostind. Resa (1757) 191; LInnE f. Suppl. (1781) 128; VoRDERMAN lc. 313; MERR. Comm. Lour. (1935) 290; HEYNE, Nutt. Pl. (1927) 1206; Ocuse, /.c. f. 47.—T. cochin- chinensis Lour. Fl. Coch. (1790) 108; BACKER, Ann. J.B.B. Suppl. 3 (1910) 418.—T. chinensis Lour. /.c.—Fig. If. Leaves green, petiole 9-20 cm long, blade 5—7 by 6-9 cm. Flowers white. Horns of the fruit very blunt, straight or subcurved, their lateral surfaces bluntly irregularly ribbed. Distr. Native of Asia, in Malaysia: introduced probably by the Chinese, cultivated mostly near Batavia by the Chinese. In Batavia markets fruits are offered for sale of avariety with black sharp- pointed curved points suggesting buffalo horns often referred to as TJ. bispinosa Roxs.; these fruits are imported apparently directly from China (Teysm. 4, p. 499; WEBSTER /.c. fig.). Vern. Lengkat, lengkong, ling, lingkok, chin, kerendan (Batavia), calthrop(s), Chinese water 2. Trapa maximowiczii KORSHINSKY, Act. Hort. Petrop. 12 (1892) 336; DE VooGD, Trop. Natuur, 21 Fig. 1. Trapa maximoviczii Korsu. a. leaf, b. ger- chestnut (Engl.), waternoot (Dutch). Notes. Already in 1879-80, the Colon. Museum, Haarlem, received fruits of T. bicornis from Atjeh sent by Mr J. ScHAAR (cf. Tijd. Ned. Mij t. Bev. Nijv. for 1880). minating fruit, c, d. fruits——e. Trapa bispinosa Roxs. (edible, imported fruits on the market at Batavia).—f. Trapa bicornis Oss. var. cochinchi- nensis (LouR.) GLUCK (edible, cultivated locally), in section, X 1/2. 44 FLORA MALESIANA (1932) 62, 63, f. 9.—T. quadrispinosa auct. non Roxs.; VORDERMAN l.c.; Trop. Natuur 9 (1920) 73, £.; HEyNE I.c. 1207; Ocuse Lc. fig. 48.— Fig. 1. Leaf blade towards the base black-brown or with 2 dark spots, 2!/2-4 by 31/2-5!/2 cm, petiole 5-15 cm. Corolla pale lilac, anthers yellow. Fruit tipped by the conical hardened style. Thorns straight or curved often unequal, their apex barbed, surface of the fruit smooth, not ribbed. [ser. I, vol. 47, Dec. 1949] Distr. SE. Asia, rare but certainly native in Malaysia: W.-NW. Java (from Indramaju to Ban- tam, e.g. Danu swamp (ANON. (1877), lc.; Vor- DERMAN, /.c.) and S. Sumatra (Palembang Res., Lake Teloko, near Kaju Agung). Ecol. Swamps and ponds, disappearing under anthropogenic influence. Vern. Salekat, salaikat, M. Notes. Possibly one of the forms of T. natans L. s. ampl. MORINGACEAE (C. G. G. J. van Steenis, Buitenzorg) 1. MORINGA Juss. Gen. (1789) 348; Endl. Gen. Pl. (1840) no 6811, p. 1321; B. & H. Gen. Pl. 1 (1862) 430; E. &. P. 3, 2 (1891) 244. Small trees, mostly deciduous, bark gummy, wood soft, roots thickened, pungent; trunk often inflated. Leaves spread, imperfectly 2-4-imparipinnate: tissue with myrosin cells; pinnae opposite, provided with stipitate glands at the base of the petiolules and pinnae. Leaflets small, opposite, entire, all articulated. Stipules represented by blunt knobs. Flowers bisexual, zygomorphic, white (or yellow streaked red), in axillary panicles. Calyx tube short, as a hypanthium; lobes 5 im- bricate, spreading or refiexed, separately dropping. Petals 5 free, anterior one largest and erect, others reflexed, posterior smallest. Disk lining the calyx tube, with a short free margin bearing the androecium. Perfect stamens 5 epipetalous; anthers dorsifixed, I-celled, oblong, when lengthwise opened broader. Staminodes 5, subulate, with or without rudimentary anthers. Ovary superior, shortly stalked, 1-celled with 3 parietal placentas. Style filiform, stigma small. Ovules «, in 2 series on each placenta. Capsule linear, beaked, 3-6-angled; valves thick, spongy, on the inside with pitted cavities in 1 row along the median line. Seeds 3-winged (or exalate), body roundish large. Embryo exalbuminous, straight, containing oil. Distr. Ca 10 spp., confined to the semi-arid countries of Somaliland, Madagascar, SW. Africa, NE. Africa, Asia Minor, 2 spp. in India. Ecol. Said to be deciduous, but M. oleifera is to my knowledge in Malaysia never wholly so, thriving both under wet and seasonal conditions. As far as is known M. oleifera never runs wild. Uses. Numerous uses, cf. under M. oleifera. Notes. Very different opinions have been advanced on the affinity of Moringa. DALZELL arranged it with the Bignoniaceae; even recently HALLIER f. (Rec. Trav. Bot. Néerl. 15 (1918) 60, 99) followed BENTHAM, MIQUEL and others in assigning Moringa to the Leguminosae as a syncarpous remote member; GRIFFITH, however, had already predicted (Posth. Pap. 2(1839) p. xiv, 1849) that polycarpous Leguminosae should be apocarpous (as now described in Archidendron &c.). The genus is now universally adopted as allied to the Capparidaceae. In Malaysia one species occurs, only cultivated. 1. Moringa oleifera LAmMK, Encycl. 1 (1785) 398; —Morunga RumpH. Herb. Amb. 1 (1741) 184 t. 74/5.—Guilandina moringa LINNE, Sp.P1. 1 (1753) 381; Horsr. Verh. Bat. Gen. 8 (1816) 107.—Hype- ranthera moringa VAHL, Symb. bot. 1 (1790) 30; TuHuns. FI. Jav. (1825) 19; JueL, Pl. Thunb. (1918) 248.—M. pterygosperma GAERTN. Fruct. 2 (1791) 314; Hassk. Pl. Jav. Rar. (1848) 413; Mia. FI. Ind. Bat. 1, 1 (1855) 350; Suppl. (1860) 115; GresuH. Schetsen (21895) 63, t. 17.—M. nux-ben PERROTTET, Mém. Soc. Linn. Paris 3 (1824) 129; C. B. Ros. Philip. J.Sc. 3 (1908) Bot. 305.—M. polygona DC. Prod. 2 (1825) 478; Mia. Fl. Ind. Bat. 1, 1 (1855) 350, Suppl. (1860) 115.—Fig. 1. Rapid-growing often crooked tree 3— 10 m, stem 10-30 cm diam., crown thin, bark with coarse fibre. Leaves somewhat crowded towards the twig-ends; twigs lenticellate, with distinct scars; innovations pubescent. Petiole thickened at the base incl. the rachis 25-60 cm, all stalks articulated hence caduc- ous, provided with linear or clavate often curved glands; jugae 8—10. Petiolules 1-46 mm, blades ovate, obovate or oblong, 9—20(—30) by 5—12(-18) mm, tip rounded or slightly emarginate, lower sur- face pale green, nerves obscure not prominent. Panicles erect, 10-30 cm; bracts small. Flowers white with greenish base, fragrant; pedicels 7-11 mm, articulated under their tip. Calyx tube 34 mm high; posterior sepal 7-10, anterior 10-14 mm long. Petals hairy towards the base, anterior one 14-17 by 6-8 mm, posterior ones 10-13 by 5-8 mm. Sta- mens and staminodes with hairy base, directed to the largest petals, apex of the filaments recurved. Gynophore 2-3 mm. Ovary 34 mm high, densely hairy; style upwards glabrous with hollow stigma. Capsule pendent, dagger-shaped, subtorulose, 18-45 cm long, valves with 3 strong blunt ribs. Seeds 10 mm diam., wings 25 by 4-7 mm, subse- quent ones partly overlapping; testa reticulate. Distr. Indigenous in NW. India, in Malaysia introduced at an early date, now planted through- out the settled areas in all tropics, not run wild, sometimes found as a culture relic in abandoned places, up to ca 500 m. Ecol. Flowers throughout the year, fr. mostly Sept._Nov. Exclusively propagated by cuttings. Germination hypogaic. Colibris, and in India hon- eysuckers, are reported to visit the flowers. Once coralloid twig-galls 10-30 cm diam. were found in W. Java. Damage of the stem by insects causes quick exudation of gum. Vern. Horse radish tree (Engl.), kélor, J, M, Md, 46 FLORA MALESIANA [ser. I, vol. 47, Dec. 1949] marongghi, Md, marunga (Timor), k(o)léntang Cfruct.), malungay, Tag. Philip., gemunggai, mering- gai (Mal. Pen.), and many others. Uses. Many: flowers, leaves and immature fruits as vegetable. Leaves and bark (specially of the roots) medicinal. Bark-gum and seed-oil not valuable. Treelet sometimes used for living fences or as supporting tree for pepper vines (cf. RuM- PHIUS, GRESHOFF, HEYNE, BURKILL, &C.). Wood anat. M. sp.: MOLL & JANSSONIUS Mikr. Holzes 2 (1908) 513; M. oleifera: TANG, Bull. Fan Mem. Inst. of Biol. 6 (1935) 153. Both microsc. Fig. 1. Moringa oleifera LAMK, X 1/3. SAURURACEAE (C. G. G. J. van Steenis, Buitenzorg) Rhizomatose, aromatic or pungent, perennial, often stoloniferous herbs. Stem articulated. Leaves simple, entire, scattered (not alternate), often oblique; leaf base mostly reniform-cordate, nervation mostly palmate. Petiole sheathing or an intrapetiolar stipule. Flowers bisexual, actinomorphic, small, in terminal spikes or racemes or opposite the leaves (sympodial), each in the axil of a bract, bract some- times connate with the pedicel; lowest bracts sometimes petaloid. Perianth absent. Stamens 3, 4, 6 or 8, sometimes partly abortive, free or adnate to the basal part of the ovary or epigynous. Anthers 2-celled, splitting lengthwise laterally or extrorse. Ovary composed of 3-4 connate carpels, or l-celled with 4-3 parietal placentas. Styles free or connate at the base, often recurved, stigmatose on the inner surface. Fruit capsular opening at the top, or consisting of tubercled indehiscent 1-seeded cocci. Distr. 4 genera, 2 in E. Asia, 1 in California, and 1 both in Asia and Atlantic N. America; the latter with 2 species, the others monotypic. Ecol. Mostly in marshy places. Uses. In China and N. America medicinal with acrid and adstringent properties. Notes. Formerly sometimes included in Piperaceae. A key to the genera is given in Blumea 6 (1948) 244-245. KEY TO THE GENERA 1. Short spike of sessile flowers subtended by 4 petaloid bracts. Intrapetiolar stipule distinct 1. Houttuynia 1. Elongating raceme. No petaloid bracts at the base of the raceme. Petiole sheathing 2. Saururus 1. HOUTTUYNIA (non Houtr. 1780) THunB. Fl. Jap. (1784) 12, nom. gen. consery. prop.; E. & P. 3, 1 (1889) 3, excl. syn. Gymnotheca DECNE, Anemopsis HooK. & ARN. Stoloniferous. Leaves palminervous. Spikes peduncled. Stamens 3, filaments ad- nate to the base of the ovary. Ovary 1-locular, consisting of 3 partially connate carpels; placentas parietal. Fruit subglobose, opening at the apex. Seeds globose. Distr. Monotypic, from India & Indo-China to Japan and Formosa, in Malaysia doubtfully native. Notes. The generic name Houttuynia THUNB. should be preserved against the older Houttuynia HouTr. and was proposed for the list of Nomina generica conservanda (Fl. Mal. Bull. 3 (1948) 73). 1. Houttuynia cordata THUNB. FI. Jap. (1748) 234. t. 26; Mia. Fl. Ind. Bat. 1, 2 (1859) 457; T. &. B. Cat. Hort. Bog. (1866) 72; MerR. Comm. Lour. (1935) 126; STEEN. Trop. Nat. 26 (1937) 147; Blu- mea Suppl. 1 (1937) 137.—Polypara cochinchinensis Louwr. Fl. Coch. (1790) 61.—P. cordata O.K. Rev. Gen. (1891) 565.—Fig. 1. Root-stock copiously branching, up to more than 1 m long. Stems erect or ascending, up to 50 cm long. Leaves broad-ovate, base reniform-cordate, apex acuminate, 3-8'/2 by 2!'/2-6!/2 cm; petiole 1-4 cm. Petaloid bracts white, oblong, + 1-11/2 cm long. Spike 1-2 cm long. Seeds !/2~7/3 mm long, ellipsoid, testa of dry seed reticulate. Distr. Under temperate conditions from the Himalaya to E. Asia, also in Formosa, in Malaysia: only once found in W. Java, ca 1250 m alt., ques- tionably native. Might be expected to occur in Luzon or Mindoro. Ecol. A tenacious and obnoxious weed; crush- ed leaves with a fishy or fleshy smell, in W. Java found in a tea-plantation and a neighbouring bam- boo grove. Uses. A Chinese drug plant and vegetable. Vern. djukut hanjir, Sd, after the peculiar smell which much intrigued the Sundanese workers in the plantation and gave rise to a legend that the plant proceeded from the flesh and blood of a man who was killed by a tiger in the same spot. On ac- count of the smell there was even trouble with Ja- vanese weeders. 2. SAURURUS LINNE, Sp.PI. (1753) 341.—Spathium Lovur. Fl. Coch. (1790) 217.—Mattuschkia GEL. Syst. 2 (1791) 589, non al.—Saururopsis TuRCZ. Bull. Soc. Nat. Mosc. 21 (1848) I, 589; Bart. Adans. 10 (1871) 69.—Saururotus ENGL. E. & P. 3, 1 (1887) 48 FLORA MALESIANA [ser. I, vol. 47, Dec. 1949] 2.—Neobiondia PAMp. Nuov. Giorn. Bot. Ital.17 (1910) 263; GAGNEpP. Not. Syst. 2 (1911) 283. Tall erect herbs up to | m or more. Leaf base reniform-cordate. Flowers in the axils of bracts, or pedicels connate with the bracts. Stamens (8-)6, or by abortion only 4, sometimes opposite the carpels, sometimes lateral; filaments + free. Ovary composed of (3-)4 carpels connate at their base, each with 2(-4) ovules, only one developing. Styles free, recurving. Fruit depressed-globose, said to be subfleshy, parting into (3-)4 tubercled cocci. Testa of roundish seed reticulate. Distr. 2 species, one in E. Asia, 1 in Atlantic N. America. Notes. Gymnotheca DECNE (S. cavaleriei LEV.) constitutes a distinct genus. The N. American S. cernuus L. is apparently distinct through a slightly different leaf-shape, very long filaments and nodding spikes. These differential characters ought to be further studied. 1. Saururus chinensis (Lour.) Batti. Adans. 10 Loup. Encycl. Pl. (1866) 289; T. &. B. Cat. Hort. (1871) 71; Loup. Hort. Brit. (1830) 144, nomen; Bog. inedit. (1854) 63, nomen; ib. (1866) 77 (sinen- sis); MERR. En. Philip. 2 (1923) 2; Grorr, Lingn. Sc. J. 11 (1932) 84; Merr. Comm. Lour. (1935) 126.—Spathium chinense Lour. Fl. Coch. (1790) 217, ed. WILLD. (1793) 270.—S. cernuus (non L.) THUNB. Fl. Jap. (1784) 154.—S. loureiri DECNE, Ann. Sc. Nat. III, 3 (1845) 102; C.DC. Prod. 16, 1 (1869) 239; F.-VILL. Novis. App. (1880) 174; VIDAL, Phan. Cuming. (1885) 138; Rev. Pl. Vasc. Filip. (1886) 219.—Saururopsis chinensis Turcz. Bull. Soc. Nat. Moscou 21, 1 (1848) 589.—S. cumingii C.DC. Prod. 16, 1 (1869) 239. Glabrous, except for pedicels & rachis. Stem ribbed, pithy, lower part terete, upper part angular; lower /eaves amplexicaulous, leaving annular scars, upper ones half-amplexicaulous, stem somewhat zigzag towards the apex, internodes 1-4 cm; lower petioles 61/2, upper ones 3—11/2 cm long. Leaf blade mostly oblique, base reniform-cordate-truncate, apex acute to slightly acuminate, palmatinervous with 5 larger and 2 feeble nerves, shape ovate-lan- ceate, 8!/2-121/2 by 3'/2-6!/2 cm. Spikes straight, bracts in bud pine-like imbricating, 5—8 in fruit to 15cm long; peduncle glabrous, 1—3 cm. Rachis and pedicels pubescent. Flowers white, fragrant. Bracts ovate to roundish, or broad-spathulate, their blade 1-2 by 1-2 mm, - ciliate, with brown dots (glands) in the parenchyma, connate with the 1'/2-3 mm long pedicels and winging these; rachis angular by the decurrent pedicels. Anthers 4 behind the carpels, or 6 (in twos lateral of the carpels), or 8. Filaments + 1/3—1/2(—3/4) mm; anthers + convex, elliptic, + 3/s-1 mm long, extrorse. Cocci 4, tu- bercled, 11/2 mm long; style recurved ca 1/2 mm. Seed ovoid, fine-reticulate, 5/6-1 by 2/3/4 mm. Distr. Indo-China & Hainan to Central China, Japan, Riu Kiu and Formosa, in Malaysia only in Luzon (Ilocos Norte, Bontoc), in swampy places, muddy banks, rice-fields, 50-1100 m. Notes. It should be studied whether there are Fig. 1. Houttuynia cordata THUNB., habit x 2/s, several sexual forms showing floral dimorphy. flower enlarged. Uses. Used as a drug in China (GrorFF /.c.). STYRACACEAE (C. G. G. J. van Steenis, Buitenzorg) Evergreen trees or shrubs. Leaves simple, spirally arranged, sometimes pseudo- alternate, margin entire or toothed, mostly with stellate or lepidote indumentum. Stipules 0. Flowers bisexual, actinomorphic, axillary or terminal. Calyx tubular more or less adnate to the ovary; lobes if present valvate. Corolla rarely of free petals, mostly united in a basal tube, 4-7, valvate or imbricate. Stamens equal and alternate, or double the number of the petals, mostly adnate to the tube. Disk absent; anthers 2-celled, introrse, splitting lengthwise. Ovary superior, rarely semi- inferior, 3—5-celled. Style 1; stigma punctiform to 3-5-lobed. Ovules l—~ in each cell, axile. Fruit capsular (rarely drupaceous) 1—~-seeded, dehiscent or not, pericarp often thick and woody or corky, with a persistent calyx. Seeds with copious endosperm and straight or slightly curved embryo. Distr. Ca 12 genera mostly in the N. hemisphere, absent in Australia and the Central Pacific, richly developed in E. Asia. No Styracacea has yet been found in the Philippines proper, Central & East Java, and the Lesser Sunda Isl.. Sumatra is the richest centre in Malaysia. Ecol. Styrax represents a northern type in the Malaysian flora but its representatives are found both in the lowland and the mountains up to ca 1600 m alt. Most peculiar galls are found in Styrax, caused by specially adapted Aphids, which surpass the European oak galls in variety (cf. Doct. v. LEEUWEN, Bull. J.B.B. III, 4 (1922) 147; Zoocec. D.E.I. (1926) 452; Tijd. Entom. 75 (1932) Suppl. 97; Ned. Kruidk. Archief 51 (1941) 217, and J. C. VAN DER MEER Monr, Natur & Museum 63 (1933) 163, 6 fig.). Uses. The wood is little used. Of some Styrax spp. the bark after having been softened by taps yields benjoin resin from incisions made in the bruised portion. This resin contains benzoic or cinnamic acid. Tapping benjoin is a common procedure in Sumatra, mainly Palembang and Tapanuli Res.. Benjoin is used as an antiseptic, in cigarettes and ceremonials, and is an important forest product of Sumatra. Wood anat. See also generic descriptions. The ground tissue by MOLL & JANssoNntus (2 (1920) 472) called fiber tracheids according to REINDERs’s definitions (Handl. 3 (ed. 1941) 145) are libriform fibers; checked by C.A.R-G. Notes. The genus Symplocos was formerly often included in the Styracaceae or Styracineae, but is accepted to represent a separate family Symplocaceae in this Flora. Styracaceae possess stellate hairs, or scales, not fasciculate stamens, linear anthers, a half or wholly superior ovary, an imperfectly celled fruit, and differ, moreover, anatomically distinctly from the Symplocaceae. KEY TO THE GENERA 1. Pedicels articulated. Flowers dull-creamy; lobes and stamens free. ae, glabrous, imperfectly 5- celled. Seeds minute, numerous. Leaf margin serrulate . : 1. Bruinsmia 1. Pedicels not articulated. Flowers white; lobes and stamens connate in a “basal tube. Ovary hairy, imperfectly 3-4-celled. Seed 1(—2), large. Leaf margin subentire See Se eS 2. Styrax 1. BRUINSMIA BoeRL. & Koorp. Nat. Tijd. N.I. 53 (1893) 68; Perk. PfAl.R. 30 (1907) 14, 88; PerK. Gatt. Styr. (1928) 8, 13, 16, 24; STEEN. Bull. J.B.B. III, 12 (1932) 215. Glabrate tree with flattened-angular branchlets through decurrent petioles. Leaves serrate, brownescent. Flowers in (mostly foliate) terminal thyrses. Pedicel with 1 bracteole, articulated at the apex. Buds solid. Calyx broad-campanulate with trun- cate margin, entire or sub-5-toothed. Corolla-lobes_5(-6), free, imbricate. Stamens 10(-12), 5 longer alternate, 5 shorter epipetalous, or 10 subequal, coherent mutually and with the base of the corolla-lobes. Ovary free for the greater part, imperfectly 5(-6)-locular. Style 5-angular, not grooved. Stigma capitate, sub-5—6-lobed. Ovules o. Fruit indehiscent, pear-shaped; style-base mostly persistent. Seeds small, pris- matic-4-angular. Distr. Monotypic, endemic in Malaysia. 50 FLORA MALESIANA [ser. I, vol. 42 1. Bruinsmia styracoides BoERL. & KoorD. &c. ll.cc.; Wood anat. Mo.iit & JANssonius, Mikr. STEEN. J. Arn. Arb. 28 (1947) 422.—B. celebica Holzes 4 (1920) 498. Koorpb. Med.’s Lands PI.T. 19 (1898) 525.—Fig. 1. Notes. The Celebes and New Guinea speci- Evergreen tree 15-37 m, diam. 30-150 cm(most- mens possess hairy anthers. ly 25 m by 40 cm); clear bole 7-11 m, without but- tresses, bark dirty orange in section. Adult leaves oval to oblong, base mostly rounded, apex acute to acuminate, sparsely hairy, 7!/2-19 by 3-10 cm; primary nerves 6-10 pairs; petiole sulcate 1—1!/2 cm long. Thyrses 10-25 cm. Pedicels 2 mm; bracteole narrow, 1-—2!/2 mm long. Calyx 11/2-2 mm high, 5-6!/2 mm diam. Corolla lobes pubescent on both sides, tip cap-shaped, ovate-oblong, acute, 9-10 by 4-41/2 mm. Stamens 5—5!/2 and 6—6!/2 mm, some- times subequal 31/24 mm; filaments glabrous or short-hairy inside and marginal; cells 11/2-3 mm long. Ovary 2-2!/2mm high, 3-3!/2mm diam. Style 3-5 mm long. Fruit 6 by 9 to 10 by 6 mm; style remnant 3/4-6 mm long. Seeds 1!/2 mm long. Distr. Sumatra, W. Java (W of Buitenzorg), Borneo, Celebes, and New Guinea, (400—)700- 1600 m alt., expected to occur in the Philippines & Moluccas. Ecol. In primary or partly devastated forests, rather rare; globose leaf-galls 1/2-1 cm diam. are found in Borneo and New Guinea. Vern. Names local and not trustworthy. Uses. Wood not durable and besides the tree is rare, though dimensions would be sufficient. Fig. 1. Bruinsmia styracoides BOERL.& KoorD., x 3/4. 2. STYRAX LINNE, Sp.PI. (1753) 444; Perk. Pfl.R. 30 (1907) 14,17; STEEN. Bull. J.B.B. III, 12 (1932) 220.—Cyrta Lour. Fl. Coch. (1790) 278.—Lithocarpus BL. ex ROYLE, Ill. (1839), 261, in syn. Shrubs or trees, at least the innovations stellate-hairy, tomentose or lepidote. Leaves mostly tomentose below; petiole sulcate. Calyx campanulate to cup-shaped, free, or the base connate with the ovary, truncate, rarely toothed. Flowers solitary, mostly in racemes or leafy panicles. Corolla lobes 5(-6-7), connate at the base with the annular staminal tube. Stamens (8-9-)10(-11-14), erect, connate at the base; anthers split lengthwise. Ovary imperfectly 3-locular, with few ovules per cell, or only 1. Style 3-angular; stigma punctiform or indistinctly 3-lobed. Fruit globular to oblong, dehiscent or not. Seeds 1(-2) attached at its base. Distr. Ca 120 spp. in the tropics and subtropics of Europe, Asia, Malaysia, and America. In Malaysia the genus is richest in Sumatra, but the widest distributed species, St.agreste, is not found in that island. Wood anat. St. benzoin: MOLL & JANssonius, Mikr. Holzes 4 (1920) 503. St. benzoin & St. paral- leloneurum: DEN BERGER, Med. Proefstat. Boschw. 13 (1926) 152 (hand lens). Notes. In Java St. tonkinensis Pierre furnishing Saigon benzoé is recently cultivated by the Forest Service. In the key belowit is to be placed nearest to St. ridleyanum but differs distinctly by its narrow buds 7-10 by 2-3 mm. 1 KEY TO THE SPECIES . Undersurface of adult leaves with a closed tomentum, concealing the parenchyma. Pamcocollanmbricateanibud) | ¥.) "na ey ae ee ee 8. St. serrulatum var. mollissimum 2. Corolla valvate in bud. 3. Indumentum glaucous without stellate brown hairs or scales in adult leaves . 5. St. oliganthes 3. Brown stellate hairs or scales present in the indumentum on the undersurface of adult leaves. 4. Tomentum thin with appressed stellate scales. Fruit large, globose to ovoid-globose, 2'!/2-3 by 23/4—31/4 cm. Seed shining brown, base broad, 11/2 cm across, hilum 1 cm. 6. St. paralleloneurum 4. Tomentum with stellate hairs; scales absent. 5. Arms of brown stellate hairs 1/2 mm long. Leaves large 12!/2-18 by 6-12 (rarely 8 by 3) cm. Flowers shortly pedicelled, in linear many-flowered racemes. Fruit obovate, tip acute-acuminate, 13/4—2Naiemidiam. (os) «0 a eee ee es a ee SY AS Srcroteneiees Dec. 1949} STYRACACEAE (V. Steenis) 51 5. Stellate hairs minute, arms much smaller than !/2 mm. 6. Inflor. much longer than the leaves. Youngest twigs dark-brown. Pedicels 4-9 mm. Buds 9-11 by 4!/2-6 mm. Calyx 2!/2-3!/2 mm high, 4!/2—5!/2 mm diam. Petals not fleshy, sparsely pubescent inside. Connective shortly pubescent. Stigma large, 3-lobed, thrice as thick as the style 7. St. ridleyanum 6. Inflor. mostly shorter than the leaves. Youngest twigs greyish. Pedicels 11/24 mm. Mature buds 12 by 3 mm. Calyx 44!/2 mm high, 3—3!/2 mm diam. Petals fleshy, median part inside impressed and glabrous. Connective glabrous. Stigma punctiform as thick as the style 1. Adult leaves not with a closed indumentum. 3. St. benzoin 7. Corolla valvate. Racemes 2-17-flowered. Petals glabrous inside, 7-10 by 2-3 mm. Style 8!/2-12!/2 mm long. Fruit 8-18 by 3!/2-7!/2 mm, 7. Corolla imbricate. 1(—2)-seeded, not dehiscent, distinctly rostrate 1. St. agreste 8. Fruit apiculate. Calyx sparsely stellate-lepidote. Corolla lobes 16-17 by 71/2-8!/2 mm, stellate- pubescent on both sides. Racemes 2-3-flowered. Style 15-20 mm. Fruit ca 12-13 by 8-10 mm, + irregularly dehiscent at the base into 3 valves. Pedicels drooping, 1—2 cm 2. St. japonicum 8. Fruit with rounded apex. Calyx tomentose. Flowers spac in paniculate rich-flowered infil. Pedicels shorter, flowers not drooping . 8. St. serrulatum 9. Pericarp smooth. Leaves mostly subtomentose below. "Panicles mostly many-flowered 9. Pericarp rugose. Leaves subglabrous below. Depauperate racemes . 1. Styrax agreste (Lour.) G. Don, Gen. Hist. 4 (1837) 5; Perk. Pfi.R. 30 (1907) 27; STEEN. Bull. wept 12) (1932) 224: J. Arn. Arb: 20 (1939) 220; ibid. 28 (1947) 423.—Cyrta agrestis Fig. 2. Fruits and seeds of Malaysian Styrax. Upper row from left to right: St. serrulatum Rox. var. mollissimum STEEN., St. serrulatum ROXB. var. rugosum STEEN., St. oliganthes STEEN., St. agreste (Lour.) G. Don, St. crotonoides CLARKE. Second row: two forms of St. paralleloneurum PERK., with seed. Third row: St. benzoin DRYAND., right: a seed of St. benzoin DRYAND. var. hiliferum STEEN., show-. ing the large hilum, x 3/s. Lour. FI. Coch. (1790) 287.—St. warburgii PERK. Pfl. R. 30 (1907) 27.—St. serrulatum (non RoOxs.) Hus. WINKL. Bot. Jahrb. 50 (1914) 195; HALL. f. B.B.C. 34, HL (1916) 42.—St. ledermannii PERK. Notizbl. 10 (1928) 457.—St. rostratum HosoKAWA, Trans. N. H. Soc. Form. 28 (1938) 65.—Fig. 2, 3. Shrub or small tree 3-12 m, up to 5 cm diam. Leaves thin, ovate-oblong to ovate-lanceolate, apex acute to acuminate, 3!/2-13 by 2-6'cm; nerves 4-9 var. mollissimum var. rugosum pairs; petiole 21!/2-6 cm. Racemes 1'/2-12'/2 cm, lowest fis often axillary. Pedicels frequently united in twos, 4-6 mm, in fr. 9 mm. Calyx 3—5 mm across. Corolla tube 2—3!/2 mm high, lobes oblong to lan- ceolate, acute, outside hairy. Stamens hairy below, upper part narrowed and glabrous, 7-10 mm; an- thers linear, as long as the filaments, not glabrous. Ovary with a 2/3-11/4 mm long beak. Fruit 1-2- seeded, often oblique, rostrate by a 1—3'/2 mm long beak. Seed acute at both ends, lengthwise grooved, lepidote, base contracted, 7-13 by 2!/2-5!/2 mm. Distr. From Annam and Hainan through Ma- lasysia to the Solomons & Micronesia, in Malay- sia: Borneo, Celebes, Moluccas, New Guinea, and Bismarcks, 10-300, rarely to 1000 m alt. in the Solomons and NE. New Guinea. = LAS ig / Z ESN EAM Fig. 3. St. agreste (Lour.) G. Don, with fruits and a stem gall intermediate between the coralloid and the alcicorniform types, x 1/2. 52 FLORA MALESIANA [ser. I, vol. 42 Ecol. Undergrowth of primary forests, in Bor- neo and New Guinea inundated in the wet season, fl. and fr. mostly simultaneous, March—Nov. From Borneo coralliform twig galls are described. Vern. Papunti (SE. Born.), kaju abu, berwewa (Daj. Koetai), no fixed name. Notes. A Hainan specimen was distributed as St. subcrenata H.M. Fig. 4. Bark gall of St. benzoin DRYAND., X 7/s. 2. Styrax japonicum Sires. & Zucc. Fl. Jap. 1 (1835) 53, t. 23; Perk. Pfi.R. 30 (1907) 73.—St. kotoensis HayatTA, Ic.P1].Form. 5(1915) 121.—St. philippinensis Me_rr. & Quis. Philip. J.Sc. 56 (1935) 313, t. 1. Shrub 2-3 m. Leaves on long-branches alternate, ovate to ovate-lanceolate, sparsely stellate-lepidote to subglabrous and often with domatia in the nerve-axils below, 4!/2-9!/2 by 2-6cm; lateral nerves 4-5 pairs; petiole 4-7 mm. Pedicels 1—1!/4 cm, base densely lepidote, upwards sparsely so, thickened towards the calyx, drooping. Calyx cam- panulate, 4-5 mm high, stellate-lepidote. Fruit el- lipsoid, apex truncate, mucronate, seed oblong, 3- sulcate, surface minutely pitted, 10-11 by 5-6 mm. Distr. Japan, N. China, Riu Kiu and Formosa, in Malaysia: N. Philippines (Camiguin & Batan Isl., N. of Luzon), only twice found, apparently at low alt. Ecol. Fl. in March, fr. in June. Notes. A distinct northern type, here accepted as a slightly differentiated outlier of the St. japo- nicum population with somewhat larger flowers. 3. Styrax benzoin DRYAND. Phil. Trans. 77 (1787) II, 308; PerK. Pfi.R. 30 (1907) 59; STEEN. Bull. J. B.B. III, 12(1932)228 ; Burk. Dict. (1935) 2105.—St. benzoé BL. Cat. (1823) 6, nomen.—St. benjoin ROxB. Fl. Ind. (1832) 415.—Lithocarpus benzoin BL. ex Roy te, Ill. (1839) 261, in syn.—Plagiospermum benzoin PIERRE, FI. For. 4 (1889) 260.—St. rauensis BOERL. ms. ex GRESH. /.c. 118.—Fig. 2, 4, 5b. Tree 8-34 m, trunk 10-100 cm diam., buttresses small or absent; bark wine-red in section, 3—7 mm, wood white. Leaves ovate to oblong or lanceolate, base rounded, apex acuminate, 8-13 by 2!/2-5 cm; petiole sulcate 5-13 mm; nerves 7—13 pairs. Flow- ers fragrant, in 6-11 cm long panicles often form- ing a leafy panicle 13-20 cm long. Buds solid, blunt. Pedicels 11/24 mm, top-shaped. Corolla lobes 9-12 by 2—3!/2 mm, tube 1-2 mm high. Stam- inal tube 2!/2-4 mm, anthers 5 and 5!/4 mm long. Ovary 8-12!/2 mm high. Fruit depressed-globose 2-21/4 by 2'/4-3 cm, indehiscent; pericarp (3—)4—5 mm diam., subtended by a rarely appressed calyx 7-13 mm diam. Seed 1(-2), ca 15 by 19 mm, dull pale brown; hilum 3-6 mm diam., in var. hiliferum 10-12 mm diam. Distr. Malaysia: Sumatra (only the main land), Malay Peninsula and West Java (rare, mostly W of Buitenzorg), 10-1500 (mostly 100-700) m, doubtful from Banka Isl. Erroneously recorded from the Philippines by F.-ViLL, Nov. App. (1880) 27. Ecol. In mixed primary forests, often com- mon, mostly on fertile soils; 77. & fr. not periodic, Jan._Dec. Germination preferably in the shadow. Several gall forms are described, all of the saccate type. The fruit is eaten by swine and deer. Uses. Cultivated in Sumatra (mostly Palem- bang and Tapanuli Res.), Java (also in monsoon climate) and W. Borneo, often in clearings; there- fore, the tree occurs frequently gregarious in old secondary forests. Sumatrans often soak the fruit before planting, or peel them. Yields red benzoin, mostly consisting of benzoic acid. Resin can be drawn from 7 years old saplings. Wood worthless. Vern. (Ke)menjan, hamindjan or kumajan, with various spelling and with various additional names. Notes. No specimens are known with certainty from the Indochinese Peninsula; those recorded belong mostly to St. benzoides CRatB which differs in the absence of brown-stellate hairs, smaller flow- ers (lobes 8 by 1'/2-13/4 mm) and different fruit (1 cm high with!/2 mm thick 3-valved pericarp). This Fig. 5. Brown stellate hairs of a. St. crotonoides CLARKE, b. St. benzoin DRYAND., and c. scales of St. paralleloneurum PERK., X 160. continental species produces Siam benzoé (cf. Hook. Ic. Pl. t. 2999). var. hiliferum STEEN. /.c.—Seed subglobose 17-18 by 20-24 mm. Testa dark-brown. Hilum 10-12 mm diam.—Fig. 2. Dec. 1949] Distr. Malaysia: Malay Peninsula (Pahang) and Sumatra (Tapanuli). Vern. Kemanjan bukit (Mal. Pen.), hamindjan minjak (Batak, Sum.). Fig. 6. Styrax crotonoides CLARKE, 1/3, flower, nat. size, bunch of sack-galls, x 1/3. 4. Styrax crotonoides CLARKE, in Hook.f. Fl. Br. Ind. 3 (1882) 589; STEEN. Bull. J.B.B. III, 12 (1923) 239; BurK. Dict. (1935) 2107.—Fig. 2, 5a, 6. Tree 13-27 m by 25-50 cm. Leaves oblong to broad-elliptic, rarely subovate, base rounded, mar- gin entire, apex acute-acuminate; nerves (7-)9-11 pairs, connected by transverse veins. Infl. mostly Fig. 7. Spiral galls of St. paralleloneurum PERK., on a fruiting twig, x 2/3. STYRACACEAE (Vv. Steenis) 53 unbranched spike-shaped, rachis stout, straight, bearing fils from the base; fis solitary or in pairs. Pedicels !/2-3!/2 mm, rusty tomentose. Calyx 4—5 mm across. Corolla lobes 10 by 2'/2 mm, tube 1-2 mm high. Staminal tube 2 mm; filaments 1 mm; connective lepidote; anthers 5-6 mm. Style 6-7 mm. Mature fruit rusty tomentose. Distr. Malaysia: Malay Peninsula, 50-300 m alt. Ecol. Mixed primary forests on low hills. Um- bellate rusty coloured sack-galls are described. As far is known it yields no benzoin. 5. Styrax oliganthes Steen. Bull. J.B.B. III, 12 (1932) 241.—Fig. 2. Tree up to 33 m, trunk 50-60 cm diam. Leaves ovate-oblong, rarely lanceolate, often oblique, base cuneate or rounded, apex blunt-acuminate, margin recurved s.s., 7—-9!/2 by 3-4 cm; primary nerves 7-8 Fig. 8. Bark gall of St. paralleloneurum PERK., x 4/s. pairs; petiole 3!/2-8 mm. Flowers unknown. Calyx in fruit 31/24 by 61!/2-7!/2 mm. Fruit obovate not dehiscent, apex broadly rounded, short-mucronate, 10-13 by 8'/2-9!/2mm. Seed ovate, dull dark- brown, without papillae, 9!/2 by 6mm; hilum 4 by 3 mm. Distr. Malaysia: Sumatra (Eastcoast Res.), ca 500 m alt. Only once collected. Ecol. Primary mixed forest, fr. Febr. Notes. By its height, non-dehiscent fruit, pau- ciflorous infl. and not papillose seeds different from St. tonkinensis Crats. and St. benzoides CRAIB from continental Asia; the flowers may yield addi- tional characters. 6. Styrax paralleloneurum PERK. Bot. Jahrb. 31 (1902) 484; Pfi.R. 30 (1907) 37; STEEN. Bull. J.B. B. III, 12 (1932) 243.—St. sumatranus J.J.S. Tect. 10 (1917) 204.—Fig. 2, 5c, 6-8. Tree,- 5-35 m by 23-60 cm; clear bole 6—25 m. Leaves ovate to lanceolate, mostly ovate-oblong, 54 FLORA MALESIANA [ser. I, vol. 42 pee Pg My NOR Fig. 9. Spiral galls of St. paralleloneurum PERK.., 3/5. These twisted galls have essentially the same structure as sack-galls, and contain a cavity. distinctly acuminate, 6-16 by 2!/2-6!/2 cm; to- mentum of undersurface thinner than in St. benzoin and not woolly, golden-brown lepidote; nerves 6-8 pairs glabrate, with distinct transverse veins, mostly cinnamon-coloured, petiole 7-15 mm. Larger infil. leafy, panicles 2-11 cm long. Coro//ain mature buds 2-4 x as long as the calyx. Fls violet-scented, drooping, 4-6 mm stalked. Calyx 3-4!/2 mm high and slightly broader. Corolla 13-15 mm, tube 4— 41/2 mm; lobes 11—13 by 3—3!/2 mm. Stamens 12-— 12!/2 and 13-14 mm; filaments 9-10 mm; anthers 4-5 mm, margin of the cells lepidote. Fruit-base included by the cupular, fleshy, thickened calyx, shell 5-9 mm diam. hard-fleshy. Seeds 1(—2) ovate- globose, with broad base; testa shining, bony, dark-brown. Distr. Malaysia: Sumatra (only main land) & Malay Peninsula, 575-1700 m. Ecol. Primary mixed forests, rarely in second- ary forest (old clearings), often common but scat- tered, mostly on slopes, sometimes on ridges. Habit as in St. benzoin, resembling nutmeg trees. Galls unique, spirally twisted sack-galls. F/. mostly March—July, fr. July—Nov. Uses. Wood of inferior quality. Tapping yields white benzoin chiefly consisting of cinnamic acid, in Sumatra (Tapanuli Res.) planted in clearings and secondary forests, in Java by the For. Serv. Vern. (Ke)menjan, in different spellings and additional epithets, as in St. benzoin. Notes. Very distinct from St. benzoin, and eas- ily recognizable in the sterile state by the transverse veins and appressed scales on the underside of the leaves. f. inutilis STEEN. /.c—Tomentum of leaf under- surface tinged light brown, scales few; upper sur- face light brown to brown s.s; benzoin said to be worthless. Distr. Once found in Tapanuli, said to be removed from plantations. Vern. Kemenjan bulu (also used for St. benzoin). 7. Styrax ridleyanum PeErRK. Pfl.R. 30 (1907) 61; STEEN. Bull. J.B.B. III, 12 (1932) 247; FiscHer, Kew Bull. (1937) 438.—St. subpaniculatus (non JUNGH. & DE VR.) BACKER ex HEYNE, Nutt. PI. (1927) 1260, in nota; BuRK. Dict. (1935) 2107. Tree up to 30m by 30cm, buttresses up to 80 cm, clear bole 15 m. Leaves generally alternate, ovate to oblong-ovate, distinctly often abruptly acuminate, 7-17 by 31!/2-8 cm; nerves 7-9 pairs; petiole '!/2-1 cm long; axillary buds 3-5 mm long, brown-tomentose. Panicle leafy at the base, 7!/2— 20 cm long. Flowers fragrant. Corolla tube 2 mm high; lobes valvate or slightly induplicate-valvate, margin thickened, 9-10 by 3 mm. Stamens sub- equal, tube 2/3-1 mm high; filaments 1!/3-2 mm long; anthers 6—6!/2mm, connective prominent and broadened above the cells. Style 8 mm. Stigma lobed to truncate or hammer-shaped. Fr.unknown. Distr. Burmaand Malaysia: Sumatra & Malay Peninsula, low alt. Ecol. Apparently rare in lowland primary mix- ed forests. Dec. 1949] STYRACACEAE (Vv. Steenis) 55 Vern. Kemenjan burong (also for St. benzoin), k. landak (Mal. Pen.). Notes. The nearest ally of St. benzoin, and only differing in the flowers, though the fruit will proba- bly show additional characters when known. Not known to yield resin. Fig. 10. Flower galls of St. serrulatum ROxs. var. mollissimum STEEN., X 1/2. 8. Styrax serrulatum Roxs. FI. Ind. ed. CAREy 2 (1832) 415; CLARKE in Hook. f. FI. Br. Ind. 3 (1882) 588, p.p.; STEEN. Bull. J.B.B. III, 12 (1932) 248. No Malaysian specimen agrees entirely with the Indian ones, and I assume that the Malaysian specimens represent distinct varieties. It is a rather variable species and it is closely related to a group of allied species in SE. Asia: St. grandiflorum GriFF., St. caudatum WALL., St. hookeri CLARKE, St. virgatum Kurz, all of them possessing a larger corolla; St. japonicum is distinct by drooping, long, glabrous pedicels and a glabrous calyx. The aes- tivation is variable, being subvalvate or imbricate in bud, even in one flower. var. rugosum STEEN. /.c.—St. serrulatum auct. p.p.; GRESHOFF, Schetsen (1896) 118; GAMBLE, Mat. Fl. Mal. Pen. (1905) 251, excl. fr.; Rv. FI. Mal. Pen. 2 (1923) 296; Burk. Dict. (1935) 2107, p.p.—St. porterianum WALL. Cat. (1828) no 4401, nomen; G. Don, Gen. Hist. 4 (1837) 5; DC. Prod. 8 (1844) 267; Mia. FI. Ind. Bat. 1, 2 (1859) 463; PerkK. Pfi. R. 30 (1907) 77.—St. floribundum GRrikF. Not. Pl. As. 4 (1854) 287.—Fig. 2. Small to medium tree up to 20 m. Leaves on both surfaces with scattered small stellate hairs, 5—11 by 3—5 cm; nerves 5—6 pairs. Racemes short, termi- nal on the shoots, or axillary in depauperate race- mes or even solitary. Corolla lobes 10-11 by 4 mm. Filaments 5 mm, glabrous towards the apex; an- thers 3!/2-S mm. Style 11-14 mm. Fruit ovate- to obovate-globose, pericarp rugose 2 mm diam. Seed 6-8 mm across; hilum 5 mm long. Distr. ? India, Andamans, Mergui, Tenasserim and Malaysia: Penang Isl. and Malay Peninsula (Malacca, once). Ecol. Mixed primary forests, up to 200 m. Fi. fr. March-May. var. mollissimum STEEN. Bull. J.B.B. /.c.—St. subpaniculatum JUNGH. & DE VR. Pl. Nov. Ind. Bat. 1 (1845) 9; Mia. FI. Ind. Bat. 1, 2 (1859) 464, Suppl. (1860) 187; Gres. Schets. (1896) 118; Perk. Bot. Jahrb. 31 (1902) 483; Pfl.R. 30 (1907) 72; Burk. Dict. (1935) 2107.—Jndet. Mia. Linnaea 26 (1853) 285.—Hopea sp. Mia. l.c. Suppl. (1860) 492, in adnot.—St. subdenticulatum Mia. l.c. 187, 474; GresH. Schets. (1896) 118; Perk. T/i.cc.; S. Moore, J. Bot. (1925) Suppl. 65; v. p. MEER Monr, Trop. Nat. 20 (1931) 158.—Styrax sp. Doct. v. L. Bull. J.B.B. III, 4 (1922) 158; Zoocec. (1926) 457. —Fig. 2, 10-12. Tree 10-30 m by 8-4S cm. Leaves stellate-pubes- cent to subtomentose on the lower surface, the parenchyma remaining visible, 7-14 by 3!/2-7!/2cem; nerves 6-8 pairs. Flowers in pyramidal brown- yellow tomentose panicles 5-17 cm long; lobes 7-81/2 by 2!/2-3!/2 mm, pubescent all over, distinct- ly imbricate. Anthers 23/4 and 3—3!/4 mm. Style 8-10 mm. Fruit obovate, not dehiscent; pericarp smooth 3/4 mm diam. Seed 9!/2 by 6 mm: hilum 3 mm long. Distr. Malaysia: Sumatra, 350-1500 m alt. Figs 11. Alcicorniform galls of St. serrulatum var. mollissimum STEEN., X 1/3. 56 FLORA MALESIANA [ser. I, vol. 42, Dec. 1949] Ecol. Scattered in primary mixed forests, 1. mostly Jan.—Febr., fr. mostly May—June. Three kinds of galls are known, viz coralliform, alci- corniform and siliqua-shaped ones (cf. v. D. MEER Monr and Doct. v. L.). Uses. Once reported to yield benzoin used as incense. Fig. 12. Coralloid galls of St. serrulatum var. mollissimum STEEN., X 2/s. Excluded Styrax ceramense Wars. Bot. Jahrb. 13 (1891) 302 = Diospyros. Styrax ellipticum JUNGH. & DE VR. Pl. Nov. Ind. Bat. (1845) 10; Mia. FI. Ind. Bat. 1, 2 (1859) 464, Suppl. (1860) 187; GresH. Schets. (1896) 118; Perk. Bot. Jahrb. 31 (1902) 484; Pfl.R. 30 (1907) 86; STEEN. Bull. J.B.B. III, 12 (1932) 253.—I have not succeeded in locating the type specimen. It is certainly not Styracaceous and possibly belongs to Xanthophyllum or Vatica. Styrax glabratum (non SCHOTT) WARB. I.c. = Diospyros. Styrax javanicum BL. Bijdr. 13 (1825) 671 = Alangium. Styrax obovatum Rip. J. As. Soc. Str. Br. 61 (1912) 28 = Symplocos. Styrax rassamala REINW. ex STEUD. Nomencl. ed. 2 (1841) 651 = Alangium. Styrax villosum Bl. Bijdr. l.c. = Alangium. Styracin. gen. nov.? ZOLL. Syst. Verz. 2 (1854) 136 = Vatica. Fig. 1. Triglochin procera R.BR. var. dubia BTH. Habit, « 2/s, flower and juvenile fr. enlarged. JUNCAGINACEAE (C. G. G. J. van Steenis, Buitenzorg) 1. TRIGLOCHIN LINNE, Sp.Pl. (1753) 338; Bucu. Pfl.R. 16 (1903) 7.—Cycnogeton ENDL. Ann. Wien. Mus. 2 (1838) 210; Gen. Suppl. 1. (1848) 1369; Hutcu. Fam. FI.PI. 2 (1934) 38; pro sect. Bru. Fl. Austr. 3 (1878) 165. Glabrous, annual or perennial herbs. Leaves distichous, radical, entire, linear, with a sheath. Inflorescence terminal, spicate or racemose. Flowers bisexual, acti- nomorphic, small, inconspicuous, mostly green. Perianth segments 6, conchiform. Stamens 6 (or partly reduced), epi-tepalous. Anthers sessile, extrorse, cells 2. Carpels 6, or less by abortion, free or united, or partly free; ovule 1 per cell, basal, erect; style mostly absent. Pericarp dry. Seed exalbuminous, embryo straight. Distr. Cosmopolitan, the majority of the ca 15 spp. known from the S. hemisphere. The Malaysian species is the only one of subg. Cycnogeton (ENDL.) BUCH., distinct by entirely free carpels. Ecol. Predominantly in marshy localities, or in the water; some spp. in semi-arid countries. Notes. The family is sometimes called Scheuchzeriaceae, but Juncaginaceae has priority. It centers in the S. hemisphere, specially in the S. Pacific. HUTCHINSON (Fam. FI.PI. 2 (1934) 36, 38) has removed Scheuchzeria from Junc. to a separate family in a separate order on account of bracteate pedicels. Only one species is hitherto reported from Malaysia: 1. Triglochin procera R.Br. var. dubia (R.BR.) Bru. Fl. Austr. 7 (1878) 168; BucnH. Pfl.R. 16 (1903) 14; Sreen. J. Arn. Arb. 28 (1947) 419.—T. dubia R.Br. Prod. (1810) 343.—T. linearis ENDL. Pl. Preiss. 2 (1846/7) 54.—T. procera R.BR. var. gracilis MICHELI in DC. Mon. Phan. 3 (1881) 108. —Fig. 1. Perennial, 30—50 cm long; rhizome short, roots partly thickened and ending in globular tubers 2-3 cm across. Culms terete, solid, sheaths at the base surrounded by the fibrous remains 1—2 cm long of the nerves of former leaves. Leaves + as long as the culms, 1—3 mm broad. Spike multiflorous, rath- er lax and relatively few-flowered, 11/22, in fruit up to 7 cm long. Flowers sessile, ebracteate, towards fruiting gradually pedicelled. Stamens 6, distinctly in 2 rows, 1 mm long, very broad, slightly shorter than the erect oblong 1!/2-2 mm long tepals round- ed at the apex. Carpels mostly 3, the slightly curv- ed outwards directed stigmatic apices protruding distinctly above the anthers; stigmas large oblique, linear, grooved. Nuts mostly 3, rarely 4, or 1-2, small, 7-9 mm long, linear-oblong mostly subfal- cate, 1'/2-2!/2 mm pedicelled, erect. Distr. Australia and Tasmania to SE. Malay- sia: Wassi Kussa area in S. New Guinea, at low alt. Triglochin belongs to a series of (sometimes world-wide) spread genera, both occurring in Aus- tralia and Asia but absent or exceedingly scarce in Malaysia, as e.g.: Aldrovanda, Alisma, Anagallis,. pumila, Astragalus, Callitriche, Eryngium, Glosso- stigma, Leptocarpus, Limonium, Oxalis, Parietaria, Philydrum, Plantago, Rothia, Tenagocharis, Tragus, Villarsia, and Zannichellia. Ecol. Savannah forest, common in streams which contain water only during the wet season, locally massed. Roots partly swollen into edible terminal tubers. Hardly an aquatic plant. FY. Dec.—Jan. Notes. The species is more robust than the va- riety: length -- 1 m, leaves 4-10 mm broad, spikes denser, gynoecium less reduced. The variety more- over occupies a peculiar habitat, and seems to represent an ecotypic raciation. HUTCHINSON /.c. recognizes Cycnogeton as a separate genus by its free carpels; some other spp. of Triglochin occupy an intermediate position. Excluded Scheuchzeria palustris Mig. Ill. Fl. Arch. Ind. (1870) 48; MrcHeti in DC. Mon. Phan. 3 (1881) 95; Bucu. Pfl.R. 16 (1903) 15; STEEN. Bull. J.B.B. Ill, 13 (1934) 221.—Sch. asiatica Miq. FI. Ind. Bat. 3 (1855) 243; Suppl. (1860) 35; Koorpb. Exk. FI. Java 1 (1911) 91.—In the Leyden Herbarium is a sheet of which the origin is doubtfully given as ‘2? Java’, and said to be collected by KORTHALS. Interchange of labels is probably the cause of this erroneous record. 58 FLORA MALESIANA [ser. I, vol. 42, Dec. 1949] Fig. 1. Trigoniastrum hypoleucum Mia. Flowering twig, X2/3, a. flower, petals removed, b-c. petals, d. bag- shaped petal, e. the same, frontal view, f. fruiting tip of inflorescence, g. bracts with glands, h. winged partial fruit, 7. opened, showing insertion of seed. TRIGONIACEAE (C. G. G. J. van Steenis, Buitenzorg) 1. TRIGONIASTRUM Mia. Fl. Ind. Bat. Suppl. (1860) 394; B. &. H. Gen. Pl. 1 (1862) 139; Mig. Ann. Mus. Bot. L.B. | (1863) 142, in clavis; BENN. in Hook. f. Fl. Br. Ind. 1 (1872) 208; BAILL. Hist. Pl. 5 (1873) 91; Kina, J. As. Soc. Beng. 59, II (1890) 133; Ann. R. Bot. G. Calc. 5, 2 (1896) 135, t. 159; BoeRL. Handl. 1, 1 (1890) 78; CHopart, Bull. Boiss. 3 (1895) 136; BARTH, ib. 4 (1896) 481; E. & P. Nachtr. (1897) 209; Rip. Fl. Mal. Pen. (1922) 141; ENDERT, Med. Boschb. 20 (1928) 65; LEMEE, Dict. 6 (1935) 704 excl. syn. Factorovskya E1G.—Isopteris WALL. CAT. (1832) no 7261, nomen, ex B. & H. Lc. Tree, wood vessels mostly solitary. Leaves simple, spread (on lateral branches), penninervous, entire, margin and leaf tip glandular; upper epidermis often double and provided with mucilaginous cells; midrib sulcate above. Stipules caducous. Indumentum of simple hairs. Flowers actinomorphic, bisexual, in axillary and terminal panicles; bracts with glandular margin. Sepals 5, free, nerved as the petals are, unequal, imbricate. Petals 5, free, very unequal, imbricate; posterior saccate with reflexed emarginate limb, lateral spathulate, spreading, anterior oblique, keel-like together, including at their base the genitals; entrance of the sac with one fleshy hairy concave gland (easily breaking into 2 parts). Stamens 6 monadelphous, tube split posteriorly, eventually with some loose minute teeth, minute upper part of filaments free; anthers oval, slightly emarginate at the base, 2-celled, opening with one slit, gaping; exine (judging from boiled fis) reticulate. Ovary hairy, easily falling into 3 parts as does the simple style; stigma small punctiform. Ovule pendul- ous solitary. Fruit composed of 3 easily detaching samaras. Seeds (n.v.) elongate, shortly hairy, exalbuminous; radicle very short. Distr. Monotypic, confined to W. Malaysia, wrongly credited to New Guinea by LEMEE, /.c. Wood anat. Herscn, Lilloa 8 (1941) 132. Notes. This genus, which is the single representative of the family in Malaysia, was mostly included in the Polygalaceae, and though differing in several respects from the 2 other neotropical genera of the Trigon. by its spread leaves &c. it seems mostly allied to Trigonia. HALLIER f. suggested in passing! affinity with the Rosac.-Chrysobal. (1918) for which indeed some arguments could be advanced. In the herbarium it is sometimes confused with Angelesia which it resembles superficially in vegetative charac- ters. It has also been compared with Dichapetalaceae. The glandular leaf margin, glandular-thickened leaf tip, sulcate midrib, and the indumentum are easy characters for distinction. The gland in the posterior petal could be interpreted both as a disk or as (a) metamorphosed stamen(s). 1. Trigoniastrum hypoleucum Mra. /.c.; BENN. /.c.; Cuopat /.c.; Foxw. Mal. For. Rec. 2 (1922) 216; RIDL. /.c.; BAKER f. J. Bot. (1924/6) Suppl. 7, 143; Merr. Pl. Elm. Born. (1929) 132; Burx. Dict. (1935) 2182; incl. var. oliganthum cum var. viride Airy SHAW, Kew Bull. (1940) 253.—Isopterys pe- nangiana WALL. ex BENN. /.c.—Fig. 1. Small to moderate-sized tree 6-30 m by 14-50 cm; wood hard, yellowish, fine-grained; bark ex- uding a yellowish, later turning reddish, gummy juice. Innovations hairy. Leaves oblong with cu- neate base, apex acuminate, glandular-thickened, often + spathulate; petiole 4-7 mm, stout, sulcate; blade 8-18 by 3-6cm; upper side often with a metallic hue s.s.; lower side whitish by a very thin arachnoid appressed indumentum consisting of twisted hairs, interspersed with sclerenchymatic longer hairs; margin with minute impressed glands; side-nerves ca 5—6, reticulations prominent on both sides. Panicle leafy, 20-40 cm, lower bracts leafy, upper ones linear acute. Flowers white, fragrant; anthers pale yellow. Buds whitish s.s. Sepals ca 2!/2 by 11/2 mm, ovate, acute, hairy. Saccate petal- base ca 2mm, sometimes compressed, reflexed part ca 2 by 2!/2 mm; base thickened; laterals un- guiculate-spathulate, ca 3!/24 by 11/2 mm; anteri- ors 4-5 mm long, oblique-oblong. Staminal tube 1 mm high, filaments over !/4 mm free; anthers ca 1/2 mm. Gland ca 3/4 mm diam. Style 11/2 mm glab- rous. Ovary '/2 mm high. Samaras pale or yellow green, connate over 1!/2—2!/2 cm, with straight inner side 2!/4-3!/2 cm long and rounded or lozenge- (1) Med. ’s-Rijks Herb. Leiden -37, p. 56. € 60 FLORA MALESIANA [ser. I, vol. 4°, Dec. 1949] shaped outer side, 1—13/4 cm broad. Seed not seen. Distr. Malaysia: Sumatra, Malay Peninsula (incl. P. Penang), Borneo (incl. P. Laut). Ecol. Evergreen non-inundated rainforest, scat- tered up to 1000 m, but mostly below 300 m; at the highest altitude found in sandy rainforest, differing slightly by blunter thicker leaves; f7. fr. Sept.—June. Uses. The wood is little estimated. Vern. kikir, mangkudor (SE. Borneo), kaju bras, tinggiran batu (Palemb.), tinga batu (Asa- han), marajali, mata pasak, suginara (Mal. Pen.), apparently no fixed name. Notes. The seedling grows out of the wing; the cotyledons are epigaic; the Ist pair of leaves is opposite. COCHLOSPERMACEAE (C. G. G. J. van Steenis, Buitenzorg) 1. COCHLOSPERMUM KunrtTH, Malvac. (1822) 6; DC. Prod. 1 (1824) 255; PLANcH. in Hook. Lond. J. Bot. 6 (1847) 139, 294, 311; BorrL. Handl. 1, 1 (1890) 70; Cat. Pl. H. B. 1 (1899) 49; Rip. FI. Mal. Pen. 1 (1922) 252; Pita. in E. & P. ed. 2, 21 (1925) 316; STEEN. Bull. J.B.B. Iff, 13 (1936) 519; BAcKer, Bekn. Fl. Java 4a (1942) no 83. Trees (or shrubs), often deciduous, producing gum and an orange juice. Leaves spread, palmatilobed, often with domatia in the axils of the main ribs; stipules caducous. Flowers actinomorphic, bisexual, showy, mostly golden-yellow, pani- culate or racemose. Sepals 5imbricate. Petals 5, imbricate or contorted, emarginate. Stamens c, with free filaments, equal or subequal; anthers 2-celled, linear, basi- fixed, opening by introrse, short, often confluent pore-like slits. Ovary 1-celled with laminal placentas projecting into the cell, or perfectly or imperfectly 3-celled, the upper portion remaining I-celled; ovules «, style simple, stigma punctiform. Capsule 3—5-valved, valves of the endocarp separating from and alternating with those of the pericarp. Seeds covered by woolly hairs, mostly cochleate-reniform; endosperm copious, rich in oil; embryo large, conforming to the shape of the seed; cotyledons broad. Distr. Ca 15 spp., mostly in trop. and subtropical America, some in trop. Africa and SE. Asia, 3 species in N. Australia, rare in Malaysia; G. gillivrayi is possibly the only native Malaysian species. LAM assumed the genus to belong to the ‘antarctic’ type (Blumea 1 (1935) 135), but it is manifestly peri-tropical. Ecol. The species prefer a semi-arid or seasonal climate, and in Malaysia occur only in regions with a dry season. Some African species are fire-resistant through a corky layer on the rhizome. Notes. The family was formerly often included in Bixaceae; HALLIER f. referred it to the Tiliaceae (Med. Rijksherb. 35 (1918) 18). Several authors describe the sepals of Cochlospermum as deciduous; in C. religiosum cultivated at Buitenzorg they are decidedly persistent. Domatia are not yet recorded in the genus but in cultivated C. regium at Buitenzorg they are present in the basal axils of the main ribs as tufts of hairs, and Acari were observed between them. The generic name is preserved against the few years earlier Maximilianea MART. & SCHRANK (1819). KEY TO THE SPECIES 1. Ovary pubescent. Stamens red in the lower half 1. C. regium 1. Ovary perfectly glabrous. Stamens yellow. 2. Leaves entirely glabrous; lobes 5—7, incised for 5/s—7/s of the length of the main ribs, 6-7 by 2-3 cm. Fls ++ 6-9 cm across. Stamens + 1 cm long with blunt hardly tapering anthers 3-4 mm long, pore rounded-triangular : ; 2. C. gillivrayi . Leaves pubescent to tomentose beneath, midribs tomentose above. Blade incised to 2/3 of its “Teas into 3—5 lobes, middle lobe -- 5-11 by 4!/2-9 cm. Fls + 12-14 cm diam. Stamens 11/2-2 cm long, anthers distinctly tapering to the acute apex 6-7 mm long, pore sagittate . 3. C. religiosum 1. Cochlospermum regium (Marr. & SCHRANK) apex depressed, finely velvety pubescent, striate- Pirc. Notizbl. 8 (1924) 716; Backer, /.c.—Maxi- milianea regia MART. & SCHRANK, Flora 2 (1819) 452; BAKH. v. d. Br. Bull. J.B.B. III, 6 (1924) 185.—C. vitifolium SPRENG. Syst. Veg. 2 (1825) 596; STEEN. /.c. 521. Small tree 3-12 m, with red-brown branches. Leaves + orbicular, 10-30 cm wide, cordate, gla- brous, incised to 2/3—3/4, lobes 5, acuminate, crenate- serrate, basal axils of main ribs with domatia; petiole 10-25 cm. Flowers in dense panicles at the end of leafless twigs, pedicellate, bright yellow. Sepals 5, tomentose, persistent. Petals obovate, 4-6 cm long. Stamens ca 2cm, inner ones shortest; anther cells with a terminal pore. Capsule 5-valvate, partly green partly red, obovoid, 4-8 by 4-6 cm, nerved. Pericarp hard, endocarp cream-coloured, parchment-like. Seeds reniform covered with cot- ton-like white hairs. Distr. NativeinCentral & South America, in Ma- laysia sometimes cultivated as an ornamental tree. Wood anat. VESTAL, Philip.J. Sc. 64 (1937) 221. Note. The ovary in C. regium is not 3-locular as is often suggested but imperfectly 5-locular. 2. Cochlospermum gillivrayi BENTH. FI. Austr. 1 (1863) 106; F.v.M. Descr. Not. Pap. Pl. 1,4(1876) 54 (gillevraei); incl. var. papuanum BAKER f.J. Bot. 61 (1923) Suppl.4; STEEN. Bull. J.B.B. III, 13 (1936) 522. Small deciduous tree, 4!/2-7 m, bark pale-grey, 62 FLORA MALESIANA [ser: vole channeled and scaly (BRAss); branches of the pa- nicle, pedicels, and calyx slightly pubescent other- wise glabrous. Leaves 5-11 cm across, on 6-13 cm long petioles; stipules 34cm long, subulate; blade divided to about !/2-1'/2 cm from the base; lobes slightly toothed-crenate, 2 outer shortest and very acuminate. Panicles short and loose. Flowers yel- oe EGGS \ EE : 3. Cochlospermum religiosum (L.) ALston, Handb. Fl. Ceyl. 6 (1931) 14; STEEN. Bull. J.B.B. III, 13 (1936) iv; CorNER, Wayside trees (1940) 174.— Bombax religiosum LINNE, Sp.Pl. (1753) 552; BAKH. v. d. Br. Bull. J.B.B. III, 6 (1924) 186.— Bombax gossypium (non L.) Cay. Diss. Bot. 5 156.—C. gossypium DC. Prod. 1 G85) 297 te Fig. 1. Cochlospermum religiosum (L.) ALSTON, after the tree cultivated in the Bot. Gardens, Buitenzorg (type of C. balicum), X 1/2. low; pedicels less than 1!/4 cm, lengthening after flowering. Sepals shortly pubescent, glabrous towards the apex, with very thin edges, outer sepals usually smaller than the inner ones. Anthers oblong, curved. Capsule obovoid-oblong, rarely exceeding 7'/2 cm, depressed-truncate. Seeds enveloped in fugacious wool. Distr. N. Australia, N. Queensland, Thursday Island and other islands near N. Australia, in Ma- laysia: only known from the Port Moresby area, Terr. of Papua, sometimes planted there, also near Boku (Kemp Welch river area). Ecol. Mostly in anthropogenic localities but apparently native, common on dry rocky places along the coast, ff. July-Sept. (1824) 527; STEEN. Bull. J.B.B. III, 13(1936) 522.— Bombax conga Bur. f. Fl. Ind.(1768) 145.—‘Bom- bax lobatum’ DescHamps MS., Brit. Mus. t. 57.— C. balicum Boer . Cat. pl. Hort. Bog. 1 (1899) 49.— Fig. 1. Small rather crooked-branched, more or less deciduous tree. Leaves cordate, 7!/2-20 cm diam., margin undulate or obscurely crenate, apex of the lobes acuminate with blunt tip; petiole articulated 8-25 cm long. Stipules linear-subulate, caducous, 7-8 mm long. Flowers in a simple raceme or a loosely branched panicle, terminal, short-hairy, one flower open at a time on each branch, facing side ways. Pedicels + 2 cm long. Bracts caducous, triangular-acute, short-hairy, 4-5mm broad at the Dec. 1949] COCHLOSPERMACEAE (V. Steenis) 63 base. Corolla yellow, with a distinct odour. Sepals partly purplish, 2—2!/2 by 1—2!/4cm. Petals thickened at the base. Stamens slightly S-curved, unequal. Anthers orange, falcate, 1 mm broad. Ovary glo- bular. Style glabrous 1!/2-2 cm. Capsule obovate, 5—7cm long, valves striate-ribbed, 2!/2-3 cm broad. Seeds reniform to cochleate, brown, 5-6 mm across. Distr. India, Cambodia, introduced in Siam and Ceylon, in Malaysia: Penang Island and Ma- lacca, E. Java and Bali, doubtless introduced. Ecol. More or less established in Penang and very conspicuous in cultivated ground round the town, near Dato Kramat growing in a native settle- ment, in Malacca GRIFFITH noted ‘near a stream at the foot of... hill’; in N. Bali near a large pagode (TEYSMANN) pr. Singaradja. At Buitenzorg flow- ering throughout the year. In Penang noted as deciduous. All specimens at low alt. It is propa- gated by cuttings or rather loppings; any branch stuck in the ground will strike in wet weather. Flowers last for 11/2 day. If the flower is setting fruit the withered petals remain for a long time (CORNER). Wood anat. By GAMBLE the wood is defined as ‘extremely soft’ (Indian Timbers 1902; here also short hand lens description). Vern. Bebaru (Penang), tjanigarah (Bali), butter- cup tree, yellow cotton-tree, yellow silk cotton-tree (Engl.). The Malay name ‘bebarw’ is used in con- fusion with Hibiscus tiliaceus. Notes. Certainly introduced in Penang, never re-collected in GRIFFITH’s spot. Among the MS.- plates left by DescHAMps no 57 is unmistakably this species, which has never been re-collected in Java after DescHAmps figured it (+ 1793-1802); acc. to his MS. he did not visit Bali Island. The locality near Singaradja was visited by Mr DE Voocp in 1936; flowering material was collected near the pagodes. The late Mr Kerr wrote me, Aug. 1936, that ‘there is no good evidence that the species is indigenous in Siam; the only place is a small hill crowned by a small deserted temple where it may quite well have been planted’. In India it is often used as a temple plant, and flowers are used in offerings. In Ceylon TriMEN found it in the dry districts, but it occurs usually near tem- ples for the sake of its beautiful flowers. For these teasons I am convinced that the species is intro- duced in Malaysia, that its introduction is due to its religious fame in India, and that is has been brought to Bali by the Hindus. BOERLAGE described the Balinese specimens er- roneously as having subglabrous leaves. ZYGOPHYLLACEAE (C. G. G. J. van Steenis, Buitenzorg) 1. TRIBULUS LINNE, Sp.PI. (1753) 386; STEEN. Bull. J.B.B. III, 13(1933) 104; BAcKer, Bekn. FI. Java 4a (1942) no 66. Prostrate hairy herbs. Leaves opposite, paripinnate, mostly anisophyllous; stipu- les present. Flowers actinomorphic, 5-merous, bisexual, solitary on pseudo-axillary peduncles, white or yellow. Sepals 5, free, imbricate, persistent or caducous. Petals 5, free, patent, imbricate, fugacious. Disk present. Stamens 10, subequal or un- equal; anthers dorsifixed. Ovary superior, sessile, hairy, 5-12-lobed, 5-12-celled; style short and thick, with 5-12 decurrent stigmas; cells with 3 or more ovules. Fruit 5-angled or 5-12-winged; cocci partly abortive, spinous or tuberculate, inde- hiscent with 3-5 superposed seeds separated by septa. Distr. & Ecol. Ca 20 spp. difficult to delimit, specially developed in the dry regions of Africa and Australia. In S. Africa the spinous fruits adhere to the wool and feet of sheep (‘hoof-burs’) and are a nuisance. The family has about 26 genera, of which 12 monotypic, and ca 250 spp., mostly of warm dry countries. In Malaysia one genus and one species. 1. Tribulus cistoides LINNE, /.c.; CHAM. Linnaea 5 (1830) 44; STEEN. /.c. p. 105; DE VooGpD, Trop. Nat. 26 (1937) 162; MeRR. & PERRY, J. Arn. Arb. 21 (1940) 511.—2T. terrestris (non L.) THUNB. Flor. Jav. pt 2, cur. WIDMARK (1825) 13.—T. terrestris Fig. 1. Tribulus cistoides L., on the beach of Nusa Penida, near Bali. (DE VooGD) var. moluccensis Bl. Bijdr. I.c.; MQ. /.c. 682; For- BES, Wand. (1885) 501.—T. moluccanus DECNE, Herb. Tim. Descr. (1835) 118.—T. lanuginosa (non L.) THuns. /.c.; BLANCO, FI. Filip. (1837) 350.—T. macranthus Hassk. Flora 48 (1865) 403.—Fig. 1, 2. Perennial herb, branches decumbent, apex as- cendent, 30—-145cm long; tap-root very long. Leaves of each pair: one 6—7- the other 4—5-jugate, 2—7!/2 cm long, midrib ending into a small mucro 1/2-2 mm long. Leaflets subsessile, base obliquely rounded-cordate, apex blunt to subacute, 6-22 by 2!1/2-9 mm, both surfaces silky. Stipules falcate, acu- cuminate, erect, 3-6 mm long. Flowers inserted laterally of the axil of the smallest leaves, bright yellow, 2!/2-4 cm diam.; pedicels hairy, 2-4 cm. Sepals narrow-lanceolate, acute, appressed-hairy, 7-11 mm long, caducous. Petals obovate-cuneate, apex broadly rounded-truncate, 1-2 by 1—1!/2 cm. Stamens subequal; anthers -- 1 mm long. Disk represented by 5 small erect scales appressed to the hirsute ovary, alternating with its lobes. Cocci 4-5 with sharp stout spines, two lateral largest; peri- carp rather thick, corky. Distr. Circumtropically ubiquitous, in Malay- sia: confined to the E. part (fig. 2), avoiding the everwet Sunda Land; in some places, near har- bours, probably an alien. Ecol. In Malaysia exclusively indigenous along sand beaches and coastal dunes, locally often gre- garious, fl. March—July. Notes. Closely allied to T. terrestris L., an an- nual or biennial with much smaller flowers. Excluded Fagonia montana Mia. olim in sched. Herb. Metz, Mia. FI. Ind. Bat. 1, 2 (1859) 596, in syn.; CLARKE, in Hook. FI. Br. Ind. 3(1882) 620; BoeRL. Handl. 1 (1890) 147 = Azima sarmentosum. Though MI- QUEL mentioned the name in his Flora, the specimen on which it was based is extra-Malaysian. Fig. 2. Localities of Tribulus cistoides L. in Malay- sia; an arrow indicates a presumably introduced occurrence. PODOSTEMACEAE (C. G. G. J. van Steenis, Buitenzorg) Mostly annual, rather small, of peculiar habit, often moss-like, gregarious, confined to swift running water in streams and cascades. Tissues with silictum. Roots often thallose, flat, stem sometimes absent. Leaves mostly alternate, sometimes scattered, decussate or distichous, base often provided with a sheath, and sometimes stipule- like appendages, often dentate or divided. Flowers terminal, often in cymose inflor., mostly ¢, actinomorphic to zygomorphic. Perianth of 3-5 free or subconnate tepals, if reduced to two small, ovate or linear appendages, the bud is enveloped by an originally closed thin ‘spathella’. Stamens hypogyn, Il-, often 2 unilateral, frequently monadelphic. Anthers mostly introrse, 2—4-locular, splitting length- wise. Pollen grains single, or in twos or fours. Ovary superior, ovate to elliptic, mostly 2-, rarely 3-locular with thickened central placenta and thin septa. Ovules ©; Styles as many as carpels, free, rarely 1. Capsule septicid (often) ribbed; seeds ~, minute, exalbuminous, epidermis mucilaginous. Distr. Principally confined to the tropics throughout the world, not yet recorded from the Pacific islands and the greater part of Australia, northward as far as S. Japan, in Malaysia apparently very rare. The locality closest to Malaysia is isthmian Siam where Dr A. KeitH found Podostemon ?algaeformis Bru. in the nineties. Ecol. All members of this aberrant stream-resistant family are confined to swift not too densely shaded streams with normally clear water. The rather numerous SE. Asiatic members are all tiny to minute plants growing socially and covering the rocks in sheets. During the rainy season they are mostly found only in the sterile thallose state and submerged. In the dry season the flowers appear and the thallus whithers. They occur often very locally, and the plants are easily overlooked. Many additional records may beexpected. Local dispersal is certainly effected by water; Cladopus seedlings have been found on rocks, a cemented dike, bamboo and an iron pipe at highwater mark. Dispersal from one river system to another unconnected is still obscure (birds, fishes, insects?). Transplanting experiments which I did with rocks covered by Cladopus from Mt Gedeh on Mt Salak in W. Java, were unsuccessful (1937). So were CIFERRI’s.in the Dominican Republic (Atti Ist. Bot. G. Briosi e lab. Crittog. Ital. Univ. Pavia V, 7 (1946) 18-21). Notes. The affinity of the family is still uncertain; opinions are offered of alliance to Saxifragaceae; ENGLER accepts it to represent a separate Group Podostemales in the neighbourhood of the Urticales. KEY TO THE GENERA 1. Flowering stems very short, hardly 1 cm. Leaves crowded, scattered, imbricate, digitately lobed 1. Cladopus 1. Flowering stems elongate, 2!/2-6!/2 cm. Leaves distichous, laterally compressed, entire, the upper 3—(4)-dentate ce tee fo: be Pil ee be eth bo pet oso) 2 Bae Ae eee ee eLOLrenticols 1. CLADOPUS H. MOLL. Ann. Jard. Bot. Btzg 16 (1899) 115; STEEN. Bull. Jard. Bot. Btzg III, 159(1936) 530 (iz). Small, flatly adhering to the substratum. Roots flat, ligulate, without stomata, side-branches mostly opposite, at last concrescent in a crust on the substratum. Leaves of the sterile sessile sprouts (? roots) partly linear partly digitate with 4-7 segments as are those of the fertile stems. Flowering stems minute, hardly 1 cm in length, densely covered with scattered, imbricate, digitately lobed leaves. Flowers solitary, pedicellate, zygomorphic, before anthesis enclosed in an oblique spathella; spathella oval-acuminate, apex nipple-shaped, mostly irregularly dehiscing, some- times slit on one side. Tepals 2, narrow triangular to linear, at both sides of the base of the filament. Stamen 1 (rarely 2 ?), anther basifixed, cells divergent at the base, filament curved; pollen grains 2-celled. Ovary smooth, oblique-ellipsoid, as long as the filament, curved towards the anther. Capsule oblique-ellipsoid, smooth, the largest valve persistent. 66 FLORA MALESIANA ser. I, vol. 42 Distr. S. Japan (Kyushu), Malaysia: Java and SW. Celebes. Ecol. Swift running streams and cascades, clinging to the rocks, also found on a cemented dike and occasionally young plants on bamboo, 5—1550 m. Notes. The Japanese species C. japonicus IMAMURA belongs in my eye doubtless to Cladopus and does not represent a separate genus (Lawiella Korbz.). Specifically it differs in 8—12-lobed leaves. 1. Cladopus nymani H. MOL t., //. cc.—Fig. 1. Roots dark green often reddish tinged, densely branched at last forming a continuous concrescent sheet exceeding 60 cm diam., branches opposite or alternate, '/2-3 mm broad, tips with a minute dorsal root cap. Sterile /eaves in rosettes, simple or digitate, narrow 2-6 by '!/6—!/3 mm. Fertile stems at the root margins, up to 5 mm long, leaves with 3-7 segments '/4-1 mm long. Spathella ca. 2 mm. Tepals narrow linear acute, 1 mm long. Stamens 1, rarely 2, anthers 3/4a-1 mm long. Ovary 2-locular ca. 1'/2 by 1 mm, stigmas 2, oblique, 2/3 mm long. Capsule smooth 1!/2-13/4 by 1 mm; pedicel 2-3 mm. Distr. Malaysia: W.-E. Java, SW. Celebes, 5— 1550 m. Ecol. See under the genus. Fig. 1. Cladopus nymani MOLL., a. piece of rock with fruiting plant, x 2, b. leaf, x 4, c. open flower with dehisced spathella, x 6, d. bud, lateral, < 6, e. stem in fruit, x 4, f. flowering stem, x 4, g. bud, dorsally, x 6, h. sterile root system, juvenile plant, x 3. 2, TORRENTICOLA DoMIN, nom. prov. Bibl. Bot. 89, 2 (vol. 20) (1925) 149, t. 35, f. 7-13; ENGL. in E. & P. ed. 2, 18a(1935) 484; STEEN, J. Arn. Arb. 28 (1947) 421.—Podost. indet. F. v. M. Syst. Cens. (1882) 23; 2nd Cens. (1889) 166; BAIL. Compr. Cat. (1913) 417.— Podostemon sp. DoMiIN, I.c., nom. alt.—Fig. 2. Roots lingulate, sparsely ramified; sprouts very close, erect, mostly simple rarely branched, thin, rigid, densely foliate. Leaves distichous, equitant, base laterally compressed, obtriangular, upper ones toothed, minute, thick, 1—2-carinate, decur- rent, lowest semi-amplexicaulous, blade + patent. Flowers single, apical, strongly asymmetric. Spathella sub-oblique, tipped, irregularly circumscissile-dehiscent. Flower shortly stalked. Tepals 2, narrow, small. Stamen 1, articulate with the pedi- cel, in bud appressed to the ovary, filament broad, anther broad, connective emar- ginate, cells introrse. Pollen grains 2-celled. Ovary + globular, smooth, 2-locular with 2 grooves where the dissepiment is attached; stigmas 2, oblong-lanceolate, in bud appressed to the ovary towards the stamen. Fruit slightly oblique, terete, the largest valve persistent, ribs 10, indistinct. Placenta globular, surrounded by a thin narrow membranous dissepiment, caducous. Seeds numerous sub-angular oblong. Distr. Monotypic, Queensland and Malaysia: SE. New Guinea. Dec. 1949] PODOSTEMACEAE (Vv. Steenis) 67 sy igh) i, iy Mt Fig. 2. Torrenticola queenslandica DOMIN. a_b. branched and unbranched stems, c. lower part of stem, d. central part, e. upper part, f-g. stem-bases attached to lingulate roots on pieces of gravel, h. leaf types, i. bud on apex of stem, j. closed spathella with bud, k. spathella opened artificially, /-m. dorsal view of flower, n—p. larger valve of fruit, q. unopened fruit. (a-b, x 3/4, c-q, X 9). 68 FLORA MALESIANA _ [ser. I, vol. 42, Dec. 1949] Notes. The first specimens were found in Queensland, Johnstone river, by WALTER HILL, in 1873; they remained a long time undescribed; OLIveR sent the specimens in 1874 to WEDDELL who recognized the species as new but could not place it. The New Guinean material exactly matches Domrin’s description. The floral structure is described here for the first time. Though manifestly distinct in the lingulate roots and in phyllotaxis, the closest ally is difficult to find, as there are several genera of which the flower- structure is very similar (Cladopus, Polypleurella) but their vegetative structure differs widely from that of Torrenticola. Vegetatively some Podostemon species from America are similar. Decussate leaves are found in Willisia from the Anamalais, but this genus possesses an androphore with 2 stamens. The tipped spathella Torrenticola has in common with Cladopus. For these reasons it stands more or less isolated and apparently represents a separate genus. Between the pollen grains I found globular bodies 1/10th the length of the grains, measuring about 3 ». They were sometimes in twos. I accept these to represent sterile pollen grains. 1. Torrenticola queenslandica Domin, /.c.; ENGLER, l.c.; STEEN. l.c.—Podostemon sp. DOIN, I.c. nom. altern.—Fig. 2. Roots ca 1/2-2 mm broad. Stems 2!/2-6!/2 cm long, densely set, darkgreen. Leaves ridged on the back, ca. 1/2-2 mm long. Spathella + sessile, to + 1/2 mm stalked, + 2 mm long. Pedicel !/2-11/4 mm in bud, 2 mm in fruit. Tepals + 1/2 mm long, filament + 14/2 mm long, thecae + 3/4 mm, cells slightly unequal, pollen grains + 30y. Ovary 11/4 mm diam.; stigmas + ‘!/2 mm; seeds ca 1/4—!/3 mm diam. Distr. Queensland and Malaysia: SE. New Guinea, Roona (Laloki river), c. 200 m alt. Ecol. On submerged rocks in the river (May 1935, CARR 12415). The stems and leaves were attacked by a fungus forming brown spots. Excluded Lemnopsis major ZOLL. Syst. Verz. 1 (1854) 75, 86, acc. to BACKER, Handb. Fl. Java pt 1 (1925) 59 (ZoLL. 3430) = Halophila ovalis Hook. f. Lemnopsis minor ZO.vL. ibid. (ZOLL. 3334) = Halophila ovata GAUD. Lemnopsis mnioides Zipp. Flora 12 (1829) I, 285; Alg. Konst- & Letterbode 1 (1829) 297, acc. to HAL ter f. Med. Rijksherb. Leiden 1 (1910) 40 = Utricularia orbiculata WALL. Tristicha bifaria PRESL. Rel. Haenk. 1 (1827) 86 is suspected to represent T. Aypnoides Spr. and was erroneously recorded from the Philippine Islands acc. to MERRILL (Philip. J. Sci. 10 (1915) Bot. 189). Sp.: D’ALBERTIS, ‘What I did and what I saw’ 2 (1880) 93 mentions a plant which he found June 17, 1876 in the Fly River (STEEN. /.c. 1947) which was advanced by Lam (Blumea 2 (1936) 117) as a possible representative of the Pod. in New Guinea; the record will remain doubtful as no material was collected. BeccARI Piant. Sum. 482 in herb. Kew (STEEN. 1936, 1c.) = Aneura tamariscina STEPH. (Hepa- ticae). AMARANTHACEAE (C. A. Backer, Heemstede) Herbs, rarely climbing or clambering shrubs. Leaves opposite or alternate, exstip- ulate, simple, entire or obsoletely dentate-serrate. Flowers 9, unisexual, or partly difformed and neutral, in clusters, heads, racemes, spikes or panicles, solitary or clustered in the axil of persistent bracts, usually bibracteolate. Tepals 3—5, mostly free; bracts, bracteoles and tepals with scarious margins or entirely scarious; brac- teoles falling off with the perianth or persistent; perianth usually enclosing the fruit and falling off with it, rarely persistent. Stamens as many as petals and op- posed to them, rarely fewer; filaments free, or connate below, or almost entirely united in a cup or tube, with or without interposed dentiform, subulate, linear or short and broad pseudo-staminodes; anthers dorsifixed or inserted in a basal cleft, 1—2-celled (2- or 4-locellate). Ovary superior, 1-celled; ovules 1 or more, basal; funicles short or long. Fruit sometimes baccate or crustaceous, usually membra- nous, very rarily corky, circumscissile, indehiscent or bursting irregularly. Seeds l—c, often lenticular or subreniform, smooth or verruculose. Distr. Worldwide, more than 60 genera and ca 850 spp., few in the tropical forests, most developed in America and Africa, in Australia a big centre of Ptilotus. In Malaysia: mostly represented by widely distributed anthropochorous spp., none endemic, several naturalized. The floristic areas occupied by the native spp. can be divided into three types viz wides, Western and Eastern elements. The wides are those of spp. widely distributed in the Old World. Their native country is often unknown. The Western element (Asiatic or/and African) is represented by: Aerva curtisii, Nothosaerva brachiata (doubtful), Digera muricata, Pupalia lappacea, and is mostly confined to the western half of Malaysia. It is about as strong as the Eastern (or Australian) element which consists of: Gomphrena canescens, G. tenella, Ptilotus conicus, Deeringia arborescens, Amaranthus leptostachyus & A. interruptus. This eastern element is for the greater part confined to New Guinea, the Moluccas, and the Lesser Sunda Islands. Ecol. Mainly annuals of open places, in Malaysia: mostly in anthropogenic localities, waste places, road-sides, fields, two aquatic (Centrostachys, Alternanthera philoxeroides), one occasionally so (Alter- nanthera sessilis), a few almost exclusively in forests. Uses. Some spp. are used as vegetables, especially Amaranthus. There are some ornamentals (Celosia, Alternanthera, Gomphrena, Iresine, Amaranthus). Several are medicinal. Alternanthera ficoides var. bett- zickiana is used against soil wash. Ashes of Achyranthes aspera are rich in alcali. Notes. Flowers and ripe fruits are necessary for identification. The foliage, in many spp. is very variable and hardly offers constant characters for specific distinction. The species are, apart from the shape of the leaves, not very variable; Amaranthus tricolor and A. lividus are exceptions. Celosia argentea and some other species have produced varieties in breeding; these are sometimes unstable. Figs. 2-3, 5-8 courtesy Pasuruan Exp. Station. (ARTIFICAL) KEY TO THE GENERA 1. Leaves alternate. 2. Scandent or clambering shrubs, unarmed, green-leaved, at least 2 m tall, often very much larger. Flowers singly or clustered along the rachis of simple or panicled racemes or spikes, 5 or unisexual, glabrous, white, yellowish or greenish, 1'/2-2'/2 mm long; filaments at their base united in a cup, without intervening pseudo-staminodes; stigmas 2-4, mostly 3, patent or recurved. Fruit an inde- hiscent berry falling out of the persistent perianth when ripe. . . . .. . 