"fl TAXONOMICAL REVISIONS REPUBLIK INDONESIA REPUBLIC OF INDONESIA LEMBAGA ILMU PENGETAHUAN INDONESIA (L.I.P.I.) INDONESIAN INSTITUTE OF SCIENCES FLORA MALESIANA BEING AN ILLUSTRATED SYSTEMATIC ACCOUNT OF THE MALESIAN FLORA / INCLUDING KEYS FOR DETERMINATION I DIAGNOSTIC DESCRIPTIONS I REFERENCES TO THE LITERATURE I S YNON YM Y I AND DISTRIBUTION / AND NOTES ON THE ECOLOGY OF ITS WILD AND COMMONLY CULTIVATED PLANTS PUBLISHED UNDER THE AUSPICES OF LEMBAGA BIOLOGI NASIONAL BOTANIC GARDENS OF INDONESIA / BOGOR / JAVA AND OF THE RIJKSHERBARIUM / LEYDEN / NETHERLANDS 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 ^^.VO^/:^-' VOLUME 9 SPERM A TOPHYTA GENERAL EDITOR: Dr C. G. G. J. VAN STEENIS DIRECTOR OF THE FOUNDATION 'FLORA MALESIANA' PUBLISHED BY: MARTINUS NIJHOFF / DR W. JUNK PUBLISHERS THE HAGUE / BOSTON / LONDON 1979-1983 Distributors: for the United States and Canada Kluwer Boston. Inc. 190 Old Derby Street Hingham. MA 02043 USA for all other countries Kluwer Academic Publishers Group Distribution Cen-ter P.O. Box 322 3300 AH Dordrecht The Netherlands AHA.. I pi- H Library of Congress Cataloging in Publication Data dP (Revised for volume 9, part 3) Main entry under title: Flora Malesiana. At head of title: Republic of Indonesia, Ministry of Agriculture. Issued in pts. Includes bibliographical references. 1. Botany — Malay Archipelago — Collected works. 2. Phytogeography — Collected works. 3. Plant collectors — Malay Archipelago — Collected works. I. Steenis, Cornells Gijsbert Gerrit Jan van, ed. II. Indonesia. Departemen Pertanian. III. Series. QK366.F56 582.09598 72-175112 ISBN 90-247-2780-4 (this volume) Publication dates Part 1 27 Dec. 1979 Part 2 23 July 1982 Part 3 28 Jan. 1983 Copyright c, 1983 by Martinus Nijhoff Publishers. The Hague. All rights reserved. No part of this publication may be reproduced, stored in a retrieval system. or transmitted in any form or by any means, mechanical, photocopying, recording, or otherwise, without the prior written permission of the publisher. Martinus Nijhoff Publishers. P.O. Box 566. 2501 CN The Hague. The Netherlands. PRINTED IN THE N'ETHERLANDS CONTENTS Tiile-page (3) Contents (5) Dedication by R.E.G. Pichi Sermolli &i C. G.G.J. \an Steenis (7) Abbreviations and signs (45) TAXONOMICAL REVISIONS in alphabetical sequence Araliaceae-I by \\".R, Philipson 1 Cyperaceae-II by J.H. Kern L H.P. Nooteboom 107 Dipterocarpaceae by P.S. .Ashton 237 Liliaceae-I by J. P. Jessop 189 ADDENDA to volumes 4-9 Addenda, corrigenda el emendanda by C. G.G.J, van Steenis et al 553 INDEX Index to scientific plant names b\ M.J. \an Steenis-Kruseman &: E.E. \an Nieuwkoop 575 1864 1906 A '4if-^,^u^ ;Li^ cj>J^, ff^ ■A'u^LAt^ ^^ ■ZA^ 1918 Dedicated to the memory of ODOARDO BECCARI DEDICATION A dedication to Odoardo Beccari, the greatest botanist ever to study in Malesia, is long overdue. Although best known as a plant taxonomist, his versatile genius extended far beyond the basic field of this branch of Botany, his wide interest leading him to investigate the laws of evolution, the interrelations between plants and animals, the connection between vegetation and environ- ment, plant distribution, the cultivated and useful plants of Malesia and many other problems of plant life. But, even if he devoted his studies to plants, in the depth of his mind he was primarily a naturalist, and in his long, lonely and dangerous explorations in Malesia he was attracted to all aspects of nature and human life, assembling, besides plants, an incredibly large number of collec- tions and an invaluable wealth of drawings and observations in zoology, anthropology and ethnol- ogy. He was indeed a naturalist, and one of the greatest of his time; but never in his mind were the knowledge and beauty of Nature disjoined, and, as he was a true and complete naturalist, he was at the same time a poet and an artist. His Nelleforeste di Borneo, Viaggi e ricerche di un naturalista ( 1 902), excellently translated into English (in a somewhat abbreviated form) by Prof. E. GiGLiOLiand revised and edited by F.H.H. GuiLLEMARoas Wanderings in the great forests of Borneo (1904), is a treasure in tropical botany; it is in fact an unrivalled introduction to tropical plant life and animals, man included. It is a most readable book touching on all sorts of topics and we advise it to be studied by all young people whose ambition it is to devote their life to