1. Deeringia 2. Otherwise. 3. Fruits all or for the greater part 2~co-seeded. Flowers 9 in simple or panicled (sometimes coxcomb- like) spikes, glabrous. Stamens 5; filaments at their base united in a cup. Erect, unarmed herbs or undershrubs. 4. Style 1, rather long; stigma 1, capitate. Perianth 6-10 mm long. Fruit membranous, circumsciss; seeds 1-9 rane ona 2 Celosia 4. Style very short or absent; stigmas 2-3, linear-clavate. Perianth less than 4 mm long. Fruit baccate, falling out of the persistent perianth when ripe; seeds 10 or more, usually more than 20 1. Deeringia 3. Otherwise. Fruit 1-seeded. 5. No pseudo-staminodes. 6. Flowers 9. Style 2-3 mm long; stigmas 2-3. Unarmed. 7. Flowers in short, dense head-like spikes. Lower flowers not accompanied by 2 palmatifid scales (sterile flowers). Filaments at their base connate in a cup. Stigma 1, entire or faintly 2-lobed. Fruit thin-walled. 70 FLORA MALESIANA [ser. I, vol. 42 8. Tepals distinctly dimorphous: 2 outer ones elliptic, their bases externally with a dense tuft of hairs; 3 inner ones with a narrow cuneate claw and patent ovate-oblong blade. Fruit inde- hiscent aie ae ja at ers woe) os ou RR lige 8. Tepals subequal. Fruit c circumsciss. . Seed arillate of Se er oso iris 7. Flowers in spiciform often long racemes, the lower part of the older racemes lax. Lower flowers usually accompanied by 2 palmatifid scales (sterile flowers). Filaments free. Stigmas 2, recurved. Fruit crustaceous, indehiscent. Seed exarillate . . Gc 5. Digera 6. Flowers unisexual, in sessile (d) (Q) or (dQ) clusters; these either collected 1 in a spike or panicle or not. Filaments free. Style very short or absent; stigmas 2-4 (often 3), linear. Unarmed or spinous. 4. Amaranthus 5. Filaments alternating with subulate pseudo-staminodes. Flowers 3 or Q, in the latter case long- pilose outside. Style very short, stigmas 1—2, minute . «. <: . =. . . J. 8. Aerva 1. Leaves, at least partly, opposite. 9. Lower flowers not accompanied by fascicled hooks (difformed flowers). 10. Flowers in panicled spikes, minute, in Malaysia exclusively 9. Perianth + 11/4 mm long, externally at the base with a dense whorl of long white hairs, otherwise subglabrous, nerveless. Stigmas 2, ascending, ‘subulate.zi '@ (fo Se. War os de Re cp 2 Oe ese 10. Otherwise. 11. Anthers 2-celled (4-locellate). 12. Filaments 5, alternating with pseudo-staminodes. 13. Pseudo-staminodes short, with a broad, truncate or subdentate apex. Flowers glabrous, united in a (finally) long spike, after anthesis widely patent or reflexed. Bracteoles, or one or more tepals, after anthesis with a very acute, often pungent tip. Stigma capitate, entire. 14. Aquatic. Bracteoles broadly ovate-orbicular, membranous, not spinous. Tepals very unequal, outermost one spine-tipped . . . . . . 10. Centrostachys 14. Terrestrial. Bracteoles consisting of a rather long spine bearing on either side of its long con- cave base a much shorther, membranous nerveless wing. esas not very unequal, after an- thesis spinous ornot . . Mec 11. Achyranthes 13. Pseudo-staminodes subulate. Blowers either whites pilose or thinly beset with short hairs and then strongly nerved, not pungent Sy its < » ge 8. Aerva 12. No pseudo-staminodes. Style at best !/2 mm lone Leaves elabrous. 15. Stamens 1-2. Spikes usually clustered, !/2-11/2 cm long. Tepals nerveless, hairy outside 9. Nothosaerva 15. Stamens 5. Spikes solitary or sometimes paired, 1—2!/2cm long. Tepals strongly nerved, glabrous .. Sako. ie 13. Psilotrichum Anthers 1-celled (2- Nocellate), or absent and replaced by spurious ovaries. We Stigma 1, capitate, entire . . oo. el hUuhCUlelCleCl, |) A Cites 16. Stigmas 2, erect or spreading, sometimes minute ; 3 15. Gomphrena 9. Flowers racemed or spicate, lower ones accompanied by fascicled hooks (difformed flowers). Hooks hairy at the base, upwards glabrous. Perianth pilose outside. Style 1, stigma capitate. Fruit falling off together with the hooks, by means of these easily adhering to passers-by. 17. Filaments alternating with short, broad pseudo-staminodes. Old flowers deflexed, accompanied by sessile or subsessile fascicles of obliquely erect, 13/4—2!/2 mm long hooks +» « 0..@yathula 17. No pseudo-staminodes. Old flowers widely patent with distinctly stalked fascicles of squarrose, 3-4 mm long hooks st few eer eh ey eS ae hee Serene = el 3) es 1. DEERINGIA R.Br. Prod. (1810) 413. Erect herbs or scandent or clambering shrubs, unarmed. Leaves alternate, peti- oled, ovate to lanceolate, acute, entire. Flowers in axillary and terminal, simple or branched, frequently panicled racemes or spikes, solitary in the axil of a bract, subtended by 2 bracteoles. Tepals 5, rarely 4, oval-oblong, 1-nerved with scarious margins, glabrous. Stamens 5, rarely 4, filaments at the base united in a cup; free parts distant, filiform-subulate; no pseudo-staminodes; anthers 2-celled (4-locel- late). Ovary sessile or shortly stalked; ovules few to many; funicles long; stigmas 2-3, rarely 4, linear or + clavate. Fruit baccate, thin-walled, indehiscent, globose, broadly ellipsoid or obovoid, falling out of persistent perianth when ripe; seeds 0 to many, on long pale funicles, circular or reniform, shining black or brownish black, very finely verruculose or almost smooth. Distr. About 7 spp. in the palaeotropics from Madagascar to Australia. Dec. 1949] AMARANTHACEAE (Backer) 71 Ecol. Unlike most Malaysian Amaranthaceae the species of this genus are not anthropochorous; as a rule they inhabit forests. Uses. Only one species used by the Malaysians, mainly medicinally. Notes. This is the only Malaysian Amaranthaceous genus of which the fruits (red or white berries) fall off unopened, leaving behind bracts, bracteoles and perianth. KEY TO THE SPECIES 1. Single flowers sessile or subsessile. Single spikes consisting of 50 or fewer flowers. Flowers 3 or (d ) (Q). Tepals appressed against the ripe living berry. Seeds distinctly verruculose. 2. Flowers 9, in simple or very sparingly branched spikes or sometimes solitary. Perianth 2!/4—-2!/2 mm long. Free parts of the filaments about as long as the staminal cup or but slightly longer. 3. Spikes rather less than 1 cm long. Flowers very few, occasionally 1. Climber Ripe berry red. Seeds 1-4. 3. D. tetragyna 3. Spikes 3-12 cm long. Flowers 4-50. Ripe berry ‘white. "Seeds ‘10-64. Erect herb or undershrub, 1-2 m high 2. D. polysperma 2. Flowers (dc) (9) (male ones ‘spuriously bisexual but < OV ary empty), in panicled spikes. Branches of the panicle widely patent. Perianth 1!/4-2 mm long. Free parts of the filaments in gd much longer than the staminal cup. Ripe berry bright red. Seeds 1-2. Very tall climber 4. D. arborescens 1. Single flowers on 2/3-2 mm long pedicels, 5. Single racemes often consisting of more than 50 flowers. Free parts of the filaments several times longer than the staminal cup. Tepals under the ripe living berry patent or reflexed. Ripe berry bright red. Seeds 0-9, almost smooth 1. Deeringia amaranthoides (LAMK) Merr. In- terpr. Herb. Amb. (1917) 211; Spec. Blanc. (1918) 136; En. Philip. F1.Pl. 2 (1923) 126.—Achyranthes amaranthoides LAMK, Enc. 1 (1785) 548.—Celosia baccata RETZ, Obs. 5 (1789) 32; BLANco, FI. Filip. (1837) 193.—Deeringia celosioides R.BR. (non HaAssk. 1848) Prod. (1810) 413; DECNE in Nouv. Ann. Mus. 3 (1834) 371; SPAN. in Linnaea 15 (1841) 345; BLANco, FI. Filip. ed. 2 (1845) 135; ed. 3, 1 (1877) 244, t. 236; Mia. FI. Ind. Bat. 1, 1 (1858) 1025; Bru. Fl. Austr. 5 (1870) 209; F.v.M. Descr. Not. pt 3 (1876) 41; Hook. f. Fl. Br. Ind. 4 (1885) 714; Bat. Queensl. Fl. pt 4 (1901) 1218; Ripv. FI. Mal. Pen. 3 (1924) 4; HEYNeE, Nutt. Pl. (1927) 604; OcuHsE & BAKH. V. D. BR. Veget. (1931) 30, fig. 19; GAGNn. in FI. Gén. I.C. 4 (1936) 1055.—Deeringia indica RETZ ex BL. Bijdr. (1825) 542; Hassk. PI. Jav. Rar. (1848) 436.—Lestibudesia philippica Werno. Syll. Ratisb. 1 (1828) 118.—Celosia phi- lippica StEUD. Nom. ed. 2 (1841) 316, Mia. in DC. Prod. 13, 2 (1849) 241, Merr. En. Philip. F1.Pl. 2 (1923) 127.—Deeringia baccata Moa. in DC. Prod. 13, 2 (1849) 236; Forpes, Wand. (1885) 515 and 502 (here doubtfully but wrongly as Gouania leptostachya DC.); Koorp. Exk. FI. 2 (1912) 194; Merr. Fl. Man. (1912) 190; Domry, Beitr. Pfian- zengeogr. Austr. 1,2 (1921) 628.—Jresine amaran- thoides Moa. in DC. Prod. 13, 2 (1849) 348.—Cla- dostachys baccata O.K. Rev. 2 (1891) 541.—Fig. 1. Scandent or clambering shrub, often with long, pendulous branches, 2-6 m (to up to 15 m ?) high; stem in the higher part obtusangular and finely appressed pubescent, at an advanced age terete and glabrous. Leaves ovate or ovate oblong-sublan- ceolate, base acute, obtuse, rounded or subtrun- cate, often unequal, frequently shortly contracted into the petiole, tapering towards the apex or acu- minate, acute, crowned by a longish, often caduc- ous mucro, at first on both surfaces very thinly patently pilose, glabrescent, 4-15 by 2-8 cm (floral leaves often much smaller); midrib in the living plant distinctly prominent beneath; petiole 1—6cm. Flowers racemed; racemes axillary and terminal, 1. D. amaranthoides very often divaricately branched, 5—35 cm (0-8 cm peduncle included), rather dense or in lower part lax, often more than 50-flowered; highest racemes usually collected in a terminal panicle of 15-75 cm Fig. 1. Deeringia amaranthoides (LAMK) MERR. from Java, x 1/s. long; rachises of inflorescence finely and not very densely appressed pubescent; bracts narrowly tri- angular, very acute, + 11/2 mm; bracteoles ovate, acute, + 1mm. Flowers solitary or clustered, malodorous. Pedicels 2/3-2 mm. Tepa/s during an- thesis widely patent or refiexed, under the fruit reflexed as are the stamens, obtuse or rounded, concave, pale green or somewhat yellowish, white- margined, often, especially under the fruit, tinged with réd, 11!/2-2'/2 mm long. Staminal cup !/4—!/3 mm:; free parts of the filaments several times longer G2. FLORA MALESIANA [ser. I, vol. 42 than the cup, for the rest variable as to length, 1'/2-23/4 mm. Stigmas 3, greenish white, 1—1!/2mm, recurved on the fruit. Berry globose-obovoid, bright red, 4-7 mm diam. Seeds 0-9, usually no more than 5, circular with emarginate base, 1—11/3 mm diam., almost smooth. Distr. From India to China, southward to Australia, in Malaysia: throughout, not yet re- ported from the Moluccas & Borneo. Ecol. In Java, especially in the drier eastern half, 1-1500 m, in teak- and open mixed forest, forest borders, secondary forests, tall brush-wood, hedges, often, though by no means exclusively, on calcareous soil. Sometimes cultivated for its medi- cinal properties. Uses. The natives press the root in diluted vinegar and add a piece of onion (Dutch: ajuin; BURKILL (Dict. 775) mistranslated alum = aluin). The juice so obtained is sniffed up; it looses the mucus and cures the head-ache caused by obstruc- tion of the nasal cavities. The leaves are applied to sores; young cooked sprouts are eaten with rice. Vern. Bayam besar, bayam pohon, pantjar luhur, tangtang angin. Notes. The specimens found in native hedges may have been planted. D. amaranthoides, D. baccata, and D. celosioides are considered specific- ally distinct by SUESSENGUTH (FEDDE, Rep. 44 (1938) 39). Lcannot agree with this view; the differ- ences mentioned by S. are hardly of any importance and moreover are very inconstant in Malaysian materials. 2. Deeringia polysperma (Roxs.) Mog. in DC. Prod. 13, 2 (1849) 236; Mia. FI. Ind. Bat. 1, 1 (1858) 1026; Koorp. Exk. Fl. 2 (1912) 194; Merr. En. Philip. Fl.Pl. 2 (1923) 126; GaGn. in FI. Gén. I.C. 4 (1936) 1056.—Celosia polyperma Roxs. FI. Ind. 2 (1824) 511.—Lestibudesia latifolia BL. Bijdr. (1825) 541.—Celosia latifolia StEuUD. Nom. ed. 2 (1840) 315; Moa. in DC. Prod. 13, 2 (1849) 244; Mia. FI. Ind. Bat. 1, 1 (1858) 1027.—Deeringia in- dica ZOLL. ex Moa. in DC. Prod. 13, 2 (1849) 236; ZOLL. Syst. Verz. (1854) 110; Mia. Fl. Ind. Bat. 1, 1 (1858) 1026; Koorp. Minah. (1898) 565; RIDL. Fl. Mal. Pen. 3 (1924) 5.—Deeringia celosioides (non R.Br. 1810) Hassk. Pl. Jav. Rar. (1848) 436. —Deeringia indica var. pubescens SCHINZ in Bull. Herb. Boiss. II, 3 (1903) 3.—Deeringia polysperma var. pubescens MeErR. En. Philip. FI.Pl. 2 (1923) 126.—Deeringia salicifolia SCHINz in E. & P. Nat. Pfi. Fam. ed. 2, 16c (1934) 27, nomen. Erect herb or undershrub, 1-2 m. Young stems, petioles, leaves, rachises of the spikes, bracts and bracteoles frequently clothed with shortish, thick- ish brown hairs, glabrescent; stem in the higher part obtusangular. Leaves ovate-oblong-lanceolate from a cuneate or shortly contracted base, nar- rowed upwards or slightly acuminate, acute, her- baceous or slightly fleshy, 3-22 by 1!/2-12 cm; midrib in the living plant slightly prominent be- neath; petiole '/2-5 cm. Flowers spicate; spikes axillary, single or sometimes paired, erect or more or less patent, simple or sometimes sparingly branched, 3-12 cm long, rather dense or, at an advanced age, rather lax, 4~-50-flowered. Flowers quite sessile; bracts broadly ovate, obtuse or rather acute, + 1!/2mm long; bracteoles ovate, 1—11/4 mm. Perianth 2'/4—2'/2 mm long. Tepals during anthesis erect or obliquely erect, afterwards appressed to the fruit, very obtuse, green, white-bordered. Stam- inal cup 3/4-1 mm high; free parts of the filaments about as long as the cup or slightly longer. Stigmas 2-3, recurved or on the fruit obliquely erect or patent, !/2-7/3 mm long. Berry globose or broadly ellipsoid, white, + 3 mm diam. Seeds 10-64, usu- ally more than 20, reniform, + 3/4 mm diam, very finely verruculose. Distr. Malaysia: Mal. Peninsula, Sumatra, Java, Philippines, Celebes, Kabaena, Moluccas, and New Guinea. Ecol. In Java, 5-800 m alt. (in the Philippines ascending to 1800 m, according to MERRILL), in thickets and shaded localities, in forests and on forest borders, much rarer than D. amaranthoides. Notes. In a fruiting state conspicuous by its white berries. 3. Deeringia tetragyna Roxs. Fl. Ind. ed. CAREY (1832) 683; WiGurT, Ic. 2 (1843) 729; Mia., Fl. Ind. Bat. I, 1 (1858) 1026. Climbing shrub, entirely glabrous; young shoots pendulous. Leaves ovate from a broad, rounded base, shortly contracted into petiole, shortly acuminate, acute, slightly un- dulate, 3-7cm by 24cm; petiole 3/4—-1!/2 cm. Flowers axillary, spicate or sometimes solitary; spikes erect, shortly stalked, rather less than 1 cm long, few-flowered, dense; bracts ovate-oblong, obtuse, + 1!/2 mm; bracteoles much shorter than perianth, oblong, cvtuse. Tepals 4—5, oval-oblong, obtuse, very concave, 2!/4-21!/2 mm long. Stamens about equalling perianth; staminal cup rather large; free parts of filaments linear from triangular base, about as long as cup or slightly longer. Ovary subglobose, with 3-4 longitudinal furrows, few- ovuled; styles 3-4, recurved, linear-subclavate. Berry (not seen) subglobose, + lobed, red, succu- lent. Seeds 1-4, mostly 1. Distr. Malaysia: Moluccas (according to Rox- BURGH). Notes. This imperfectly known species of which I could examine the type specimen I find to represent a distinct species which is tolerably well figured by WiGurT. Its native country is said to be the Moluccas on the authority of ROXBURGH l.c. who states that it was accidentally introduced from there into the Botanic Gardens, Calcutta. It is strange that it has never been re-collected. 4. Deeringia arborescens (R.Br.) DRruce, Rep. Bot. Exch. Club Br. Isl. 1916, 619 (1917); Domin, Bibl. Bot. 89 (1921) 74; Beitr. Pflanzengeogr. Austr. 1, 2 (1921) 628.—Lestibudesia arborescens R.Br. Prod. (1810) 414.—Celosia arborescens SPRENG. Syst. 1 (1825) 815; Moa. in DC. Prod. 13, 2 (1849) 243.— Lagrezia altissima Moa. in DC. Prod. 13, 2 (1849) 253.—Deeringia altissima F.v.M. Fragm. Phyt. Austr. 2 (1861) 92, Lc. 6 (1864) 251; Bru. Fl. Austr. 5 (1870) 210; HEMsL. Rep. Bot. Chall. 1, 3 (1884) 182; F.v.M. Descr. Not. Dec. 1949] Pap. Pl. 7 (1886) 28; BAILEY, Queens]. FI. pt 4 (1901) 1219.—Cladostachys altissima O.K. Rey. 2 (1891) 541. Woody climber, ascending to top of tallest trees (ex Bru.), glabrous or on young vegetative parts clothed with brown, +- crisped hairs. Leaves oblong or lanceolate from a cuneate or contracted base, obtuse or rather acute, firmly herbaceous, 5—20 by 1'/2-10 cm; petiole 1!/2-3'/2cm. Flowers (6d) (9) (male ones pseudo-hermaphrodite), panicled; sin- gle panicles 4-15 cm long, axillary and terminal; highest often collected in a terminal, rather large panicle; branches of single panicles widely patent, spiciform, dense or rather lax, 1-3 cm long. Flow- ers sessile or subsessile, glabrous; bracts and brac- teoles thinly membranous, nerveless.—d: Bracts ovate-orbicular, rounded, -+ 1 mm long; bracteo- les slightly smaller than the bract, much shorter than the perianth, + 3/4 mm diam; perianth white. Tepals oblong or oblong-obovate, rounded at apex, very convex, nerveless, 1'/2-2 mm long. Adult sta- AMARANTHACEAE (Backer) 73 mens slightly exceeding the perianth; filaments thin, finally much longer than the short staminal cup; anthers oblong, yellow. Ovary shortly stalked, conical, glabrous, empty; style very short; stigmas 3, linear-clavate, short, thick, recurved.—9: Bracts and bracteoles much shorter than perianth, Imm long; bracts ovate-triangular, bracteoles oval..Perianth greenish, 1!/4—1'/2 mm long, at last appressed against ripe fruit. Staminodes 5, varying from slightly shorter to slightly longer than peri- anth; filaments at base connate in a comparatively large and wide cup; their free parts slightly longer than cup; their anthers deformed, small, empty. Ovary subglobose, 6—15-ovuled; style short; stig- mas 3, recurved, shortly linear, thick. Berry glo- bose, red, with the top exserted from the perianth, 3-4 mm diam. Seeds 1—2, reniform, black, densely verruculose, -- | mm diam. Distr. NE. Australia, in Malaysia: collected in Buton Isl. (SE. Celebes), the Tanimbar Isl. (S. Mo- luccas), and SE. New Guinea (Saibai Island) 2. CELOSIA Pmne opel. 1 (1753) 205. Erect, entirely glabrous annuals. Stem angular-ribbed. Leaves alternate, peti- oled, ovate to linear, entire or subentire, often with small semilunar leaves in the axils. Flowers 9, in simple or branched, dense or interrupted, sometimes deformed, terminal or axillary spikes, solitary in axil of bract, subtended by 2 bracteoles. Tepals 5, free, during anthesis erecto-patent or spreading, before and after anthesis erect, ovate-oblong, acute, scarious, longitudinally nerved. Stamens 5; filaments at the base connate in a cup; free parts linear from a triangular base, often alter- nating with minute, triangular pseudo-staminodes; anthers oblong-linear, 2-celled (4-locellate). Ovary sessile with broad base; ovules ~, on short funicles; style 1, filiform, persistent; stigma capitate, faintly 2—3-lobed. Utricle thin-walled, cir- cumsciss in or about the middle. Seeds 1-o, lenticular, shining black. Distr. About 60 spp. mainly in the subtropics and temperate regions of Africa and America, in Malaysia: no indigenous species. In Malaysia one wild species, and several more or less deviating forms which are cultivated for orna- mental purposes. 1. Celosia argentea LINNE, Sp.Pl. (1753) 205; BL. Bijdr. (1825) 543; DecNE in Nouv. Ann. 3 (1834) 372; BLANCO, FI. Filip. (1837) 192; ed. 2 (1845) 135; ed. 3, 1 (1877) 243; SPAN. in Linnaea 15 (1841) 345; Moa. in DC. Prod. 13, 2 (1849) 242; Hassk. in Pl. Jungh. (1852) 128; Mia., Fl. Ind. Bat. I, 1 (1858) 1028; Hook. f. Fl. Br. Ind. 4 (1885) 714; BaAILey, Queensl. Fl. pt 4 (1901) 1218; Laur. & ScHum. Fl. D. Schutzg. (1901) 303; PULLE in Nov. Guin. 8 (1910) 351; Koorp. Exk. FI. 2 (1912) 194; MeRR. Fl. Man. (1912) 190; Ascn. & Gr. Syn. 5, 1 (1913) 222; MERR. Interpr. Herb. Amb. (1917) 212; En. Born. Pl. (1921) 245; En. Philip. FI.Pl. 2 (1923) 127; Ripe. Fl. Mal. Pen. 3 (1924) 5; HEYNgE, Nutt. Pl. (1927) 604; Backer, Onkr. Suiker. (1930) 216, Atl. t. 226; OCHSE & BAKH. V. D. B. Veget. (1931) 27, fig. 17, 28, fig. 18; GAGN. in Fl. Gén. I.C. 4 (1936) 1056.—Celosia cristata LINNE Sp.Pi. (1753) 205, BLANCO, FI. Filip. (1837) 191, &c.; Koorpb. Exk. Fl. 2 (1912) 195; HEYNE, Nutt. Pl. (1927) 605. —Celosia coccinea LINNE, Sp.Pl. ed. 2 (1762) 297; BLANco FI. Filip. ed. 2 (1845) 134; op cit. ed. 3, 1 (1877) 241, t. 64, &c.—Celosia pyramidalis BURM. f. FI. Ind. (1768) 65, t. 25, fig. 1.—Celosia huttonii MAsT. in’) Gard: ‘Chront, 18725) 215s ASCH-mas GRAEBN. Syn. 5, 1 (1913) 222. Forma spontanea: Annual, 0.4-1'/2 m; stem erect, green or red, strongly ribbed, often much branched. Leaves on petioles of '/4-13/4 cm or highest almost sessile, oblong-lanceolate or lan- ceolate-linear, rarely ovate-oblong, acute at both ends, herbaceous, often tinged with red, 4-18 by 3/4—61/2 cm; highest often very small; leaf axils often provided with + falcate small leaves. Spikes solitary or sometimes paired, erect, stalked or partly subsessile, often much lengthening during anthesis, at length cylindrical with a conical apex, very dense, throughout their length (when pure- 74 FLORA MALESIANA [ser. I, vol. 42 bred) with 9 flowers, usualy simple, sometimes bifid or trifid at apex, 2—22 by 1—13/4 cm; their stalk ribbed-furrowed, often ri ict during anthe- sis, finally !/2-21 cm. Flowers solitary, sessile, obli- quely patent; bracts and bracteoles persistent after fall of the perianth, ovate-oblong, pellucid, 1-nerv- ed, mucronate, 3-7 mm long. Perianth 6-10 mm, at first shining white with a pink tip or almost en- tirely pink, withering white. Adult staminal cup 1!/2-2 mm high; free part of filaments 2!/2-3 mm; pseudo-staminodes minute, triangular. Style violet, 3'/4a—5 mm. Utricle included by the perianth, obo- void with rounded apex, + 3!/2 mm long. Seeds 1-9, 1!/4-1!/2 mm diam. Distr. Ubiquist, in Malaysia: wild but not na- tive, introduced here perhaps very long ago (from trop. Africa?), thoroughly established in the settled areas throughout the Archipelago. Ecol. In Java, 1-700 m alt., a rather common weed of open dry localities, field, gardens, waste places, locally often numerous. Uses. Seeds used by the Chinese for poultices and for adorning cakes. The leaves furnish an inferior vegetable. Vern. Borotjo, S. Notes. Forms cultivated for ornamental pur- poses are found throughout Malaysia; not unfre- quently such forms are met with as garden escapes. These latter may breed true but often also they display to a great and variable degree a regression to the wild form, possibly due to intercrossing with it. One branch of a plant may bear the marks of cultivation (in the shape of sterile flowers) whilst another is quite alike the wild form. In cultivated forms the higher flowers of the in- florescence are sterile, the rest fertile; the fertile flowers being usually somehat smaller than those of the wild form; the entire inflorescence is usually red, violet, yellow or orange. It may have the shape of an almost sessile cock’s comb with a thick sinuous crest; this form usually goes under the name of C. cristata L. This is a very common orna- mental plant in Malaysia where it reaches a height of 0.4—-1'/2 m; the inflorescence is usually bright red; the ovate leaves often bear a large bright red blotch. In another series of cultivated forms (forma plumosa (Voss.) BACK.) the inflorescence is loosely branched; the branches are cylindric and have a much constricted tail-like sterile upper part. The two forms may be found on a single plant; the terminal inflorescence being more or less cock’s- comb-like, the lower ones cylindrical, tail-ended. Excluded Celosia nana BLANCO, FI. Filip. (1837) 192 = Al/- ternanthera celosioides Mog. in DC. Prod. 13, 2 (1849) 360 are according to MERRILL (En. Philip. FLPl. 3 (1923) 135) = Ammannia baccifera L. (Lythr.) Doubtful Celosia cernua Juss.; F.-VILL. Noviss. Append. (1880) 168. Celosia phytolaccaefolia Juss.; F.-VILL. Noviss. Append. /.c. Celosia bicolor BLANCO, FI. Filip. (1837) 191; Me_erR. Spec. Blanc. (1918) 383.—Celosia glauca (non ROTTL.) BLANCO, op. cit. ed. 2 (1845) 135, ed. 3, 1 (1877) 242. The form BLANCco described was according to MERRILL (Enum. Philip. FI.Pl. 2 (1923) 127) certainly no amaranthaceous plant, and may possibly have been a species of Ammannia (Lythrac.) 3. ALLMANIA R.Br. ex WIGHT, in Hook. J. Bot. (1834) 226, t. 128. Erect or ascending annual. Leaves alternate, varying from linear to obovate, entire. Flowers 9, in terminal or (by development ofan axillary branch) leaf-opposed heads; heads stalked or subsessile, composed of crowded, 3—7-fiowered cymes; flowers solitary in the axil of a bract, subtended by 2 bracteoles which partly act as bracts to branches of a cyme. Tepals free or nearly so, erect or erecto-patent, subequal, ovate-lanceolate, pellucid with green or purple midrib. Stamens 5; fila- ments at base connate in a short cup; anthers 2-celled (4-locellate); no pseudo- staminodes. Ovary ovate, compressed, narrowed into style; ovule 1, erect on short, broad funicle; style filiform, stigma capitate, faintly 2-lobed. Utricle enclosed by the perianth, ovoid, compressed, thin-walled, circumsciss below the middle. Seed erect, lenticular, at base with a cupular, 2-lobed, thin aril. Distr. Monotypic, trop. Asia and Malaysia. Notes. There is only one variable species, described under several names. All forms pass gradually into each other. 1. Allmania nodiflora (L.) R.Br. in WALL. Cat. (1832) 6890, nomen; Hook. f. Fl. Br. Ind. 4 (1885) 716; Koorb. Exk. FI. 2 (1912) 195; RipL. Fl. Mal. Pen. 3 (1924) 5; BAcKER, Onkr. Suiker. (1930) 217, Atlas t. 227; GAGN. in FI. Gén. I.C. 4 (1936) 1059. —Celosia nodiflora LINNE, Sp.Pl. (1753) 205; Burm. f. Fl. Ind. (1768) 66.—Allmania albida R. Br. in WALL. Cat. 6981 (1832), nomen; Hook. f. Dec. 1949] Fl. Br. Ind. 4 (1885) 717; Merr. En. Philip. FI.PI. 2 (1923) 127; GaGn. in FI. Gén. I.C. 4 (1936) 1059. —Allmania esculenta R.Br. in WALL. /.c. 6892.— Chamissoa brownei STEUD. Nom. ed. 2 (1841) 344. —Chamissoa javanica HAssk. Fl. Jav. Rar. (1848) 434; Moa. in DC. Prod. 13, 2 (1849) 249.—Cha- missoa albida Moa. in DC. Prod. 13, 2 (1849) 248. —Chamissoa esculenta Moa. l.c. 249.—Chamissoa nodiflora Mart. ex Moa. /.c.; HASsK. in PI. Jungh. (1852) 129; Mra. Fl. Ind. Bat. 1, 1 (1858) 1029; Suppl. (1860) 149.—Chamissoa pyramidalis Moa.' lc. 248.—Allmania_ pyramidalis Koorp. Exk. Fl. 2 (1912) 195. Erect or ascending annual 0.1—0.8 m_ long; taproot long; stem branched from the base or nearly so, solid with thickened nodes, glabrous or obscurely thinly pubescent. Leaves linear, spathu- late, oblong or obovate, narrowed into petiole, acute, obtuse, rounded or abruptly shortly acumi- nate, mucronate, glabrous or on undersurface thinly pubescent, rather fleshy, 1!/2-6!/2 by !/3- 21/2 cm; petiole 2-10 mm. Heads terminal or leaf- opposed, at first subglobose, afterwards somewhat lengthened, 3/s-2 cm long; their stalk 2-35 mm, rather robust, scantily hairy or glabrous; single cymes sessile, 3—7-flowered; bracts and bracteoles ovate-lanceolate, long acuminate, keeled, 3-5 mm long; midrib green or purple; margins shining AMARANTHACEAE (Backer) i i white; keel scaberulous outside. Tepals before and after anthesis erect, during anthesis obliquely spreading, with strong, green or purple midrib and pellucid white, shining margins, glabrous or scab- erulous on back, 4-5 mm long. Stamens shorter than perianth. Ovary and style glabrous; style dur- ing anthesis + 2mm, on fruit (conical base in- cluded) + 3 mm; stigma about as high as anthers. Utricle glabrous, +- 3'/2 mm; seed shining black, + 2'/2 mm diam.; aril enclosing base of seed, pale pink or pale brown. Distr. Tropical Asia; in Malaysia: Singapore, Sumatra, Java, Madura, Sumba, Wetar & the Phil- ippines. Ecol. In Java from the plains up to + 100m alt. (very rarely higher), locally often a rather com- mon weed on light especially sandy soils, sandy shores, fields, roadsides and dunes. Uses. Long ago reported to be eaten at Singa- pore. Notes. Often split into 2 species: A. nodiflora with sessile heads and A. pyramidalis with pedun- cled ones. This difference exists only on paper, not in nature where the two forms pass into each other. I never saw a form with quite sessile heads. SCHINZ (in E. & P. Nat. Pfl. Fam. ed. 2, 16c, p. 33) figures A. nodiflora with clearly peduncled heads but he described them as sessile. 4. AMARANTHUS LINNE, Sp.Pl. 1 (1753) 989. Annuals, erect or wholly or partly decumbent, unarmed or spinous. Leaves alter- nate, entire. Flowers (d Q) in sessile, small, dense clusters, clusters axillary or collected in axillary and terminal, solitary or panicled spikes. Flowers solitary in the axil of a bract, sustended by 2 bracteoles; bracts and bracteoles small, scarious. Tepals 3 or 5, rarely 4, erect or obliquely patent, free, subequal, membranous, green, purple, or pellucid with a green or purple median band, after anthesis sometimes indurate at base. Stamens as many as tepals; filaments free, filiform, no pseudo- staminodes; anthers 2-celled (4-locellate). Ovary ovate or oblong, ovule 1, sessile, erect; style short or none; stigmas 2-4, often 3, erect or spreading-recurved, linear. Utricle laterally compressed, membranous, circumsciss when ripe or bursting irregularly or falling off unopened together with the perianth; seed erect, lenticular, shining black or brown. Distr. About 40 spp. aliover the world, specially developed outside the tropics, several elsewhere intro- duced. Most of the Malaysian species are ubiquists; of some the native country is unknown. Of the 7 wild Malaysian species one is frequently also cultivated; the two others are almost exclusively cultivated but occasionally met with as strays from gardens. Ecol. Weeds of waste places, roadsides, fields and gardens, locally sometimes gregarious. Uses. Some species serve as vegetables; some are used medicinally or for ornamental purposes. Vern. In the Malay language all species are called bayam (with various additions). Notes. Probably several of the Malaysian species have been introduced. KEY TO THE SPECIES 1. Utticles,? also when adult and quite ripe, indehiscent or at last bursting irregularly, falling off together with the perianth. Tepals very shortly mucronate. Stigmas very short ('/4—/3 mm), erect or suberect. Bracts and bracteoles shorter than the perianth. Unarmed. (1) Not Celosia pyramidalis Burm. f. which is Celosia argentea L. (2) See footnote next page. 76 FLORA MALESIANA [ser. I, vol. 42 2. Tepals in all flowers 3, exceptionally 4. 3. Ripe utricle very strongly corrugated, seed-containing part entirely included by the perianth, from which only the seedless conical beak emerges 1. A. gracilis 3. Ripe utricle smooth or as rugulose, top of the seed- ‘containing part slightly emereing from the perianth 2. A. lividus 2. Tepals 5. Ripe utricles ina dried state faintly ‘longitudinally ribbed and + + rugulose 3. A. interruptus 1. Adult guite ripe utricles! circumsciss a little below the middle. Perianth and cup-shaped base of the utricle persisting till after the fall of the lid and the seed. Stigmas 3/4—2'/2 mm, often recurved when long. 4. Tepals in all or most flowers 5, less often 4, rarely (only in a few flowers) 3, shortly mucronate. 5. Flower clusters for a great part solitary in the higher leaf axils, for the rest collected in spikes or panicles which terminate the main stem and its branches. Bracts not longer than the perianth. 6. Axillary flower-clusters exclusively 9; those of the spikes and panicles for the greater part or almost entirely ¢. Midrib of the tepals not much broadened upwards. 7. Style 11/s-11/2 mm long. 9 clusters usually armed with 2 very sharp spines, rarely with 1 spine only or unarmed. Midrib of the tepals green or purple 5. A. spinosus 7. Style 1/4-!/2 mm long. Flower-clusters unarmed. Midrib of the tepals green. Small plant 6. A. leptostachyus 6. Terminal spike of the main stem almost entirely 9, other spikes entirely 6, or at their base d, higher up 9. Midrib of the tepals green, in the lower half very thin, in the upper half (up to quite near the apex) much thickened. Unarmed 7. A. dubius 5. Flower clusters all collected in terminal and axillary panicles of spikes: Bracts often longer than the perianth. 8. Tepals of 9 flowers not or hardly overlapping, 1/3—!/2 mm wide. Panicles erect, or nodding only in their upper half. Cultivated and occasionally met with as an escape from gardens 8. A. hybridus 8. Tepals of 9 flowers distinctly overlapping for the greater part of their length, 2/3—/4 mm wide. Panicles (in Malaysian specimens) drooping almost from the very base. Possibly exclusively cultivated. 4. Tepals in all flowers 3, Sravided With a lone apicalh awn 1. Amaranthus gracilis Desr. Tabl. Ec. Bot. (1804) 43; THELLUNG in AscH. & Gr. Syn. 5,1 (1914) 335; Domimn. Beitr. Pflanzengeogr. Austr. 1, 2 (1929) 634; Heyne, Nutt. Pl. (1927) 605; BAcKER, Onkr. Suiker. (1930) 222, Atl. t. 232; OCHSE & BAKH. v. D. BR. Veget. (1931) 18.—Amaranthus viridis LINNE, Sp.Pl. ed. 2, 2 (1763) 1405, ex parte; Bru. FI. Austr. 5 (1870) 215; Hook. f. Fl. Br. Ind. 4 (1885) 720; BAILEY, Queensl. Fl. pt. 4 (1901) 122; PULLE, Nova Guinea 8 (1910) 351; Koorp. Exk. Fl. 2 (1912) 197; MerR. Interpr. Herb. Amb. (1917) 212; En. Born. (1921) 246; En. Philip. F1.Pl. 2 (1923) 128; GAGNn. in Fl. Gén. I.C. 4 (1936) 1064.—Che- nopodium caudatum JAcQ. Coll. 2 (1788) 235 (non Amaranthus caudatus L.).—Amaranthus polysta- chyus WILLD. Sp.Pl. 4 (1805) 385; BrLume, Bijdr. (1825) 538; Koorp. Exk. Fl. 2 (1912) 197.— Euxolus caudatus Moa. in DC. Prod. 13, 2 (1849) 274; Mia. Fl. Ind. Bat. 1, 1 (1858) 1036.—Euxo- lus polystachyus Mia. Fl. Ind. Bat. 1, 1 (1858) 1036. Annual, erect, ascending or rarely prostrate, 10-75 cm long, often much branched, unarmed; stem terete-obtusangular, glabrous or thinly pub- escent. Leaves (larger ones at least) rather long- petioled, ovate-rhomboid-oblong from obtuse or cuneate, often decurrent base, acute, obtuse, round- ed or retuse, glabrous or on stronger nerves spar- 9. A. caudatus 4. A. tricolor ingly pubescent, green; larger ones 3-9 by 2-6!/2cm. Flowers green; lower clusters axillary; upper ones in terminal, rather dense, continuous or interrupted (39) spikes or panicles; bracts and bracteoles ovate, minutely mucronate, shorter than adult peri- anth; tepals 3 (exceptionally 4), very shortly mu- cronate with transparent white margins and green median band, glabrous, very convex, in d oblong- linear, + 1!/2mm long, in 9 narrowly oblong- spathulate, during anthesis 2/3-1 mm, when fruit- ing 11/4-13/s mm long; co flowers cften with rudi- mentary filiform ovary; ovary in Q oblong; stigmas 2-3 on conical top of ovary, erect or suberect, !/4— 1/3 mm. Utricle falling off together with the peri- anth, about as long as this, only the subconical short beak emerging, very strongly corrugated, + 11/2 mm long, indehiscent or at last bursting irre- gularly. Seed with a blunt margin, shining brown or black, 1—11!/4 mm diam. Distr. Tropical ubiquist, in Malaysia: through- out the Archipelago. Ecol. In the lower regions, especially below 600 m alt., a very common weed of cultivated areas, also in waste places, locally often abundant. To my knowledge never cultivated in Malaysia. Uses. In the Moluccas used as a food. Vern. Besides the general name bayam many local names. (1) Herbarium specimens have not rarely been collected before the utricles were quite ripe, some- times when they were still very young. In such cases a normally circumsciss utricle often seems to burst irregularly (by pressure). J Dec. 1949] AMARANTHACEAE (Backer) 3 2. Amaranthus lividus LINNE, Sp.PI. 1 (1753) 990; THELLUNG in AscH. & Gr. Syn. 5, 1 (1914) 319; HeyYNeE, Nutt. Pl. (1927) 605; BACKER & SLoor. Theeonkr. (1924) 107, t. 107; BACKER, Onkr. Sui- ker. (1930) 220, Atl. t. 231.—Amaranthus viridis LiInne, Sp.Pl. ed. 2, 2 (1763) 1405, ex parte.— Amaranthus blitum (non L.) Mia. Fl. Ind. Bat. 1 (1858) 1033; Bru. FI. Austr. 5 (1870) 213; Hook. f. Fl. Br. Ind. 4 (1885) 721; OCHse & BAKH. V. D. Br. Veget. (1931) 17, fig. 11.—Euxolus lividus Moa. l.c. 273. Annual, erect of prostrate, 5-80 cm long, often much branched (frequently from very base), un- armed; stem terete-obtusangular, quite glabrous. Leaves (larger ones at least) rather long petioled, obovate or + rhomboid, rarely oblong, from cune- ate base, with broadish, usually deeply emarginate mucronate apex, green or more or less suffused or blotched with purple or entirely purple; larger ones 3-6 by 2-4 cm. Lower flower-clusters axillary, higher ones on older vigorous plants always col- lected in terminal and axillary spikes or panicles; bracts and bracteoles ovate, acute, much shorter than adult perianth. Tepals 3, very shortly mu- cronate, with transparent margins and green or purple median band, very concave, in d oblong, + 11/4 mm long, in 9 oblong-spathulate, slightly accrescent with age, 1!/4-11/4 mm. Filaments equal- ling the perianth or slightly shorter. Rudimentary ovary in 6 often present, filiform; ovary in 9 oblong; stigmas 2-3, erect or suberect, !/3—*/3 mm long; adult utricle broadly ellipsoid, acute, laterally compressed, slightly exceeding perianth, smooth or faintly rugulose when ripe, 1!/2-2 mm long, fal- ling off together with perianth, indehiscent or at last bursting irregularly; seed wit a rather blunt margin, shining black or blackish brown, 1-1'/4 mm diam. Distr. In Malaysia: Sumatra, Java, Celebes & Philippines, probably also elsewhere. Ecol. In Java from the lowlands up to + 2000 m, a very common weed in cultivated and waste places. Uses. Used by the Indonesians as a food. Vern. Bayam. Moreover, many local names. Notes. This is a very variable species. 3. Amaranthus interruptus R.Br. Prod. (1810) 414; Bru. Fl. Austr. 5 (1870) 215; BAILEY, Queensl. FI. pt 4 (1901) 1221.—Amaranthus spiratus Zipp. ex SPAN. in Linnaea 15 (1841) 345, nomen.—Euxolus interruptus Moa. in DC. Prod. 13, 2 (1849) 275; F.v.M. Descr. Not. pt 5 (1878) 87. Annual, erect or ascending, 40-60 cm long, al- - most simple or in higher part with (sometimes many) obliquely erect branches, unarmed; stem straight or slightly flexuous, obtusangular, glabrous or very thinly clothed with minute patent hairs. Leaves (larger ones at least) rather long petioled, oblong from long-cuneate base, much narrowed in upper half, obtuse or slightly emarginate, shortly - mucronate, with oblique prominent primary lateral nerves; glabrous; larger leaves 3-6 by 11/4—2!/2 cm. Flower-clusters dense; lower ones axillary; higher ones collected in continuous or more or less inter- rupted spike; terminal spike simple or branched; flowers in upper part of panicle-branches often exclusively d, lower down largely or exclusively 2; bracts and bracteoles shorter than perianth, mucronate on broadly oval transparent base. Flowers green, all of them (in the Timor-specimen) 5-merous. Tepals of ¢ oblong, acute or minutely mucronate, 1!/3-1'/2 mm long; tepals of 9 narrowly spathulate, very shortly mucronate, slightly ac- crescent with age, 1!/s—-13/4 mm long. Styles 2-3, erect !/3-!/2 mm. Urricle falling off together with the perianth, broadly ellipsoid, rather thick, in a dried state faintly longitudinally ribbed and rugu- lose, tipped above the seed with a sharply delimi- tated, broad, obtuse cone (bearing the unaltered styles), indehiscent or in dried materials bursting irregularly. Seed thick, with an obtuse margin, shining brownish black, + 1 mm diam. Distr. Eastern Australia, in Malaysia: Timor (SPANOGHE) and SE. New Guinea (LAWEs; Port Moresby, CHALMERS, TURNER; Rigo distr. MAC- GREGOR). Notes. Sometimes confused with unarmed forms of Amaranthus spinosus L. which may at once be recognized by the much longer (1 '/4—1!/2mm) styles. It has also been confused with Amaranthus leptostachyus BTH. with differs by the circumsciss smooth utricles. 4. Amaranthus tricolor LINNE (sens. ampl.) Sp.PI. (1753) 989; THELLUNG in AscH. & Gr. Syn. 5, 1 (1914) 272: Merr. Interpr. Herb. Amb. (1917) 213; En. Philip. FI.Pl. 2 (1923) 128; Herne, Nutt. Pl. (1927) 606; BACKER, Onkr. Suiker. (1930) 220, Atl. t. 230; OcHsE & BAKH. V. D. BR. Veget. (1931) 25.—Amaranthus melancholicus LINNE, Sp.PI. (1753) 989; Moa. in DC. Prod. 13, 2 (1849) 262; Mio. FI. Ind. Bat. 1, 1 (1858) 1032.—Amaranthus mangostanus LINNE, Cent. PI. 1 (1755) 32; Moa. in DC. Prod. 13, 2 (1849) 261; Mia. FI. Ind. Bat. 1, 1 (1858) 1032; Hook. f. FI. Br. Ind. 4 (1885) 720; Merr. Interpr. Herb. Amb. (1917) 213; En. Born. (1921) 245; Ripv. Fl. Mal. Pen. 3 (1924) 6; GAGN. in Fl. Gén. I.C. 4 (1936) 1062.—Amaranthus poly- gamus LINNE, Cent. Pl. 1 (1755) t. 32.—Amaran- thus gangeticus LINNE, Syst. ed. 10, 2 (1759) 1268; Moo. in DC. Prod. 13, 2 (1849) 261; Mia. FI. Ind. Bat. 1, 1 (1858) 1032; Hook. f. Fl. Br. Ind. 4 (1885) 719; Koorp. Exk. FI. 2 (1912) 196; Ripi. Fl. Mal. Pen. 3 (1924) 6; GAGN. in Fl. Gén. I. C. 4 (1936) 1063.—Amaranthus oleraceus (non LINNE) BURM. f. FI. Ind. (1768) 198 (sphalm. 298); BLUME, Bijdr. (1825) 539; DEcNE in Nouv. Ann. Mus. 3 (1834) 371; SPAN. in Linnaea 15 (1841) 345; Mia. Fl. Ind. Bat. 1, 1 (1858) 1033; Suppl. (1860) 149.—Ama- ranthus salicifolius Hort. VEITCH ex Gard. Chron. (1871) 1550, fig. 331; Merr. En. Philip. F1.Pl. 2 (1923) 129.—Fig. 2. Annual; stem under cultivation erect and often very robust, up to 1!/2-2!/2 m high, in a wild state usually much smaller, erect or ascending, angular, glabrous or in higher part thinly pubescent. Leaves (larger ones at least) long-petioled, rhomboid- ovate-oblong-lanceolate from a cuneate or acute, often decurrent base, narrowed in upper part, acute, 78 FLORA MALESIANA [ser. I, vol. 42 obtuse, rounded, retuse or emarginate, glabrous or on larger nerves thinly pubescent, in wild specimens entirely green, in cultivated (for orna- : NAPA \ te Cra Y¥ FO je. ie Ny J a ee yi NW A Tt St NI] SS S A W ISN ISSA yy LI EZ ES NT SSIs ™~ Ss AN RSS Fig. 2. Amaranthus tricolor L. from Java, x 1/6. ment) forms often tinged or blotched with purple or entirely purple, sometimes bright red with yel- low; larger leaves 10-25 by 3-12 cm. Flower-clus- ters dense; lower ones axillary, higher ones often collected in rather thick spike; ¢ and 9 flowers intermixed; bracts and bracteoles long-awned from broad base, as long as adult perianth or shorter. Tepals 3, long-awned from broad base, with broad transparent margins and green or purple median band, in d 3!/2-6 mm long, in 9 at first 2-3 mm, under the ripe fruit 3-5 mm. Filaments about as long as perianth or shorter, often much shorter; ovary cylindrical or obconical; styles 3, 2—2!/2 mm long. Urricle flask-shaped, circumsciss somewhat below middle; lid with thickened base and sud- denly contracted, conical, obtuse apex. Seed with a rather obtuse margin, shining blackish brown or brown, 1—1!/4 mm diam. Distr. Ubiquist, possibly native in trop. Asia, in Malaysia: throughout the Archipelago. Ecol. Very frequently cultivated as a pot-herb, often run wild in waste places, in fields, along road- sides, locally often abundant, 1—700 m. Uses. Cultivated green-leaved forms very fre- quently eaten by Europeans and non-Europeans as a Substitute for spinach. Variegated-leaved forms sometimes kept in gardens as ornamentals. Vern. Bayam. Moreover, local names. 5. Amaranthus spinosus LINNE, Sp.Pl. (1753) 991; Bur. f. Fl. Ind. (1768) 200 (sphalm. 300); BLUME, Bijdr. (1825) 540; DecNeE, Nouv. Ann. Mus. 3 (1834) 371; BLaNnco, FI. Filip. (1837) 710; ed. 2 (1845) 491; ed. 3, 3 (1879) 113; SPAN. in Linnaea 15 (1841) 345; Hassx. Pl. Jav. Rar. (1848) 432; Moa. in DC. Prod. 13, 2 (1849) 260; Mia. FI. Ind. Bat. 1, 1 (1858) 1031; Hook. f. Fl. Br. Ind. 4 (1885) 718; Koorp. Minah. (1898) 564; BAILEY, Queensl. Fl. pt 4 (1901) 1220; Koorp. Exk. Fl. 2 (1912) 195; THELLUNG in AscH. & Gr. Syn. 5, 1 (1914) 267; Merk. Interpr. Herb. Amb. (1917) 213; En. Born. (1921) 245; En. Philip. FI.PI. 2 (1923) 128; Domin, Beitr. Pflanzengeogr. Austr. 1, 2 (1929) 629; RID. Fl. Mal. Pen. 3 (1924) 6; HeyYNe, Nutt. PI. (1927) 606; BAcKeErR, Onkr. Suiker. (1930) 220, Atl. t. 229; OCHSE & BAKH. V. D. Br. Veget. (1931) 23; GAGN. in Fl. Gén. I.C. 4 (1936) 1062. Annual, erect, often much branched, 15-100 cm high; stem terete or obtusangular, green or more or less suffused with purple, glabrous or slightly pu- bescent. Leaves (largerones at least) rather long petioled, ovate-oblong-lanceolate, from acute, often slightly decurrent base, in their upper part gradually narrowed, obtuse, rounded or slightly retuse, often shortly mucronate, glabrous or, when young, slightly pubescent on the nerves; larger ones 3!/2-11 by 11/4—4!/2 cm. Flower-clusters dense; lower ones axillary; higher ones often collected in axillary and terminal spikes; spikes often branched in their lower part; terminal spike above base usu- ally wholly 3, with weak spines or quite unarmed, finally often with a drooping apex; axillary clusters and those on the base of the spike (rarely also the higher ones) usually armed with 2, or sometimes more, obliquely erect or patent, straight, thin, very Dec. 1949] sharp, !/2-2cm long spines (metamorphosed bracts), sometimes with one spine only, rarely (var. inermis SCHINZ) unarmed; bracts and bracteoles mucron- ate from a broad base, shorter than the adult peri- anth or at best as long. Tepa/ls 5, shortly mucron- ate, very convex, with transparent margins and green or purple median band, in d ovate-oblong, 2—2!/2 mm long, in 9 oblong-spathulate, at first 1!/4-11/2 mm, under the ripe fruit 13/4—23/4 mm long. Filaments about equalling perianth or slightly longer or shorter. Ovary oblong; styles mostly 3, sometimes 2, when adult recurved and 1!/4—1!/2 mm long. Utricle oblong, with a 3-lobed apex, circum- sciss a little below the middle. Seed with a thin margin, shining black or brownish black, + 1 mm diam. or slightly larger. Distr. Ubiquist, in Malaysia: throughout the Archipelago. Ecol. At present throughout Java, from the lowlands up to + 1400 m, a very common weed of waste places, railway-yards, waysides, fields and gardens, often gregarious. Uses. Used as a diuretic, an emmenagogue and a lactagogue, further for poultices and against gonorrhea. Vern. Bayam duri (i.e. spinous bayam). More- over many local names. Notes. Possibly an introduced species. In Ma- laysian specimens ripe adult utricles always open circumsciss a little below the middle. 6. Amaranthus leptostachyus BrH. Fl. Austr. 5 (1870) 214; BAmey, Queens]. Fl. part 4 (1901) 1220; Domin, Beitr. Pflanzengeogr. Austr. 1, 2 (1921) 630. Annual, not or sparingly branched, 7!/2-25 mm high; stem obtusangular, glabrous or very sparing- ly beset with patent minute hairs. Leaves (larger ones at least) long petioled, ovate-oblong from a cuneate base, in their upper part gradually nar- rowed to an obtuse or acute, minutely mucronate apex, glabrous or beneath on the nerves with few scattered, very minute, patent hairs, 1-6 by 1/3-21/2 cm; primary nerves in dried specimens distinctly prominent beneath; petiole ‘/4—6 cm. Flower-clusters rather dense; lower ones axillary, consisting almost or entirely of Q flowers; higher clusters collected in axillary and terminal spikes, often forming together a terminal panicle, con- sisting mainly of d flowers, in the lower part often intermixed with Q ones; bracts and bracteoles mucronate, shorter than the perianth. Tepals usu- ally 4-5, sometimes 3, erect or obliquely patent, oblong-spathulate with a distinct mucro, scarious with a conspicuous but rather thin green midrib, 11/3-11/2 mm long, 1/3—'/2 mm wide. Stamens in d 4-5, not rarely 3. Styles in 9 2—3, erecto-patent or recurved, !/3—!/2 mm long. Utricle tipped by a thick conical beak, crowned by the styles, not rugulose, circumsciss; upper part falling away, leaving the much shorter cup-shaped persistent base in the perianth. Seed erect, lenticular, shining blackish brown, somewhat less than 1 mm diam. Distr. N. Australia, in Malaysia: SE. New Gui- nea (Lorne Range, CHALMERS; Jimari, FITZGERALD). AMARANTHACEAE (Backer) 79 Notes. Sometimes confused with A. interruptus R.Br. which may be easily recognized by the in- dehiscent, in a dry state slightly rugulose utricle and constantly 5-merous flowers. From unarmed forms of A. spinosus L. easily distinguishable by the much shorter styles (in A. spinosus \'/4— 1'/2 mm). 7. Amaranthus dubius MArt. Hort. Erl. (1814) 197; THELL. in Ascu. & Gr. Syn. 5, 1 (1914) 265. Annual, erect, !/2-1 m high, in its upper part often with many obliquely erect branches, unarm- ed; stem obtusangular, green, glabrous or very thinly beset with minute patent hairs. Leaves (larger ones at least) long-petioled, ovate-oblong or ovate- rhomboid from broadly cuneate base, much nar- rowed in the upper part, slightly emarginate, mi- nutely mucronate, glabrous; larger leaves 6—20 by 4-10 cm. Flowers-clusters dense, green, lowest ax- illary; higher ones collected in dense spikes; spikes (especially the terminal ones which frequently reach 10-25 cm in length) in their lower part fre- quently with few or many obliquely patent branch- es, often sinuous, either almost entirely d or at base 6, higher up 9; terminal spike often entirely 9; larger bracts broad, thinly membranous, oval with a long mucro, totalling + 2mm. Perianth 1'/2- 2!'/2 mm long; that of the 9 flowers somewhat in- creasing with age; tepals 4—5, rarely 3, oval-oblong; their midrib in lower half very thin, in upper half (up to quite near apex) much thickened, produced into a short (often very short) mucro. Filaments short. Ovary shortly 3-lobed; styles 3, 1—1!/4 mm long, often recurved. Utricle ellipsoid, when fully ripe circumsciss in or slightly below middle. Seed with a rather thin margin, brownish black, shining + 11/6 mm diam. Distr. Native of tropical America, of rather recent introduction in Java; collected for the first time in 1922 and repeatedly afterwards at Buiten- zorg, later also at Bandoeng. Not yet found in any other part of Malaysia. Ecol. Weed of gardens, road-sides and waste places, abundantly fruiting, may be expected to spread rapidly. 8. Amaranthus hybridus LINNE subsp. cruentus (L.) THELL. var. paniculatus (L.) THELLUNG in Ascu. & Gr. Syn. 5, 1 (1914) 247; Heyne, Nutt. Pl. (1927) 605; BACKER, Onkr. Suiker. (1930) 219, Atl. t. 228.—Amaranthus hybridus L. Sp.P1. (1753) 990.— Amaranthus cruentus LINNE, Syst. Pl. ed. 10, 2 (1759) 1269, non WILLD. ex Roxs. quod est A. cau- datus L.—Amaranthus paniculatus LINNE, Sp.PI. ed. 2 (1763) 1406; Moa. in DC. Prod. 13, 2 (1849) 257; Mia. FI. Ind. Bat. 1, 1 (1858) 1030; Bru. FI. Austr. 5 (1870) 213; Hook. f. Fl. Br. Ind. 4 (1885) 718; BAILEY, Queensl. Fl. pt 4 (1901) 1220; Koorp. Exk. Fl. 2 (1912) 196; Merr. En. Philip. FI.Pl. 2 (1923) 128; GaGn. in Fl. Gén. I.C. 4 (1936) 106.— Amaranthus speciosus Sims, Bot. Mag. (1821) t. DOT Annual, erect, in higher part often much branch- ed, 0.15—3 m, unarmed; stem obtusangular, strong- ly suffused with purple; younger parts more or less 80 FLORA MALESIANA {[ser. I, vol. 42 densely clothed with short hairs or almost glabrous. Leaves (larger ones at least) long-petioled, ovate- oblong-lanceolate from an acute often short-decur- rent base, in their upper half gradually narrowed, obtuse, shortly mucronate, often with wavy mar- gins, dark green above, strongly tinged with purple beneath; larger ones 10-30 by 3-12 cm; nerves beneath or on both sides more or less densely hairy. Flower-clusters crowded, paniculate or in feeble specimens spicate: panicles (spikes) terminal and frequently also in the higher leaf axils, erect or at the top more or less drooping, puberulous, (dQ); terminal panicle in well-developed specimens 15-40 cm long. Flowers dark purple, 5-merous; bracts and bracteoles long-pointed distinctly long- er than perianth. Tepals oblong, not or hardly overlapping, with a short or very short mucro, 1/3-1/2 mm wide, in d 13/4s—-2 mm long, in 9 during anthesis (mucro excluded) 1!/3—-1!/2 mm long, after- wards up to 2 mm. Filaments white, equalling peri- anth or slightly longer. Ovary in 6 rudimentary, hardly perceptible, in 9 oblong; styles 3, less often 2, + recurved, + 3/4 mm long; utricle exceeding perianth, + urceolate, in the lower half pale, in the upper half purple, circumsciss between the pale and the purple parts. Seed dark brown, shining, 1—11/4 mm diam. Distr. Native country unknown, introduced into Malaysia very long ago: Sumatra, Java, Lesser Sunda Islands. Ecol. In Java from the lowlands up to + 1300m cultivated for ornamental purposes and sometimes met with as a stray from gardens, but nowhere firmly established, not truly naturalized. Uses. in Malaysia unknown. Vern. Bayam kéjong, J, and a few local names. 9. Amaranthus caudatus LINNE, Sp.P!. (1753) 990; Moa. in DC. Prod. 13, 2 (1849) 255; Mia. Fl. Ind. Bat. 1, 1 (1858) 1030; Hook. f. Fl. Br. Ind. 4 (1885) 719; BAILEY, Queens]. Fl. pt 4 (1901) 1220; Koorp. Exk. Fl. 2 (1912) 196; THELLUNG in Ascu. & Gr. Syn. 5, 1 (1914) 231; Ripv. Fl. Mal. Pen. 3 (1924) 6; GAGN. in FI. Gén. I.C. 4 (1936) 1061. Annual, erect, not or sparingly branched, 0.3— 1'/2 m, unarmed; stem obtusangular, usually suf- fused with purple, thinly beset with patent short hairs. Leaves (larger ones at least) long petioled, rhomboid-ovate-lanceolate from cuneate base, in their upper half gradually narrowed, obtuse, mu- cronate, green, often bordered with purple; dimen- sions?; nerves beneath usually pale. Flower-clusters very densely spicate; lower panicled; panicle in Malaysian specimens borne by a flaccid peduncle, drooping from the very base; terminal spike fre- quently much longer than the others; bracts and bracteoles broad, with a long apical point; many slightly exceeding the flowers. Perianth 5-merous. Tepals in gd ovate-oblong, shortly mucronate, in 9 oblong-obovate-subspathulate, with at least partly overlapping margins, 2—2!/2 mm (including the often rather long mucro) by 2/3—3/4 mm, purple. Styles 3, + 3/4 mm long. Urtricle slightly exceeding perianth, lageniform, circumsciss. Seed 1—11/4 mm diam; dark brown, shining. Distr. Ornamental plant of old, native coun- try not known with certainty, in Malaysia but rare- ly cultivated. Notes. The only Malaysian specimen I have seen had been collected in NE. Sumatra above Si- bolangit, -- 1350 m); it may have been taken from a cultivated plant. 5. DIGERA Forsk. Fl. Aeg.—Arab. (1775) 65. Annual. Leaves alternate, petioled, entire or subentire. Flowers in axillary pe- duncled spiciform racemes; lower part of raceme in each axil of persistent bracts with 3 flowers on very short common stalk; central flower of triad perfect, 9, 2- bracteolate; tepals 5, almost free; 2 outer ones larger than the 3 other ones and together embracing them. Stamens 5; filaments free, filiform; no pseudo-stami- nodes; anthers oblong, 2-celled (4-locellate). Ovary obovoid, truncate; ovule 1, erect; style filiform, rather long; stigmas 2, recurved, linear, short. Lateral flowers in the axil of bracteoles of the fertile flower, reduced to a stalked palmatifid scale; scales towards the apex of the raceme gradually smaller, in the highest flowers absent. Utricle falling off together with the enclosing perianth, bracteoles and scales, rugulose-tuberculate, with keeled sides ending at the top in a small hornlet, crustaceous, indehiscent. Seed erect, exarillate. Distr. Monotypic; northern Africa through the Orient and S. Asia to Malaysia. Notes. In feeble specimens the sterile lateral flowers are sometimes absent. 1. Digera muricata (L.) MArT. Beitr. Amar. (1825) 77, no 2.—Achyranthes muricata LINNE, Sp.Pl. ed. 2 (1762) 295.—Achyranthes alternifolia Mant. (1767) 50; Roxs. Fl. Ind. ed. CAREy 1 (1832) 674.—Digera arvensis Forsk. Fl. Aeg.—Arab. (1775) LINNE, 65; Moa. in DC. Prod. 13, 2 (1849) 324; HAssk. in Pl. Jungh. (1852) 132; Zoi. Syst. Verz. (1854) 109; Mia. FI. Ind. Bat. 1, 1 (1858) 1044; Hook. f. Fl. Br. Ind. 4 (1885) 717; Koorp. Exk. Fl. 2 (1912) 197.—Cladostachys frutescens D. Don, Prod. Dec. 1949] (1825) 76; Mia. Fl. Ind. Bat. 1, 1 (1858) 1025.— Digera forskaoli Br. Bijdr. (1825) 542; HAssk. PI. Jav. Rar. (1848) 425; Mor. Syst. Verz. (1845/6) 73; ZOLL. Syst. Verz. (1854) 109.—Desmochaeta muri- Fig. 3. Digera muricata (L.) Mart. from Java, x 1/4. AMARANTHACEAE (Backer) 81 cata WiGHurT, Icon. (1843) t.732.—Cladostachys mu- ricata Mog. in DC. Prod. 13, 2 (1849) 235.—D. alternifolia AscH. in SCHWEINF. Beitr. Fl. Aeth. (1867) 180; AscH. & Gr. Syn. 5, 1 (1914) 357; BACKER, Onkr. Suiker. (1930) 223; Atl. t. 233.— Fig. 3. Annual, often branched from base; small speci- mens: erect; larger ones prostrate-ascending or with widely patent, prostrate-ascending, often long branches, 0.15—1.6 m long; stem often flexu- ous, glabrous or slightly pubescent. Leaves ovate or ovate-oblong from a cuneate, obtuse, rounded or subcordate base, with an acute, obtuse or rounded apex, entire or obsoletely crenulate, her- baceous, glabrous, 1!/2-7!/2 by 3/s—-5!/2 cm; petiole 3/4-5 cm. Racemes solitary, widely patent-ascend- ing, in upper part dense, lower down rather lax, 1-30 cm_ long ('/4-7!/2 cm of peduncle included), glabrous or subglabrous; lowest fruits often falling off before expansion of the highest flowers; bracts widely patent, ovate-lanceolate, concave with a strong midrib and broad scarious margins, glabrous (as are pedicels, bracteoles, scales and perianth), 2!/2-3 mm long, persistent; pedicels very short; bracteoles appressed against sterile flowers or in the absence of these against the perianth, ob- long with scarious margins, 2—2!/2 mm long. Sterile flowers appressed against the fertile flower, flat, much dilated from a stalk-like base, palmatifid, towards the apex of the raceme gradually smaller, in the highest flowers absent. Tepals during anthesis more or less patent, afterwards erect; 2 outer ones 31/2-41/4 mm long, concave, 5—7-nerved, green with whitish or pink borders; 3 inner ones shorter, much narrower, thinner, obtuse, pink, 1—-2-nerved. Adult filaments much longer than anthers. Style (short stigmas excluded) 2—2!/2 mm; perianth after anthesis not or hardly accrescent. Fruit compressed- globose, between the apical hornlets slightly depressed, bearing a persistent style-base, 22114 mm diam. Distr. N. Africa through the Orient to southern Asia, in Malaysia: Java, Madura, Kangean Arch., Celebes, Moluccas, Sumba and Sumbawa. Ecol. Obviously preferring the drier areas, 1-250 m, in fields (especially when sandy), along road-sides, railway-embankments, waste places, usually in scattered specimens. Vern. Bayam sidit, J. Notes. The specimen preserved in BURMAN’S collection (Herb. DeLesserT, Geneva) named Achyranthes muricata and mentioned in BURM. F- FI. Ind. (1768) 63 is Amaranthus gracilis DEsF. The true Digera muricata lies in Herb. BURMAN as ‘Blitum, malaice Baian clatek’ (bajam glatik). 6. CYATHULA BLumE, Bijdr. 11 (1825) 548, nom. consery., non LOUR. Perennial herbs or undershrubs. Leaves opposite, entire. Flowers clustered; clus- ters either singly along the rachis of a long raceme on short, jointed stalks, deflexed after anthesis, or (not in Malaysia) in dense globose heads; perfect flowers in each cluster 1-3, ©, at least partly accompanied by imperfect sterile ones (reduced to 82 FLORA MALESIANA [ser. I, vol. 4? fascicled hooks. Tepals of perfect flowers 5, oblong, shortly acuminate, with scari- ous margins, longitudinally nerved. Stamens 5; filaments at the base connate into a short cup; free parts alternating with shorter, dentate or lacerate pseudo-stami- nodes; anthers 2-celled (4-locellate). Ovary obovoid; ovule 1, pendulous from a long funicle; style filiform; stig- ma capitellate. Utricle ellipsoid, thin- walled, indehiscent, by means of the hooks easily adhering to passers-by. Distr. Pantropic, probably two dozen species, centering in Africa, in Malaysia only one widely distributed species and an endemic variety. KEY TO THE SPECIES 1. Leaves rhomboid-obovate or rhomboid-oblong; larger ones less than twice as long as broad. 1. C. prostrata 1. Leaves lanceolate or linear-lanceolate; larger ones more than 2!/2 times as long as broad. 1. C. prostrata var. lancifolia 1. Cyathula prostrata (L.) BLume, Bijdr. (1825) 549: Moa. in DC. Prod. 13, 2 (1849) 326; HAssk. in Pl. Jungh. 2 (1852) 133; Hook. f. Fl. Br. Ind. 4 (1885) 723; BaAILey, Queensl. Fl. 4 (1901) 1230; Koorpb. Exk. Fl. 2 (1912) 197; Merr. Fl. Man. (1912) 194; Interpr. Herb. Amb. (1917) 214; En. Born. (1921) 246; Domin, Beitr. Pflanzengeogr. Austr. 1, 2 (1929) 639; BAcKER, Trop. Natuur 11 (1922) 81, cum ic.; Merr. En. Philip. FI.Pl. 2 (1923) 129; RipL. Fl. Mal. Pen. 3 (1924) 7; HEYNE, Nutt. Pl. (1927) 606; BAcKeEr, Onkr. Suiker. (1930) 224, Atl. t. 234; GaAGNn. in Fl. Gén. I.C. 4 (1936) 1070.—Achyranthes prostrata LINNE, Sp.Pl. ed. 2 (1762) 296; Burm. FI. Ind. (1768) 64.—Pupalia prostrata Mart. Beitr. Amar. (1825) 113; HAssk. Pl. Jav. Rar. (1848) 427.—Cyathula geniculata (non Lour.) Mia. Fl. Ind. Bat. 1, 1 (1858) 1045; Koorpb. Minah. (1898) 565.—Fig. 4. Perennial herb, ascending or erect from a rooting base, 30-50 cm high; stem obtusely quadrangular, thickened above the nodes, often tinged with red, rather densely clothed with fine hairs. Leaves rhomboid-obovate or rhomboid-oblong from a contracted or narrowed, rounded, obtuse or acute base and a mostly triangular, acute or rather obtuse apex, entire, ciliate, bordered with red, otherwise green or, especially in a young state, more or less tinged with red, herbaceous, on both surfaces more or less densely patently hairy, 1'/4-15 by 3/4-6!/2 cm; petiole 1-12 mm. Flowers racemed; racemes terminal and often also in the highest leaf- axils, erect, straight or + sinuous, 19-45 cm (in- cluding 1-12 cm peduncle); rachis rather densely pubescent; bracts ovate, acuminate very acute, reflexed after anthesis; clusters shortly stalked, in the lower part of the inflorescence more or less distant, in the higher part crowded, at first erect, afterwards patent, at last reflexed; lower clusters composed of 2-3 perfect 3 flowers and several im- Fig. 4. Cyathula prostrata (L.) BL. from Java, perfect sterile ones; imperfect flowers towards the x< WEE Dec. 1949] apex of inflorescence gradually fewer; apex of in- florescence bearing solitary perfect flowers without imperfect ones; stalks of clusters joined just above the base; ripe clusters falling off as a whole. Tepals of 3 ovate-oblong, strongly mucronate, 2!'/s-3 mm long, dull pale green, glabrous within, externally clothed with appressed or patent, rather long, white hairs; outer ones 5-, inner ones 3-4 nerved. Free parts of the filaments + 1 mm; anthers minute; pseudo-staminodes rectangular-cuneate with a truncate, shortly dentate apex. Ovary with a broad, flat top; style + 2/3mm. Utricle glabrous 1'/2 mm; seed shining brown. Imperfect flowers 13/s—2!/2 mm long, sessile or almost so; hooks obli- quely patent, above the hairy base glabrous and usually red. Distr. Africa to China and Australia, prob. in- troduced in Central America, in Malaysia: through- out the Archipelago. Ecol. In the settled areas, 1-1650 m, common in shaded localities, along roadsides, teak- forests, forest-borders, secondary forests, often gregarious. Uses. Used for some medicinal purposes, against cough, dysentery, cholera and intestinal worms. AMARANTHACEAE (Backer) 83 Vern. Rumput djarang-djarang, M, ranggitan, J, and several local names. Notes. The type specimen of Cyathula genicu- lata Lour. is Achyranthes aspera L. var. lancifolia (MERR.) comb. nov.—Cyathula lancifolia Merr. in Philip. J.Sc. 11 (1916) Bot. 179; En. Philip. F1.Pl. 2 (1923) 129.—C. lancifolia var. stenophylla MeRR. in Philip. J.Sc. 29 (1926) 478. Leaves lanceolate or linear-lanceolate from a narrowed, acute base, towards the apex tapering or slightly acuminate, acute or rather obtuse, api- culate, entire, ciliate, herbaceous, on both surfaces rather thinly clothed with appressed long thin hairs or, barring the nerves, subglabrous beneath, 1!/2-7 by !/3—2!/2 cm; petiole !/4—!/2 cm. Distr. Malaysia: Philippines (Luzon, Samar, Bohol, Mindanao). Ecol. Damp forests at low and medium alti- tudes. Notes. Except the narrow leaves I can find no differences with the species; there are no transitions. MERRILL’s var. stenophylla is apparently a dwarf, characterized by a much slenderer habit and short racemes 5—10 cm long incl. the peduncle. 7. PUPALIA Juss. in Ann. Mus. 2 (1803) 132, nomen consery.; Mog. in DC. Prod. 13, 2 (1849) 331. Herbs, sometimes woody at the base. Leaves opposite, entire. Flowers spicate or racemed; spikes (racemes) terminal and axillary; lower part of spike (raceme) in axils of persistent bracts with a flower-cluster consisting of 2-3 bibracteolate © flowers accompanied by some rudimentary ones; highest flowers often solitary and ©, without rudimentary flowers. Tepals 5, free, with scarious margins, 3—S-nerved, not indurate at base. Stamens 5, at the base connate in a very short cup; no pseudo- staminodes; anthers oblong, 2-celled (4-locellate). Ovule 1, pendulous from a long funicle; style filiform; stigma capitate. Utricle oblong-obovoid, subcompressed; pericarp thin with a sharply delimitated somewhat thicker apex, which finally falls off letting out the seed; rudimentary flowers consisting of fascicled squarrose hooks; clusters falling off (with the bracteoles) as a whole, by means of the hooks, readily and firmly adhering to passers-by. Distr. Few spp., from Africa to India and Malaysia. Hort. Monsp. (1813); DECNeE in Nouy. Ann. Mus. 3 (1834) 372; SPAN. in Linnaea 15 (1841) 345. 1. Pupalia lappacea (L.) Juss. in Ann. Mus. 2 (1803) 132; Moa. in DC. Prod. 13, 2 (1849) 331; Mia. FI. Ind. Bat. 1, 1 (1858) 1046; Hook. f. FI. Br. Ind. 4 (1885) 724; Fors. Wand. (1885) 515. Koorpb. Exk. Fl. 2 (1912) 198; Merr. in Philip. J. Sc. 11 (1916) Bot. 269; En. Phil. F1.P1.2 (1923) 129.—Achyranthes lappacea_ LINNE. Sp.Pl. (1753) 204; Roxs.Fl. Ind. ed. CAREY (1832) 673.—Achy- ranthes atropurpurea LAMK. Enc. 1 (1785) 346.— Pupalia atropurpurea Moa. I.c. 331; ZOOL. Syst. Verz. (1854) 109; Mia. Fl. Ind. Bat. 1, 1 (1858) 1046; Hook. f. Fl. Br. Ind. 4 (1885) 723; Fors. Wand. (1885) 515; Koorp. /.c. 198.—Desmochaeta atropurpurea DC. and D. flavescens DC. ex Cat. Perennial herb, erect or clambering, often much branched, !/2-2m high; stem obtusely quadran- gular or subterete, thickened above the nodes, finely pubescent. Leaves ovate-oblong from a rounded, obtuse or acute base, contracted into the petiole, acutely acuminate, on both surfaces glabrous or more or less densely clothed with shorter or longer, appressed or obliquely patent, white hairs, 2-12 by 1'/4—-7 cm; petiole 2-25 mm, finely pubescent. Spikes (racemes) terminal and in the highest leaf axils, erect of obliquely erect, 84 FLORA MALESIANA [ser. I, vol. 4? 6-35 cm long (1-10 cm peduncle included), finely and more or less densely patently pilose. Flower- clusteis sessile or very shortly stalked, lower re- mote, higher ones crowded, all clusters at first erect, afterwards patent, consisting of 2-3 6 flowers and several rudimentary ones; highest 0 flowers solitary without accompanying rudimentary flow- ers; bracts and bracteoles ovate-oblong. acute, pilose; bracts 2!/2-5 mm long, soon widely patent or reflexed; bracteoles 2!/2-4 mm. Tepals oblong or ovate-oblong, with a short acicular apical point, 4-5 mm long, dull pale green, albido-pilose out- side, inside glabrous. Staminal cup !/3—!/2 mm high, slightly fleshy; free parts of the filaments + 2mm, often pink; anthers + 1/2 mm. Ovary with a narrow base and a clearly delimitated broadly rounded apex; style 1!/2-13/4 mm. Utricle 2—2!/2 mm long; seed shining dark brown; rudimentary flow- ers at first subsessile and small, afterwards borne on a distinct, villous pedicel and much accrescent; hooks finally 3-4 mm long, pale green or purple, glabrous above the villous base. Distr. Africa to India, in Malaysia: eastern part of Java, Madura, Bali, Kangean Arch., Phil- ippines, Celebes, Saleier, Lombok, Sumba, Sum- bawa & New Guinea. Ecol. Distinctly preferring the periodically dry areas, 3-300 m, in sunny or slightly shaded local- ities, grassy wilds, jungles, hedges, teak-forests, in many regions rather common but usually as scattered specimens. Vern. Djembrengan, J, tjaki, J 8. AERVA (4erua AUCTT.) Forsk. Fl. Aeg.—Arab. (1775) 170, nomen consery. Herbs or undershrubs, erect, straggling or clambering. Leaves alternate or op- posite, quite entire. Flowers in axillary and terminal spikes, small 9 or (¢) (9), soli- tary in the axil of a persistent bract, sustended by 2 bracteoles: the latter either falling off with the perianth or not; rachis of the spike remaining whole after fruit- ing, or breaking up. Tepals 5, free, thin and tender or rather firm, hairy, 1- or more- nerved. Stamens 5; filaments at base connate in a short cup; free parts subulate, alternating with shorter, subulate, pseudo-staminodes; anthers 2-celled (4-locellate). Ovary compressed, glabrous; ovule 1, pendent from apex of long funicle; style very short, stigmas 1—2. Utricle falling off with the perianth, much compressed, bursting irregularly. Seed vertical, reniform, shining black. Distr. About 10 spp. in the Old World, centering in Africa. Notes. Aerva javanica (BuRM. f.) Juss. does not occur in Java, nor for that matter elsewhere in Ma- laysia. It is reduced to the African-Asian Aerva persica (BURM. f.) MERR. KEY TO THE SPECIES 1. Spikes very dense. Tepals very thin, l-nerved, externally clothed with many long white hairs, often densely woolly. Bracteoles not falling off together with the fruiting perianth. 2. Stigmas 2, very distinct, shortly linear, obliquely spreading. Spikes for the greater part 1/2-2!/2 cm long, with a usually rounded apex, mostly 2-4 together in the axils of normal leaves, never forming a loosely branched panicle. Tepals 11!/4—1!/2 mm. Leaves !/2—5 by !/4-3 cm. Erect herb, 10-110 cm high. 1. Ae. lanata 2. Stigma 1, entire or very obscurely 2-lobed with broadly rounded lobes. Spikes 3/4—-5 cm long, usually with a conical apex, partly in the axils of ordinary leaves, partly in the axils of bracts, often forming a + paniculate inflorescence. Tepals 2—2!/2 mm. Leaves 1!/2—-7!/2 cm. Erect or more or less clambering, 3/42 m high eng 2. Ae. sanguinolenta 1. Spikes not very dense. Tepals rather firm, qu 5- nerved, extemially) thinly ‘beset with very short hairs. Bracteoles falling of together with the fruiting perianth. Bract-bearing rachis remaining whole for a long time after the fall of the flowers 3. Ae. curtisii 1. Aerva lanata (L.) Juss. in Ann. Mus. Paris 11 vided from near the base into ascending or erect (1808) 131; BLume, Bijdr. (1825) 547; Hassk. PI. Jav. Rar. (1848) 423; Moa. in DC. Prod. 13, 2 (1849) 303; HAssk. in Pl. Jungh. (1852) 132; Mia. Fl. Ind. Bat. 1, 1 (1858) 1039, Suppl. (1860) 149, 365; Hook. f. Fl. Br. Ind. 4 (1884) 728; PULLE in Nova Guinea 8 (1910) 352; Koorp. Exk. Fl. 2 (1912) 198; Merr. En. Philip. Fl.Pl. 2 (1923) 130; Heyne, Nutt. Pl. (1927) 607; BAcKER, Onkr. Sui- ker. (1930) 225, Atl. t. 235.—Achyranthes lanata . LINNE, Sp.Pl. (1735) 204.—Fig. 5. Perennial erect herb, 10-110 cm long, often di- branches; main branches and upper part of the stem often unbranched for a considerable length, leafy and flowering almost throughout; stems terete, hard, densely clothed with appressed and patent white hairs; internodes usually shorter than 2 cm. Leaves alternate, oval-elliptic-obovate from a cuneate or contracted base, acute or rather ob- tuse, with a very short mucro, on both surfaces (especially so beneath) rather densely appressed white-pubescent, 6-50 by 3-30 mm; highest leaves often very small; petiole 2-15 mm. Spikes mostly Dec. 1949] AMARANTHACEAE (Backer) 85 2-4 together, patent or obliquely erect, cylindric with a usually rounded apex, pure white, !/2—1!/2cm, sometimes up to 2!/2 cm long; highest often col- lected into a dense leafless inflorescence, but never forming a loosely branched panicle. Flowers softly membranous; all 0; bracts and bracteoles ovate-oval, mucronate, white, externally hairy, 3/4— 1mm long. Tepals 1!/3—-1'/2 mm, densely white- woolly outside, oval-oblong, rounded or very ob- tuse; 2 outer ones with minute mucro, entirely white, 3 others with green midrib. Stamens + half as long as perianth; style totalling !/4—'/3 mm, bifid Fig. 5. Aerva lanata (L.) Juss. from Java, x 1/s. nearly halfway down; arms very distinct, obliquely spreading, shortly linear. Utricle + 1mm diam. Seed 2/3—3/4 m. Fruiting spike easily breaking up (in a dried state). Graceful plant! Distr. Africa to Asia, in Malaysia: Sumatra, Banka, Java, Madura, Philippines, Aru Isl., Timor, and New Guinea. Ecol. Especially in periodically dry areas, in Java '/2-100 m, in dry localities, along roadsides, on neglected premises, waste places, locally often frequent. Uses. Leaves steeped in hot water used as a remedy against sudden swellings. Vern. Katumpangan ayer, M. Notes. The sheet conserved under the name Achyranthes lanata in Herb. BURMAN at Geneva is Ae. sanguinolenta (L.) BL. A second specimen of Ae. sanguinolenta in the Herb. BURMAN bears the wrong name of Celosia lanata L. (= Aerva java- nica JUSS.). 2. Aerya sanguinolenta (L.) BL. Bijdr. (1825) 547; DEcNE in Nouv. Ann. Mus. 3 (1834) 371; SPAN. in Linnaea 15 (1841) 345; Moa. in DC. Prod. 13, 2 (1849) 300; Hassk. in Pl. Jungh. (1851) 132; Mig. Fl. Ind. Bat. 1, 1 (1858) 1038; Koorp. Exk. FI. 2 (1912) 198; Merr. Interpr. Herb. Amb. (1917) 214; HeEyNe, Nutt. Pl. (1927) 607; BAcKER, Onkr. Suiker. (1930) 226, atl. t. 236.—Achyranthes san- guinolenta LINNE, Sp.Pl. ed. 2 (1762) 294; Burm. f. Fl. Ind. (1768) 63.—Achyranthes scandens Roxs. Fl. Ind. 2 (1824) 509.—Aerva scandens WALL. Cat. (1829) 6911, nomen; Moa. lI.c. 302; Mia. L.c. 1039; Hook. f. Fl. Br. Ind. 4 (1884) 727; Koorp. I.c. 198; Merr. En. Philip. FI1.Pl. 2 (1923) 130; Gaan. in Fl. Gén. I.C. 4 (1936) 1034.—Aerva timorensis Moa. I.c. 301; Mita. lc. 1039. Perennial herb, often more or less woody at base, 3/4-2 m high, erect or + clambering, branch- ed or not; stem terete, its upper part densely cloth- ed with appressed or patent soft white hairs, gradu- ally glabrescent downward; internodes often longer than 3cm. Leaves opposite or alternate (often ona single specimen), ovate-elliptic, oblong or lanceolate from a cuneate or contracted base, usually acute, mucronate, on both surfaces (espe- cially so beneath) rather densely clothed with ap- pressed white hairs, more or less tinged with purple (type) or green, 15-75 by 645mm; petiole 3-10 mm. Spikes solitary or fascicled, partly in axils of ordinary leaves, partly in those of bracts and then often collected into a lax terminal spike or raceme with a well-developed terminal spike, cylindric, usually with a conical apex, 3/4—-5 cm long, more or less tinged with purple or sordidly white, rarely pure white. Flowers softly membra- nous, (0) (Q); bracts and bracteoles ovate, mucron- ate, externally pilose but not very densely so, rarely glabrous, acute 1—-1'/2 mm long. Tepals ob- long, acute, externally pilose but not very densely so, 2—2!/2 mm long; 2 outer ones minutely mucro- nate. Stamens slightly more than half as long as perianth. Style totalling -+- '1/2 mm; stigma entire or very slightly 2-lobed with rounded lobes. Fruit- ing spike very dense and rather thick, not easily 86 FLORA MALESIANA [ser. I, vol. 42 breaking up. Utricle fully 1 mm diam. Seed 3/4— 1 mm diam., shining brownish black. Distr. India to China, in Malaysia: Java (incl. Madura & Kangean), Celebes (incl. Saleier & Muna), Philippines, Moluccas, (Ambon, Tanim- bar), Lesser Sunda Isl. (Lombok, Sumbawa, Flo- res, Timor). Ecol. Especially in periodically dry areas, in Java 5—200 m, in sunny or moderately shaded dry localities, brushwood, hedges, neglected premises, locally often numerous. The purple-tinged typical form is sometimes cultivated for medicinal purpose. Uses. The red-leaved form used internally against haematuria and irregular or painful men- struation (doctrine of the signature). Vern. Ki sambang, sambang tjalak. Notes. In the Kew-herbarium I found 2 abnor- mal specimens collected in 1884 by J. G. Fr. RIEDEL in Timor Laut (=Tanimbar Archipelago; 131—132° E, 7—-8° S), where also normal plants have been gathered. These abnormal specimens had panic- ulate short spikes of deformed hairy flowers; most of these were asexual and consisted of insufficiently differentiated bracts, bracteoles and tepals. But several flowers were pseudo-bisexual; stamens 5, at the base connate in a short cup; filaments alter- nating with short subulate pseudo-staminodes; anthers 2-celled (4-locellate); ovary much com- pressed, glabrous, empty; stigma capitate, entire. Leaves on both surfaces pubescent; inflorescence pubescent. They had been reported to represent Nothosaerva brachiata by HEMSLEY (Rep. Bot. Chall. Exp. 1, 3 (1884) 183). 3. Aerva curtisii OLIV. in Hook. f. Ic. Pl. 23 (1892), t. 2201; RipL. Fl. Mal. Pen. 3 (1924) 8; Burk. & HEND. in Gard. Bull. 3 (1925) 409. Straggling herb, sometimes slightly woody at base, 30 cm of usually much more; stem rather robust, rather thinly pilose. Leaves opposite, ob- long-lanceolate or lanceolate from a gradually narrowed acute base, very acute, herbaceous, on both sides (especially beneath) thinly clothed with patent, rather long thin hairs, 5-15 by 2-4 cm; petiole 3/4-2 cm. Spikes terminal or in higher forks of stem, racemed, umbellate or subpaniculate on a 1—2!/2 cm common peduncle, erect or patent, 1— 41/2 cm long, much less dense than in the 2 other Malaysian species; rachis rather thinly clothed with patent short hairs; bract-bearing rachis thin, not breaking up after fall of flowers or fruits. Flowers 3-farious or in the higher part of the spike bifarious, 5, bracts persistent after the fall of flow- ers, patent, ovate, acuminate, acute, glabrous, very concave, l-nerved, thinly membranous, + 1!/2 mm long; bracteoles falling off together with the peri- anth, ovate, acute, very thin, nerveless, + 1 mm. Tepals oblong, acute, rather firm, strongly 3—S- nerved, externally thinly clothed with very short hairs, + 3 mm long. Free parts of filaments subul- ate from a broad base. Style + '/2 mm; stigma subcapitate, entire. Utricles obovoid-oblong. Distr. Malaysia: Malay Peninsula (Perak). Ecol. On rocks in forests, 150-1000 m, fr. May. Notes. This species is distinctly allied to some continental SE. Asiatic species, e.g. Aerva cochin- chinensis GAGN. 9. NOTHOSAERVA WIGHT, Icon. 6 (1853) 1. Erect annual herb. Leaves opposite. Flowers in axillary, solitary or usually clus- tered, very dense, short spikes, minute, 3—5-merous, 9; flowers solitary in the axil of a bract, subtended by 2 bracteoles; bracts and bracteoles minute, very thin, hyaline, persistent till after fall of fruiting perianth; rachis of spike not breaking up after fruiting. Tepals free acute, hyaline, 1-nerved, outside villous. Stamens 1—2, minute, free; no pseudo-staminodes; anthers 2-celled (4-locellate). Ovary oblong, com- pressed, glabrous; ovule 1, pendulous from the apex of along funicle; style very short, stigma capitate, entire. Utricle compressed, thin-walled, indehiscent. Seed lenticular. Distr. Monotypic, distributed through tropical Africa, Mascarenes & Comores to tropical Asia, and possibly Malaysia. 1. Nothosaeryva brachiata (L.) WIGHT, Icon. 6 (1853) 1; Hook. f. Fl.Br. Ind. 4 (1885) 726; Hus. WINKEL. in Bot. Jahrb. 49 (1913) 366; MerrR. En. Born. (1921) 246.—J/lecebrum brachiatum LINNE, Mant. (1767) 23.—Aerva brachiata Mart. Beitr. Amar. (1825) 83, no 3; Moa. in DC. Prod. 13, 2 (1849) 304; Mia. Fl. Ind. Bat. 1, 1 (1858) 1040. Stem usually much branched, often so from near the base, 0.10—-0.60, glabrous. Leaves elliptic or ovate-elliptic from an acute base, obtuse or acute, minutely mucronate, thinly herbaceous, green, quite glabrous, 2—5S by 1/2-2 cm; petiole 4-8 mm. Spikes yery numerous, for the greater part in clusters of 3-8, rarely solitary or paired, erect or patent, sessile or shortly stalked, '/2-1'/2 cm long, cylindric with rounded apex, white; bracts and bracteoles glabrous, in sicco white, patent; bracts ovate acute, + 1mm long; bracteoles slightly smaller. Tepals + 11/4 mm long, oblong, acute, nerveless, bluish red (ex WINKLER; in the specimens seen by me in sicco white), patently vil- lous outside. Stamens 1-2; filaments very thin, an- thers minute, style hardly perceptible. Utricle fall- ing off with the enclosing perianth. Seed shining, brownish black, + 3/4 mm diam. Distr. Trop. Africa & Mascarenes to SE. Asia, Dec. 1949] AMARANTHACEAE (Backer) 87 in Malaysia: SE. Borneo (between Kuma and Sa- linahu, fis bluish rosa, HUB. WINKLER 2950, fl. July). Notes. I have not seen WINKLER’s number which is the sole record in Malaysia. Neither have I seen a specimen with hairy stems or leaves. This species is sometimes confused with Aerva lanata (L.) Juss. which may be easily distinguished by the distinct hairiness of the stem and leaves and the manifestly bifid, longer style. The specimens cited by HemsLey (Rep. Bot. Chall. 1, 3 (1884) 183) are abnormal plants and have proved to belong to Aerva sanguinolenta (L.) BL. 10. CENTROSTACHYS WALL. in Roxs. FI. Ind. 2 (1824) 497. Aquatic herb. Leaves opposite, petioled, entire, herbaceous. Flowers 9, spicate; spikes terminal, erect many-flowered, at last elongate; only few flowers open at the same time. Flowers solitary in the axil of a thin bract, subtended by 2 membranous bracteoles, after anthesis deflexed. Tepals 5, spreading during anthesis; before and after anthesis erect, firmly membranous, after anthesis hardened at base; outer- most one rigid, pungent, 1-nerved; others distinctly shorter, plurinerved. Stamens 5, much shorter than perianth, at the base connate in a short cup, alternating with short, cuneate pseudo-staminodes; these dorsally, just below the truncate or sub- dentate top, with a fimbriate scale much surpassing them. Anthers oblong, 2-celled (4-locellate). Ovary glabrous; ovule 1, pendent from a long funicle; stylefili form, short, persistent; stigma capitate. Utricle ovoid, thin-walled, indehiscent; seed erect. Distr. Monotypic, from trop. Africa to SE. Asia, Java and Norfolk Island (+ 168° E, 29° S). 1. Centrostachys aquatica (R.BR.) WALL. Cat. (1829) 6932, nomen; Mog, in DC. Prod. XIII, 2 (1849) 321; Mia. FI. Ind. Bat. 1, 1 (1858) 1043.— Achyranthes aquatica R.Br. Prod. (1810) 417; Roxs. FI. Ind. ed. Carey (1832) 673; Hook. f. FI. Br, Ind. 4 (1885) 730; Koorp. Exk. Fl. 2 (1912) 149; GaGwn. in Fl. Gén. I. C. 4 (1936) 1073. Stem floating or ascending, rooting, terete, thick, striate-ribbed, densely appressed pubescent, me- dullate, 0.75—1.50 m long. Leaves oblong-lance- olate from an acute base, acuminate, acute, 7!/2-15 by 2-3 cm, on both surfaces clothed with appressed long, white hairs; indumentum of young leaves very dense, growing thinner with age; petiole 1-3 cm. Spikes erect, 71/245 cm; rachis rather den- sely appressed pilose; bracts, bracteoles and flow- ers glabrous; flowers at first crowded, afterwards remote; bracts soon reflexed, ovate, acuminate, 3— 4mm; bracteoles embracing together the foot of the perianth, broadly ovate-orbicular, very con- cave, 3—3!/2 mm diam. Outermost tepal with very narrow transparent margins and a firm, slightly re- curved, subulate tip, 6-8 mm long; other ones with broader transparent margin 5-7 mm long. Fila- ments (staminal cup included) 2-3 mm long; an- thers 11/4-11/2 mm. Style 13/s-2!/2 mm. Utricle (not seen) rather acute, + 4 mm long. Distr. Trop. Africa to SE. Asia, in Malaysia: E. & Central Java. Ecol. In Central Java at 450, in E. Java at 20 m, in swampy or inundated localities, locally abun- dant but, on the whole, very rare. 11. ACHYRANTHES LINNE, Sp.Pl. 1 (1753) 20. Erect or ascending terrestrial herbs. Leaves opposite, petioled, entire, herba- ceous. Flowers © spicate. Spikes terminal or axillary, erect, many-flowered, at last elongate; only few flowers open at the same time; flowers solitary in the axil of an acuminate, acute, membranous, persistent bract, subtended by 2 bracteoles, after anthesis deflexed; bracteoles consisting of a rather long spine bearing on either side of its thick concave base a much shorter, membranous nerveless wing. Tepals 5, spreading during anthesis, before and after anthesis erect, membranous or herba- ceous-coriaceous, 1- or more-nerved, very acute, in fruit pungent or not. Stamens 5, much shorter than the perianth; filaments at the base connate in a short cup, alternating with short broad pseudo-staminodes; anthers oblong, 2-celled (4-locel- late). Ovary glabrous; ovule 1, pendent from a long funicle; style filiform, short, persistent; stigma capitate. Utricle falling off together with perianth and bracteoles, 88 FLORA MALESIANA [ser. I, vol. 42 by means of bracteoles or tepals easily adhering to passers-by, ellipsoid with trun- cate or depressed apex, thin-walled, indehiscent. Seed erect. Distr. Few spp., mostly in the Old World, often introduced. Notes. In Malaysia transitions between the 2 species described below have not yet been found. KEY TO THE SPECIES 1. Back of the pseudo-staminodes just below the apex with a long-fringed scale much surpassing the pseudo-staminode itself. Basal wings of the bracteoles (when not damaged) adnate throughout their length to the spine, 1!/s—-2 mm long . 1. A. aspera 1. Back of the pseudostaminodes without a scale. ‘Basal wings of the bracteoles inserted on the thick base of the spine, otherwise free from it, !/2-11/4 mm Jong . Achyranthes aspera LINNE, Sp.PIl. (1753) 204; aca Fl. Filip. (1837) 188, ed. 2 (1845) 133; ed. 3, 1 (1877) 239; Moa. in DC. Prod. 13,2(1849) 314; Mio. FI. Ind. Bat. 1, 1 (1858) 1042; Suppl. (1860) 365; Bru. FI. Austr. 5 (1870) 240; Hook. f. FI.Br. Ind. 4 (1885) 730; Bogert. Ned. Kruid. Arch. II, 5 (1891) 421; Koorp. Minah. (1898) 563; BAILEy, Queens]. Fl. pt 4 (1901) 231; Koorp. Exk. Fl. 2 (1912) 199; Merr. Interpr. Herb. Amb. (1917) 214; En. Born. (1921) 246; En. Philip. Fl.Pl. 2 (1923) 31; Domin, Beitr. Pflanzengeogr. Austr. (1929) 638; RIDL. Fl. Mal. Pen. 3 (1924) 8; Heyne, Nutt. PI. (1927) 607; BACKER, Onkr. Suiker. (1930) 228, Atl. t. 238; OCHSE & BAKH. Vv. D. BR. Veget. (1931) 40, fig. 7; GAGN. in FI. J.C. 4. (1936) 1071.—Achy- ranthes canescens R.Br. Prod. (1810) 417; Moa. l.c. 315; Hassk. Pl. Jav. Rar. (1848) 412.—Achy- ranthes argentea DECNE in Nouv. Ann. Mus. 3 (1834) 372; SPAN. in Linnaea 15 (1841) 345.—Achy- ranthes grandifolia Moa. I.c. 313; Mia. l.c. 1042. Erect rather stiff herb ‘/4-1!/4 m high, usually branched from near the base; branches obliquely erect or ascending; stem angular-ribbed, thickened above the nodes, hard, more or less densely hairy. Leaves oval-obovate or elliptic-oblong, from an acute or obtuse base, acuminate or not, acute, ob- tuse or rounded, entire, flat or more or less wavy, more or less densely hairy or, barring the nerves, glabrous or subglabrous, 11/2-10 cm by 3/4—51/4 cm; petiole !/2-1!/2cm. Spikes terminal, erect, 10-75 (‘/2-15 cm peduncle included); rachis rather robust, stiff, angular-ribbed, more or less densely clothed with appressed or more or less patent, rather long white hairs; bracts long-acuminate, not pungent, 2—31/2 mm Jong, before anthesis erect, afterwards spreading, at last quite reflexed as often is the fruit- ing perianth also, + silvery; bracteoles appressed against base of perianth. Spines 23/4—4!/2 mm, shining, often tinged with purple, sharp; basal wings almost throughout their length adnate to spine but most easily separating from it, 1!/4-2 mm long. Tepals ovate-lanceolate, very acute, green with pale margins, with 3 or more rather strong nerves, during anthesis 31!/2-5!/2 mm long, after- wards up to 4!/2-6!/2 mm, hardening and becoming pungent. Filaments (staminal cup included) 2!/4— 31/2 mm; pseudo-staminodes truncate or crenulate, just below the apex with a dorsal long-fringed scale far exceeding the top of the pseudo-staminode itself; ovary turbinate; style 1-2 mm. Urricle roun- ded at the base, 21/2-23/4 mm long. Distr. Ubiquist, in Malaysia: throughout the 2. A. bidentata Archipelago, possibly not truly indigenous in Ma- laysia. Ecol. Sunny dry localities especially in regions with a well-marked dry monsoon: road-sides, waste places, a typical ruderal, 1—-2300 m. Uses. Rubbed on the body of young children against convulsions. Vern. Djarong, njarong, etc. Notes. In Journ. Ind. Bot. Soc. 11 (1932) 335 A. C. Josui describes and figures a case of dedoub- lement of stamens in this species. 2. Achyranthes bidentata Bri. Bijdr. (1825) 545; Moa. in DC. Prod. 13, 2 (1849) 312; Mra. FI. Ind. Bat. 1, 1 (1858) 1040; BoerL. in Ned. Kruid. Arch. II, 5 (1891) 427; Koorp. Minah. (1898) 564; ScHum. & Laut. Fl. Deut. Sch. Geb. (1901) 306; Koorpb. Exk. Fl. 2 (1912) 199; PULLE in Nova Guinea 8 (1912) 627; Merr. En. Philip. FI.Pl. 2 (1923) 545; HEYNE, Nutt. Pl. (1927) 607; GAGN. in Fl. Gén. I.C. 4 (1936) 1072.—Achyranthes javanica Moa. /.c. 312; Mia. lc: 1041. Erect or ascending, rather flaccid herb, 3/4—1!/2 m long; stem -++ quadrangular with longitudinally furrowed sides, often purple in higher part, thinly or moderately densely clothed with appressed or patent long, fine, white hairs. Leaves elliptic-oblong- lanceolate or ovate-lanceolate from an acute or obtuse base, long acuminate, on both surfaces thinly or moderately densely clothed with appress- ed or patent longish hairs, 5-20 by 1-8 cm; petiole 1/2-31/2 cm. Spikes terminal and in the higher leaf axils, 4-45 cm long (including 1-15 cm peduncle); rachis rather thin and flaccid, often somewhat tor- tuous, rather densely clothed with appressed of more or less patent long white hairs; bracts long- acuminate, not pungent, 3—3!/2 mm long, before anthesis erect, afterwards patent or reflexed; brac- teoles appressed against base of perianth, often slightly recurved at the apex, very variable in length, 2!/2-51/2 mm, basal wings inserted on the thick base of a spine, otherwise free, erect or more or less patent, !/2-1!/4 mm long. Tepals (midrib ex- cepted) thinly membranous and nerveless, very acute, during anthesis often violet, often distinctly unequal, variable as to length, 4'/2-7 mm, not be- coming pungent. Filaments (staminal cup included) 2-2!/2 mm long; pseudo-staminodes truncate, en- tire or mostly irregularly dentate, without dorsal scale. Style '/2-2mm. Urricle subtruncate, 2- 21/2 mm long. Distr. Tropical Africa and Asia, in Malaysia: Dec. 1949] Sumatra, Java, Celebes, Philippines, Moluccas, Lombok and New Guinea. Ecol. Throughout Java, 350-2500 m, in forests and well-shaded localities, often abundant along trails under everwet conditions. Uses. Internally used as an anthelmintic; masti- AMARANTHACEAE (Backer) 89 cated against malignant ulcers of the cavity of the mouth. Notes. It is remarkable that this species has not yet been collected in the Mal. Peninsula, Borneo and most of the Lesser Sunda Islands. Unlike A. aspera it is certainly indigenous in Malaysia. Excluded Achyranthes linearifolia Sw. in WIKSTROM, Vet. Akad. Handl. Stockholm fOr 1825 (1826) 428 (not: 48) is quoted by Ind. Kew. as described from the Moluccas. It came from St Barthélemy, E of Porto Rico, in the West Indies, as I was kindly informed by Dr FLorIn. Achyranthes spiciflora Burm. f. Ind. Alt. Herb. the Globe 4 (1800) 257 is based on Cauda felis agrestis RUMPH. Herb. Amb. 4, p. 84. According to MERRILL (J. Arn. Arb. 29 (1948) 188) this is Acalypha amentacea Roxs. (Euph.). Achyranthes hispida PENNANT, lc. is according to MERRILL, /.c., Acalypha hispida Bur. f. Fl. Ind. (1768) 303, pl. 61, f. 1 (Euph.). Amb. (1769) 5, non ibid. 203; PENNANT, Outl. of 12, PTILOTUS R.Br. Prod. (1810) 415. Annual herbs. Leaves alternate, entire, often narrow. Flowers 9, terminal, in glob- ular conical or cylindrical dense heads or short spikes, solitary in the axil of a bract, subtended by 2 bracteoles; bracts and bracteoles scarious; tepals free or at the base shortly connate, usually dimorphous, after anthesis indurated at the base or not, glabrous or clothed on the back with denticulate, often long hairs, inside glabrous or woolly. Stamens 5; filaments free or connate in short cup, with or without interposed, small, translucent pseudo-staminodes, all perfect or 1—2 sterile. Anthers 2-celled (4-locellate). Ovary sessile or shortly stalked, glabrous or hairy; ovule 1, pendent from basal erect long funicle; style central or slightly excentrical, long, thin; stigma capitate. Utricle enclosed by perianth, indehiscent. Seed vertical, sometimes arillate. Distr. If united with Trichinium, which is now almost universally done, this large genus is practically confined to Australia and Tasmania, in Malaysia: one Australian species. Ecol. Mainly confined to semi-arid regions. 1. Ptilotus conicus R.Br. Prod. (1810) 415; Moa. in DC. Prod. 13, 2 (1849) 282; Bru. Fl. Austr. 5 (1870) 242; BAILEY, Queensl. FI. pt 4 (1901) 1222.— Ptilotus corymbosus (non R.BR.), BLUME, Bijdr. (1825) 543; Mia. Fl. Ind. Bat. 1, 1 (1858) 1036.— Ptilotus amabilis SPAN. in Linnaea 15 (1841) 345, ic. ined. 59 in Herb. Lugd. Bat.—Ptilotus conicus var. timorensis ENGL. Bot. Jahrb. 7 (1885) 454. Stem erect, 40-60 cm high, slender, glabrous, often already quite near the base divided into as- cending main-branches, in higher part with erecto- patent branchlets. Leaves rather distant, narrowly linear, acute, with (in sicco) recurved margins, glabrous, 2-5 cm by !/2-2!/2 mm. Heads in higher portion of plant loosely paniculate-corymbose, at first globular, -+ 3/4 cm diam., growing more. cylindrical with age, finally up to 11/2 cm long; bracts and bracteoles with short acicular tips, l-nerved, 2!/2-3!/2 mm long, persistent after the fall of perianth; flower-axis articulated above the bracteoles. Tepals shortly united at the base, purple, + 4!/2 mm long; 2 outer ones elliptic, outside at base with dense tuft of erecto-paten long hairs, otherwise glabrous, 3—3!/2 mm long; 3 inner ones with an erect, narrowly cuneate, 3-nerved claw and a slightly longer and (at the base) broader, ovate-oblong, l-nerved, patent blade; claw along either margin densely clothed with intricate, long, thin, dentate hairs. Perfect stamens 5, glabrous, + 41/2 mm, filaments narrowly ligular, at base con- nate into a short cup; free parts tapering upwards but at 3/41 mm below top abruptly broadened into a subcircular disk, above this dilatation much con- tracted, filiform; anthers short; no pseudo-stami- nodes. Style glabrous, + 21/2 mm. Utricle narrowly ovate compressed, + 2!/2 mm. Distr. N. Australia (Gulf of Carpentaria), in Malaysia: Lesser Sunda Islands (Flores, Roti, Ti- mor, Wetar), and S. Moluccas (Tanimbar, Key); a distinct Australian element in the Malaysian flora. Ecol. Apparently confined to open country in periodically dry regions, at low alt., in Timor on calcareous hills. 13. PSILOTRICHUM BLumE, Bijdr. (1825) 544. Dwarf shrubs or herbs. Leaves opposite, entire, Flowers 9, interminal or axillary 90 FLORA MALESIANA [ser. I, vol. 4? heads or narrow spikes, solitary in the axil of a bract, subtended by 2 bracteoles. Tepals 5, free, narrow, strongly longitudinally plurinerved, membranous, after an- thesis indurate or not. Stamens 5, filaments + unequal, at the base connate in a short cup; anthers small, 2-celled (4-locellate); no pseudo-staminodes. Ovary ellipsoid or globose; ovule 1, pendulous from a long erect funicle; style thin; stigma capitate. Fruit enclosed by the perianth, thin-walled, indehiscent; seed erect, len- ticular. Distr. About 14 spp. in Africa, SE. Asia and Malaysia, centering in Africa. 1. Psilotrichum ferrugineum (RoxsB.) Moa. in DC. Prod. 13, 2 (1849) 279; Hook. f. Fl. Br. Ind. 4 (1885) 725.—Achyranthes ferruginea Rox. FI. Ind. ed. WALL. 2 (1824) 502.—Psilotrichum trichoto- mum BLUME, Bijdr. (1825) 545; Mor. Syst. Verz. (1845/6) 73; HAssk. Pl. Jav. Rar. (1848) 419; Moa. I.c. 280; HAssk. in PI. Jungh. (1852) 130 (sphalm. tricholonum); ZOLL. Syst. Verz. (1854) 109; Hoox. f. Fl. Br. Ind. 4 (1885) 725; Koorp. Exk. Fl. 2 (1912) 199; Merr. En. Born. (1921) 246; En. Philip. 2 (1923) 131; Riv. Fl. Mal. Pen. 3 (1924) 8; BAcKER, Onkr. Suiker. (1930) 227, Atl. t. 237; GAGN. in FI. Gén. I.C. 4 (1936) 1066.—Leiosper- mum ferrugineum WIGHT, Ic. (1843) 721.—Ptilotus trichotomus Miq. Fl. Ind. Bat. 1, 1 (1858) 1037. —Fig. 6. Annual, erect or ascending, often much branch- ed from near the base, 5—50 cm long, in sunny lo- calities often strongly tinged with purple; stems thin, in the leaf axils and on the young nodes often + hairy, otherwise glabrous; pairs of leaves often distant. Leaves patent, variable in shape, lance- olate, oblong, elliptic or obovate, minutely mucron- ate, rather fleshy in a living state, glabrous, 3/4— 71/2 by 1/2-2 cm; petiole 3-10 mm. Spikes usually terminal or in forkings of stem, sometimes also axillary, solitary or rarely paired, sessile or on thin and often rather long peduncles, at first shortly conical, afterwards lengthened, rather acute, dense, 1—21/2 cm long; rachis pilose; lowest fruits often falling off before expansion of the highest flowers; bracts and bracteoles thinly membranous; bracts widely patent or subreflexed, ovate-lanceolate, concave, very acute, 1!/4—-1!/2 mm long, persistent; bracteoles ovate-triangular, nerveless, 1!/2-3/4 mm long, falling off together with the perianth and the fruit. Perianth much longer than the bract and the BN bracteoles. Tepals lanceolate, very acute, through- A out their width with 3—5 strong longitudinal ner- ves, glabrous, 2—2!/2 mm long, not or hardly indu- rate after anthesis. Filaments very thin, !/2-3/4 mm long (cup included). Style !/2/s mm, persistent. \ Utricle ellipsoid, compressed-pellucid but opaque \ at the apex, 1—11/2 mm long. Seed vertical, shining / \ black or blackish brown, + 1 mm long or slightly longer. \ Distr. SE. Asia, in Malaysia: throughout the Archipelago, not yet reported from the Lesser a ee { Sunda Islands and the Moluccas. \ ? Ecol. Humid clayey fields, along ditches and trenches, locally often numerous, 5—1200 m, on the whole not a very common species. Fig. 6. Psilotrichum ferrugineum (RoxsB.) Moa. Notes. The shape of the leaves is very variable. from Java, X 1/3. Dec. 1949] AMARANTHACEAE (Backer) 9] 14. ALTERNANTHERA Forsk. Fl. Aeg.—Arab. (1775) 28. Annual or perennial, erect, ascending, trailing, creeping, floating or clambering herbs, often hairy; hairs dentate or smooth. Leaves opposite, entire. Flowers © or by malformation Q, in axillary or rarely terminal, sessile or peduncled heads or short spikes, solitary in axil of bract, subtended by 2 bracteoles; bracts and brac- teoles scarious. Perianth often dorsally compressed. Tepals 5, free, equal or un- equal, glabrous or hairy. Stamens normally 2-5, sometimes partly anantherous; fila- ments at the base united in a tube or a short cup; free part short, usually alter- nating with (sometimes very minute) pseudo-staminodes; anthers small, 1-celled (2-locellate); pseudo-staminodes entire, dentate or laciniate. Ovary compressed or not; ovule 1, pendulous from a long funicle; style short; stigma capitate. Utricle indehiscent, sometimes corky, falling off with the perianth and with or without the bracteoles. Seed vertical. Distr. Large genus, centering in America, some species in other parts of the World, in Malaysia: 6 species, of which 1 indigenous (one variety endemic); the 5 others introduced from trop. America. Three of these are naturalized and, often on a large scale, locally firmly established. The other two are cultivated. Uses. The gregarious matted growth of some species is sometimes used for protecting soils from rain- wash. Others are cultivated for ornamental purposes. Notes. In some species the hairs are, under the microscope, smooth, in others they are minutely but distinctly dentate. This neglected character is of importance for specific delimitation; it proved constant in the Malysian species treated here. KEY TO THE SPECIES 1. Heads all sessile (after the fall of the lower flowers often seemingly stalked, but in this case the spurious peduncle is at once recognized as the rachis of the head by the presence of bracts). Perianth sessile between the bracteoles. Filaments at the base united in a very short cup. 2. Bracts and tepals not spinescent. 3. All tepals 1-nerved, or only at the very base obscurely 3-nerved, not with indurate bases. Bracts not or shortly acuminate. Anthers 3, oval or oblong. Pseudo-staminodes minute, entire, perhaps some- times wanting. Leaves green. Hairs smooth. 4. Leaves variable as to shape but not narrowly linear-filiform, 3/s-15 by '/s-3 cm . 2. A. sessilis 4. Leaves very narrowly linear-filiform, 2-6 cm by !/2~-2mm. . ._ 2. A. sessilis var. tenuissima 3. Three outer tepals in their lower !/3—!/2 distinctly 3-nerved, their bases at last indurate. Bracts rather long-acuminate. Anthers 5, linear, 1-2 of them sometimes shorter than the others. Leaves often coloured. Hairs dentate . . . 3%. A. ficoides 2. Bracts and 2 outer tepals with aneeeae tips. Entire head conspicuously prickly. Stems trailing or creeping. Hairs dentate . . re 1. A. repens 1. Heads usually borne on a distinct bractless peduncle (if sessile then ‘the perianth is 5—7 mm long!). Filaments united at the base into a distinct tube. Pseudo-staminodes distinct, divided into narrow apical straps. 5. Stems solid, their young parts hairy all round; hairs minutely dentate. Bracts, bracteoles and peri- anth hairy. Leaves, when not too old, densely appressed-hairy all over the lower surface. Not aquatic. 6. Perianth yellowish-white, placed between the bracteoles on a short but distinct stalk, falling off together with this stalk. Top of the bracteoles with a distinct dorsal hairy crest. Bracts with a rather long mucro. Robust plant . . zine 5. A. brasiliana 6. Perianth red, sessile between the Briciealcs Bracteoles nat crested. ‘Bracts with a very short mucro or without one. . . . 6. A. porrigens 5. Stems in their lower part ‘fistulose. Leaf axils with white smooth hairs, on two opposite sides with a longitudinal hairy groove, otherwise glabrous. Plant of very humid localities, often growing in Riatlow. water. cu. Sc nce Pele te les) he ae A ee ee ee ecereites 1. Alternanthera repens (L.) StEuD. Nomencl. ed. Prod. 13, 2 (1849) 358; Mig. Fl. Ind. Bat. 1, 1 2, 1 (1840) 65; O.K. Rev. Gen. 2 (1891) 540; (1858) 1049; Backer, Bull. J.B.B. II, 12 (1913) 7. Ascu. & Gr. Syn. 5, 1 (1914) 362; BACKER, Onkr. Prostrate herb, often rooting; taproot robust; Suiker. (1930) 229, atl. t. 239.—Achyranthes repens _ stem terete, hard, on the younger parts densely Linné, Sp.Pl. (1753) 205.—Illecebrum achyrantha clothed with appressed, minutely dentate, white LINNE, Sp.Pl. ed. 2 (1762) 299.—Alternanthera hairs, 10-50 cm long. Leaves elliptic-obovate, nar- achyrantha R.Br. Prod. (1810) 417;-Mog. in DC. rowed into the petiole, with an obtuse or roundéd 92 FLORA MALESIANA [ser. I, vol. 4? top, on the upper surface glabrous or thinly ap- pressed-pilose, on the lower surface glabrous or on the nerves appressed-pilose, 1'/4—-4!/2 by !/3— 2 cm, in a single pair often of very unequal size; petiole 2-10 mm. Heads 1-3 in the leaf-axils, ses- sile, globose or oblong, white, !/2-1'/2 cm long; bracts spine-tipped, -+- 4mm (including spine); bracteoles acuminate, very acute but not spinous, 3-4 mm long. Tepals very unequal: 2 abaxial ones much larger than the others, convex, spine-tipped, + 5 mm long, above base right and left with hair- tuft, in the lower half with 3 upwards confluent nerves, hardening after anthesis; adaxial tepal rather flat, oblong, dentate near tip, mucronate, not spiny, at base right and left with hair-tuft, + 31/2 mm long, two inner tepals much smaller than the others, their lower halves very concave, em- bracing the ovary and afterwards the fruit, on the middle of their back with a patent hair-tuft. Sta- mens 5, all perfect; filaments (basal cup included) 1/>-3/4 mm; anthers oblong, minute; pseudo-stami- nodes much shorter than filaments, broad, entire, emarginate or irregularly dentate. Style very short. Fruit falling off with perianth and_ bracteoles, broadly oval, much compressed, truncate or retuse, brown + 1!/2 mm long. Distr. Native of trop. America, introduced in Java, collected there for the first time in 1912, now established in some waste places on and near the northern coast, at low altitude, locally abundant, but on the whole still rare. Notes. The specimen mentioned by BURMAN f. in his FI. Ind. (1768) 66 as I/lecebrum achyranthes, and conserved at Geneva in Herb. DELESSERT, is Alternanthera sessilis. 2. Alternanthera sessilis (L.) R.Br. ex R. & S. Syst. 5 (1819) 554; Moa. in DC. Prod. 13, 2 (1849) 357; Mio. Fl. Ind. Bat. 1, 1 (1858) 1048; Hook. f. FI. Br. Ind. 4 (1885) 731; O.K. Rev. Gen. 2 (1891) 540; Koorp. Exk. Fl. 2 (1912) 200; AscH. & Gr. Syn. 5, 1 (1914) 361; Merr. Interpr. Herb. Amb. (1917) 215; En. Born. (1921) 246; En. Philip. 2 (1923) 132; Rip. Fl. Mal. Pen. 3 (1924) 10; BAcK. & SLoor. Handb. Thee. (1924) 109, t. 109; HEYNE, Nutt. Pl. (1927) 608; Backer, Onkr. Suiker. (1930) 230, Atl. t. 241; GaAGN. in Fl. Gén. I.C. 4 (1936) 1077.—Gomphrena sessilis LINNE, Sp.Pl. (1753) 225.—Illecebrum sessile BURM. f. Fl. Ind. (1768) 166.—Illecebrum indicum Houtt. Nat. Hist. 2, 7 (1777) 713, t. 43, f. 3.—Alternanthera triandra LamMkK. Encycl. 1 (1783) 95; Burk. Dict. 1 (1935) 116.—Alternanthera_ denticulata R.Br. Prod. (1810) 417; Brume, Bijdr. (1825) 546; Mia. L.c. 1048; Bru. Fi. Austr. 5 (1870) 249; Hook. ff. lc. 731; BAILEY, Queensl. Fl. pt 4 (1901) 1233, Domin, Beitr. Pflanzengeogr. Austr. 1, 2 (1929) 639.— A. nodiflora R.Br. Prod. (1810) 417; Mia. Le. 1047; Bru. Jc. 249; Battey, /.c. 1233; KoOorD. Exk. Fl. 2 (1912) 200; Domin, /.c. 639.—Achyran- thes villosa (non Forsk.) BLANCO, FI. Filip. (1837) 189; ed. 2 (1845) 134; ed. 3, 1 (1877) 240.—Adeno- gramma oppositifolia HAssk. in Flora 31 (1851) 754._Fig. 7. Herb, perennial or under unfavourable circum- stances short-lived, often pluricaulous, 10-100 cm long; taproot robust; stem either erect, ascending or creeping and solid, (or, in inundated localities, floating and in the lower part fistular), green or more or less tinged with purple, on 2 opposite sides with a longitudinal row of hairs, across the nodes Fig. 7. Alternanthera sessilis (L.) R.BR. from Java, x !/s. with a transverse row of hairs, otherwise glabrous; hairs articulate, smooth. Leaves variable as to shape and size, varying from linear-lanceolate via oblong to oval or obovate, acute at the base, acute, obtuse or rounded at the apex, green, glabrous or thinly and finely pilose, 3/4-15 cm by ‘/4—3 cm, in wet localities comparatively large, in dry localities much smaller; petiole 1-5 mm. Heads 1—4 in the axil of present or fallen leaves, sessile but by the fall of lower flowers often spuriously peduncled, at first globose, growing more cylindrical with age, 1/2~-11/2 em long; rachis densely white-hairy; bracts and bracteoles not or shortly acuminate, glabrous, white, persistent after the fall of the flowers; bracts 2/3-1 mm; bracteoles 1—1!/2 mm. Tepals shortly acuminate or not, glabrous or on the back with few or several hairs, white or purplish, shining (in a living state), l-nerved or only at the very base obscurely 3-nerved, after anthesis not hardening in the lower part, 2'/-3 mm long. Filaments 5 of which only 3 antheriferous, ++ 3/4 mm long (basal cup included); anthers oval-oblong, '/6—'/4 mm; pseudo-staminodes subulate-filiform, entire, very minute, not exceeding filaments, often shorter. Style during anthesis very short, afterwards slightly + Dec. 1949] lengthened. Fruit falling off with the perianth, obreniform, deeply emarginate, corky, dark brown, glabrous, 2—2'/4 by 2!/2-3 mm; lobes finally longer than the interposed style. Distr. Throughout the Old World, in Malaysia: throughout the Archipelago. Ecol. Acommon plant, !/2-1250 m, in constant or periodically humid or even inundated, open localities: fallow rice-fields, road-sides, gardens, shallow ditches, swamps, tea-plantations. Fruits often floating in great quantities upon the water. Uses. An infusion of the entire plant is used as a remedy against intestinal cramps and as a cooling hair-wash. Vern. Daun tolod, M, kremak, J. Notes. The habit of the species varies exceed- ingly with the habitat. var. tenuissima (SUESS.) comb. nov.—Alternan- thera tenuissima SUESSENGUTH in Bot. Arch. 39 (1939) 382. Leaves from a narrowed base very narrowly linear or almost filiform, acute, shortly mucronate, with a strongish midrib, when very young sparsely beset with longish white hairs, soon becoming glabrous, 2-6 cm by !/2-2 mm. Pseudo-staminodes not found. Distr. Malaysia: NE. New Guinea (Morobe distr., CLEMENS 6339). Ecol. The only collection known was made in a garden, as a weed, at c. 1800 m alt. Notes. In contrast to SUESSENGUTH who says that it is not closely allied to any Malaysian spe- cies, I find this aberrant form in all essential char- acters agreeing with A. sessilis from which it differs only by the peculiar shape and width of the leaves. It might be an etiolated or depauperated form. A specimen closely resembling this variety was figured by DomIn (/.c.) under the name of A. nodi- flora R.Br. from Queensland. 3. Alternanthera ficoides (L.) R.Br. ex R. & S. Syst. 5 (1819) 555; Merr. En. Philip. FI.Pl. 2 (1923) 131.—Gomphrena ficoidea LINNE, Sp. PI. (1753) 235.—Gomphrena polygonoides LINNE, Sp. Pl. (1753) 225, ex parte——Alternanthera polygo- noides R.Br. Prod. (1810) 416; AscH. & Gr. 5, 1 (1914) 364.—Teleianthera manillensis WAP. in Nov. Act. Ac. Nat. Cur. 19 (1843) Suppl. 1, 404.— Telanthera polygonoides Moa. in DC. Prod. 13, 2 (1849) 363. var. bettzickiana (NICH.) comb. nov.—Alternan- thera bettzickiana Nicw. Gard. Dict. ed. 1 (1884) 59; AscH. & Gr. Syn. 5, 1 (1914) 365.—Alternan- thera amoena (non Voss.?) BACK. & SLooT. Handb. © _ Thee. (1924) 108, t. 108; HEYNE, Nutt. Pl. (1927) 608. Perennial herb, 20-50 cm high; stem erect or at the base for a greater or smaller part decumbent and rooting, often much branched and forming dense tufts, terete in the lower part, quadrangular upwards, on 2 opposite sides with a longitudinal furrow, appressed-pilose at the top and on the nodes. Leaves oblong, oblong-obovate or spathul- ate from an acute base, acute or obtuse, finely mucronate, often + crisp, not rarely entirely AMARANTHACEAE (Backer) 93 green, but mostly in different ways variegated with brownish red, bright red, pink or yellow, when young clothed with fine dentate hairs, glabrescent, 1-6 by '/2-2 cm; petioles 1-4 cm, + hairy. Heads terminal and axillary, often in dense clusters of 2-5, globular or oblong, '/2-1 cm long; hairs of bracts, bracteoles and floral parts, when present, minutely dentate; rachis hairy; bracts and bract- eoles rather long-acuminate, very acute, glabrous or on the back with long hairs; bracts 13/s-3 mm; bracteoles 2—23/4 mm. Perianth often + difformed and then consisting of more than 5 tepals; normal tepals white or yellowish, shiny; 3 outer ones in their lower '/3—!/2 with 3 close-set strong, upwards convergent nerves; their bases finally indurate; 2 abaxial tepals ovate-oblong, 3-4 mm long, con- cave, in their lower halves rather densely patently pilose on the back; their upper halves acutely acuminate, glabrous; adaxial tepal ovate-oblong, faintly concave or almost flat, sparingly hairy or glabrous, 23/4-3!/2 mm long; 2 inmost tepals very concave, narrower and shorter than the others, sparingly hairy of glabrous. Stamens (basal cup included) 11/4—2!/4 mm long; anthers 5, linear; 1-2 often somewhat shorter than the others and sterile; fertile ones 3/s-1 mm; pseudo-staminodes strap- shaped, reaching up to the middle or the top of the anthers, at the apex cleft into 3-5 very narrow Strips. Ovary glabrous; style subconical rather thick + 1/2 mm. Urricle in Java not developing (in typical A. ficoides faintly notched). Distr. Native of Brazil, already long ago intro- duced into Java, in Malaysia (at least in Sumatra and Java) frequently cultivated as an ornamental plant, or in tea-plantations along the borders of terraces as a protector from rain-wash, for which purposeitis, by its densely tufted growth, eminently adapted, 1—2000 m. Ecol. In Malaysia fruits are never produced. Uses. See above. Vern. Djoekoet sélon (Ceylon-grass), ketjitjag abang, J. Notes. A form of A. ficoides with entire, subu- late pseudo-staminodes and shorter petioles has been collected in Banka, probably also in a culti- vated state: var. versicolor (REGEL) BACK. 4. Alternanthera philoxeroides (MART.) GRISEB. in Goetr. Abh. 24 (1879) 36; O.K. Rev. Gen. 2 (1891) 540; Koorp. Exk. FI. 2 (1912) 200; BACKER, Onkr. Suiker. (1930) 230, Atl. t. 240; OcHsE & BAKH. V. D. Br. Veget. (1931) 15, f. 10.—Bucholzia philoxeroides Marv. Beitr. Amar. (1825) 107.— Telanthera philoxeroides Mog. in DC. Prod. 13, 2 (1849) 362; BACKER in Ann. J.B.B. II, Suppl. 3 (1909) 417. Perennial herb, ascending from a creeping or floating, rooting base, often much branched and forming dense masses, !/2-1 m long; stem fistular, in the leaf-axils with a transverse row of white, smooth hairs, on 2 opposite sides with a longitu- dinal hairy groove. Leaves oblong or oblong-oboy- ate from a tapering base, acute or rather obtuse, submucronate, glabrous or ciliate, 2!/s-8 by 3/4— 21/2 cm; petiole 3-6 mm. Heads terminal and ses- 94 FLORA MALESIANA [ser. I, vol. 42 sile, or usually axillary and then mostly peduncled, solitary, ovoid-globular-ellipsoid, 3/4—1'/2 cm long; peduncle usually 1—4!/2 cm, not rarely shorter, sel- dom almost wanting, on the adaxial side with a longitudinal hairy groove, otherwise glabrous; bracts and bracteoles 1-nerved, glabrous, white, persistent after fall of perianth; bracts ovate-trian- gular, 2—23/4mm; bracteoles ovate, acuminate, very acute, 2!/4—-21!/2 mm. Perianth almost sessile above the bracteoles, dorsally compressed, shining white, glabrous, 5-7 mm long. Tepals oblong, acute or rather obtuse, 1-nerved. Stamens in nor- mal flowers (see beneath) 5; filaments 3!/2-4 mm (including short staminal cup); anthers linear, 1— 1'/4 mm; pseudo-staminodes about as long as sta- mens, oblong-linear; their tips divided into a few narrow strips. Ovary shortly stalked, broadly obov- ate-cuneate, dorsally compressed, rounded at the apex; style short, thick. Fruit in Malaysia never produced. Distr. Native of Brazil, introduced long ago in Java, in Malaysia found for the first time in 1875 near Batavia by O. KUNTZE, at present quite na- turalized in many localities in W. Java, not yet re- corded from elsewhere in Malaysia. Ecol. In stagnant or slow-moving shallow wa- ter, pools, ditches, often gregarious. In Malaysia fruits are unknown. Notes. In Java the stamens are very often re- placed by as many sterile spurious ovaries sur- rounding the true ovary. This malformation was pictured and described by CosTerRus in Ann. Jard. Bot. Btzg 23 (1910) 12, t. III. 5. Alternanthera brasiliana (L.) O.K. Rev. Gen. 2 (1891) 537.—Gomphrena brasiliana LINNE, Amoen. 4 (1759) 310.—Alternanthera strigosa HAssk. in Tijdschr. Nat. Gesch. & Phys. 5 (1838/9) 259; in Flora (1842) II /it. p. 19; Plant. Jav. Rar. (1848) 420.—Telanthera strigosa Moa. in DC. Prod. 13, 2 (1849) 370; Koorp. Exk. FI. 2 (1912) 201.—Mogi- Phanes jacquinii (non SCHRAD.) BOERL. Hand. Ge- nees- & Nat. Congr. Leiden (1889) 148.—Telan- thera praelonga (an Mog. ?) BACKER in Ann. J.B.B. Suppl. 3 (1909) 416. Perennial herb, decumbent at the base, higher up ascending-erect or clambering among and over other plants, often widely branched, 1!/2-3 m high; all hairs minutely dentate; stem obtusangular or subterete, thickened and articulate above the nodes, on the younger parts densely clothed with appressed acroscopic long white hairs, gradually glabrescent. Pairs of /Jeaves rather distant, blade ovate-lanceolate from a cuneate or contracted base, acuminate, acute, at first densely appressed pilose on both surfaces, slowly glabrescent, 31/2-10 by 3/44 cm; petiole 1/2-1!/4 cm. Heads terminal, often in bifurcations of the stem, solitary or rarely 2-3 together, stalked, at first hemispheric-ovoid and 6-8 mm diam., afterwards increasing in size, ovoid-oblong, 12-16 by 10-12 mm; stalks at first short, lengthening with age, finally 6-16 cm, more or less densely clothed with appressed acroscopic long white hairs; rachis densely hairy; bracts and bracteoles long-ovate, acuminate, acute, yellowish white, 1-nerved; bracts 2!/2—-31/4 mm, at first den- sely clothed with appressed long, white, for the greater part deciduous hairs; bracteoles consider- ably shorter than perianth, 3—3!/2 mm, on back long-hairy; persistent. Perianth distinctly stalked between bracteoles, falling off with the stalk; stalk 1/23/4 mm long, at base with a whorl of patent longish hairs, thick, with 5 longitudinal ribs; ribs strong, at the base thickened and shining (not glan- dular). Tepals oblong-lanceolate, acute, strongly 3-nerved, yellowish white, at first on back appress- ed long-hairy, gradually losing most hairs, 4-5 mm long. Filaments (staminal cup included) 2—2!/2 mm; anthers linear, 1'/3-11/2 mm; pseudo-staminodes narrow, slightly longer than stamens, shortly den- tate at apex. Ovary obovoid; style 1/3—!/2 mm. Utricle ellipsoid, its top on both sides of the style with an obtuse knob, + 2!/4 mm long. Distr. Native of trop. America, introduced into Java more than a century ago, in Malaysia: at present naturalized in a wide circle around Buiten- zorg, and also collected in Central Java, as yet not found in other islands. Ecol. Moist, shaded localities, 200-600 m, steep ravine slopes, stream banks, locally often gregarious. 6. Alternanthera porrigens (JAca.) O.K. Rev. Gen. 2 (1891) 538; Ascn. & Gr. Syn. 5, 1 (1914) 366.— Achyranthes porrigens JACQ. Hort. Schoenbr. 3 (1798) 54, t. 350.—Telanthera porrigens Moa. in DC. Prod. 13, 2 (1849) 377. Erect or ascending with erecto-patent branches, 1/2-11/4 m high; all hairs minutely dentate; stems thin, firm; young parts densely clothed with ap- pressed acroscopic rather long hairs. Leaves ellip- tic-oblong from an acute or contracted base, acute, mucronate, 1!/2—6!/4 by 1/3-31/4 cm, at first on both surfaces densely clothed with appressed long hairs; hairs subpersistent or those of upper surface gradu- ally disappearing; petiole 2-10 mm. Heads terminal and near the tops of the branches also axillary, not very numerous, highest often in lax umbelliform inflorescences, often 2—3 together on a common pe- duncle and then 1-2 sessile; rest shortly peduncled; peduncle thin, firm, up to 10 cm long but often much shorter; axes of inflorescence and stalks of heads densely clothed with appressed acroscopic hairs; heads at first ovoid, afterwards more cylin- drical, 6-15 by 4-6 mm; lowest flowers already fallen before expansion of highest; axis of head densely hairy; bracts acute or very shortly mucro- nate, + 2 mm long, bracteoles very acute, densely hairy on back, + 2!/4 mm. Perianth sessile between the bracteoles, red, 3—3!/2 mm long. Tepals oblong or ovate-oblong, 1-nerved, in lower half rather densely hairy on the back. Filaments (staminal cup included) 13/4-2 mm; anthers linear, 3/s-1 mm; pseudo-staminodes strap-shaped, about as long as stamens, at top divided into a few narrow strips. Style !/4s—!/2 mm. Fruit broadly oblong, dorsally compressed, 1—1!/4 mm long. Distr. A native of Peru, in Malaysia: once found in West Java as an introduced ornamental, and already met with as a garden-escape, probably not fit for naturalization. Dec. 1949] AMARANTHACEAE (Backer) 95 15. GOMPHRENA LINNE, Sp.Pl. 1 (1753) 224. Annual, or less often perennial herbs. Leaves opposite, sessile or on short petioles. Flowers 9, in terminal, solitary, sessile or subsessile heads or short spikes; recep- tacle cylindric or swollen. Flowers solitary in the axil of a persistent bract, sub- tended by 2 bracteoles; bracts and bracteoles scarious, glabrous; bracteoles erect, navicular, acute, often coloured, with or without dorsal crest, glabrous, falling off with perianth. Tepals 5, erect, free or nearly so, on back long-woolly. Stamens monadelphous; staminal tube long or short, shortly 5-lobed; free parts of filaments with or without intervening pseudo-staminodes, entire, retuse or distinctly 2-lobed; anthers introrse, 1-celled (2-locellate). Ovary compressed, glabrous; ovule pendu- lous from long erect funicle; style short or long; stigmas 2, erect or spreading, short, sometimes almost inconspicuous. Utricle compressed, indehiscent. Distr. Large genus, centering in trop. America, some spp. nativein Australiaand SE. Malaysia, one an introduced ubiquist. Of the species collected in Malaysia 3 are natives of America and have been introduced. The 2 other species are native both in Australia and in SE. Malaysia, and represent a distinct Australian element in the Malaysian flora. Notes. The flowers are in this genus 5, at least in the Malaysian spp. In those of an Australian speci- men of G. brownii Mog. I could find no trace of an ovary. SCHINZ in ENGL. & PR. Nat. Pfi. 2e Aufl. 16c (1934) 26 describes this genus as lacking pseudo-staminodes. But on p. 79 of the same volume he figures 2 species with very distinct pseudo-staminodes. KEY TO THE SPECIES 1. Bracteoles with a (sometimes very narrow) dorsal crest. 2. Dorsal crest of the bracteoles well-developed, very distinct, dentate-serrate on the back. 3. Head globose or depressed-globose, 13/4—2!/4 cm long when adult. Bracteoles deep purple, pink or white, 2—3!/4 mm broad (crest included). Lobes of the staminal tube at least partly obtuse or rounded, not or hardly longer than the unwithered anthers. : 1. G. globosa 3. Heads at first ovoid, afterwards more spiciform, cylindric, 2!/2-5 cm long when adult. Bracteoles orange or reddish, 4-5 mm broad (crest included). Lobes of the staminal tube very acute, much longer than the unwithered anthers. Exclusively cultivated . : 3. G. haageana 2. Dorsal crest of the bracteoles very narrow, dorsally entire, not reaching up to the apex of the bracteole. Flowers white . a : 2. G. celosioides 1. Bracteoles without a dorsal crest. 4. Adult heads 3-4 cm diam. Perianth much compressed, 12-14 mm long, much longer than the bracte- oles. Tepals acute. Staminal tube with long filiform pseudo-staminodes between the filaments, free parts of the latter 5-6 mm long. Style 8-9 mm (spreading stigmas included), exceeding the stamens. Robust herb 2 4. G. canescens 4. Adult heads 3/4-1 cm diam. Perianth not or hardly compressed, 31, a mm long, about equalling the bracteoles or slightly longer. Tepals rather obtuse. Staminal tube without pseudo-staminodes between the filaments, free parts of the latter + '/2 mm long. Style + ‘/2 mm (the erect minute stigmas included), not exceeding the stamens. Small herb 5. G. tenella 1. Gomphrena globosa LINNE, Sp.Pl. (1753) 224; Burm. FI. Ind. (1768) 72; BL., Bijdr. (1825) 548; BLAnNco, FI. Filip. (1837) 198; ed. 2 (1845) 139; ed. 3 (1877) 251, t. 78; DECNE in Nouv. Ann. Mus. 3 (1834) 372; SPAN. in Linnaea 15 (1841) 346; Moa. in DC. Prodr. 13, 2 (1849) 409; Mia. Fl. Ind. Bat. 1, _ 1 (1858) 1050; Suppl. (1860) 150; Hook. f. Fl. Br. Ind. 4 (1885) 732; BAILEY, Queensl. Fl. pt 4 (1910) 1235; PULLE in Nova Guinea 8 (1910) 352; Koorp. Exk. FI. 2 (1912) 201; STUCHLIK in FEDDE, Rep. XII (1913) 337 seg.; ASCH. & Gr. Syn. 5, 1 (1914) 367; MerR. Interpr. Herb. Amb. (1917) 215; En. Born. (1921) 246; En. Philip. Fl.Pl. 2 (1923) 132; Ript. Fl. Mal. Pen. 3 (1924) 10; Heyne, Nutt. Pl. (1927) 609; BAcKerR, Onkr. Suiker. (1930) 231, Atl. t. 242; Gacn. in Fl. gén. I. C. 4 (1936) 1067.— Flos globosus RumMpH. Herb. Amb. 5, 289, t. 100, fig. 2.—Fig. 8. Annual, erect or at base decumbent and rooting, 15-60 cm high; stem thickened at base of inter- nodes, often tinged with red, on young parts ap- pressed pilose. Leaves oblong or oblong-obovate from an acute base, obtuse, often + undulate, thinly pilose on both surfaces, 5-15 by 2-6 cm; petiole 1—1!/2 cm; 2 topmost leaves sessile or nearly so. Heads sessile or subsessile above the topmost pair of leaves, solitary or sometimes in clusters of 2-4, globose or depressed globose, 13/4—2!/4 cm diam.; bracts ovate-triangular, acuminate, acute, 3-6 mm:; bracteoles with very distinct dentate-ser- rate dorsal crest, much surpassing the bract, 7— 12 mm by 2-3!/s mm (crest included), deep purple, 96 FLORA MALESIANA [ser. I, vol. 42 pink or white. Perianth shorter than the bracteoles, yellowish green, white-woolly outside, 6—6!/2 mm long. Staminal tube about equalling the perianth; its lobes at least partly obtuse or rounded, not or hardly longer than unwithered anthers. Style much shorter than staminal tube, bifid. Fruit ovoid, + 2!/2 mm; seed reniform, swollen. Distr. Native of trop. America; long ago intro- duced into Malaysia: found throughout the Archi- pelago either cultivated or semi-naturalized. Ecol. In settled areas in waste places not rarely found as a stray from gardens but nowhere firmly established, and not truly naturalized, 1--+- 1300m. FL. fr. thoughout the year. Uses. Cultivated in gardens as an ornamental. Cooked leaves may be eaten. Vern. Bunga knop, M. Fig. 8. Gomphrena globosa L. from Java, 1/4. 2. G. celosioides MArT. Beitr. Amar. (1825) 93; Moa. in DC. Prod. 13, 2 (1849) 410; SeUBERT in Mart. FI. Bras. 5, 1 (1875) 218; SANDW. in Kew Bull. (1946) 29; STEEN. in Bull. J.B.B. III, 17(1948) 402. Annual, erect or ascending, 10-25 cm long, often branched from the base; stem finely appressed- white-pilose; internode immediately beneath 2 top- most leaves often long. Leaves shortly petioled or subsessile, spathulate or oblong-lanceolate from an acute base, acute or rather obtuse, ending in a short rather hard point, glabrous or sparingly ap- pressed pilose above, thinly or rather densely ap- pressed pilose beneath, 2—4'/2 by !/2-1'/4 cm; 2 top- most leaves subsessile. Heads sessile above highest pair of leaves, at first subglobose, -- 1 cm diam., gradually lenghthening into a spike, finally up to 4cm long; receptacle long-white-woolly; bracts ovate, acuminate, very acute, 3-4 mm; bracteoles long-ovate, very acute, white, -- 6 mm, in the high- er part with a dorsal crest; crest narrow, abruptly ending below top of the bracteole and there spar- ingly shortly, irregularly dentate, otherwise entire. Perianth inserted on a minute knob, pure white, somewhat shorter than the bracteoles, 4!/2—5!/2mm. Tepals narrowly lanceolate, acute, in lower half externally densely clothed with long fine white hairs. Stamens slightly shorter than perianth; fila- ments nearly entirely connate; apical teeth of the staminal tube varying from rather obtuse to rather acute, not or shortly exceeding unwithered anthers, -- 2/3 mm. Style much shorter than the staminal tube, bifid somewhat deeper than halfway down, totalling 3/s-1 mm. Ripe fruit not seen. Distr. Native of trop. America, introduced in trop. and S. Africa, India and Australia, in Ma- laysia: in Batavia, Manila (CoerT no 1400, a. 3.6.36), and Singapore, well on its way to spread over the Old World. The statement of SANDWITH /.c. that the plant has run wild all over the Botanic Gardens at Buitenzorg was due to misinformation. Ecol. Roadside weed, between grass, locally gregarious, tufted in mats (Singapore, harbour- yard, 1926; Batavia, 1946), well adapted to hot, dry, dusty situations at low elevations. 3. Gomphrena haageana KLOTZSCH in OTTO & Dieter. Allg. Gartenz. 21 (1853) 297. For differentiating characters see the key to the species. Distr. Native of Mexico, in Malaysia: exclu- sively and occasionally cultivated. 4. Gomphrena canescens (PorrR). R.BR. Prod. (1810) 416; Moa. in DC. Prod. 13, 2 (1849) 398; Bru. Fl. Austr. 5 (1870) 253; HeMsL. Rep. Bot. Chall. 1, 3 (1884) 184; BarLey, Queensl. Fl. pt 4 (1901) 1235; SrUCHLIK in FepDpDE, Rep. XII (1913) 345; Domin, Beitr. Pflanzengeogr. Austr. 1, 2(1929) 644.—Philoxerus canescens Porr. in LAMK. Tabl. Enc. Bot. Suppl. 4 (1816) 393.—Gomphrena lanu- ginosa SPAN. in Linnaea 15 (1841) 346, ic. ined. 51 (non vidi); Miq. Fl. Ind. Bat. 1, 1 (1858) 1051. Erect annual, 60-90 cm high, not or sparingly branched, stem hard, densely clothed with erect appressed long white hairs. Leaves sessile, linear Dec. 1949] or narrowly linear-lanceolate, acute, firmly herba- ceous with a strong midrib, on both surfaces den- sely clothed with appressed long white hairs, 2!/2— 5 cm by 2-5 mm. Heads subtended by an involucre of 5-9 patent cauline leaves, depressed globose, 3-4 cm diam.; involucral leaves narrowly linear, tapering to a very acute apex, l-nerved, at the base long-ciliate, on both surfaces densely clothed with appressed long white hairs, 2-3 by 4-6 mm; recep- tacle densely long-white-woolly; bracts and bract- eoles tapering from a broad base, bracts 6-7 mm long; bracteoles very acute + 8 mm. Flowers be- tween the bracteoles on a very short thick pedicel; perianth strongly compressed, purple, greatly sur- passing the bracteoles 12-14mm long. Tepals nearly free, lanceolate, acute, |-nerved, in the lowest 1/3-1/2 of the back (especially at the base) clothed with long hairs. Stamens 7—9 mm long, at the base for a length of 2-4 mm connate into a tube; free parts of filaments flat, glabrous 5-6 mm, alternat- ing with shorter subulate acute anantherous pseu- do-staminodes. Ovary glabrous; style glabrous, 8—9 mm, longer than the stamens, shortly bifid; arms spreading. Fruit? Distr. N. Australia, in Malaysia: Lesser Sunda Isl. (Timor, SPANOGHE), Moluccas (Tanimbar, RIEDEL). Ecol. Probably in sunny dry localities, in Timor on calcareous rocks, apparently rare, only twice collected. 5. Gomphrena tenella (Moq.) Bru. Fl. Austr. 5 (1870) 256.—Iresine tenella Mog. in DC. Prod. 13, 2 (1849) 343. Erect annual, Malaysian specimens only 2!/2— 6 cm high, unbranched; stem thin, hard, in higher AMARANTHACEAE (Backer) 97 part densely woolly. Leaves in 2—4 distant pairs, narrowly linear, glabrous, above, thinly patently pilose beneath, !/2—3/4 cm long. Heads sessile above the topmost pair of leaves, globose, 3/4—1 cm diam.; receptacle very densely clothed with longish white hairs; bracts ovate, shortly apiculate, -- 2!/2 mm long, much shorter than the bracteoles; axis of flower densely woolly; bracteoles ovate, acute, ++ 3'/2mm long. Tepals nearly free, oblong, rather obtuse, 3!/2-4 mm long, in the lower half with an oblong herbaceous central field; this field on the back densely clothed with long, entangled, in dried specimens brownish hairs. Filaments connate up to near apex; staminal tube not shorter than ovary and style together; free parts of filaments + !/2 mm long, broad, slightly narrowed upwards, without intervening pseudo-staminodes. Style minute; stig- mas 2, erect, subulate; style and stigmas together + 1/2 mm long. Fruit? Distr. N. Australia, in Malaysia: Moluccas or the Lesser Sunda Islands. The only specimen on which this record is based was collected by REIN- WARDT in 1821, and is preserved in the Rijksher- barium, Leyden. According to an accompanying note by BLUME it was collected somewhere in the Moluccas, but it may have been gathered in Flores Island. Notes. I am not wholly satisfied that my iden- tification of the specimen described above is cor- rect. The dimensions of its vegetative parts are very much smaller than those given by MOQUuUIN and BENTHAM for Australian specimens. However, it is possible that REINWARDT collected the plant in a very sterile or arid locality. I have not examined any other specimen of this species. 16. IRESINE P. BROWNE, Hist. Jamaica (1756) 358. Erect, ascending or scandent herbs or undershrubs. Leaves opposite, petioled, entire or subentire. Flowers (d)() or 9, paniculate; panicles terminal, often also in the highest leaf axils, many-flowered; their ultimate branches spiciform, bearing solitary or clustered, minute flowers; flowers solitary in axil of bract, subtended by 2 bracteoles, membranous, usually shining, subglabrous or woolly. Tepals 5, oblong or ovate-oblong, acute; stamens (in 9 reduced to very minute staminodes) 5; filaments at the base connate in shallow cup, filiform, with or without inter- posed pseudo-staminodes; these usually short, broadly triangular, rarely long; anthers oblong, 1-celled (2-locellate); ovary (wanting in d) compressed, ovule 1, pendulous from an erect funicle; style very short; stigmas 2, subulate, erect-ascend- ing, short; utricle compressed, orbicular, thin-walled, indehiscent; seed lenticular or reniform, shining. Distr. Large genus, centering in America, also in the Galapagos Isl.,in Malaysia: one Brazilian species cultivated and locally naturalized. 1. Iresine herbstii Hook. f. in Gard. Chron. (1864) 654, 1206; Bot. Mag. (1865) t. 5409; AscH. & GR. Syn. 5, 1 (1914) 369; Merr. En. Philip. 2. (1923) 132; HeyYNe, Nutt. Pl. (1917) 609; BatLey, Stand. Cycl. 2 (1935) 1662; BAcKER, Bekn. FI. Java (emerg. ed.) IV A, fam. 63 (1942) 12.—Jresine celosioides (non LINNE!) BOERL. in Verh. 2e Ge- nees- en Nat. Congr. Leiden (1889) 148; BACKER, Ann. J. B. B. Suppl. 3 (1909) 405; Koorp. Exk. FI. 2 (1912) 201. 98 FLORA MALESIANA [ser. I, vol. 42 Perennial herb, erect or ascending, often much branched, slightly fleshy; nodes thickened; base of the internodes in dried specimens strongly con- stricted; young stems pubescent, especially on and near nodes. Leaves broadly ovate-orbicular, broad- ly oval or broadly obovate, at the frequently slightly unequal base very obtuse, rounded or trun- cate but shortly contracted into the petiole, with a usually more or less deeply emarginate but some- times rounded or shortly acuminate and then very acute apex, entire or subentire, concave, thinly fleshy, either shining dark red with lighter coloured bands along the main-nerves or (var. aureo-reticu- lata NicH.) green with golden yellow bands, on both surfaces very thinly clothed with shining brown or yellowish, partly bifid and appressed hairs, 2!/2-8 cm long and wide; petioles 1—5 cm; those of a single pair connected at the base bya transverse row of longish, thickis, harticulate hairs. Panicles terminal and often also in axils of the highest and then small leaves, 5—5SO cm long, united in a terminal, erect, 8-60 cm long, often much branched and rich-flowered panicle; primary branches of the panicle erecto-patent, branched from base or nearly so; ultimate branchlets spici- form, rather dense-flowered, 1—4 cm long; rachises of the panicle thin, red, at first rather densely cloth- ed with longish shining hairs; glabrescent. Flowers not concealing the rachis, in Malaysia exclusively 9; bracts, bracteoles and perianth greenish white or yellowish white; bracts and bracteoles persistent after fall of the perianth, ovate, rather acute, con- cave, nerveless, glabrous; bracts + 1 mm long; bracteoles slightly wider, + 11/4 mm long. Perianth + 11/4 mm long, at the base externally with a dense whorl of long, very thin, white hairs; tepals dor- sally with a few short hairs, otherwise glabrous, oblong, rather acute, nerveless; pseudo-staminodes minute. Ovary suborbicular, much compressed, glabrous; stigmas obliquely patent-ascending, -+ 1/2 mm. Fruit never produced in Malaysia. Distr. Native of Brazil, in Malaysia: exclusive- ly cultivated for ornamental purposes (already be- fore 1894) and locally naturalized in e.g. Java, Celebes, and Ceram. Ecol. Naturalized in forest borders and along forest paths, 500-1500 m, locally abundant. Uses. The leaves are squeezed in water in order to obtain a red dye used for colouring agar agar jellies. “Vern. Bayam merah, M. Notes. In Malaysia exclusively 9 flowers are found. In none of the many specimens I examined, I have found d flowers nor have I traced any de- scription of them. Fruit is never produced in Ma- laysia. Excluded Nevrolis fuscata RAFINESQUE, Autikon Bot. (1840) 150. This new genus was based by RAFINESQUE on Celosia virgata Hort. (non JAcQ.). Its native origin was cited as: ‘Borneo or Moluccas?’ I cannot iden- tify this with certainty. It is possible that MERRILL (Rafin. p. 119) is right in reducing it to Celosia. RAFINESQUE might have had a specimen belonging to a garden form of JACQUIN’s species which is na- tive in South America. CHENOPODIACEAE (C. A. Backer, Heemstede) Annual or perennial herbs or shrubs, often fleshy, glabrous, papillate or hairy. Leaves opposite or alternate, exstipulate, sometimes seemingly wanting, stalked or sessile, entire, dentate-serrate-lobed or irregularly gashed. Flowers solitary, 2—3-nate or glomerate, usually sessile, either axillary or in terminal or axillary dense or inter- rupted spikes or panicles, 9 or unisexual, monochlamydous, rarely achlamydous, small; bracts present or absent, usually small, rarely leafy. Perianth herbaceous or sometimes scarious, rarely (in 9) absent, 3—5-partite with (in bud) imbricate seg- ments, or sometimes almost entirely gamophyllous and then shortly lacerate-den- tate or unilaterally cleft, persistent, after anthesis accrescent or not. Stamens often the same number as tepals and opposite to them, sometimes fewer, usually inserted on or near base of perianth; filaments free or shortly connate; anthers dorsifixed or inserted in a basal cleft, 2-celled (4-locellate); cells bursting longitudinally. Ovary free or at the base adnate to the perianth, |-celled; ovule 1, basal, sessile and erect or suspended from a funicle; styles or stigmas 2-5, linear. Utricle either enclosed by the perianth or not, indehiscent or rarely operculate; seed erect, oblique or horizontal, usually compressed; endosperm mostly present, peripheral, sur- rounding the embryo; embryo annular or spirally twisted. Distr. Species numerous, inhabitants of the temperate and tropical zones of both hemispheres. Ecol. Often in maritime, saline, or in permanently or periodically dry regions. Many are weeds of cultivation. In Malaysia restricted to maritime and mountainous districts. Uses. In Malaysia some species are cultivated as vegetables, others as medicinal or ornamental plants, but not for the production of sugar. (ARTIFICIAL) KEY TO THE GENERA 1. Apparently leafless; stem spuriously articulate. Leaves decussate; those of each pair connate through- out their length into a tubular fleshy sheath which tightly and entirely encloses the appertaining inter- node and is widened at the apex into a shallow cup. Flowers spicate; bracts decussate, crowded; flowers in the axil of one bract + collateral; perianth gamophyllous, scarious, + flagon-shaped; stamen 1 (sometimes 2?). 2. Floral bracts almost entirely connate into a cup; base of the cup on 2 opposite sides provided with collateral apertures for the protrusion of stamens and stigmas. Apex of perianth laterally split. 5. Arthrocnemum 2. Floral bracts almost free; their margin dilated into a patent semi-orbicular scale, their base without apertures. Apex of perianth shortly dentate-lacerate . . . . . . . . +. £4, Tecticornia 1. Leafy in a normal way; leaves alternate. 3. Leaves spine-tipped. Bracts exceeding the perianth. Tepals about the middle with a transverse thickening, above this thickening scarious . . dh Ue ote ee ee SD 3. Leaves not spine-tipped. Tepals without a transverse Pickenioe 4. Leaves sessile, narrowly linear, semi-terete, very succulent. Bracteoles distinct though small, trans- parent, after anthesis Stellately Spreading =... s 46 se me ee ee 4. Leaves distinctly petioled, flat. 5. Flowers (3d) (9); 6 flowers in spicate or panicled clusters; 9 flowers iomey in leaf axils. No bracteoles. In Malaysia exclusively cultivated. . . . . . . 3. Spinacia 5. Flowers all clustered, all or for the greater part 0; the Q, when present, in the same cluster as the Od ones. 6. Ovary free. Flowers after anthesis not coalescent. No bracteoles pei eae Dae Chenopodium 6. Ovary adnate to base of perianth. Flowers coalescent after anthesis. Bracteoles minute. In Ma- faysia- only cultivated: 0 9 sa ec ete yn 1. BETA LINNE, Sp. Plant (1753) 222. In Malaysia not wild; 1 species frequently cultivated in the mountainous regions of Java. 100 FLORA MALESIANA [ser. I, vol. 4 1. Beta vulgaris LINNE, Sp. Plant. (1753) 222; f. cicla [LINNE, Sp. Plant. (1753) 222, pro sp.]. Ascu. & Gr. Syn. 5, 1 (1913) 12; OCHsE & BAKH. Dutch: Snijbiet. Leaves eaten as spinach; root Vv. D. BR. Veget. (1931) 102. rather tough, unfit for human food. In Java 2 forms cultivated, exclusively for the f. rapa [Dum., Fl. Belg. (1827) 21 pro sp.]. Dutch: European table, not eaten by the Indonesians, Kroot, roode biet. Leaves not eaten. Root fleshy, both of them unfit for the manufacture of sugar: dark red, edible after cooking. 2. CHENOPODIUM EINNE, Sp-PIt ed: 1 (1753) 218: Annual or perennial herbs, sometimes strongly smelling; young parts often more or less densely clothed with minute, powdery, white or pink vesicles which, when young, contain a watery liquid but usually soon shrivel and lose their colour. Leaves alternate, petioled, herbaceous, variable as to shape, entire, dentate-serrate or irregularly gashed. Flowers 9 or by abortion Q, sessile, clustered; clusters solitary in the leaf axils or in axillary and terminal cymes, spikes or panicles; no bracteoles. Tepals 5 or sometimes 4, free or shortly connate, vaulted, herbaceous, often longi- tudinally thickened or keeled on the back, in the Malaysian species not fleshy after anthesis. Stamens in 9 the same number as tepals, inserted on the base of the peri- anth or free from it, at the base sometimes connate into a fleshy disk. Ovary de- pressed globose; style short; stigmas 2-5. Fruit often embraced by conniving tepals, thin-walled indehiscent. Seed usually horizontal, sometimes oblique or vertical, shining or dull, smooth or finely tuberculate, lenticular; its margin keeled or not; testa thinly coriaceous; embryo annular, surrounding the usually mealy en- dosperm. Distr. Species about 60 in the temperate zones of both hemispheres. A small number naturalized in the mountainous districts of the tropics. Ecol. For the greater part weeds of cultivation; many prefer a fertile soil. Uses. A few species cultivated either for their oil-producing fruits, as a substitute for tea, or for orna- mental purposes. Notes. Most species flower and fruit freely. For the determination ripe fruits are of value. The colour of young leaves should be noted by Collectors. BurRMAN (FI. Ind. (1768) 72) records Chenopodium urbicum L. as having been sent to him from Java. The specimen mentioned by him is conserved at Geneva and has been correctly named. Nevertheless I have omitted the species from my key as I feel not at all convinced that it was really collected in Java; BURMAN was often not careful in his records. Ch. urbicum was never afterwards collected in Malaysia. KEY TO THE SPECIES 1. Top of ovary and fruit studded with (in vivo yellow) glands; stigmas 2-5; embryo encircling only 1/2-2/3 of the seed. Young vegetative parts and outside of perianth without powdery white or pink vesicles. Undersurface of leaves with (sometimes rather indistinct) yellow glands. Strongly smelling. 1. Ch. ambrosioides 1. Top of ovary and fruit glandless; stigmas 2; embryo encircling almost the entire seed. Young vege- tative parts and outside of perianth with powdery white or pink vesicles. Leaves without any yellow glands. Not or faintly smelling. 2. Segments of perianth after anthesis widely a not covering the fruit. Young parts without paeoutery vesicles, quite glabrous ae: . . .2. Ch. polyspermum . Segments of perianth after anthesis connivent, covering the fruit. Young, or at least very young, parts with powdery white or red vesicles. 3. Only very young parts with distinct vesicles; these white, very soon shrivelling and losing their colour; old leaves on both surfaces dark green and feebly shining. Seed (after removal of pericarp) dull black, rather obscurely Sa Panicles small; terminal panicle not much larger than the axillary ones... . . 3. Ch. murale 3. Vesicles white or red, not very soon Josing ‘their colour; old leaves “Ae dark green on both surfaces. 4. Larger leaves irregularly and rather coarsely serrate-dentate-laciniate or deeply gashed, often longer than 5 cm and wider than 3 cm; undersurface of leaves not very densely and persistently vesiculose. Seed after removal of pericarp shining blackish brown, almost smooth 4. Ch. album Dec. 1949] CHENOPODIACEAE (Backer) 101 4. Leaves quite entire, small (mostly 2-4 by 1'/2-3 cm), on the undersurface very densely and per- sistently vesiculose; hence in a dried state very pale beneath; their nerves and margins often reddish. Flower-clusters spicate; higher spikes united in a terminal, small, leafless, paniculate inflorescence; rachises of the inflorescence and outside of the perianth densely clothed with patent, oblong, reddish vesicles 1. Chenopodium ambrosioides LINnng£, Sp.PI!. (1753) 219: BLANCO, FI. Filip. (1837) 200; ed. 2 (1845) 140; ed. 3, 1 (1877) 253, tab. 69; Moa. in DC. Prod. 13, 2 (1849) 72; WIGHT, Ic. 5 (1852) tab. 1786; Mia. Fl. Ind. Bat. 1, 1 (1858) 1017; Bru. FI. Austr. 5 (1870) 162; Hook. f. Fl. Br. Ind. 5 (1886) 4; BAILEy, Queensl. FI. pt 4 (1901) 1243; Koorp. in Nat. Tijd. N.I. 60 (1901) 256; Back. in Ann. J.B.B. Suppl. 3 (1909) 398; Courcn. in FI. gén. I.C. 5 (1910) 5; Merr. Fl. Man. (1912) 189; Koorp. Exk. Fl. 2 (1912) 189; Ascu. & Gr. Syn. 5, 1 (1913); MERR. Sp. Blanc. (1918) 136; En. Philip. FI.Pl. 2 (1923) 125; Hear, Ill. Fl. Mitt. Eur. 3 (1912) 233; HEYNE, Nutt. Pl. 2 (1927) 602; Domin, Beitr. Pflanzen- geogr. Austr. 1, 2 (1929) 618; Burk. Dict. Ec. Pr. 1 @935) 523. Erect or ascending annual, often very much branched; entire plant strongly smelling, without powdery vesicles, 15 cm to 1 m high; stem angular- ribbed, glabrous or finely pubescent. Leaves ob- long-lanceolate from a narrowed or contracted, acute, often more or less decurrent base, acute or rather obtuse; larger ones coarsely or shallowly serrate-dentate; smaller ones less deeply incised; highest entire, gradually changing into bracts; all leaves herbaceous, bright green, on the undersur- face more or less densely studded with in vivo yel- low (sometimes very inconspicuous) glands, other- wise subglabrous or sparingly beset with short white hairs, 11/2-15 by !/2-5 cm; midrib prominent beneath; other nerves thin; petioles short or me- dium-sized: those of highest leaves obsolete. Flow- er-clusters small, 3—25-flowered, in the axils of suc- cessive, conspicuous, narrow, + bractlike leaves, united in short or longish, rather lax spikes, form- ing together a leafy panicle; floral leaves much sur- passing the clusters, acute, with a strongish midrib. Flowers 9 or partly 9, sometimes partly d and then with 4-5 stamens and a rudimentary, densely glan- dular ovary. Perianth light green with a pale base, 11/4-11/2 mm long, 4~S-cleft to near the base; seg- ments ovate-triangular, rather acute, very concave, not or indistinctly keeled. Stamens in 5 4—5, rarely 1-3; filaments slightly exceeding the perianth. Ovary depressed globose, on top with many small, in vivo yellow glands; stigmas 2—5, usually 3 or more. Ripe fruit entirely concealed by the conniv- ing tepals. Seeds horizontal, or a few (rarely many or all) erect, broadly oval-obovoid, shining brown- ish black, 2/3-+/; mm diam.; embryo encircling 1/3_2/3 of the seed. Distr. Native of tropical America, introduced in many other regions: Europe, Asia, Africa, Aus- tralia, in Malaysia: naturalized throughout the Philippines, in Java and in N. Celebes (Minahasa). At present found in a wild state in the western and the eastern part of Java between 1600 and 2000 m. Ecol. Road-sides, locally often very numerous. In Celebes collected in dry ricefields. F/. Jan.—Dec. 5. Ch. acuminatum Uses. The slightly poisonous oil distilled from the seeds is used as a remedy against ankylostomi- asis. For this purpose the plant has been cultivated in Java, but it cultivation was abandoned as being insufficiently remunerative. Notes. In Java the seeds are all or nearly all hori- zontal. The Celebian specimen has vertical seeds (see also the description by Moquin, I.c. p. 73) but in all other respects it completely agrees with the Javan material. It was collected in 1840 by For- STEN and had as late as 1949 still distinctly retained its peculiar smell. 2. Chenopodium polyspermum LInné, Sp.P1.(1753) 220; Moa. in DC. Prod. 13, 2 (1849) 62; AscnH. & Gr. Syn. 5, 1 (1913) 26; Heat, Ill. Fl. Mitt. Eur. 3 (1912) 222; Merr.in Philip. J.Sc. 5 (1910) Bot. 145; En. Philip. Fl.Pl. 2 (1923) 126. Annual, erect or ascending, quite glabrous, glandless, without a distinct smell, without powdery vesicles, 4-100 cm (mostly 15-75 cm) long, dull green, rarely shining, in a living state often more or less strongly tinged with purple, frequently much branched (often from quite near the base); branches obliquely erect or the lower widely patent-ascending, not rarely partly pros- trate; stem and branches angular. Leaves herba- ceous, rarely fleshy, entire or the larger ones some- times with one or a few triangular teeth near the base; lower leaves on rather long petioles, ovate of ovate-oblong from a cuneate or obtuse, rarely hastate base, shortly contracted into the petiole, acute or obtuse, rarely subretuse, frequently very shortly mucronate, 4-11 by 1'/2-6cm; highest leaves much smaller, oblong-lanceolate. Cymes very numerous, most variable as to size, dense or rather loose, few- to many-flowered; highest very often collected in a narrow, paniculate inflores- cence (reminding of Amaranthus gracilis). Flowers sessile; tepals oval-obovate or oval-oblong, obtuse, thin, with a very distinct, not-keeled midrib, 11/3— 11/2 by !/3-!/2 mm, after anthesis widely patent, not concealing fruit; stamens 5, on the base of the peri- anth, + equalling tepals; stigmas 2, erect or suber- ect, minute, persistent. Fruit depressed-globose, + 11/3 mm diam., very thin-walled. Seed blackish brown, feebly shining, very faintly striolate. Distr. Europe, continental Asia; elsewhere lo- cally introduced. Ecol. Weed of fields and gardens; waste places. Notes. I have seen no Malaysian specimen but MERRILL /.c. records the plant as a casual weed in the Philippines. The species is very variable. 3. Chenopodium murale LINNE Sp.PI. (1753) 219; Moo. in DC. Prodr. 13, 2 (1849) 69; Mia. FI. Ind. Bat. 1, 1 (1858) 1016; Bru. Fl. Austr. 5 (1870) 160; Hook. f. Fl. Br. Ind. 5 (1886) 4; Bart. Queens]. FI. pt 4 (1901) 1243; Koorp. in Nat. Tijdsch. N.I. 60 102 FLORA MALESIANA [ser. I, vol. 42 (1901) 256; BAck. in Ann. J.B.B. Suppl. 3 (1909) 398; Koorp. Exk. FI. 2 (1912) 190, 191; AscH. Gr. Syn. 5, 1 (1913) 33; Heat, Il. Fl. Mitt. Eur. 3 (1912) 223; Domin, Beitr. Pflanzengeogr. Austr. 1, 2 (1929) 618. Annual, erect or ascending, often much branch- ed, slightly fetid, without any yellow glands, 15— 80 cm long; all vegetative parts and outside of peri- anth, when very young, clothed with white, pow- dery vesicles, otherwise glabrous; vesicles soon shrivelling up and losing their colour; stem and leaves then darkgreen, somewhat shining; stem angular or ribbed; leaves ovate-rhomboid-elliptic- oblong from a cuneate base, acute, irregularly coarsely acutely dentate-serrate, herbaceous, 1!/2— 12 by 3/4-7!/2 cm; petioles of lower leaves 3-5 cm, of higher leaves gradually smaller, of topmost ones often very short. Flowers in axillary and terminal, 1-4 cm Jong panicles with erecto-patent branches, densely clustered, S-merous; 9; terminal panicle not or hardly longer than the axillary ones; peri- anth herbaceous. Tepa/s distinctly connate at the base, oval, with rounded or very obtuse tips, faintly keeled on the back, 11/2-2 mm long; stamens 5, on the very base of the perianth; stigmas 2, short. Fruit tightly enclosed by incurved tepals, depressed globose, 1'/s—-1'/2 mm diam.; pericarp very thin, not readily separating from the seed. Seed hori- zontal, encircled by a very distinct rather sharp keel; testa (after removal of the pericarp) dull black, very finely papillate; embryo encircling al- most the entire seed. Distr. Possibly originating from continental Asia but since long widely spread in Europe, Ame- rica, Africa and Australia. In Malaysia: introduced in Java already more than 90 years ago, but as yet still confined to Mt Tengger in the eastern part between 1900 and 2300 m. Ecol. Fields and gardens, road-sides, locally very common. Vern. Dieng, J, dieng dempo, J, dieng idjo, J. 4. Chenopodium album LINNE£, Sp. Plant. (1753) 219; Moa. in DC. Prod. 13, 2 (1849) 70; Mia. FI. Ind. Bat. 1, 1 (1858) 1017; Bru. Fl. Austr. 