tropical research. In the last years of his life, Beccari was rearranging his diaries, notes and observations of the expeditions to eastern Malesia with the intention of publishing a second book on his explorations, but very unfortunately death did not allow him to carry out his wish. He left only a revised copy of his diaries and field notes which formed the bulk of the book Nuova Guinea, Selebes e Moluc- che, published posthumously by his son Nello Beccari in 1924. It is neither well known nor duly appreciated outside Italy, since no translation has been published. Undoubtedly it lacks the glam- our and freshness of the previous book, being devoid of the original and acute observations deriv- ed from his field research, which are largely profuse in his Nelle foreste di Borneo, but it offers a good and fascinating description of his adventurous travels with a wealth of interesting remarks, and it is an invaluable documentation of the natural features of those almost unexplored coun- tries, particularly of the characteristics and customs of their inhabitants. This synthesis, however, is only part of his oeuvre and before entering on his achievements, let us first look at his life and the development of his ideas and ideals. Beccari's early youth was ill fated. He was born in Florence, in his father's home in the Via dei Benci at the corner of Borgo dei Greci, on November 16, 1843. His mother, Antonietta Mi- Nucci, from Radda in Chianti in Tuscany, died soon after his birth, and his father Giuseppe Bec- cari, from an ancient family native of Rimini (Romagna), died in 1849 when Odoardo was six years old; he was brought up by his maternal uncle Minuccio Minucci. In April 1853 he entered the Collegio 'Ferdinando' in Lucca, where his love for botany was nurtured by the Vice Rector and Prefect of Studies, the Abbe Ignazio Mezzetti ' and by his Professor of Botany in the Lyceum of Lucca and Director of the Botanic Garden, Cesare Bicchi. The latter, aware of the talent of his pupil and perhaps foreseeing his glorious future, in 1860 dedicated to him a new spe- cies, Tulipa beccariana^, the first of the numerous plants and animals to be named in his honour. Beccari's first collections date back to 1856, when he was a student of the College of Lucca and still only 13 years old. During his stay there he assembled a herbarium, which was still in exis- (1) In his honour, in 1871, Beccari named a new genus of Annonaceae Mezzettia. (2) Tulipa beccariana Bicchi, Agg. Fl. Lucch. (1860) 21, nom. nud.; 1 Giardini 8 (1861) 50, t. 2. (7) Flora Malesiana tence at the beginning of the present century. A search for it in Lucca as a separate herbarium was unsuccessful, but several specimens with labels headed 'Erbario Beccari' are present in the herbarium of Ignazio Mezzetti, now kept in the Lyceum Machiavelli of Lucca, where Beccari attended secondary school until July 1861. Whether these specimens are part of the separate Bec- cari herbarium included in Mezzetti's herbarium or duplicates of it, is uncertain. Other plants collected in the period in which Beccari was a student in Lucca are kept in Webb's herbarium in Florence. In August 1861 he published his first paper and in the autumn of the same year, when he was 18 years old, Beccari commenced his studies in the Faculty of Natural Science at the University of Pisa. At first, perhaps under the influence of Bicchi, he devoted his attention to the Cryp- togams and already in 1 862 Beccari's name appears together with those of several eminent botan- ists of the time, among the collectors of the 'Erbario Crittogamico Italiano', the classical exsiccata with printed labels, founded in 1858 by Giuseppe de Notaris of Genoa. In the University of Pisa, Beccari distinguished himself so much in botany, that the celebrated botanist Pietro SAVimade him an assistant to the chair of botany in January 1863, while he was still an undergraduate. Dissatisfied with the conservatism of Savi, however, he gave up his assis- tantship and moved to the University of Bologna from where he took his degree in Natural Science on July 1, 1864, by disputing his thesis on the structure of the lichen Arnoldia cyathodes Mass. [ = Plectopsora cyathodes (Mass.) Korber] with the famous professor in botany Antonio Berto- LONI. Before his graduation, Beccari had already planned a long journey to far away regions, when in June 1864, in the laboratory of