5 (1870) 159; Hook. f. Fl. Br. Ind. 5 (1886) 3; BAIL. Queensl. Fl. pt 4 (1901) 1242; Koorp. in Nat. Tijd. N.I. 60 (1901) 256; BAck. in Ann. J.B.B. Suppl. 3 (1909) 398; Koorp. Exk. FI. 2 (1912) 191; AscH. & Gr. Syn. 5, 1 (1913) 38; Hear, Ill. Fl. Mitt. Eur. 3 (1912) 225; Domi, Beitr. Pflanzen geogr.Austr. 1, 2 (1929) 617. Erect annual, often much branched, not mark- edly smelling, without any yellow glands, 15 cm— 1!/2m high; all vegetative parts and outside of perianth, when young, densely clothed with white or partly amaranthine powdery vesicles, otherwise glabrous; the amaranthine vesicles soon turning white; all old vesicles shrivelling up and losing their colour. Stem angular, ribbed, with longitu- dinal dark green or red streaks. Lower /eaves long- petioled, ovate-rhomboid, irregularly and rather coarsely dentate-serrate-laciniate or deeply gashed; _ higher ones gradually shorter-petioled, elliptic- oblong-lanceolate from an acute or contracted base, acute or obtuse, less deeply incised or entire; all leaves herbaceous, 1!/2-15 by !/2-13 cm. Flowers in panicled clusters, 5-merous, 5; panicles often collected in a large terminal leafy paniculate in- florescence. Perianth herbaceous, not becoming fleshy after anthesis. Tepals distinctly connate at the base, oval, very concave, obtuse, with a strong, rounded midrib, 11/2-21/4 mm long. Stamens slight- ly longer than the perianth. Ovary depressed glo- bose; stigmas 2, short. Fruit in the living plant en- a diay y ip Rim, i Kee Wi my ‘i, ff cy oF ¥ % Ri Ly i « Fig. 1. Tecticornia cinerea (F.v.M.) BAIL. from saline coastal flats N of Mt Baluran; drawn after a living specimen by A. HAMzAH, Nov. 1941, x 4/2. Dec. 1949] CHENOPODIACEAE (Backer) tirely enclosed by the incurved tepals, depressed- globose, finely papillate. Seed horizontal, lenticu- lar, surrounded by an obtuse keel, shining blackish brown, smooth or nearly so, 1!/4—13/4 mm diam. Distr. Europe, continental Asia, Africa, Aus- tralia, America, in Malaysia: in Java introduced already very long ago, naturalized in the eastern part of the island, 800-1800 m. Ecol. Fields, gardens, locally common. Vern. Dieng, J, dieng putih, J. Notes. Extremely polymorphous, buried by fanatic species-splitters under a mountain of unnecessary and useless names. subsp. amaranthicolor CostE & REYN. in Bull. Herb. Boiss. II, 5 (1905) 979; HEYNgE, Nutt. Pl. (1927) 602. —Chenopodium amaranticolor (CosTE & REYN.) CostTE & REYN. in Bull. Soc. Bot. Fr. 54 (1907) 181; Ascu. & Gr. Syn. 5, 1 (1913) 66. Young parts and outside of perianth densely clothed with amaranthine vesicles, retaining their colour during a long time but at last turning pale and shrivelling. Plant up to 2!/2 m high. Otherwise like the main species and in a dried state indistinguishable from it. Introduced in Java long ago, at present naturalized in the western and the eastern part between 1200 and 2300m above sea-level. Roadsides, fields, locally often numerous. By the Indonesians sometimes cultivat- ed as a vegetable. Note. In a living state very conspicuous by the bright amaranthine colour of the young parts. A specimen (LORZING 7168) of Ch. album was in 1920 collected in N. Sumatra. I have seen it only in a dried state and it bears no note indicating the colour of the young leaves, so that it is impossible to make out whether it belongs to the typical species or to its subsp. amaranticolor. 103 5. Chenopodium acuminatum WILLD. (non SCHUR) Acta Nat. Cur. 2 (1799) 124, tab. 5, fig. 2; Moa. in DC. Prod. 13, 2 (1849) 62; Merr. in Philip. J. Sc. 3 (1908) Bot. 405; En. Philip. Fl. Pl. 2(1923) 125. Annual, erect or ascending, 20-30 cm _ high, branched or not; all young parts, especially under- surface of leaves, clothed with oblong, red or white vesicles; stems and branches angular; branches obliquely erect; their vesicles, except on the an- gles, subpersistent. Leaves entire, in sicco thickish, retaining their indumentum of vesicles during a long time especially on the undersurface where it is very dense, hence in sicco very pale beneath, often with a reddish margin and a reddish undersurface of nerves; lower leaves on rather long, reddish vesiculose petioles, ovate from a very obtuse or rounded-subtruncate, shortly contracted base and an obtuse or rounded, very shortly pointed apex, 2-4 by 11/2~-3 cm; higher leaves on shorter petioles, shorter, narrower, acute, distinctly and finely acu- minate. Rachises of inflorescence densely clothed with reddish vesicles. Flower-clusters subglobose, small, dense, spicate, crowded or the lower rather distant; higher spikes united in a terminal, rather small leafless paniculate inflorescence. Flowers sessile, outside densely reddish-vesiculose. Tepals 5, broadly oval, obtuse or rounded, very concave thin, 1-nerved, 1!/3—-1!/2 mm long, before and after anthesis connivent. Stamens 5, about equalling the perianth; anthers thick; ovary (in our specimens) minute. Style short, 2-armed. Fruit (not seen) finely wrinkled; seed dull black or shining. Distr. Siberia, China, Japan, Formosa; in Ma- laysia: Philippines (Batan Isl., between Formosa and Luzon). Ecol. Waste places. Note. I could examine only 2 specimens, neither of them collected in Malaysia. 3. SPINACIA Linné, Sp. Pl. (1753) 1027. In Malaysia not wild; 1 species, the true spinach, occurs but rarely cultivated. 1. Spinacia oleracea LINNE, Sp.Pl. (1753) 1027, the true spinach, is but very rarely cultivated in Malaysia and only by way of experiment. It is sub- stituted by two quite different species, Amaranthus tricolor L. (Amaranthaceae), bayam, and Tetra- gonia expansa Murr. (Aizoaceae), New-Zealand spinach, kabak, M. 4. TECTICORNIA Hook. f. in Bro. & Hook. f. Gen. Pl. 3 (1880) 65. Only seemingly jointed and leafless, glabrous, softly succulent herb. Leaves decussate; those of each pair connate throughout their length in a tubular fleshy sheath tightly and entirely enclosing the appertaining internode and widened at the apex into a shallow, faintly bilobed cup, which embraces the slightly attenuate base of the next higher sheath. Flowers minute, spicate; spikes terminal and frequently also in highest leaf axils, oblong or shortly cylindrical, very obtuse, bracteate; -bracts decussate, densely crowded, almost free, not provided with basal apertures, fleshy; their margin dilated into a semi-orbicular, + patent, thin-margined scale. Flowers often in threes, sessile, collateral, 9; perianth flask-shaped, very thin, with 104 FLORA MALESIANA [ser. I, vol. 4 a narrow, dentate-lacerate mouth. Stamen 1; filament short; anther exsert, oblong- linear. Ovary ovoid-oblong, compressed, narrowed into a longish, shortly bifid style; ovule subsessile. Utricle erect, pericarp very thin, hyaline. Seed oblong, compressed, densely papillate; albumen hard; embryo slender, radicle inferior. Distr. Monotypic, Australia and S. Malaysia. 1. Tecticornia cinerea (F.v.M.) Bam. Queensl. Fl. 4 (1901) 1261; VALET. Bull. Dép. Agr. Ind. Néerl. 10 (1907) 9; Kocu in ZW. Nieuw-Guin. Exp. 1904/5 (1908) 505; PULLE in Nova Guin. 8 (1910) 349.—Halocnemum cinereum F.v.M. Fragm. 1 (1858) 140.—Salicornia cinerea F.v.M. Fragm. 6 (1868) 251.—Fig. 1. Perennial (?), much branched, erect or prostrate with erect branches, 15-30 cm long; young stems after removal of leaf-sheaths thinly wiry, tough; old stems rather robust, cylindrical, woody; adult internodes 1-2 cm; whole plant in a dried state greyish. Spikes usually 1 terminal and 2-6 in higher axils; the latter spikes opposite, widely patent, 1/2-21!/4 cm long. Utricle brown, 11/4—13/4 mm long; seed yellowish brown. Distr. N. Australia, in Malaysia: SW. New Guinea, S. coast near Merauke, (long. 140° E, G. M. VERSTEEG 1895; 8.9.07, KOCH s.n.) and coast near Mt Baluran, E. Java (long. 114°25’ E, Hoo- GERWEREF s.n. Novy. 1941). Ecol. Low, clayey, seasonally swampy and again completely desiccating localities, cut up after a prolonged dry period in their upper layers into hard clumps by criss-crossing cracks, locally gre- garious. According to Kocu the loamy plain was inundated in the rainy season, and had (had) prob- ably connection with the mouth of the Digul river; possibly the soil was, therefore, saline. It is also said to have been collected in Jmperata-fields, but we should bear in mind that non-botanists are apt to confound Jmperata with other grasses. Notes. Making an accurate detailed description of this species based on herbarium-specimens is impossible, owing to the extreme shrinking of dried materials and their pappiness after soaking. A new description after the living plant is urgently needed. Though Hooker based his new genus Tecticornia on this species only, he did not create the binomium, which is often wrongly ascribed to him. 5. ARTHROCNEMUM Moa. Chenop. Enum. (1840) 111. Only seemingly jointed and leafless herbs or undershrubs. Leaves decussate; those of each pair connate almost throughout their length in a tubular fleshy sheath, tightly and entirely enclosing the appertaining internode and widened at the apex into a rather shallow, faintly 2-lobed cup which embraces the shortly at- tenuate base of the next higher sheath. Flowers minute, spicate; spikes terminal, on often very short lateral branches, bracteate; bracts crowded, decussate, pairwise connate into a very faintly 2-lobed cup, embracing the base of the next higher cup; cups very fleshy at base, gradually thinner upwards, at the base on 2 opposite sides with 3 small collateral apertures for the protrusion of the adult anthers and stigmas (over the margin of the next lower cup); pairs of aperture-triads decussate. Flower usually 1 behind each aperture, 9 or unisexual; bracteoles narrowly linear- spathulate, thinly membranous. Perianth thinly membranous, gamophyllous, at the apex or also deeper unilaterally cleft. Stamen 1 (sometimes 2?); filament short; anther thick. Ovary membranous; stigmas 2, subulate, rather long. Utricle (not seen In a ripe state) membranous or hard. Seed compressed; albumen well-devel- oped, one-sided, mealy; embryo comma-shaped. Distr. Species + 12, along the Mediterranean coast and in the warmer regions of N. America, Asia and Australia, in the coastal districts and salt-marshes, in Malaysia 1 species. 1. Arthrocnemum indicum (WILLD.) Mog. Chenop. Decne in Nouv. Ann. 3 (1834) 370; SPAN. in Lin- Enum. 113; in DC. Prod. 13, 2 (1849) 151; Mia. Fl. Ind. Bat. 1, 1 (1858) 1020; Fors. Nat. Wand. (1885) 515; Hook. f. FI.Br. Ind. 5 (1886) 12.— Salicornia indica WILLD. in Nova Acta Hist. Nat. 5, 111, t. 4, fig. 1; Roxs. Fl. Ind. ed. 1, 1 (1820) 185; naea 15 (1841) 345; WiGurt, Ic. 3 (1845) 737.—S. fruticosa DECNE (non L.) in Nouv. Ann. 3 (1834) 370; SPAN. in Linnaea 15 (1841) 345.—Salicornia spec. in ZOLL. Syst. Verz. (1854) 108, Z. 3339 (sphal- mate 3329).—S. brachiata MiqQ. (non Roxs.) FI. Dec. 1949) Ind. Bat. 1, 1 (1858) 1019; Koorb. Exk. Fl. 2 (1912) 192.—Salicornia australasica Mog. (ubi?) ex SCHINZ in E. & P. ed. 2, 16c (1934) 552. Rather robust perennial herb, woody beneath, divaricately branched from the base; branches as- cending, much divided; young shoots after removal of leaf sheaths very thin, wiry, afterwards growing thick and woody, not breaking up into joints. Leaf sheaths on young branches 7-10 mm long; their widened top with a thin, very minutely den- tate upper margin, gradually drying up, long per- sistent. Spikes erect or erecto-patent, cylindrical, very obtuse, 1!/2-4!/2 cm long, distinctly thicker than the branches by which they are borne; brac- teal cups 12-30, from the narrowed base to the upper margin 2!/2-3 mm long, at last separating CHENOPODIACEAE (Backer) 105 from the thin rachis on which they seem to be strung. Spikes apparently unisexual, but both sexes present on a single plant. Flowers free from each other; d perianth obtriangular, more or less deeply split; stamen 1, anther exsert; 9 perianth obliquely flagon-shaped, unilaterally split at the apex, 1'/2-2 mm long. Utricle compressed, indu- rate. Seed erect, orbicular; testa membranous. Distr. Coasts of tropical Africa, Hindustan, Bengal, in Malaysia: Java, Madura, Kangean Ar- chipelago, Sumbawa, Sumba, Timor. Ecol. Near the sea on salt clayey soils, which may be covered by a thin layer of sand, often more or less gregarious. F/. Jan.—Dec. Vern. Kemalahala (Sumba). 6. SUAEDA Forsk. Fl. Aeg.—Arab. (1775) 69, 18. Annual, biennial or perennial herbs, or shrubs, erect, ascending or prostrate; stems not conspicuously articulate. Leaves alternate, sessile, narrowly linear, terete or semiterete, succulent. Flowers small, 9 or partly unisexual, sessile, clustered or higher ones solitary; clusters in the axil of a leaf or a bract, 2-~-flowered, often united in paniculate inflorescences. Flowers 2—3-bracteolate; perianth herbaceous, deeply 5-cleft; segments subequal or 2—3 outer ones swollen on the back into tubercle; rarely all of them winged. Stamens 5, on base of perianth. Ovary free or at the base adnate to the base of the perianth, globose, ovoid orflask-shaped; styles 2-5, subulate or filiform. Fruit enclosed by the enlarged, more or less succulent perianth; pericarp membranous or spongy. Seed horizontal, oblique or erect, smooth; embryo planospiral, often green. Distr. Few spp., all over the world. 1. Suaeda maritima (LINNE) Dum. FI. Belg. (1827) 22; Bru. Fl. Austr. 5 (1870) 206; Hook. f. Fl. Br. Ind. 5 (1886), 14; BAmEY, Queensl. Fl. 4 (1901) 1263; Koorp. Exk. FI. 2 (1912) 192; Ascu. & Gr. Syn. 5, 1 (1913) 240; Hear, Ill. Fl. Mitt. Eur 3 (1912) 257; Domin, Beitr. Pflanzengeogr. Austr. (1929) 626; HEYNE, Nutt. Pl. (1927) 604; BACK. Onkr. Suiker. (1930) 214; Atlas t. 225; OCHSE & BAKH. V. D. BR. Veget. (1931) 105 cum icone.—Che- nopodium maritimum LANNE, Sp.Pl. (1753) 221.— Salsola indica WILLD. Sp. Plant. I (1797) 1317; BL. Bijdr. (1825) 536.—Chenopodium australe R.BR. Prodr. (1810) 407.—Suaeda indica Moa. in Ann. Sc. Nat. 23 (1831) 316.—S. australis Moa. in Ann. Sc. Nat. 23 (1831) 318; Courcu. in Lec. FI. Gén. I.C. 5 (1910) 8.—Chenopodina maritima Moa. in DC. Prodr. 13, 2 (1849) 161.—Ch. australis Moa. in DC. Prodr. 13, 2 (1849) 163.—Suaeda nudiflora Moa. in DC. Prodr. 13, 2 (1849) 155; VAL. in Bull. Dept. Agr. 10 (1907) 9; Koorp. Exk. fl. 2 (1912) B -192. Perennial glabrous herb, 7-45 cm long, at an ad- vanced age often woody at the base, usually divid- ed from the base into obliquely erect or ascending branches, often rooting from the lower joints; old stems tuberculate by scars of fallen leaves. Leaves rather crowded, linear, semiterete, with well-devel- oped aquiferous tissue, glaucous or more or less tinged with purple or entirely purple, 1—4!/2 cm long, with a saltish taste. Inflorescence usually pan- iculately branched, 2!/2-15 cm; flowers in clusters of 2-5 or highest solitary, all 6; lower bracts rather large, foliaceous; higher ones gradually smaller; bracteoles at the base of perianth 2-3, oblong ob- tuse, transparent, 2/3-1 mm long, persistent, after fall of the fruit + stellately spreading. Perianth green of more or less suffused with purple; seg- ments at first (Q stage) conniving, leaving only at top a small orifice for the protrusion of the styles, afterwards (d stage) spreading, after anthesis once more connivent and enclosing the fruit, ovate, ob- tuse, with + transparent margins and top; perianth in the Q stage + 11/4 mm diam., in the d stage + 2!/2mm; anthers broad, bilobed at the base, + 1/2 mm long. Ovary free from perianth, ovoid-con- ical; styles 2, rarely 3, rather long. Fruiting perianth depressed, without either tubercles or wings, in the living plant 2—2!/2 mm diam., succulent. Seed usu- ally horizontal, rarely vertical, shining brown; al- bumen very scanty. Distr. Europe, N. Africa, Asia, Australia, N. America, in Malaysia: West and East Java, Ma- dura, New Guinea (?). Ecol. Moist or swampy, clayey, saltish soils near the sea, often gregarious, frequently very conspi- cuous by its purple colour. Red and green speci- mens often grow intermixed. Vern. Alur, J. 106 FLORA MALESIANA [ser. I vol. 47, Dec. 1949] Use. Leaves sometimes eaten by the Indonesians as a vegetable. Notes. Very polymorphous species. The two specimens recorded as Suaeda nudiflora by Koorpers /.c. for the Karimon Djawa Isl. be- long both to Salsola kali. ZOLLINGER 2909 from Banjuwangi (E. Java) is Suaeda maritima; Zo.- LINGER (Syst. Verz. (1854) 108) could not place it. 7. SALSOLA LINNE, Sp.Pl. (1753) 222. Herbs or shrubs, glabrous or hairy; stems not conspicuously jointed. Leaves alternate or lowest opposite, sessile, linear or triangular, often spine-tipped. Flowers axillary, solitary or glomerate, sessile, 9, bibracteolate at the base; bracteoles ex- ceeding perianth. Perianth 5-partite down to the base; segments ovate-oblong, in or below the middle with a transverse thickening, above the thickening scarious, after anthesis completely embracing the fruit; the thickening after anthesis often excrescent into a horizontal wing; basal part of perianth unchanged or slightly indurate. Stamens 5, inserted on annular disk; filaments linear or subulate; anthers short or long; connective either produced into a point or not. Ovary globose-ovoid; style short or long, split into 2 long arms. Fruit falling off together with perianth; pericarp membranous or fleshy. Seed usually horizontal, exalbuminous; embryo coiled in a conical spire, often green. Distr. Species + 100, in Europe, Africa and Asia; a few introduced species in America and Australia, in Malaysia only the ubiquist S. kali L. 1. Salsola kali LINNE, Sp.PI. (1753) 222; Moa. in DC. Prodr. 13, 2 (1849) 187; Bru. Fl. Austr. 5 (1870) 207; Hook. f. Fl. Br. Ind. 5 (1886) 17; Bat- LEY, Queensl. Fl. 4 (1901) 1264; PULLE in Nova Guin. 8 (1910) 349; Koorp. Exk. Fl. 2 (1912) 192; AscH. & Grp., Syn. 5, 1 (1913, 1914) 207; HeEG1, Ill. Fl. Mitt. Eur. 3 (1912) 258; BAck. Trop. Natuur 11 (1922) 135, tab. 2, 3.—Salsola tragus LINNE, Sp. Pl. Ed. 2 (1763) 322; DcnE in Nouv. Ann. Mus. 3 (1834) 370; SPAN. in Linnaea 15 (1841) 345.—S. australis R.Br. Prodr. (1810) 411; Moa. in DC. Prodr. 13, 2 (1849) 188; Mig. Fl. Ind. Bat. 1, 1 (1858) 1022; Fors. Nat. Wand. (1885) 515.—S. brachypteris Moa. in DC. Prodr. 13, 2 (1849) 189; ZOLL. Syst. Verz. (1854) 168 (specimen not seen); Mia. Fl. Ind. Bat. 1, 1 (1858) 1022; Fors. Nat. Wand. (1885) 515. Psammophilous, glabrous or slightly pubescent annual, usually branched from the base and form- ing dense tufts; stems erect, or prostrate beneath, often woody at the base, 30-60 cm long. Lower leaves of young Malaysian specimens narrowly linear, rather flaccid, 2—71/2 cm long; higher leaves gradually shorter, proportionally broader; highest tapering from a broad amplexicaulous membra- nous-margined base, channelled, rigid, recurved, 1/2-1 cm long; all leaves spine-tipped, fleshy, + glaucous. Flowers solitary in axil of floral leaves, remote or in small numbers densely crowded on short axillary branchlets and then seemingly in axillary fascicles, which, when fruits are ripe, fall off as a whole; bracteoles recurved, rigid, tapering from a broad base, channelled, pungent, 4-5 mm long; perianth-segments ovate-oblong acute, 3- 31/2 mm. Filaments linear; anthers short, bilobed at the base, ++ 1!/2 mm; connective not or hardly produced. Style arms far exserted from perianth; base of fruiting perianth campanulate, pergame- neous, -+ 2 mm high, closely embracing the fruit; wings varying from very short to well-developed, often pink; 3 of them usually much broader than the rest, often emarginate; tips of the perianth- segments conniving above the fruit into a cone, thinly scarious. Fruits shortly obconical, truncate, tipped by the style-base, + 11/2 mm diam., at last circumsciss; apical part falling away. Seed hori- zontal, subglobose, shining black. Distr. Europe, N. Africa, Asia, Australia, N. America, in Malaysia: Java (said to have been col- lected in 1828 by BELANGER; I saw no Javan speci- mens), Karimon Djawa Arch., Madura, Bali, Kan- gean Arch., Celebes, Saleier, Buton, Timor, Key Isl., New-Guinea. Ecol. Sandy sea-shores, very local. Flowers in dry regions in the rainy season; disappears in the latter half of the dry monsoon. Vern. Landep, J, sundepan, J. Note. Very polymorphous plant. TEYSMANN (in Nat. Tijd. Ned. Ind. 11 (1856) 202) records Salsola for the region between Sura- baya and Tuban, but in a manner which sug- gests that he was mistaken. He says: “The road crosses coastal swamps in which grow Rhizophora, Avicennia, Salsola, Trianthema and other marsh- plants.’ Asa matter of fact, Salsola never grows in marshes; TEYSMANN, whose knowledge of plants was ample but superficial, may have confounded it with Suaeda. PLUMBAGINACEAE (C. G. G. J. van Steenis, Buitenzorg) Boiss. in DC. Prod. 12 (1848) 617; Mig. Fl. Ind. Bat. 2 (1859) 993; B. & H. Gen. Pl. 2 (1876) 623; CLARKE, in HOOK. f. Fl. Br. Ind. 3 (1882) 478; BENTH. Fl. Austr. 4 (1869) 265; Pax, in E. & P. 4, 1 (1889) 116; Boer. Handl. 2, 1 (1891) 274; Batt. Hist. Pl. 11 (1892) 359; Batt. Queensl. Fl. (1900) 943; GamBLe, J. As. Soc. Beng. 74, II (1906) 84; Rip. Fl. Mal. Pen. 2 (1923) 224; SpraGueE, J. Bot. 62 (1924) 267; Gams, in Hear, Ill. Fl. Mitt. Eur. 5, 3 (1926) 1877; Hutcu. Fam. FI.Pl. 1 (1926) 290; BAcKER, Onkruidfl. Jav. Suik. (1931) 477. Herbs or undershrubs. Leaves simple. Stipules absent. Flowers bisexual, actino- morphic, often in unilateral inflorescences, or subumbellate. Bracts often sheath- ing, dry and membranous. Bracteoles 2. Calyx tubular, gamosepalous, often con- spicuously ribbed, folded, the membranous folds often hyaline, lobes 5, often scari- ous. Petals free, but mostly connate at the base, contorted. Disk 0. Stamens 5, epipetalous, and connate with their base. Anthers 2-celled, opening lengthwise. Ovary superior, mostly sessile, often angled, i-celled with 1 ovule pendulous from a basal funicle; styles 5, free or variously connate; stigma subcapitate. Capsule membranous, mostly included, circumscissile near the thin base, rarely valvate from the base upwards. Seed 1, with or without endosperm, cylindric. Distr. Throughout the world, ca 10 genera. Ecol. Mostly in salty steppes, or littoral. Leaves variously reduced to needle-shape, often provided with glands secreting water, mucus, salts or CaCOs. Uses. Several species are medicinal; see Plumbago. KEY TO THE GENERA 1. Leaves scattered or alternate on the stems, not rosulate. 3. Petiole alate throughout, clasping the branch. Blade orbicular . . . . . . . 1. Aegialites peecwole sot so. Blade not:orbicular 2. =: = = = <= « =» « « « « +» = «ds Pimubago ae Scapose herbs. Leaves rosulate. Rootstock. . Flowers in heads. Scape unbranched. Leaves 1-nerved. . . «a0 =) Ss -Annema 2 Flowers in wide-branched corymbs, unilateral, spicate. Leav es large, pinnatisect- -spathulate 4. Limonium 1. AEGIALITES R.Br. Prod. (1810) 426 (not of TRIN. 1820); Roxs. Fl. Ind. 2 (1832) 111 (Aegelatis); Boiss. /.c. 621; Grirr. Not. 4 (1854) 207 (Aegiatilis); Mia. I.c. 994; B. & H. Le. 624; Kurz, J. As. Soc. Beng. 46, II (1877) 217; For. Fl. Burma 2 (1877) 96; CLARKE, /.c. (Aegianilites); BOERL. I.c. 278; PAx /I.c.; BAILL. /.c.; GAMBLE L.c.— Aegialinites PRESL, Bot. Bem. (1844) 103. Simple-stemmed shrub, or undershrubs, '/3-3 m tall, branches with pith. Leaves alternate, orbicular, glabrous, coriaceous, entire, smooth, gland-dotted; nerves and veins parallel; petiole long, thick, winged, clasping the stem, leaving annular scars, glandular inside. Flowers solitary in the axil of a bract, erect, in leafy panicled “often fork-branched racemes not much exceeding the leaves. Bracts sheathing, en- closing 2 similar but smaller boat-shaped bracteoles, all persistent, glandular in- side. Calyx persistent, wholly or only upwards with 5 prominent ribs, folded be- tween, tubular, lobes short. Corolla white, longer than the calyx, easily detached circumscissile at the base, lobes elongate-spathulate, subconcave 3-nerved, mutu- ally connate at the base in a short tube together with the stamens; tube annular or barrel-shaped, after anthesis when pushed out splitting upwards from the base. 108 FLORA MALESIANA [ser. I, vol. 42 Stamens inserted on the apex of the tube, anthers basifix c. as long as the corolla, sagittate at the base; cells + parallel, halfway free, latrorse; connective a narrow furrow between the cells; pollen 90-120 diam. Styles free, articulate at the base; stigma small, punctiform-capitate, reaching just above the stamens. Capsule linear, long-exserted, pentagonal, dehiscing finally along the angles. Distr. 2 spp., SE. Asia to Australia, absent in many districts of Malaysia (fig. 1). Ecol. In the open mangrove, according to TEYSMANN and Brass also on rocky beaches and exposed shorelines, on sandy soil (WARBURG), locally in small pure groves. The stembase in Ae. rotundifolia is said to be swollen and conical (GRIFFITH, CRAIB, BRANDIS). Aerophores are not found (WARBURG). GRIFFITH says that the petioles, bracts, and bracteoles exude a viscose fluid. This is due to muci- laginous glands which are widely distributed in the family (cf. Witson, Ann. Bot. 4 (1890) 244). In the Ceram specimens I found sand adhering to calyx and corolla. The exocarp is coriaceous; the mesocarp is swollen and spongy and contains air in the herbarium; the endocarp is thin. The mesocarp therefore seems to contribute the means by which the fruit can drift. The elongated fruit reminds of viviparous Rhizophoraceae, Aegiceras, &c. The flowers seem not to be ephemeral; therefore measurements of the ovary (proportion length/diameter) are not constant, as it enlarges during anthesis. The corolla is after anthesis circumscissile at the base and is slowly pushed out, and its basal obconical part enclosing the ovary is partly slit upwards. Field study is urgently needed to clear the life-history and variability of this peculiar genus. Notes. The genus consists of two widely separated populations which show rather unimportant dif- ferences in the flower. As no intermediates are known and the ecological behaviour is different I accept them provisionally as two distinct species. The disjunction is similar to that in Aldrovanda, Philydrum lanuginosum, &c; the absence in the intervening area is unexplained. KEY TO THE SPECIES 1. Leaves shining above, reticulations prominent s.s. Flowering parts larger than in next species, longer pedicelled. Calyx 13 mm, lobes hardly imbricating, acute-triangular. Corolla tube (3!/2-)5 mm high, lobes 12 by 3-4 mm below the apex. Anthers 3 mm; filament inserted in the middle. Ovary 5-10 by 2!/2-5 mm, little acuminate towards the apex ae 2. Ae. rotundifolia 1. Leaves dull above, reticulations indistinct s.s. Flowering parts smaller than i in the prec. species. Flower sessile or up to 5 mm pedicelled. Calyx 7-8 mm, lobes distinctly imbricating, their apex rounded to subtruncate, mucronulate. Corolla tube 1-2 mm high, lobes 9-10 by 11/2-2 mm. Anthers 2 mm long, filament inserted at 1/3 of the length. Ovary 3—4 by 1!/2-2 mm, oval . 1. Ae. annulata Stembase swollen, conical. Leaves rotundate to oval, 4-8 by 2!/2-7.cm. Petiole 3!/2-8 cm long. Bracts 7-9 by 6 mm, bracteoles 6-8 by 2 mm. Flowers sessile or up to 5 mm pedicelled. Stamens 7-8 mm long. Styles 6-7 mm. Fruit 2 mm diam., first straight, gradually elongating, later falcate up to 5 cm long, but not thickening. Distr. Queensland, NW. Australia, Thursday Isl. and E. Malaysia: S. New Guinea, Moluccas (Aru, Ceram) and Lesser Sunda Isl. (Timor, Se- mau, Alor, Solor, Adonara, E. Flores (Laran- tuka)).—Fig. 1. Ecol. Brass’s Papuan specimens were only 30-45 cm tall. Grows in isolated specimens or small colonies in more sandy and rocky places than the following species. Fig. 1. Distribution of Aegialites. 1. Aegialites annulata R.Br. Prod. 1 (1810) 426; GAUD. in Freyc. Voy. (1826) t. 51; Borss. in DC Prod. 12 (1848) 621; Mig. FI. Ind. Bat. 2 (1859) 995; BENTH. FI. Austr. 4 (1869) 266; TEysM. Nat. Tijd. N.I. 34 (1874) 456; Hemsv. Rep. Chall. Bot. pt 3 (1884) 161; Wars. Bot. Jahrb. 13 (1891) 400; BAILL. Hist. Pl. 11 (1892) 359; Batt. Queensl. FI. (1900) 943; PuLLE, Nova Guinea 8 (1910) 397; BariL. Compr. Cat. Q. Pl. (1913) pl. 298; Wuite, J. Arn. Arb. 10 (1929) 258.—Aegianilites annulata Pres_, Bot. Bem. (1844) 103. 2. Aegialites rotundifolia Roxs. FJ. Ind. 2 (1832) 111 (Aegelatis); GrirF. Not. 4 (1854) 207, t. 461, f. 2; PRAIN, Beng. Pl. 1 (1903) 638; CLARKE in Hook. f. Fl. Br. Ind. 3 (1882) 479; BRANDis, Ind. Trees (1906) 413; GAmBLeE, J. As. Soc. Beng. 74, II (1906) 85; Rip. Fl. Mal. Pen. 2 (1923) 225; Cralis, Fl. Siam 2 (1938) 320.—Aegianilites rotun- difolia PREsL, Bot. Bem. (1844) 103; Grirr. Ic. pl. As. (1854) t. 654.—Ae. annulata var. rotundifolia Boiss. in DC. Prod. 12 (1848) 621; Kurz, J. As. Soc. Beng. 46, II (1877) 217; For. Fl. Burma 2 (1877) 96. Leaves orbiculat, base mostly rounded or broad- Dec. 1949] PLUMBAGINACEAE (Vv. Steenis) 109 cuneate to slightly cordate, apex truncate or slight- ly obtuse, protruding, 4!/2-7 cm diam. Petiole 5— 7'/2 cm long. Bract 10-11 mm. Bracteoles 8—9 mm. Flowers pedicelled up to 13 mm. Stamens 13-— 18 mm. Styles 9-10 mm. Ripe fruit not described. Distr. Bengal, Burma, Siam, Mergui, Tenasse- rim, Andaman Isl., according to RIDLEY errone- ously recorded for the Malay Peninsula, but likely to occur there somewhere orin N.Sumatra.—Fig.1. Ecol. Low muddy mangrove. 2. PLUMBAGO LINNE, Sp.PI. 1 (1753) 151; cf. lit. under family; STEEN. Trop. Natuur 26 (1937) 12. Perennial herbs or undershrubs, rarely annual, often straggling or subscandent. Leaves spread, entire, older ones often below pale-lepidote by excreted carbonates, or reduced on the flowering stems; petiole often semi-amplexicaulous-auriculate at the base. Flowers in terminal racemes or spikes, often united in a leafy panicle, blue, rosa, white or violet, ephemeral, not caducous. Calyx tubular, outside often with sessile or stalked glands, teeth erect, not enlarged in fruit. Corolla funnel- shaped, lobes spreading. Stamens free, broadened at the base. Style short, with 5 branches. Capsule included in the persistent calyx and (often twisted) corolla; pericarp thin, hardened above, circumscissile near the base, caducous part often splitting towards the apex with 5 valves. Distr. About 10 spp. in all tropics, often used as ornamentals. Ecol. Not limited to saline localities, but preferably under semi-arid conditions. Of P. indica no fruit has ever been found, and P. aphylla and P. auriculata never produce fruit in Java; they are propagated vegetatively. The leaves of the latter are often covered below with greyish scales of excreted CaCOs. Uses. Plumbago, leadwort, derives its name from the colour adapted by the skin after the medicinal use of the European P/. europaea, a plant used in historic time for curing p/umbum, an illness of the eyes. The Indian species too are often used medicinally. The active substance, plumbagin, is known as a nar- cotic and specially as a vesicatory and anthelmintic. It is specially extracted from the roots; radix vesica- toria was already figured by Rumpuius, Herb. Amb. 5, p. 453, t. 168 (P. indica). Several spp. are known as ornamentals, mostly P. auriculata. In a strict sense none is native in Malaysia, though P. zeylanica might be accepted as such. KEY TO THE SPECIES 1. Corolla white, tube less than 2!/2 cm long, limb 12-15 mm diam. Rachis of the raceme or spike with sessile or stalked glands. 2. Leaves well-developed on flowering stems, 3—12!/2 by 2- 5 cm. Rachis with sessile glands 1. Pl. zeylanica . Leaves mostly only present on young stems, !/2—-1'/2 cm long, absent on flowering stems. Rachis with stalked glands 2. Pl. aphylla 1. Corolla red or blue, tube 21f-4¢ cm fone! limb 2 2 aie cm diam. Rachis hairy or ‘glabrous but without glands. 3. Corolla light blue, lobes without a distinct mucro. Sats greenish-yellow, 10-14 mm long, lower 1/32/3 part without glands, finely but short-hairy . 3. Pl. auriculata 3. Corolla red, lobes distinctly mucronate. Calyx red, 8-9 mm long, glandular all over, glabrous 1. Plumbago zeylanica LINNF, Sp.PI. (1753) 151, cf. lit. under family; BENTH. Fl. Austr. 4 (1869) 267; MerR. FI. Man. (1912) 362; Sp. Blanc. (1918) 300; En. Philip. 3 (1923) 275; HEYNE, Nutt. Pl. (1927) 1222; MerRR. Comm. Lour. (1935) 301; Backer, Bekn. FI. Jav. 8 (1949) fam 181, p. 2.— PI. auriculata (non LAMK) BI. Bijdr. 14 (1826) 736. —P. viscosa BLANCO, FI. Filip. (1837) 78; ed. 2 (1845) 58; ed. 3, 1 (1877) 111. Straggling shrub. Twigs long, not rooting, 1!/2- 21/2 m. Young lJeaves with caducous small auricles. Racemes 6-30 cm, glands green often red-tipped. Calyx glabrous, glandular all over, green. Corolla tube 18-22 mm, lobes obovate 6-7 mm. Anthers blue-purple. Ovary and style glabrous. Fruit ob- long, acute with 5 furrows, calyx patent and recurved. Distr. Tropics of the Old World to Hawaii, 4. Pl. indica possibly only indigenous in SE. Asia, in Malaysia: not yet found in Borneo and the Moluccas, also cultivated. Ecol. Both under feeble and strong dry-season conditions, preferably in the latter, always in an- thropogenic localities, savannahs, thickets, &c, up to 1000 m, f7. Jan.—Dec. Vern. Bama, bantji, poksor, godong eéntjok, J, karéka, Md, ki éntjok, Sd, daun éntjok, M, bama (Bali), oporio (Timor), jarak, cheraka (Mal. Pen.); Philippines: bangbang, Ilk., sangdikit, Tag., talankan, Ik. 2. Plumbago aphylla Boser ex Boiss. in DC. Prod. 12 (1848) 694; Pax in E. & P. 4, 1 (1889) 117; STEEN. Trop. Natuur 26 (1937) 13; BACKER, Bekn. Fl. Jav. 8 (1949) fam. 181, p. 2. Strong taproot producing few-branched, rather 110 FLORA MALESIANA [ser. I, vol. 42 Fig. 2. Plumbago indica L., x 1/2. (Courtesy Pasuruan Exp. Station) Dec. 1949] PLUMBAGINACEAE (Vv. Steenis) straight, rod-shaped stems '/2-1 m long, often root- ing and producing new plants. Leaves without auricles. Racemes 2-6cm long. Calyx 7-8 mm high, hairy between the glands. Corolla-tube 14- 16 mm, much broader than in the prec. sp., lobes emarginate. Anthers green-yellow. Style base long- hairy. Distr. Indigenous in Madagascar, in Java some- times in gardens but not very recommendable as an ornamental; no fruit is produced in Malaysia. Fl. Jan.—Dec. 3. Plumbago auriculata LAMK, Encycl. 2 (1876) 270; Merr. FI. Man. (1912) 361; En. Philip. 3 (1923) 275, non BL. (1825).—P. capensis THUNB. Prod. Fl. Cap. (1794) 33; DC. Prod. 12 (1848) 693; NAVES in BLAnco, FI. Filip. ed. 3 (1877) 83, t. 27; F.-VILL. Noy. App. (1880) 122; STEEN. Trop. Na- tuur 26 (1937) 13; BACKER, Bekn. FI. Jav. 8 (1949) fam. 181, p. 2. Erect shrub or halfshrub, !/3-1!/2 m. Leaves ob- long to obovate, 11/2-5 by 3/42 cm, auricles mostly large. Upper axils witha bundle of leaves. Racemes mostly combined to leafy corymbs; rachis 1— 6 cm long, densely puberulous. Calyx green, 10— 14 mm long, lower '/3—/3 puberulous but without glands. Ovary pear-shaped, style base glabrous. Distr. Native of S. Africa, in Malaysia: a com- mon ornamental, up to 1100 m, does not run wild. Fl. Jan.—Dec. Vern. Mannentrouw, (Dutch). verliefde luitenantjes 4. Plumbago indica LINNE in Stickm. Herb. Amb. (1754) 24; Amoen. Acad. 4 (1759) 133; Horsr. Verh. Bat. Gen. 8 (1816) 108; Merr. Int. Rumph. (1917) 414; En. Philip. 3 (1923) 275; HeyNe, Nutt. Pl. (1927) 1221; Backer, Onkruidfi. Suik. (1931) 478; STEEN. Trop. Natuur 26 (1937) 13; BACKER, Bekn. Fl. Jav. 8 (1949) fam. 181, p. 3.—Radix vesicatoria RUMPH. Herb. Amb. 4 (1750) 453, t. 168 (type).—PI. rosea LINNE, Sp.Pl. ed. 2 (1762) 215; BuRKILL, Dict. (1935) 1774; Mig. Fl. Ind. Bat. 2 (1859) 993; CLARKE in Hook. f. Fl. Br. Ind. 3 (1882) 481; GresH. Schets. 210; MerrR. Fl. Man. (1912) 362; PELLEGRIN, FI. Gén. I.C. 3 (1930) 752; Ripi. Fl. Mal. Pen. 2 (1923) 225.—PIl. coccinea (Lour.) SALiIss. Prod. (1796) 122.—Fig. 2. Branched from the base, stems flaccid, some- times rooting, '/2-1!/2 m. Leaves oblong, 5-15 by 2-8 cm, without fascicled leaves in the axils, petiole not auriculate. Ca/yx red, 8-9 mm high, glabrous, glandular all over. Racemes not corymbiform, rather sparse. Rachis glabrous, 10-30 cm long. Ovary ovate-oblong. Style base short-hairy. Fruit unknown. Distr. Widely distributed in the Old World tropics, probably native in SE. Asia, certainly not a sport or variety of P/. zeylanica, in Malaysia: not yet recorded from the Malay Peninsula, Borneo, and New Guinea, ascending to 1000 m, also cul- tivated. Ecol. Always in anthropogenic localities, lo- cally run wild or semi-spontaneous, often persist- ent in abandoned cultivations, also in teak-forests, fi. Jan.—Dec. Uses. Ornamental, sometimes medicinal as a substitute for Rauwolfia serpentina BTH. Vern. Poelé pandak (lalaki), tjeraka mérah, akar binasa, M, bama, J, daun srunén, Md, vuurwortel, Dutch, akar binasa, Mol., setaka, Mol., mehula- toe, Amb., auwarian, Banda, mehutana hane, Mol., nehulatu, Mol.: Philippines: /aurel, Bik., Sp., Tag., panting panting, Mag., sutungau, Tagb., ulanda, Sul. 3. ARMERIA WILLD. En. Hort. Berol. 1 (1809) 333, nom.cons.—Statice L. p.p. Perennial tufted scapose herbs with narrow rosulate /eaves and a subterraneous, branched rootstock. Flowers in heads subtended by dry bracts, bases of the outer ones coalescent into a tubular sheath. Calyx obconical, 5-lobed, lobes mucronate. Petals united at the base. Stamens inserted on top of the tube. Style + free, stigma cylindric. Fruit circumscissile at the base. Distr. Spp. 10 or 60 depending on the specific concept, cosmopolitan, not native in Australia and Malaysia. 1. Armeria maritima (MILL.) WILLD. /.c.—Statice armeria LINNE, Sp.Pl. (1753) 274.—Statice mari- _ tima Mi. Gard. Dict. ed. 8 (1768) no 3.—Armeria vulgaris WILLD. l.c.—Statice armeria var. maritima (MILL.) Gams, in Hear, Ill. Fl. Mitt. Eur. 5, 3 (1926) 1888; Backer, Bekn. Fl. Jay. 8 (1949) fam. 181, p. 3. Leaves 1-nerved, slightly puberulous on margins and midrib, 5-7 by 1 mm. Culms 10-20 cm, thin and hard, puberulous, apex hollow. Heads 2- 23/4 cm diam., sheath 10-18 mm. Calyx 6 mm high. Petals cuneate-spathulate, emarginate, rosa to red, their base pale, rarely white, 8-10 mm long. Style bases hairy. Distr. Native of Europe, sometimes cultivated in the mountains of Java (acc. to BACKER), fil. Aug. 112 FLORA MALESIANA [ser. I, vol. 47, Dec. 1949] 4. LIMONIUM MILL. Gard. Dict. Abridg. ed. 4 (1754); ADANs. Fam. 2 (1763) 283; DRUCE, Rep. Bot. Exch. Club Br. Isl. (1913) I, 433.—Statice L. em. WILLD. non MILL. Stiff, erect herbs. Leaves rosulate, spathulate, entire or lobed, often large. Flowers in widely branched corymbs, consisting of unilateral spikes. Bracteoles unequal. Calyx tube narrow, 5-ribbed; limb scarious, coloured, teeth often mucronate. Pet- als mostly + free, at the base connate with the stamens. Styles + free, stigma cylindric, rarely (§ Goniolimon) capitate. Fruit indehiscent or capsular at the apex. Distr. More than 100 spp., cosmopolitan, not native in Malaysia. This absence is surprising as the genus occurs both in Australia and Asia. This distributional discontinuity is found in several plants, e.g. Philydrum lanuginosum, Rothia trifoliata, Samolus valerandi, &c. Some of the plants of this type have occasionally been found in Malaysia, e.g. Eriocaulon setaceum, Hydrocotyle peltata, Aldrovanda vesicu- losa, Montia, Anagallis pumila, but remain exceedingly scarce. 1. Limonium sinuatum (L.) MIL. Gard. Dict. ed. 8 (1768) no 6; Gams in Hea, Ill. Fl. Mitt. Eur. 5, 3 (1926) 1881; BACKER, Bekn. Fl. Jav. 8 (1949) fam. 181, p. 4.—Statice sinuata LINNE, Sp.Pl. (1753) 276; CurT. Bot. Mag. t. 71; Borss. in DC. Prod. 12 (1848) 635. Robust, !/2-1_m. Leaves lanceolate-spathulate, deeply pinnatisect, 20-50 by 2!/2-6 cm (incl. the long petiole), sparsely long-hairy. Peduncle and branches distinctly winged, wings + crisped, on each fork with 3 erect, leafy, linear, 1-12 mm long appendages. Unilateral spikes dense, stalk between the bracts 3-alate, one wing narrow, the others broadening. Large bracteole folded, short-3-lobed, apex on the back with 2-3 teeth, narrow bracteole needle-tipped. Calyx tube 6 mm long, limb entire, fine-plaited, blue, white or rosa, 6-7 mm long. Distr. Native of the Mediterranean, sometimes ornamental in the mountains of Java (acc. to BACKER), ff. July, Nov. Excluded Limonium billardieri (GIRARD) O.K. Rev. Gen. (1891) 394.—Stratice billardieri GiRARD, Ann. Sc. Nat. III, 2 (1844) 325; Boiss. in DC. Prodr. 12 (1848) 658; Mra. FI. Ind. Bat. 2 (1859) 995; HatL- LIER f. in ELBERT, Sunda Exp. 2 (1912) 294; Koorb. Exk. Fl. Java 3 (1912) 36 (sic). This species was based on a specimen said to have been collected by LA BILLARDIERE in Buru Is- land (Moluccas) on the expedition of ‘La Recher- che et l’Espérance’ which went along the Cape Verde’s to Cape of Good Hope, Australia and Me- anesia to the Moluccas, and stayed in Buru from Sept. 18-30 at Kajeli, the principal harbour. A co- type kindly sent by Dr BAEHNI agrees with Bur- CHELL 512 at Leyden and is identified as L. equiseti- num (Bolss.) DYER var. depauperatum (Botss.) from the Cape. There are one developed flower and one reduced flower in each spikelet. It is closely allied to L. scabrum (THUNB.) O.K. The occurrence of minute tufts of hairs on the scape is typical. Cf. also WRIGHT, in FI. Cap. 4, 1 (1909) 422. This is the 2nd instance in which a species of this genus was mislabelled (cf. Kew Bull. 1948, 368). UMBELLIFERAE (* P. Buwalda, Groningen) Annual or perennial herbs, never woody shrubs (in Malaysia). Stems often fur- rowed and with soft pith. Leaves alternate along the stems, often also in rosettes; petiole usually with a sheath, sometimes with stipules at the base; lamina usually much divided, sometimes entire. Flowers polygamous, in simple or compound um- bels, sometimes in heads, terminal or leaf-opposed, beneath with or without in- volucres and involucels. Calyx teeth 5, often obsolete. Petals 5, alternate with the calyx teeth, equal or outer ones of the inflorescence enlarged, entire or more or less divided, often with inflexed tips, inserted below the epigynous disk. Stamens alter- nate with the petals, similarly inserted. Disk 2-lobed, free from the styles or con- fluent with their thickened base, forming a stylopodium. Ovary inferior; styles 2. Fruits with 2 one-seeded mericarps, connected by a narrow or broad junction (commissure) in fruit separating, leaving sometimes a persistent axis (carpophore) either entire or splitting into 2 halves; mericarps with 5 longitudinal ribs, 1 dorsal rib at the back of the mericarp, 2 lateral ribs at the commissure; 2 intermediate ribs between the dorsal and the lateral ones; sometimes with secondary ribs be- tween the primary ones, these without fascicular bundles; often vittae in the ridges between the ribs or under the secondary ribs, and in the commissure, seldom under the primary ribs. Distr. Numerous genera and species, all over the world. The representatives native in Malaysia belong geographically to five types. (1) Ubiquitous genera (Hydrocotyle, Centella, Oenanthe); one species, Hydrocotyle vulgaris, shows a remarkable disjunction, occurring in Europe & N. Africa and also in New Guinea, Australia and the Marshall Islands. (2) Western elements are Sanicula (wide-spread in the N. hemisphere but absent from New Guinea and Australia), Heracleum and Pimpinella; though some spp. are endemic their close relatives are found in SE. Asia. (3) A distinctly N. element is the Japano-For- mosan Peucedanum japonicum in the islands N. of Luzon. (4) A distinct Australian element is Trachymene which centers in Australia and occurs also in New Caledonia and Fiji; this genus shows a relatively rich secondary centre in East Malaysia; another Australian alliance is found in ubiquitous Eryngium of which the only native Malaysian species hitherto known is allied to Australian spp. (5) A distinct Subantarctic- distributed genus is Oreomyrrhis which centers in New Guinea by 4 spp.; one of these occurs from Kina- balu to Australia, New Zealand to Andine South America as far as Mexico; a marked instance of the ancient alpine-Papuan South Pacific plant refuge (v. ST.). Ecol. As to altitude both the cultivated and native spp. prefer microtherm localities: Pimpinella, Trachymene, Heracleum, Oreomyrrhis, Eryngium, and Hydrocotyle vulgaris are confined to the montane or subalpine zones, Sanicula descending to the colline subzone. Some Umbelliferae are found in the alpine zone above 4000 m alt., e.g. Hydrocotyle sibthorpioides, and Oreomyrrhis andicola. Several oreophytes show a remarkable reduction of the leaf surface, and some are reduced to cushions (Oreomyrrhis andicola) or true pin-cushions (Trachymene pulvilliforma and Oreomyrrhis azorellacea). There are only very few spp. preferring a shaded locality, e.g. Sanicula europaea, Hydrocotyle javanica, Trachymene erodioides, ? He- racleum sumatranum. Pimpinella javana and P. pruatjan are often found in Casuarina forests. Oenanthe javanica, Centella,and the Hydrocotyles decidedly prefer moist or marshy places. In the semi-arid regions of Malaysia Umbelliferae are exceedingly scarce; Hydrocotyle javanica is a typical indicator for everwet conditions. Many spp. show a remarkable adaptive capacity to wide altitudinal limits, e.g. Hydrocotyle sibthorpioides 14050 m, H. javanica 1-2900 m, Oenanthe javanica 1—2800 m, Centella asiatica 1—2500 m, Sanicula europaea 500-3060 m (v. ST.). Uses. Quite a number of Umbelliferae are cultivated for the essential oil contained in their fruit and are used as condiments. The leaves of some spp. are eaten as vegetables or for medicinal purposes. Under the several species data are mentioned drafted from HEYNE, De Nuttige Planten ed. 2 (1927), OCHsE & BAKHUIZEN VAN DEN BRINK, Indische Groenten (1931) and BuRKILL, A dictionary of the economic prod- ucts of the Malay Peninsula (1935). Notes. The extensive treatment in my former revision (Blumea 2, 1936) is mostly followed; several novelties are added. Of the genera which are represented only by introduced or cultivated spp. generic characters are not given. When collecting Umbelliferae it is to be observed that ripe fruits and basal leaves are essential for identification. - PS. The MS. of the present revision was made before the author received the rich New Guinean col- lections of L. J. Brass, M. S. CLEMENS, C. E. CARR; on the latter he prepared a separate paper to be 114 FLORA MALESIANA [ser. I, vol. 42 published in the Journal of the Arnold Arboretum which he finished just before his sudden and lamented death (cf. Bull. Bot. Gard. Btzg III, 17 (1948) 377). I am responsible for fitting the new data to the original MS. as well as for the description of Peucedanum (v. ST.). Figures 1 & 5 courtesy Pasuruan Exp. Station, fig. 2-4, 6 & 10 courtesy Blumea. KEY TO THE GENERA 1. Flowers in simple umbels! or heads often united in more compound infloresceaces, but not in com- pound umbels. 2s eaves and inyolucres pricklys Flowersanvheadswe, wean). 4) Se eee 5. Eryngium 2. Leaves and involucres not prickly. Flowers in umbels. 3. Fruits with uncinate bristles Sn oe toe ho noe > 4 4 Sanne 3. Fruits without uncinate bristles. 4. Fruit at least twice as long as broad; generally not laterally flattened . . . 9. Oreomyrrhis 4. Fruit not longer than broad, laterally flattened. 5. Mericarps 7—9-ribbed, with connecting veins between the ribs. Leaves simple, crenate, reniformous 2. Centella 5. Mericarps 3-ribbed. Leaves otherwise. 6. Leaves without sheaths but with distinct entire stipules. Corolla valvate . . 1. Hydrocotyle 6. Leaves with sheaths, with or without lacerate stipule-like appendages. Corolla imbricate 2. Trachymene 1. Flowers in compound umbels which are sometimes united in more compound inflorescences. 7. Mericarps winged on the margin. 8. Fruit not strongly dorsally flattened, more than twice as One as broad. Leaves tripinnate, ultimate segments nearly filiform : 2 2) eos Aneto: 8. Fruit strongly dorsally flattened, at most twice ; as long as broad. Leaves pinnate to bipinnate, extreme segments not filiform. 9. Leaflets or ultimate leaf-segments cuneate-obovate, only dentate or incised at the broadened apex. Mericarps about twice aslongas broad . . . . ee ieee 22. Peucedanum 9. Leaflets or ultimate leaf-segments not cuneate- obovate, margin serrate or crenate. Mericarps at most 11/2 times as long as broad. 10. Ovary hairy. Involucels 6-7. ae of the mericarps 2!/2 mm broad. Corolla white or reddish, radiating . . . 21. Heracleum 10. Ovary glabrous. Involucels 0-2, “Wing of the mericarps Glow mm ‘broad. Corolla yellow, not radiating . . eid Dt et gos, <2 8 Sa 20. Pastinaca 7. Mericarps not winged at the margin. 11. Fruit Jaterally flattened. Leaves simple, roundish . . . . . . . +. +. 41. Hydrocotyle 11. Fruit not laterally flattened. Leaves usually compound. 12. Fruit with a sterile neck or a short beak visible on the ovary as a dark green ribbed neck 6. Chaerefolium 12. Fruit without a sterile neck or beak. 13. Calyx teeth distinct. 14. Ovary and fruit bristly. 15. Fruit with uncinate bristles; stems and leaves hairy; leaf-segments not very narrow; flowers notradiating . . : 7. Torilis 15. Fruit with stellate hairs: stems ‘and leaves glabrous: extreme leaf-segments linear to filiform; flowers radiating . Se ee ee mre shee 31), (Comin 14. Ovary and fruit entirely glabrous. 16. Mericarps hollow at the ventral side; primary ribs visible as undulate lines, secondary ribs somewhat more prominent; flowers radiating . . . §. Coriandrum 16. Mericarps not hollow at the ventral side; marginal ribs thicker ‘than the lateral ones, secondary nls Goce TONERS mObimtmye = 5 5 4G 5 6 oF OG 6 o G ¢ 17. Oenanthe 13. Calyx teeth not distinct. 17. Ovary and fruit entirely glabrous. 18. Leaves ternate; umbels and umbellules few-rayed. Oo ed he Uae 14. Cryptotaenia 18. Leaves pinnate or bipinnate. 19. Flowers yellow or yellowish green. 20. Involucels many-leaved; leaves 3—4-pinnate with + filiform segments . 18. Foeniculum? 20. Involucels 0-2-leaved; lower leaves 3-pinnate with nearly obovate or cuneate leaflets 19. Petroselinum 19. Flowers white or reddish. (1) Rarely solitary (1-flowered umbels). (2) Foeniculum and Anethum are very alike with the exception of their fruits, which in Anethum are very distinctly winged, in Foeniculum not. They may further be distinguished, besides by their char- acteristic odor, by fine-puncticulate stems in Foeniculum which are absent in Anethum. Dec. 1949] UMBELLIFERAE (Buwalda) 115 . Ripe fruits 1!/2-2 mm through, roundish when seen from the lateral side; carpophore entire or very shortly bifid at the apex . - . 11. Apium 21. Ripe fruits 4-5 mm long and half as ; broad: carpophore bifid to ) nearly 2/3 of its length 15. Carum 17. Ovary and fruits bristly, hairy, or with scale-like trichomes. 22. Involucres pinnatifid . 22. Involucres not pinnatifid. 23. Leaves simple, or pinnate with simple leafiets; leaves and stems hairy 23. Leaves pinnate with divided leafiets; leaves and stems glabrous 23. Daucus 16. Pimpinella 13. Trachyspermum 1. HYDROCOTYLE LINNE, Sp.Pl. 1 (1753) 234; Buw. Blumea 2 (1936) 122 (Jit.). Perennial, stems prostrate or rooting at the nodes, sometimes suberect. Leaves petiolate and stipulate, in outline rhomboid, peltate or cordate, palminerved, en- tire, lobed or divided, crenate to crenate-serrate. Umbels simple, sometimes irreg- ularly subcompound. Involucral bracts few or 0. Calyx teeth minute or obsolete. Petals entire, valvate in bud. Disk plane, margin elevated. Styles from the base filiformous or with thickened base. Fruit laterally flattened, commissure narrow; vittae 0; mericarps with dorsal ribs, marginate, lateral ribs in the commissure, intermediate ones straight or arcuate. Distr. About 100 spp. all over the world, mainly in the S. hemisphere. KEY TO THE SPECIES 1. Leaves peltate 1. Leaves not peltate. 3. H. vulgaris 2. Leaves usually less than 3 cm in diam. Stems creeping, sometimes with ascending extremities. Inflores- cences single, sessile or short-peduncled, along the creeping stems and the ascending tips. Fruits up to 15 in each inflorescence, yellow to dark-brown when ripe 2. H. sibthorpioides 2. Leaves usually more than 3cm in diam. Stem creeping with ascending branches. Inflorescences single or in bundles, sessile to long-peduncled, usually along the ascending branches only. Fruits more than 15 in each inflorescence, blackish brown when ripe 1. Hydrocotyle javanica THUNB. Diss. Hydroc. (1798) p. 3 no 17, p. 6, t. 2; Buw. Blumea 2 (1936) 122 (lit. —H. hirta R.Br. ex Ric. Ann. Gén. Sc. Phys. 4 (1820) 64.—H. nepalensis Hoox. Exot. Fl. 1 (1823) t. 30.—H. sundaica BL. Bijdr. 15 (1826) 883.—H. globata BL. Bijdr. 15 (1826) 883.—H. zeylanica DC. Prod. 4 (1830) 67.—H. podantha MOLKENB. in Mio. PI. Jungh. (1851) 89.—H. rotun- difolia (non DC., 1830) Wars. Bot. Jahrb. 13 (1891) 397.—H. novo-guineensis WARB. Bot. Jahrb. 16 (1892) 24——H. versteegii HemMst. Kew Bull. (1909) 259. Stems rarely entirely erect, 10-50 cm, terete, glabrous or short-hairy. Stipules 3-8 by 4-6 mm broad-ovate, roundish to acute, membranaceous, entire or the apex fringed. Petioles 2-20cm, short- hairy; /Jamina usually 3-8 cm through, rarely only 11/2 cm, roundish to 5—8-angular in outline, cor- date, 5—8-lobate, lobes crenate to crenate-serrate, _ more or less triangular, glabrous, rarely sparsely hairy. Inflorescences single or in groups, opposite to the leaves, sometimes united to an umbel with an involucre of few small bracts, sometimes also terminal. Peduncles 1—7 cm, rarely absent, glabrous or short-hairy. Involucres many around and be- ween the flowers, 1 by 3/4 mm, ovate-acute, en- ire or base with small teeth, outer ones refiexed in tuit. Pedicels 15-50, 0—'/2 mm, rarely longer. Pe- als 1 by 1/2 mm, lanceolate, acute. Mericarps 1— 1. H. javanica 11/4 by nearly 3/4 mm, glabrous or short hirsute or even with short curved hairs, sometimes red-punc- tulate when young, red-brown to blackish when ripe. Distr. SE. and E. Asia to the Solomon Islands, Australia, Tasmania, and in tropical Africa, in Malaysia: all over the Archipelago, not yet found in the Lesser Sunda Islands, Madura and Kangean Islands. Ecol. In shaded and forested places, 1-2900 m, but in the periodically dry parts of Central and E. Java not below 1000 m, rarely descending to 700 m near hot springs or along stream banks, decidedly avoiding the semi-arid regions. Uses. Leaves as a fish-poison. Vern. Pegagah gajah (Mal. Pen.); pegagoh, pe- gagan, mangi-mangi (Sum.); dulang sontak, daun sontok, S; variable in Javanese. Notes. Small forms from high altitudes are dif- ficult to separate from H. sibthorpioides LAMK. These specimens have at least 15 fruits in the um- bellule, and for that reason I refer them to H. javanica. In New Guinea some specimens have very long-pedicelled flowers; there is a series of transitions to subsessile and sessile flowers. 2. Hydrocotyle sibthorpioides Lamx, Enc. Méth. Bot. 3 (1789) 153; Buw. Blumea 2 (1936) 128 (Jit.). —AH. nitidula Ricu. Ann. Gén. Sc. Phys. 4 (1820) 116 FLORA MALESIANA [ser. I, vol. 4 60, t. 63, fig. 33.—H. ranunculoides var. incisa BL. Bijdr. 15 (1826) 884.—H. splendens Bu. Bijdr. 15 (1826) 884.—H. hirsuta var. minuta BL. Bijdr. 15 (1826) 884.— H. rotundifolia DC. Prod. 4 (1830) 64.—H. hirsuta (non Sw., nec SPRENG.) DC. Prod. 4 (1830) 67.—H. latisecta ZOLL. Syst. Verz. (1854) 138, 140.—H. zollingeri MOLKENB. in Mra. PI. Jungh. (1851) 91.—H. puncticulata Mia. Fl. Ind. Bat. I, 1 (1856) 732.—H. benguetensis ELM. Leafl. Philip. Bot. 2 (1909) 628.—H. delicata Ev. Leafl. Philip. Bot. 2 (1909) 629.—Fig. la-b. Stems long-creeping or with ascendent extremi- ties, sometimes almost caespitose, terete, thin or almost filiformous, glabrous or sparsely hairy. Stipules !/2-1 by nearly 11/2 mm, ovate to obovate, acute, entire or fringed. Petioles 1/2-6 cm, or even shorter in the uppermost /eaves, more or less Fig. 1. a—b. Hydrocotyle sibthorpioides LAMK, X '/2, c. Centella asiatica URBAN, X 1/s. hairy; lamina !/3—2!/2 cm through, roundish to 5- angular in outline, deeply cordate, 3—5-lobate to 3—5-partite; segments crenate to serrate, more or less pilose to hirsute. Inflorescences single along the creeping stems; peduncles 0-3 cm, filiformous, glabious or short hairy; involucres 4-10 around and between the flowers, nearly 1 by !/2 mm, ovate, lanceolate, acute, base with 2 acute teeth, some- times filiformous, lower ones reflexed in fruit. Pe- dicels 10-15; petals greenish white, nearly 3/4 by 1/2 mm, ovate, acute. Mericarps 1—-1'/4 by 3/4 mm, yellow to brown, glabrous or with short stiff hairs, sometimes red-punctulate. Distr. Australia, tropical Asia, tropical Africa, S. America (?), in Malaysia: all over the Archi- pelago. Ecol. Sunny or slightly shaded, damp, fertile localities, along streambanks, between stones of pathways and alongside walls, from 1—4050 m. Uses. Raw or steamed eaten with rice; medici- nal against skin diseases. Vern. Kurawet galeng, antanan in several com- binations, S, sumud, samangi in several combina- tions, J, but Javanese names are rather variable; salatun, patekan tjéna, Md. Notes. Very variable as to leaf shape, depth of incisions, and hairiness; the numerous forms are connected by series of transitions. 3. Hydrocotyle vulgaris LINNE, Sp.Pl. 1 (1753) 234; Buw. Blumea 2 (1936) 133 (Jit.), Stems thin, creeping. Petioles 1-17 cm, with spreading hairs to the apex; lamina 3/4—3!/2 cm through, orbicular, peltate, 8—13-nerved, coarsely crenate to slightly lobed. Inflorescences solitary or few together on the nodes; peduncles !/2-14 cm, filiformous; with 1-10 whorls of flowers, each flow- er with an ovate membranous, acute bract. Petals nearly 3/4mm, ovate, white or reddish. Fruits smooth, 1!/2-2 by 13/4—21/2 mm, transversely ellip- tical. Distr. Europe, N. Africa, Australia, and the Marshall Islands, apparently absent in continental Asia; in Malaysia only in W. New Guinea (Arfak Mts). Ecol. In open marshes, -+ 2000 m. Notes. Description after European and New Guinean materials. In Malaysian specimens the fruits are smooth, not covered with reddish warts as in the European form. 2. CENTELLA Linng, Pl. Afr. Rar. (1760) 28; Buw. Blumea 2 (1946) 133.—Hydrocotyle sect. Cen- tella Bth. Fl. Austr. 3 (1866) 338. Perennial, sometimes suffruticose, erect, prostrate, or rooting at the nodes. Leaves entire, crenate or lobate, palminerved; petioles with sheaths. Umbels sim- ple, sessile or subsessile. Involucres few or 0. Calyx teeth obsolete. Petals entire, imbricate in bud; disk plane, margin elevate; styles from the base filiformous. Fruits laterally flattened, vittae 0; commissure narrow; mericarps with dorsal ribs marginate, lateral and intermediate ribs arcuate, all connected by veins, sometimes with 2—4 secondary ribs. Dec. 1949] Distr. All over the world. UMBELLIFERAE (Buwalda) Note. Solandra L. Syst. ed. 10 (1759) 1269 is an older name for Centella but is rejected against the Solanaceous Solandra Sw. (1787); it is, therefore, not available in the Umbelliferae. 1. Centella asiatica (L.) Urs. in MArrt. FI. Bras. 11, 1 (1879) 287, t. 78, fig. 1; Buw. Blumea 2 (1936) 134.—Pes equinus RUMPH. Herb. Amb. 5, p. 455, t. 169, f. 1.—Hydrocotyle asiatica LINNE, Sp.PI. 1 (1753) 234.— Trisanthus cochinchinensis Lour. Fl. Coch. 1 (1790) 176.—Hydrocotyle hebecarpa DC. Prod. 4 (1830) 63.—H. asiatica var. hebecarpa Hassk. Pl. Jav. Rar. (1848) 459.—H. asiatica yar. pedunculata O.K. Rev. Gen. 1 (1891) 268.—Fig. Ic. Stems creeping with long stolons, more or less puberulous in the young state. Leaves in rosettes; petioles 1-40 cm, sometimes puberulous; lamina 1-7 cm diam., roundly reniform, crenate or crenate-dentate. Umbels solitary or 2—5 together in the axils of nearly 3 mm long bracts; peduncles 1/2-5 cm, shorter than the petioles. Flowers usually volucres 2, 3-4 by nearly 1'/2 mm, ovate. Petals red, 1—1'/2 by 3/4mm. Mericarps about 2 by 1!/2mm, subhairy when young. Distr. Pantropie, in Malaysia: all over the Ar- chipelago. Ecol. Sunny or slightly shaded, fertile, damp localities, along streambanks, also between stones of pathways and alongside walls, 1—2500 m. Uses. Leaves raw or steamed eaten with rice. Medicinal uses many, especially against skin dis- eases and as a diuretic. Capacity for holding earth against erosion. Vern. (daun) Pegaga (Mal. Pen., Sum., Born.) daun kaki kuda, antanan in several combinations, S; patjul gowang, rendeng, gagan-gagan, J (but rather variable), kolotide manora (Tern.), dogauke, 3, middle one sessile, Jateral ones pedicellate; in- gogauke, andanan (New Guinea). 3. TRACHYMENE RupGgE, Trans. Linn. Soc. Lond. I, 10 (1811) 300; NorMan, J. Bot. 69 (1931) 287; Buw. Blumea 2 (1936) 138.—Didiscus DC. in Curt. Bot. Mag. 55 (1828) t. 2875; Domi, Sitz. Ber. Bohm. Ges. Wiss. (1908) 2. Annual or perennial, hirsute to glabrous, sometimes glandular-hairy. Stems erect, procumbent or ascendent, branched, often in a sympodial way. Leaves alternate along the stems, sometimes also in rosettes, roundish cordate to broadly cuneate, ternately divided, or entire, narrow-cuneate to subspathulate; petioles with sheaths. Umbels simple, terminal or opposite the leaves, sometimes in a corymbiform di-monochasium. Involucres linear. Calyx teeth minute, rarely subulate. Petals en- tire, imbricate in bud. Disk plane. Styles from the base filiform. Fruits laterally flattened, vittae 0; commissure narrow; mericarps with dorsal ribs, marginate, lateral ribs in the commissure, intermediate ribs arcuate, subprominulent. Car- pophore persistent, undivided. Distr. This genus is chiefly Australian; outside Australia it is spread to New Caledonia and the Fiji Islands, in Malaysia it occurs in New Guinea, Timor, Flores, Celebes, Borneo, and the Philippines. KEY TO THE SPECIES 1. Plant glandular-hairy. 2. Ovary hairy; ripe fruits roughly tuberculate with glandular hairs; annual, erect, cultivated 16. T. caerulea 2. Ovary glabrous; ripe fruits smooth; wild mountain species 11. T. adenodes 1. Plant not glandular-hairy. 3. Leaves nearly triangular and somewhat hastate in outline, tripartite or ternate with the middle seg- ment longer than the lateral ones 5 . . . 6. T. erodioides 3. Leaves never triangular hastate, more roundish or more cuneate in ‘outline, if tripartite or ternate, then the middle segment hardly longer than the lateral ones. 4. Leaves about as long as broad, base cuneate. 5. Petiole at least twice as long as the lamina. Prolongated leafy stems absent. Stem not papillose. 2. T. novoguineensis 5. Petiole at most as long as the blade. Leafy branched stems present. Stems, petioles, and peduncle densely papillose ct 1A eee 13. T. flabellifolia 4. Leaves broader than long or - longer than broad. 6. Leaves longer than broad, all of them cuneate to spathulate. 7. Pedicels 1-2. Leaf-blade spoon-shaped, 1-nerved, 2-2/2 by 1/2-3/4 mm, tip mucronate. Leaves densely imbricate. Cushion plant Se ee ee SE pulvilliforma 118 FLORA MALESIANA [ser. I, vol. 42 7. Pedicels at least 10. Leaf-blade otherwise, at least 7 by 3 mm, mostly with at least 3 apical teeth. 8. Leaves coriaceous and stiff, the lamina at least 5 times as long as broad . 4. T. rigida 8. Leaves not coriaceous and stiff, the lamina at most 3 times as long as broad. 9. Petiole at least twice as long as the lamina; prolongated leafy stems absent. 2. T. novoguineensis 9. Petiole as long as the lamina or shorter; leafy stems present. 10. Peduncles shorter than the leaves. Leaves subspathulate, not in rosettes but somewhat crowded towards the extremities of the stems Reaoel 2 14. T. rosulans 10. Peduncles longer than the leaves. Leaves cuneate. 11. Leaves broad-cuneate, 1—2!/2 by 1-3 cm, single or few together in axillary clusters . Leaves narrow-cuneate, base of the stem and the meats Sp 13. T. flabelliformis 2 by !/2-1 cm, about twice as long as broad, in rosettes at the 3. T. koebrensis 6. Leaves broader than long, sometimes the upper ones cuneate, ‘xarely also the lower broadly cuneate. 12. Stems procumbent; leaves to 1 cm long and broad, their teeth with apical hairs. 5. T. acrotricha 12. Stems erect or ascendent, sometimes caespitose; leaves generally more than 1 cm long and broad, their teeth not with apical hairs. 13. Plants with rosettes at the base of the stems, sometimes also in the upper leaf axils and at the bases of the branches. 14. Umbels single from the rosettes, or moreover from the PLCS stems, but never forming a corymbiform dichasium . 1. T. saniculaefolia 14. Umbels forming a corymbiformous dichasium on more or less erect stems. 15. Calyx teeth at most 3/4 mm long; leaves more’ or less divided, but not ternate. 7. T. celebica 15. Calyx teeth up to 2!/2 mm long; leaves ternate 8. T. sarasinorum 13. No rosettes at the base of the stems and the branches, or if ‘small rosettes are present at the base of the stems, these rosettes have disappeared before flowering and the umbels do not form a terminal corymb. 16. Leaves to 1-2 cm long and 3-4 cm broad; surface of stems, sheaths, and petioles densely papil- lose; fruit with knob-shaped trichomes 16. Leaves more than 2 cm long and 3 cm broad; fruits smooth. 12. T. papillosa stems, ‘sheaths and petioles not papillose; 17. Leaves not in rosettes but more densely placed in the lower thicker portion of the stem, 3-fid to ternate, biserrate with acute teeth 9. T. acerifolia 17. Probably small rosettes at the very base of the stem, the latter very ‘slender i in its lower portion. Leaves 21/2-7 by 4-8 cm, ternate with petiolulate leaflets, the latter serrate with broad, shortly acuminate teeth 1. Trachymene saniculaefolia STAPF in Hook. Ic. Pl. 24 (1894) t. 2308; Buw. Blumea 2 (1936) 141 (lit.).—Didiscus saniculaefolius MERR. Philip. J.Sc. Bot. 2 (1907) 255, 256, 292.—Hydrocotyle azorel- lacea F.v.M. J. Bot. 31 (1893) 324, nomen. Perennial, reddish, more or !ess hirsute to glabrous; caudex with rosettes from which sym- podial leafy stems with terminal inflorescences and axillary rosettes, flower-bearing or not. Stems te- rete, striate. Sheaths 5-10 by 2-3 mm, tapering into the petiole, ciliate; petioles 3-13 cm; lamina 3/4-4 by 1-6 cm, roundly reniform to broadly cune- ate, trifid to tripartite, even ternate, segments broadly rhomboid or narrower, sometimes divided again, apical part: ultimate segments serrate to lo- bate. Umbels terminal in the rosettes or on elon- gated stems opposite the leaves; peduncles 3— 29 cm; involucres 7—25, 5-15 by 1—3 mm, lanceo- late, acuminate, sometimes dentate, spreading, ap- pressed in fruit; pedicels 5—more than 30, S-I15mm, inner ones gradually shorter, spreading, incurved in fruit. Calyx teeth 1/2-2 by 1—1!/2 mm, triangular, acute, equally developed or one larger. Petals 2— 2!/2 by '/2-11/2 mm, white to dark pink, ovate to lanceolate; styles !/2-11/2 mm. Mericarps 11/2-3 by 1-2 mm, glabrous, reddish purple to purple. Distr. Australia (N.S. Wales), in Malaysia: 10. T. arfakensis Philippines (Mindoro), Br. N. Borneo and E. New Guinea. Ecol. Open shallow damp places, at the base of rock walls at falls, rock crevices, grasslands, on burnt over ground and in forest glades, 1800— 4020 m. Uses. In Borneo used as Dusan medicine. Notes. Rather variably as to the length of the stems, hairiness, shape and incision of the lamina. 2. Trachymene noyoguineensis (DOMIN) BUw. Blu- mea 2 (1936) 144. fig. 2a.—Didiscus saniculifolius var. novoguineensis DOMIN, Sitz. Ber. BGhm. Ges. Wiss. (1908) 67.—Fig. 2a. Perennial, often reddish, more or less hirsute to glabrous; caudex with rosettes, bearing again axil- lairy rosettes; dwarf forms only 2-3cm. Leaf sheaths 2'/2 by 3 mm, ciliate, tapering in the petiole; petioles 1-13 cm; lamina (3)—7—30 by (2!/2)—5- 17mm, cuneate, trilobate or trifid, apical part: seg- ments with 2—3 subacuminate teeth. Peduncles (1!/2)—3!/2-37 cm, terete, striate or subsulcate; in- volucres 7—13, 8-12 by 1/2-1!/4 mm, lanceolate, acute, glabrous or margin ciliate, spreading, ap- pressed in fruit; pedicels 12-30, to 5 mm, inner ones gradually shorter, spreading, to 14mm and incurved in fruit, glabrous. Calyx teeth 1/4—!/2 mm, Dec. 1949] UMBELLIFERAE (Buwalda) 119 RHoeksema Fig. 2. a. Trachymene novoguineensis (DoMIN) Buw., b. Trachymene arfakensis (GiBBs) Buw., X 2/3. 120 FLORA MALESIANA [ser. I, vol. 42 narrow or broad-triangular, somewhat enlarged in fruit. Petals c. 11/2 by 3/4 mm, creamy white, white or violet, elliptical. Styles to 3/4.mm. Mericarps to 3 by 1'/2mm, glabrous, brown-yellow or tinged with red to dark violet. Distr. Malaysia: W._E. New Guinea. Ecol. Open, stony localities, brook banks, marshy grasslands, 2700-3720 m. 3. Trachymene koebrensis (Gipps) Buw. Blumea 2 (1936) 146, fig. 1la.—Didiscus koebrensis GrBBs, Contr. Arfak Mts (1917) 165.—Fig. 3a. Perennial, entirely glabrous. Stems to 40cm, erect or prostrate, sometimes finely papillose, with leaves over the whole length and rosettes in the leaf axils, densely set with swollen petiole-bases in the basal part and below the rosettes, with branches from the axillary rosettes, again with rosettes in the leaf axils. Sheaths nearly 2 by 3 mm, tapering into the petiole; petiole 1-2 cm, canaliculate above; lamina 1-2 by 1/2-1 cm, cuneate, tapering into the petiole, apex with 3—5 triangular, acute teeth. Pe- duncle 6!/2-8!/2 cm, terete, striate; involucres 5—13, nearly 6 by !/2 mm, lanceolate, acute; pedicels 15— 30, to 5mm, the inner ones shorter, spreading, somewhat incurved in fruit. Calyx teeth 3/4-1!/2mm, narrowly triangular or subulate, sometimes un- equally developed. Petals nearly 11/4 by 3/4 mm, obovate, white or white with purple tinge; styles nearly 1!/smm. Mericarps to 2'/2 by 1!/2 mm, tuberculate or smooth. Distr. Malaysia: W. New Guinea. Ecol. Open burnt localities and sterile lime- stone slopes, 2400-3225 m. 4. Trachymene rigida Buw. Blumea 2 (1936) 147, fig. 1 b-d.—_Didiscus odontocoleus BUW. ex STEEN. Bull. Jard. Bot. Btzg III, 13 (1934) 255, nomen. Fig. 3b-d. Perennial, entirely glabrous; branched caudex with rosettes from which sympodial leafy stems with terminal inflorescences and axillary few- leaved rosettes from which stems branched in the same mode. Stems procumbent, angulate, sulcate, at the nodes incrassate. Sheaths 1-3 mm long and broad with subulate thick coriaceous to 3 mm long appendages; petioles 2 by 0.1 cm, difficult to dis- tinguish from the lamina; /eaves with petiole 2— 9 cm; lamina 4~7 mm broad, thick and stiff-cori- aceous, narrow-cuneate-spathulate, apex with 1-5, mostly 3, obtuse, triangular teeth 3 by 1-2 mm; margin entire, subrecurved. Peduncles 5!/2-11 cm, 1/2-11/2 mm _ thick, angulate, sulcate; involucres 10-12, 5—10 by !/2-11/2 mm, lanceolate, acute or sub- obtuse; pedicels 20 or more, 2-4 mm, somewhat spreading, erect in fruit. Calyx teeth 1/4-/4 mm, obtuse, persistent. Petals 11/2-2 by 1 mm, oblong- ovate, inside white; styles 11/2-2 mm. Mericarps to 3 by 21/2 mm, dark violet or brownish. Distr. Malaysia: W. New Guinea. Ecol. Open localities, 3000 m. 5. Trachymene acrotricha Buw. Blumea 2 (1936) 148, fig. le-f.—Fig. 3e-f. Perennial; caudex at the top densely set with in- crassate leaf rudiments. Stems to 13 cm, prostrate, sulcate, densely-hirsute towards the tip, hairs divar- icate, to 11/2 mm. Leaves sparse, rosettes absent; sheaths 4 by 2 mm, tapering into the petiole, mar- gin long-ciliate, hairs to 3 mm; petiole to 7 cm, canaliculate, densely long-hirsute, hairs to 2 mm; lamina to 7 by 10-15 mm, of the lower leaves reni- form, of the upper leaves rhomboid in outline, tripartite or trifid with cuneate segments, apical portion broad-dentate, teeth ending in an apical hair, subcoriaceous, involute when dried up, pal- minerved, glabrous above, beneath sparsely hirsute on the nerves. Umbels opposite the leaves in the upper portion of the stem; peduncles !/2-2 cm, den- sely hirsute; involucres 8-10, 4mm, lanceolate, acute, canaliculate, glabrous with hairy teeth and tip; pedicels 10-20, to 4 mm, the inner ones shorter, glabrous, somewhat dilatate at the tip. Calyx teeth 1/2] by 1 mm, elliptical, triangular, acute. Petals c. 11/2 by 1 mm, apiculate. Styles to 3/s mm. Meri- carps 2'/4-3 by 11!/2-2 mm, glabrous, ribs indistinct. Distr. Malaysia: SW. Celebes. _ Ecol. Stony localities, mountain heaths, 3100 m. 6. Trachymene erodioides Buw. Blumea 2 (1936) 149, fig. 1 g-h.—Didiscus erodioides Buw. ex STEEN. Bull. Jard. Bot. Btzg III, 13 (1934) 255, nomen.—Fig. 3g-h. Small; stems creeping, terete, subincrassate at the nodes, hirsute or subhirsute, with hairs to 2mm. Leaves single and in few-leaved axillary rosettes; sheaths ca 2 by 1 mm, tapering into the petiole, outside densely hirsute, hairs 1-5 mm; pe- tioles 11/24 cm, canaliculate, pilose with divar- icate crisp hairs; lamina 11/2-2!/2 by 1-2 cm, in out- line ovate-triangular, somewhat hastate, tripartite or ternate, terminal segment 1-2 by !/2-1!/2 cm, triangular-rhomboid, lateral segments 5—12 by 5-7 mm, ovate, all segments pinnatifid at the base, crenate towards the apex with short acuminate tips. Umbels teiminal or from the axillary rosettes; peduncles 2!/2-3!/2 cm, ascendent, slender, terete, striate, densely hairy, hairs 1-2 mm, crisp; invol- ucres 5-6, 3-5 mm, lanceolate, glabrous or sub- ciliate; pedicels 12-15, to 7 mm, inner ones shorter, glabrous. Calyx teeth to 1/s mm or absent (on the fruits). Styles + 1/2 mm. Mericarps 2'/2-3 by nearly 2 mm, glabrous, equal or subequal. Distr. Malaysia: SW. Celebes. Ecol. Stony localities in mossy forest, 2700 m. 7. Trachymene celebica HemMsL. Kew Bull. (1896) 37; Buw. Blumea 2 (1936) 149.—Didiscus celebicus SARASIN, Reisen in Celebes 2 (1905) 337.—Didiscus buginensis WOLFF in Feppe, Rep. 17 (1921) 439. Perennial; caudex with rosettes from which flow- er-bearing stems and lateral rosettes, sessile or on short stolons from the upper leaf axils. Stems 20— 50 cm, erect or ascending, nearly terete, more or less ribbed, densely hirsute to subhirsute, little branched and few-leaved beneath, terminated by a corymbiform inflorescence of umbels. Sheaths of the rosette leaves 2 by 3/4-11/3 cm, outside glabrous, towards the apex hirsute and ciliate, hairs 2-4 mm; petioles 5-15 cm, hirsute; lamina Dec. 1949] UMBELLIFERAE (Buwalda) 121 Fig. 3. a. Trachymene koebrensis (Gipss) Buw., b-d. Trachymene rigida Buw., e-f. Trachymene acrotricha Buw., g-h. Trachymene erodioides Buw. (plants x 2/3, fruits 4). 122 5-13 by 7-14 cm, in outline roundish, deeply cord- ate, 3—7-palmatifid, segments obovate, 3-lobed, biserrate, densely hirsute; cauline leaves and inflo- rescence bracts gradually smaller and shorter peti- oled, uppermost ones subsessile with less numerous and narrower segments and smaller sheaths. Pe- duncles 2'/2-6 cm, upper ones shorter; involucres numerous, nearly 10mm, narrowly lanceolate, long-acuminate, appressed; pedicels more than 50, up to 17 mm, inner ones shorter, spreading, in- curved in fruit. Calyx teeth small, acute, persistent. Petals white, nearly 21/2 by 1!/2 mm, elliptic, acute. Styles nearly 3 mm, persistent. Mericarps nearly 4 by 3 mm, red. Distr. Malaysia: SW. Celebes. Ecol. Open, stony localities, 2300-3000 m. Uses. Roots (raw) as medicine against stomach- ache. Vern. Kriongo, djahé merah. 8. Trachymene sarasinorum (WOLFF) Buw. Blumea 2 (1936) 151.—Didiscus sarasinorum WOLFF in FEDDE, Rep. 17 (1921) 440. Perennial; caudex with rosettes from which flower-bearing stems and lateral rosettes, sessile or on short stolons from the upper leaf axils. Stems erect, 30-40 cm, terete, striate, sparingly hirsute, more densely at the nodes. Leaves nearly all in a rosette; sheaths 3-6 by 5—8 mm, at the back and the margin with 1-2 mm long hairs, abruptly con- tracted into the petiole; petiole 6-8 cm, hirsute, more densely towards the lamina; Jamina 4-4!/2 by 6-7 cm, roundish in outline, deeply cordate, ter- nate, the central leafiet nearly 4 by 31/2 cm, tripar- tite, lateral leaflets hardly smaller, obliquely trifid, all ultimate segments 2—3-lobed, coarsely serrate, rather sparingly appressedly hirsute; cauline leaves smaller, shorter petioled, bracts of the dichasium nearly sessile. Peduncles 34cm, terete, striate, shortly hirsute; involucres numerous, 7—10 by 1/2 mm, narrowly lanceolate, acuminate with few hairs at the margin and on the midrib, spreading, appressed in fruit; pedicels 7-11 mm, spreading, suberect in fruit. Calyx teeth nearly 2!/2 mm, subu- late. Petals 2—2!/2 by 1 mm, ovate, white. Styles nearly 2mm. Mericarps nearly 4'/2 by 31/2 mm, entirely glabrous. Distr. Malaysia: SW. Celebes. Ecol. Probably open, stony localities, 1100 m. 9. Trachymene acerifolia NORMAN, Journ. Bot. 69 (1931) 287; Buw. Blumea 2 (1936) 151, fig. 3.— Didiscus acerifolia STEEN. Bull. Jard. Bot. Btzg Ill, 13 (1934) 255. Stems 20-45 cm, erect and terete in the lower portion, terete or subangular in the upper part, with spreading branches, the whole densely vel- vety hairy with yellowish brown indumentum, glabrescent; glabrous in the young state. Leaves rather densely set in the basal part, more remote upwards, hirsute to glabrous; petioles of the lower leaves to 13!/2cm, gradually shorter upwards, in the upper leaves nearly absent, all of them slightly sheathing at the base, hairy like the stems; lamina~ palmatifid to ternate, segments rhomboid to obo- FLORA MALESIANA [ser. I, vol. 4 vate, the middle one 3-lobate to trifid, besides, all biserrate with acuminate teeth; lamina of the upper leaves smaller and more cuneate. Umbels opposite the leaves; peduncles 1-5 cm, to 7 cm in fruit, terete, grooved, hairy like the stems; involucres 7-10, to 3 mm broad, shorter as or as long as the pedicels, lanceolate, hairy like the leaves; pedicels 25-40, to 7mm, to 15 mm in fruit, inner ones shorter. Calyx teeth to !/2mm, acute or obtuse. Petals to 2 by 1 mm, cream to pink, elliptical, acute. Styles nearly '/2 mm, to 1!/2 mm in fruit. Mericarps to 6 by 4mm, equal. Distr. Malaysia: Lesser Sunda Islands (Timor, Flores), SE. Celebes. Ecol. In mountain forests, damp places, 1800- 2600 m. Notes. The specimens from Celebes differ from those from the Lesser Sunda Islands by stronger developed leaf sheaths, by broader involucres which enclose the flower when young, and by pink to purple flowers. 10. Trachymene arfakensis Buw. Blumea 2 (1936), 154, fig. 2b.—Didiscus arfakensis GripBs, Contr. Arfak Mts (1917) 166.—Fig. 2b. Stems to 60cm, erect or suberect, glabrous, slender, base thickened with scars and remnants of leaf sheaths, unbranched in the lower portion, terete, striate to subsulcate, several times dichoto- mously branched in the upper part, branches spreading and sympodial. Sheath 2-7 by 1!/2- 4 mm, tapering into the petiole, ciliate with hairs to 2 mm; petioles 2-7 cm in the lower /Jeaves, grad- ually shorter upwards, canaliculate, glabrous or towards the lamina with few to 2 mm long hairs; lamina 2!/2-7 by 4-8 cm, roundish-cordate, ternate, leaflets with petiolules to 11/2 cm, 2-3-fid to 2-3- partite, segments 3-lobed and coarsely serrate, teeth acuminate and apiculate, upper surface sub- glabrous, beneath sparingly hirsute, especially on the nerves, base long-ciliate. Umbels opposite to each other leaf; peduncles 11/4-7 cm, terete to sul- cate, involucres 5-10, 5-10 mm, linear to filiform, broadest ones with few filiform teeth; pedicels 20-30, 5-8 mm, 10-15 mm in fruit, spreading, inner ones shorter. Calyx teeth hardly any. Petals 1—11/2 by 3/4-1 mm, white, ovate, acute. Styles 1—1'/2 mm. Mericarps to 5 by 3 mm. Distr. Malaysia: Celebes, W. New Guinea (Mt Arfak). Ecol. Marshy, muddy localities, often in groups, 1500-2400 m. 11. Trachymene adenodes Buw. Blumea 2 (1936) 155, fig. 4a—b.—Fig. 4a—b. Stems to 50 cm and more, terete, striate, lower portion ascendent with rosettes at the base, glabrous, unbranched, upper portion dichotom- ously later sympodially branched, densely hirsute, hairs to 2 mm. Leaves rather densely in the lower portion, upwards more remote; sheaths 5-8 by 3-5 mm, semi-amplexicaulous, tapering into the petiole, glabrous, margin long-ciliate with partly glandular hairs to 3 mm; petioles 7-20 cm, up- wards gradually shorter, uppermost ones nearly Dec. 1949] UMBELLIFERAE (Buwalda) F Hoekse Fig. 4. a-b. Trachymene adenodes Buw., c. Trachymene papi b. glandular hair of the petiole < 16, d. C ti Mma llosa Buw. (plants x 2/3, x 4.) 123 124 FLORA MALESIANA [ser. I, vol. 4? absent, towards the lamina densely hirsute with glandular hairs to 3mm; lamina of the rosette leaves and lower leaves 4!/2 by 71/2 cm, roundish- cordate to subreniform, 3—5-partite to ternate, segments rhomboid-ovate, middle ones trifid with 2-3-lobate parts, all segments moreover serrate with broad, subacuminate to subapiculate teeth, sparsely pilose with partly glandular appressed hairs, to 2 cm especially towards the margin and in the incisions. Umbels opposite the leaves and in the bifurcations; peduncles to 10cm, shorter towards the end of the stems, towards the top with glandular hairs; involucres 6-8, 7-10 by 1-1!/2 mm, lanceolate, acute, glabrous or long- ciliate; pedicels -- 30, 7-9 mm, inner ones shorter, glabrous. Calyx teeth nearly '/4 mm, broad-trian- gular. Petals 11/2-2 by 1-11/2mm, obovate. Styles + 1!/2mm. Mericarps (unripe) 2 by 3!/4 mm, glabrous. Distr. Malaysia: NE. New Guinea. Ecol. Mountains, 2400-3000 m. Notes. When more material will come to hand this species might possibly appear to be referable to T. arfakensis which it resembles in leaf shape but from which it differs by its longer petioles, longer peduncles and the glandular indumentum of the petioles, stems and peduncles. As yet I have seen no transitional forms. 12. Trachymene papillosa BUw. Blumea 2 (1936) 157, fig. 4c-d.—Didiscus scabriusculus Buw. ex STEEN. Bull. Jard. Bot. Btzg III, 13 (1934) 255, nomen.—F¥ig. 4c. Stems 20-40 cm, probably ascendent, terete, densely papilose and densely hirsute with 1 mm long, stiff hairs, branched in the upper portion. Leaves sparse; sheaths 2-3 by 2 mm, semi-amplex- icaulous, tapering into the petiole, papillose like the stem, above long-ciliate, hairs 1-2 mm; petioles 1/2-21/2 cm, hirsute and papillose like the stems; lamina 1-2 by 3-4 cm, orbicular-reniform in outline, ternate; leaflets 1-2 by 1—1!/2 cm, attenuate towards the base, 2—3-fid or 2—3-partite, terminal segments often biserrate, at the base papillose to glabrous, on both sides sparsely hirsute to glabrous. Umbels terminal, often opposite the leaves; pe- duncles 3—8!/2 cm, terete, striate, hirsute and papil- lose like the stem or glabrous; involucres 6-12, 4-5 by !/2-1 mm, linear-lanceolate, acute, glabrous or long-ciliate; pedicels 30-50, to 5 mm, inner ones shorter, glabrous, spreading, erect to reflexed in fruit. Calyx teeth !/4—-1/2 mm long and broad, trian- gular, equal. Petals circa 11/2 mm, reddish white, ovate, acute. Style 1—-1!/2 mm. Mericarps to 2 mm long and broad, equal, black, with tuberculiform scales, especially between the intermediate ribs and the commissure, to entirely smooth. Distr. Malaysia: SW. New Guinea (Central Range). Ecol. Grassy, deforested slopes, on sandy soil, also forest edges, 1600—3500 m. 13. Trachymene flabellifolia n. sp. Species nova ad Tr. papillosam Buw. accedens tamen foliis cuneatis flabelliformibus distinguenda. Perennial herb. Rhizome 7—12 mm thick; stems up to 30 cm long, branching from the base, ascend- ent, furrowed or subangulate, near the base black- corky, roughish, upper part densely papillose and hirsute (hairs up to 1 mm long, appressed, and stiffish). Leaves single or few together in axillary clusters, 5-15 mm apart; sheath 2-3 mm long, 11/2-2 mm wide, narrowing to the petiole and on the margin ciliate (hairs up to 2 mm long); petiole 6-11 mm long, sparsely hirsute (hairs up to 1 mm long); blade 1—2!/2cmlong, 1—3cm wide, fan-shaped, incised or split into 2—3-parted wedge-shaped seg- ments, at the tip sharply toothed; main nerves on both surfaces distinctly flabellate. Umbels placed in the upper part of the stem opposite the leaves; ped- uncle terete, striate, subpapillose, ?—5!/2 cm long, sparsely hirsute (hairs up to 1!/2 mm long, spread- ing); bracts forming an involucre, 11 or less, up to 11 mm long and c. 3/4 mm wide, acute, lanceolate, at the base connate; pedicels 30—SO, terete, slender, glabrous, the outer up to 5 mm long, the inner shorter, slightly longer when in fruit. Calyx with c. 1/4mm long, sharp teeth. Petals 3/4-1 mm long, -c. 1 mm wide, ovate, subconcave. Stamens 5, an- thers dorsifix, rounded-elliptic. Styles 2, subfili- form, 1!/4-1!/2 long, curved inwards. Fruit much flattened, kidney-shaped. Mericarps up to 3 mm long and up to 2!/2 mm wide, median rim c. 1 mm distant from the commissure, slightly crowned by the styles, densely covered by knobbly hair-like outgrowths; carpophore entire, 11/2-13/4 mm long, hardly two-tipped. Distr. Malaysia: West New Guinea (Central Range, near Lake Habbema, 3225 m camp, BRASS 9586, type). Ecol. Common in mossy glades, 3225 m alt. Notes. Allied to T. papillosa but different by its cuneate to flabelliform leaves. 14. Trachymene rosulans (DANs.) Buw. Blumea 2 (1936) 158.—Didiscus rosulans DANS. Brittonia 2 (1936) 135, cum icone. Perennial (or annual). Main root fusiform, branched. Main stem erect, to 13 cm, at the base to 2!/2 mm thick, almost covered with the thick- ened bases of leaf sheaths, producing procumbent or ascendent branches, with scale-like leaves, to 14 cm; upper portion with branches that are like the upper portion of the main stem, whole plant a semi-globose whole. Leaves scattered, more den- sely set towards the extremities, somewhat forming terminal rosettes; sheaths 2-12 by 3mm, with membranous margin, tapering into the lamina; lamina 8-18 by 3—9 mm, spathulate, apical portion with 3 acute or obtuse teeth. Umbels opposite the leaves; peduncles 5-12 mm, terete, striate; invol- ucres 8-12, 6-9 by 11/2 mm, lanceolate; pedicels 10-20, 4-7 mm, inner ones shorter, hardly elon- gated in fruit. Calyx teeth to 3/4 mm, triangular, acute. Petals nearly, 1!/2 by 1 mm, pink or pale pink, roundish-elliptical; styles nearly 1!/4 mm. Mericarps to 31/2 by 3 mm, entirely glabrous. Distr. Malaysia: SE. New Guinea. Ecol. Open places, burnt fringes of forest, grasslands, 2840 m. Dec. 1949] 15. Trachymene pulvilliforma n. sp. Ab omnibus speciebus generis Trachymene differt modo crescendi, pulvillos densos formante, foliis densissime imbricatis apicem caulis versus et magis minusve in rosulas confertis. Pars inferior folii semi- amplexicaulis et vaginans, lamina cochlearis 2- 2!/2 mm longa, '/24/4 mm lata, apice mucronata, uninervata. Umbellae simplices, brevissime pedun- culatae bracteis 2-4 involucratae, pedicellis 1-2. Ex affinitatis T. rosulans (DANSER) BuUW. A herb, probably perennial, entirely glabrous; root fusiform, branching, fibrous; stem prostrate and much branching from the base, branches close, the slightly ascending upper ones forming a half- spherical cushion which is up to 1 cm deep and 5—15 cm in diam. Leaves very densely imbricate, near the top more or less in clusters together, the basal part half amplexicaulous and sheathing; sheath appressed, 1!/2-3 mm long, !/2-1 mm broad, narrowing towards the blade; blade spoon-shaped, 2—2!/2 mm long, '/2-3/4 mm wide, one-nerved, tip mucronate. Umbels terminal or opposite the leaf clusters, when flowering hidden among the leaves, when in fruit slightly exserted; peduncle somewhat flattened c. 2 mm long; bracts 2—4, forming an in- volucre, 1!/2-2 mm long, up to !/2 mm wide, lan- ceolate or spathulate-lanceolate. Pedicels 1-2, slightly flattened, c. 1 mm long, in fruit up to 2'/2 mm long. Calyx indistinctly toothed. Petals 5, c. | mm long, c. 3/4 mm wide, white. Stamens 5, anthers dorsifix, rounded-elliptical, pink. Styles 2, subfiliform, c. '/2 mm long. Fruit glabrous, strongly flattened, kidney-shaped. Mericarps up to 2 mm long and up to 1'/2 mm wide, rim indistinct, the median 3/4mm distant from the commissures, slightly crowned by the styles, carpophore entire, c. 1 mm long, hardly two-tipped. Distr. Malaysia: West New Guinea, (3 miles E UMBELLIFERAE (Buwalda) 125 of the summit of Mt Wilhelmina, BRAss 9426, type). Ecol. Forming bright green cushions, c. 5- 15 cm in diam. on old camp site, 3650 m alt. Notes. Different from all other spp. by its dense mode of growth, its peculiar leaf-shape, and its umbels being 1—2-pedicelled. In mode of growth it comes nearest to 7. rosulans. 16. Trachymene caerulea GRAH. Edinb. New Phil. Journ. 5 (1828) 380; Buw. Blumea 2 (1936) 158. —Didiscus caeruleus Hook. in Curt. Bot. Mag. 55 (1828) t. 2875.— D. cyaneus DC. Mém. Omb. 11 (1829) 28.—Huegelia caerulea REICHENB. Iconogr. Exot. (1829) t. 20. Annual, pilose and glandular in nearly all parts. Main root fusiform with fibrous branches. Stems single, erect, upper portion with usually simple branches not overtopping the main stem. Lower /eaves petioled, petioles 1!/2-4 cm, hardly sheathing; lamina roundish in outline, ternate, leaflets bipinnatifid to bipinnatipartite, segments narrow with subacute to subobtuse apiculate tips; upper leaves sessile or subsessile, less divided, uppermost ones with only 3 narrow segments. Um- bels terminal on the main stem and the branches, co -flowered; involucres oo, linear, nearly filiform towards the tip, nearly as long as the flowers; pedicels 10-25 mm, inner ones gradually shorter, spreading, more erect in fruit. Outermost flowers larger than the other ones, not fruit-bearing, prob- ably male. Calyx teeth very short, subulate. Petals 23/4-3 by 2—21/4 mm, ovate to obovate, shortly un- guiculate at the base, outside with short glandular hairs. Styles nearly 1 mm; ovary glandular hairy. Mericarps 3'/4 by up to 23/4 mm, roughly tuber- culate with glandular hairs. Distr. Australia, in Malaysia: cultivated as a garden plant. 4. SANICULA LINNE, Sp.Pl. 1 (1753) 235; Buw. Blumea 2 (1936) 159 (Jit.). Erect herbs; /eaves palmately 3—5-partite, segments dentate, lobed or pinnately dissected. Flowers in irregular compound umbels with few rays; involucres leaf- like; umbellules usually small, with small involucels. Calyx teeth subherbaceous or membranaceous. Petals white, emarginate with inflexed tip, slightly imbricate. Disk flat with raised margin encircling the styles. Styles from the base filiform or subincrassate. Fruits echinate, ovoid, subterete or laterally subflattened; com- missure broad; mericarps with obscure ribs, lateral ones in the commissure, inner surface flat; vittae slender, solitary in each ridge, some very slender scattered in the endosperm. Distr. About 40 spp. (depending on specific delimitation) distributed throughout the world and in Hawaii and Patagonia, but not in Australia and New Zealand. 1. Sanicula europaea LINNE, Sp.Pl. 1 (1753) 235; Buw. Blumea 2 (1936) 159 (/it.).—S. elata D. Don, Prod. Fl. Nep. (1825) 183.—S. javanica BL. Bijdr. 15 (1826) 882.—S. montana BL. Bijdr. 15 (1826) 882; MOLKENB. in Mia. Pl. Jungh. (1851) 93, cum var. genuina, javanica, divaricata.—S. montana var. genuina, var. javanica ZOLL. Syst. Verz. (1854) 138. —S. elata var. normalis, var. partita O.K. Rey. Gen. Pl. 1 (1891) 269.—S. europaea var. javanica WoLFF in Pfl.R. 61 (1913) 64. Perennial with more or less creeping rhizomes. Stems 15-75 cm, slender, deeply grooved, glabrou 126 FLORA MALESIANA [ser. I, vol. 42 or rarely like the whole plant hairy. Petiole of the lower leaves 3—20 cm, lamina tripartite to ternate, segments incised and serrate-crenulate, teeth mu- cronulate. Umbels in a dichasium, ending in mono- chasia, sessile or on peduncles up to 1!/2 cm, with 5-8 involucres, 4—6 flowered, with 2—3 outer male flowers on pedicels 1/2-1 mm, and 2-4 female flow- ers, sessile or on pedicels up to 1/2 mm. Calyx teeth 11!/2-1 by 1/4 mm, oblong, acute. Petals nearly 11/4 by !/2 mm, with inflexed tip. Mericarps nearly 2 by 1 mm, densely covered with about 1'/2 mm long uncinate bristles. Distr. Temperate and tropical parts of Europe, Africa and Asia; in Malaysia: all over the Archi- pelago eastwards to Ceram. Ecol. Mountain forests, 500-3060 m. Vern. Daun ketapan, kundje, tespong, tetespon- gan, S; Javanese names rather variable. Notes. The Malaysian material is rather uni- form; it differs from the European form by its um- bels arranged in widely branched di-monochasia, whereas the latter has more crowded inflorescences and consequently nearly compound umbels. 5. ERYNGIUM LINNE, Sp.PI. 1 (1753) 232; Buw. Blumea 2 (1936) 164 (Jit.). Erect herbs, often spinescent. Leaves spiny dentate, entire, lobed or dissected. Flowers in heads or compact spikes, all bracteolate. Calyx teeth rigid, acute or prickly. Petals erect, white, with inflexed tip, scarcely imbricate. Disk flat, raised margin encircling the styles. Styles from the base filiform. Fruit ellipsoid, nearly cylindrical, commissure broad; mericarps with subprominent ribs, inner surface subconcave; vittae inconspicuous or 0; carpophore deciduous. Distr. Over 200 spp. described, distributed throughout the world with exception of trop. & S. Africa, centering in the New World specially Mexico, in the tropics usually on the mountains, E. rostratum cir- cum-S.Pacific. KEY TO THE SPECIES 1. Radical leaves simple, with a dentate margin 1. Radical leaves distinctly 2-pinnately lobed AOA AES > OVA PFEL LE Gee == >. Fig. 5. Eryngium foetidum L., x 1/s. 1. E. foetidum 2. E. moluccanum 1. Eryngium foetidum LINNF, Sp.Pl. 1 (1753) 232. —Buw. Blumea 2 (1936) 164.—Fig. 5. Roots fusiform. Stems 15-60 cm, many times di- monochasially branched with spreading branches, subglabrous, grooved. Leaves nearly all in a rosette, 3-32 by 1-4 cm, glabrous, lanceolate- spathulate, obtuse, sessile, base more or less narrow- ly sheathy, margin dentate, teeth with a spiny hair. Bracts of the inflorescences 1-6 cm, palmatilobate to -partite, with spiny tips and teeth, strongly nerv- ed, lowermost often like normal leaves. Peduncles 1-10 mm; heads 5—10 mm, cylindrical; involucres 5-7, spreading, nearly lanceolate with few spiny teeth. Flowers sessile in the axils of narrow mem- branous-margined bracts 1!/4-1!/2 mm long. Calyx teeth nearly 3/4 mm, lanceolate, acute with narrow membranous margin. Petals !/2-3/4 by + 1/4 mm, greenish white. Mericarps 1-1'/2 by 1/23/4 mm, densely warty, glabrous, ribs indistinct. Distr. Indigenous in tropical America, intro- duced in some parts of tropical Africa and Asia; in Malaysia: Malay Peninsula, Sumatra, Java. Ecol. In not too dry regions, in shaded or sun- ny, fertile localities, arable lands and grasslands, sawah-dikes, forest edges and along streambanks. Uses. Raw or steamed eaten with rice. Vern. Kangkong kerbau, jeraju gunung (Mal. Pen.); umbu palembang, ketumbor djawa (Sum.); (rumput) walang, katuntjar walanda, katuntjar blanda walang in several combinations, S, tumbaran unga, ketumbar landa, ketul kebo, djinten, djintenan, J; stinkdistel (Dutch). Notes. In the Malay Peninsula for the first time collected in 1888; in Java observed as early as 1896 [EDELING, Nat. Tijd. Ned. Ind. 31 (1870) 294], in Sumatra in 1915. Dec. 1949] UMBELLIFERAE (Buwalda) 127 The plants smell of bugs caused by an aromatic oil containing the aldehyd dodecen-2-al-1 (KOoL- HAAS, Rec. trav. chim. Pays-Bas 51 (1932) 460). 2. Eryngium moluccanum 7. sp. prov. Habit of E. campestre. Leaves petioled, broad triangular-ovate in outline, 10-15 cm _ through, prickly. Stem little branched. Heads oblong. Distr. Malaysia: Moluccas (Ceram, summit zone of Mt Pinaja, ca 3000 malt., EYMA a. 1937/8). Notes. A provisional examination in 1940 of the rather few specimens of this most remarkable discovery by the late Dr EYMA gave as a result that it was a new species, probably allied to some Aus- tralian species. Unfortunately the specimens can- not be traced, and are probably lost. (v. ST.). 6. CHAEREFOLIUM HALL. Hist. Stirp. Helv. 1 (1768) 327; Buw. Blumea 2 (1936) 168.—Anthriscus Pers. Syn. 1 (1805) 320. 1. Chaerefolium cerefolium SCHINZ & THELL. Vier- teljahrsschr. Naturf. Gesell. Ziirich 53 (1909) 554; Buw. Blumea 2 (1936) 168.—Scandix cerefolium Linn£E, Sp.Pl. 1 (1753) 257.—Anthriscus cerefolium Horem. Gen. Pl. Umbell. (1814) 41. Herb, somewhat hirsute. Stems 25—50 cm, striate and grooved. Lower /eaves with petioles to 7 cm with sheathing base, upper leaves subsessile to ses- sile on the sheaths; lamina 4-11 by 3-15 cm, tri- angular in outline, bi- to tripinnate, primary leaf- lets ovate, obtuse, !/2-2!/2 cm petiolulate, second- ary leaflets ovate, pinnatipartite, with obtuse tips. Compound umbels sessile in a di-monochasium; rays 3—5, 5—25 mm; pedicels 4-9, 2-4 mm, to 5 mm in fruit; involucres absent; involucels 3—4, nearly 2 by 3/4 mm, lanceolate, acute with narrow mem- branous margin. Petals nearly 1—1'/2 by '/2-1 mm, white, obcordate with short inflexed tips. Mericarps 5-6 by 1 mm, sometimes antrorsely hirsute when unripe, black and finely granular when ripe, grooy- ed inside, with a beak to 2!/2 mm; disk flat. Distr. Indigenous in SE. Europe and W. Asia, cultivated and subspontanous in all parts of the world; in Malaysia stated to be cultivated in Java [Mia. Fl. Ind. Bat. I, 1 (1856) 744]. Notes. Description after European materials; no specimens from Malaysia seen by me. 7. TORILIS Apans. Fam. Pl. 2 (1763) 99; Buw. Blumea 2 (1936) 169.—Caucalis BENTH. & Hook. f. Gen. Pl. 1 (1867) 928, pro parte. Annual to perennial erect herbs, retrorsely pubescent. Leaves pinnately divided. Umbels compound, rays often few. Involucres many to 0, involucels many. Flowers white or reddish, outer ones radiating. Calyx teeth triangular, acute. Petals cuneate or obovate, emarginate, with inflexed tips. Disk continuous with the base of the styles. Fruits ovate to oblong, laterally subflattened, constricted at the broad com- missure; mericarps with primary and secondary ribs obscure or subprominent, lateral ribs in the commissure, muricate-setose, inner face sulcate; carpophore entire or 2-fid. 1. Torilis japonica DC. Prodr. 4 (1830) 219; Buw. Blumea 2 (1936) 169.—Tordylium anthriscus LYNNE, Sp.Pl. 1 (1753) 240.—Caucalis anthriscus Hubs. Fl. Angl. ed. 1 (1762) 99.—Caucalis japonica Houtt. Nat. Hist. II, 8 (1777) p. 42, t. 45, 1.— Torilis anthriscus (non GAERTN. 1788) GMEL. FI. Bad. 1 (1806) 615.—Torilis scabra (non DC. 1830) ZOLL. Syst. Verz. (1854) 139. Stems to more than 1 m, finely striate, rough by appressed bristles. Leaves triangular in outline, sparingly appressedly hirsute, pinnate, leaflets pin- natipartite, segments pinnatifid to serrate. Umbels terminal and axillary; peduncles 5—20 cm; rays 4— 12, '/2-3 cm; pedicels 4-10, 1-4 mm, all antrorsely hirsute; involucres 2-6; involucels 3-7, nearly fili- form. Calyx teeth nearly ‘/2 mm, triangular- lanceolate, mucronulate. Petals 1/2-1 mm through, appressedly hairy outside. Mericarps about 4 by 11/2 mm, oblong, ribs obtuse, grooves with densely placed uncinate bristles. Distr. Indigenous in Europe, N. Africa, tem- perate Asia, Himalayan Mts, introduced rho) asp Asia and America, in Malaysia: N. half of Suma- tra and E. Java (Mt Tengger). Ecol. Mountains, 1225-2500 m. Vern. Ambo-ambo (Sum.), tumbaran alas, J. Notes. This species is said to have been intro- duced in S. Asia and America among clover seed. 8. CORIANDRUM LinngE, Sp.Pl. 1 (1753) 256; Buw. Blumea 2 (1936) 170. 128 FLORA MALESIANA [ser. I, vol. 42 1. Coriandrum sativum LINNE, Sp-Pl. (1753) 256; Merr. Comm. Lour. (1935) 294; Buw. Blumea 2 (1936) 171.—Coriandrum testiculatum (non L.) Lour. Fl. Cochinch. (1790) 180.—Bifora loureirii Kostec. Allg. Med.-Pharm. FI. 4 (1835) 1183.— Atrema testiculatum Mia. FI. Ind. Bat. 1, 1 (1856) 744. Annual, entirely glabrous. Stems to 75 cm, te- rete, striate. Lower /eaves palmatilobate to -partite, middle leaves pinnate, segments gradually nar- rower, obtuse, upper leaves pinnate to bipinnate, segments !/2mm broad. Compound umbels ter- minal or seemingly lateral, peduncles 2-10 cm; trays 3-5, 1—2!/2 cm; pedicels 3-5 mm; involucres 0-1, to 5 mm; involucels 3—5, to 5 by 1/2 mm, linear. Calyx teeth nearly 1 mm, triangular-lanceolate to oblong-lanceolate, somewhat radiating. Petals white, radiating, outer ones 34 mm, all deeply bipartite with inflexed tips. Mericarps 4 by 2 mm, inside hollow, forming a nearly globose fruit; pri- mary ribs undulated lines, secondary ribs subprom- inent, filiformous. Distr. Indigenous from the Mediteranean re- gion to central Asia; cultivated in nearly all parts of the world, in Malaysia: cultivated from 0—2200 m. Uses. Fruit as spice; mixed with rice in prepar- ing yeast; leaves for flavouring foods; medicinal as a mild stimulant. Vern. Wansui, katuntjar, ketumbar, J, koriander (Dutch). Notes. Unripe fruits smell of bugs. The coriander seeds sold in the markets are stated to have been introduced from India. 9. OREOMYRRHIS ENDL. Gen. Pl. (1839) 787; Buw. Blumea 2 (1936) 172. Perennial herbs, often caespitose, often pubescent or villose. Leaves with sheaths, pinnately dissected, or entire, often all in rosettes. Umbels simple. Invol- ucres many. Calyx teeth inconspicuous. Petals entire, imbricate in bud. Disk con- tinuous with the base of the styles. Fruit oblong or narrow, slightly laterally flat- tened, usually tapering towards the end; commissure broad; mericarps subterete, ribs obtuse, prominent, lateral ones close to the commissure; vittae 1 under each groove and usually 2 at the commissure. Carpophore often bipartite. Distr. Few spp. circum-S.Pacific from Mexico to N. Borneo. KEY TO THE SPECIES 1. Leaves compound. 2. Leaves pinnate to bipinnate; primary leaflets in several pairs. Peduncles 0.7—-8 cm. Involucres 24 mm. Pedicels 1-9. Fruits 2!/2-31!/2 cm 1. O. andicola 2. Leaves subternate; lateral leaflets 3—5- -partite; ‘terminal leaflet ternate with tripartite segments. Pe- duncles 12—33 cm. Involucres 5-8 mm. Pedicels 15-30. Fruits 4-6 mm . 2. O. papuana 1. Leaves not compound, linear to narrowly spathulate. 3. Pedicels 6-8. Fruits 5 by c. 1—-1!/2 mm. Leaves 2-20 cm 3/4 mm. Leaves 3-4 mm 3. Pedicels solitary. Fruits c. 1'/2 by 1. Oreomyrrhis andicola Hook. f. Fl. Antarct. 2 (1844-47) p. 288, t. 101; Buw. Blumea 2 (1936) 173.—Mpyrrhis andicola KUNTH in Humps. & BonPL. Noy. Gen. & Sp. 5 (1821) p. 13, t. 419.— Caldasia andicola DC. Mém. Ombell. (1829) 60.— Oreomyrrhis colensoi Hoox. f. Fl. Nov. Zel. 1 (1853-55) 92.—O. haastii Hoox. f. Handb. N. Zeal. Fl. (1864) 91.— O. borneensis Merr. Am. Journ. Bot. 5 (1918) 515, ic. 36.—O. pumila RIDL. Trans. Linn. Soc. II, Bot. 9 (1916) 63. Small tufts to spreading soft-haired cushions. Main root fusiform. Caudex with few erect branches bearing one or more rosettes. Leaves 0.8-16 cm; sheath 3-30 by 1—3!/2mm, tapering into the petiole, margin membranaceous, dorsally glabrous or short-hirsute, inside short-hirsute or sparingly pilose, rather densely ciliate; petiole 0.3— 10 cm, canaliculate, glabrous or short-hirsute; la- mina 1/24 by 0.41.6 cm, triangular-ovate, pinnate to bipinnate, leaflets 5-11, lower ones with 3-5 secondary leaflets, all leaflets pinnatifid to pinna- 3. O. linearis 4. O. azorellacea tipartite, segments 1—2!/2 by 1/4—1/2 mm, thinly cori- aceous, glabrous or shortly hirsute, sometimes mu- cronulate, finely ciliate, margins sometimes re- curved; small leaflets only with acute, broad-trian- gular teeth. Umbels 1 to several in each rosette; peduncles 0.7-8 cm, terete, densely hairy with spreading somewhat silky hairs, or short hirsute towards the apex, sometimes glabrescent, some- times exceeding the leaves; involucres 5-10, 2— 4mm, ovate-lanceolate with broad base, outside densely sericeous or shortly hirsute, sometimes finely ciliate; pedicels 1-9, 0-1/2 mm, to 3 mm in fruit. Petals nearly 1 mm, broad elliptic-ovate, base shortly ciliate or glabrous, white or reddish. Fruits 21/2-31/2 by 3/4—11/4 by 3/4-11/2 mm, oblong ovate, somewhat incurved, densely short-hirsute to glabrous. Distr. Central and S. America from Mexico to - the Falkland Islands, moreover in Australia and New Zealand, in Malaysia: Br. N. Borneo (Kina- balu) and New Guinea. Dec. 1949] UMBELLIFERAE (Buwalda) 129 Ecol. Open places, rock crevices, alpine grass- land, open bogs, among shrubs, 3150-4240 m. Notes. Oreomyrrhis andicola isvery polymorph- ic. In the vast area which it covers it is glabrous to white-tomentose; the height varies from 1!/2- Fig. 6. Oreomyrrhis papuana Buw. (plant x 2/3, fruit x 4). 50 cm; the rosettes are dense or loose bearing sim- ple umbels or slightly branched stems with few leaves and several umbels arranged again nearly in an umbel; the leaves are bi- to tripinnate, rarely simply pinnate with pinnately divided leaflets; the petioles are longer or shorter than the lamina. 2. Oreomyrrhis papuana Buw. Blumea 2 (1936) 175, fig. 5.—Fig. 6. Perennial, main root fusiform. Caudex with few erect branches with rosettes. Leaves to 18 cm; - sheaths 1-4 by !/2 cm; tapering into the petioles, glabrous, margin scarcely membranaceous; peti- oles 4-12 cm, canaliculate, subglabrous, apical part subhirsute; lamina 11/2-2!/2 by 0.8-2 cm, rhom- boid-ovate in outline, subternate to pinnate, seg- ments 2-3-partite, ultimate segments to 7 by 1 mm, lanceolate-cuneate to linear-lanceolate, thick-cori- aceous, at the thick margin and beneath on the nerves with antrorse bristles. Umbels 1 or more in each rosette; peduncles 12-33 cm, terete, subsul- cate, towards the apex densely hirsute with small retrorsely appressed bristles; involucres 6-9, 5-8 by 1-2 mm, oblong-spathulate, broad at the base, texture and indumentum as the leaf segments, re- flexed in fruit; pedicels 15-30, very short when flowering, to 5 mm in fruit, inner ones shorter, scabrous with short retrorsely appressed bristles. Petals \—1'/4 by + 3/s mm, oblong-ovate or oblong- obovate. Fruits 4—6 by ca 3/4 by 11/4 mm, glabrous. Carpophore undivided. Distr. Malaysia: W. New Guinea (Central Range). Ecol. Open swampy, grassy localities, 3200— 3500 m. 3. Oreomyrrhis linearis HEMsL. in Hook. Ic. PI. 26 (1899) t. 2590; Buw. Blumea 2 (1936) 176. Perennial, caespitose. Roots with fibrous branches. Caudex with numerous erect branches, to 12 cm, beset with fibrous leaf rudiments, at the ex- tremities bearing dense or lax rosettes. Leaves 2- 20 cm; sheaths 5—35 by 1—4'/2 mm, tapering into the petiole, margin yellowish, 1/2 mm; petiole 3/4— 1 mm broad, gradually widening to the narrowly linear blade; lamina to 5 mm broad, glabrous or the upper surface retrorsely appressedly hirsute, apical portion on each side 1-4-dentate, margin thickened, in narrow leaves revolute, finely retror- sely ciliate; nervation pinnate, lateral nerves strong- ly ascending, hardly visible above, strongly promi- nent beneath. Peduncles 1 to several, 6-30 cm, erect or subcurved, rarely with one leaf, triangular, an- JNouk Fig. 7. Oreomyrrhis azorellacea Buw. a—b. mode of branching, x 1, c-d. flowering and fruiting twigs, x 4, e. fruit, x 8, f. mericarp in cross- section, x 24. 130 MALESIANA [ser. I, vol. 42 "4 £ ee: Fig. 8. Oreomyrrhis azorellacea BUW., a cushion plant of the alpine grassland on summit of Mt Albert Edward (Papua), + 4000m alt. (BRAss, ARCHBOLD expeditions) gles subincrassate, retrorsely hirsute towards the apex with appressed whitish bristles to !/2 mm, fin- ally glabrescent; involucres 5—6, 2—S by 1/2-1!1/4 mm, lingulate, obtuse, towards the connate base retror- sely hirsute; pedicels 6-8, to 11/2 mm, inner ones sessile, in fruit 2!/2-10 mm, densely retrorsely hir- sute. Petals 1.1-1.2 by 0.8 mm, violet, triangular. Fruits to 5 by nearly 1 by 1!/2 mm, slightly curved, entirely glabrous. Distr. Malaysia: SE. New Guinea. Ecol. Open sunny localities, open grasslands, grassy creek banks in open country, 3000-3900 m. 4. Oreomyrrhis azorellacea n. sp.—Fig. 7-8. Species nova glabrata suberecta, habitu maxime ut aliquis species generis Azorella. Faciliter distin- guenda caule 3—4'/2 cm longo, foliis 2—4-fariis, den- sissime imbricatis, 3-4 mm _ longis, inflorescentia uniflora, pedunculo 2 mm longo, bracteis involucratis 4, pedicello 1/2 mm longo, fructu \'/2mm longo et 3/4 mm lato. Forma foliorum valde ad O. linearis HEMSL. accedens tamen planta in toto distincte minor. A herb, probably perennial, entirely glabrous; stem nearly erect, 3—41/2 cm long, much branching from the base, branches up to 3 cm long, closely packed and rather stiff, the upper gradually shorter and forming a comparatively dense cushion. Leaves very densely imbricate in 2-4 rows, 3-4 mm long, the lower part amplexicaulous and sheathing; sheath appressed, up to 1 mm wide, slightly ciliate on the pellucid margin and gradually narrowing in the petiole; blade up to 11/2 mm long, up to 1 mm wide, stiffish, coriaceous, spreading, concave, l-nerved, ciliolate along the submembranaceous margin, tip blunt. Jnflorescences uniflorous, on top of short branches along the stem, hidden among the foliage during flowering and when in fruit. Peduncle 2 mm long, more or less flattened, bracts 4, forming an involucre, spreading, lanceolate, 11/2-2 mm long, !/2-1!/4 mm wide, connate at the base, suggesting leaves. Flowers on up to !/2mm long pedicels. Calyx tube shortly campanulate, laterally slightly compressed, c. 1mm long, 1!/2 mm wide, indistinctly furrowed; teeth absent. Petals red, 1 mm long, 1/2 mm wide, single-nerved, ellip- tical, acute. Stamens c. 1/2 mm long, anthers dor- sifix, rounded-ellipsoid, up to !/2 mm long and as wide. Styles 2, conical, up to !/2 mm long. Fruit c. 11/2 mm long, 3/4 mm wide, slightly narrowed near the top and slightly compressed laterally; rims ob- tuse, equally slightly prominent, beside the com- missures, joining near the top of the fruit; stylopo- dium conical, up to '/2 mm long; mericarp nearly terete. Distr. Malaysia: East New Guinea (Mt Albert Edward, Brass 4306, type). Ecol. Tiny vivid-green plant occurring in dense pin-cushion masses, common on alpine grasslands, 3680 m alt. Notes. In mode of growth and inflorescence it resembles fallaciously some spp. of Azorella. How- Dec. 1949] ever, it has a parenchymatic endocarp, the vittae are solitary in the furrows and 2 at the commissure; the endosperm is furrowed at the commissure on UMBELLIFERAE (Buwalda) 131 cross section. Therefore, it is an Oreomyrrhis. Its leaf shape comes very close to O. linearis, but it is much smaller in all parts. 10. CUMINUM LINNE, Sp.Pl. 1 (1753) 254; BisscHop-Grev. Plant. Ned. Ind. (1883) 204; Buw. Blumea 2 (1936) 178. 1. Cuminum cyminum LInné, Sp.PI. 1 (1753) 254; Buw. Blumea 2 (1936) 178. Annual. Stems 15-50 cm, erect, strongly diver- gently branched from the base, terete, striate, en- tirely glabrous. Leaves short-petioled or sessile on a sheath to 1 by !/2 cm with membranaceous white margins, auriculate at the apex or tapering into the petiole; lamina 3-10 cm, bipinnate, segments to 11/2 mm broad, linear. Compound umbels opposite to the leaves or terminal; peduncles 2-4 cm; rays 4-6, 1—-1'/2 cm; pedicels 3—7, 4-5 mm; involucres 3-5, 2—3!/2 cm, tripartite or twice tripartite, seg- ments filiform, sessile on a !/2 mm long sheath with membranaceous white margins; involucels 2-4, to 9mm, margin white-membranaceous. Calyx teeth 1-1!/2mm, linear to subulate, persistent. Petals nearly 1 by '/2 mm, white to reddish, obcor- date with inflexed tips. Mericarps 5—7 by nearly 3 mm, somewhat laterally flattened, main ribs fili- form, bristly, ridges with a stellate-hairy line. Distr. Indigenous in Turkestan, cultivated in all parts of the world; in Malaysia: stated to be cultivated in the mountains of Java (HEYNE, OCHSE & BAKH., /l.cc.). Uses. Oil distilled from the seeds for making liqueurs; seeds for seasoning curries; medicinal ex- ternally and internally as stomachic and astringent. Vern. Djinten putih, M, djinten bodas, S, djinten poté, Md, komijn (Dutch), cumin (Engl.), Kiimmel (Germ.). Notes. Description after plants from the Ori- ent; no specimens from Malaysia seen by me. The Cuminum sold in the markets is stated to have been introduced from India. 11. APIUM LINNE, Sp.Pl. 1 (1753) 264; Buw. Blumea 2 (1936) 179. KEY TO THE SPECIES 1. Leaves pinnate, with broad tripartite to trilobate petiolate leaflets 1. Leaves bi- to tripinnate, with very narrow or filiform segments 1. Apium grayeolens LINNE, Sp.Pl. 1 (1753) 264; Buw. Blumea 2 (1936) 179. Main root fusiform or tuberiform. Stems 25-90 cm, angular, striate and grooved. Petioles rather long; sheaths to 2 cm in the lower J/eaves, white-margined; lamina pinnate, leaflets 2—2!/2 by to 3 cm, trilobate to tripartite, petiolulate, in the upper leaves smaller, ternate to 3-partite. Com- pound umbels opposite the leaves; peduncles 0— 2 cm; rays 10-15, 1-3 cm; pedicels 6-10, 2-3 mm; involucres and involucels absent. Calyx teeth ab- sent. Petals 1/2 mm through, white or greenish, with inflexed tips. Mericarps 1 by up to 3/4 mm, ribs narrowly winged; stylopodium nearly 1/4 mm high, halves conical. Carpophore emarginate. Distr. Indigenous in the temperate parts of Europe, Africa and Asia, also in S. America, cul- tivated elsewhere, in Malaysia: cultivated from 1-2100 m. Uses. Leaves, petioles and tuberiform roots for flavouring dishes; seeds as spice and medicinal. Vern. Saladri, S, celery (Engl.), selderie (Dutch). 1. A. graveolens 2. A. tenuifolium 2. Apium tenuifolium THELL. in Heart, Ill. Fl. Mit- teleur. 5, 2 (1926) 1140; Buw. Blumea 2 (1936) 181.—Sison ammi (non L. 1753). Jaca. Hort. Vin- dob. (1773) t. 200 excl. syn. ex THELL. in HEGI L.c.—Cnidium tenuifolium MoENCH, Meth. (1794) 98, excl. syn.—Pimpinella leptophylla Pers. Syn. 1 (1805) 324.— Helosciadium leptophyllum DC. Mém. Soc. Phys. Genéve 4 (1828) 493.—Apium leptophyl- lum BENTH. FI. Austr. 3 (1866) 372.—Apium ammi Urs. in Mart. FI. Bras. 11, 1 (1879) 341, t. 91. Main root fusiform. Stems 40—50 cm, striate, nearly glabrous. Leaves bi- to tripinnate, segments 1/21 mm_ broad, nearly filiform. Compound umbels opposite the leaves; peduncles 0—2 cm; rays 3—5, 1/2-1 cm; pedicels 5-10, 24 mm; invol- ucres and involucels absent. Calyx teeth 0. Petals nearly 0.4-0.6 by 0.2 mm, with strongly inflexed tips, white. Mericarps + 11/2by'!/2mm, ribs obtusely keeled. Stylopodium bipartite, halves small, conical. Carpophore to + 1/7 from the apex bipartite. Distr. Central and S. America, Australia, New Zealand, cultivated and adventive in Europe and Asia, in Malaysia: subspontaneous, 700—750 m. 12. PETROSELINUM Hit, Brit. Herbal (1756) 424; Buw. Blumea 2 (1936) 191.—Carum sect. Petrose- linum BENTH. & Hook. f. Gen. Pl. 1 (1879) 891. 132 FLORA MALESIANA [ser. I, vol. 42 1. Petroselinum crispum (MILL.) NYMAN ex auctt. Kew in Handlist Herb. Pl. Kew ed. 3 (1925) 122.— Apium petroselinum LINNE, Sp.Pl. 1 (1753) 263.— Petroselinum vulgare HIL1, Brit. Herb. (1756) 424, ic. p. 60; Buw. Blumea 2 (1936) 182.—Apium crispum MiLL. Gard. Dict. ed. 8 (1768) no 2; SHaw, Kew Bull. (1938) 257; ibid. (1939) 168.— Petroselinum hortense HorrM. Gen. Pl. Umbell. (1814) 163, t. Il, 7—Petroselinum sativum HOFFM. op. cit. 177.—Carum petroselinum BENTH. & HOOK. f. Gen. Pl. 1 (1867) 891.—Petroselinum petroselinum Karst. Fl. Deutschl. 2 (1895) 394. Stems 25-100 cm, erect, grooved. Lower leaves to tripinnate, leaflets obovate to cuneate, tripartite; upper Jeaves ternate. Compound umbels terminal and axillary; peduncles 2-12 cm; rays 5-10, 1- 3 cm; pedicels 3-15, 2-5 mm; involucres 1-3, in- volucels 3-8. Calyx teeth absent. Petals nearly 1 by 1/2 mm, with inflexed tips, greenish yellow. Meri- carps 2-2'/2 by nearly 1 mm, ribs filiformous. Distr. Indigenous in S. Europe and N. Africa, cultivated and subspontaneous elsewhere, in Ma- laysia: cultivated up to 2000 m. Uses. Leaves for flavouring dishes, and as a diuretic. Vern. Potrasoli, M, parsley (Engl.), peterselie (Dutch). Notes. According to SHAW the names in HILL’s British Herbal are nomenclaturally not eligible since Hitt did not accept the binary system of nomenclature in this work. 13. TRACHYSPERMUM Link, Enum. Hort. Berol. 1 (1821) 267; Buw. Blumea 2 (1936) 183.—Carum sect. Trachyspermum BENTH. & Hook. f. Gen. Pl. 1 (1867) 891. KEY TO THE SPECIES 1. Leaves 2-3-pinnatisect, the ultimate segments of the lower leaves to 1 mm broad. Calyx teeth distinct. Fruit with broad, roundish, scale-like hairs . 1. T. ammi 2. Leaves 2-pinnatisect, ultimate segments of the lower leaves more than 2 mm broad. Calyx teeth obsolete. Fruit with narrow, obtuse, nipple-shaped hairs . 1. Trachyspermum ammi SprRAGUE, Kew. Bull. (1929) 228; Buw. Blumea 2 (1936) 183.—Carum Rumpu. Herb. Amb. 5, p. 270.—Amudium RUMPH. l.c.—Sison ammi_ LINNE, Sp.Pl. 1 (1753) 252.— Ammi copticum LINNE, Mantissa 1 (1767) 56.— Ligusticum ajouan Roxs. Hort. Beng. (1814) 21, nomen.—L. ajowan Roxs. Fl. Ind. ed. CAReEy, 2 (1832) 91.—Trachyspermum copticum LINK, Enum. Hort. Berol. 1 (1821) 267.—Ptychotis coptica DC. Mém. Soc. Phys. Genéve 4 (1828) 496.—P. ajowan DC. Mém. Soc. Phys. Genéve 4 (1828) 497.—Am- mi glaucifolium (non L.) Buico, FI. Filip. (1837) 213.—Daucus anisodorus BLCOo, op. cit. ed. 2 (1845) 150, ed. 3, 1 (1877) 269.—Carum copticum HIERN, FI. Trop. Afr. 3 (1871) 12. Stems 25-45 cm, striate, glabrous, usually strongly branched. Leaves 2-3-pinnate, ultimate segments to 1 mm broad, narrow-oblong. Com- pound umbels terminal or seemingly lateral; ped- uncles 1-61/2 cm; rays 5-9, 1/2-1 cm, to 2cm in fruit; pedicels 4-15, 1-6 mm; involucres 3—5, ob- long, sometimes divided; involucels 4-5, oblong; the bracts of both very unequal in length, hirsute with membranaceous margin. Calyx teeth nearly 0.2mm, thickly subulate. Peta/s 0.6-0.7 mm through, obcordate with inflexed tips. Fruits to 2 by 1 mm, along the ribs with broad scale-like hairs. Distr. Indigenous and cultivated in Egypt, Abyssinia, SW. Asia to E. India, subspontaneous in Europe, in Malaysia: stated to be formerly cul- tivated in Java (HEYNE /.c.). Uses. Seeds medicinal as a carminative and in plasters; their medicinal agent is thymol. The seeds sold in the native drugstores are stated to have been introduced from India. 2. T. roxburghianum Vern. Mungsi, M, mosé, Md, musi (Bali). Notes. Description after materials cultivated by HEyYneE in his garden. 2. Trachyspermum roxburghianum Crars, FI. Siam. Enum. 1 (1931) 788; Wo Fr, Pflanzenr. 90 (1927) 129, nom. altern.; Buw. Blumea 2 (1936) 184.—Apium involucratum RoOxB. ex FLEM. Ind. Med. Pl. in As. Research. 11 (1810) 157.—Ptycho- tis roxburghiana DC. Prod. 4 (1830) 109.—P. in- volucrata ROYLE, Ill. Bot. Himal. 1 (1839) 229.— Carum roxburghianum Kurz, J. As. Soc. Beng. 46, II (1877) 114.—Carum involucratum MerRR. En. Philip. Fl. Pl. 3 (1923) 239.—Trachyspermum in- volucratum (non Marre, 1922) WoLFF in Pflanzenr. 90 (1927) 89. Stems 15-90 cm, striate, subglabrous, usually strongly branched. Leaves pinnate; leaflets pinna- tifid to pinnatipartite, extreme segments to 3 mm broad, those of the upper leaves gradually nar- rower to nearly filiform. Compound umbels terminal and axillary; peduncles 2-8 cm; rays 2-6, 1—2!/2 cm; pedicels 5-15, 2-6 mm; involucres 2-5; involucels 5-8; both very narrow, finely ciliate. Calyx teeth hardly 0.1 mm. Petals nearly 11/4 by 3/s mm, obcordate with inflexed tips, white or greenish white. Mericarps nearly 2!/2 by 3/4 mm, oblong, with very short obtuse spreading hairs. Distr. Of unknown provenance, now cultivated and subspontaneous in tropical SE. Asia, in Ma- laysia: throughout the Archipelago. Uses. Raw or steamed eaten with rice; also for flavouring dishes. Vern. Renggirung, adas, djentoa (Sum.), suragé, S, pletikapu, J, sorowai (Halmaheira). Dec. 1949] UMBELLIFERAE (Buwalda) 14. CRYPTOTAENIA DC. Mém. Ombel. (1829) 42; Buw. Blumea 2 (1936) 185. 1. Cryptotaenia canadensis DC. Prod. 4 (1830) 119; Buw. Blumea 2 (1936) 185.—Sison canadense LInng£, Sp.PI. 1 (1753) 252.—Cryptotaenia japonica HAssk. Retzia 1 (1855) 113. Rhizome 1-2 by to 1 cm, chambered. Stems erect, to 90 cm, terete, striate. Petioles to 10 cm, upper ones gradually shorter; sheaths with mem- branous margins, apex auriculate; lamina ternate, leaflets sessile or short-petiolulate, ovate to rhom- boid, irregularly biserrate to bidentate, lateral ones often bifid to bipartite. Compound umbels terminal on the stems and the branches, united to leafy panicles; peduncles 1-8 cm; rays 5-7, 3-50 cm, those of one umbel very different in length; pedi- cels 6-10, 11/2-15 mm, those of one umbellule very different in length; involucres 0-2, to 4 mm, subul” ate; involucels 2—5, to 1 mm, subulate. Calyx teeth absent, short in fruit. Petals nearly 1 by !/2-3/4 mm, obcordate with inflexed tips, white. Mericarps 4-6 by 1!/2 mm, oblong-ellipsoidal, attenuate at both ends, somewhat laterally flattened, distinctly rib- bed; stylopodium conical, bipartite, the halves together with the styles forming nearly 3/4 mm long beaks. Distr. Indigenous in eastern N. America, China and Japan, in Malaysia: stated to be cultivated by the Japanese (OCHSE & BAKH., /.c.). Vern. Salderi djepang, M. Notes. Description after materials cultivated in the Bot. Gardens of Buitenzorg and Groningen. 15. CARUM EINNE, Sp.Pl. 1 (1753) 263; 1. Carum carvi LINNE, Sp.Pl. 1 (1753) 263; Buw. Blumea 2 (1936) 186. Stems to 55 cm, erect, terete, striate. Petioles to 13 cm, upper ones gradually shorter, uppermost ones absent, all of them with a sheath with mem- branous margin and auriculate apex; lamina to 13 by 5cm, oblong, bipinnate, segments divided. Compound umbels terminal to the stems and its branches; peduncles 1-11 cm; rays 5-8, !/2-2cm; pedicels 6-14, 1!/2-5 mm, to 9 mm in fruit; invol- ucres none or 1, subulate; involucels none. Calyx teeth none. Petals to 11/4 by 1 mm, obcordate with short inflexed tips, white. Mericarps 4-5 by 1 mm, Buw. Biumea 2 (1936) 186.—Carum sect. Carvi BENTH. & Hook. f. Gen. Pl. 1 (1867) 890. often falcate, ribs distinct, yellowish. Stylopodium bipartite, halves low-conical. Distr. Indigenous in Europe, temperate Asia, cultivated elsewhere, in Malaysia: stated to be cul- tivated in the mountains of Java (MIQUEL, BU- WALDA, /.c.). Uses. Seeds in confectionery, also as medicine; oil from the seeds for making liqueur. Vern. Karwij (Dutch). Notes. Description after European materials, no Malaysian specimens seen by me. The seeds sold in native drugstores are stated to have been introduced from India. 16. PIMPINELLA LINNE, Sp.Pl. 1 (1753) 263; Buw. Blumea 2 (1936) 187.—Murrithia ZOLL. Nat. & Geneesk. Arch. 2 (1845) 576.—Heterachaena Zoi. Nat. & Geneesk. Arch. 2 (1845) 577.—Anisometros Hassk. Flora 30 (1847) 602. Annual to perennial herbs. Leaves 1-2-pinnate, 1—2-ternate or decompound, rarely undivided and only dentate. Umbels compound; involucres and involucels few or none. Calyx teeth obsolete or small. Petals usually emarginate, often with inflexed tips. Disk continuous with the base of the styles. Fruits ovate or broader than long, laterally flattened, usually constricted at the broad commissure; meri- carps terete to subpentagonal, often dorsally flattened, ribs slender, obscurely prominent, inner face flat; ridges with 2-3 vittae. Carpophore entire, 2-fid or 2-partite. Distr. Over 100 spp. described from Africa, Europe and continental Asia, in Malaysia: Java, Bali, ae zon: KEY TO THE SPECIES 1. Involucres 3 to more. Lower leaves usually imparipinnate. 2. Fruits densely warty RM see. «ts 2. Fruits hairy . 1. Involucres 1-2. Lower leaves mostly ean 3. Lower and middle leaves simple, with serrate margin, not lobed. Umbel rays 20-30 2. P. javana 3. Lower leaves orbicular to reniform, often crenate, sometimes lobate, middle leaves ternate to pinnate with incised leaflets. Umbel rays 8-14 1. P. anisum 3. P. pruatian 4. P. ascendens 134 FLORA MALESIANA [ser. I, vol. 4 \. Cee Fig. 9. Pimpinella javana DC. on Mt Tengger (E. Java), in open Casuarina junghuhniana forest. (DE VOOGD) 1. Pimpinella anisum LINNE, Sp.PIl. 1 (1753) 264; Buw. Blumea 2 (1936) 187. Annual. Stems erect, terete, grooved, pubescent. Petioles of the lower /eaves 4-10 cm, upper ones gradually shorter, uppermost leaves sessile, all with membranously margined sheaths, lower lami- na cordate, crenate to serrate, subsequent ones suc- cessively incised, ternate and nearly pinnate, leaflets dentate to incised. Umbels terminal to the stems and its branches; peduncles 2!/2-7 cm; rays 8-14, 4-25 mm; pedicels 7-13, 1-5 mm; involucres 0-2, 3-4 mm, narrow; involucels 0-2, 1 mm, subulate. Petals nearly 1 mm, obcordate with inflexed tips. Mericarps to 5 by 2 mm, ellipsoid, short-hairy by antrorse hairs. Distr. From an unknown provenance, probably from the Orient, cultivated and subspontaneous through the world, especially in the Mediterra- nean region and Central Europe, in Malaysia: stated to be sometimes cultivated in Java (Mia. FI. Ind. Bat. I, 1 (1856) 740; BisscHop-Grev. Plant. Ned. Ind. (1883) 204; Koorp. Exkurs. FI. Java, 2 (1912) 727; WIGMAN in VAN Gork. O.lI. Cult. 2 (1913) 883). Uses. Fruits carminative and as medicine. Notes. Description after European materials; no specimens from Malaysia seen by me. The fruits sold in native drugstores are stated to be from In- dia, which gets its supply from Persia. 2. Pimpinella javana DC. Prod. 4 (1830) 122; MOLKENB. in Mia. Pl. Jungh. (1851) 96, cum var. macrophylla, sylvestri, microphylla; Buw. Blumea 2 (1936) 188.—Murrithia cordata ZOLi. Nat. & Geneesk. Arch. 2 (1845) 576.—Fig. 9. Stems erect or ascendent, 50-150 cm, terete, striate, shortly and densely hairy, almost tomen- tose in the youth, later glabrescent. Lower /eaves nearly in a rosette; petioles to 10 cm, sheaths 3- 6 cm; lamina to 12 by 10 cm, entire, ovate in out- line, deeply cordate, subobtusely to acutely serrate; upper leaves gradually smaller and shorter peti- oled, more acutely serrate or even dentate, upper- most ones with branches in their axils, often tripar- tite, all of them more or less hairy above, white- tomentose beneath in the youth, glabrescent later. Umbels in an oblong panicle, terminal on the stems and its branches or seemingly opposite the ieaves; peduncles 4-15 cm; rays 20-30, 2-4 cm; pedicels 12-16, 3-8 mm; involucres 0-4, involucels 1|—4, nearly filiform, shorter than the outer pedicals. Petals nearly 11/2 by 1 mm with small inflexed tips. Mericarps nearly 2 by 1 mm, densely hairy with short spreading hairs. Distr. Malaysia: Java (from Mt Sindoro east- ward) and Lesser Sunda Islands (Bali). Ecol. Open or lightly forested localities, be- ‘tween 1200 and 3125 m. Dec. 1949] Vern. Glongong, gembogan, kemboan, pungan, J. Notes. P. javana is closely allied to the group of allied forms from SE. Asia described as P. candol- leana W. & A., P. leschenaultii CLARKE, P. pul- neyense GAMBLE, P. yunnanensis WOLFF, P. cam- bodgiana DE Botss., P. coriacea DE Boiss.; it differs only by its densely hairy fruits which in the above- mentioned species is more or less covered with scale-like papillae. In Bali I found in a moist locality, some umbels which had produced small roots; this shows the relative value of this character which was used in P. pruatjan for specific distinction (v. ST.). Sum- 3. Pimpinella pruatjan MOoLKENB. in Mia. PI. JUNGH. (1851) 97, cum var. depressa; Buw. Blumea 2 (1936) 191.—Heterachaena alpina ZOLL. Nat. & Geneesk. Arch. 2 (1845) 577.—Anisometros alpina HAssk. Flora 30 (1847) 602.—Pimpinella panatjan Murs. ex ROSENTH. Syn. Pl. Diaphor. (1862) 533. —Carum panatjan BAILL. Hist. Pl. 7 (1880) 178.— Pimpinella alpina (non Host, 1827); Koorp.- ScHum. Syst. Verz. I, 1, fam. 228 (1911) 98.—P. leeuwenii WOLFF in FEDDE, Repert. 20 (1924) 159. —P. pruatjan var. prolifera STEEN. Bull. Jard. Bot. Btzg III, 13 (1935) 349. Perennial. Stems several, 5-50 cm, ascending, sometimes spreading, rooting and forming roset- tes, terete, striate, puberulous when young, later glabrescent. Leaves mostly in rosettes; petioles to 10cm; sheaths to 3 cm; lamina imparipinnate, leaflets usually 5, rarely to 11 or only 1, 1—2!/2 cm, sessile or subsessile, roundly cordate, crenate-ser- rate to bicrenaté-serrate, or slightly lobed; upper leaves shorter petioled and smaller, leaflets less densely incised with narrower more acute segments; all sparingly hairy above, densely so beneath in the youth, later glabrescent. Umbels terminal to the stems and branches, often seemingly opposite the leaves; peduncles 1—7 cm; rays 4-8, 7-25 mm; pedi- cels 4-8, 1-4 mm; involucres 3-6, nearly filiform; UMBELLIFERAE (Buwalda) 135 involucels 3—6, nearly filiformous, shorter than the pedicels. Peta/s nearly 1|'/4 by 1 mm, with inflexed tips. Mericarps nearly 2 by | mm, warty. Distr. Malaysia: Java (from Mt Pangrango in W. Java to Mt Argapura (Jang) in E. Java). Ecol. Slightly shaded localities, on grasslands, in Casuarina-forests, along pathways and stream- banks, 1800-3300 m. Uses. Roots medicinal as a diuretic. Vern. Antanan kawat, antanan gunung, S, pur- wotjeng, tjumbuan, rumput demooh, J. 4. Pimpinella ascendens DALz. in Hook. J. Bot. & Kew Gard. Misc. 2 (1850) 261.—P. pinetorum Merr. Philip. J.Sci. 29 (1926) 482; Buw. Blumea 2 (1936) 192. Perennial herb. stems erect or ascendent, to 40 cm, terete, striate, shortly and densely white- hairy, later glabrescent. Leaves in a rosette at the base of the stems and few along the stems; petioles 2-10 cm, sheaths to 12 mm; lamina imparipinnate, leaflets 5-7, 6-15 by 6-18 mm, sessile or subses- sile, subcordate to suborbiculate, to truncate; cren- ate-serrate to bicrenate-serrate; upper leaves short- er petioled and smaller; all densely white-hairy above and beneath. Umbels terminal to the stems and its branches; peduncles 11-17 cm; rays 5-6, 11-30 mm; pedicels 4-10, 2-5 mm; involucres 1—2, 4-8 mm; involucels 1-4, 11/2-2 mm, shorter than the outer pedicels. Petals nearly 11/4 by 3/4 mm, with inflexed tips. Mericarps nearly 13/4 by 1 mm, sparsely hairy. Distr. Peninsular India, in Malaysia: Philip- pines (Luzon, Benguet, Mt Pulog). Ecol. Along trails in pine-forest, c. 2000 m alt., locally rather abundant. Notes. Philippine material is in the herbaria very scarce. According to MERRILL it is in all probability most closely allied to P. niitakayamensis HayaATaA of Formosa, from which it is at once dis- tinguishable by its indumentum. 17. OENANTHE LINNE, Sp.PI. 1 (1753) 254; Buw. Blumea 2 (1936) 194.—Sium sect. Drepanophyl- lum Bx. Biydr. 15 (1826) 881. Glabrous herbs. Roots fusiform or fasciculate-tuberculate. Leaves 1—3-pin- nate, ultimate segments large, linear or minute, rarely reduced to sheaths. Umbels compound. Involucres and involucels several, sometimes few or none. Calyx teeth small, acute. Petals emarginate, with long inflexed tips, white; outer ones of the inflorescence often enlarged. Disk continuous with the base of the styles. Fruits glabrous, ellipsoid or globose, nearly terete; commissure broad; mericarps dorsally flattened, inner face flat; lateral primary ribs broad, corky; dorsal and interme- diate ones often much smaller, sometimes obsolete, seldom all ribs subequal; ridges with | vitta. Carpophore 0. Distr. Some dozens of spp. (depending on specific delimitation), mostly in the N. hemisphere, also in S. Africa and trop. Australia. 136 FLORA MALESIANA [ser. I, vol. 42 1. Oenanthe javanica DC. Prod. 4 (1830) 138; Buw. Blumea 2 (1936) 194.—Sium javanicum BL. Bijdr. 15 (1826) 881.—Sium laciniatum Bu. Bijdr. 15 (1826) 881.—Falcaria javanica DC. Prod. 4 (1830) 110.—F. laciniata DC. Prod. 4 (1830) 110.— Oenanthe stolonifera DC. Prod. 4 (1830) 138.— O. linearis DC. Prod. 4 (1830) 138.—Dasyloma ben- ghalensis DC. Prod. 4 (1830) 140.—Phellandrium stoloniferum Roxs. Hort. Beng. (1814) 21, nomen. —Oenanthe laciniata ZOLL. Syst. Verz. (1854) 139. —Dasyloma javanicum Mia. Fl. Ind. Bat. I, 1 (1856) 741.—Dasyloma laciniatum Miq. FI. Ind. Bat. I, 1 (1856) 741.—Dasyloma japonicum Mia. Ann. Mus. Bot. Lugd. Bat. 3 (1867) 59.—Dasyloma subbipinnatum Mia. Ann. Mus. Lugd. Bat. 3 (1867) 59.—Oenanthe benghalensis BENTH. & Hook. /f. Gen. Pl. 1 (1867) 906.—Oenanthe thomsonii CLAR- KE in Hook. f. Fl. Br. Ind. 2 (1879) 697.—Oenanthe stolonifera var. javanica KUNTZE, Rey. Gen. Pl. 1 (1891) 269.—Oenanthe schlechteri WoOLLF in Scuum. & Laut. Nachtr. Fl. Deut. Schutzgeb. (1905) 333, t. 14.—O. rivularis DUNN, J. Linn. Soc. Bot. 35 (1907) 496. Perennial. Stems 10-100 cm, erect or ascending from a creeping base, terete, ramose. Petioles to 10 cm, often with sheaths; lamina pinnate to bipin- nate, segments ovate, serrate to narrowly oblong, or divided again, this making the leaf 4—5S-pinnate. Umbels terminal and opposite the leaves; peduncles 1-20 cm, rarely none; rays 5—15, 1/2-3 cm; pedicels 10—25, 2-5 mm; involuctes none or 1; involucels 2-8, 2-4 mm, linear. Calyx teeth nearly 1/2 mm, acute. Petals nearly 1 by 3/4 mm. Mericarps 2—3 by 1/2-1 mm, ribs swollen, marginal much more than the dorsal ones, the latter if strongly swollen nearly confluent. Distr. SE. and E. Asia, Formosa, Japan, Queensland, in Malaysia: all over the Archipelago. Ecol. Swampy places, along streams, wet grass- lands, and clearings, 1—2800m, but especially above 1000 m, sometimes cultivated. Uses. Vegetable, raw or steamed eaten with rice. Vern. Lampong, selemor, shelum (Mal. Pen.), batjarongi, piopo (Sum.), tespong, S, pambong, pam- pung, pangpung, matjen salade ajer, sladri gunung J, rukut teleme (Celebes). Notes. Very variable as to the dimensions of all its parts, the compoundness of its leaves, the -length of the peduncles, the number of pedicels in the umbellules, and the dimensions of the fruits. Many forms formerly described as distinct species; all are connected by intermediates. 18. FOENICULUM ADANS. Fam. Pl. 2 (1763) 101; Buw. Blumea 2 (1936) 200. 1. Foeniculum vulgare MILLER, Gard. Dict. ed. 8 (1768); Buw. Blumea 2 (1936) 200.—Anethum foe- niculum LINNE, Sp.Pl. 1 (1753) 263.—Foeniculum capillaceum Gi.is. Fl. Lithuan. Inchoat. coll. IV (1782) 40.—F. officinale ALLIONI, Fl. Pedem. 2 (1785) 25.—Ozodia foeniculacea W. & ARN. Prod. (1834) 375. Perennial, entirely glabrous. Stems erect, to 2m. Sheaths 4-12 cm in the lower /eaves, shorter upwards, apex with cucullate-connate auricles; lamina usually 3—4-pinnate, segments filiform. Compound umbels terminal to the stems and the branches; peduncles 5-16 cm; rays 30-70, 5—7 cm; pedicels 5—30, 1/2-1 cm; involucres and involucels none. Calyx teeth none. Petals yellow, strongly curved inward. Mericarps nearly 8 by 2 mm, ribs filiformous, nearly equal, not at all winged. Distr. Indigenous in the Mediterranean region, cultivated all over the world, in Malaysia: culti- vated throughout the Archipelago, subspontaneous on several mountains in E. Java, common on Mt Tengger (Sand Sea). Uses. Young leaves and fruits for flavouring dishes; medicinal for giving agreeable flavour to medicines; in European confectionery. The seeds sold in Java are stated to be introduced from India. Vern. Adas, adas manis, J, venkel (Dutch). 19. ANETHUM LINNE, Sp.Pl. 1 (1753) 263; Buw. Blumea 2 (1936) 202.—Peucedanum sect. Ane- thum BENTH. & Hook. f. Gen. Pl. 1 (1867) 919. 1. Anethum graveolens LINNE, Sp.PI. 1 (1753) 263; Buw. Blumea 2 (1936) 202.—Anethum sowa DC. Prod. 4 (1830) 186.—Peucedanum graveolens Hiern, FI. Trop. Afr. 3 (1871) 19. Annual. Stems 50-100 cm, terete, striate; sheaths 1!/2-2 cm in the lower /eaves, shorter up- wards, white-margined, apex with cucullate-con- nate auricles; lamina 3-pinnate, segments filiform. Compound umbels terminal to the stems and its branches: peduncles 4-13 cm; rays 5—15, 2—4 cm; pedicels 5—25, 1/2-1 cm; involucres and involucels- none. Calyx teeth ncne. Petals yellow, strongly curved inward. Mericarps nearly 5 by 3 mm, more- over with a wing !/4—!/2 mm, oblong. Distr. Indigenous in S. and SW. Asia, culti- vated in most parts of the world, in Malaysia: cul- tivated throughout the Archipelago from 10-— 1050 m. Uses. Raw or steamed eaten with rice; fruits for flavouring drinks, for native confectionery, in soups, sauces, etc. The fruits sold in Java are stated to be introduced from India. Vern. Adas (Sum.), walahandji (Sumba), djinten, adas sowa, M, dille (Dutch). Dec. 1949] UMBELLIFERAE (Buwalda) 137 > WNW, a rs e7 nee \ Ah J IE NSE. Th Weis I Pes,” AS? ey > » PER? AOR ZING TP L_ se\loer ? iy rr ie vd pip ae Re ER Se A — re OW AGS Z tN Se me / Zi ‘ f ace & Sess Wy \ Rag, Si { oF eyes) ‘ AK wee “ j AYER, . \ \ RHoeksema Fig. 10. Heracleum sumatranum Buw. (a—c. X 1/2, d-f. x 3). 138 FLORA MALESIANA [ser. I, vol. 42 20. PASTINACA LINNE, Sp.Pl. 1 (1753) 262; Buw. Blumea 2 (1936) 203. 1. Pastinaca sativa LINNE, Sp.Pl. 1 (1753) 262; Buw. Blumea 2 (1936) 203. Main root fusiform. Stems angular and strongly grooved. Leaves pinnate, leaflets 2-13 by 1-5 cm, oblong-ovate, often 3-lobate to 3-partite, irregularly crenate. Compound umbels terminal on the stems and its branches; peduncles 3—7 cm; rays 5—12, 1-4 cm; pedicels 10-20, 2-7 mm; involucres and involucels none or 1-2. Calyx teeth none. Petals yellow, with inflexed tips. Mericarps inclu- sive the 1/4—1/2 mm broad marginal wing 5—7 by 4-5 mm, broad-elliptic. Distr. Spontaneous in Europe and temperate Asia, elsewhere cultivated and subspontaneous; in Malaysia cultivated in W. Java, ca 900 m. Uses. Medicinal as a diuretic. Notes. Description after European materials; Malaysian specimens scanty. 21. HERACLEUM LINNE, Sp.Pl. 1 (1753) 249; Buw. Blumea 2 (1936) 204. Perennial or biennial herbs, seldom glabrous. Leaves broad-lobate to tripinnate, rarely ternately dissected, segments broad. Umbels compound with many rays. Involucres few or 0, simple, rarely many; involucels many, rarely divided. Flowers polygamous, often radiating, white or yellowish. Calyx teeth obsolete, seldom small, linear, lanceolate. Petals obovate, cuneate-rhomboid or unguiculate, emar- ginate to 2-fid with inflexed tips. Ovary hairy or pubescent. Fruits orbicular-obo- vate or elliptical, strongly dorsally flattened; dorsal and intermediate ribs thin- filiformous, lateral ribs usually with a broad wing; vittae usually solitary in each ridge, as long as the mericarps or abbreviated and dilatated below. Carpophore 2-partite. Distr. Over 70 spp. confined to the N. hemisphere. 1. Heracleum sumatranum Buw. Blumea 2 (1936) 204, fig. 6.—Fig. 10. Stems probably erect, striate or subsulcate, up- wards more or less hirsute, incrassate at the nodes; internodes 20-55 cm, upwards shorter. Leaves few, with small rosettes in the axils, sessile on a sheath. Sheaths 40 by 15 mm, amplexicaulous, margins membraneous, tips auriculate or narrowed. La- mina to 20 by 28 cm, deltoid in outline, biternate to ternate; terminal leaflet with a to 10cm long petiole, tripartite or ternate; Jateral ones with to 3!/2 cm long petioles; petioles of the secondary and tertiary leaflets gradually shorter to 0; leaflets ob- long-ovate, long-acuminate, all serrate to biserrate with short-acuminate apiculate teeth, beneath thin- hirsute on the nerves. Peduncles 10-20 cm; invol- ucres none; involucels 6—7 by circa !/2 mm, lance- olate, long-acuminate, somewhat hirsute with nar- row, membranous margin; rays 9-12, 11/2-3 cm, sulcate, shortly hirsute, to 4-6 cm in fruit, spread- ing, reflexed later; pedicels to 20, 2-5 cm, densely thin-hirsute, to 8-12 cm in fruit, spreading, later reflexed. Calyx teeth inconspicuous or to 1!/4 mm in the marginal flowers. Petals of the central flow- ers to 11/2 by 1 mm, elliptical to obovate, inflexed tip 1 mm; marginal flowers radiating, outer petals to 3 by 4mm, broad-obcordate. Mericarps 61/2-8 by 51!/2-7 mm, roundly ovate, glabrous, with a 2!/2mm broad wing; marginal ribs '/2 mm from the margin, vittae transversely septate. Distr. Malaysia: Central Sumatra (Mt Singa- lang). Ecol. Mountain forests, 2400 m. Notes. Closely allied to the group enumerated in the Fl. Br. Ind. from H. wallichii DC. to H. barmanicum Kurz. As these species show only slight differences, and H. sumatranum Buw. differs more from them than these species inter se, it is maintained as a different species; however, it might perhaps be better to unite them all to one poly- morphic species. 22. PEUCEDANUM LINNE, Sp.Pl. (1753) 244. Often robust herbs, root fusiform, sometimes tuberous. Leaves pinnate. Flowers bisexual, upper ones sometimes d, white or yellow, rarely purple. Compound umbels without involucres. Involucels present. Calyx rim abbreviate, shortly or distinctly toothed. Petals broad-ovate, with a long, inflexed tip. Stylopodium thick-conical, surrounded by the calyx rim. Fruit strongly dorsally compressed, narrow- to broad- elliptic, sometimes emarginate; marginal wings coherent, loosening when the meri- Dec. 1949] UMBELLIFERAE (Buwalda) 139 Fig. 11. Peucedanum japonicum Tuuns. Basal leaf and flowering stem, x 1/2, mericarps, x 5 (SIEBOLD H.L.B. 908. 260.495), root, < 1/2 (MAxIMovicz Iter 2, H.L.B. 908. 260.483). 140 FLORA MALESIANA [ser. I, vol. 42, Dec. 1949] carps areripe. Mericarps rather thin and slightly concave, with a distinct marginal wing in the base of which the marginal ribs are merging; dorsal side with 3 distinct ribs; vittae narrow, 1-3 between the ribs and 2-6 at the commissure. Carpophore split to the base, filiform. Distr. Large polymorphous genus, sometimes split into several genera formerly recognized as sections, about 200 spp., centering in the Orient, NE. Africa and W. North America, not or scarcely in S. America and absent in Australia, in Malaysia only known from the islands N of Luzon. 1. Peucedanum japonicum Tuuns. FI. Jap. (1784) in flower 1-2 and 4-5 mm, outer largest, in fruit 117; HayaTa, Ic. Pl. Form. 2 (1912) 57; MERR. hardening but not elongating. Flowers not radiant. Philip. J.Sc. 30 (1926) 418.—Fig. 11. Calyx indistinct. Petals ? white, c. 1!/4 by 1 mm, Taproot elongated fusiform, apex with erect emarginate through inflexed tip. Stamens inflexed remnants of sheath nerves 1 cm long; flowering in bud; filaments’°2 mm. Stylopodium blunt, very parts puberulous otherwise glabrous. Stem terete, _ thick, cap-like covering tip of ovate ovary and as grooved, slightly flexuose, erect, often branched, broad as it, margin crenate. Styles 2, exceedingly 1/2-3/4 cm diam., solid, 15-100 cm tall. Branches — short. Fruit partly abortive, 5-6 by 2!/2-3 mm, el- with distinct nodes, alternate, rather erect. Basal liptic, crowned by the stylopodium. Mericarps leaves in large specimens long-petioled with 3 bi- | dehiscing from the base, pendent from the filiform jugate, long-stalked, ternate segments, blade c.30-_ _carpophore halves, minutely puberulous to sub- 40 cm diam.; leaves of small specimens and flower- glabrous, marginal wing ca !/3—'/2 mm broad, ven- ing stems much smaller, 7-10 cm diam., biternate; _ tral side rather flat, through wing subconvex; body petiole 3-5 cm sheathing over its entire length, darkish, wings and ribs pale brown (description amplexicaulous at the base, striate; lateral seg- after Japan. specimens in Rijksherbarium). ments !/2-2 cm petiolulate, terminal ones 1!/2- Distr. Japan, Taiwan, in Malaysia: Philippines 4 cm, all 3-parted; leaflet-segments sessile, often (Batan Islands, N of Luzon: Sabtan Island, connate at the base, lateral ones oblique, all obo- © MERRILL 11755, not seen). vate-cuneate, margins entire, apex broadened 3-—5- Ecol. In crevices of cliffs along the seashore. toothed or -lobed, lobes often dentate, about equal Notes. The insertion of this species is wholly in size 21/2-4 by 1-3 cm; uppermost leaves reduced. on Dr MERRILL’s authority who collected and Inflorescense corymbiform. Compound umbels identified it. His identification was checked with terminal, 4-7 cm wide; peduncle 6—7 cm, in fruit the late Dr HAyATA while the late Dr JuEL com- to 10 cm, stout, striate, erect. Rays 15-25, unequal pared a fragment with THUNBERG’s type. The only in length, 1/2-2!/2 cm. Involucres 0. Secondary difference with the type was that the Philippine peduncles in flower 1-2, in fruit 2-3 cm long, har- _ specimen was glabrous. By Ind. Kew. it is wrongly dening. Involucels 7-10, subequal or very unequal, reduced to Ligusticum acutifolium, an error for lanceolate-oblong, acute, 2-6 mm. Pedicels 17-20, __L. acutilobium S. & Z. (v. ST.). 23. DAUCUS Linné, Sp.Pl. 1 (1753) 242; Buw. Blumea 2 (1936) 207. 1. Daucus carota LINNE, Sp.Pl. 1 (1753) 242; Buw. with inflexed tips, peripheric ones radiating. Meri- Blumea 2 (1936) 208. carps 3 by 11/2-2 mm, nearly oblong; primary ribs Annual, biennial or perennial. Main root fusi- filiform with rather few nearly 1/4 mm long, fine form. Stems erect, striate or grooved, hirsute. bristles, secondary ribs beset with nearly 1 mm Leaves 2-3-pinnatipartite, segments lanceolate. long, rigid bristles. Compound umbels with flat or round surface when Distr. Spontaneous in Europe, N. Africa, and flowering, with incurved peduncles and pedicels temperate Asia, cultivated in all parts of the world, and hollow surface in fruit; peduncles 2-25cm; in Malaysia: cultivated from 0-1800 m, subspon- rays 15-30, 1-6 cm; pedicels 20-30, !/2-1!/2cm; taneous on grassy mountain sides. involucres 3-5 cm, pinnatipartite, white-margined Uses. Roots as vegetable, young leaves raw or towards the base; involucels 5—7, 1/2-2 cm, entire steamed eaten with rice. to pinnatipartite, lanceolate. Calyx teeth 1'/4— Vern. Boktel, S, peen, wortelen (Dutch), carrot 1/2 mm, triangular, acute. Petals white or dark red (Engl.). in 5—7 central sterile flowers of the central umbel, Excluded & doubtful Conium maculatum L.; BorrRL. Handl. 1, 2 (1890) Hydrocotyle nuanavoides F.v.M. Bot. Centr. 616. This species has never been foundin Malaysia. BI. 1 (1892) 194, nomen. New Guinea. Hydrocotyle monopetala Buco, FI. Filip. (1837) Hydrocotyle villosa (non L. f.) KoorD. Teysman- 213. MERRILL places this with doubt in Ophiorrhiza nia 11 (1901) 252. Prob. a writing mistake; H. vil- (Rub.), cf. En. Philip. F1.Pl. 3 (1923) 502. losa L. f. is a native of S. Africa. FLORA MALESIANA UNDER THE AUSPICES OF THE KONINKLIJKE PLANTENTUIN VAN INDONESIE ‘ROYAL BOTANIC GARDENS OF INDONESIA’ BUITENZORG, JAVA SCIENTIFIC COMMUNICATIONS concerning Flora Malesiana must be addressed to the general editor Dr C. G. G. J. VAN STEENIS, c/o Roy. Botanic Gardens, Buitenzorg, Java NOTICE FOR SUBSCRIBERS The issue of this work in 1950 will consist only of vol. 1: ‘Cyclopaedia of Collectors, Collections’ &c. by Mrs M. J. VAN STEENIS- KRUSEMAN APPLICATIONS TO-BE DIRECTE Dike N.V. ERVEN P. NOORDHOFF NOORDHOFF-KOLFE N.Y. P.O. Box 39 P.O. Box 103 Groningen, Holland Batavia-C., Java and in the Americas to Dr FRANS VERDOORN Chronica Botanica Co. P.O. Box 151, Waltham 54 (Mass.), U.S.A. For sale only , Not for exchange INDEX to revised families in vol. 4 parts 1-2 Aceraceae SS ot es es 3 Nyssaceae . 4) Se Actinidiaceaes. str)5 (6°20 ° > BT? Philydraceae’....-3. tee 5 Amaranthaceae’ .%. . . . 69 . Plumbaginaceae = .. 9 2 a Ancistrocladaceae’.- > \. “x. 8 Podostemaceae .. -/ . Je age Aponogetonaceae. . °.. . 11. Sarcospermaceae . . . age Burmanniaceae i . .. 13. Saururaceae’ 55") ac) eee Ceratophyllaceae . -. . ~~. 41 Sphenocleaceae . %. ° ~) “Sees Chenopodiaceae . . . . 99 Stackhousiaceae . . . se Cochlospermaceaeé. . . .. 61, Styracaceae © . §. | ee Hydrocaryaceae .. . . .:+ 43. Trigoniaceae . . . See Juncaginaceae pee og STS Umbelliferae i’ a2 2 2 Moringaceae =. 1» .. |. “45 Zygophyllaceac << °° aeeaeee PUBLISHED BY NOORDHOFF-KOLFF N.V./ BATAVIA PRINTED IN THE NETHERLANDS BY JOH. ENSCHEDE EN ZONEN » HAARLEM JAN 41952 Pi) Rey mA oOLAN A DILLENIACEAE (R. D. Hoogland, Leyden) Trees, shrubs, lianas or perennial herbs. Leaves spirally arranged, opposite in one species only (Madagascar). Blade simple or, rarely, (only in Acrotrema) to threefold pinnatisect. Stipules absent, but in Acrotrema and a number of species of Dillenia petiole with stipule-like, often wholly or partly caducous wings. Inflorescence cymose or racemose, sometimes reduced to a single flower, terminal or axillary. Flowers 3, actinomorphic to (mainly in the androecium) zygo- morphic, hypogynous, mostly yellow or white. Sepals (3—) 4-5 (—20), imbricate, persistent in fruit. Petals (2—) 3-5 (—7), caducous usually within half a day after opening of the flower, imbricate in bud, all equal, apex rounded or emarginate. Stamens ~-3, often partly staminodial, free or partly coherent by their filaments, centrifugal. Anthercells basifix, oblong to linear, opening with an apical pore or a longitudinal slit. Carpels 1-+20, free or connate along the central axis only, with free styles. Ovules ~-—l, anatropous, apotropous, on an axile placenta. Fruit dehiscent or indehiscent, in the latter case permanently enclosed by the sepals. Seeds arillate or with a rudimentary aril, with abundant endosperm and a minute, straight embryo. Distr. Ca 10 genera, of which one circumtropical (Tetracera), 3 confined to tropical S. America, one in the Old World tropics from Madagascar to the Fiji Islands (Di/lenia), one endemic in Ceylon (Schumacheria), one in S. India, Ceylon, and the Malay Peninsula (Acrotrema, fig. 5), one endemic in Borneo (Didesmandra), one endemic in Australia (Pachynema), and one on the southern hemisphere from Madagascar to the Fiji Islands, mainly in Australia (Hibbertia, fig. 3). Many species are relatively limited in distribution, none is distributed throughout Malaysia. Ecol. Most Malaysian species occur in primary or secondary evergreen forests. A few deciduous species of Dillenia are found in monsoon forests or in savannahs, some Tetraceras in open vegetation and thickets and both species of Hibbertia in open scrub. In size Hibbertia shows the biggest development among the genera with over 100 spp., next in sequence are Dillenia with ca 55 spp. and Tetracera with over 30 spp. Most representatives are found in the everwet parts of the tropics, but the ability of the family to produce drought-resistant forms is distinctly shown by various xerophilous species of Hibbertia and Pachynema, showing remarkable adaptations and reduc- tions in foliage and habit. As to altitudinal distribution the tropical representatives are generally confined to the tropical zone proper, i.e. below 1000 m. Exceptions are Hibbertia scandens, found both at low altitude in the Aru Islands and at ca 2300 m on Mt Arfak, and some species of Dillenia, which may occasionally be found up to 1800 m. Notes. A more extensive treatment of Dillenia, including all extra-Malaysian species, will be pub- lished in Blumea vol. 7, a similar treatment of Tetracera for Asia, Malaysia, Australia, and the Pacific will be published in Reinwardtia vol. 1. Latin diagnoses of new taxa will be given there. Additional species of Hibbertia, and possibly also of the other Australian genus Pachynema, can be expected to occur in favorable localities in the extreme Eastern part of Malaysia. For a proper identification of Dilleniaceae it is of primary importance to have flowering material, fruits is less important. Sterile material is almost worthless. KEY TO THE GENERA 1. Receptacle flat. Carpels completely free or loosely coherent along their adaxial side. 2. Stamens , free. 3. Anthercells parallel, connective linear. 4. Small perennial herbs. Petiole with amplexicaul wings : :, »| <3; Acrotrenia 4. Small shrubs or climbers. Leaves clasping, but not completely amplexicanl . . 2. Hibbertia 3. Anthercells diverging towards the base, connective broadened Sone Jae 1. Tetracera 2. Androecium consisting of 2 groups, each of 1 stamen and 4 staminodes with coherent filaments 4. Didesmandra 1. Receptacle between the carpels conical. Carpels distinctly coherent along their adaxial side 5. Dillenia 1. TETRACERA LInngE, Sp. Pl. (1753) 533; Gen. Pl. ed. 5 (1754) 237.—Delima LINNE, Gen. PI. ed. 5 (1754) 231.—Assa Houtt. Nat. Hist. 5 (1776) 275.—Wahlbomia THUNB. Vet. 142 FLORA MALESIANA [ser. I, vol. 4 Akad. Handl. Stockh. (1790) 215.—Eleiastis RAFIN. Sylva Tellur. (1838) 165.