Prof. Giovanni Capellini, geologist at the University of Bo- logna, he met Marquis Giacomo Doria, a young, impassioned naturalist, later patron and Maece- nas of science, and founder of the Museo Civico di Storia Naturale at Genoa which bears his name. The two young men, united by the same enthusiasm for scientific exploration in unknown countries, soon fraternized. Shortly after his graduation, Beccari visited Doria at Genoa, where they decided to undertake a long exploration together and, counselled also by the celebrated British naturalist John Ball, they chose the Kingdom of Sarawak, in Borneo, as the destination of their enterprise. As part of his preparations, Beccari spent the period from February to April 1865 in the great British centres of botany, the British Museum in London and Kew Gardens, ob- viously to acquaint himself roughly with the plants of Borneo. He met the Hookers, Charles Darwin and Sir James Brooke, the Rajah of Sarawak, who assured him of the assistance of his nephew, the Tuan-muda, Sir Charles Brooke, then governing the territory in his absence. Thus prepared, young Odoardo, at the age of 22, commenced his studies on the flora of the Malesian tropics, which was to become his main life occupation, and in which he would rise to the greatest heights as a scientific explorer, naturalist, and botanist. He sailed from Southampton on April 4, 1865, and met DoRL\and his own brother, Giovanni Battista Beccari (who was on his way to Japan), at Alexandria. From there they travelled by train to Suez and by boat to Aden and then to Ceylon, where they spent a fortnight. There Beccari visited the famous Botanic Gar- dens at Peradeniya and climbed Mt Petrotallagalla, where he made his first personal acquaintance with the tropical flora and started collecting. Via Penang island and Singapore, the voyagers arriv- ed on June 19, 1865 at Kuching, capital of Sarawak, which they had selected as their base of opera- tions. At the beginning they were guests of the Tuan-muda, Sir Charles Brooke; later they settled in a house of their own with servants, and also bought a small boat ('sampan') for their excursions in the forests along the river. Soon Beccari and Doria took up their botanical and zoological col- lecting in the dense and primitive forests which at that time surrounded Kuching. Beccari was anxious to know the mountains and to collect intensively; thus he undertook the construction of a (8) Dedication big hut in the forest of Gunong Mattang at an altitude of about 300 m with the intention of making it the base for the explorations and collections in the primitive forests of Borneo. However, after some months the health of Doria deteriorated to such a point that at the beginning of March 1866 he was forced to return to Italy. Thus, Beccari, having accompanied his friend to Singapore, re- mained alone to carry out the programme which he had planned with him and had hoped to realize together. At the beginning of April he moved to his house in the forest of Gunong Mattang which he called 'Vallombrosa', after the great monastery hidden in the dense mountain forest of Prato- magno, east of Florence. Together with a Chinese cook and four Malesian boys he spent nearly all the remaining months of 1866 there, except for four excursions and a period in October-No- vember at Kuching to pack his collections. On one of these excursions to Gunong Poe he discov- ered a new species of Rafflesia, the famous parasitic plant with vegetative parts extremely reduced and a gigantic flower, the largest in the plant kingdom, apparently arising directly from the stem of the host liana. The largest flower of the new species, Rafflesia tuan-mudae, so named in honour of the Tuan-muda of Sarawak, Sir Charles Brooke, attained about 56 cm in diameter. In Nelle foreste di Borneo Beccari gives many details of his hut at Mattang. He had cleverly designed it for drying plants and preparing zoological specimens rather than for lodging; soon it became an active and efficient laboratory, full of all sorts of products of nature. In his book he described his primitive life there as very happy and fully suited to his temperament. With only a cotton coat, trousers and a Chinese straw hat, mostly bare-footed, he carefully explored the sur- rounding primary forest, assembling marvellous collections of plants and animals. Back at his hut, he devoted many hours to arranging