— Delimopsis Miq. Fl. Ind. Bat. 1, 2 (1859) 9. Shrubs, sometimes straggling, or lianas. Leaves simple, often scabrid on one or both sides. Petiole short, furrowed. Panicles axillary or terminal, few- to many- flowered. Flowers actinomorphic, fragrant. Sepals 4-5, persistent, often reflexed when in fruit. Petals 3-5, caducous, with emarginate apex, whitish or slightly red- dish. Stamens «, with broadened connective, thereby anthercells divergent towards the base. Carpels 1-4, free, with a short style, ending in a simple stigma, with 4-20 ovules. Capsule coriaceous, opening with 1—2 longitudinal slits, ovoid with a short beak, one- to few-seeded. Seed glossy dark brown to black. Aril fleshy, cup-shaped, reddish or purplish, enveloping at least the base of the seed, equal- or unequal-sided with fimbriate or laciniate apical margin. Distr. The only pantropical genus in the family, absent, however, from the Pacific area except New Caledonia. The Malaysian species are confined to the Asiatic-Australian region. There are two local endemics, viz T. maingayi HOOGL. in the Malay Peninsula and 7. /Januginosa DiELS in New Guinea. The other Malaysian species have rather large areas. Ecol. Lianas in rain-forests, on forest margins and in hedges; some species occur as shrubs in open country. Usually below 500 m alt., rarely up to 1300 m. None of the Malaysian spp. shows a distinct flowering-time. Vern. The Malay name ampalas (ampélas, émpélas, mémpélas, mumplas) is in use throughout W. Malaysia for the whole genus, some species being distinguished by epithets. Akar = root or liana is also found in a number of combinations. Uses. The scabrid leaves of some species are used as a substitute for sandpaper; the Malay name for sandpaper has been derived from the plantname: mémpélas. The stems can be used as cordage. Notes. The genus Delima L., still kept apart by RmpLey (FI. Mal. Pen. 1, 1922, 3), is included here in Tetracera, as has been done already by VAHL (Symb. Bot. 3, 1794, 70). GILG & WERDERMANN (in ENGL. & Pr. Nat. Pfl. Fam. 2nd ed., 21, 1925, 18) retain Delima as a section of Tetracera. It should be noted, however, that the differences between Delima and Eutetracera are only found in the number of carpels, being 2—6 in Eutetracera, 1 in Delima, usually however on the same plant in a number of flowers 2. Of the 4 Malaysian species, belonging to the section Delima, 2 show close relationships to species belonging to the section Eutetracera, viz T. glaberrima MARTELLI to T. akara (BURM. f.) MERR. and T. maingayi Hooct. to T. fagifolia BL. Delima can be retained as a section on practical grounds only; it does not form a natural taxon. KEY TO THE SPECIES 1. Carpels 1, in a few flowers on the same plant 2 (section Delima). 2. Carpels and fruit hirsute . . o @ ow w cep o os 2) 18% 2Eescandens 2. Carpels and fruit glabrous or with minute scales. 3. Inflorescence up to 5-flowered, usually axillary. Flower ca 2'/2 cm diam. Sepals4 . 2. T. glaberrima 3. Inflorescence at least 15-flowered, terminal. Flower ca 1—1!/2 cm diam. Sepals 5. Are SepalseelabnOuUswMSides me Vaux cn Team eG Dg che 0. -s/= ce, See 3. T. asiatica Ass SepalsascniceOusmmside . Ges “So 8 ee eee a i ee i A (5 Bee ae 1. Carpels 24 (section Eutetracera). : 5. Carpels and fruit hairy over the whole surface. 6. Indumentum of the carpels consisting of rather thin villose hairs, caducous. Species from W. Malaysia . . . . . 12. T. arborescens 6. Indumentum of the carpels consisting of rather rigid, persistent, straight hairs. Species from E. Malaysia. 7. Hairs of the carpels ca 2 mm long. Inflorescence 24-flowered . . . . . 5. T. lanuginosa 7. Hairs of the carpels ca 1/2 mm long. Inflorescence 15—50-flowered . . . . 6. T. nordtiana 5. Carpels and fruit glabrous or with minute scales or with few rigid hairs on the back only. 8. Sepals 4. Inflorescence up to 12-flowered, terminal or axillary, without leaves in the basal part. Flower ca 2!/2-3 cm diam. ), Sepals glabrous inside ee es ck Be he pe Re a) eee 9. Sepals sericeous inside . . st) Sdicrakaral 8. Sepals 5—6. Inflorescence at least 15- flowered, rarely less- flowered, terminal, often with small leaves in the basal part, or axillary, but then always on a few-leaved branch. Flower ca 1-2'/2 cm diam. 10. Sepals glabrous inside. 11. Branches of the inflorescence strigose, without stellate groups of shorter hairs. 9. T. loureiri 11. Branches of the inflorescence with single strigose to hirsute hairs, together with stellate groups Oleshortersnalnsae Renae Rares Boel tee ee) EG ORNs) rss CMs a 10. T. korthalsii Dec. 1951] DILLENIACEAE (Hoogland) 143 10. Sepals, at least the 3 inner ones, sparsely to densely sericeous inside. 12. Two outer sepals glabrous inside . 12. All sepals sericeous inside. 11. T. macrophylla 13. Younger branches villose. Leaves under the inflorescence small (ca 4 by 3. cm), obovate. 12. T. arborescens 13. Younger branches strigose. Leaves under the inflorescence larger, elliptic to oblong. 1. Tetracera scandens (L.) Merr. Int. Rumph. (1917) 365; BAck. Bekn. Fl. Java em. ed. 4 (1942) fam. 80, 2.—Funis urens aspera RUMPH. Herb. Amb. 5 (1747) 13, t. 9.—Tragia scandens L. in StickM. Herb. Amb. (1754) 18; Amoen. Acad. 4 (1759) 128.—Delima sarmentosa L. Syst. ed. 10 (1759) 1076; Mia. Fl. Ind. Bat. 1, 2 (1859) 7; ViL- LAR, Nov. App. (1880) 2; Rip. Fl. Mal. Pen. 1 (1922) 3.—Tetracera sarmentosa VAHL, Symb. Bot. 3 (1794) 70; Roxs. FI. Ind. ed. CAREy 2 (1832) 645, p.p.; BLANCO, FI. Filip. ed. 2 (1845) 320; op. cit. ed. 3, 2 (1878) 227; Merr. Govt Lab. Publ. Philip. 27 (1905) 15.—Delima hebecarpa DC. Syst. 1 (1818) 407; De ess. Ic. Sel. Pl. 1 (1821) t. 72.—Delima intermedia Bu. Bijdr. 1 (1825) 4; Hassx. Pl. Jav. Rar. (1848) 176.—Delima frangulaefolia PRESL, Rel. Haenk. 2 (1835-6) 73; VILLAR, Nov. App. (1880) 2.—Delima aspera BLANCO, FI. Filip. (1837) 429; op. cit. ed. 2 (1845) 299; op. cit. ed. 3, 2 (1878) 191, t. 190.—Tetracera monocarpa BLANCO, FI. Filip. (1837) 459.—Delima sarmentosa var. hebe- carpa HK. f. & Tu. FI. Ind. 1 (1855) 61; Mia. FI. Ind. Bat. Suppl. 1 (1860) 618; Mia. Ann. Mus. Bot. Lugd. Bat. 4 (1868) 73; Hx. f. & Tu. FI. Br. Ind. 1 (1872) 31; Kina, J. As. Soc. Beng. 58, 2 (1889) 362.—Delima sarmentosa var. Mia. Fl. Ind. Bat. 1, 2 (1859) 7.—Tetracera sarmentosa var. hebecarpa MARTELLI in Becc. Malesia 3 (1886) 150; VIDAL, Rey. Pl. Vasc. Filip. (1886) 36.—Tetracera hebe- carpa BOERL. Cat. Hort. Bog. (1899) 3; Back. FI. Batavia 1 (1907) 4; Schoolfl. Java (1911) 8; Koorpb. Exk. Fl. Java 2 (1912) 600.—Tetracera volubilis Merr. Spec. Blanc. (1918) 362, sphalm., non L.— Tetracera scandens var. hebecarpa HEYNE, Nutt. Pl. (1927) 1070.—Delima scandens BuRK. Dict. (1935) 776. Liana (up to 30 m) or small shrub (up to 2 m). Leaves oblong to obovate, ca 6-15 by 3-7 cm, scabrid, apex and base rounded to obtuse. Petiole ca 6-12 mm. Inflorescence terminal, many-(up to ca 200)-flowered, with in the basal part 1—5 leaves, up to ca 40 by 20cm. Flower ca 6-8 mm diam. Sepals 4, on the same plant in some (ca 5%) of the flowers 5, ca 3 by 2 mm, scabrid outside. Petals 3. Stamens 3mm long; anthercells reaching each other at the apex. Carpels with 0.4-0.7 mm long rigid hairs; ovules ca 10. Capsule ovoid, ca 10 by 6 mm, 1(—2)-seeded. Seeds 4 by 3mm. Aril 2-3 mm long, fimbriate for 3/4/10 of its length. Distr. S. China (Yunnan), Burma, S. Siam, S. Indo-China, Andaman Islands, in Malaysia: throughout W. Malaysia, not E of the Philippines, Celebes and the Lesser Sunda Islands (Flores). Ecol. Creeper or climber in thickets and sec- ondary forests, especially on riverbanks and near the seacoast; in more open vegetation forming small shrubs. From sea-level up to 1000 m, rarely above 500 m. 13. T. fagifolia Vern. Sumatra: akar ampala (Priaman), a. (rem)pélas (Palemb.), baik sipi hendak (Lamp.), galinggin (Asahan), hasahan (Lamp.), mémpélas padang (Bengkalis), ompe (Atjeh). Malay Penin- sula: akar mémpélas hari bétina, a. m. tikus, a. m. puteh, a. pélah, ampélas hari (bétina), a. puteh, a. kasap, a. rimau, a. tikus. Java: (akar) mémpélas hari bétina, a. m. ojod, a. m. putih, a. m. tikus, kaju as(s)ahan, ki asahan (Mal.); areuj ki assahan lalaki, a. ki assahan, ki asahan, asahan areuj, kroko ojod (Sund.); bo, débo, démbo, kroko, ojal, roko, roko- kan, singaran (Jav.). Kangean: ampélas. Borneo: agupit (Bajau), akar ampalas, ampalas (Malay), kérub kérub (Bajau), panpan (Sungei). Philippines: dangilian (Bag.), malakatmon (Tag., Pamp.), mal- bastigbalang (Vag.), pakiling (Sbl.). Celebes: /um- piwi apaélaé. Bali: bun api-api. Uses. The leaves are used for polishing wood and metal. The stems may be used as cordage. Medical use is unimportant, cf. BURKILL, /.c. Notes. The glabrous-fruited form, described as Delima sarmentosa var. glabra by HK. f. & Tu. FI. Ind. 1 (1855) 61 is considered here as a separate species; cf. T. asiatica (LoUR.) HOOGL. 2. Tetracera glaberrima MARTELLI in Becc. Ma- lesia 3 (1886) 150. Scandent shrub. Branches glabrous. Leaves el- liptic to obovate, ca 5—9 by 2-4 cm, rather cori- aceous, bright glossy green above, not scabrid, with acuminate apex. Petiole 3-8 mm. Inflorescence ax- illary, 1—5S-, usually 3-flowered. Flower ca 2'/2 cm diam. Sepals ca 9 by 6 mm, glabrous to sparsely sericeous inside. Petals 4. Stamens 5 mm long; anthercells reaching each other at the apex. Carpels with ca 10 ovules. Capsule ovoid, ca 15 by 10 mm with a 2-3 mm long beak, 1—3-seeded. Seeds 3-4 by 2-3 mm. Aril 5 mm long, laciniate to about halfway its length. Distr. Malaysia: Borneo (Kuching); has been cultivated in the Botanic Gardens at Bogor. Notes. The species is most closely related to Tetracera akara (BuRM. f.) MERR., from which it differs by the single carpel, the less densely seri- ceous inside of the sepals, and the relatively broader leaves. 3. Tetracera asiatica (LOUR.) HOOGL. comb. nov.— Seguieria asiatica Lour. Fl. Coch. (1790) 341.— Delima sarmentosa (non L.) Burm. f. Fl. Ind. (1768) 122, t. 37; DC. Syst. 1 (1818) 407; Prod. (1824) 69; Trim. Handb. FI. Ceyl. 1 (1893) 5.—Tetracera sar- mentosa (non VAHL) Roxs. FI. Ind. ed. CAREY 2 (1832) 645, p.p.—Leontoglossum scabrum HANCE in Wacp. Ann. 2 (1851) 18.—Delima sarmentosa var. glabra Hk. f. & Tu. FI. Ind. 1 (1855) 61; Hx. f. & Tu. FI. Br. Ind. 1 (1872) 31.—Delimopsis hirsuta Mia. FI. Ind. Bat. 1, 2 (1859) 10; Suppl. 1 (1860) 144 FLORA MALESIANA [ser. I, vol. 43 152, 618.—Delima sarmentosa f. hirsutior Mita. Ann. Mus. Bot. Lugd. Bat. 4 (1868) 73.—Tetracera hirsuta BoerRL. Cat. Hort. Bog. (1899) 3.—Tetra- cera levinei MerR. Philip. J. Sc. Bot. 8 (1918) 147. —Tetracera scandens (non MerR.) MERR. Lingn. Sc. J. 5 (1927) 128.—Davilla hirsuta Teysm. & BINN. ms. Small shrub (to 3m) or liana (to 12m) with scabrid branches. Leaves oblong, 6—12 by 3—5!/2 cm, scabrid. Petiole 5-10 mm. Inflorescence terminal, 30-150-flowered, in the basal part often with 14 leaves, 10-25 by 5—15 cm; branches scabrid. Flower ca 8-10 mm diam. Sepals 5, the outer two 2 by 1'!/2 mm, the inner three 4 by 3 mm, scabrid out- side. Petals 3. Stamens 3-4 mm long; anthercells slightly separated at the apex. Carpels with ca 10-12 ovules. Capsule ovoid, ca 6-10 by 4-6 mm with a 2-5 mm long beak, 1(—2)-seeded. Seeds 4 by 3 mm. Aril 5 mm long, fimbriate for 1/2-7/3 of its length. Distr. Ceylon, Assam, Bengal, Andaman Is- lands, E. Siam, Indo-China, S. China, in Malaysia: Sumatra, Malay Peninsula, and ?Borneo. Notes. The species has been separated from Tetracera scandens (.) MERR., in which it has been included until now as a glabrous-fruited variety, as it appeared that the difference in the carpel is con- nected with the number of sepals. The calyx in T. scandens is tetramerous, only in a few flowers on the same plant pentamerous, in 7. asiatica it is in all flowers pentamerous. Within the species 4 subspecies can be recog- nized. They differ in the shape of the leaf, but the most characteristic and constant difference is found in the structure of the indumentum. As to this character no intermediate forms have been found. The 2 extra-Malaysian subspecies are geographi- cally isolated; these are ssp. zeylanica HOOGL. in Ceylon, and ssp. asiatica in E. Siam, Indo-China and S. China (Kwangsi, Kwangtung and Hainan). The areas of the two Malaysian subspecies overlap in the Malay Peninsula. ssp. andamanica HOOGL. ssp. nov. Young branches sparsely strigose together with sparsely to profusely distributed divergent tufts of ca 3—12 shorter (0.3—0.5 mm long) hairs; branches of the inflorescence similar. Leaves generally nar- rower than in the second Malaysian subspecies, with acute to obtuse apex and base. Distr. Assam, Bengal, Andaman Islands, in Malaysia: Malay Peninsula (rare). ssp. Ssumatrana HooGL. ssp. nov.—Delimopsis hirsuta Mig. l.c—Delima sarmentosa f. hirsutior Mio. /.c.—Tetracera hirsuta BOERL. |.c.—Davilla hirsuta TEYSM. & BINN. ms. Young branches hirsute with 1'/2-2!/2 mm long hairs together with profusely distributed divergent tufts of ca 3-12 shorter (0.3—0.5 mm long) hairs; branches of the inflorescence similar. Leaves with rounded apex and rounded to obtuse base. Distr. Malaysia: Sumatra, Malay Peninsula and ?Borneo. Ecol. In forests, up to 1300 m. Notes. The Borneo record is based on sterile material only (Tetracera setigera KORTH. ms.). 4, Tetracera maingayi HOOGL. nom. nov.—Delima laevis MAING. ms. ex KING, J. As. Soc. Beng. 58, 2 (1889) 362.—Tetracera borneensis (non MiQ.) RIL. Fl. Mal. Pen. 1 (1922) 6. Scandent shrub. Leaves oblong, ca 7'/2-15 by 3-6 cm, with acute, somewhat acuminate apex, coriaceous, shining above. Petiole 10-20 mm. In- florescence terminal, rather many-flowered (up to 250), basal part often with 1-3 leaves. Flower ca 12-15 mm diam. Sepals 5, the outer two 31/2 by 3 mm, the inner three 5 by 3!/2-4!/2 mm, scabrid Outside. Petals 3. Stamens 2!/2 mm long; anther- cells manifestly separated at the apex. Carpel 1, with ca 6 ovules. Capsule oblong, 8-12 by 3-4 mm, with a 2-3 mm long beak. Seeds unknown. Distr. Malaysia: Malay Peninsula (Malacca, Selangor, Penang) and ?Borneo. Ecol. Seems to be rare in lowland forests up to 200 m. Vern. Malay Peninsula: akar mémpélas, a. m. bétina. Borneo: akar amplas. Notes. The specific epithet /aevis is already occupied in Tetracera, hence a new name is needed. The single Borneo record is based on a specimen in the Singapore herbarium bearing the inadequate indication ‘Borneo, Remow, 1703’. 5. Tetracera lanuginosa DieELs, Bot. Jahrb. 57 (1922) 440. Liana. Branches hirsute with up to 2 mm long hairs, together with groups of very small, stellately grouped hairs (0.1 mm long). Leaves elliptic, 5—7 by 3!/2-4!/2 cm, slightly scabrid, with rounded apex and base, rather sparsely hirsute with up to 2mm long, rather rigid hairs. Petiole 10-15 mm. Inflo- rescence terminal on a few-leaved lateral branch, few (2-4)-flowered, ca 34cm long. Flower ca 15 mm diam. Sepals 5, approximately circular, the outer two ca 4mm diam., the inner three ca 6 by 7mm, lanuginose outside together with stellate groups of short hairs, glabrous to very sparsely shortly strigose inside. Petals 3, ca 9 by 6mm. Stamens 2!/2-3 mm long; anthercells strongly separated at the apex. Carpels 2-3, with ca 2 mm long, rather thin, ferrugineous hairs. Fruit unknown. Distr. Malaysia: NE. New Guinea (April River, Sepik District), once collected. Ecol. Primary forest; f/. in September. 6. Tetracera nordtiana F.v.M. Fragm. 5 (1865) 1; Bait. Queensl. Fl. 1 (1899) 9; ScHuM. & HOLLR. Fl. Kais. Wilh. (1899) 47; ScHum. & Laut. FI. Deut. Schutzgeb. Siidsee (1901) 444; Bart. Compr. Cat. Queensl. Pl. (1909) 18, f. 3.—Tetracera eury- andra (non VAHL) Roxs. FI. Ind. ed. CAREY 2 (1832) 646.—Tetracera wuthiana F.v.M. Fragm. 10 (1876) 49; Batt. Queensl. Fl. 1 (1899) 10.—Tetra- cera everillii F.v.M. Descr. Notes Pap. Pl. 7(1886) ’ 25.—Tetracera moluccana MARTELLI in BEcc. Ma- lesia 3 (1886) 153.—Tetracera cowleyana BAIL. Dept Agric. Brisb. Bot. Bull. 5 (1892) 7; Queensl. Dec. 1951) * 1 (1899) 9; Compr. Cat. Queensl. Pl. (1909) 18, . 3bis.—Tetracera boerlagei MERR. Int. Rumph. tes 366.—Tetracera pilophylla Diets, Bot. Jahrb. 57 (1922) 440.—Tetracera volubilis (non L.) RENDLE, J. Bot. 59 Suppl. (1923) 2. Shrub or large climber, up to 10m high, with slightly scabrid branches. Leaves elliptic to lance- olate, ca 5-10 by 3—5 cm, with rounded to acute apex and base. Petiole 5-15 mm. Inflorescence ter- minal, 15—50-flowered, often with 1—3 leaves in the basal part. Flower ca 6-10 mm diam. Sepals 4—S, the outer two circular, ca 1!/2-2 mm diam., the inner two or three ovate, ca 3—4!/2 by 2—3!/2 mm, glabrous or rarely sericeous inside, scabrid outside. Petals 3. Stamens 4—5 mm long; anthercells strong- ly separated at the apex. Carpels 2-4, usually 3, with rather rigid, 0.5 mm long hairs thinner than DILLENIACEAE (Hoogland) 145 in 7. scandens (L.) MERR., with ca 10 ovules. Cap- sules ovoid, 5-8 by 3—S mm with a 1—2 mm long beak, 1-seeded. Seed ca 3 by 2!'/2 mm. Aril 5mm long, laciniate to 2/3-3/4 of its length. Distr. Queensland (E. coast), in Malaysia: SE. Celebes (Kabaéna Island), Moluccas (Mangoli of the Sula group, Ceram, Ambon, Aru Islands), New Guinea, and Louisiades. Notes. A very variable species, of which 6 va- rieties are distinguished here between some of which intermediate forms occur. Most of these have been described as separate species, but to my opinion the differences do not justify to keep them apart. As they show no geographical exclusion I have provisionally accepted them as varieties instead of subspecies. For further study more ma- terial, with notes on the habitat, is highly desirable. KEY TO THE VARIETIES i Sepals slightly sericeous inside Sepals glabrous inside. 2 Leaves glabrous above 2. Leaves more or less hairy above. var. celebica var. wuthiana 3. Leaves relatively small, ca 5—7 by 3-4 mm. Capsules relatively small, ca 5 by 3 mm. 4. Indumentum consisting of strigose or strigose-hirsute hairs together with stellate groups of shorter hairs var. nordtiana 4. Indumentum consisting of strigose- hirsute hairs together with shorter, villose hairs. 3. Leaves relatively large, ca 8-10 by 4-5 cm. ees relatively large, ca 8 by 5 mm. 5. Sepals villose outside 5. Sepals strigose outside, with hairs single c or in sroups of 2- 5 var. nordtiana.—T. nordtiana F.v.M. l.c.—T. pilo- phylla Diets, I.c.—T. volubilis (non L.) RENDLE, ie: Young branches, branches of the inflorescences, leaves and outer side of the sepals with strigose or strigose-hirsute hairs, together with stellate groups of shorter hairs. Sepals glabrous inside. Capsules rather small, ca 5S by 3 mm. Distr. Queensland, in Malaysia: (Aru Islands), E. New Guinea. Ecol. Rain-forest climber. Notes. The Malaysian specimens show transi- tions to var. moluccana (MARTELLI) HOOGL. Moluccas var. everillii (F.v.M.) HooGL. stat. nov.—T. everillii F.v.M. Lc. Young branches, branches of the inflorescences, and leaves hirsute with up to 2 mm long hairs. Sepals villose outside, glabrous inside. Capsules rather large, ca 8 by 5 mm. Distr. Malaysia: SE. New Guinea (Fly River, once collected). var. wuthiana (F.v.M.) HooGL. stat. nov.—T. wuthiana F.v.M. L.c. Young branches, branches of the inflorescences, and nerves on the lower side of the leaves strigose, leaves glabrous above. Sepals sparsely strigose out- side, glabrous inside. Distr. Queensland, in Malaysia: an interme- diate form to var. nordtiana in New Guinea (CARR 12832, Koitaki). Ecol. In forest ca 300 m alt. var. louisiadica var. everillii var. moluccana var. moluccana (MARTELLI) HOOGL. stat. noy.— T. euryandra (non VAHL) Roxs. /.c.—T. moluccana MARTELLI /.c.—T. cowleyana BAIL. /.c.—T. boer- lagei MERR. l.c. Young branches and branches of the inflores- cences strigose to hirsute with up to 11/2 mm long hairs. Leaves sparsely hirsute with rather rigid, up to 11/2 mm long solitary hairs. Sepals strigose with up to 0.7 mm long hairs, single or in groups of 2-5, outside, glabrous inside. Capsules rather large, ca 8 by 5mm. Distr. Queensland, in Malaysia: Aru Islands, New Guinea. Vern. Amboina: talir hassat, hassat and hassat cotel. Probably also: gumi uccu (fide RUMPHIUS) in Ternate. Notes. Funis urens glabra RumMpH. Herb. Amb. 5 (1747) 13 may be this species, as has been sug- gested by MERRILL. The Rumphian description, however, is very defective. Moluccas, var. louisiadica HOOGL. var. nov. Young branches and branches of the inflores- cences strigose-hirsute with up to 0.8mm long hairs, together with a rather dense indumentum, closely appressed, of shorter, villose hairs. Leaves hirsute with up to 0.5 mm long hairs, most densely so on the nerves above, densely shortly villose on the intervenium, strigose-hirsute on the nerves beneath. Sepals densely villose outside, glabrous inside. Capsules rather small, ca 5 by 3 mm. Distr. Malaysia: Louisiades (Joannet Island, once collected). 146 FLORA MALESIANA var. celebica HOOGL. var. nov. Young branches and branches of the inflores- cence rather densely hirsute with up to 0.7 mm long hairs. Leaves hirsute with up to | mm long hairs. Sepals slightly sericeous in the central part inside, strigose-hirsute with 0.1-0.3 mm _ long hairs, partly in groups of 2-5, outside. Distr. Malaysia: SE. Celebes (Kabaéna Island). 7. Tetracera indica (CHRISTM. & PANZ.) MERR. Int. Rumph. (1917) 367; BAcK. & SLoot. Theeonkr. (1924) no 174; HeYNE, Nutt. Pl. (1927) 1070; Burk. Dict. (1935) 2143; Merr. J. Arn. Arb. 19 (1938) 354: Back. Bekn. Fl. Java em. ed. 4 (1942) fam. 80, 3.—Eugenia malaccensis (non L.) BURM. f. FI. Ind. (1768) 114; cf. Steen. Bull. Bot. Gard. Btzg 18 (1950) 459.—Assa Houtr. Nat. Hist. 5 (1776) 275, t. 26, f. 1.—Assa indica Houtt. ex CHRIST. & PANZ. Pfi. Syst. 4 (1779) 40, t. 26, f. 1.—Wahlbo- mia indica THUNB. Vet. Akad. Handl. Stockh. (1790) 215, t. 9.—Assa exotica GMEL. Syst. (1791) 839.—Tetracera laevis VAHL, Symb. Bot. 3 (1794) 71, non al.—Tetracera assa DC. Syst. 1 (1818) 402; Hk. f. & Tu. Fl. Ind. 1 (1855) 63; Mig. FI. Ind. Bat. 1, 2 (1859) 8; Mia. Ann. Mus. Bot. Lugd. Bat. 4 (1868) 74; Hx. f. & Tu. Fl. Br. Ind. 1 (1872) 31; Kina, J. As. Soc. Beng. 58, 2 (1889) 362; BACK. Fl. Batavia | (1907) 3; BAcK. Schoolfl. Java (1911) 9: Koorp. Exk. FI. Java 2 (1912) 600; RipL. FI. Mal. Pen. 1 (1922) 5.—Tetracera wahlbomia DC. Syst. 1 (1818) 403.—Tetracera malabarica LAMK, Illustr. 3 (1823) 32, t. 485, f. 1.—Tetracera dicho- Fig. 1. Tetracera indica (CHRISTM. & PANZ.) MERR. with buds, flowers, and fruits (right lower corner). Palembang (DE VooGD). [ser. I, vol. 43 toma BL. Bijdr. 1 (1825) 4.—Tetracera gracilis BL. ibid.; Miq. Fl. Ind. Bat. 1, 2 (1859) 9.—Tetracera trigyna Roxs. FI. Ind. ed. CAREY 2 (1832) 645; HUNTER (ed. by RIDL.) J. Str. Br. R. A. S. 53 (1909) 98.—Eleiastis laevis RAFIN. Sylva Tellur. (1838) 165.—Fig. 1. Shrub to 2 m or liana to 5 m. Leaves elliptic to oblong or obovate, ca 6-10 by 3-5 cm, usually rather thin, slightly glossy above, not scabrid, with acute to obtuse apex. Petiole ca 6-10 mm. Inflores- cence terminal on the main branch or on few- leaved axillary branches, few-(usually 4—7)-flow- ered, up to 8 by 6 cm. Flowers ca 2'/2-3 cm diam. Sepals 4, 8-10 by 7-9 mm, glabrous. Petals 3-5, usually 4, reddish white. Stamens 6-8 mm long; anthercells reaching each other at the apex. Car- pels 3-4, with a few rigid hairs on the back, ovules ca 10-15. Capsules globular, ca 10 mm diam., with a 2-6 mm long beak, 1—7-, usually 2-seeded. Seeds 3'/2 by 2!/2mm. Aril 8-10 mm long, finely fim- briate to nearly its base. Distr. From Chittagong to S. Burma, S. Siam and S. Indo-China, once collected in China (Fu- kien), in Malaysia: Sumatra, Malay Peninsula, Java, Madura, and Kangean. Ecol. Small shrub in open places; low liana, climbing over low shrubs, in brushwood and open forest. From sea-level to 600m. According to BACKER & VAN SLOOTEN, /.c., propagation mainly by suckers. Vern. Sumatra: aplas kédjong (Djambi), baih siepiek, bait sipiek suloh (Lamp.), djélati (Pal.), mémplas gadja (Sum. E. Coast), sipik suluh (Lamp.) wajit sipit (Mengg.). Malay Peninsula: akar pulas duyio, ampalas (ampélas, émpélas, hémpélas, mém- pélas), ampalas lichin, a. mihsak, a. minyak, a. payah, kalintat niamok, ma ampalasu akar. Banka: akar témpélas. Java: akar mémpélas (témpélas), asahan, émpé as (mémpélas) akar, kaju as(s)ahan (Mal.); (areuj) ki as(s)ahan (Sund.); bo (Jav.). Kangean: buko-buko. Uses. For cordage; leaves (BURKILL, /.c.) as sandpaper? Medical use unimportant, cf. BurR- Kini /-G: 8. Tetracera akara (BurRM. f.) MeRR. Philip. J. Sc. 19 (1921) 366.—Akara-Patsjoti RHEEDE, Hort. Mal. 5 (1685) 15, t. 8.—Calophyllum akara BurM. f. FI. Ind. (1768) 121.—Tetracera laevis (non VAHL) DC. Syst. 1 (1818) 401; Hx. f. & Tu. FI. Ind. 1 (1855) 62; Mig. Ann. Mus. Bot. Lugd. Bat. 4 (1868) 74; Hx. f. & Tu. FI. Br. Ind. 1 (1872) 31; BAcK: Schoolfl. Java (1911) 9; Bekn. Fl. Java em. ed. 4 (1942) fam. 80, 2.—Tetracera sericea BL. Bijdr. 1 (1825) 3; Mia. FI. Ind. Bat. 1, 2 (1859) 9.— Tetra- cera assa (non DC.) HAssk. Pl. Jav. Rar. (1848) 177.—Tetracera axillaris MARTELLI in Becc. Ma- lesia 3 (1886) 151.—Tetracera assa var. RIDL. J. Str. Br. R. A. S. 33 (1900) 37.—Tetracera syl- vestris RIDL. J. Str. Br. R. A. S. 54 (1910) 8; FI. Mal. Pen. 1 (1922) 5. High climbing or creeping liana, up to 25 m. Leaves oblong to lanceolate, ca 8-13 by 3'/2-6 cm, rather coriaceous, bright glossy green above, not scabrid, with acuminate apex. Petiole 5-7 mm. Dec. 1951] Inflorescence terminal or axillary, few-(usually 5-8)-flowered, up to 8 by 6cm. Flowers ca 2!/2—-3 cm diam. Sepals 4, 8-10 by 6-8 mm, glabrous outside, densely sericeous on the centre inside. Petals 3-4, white or greenish white. Stamens 7-8 mm long; anthercells reaching each other at the apex. Car- pels 3-4, with ca 10 ovules. Capsules globular, ca 10mm diam., with a 1-3 mm long beak, 1-2- seeded. Seeds 3 by 2mm. Aril 6 mm long, fim- briate for !/2-3/4 of its length. Distr. S. India, Ceylon, in Malaysia: Sumatra, Malay Peninsula, W. Java, Borneo, and Celebes. Ecol. In lowland forests, up to 750 m alt. Vern. Sumatra: daun amplas (Pal.). Malay Peninsula: akar, akar rusa-rusa, mumplas rimba. Java: aroy péngasaman (Sund.). Notes. 7. axillaris MARTELLI represents a small-leaved form, to my opinion of no taxonom- ical value. The acuminate, glossy leaves are typical for the species. 9. Tetracera loureiri (FIN. & GAGNEP.) CRAIB, Kew Bull. (1922) 165; Fl. Siam. En. 1 (1925) 20.—Tetra- cera assa var. loureiri FIN. & GAGNEP. Bull. Soc. Bot. Fr. Mém. 4 (1906) 3.—Tetracera sarmentosa var. loureiri FIN. & GAGNEP. FI. Gén. J.C. 1 (1907) 16.—Tetracera fragrans RiDL. J. Str. Br. R. A. S. 59 (1911) 62; Fl. Mal. Pen. 1 (1922) 6, non WILDEM. & Dur. (1899). Low climber. Leaves elliptic to oblong, ca 5-11 by 2!/2-6cm, rather coriaceous, not or slightly scabrid, with rounded or obtuse apex. Petiole ca 7-10 mm. Inflorescence terminal, many-(usually 40-80)-flowered, often with 1—2(-4) leaves in the basal part, 10-20 by 5-10cm. Flowerca10mmdiam. Sepals 5, the outer two ca 4'/2 by 3!/2 mm, the inner three 6!/2 by 4!/2 mm, slightly scabrid outside. Pet- als 3. Stamens 4-5 mm long; anthercells manifestly separated to nearly reaching each other at the apex. Carpels 2-3, with 8-16 ovules. Capsules ovoid, T'/2 by 5'/2 mm with a 2 mm long beak, 1—2-seeded. Seeds 2-3 mm diam. Aril unequal-sided, 2-6 mm long, laciniate to !/s—'/4 of its length. Distr. Siam and S. Indo-China, in Malaysia: only in the N. of the Malay Peninsula. Ecol. In thickets and hedges, from sea-level up to 400 m. Vern. In the Malay Peninsula noted only: mémpeélas. 10. Tetracera korthalsii Mig. Ann. Mus. Bot. Lugd. Bat. 4 (1868) 75; Merr. En. Born. (1921) 381.—Tetracera subrotunda Em. Leafl. Philip. Bot. 5 (1913) 1771.—Tetracera elmeri MERR. Univ. - Calif. Publ. Bot. 15 (1929) 194. Large climber or creeper with scabrid branches. Leaves broadly elliptic to elliptic-oblong or obovate, smooth to slightly scabrid on both sides. Inflorescence 40—200-or more-flowered, terminal, basal part often with 1—2 leaves, 10—30(—100) by 6-20 cm, with more or less scabrid branches. Flower ca 10 mm diam. Sepals 5(-6), the outer two 4 by 3, the inner three (or four) 5 by 4 mm, scabrid outside. Petals 3. Stamens 31/24 mm long; anther- cells strongly separated at the apex. Carpels 3, DILLENIACEAE (Hoogland) 147 with ca 9 ovules. Capsules ovoid, ca 7 by 4mm with a 1-2 mm long beak, 1-seeded. Seeds 4!/2 by 3'/2 mm. Aril unequal-sided, 2!/2-5 mm long, laci- niate to !/3—!/2 of its length. Distr. Malaysia: Borneo, Palawan, Celebes, W. Moluccas (Taliabu of the Sula group). Notes. The species can be divided into 2 va- rieties on account of the leaf-shape; there are no differences in floral characters and intermediate forms are unknown. var. korthalsii. Leaves elliptic-oblong, ca 6'/2-17 by 4-8 cm; apex acute or slightly acuminate, base acute. Peti- ole ca 8—20 mm. Distr. Malaysia: Borneo, Celebes, W. Moluc- cas (Taliabu). Ecol. Climber in forests, up to 700 m alt. Vern. Borneo: émpélas (Mal.), pampad (Du- sun). Uses. Used for polishing wood. var. subrotunda (ELM.) HOOGL. stat. nov.—Tetra- cera subrotunda EM. l.c.—Tetracera_ elmeri MerR. l.c. Leaves broadly elliptic, ca 81/2-22 by 5!/2-13 cm; apex and base rounded. Petiole 12-30 mm. Distr. Malaysia: Br. N. Borneo (Tawao), Phil- ippines (Palawan). Ecol. A large climber in primary forest at low alt. Notes. Tetracera elmeri MERR. represents a hirsute form. 11. Tetracera macrophylla WALL. ex HK. f. & TH. Fl. Ind. 1 (1855) 63; Mia. Fl. Ind. Bat. 1, 2 (1859) 8; Hx. f. & Tu. FI. Br. Ind. 1 (1872) 32; Kine, J. As. Soc. Beng. 58, 2 (1889) 363; Ripi. Fl. Mal. Pen. 1 (1922) 4; Burk. Dict. (1935) 2143.—Tetra- cera macrocarpa WALL. Cat. (1828) no 6628, nomen.—Tetracera scaberrima Mia. FI. Ind. Bat. 1, 2 (1859) 8; Ann. Mus. Bot. Lugd. Bat. 4 (1868) 75; Merr. En. Born. (1921) 382.—Tetracera teys- mannii MARTELLI in Becc. Malesia 3 (1886) 150.— Tetracera radula MARTELLI in Becc. Malesia 3 (1886) 153; Merr. En. Born. (1921) 382, non Martius (1863).—Tetracera grandis KING, J. As. Soc. Beng. 58, 2 (1889) 363; Ann. Roy. Bot. Gard. Calc. 5 (1896) 115, t. 129; Rip. Fl. Mal. Pen. 1 (1922) 4.—Tetracera havilandii Riot. Kew Bull. (1912) 381.—Tetracera scabricaulis RipL. Kew Bull. (1912) 381. Liana up to 10m or ?tree; younger branches usually more or less scabrid. Leaves elliptic to oblong, ca 8-15 by 5—10 cm, little to very scabrid, with rounded to obtuse apex and base. Petiole ca 15-30 mm long, 24 mm broad, up to 8 mm broad in the leaves at the base of the inflorescences. Inflo- rescence terminal, 25—200-flowered, often with 1-4 leaves in the basal part, 10-40 by 4-15 cm; branches scabrid with stellate groups of hairs. Flower ca 2-21/2 cm diam. Sepals 5—6, the outer two 8-9 by 7-8 mm, the inner 3-4 11-12 by 8-9 mm, scabrid outside. Petals 3, rather thick, apex not emar- ginate. Stamens 5—7 mm long; anthercells mani- 148 FLORA MALESIANA [ser. I, vol. 4 festly separated at the apex, connective strongly emarginate between them. Carpels 3—4, with a few rigid hairs on the back, with ca 14 ovules. Capsules ovoid, 8-10 by 6-8 mm with a 2-3 mm long beak, 1—2-seeded. Seeds 6!/2 by 4!/2 mm; aril unequal- sided, 5—9 mm long, slightly laciniate over ca !/s of its length. Distr. Malaysia: Sumatra, Malay Peninsula, Banka, Borneo. Ecol. Climber in dry as well asin swampy for- ests, up to 300 m alt. Seems to be rarely in flower. Vern. Sumatra: akar ampaleh riembu (Sum. W. Coast). Malay Peninsula: ampalas gajah (= ele- phant or big a.), a. rimau (= tiger a.), a. rimbah, a. lidah kuching (= cat’s tongue a.). Borneo: akar tembara (W. Kutei), ampalas (Saraw.). Uses. Medicaluseunimportant, cf. BURKILL, /.c. Notes. The species is very variable as to leaf- size and degree of scabridness; it is most easily recognized by its calyx (outer two sepals glabrous, inner sepals densely sericeous inside) and rather large flowers. 12. Tetracera arborescens JACK, Mal. Misc. 1, 5 (1820) 45; Mra. Fl. Ind. Bat. 1, 2 (1859) 9; GAGE & Burk. J. Str. Br. R. A. S. 73 (1916) 242.—Tetra- cera euryandra (non VAHL) HK. f. & TH. Fl. Ind. 1 (1855) 63; Mia. FI. Ind. Bat. 1, 2 (1859) 8; Ann. Mus. Bot. Lugd. Bat. 4 (1868) 75; Hk. f. & Tu. FI. Br. Ind. 1 (1872) 32; Kina, J. As. Soc. Beng. 58, 2 (1889) 362; Back. Schoolfl. Java (1911) 9.—Tetra- cera laevigata Mia. FI. Ind. Bat. 1, 2 (1859) 8; Ann. Mus. Bot. Lugd. Bat. 4 (1868) 74.—Tetracera sub- cordata BoERL. Cat. Hort. Bog. (1899) 3.—Tetra- cera lucida WALL. Cat. (1828) no 6631, nomen; Rip. Fl. Mal. Pen. 1 (1922) 5.—Tetracera lucida var. lanuginosa RIDL. ibid. Strong woody climber, shrub or ?small tree. Younger branches villose to densely villose-floc- cose. Leaves obovate to elliptic-oblong, ca 6-10 by 3—5 cm, coriaceous, not scabrid, the upper ones under the inflorescence 3-4 by 2-3 cm, often vil- lose-floccose when young. Petiole 3-5 mm. Inflo- rescence terminal, (6—)10—50-flowered, up to 15 by 6cm. Flower ca 15mm diam. Sepals 5—6, ca 5 by 3 mm, slightly scabrid outside. Petals 3, white. Stamens 4—5 mm long; anthercells slightly to mani- festly separated at the apex. Carpels 3, with ca 10-12 ovules. Capsules ovoid, 7 by 4mm with a 2-3 mm long beak, 1-seeded. Seeds 21/2 by 11/2 mm. Aril 3—5 mm long, laciniate to half its length. Distr. Malaysia: Sumatra (Tapanuli, East Coast), Malay Peninsula, Banka, Billiton, Borneo, and ?W. Java (Papandajan, KORTHA‘iS). Ecol. Swampy forests, riverside scrubs, only at low altitudes. Vern. Sumatra: andor ruhas igung (Tapan.), mohi-mohi (Sibolga). Banka: akar témpélas. Bil- liton: akar mémplas. Notes. JAcK’s description is insufficient. From study of a specimen of JACK, present in the Leyden herbarium, it is clear that the present species was meant. 13. Tetracera fagifolia BL. Bijdr. 1 (1825) 4; Mia. Fl. Ind. Bat. 1, 2 (1859) 9; Ann. Mus. Bot. Lugd. Bat. 4 (1868) 75: Ripv. J. Str. Br. R.A.S.54(1909) 10; Fl. Mal. Pen. 1 (1922) 6, non WILLD. ex SCHLECHT. (1833).—Tetracera rigida Bu. Bijdr. 1 (1825) 4; Mia. FI. Ind. Bat. 1, 2 (1859) 9; BAck. Schoolfl. Java (1911) 9; Bekn. Fl. Java em. ed. 4 (1942) fam. 80, 2.—Tetracera blumei WALP. Rep. 1 (1842) 67.—Tetracera sumatrana Mia. FI. Ind. Bat. Suppl. 1 (1861) 618, 619.—Tetracera fagifolia f. subintegerrima Mia. Ann. Mus. Bot. Lugd. Bat. 4 (1868) 75.—Tetracera borneensis Miq. Ann. Mus. Bot. Lugd. Bat. 4 (1868) 76; Merr. En. Born. (1921) 381.—Tetracera obovata BOERL. Cat. Hort. Bog. (1899) 3.—Tetracera philippinensis MeERR. Philip. J. Sc. Bot. 9 (1914) 375; En. Philip. 3 (1923) 58.—Tetracera obliquinervia ELM. Leafl. Philip. Bot. 7 (1915) 2621.—Fig. 2. Liana, up to 14m high, branches slightly sca- brid. Leaves elliptic to lanceolate, more or less coriaceous, usually shining. Petiole ca 7-20 mm. Inflorescence terminal, ca 40—250-flowered, often with 1(—3) leaves in the basal part, ca 15-40 by - 8-25 cm; branches scabrid, with small tufts of 0.2-0.4 mm long hairs on the extreme branches only. Flower ca 8-12 mm diam. Sepals 5(—6), the outer two 4 by 4mn, the inner 3(—4) 5!/2-7 by 4!/2-51/2 mm, scabrid outside. Petals 3. Stamens 4!/2-6 mm long; anthercells manifestly separated at the apex, connective somewhat emarginate between them. Carpels 3, with ca 10 ovules. Cap- sules ovoid, 5-8 by 4-6 mm, with a 1-3 mm long beak, 1(—2)-seeded. Seeds 5 by 3mm. Aril ca 7mm long, unequal-sided, laciniate to !/4—!/2 its length. Distr. Malaysia: Sumatra, Malay Peninsula, Banka, W. Java, Borneo, Philippines. Notes. The 2 varieties, distinguished here, dif- fer only in their vegetative parts. There is a rela- tively small number of intermediate forms. The first to consider 7. fagifolia BL. and T. rigida BL. as conspecific was Mique (1868) /.c., who used the first name. var. fagifolia. Leaves ca 12-16-nerved, 1.42.25 times as long as broad, ca 7—20 by 5-10 cm; the lateral nerves curving upward, ending in the margin. Distr. Malaysia: Sumatra (incl. Simalur and Mentawei), Malay Peninsula (Johore), W. Java, Borneo, Philippines. Ecol. Climber in primary forest, in scrub or in bamboo forest, 100-750 m alt. Vern. Sumatra: alor ampaleh (Simalur), am- palu riembu (Lamp.), sapbet (Siberut). Java: aroy (ki) assahan, kiassahan, ki saun (Sund.). Philip- pines: balau-balau (Mbo). var. borneensis (M1Q.) HOOGL. stat. nov.—Tetra- cera borneensis Mia. l.c. Leaves ca 8—10-nerved, 1.4—3.5 times as long as broad, ca 6-13 by 2.7—5.5 cm; lateral nerves not quite reaching the margin. Distr. Malaysia: Sumatra (Muara Mengku- lem, once collected), Banka, Borneo, SE. Celebes (Kendari). Ecol. Primary forest; on Mt Kinabalu up to 1300 m. Dec. 1951] DILLENIACEAE (Hoogland) Fig. 2. Tetracera fagifolia BL. a. Flowering branch, 2/3, b. flower, x 2, c.stamen, x 7, d. fruits, x 2, ~ e. seed enveloped by aril, x 4. 150 FLORA MALESIANA [ser. I, vol. 4% 2. HIBBERTIA Anpr. Bot. Rep. (1800) t. 126; B. & H. Gen. Pl. | (1862) 14; Bru. FI. Austr. 1 (1863) 17; Batt. Queensl. FI. 1 (1899) 11. Small ericoid or erect shrubs, rarely trees or lianas, mostly much branched. Leaves spirally arranged, rarely opposite (Madagascar), simple, often with reflexed margins. Inflorescence dichasial, usually reduced to a few-flowered pseudo-raceme, spike or to solitary flowers. Bracts and bracteoles often present. Flowers sessile or shortly pedicellate. Sepals 5. Petals 5(—3). Stamens ~~3, often partly staminodial, either surrounding the carpels regularly or reduced on one side, free or slightly connate at the basis, introrse, opening with longitudinal slits, rarely with apical pores. Carpels ~—1, with 15-1 ovules, free. Style filiform, usually curved or re- curved. Follicles usually one-seeded. Seed arillate. Distr. A large genus with ca 100 species in Australia and Tasmania, 20 in New Caledonia, one in the Fiji Islands, two in extreme Eastern Malaysia, and one in Madagascar (fig. 3). KEY TO THE SPECIES 1. Stamens and staminodes all placed on one side of the carpels. Carpels 2. Shrubs 1. H. banksii 1. Stamens placed regularly round the carpels, without staminodes. Carpels 5. Scandent 2. H. scandens Fig. 3. Distribution of the genus Hibbertia. Separately indicated are the areas of H. banksii (R.BR. ex DC.) Bru. (...), H. scandens (WILLD.) DRYAND. (3 black spots), and H. scandens var. novoguineensis (Gress) Hooct. (+). 1. Hibbertia banksii (R.Br. ex DC.) Bru. FI. Austr. 1 (1863) 20; BAIL. Queensl. Fl. 1 (1899) 12; BANKS & SOLANDER, Illustr. Bot. Cook’s Voy. 1 (1900) 5, t. 2; A. C. Smiru, J. Arn. Arb. 22 (1941) 497.—Hemistemma_ banksii R.BR. ex DC. Syst. 1 (1818) 414. Shrub 1-2 m. Leaves oblong-lanceolate with a few obtuse teeth, stiff with recurved margins, glossy green above, pubescent below with a rusty brown tomentum on the nerves and a yellowish tomentum on the intervenium, 5—15 by 0.7—3 cm. Raceme axillary, 5—12-flowered, 4-6cm _ long. Flowers almost sessile. Sepals oblong, acute, dense- ly hairy outside, 8 by 4mm. Petals yellow, obo- vate with emarginate apex, 14 by 7 mm. Stamens ca 25, 3'/2 mm long; staminodes ca 12, on the out- side of the stamens, 2.7 mm long. Carpels densely hairy, with 3 ovules. Follicles one-seeded, thin- walled. Seed with membranaceous aril. Distr. N. Queensland, E. coast, rare, in Ma- laysia: S. New Guinea, Wassi Kussa River region, once collected (fig. 3). Ecol. The only New Guinea specimen was found in Agonis scrub. 2. Hibbertia scandens (WILLD.) DRYAND. in KON. & Sims, Ann. Bot. 2 (1807) 525.—Dillenia scandens WILLD. Sp. Pl. 2 (1799) 1351.—Hibbertia volubilis Anpr. Bot. Rep. (1800) t. 126; Bru. Fl. Austr. 1 (1863) 37; BAL. Queensl. Fl. 1 (1899) 16.—Dillenia volubilis VENT. Choix (1803) 11.—AHibbertia novo- guineensis GiBBS, Phytogr. & Fl. Arfak Mts (1917) 148; KANEH. & Hatus. Bot. Mag. Tokyo 57 (1943) 63.—Fig. 4. Low scandent shrub. Leaves lanceolate, acute to acuminate, slightly hairy above, rather densely to densely hairy beneath, 3-12 by 0.4—2!/2 cm. Flowers solitary on short few-leaved side-stalks, ca 4 cm diam. Sepals ovate, acute, densely hairy out- side, woody, 15-18 by 8-9 mm. Petals yellow, obovate with slightly emarginate apex, ca 2 by -1.4cm. Stamens co, 7-9 mm long. Carpels glabr- ous, with 6 ovules. Follicles 1-6-seeded. Seed with membranaceous aril. Dec. 1951] Distr. Australia along the E. coast except the extreme North, S to ca 35°, in Malaysia: Aru Islands (Trangan: BUWALDA 5510) and NW. New Guinea (Arfak Mts) (fig. 3). Fig. 4. Hibbertia scandens (WILLD.) DRYAND. var. novoguineensis (GIBBS) HOOGL., nat. size. DILLENIACEAE (Hoogland) 151 Ecol. In Trangan at low alt., on the Arfak Mts between 2000 and 3000m in low spinneys on burnt Open summit. Notes. In Malaysia two varieties can be distin- guished on the leaf-shape: var. scandens with leaves rather densely hairy beneath, 6-12 by 1.2—2!/2 cm (Trangan Island), and var. novoguineensis (GIBBS) HOoGL. stat. nov. (H. novoguineensis GiBBs) with leaves densely hairy beneath, smaller and nar- rower, 3—8!/2 by 0.4-1 cm (Arfak Mts). The sta- mens in the type specimen of H. novoguineensis are 7 mm long, in the collection of KANEHIRA & HATUSIMA 9 mm, in var. scandens they are 9 mm. In var. novoguineensis the anthercells are rela- tively longer than in var. scandens. These differ- ences, however, do not justify to my idea the distinction of the two forms as separate species. Fig. 5. Geographical distribution of the genus Acrotrema; the dots represent the localities of A. costatum JACK, the question marks doubtful re- cords of this species. 3. ACROTREMA WAck, Mal. Misc. 1, 5 (1820) 36; Hk. f. & Tu. FI. Ind. 1 (1855) 64; Bide Gen: PL 1 (1862) 13: Hx: f & Tu. Fl. Br. Ind. 1-(1872) 32; Trim; Handbs pe Ceyle i (1893) 6. Perennial herbs with a horizontal, woody rhizome. Leaves all radical or on a short stem, simple, pinnatisect or pinnate, the petiole with sheathing, membra- naceous, caducous wings. Inflorescence a terminal raceme, sometimes reduced to a single flower, with membranaceous bracts. Flowers 5-merous. Stamens I15-~, usually in 3 bundles alternating with the carpels. Carpels usually 3, slightly coher- ent in the centre, with linear, recurved styles, with 2-6 or 10-20 ovules. Follicles irregularly dehiscing, with 1-15 seeds. Seeds with a white membranaceous aril. Distr. Ca 10 spp., 1 in the Deccan Peninsula and 8 in Ceylon, 1 in Lower Burma, Peninsular Siam and the Malay Peninsula (fig. 5). 1. Acrotrema costatum JACK, /.c.; KING, J. As. Soc. Beng. 57, 2 (1890) 361; Hx. f. & Tu., l.c. (1855) 65; Kurz, Nat. Tijd. Ned. Ind. 27 (1864) 175; Hx. f. & Tu., /.c.. (1872) 32; Gace & BURK. J. As. Soc. Str. Br. 73 (1916) 242; Merr. En. Born. (1921) 382; Riwv. Fl. Mal. Pen. 1 (1922) 7; BurRK. Dict. (1935) 41; HENDERS. Mal. Wild FI. 1 (1949) 20, f. 9.—Fig. 6. FLORA MALESIANA Bye [ser.-lj voliee2 Fig. 6. Acrotrema costatum JACK. a. Habit, x 2/3, b. leafbase, x 2/3, c. flower, x 4/3, d. stamen, X 7. Stem from very short up to 25 cm long. Leaves obovate, dentate, base auriculate, hairy, deep green, often with a whitish line along the midrib, 7-25 by 3-10cm; petiole 1—2(-6) cm. Raceme 9-13 cm, erect, ca 10—-12-flowered; bracts ca 6 by 2mm. Flowers yellow, diam. 3 cm, opening singly; pedicel 5-15 mm long. Carpels 3, with 2-6 ovules. Follicles enclosed by the sepals. Seeds finely echinate. Distr. Lower Burma, Peninsular Siam, in Ma- laysia: Malay Peninsula, N. Sumatra?, Banka? Ecol. In dense, wet forests or on moist shady rocks; up to 1000 m alt. Vern. Once noted: punai tanah (Pahang). Notes. One Sumatra record is based on a speci- men, collected by BATTEN POOLL, labeled only: Sumatra, 1939; the other is based on a note by JACK (cf. GAGE & Burk. /.c.). Of the Banka re- cord (cf. Kurz, /.c.) I have seen no material. The Borneo record of MERRILL, /.c., was based on a specimen of Neurocalyx sp. (Rubiaceae). 4. DIDESMANDRA STAPF in HOOK. Icon. t. 2646 (1900). Woody plants with scabrid, hairy branches. Leaves with amplexicaul sheath and short petiole. Flowers regular in calyx and corolla, zygomorphic in androecium and gynoecium, placed one-sided on simple or bifurcate branches of a 4—6- branched panicle with reduced central axis, almost sessile. Sepals and petals 5. Stamens in 2 bundles on the adaxial side of the carpels, each bundle consisting of 1 stamen and 4 staminodes with connate filaments, the stamen uncinate-curved, exceeding the staminodes, with longitudinally dehiscent linear anthercells, the con- nective forming a deltoid membrane above the cells, the staminodes only slightly curved. Carpels 2, with a long filiform style; ovule 1, inserted at the base. Fruit a nut. Seed with a thin membranaceous aril. Distr. Monotypic, known only from Borneo. Dec. 1951] DILLENIACEAE (Hoogland) 153 Wty Ze; XY WS neg wi ij WV. y ~| pe . in Fig. 7. Didesmandra aspera Starr. a. Habit, x 1/2, b. flower, x 1/2 (after STAPF), c. stamen and 4 stami- nodes, x 3, d. tip of staminode, x 10, e. ditto of stamen, x 10, f. ovary, x 3, g. seed (black) with unilateral aril, < 3. 154 FLORA MALESIANA [ser. I, vol. 43 Notes. Most closely allied to Schumacheria VAHL from Ceylon, which has many stamens in one bundle, without staminodes. 1. Didesmandra aspera STAPF, /.c.; MeRR. En. Born. (1921) 382.—Fig. 7. Plant ca 2!/2 m high. Leaves scabrid, ovate, apex acute, base rounded, margin slightly toothed, 15-30 by 7-13 cm, 12—-16-nerved; nerves prominent and hairy beneath; petiole channelled, 1!/2-3 cm. diam. Sepals elliptic-oblong, the 2 outer ones smal- ler. Petals obovate, 25 by 13 mm, yellow. Filament 2 mm long, | mm thick, anther 17 mm long, sterile anthers 8-10 mm. Carpels and fruit glabrous; style 15-20 mm. Distr. Malaysia: Borneo (Sarawak), twice col- lected. Inflorescence 15—20cm long. Flowers ca 5 cm 5. DILLENIA LINNE, Sp. Pl. 1 (1753) 535; Gen. Pl. ed. 5 (1754) 239.—Wormia Rotts. Nye Samml. Danske Vid. Selsk. Skrift. 2 (1783) 532.—Lenidia THou. Gen. Nov. Madag. (1806) 17.—Colbertia SALIsB. Parad. Lond. (1807) sub t. 73.—Capellia BL. Bijdr. (1825) 5.—Reifferscheidia PRESL, Rel. Haenk. 2 (1836) 74.—Capellenia BL. ex Hassk. Cat. Pl. Hort. Bog. (1844) 187. Trees or shrubs, often with reddish bark peeling off in thin papery scales. Leaves spirally arranged, simple. Petiole in a number of species with usually wholly ca- ducous, rarely partly or wholly persistent, broad wings, amplexicaul in the young leaf and then enclosing and protecting the terminal bud. /nflorescence a composed or simple raceme, in a number of species reduced to solitary flowers, usually ter- minal on consequently sympodial branches, in one species axillary; one species with terminal and cauline inflorescences, some other (mainly extra-Malaysian) ramiflorous with fascicled flowers. Bracts small, caducous, or obsolete. Bracteoles well developed in some spp., in others obsolete. Flowers actinomorphic. Sepals (4-)5(-6), in a few species more, concave. Petals 5, in some species absent, in one species 4-6, usually obovate with rounded apex, yellow or white, rarely reddish. Stamens «, all of approximately the same length or of different lengths arranged in 2 or more, not always sharply separated groups; occasionally part of the outer- most stamens staminodial, in some species a wholly staminodial outer group, in one species a wholly staminodial inner group. Anthercells parallel, opening usually with a terminal pore, less often with longitudinal slits. Carpels 4-20, coherent along the cuneate central part of the receptacle, with filamentous or linear, more or less spreading styles; stigma in most spp. indistinct, only in 2 species (D. serrata TuHungB. and D. celebica HOOGL.) distinct, knoblike. Ovules 6 to ca 60. Fruit either dehiscent, the rather fleshy carpels spreading like a star, or indehiscent, enclosed by the more or less enlarged and thickened sepals. Seeds arillate or exarillate, glabrous or rarely finely echinate. Distr. Ca 60 spp., from Madagascar and the Seychelles to the Fiji Islands, in the North to the S. slopes of the Himalayas, Yunnan, Kwangsi and Kwangtung, in Australia only one species on the E. coast of Queensland; not in New Caledonia. The most widespread species are D. indica L. (from India to Borneo and Java) and D. pentagyna Roxs. (cf. fig. 12). There are a number of local endemics, partic- ularly in the Philippines, New Guinea, and the Pacific Islands. Ecol. Most species occur in evergreen forests on dry to very wet soil. Some deciduous species are found in monsoon forests or savannahs, one evergreen species in savannahs of New Guinea and N. Australia. As to altitudinal distribution the species generally occur below 1000 m, but some are occasion- ally found above this altitude, up to 2000 m; one species (D. montana Diets) has been collected only above 1000 m. Stilt-roots occur constantly only in a few species, viz D. borneensis HOOGL., D. eximia Miq., and D. reticulata KING; in a few others they may be occasionally developed, e.g. in D. papuana MARTELLI and D. albiflos (RIDL.) HOOGL. The leaves of saplings and young plants are often considerably larger than those of the full-grown plants. In most cases these leaves are relatively narrower, without showing a distinct dimorphism. In DILLENIACEAE (Hoogland) Dec. 1951] 155 2/5, fee se c. inflorescence, x 2/s, d. longitudinal section of flower during full anthesis, x 4/s, e. stamens, x 2, f. apex of anther, x 4, g. seed with aril, x 2. Fig. 8. Dillenia papuana MARTELLI. a. Fruiting branch, x 2/s, b. winged petiole of young lea 156 FLORA MALESIANA [ser. I, vol. 43 D. ferruginea (BAILL.) GILG from the Seychelle Islands, however, a distinct leaf-dimorphism is found between these leaves. In a less degree this is found in Malaysia in D. pentagyna Roxs. and probably in D. pteropoda (MiQ.) HOOGL. In some species the petals drop without having opened in anthesis. In Malaysia this is found only in D. papuana MARTELLI and some other New Guinean species. The dispersal of fruits and seeds is effected mainly by animals. The indehiscent fruits would be eaten mainly by mammals, the arillate seeds of the species with dehiscent fruits by birds. Transport by water is a means of dispersal in D. indica L. Uses. Though sometimes used for light constructions, the wood is generally of low value because of the short durability. D. pentagyna Roxs. is used for making a charcoal of good quality. The indehiscent fruits of some species are eaten, in particular in curries and jellies; they have an acid taste. Mixed with syrup they make a cough cure, and they are sometimes used for washing the hair. Because of the beautiful flowers and foliage a number of species are suitable as ornamental trees or shrubs, e.g. D. indica L., D. philippinensis ROLFE, and D. suffruticosa (GRIFF.) MARTELLI. Wood anat. MoLt & JANssonius, Mikr. Holzes 1 (1906) 67: ground tissue originally described as libriform fibers which error was later corrected into fibertracheids, see also REINDERS, Hand]. Planten- anatomie 4 ed. Wageningen (1915) 142-147; PEARSON & BROWN, Comm. Timb. | (1932) 1; VESTAL, Philip. J. Sc. 64 (1937) 205; DEN BERGER, Med. Proefstat. Boschwezen 13 (1926) 118 (handlens). D. excelsa (= Wormia excelsa): M.& Js p. 69. D. indica: M. & Js p. 71; P. & B. p. 3. D. pentagyna: M. & Js p. 78; P. & B. p. 7. D. obovata (= D. aurea, non SMitH): M. & Js p. 76. D. parviflora GRIFF. (extra- Malaysian): P. & B. p. 6. Vern. The Malay name ‘simpur’ (simpoh, sémpur, etc.) is in general use throughout W. Malaysia, some species being distinguished by epithets. In the Philippines ‘katmon’ is in general use. Notes. Several classifications of the genus have been proposed, the basic one usually being in Dillenia and Wormia as separate genera or subgenera on account of the fruit (indehiscent versus dehiscent) by some authors, on account of the seed (exarillate versus arillate) by others. Other subgeneric classifica- tions are based on the base of the petiole (amplexicaul versus non-amplexicaul) or on the structure of the androecium. It is impossible, however, to distinguish groups, characterized by the combination of more than one character; relationships within the genus are reticulate. MarTELLI in Becc. Malesia 3 (1886) 150-167 was the first to unite the genera Dillenia and Wormia. He listed the species of both genera, but failed to make formally the new combinations, which have been ascribed to him by Dur. & JACKS. in the first supplement of Index Kewensis. An important character of the androecium is the position of the anthers in bud; in a number of species the inner row is reflexed outwards. This may be permanent in anthesis. I have used the term ‘straight’ or ‘straight or slightly curved’ in all other cases. The sequence of the species as adopted here reflects as far as possible their affinity. KEY TO THE SPECIES 1. Basis of the petiole, later leaf-scar, completely amplexicaul. 2. Petiolar wings not constricted below the blade. Nervation of the wings not sharply separated from that of the blade, though often less distinct. 3. All stamens of approximately the same length. Flowers white . . ‘ . 1. D. pteropoda 3. Androecium consisting of small staminodes on the outside and stamens of different lengths, the length increasing towards the centre. Flowers yellow. . . « » 15. Desotiriticosa 2. Petiolar wings more or less constricted below the blade. Wings without distinct nervation or nerva- tion different and independent from that of the blade. 4. Sepals 8 or more. 5. Sepals 8-9, not much different in size : » « « a £ SSD Simarsopalis 5. Sepals 11-17, distinctly increasing in size towards the centre . . . . 14, D. reifferscheidia 4. Sepals 4-6, usually 5. 6. Stamens gradually decreasing in size towards the centre of the flower, the innermost ones staminodial . . . eh ee 12. D. fagifolia 6. Stamens all of approximately the s same e length or r the innermost ones onger than the outer ones. If present, staminodes on the outside. 7. All stamens of approximately the same length. 8. Carpels hirsute. 9. Inflorencence simple, distinctly zigzag. Flower ca 6'/2 cm diam., yellow . 2. D. beccariana 9. Inflorescence usually composed, with only slightly zigzag axes. Flower ca 4 cm diam., white. 3. D. albifios 8. Carpels glabrous. 10. Stigma distinct, knoblike. Flowers apetalous. 11. Petiolar wings semi-obcordate. Flowers rather large (e.g. stamens ca 9-11 mm long). 4. D. serrata 11. Petiolar wings broadly linear. Flowers smaller (e.g. stamens ca 6—7 mm long). 5. D. celebica Dec. 1951] DILLENIACEAE (Hoogland) 1577 10. Stigma indistinct. Flowers with (caducous!) petals. 12. Upper part of petiolar wings persistent . . . . . . =. =. ~. . 6. D. ovalifolia 12. Petiolar wings wholly caducous. een @annelsr4—OnlISually Sits Maen i sw eel tel ua ts le | eck eae n Ea) MLO ISChers 13. Carpels 7-15. 14. Stamens with a 1-2 mm long acumen at the apex. . . . . . . 7. D. papuana 14. Connective at most slightly exceeding the anthercells. 15. Younger parts with dense villose indumentum . . . . . . . 8. D. montana 15. Younger parts glabrous or slightly hirsute. 16. Leaves ovate, ca 15—19-nerved. Sepals ca 25 by 22 mm 2) oD Schechter 16. Leaves elliptic, ca 6—-9-nerved. Sepals ca 15 by 12mm. . . . II. D. quercifolia 7. Innermost stamens longer than the outer ones, with the apical part reflexed outward in bud. 17. Apex of petiolar wing for ca '/4 or more of its length exceeding its insertion to the petiole. 18. Flowers solitary on a pedicel, terminal. Apex of the petiolar wings rounded. [Seblowersica 6—Sicmidiam. Canpelsi6—9) = = 3.) 2 16. D. ochreata 19. Flowers ca 20 on diam. Carpels 14-16. . . . . . . 17. D. megalantha 18. Flowers in a 2—3-flowered raceme. Apex of the petiolar wings obtuse . 18. D. talaudensis 17. Apex of the petiolar wing not or hardly exceeding its insertion to the petiole. 20. Upper part of petiolar wings persistent . . Je |. 19D yalata 20. Petiolar wings wholly caducous, or at most small auricles persistent near the basis of the blade. 21. Petiolar wings elliptic-oblong. Flowers large, more than 10 cm diam. 20. D. philippinensis 21. Petiolar wings narrower. Flowers smaller, up to 10 cm diam. 22. Leaves elliptic to oblong, rather coriaceous, 5-8-nerved . . . . . 21. D. diantha 22. Leaves oblong to lanceolate, not coriaceous, 8—20-nerved. 23. Apex of the leaf acuminate. Plant often cauliflorous. Flowers whitish . 22. D. bolsteri 23. Apex acute. Plant not cauliflorous. Flowers eae 24. Leaves glabrous, 8—12-nerved hae . . . 23, D. auriculata 24. Leaves, at least the young ones, Strigose on . the nerves s beneath, 10—18-nerved. 24. D. castaneifolia 1. Basis of the petiole, later leaf-scar, not completely amplexicaul, clasping up to 3/4 of the branch. 25. Inflorescence or solitary flowers typically terminal on leaf-bearing branches, often later lateral, leaf-opposed. Branches consequently sympodial. 26. Deciduous trees, with stilt-roots. Inflorescence appearing with the leaves, often immediately at its base with 2 or 3 branches, forming a loose cluster. 27. Carpels 4—6. Stamens all of approximately the same length. Flowers apetalous . 25. D. eximia 27. Carpels 7-10. Stamens not all of the same length. Flowers with (caducous!) petals. 28. Carpels 7-8. Petiole densely hirsute above, nearly glabrous beneath . . 26. D. borneensis 28. Carpels 9-10. Petiole more densely hairy beneath than above . . . . 27. D. reticulata 26. Evergreen trees, without stilt-roots. Inflorescence never branched immediately at its base. Flowers either solitary or in a raceme. 29. Stamens all of approximately the same length. Pedicel with 3 large verticillate bracteoles. Leaves densely velvety tomentose beneath . ~ 2 J. 285Dehookert 29. Stamens in 2 sharply separated groups of different lenght. No ‘such bracteoles. Leaves not densely velvety tomentose beneath. 30. Carpels less than 12. 31. Carpels 6 or more. 32. Flowers in 4 to many-flowered inflorescences. 33. Inflorescence continuously growing. Upper internodes of inflorescence up to 1 cm long. Leaves 6-15 by 3-7cm . . . . . 29. D. luzoniensis 33. Inflorescence with only short fe of erowile Internodes of inflorescence longer. Leaves 15-30 by 7-10cm_. . S OS 6 ees 0D Sexcelsa 32. Flowers solitary or in up to 3- flowered inflorescences. 34. Leaves small (up to 7 by 2!/2 cm), lanceolate, coriaceous . . - 31. D. sibuyanensis 34. Leaves much larger. 35. Leaves elliptic with rounded apex. Flowers ca 16cmdiam. . . . . . 32. D. ovata 35. Leaves oblong with obtuse to acute apex. Flowers ca6cmdiam . . 33. D. sumatrana Bile, Calg Sous Ai ee ie ic mr oe oe de hk 34. D. monantha 30. Carpels 15—20 Aas Se 52D indica 25. Flowers either axillary on leaf-bearing branches ¢ or solitary or in fascicles on leafless branches. 36. Carpels 5—6. Flowers small, ca 2'/2-3!/2 cm diam. 37. Flowers axillary. Evergreen species with small leaves... ee a 2 3020D)> pulchella 37. Flowers in twig-born fascicles. Deciduous species with large leaves . . 37. D. pentagyna 36. Carpels 9-11. Flowers large, ca 15cmdiam. . . . . . +. + « - - 38D. obovata 158 FLORA MALESIANA [ser. I, vol. 4 1. Dillenia pteropoda (Mia.) HOooGL. comb. nov. —Wormia pteropoda Mig. Ann. Mus. Bot. Lugd. Bat. 4 (1868) 77.—Dillenia papyracea MERR. Philip. J.Sc. Bot. 9 (1915) 520; En. Philip. 3 (1923) 61.—Dillenia megalophylla Merr. Philip. J.Sc. 14 (1919) 421; En. Philip. 3 (1923) 60.—Wormia papyracea GiLG & WERDERM. in ENGL. & PR. Nat. Pfi. Fam. ed: 2, 21 (1925) 35. Large tree, up to ca 40m high, up to 1!/4m diam. Bark red-orange, peeling off in plates. Leaves elliptic, subcoriaceous, ca 17—21-nerved, 30-100 by 16-60 cm, blade with rounded to obtuse apex, obtuse to acute base and entire to slightly undulate-dentate margin. Petiole ca 5—10 cm long, wings up to 2!/2 cm broad, often caducous. Raceme simple, ca 4-7-flowered, up to 40 cm long. Flowers ca 10 cm diam. Sepals 5, ca 25-33 by 20-22 mm, slightly to densely velvety outside. Petals white, ca 45—50 by 30-32 mm. Stamens ca 700, 14-18 mm long. Carpels 8-12, ca 10 by 3!/2mm, with ca 1 cm long styles, each with ca 25 ovules. Fruit dehiscent. Seeds surrounded by a basal, loose, somewhat cup-shaped aril. Distr. Malaysia: Philippines (N. Luzon, Min- danao) and N. Moluccas (Halmahera, Batjan). Ecol. In primary forests, often along streams, from sea-level up to 500 m. Vern. Philippines: tukoran (Lan.), malaigang (Sul.). Moluccas: several names recorded, but none constant. Notes. TEISMANN 5886 from Batjan probably represents leaves from a coppice or from a, young plant. The leaves are manifestly dentate and have a very short petiole. They are narrower than in the other specimens, ca 90 by 28 cm, with more lateral nerves (ca 40—50). Dillenia papyracea Merr. and D. megalophylla Merk. differ only in the length of the pedicel; this does not seem sufficient argument to keep the two separated. From the Moluccas (including the type specimen) only sterile material is available. 2. Dillenia beccariana MARTELLI in Becc. Malesia 3 (1886) 158; Merr. En. Born. (1921) 382.— Wor- mia heccariana RipvL. Saraw. Mus. J. 1 (1913) 71. Small tree, up to 6 m high. Leaves oblong to nar- rowly obovate, ca 20—30-nerved, 18-45 by 8-16 cm, with obtuse, often acuminate apex, rounded to obtuse base and slightly to manifestly dentate margin. Petiole ca 3-6 cm long; wings near their base 4-9 mm broad, gradually narrowing towards the base of the blade, coherent with the blade over a breadth of 1!/2-2mm, when older loosening from the base of the blade and near the base of the petiole, but not caducous. Raceme up to 20-flow- ered, up to 60 cm long, distinctly zig-zag. Flowers ca 61/2 cm diam. Sepals 5, ca 17-20 by 14-15 mm. Petals yellow, ca 33 by 22 mm. Stamens ca 130, 11-13 mm long. Carpels 5-6, ca 8-10 by 2mm with ca 5 mm long styles, the carpels and the base of the styles densely covered with rigid, up to 2mm long hairs; each with ca 20 ovules. Fruit dehiscent, but the carpels possibly only slightly spreading, the young fruit not enclosed by the sepals. Carpels 25 by 16 mm, 1—3-seeded. Seeds 4 by 2!/2 mm, with a 0.2-0.4 mm long membranac-, eous aril. Distr. Malaysia: Borneo (Sarawak). Ecol. In low altitude forests. Vern. Simpoh, s. délaki (= male s.), pétasi. Notes. Closely related to D. albiflos (R1DL.) HooGL. 3. Dillenia albiflos (RipL.) HooGL. comb. nov.— Wormia albiflos Ripv. J. Str. Br. R. A. S. 54 (1910) 6; Fl. Mal. Pen. 1 (1922) 9.—Wormia beccariana (non Rip.) Corn. Gard. Bull. S.S. 10 (1939) 4; Wayside Trees Malaya (1940) 205. Tree, up to 17 m high, with red bark, rarely with few stilt-roots. Leaves elliptic to oblong, ca 15—35- nerved, 20-40 by 9-20 cm, with rounded to acute, often slightly acuminate apex, rounded to obtuse base, and slightly undulate to manifestly dentate margin. Petiole ca 2'/2-4 cm long; wings near the base 5-11 mm broad, gradually narrowing towards the base of the blade, coherent with the blade over a breadth of 1-2 mm, when older loosening from the base of the blade and near the base of the petiole, but not caducous. Inflorescence pendent, up to 30-flowered, composed, being a raceme with the second and often third flower replaced by a secondary raceme, the branches slightly zig-zag. Flowers ca 4cm diam. Sepals 5, ca 15-20 by 12-14 mm. Petals pale cream white, ca 20 by 13 mm. Stamens ca 160, 5!/2-8 mm long. Carpels 5-6, ca 4-5 by 11/2 mm with ca 5 mm long styles, the carpels and the lower half of the styles rather densely covered with rigid, 0.4-0.7 mm long hairs, each with ca 10 ovules. Fruit dehiscent, the sepals in fruit enlarged to 25 by 15 mm. Carpels 10 by 12 mm, 1—2-seeded. Seeds 4 by 3 mm, finely echi- nate, with a 1 mm long membranaceous aril. Distr. Malaysia: S. Malay Peninsula (E. Jo- hore). Ecol. In dry forest and on dry hillocks in swamps, at low altitude. Vern. Only noted: simpoh. Notes. Closely related to D. beccariana MAR- TELLI, from which it differs by the composed in- florescence, white, slightly smaller flowers and less densely hairy carpels with distinctly shorter hairs. According to CorNneER (fieldnote) the biggest trees have the biggest leaves. 4. Dillenia serrata THUNB. Trans. Linn. Soc. 1 (1791) 201; Mig. Fl. Ind. Bat. 1, 2 (1859) 685; Merr. Int. Rumph. (1917) 368.—Sangius RUMPH. Herb. Amb. 2 (1741) 142, t. 46.—?Songium RUMPH. Herb. Amb. 2 (1741) 140, t. 45.—?Dillenia elliptica THUNB. Trans. Linn. Soc. 1 (1791) 200; MARTELLI in Becc. Malesia 3 (1886) 161; Merr. Int. Rumph. (1917) 367; Heyne, Nutt. Pl. (1927) 1071. Rather large tree, up to 30m high and 70cm diam., with thinly scaling, reddish gray bark. Leaves oblong to lanceolate, 16—35-nerved, 20-45 by 8-19 cm, with rounded to acute apex, obtuse to acute base, and nearly entire to manifestly den- "tate margin. Petiole 2!/2-6'/2 cm long, with half- obcordate, caducous wings. Wings broadening towards the apex, the apex rounded, extending Dec. 1951] distinctly above its insertion, at the basis 5-8, near the apex 15-30 mm broad, glabrous to densely sericeously hirsute beneath. Raceme 2—6-flowered, up to 15 cm long with straight to rather tortuous axis. Flowers probably apetalous, ca 7!/2 cm diam. Sepals 5, ca 40 by 25 mm, densely sericeously hir- sute outside. Stamens ca 750, 9-11 mm long. Car- pels ca 18-19, ca 4!/2 by 1'/2 mm, with ca 8 mm long, in the basal half parallel, in the apical half slightly spreading styles with a cushion-shaped stigma, stigma ca 0.4 mm thick and 1 mm diam.; each carpel with 5—9 ovules. Fruit indehiscent, yellowish, appressed globular, 3'/2 cm high, 6 cm diam. including the enclosing sepals. Sepals in fruit enlarged to 6!'/2 by 5!/2 cm, at the base up to 3mm thick, not completely covering the carpels at the apex. Carpels 25 by 16 mm, up to 5-seeded. Seeds black with reticulate surface, exarillate. Distr. Malaysia: Celebes, Buton, and Muna Islands. Ecol. In primary forests up to 180 m. Uses. The fruit is eaten. Vern. Celebes: déngén, d. bolusu, déngilo, dongi, wuadéngi. Muna: sonih. Notes. THUNBERG’s binomial is wholly based on Sangius RuMPH. RUMPHIUS’ description and plate are sufficient for the recognition of the spe- cies. Dillenia elliptica THUNB. is wholly based on Songium Rumpu. The identification of this species is much less certain, but the present species is the only one, that Rumpuius’ description and plate can be compared with, except that the flowers in RUMPHIUS’ species are solitary. 5. Dillenia celebica HOOGL. spec. nov. Tree, up to 30 m high, 50 cm diam., with small buttresses, greyish brown bark, slightly flaky in large plates, and greyish red heartwood. Leaves elliptic-oblong, ca 15—20-nerved, 13-18 by 6- 10 cm, with obtuse to acute, often slightly acumin- ate apex, rounded-obtuse to obtuse base, and undulate to dentate margin. Petiole 4-8 cm long, with usually 2!/2-5 mm, rarely up to 10 mm broad wings; wings with slightly auriculiform apex, caducous. Raceme 1-S-flowered, up to at least 4cm long. Flowers apetalous, ca 4!'/2cm diam. Sepals 5, ca 21-25 by 16-19 mm, sericeous outside. Stamens ca 300, 6-7 mm long. Carpels ca 11, ca 8 by 3!/2 mm, with ca 7 mm long spreading styles with a cushion-shaped stigma, stigma ca 0.3 mm thick and 0.8mm diam.; each carpel with 3-4 ovules. Fruit unknown. Distr. Malaysia: N. and C. Celebes. Uses. The wood is used for house-building. Notes. Closely related to D. serrata, also en- demic of Celebes, from which it differs by the much longer petioles, the shape of the petiolar wings, and the much smaller number of carpels. The 2 species are the only ones, where a distinct, knoblike stigma is found in Dillenia. 6. Dillenia ovalifolia HooGL. spec. nov.—Dillenia alata MARTELLI in Becc. Malesia 3 (1886) 157, quoad descr. Tree, up to 30 m high, 60 cm diam., with abso- DILLENIACEAE (Hoogland) 159 lutely glabrous branches. Leaves rather coriaceous, elliptic to nearly orbicular, 8—-13-nerved, 7—27 by 6—22 cm, with rounded apex and base and slightly undulate margin; in bud folded thus as to leave a faint, longitudinal line between each pair of lateral nerves. Petiole 4'/2-10cm long, with 5-10 mm broad wings, with a horseshoe-shaped cushion at the insertion. Wings linear with rounded apex, partly caducous, the part falling off being the whole breadth at the base of the petiole, gradually narrowing to a wholly persistent wing at !/3—3/4 of the petiole; apex auriculate, the auricles of both wings coherent above the petiole, extending slight- ly above the blade. Flowers solitary, terminal, soon lateral, leaf-opposed. Peduncle 5!/2-7 cm long, at the base triangular in transverse section. Sepals 5, the outer 2 ca 25-28 by 19-20 mm, the inner 3 35-40 by 20-22 mm, glabrous or not. Petals white, at least 25 by 18 mm. Stamens ca 900, 5-7 mm long, with a 0.7-0.8 mm long acumen. Carpels 7-8, glabrous, ca 9 by 2!/2 mm with 13 mm long styles, each with ca 8 ovules. Fruit dehiscent. Car- pels ca 17 by 10mm, 1-seeded. Seeds ca 5 by 3!/2mm, enclosed by a 3 mm long aril. Distr. Malaysia: Moluccas (Halmahera, Mo- rotai) and Japen Island near NW. New Guinea (Geelvink-Bay). Ecol. In primary forest, from sea-level to 1000 m. Vern. Japen Island: karoe ai and wadajouw. Notes. For the identity of Dillenia alata MAR- TELLI see p. 164. The impression on the leaf, caused by the folding in bud, has not been noted so clearly in any other Dillenia. The specimens from the Moluccas differ from those from Japen Island, which are entirely glabr- ous, by a dense short sericeous indumentum on the young branches, on the basal part of the nerves on the lower surface of the leaves, on the lower side of the petiole, on the peduncle, and on the outer side of the sepals. They represent a distinct variety, var. sericea HOOGL. var. nov. 7. Dillenia papuana MARTELLI in Becc. Malesia 3 (1886) 156.—Wormia pteropoda (non Miq.) BOERL. Cat. Bog. (1899) 5.—Dillenia calothyrsa DIELS, Bot. Jahrb. 57 (1922) 437.—Wormia calothyrsa GiLG & WERDERM. in ENGL. & PR. Nat. Pfl. Fam. ed. 2, 21 (1925) 35.—Wormia papuana GILG & WERDERM. /.c.—Wormia macrophylla (non G.&W.) A. C. Smitu, J. Arn. Arb. 22 (1941) 498.—Fig. 8, 9. Tree, up to 30m high, 1 m diam., often with buttresses, with pale reddish brown bark peeling off in very thin papery scales. Leaves elliptic to ovate, ca 20—25-nerved, 15-40 by 10-35 cm, on young trees and saplings narrower, ca 30—35- nerved, 50-100 by 25-50cm, with rounded to obtuse apex, rounded base, and undulate to slightly dentate margin. Petiole 4-8 cm long, on young trees and saplings up to 10cm, with half-obcor- date, rarely oblong, caducous wings. Wings usually broadening towards the apex, the apex rounded, extending distinctly above its insertion, at the basis 6-10, near the apex up to 35 mm broad. Raceme 4-7-flowered, up to 8 cm long with tortuous axis. 160 FLORA MALESIANA [ser. I, vol. 4 Fig. 9. Dillenia papuana MARTELLI. Trees ca 30 m tall, ca 1 m diam., in 2 m deep inundated flood-plain forest. This type of forest was inundated Jan. to May 1939. Bernhard Camp, Idenburg River, West New Guinea (BRAss, ARCHBOLD Expeditions). Dec. 1951] Flowers probably never quite expanding, the sepals only slightly diverging, the petals falling off without spreading. Sepals 5, ca 30-40 by 25-40 mm. Petals yellow, cucullate when falling, 25—45 by 12—-25mm. Stamens ca 185-250, 13-24 mm long (but in a single flower all of approximately the same length), with a 1-2 mm long, acute acumen above the anthercells. Carpels 10-15, ca 10-15 by 2-3 mm, with 11-15 mm long styles, each with ca 25 ovules. Fruit dehiscent. Carpels 25-35 by 16-24 mm, one-seeded. Seeds 6 by 5 mm, black, enclosed by a rather thick fleshy aril. Distr. Malaysia: Tanimbar and Aru Islands, New Guinea, and islands in the Geelvink Bay. Ecol. In primary forest at low altitudes, on dry or temporarily flooded soil (fig. 9), once collected at ca 2000 m. Uses. Said to supply a good timber. Vern. Tanimbar Islands: kamjemeje. Guinea: several noted, but none constant. New 8. Dillenia montana DieLs, Bot. Jahrb. 57 (1922) 437.—Wormia montana GILG & WERDERM. in ENGL. & Pr., Nat. Pfl. Fam. ed. 2, 21 (1925) 35. Tree, ca 35 m high with 15 m clear trunk, ca 40 cm diam., with reddish brown papery scaly bark and brown wood. Younger branches densely villose. Leaves elliptic to ovate-elliptic, 9-14-nerved, ca 10-21 by 6-17!/2 cm, with rounded, slightly acumin- ate apex, rounded base, and dentate margin, more or less villose on both sides. Petiole 2!/2-7 cm long, with linear-oblong, caducous wings. Raceme 2- flowered, up to 6 cm long, with densely villose axis. Flowers incompletely known, possibly never quite expanding. Sepals 5, the two outermost ones ca 30 by 30 mm, the three innermost ones ca 35 by 30 mm, the outer ones slightly villose outside. Petals 5, yellow. Stamens ca 90, 20 mm long. Carpels 8-9, ca 18 by 4 mm, with 9 mm long, only slightly spreading styles; each carpel with 13-14 ovules. Fruit unknown. Distr. Malaysia: NE. New Guinea (Central Highlands and Hunstein Range). Ecol. In forests, 1350 and 2000 m. Vern. Burra (Arona), warawaka (Aiyura). Notes. The species is closely related to Dil- lenia schlechteri DiELs and D. papuana Mar- TELLI. 9. Dillenia schlechteri DreLs, Bot. Jahrb. 57 (1922) 438.—Dillenia alata var. macrophylla Laur. Bot. Jahrb. 45 (1911) 362.—Dillenia macrophylla DiELs, Bot. Jahrb. 57 (1922) 437.—Wormia schlechteri GILG & WERDERM. in ENGL. & Pr. Nat. Pfi. Fam. _ed. 2, 21 (1925) 35.—Wormia macrophylla GILG & WERDERM. in ENGL. & Pr. Nat. Pfl. Fam. ed. 2, 21 (1925) 35.—Wormia nitida A. C. SmitH, J. Arn. Arb. 22 (1941) 499. Large tree, up to 35 m high, 2 m diam., with buttresses to 3 m tall, 1 m long; bark reddish brown, flaky; wood reddish brown, hard and heavy. Leaves broadly ovate, ca 15—21-nerved, 11- 30 by 7!/2-21 cm, with rounded apex, rounded, slightly cordate base, and slightly undulate margin. Petiole 4-9 cm long, with 4-8 mm broad wings. DILLENIACEAE (Hoogland) 161 Raceme 2-6-flowered, up to ca 7 cm long. Flowers probably never quite expanding, the sepals only slightly diverging, the petals falling off without spreading. Sepals 5, ca 25 by 22 mm. Petals bright yellow, cucullate when falling, ca 33 by 28 mm. Stamens ca 100, 14-17 mm long. Carpels 8-11, ca 8-9 by 2-2!/2 mm, with 10-12 mm long styles, each with ca 14-18 ovules. Fruit dehiscent, the sepals enlarged to ca 35 by 30 mm. Carpels ca 25 by 18 mm, I-seeded. Seeds ca 4 by 3 mm, enclosed by a 2!/2 mm long aril. Distr. New Ireland, in Malaysia: Guinea. Ecol. In primary forests, in New Guinea found only between 1300 and 1700 m, in New Ireland at low altitude. Uses. The durable wood is used for building purposes. Vern. Once noted: manaya (Kuni language). E. New 10. Dillenia fischeri MeERR. Philip. J. Sc. Bot. 9 (1915) 518; En. Philip. 3 (1923) 60.—Wormia fischeri GILG & WERDERM. in ENGL. & PR. Nat. Pfi. Fam. ed. 2, 21 (1925) 35. Tree, up to 20 m high. Leaves elliptic-oblong to oblanceolate, coriaceous, ca 7—11-nerved, 6'/2-14 by 2.8-6!/2cm, with rounded apex, obtuse to rounded base, and nearly entire to dentate, mainly in the upper part of the leaf, margin. Petiole 1!/2-3 cm long, with linear, 1 mm broad, caducous wings. Inflorescence terminal on the leaf-bearing branches or lateral on the older branches in the axil of a leaf-scar; raceme 2—6-flowered, sometimes with one lateral branch, up to 5 cm long. Flowers ca 6cm diam., on a 3-6 cm long pedicel. Sepals 4-6, usually 5, ca 13-16 by 8-12 mm. Petals as many as sepals, white, ca 30 by 14 mm. Stamens ca 120-160, the outer ones slightly longer, 9 mm, than the inner ones, 7 mm. Carpels 4-6, usually 5, ca 7 by 2!/2mm, with 5-8 mm long styles, each with 8-10 ovules. Fruit unknown. Distr. Malaysia: Philippines (known only from Mindanao, Butuan subprovy.). Ecol. In semi-open forests at low altitude. 11. Dillenia quercifolia (WHITE & FRANCIS ex LANE Poote) HooGt. comb. nov.—Wormia quercifolia WuHuitE & FRANCIS ex LANE POOLE, For. Res. Terr. Papua & N. Guinea (1925) 116; WHITE & FRANCIS, Proc. Roy. Soc. Queensl. 38 (1926) 242, f. 9 (1927). Large tree, ca 35 m high with 28 m clear trunk ca 1.2m diam., with reddish, papery scaly bark and yellow to rose-brown wood. Leaves elliptic 6—9-nerved, 8-15 by 4!/2-12 cm, with obtuse to rounded, minutely acuminate apex, obtuse to rounded base, and more or less undulate margin, glabrous. Petiole 3-4 cm long, the wings 3-7 mm broad. Raceme 2-4-flowered, up to ca 5 cm long with tortuous axis. Flowers ca 4-5 cm diam. Sepals 5, 12-16 by 10-13 mm. Petals not known from open flowers. Stamens ca 60, 10 mm long. Carpels 7-10, usually 8, ca 5 by 2mm, with 7 mm long style, each with 8-9 ovules. Fruit unknown. Distr. Malaysia: New Guinea. 162 FLORA MALESIANA [ser. I, vol. 43 Ecol. ‘Flowers July to August in N. Division’ (LANE POOLE). Vern. Lalagi (Buna & Binandele). 12. Dillenia fagifolia HOOGL. spec. nov. Large tree, nearly 50m high with 30m clear trunk, ca 1 m diam., with branched buttresses and reddish brown, papery scaly bark; wood pinkish or pale red brown. Leaves elliptic, 17—19-nerved, 12!'/2-16 by 8-10 cm, with obtuse apex, obtuse to rounded base, and entire to very slightly undulate margin. Petiole 5—S'/2cm long, with 7-8 mm broad wings. Raceme 6-flowered, with tortuous, ca 6cm long axis. Flower known only in bud. Sepals 5, the outermost one (and probably in the open flower the other ones of approximately the same size) 22 by 25 mm. Petals present. Stamens in 2 rather distinct groups, the outer ones, ca 60, fertile, decreasing in size towards the centre, 5—2'/2 mm long, with 0.3 mm long acute acumen; the inner ones, ca 90, sterile, ca 1 mm long. Carpels 12, in bud ca 2 by 1 mm, with 1 mm long style, each with 12-14 ovules. Fruit unknown. Distr. Malaysia: E. New Guinea (once col- lected near Aitape, L. S. SmirH, N.G.F. 1229). Vern. Ainedin (But near Wewak). Notes. The species is only very imperfectly known. The structure of the androecium, however, is so characteristic, that the said specimen certainly represents a new species. This type of androecium was not yet known in Dillenia. 13. Dillenia marsupialis HOoGL. spec. nov.—Dil- lenia ochreata (non MARTELLI) MeERR. En. Philip. 3 (1923) 61. Small tree. Leaves elliptic to oblong, ca 13-16- nerved, 12-20 by 5!/2-10cm, with rounded or obtuse, + distinctly acuminate apex, obtuse to acute base, and nearly entire to distinctly dentate margin, entirely glabrous. Petiole 11/2-4 cm long, with nearly circular to obovate wings. Wings up to 35 by 22mm with rounded apex and entire margin, glabrous, wholly caducous. Flowers ter- minal, solitary, ca 10cm diam. Sepals 8-9, the outermost ones only slightly smaller than those towards the centre, 3!/2-4 by 2!/2-3 cm. Petals unknown. Stamens in 2 groups, the outer group ca 260, ca 13 mm long, straight in bud, the inner group ca 75, ca 23 mm long, with their apex reflex- ed in bud. Carpels ca 15, ca 12 by 2mm, with ca 20mm long, recurved styles, each with 7-10 ovules. Fruit indehiscent, subglobose, 4-5 cm diam. including the enclosing sepals. Carpels ca 25 by 12mm, 1-—2-seeded. Seeds enclosed by a membranaceous aril. Distr. Malaysia: Philippines (Luzon, Panay, and Catanduanes). Ecol. On forested slopes, up to 1200 m alt. Notes. Closely related to Dillenia reiffer- Scheidia VILLAR, not to D. ochreata, though its leaves resemble very much that species. It is easily recognized from the latter species by the larger number of sepals and larger flowers and fruits. 14. Dillenia reifferscheidia VILLAR, Novy. App. (1880) 3; BLANco, FI. Filip. ed. 3 (1880) t. 344; VIDAL, Rev. Pl. Vasc. Filip. (1886) 38; MerRR. En. Philip. 3 (1923) 61.—Reifferscheidia speciosa Pres_, Rel. Haenk. 2 (1836) 74, t. 62.—Dillenia speciosa GILG in ENGL. & Pr. Nat. Pfl. Fam. 3, 6a (1893) 124, non THUNB. (1790).—Dillenia reiffer- scheidia var. rosea ELM. Leafl. Philip. Bot. 3 (1923) 61. Tree, up to ca 11 m high, 60 cm diam. Leaves elliptic to obovate, ca 14—20-nerved, ca 15-40 by 10-30 cm, with rounded apex and base and slightly to manifestly dentate margin. Petiole 3-4 cm long, strigose beneath, with obovate wings. Wings ca 3-5 by 2!/4-4 cm, with rounded apex and entire margin, more or less woolly hairy beneath, wholly caducous. Flowers terminal, solitary or less often 2-3 together, if solitary with 2-5 or without brac- teoles, if 2 (resp. 3) 1 (resp. 2) with and 1 without bracteoles; ca 17'/2cm diam. Sepals 11-17, the outer ones ca 20 by 22 mm, the inner ones ca 65 ~ by 40 mm. Petals white or, rarely, rose red (var. rosea ELM.), ca 9 by 5cm. Stamens in 2 groups, outer group ca 375, 11 mm long, straight in bud, inner group ca 60, 25mm long, with their apex reflexed in bud. Carpels ca 15, ca 12 by 3 mm, with ca 18mm long, recurved styles, each with 9-12 ovules. Fruit indehiscent, subglobose, 5-8 cm diam. including the enclosing sepals. Carpels 20-25 by 10-12 mm, 1—4-seeded. Seeds black, enclosed by a membranaceous aril. Distr. Malaysia: Philippines (from the S. half of Luzon to Mindanao, not in the Westernmost islands). Ecol. In primary forests in humid regions with abundant rainfall, from low altitude to ca 1000 m. Uses. Construction timber tree. The fruit makes a good preserve. Vern. Balali (Bik.), katmon (Tag., P. Bis.), k. kadlagan (Bik.), k. kalabau (Yag.), palali (Bik.). 15. Dillenia suffruticosa (GriFF.) MARTELLI in Becc. Malesia 3 (1886) 163; Merr. En. Born. (1921) 384; pe Wit, Bull. Bot. Gard. Btzg, III, 18 (1949) 208.—Wormia suffruticosa GrirF. Not. 4 (1854) 706; Ic. Pl. As. (1854) t. 649; Hk. f. & TH. Fl. Br. Ind. 1 (1872) 35; Kina, J. As. Soc. Beng. 58; 2 (1889) 364: Ripy. J. Str Brekeaesms4 (1910) 5; BAck. Schoolfl. Java (1911) 10; Koorpb. Exk. Fl. Java 2 (1912) 601; BLAAuw, Trop.: Nat. Schets. Kleur. (1913) 17, t. 12; Rip. Fl. Mal. Pen. 1 (1922) 8; Burk. Dict. (1935) 2265; Corn. Gard. Bull. S. S. 10 (1939) 9; Wayside Trees Malaya (1940) 207, pl. 53; Back. Bekn. Fl. Java em. ed. 4 (1942) fam. 80, 4.—Wormia excelsa (non JACK) Hk. f. & Tu. Fl. Ind. 1 (1855) 67.—Wormia sub- sessilis Mia. Fl. Ind. Bat. Suppl. (1860) 619; Ann. Mus. Bot. Lugd. Bat. 1 (1864) 315, t. 9; ibid., 4 (1868) 77; Rip. J. Str. Br. R.A.S.54 (1910) 4; FI. Mal. Pen. 1 (1922) 7, f. 2.—Wormia burbidgei Hook. f. Bot. Mag. (1880) t. 6531.—Dillenia bur- bidgei GILG in ENGL. & Pr. Nat. Pfl. Fam. 3, 6 (1893) 123.—Wormia subsessilis var. borneensis Rip. J. Str. Br. R.A.S. 54 (1910) 6.—Dillenia suffruticosa var. borneensis Rip. Saraw. Mus. J. 1 Dec. 1951] (1913) 71; Merr. En. Born. (1921) 384.—Dillenia sp. DE VooGp. Trop. Natuur 21 (1932) 61 —Fig. 10. Large shrub, up to 10 m high. Leaves elliptic to obovate, ca 12—20-nerved, ca 15-25 by 8-12 cm, blade with -+ obtuse apex and base and entire to dentate margin, glabrous above or sometimes Fig. 10. Dillenia suffruticosa (GRIFF.) MARTELLI, Palembang (DE VooGD). slightly woolly on young leaves, beneath slightly to densely woolly on the lateral nerves, on both sides of the central nerve (continuing on the peti- ole), and along the line which delimits the bud- enclosing part of the leaf-basis. Petiole ca 2-6 cm long with up to 1!/2cm broad, usually persistent wings. Raceme usually simple, sometimes com- posed, ca 5—12-flowered. Flowers ca 8-11 cm diam. Sepals 5, ca 15—22 by 8-12 mn, in fruit enlarged to ca 18-25 by 10-15 mm. Petals bright yellow, ca 40-50 by 25-30 mm. Staminodes ca 100, 6 mm long. Stamens ca 175, the outer ones 8 mm long, straight in bud, the inner ones 13 mm long, with their apex reflexed in bud, with intermediate lengths. Carpels 5—8, usually 7, ca 5 by 2 mm, with yellowish white, ca 1 cm long styles, each with 7-10 ovules. Fruit dehiscent. Carpels red, ca 20-25 by 10-16 mm, each 1-4-seeded. Seeds brown or black, with a membranaceous, scarlet aril. Distr. Malaysia: Sumatra (Palembang), Malay Peninsula, Riouw- and Lingga-Archipelagos, Na- tuna, Banka, Billiton, W.Java,and Borneo. Con- trarily to the opinion of DE Wit I do not believe that the species is indigenous in W. Java. All col- lections are made within a relatively small area DILLENIACEAE (Hoogland) 163 around Bogor and of fairly recent date, all made at least 30 years after the introduction of the spe- cies into the Botanic Garden. Moreover, the spe- cies easily naturalizes, e.g. in Jamaica, where it has become abundant. Ecol. In marshes, along streams, and on the margin of forests, often forming thickets, from sea-level up to 500 m. Flowering continuously, each flower open for one day only, between 2 flowers of the same raceme a difference of ca 3-4 days. Fruit ripe after 36 days (CORNER, 1940); seeds eaten by birds. Uses. Sometimes planted as an ornamental. Vern. Malay Peninsula: simpoh, s. ayer (= water s.), s. gajah (= elephant or big s.), s. pasir (= sand s.). Natuna: simpor. Banka: simpong, sipor, kaju simpur (= tree s.), kémbang mésimpur or masimpur (= flower s.), mininpor, simpor pram- puan (= female s.). Billiton: simpur. Borneo: dungin (Dusun), simpor, simpur, s. bini (= female s.), s. ayer (Mal.). 16. Dillenia ochreata (Miq.) TElsM. & BINN. Cat. Hort. Bog. (1866) 178, nomen, ex MARTELLI in Becc. Malesia 3 (1886) 178; HryNe, Nutt. PI. (1927) 1072.—Wormia ochreata Mia. Ann. Mus. Bot. Lugd. Bat. 4 (1868) 77, t. 1; Koorp. Meded. Pie 19N(898)rs272 Tree, up to 15 m high and 35 cm diam. Leaves ovate or elliptic-oblong, ca 15—18-nerved, 10-20 by 4'/2-10 cm, with acute, often slightly acuminate, less often obtuse apex, rounded to acute base, and nearly entire to slightly dentate margin, glabrous. Petiole ca 16-20 mm long, with broad half-obcor- date wings; wings ca 16-18 by 13-15 mm, with entire margin, glabrous, caducous. Flowers solitary, terminal, ca 6-8 cm diam. Sepals approximately circular, 2—2!/2 cm diam., glabrous. Petals yellow, 4 by 3!/2 cm. Stamens ca 165, the outermost ones ca 7'/2mm long, straight in bud, the innermost ones ca 12 mm long, with the apex reflexed in bud, with intermediate lengths. Carpels 6-9, ca 7 by 2!/2 mm, with 7 mm long styles; each with 8 ovules. Fruit indehiscent, pale green, slightly depressed- globular, ca 32 mm diam., 26 mm high. Carpels very slightly spirally twisted, ca 15 by 12 mm, 1-seeded. Seeds ca 5 by 4mm, very finely densely echinate, enclosed by a 10 mm long, rather fleshy aril. Distr. Malaysia: NE. Celebes (Minahasa). Ecol. In forests from low altitude to 800 m, often found on volcanic sand. Vern. Kelemur (Alf.); some other names noted, but not constant. 17. Dillenia megalantha Merr. Philip. J. Sc. Bot. 9 (1914) 519; Wester, Philip. Agr. Rev. 14 (1921) 242; Merr. En. Philip. 3 (1923) 60.—Dillenia mindanaensis ELM. Leafl. Philip. Bot. 7 (1915) 2611; WesTER, Philip. Agr. Rev. 14 (1921) 287, t. 27b; Merr. En. Philip. 3 (1923) 60.—Wormia megalantha GiLG & WERDERM. in ENGL. & PR. Nat. Pfl. Fam. ed. 2, 21 (1925) 35. Tall tree, up to 20(-40?) m high, 40 cm diam., with brown and gray bark, scaling offin thin plates. 164 FLORA MALESIANA [ser. I, vol. 4 Leaves oblong to oblanceolate, ca 25—35-nerved, 25-70 by 8-25 cm, with acute, often slightly acu- minate apex, rounded base, and manifestly dentate margin. Petiole 2!/2-5 cm long, with obovate wings; wings 2!/2-5 by 1!/2-3 cm (on saplings up to 10 by 6 cm), with rounded apex and slightly to manifestly dentate margin, caducous. Flowers solitary, ter- minal, ca 20cm diam. Sepals elliptic, 25-40 by 20-30 mm. Petals yellow, ca 10 by 7!/2 cm. Sta- mens in 2 distinct groups, the outer ones ca 600, 10-12 mm long, straight in bud, the inner ones ca 100, 17-25 mm long, with their apex reflexed in bud; on the outside a small number of stami- nodes, 6-14 mm long. Carpels 14-16, ca 10 by 3mm with 22 mm long styles, each with ca 25 ovules. Fruit indehiscent, 5—7!/2 cm diam. including the enclosing sepals, which are ca 7 by 5 cm, at the basis 8 mm thick. Carpels ca 25 by 11 mm, 1-seeded. Seeds obovoid, 6 by 4 mm, finely shortly echinate, with an 8 mm long, membranaceous aril. Distr. Malaysia: Philippines (S. Luzon to Min- danao, absent from N. Luzon to Palawan). Ecol. In primary forests at low altitude, up to 1000 m. Uses. The fruit is eaten. Vern. Kalambog (Bag.), katmon (Bik., S.L. Bis., Bag.), kKatmén-bayani (Tag.), lumbdg (Sub.), palali (Sub.). 18. Dillenia talaudensis HOOGL. spec. nov. Small tree, ca 8 m high, 11 cm diam.; branches glabrous. Leaves elliptic or ovate, 18—20-nerved, 20-30 by 14-21 cm, with obtuse to rounded apex with small acumen, rounded base, and slightly dentate margin. Petiole 3!/2-5 cm long, with caduc- ous wings. Wings obovate, up to 12 mm broad, the apex extending about !/2cm above the inser- tion, ending into a mucronate acumen. Raceme 2-3-flowered, with 10-12cm long axis. Flowers known only in bud and as young fruit, on a short, thick pedicel. Sepals 5, the outer 2 ca 21 by 19 mm, the inner 3 ca 30 by 23 mm. Petals in bud up to 22 by 15 mm. Stamens in 2 distinct groups, the outer ones, ca 250, ca 6 mm long in bud, 10 mm in the young fruit, straight in bud, the inner ones, ca 80, ca 11 mm long in bud, 19 mm in the young fruit, with their apex reflexed in bud. Carpels 14, ca 5 by 1.3 mm in bud, with 11 mm (in the young fruit 20 mm) long styles, each with 8-11 ovules. Fruit unknown. Distr. Malaysia: Salibabu Island, Talaud group N of Celebes, once collected. Ecol. Common in secondary forest. Uses. The fruit is eaten uncooked. Vern. Luaran’a. 19. Dillenia alata (R.BR. ex DC.) MARTELLI in Becc. Malesia 3 (1886) 157.—Wormia alata R.BR. ex DC. Syst. 1 (1818) 434; Prod. 1 (1824) 75; Bru. Fl. Austr. 1 (1863) 16; Mia. Ann. Mus. Bot. Lugd. Bat. 4 (1868) 78; F.v.M. Fragm. 7 (1871) 124; BaIL. Queensl. Fl. 1 (1899) 10; BANKs & Sot. Ill. Bot. Capt. Cook’s Voy. 1 (1900) 5, t. 1; Bam. Compr. Cat. Queensl. PI. (1909) 18, pl. 2; BRAss, J. Arn. Arb. 19 (1938) 186, pl. 222.—Lenidia alata Porr. Dict. Sc. Nat. 25 (1822) 448.—Wormia ape- tala GAUD. in FReyc. Voy. Bot. (1826) 476, t. 99.— Dillenia apetala MARTELLI ex Dur. & JACKS. Ind. Kew. Suppl. 1 (1902) 136. Tree, up to 20 m high, 60 cm diam., with reddish brown bark peeling off in thin papery flakes, and crooked branches. Leaves ovate to elliptic, ca 8—14- nerved, 8—25 by 5-15 cm, with rounded apex and base and entire, slightly recurved margin. Petiole 21/24 cm long, with 2-6 mm broad wings; wings narrowing towards the base of the blade, partly caducous, leaving behind a !/2 mm broad part on the lower !/2-3/s of the petiole, the wing on the upper 2/s—!/s permanent. Raceme 2-4-flowered. Flowers ca 7\/2 cm diam. Sepals 5, the outer 2 cir- cular, 1.3 cm diam., the inner 3 elliptic-ovate, 2 by 11/2 cm. Petals yellow, 40 by 25 mm. Stamens in 2 distinct groups, the outer ones ca 100, 7-8 mm long, straight in bud, the inner ones ca 18, 11-13 mm long, with their apex reflexed in bud. Carpels 6-8, deep crimson, ca 10 by 4mm, with red, ca 14 mm long styles, each with ca 8 ovules. Fruit dehiscent. Carpels 18—20 by 10-14 mm, 1—3-seeded. Seeds 4 by 3 mm, black, enclosed by a membran- aceous, waxy white aril. Distr. E. Coast of N. Queensland, S to about 20° S, in Malaysia: Waigeo and S. New Guinea. Ecol. In Malaysia found in savannah forests. Notes. The first description of the species was that by DC. (1818) /.c. under the name Wormia alata R.BR., citing Dillenia alata BANKS as a syno- nym. The first legitimate transfer to Dillenia was effected by MARTELLI (1886), who, however, de- scribed a specimen of Dillenia ovalifolia HOOGL. 20. Dillenia philippinensis Roire, J. Linn. Soc. 21 (1884) 307; VIDAL, Rev. Pl. Vasc. Filip. (1886) 37; Mera. FI. Manila (1912) 331; WesTER, Philip. Agr. Rey. 8 (1915) 104, t. 7a; MERR. Sp. Blanc. (1918) 263; BROWN, Minor Prod. Philip. For. 2 (1921) 338, f. 62, 63; MeRR. En. Philip. 3 (1923) 61.— Dillenia indica (non L.) BLANCO, FI. Filip. (1837) 472; VILLAR, Nov. App. (1880) 3.—Dillenia spe- ciosa (non THUNB.) BLANCO, FI. Filip. ed. 2 (1845) 329, ed. 3, 2 (1878) 244, atlas t. 199.—Dillenia cat- mon ELM. Leafl. Philip. Bot. 7 (1915) 2610; MeRR. En. Philip. 3 (1923) 59. Tree, up to 17 m high, 60 cm diam., with rather low-attached crown, reddish bark, and dark wood. Leaves elliptic or ovate to lanceolate, ca 10-15- nerved, 8—25 by 6-16 cm, chartaceous, with round- ed to obtuse, often slightly acuminate apex, round- ed to obtuse base, and slightly dentate or undulate margin. Petiole 3!/2-5 cm long, with half-elliptic to half-oblong, 3-12mm broad, caducous wings. Inflorescence a 1—2-, rarely 3-flowered raceme with 5-16cm long axis. Flower ca 10-15cm diam. Sepals 5, the two outer ones ca 1.8 by 1.8 cm, the 3 inner ones 2—2!/2 by 1.7—2 cm. Petals white, 4~7 by 2-5 cm. Stamens in 2 distinct groups, the outer ones, ca 230, yellow, 11 mm long, straight in bud, the inner ones, ca 40, purplish, 15-23 mm long, with their apex reflexed in bud. Carpels 10-12, 7-9 by 2!/2-3 mm, with linear, ca 17 mm long, spreading styles, each with 10-12 ovules. Fruit in- Dec. 1951] dehiscent, depressed-globose, 4—5 cm high, 5-6 cm diam. including the enclosing slightly fleshy sepals. Carpels slightly spirally twisted, fleshy, 20 by 13 mm, 1-4-seeded. Seeds 5 by3 mm, very finely echin- ate, at the base enclosed by a 2 mm long, mem- branaceous aril. Distr. Malaysia: Philippines from the Babuyan Islands to the Sulu Archipelago, not in Palawan. Ecol. Common in forests at low and medium altitudes, rarely above 1000 m, once collected at 1800 m. Uses. The pulp from the fruit is eaten; it makes an excellent sauce or jam and is used, mixed with sugar, as a cough cure. It is also used for cleansing the hair. A red dye is obtained from the bark. Vern. The following vernaculars are in use: balale, balobaydauak, bihis, biskan, bolobayduak, cachuchio, dingin, kalambugui, kambug, katmon, kulambig, palali, palale, and pamamalien, the most frequently used ones being katmon and palale. Notes. Merr. (1923, /.c.) distinguishes a var. pubifolia, which differs from the typical form by the hirsute inflorescences and sepals and the pilose underside of the leaves. The differences are only very slight. 21. Dillenia diantha HOOGL. spec. nov. Tree, up to 25 m high, 60 cm diam. Leaves ellip- tic to oblong, 5—8-nerved, ca 6-12 by 4!/2-7'/2 cm, with rounded to obtuse or slightly emarginate apex, rounded to obtuse base, and slightly undulate to dentate margin, glabrous. Petiole 11/24 cm long, with linear-lanceolate, 1-2 mm broad wings; wings with rounded or-auriculiform apex, caducous. In- florescence a 2-flowered raceme, less often flowers solitary; axis 11/24 cm long. Flowers ca 9 cm diam. Sepals 5, elliptic, ca 22 by 15 mm. Petals yellow, ca 45 by 29 mm. Stamens in 2 distinct groups, the outer ones ca 155, 9-10 mm long, straight in bud, the inner ones ca 20, 13-15 mm long, with their apex reflexed in bud. Carpels (5S—)7-9, glabrous to sparsely shortly hirsute, mainly in the apical part, ca 9 by 2!/2 mm, with ca 15 mm long styles. Cell- wall inside the apical part and wall of the stylar canal hirsute. Each carpel with 9-11 ovules. Fruit dehiscent, the sepals enlarged to ca 28 by 20 mm. Carpels 18 by 16 mm, showing the hirsuteness in- side the apical part. Seeds unknown. Distr. Malaysia: Philippines (Luzon). Ecol. In forests at low altitude. Uses. The wood is used for building purposes. Vern. Babacao (Ibanag dial.), malacatmon (Tag.), marapalali (Ilocane). Notes. The leaves of the species are similar to those of D. luzoniensis and D. monantha; the spe- cies is easily distinguished by the winged petiole and consequently amplexicaul leaf-scar. 22. Dillenia bolsteri Merr. Philip. J.Sc. Bot. 7 (1912) 305; En. Philip. 3 (1923) 59.—Dillenia cauli- flora MerRr. Philip. J.Sc. Bot. 9 (1915) 517; En. Philip. 3 (1923) 60. Tree, up to 20 m high. Leaves elliptic-oblong, ca 15—20-nerved, 10-25 by 4-11 cm, with acuminate apex with ca 1 cm long acumen, acute to obtuse DILLENIACEAE (Hoogland) 165 base, and slightly dentate margin, most strongly so in the upper part of the leaf. Petiole 2-4'/2 cm long, with lanceolate, 4-6 mm broad wings; wings vil- lose-pubescent beneath, caducous. Inflorescence either a terminal, 2-4-flowered, up to 10 cm long raceme, or cauline with a woody, sparingly branched axis with flowers solitary or in 2-flowered racemes terminal on this sympodially built axis. Cauline inflorescence with ovate bracts, represent- ing the winged petiole of the normally developed leaf, conduplicate, ca 10-20 mm long, 2 x 5-10 mm broad, amplexicaul, leaving amplexicaul scars on the axis. Flowers ca 6 cm diam. Sepals 5, elliptic, ca 20 by 16 mm. Petals whitish, 30 by 13 mm. Sta- mens in 2 distinct groups, the outer ones ca 160, 5—S!/2mm long, straight in bud, the inner ones ca 25, 7-8'!/2 mm long, with their apex reflexed in bud. Carpels 8-10, ca 6 by 3 mm with cylindrical, 7 mm long styles, each with 6-8 ovules. Fruit in- dehiscent, globular, ca 2!/2 cm diam. including the enclosing sepals, which are up to 30 by 30 mm, at the base 0.4 mm thick. Carpels ca 15 by 9mm, 1-seeded. Seeds obovate, 5 by 31/2 mm, exarillate. Distr. Malaysia: Philippines (Samar, Leyte, and Surigao Prov. of Mindanao). Ecol. In primary forests at low altitude. Notes. The present species is the only one in Dillenia where these sympodially built cauline in- florescences are found. The twig-born and cauline inflorescences of Dillenia pentagyna and related species most probably are monopodial and com- parable with the inflorescences of Dillenia pul- chella. 23. Dillenia auriculata MARTELLI in Becc. Malesia 3 (1886) 159.—Wormia auriculata GILG & WER- DERM. in ENGL. & PR. Nat. Pfl. Fam. ed. 2, 21 (1925) 35; A. C. Smitn, J. Arn. Arb. 22 (1941) 501. Slender, tall tree, up to 30m high, with flaky reddish brown bark. Leaves oblong, ca 8—12- nerved, 7-16 by 3-7 cm, with rounded-obtuse to acute apex, rounded to obtuse base, and entire to slightly undulate margin, glabrous. Petiole 1!/2—-5 cm long, with up to 7 mm broad, linear-lanceolate wings: wings caducous except often a small, up to 5 mm long, upper part, remaining as 2 small auri- cles below the blade. Raceme 1—3-flowered, up to 11 cm long. Flowers ca 8-10 cm diam. Sepals 5, the outer 2 ca 17-19 by 15-17 mm, the inner 3 ca 25-30 by 20-24 mm. Petals narrowly obovate, yel- low, ca 40 by 16 mm. Stamens in 2 distinct groups, the outer ones ca 225, 8-10 mm long, straight in bud, the inner ones ca 28, ca 18 mm long, with their apex reflexed in bud; yellow. Carpels 6-10, ca 8 by 3 mm, with linear, red, 16 mm long styles; each with 8-11 ovules. Fruit unknown. Distr. Malaysia: New Guinea. Ecol. On riversides and in ridge forests. Ca 1 m long proproots are noted for this species. 24. Dillenia castaneifolia (M1Q.) MARTELLI ex Dur. & JAcks. Ind. Kew. Suppl. 1 (1902) 136 (castaneaefolia); Diets, Bot. Jahrb, 57 (1922) 438. —Wormia castaneifolia Mia. Ann. Mus. Bot. Lugd. Bat. 4 (1868) 78; MARTELLI in BEcc. Malesia 166 FLORA MALESIANA [ser. I, vol. 4 3 (1886) 164 (castaneaefolia); MiQ. ex Hook. f. & JAcKs. Ind. Kew. 2 (1895) 1233 (castanaefolia).— Wormia macdonaldi F.v.M. Vict. Natural. 2 (1886) 146; Bot. Centralbl. 26 (1886) 114.—Dillenia misorensis MARTELLI in Becc. Malesia 3 (1886) 160.—Dillenia albertisiana MARTELLI in BEcc. Ma- lesia 3 (1886) 161.—Wormia longepetiolata WARB. Bot. Jahrb. 13 (1891) 378.—Dillenia pedunculata K. ScHum. & Laut. Fl. Deut. Schutzgeb. Siidsee (1901) 445.—Dillenia macdonaldi MARTELLI ex Dur. & Jacks. Ind. Kew. Suppl. 1 (1902) 136.— Wormia hirta R1iDL. Trans. Linn. Soc. Bot. 9 (1916) 13.—Dillenia longepetiolata Diets, Bot. Jahrb. 57 (1922) 436.—Dillenia hirta Diets, Bot. Jahrb. 57 (1922) 436.—Dillenia castaneifolia var. dolichobo- trys Diets, Bot. Jahrb. 57 (1922) 439.—Wormia misorensis GILG & WERDERM. in ENGL. & PR. Nat. Pfl. Fam. ed. 2, 21 (1925) 35.—Dillenia alata var. macrophylla (non LAUT.) LANE POOLE, Rep. For. Res. Papua (1925) 116. Tree, up to 20 m high, 50 cm diam., with light brown heartwood. Leaves oblong to elliptic- oblong, ca 10—-16-nerved, 10-25 by 5-15 cm, with acute to rounded apex and base and entire to slightly dentate margin. Petiole 1!/2-6 cm long, with 4-7 mm broad wings; wings narrowing towards the base of the blade, caducous. Raceme 1—6-flowered, up to 30 cm long. Flowers 61/2-9 cm diam. Sepals 5, the outer 2 elliptic to circular, 15 by 12-15 mm, the inner 3 obovate, 20-30 by 15-18 mm. Petals deep lemon yellow, 40-50 by 25-30 mm. Stamens in 2 distinct groups, the outer ones ca 250-300, 6!/2-8 mm long, straight in bud, the inner ones ca 23-35, 11-14 mm long, with their apex reflexed in bud; on the outside often a number of staminodes (up to 25). Carpels 8-10, red, 6—7 by 1!/2-2 mm, with 9-11 mm long styles, each with ca 6-12 ovules. Fruit dehiscent, when ripe with carmine red sepals enlarged to 30-40 by 17—20mm. Carpels 15 by 10 mm, 1-seeded. Seeds 4 by 3!/2mm, enclosed by a membranaceous aril. Distr. Malaysia: New Guinea and islands in the Geelvink Bay. Ecol. In primary and secondary forests, usu- ally on riversides. Vern. Oesang, wesang (Dutch New Guinea). Notes. The species is very variable in size of the leaves, less so in the leaf-shape, and in the length of the inflorescence. These forms, described as separate species or varieties, are connected with each other by intermediate forms, and considered here to represent one, polymorphic species, the most polymorphic one found in the genus. 25. Dillenia eximia Mra. FI. Ind. Bat. Suppl. (1860) 620; Ann. Mus. Bot. Lugd. Bat. 4 (1868) 79; MERR. En. Born. (1921) 383.—Dillenia crassisepala MAR- TELLI in Becc. Malesia 3 (1886) 156; Merr. En. Born. (1921) 383.—Wormia scortechinii KING, J. As. Soc. Beng. 58, 2 (1889) 365.—Wormia kunstleri Kina, J. As. Soc. Beng. 58, 2 (1889) 366.—Dillenia scortechinii Rip. J. Str. Br. R.A.S. 54 (1910) 7; Fl. Mal. Pen. 1 (1922) 12. Deciduous tree, up to 40 m high, 70 cm diam., with large buttresses, passing into stilt-roots. Leaves elliptic to obovate, ca 18—28-nerved, 15—25 by 9-15 cm, with rounded to obtuse apex, obtuse to somewhat cordate base, and entire to undulate- dentate margin; in saplings longer and narrower, 45—80-nerved, 35—75 by 13-25 cm. Petiole 3-7 cm long, in saplings up to 17 cm. Inflorescence a (3-) 5—12(-18)-flowered composed raceme, forming a loose cluster. Flowers appearing with the leaves, apetalous, ca 2!/2 cm diam. Sepals 5, about circu- lar, 9-12 mm diam. Stamens all of approximately the same length, ca 150-180, 4!/2-5!/2 mm long; margin of the anthercells ciliate. Carpels 4-6, usu- ally 5, white, ca 31/2 by 11/2 mm, with 5 mm long, white styles, each with ca 30 ovules. Fruit inde- hiscent, dark green, slightly flattened-globular, ca 30 mm diam., 25 mm high including the enclosing sepals, which are up to 40 by 27 mm, at the basis up to 8 mm thick. Carpels 10 by 7 mm, 1—2-seeded. Seeds 5!/2 by 3!/2 mm, with a rudimentary, about 0.2 mm long aril. Distr. Malaysia: Sumatra, Malay Peninsula, -and Borneo. Ecol. In primary forests at low altitude (to 300 m), on wet to rather dry soil. F/. in the Mal. Pen. from March to May, fr. from April to July; in Sumatra and Borneo ff. from July to Nov., /r. from Nov. to Jan. Uses. The wood is rarely used in house- building. Vern. Mal. Pen.: simpoh, s. jangkang (=stilted s.), merah. Sumatra: bawal (Djambi), bira (Indr.), djangkang (Pal.), gawal, gawar (Indr.), mempélu (Sum. E. Coast), sédjérangkong (Pal.), simar tim- baho darat (Tapan.), simpur, s. kidjang, s. rawang (Pal.). Borneo: bériga, riga (Daj.), djongong (Mal.), entepung rimba (Mal.), ketang bajut (Daj.), kadjang (Mal.), markadjang (Daj.), suretang (Daj.), tém- puru (Daj.). 26. Dillenia borneensis HOOGL. spec. nov. Tree, probably deciduous, up to 40m high, 70cm diam., with stilt-roots. Leaves elliptic to elliptic-oblong or obovate, ca 25—35-nerved, 25—40 by 12-20 cm, with rounded apex, obtuse base, and minutely undulate-dentate margin. Petiole 4~9 cm long, densely sericeously hirsute above. Infloresc- ence a 3-10-flowered composed raceme, forming a loose cluster. Flowers appearing with the leaves, ca 6 cm diam. Sepals 5, elliptic, 1.2-2 by 0.8—1.4 cm. Petals yellow, ca 2!/2 by 1'/2cm. Stamens ca 335, in 2 distinct groups, either the outer group all of the same length, ca 8 mm long, straight in bud, the inner group ca 14mm long, with their apex reflexed in bud, or the stamens of the outer group gradually decreasing in size towards the centre from 11 to 4mm, with their apex more or less inflexed in bud, those of the inner group ca 13 mm long, with their apical part (ca 2 mm) reflexed. Carpels 7-8, ca 8 by 1 mm, with ca 8 mm long styles, each with ca 25-50 ovules. Fruit unknown. Distr. Malaysia: Borneo. Ecol. In primary and secondary forest at low altitude. Vern. Gérige, (Mal.). riga, rogung (Daj.), simpur Dec. 1951] DILLENIACEAE (Hoogland) 167 ee | 4 BR ie | Hn 4 Te | KING with stilt-roots in the Mal. Peninsula. Courtesy For. Res. Fig. 11. Dillenia reticulata Institute, Kepong. 168 FLORA MALESIANA [ser. I, vol. 43 Notes. Most specimens have the second type of androecium:; the first type was found only in one collection, which agrees with the species in all other characters. As, however, the flowers in this specimen (JAHERI 840) are not attached to the spe- cimen, but added loose, I have not given this form varietal rank, though I am convinced that the flowers belong to the specimen. When better known, the two forms may have to be distinguished as separate varieties. The species is strongly char- acterized by the densely hirsute upper side of the petiole. 27. Dillenia reticulata KING, J. As. Soc. Beng. 58, 2 (1889) 367; Ripxt. Fl. Mal. Pen. 1 (1922) 11; Foxw. Mal. For. Rec. 3 (1927) 148 (with 2 plates); Burk. Dict. (1935) 810.—Wormia_ mollissima BoerRL. Cat. Hort. Bog. (1899) 5.—?Dillenia rhizo- phora BoerRLt. & Koorp. in Koorp.—SCHUM. Syst. Verz. 2 (1910) 36.—Fig. 11. Deciduous tree, up to 40 m high, 17 cm diam., with conspicuous stilt-roots. Leaves elliptic or ellip- tic-oblong to obovate, ca 25—35-nerved, 15—30 by 10-20 cm, with rounded to slightly emarginate apex, obtuse to rounded or cordate base, and en- tire to slightly undulate-dentate margin. Petiole 4-10 cm long. Inflorescence a (3—)5—10(—15)-flow- ered composed raceme, forming a loose cluster. Flowers appearing with the leaves, ca 8 cm diam. Sepals 5, broadly elliptic, ca 20-25 by 16-20 mm. Petals yellow, ca 35 by 16 mm. Stamens ca 400-440, all straight in bud, those of the outer whorl ca 11 mm long, those immediately within this whorl 5 mm long, the size gradually increasing towards the centre of the flower to 9 mm in the innermost ones. Carpels 9-10, ca 6 by 11/2 mm, with 5-6 mm long styles, each with 50-70 ovules. Fruit inde- hiscent, greenish yellow, slightly flattened-globular, ca 35 mm diam, 30 mm high including the enclos- ing sepals, which are up to 45 by 42 mm, at the base 6mm thick. Carpels 16 by 8!/2mm, 1-3- seeded. Seeds 31/2 by 2!/2 mm, with a rudimentary, ca 0.2 mm long aril. Distr. Malaysia: Sumatra, Malay Peninsula, and Borneo. Ecol. In primary forests at low altitude, on wet to rather dry soil. The stilt-roots are also developed when the tree grows in a never-flooded habitat. As far as can be derived from the few collections, the flowering- and fruiting-time agree with those of Dillenia eximia Mia. Vern. Mal. Pen.: simpoh, s. jangkang (=stilted S.), S. paya (= marshs.). Borneo: simpur, témpuran. Notes. In the vegetative state the species is not distinguishable with certainty from hirsute forms of Dillenia eximia Mia. Dillenia rhizophora BoeERL. & KoorD. is such a status, most probably referable to the present species. On account of the hairyness of the carpels two varieties can be distinguished: var. reticulata with the carpels hirsute in the apical part, and var. psilocarpella HOoGL. var. nov. with glabrous car- pels. The first variety is known only from the Malay Peninsula, the second from the whole area of the species. 28. Dillenia hookeri Pierre, Fl. For. Coch. 1(1879) t. 5; Fin. & GaGnep. FI. Gén. I.C. 1 (1907) 20; Crals, Fl. Siam. En. 1 (1925) 22. Tree, 10-15 m high, or, more often, low shrub, 1/22 m high, with reddish wood. Leaves oblong to oblanceolate, ca 30-40-nerved, 17—22 by 7-9 cm, with rounded, sometimes slightly acuminate apex, acute base, and entire to slightly dentate margin, densely velvety-tomentose beneath. Petiole 1!/24 cm long. Flowers single, rarely 2, terminal, 4-5 cm diam., the pedicel with 3 verticillate bracteoles; bracteoles lanceolate, 20-35 by 7-10 mm. Sepals 5-6, ovate to elliptic, ca 15 by 10 mm, densely silky hairy outside. Petals yellow, ca 25 by 13 mm. Sta- mens ca 200, all of approximately the same length, 8-10 mm long, with sharply emarginate apex. Car- pels 6-7, ca 5 by 11/2 mm with 11 mm long styles, each with ca 18 ovules. Fruit indehiscent, 2—2!/2cm diam. including the enclosing sepals, which are up to 25 by 14mm. Carpels 14 by 6 mm, 1—5-seeded. Seeds obovate, 31/2 by 3 mm, exarillate. Distr. Indo-China and Siam, N. to ca 17° N, in Malaysia: only in Peninsular Siam. Ecol. In open deciduous forests and savannahs. Notes. Prerre /.c. describes and figures the sta- mens as being of different lengths, arranged in 2 groups. I analysed several flowers, but always found all stamens of approximately the same length. This species is the only Malaysian one, which combines this character with a not amplexicaul leaf-basis. 29. Dillenia luzoniensis (VIDAL) MARTELLI ex DUR. & JAcks. Ind. Kew. Suppl. 1 (1902) 136; Merr. Philip. J. Sc. 1 (1906) Suppl. 95; En. Philip. 3 (1923) 60.—Wormia luzoniensis VIDAL, Rev. Pl. Vasc. Filip. (1886) 36; E-mer, Leafl. Philip. Bot. 7 (1915) 2622.—Tetracera borneensis (non MiQ.) VIDAL, Rev. Pl. Vasc. Filip. (1886) 36. Small tree, ca 5m high. Leaves subcoriaceous, elliptic to oblong, ca 6—12-nerved, 6-15 by 3-7 cm, with rounded to slightly emarginate apex, rounded base, and entire to slightly undulate margin, glossy. Petiole ‘/2-1cm long. Raceme terminal, later lateral, leaf-opposed, with only one flower at the same time, up to 40 cm long with 30 scars of fallen flowers; axis more or less zig-zag. Flowers 6-8 cm diam. Sepals 5, elliptic, 15-18 by 10-14 mm. Petals yellow, 4-5 by 3-4cm. Stamens in 2 distinct groups, the outer ones ca 120, 8 mm long, straight in bud, the inner ones ca 50, 14-20 mm long, with their apex reflexed in bud; on the outside often a few staminodes, 7 mm long. Carpels 7-8, ca 8 by 2!/2 mm, with 10 mm long styles, each with ca 16 ovules. Fruit dehiscent, the sepals pink, enlarged to ca 25 by 15mm. Carpels 17 by 14mm, 1-2- seeded. Seeds ovoid, 3 by 2mm, enclosed by a membranaceous, up to 5 mm long aril. Distr. In Malaysia: Philippines Luzon). Ecol. On riverbanks in forests at low altitude. Notes. As MERRILL (1923) included under the species a number of specimens of Dillenia diantha Hooct. his data on vernacular names and alti- tudinal distribution are not reliable. He gives as the altitudinal limit 1200 m. (Palawan, Dec. 1951] 30. Dillenia excelsa (JACK) GILG in ENGL. & PR. Nat. Pfl. Fam. 3, 6 (1893) 123; Merr. En. Born. (1921) 383; En. Philip. 3 (1923) 60; Heyne, Nutt. Pl. (1927) 1072.—Wormia excelsa JACK, Mal. Misc. 2, 7 (1822) 69; pE Vriese, Pl. Ind. Bat. Or. (1856) 79, t. 6-7; Mia. Fl. Ind. Bat. 1, 2 (1859) 10; Mia. Ann. Mus. Bot. Lugd. Bat. 4 (1868) 76; Koorb. & VAL. Bijdr. 1 Booms. Java (1894) 168; MoLL & JANss. Mikr. Holz. Jav. Baumart. | (1906) 71; Back. Schoolfl. Java (1911) 10; Koorp. Exk. FI. Java 2 (1912) 600; Koorb. & VAL. Atl. Baumarten Java 1 (1913) f.4; Gace & Burk. J. Str. Br. R. A. S. 73 (1916) 243; Corn. Gard. Bull. S. S. 10 (1939) 5; Wayside Trees Malaya (1940) 206; BAcK. Bekn. FI. Java em. ed. 4 (1942) fam. 80, p. 3.— Capellia multiflora BL. Bijdr. 1 (1825) 5.—Capel- lenia multiflora BL. ex Hassk. Cat. Hort. Bog. alt. (1844) 178.—Capellenia pauciflora ZOLL. & Mor. Syst. Verz. (1845-6) 35.—Wormia oblonga WALL. Cat. (1828) no 951, nomen; Hx. f. &. Tu. FI. Ind. 1 (1855) 67; Mia. FI. Ind. Bat. 1, 2 (1859) 11; Hk. Sf. & Tu. FI. Br. Ind. 1 (1872) 35; Kina, J. As. Soc. Beng. 58, 2 (1889) 364; Ripi. Fl. Mal. Pen. 1 (1922) 9.—Wormia grandifolia Mia. FI. Ind. Bat. Suppl. (1860) 619.—Wormia excelsa f. grandifolia Mig. Ann. Mus. Bot. Lugd. Bat. 4 (1868) 76. —Wormia excelsa var. borneensis MiQ. Ann. Mus. Bot. Lugd. Bat. 4 (1868) 77.—Dillenia mag- oliaefolia MARTELLI in Becc. Malesia 3 (1886) 155.—Dillenia glabra MARTELLI in Becc. Malesia 3 (1886) 157; Merr. En. Born. (1921) 383.—Dil- lenia tomentella MARTELLI in Becc. Malesia 3 (1886) 159; Merr. En. Born. (1921) 383.—Dillenia mattanensis MARTELLI in Becc. Malesia 3 (1886) 160; Merr. En. Born. (1921) 383.—Dillenia oblonga GILG in ENGL. & Pr. Nat. Pfl. Fam. 3, 6 (1893) 123.—Dillenia pauciflora GILG in ENGL. & Pr., Nat. Pfil. Fam. 3,6 (1893) 123.—Wormia pau- ciflora KoorD. & VAL. Bijdr. 1 Booms. Java (1894) 169; BAcK. Schoolfl. Java (1911) 10.—Capellia pauciflora ZOLL. & Mor. ex Hook. f. & JACKS. Ind. Kew. 1 (1895) 415.— Wormia tomentella RIDL. JeSte bre RA. S- 33) (1900) 37; 5. Str. Bro R.A-S.- 54 (1910) 5; Fl. Mal. Pen. 1 (1922) 9; Burk. Dict. (1935) 2265.—Dillenia secunda HUNTER (ed. by Rpt.) J. Str. Br. R. A. S. 53 (1909) 98.—Dillenia excelsa var. borneensis MeERR. En. Born. (1921) 383.—Wormia excelsa var. pubescens CORN. Gard. Bull. S.S. 10 (1939) 7.— Wormia excelsa var. tomen- tella Corn. Gard. Bull. S. S. 10 (1939) 8.—Dillenia excelsa var. pubescens CORN. ex MASAMUNE, En. Phan. Born. (1942) 463.—Dillenia excelsa var. tomentella CORN. ex MASAMUNE, En. Phan. Born. (1942) 463. Tree, up to 40 m high with 20 m clean trunk, * 75 cm diam., with gray to brown bark, scurfy with reddish scales, and red to dark brown heartwood. Leaves coriaceous, elliptic to oblong, ca 10—13- nerved, 15-30 by 7-10 cm, with rounded to acute, sometimes slightly acuminate apex, acute, often unequal-sided base, and slightly undulate, entire or dentate margin, glossy. Petiole 2-5cm long. Raceme simple or composed, the only lateral branch at the place of the second flower, up to 12-flowered, sometimes some inflorescences crowd- DILLENIACEAE (Hoogland) 169 ed at the end of a branch and inflorescence there- fore seemingly up to 30-flowered. Flower 7-10 cm diam., the pedicel distinctly thickened at the apex. Sepals 5, elliptic to ovate, 20-25 by 13—16mm, glabrous to rather densely tomentose outside. Petals bright yellow, 40-50 by 25—33 mm. Stamens in 2 distinct groups, the outer ones ca 300, 10-11 mm long, straight in bud, the inner ones ca 30, 16-20 mm long, with their apex reflexed in bud. Carpels 5—10, usually 6-8, 12-14 by 2!/2 mm, with 16-20 mm long, pink styles, each with 20-25 ovules. Fruit dehiscent. Carpels 18-20 by 12-16 mm, yel- lowish green outside, whitish inside, 1—3-seeded. Seeds 6 by 4 mm, dark brown, with a 2-3 mm long, red, membranaceous aril. Distr. Malaysia: Sumatra, Malay Peninsula, Banka, W. Java (E. to Nusa Kambangan), Borneo, and Philippines (Balabac Island, S of Palawan). Ecol. In forests on rather dry to swampy soil, often along streams. Uses. The wood, which has a durability indoors of 15—20 years, is sometimes used in housebuilding. Vern. A large number of vernaculars has been noted: Sumatra: ampalu, a. rimbo (Tapan., Sum. W. Coast), bahah falah (Simalur), boengah-simpur (= flower-s.) (Indrag.), enéhé (Enggano), kalek bakoerok (Tapan., Sum. W. Coast), nilau birrih (Djambi), peileggoa (Mentawei), pisang mawe (Sum. E. Coast), simpur (Djambi, Sum. E. Coast, Palemb., Benk.), s. aréng (Lamp.), s. ayer (Lamp.), s. kimbe (Palemb.), s. Jakki (Banka), s. pérampuan (Lamp.), s. rawang (Lamp., Palemb.), s. rimba (Benk., Banka, Palemb.), s. talang (Palemb.), s. tjipuh (Sum. E. Coast), s. way (Lamp.), sisitoeé (Atjeh). Malay Peninsula: kambai hutan, penaga hlui, simpur, s. ayer, s. padi, s. pagar, Ss. pasir, Ss. paya. Java: djadjatian (Sund.), drégél (Jav.), ki ségél, ségél, sémpur ségél, s. tjai (Sund.), s. batu, s. lanang (Jav.), simpur (Mal.), wuru (Jav.). Borneo: alang-alung (Daj., Punan), djamihing (Mal., Bon- darese), djelangin (Tidung, Batajan), djengin (Mal., Berauw.), djiing, djochin (Daj.), gara (Daj., Ka- puas), kadjamihing (Daj., Kapuas; Mal., Sampit), kajuringin (Daj.), kandikara (kéndikara, kandik- kara) (Mal., Kutinese), kanigara (kénigara, kali- gara) (Bandjar., Bondar., Mal.), kégihing, urib (Daj.), pampan (Dusun rungus), randaman (Banggi), ringin (Daj., Dusun), simporotan (Mal., Br. N. Borneo), simpur (Mal., Br. N. Borneo), s. bukit (Mal., Sarawak), s. Jaki (Mal., Brunei, Sara- wak and Sandakan), tidahura, tjumihing (Daj., Siangese). In the Bahasa Indonesia kéndikara has been accepted. Notes. CORNER (1939, /.c.) distinguishes 2 va- rieties besides the typical form. Dillenia excelsa var. pubescens (CORN.) CORN. ex MASAMUNE is distinguished by the tomentose-hirsute indumen- tum on the outer side of the sepals, continuing on the pedicel and the axis of the inflorescence. This is the commonest form of the species in the Malay Peninsula and is also found in Sumatra, Borneo, and the Philippines. In Borneo intermediates to the typical form with glabrous sepals are frequent. Dillenia excelsa var. tomentella (MARTELLI) CORN. ex MASAMUNE is distinguished by the hirsute indu- 170 FLORA MALESIANA [ser. I, vol. 43 mentum on the intervenium on the lower side of the leaf; it is found in Sumatra, the Malay Pen- insula, Banka, and Borneo. The species is rather variable, but is easily rec- ognised by the rather large, coriaceous, lucid leaves, and the typically thickened apex of the pedicel, which occurs in most species, but is in none so pronounced. 31. Dillenia sibuyanensis (ELM.) MerR. Philip. J.Sc. Bot. 9 (1914) 332; En. Philip. 3 (1923) 62.— Wormia sibuyanensis ELM. Leafli. Philip. Bot. 5 (1913) 1772. Shrub, 3 m high or higher, with smooth bark, scaling off in plates. Leaves rigidly chartaceous, narrowly obovate to oblanceolate, ca 7—9-nerved, 3!/2-7 by 1.1—2!/2 cm, with acuminate apex. acute base, decurrent along the petiole, and entire to slightly undulate margin; glossy above. Petiole 1-1'/2cm long. Flowers solitary or, rarely, in a 2-3-flowered raceme, ca 5cm diam. Sepals 7-9, elliptic to obovate, the outer ones ca 12 by 7mm, the inner ones ca 15 by 10 mm. Petals lemon yel- low, ca 3 by 1.3 cm. Stamens in 2 distinct groups, the outer ones ca 70, 7-8 mm long, straight in bud, the inner ones ca 20, 12-16 mm long, with their apex reflexed in bud. Carpels 5—8, usually 7, bright red, ca 6-8 by 1'/2-1.8 mm, with 12 mm long, white styles, each with 6 ovules. Fruit dehiscent. Carpels 15 by 10mm, 1-seeded. Seeds obovate, 3!/2 by 2.8 mm, enclosed by a yellow, membran- aceous aril. Distr. Malaysia: Philippines (Sibuyan Isl.). Ecol. Scattered along forested river banks from about sea level to 300 m. 32. Dillenia ovata WALL. Cat. (1828) no 945, nomen; HK. f. & Tu. Fl. Ind. 1 (1855) 70; Mia. FI. Ind. Bat. 1, 2 (1859) 12; Hx. f. & Tu. FI. Br. Ind. 1 (1872) 36; PrerRE, Fl. For. Coch. 1 (1879) t. 10; K1na, J. As. Soc. Beng. 58, 2 (1889) 366; BAck. FJ. Batavia 1 (1907) 5; Back. Schoolfl. (1911) 11; Koorb. Exk. FJ. Java 2 (1912) 601; RipL. FJ. Mal. Pen. 1 (1922) 11; Burk. Dict. (1935) 810; Corn. Wayside Trees Malaya (1940) 204; Back. Bekn. Fl. Java em. ed. 4 (1942) fam. 80, 5. Tree, up to 30 m high and | m diam. but usually smaller, 6-12 m high and 30-40 cm diam., with rather knotted trunk and low-attached crown. Leaves ovate or elliptic, ca 18—25-nerved, 10-25 by 7-13 cm, on saplings ca 50-nerved, 30-60 by 14-18 cm, with rounded to obtuse, on saplings acute apex, rounded to acute, often unequal-sided base, and entire to obscurely dentate margin. Petiole ca 2'!/2-5 cm long. Underside of the leaf and petiole hirsute-tomentose. Flower ca 16 cm diam. Sepals 5, elliptic, 25-30 by 16-27 mm, the innermost ones the narrowest, the two outer ones densely villose outside, the third one with one glabrous margin, the inner ones densely villose on the central part only. Petals yellow, 6'/2-7!/2 by 5-6 cm. Stamens in 2 distinct groups, the outer ones, ca 450, 16-20 mm long, straight in bud, the inner ones, ca 25, 25 mm long, with their apex reflexed in bud. Car-~ pels 8-12, usually 10, 6-7 by 2-2!/2mm, with 20 mm long, yellowish white styles; each with ca - Nutt. Pl. (1927) 1072; 50 ovules. Fruit indehiscent, dull yellow, depressed- globose, ca 4'/2 cm high, 6 cm diam. including the enclosing sepals, which are ca 7!/2 by 5!/2 cm, at the base 1!/2 cm thick. Carpels ca 22 by 10 mm, each with 1-7 seeds in glutinous pulp. Seeds obo- void, 5 by 4 mm, black, glabrous, exarillate. Distr. Siam and Indo-China to ca 17°30’ N, in Malaysia: Sumatra, Malay Peninsula (except Ma- lacca and Johore), Banka. In Java cultivated only. Ecol. On sandy or rather dry spots, in belukar, at low alt. Uses. The wood is used in Indo-China for making small objects. Vern. Simpoh ajer (Mal. Pen.), simpur laki (la- laki) (Banka) (= male s.), s. minjak, ranggang wakka (S. Sum.). 33. Dillenia sumatrana Mig. Fl. Ind. Bat. Suppl. (1860) 620.—Dillenia meliosmaefolia Hx. f. & Tu. Fl. Br. Ind. 1 (1875) 36; Ripi: J, Stra Bikeaes: 54 (1910) 7; Fl. Mal. Pen. 1 (1922) 12; HEyYNE, Burk. Dict. (1935) 810; Corn. Gard. Bull. S. S. 10 (1939) 3; Wayside Trees Malaya (1940) 204.— Wormia meliosmaefolia Kina, J. As. Soc. Beng. 58, 2 (1889) 365; Ann. Roy. Bot. Gard. Calc. 5 (1696) 115, t. 130.—Wor- mia parviflora Riv. J. Str. Br. R.A.S. 54 (1910) 6; Fl. Mal. Pen. 1 (1922) 9.—Dillenia elmeri MERR. P]. Elm. Born. (1929) 195. Tree, up to 20 m high, 35 cm diam., with reddish brown bark. Leaves oblong, ca 15-—20-nerved, 16-30 by 7-16 cm, with acute to acuminate apex, acute to obtuse base, and nearly entire to manifestly dentate margin. Petiole 11/2-5 cm long. Flowers solitary or in a 2(—3)-flowered raceme with the flowers attached close to each other, ca 6 cm diam. Sepals 5, ovate, 10-14 by 9-11 mm, densely silky hirsute outside. Petals yellow, ca 25 by 15 mm. Stamens in 2 distinct groups, the outer ones ca 200, 6—7!/2 mm long, straight in bud, the inner ones ca 18-20, 10-11 mm long, with their apex reflexed in bud. Carpels 7-10, white, ca 4-5 by 1'/2 mm, with white, 5-6 mm long styles, each with 15—20 ovules. Fruit indehiscent, yellow or orange-yellow, about globular, 2—2!1/2cm diam. including the enclosing sepals, which are up to 35 by 20 mm, up to 2!/2 mm thick at the base. Carpels ca 12 by 7mm, 1-2- seeded. Seeds 5 by 3!/2 mm, exarillate. Distr. Malaysia: Sumatra (incl. Nias and Siberut Islands), Malay Peninsula (W.-side only), and Borneo (Sarawak and Br. N. Borneo). Ecol. In forests at low altitude to ca 350 m. Uses. The wood is sometimes used in house- building. Vern. Malay Peninsula: simpur, s. bukit, and s. padi. Sumatra: sipang-sipang and duwa saopang (Sum. E. Coast), wahom bouho (Nias), surumak délok (Simalur). 34. Dillenia monantha Me_rr. Philip. J. Sc. Bot. 9 (1914) 321; En. Philip. 3 (1923) 60. Smooth-barked tree, up to 17m high. Leaves subcoriaceous. elliptic-oblong to lanceolate, ca 7—-9- nerved, 8—14 by 3!/2-7 cm, with rounded to obtuse, rarely acute apex, obtuse to acute base, and slightly Dec. 1951] undulate-dentate margin, glossy. Petiole 5-25 mm long. Flowers solitary, terminal, ca 10 cm diam. Sepals 5, 15-22 by 12-15 mm. Petals yellow, ca 50 by 28 mm. Stamens ca 110, the outer ones from 9 mm long, straight in bud, the inner ones up to 15 mm long, with their apex reflexed in bud, with intermediate lengths. Carpels 4-5, ca 7-8 by 1!'/2 mm, glabrous to sparsely hirsute with 0.2-0.4 mm long, rather rigid hairs, each with 11-16 ovules. Styles 8 mm. Fruit dehiscent, carpels ca 15 by 13 mm. Distr. Malaysia: Philippines (from Palawan to Busuanga). Ecol. In dry secondary forests and in open grasslands at low altitudes. Vern. Malacatmon (Palawan). 35. Dillenia indica LINNE, Sp. Pl. (1753) 535; Burm. f. Fl. Ind. (1768) 124; Mia. Ann. Mus. Bot. Lugd. Bat. 4 (1868) 79; Hk. f. & Tu. FI. Br. Ind. 1 (1872) 36; Kina, J. As. Soc. Beng. 58, 2 (1889) 366; Koorp. & VAL. Bijdr. 1 Booms. Java (1894) 161; GAMBLE, Man. Ind. Timb. 2nd ed. (1902) 4; MOLL & Janss. Mikr. Holz. Jay. Baumart. 1 (1906) 71; BaAcK. Schoolfi. Java (1911) 10; Koorp. Exk. FI. Java 2 (1912) 601; Koorp. & VAL. Atl. Baumart. Java 1 (1913) f.2; Merr. En. Born. (1921) 383; Troup, Sylvicult. Ind. Trees 1 (1921) 3; WeEsTER, Philip. Agr. Rev. 14 (1921) 277; Rip. Fl. Mal. Pen. 1 (1922) 10; GitG & WERDERM. in ENGL. & Pr. Nat. Pfl. Fam. ed. 2, 21 (1925) 34, f. 25; HEYNE, Nutt. Pl. (1927) 1072; Burx. Dict. (1935) 809; BLATTER & MILLARD, J. Bomb. Nat. Hist. Soc. 38 (1936) 415, pl. 29, 31 & 32; Some Beautiful Ind. Trees (1937) 45 and 3 pl.; CorN. Wayside Trees Malaya (1940) 204, pl. 52; BAck. Bekn. Fl. Java em. ed. 4 (1942) fam. 80, 4.—Syalita RHEEDE, Hort. Mal. 3 (1683) 39, t. 38-39; ADANs. Fam. PI. 2 (1763) 364.—Dillenia speciosa THUNB. Trans. Linn. Soc. 1 (1791) 200; SM. Exot. Bot. 1 (1804) 3, t. 2, 3; Roxs. FI. Ind> ed. CAREY 2 (1832) 650; Hk. f. & Tu. FI. Ind. 1 (1855) 69; Mia. FI. Ind. Bat. 1, 2 (1859) 11.—Dillenia elongata Mia. FI. Ind. Bat. 1, 2 (1859) 12.—Dillenia indica f. elongata Mia. Ann. Mus. Bot. Lugd. Bat. 4 (1868) 79. Tree, up to 30 m high, 120 cmdiam., with rather crooked trunk and irregular, usually rather lowly (up to 15 m) attached crown. Leaves bright green above, oblong, ca 25—50-nerved, 10-35 by 5—13cm, on saplings and young trees up to 70 by 18 cm, with acute to acuminate apex, rounded to acute base, and slightly to manifestly dentate margin. Petiole 2!/2-7!/2 cm long, on saplings and young trees up to 15 cm. Flower 15—20 cm diam. Sepals — __5, elliptic, 4-6 by 3—5 cm, up to 1 cm thick at the base. Petals white, 7-9 by 5—6!/2 cm. Stamens in 2 distinct groups, the outer ones, ca 550, 13-15 mm long, straight in bud, the inner ones, ca 25, 20—22 mm long, with their apex reflexed in bud; yellow. Carpels 14-20, ca 14 by 3 mm, yellowish green, with linear-lanceolate, white, 25 mm long, up to 31/2 mm broad styles; each with 40-80 ovules. Fruit indehiscent, yellowish green, 8-10 cm diam. including the enclosing sepals, which are up to 15 by 12 cm, 2!/2 cm thick at the base. Carpels ca 35 DILLENIACEAE (Hoogland) 171 by 15 mm, each with 5 or more seeds in colourless glutinous pulp. Seeds reniform, 4 by 6 mm, black, finely echinate, exarillate. Distr. Ceylon, India (Deccan Peninsula, Ben- gal, Assam), S. China (Yunnan), Burma, Siam, Indo-China, in Malaysia: Sumatra, Malay Pen- insula, W. and M. Java, Borneo. Ecol. Mainly on stony banks of rivulets (Sa- raca-streams, CORNER) and rivers, mainly at low elevation up to 500 m. Once collected at 1700 m (Atjeh, Sumatra). Uses. The fruit, in particular the enclosing sepals, is eaten fresh and in curries and jellies. With syrup a cough mixture is made from it. The wood, which is moderately hard and has a dura- bility under water of ca 3 years, is sometimes used for house-building or gunstocks (India). The spe- cies is planted rather often as an ornamental tree. Vern. The Malay name, in use in Sumatra, the Malay Peninsula and Java is simpur or s. ayer (= water s.). In Sundanese the name is sémpur, s. batoe (= stone s.) or s. tjai (= water s.), in Java- nese sémpoe or kosar. In the Philippines, where the species is cultivated, it is known under the Singha- lese name hondapara. Engl.: elephant-apple. Notes. The dispersal of the seeds is said to be effected by animals, among others by elephants (BLATTER & MILLARD), or with current water. In the latter case the seeds may germinate in the fruit, which is left behind on the bank of a river, often partly filled up with mud, which gives a favorable substratum for the germination. 36. Dillenia pulchella (JACK) GILG in ENGL. & Pr. Nat. Pfl. Fam. 3, 6 (1893) 123; Merr. En. Born. (1921) 383; HEYNE, Nutt. Pl. (1927) 1072.— Wormia pulchella Jack, Mal. Misc. 2, 7 (1822) 70; Hk. f. & Tu. FI. Ind. 1 (1855) 68; Mig. Fl. Ind. Bat. 1, 2 (1859) 11; Hx. f. & Tu. FI. Br. Ind. 1 (1872) 36; Kina, J. As. Soc. Beng. 58, 2 (1889) 365; GAGE & Burk. J. Str. Br. R.A.S. 73 (1916) 243; RIDL. Fl. Mal. Pen. 1 (1922) 9; Burk. Dict. (1935) 2265; CorNER, Wayside Trees Malaya (1940) 206.—Dil- lenia micrantha MARTELLI in Becc. Malesia 3 (1886) 156; Merr. En. Born. (1921) 383.—Dillenia parvifolia MARTELLI in Becc. Malesia 3 (1886) 158; Merr. En. Born. (1921) 383. Tree, up to 40 m high with 25 m clean trunk, 90 cm diam., with reddish bark. Leaves elliptic to obovate, 4-8-, usually 6—7-nerved, ca 5-12 by 31/2-61/2 cm, with rounded, sometimes slightly emarginate, to obtuse apex, acute base, decurrent along the petiole, and entire margin. Petiole 1—-1'/2 cm long. Flowers axillary, solitary or 2(—3) serially placed, less often 3-6 on the apical part of a branch in the axil of distinct leaf-scars. Flowers ca 31/2 cm diam., on 2!/2-7'/2 cm long pedicels. Sepals 5, ovate to elliptic, the outer one ca 81/2 by 7 mm, the inner four 12-16 by 8—11 mm. Petals yellow, about oblong, 20 by 9 mm. Stamens in 2 distinct groups, the outer ones ca 110, 4-4'/2 mm long, straight in bud, the inner ones ca 10, 10-11 mm long, dis- tinctly thicker than those of the outer group, with their apex refiexed in bud; opening with lateral longitudinal slits. Carpels 4-6, usually 5, ca 44"/2 72 FLORA MALESIANA [ser. I, vol. 43 by 1-1!/2 mm, with 5!/2-6!/2 mm long styles; each with 7-10 ovules. Fruit dehiscent. Carpels ca 15 by 12mm, 1—2-seeded. Seeds obovate, 3 by 2 mm, enclosed by a red, membranaceous aril. Distr. Malaysia: Sumatra (not in the Lam- pong Distr.), Malay Peninsula, Riouw- and Lingga Archipelagos, Banka, and Borneo (only W. Borneo, Sarawak, and Br. N. Borneo). Ecol. In primary or old secondary lowland fer- ests, on wet, often peaty soil. Seems to be flowering at irregular intervals. Uses. The wood, which is hard and heavy, is of good quality and is used in house-building. Vern. Sumatra: djaha keling (Pal.), dungun (Sum. E. Coast), simar pimasa (Tapan.), simpur paja (Sum. E. Coast). Mal. Pen.: simpur ayer (= water S.), s. paya (= marsh s.), s. paya hitam (= black marsh s.). Riouw-Arch.: pérépat darat. Bor- neo: debak lulus, ilas (Daj.), simpur bukit, s. laki, 5S. paya, s. rimba (Brunei and Br. N. Borneo). Notes. The present species is the only one in Dillenia with axillary flowers. Probably the inflor- escences of a group of cauli- and ramiflorous spe- cies, of which Dillenia pentagyna Roxs. is the only Malaysian representative, are comparable to the 3-6-flowered leafless branches, which sometimes occur in the present species, in which the axis is reduced. 37. Dillenia pentagyna Roxs. Pl. Corom. | (1795) 21, t. 20; Fl. Ind. ed. Carey 2 (1832) 652; Hk. f. & Tu. Fl. Ind. 1 (1855) 71; Mua. Fl. Ind. Bat. 1, 2 (1859) 12; Hx. f. & Tu. FI. Br. Ind. 1 (1872) 38; Kurz, For. Fl. Br. Burma 1 (1877) 21; Pierre, Fl. For. Coch. 1 (1879) pl. 6-8; Koorp. & VAL. Bijdr. 1 Booms. Java (1894) 163; Fin. & GAGNEP. Fl. Gén. I.C. 1 (1907) 19; Back. Schoolfl. Java (1911) 11; Koorp. Exk. Fl. Java 2 (1912) 601; Koorb. & VAL. Atl. Baumart. Java 1 (1913) f. 3; Back. Bekn. Fl. Java em. ed. 4 (1942) fam. 80, 5.—Colbertia coromandeliana DC. Syst. 1 (1818) 435.—Wormia coromandelina SPRENG. Syst. 2 (1825) 631.—Colbertia augusta WALL. Cat. (1828) no 948, nomen; G. DON, Gen. Syst. 1 (1831) 77 (Wormia augusta STEUD. Nomencl. ed. 2, 2, 1821, 789, nomen).—Dillenia augusta Roxs. Hort. Beng. (1814) 43, nomen; Fl. Ind. ed Carey 2 (1832) 652.—Colbertia minor ZoLL. & Mor. Nat. Gen. Arch. N.I. 2 (1845) 579.—Dillenia floribunda HK. ace Eee ind (S55) ile Hike jade Dee Br Ind. 1 (1872) 38 (Wormia floribunda StEUD. Nom. ed. 2, 2, 1821, 789, nomen; Colbertia flori- bunda WALL. Cat., 1828, no 950, nomen).—Dil- lenia pentagyna var. augusta KURZ, J. As. Soc. Beng. 43, 2 (1874) 46.—Dillenia baillonii PIERRE ex LANESS. PI. Util. Colon. Fr. (1886) 281, 702; Fin. & GaGneP. Fl. Gén. I.C. 1 (1907) 19.—Dillenia minor GILG in ENGL. & Pr. Nat.Pfl. Fam. 3, 6 (1893) 125. —Dillenia hainanensis Merr. Lingn. Sc. J. 13 (1934) 64. Deciduous tree, up to 25 m high, with usually rather crooked, up to 12m high clear trunk, to 1m diam. Bark smooth, grayish, peeling off in thin scales. Leaves elliptic-obovate, ca 25-—50- nerved, 20-50 by 10-20 cm, with rounded to obtuse apex, acute base, decurrent along the petiole, and manifestly dentate to nearly entire margin. Petiole 1'/2-5 cm long, slightly winged with permanent, not amplexicaul wings. Leaves on saplings and young plants oblanceolate, up to 75-nerved, up to 120 by 40 cm, with up to 15 mm broad petiolar wings, not sharply separated from the blade, dis- tinctly nerved like the blade. Flowers 2-7 in a fas- cicle on an up to 3 mm long, short shoot with hairy bracts, on branches of 6mm and more diam.; flowering mainly when leafless. Flowers 2!/2-3 cm diam., on a 2!/2-6 cm long pedicel without brac- teoles. Sepals 5, elliptic, 8-12 by 5-9 mm. Petals yellowish or whitish, 15-20 by 5-10 mm. Stamens in 2 distinct groups, the outer ones 60-90, 2!/2-4 mm long, straight in bud, the inner ones 10, 6-9 mm long, with their apex reflexed in bud, opening with longitudinal slits. Carpels 5(-6), ca 3'!/2-4 by 1.2- 1.5mm, with 4mm long styles; each with 5-20 . ovules. Fruit indehiscent, approximately globular, yellow, orange or red, 15 mm diam., 13 mm high including the enlarged, fleshy sepals, which are up to 16 by 14mm, up to 3 mm thick at the base. Carpels ca 8 by 5 mm, 1(—2)-seeded. Seeds ovoid, 5 by 31/2 mm, exarillate. Distr. In monsoon-regions, in continental Asia: India (Deccan Peninsula, N. to United Prov. and Assam), Burma, Andamans, Siam, Yunnan, Indo-China, and Hainan, in Malaysia: M. and E. Java (W to Cheribon), Kangean, S. Celebes (Ma- kassar and Muna Isl.), and Lesser Sunda Islands (Timor, Sumba, and Wetar) (cf. fig. 12). distribution of Dillenia pentagyna Roxs. (disjunct shaded areas). Fig. 12. Geographical Ecol. In primary and teak forests, also in fire- savannahs, up to 1000 m alt., on periodically often very dry localities. Seems to be rather fire-resistant. FI. June-November, at one locality all trees ap- proximately at the same time. Uses. The wood, though of rather good qual- ity, is rarely used because of its crookedness; a rather good charcoal is made of it. Vern. Djunti (Sund.), sémpu (Jav.) and sumpor (Mad., also in Kangean) in Java, rondemiin Muna, _kahalalé and papunuk in Wetar. Notes. The species is the only Malaysian representative of a small group of deciduous trees Dec. 1951] DILLENIACEAE (Hoogland) 173 ss Fig. 13. Dillenia obovata (BL.) HooGL. G. Pantjar, West Java (DE VooGpD). with fascicled flowers; the other species are found only in the monsoon-regions of continental Asia. 38. Dillenia obovata (BL.) HooGL. comb. nov.— Colbertia obovata BL. Bijdr. 1 (1825) 6; HAssk. PI. Jav. Rar. (1848) 175.—Wormia obovata SPRENG. Cur. Post. (1827) 213.—Dillenia ornata WALL. PI. As. Rar. 1 (1830) 21, t. 23; PARKINSON, Ind. For- - ester 61 (1935) 451, pl. 28, f. 1.—Dillenia aurea (non Sm.) Hx. f. & Tu. FI. Ind. 1 (1855) 70, p.p.; DE VRIESE, Pl. Ind. Bat. Or. (1856) 80; Mia. Fl. Ind. Bat. 1, 2 (1859) 12; Ann. Mus. Bot. Lugd. Bat. 4 (1868) 80; Hx. f. & Tu. FI. Br. Ind. 1 (1872) 37, p.p.; PIERRE, Fl. For. Coch. 1 (1879) pl. 11-13; Kina, J. As. Soc. Beng. 58, 2 (1889) 367; Koorpb. & VAL. Bijdr. 1 Booms. Java (1894) 165; FIN. & GaAGneP. FI. Gén. I.C. 1 (1907) 21; Back. FI. Batavia 1 (1907) 21, 385; Schoolfl. Java (1911) 11; Koorp. Exk. Fl. Java 2 (1912) 601; Koorb. & VAL. Atl. Baumart. Java 1 (1913) f. 1; Ripv. FI. Mal. Pen. 1 (1922) 11; HeYNE, Nutt. Pl. (1927) 1071; BurK. Dict. (1935) 809, p.p.; CORN. Wayside Trees Malaya (1940) 201, pl. 50, 51; BAck. Bekn. FI. Java em. ed. 4 (1942) fam. 80, 5.—Dillenia har- mandii GAGNEP. Not. Syst. 6 (1937) 39.—Fig. 13. Deciduous tree, with rather crooked trunk, up to 35 m high with up to 14 mclear trunk, to 70 cm diam. Leaves obovate, dull with a waxy coating beneath, ca 30—45-nerved, 20-40 by 12-20cm, with rounded, on young plants and saplings acute, apex, acute, gradually narrowing base, and entire to slightly dentate margin. Petiole 11/2-4 cm long. Flowers solitary, rarely 2 or 3, at the end of short side-branches, near the basis of the peduncle with a number of bracts, flowering mainly before the appearance of the leaves. Flowers 14-16 cm diam. Sepals 5, elliptic, ca 30-38 by 20-30 mm. Petals bright yellow, 61/2-8 by 41/2-51/2 cm. Stamens in 2 distinct groups, the outer ones yellow, ca 200-240, 12-13 mm long, straight in bud, the inner ones 174 FLORA MALESIANA [ser. I, vol. 4, Dec. 1951] yellowish white, ca 40-55, 22-24 mm long, with their apex reflexed in bud. Carpels ca 9-11, ca 13 by 2 mm, with 20 mm long styles, each with 25-35 ovules. Fruit indehiscent, yellow or orange-col- oured, approximately globular, 35-40 mm diam. including the enlarged fleshy sepals, which are up to 50 by 40 mm, 6 mm thick at the base. Carpels ca 25 by 10mm, 1- to few-seeded. Seeds 5 by 3!1/2 mm, exarillate, embedded in transparent slime. Distr. Lower Burma, Indo-China, and Siam, in Malaysia: Northern part of the Malay Peninsula, S. Sumatra, and W. Java. Ecol. In secondary forests, on stony, period- ically rather dry soils. F/. at the end of the relatively dry period, about April in the Malay Peninsula, July to October in S. Sumatra and W. Java. Uses. The fruit is eaten in curries, the wood sometimes used in house-building. Vern. Mal. Pen.: simpur. Sumatra: simpur, s. bénér, s. rimba, s. talang. Java: sémpu (Jav.), sém- pur, s. batu (Sund.). Notes. Dillenia aurea SM. is a closely re- lated species, occurring in NE. India and N. and Central Burma; cf. PARKINSON (1935). The differ- ences in reproductive parts are slight, but the leaf- shape in adult state is very characteristic. Excluded and doubtful Dillenia grandifolia WALL. Cat. (1828) no 946, no- men; HK. f. & TH. Fl. Ind. 1 (1855) 71; Mia. FI. Ind. Bat. 1, 2 (1859) 12; Hx. f. & Tu. FI. Br. Ind. 1 (1872) 38; KiNG, J. As. Soc. Beng. 58, 2 (1889) 368; RipL. J. Str. Br. R.A.S.59 (1911) 61; Fl. Mal. Pen. 1 (1922) 11; Cram, Fl. Siam. En. 1 (1925) 22; Corn. Wayside Trees Malaya (1940) 203. The type-collection consists of leaves from a young tree or sapling and is referable either to Dillenia ovata WALL. ex HK. f. & TH. or to Dillenia reticulata KING. Most of the later literature refers to specimens belonging to the latter species. From study of the leaves of saplings of both species, which come into consideration, the identity must be decided. Dillenia indica var. aurea (SM.) O.K. Rev. Gen. Pl. 1 (1891) 4. KUNTZE, when making this new combination, had a specimen of Dillenia ovata WALL. ex. HK. f. & Tu., from cultivation in the Botanic Gardens at Bogor. Dillenia aurea SM. is an Indian species; cf. sub D. obovata (BuL.) HOOGL. Tetracera tripetala Turcz. Bull. Soc. Nat. Mosc. 36, 1 (1863) 547. The description is insufficient for identification. Of the Javanese species Tetracera fagifolia BL. comes nearest to it; most probably the species will have to be reduced to it. CAPRIFOLIACEAE (J. H. Kern, Bogor, and C. G. G. J. van Steenis, Leyden) Small trees, shrubs or twining woody plants, rarely herbs; branches terete. Glands present in various parts. Indumentum consisting of simple hairs, or in Viburnum sometimes lepidote; glandular hairs mostly present. Stems often pithy. Leaves decussate, simple or deeply divided (Sambucus), sometimes provided with pitted or cup-shaped glands exuding resin. Stipules absent or very small. Flowers 9, actino- morphic or zygomorphic, mostly cymosely arranged, 4-5-merous; outer flowers in an inflorescence sometimes differing from the normal ones, rarely (Sambucus p.p.) some fis aborted into extra-floral nectaries. Calyx adnate to the ovary, (4-)5-fid or -toothed, mostly constricted below the limb; sepals often enlarged in fruit. Corolla epigynous, gamopetalous, sometimes 2-lipped, lobes mostly imbricate in bud. Stamens inserted on the corolla tube, alternating with the lobes, extrorse or introrse. Anthers free, 2-celled, dorsifixed, versatile, cells parallel, opening length- wise, mostly introrse; filaments sometimes reflexed or curved in bud. Ovary infe- rior, 1-(2-)3—5(—8)-celled, in fruit cells sometimes partly abortive. Style terminal, often slender with one knoblike stigma, or 3 short partly connate styles. Ovules 1(—~), pendulous or axile. Fruit a drupe or berry, rarely a capsule. Seeds often only one per fruit, often with bony testa. Endosperm copious, sometimes ruminate; embryo straight, often small and linear, axial, cotyledons oval or oblong. Distr. Ca 10-14 genera, mainly distributed on the N. hemisphere, in the tropics mostly confined to the mountains, on the S. hemisphere only Viburnum and Sambucus, an endemic genus in New Zealand, two monotypic endemic genera in New Caledonia, in Australia only Sambucus in the eastern part. Ecol. Caprifoliaceae do not play an important role in Malaysian vegetation as to numbers: both Lonicera and some spp. of Sambucus may predominate locally in mountain thickets and clearings. Most members of the family are insect-pollinated; Lonicera has a nocturnal fragrance; the exact function of the peculiar metamorphozed flowers in Sambucus javanica is not known. Uses. Many spp. of all three genera treated here are used as ornamentals. The honey-suckles (Loni- cera) have mostly fragrant flowers. _ Notes. The family seems to be most related to the Valerianaceae to which the foliage of Sambucus and the occurrence of valerianic acid in Viburnum add in importance, but there is no unanimity on the delimitation of the family. BAILLON (Hist. Pl. 7, 1880, 352 seg.), MCATEE (Bull. Torrey Bot. Club 48, 1921, 149), and FritscH (Bot. Centr. BI. 1892, ii, p. 169) are of opinion that Caprifoliaceae ought to be merged into Rubiaceae. The occurrence of interpetiolar stipules or appendages accepted to represent stipules obscure a clear distinction. I fail to understand why Kurz can eliminate Scyphiphora from the Rubiaceae and refer it to Caprifoliaceae (J. As. Soc. Beng. 45, ii, 1876, 133; For. Fl. 2, 1877, 4) ‘by the structure of the ovary and position of the ovules which are tell-tale marks of its caprifoliaceous descent’, as the gynoecium is very variable in Rubiaceae. Moreover, Scyphiphora possesses stipules bearing colleters (see below). With Carlemannia and Sylvianthus (the latter absent from Malaysia) the case is different; HOooKErR f. and PirarD included them in Rubiaceae. SOLEREDER in his valuable study on the anatomy of the rubi- aceous complex showed (Bull. Herb. Boiss. 1, 1893, 171, 173, 174) that they belong to Caprifoliaceae, by absence of stipules and presence of capitate-glandular hairs typical of Caprifoliaceae. He is followed by HALLER f. and BREMEKAMP. However, a general merging of Caprifoliaceae into Rubiaceae seems to us undesirable, as by that procedure the various caprifoliaceous genera would be assigned to various rubiaceous tribes, and come to remote positions not reflecting their affinity. Another question is whether, contrarily, genera hitherto accepted as rubiaceous ought to be removed to Caprifoliaceae. SOLEREDER (i.c.), followed by KRAusE (Ber. D.B.G. 28, 1909, 446-452) and GLUck (Blatt- u. Bliitenmorph. Stud. 1919, 135) have focussed attention to the regular and typical occurrence of resin-glandular bodies, or trichomes, on the adaxial basis of Rubiaceous stipules, called colleters. Col- leters are known in a limited number of other families and genera such as Cunoniaceae, Rhizophoraceae, Caesalpiniaceae, and, as was found recently, are also typical for Nothofagus. Dr BAKHUIZEN VAN DEN BRINK Jr, who is now revising Rubiaceae for this Flora, has found them in all genera hitherto studied except a few, viz: Dentella, Hedyotis, Borreria, Spermacoce, Richardsonia, Anotis, Allaeophania, and Diodia, of which genera some appear to be aberrant in Rubiaceae on account of their ovules, by which character they come now wide apart in rubiaceous taxonomy, whereas in other points they appear 176 FLORA MALESIANA [ser. I, vol. 43 closely allied (cf. also BOERLAGE, Handl. 2, 1891, 9, 10 & 19). The research on the problem whether these are really rubiaceous is one of time-consuming nature and of too great importance to be solved prematurely. Pending continued research the present revision is published in the old circumscription of Caprifoliaceae. It seems desirable to make additional karyological investigations on genera eventually to be joined to Caprifoliaceae as in the old circumscription the basic chromosome numbers seem to be rather constantly 8 and 9 (cf. SAx & Kriss, J. Arn. Arb. 11, 1930, 147-152). An account of pollen struc- ture seems also desirable. The genus Gaertnera is distinct from Rubiaceae by a superior ovary and absence of colleters, and will be treated, in this Flora, as loganiaceous, though SOLEREDER (Ber. D.B.G. Gen.-Vers. Heft 1890, p. 70; Bull. Herb. Boiss. 1, 1893, 169) assigns it to Rubiaceae. In both families it keeps an isolated position. The family appears to us a coherent and natural one: minor though by no means neglectable charac- ters common to all are: glandular hairs and glands in other parts, enlarging sepals, a distinct constriction between ovary and sepals, and dorsifixed anthers, a character hitherto not specially stressed in literature (in Sambucus the cells are free but fixed in the middle!). Leaves of juvenile forms of some species are occasionally distinctly lobed, e.g. in Symphoricarpus and Lonicera. In collecting Caprifoliaceae it is urgent to collect both full-grown flowers and ripe fruits. The first author is responsible for the revision of Viburnum, the second author for the rest. KEY TO THE GENERA ipelcavesinimnatery a gee vou tebaay Pic HenMe de ee OSE iA yr AOS SMe oe 3. Sambucus 1. Leaves simple. 2. Erect herbs. Corolla 4-merous. Stamens 2. Capel dehiscing; 3) e . . . 4, Carlemannia 2. Shrubs, small trees, or lianas. Corolla 5-merous. Stamens 5. Drupe or berry. 3. Ovary-cells with more than 1 ovule. Corolla 2-lipped, at least 12 mm long. Style simple, long, with a capitate stigma. Berry . . . . 1. Lonicera 3. Ovary-cells with one ovule. Flowers actinomorphic, at most 10. mm i long. Style 3-fid, short. Drupe. 2. Viburnum 1. LONICERA LINNE, Sp.PI. (1753) 173; Gen. Pl. (1754) no 210; DC. Prod. 4 (1830) 330; REHDER, Rep. Mo. Bot. Gard. 14 (1903) 27-232, pl. 1-20; STEEN. J. Arn. Arb. 27 (1946) 443.—Caprifolium ZINN. Cat. Pl. Gott. (1757) 10. Shrubs or (in Malaysia exclusively) woody climbers twining to the right, or, in the absence of sufficient support, locally scrambiing shrubs; bark at last lengthwise splitting, twigs mostly contorted. Leaves hairy or glabrous, entire (in the Malaysian spp.), free (or in extra-Malaysian subg. Periclymenum) connate. Stipules absent, but leaf-bases connected by a raised line on the node. Flowers 5-merous, mostly sessile, in 2-flowered, axillary, peduncled or rarely almost sessile cymes, not rarely specially developed towards the end of the branches and forming a leafy terminal panicle. Cymes occasionally 3-flowered. Each pair of flowers subtended by 2 bracts and 4 bracteoles, the latter sometimes covering the ovary. Calyx-tube ovoid or subglobose, teeth mostly small. Corolla elongated, mostly 2-lipped, upper lip repre- senting 4 lobes, and mostly 4-lobed at its apex, lobes imbricate in bud. Stamens 5, anthers introrse, mostly exserted; filaments inserted near the apex of the tube. Ovary 2—3-celled, style filiform, elongated, usually exsert, and exceeding the sta- mens, stigma capitate. Ovules axile, 3-8 per cell, pendulous. Berries mostly few- seeded, free or (in extra-Malaysian spp.) connate in pairs. Seeds generally ovoid with fleshy albumen and a terete embryo. Distr. Ca 150 spp. described from the N. hemisphere, centering in the Himalayas to Central and E. Asia, not in Ceylon, crossing the equator only in Malaysia. The Malaysian spp. belong to § Nintooa DC. comprising -+ 30 spp. centering in SE.-E. Asia, with 1 sp. in the Mediterranean. Ecol. In Malaysia only found in the W. part, confined to the zones above 1000 m alt. Uses. Two spp. are used widely as ornamentals. Wood anat. CHALK & SCHATTAWAY, Proc. Roy. Soc. B 113 (1933) 82. Notes. In the absence of sufficient support specimens may turn to scrambling, but they are in the forest true lianas. Dec. 1951] CAPRIFOLIACEAE (Kern & vy. Steenis) La For measurements of flowers, full-grown open flowers should be used. All Malaysian spp. including the cultivated ones belong to § Nintooa. In my eye REHDeR has distinguished too many spp. in this section. His subsections Longiflorae and Breviflorae 1 think are rather artificial. KEY TO THE SPECIES 1. Bracts below each pair of flowers foliaceous 1. Bracts subulate to lanceolate, not leafy. 2. Ovary hairy all over 2. At most the extreme apex of the ovary hairy. 1. L. japonica 2. L. confusa 3. Flowers in the dried state at most -| 2!/2 cm long. 4. Flowers slender, 2-4 on slender axillary peduncles. Flowering parts hairy and besides with capitate glandular hairs. In anthesis both lips recurved. Internodes slender, twigs soon shiny-brown. 7. L. javanica 4. Flowers thickish, crowded towards the twig-ends, peduncles short, congested towards the twig- ends, flowers not exceeding the leaves. Capitate glandular hairs absent or negligible. In anthesis only the lower lip recurved. Internodes stiffish, twigs not shiny-brown 6. L. acuminata 3. Opened flowers in the dried state at least 3 cm long. 5. Plant entirely glabrous (except few negligible hairs on stamens and style and some ciliae on bracts and calyx). Nerves few, not prominent, reticulations indistinct . 3. L. sumatrana 5. Plants hairy, nerves and reticulations distinct. 6. Inflorescences many-flowered, mostly congested into a globose or oval leafy panicle. Floral parts with subsessile, capitate, glandular hairs. Leaves glaucous-tomentose underneath, nerves and reticulations impressed above . 5. L. pulcherrima 6. Peduncles axillary, 2-flowered. Capitate glandular hairs absent. Nerves and reticulations hirsute underneath, not distinctly impressed above 1. Lonicera japonica THUNB. FI. Jap. (1784) 89; Mia. Ann. Mus. L.B. 2 (1866) 269; REHDER, Ann. Mo. Bot. Gard. 14 (1903) 159; Bor & RAIZADA, J. Bombay N. H. Soc. 44 (1943) 76, f. 3; STEEN. J. Arn. Arb. 27 (1946) 444, in clavis; BACKER, Bekn. Fl. Java, em. ed. 8 (1949) fam. 175, p. 6.— Lonicera chinensis WATSON, Dendr. Brit. 2 (1825) t. 117; HaAssk. Flora 28 (1845) 242; Fier, Pl. Bot. Tuin Weltevreden (1855) 60; Mia. FI. Ind. Bat. 2 (1856) 127; Kurz, Nat. Tijd. N.I. 27 (1861) 198.— L. repens Zipp. ex Hassk. Cat. Hort. Bog. (1844) 116, in syn.; Miq. FI. Ind. Bat. 2 (1856) 128; STEEN. I.c. 451; Blumea 6 (1948) 243. Twining. Twigs short-pubescent, glabrescent, sparsely glandular-hairy, shiny-brown, ultimate internodes short-tomentose. Leaves ovate-oblong to .ovate-lanceolate, base rounded to truncate, apex acute, acutish or subacuminate, midrib and edges pilose, darkgreen and shiny above; 3—8!/2 by 11/2-4 cm, petiole 3/4-1 cm. Flowers fragrant, 3—5 cm. Peduncle !/4-11!/2 cm, tomentose. Bracts pet- ioled, leafy, oblong-ovate, or oblong-obovate, 10-18 by 4-8 mm. Bracteoles roundish, 3/4 mm, hairy, glandular, ciliate. Calyx-lobes linear from a triangular base, haired, + 11/2 mm, ciliate. Corolla 4-5 cm, creamy, or pale rosa-tinged towards the base, fading orange-yellow, lobes as long as the tube, hairs mixed with capitate glandular hairs, tube + 2!/2 cm, lobes 1/3 of the limb of upper lip. Style glabrous, + as long as the corolla. Stamens glabrous. Berry globular, 6-7 mm diam., black. Distr. From Yunnan to Formosa & Japan, in Malaysia: cultivated as an ornamental up to + 1000 m. Ecol. Fruitisrarely set in Malaysia, fl. Jan._Dec. Uses. BURKILL (Dict. 1935, 1363) says that at Singapore Chinese import flowers from China; they are said to possess antifebrile, corrective and 4. L. malayana astringent properties. The vegetative parts contain a saponin. Notes. An early introduction, possibly via the Botanic Gardens at Bogor, as far as known never run wild. Many varieties have been distinguished (REHDER, /.c.). 2. Lonicera confusa DC. Prod. 4 (1830) 333; ReHpDeER, Ann. Mo. Bot. Gard. 14 (1903) 156; Merr. En. Born. (1921) 582; Bor & RAIZADA, J. Bomb. N.H. Soc. 44 (1943) 75, f. 2; STEEN. J. Arn. Arb. 27 (1946) 444 in clavis; BACKER, Bekn. FI. Java. em. ed. 8 (1949) fam. 175, p. 6.—L. japo- nica (non THUNB.) ANDR. Bot. Rep. 9 (1809) t. 583. —L. multiflora CHAMP. in Hook. J. Bot. Kew Misc. 4 (1852) 167. Whole plant short-tomentose, with the exception of the upperside of the leaves which is sparsely pubescent. Twigs + eglandular. Leaves ovate- oblong to oblong, base rounded, truncate, or sub- cordate, apex acute or acutish, margin tending to recurve, texture tending to be bullate by impressed nerves and reticulations, 3—6 by 11/2-3 cm. Petiole 1/>-11/4 cm. Inflorescences lateral, condensed, + 7-flowered or at the end of twig multiflorous, pro- vided with reduced leaves, exceeding the leaves, on 1-2 cm long stalks; cymes + sessile or some mm peduncled. Bracts linear, as long as or longer than the ovary, rarely subspathulate. Bracteoles sub- orbicular, 1 mm, hairy. Calyx-lobes narrow-trian- gular, hirsute, shorter than the ovary. Corolla 4—5 cm, pubescent, lobes + as long as the slender tube, provided with capitate glandular hairs; apex of the upper lip split for '/s of its length, lobes ovate. Style glabrous. Stamens only sligthly hairy towards the base. Distr. China, Hainan, in Malaysia rarely cul- tivated as an ornamental. 