his collection, making drawings and descriptions and recording those notes and observations which later became the basis of his fascinating book Nelle foreste di Borneo. At the beginning of 1867 Beccari abandoned the hut at Mattang and spent the first two months at Kuching collecting in the surroundings, but chiefly arranging and packing his large collections. In March 1867 he again undertook his adventurous wanderings with the intention of visiting the interior of Sarawak. One of his trips from mid-March to the last days of xMay was devoted to the exploration of Batang-Lupar and the lakes of Kapuas with the main purpose of hunting orang- utan. He assembled there one of the best collections of these animals (skin, skeletons, heads and skulls, and even a foetus) and a wealth of observations which allowed him to express the opinion that the hominids did not originate in dense forest, like that of Borneo, and that the orang-utan, particularly well adapted to an arboreous environment, would be, not an ancestor, but a collateral of man. In his opinion, the hominids were derived from forerunners, allied to the great anthro- poids of tropical Africa, with an anatomical conformation, particularly of the limbs, more suited to evolve towards a biped gait and an erect habit and they had their origin in more open vegetation, like that of some regions of tropical Africa, where we find the greatest number of large mammals with rapid locomotion. Recent research in south-western Ethiopia seems to support this hypo- thesis. From August 12 to September 14, 1867, Beccari collected in the district of Bintulu and in the country of the Kayan. From there, he was looking forward to organizing an expedition to the inte- rior regions of Sarawak, which at that time were still nearly unexplored and hardly visited by Europeans; but his project found every possible difficulty and obstacle. Despite them, without guide or interpreter, but with only four men and a small boat, he set out on September 15 from Bintulu on his journey through the interior of Sarawak along the basins of the main rivers Bintulu, Redjang and Batang Lupar and their tributaries, across the ridges of hills and mountains which represent the watershed between them. The journey was made mostly sailing up, or down, the rivers in various native paddle boats obtained from time to time from the natives, but also on foot (9) Flora Malesiana to overcome some impassable rapids or to cross tlie ridge between two adjacent basins; often he was forced to walk with difficulty in the stream beds, or to proceed slowly with a compass through the dense forests; more than once he was in real danger, even near the end of his travels when, having lost his compass, without food, in an unhabited region, he got lost for two days in a dense forest. On November 20, 1867, Beccari arrived at Kuching where he concluded this long, hard and risky enterprise. In the first two years of his stay in Borneo Beccari's health remained excellent, but in the last months it had been deteriorating. Already in June 1867 he had suffered the first attack of malaria and later many others followed. Furthermore, in July of the same year he had observed the first symptoms of elephantiasis on his right ankle. After the expedition to the interior of Sarawak he spent two months in Kuching arranging and packing the collections he had made. He had planned another long journey crossing the inland of Borneo from Kuching to Pontianak; but in January his health worsened, and being unable to subdue the high fever which had troubled him for some days, BECCARiwas forced to undertake his homeward-bound voyage. He left Kuching on January 29, 1868 and arrived in Italy on March 2, after explorations in Borneo which had lasted almost three years. In Florence Beccari was the guest of his old friend from the College of Lucca, Emilio Marcuc- ci, who had taken up the profession of architect but had not given up his love of botany, and who greatly assisted Beccari in that period in recovering his health. The house was located in Borgo Tegolaio 48, very close to the Museum of Physics and Natural History, where Beccari had prob- ably assembled his collections. The house soon became a meeting place for young lovers of natural history including Levier and Sommier. Soon after his return from Borneo G. Doria and R. Gestro, from the Civic Museum of Natural History of Genoa, were