178 FLORA MALESIANA [ser. I, vol. 43 Note. Closely allied to L. japonica, in one spe- cimen I found subspathulate bracts tending to become foliaceous. 3. Lonicera sumatrana Mia. FI. Ind. Bat. Suppl. (1860) 213, 537; REHDER, Rep. Mo. Bot. Gard. 14 (1903) 149; Steen. J. Arn. Arb. 27 (1946) 445.—L. leiantha Kurz, J. As. Soc. Beng. 43, II (1874) 188; For. Fl. Burma 2 (1877) 3; REHDER, /.c. 163; CRAIB, Fl. Siam. En. 2 (1932) 5.— Caprifolium sumatra- num O.K. Rev. Gen. 1 (1891) 274.—C. leianthum O.K. /.c.—L. jasminifolia MERR. Pap. Michigan Ac. Sc. 19 (1934) 199. Slender, + entirely glabrous. Internodes 4~7 cm. Leaves subtriplinerved at the base, without dis- tinct reticulations, thickish, narrowed towards the base, elliptic to oblong-elliptic, acuminate, 4-9 by 2-3 cm, nerves 4-5 on either side, not prominent; petiole 1/3-1 cm long. Peduncles axillary and ter- minal, 1—11/2 cm long, 2-flowered; at the twig-ends sometimes 4 + together. Flowers slender, glabrous except some small negligible ciliae on the edge of the calyx and bracts, white or rosa-like beige. Bracts half as long as the ovary, lanceolate-acumin- ate, 1!/2-2 mm. Bracteoles suborbicular-ovate, 1 mm, blunt. Calyx-tube (ovary) constricted at the apex, free part shorter than the ovary, split halfway down, lobes ovate, subacute, 1 mm. Corolla slen- der, 5'/2-6!/2 cm long; tube 31/24 cm, upper lip split into 4 lobes + 1/s—'/6 of its length. Style sub- glabrous with reflexed, sparse, white hairs, 6-7 cm long. Stamens subglabrous, distinctly exsert. Distr. S. Siam (Puket), Ava (Burma), Yunnan, in Malaysia: Sumatra (rare). Ecol. Thickets and forest borders in the moun- tain regions, 1000-1250 m, f7. Apr.—June. Vern. Kaju kaleh simienjak (Alahanpandjang). Notes. REHDER places this well-defined sp. in- correctly in subsect. Breviflorae because of the fact that Mique. described the flowers after specimens in the immature bud state. Although I have not seen the type specimen of L. leiantha Kurz the description is wholly fit for Sumatran plants, at which MERRILL has already hinted. 4. Lonicera malayana HENDERSON, J. Fed. Mal. Stat. Mus. 11 (1924) 187; Rip. Fl. Mal. Pen. 5 (1925) 313; STEEN. Blumea 6 (1948) 243; HEN- DERSON, Mal. Nat. J. 6, 1 (1950) 189, f. 170. Scrambling shrub, branches hirsute by long spreading, and short hairs, intermixed with very few + sessile glandular-capitate hairs. Leaves oblong- elliptic, not bullate, but nerves and reticulations im- pressed above, base rounded, apex short-acumin- ate, 6-11 by 23/4-5 cm, glabrous above except the base of the midrib, spreading-hirsute underneath on nerves and reticulations, and with short-stalked, sparse capitate-glandular hairs, edge + flat, fim- briate by + 2mm long hairs; nerves 5-6 pairs; petiole 5-7 mm, yellow-hairy. Peduncles axillary, 11/2-31/2 cm, yellow-hairy, 2-flowered. Bracts linear-lanceolate, + 3 mm, hairy, as long as the ovary, twice as long as the ovate-acuminate, 1!/2-2 mm similar bracteoles. Flowers hairy, white fading to pale yellow, 5-6 cm long. Calyx-tube urceolate, drying bluish, glabrous, teeth linear, hirsute, 11/2-2 mm long. Corolla-tube 3!/2-41/2 cm long, very thin (i-1'/2 mm diam.), straight, hirsute by reflexed rather appressed, setaceous yellow hairs, eglan- dular, yellow-strigose inside, upper lip strongly in- rolled-recurved, + 15 by 31/24 mm, lobes ovate blunt, -+- 2 mm, lower lip spreading, ca 6 by 1 mm. Filaments glabrous, + 2!/2cm exserted, anthers 3 mm, linear. Style glabrous, long-exserted, 71/2~8 cm, stigmatic knob distinct. Distr. ?Hainan, in Malaysia: Malay Peninsula (Pahang), twice collected. Ecol. Scrambling over bushes by Bertam River, ca 1000 m, ff. April-June. Notes. My former referring this sp. provision- ally to L. pulcherrima has been entirely wrong; it is perfectly distinct from all other Malaysian spp. The affinity of this sp. seems to be with the E. Asiatic L. affinis Hook. & ARN., L. similis HemsL., and L. macrantha Spr. From L. siamensis GAMBLE differing by long-exsert stamens and -style, from L. similis and macrantha by thinner, eglandular corolla. F. C. How & N. K. CHUN 70187 from Hainan seems an exact match. 5. Lonicera pulcherrima RipL. J. Mal. Br. R. As. Soc. 1 (1923) 64; Merr. Contr. Arn. Arb. 8 (1934) 165; Steen. J. Arn. Arb. 27 (1946) 449, excl. syn.; Blumea 6 (1948) 244. Climbing or scrambling, branches short-tomen- tose. Leaves ovate, blunt or acute, base shortly narrowed, glabrous above, short glaucous-tomen- tose beneath, nerves and reticulations impressed above, 3-7'/2 by 1!/2-5!/2 cm. Petiole tomentose, 1/2-]1/2 cm. Flowers in terminal-congested, many- flowered leafy panicles exceeding the leaves, stalks and flowering parts with subsessile, red, capitate- glandular hairs interspersed in the tomentum. Bracts narrow-lanceolate, 2 mm, tomentose. Brac- teoles + orbicular, + 1 mm, shorter than the ovary, tomentose. Calyx-tube ellipsoid, glabrous except near the apex, lobes lanceolate, as long as the tube, 1!/2-2 mm. Corolla pale yellow turning orange with age, tube 2-2!/2cm, slender, subangular, tomentose, lips -+_ 1!/4-13/4 cm, upper lip incised to ‘/4-!/5. Stamens far exsert, filaments sparsely hairy towards the base, anthers 3 mm. Style far exsert, glabrous or occasionally with a few negli- gible, spreading hairs. Distr. ?China, in Malaysia: Sumatra (Atjeh, Tapanuli, Eastcoast). Ecol. Streamborders, forest borders, rocky places, 850-1400 m, ff. Sept.—Febr. Notes. Seems to occur in continental Asia and should be compared with L. leschenaultii WALL. from India. L. reticulata (non RAFIN.) CHAMP. from Kiangsi (LAU 4649) seems to be an exact match. open 6. Lonicera acuminata WALL. in Roxs. FI. Ind. 2 (1824) 176; DC. Prod. 4 (1830) 334; Mia. Ann. Mus. L.B. 2 (1866) 270; CLARKE in Hook. f. FI. Br. Ind. 3 (1880) 10; REHDER, Ann. Rep. Mo. Bot. ~ Gard. 14 (1903) 150; STEEN. J. Arn. Arb. 27 (1946) 445: Blumea 6 (1948) 244.—Caprifolium loureiri BL. Bijdr. 13 (1826) 653.—L. loureiri DC. Prod. Dec. 1951] CAPRIFOLIACEAE (Kern & v. Steenis) 179 Fig. 1. a. Lonicera acuminata WALL. 4 (1830) 334; Hassx. Flora 28 (1845) 241, incl. var. oblonga; Hook. Ic.P1. 9 (1852) t. 806, incl. var. major; Hook. f. & Tu. J. Linn. Soc. Bot. 2 (1858) 172; REHDER, Ann. Rep. Mo. Bot. Gard. 14 (1903) 149; Koorpb. Exk. Fl. Java 3 (1912) 287; BACKER, Bekn. FI. Java em. ed. 8 (1949) fam. 175, p. 2.—L. flavescens JUNGH. Java ed. 2, neerl. 2 (1854) 407, nomen.—L. oxylepis Mia. Fl. Ind. Bat. 2 (1856) 125, incl. var. oblonga (HAssK.).—L. leschenaultii (non WALL.) Mia. /.c. 126.—L. henryi HEMsL. J. Linn. Soc. 23 (1888) 359; REHDER, /.c. 148.—Caprifolium henryi O.K. Rev. Gen. 1 (1891) 274.—C. acumina- tum incl. var. normale, loureiroi, oxylepis O.K. Rev. Gen. 1 (1891) 273.—L. giraldi REHDER, /.c. 150.— L. rehderi Merr. Govt Lab. Publ. (Philip.) 29 (1905) 49; Philip. J.Sc. 5 (1910) Bot. 391; En. Philip. 3 (1923) 578.—L. philippinensis MEeERR. Philip. J.Sc. 1 (1906) Suppl. 240.—L. transarisa- nensis HAyYATA, Ic. Pl. Form. 6 (1916) 25.—L. vestita W.W.SM. Not. R. Bot. Gard. Edinb. 10 (1917) 49.—L. mindanaensis MeErR. Philip. J.Sc. 20 (1922) 471.—L. ‘javanica DC.’, Doct. v. LEEUWEN, Verh. Kon. Ak. Wet. A’dam 31 (1933) 235-237, f. 61-62, pro forma 9.—Fig. 1a, 2. Woody climber or scrambling shrub. Twigs patently hirsute. Internodes rather short. Leaves ovate-oblong to lanceolate, variable in shape, base rounded, truncate or subcordate, apex acute to acuminate, texture often bullate by impressed nerves and reticulations, both sides green, hairiness in various degrees but midrib above always hairy, x 3/4, b. Lonicera javanica DC. x 3/4, c. Carlemannia tetragona Hook. f., fruit x 4,.d-g. seeds of Sambucus: d. S. canadensis L., e. S. adnata WALL., f. S. javanica Bu., g. S. coerulea RAFIN., all x 5. 3-8 by 11/24 cm; petiole patent-yellow hairy, '/4-1 cm. Cymes mostly contracted towards the twig- ends, infil. not exceeding the leaves. Peduncle densely patent-hirsute, sometimes with few sessile glands. Bracts 5-8 mm rarely shorter, narrow tri- angular, as long as or longer than the ovary. Brac- teoles 2 mm, acute oblong, shorter than the ovary. Calyx green, lobes ciliate, with a few stiff hairs on the back. Corolla 2—2!/2 cm long, sulphureous, not fragrant, rather thick and club-shaped in bud, in anthesis upper lip erect, lower lip refiexed; tube glabrous or with few reflexed strigose hairs, some- times with few sessile glands. Style + as longas the corolla, mostly hairy. Stamens mostly hairy. Berry black. Distr. India to S. China and Formosa, in Ma- laysia: Sumatra, Java, Bali, Philippines (Luzon). Ecol. Forest borders, thickets, ericoid crooked forest, mossy forest, 1800-3300 m, ff. Jan.—Dec. Vern. Ki seroh, S. Notes. Both L. acuminata and L. javanica show the remarkable ‘mass-elevation effect’ (cf. J. Arn. Arb. 27 (1946) 447; Fl. Mal. 4 (1949) xlix, f. 37). Cf. fig. 3. In Java the species exclude each other altitudinally, and represent vicarious species (fig. 3). On the summit of Mt Pangrango, W. Java, DocTERS VAN LEEUWEN studied the variability (/.c.). He noted the occurrence of 9 fis the anthers of which do not contain pollen. He also mentioned the insect pollination by Bombus. Part of his ob- servations belong to L. javanica. Aberrative forms 180 FLORA MALESIANA [ser. I, vol. 4 I found in the herbarium are with 3-verticillate leaves, and others with occasionally flowers in triads. A duplotype in fruit at Kew leaves no doubt about the identity of L. mindanaensis MERR. Fig. 2. Lonicera acuminata WALL. on summit of Mt Pangrango, W. Java (DOCTERS VAN LEEUWEN). 7. Lonicera javanica (BL.) DC. Prod. 4 (1830) 333; Mio. FI. Ind. Bat. 2 (1856) 125; REHDER, Ann. Rep. Mo. Bot. Gard. 14 (1903) 157; Koorp. Exk. FI. Java 3 (1912) 287; Doct. v. LEEUWEN, Proc. Kon. Ak. Wet. A’dam 31 (1933) 237, f. 60 pro forma ¢; STEEN. J. Arn. Arb. 27 (1946) 450; BACKER, Bekn. Fl. Jav. em. ed. 8 (1949) fam. 172 p.2.—Caprifolium javanicum BL. Bijdr. 13 (1826) 653.—Fig. 1b. Slender, twining. Ultimate twigs short-hairy, mixed with capitate-glandular hairs, internodes soon glabrate, shiny-brown, rather long. Leaves ovate to obovate, or ovate-oblong, base truncate, rounded or cordate, apex acute to acuminate, upperside glabrous or the base of the midrib hairy, beneath glaucous, short-pubescent, 4-11 by 2— 6!/2 cm; petiole !/2-1 cm, pubescent. Inflorescences short-grey-hairy, mixed with distinctly stalked capitate-glandular hairs, terminal and in the upper axils so as to form a rather lax, leafy panicle, exceeding the leaves; lower branches up to 6 cm, with reduced leaves, upper internodes not abbreviated. Flowers creamy, fragrant, later fading into yellowish. Bracts linear-lanceolate, as long as or longer than the calyx. Bracteoles ovate, shorter than the ovary. Calyx purplish, lobes lanceolate, 11/2-2 mm, the lobes mostly rather densely grey-hairy, sometimes only ciliate. Bud club-shaped, thickened end + '/3—!/s of the mature bud, often acute. Corolla 18-30 mm (fresh 3—31/2 cm) long, the tube slender, 1 mm or thinner, 1-2 cm, lobes narrow, 8-14 mm long, -+ 1—2!/3 mm broad, during anthesis both lips recury- ed, lobes as long as the tube or shorter; hairs on the corolla recurved, except the capitate glandular ones. Anthers thin, 2-4 by 1/3!/2 mm. Stamens protruding, glabrous. Style often longer than the corolla, glabrous, exserted. Berry dark purple, ripe prob. black. Distr. Malaysia: Java, Lesser Sunda Islands (Bali), Philippines (Mindanao). Ecol. Forests and forest borders, 1000-2000 m; cf. the note under L. acuminata. Vern. Ki seroh, S, gauod-bukid (Buk.). Notes. Apparently closely allied to L. glabrata DC. and L. affinis Hook. & ARN. from continental Asia, and in some respects to L. macrantha. The density of pubescence is very variable, ranging from hairy forms to almost glabrous ones; leaf- shape and size also vary, even on one twig from ovate to obovate. Pubescence of calyx-teeth varies from ciliate to tomentose. Occasionally 3-lobed leaves occur on young shoots. Sometimes 4-flow- ered cymes are observed. Gedeh Patoeha Papandajan |Goentoer TjiKoerai |Malabar Tjeremai Slamat Soembing ~| Sindoro =| Telemojo — Dieng +-—| Merbaboe 2 S o — 1200 1100 Fig. 3. Altitudinal localities (in metres) of Loni- cera acuminata WALL. (xX) and L. javanica (.) in Java and Bali. Mountains arranged from W towards E. Thickened vertical lines indicate the altitude of the summits. 2. VIBURNUM LINNE, Sp.Pl. (1753) 267; BI. Bijdr. 13 (1826) 655; Mia. FI. Ind. Bat. 2 (1856) 119; BoerL. Handl. 2? (1891) 3; K. & V. Bijdr. 5 (1900) 36; KING & GAMBLE, J. As. Soc. Bengal 72? (1903) 112; Koorpers, Exk. Fl. Java 3 (1912) 285; MErR. En. Dec. 1951] CAPRIFOLIACEAE (Kern & vy. Steenis) 181 Philip. 3 (1923) 577; Rip. Fl. Mal. Pen. 2 (1923) 1; BACKER, Bekn. Fl. Java, em. ed. 8 (1949) fam. 175, p. 2; KERN, Reinw. 1 (1951) 107. Shrubs or small trees. Leaves petiolate, simple, entire or serrate-dentate or trifid, pinnate- or palmate-nerved, (in the Malaysian species) exstipulate. Inflorescence terminal, compound, corymbiform or paniculate, primary rays usually whorled, flowers cymosely arranged. Bracts and bracteoles usually small, caducous. Flowers actinomorphic, the marginal ones sometimes (not in Malaysia) radiant, neutral. Calyx 5-lobed or 5-partite. Corolla white, creamy or pink, rotate, campanulate, hypocrateriform or tubular; lobes 5, imbricate in bud. Stamens 5; filaments narrow. Ovary 1-celled. Ovule anatropous, pendulous from the apex, solitary. Style short, conical; stigmas 3, often connate. Fruit a drupe, crowned by the persistent calyx and style, l-seeded. Endocarp horny or stony, in cross-section often undulate or with inflexed edges. Albumen often ruminate. Distr. Large genus; several hundred, often polymorphous species in Europe, Asia, and America, 16 spp. in Malaysia. Uses. None of the Malaysian species are known to be of importance economically. Wood anat. MoLi & JANssonius, Mikr. Holzes 4 (1920) 5. V. coriaceum: M. & Js. p. 19, V. sam- bucinum: p. 20, V. lutescens: p. 11. Notes. For a satisfactory identification both flowers and ripe fruits are required, a condition rarely found in the herbarium. From 3 spp. flowers are unfortunately hitherto unknown. The stellate hairs mentioned to occur in some spp. are apparently not properly stellate but fasciculate hairs (cf. McArtee, Bull. Torrey Bot. Cl. 48, 1921, 149 seq.). KEY TO THE SPECIES (for flowering specimens) As the flowers of V. amplificatum, V. cornutidens and V. clemensae are unknown, these species have been omitted. 1. Leaves triple-nerved. Tube of corolla hairy within. . . . . . . . . 15. V. propinquum 1. Leaves penninerved. Tube of corolla glabrous within. 2. Corolla squamulose without. All young parts densely covered with minute rusty-coloured peltate scales. Leaves entire, the underside at first densely covered with minute scales, later on densely punctulate . . a 6. V. punctatum 2. Corolla quite glabrous. or pubescent without, sometimes gland- dotted, but not squamulose. Young parts without peltate scales. 3. Corolla pubescent without, rotate, tube very short, about 1/2 mm long, lobes 1 mm. Filaments 11/2-2 mm long. Young branchlets Saad aac ae ee See 16. V. luzonicum 4. Adult leaves pubescent. . « doar 6S erie 4. Adult leaves nearly glabrous except for the midrib. Spubeaves nearly entire = <= - Re ee ne Oe See ens, ph var. apoense 5. Leaves strongly dentate. 6. Apex of leaves acute to shortly acuminate. . . .- - . -. .. . var. floribundum 6. Apex of leaves slenderly acuminate. . ie i en < var. sinuatum 3. Corolla glabrous or gland-dotted without, tube at least 1 mm long. 7. Filaments adnate to the throat of the corolla. Corolla shortly salvershaped-campanulate, limb horizontally spreading, finally reflexed; tube 2-3 mm long, lobes 2mm . 13. V. odoratissimum 7. Filaments adnate to the base or the tube of the corolla. Shape of the corolla different. 8. Ovary pubescent. Corolla glabrous, tube 1—1!/2 mm long, lobes 1-2 mm. Filaments 5—-7(-9) mm. Leaves large, 10-25 by 5-10 cm, coriaceous, entire . . - - -. . .- 7. V.sambucinum 9. Ovary and axes of inflorescence densely pubescent. 10. Underside of leaves glabrous except for a few hairson the nerves . . var. sambucinum 10. Underside of leaves villous. oe t3a°,” 3,0 eee var. tomentosum 9. Ovary and axes of inflorescence subglabrous Oe ee ee Pe var. subglabrum 8. Ovary not pubescent. 11. Corolla distinctly tubular, tube at least 3 times as long as lobes, the latter about 1 mm, erect. 12. Leaves quite entire, underside with distinct glandular pit at the base on both sides of midrib, apex obtuse or shortly and bluntly acuminate. Filaments inserted at base of corolla, 7-8 mm long. . 2. “2. Vibeccari 12. Leaves usually distinctly dentate, ‘sometimes nearly entire, underside bearded in nerve-axils, but without glandular ie san pire Meee) Se eee Filaments usually about Avmm IOng- tas . Ss. 1. VY. coriaceum 182 FLORA MALESIANA [ser. I, vol. 43 13. Tube of corolla 3-4 mm ae Filaments about 4mm ue adnate !/2-1 mm above base of corolla . var. coriaceum 13. Tube of corolla about 6mm lone. Filaments tp to 6 mm long, adnate 2-3 mm above base Ofacorollameaniaeee : var. longiflorum 11. Corolla not distinctly tubular, tube less than 3 times as ; long as lobes, the latter usually more than 1 mm long. 14. Inflorescence shortly pyramidal, paniculate. Corolla rotate-campanulate, tube about 1 mm long. Filaments 2-3 mm long. 15. Leaves thinly coriaceous. Corolla 2—2!/2 mm long. Filaments inserted at base of the corolla. (Between 500 and 1500 m altitude, sometimes up to 2300 m) . . . 10. V. lutescens 15. Leaves manifestly coriaceous. Corolla 3 mm long. Filaments adnate to the corolla !/2-1 mm above the base. Without fruits hardly distinguishable from V. lutescens. (Altitude at least 2300 m) ay eee 11. V. junghuhnii 14. Inflorescence corymbiformt Tbe! of the corolla usually exceeding 1 mm. Filaments at least 6 mm long. 16. Filaments 6(—7) mm long, in bud with inflexed top. 17. Corolla shortly tubular-turbinate, globular in bud, tube about 2mm, lobes 1!/2-2 mm. Leaves gradually long-acuminate . . é 4. V. platyphyllum 17. Corolla broad-tubular, obovoid in bud, tube about 21/2 mm, lobes about 1'/2 mm. Leaves obtusevor shortlyand bluntlyiacuminate 3-2 | 3. V. glaberrimum 16. Filaments (8—)9—-10 mm long, serpentine in bud. 18. Leaves dull, hispidulous on the midrib and the primary side-nerves at the underside, with- out glandular pits. Corolla rotate-cupular, tube 1(—11/2) mm, lobes 2—2!/2 mm. 8. V. hispidulum 18. Leaves shining, glabrous, with distinct glandular pits at the base. Corolla turbinate, tube Apri lobes Wein “4. 6 9s 5 6 & 8 &@ o o 6 « 9. V. vernicosum KEY TO THE SPECIES (for fruiting specimens) 1. Leaves triple-nerved. Fruits nearly globose, 4-5 mm long,4mm wide . . . 15. V. propinquum 1. Leaves penninerved. Fruits usually compressed, if not, more than 5 mm long. 2. Endocarp with strongly incurved edges, ventral side deeply intruding, embracing an internal split or cavity. 3. Fruit oblong, 16 by 7mm. Internal cavity of the fruit bilobate in cross-section. Leaves entire, glabrous, elliptic-oblong to slightly obovate, up to 26 by 12-13 cm... 12. VY. amplificatum 3. Fruit smaller. 4. Leaves minutely papillose- rugulose, entire. Ventral side of endocarp forming a narrow split, central cavity at most '/2mmindiameter . . ; 14. V. clemensae 4. Leaves smooth, usually dentate or serrate. Central cavity of fruit about 2 mm wide. 5. Fruit ovoid, 6-7 by 4-5 mm. Internal cavity of fruit nearly circular in cross-section. Leaves dentate to nearly entire . . . . . 13. V. odoratissimum 5. Fruit obovoid, 7-9 by 5—7 mm. ‘Internal cavity of the fut broad, bilobate in cross-section. Leaves closely crenate-dentate . . . . 1. V. junghuhnii 2. Endocarp undulate in cross-section, i.e. with shallow ‘grooves on both ‘sides, without internal split or cavity. 6. Leaves quite entire. 7. All young parts densely covered with minute peltate scales. Underside of the leaves at first densely covered with minute scales, later on densely punctulate. Fruit elliptic to somewhat obovoid, 9-11 by 6-7 mm ; Seer a a oe) See Pt Oe Ve Det enD 7. Young parts without peltate scales. 8. Ripe fruit small, 5—6(—6!/2) mm long. 9. Leaves usually ovate-lanceolate, aa oe long-acuminate. Fruit ovoid to broad- ellipsoid, 6(—6!/2) by 5-6 mm. ; =. . 1eVeconacenm 9. Leaves broader, ovate to obovate, obtuse | or shortly and bluntly acuminate. Fruit about 5 by 5mm, ovate to nearly orbicular . . RiP GE: pte 3. V. glaberrimum 8. Ripe fruits larger (without flowers hardly determinable!). 10. Nerve-axils and leaf-base without glandular pits or spotty glands. Midrib and primary nerves somewhat hispidulous beneath. Young branchlets and axes of infructescence glabrous or sparingly hispidulous. Young fruits glabrous . . + LAs 8. V. hispidulum 10. Nerve-axils and/or leaf-base at underside glandular pitted’ « or with (sometimes indistinct) spotty glands. Midrib and primary nerves not hispidulous. 11. Young branchlets densely pubescent. Axes of the inflorescence pubescent, glabrescent. Young fruits thinly pubescent, soon glabrescent: Underside of leaves at the base with a (sometimes indistinct) spotty gland on both sides of midrib. . . . 7. VY. sambucinum 12. Axes of the infructescence densely pubescent, later on glabrescent. Dec. 1951] 13. Underside of leaves glabrous except for a few hairs on the nerves 13. Underside of the leaves villous a Axes of the infructescence nearly glabrous CAPRIFOLIACEAE (Kern & vy. Steenis) 183 var. sambucinum var. tomentosum var. subglabrum . Young branchlets and axes of infructescence glabrous to somewhat ‘(not densely) pubescent. Young fruits gabteus. 14. Leaves large, up to 2 elongated and slender acumen 14. Leaves smaller, obtuse or shortly and bluntly acuminate. 2 by 10 cm, ovate to oblong-ovate, apex rane narrowed into an 4. V. platyphyllum 15. Young axes of the inflorescence pubescent, glabrescent. Young parts not vernicose. Primary nerves at the underside of the leaves little prominent 2. V. beccarii 15. Infructescence quite glabrous. Young parts vernicose. Primary side- -nerves at the underside of the leaves prominent 9. V. vernicosum 6. Leaves dentate or serrate (sometimes superficially). 16. At least the midrib of the leaves pubescent. ovate, much compressed, 5—7 by 5-6 mm, ripening red (always?) 17. Adult leaves pubescent Leaves chartaceous, rarely subcoriaceous. Fruit 16. VY. luzonicum var. luzonicum 17. Adult leaves nearly glabrous except ‘for the midrib. 18. Leaves nearly entire 18. Leaves strongly dentate. 19. Apex of the leaves acute to shortly acuminate 19. Apex of the leaves slenderly acuminate . var. apoense . var. floribundum var. Sinuatum 15. Leaves glabrous (sometimes bearded in the nerve- pealay en more or r less coriaceous. Fruit ripening purplish or bluish black. 20. Leaves thickly coriaceous, beneath with shallowly sunken glands in the nerve-axils, margins conspicuously corniculate-dentate. Fruit broadly ovate, 8 by 6-7 mm. 20. Not combining these characters. 5. V. cornutidens 21. Infructescence shortly paniculate. Fruit oblong-elliptic, 7-10 by 4-5 mm or still larger. 10. V. lutescens 21. Infructescence corymbiform. Fruit more ovate or ovoid, smaller. 22. Leaves usually ovate-lanceolate, gradually ge eremaaca distinctly dentate. Fruit 6(—6!/2) by 5-6 mm . 1. V. coriaceum 22. Leaves broader, ovate ‘to obovate, obtuse or shortly ‘and bluntly acuminate, nearly entire. Fruit 5 by 5 mm 1. Viburnum coriaceum BL. Bijdr. 13 (1826) 656; DC. Prod. 4 (1830) 329; Hassx. Flora 3 (1845) 242; Mia. FI. Ind. Bat. 2 (1856) 120, Suppl. (1860) 213; OERSTED, Vid. Meddel. Kjéb. 1860 (1861) 300, t. 6 f. 5-6; CLARKE in Hook. f. Fl. Br. Ind. 3 (1880) 5; K. & V. Bijdr. 5 (1900) 38; Koorp. Exk. FI. 3 (1912) 285; Koorp. Fl. Tjib. 32 (1918) 37; DANGuy in Fl. Gén.I.C. 3 (1922) 8; KERN, Reinw. 1 (1951) 115. —V. forbesii var. FAwc. in Forses, Wand. (1885) 507.—V. cylindricum Ham. ex D. DON sensu REHDER in SARGENT, Trees & Shrubs 12 (1908) 112: BACKER, Bekn. Fl. Java em. ed. 8 (1949) fam. 175, p. 3.—Fig. 4a-f. Shrub or small tree up to 15 m. Young parts thinly stellate-pubescent. Leaves coriaceous, some- what shining, glabrous above, often bearded in the nerve-axils on the underside, ovate to lanceolate, 10-24 by 4-8 cm, apex mostly gradually long-acumi- nate, base rounded or somewhat acute, margins su- perficially remotely dentate to rather densely serrate- dentate, sometimes almost entire; primary nerves 5-7 on each side, indistinctly anastomosing; petiole 2!/24 cm. Inflorescence umbellate, corymbiform, up to 10 cm across; axes thinly stellate-pubescent; peduncle short, up to 2!/2 cm; primary rays 5-7. Bracts and bracteoles small, ovate. Flowers some- what scented. Calyx-limb obscurely toothed, 1!/2-2 mm diam., teeth triangular, acute. Corolla tubular, ellipsoid-obovoid in bud, creamy white to white, sometimes pink without, usually dotted with brown; tube 3—4 mm, lobes erect, rounded trian- 3. VY. glaberrimum gular, 1 mm. Stamens exserted; filaments with inflexed top in bud, white, adnate to the corolla- tube !/2-1 mm above the base, about 4 mm long; anthers oblong, purplish, 1—-1!/2 mm. Ovary cylin- dric, glabrous or lepidote, 11/2-2 mm long. Drupe ovoid to broad-ellipsoid or nearly spheric, slightly compressed, bluish black, 6—6!/2 by 5-6 mm; endo- carp undulate in cross-section, dorsally 2-grooved, ventrally 3-grooved, lateral grooves often obsolete. Distr. SE. Asia, in Malaysia: Sumatra, Java, Lesser Sunda Islands (Bali, Lombok, Flores, Timor). Ecol. In open primary and secondary forests, especially in forest borders, sometimes in brush- wood or in grassy plains, from 1000 (especially 1500) m upward, often common and one of the pioneers in natural reafforestation. F/. fr. Jan.-Dec. Vern. Often noticed names: kiapu, kikukuran, S, tementilan, meniran, J. Notes. Extremely variable in all its parts, mainly in leaf shape. var. longiflorum KERN, /.c. Tube of corolla 6-7 mm long, lobes 1 mm. Filaments about 6 mm long, adnate to the corolla 2-3 mm above the base. Distr. Malaysia: Sumatra (Eastcoast). 2. Viburnum beccarii GAMBLE, J. As. Soc. Bengal 722 (1903) 114; Hartrer f. Med. Rijksherb. 14 (1912) 37; Rip. J. Fed. Mal. St. Mus. 8 (1917) 44; Fl. Mal. Pen. 2 (1923) 2; KERN, Reinw. 1 (1951) 184 FLORA MALESIANA [ser. 5 vole4? 4 Fig. 4. Viburnum coriaceum BL. a. Flowering twig, b. flower, c. ovary, d. aestivation of stamen, e. fruit, f. cross-section through fruit,—V. sambucinum BL., g. flower, A. aestivation of stamen,—V. odoratissimum ~ Kgr, i. flower, j. cross-section through fruit,—V. Jutescens BL., k. cross-section through fruit. Dec. 1951] 120, f. 2.—Viburnum sp. Merr. Contr. Arn. Arb. 8 (1934) 164. Sprawling or climbing shrub or small tree up to 8 m. Leaves coriaceous, upperside glabrous, under- side minutely gland-dotted and with a distinct glandular pit at the base on both sides of the midrib, often smaller glands in the nerve-axils; elliptic, obovate or ovate, 8-15 by 5-8 cm, apex obtuse or shortly and bluntly acuminate, rarely somewhat emarginate, base cuneate to nearly rounded, margins entire; midrib prominent be- neath, primary nerves much less prominent, 4-8 on each side, arcuately anastomosing; petioles 1!/2-3'/2 cm. Inflorescence corymbiform, umbella- tely branched, 6—12 cm across (in fruit up to 15cm), young axes rather densely brown stellate-pu- bescent, glabrescent; peduncle rather stout, up to 12cm; primary rays 5-8. Bracts and bracteoles small, linear-lanceolate, rusty stellate-pubescent. Calyx-limb cupular, obscurely lobed, minutely gland-ciliolate, otherwise glabrous. Corolla tubu- lar, obovoid-ellipsoid in bud, white; tube 3-4 mm, lobes erect, rounded to rather acute, minutely gland-ciliolate, 1 mm. Stamens long-exserted, with inflexed top in bud, inserted at base of corolla, 7-8 mm; anthers oblong, purplish, about 2 mm. Ovary cylindric, glabrous, 2 mm long. Drupe ovate, com- pressed, (young) bluish green, 9-10 by 6-7 mm. Endocarp undulate in cross-section, dorsally 2- grooved, ventrally 1-grooved. Distr. Malaysia: Malay Peninsula (Perak, Pa- hang) and Sumatra (Atjeh, Eastcoast, Westcoast). Ecol. In forests and thickets, 1100-1900 m. 3. Viburnum glaberrimum Merr. Philip. J.Sc. 4 (1909) Bot. 329; En. Philip. 3 (1923) 577; KERN, Reinw. 1 (1951) 122. Small tree, nearly glabrous. Leaves coriaceous, shining, glabrous except for the bearded (or glan- dular pitted) nerve-axils on the underside, ovate or elliptic, (6-)8-15 by (3—)6-8 cm, apex broadly and obtusely acuminate to nearly rounded, base round- ed or slightly decurrent-cuneate, margins entire or remotely undulate-dentate; nervation rather prom- inent beneath; primary nerves 6-8 on each side, anastomosing; petiole 24cm. Inflorescence um- bellate, corymbiform, 5-10 cm across, axes thinly stellate-pubescent, glabrescent; peduncle stout, 2-4 cm long; primary rays 5—7. Bracteoles very small, caducous before anthesis. Calyx-limb ob- scurely lobed. Corolla broad-tubular, obovoid in bud, glabrous, tube about 2!/2 mm, lobes erect, rounded, about 1!/2 mm. Stamens exserted; fila- ments adnate to base of corolla, in the flower-bud inflexed at the top, about 6 mm; anthers oblong, 11/2-2 mm. Ovary cylindric, glabrous, 1 mm long. Drupe ovate to nearly orbicular, compressed, 5 by 5 mm. Endocarp obscurely undulate in cross-sec- tion, with 2 shallow dorsal grooves and 1 shallow ventral groove. Distr. Malaysia: danao). Ecol. In primary forests; altitude according to MERRILL (1909 /.c.) 300-450 m, according to MER- RILL (1923) /.c., 1000-1400 m. Philippines (Luzon, Min- CAPRIFOLIACEAE (Kern & v. Steenis) 185 Note. Closely allied to V. coriaceum, which is unknown from the Philippines. The differences are presumably sufficient to justify specific separation. The corolla of V. coriaceum is more distinctly tu- bular, the corolla-bud more ellipsoid. The stamens of V. glaberrimum are inserted at the base of the corolla, those of V. coriaceum somewhat adnate to the tube; the filaments are 6 mm long, those of V. coriaceum reach this length only in some large- flowered specimens. 4. Viburnum platyphyllum Merr. Philip. J.Sc. 10 (1915) Bot. 284; Kern, Reinw. 1 (1951) 123, f. 3.—V. pachyphyllum Merr. sphalm. En. Philip. 3 (1923) Sle Tall tree, nearly glabrous. Leaves firmly chart- aceous to subcoriaceous, somewhat shining, pale olivaceous when dry, glabrous, ovate to oblong- ovate, 9-22 by 4-10 cm, apex gradually narrowed to the usually elongated and rather slender acumen, base obtuse to somewhat acute, margins entire to obscurely undulate; primary nerves 6-7 on each side, somewhat prominent beneath, indistinctly anastomosing, axils on the lower surface (often also axils of coarser secondary nerves) glandular- pitted; petioles 4-5 cm, of the smaller leaves 1-2 cm. Inflorescence large, umbellate, corymbiform, up to 18 cm across, axes thinly stellate-pubescent, glabrescent; peduncle 4-6 cm long; primary rays about 7. Flowers numerous, fragrant. Calyx-limb with short but distinct triangular teeth. Corolla globular in bud, when open shortly tubular to somewhat turbinate, gradually slightly widened towards the top, white, glabrous, tube 2—2!/2 cm, lobes erect, rounded, 1!/2-2 mm. Stamens exserted; filaments inserted at base of corolJa, in the flower- bud with inflexed top, sometimes moreover with a distinct fold in the lower part, 6-7 mm; anthers oblong, 2mm. Ovary cylindric, glabrous, 1 mm long. Drupe oblong-ovate, compressed, 8-9 by 6 mm. Endocarp slightly undulate in cross-section, with 2 dorsal grooves and 1 ventral groove. Distr. Malaysia: Philippines (Leyte). Ecol. In forests, at about 500 m. Note. Very closely allied to V. glaberrimum, from which it is possibly not specifically different. The main differences are the much larger leaves with slender acumen (also smaller-leafed forms occur), the more distinct calyx-teeth, the shape of the corolla (widened to the top, in V. glaberrimum broad-tubular) and the larger fruits. 5. Viburnum cornutidens Merr. Philip. J.Sc. 26 (1925) 491; En. Philip. 4 (1926) 251; Kern, Reinw. 1) 9ST) 25: Small, glabrous tree, about 5m high. Leaves thickly coriaceous, shining, olivaceous or brownish olivaceous, obovate to elliptic, 10-14 by 7-10 cm, apex rounded to shortly and obtusely acuminate, base acute, margins conspicuously corniculate- dentate, teeth straight, obtuse, 1-2 mm long, chiefly terminating the primary nerves, these about 9 on each side, nearly straight, once (sometimes twice) forked, beneath with glandular pits in the axils; petioles 2-3!/2 cm long. Infructescence umbellate, 186 FLORA MALESIANA [ser. I, vol. 43 corymbiform, about 15 cm across; peduncle stout, about 5 cm long; primary rays 5—7. Drupe broadly ovate, compressed, 8 by 6-7 mm. Endocarp slight- ly undulate in cross-section, dorsally 2-grooved, ventrally 1-grooved. Distr. Malaysia: Philippines known from Mt Baudan). Ecol. On forested slopes at + 1800 m. Vern. Manano (Igorot). Note. Flowers unknown. Like V. platyphyllum an ally of V. glaberrimum. The species is (suffi- ciently?) characterized by the thickly coriaceous, broad, dentate leaves and the broadly ovate fruits. (Luzon, only 6. Viburnum punctatum Ham. ex D. Don, Prod. Fl. Nepal. (1825) 142; DC. Prod. 4 (1830) 324; CLARKE in Hook. f. FI. Br. Ind. 3 (1880) 5; DANGvuy in FI. Gén. I.C. 3 (1922) 12; KERN, Reinw. 1 (1951) 127.—V. acuminatum WALL. ex DC. Prod. 4 (1830) 325; W. & A. Prod. 1 (1834) 388; Wicurt, Icones 3 (1845) 13, t. 1021. Shrub or small tree up to 18m. Young parts densely covered with minute, rusty-coloured, pelt- ate scales, leaving numerous punctiform scars when they fall off. Leaves coriaceous, upper surface glabrous, underside densely covered with minute scales, neither bearded nor glandular-pitted in the nerve-axils, elliptic-lanceolate to lanceolate, 5-13 by 2-4'/2 cm, apex bluntly acuminate, base attenuate, margins entire; primary nerves 5—7 on each side, rather prominent beneath, anastomos- ing; petioles 1—1!/2cm. Inflorescence umbellate, corymbiform, 5—10 cm across (infructescence up to 15 cm), axes densely squamulate; peduncle very short, 1-3 cm; primary rays 3—5. Bracts and bract- eoles minute, ovate-lanceolate, fimbriate. Flowers fragrant, about 5mm wide. Calyx-teeth ovate- triangular, obtuse, light-margined, squamulate, about 3/4 mm long. Corolla white, glabrous within, squamulate without, globular in bud, nearly rotate (only slightly campanulate) when open, tube 1 mm, lobes broad-ovate, rounded, somewhat overlap- ping, 2 mm. Stamens somewhat exserted; filaments in bud with inflexed top, inserted near base of corolla, 3-4mm; anthers elliptic, 1 mm. Ovary cylindric, lepidote, 1!/2-2 mm long. Drupe elliptic or slightly obovate, much compressed, young squamulose, ripening black, 9—11(—12) mm by 6-7 mm. Endocarp undulate in cross-section, with 2 dorsal and 3 ventral grooves. Distr. SE. Asia, from Nepal, Kumaon and the Deccan to Siam and Indochina, in Malay- sia: N. Sumatra (Atjeh), twice collected. Ecol. Secondary growths, 800—1000 m. 7. Viburnum sambucinum BL. Bijdr. 13 (1826) 656; DC. Prod. 4 (1830) 325; Mia. FI. Ind. Bat. 2 (1856) 120;HAssk. Bonpl.7 (1859) 170; OERSTED, Vid. Med- del. Kj6b. 1860 (1861) 299, t. 7, f. 11-13; CLARKE in Hook. f. Fl. Br. Ind. 3 (1880) 5, 671; K. & V. Bijdr. 5 (1900) 40; GamBLeE, J. As. Soc. Bengal 722 (1903) 113; Koorp. Exk. Fl. 3 (1912) 285; Koorp. FI. Tjib. 32 (1918) 38; Merr. J. Str. Br. Roy. As. Soc. 86 (1921) 582; DANGUy in FI. Gén. I.C. 3 (1922) 11; Rit. Fl. Mal. Pen. 2 (1923) 1; CoRNER, Way- side Trees (1940) 183; BACKER, Bekn. FI. Java, em. ed. 8 (1949) fam. 175, p. 4; KERN, Reinw. 1 (1951) 129.—V. integerrimum WALL. Cat. 457; Hoox. fi & Tu. J. Linn: Soc. 218538) aige— V. forbesii FAwc. in Forpes, Wand. (1885) 506, excl. var. —Fig. 4 g-h, 5. Shrub or small tree, up to 10-(15) m. Young branchlets densely stellate-pubescent. Leaves more or less coriaceous, glabrous except for a few hairs on the nerves and the bearded nerve-axils at the underside (see var. 2), here with an often indistinct spotty gland at the base on both sides of midrib, elliptic-oblong to oblong-lanceolate, 10-25 by 5-10 cm, apex abruptly short-acuminate, base cuneate, margins entire; nervation rather promi- nent beneath; primary nerves 5—7 on each side, anastomosing; petioles up to 4cm long. Jnflores- cence densely many-flowered, umbellate, corym- biform, up to 15-18 cm across, axes densely stel- late-pubescent; peduncle stout, 4-6 cm long; pri- mary rays 6-8. Bracts and bracteoles small, linear- lanceolate, stellately pubescent. Flowers small, 3-4 mm diam., very fragrant. Calyx-limb with ovate- triangular, acute, ciliate lobes, 3/s-1 mm long. Co- rolla rotate or campanulate, globular in bud, white or creamy, tube 1(—1'/2) mm, lobes spreading, ovate, rounded, (1-)1'/2(-2) mm, usually tube somewhat shorter than lobes, not rarely the reverse. Stamens long-exsert; filaments almost filiform, serpentine in bud, inserted at base of corolla, (4-)5—7(-9) mm; anthers elliptic to oblong, 3/4-1 mm. Ovary cylin- dric, usually densely hairy, 1—-1'/2 mm long. Drupe ovate, much compressed, young thinly hairy, gla- brescent, ripening bluish black, (7—-)9(-10) by (5—)6(—7) mm. Endocarp undulate in cross-section, dorsally 2-grooved, ventrally 3-grooved, lateral ventral grooves often obsolete. Distr. Cambodia, in Malaysia: chiefly in the western part, often frequent, Malay Peninsula, Su- matra, Borneo, Java (rare in the central and eastern part), Lesser Sunda Islands, Celebes and the Moluccas, but here presumably very rare (Ceram). Ecol. Open primary and secondary forests, in brushwood, particularly at forest-edges in the lower mountain zone (up to 1800 m), occasion- ally in swampy places in the lowlands. Fi. fr. Jan.—Dec. Vern. Ki kukuran, ki bewog, S, bleber, J, and many other local names. var. subglabrum KERN, Reinw. 1 (1951) 130.—V. longistamineum RIDL. J. Fed. Mal. St. Mus. 6(1915) 151; Fl. Mal. Pen. 2 (1923) 2; SymincToN, J. Mal. Br. R. As. Soc. 143 (1936) 353. Inflorescence small, 5-7 cm across. Axes of inflorescence and ovary subglabrous. Distr. Malaysia: 1800 m. Malay Peninsula, 1100- var. tomentosum HALLER f. Med. Rijksherb. 14 _ (1912) 36; KERN, Reinw. 1 (1951) 130, f.4.—V. suma- tranum Mia. Fl. Ind. Bat. Suppl. (1860) 537.—V. villosum RiDL. J. Str. Br. Roy. As. Soc. no 61 (1912) 10; Fl. Mal. Pen. 2 (1923) 2.—V. inopinatum CRAIB, Dec. 1951] Fig. 5. Viburnum sambucinum Bu Kew Bull. (1911) 385; DANGUuy in FI. Gén. I.C. 3 (1922) 10. Underside of the full-grown leaves softly villous by simple, forked and stellate hairs. Distr. Siam, in Malaysia: Malay Peninsula (Se- langor), Sumatra (especially in the northern half), the typical form probably restricted to higher alti- tudes, 800-1200 m. 8. Viburnum hispidulum KERN, Reinw. 1 (1951) ISG, iis 5 Tree up to 24 m. Leaves coriaceous, dull, glabr- ous above or somewhat hispidulous on midrib and primary nerves, hispidulous beneath, especially on midrib and primary side-nerves, punctulate, neither glandular-pitted nor bearded in nerve-axils, elliptic to elliptic-oblong or obovate, 12-17 by 8-9 cm, apex abruptly shortly and bluntly acuminate (acu- men up to 5 mm), base cuneate to broadly cuneate, margins entire; nervation prominent beneath; pri- mary nerves 5S—7 on each side, anastomosing; peti- oles 2-6cm. Inflorescence umbellate, corymbi- form, 10-15 cm across; peduncle short, up to 5 cm; primary rays 6-7; bracteoles firm, ovate to lanceolate, gland-dotted, ciliate, 4-6 mm long. Ca- lyx-limb cupular, glabrous, obscurely lobed, 1 mm long. Corolla (creamy) white, globular in bud, rotate-cupular when open, tube 1(-1'/2) mm, CAPRIFOLIACEAE (Kern & v. Steenis) 187 . G. Pantjar, West Java (DE VooGpD). lobes ovate to oblong, 2—2!/2 mm. Stamens much exserted; filaments thick, serpentine in bud, in- serted at base of corolla, 9-10 mm; anthers oblong, 2 mm. Ovary cylindric, lepidote, 11/2 mm long and thick. Drupe oblong-elliptic to slightly obovate, compressed, 9-10(-11) by 6-7 mm. Endocarp un- dulate in cross-section, dorsally 2-grooved, ven- trally 3-grooved. Distr. Malaysia: Br. N. Borneo (Mt Kinabalu), -+- 2000 m. Note. Differs from V. vernicosum mainly by its dull hispidulous leaves, the absence of glandular pits at the leaf-base and the rotate-cupular corolla. 9. Viburnum vernicosum GiBBs, J. Linn. Soc. 42, Bot. (1914) 86; Merr. En. Born. (1921) 582; KERN, Reinw. | (1951) 139, f. 6. Shrub or small glabrous tree, up to 10 m. Young parts very shining, vernicose. Leaves coriaceous, shining, chiefly on underside densely punctulate and gland-dotted, beneath with distinct glandular pit at the base on both sides of midrib and often smaller ones in higher nerve-axils, elliptic to slight- ly obovate, 12-18 by 6-11 cm, apex shortly and abruptly acuminate (acumen up to 1 cm long), base acute, margins entire; primary nerves 5-7 on each side, prominent beneath, anastomosing. Petioles 1-3 cm. Inflorescence umbellate, corymbiform, up 188 FLORA MALESIANA [ser. I, vol. 43 to 11 cm across; peduncle stout, up to 6 cm long; primary rays 5—7; bracteoles firm, oblong to lan- ceolate, with membranous margins, gland-dotted, 5-7 mm long. Calyx-limb obscurely lobed. Corolla creamy white, gland-dotted on outside, turbinate, obovoid in bud, tube 2!/2-3 mm, lobes erect, rounded triangular, 1!/2-2 mm. Stamens much ex- serted; filaments thick, serpentine in bud, inserted at base of corolla, (8-)9-10 mm; anthers oblong, 2mm. Ovary cylindric, lepidote, vernicose, 1—1!/2 mm long. Drupe ovate, compressed, purplish black, 10 by 7-8 mm. Endocarp undulate in cross- section, (often irregularly) 2-grooved on dorsal side, 3-grooved on ventral side, lateral grooves often nearly absent. Distr. Malaysia: Borneo. Ecol. Primary and secondary forests, 900- 2700 m. Note. Very polymorphous species. The above description refers to what presumably may be called the common form. Specimens with larger, elliptic fruits (12-14 by 7-8 mm), others with broad-elliptic leaves (14-16 by 9-11 cm) were also collected. 10. Viburnum lutescens BL. Bijdr. 13 (1826) 655; DC. Prod. 4 (1830) 325; Maxim. Mél. Biol. 10 (1880) 651; GaAmBste, J. As. Soc. Bengal 72 (1903) 114; Koorp. Exk. Fl. 3 (1912) 286; Koorpb. FI. Tjib. 32 (1918) 38; Rip. Fl. Mal. Pen. 2 (1923) 2; CorRNER, Wayside Trees (1940) 183; BACKER, Bekn. Fl. Java em. ed. 8 (1949) fam. 175, p. 5; KERN, Reinw. | (1951) 142.—V. monogynum BL. Bijdr. 13 (1826) 655.—V. sundaicum Mia. Fl. Ind. Bat. 2 (1856) 121; K.&V. Bijdr. 5 (1900) 43.— V. colebrooki- anum (non WALL.) DANGuy, FI. Gén. I.C. 3 (1922) 9.—?V. elegans JUNGH. Nat. Gen. Arch. 2 (1845) 36.—Fig. 4k. Shrub or small tree, up to 10 m, usually much lower. Youngest parts thinly stellate-pubescent. Leaves thinly coriaceous, upperside glabrous, un- derside thinly stellate-pubescent to almost glabr- ous, very variable in shape, broad-elliptic, ovate or oblong-elliptic, up to 18 by 10 cm, apex short- acuminate, base nearly rounded to cuneate, mar- gins in upper 2/3 coarsely crenate-serrate to finely serrate, teeth shortly mucronate, lower !/3 entire or superficially dentate; nervation rather prominent beneath; primary nerves 5—8 on each side, indis- tinctly anastomosing; petioles 1—2 cm. Inflorescence terminal or spuriously lateral, paniculate, short- pyramidal, 5-9 cm across, 5—7(-10) cm _ long; axes stellate-pubescent, glabrescent; primary branches 4~7, verticillate. Bracts and bracteoles minute, stellate-pubescent. Flowers somewhat frag- rant, 4-5 mm wide. Calyx-teeth ovate-triangular, 3/4 mm long. Corolla globular in bud, nearly rotate (only slightly campanulate) when open, creamy white, glabrous, tube 3/4-1 mm, lobes ovate, rounded, 11/4—1!/2(-2) mm. Stamens somewhat ex- serted, filaments inserted near base of corolla, in bud with inflexed top, white, 2~-3 mm; anthers el- liptic, sordidly white, 1 mm. Ovary cylindric, glabr- ous, 1-1'/2 mm long. Drupe oblong-ellipsoid, somewhat oblique, slightly compressed, ripening purplish-black, 7-10 by 4-5 mm (rarely 11-12 by 7-8 mm: Borneo, Malay Penins.). Endocarp undul- ate in cross-section, with 2 dorsal and 1 ventral groove. Distr. SE. Asia, in Malaysia: Malay Pen- insula (local), Borneo (few times), Sumatra, Java, Lesser Sunda Islands (Bali, Lombok). Ecol. Primary and secondary forests, brush- wood, often common but scattered, usually 500- 1500 m, rarely lower (—150 m) or higher (one re- cord of 2400 m from Mt Patuha). FI. fr. Jan.—Dec. Vern. Many native names have been recorded, e.g. ki rantja, ki kukuran, ki bewog, Sd, tjéré, wuru watu, J, kapor-kaporan, porkaporan, Md, kaju nassi (Sum.). Uses. Sometimes cultivated as a hedgeplant, easily propagated by cuttings. Ripe fruits are read- ily eaten by birds. 11. Viburnum junghuhnii Mia. Fl. Ind. Bat. 2 (1856) 123; K.& V. Bijdr. 5 (1900) 47; Koorp. Exk. FI. 3 (1912) 286; BACKER, Bekn. FI. Java 8 em. ed. (1949) fam. 175, p. 4; Kern, Reinw. 1 (1951) 147, f. 7.—V. lutescens (non BL.) HALLIER f. Med. Rijks- herbs eC OiUl) Sips py: Subarborescent shrub or small tree, up to 18 m. Leaves coriaceous, broad-elliptic, obovate-elliptic to nearly lanceolate, apex rounded or shortly acu- minate, base more or less attenuate, closely cren- ate-dentate, teeth apiculate, glabrous, (4—)8—12!/2 by 3-6 cm; nervation very prominent beneath; pri- mary nerves 5—7 on each side, anastomosing; peti- oles 11/2-21/2(-4) cm. Inflorescence terminal or pseudolateral, paniculate, short-pyramidal, up to 5 cm long and 8 cm wide; lower ramifications 3—5- nately whorled, upper ones alternate; peduncle 3-6 cm; bracteoles small, lanceolate to ovate. Flowers fully 5mm wide, fragrant. Calyx-limb 1 mm long, distinctly toothed, teeth ovate-trian- gular. Corollacreamy white, globular in bud, rotate- campanulate when open, glabrous, tube short, fully 1mm, lobes up to 2mm, ovate, rounded. Stamens hardly exserted; filaments in bud with in- flexed top, inferior part adnate to corolla !/2-1mm, free part 1!/2-2 mm; anthers elliptic, 1 mm. Ovary cylindric, glabrous, 2mm long. Drupe obovoid, 7-9 by 5-6 mm, compressed. Endocarp in cross- section slightly undulate and with strongly incurv- ed edges, ventral side therefore deeply intruding, embracing a broad, bilobate cavity. Distr. Malaysia: Sumatra (Westcoast: Mt Korintji), Java. Ecol. Forests, 2300-2600 m. 12. Viburnum amplificatum KERN, Reinw. | (1951) 150, f. 8.—Viburnum sp. Merr. Pl. Elm. Born. (1929) 297. Shrub-like tree. Leaves thinly coriaceous, dull, dark olivaceous above, brown beneath, glabrous, neither glandular pitted at the base nor bearded in the nerve-axils, elliptic-oblong to slightly obovate, up to 26 by 12-14 cm, apex abruptly short-acumin- ate (acumen rather blunt, !/2-1 cm), base cuneate to somewhat rounded, margins entire, sometimes distantly and obscurely undulate; nervation prom- Dec. 1951] CAPRIFOLIACEAE (Kern & y. Steenis) 189 inent beneath; primary nerves 5-7 on each side, anastomosing; petioles 2-4 cm. Infructescence um- bellate, corymbiform, about 13 cm diam.; peduncle stout, about 7 cm; primary rays 7-8. Drupe oblong, very slightly dilated upwards, much flattened, with a distinct groove on both sides, black, 16 by 7 mm. Endocarp with broad longitudinal groove on the dorsal side, the incurved edges forming a deep, broad, in cross-section bilobate furrow on the ventral side. Distr. Malaysia: Br. N. Borneo (Tawao). Note. Flowers unknown. The species is read- ily recognizable by its large leaves and fruits and by the cross-section of the endocarp, reminding one of V. junghuhnii. 13. Viburnum odoratissimum Ker, Bot. Reg. 6 (1820) t. 456; DC. Prod. 4 (1830) 326; CLARKE in Hook. f. Fl. Br. Ind. 3 (1880) 7; Maxim. Meél. Biol. 10 (1880) 645, 649: REHDER in SARGENT, Trees and Shrubs 2 (1908) 107; DANGuy in FI. Gén. I.C. 3 (1922) 5; Merr. En. Philip. 3 (1923) 577; BACKER, Bekn. FI. Java em.ed.8 (1949) fam. 175, p.3; KERN, Reinw. 1 (1951) 152, f. 9.—V. hasseltii Mia. FI. Ind. Bat. 2 (1856) 123; K.& V. Bijdr. 5 (1900) 46; Koorp. Exk. FI. 3 (1912) 286.—V. zambalense ELM. Leafi. Philip. Bot. 9 (1934) 3181.—Fig. 4i-j. Shrub or small tree, sometimes up to 20m, glabrous, only youngest parts with few stellate hairs. Leaves more or less coriaceous, dull or some- what shining, glabrous or underside with few scattered stellate hairs and somewhat bearded in the nerve-axils, elliptic-oblong to oblong-lanceolate or obovate, 8-15 by 3-7 cm, apex shortly and bluntly acuminate, rarely rounded or emarginate, base attenuate, margins in the upper half obsole- tely (sometimes distinctly) toothed or nearly entire, cartilagineous; nervation prominent beneath; pri- mary nerves 5—7 on each side, indistinctly anasto- mosing; petioles (1/2-)1—2 cm. Inflorescence panicul- ate, more or less elongated pyramidal, up to 10 cm long and wide, many-flowered; axes slightly stellate-pubescent; peduncle 2-5cm, primary branches verticillate, divaricate; bracteoles linear- lanceolate, ciliate, 1-3 mm long. Flowers fragrant, 5-6 mm wide. Calyx-limb cupular, 1 mm long, teeth broad-triangular, glabrous or sparsely ciliate. Corolla obovoid to ellipsoid-obovoid in bud, shortly funnel-shaped-campanulate when open, creamy white, glabrous or sometimes with some stellate hairs on outside, tube gradually widened upwards, 2-3 mm, limb horizontally spreading, finally reflexed, lobes broad-ovate, rounded, somewhat overlapping, 2 mm. Stamens exserted, filaments adnate to corolla-throat, in bud with inflexed top, 2-3 mm; anthers oblong, 1!/2- 13/s mm. Ovary glabrous, 11/2-2 mm long. Drupe ovoid, 6—7 by 4-5 mm, purplish-black. Endocarp dorsiventrally compressed and strongly incurved, edges nearly touching, ventralsidestrongly concave, embracinga cavity of 1!/2-2 mm diam. Seed strongly compressed, falcate in cross-section, 1 mm thick. Distr. Eastern India to Indo-China, E. China & Formosa, northward to Japan, in Malaysia: Philippines (Luzon, Mindoro, Panay, Negros, Ley- te, Mindanao), Celebes. The record for W. Java (Mt Pangrango, KUHL & VAN HASSELT) is presuma- bly a misstatement; it has not been retraced there. Ecol. Primary forests, ravines, thickets, efc., 1000-2000 m. Fi. fr. Jan.—Dec. Note. The above description has only reference to the Malaysian form. The widespread species is extremely variable both in the shape of the leaves and in the flowers. 14. Viburnum clemensae KERN, Reinw. 1 (1951) Santen LO: Small glabrous tree. Leaves coriaceous, dull, in dry state olivaceous above, yellowish or brownish green beneath, minutely papillose-rugulose all over (under the lens), underside glandular-pitted both in the axils of the primary and secondary nerves, elliptic to oblong-lanceolate or slightly obovate, 9-11!/2 by 31/2-5!/2cem, often inequilateral and somewhat falcate, apex abruptly short-acuminate to nearly caudate, rarely rounded, base cuneate, margins entire, cartilaginous; nervation prominent beneath, primary nerves 4-5 on each side, anasto- mosing; petioles 1—2!/2 cm. Jnfructescence nearly sessile, small (4-8 cm long), paniculate, lowest branches ternate, middle ones opposite, upper ones alternate. Bracteoles minute, lanceolate. Drupe el- lipsoid or oblong-ovoid, not compressed, 10 by 6-7 mm, shining, calyx-teeth persistent, triangular, 3/s mm long. Mesocarp thin, scanty fleshy. Endo- carp strongly incurved, dorsal side orbicular in cross-section, edges touching, ventral side folded to an internal longitudinal crest 2—2!/2 mm broad, slightly widened at the upper margin, here em- bracing a cavity of about '!/2 mm diam. Seed com- pressed, reniform in cross-section, 2 mm thick. Distr. Malaysia: Br. N. Borneo (Mt Kinabalu). Note. Flowers unknown. Easily distinguish- able from V. odoratissimum by the minutely rugu- lose leaves and the smaller number of primary side- nerves, but mainly by the size of the fruit and the quite different cross-section of the endocarp. 15. Viburnum propinquum Hens. J. Linn. Soc. 23 (1888) 355; REHDER in SARGENT, Trees and Shrubs 2 (1908) 33, 133, pl. 115; MerR. En. Philip. 3 (1923) 578; KERN, Reinw. 1 (1951) 160.—V. valerianicum Evo. Leafi. Philip. Bot. 7 (1915) 2578. Shrub. Leaves thinly coriaceous, glabrous except for the bearded axils of primary side-nerves on the underside, ovate to ovate-lanceolate, often some- what falcate, 6-8—(11) by 2!/24(5!/2) cm, apex acute to long-acuminate, base cuneate to broadly cuneate, margins on both sides !/2-1(—2) cm above the base with an impressed small gland, nearly entire, only minutely and distantly serrulate, teeth reduced to mucros hardly !/2 mm long; nervation prominent beneath, basal primary nerves nearly as prominent as the midrib (leaves therefore triple-nerved), more apical 2-3 pairs less prominent, all side-nerves anas- tomosing; petioles 1-2cm. Inflorescence umbellate, corymbiform, (3—-)5-7 cm across, axes glabrous; peduncle short, 2(—5) cm; primary rays 5—7. Bracts and bracteoles minute, ovate, ciliate. Calyx-limb 1 mm long, obscurely lobed; lobes ovate, obtuse, 190 FLORA MALESIANA [ser. I, vol. 4 glabrous, about !/2 mm. Corolla whitish or yellow- ish green, 4mm wide, campanulate-rotate, globular in bud; tube broad, scarcely 1 mm long, hairy within, lobes ovate to oblong, rounded, recurved in anthesis, 11/2 mm. Stamens exserted:; filaments adnate near base of corolla, in bud with inflexed top, 2-2!/2mm; anthers broadly elliptic, 3/4 by 1/2 mm. Ovary 3/4-1 mm long and wide, glabrous. Drupe globose-ovoid, bluish black, 4-5 by 4 mm. Mesocarp thin, scanty fleshy; endocarp thin, orbi- cular in cross-section, ventrally slightly 1-grooved. Seed ovoid; albumen deeply ruminate. Distr. China (Hupeh) and Formosa, in Ma- laysia: Philippines (Luzon: Rizal, Benguet). Ecol. Mossy forest, 2200-2450 m. Notes. The triplenerved leaf with marginal glands (reminding one of those of Vaccinium spp.), the hairy corolla-tube, the achenoid drupe and the peculiar cross-section of the endocarp characterize this species; it stands isolated among the other Malaysian Viburnums. According to ELMER /.c. the strong odour, especially in cured specimens, is that of Valeriana. The same fact had already been stated by HALLIER in V. sambucinum, V. coriaceum and V. lutescens. VAN ITTALIE ascertained the presence of valerianic acid in V. sambucinum. On this ground HALLIER supposed the close relationship of Vibur- num with the Valerianaceae (Med. Rijksherb. 14 (1912) 36; op. cit. 37 (1918) 92). 16. Viburnum luzonicum Ro tre, J. Linn. Soc. 21 (1884) 310; VipaL, Phan. Cuming. (1885) 117; Rev. Pl. Vasc. Philip. (1886) 147; REHDER in SARGENT, Trees & Shrubs 2 (1908) 97, pl. 146; Merr. Philip. J.Sc. 5 (1910) Bot. 391; DANGuy in Fl. Gén. I.C. 3 (1922) 13; MerR. En. Philip. 3 (1923) 577; KERN, Reinw. 1 (1951) 161.—V. laxum Exo. Leafl. Philip. Bot. 7 (1915) 2576. Shrub 3-6 m high. Ultimate ramifications (often densely) ferrugineous-pubescent. Leaves extremely variable, chartaceous to subcoriaceous, young ones pubescent by simple antrorse and stellate hairs, more or less glabrescent, 3—-8(—13) by 2-5 cm, ovate to lanceolate, apex acute to long acuminate, base broadly cuneate to rounded, often inequilateral, margins almost entire to sinuate-dentate in upper part; nervation prominent beneath, often hidden by indument; primary nerves 5—7 on each side, usually terminating in teeth, lower ones anasto- mosing; petioles densely pubescent, 1/2-1 cm. Jn- florescence terminal or spuriously lateral, umbel- late, corymbiform to semi-globose, 3—5 cm across, axes densely ferrugineous pubescent; pedunclevery short, up to 2 cm; primary rays 3-6. Bracts and bracteoles small, lanceolate, pubescent and ciliate, 1-2 mm. Flowers slightly odorous, 3—5 mm wide. Calyx hardly 1mm long, deeply lobed, lobes ovate-lanceolate, pubescent. Corolla rotate, glo- bular in bud, creamy white or somewhat pink, strigose and stellate-pubescent without, glabrous within, tube very short (1/2 mm), lobes elliptic-ob- long, rounded, 1!/2-2 mm. Stamens exserted, but shorter than corolla-lobes; filaments adnate to base of corolla, 1'/2-2 mm; anthers broadly elliptic, yellow, !/2-3/s mm long. Ovary cylindric, densely pubescent, 1 mm long. Drupe ovate, much com- pressed, red (or ripening black?), (5S—)6—7 by 5-6 mm. Endocarp slightly undulate in cross-section, the 2 dorsal and 3 ventral grooves often ob- solete. Distr. China & Indo-China to Formosa, in Malaysia: Philippines, Moluccas (Buru). Ecol. Thickets and forests, 800-2200 m. Fi. fr. Jan.—Dec. Vern. Many local names have been noticed, e.g. atalba, tilba (Ig.), atiba (Bon.), bangas-bangas ‘(Bag.), bagiroro (Bik.), putud (Gad.). var. apoense Eom. Leafl. Philip. Bot. 7 (1915) 2577; Merr. En. Philip. 3 (1923) 577; KERN, Reinw. 1 (1951) 162. Leaves subcoriaceous, nearly glabrous except for midrib, ovate-lanceolate, apex long-acuminate, acumen often falcate, margins nearly entire; pri- mary nerves often anastomosing. Distr. Malaysia: Philippines (Luzon?, Min- danao). Vern. Angganasi, bangas (Bag.). atadatud (Buk.), bangas- var. floribundum (MERR.) KERN, Reinw. 1 (1951) 163.—V. floribundum MeErr. Philip. J.Sc. 4 (1909) Bot. 328; En. Philip. 3 (1923) 577. Leaves chartaceous, nearly glabrous except for midrib and primary nerves, ovate, apex acute to shortly acuminate, margins rather strongly dentate in upper half, primary nerves for the greater part terminating in teeth. Distr. Malaysia: Philippines (Luzon). var. Sinuatum (MERR.) KERN, Reinw. 1 (1951) 163. —V. sinuatum MERR. Gov. Lab. Publ. 35 (1906) 65; Philip. J.Sc. 1, Suppl. (1906) 137; En. Philip. 3 (1923) 578. Leaves oblong-ovate, apex slenderly acuminate, acumen usually falcate, margins coarsely sinuate- dentate; otherwise as var. floribundum. Distr. Malaysia: Philippines (Luzon, Negros), Moluccas (Buru). Vern. Taringongog (Neg.). 3. SAMBUCUS LINNE, Sp.Pl. (1753) 269; Gen. Pl. (1754) p. 130, no 334; MILLER, Ber. D.B.G. 2 (1884) 452; Fritscn, Oesterr. B.Z. 39 (1889) 214; DAmmer, ibid. 40 (1890) 261; Hock. Bot. Centr. BI. 51 (1892) 233; Fritscu, ibid. p. 81; in E.& P. 4, 4 (1897) 157; SCHWERIN, Mitt. Deut. Dendr. Ges. (1909) 1-56; op. cit. (1920) 194. Trees, shrubs, or erect herbs. Stem and branches pithy, nodes often with stipule- Dec. 1951] CAPRIFOLIACEAE (Kern & v. Steenis) 191 like appendages. Leaves imparipinnate, or incompletely bipinnate, rarely laciniate. Leaflets serrate or divided, opposite or alternate. Flowers actinomorphic, articu- lated with the pedicel, mostly white, in terminal, flat or convex corymbs; flowers sometimes dimorphic: gynodioecious, or part of them aborted into nectarial glands. Bracts mostly absent. Bracteoles 1 or absent. Calyx-tube short, limb 5- parted. Corolla rotate, 5-lobed, lobes valvate or imbricate(?). Stamens 5, inserted on the base of the corolla; filaments filiform, erect; anthers oblong, latrorse, cells free, attached in the middle. Ovary 3—S-celled, each cell with 1 ovule; stigmas 3 or 5, short, broad, on a cushion-like style. Berry 3—5-seeded. Seeds with - flat sides, back convex, granulate, embryo + as long as the seed, terete. Distr. Ca 10-20 spp., throughout the globe, absent from Oceania, S. & Central Africa, and Australia (except its E. part and Tasmania), centering in the N. hemisphere. Uses. None of the various medicinal uses ascribed to the elder in Europe, Asia, and America is known from Malaysia. Notes. The discrimination of the specific characters as given by VON SCHWERIN is not very satis- factory, specially if only herbarium materials are available. Though the colour of the berry varies rather in several species, it is sometimes accepted for specific delimination between others. Leaf-shape and indumentum represent variable characters. The size and shape of the seeds may furnish additional specific characters (cf. fig. 1). Among Caprifoliaceae it seems that Sambucus represents the closest relation to Valerianaceae. Some specimens show well-developed stipules; these are hairy inside towards the base but do not possess colleters. KEY TO THE SPECIES 1. Corymbs with a number of flowers aborted into top-shaped, mostly yellow nectaries. Seeds ovate. 1. S. javanica 1. Corymbs without such nectaries. Seeds oblong. 2. Upper 1-2 pairs of leaflets sessile-adnate. Leaves simply pinnate. Berry oblong 2. Upper leaflets not adnate. Leaves often bipinnate. Berry globular 1. Sambucus javanica REINW. ex BL. Bijdr. 13 (1826) 657; DC. Prod. 4 (1830) 322; Hassx. Flora 28 (1845) 243; BLANCco, FI. Filip. (1845) 151, ed. 3, 1 (1877) 271; Mia. FI. Ind. Bat. 2 (1856) 124; Hk. f. & Tu. J. Linn. Soc. Bot. 1 (1857) 180; Hook. f. Fl. Br. Ind. 3 (1880) 2; Boerv. Handl. 2 (1891) 3; Koorp. Minah. (1898) 492; Hutrcu. Kew Bull. (1909) 193; HALL. f. Med. Rijksherb. 14 (1912) 38; Koorpb. Exk. Fl. 3 (1912) 285; BuysMANn, Flora 106 (1914) 115; BoLp. Zakfl. (1916) 43; Merr. Sp. Blanc. (1918) 370; ScHweRIN, Mitt. Deut. Dendr. Ges. 18 (1909) 41, ibid. (1920) 222, incl. var. bore- alis SCHWER., var. formosana (NAKAI) SCHWER. & var. meridionalis SCHWER.; LORZING, Trop. Natuur 10 (1921) 103, f. 6; WELSEM, op. cit. 181-183; BACK. & Stoor. Theeonkr. (1924) 211; Moore, J. Bot. (1924) 46; Merr. En. Philip. 3 (1923) 576; Contr. Arn. Arb. 8 (1934) 164; Comm. Lour. (1935) 375; P’e1, Bot. Bull. Ac. Sin. 1 (1947) 8; BAck. Bekn. Fl. Java em. ed. 8 (1949) fam. 175, p. 1.—S. angustifolia NORONHA, Verh. Bat. Gen. 5 (1790) 85, nomen, cf. Hassk. l1.c.—S. canadensis (non L.) Toons. Fl. Jap. (1784) 126; Fl. Jav. (1825) 5; HAssk. Cat. Hort. Bog. (1844) 117, an T. & B. ibid. (1866) 120?—S. chinensis LINDL. Trans. Hort. Soc. Lond. 6 (1826) 297; DC. /.c.; ScHEFF. Nat. Tijd. N.I. 34 (1874) 41.—S. thunbergiana BL. ex Mia. Ann. Mus. L. B. 2 (1866) 265; Kurz, For. FI. Burma 2 (1877) 3.—S. hookeri REHDER in SARG. Pl. Wils. 2 (1912) 308.—S. formosana NAKAI, Bot. Mag. Tokyo 31 (1917) 211.—Ebulum formosanum . 3. S. adnata . 2. S. canadensis Nakal, J. Coll. Sc. Tokyo 31 (1917) 211.—Fig. If. Little branched, more or less weedy, erect shrub 1—3!/2 m. Pith white. Twigs terete, nodes between the petioles often with recurved, rarely foliaceous, stipular appendages. Leaflets 2—6-jugate, higher ones often 1-—3-foliolate, rachis often with foli- aceous appendages at the base of the leaflets, upper- most leaflets very rarely adnate to the terminal leaflet and the rachis. Leaflets oblong to linear- lanceolate, shortly stalked to sessile, sometimes adnate to the rachis, base cuneate to cordate, sym- metric to oblique, mostly opposite, rarely alternate, apex acuminate, shallowly serrate, lower teeth glandular-swollen, 7-22 by 1'/2-6!/2cm; midrib and base of side-nerves hairy above, hairy or glabr- ous beneath. Corymbs fiat-topped, 3—-5—(7-) rayed, mostly hairy to subglabrous, with a foliate base, near the branching often with stalked glands. Part of the fis deformed into erect, stalked, yellow, orange or sometimes (?) green, top-shaped, per- sistent, nectaries, 3 mm diam., impressed at the top.! Pedicels 0-2 mm. Calyx-tube sulcate length- wise, lobes triangular, acute, persistent, !/2 mm. Corolla white or creamy, fragrant (as in S. nigra), (1!/2-)21/2-31/2 mm long, lobes acute, valvate in bud, (1—)2—2!/2 mm. Filaments mostly + !/2 mm, as long as the yellow anthers not exceeding the co- rolla-lobes, the latter spreading, rarely reflexed. (1) Not galls as DANGUuy (F. Gén. I.C. 3, 1922, 4) supposes (sic/). 192 FLORA MALESIANA [ser. I, vol. 43 Stigmas 3, narrowed towards the apex. Ovary 3- celled, sulcate. Ripe berry ovoid, black, 3-seeded, 3-4 mm diam. Seeds ovate, pointed 2 mm long. Distr. SE.-E. Asia, Japan, Formosa, in Ma- laysia: not E. of the line Philippines-Celebes-Lom- bok, but not yet found in the Malay Peninsula! Ecol. Everwet primary and secondary forests, thickets, clearings, mostly in -+- shaded localities, (350-)700-2000 m, f/. fr. Jan._Dec. The nectaries have been actually found to contain honey. The habit has been described as ‘straggling’, but this is certainly incorrect. Vern. Ki katumpang, ki tamblég, ki téspong, bébédjaran, S, mantjo, séngitan, J, galamat, kala- mat (Ig.), sauco (Sp.), sélando (Karo-Batak), Ja- vaanse ylier, D, tatamaikang (Minah.) Notes. P’er describes the fruits as red from China. It is quite possible that Malaysian speci- mens represent some varieties; e.g. I found in a specimen from Sumatra (Doct. v. LEEUWEN 12831) the petals recurved and filaments twice as long as the anthers instead of as long as these. In hairy young infl. the flowers are mostly subsessile. Along with normal flowers I found sometimes sta- mens exposed in flowers with apparently aborted corollas. S. javanica is, apparently, the only repre- sentative of the section Scyphidanthe Mia. S. hoo- keri REHD. does not appear to be different. 2. Sambucus canadensis LINNE, Sp.PI. (1753) 269; DC. Prod. 4 (1830) 322; ScHwerIN, Mitt. Deut. Dendr. Ges. (1909) 35; ibid. (1920) 215; v. WELSEM, Trop. Natuur 10 (1928) 181-183, cum ic.; BRUGGE- MAN, Ind. Tuinb. (1939) 263, f. 268; BACKER, Bekn. Fl. Java em. ed. 8 (1949) fam. 175, p. 2.—S. bipin- nata MOENCH. Meth. PI. (1794) t. 506, ?non SCHL. & CHAM. (1830).—Fig. 1d. Shrub 1—3!/2 m, forming subterranean sprouts. Twigs with stipular glandular appendages on the nodes or not, 3—5-jugate, in the larger /eaves the lowest 1-2 pairs of leaflets mostly 2-3-foliolate, rachis not rarely with small, gland-tipped, leafy appendages; leaflets opposite, ovate to lanceolate, base variable, apex acuminate, base of margin entire, above serrate, 4!/2-12 by 11/24 cm; midrib hairy on both sides, underside hairy on the nerves. Corymbs 20-45 cm diam., convex, (4~)5-rayed, + glabrous, axes purple. Pedicels 5-6 mm. Flowers feebly fragrant. Calyx-tube 1 mm high, hardly sulcate, lobes ovate, blunt, after anthesis appressed, in fruit dark purple. Corolla creamy, 4—4!/2 mm, lobes blunt, often toothed, 3—3!/2 mm, imbricate in bud. Filaments small in bud, in anthesis 3 mm, spreading. Anthers yellow, oblong. Ovary 5-celled, stigmas 5. Berry globular, subapplanate, shiny black-purple, 4-5 mm diam., (4-)5-seeded. Seeds + 2!'/2mm long, oblong-elliptic. Distr. N. America, said to have been imported in Java as late as 1918 from Indochina by the Botanic Gardens, Bogor (vy. WELSEM, /.c.), but rapidly dispersed in gardens throughout Malaysia as an excellent, often rich-flowering, ornamental. Ecol. Cultivated, 200-1300m, ff. Jan.—Dec. The flowers remain opened for several days, they do not produce honey in Java; easily propagated by cuttings; if cut twigs are partly defoliated, the infl. remains fresh for several days. Vern. Amerikaanse vlier, D, American elder, E. Notes. The differences between S. canadensis L. and S. mexicana PRest ex DC. [Prod. 4 (1830) 322; STANDLEY, Tr. & Shr. Mexico (1926) 1395;— ‘S. bipinnata SCHL. & CHAM. Linnaea 5 (1830) 171; CORNER, Wayside Trees (1940) 182, t. 40] seem slight and both spp. were considered conspecific by SARGENT. The main differences are apparently the oblong shape and sulcate surface of the berry in S. mexicana. The 2 specimens of S. mexicana 1 could examine had smaller flowers and more roundish anthers than those of S. canadensis. 3. Sambucus adnata WALL. ex DC. Prod. 4 (1830) 322; Hook. f. Fl. Br. Ind. 3 (1882) 3; SCHWERIN, Mitt. Deut. Dendr. Ges. (1909) 41; P’Er, Bot. Bull. Ac. Sin. 1 (1947) 7.—Fig. le. Distr. Himalaya to China, ?E. Africa; mountains. This species does not belong to § Scyphidanthe Mig. though SCHWERIN treated it as such; it is more related to S. canadensis. Entirely different from S. javanica by absence of nectaries, oblong- elliptic 2!/2 -3 mm long seeds, apparently constant- ly 1-2 pairs of upper leaflets connate with the rachis, oblong and larger berries (type at Kew!). There is one specimen in Herb. Leiden (899.-69— 80) said to come from Celebes, but the specimen consists of one leaf only and is too poor for proper identification. Its upper leaflets are adnate, but this is a feature also occurring, though very rarely, in S. javanica. | doubt the occurrence of S. adnata, which is a native of continental SE. Asia (India to China), in Malaysia. 4. CARLEMANNIA BENTH. in Hook. J. Bot. 5 (1853) 307; B. & H. Gen. Pl. 2 (1873) 63; Hook. f. FI. Br. Ind. 3 (1880) 85; BAILL. Hist. Pl. 7 (1880) 468; PiTaRD, FI. Gén. I.-C. 3(1923) 167; SOLEREDER, Bull. Herb. Boiss. 1 (1893) 173-8; K. Scu. in E. & P. 4, 4 (1897) 31; SOLEREDER, Syst. Anat. Dik. Erganz. (1908) 173, HALLIER f. in Just’s Bot. Jahrb. 36, 3 (1910) 221; Med. Rijksherb. no 1 (1910) 40; ibid. 14 (1912) 38; Arch. Néerl. Sc. Ex. Nat. IIB, 1 (1912) 224; BREMEKAmMP, Rec. Trav. Bot. Néerl. 36 (1939) 372. Generally little-branched herbs, subglabrous or hairy, apparently erect. Inflo- rescence with sparse sessile or stalked, capitate-glandular hairs. Leaves mostly 193 Dec. 1951] CAPRIFOLIACEAE (Kern & v. Steenis) obliquely elliptic, crenate-dentate, parenchyma apparently with glands at the un- derside. Petioles connected by a raised line, in which no interpetiolar stipule can be distinguished. Flowers white, yellowish or pink, in short-peduncled terminal sometimes also axillary corymbs. Calyx-tube globose, constricted at the apex; lobes 4-5, subequal or unequal, persistent. Corolla tubular, club-shaped in bud, circumsciss at the base, lobes 4, rather narrowly imbricate in bud, two outer two inner. Stamens 2, inserted in the middle of the tube, alternating with the lobes, dorsifix, cohering in bud and enclosing the style; filaments short, terete or some- what flattened, extending towards the corolla base but distinctly forming part of the corolla tube; anthers linear, latrorse, included in the tube, relatively large, opened already in bud, connective apiculate. Pollen granular. Disk annular-short- cylindric, consisting of two parts alternating with the ovarial cells, placed below the anthers. Ovary 2-celled, each cell with ~ ovules inserted on a thick basal pla- centa. Style filiform, included, stigma clavate, erect, 2-fid. Capsule membranous, + globular, or more or less pyramidal, + inflated, and 4-lobed, each lobe with a few veins, 2-celled, loculicidally 2-valved. Seeds ~, minute, oval, testa smooth(?) or reticulated, albumen granular or horny, embryo minute at the top of the albu- men (ex auct.). Distr. Three spp., Himalaya & Mishmi Hills, Tonkin, and Yunnan, in Malaysia: Sumatra. Ecol. Montane forest plants. Notes. For a discussion on the systematic position compare the notes at the head of the family. HAL.igR, after having switched in a few years his opinion from Gesneraceae to Saxifragaceae, finally agreed with SOLEREDER assigning this genus to Caprifoliaceae. Few figures exist of the genus; that of SCHUMANN is wrong. The flowers are 4-merous and the corolla is definitely imbricate in bud; the number of sepals may vary to 5. The relative large size of the stamens, their coherence in bud enclosing the style, the anther-cells shedding pollen in bud, point to an unusual anthesis, on which no field observations are available. The typical ‘glassy’ hairs, constricted apex of the calyx, and dorsifixed anthers remind of a longitudinal section of the flower of Dentella, also a plant without interpetiolar stipules. The capitate-glandular hairs, typical cymose infl. and symmetric flower point towards Lonicera, the reduction in the androecium, unknown or extremely rare in Rubiaceae, to Valerianaceae acc. to BREMEKAMP. 1. Carlemannia tetragona Hook. f. Fl. Br. Ind. 3. and inside glabrous, prob. pale, 6—6!/2 mm, lobes (1880) 85; DRAKE in Morot, J. de Bot. 9 (1895) slightly unequal, the larger ones ovate, rounded, DS EImARD. Els Gen I-©, 3) (1923)) 1680159s £ + 13/4 by 11/2 mm. Disk short-cylindric, !/2 mm 16477 incl. var. tonkinensis—Carlemannia suma- trana RID. J. Str. Br. R. As. Soc. 1 (1923) 66; STEEN. Bull. J.B.B. III, 13 (1934) 246; BRemE- KAMP, Rec. Trav. Bot. Néerl. 36 (1939) 372.— C. henryi LEv. in FeppE, Rep. 13 (1914) 178. —Fig. Ic. Puberulous delicate herb, 20-50 cm. Stems terete, older nodes articulated. Leaves membra- nous, ovate-oblong to lanceolate, base and apex acute to subacuminate, sparsely white-hairy on both sides, hairs relatively coarse, 4—9 by 1!/2—33/4 cm, primary nerves 5 pairs; petiole 2!/2 cm. Co- rymbs terminal, congested to dense, above 2 re- duced spathulate leaves, + 2!/2 cm diam., puber- ulous, not or hardly exceeding the leaves, 7— to many-flowered, each pair with a reduced cyme at the base; peduncle 1—2 cm. Fis in pairs, only 1 nar- row bract resembling the sepals; pedicels 1 mm; no bracteoles observed. Calyx tube hairy, 1 mm; lobes 4—5, linear, erect, hairy, 3-4 mm, exceeding the middle of the mature bud, in fruit 3-nerved, equal. Corolla hairy towards the tip by thick glassy hairs on a raised bulbous base, imbricated margins high, consisting of two parts, margin slightly cre- nate, lobes below the anthers. Filaments 1 mm, anthers 2!/2mm, linear, opened in bud, their tip just reaching the throat of the corolla. Style fili- form, its base fusiform-swollen above the disk, glabrous, 5 mm; stigmatic arms narrow, - 1 mm, fibrous, also in opened flowers often sticking to one of the anthers, not exceeding the stamens. Capsule 4—5 mm through, distinctly 4-lobed or 4- horny at the base, broader than high. Seeds ovate- oblong, hardly 1 mm, testa reticulate. Distr. Mishmi Hills (NE. Himalaya), Tonkin, Yunnan in Malaysia: NE. Sumatra (mountains above Medan-Deli), apparently local. Ecol. In forests, 1000-1300 m, f7. Dec.—July. Notes. Allied to C. griffithii Hoox. f. which differs by oblong sepals, 1-2 mm, distinctly unequal, and not reaching the middle of the corolla, larger fils, shorter filaments and + globular fruit without distinctly stellate-spreading basal lobes. I have not seen the fruit lobes dehiscing as RIDLEY mentions. The size of the leaves by which the Indochinese variety was distinguished seems of little value. The 194 FLORA MALESIANA [ser. I, vol. 44, Deceaig@sa ti type material is in a slightly older state than the Sumatran specimens are, but I am perfectly satis- fied about their identity. Excluded Lonicera GAERTN. = Loranthaceae. Lonicera chinensis Wats., L. confusa DC., L. ja- vanica DC., and L. macrantha DC. were mentioned to occur in the Philippines by F.—-VILLAR, Nov. App. (1880) 104, but these records have been ex- cluded by MERRILL, Gov. Lab. Publ. (Philip.) 29 (1905) 50; En. Philip. 3 (1923) 578. Lonicera symphoricarpus (non L.) BLANCO, FI. Filip. (1837) 161. = Scurrula philippinensis (CHAM. & SCHLECHT.) G. Don, cf. DANSER, Philip. J.Sc. 58 (1935) 121. Viburnum zippelii Miaq. Fl. Ind. Bat. 2 (1856) 122; SCHEFFER, Ann. Jard. Bot. Btzg-1 (1876) 28; Hat- LIER f. Med. Rijksherb. 14 (1912) 36 = V. japoni- cum (THUNB.) SPR., wrongly recorded for New Guinea. Cf. Kern, Reinw. 1 (1951) 165. Viburnum alternifolium Zot. & Mor. Syst. Verz. (1845-1846) 59 = Ilex alternifolia (ZOLL. & Mor.) Logs. POLEMONIACEAE (C. A. Backer, Heemstede) Annual or perennial herbs or shrubs, sometimes climbing by means of foliar tendrils, rarely small trees. Leaves spirally arranged or opposite, exstipulate, sessile or petioled, entire or more or less deeply divided, or compound. Flowers axillary or termi- nal, solitary, geminate, corymbose or capitate, actinomorphic or slightly zygo- morphic. Calyx 5-lobed or 5-partite, with or without transparent fields, persistent. Corolla gamophyllous, 5-lobed or 5-partite; lobes contorted in bud. Stamens 5, on the corolla-tube, inserted at equal or unequal height, alternating with the segments; filaments free from each other, included or exserted; anthers dorsifixed, 2-celled; cells opening longitudinally. Ovary superior, sessile on a disk, 3 (rarely 2)-celled; ovules in each cell l—-., inserted in the inner angle; style 1, filiform, 3 (rarely 2)-fid. Fruit a loculicidal or septifragal capsule, rarely indehiscent. Endosperm mostly copious; embryo straight or slightly curved. Distr. N. America and the Andes, rare in the Old World, absent from Africa and Australia. Genera 12, represented by upwards of 250 species. In Ma/aysia one American genus is more or less naturalized; a few other species are cultivated in gardens. Uses. The Indonesian species are exclusively of horticultural value. KEY TO THE GENERA 1. Climbing. Leaves pinnate; rachis terminating in a branched tendril. Flowers solitary or pairwise in the leaf-axils, long-peduncled, rather large. Fruit septifragal. . . . =o)