also his guests for several days, evidently to be informed, in detail, about the large zoological collections he had assembled in Borneo. At that time Beccari was very busy sorting out and working on his collections; he also made agreements with collaborating specialists to study particular groups such as seagrasses (Ascher- SON 1871), pteridophytes (Cesati1876), mosses (Hampe 1872), lichens (von Krempelhuber 1875) and hepatics (de Notaris 1876), etc. (see Appendix 4); he probably also started distributing dupli- cate specimens of his Bornean plants. However, together with his technical work, he carried on with the study of his collection. In March 1869, Beccari started, at his own expense, the publication of a new periodical, the Nuovo Giorna/e Botanico Italiano, w hich was intended as a replacement of the Giornale Botanico Italiano founded in 1844 by F. Parlatore, but interrupted in 1852. Beccari edited three volumes of the new periodical (1869, 1870, 1871) with the help of his friend Marcucci, to whom he dedi- cated the new genus Marcuccia (Annonaceae) as a sign of gratitude for the help received in editing these volumes, particularly during his travels in Ethiopia in 1870. His first accounts of Bornean plants appeared in early volumes of his journal and many other papers dealing with his collections written by himself and other botanists were published in subsequent volumes. However, in spring 1871, when preparing for his expedition to New Guinea, Beccari became aware of the difficulties of editing a journal when abroad making long expeditions in distant regions and handed the man- agement of the Nuovo Giornale Botanico Italiano over to T. Caruel, who edited it until the end of 1893, when the journal became the official publication of the Italian Botanical Society, which it continues to be. Fascinated by his primitive life in Borneo, Beccari was not satisfied with city life. Probably he had already developed the idea of undertaking a second journey to Malesia, when he received an offer to join an Italian expedition to Ethiopia. He sailed on February 14, 1870, from Genoa and (10) Dedication together with the zoologist Marquis Orazio Antinori and Prof. Arturo Issel, geologist at the University of Genoa, visited the Bay of Assab and later, on behalf of the Italian Geographical Society, the country of Bogos. There he assembled a rich collection of plants (3 1 5 species of sper- matophytes and pteridophytes and 289 species of mosses, algae, fungi and lichens) enumerated and partly described in Martelli's Florula Bogosensis (1886). He came back to Italy on October 20, 1870. Soon after his return to Florence, Beccari materialized his project to visit Malesia again and after careful preparations, training himself in geodetics, astronomy and meteorology, he set out on November 24, 1871 from Genoa for the island of New Guinea, accompanied by Count Luigi Maria d'Albertis, an Italian nobleman who was passionately fond of hunting and natural history. Their first visit was to West Java where they stayed for some time in the Botanic Gardens at Bogor. The young director. Dr. Rudolph Scheffer, must have facilitated his exploring for a few days on Mt Gedeh, with its Tjibodas mountain garden, and adjacent primary forest on Mts Pangerango and Megamendong. Further stops were made at Flores and Timor in the Lesser Sunda Islands, and the islands of Banda and Ambon, where they arrived on March 7, 1872, and enjoyed the kind and helpful hospitality of Captain P.F. Kraal and his wife, the Italian lady Amall\ Ma- LAN. After a short journey to Buru and Ceram for information they came back to Ambon, where they organized the expedition to western New Guinea renting a small schooner, the 'Bu- rung-Laut', of 25 tons with a crew of eight men. On March 21, 1872, Beccari and d'Albertis sailed from Ambon to New Guinea, and having touched the islands of Geser and Goram reached Kapaor on the west coast of the mainland. Later, on April 30, they arrived at the small island of Sorong where they dismissed the Burung-Laut and rented a hut in which they established their first scientific station, collecting chiefly along the Ramoi river. They remained on Sorong island till July 15 when, having left their collections there with one of their men as keeper, they sailed in a ramshackle indigenous sailing boat with a crew of eleven Papuas to Dorei and later to Andai, where they arrived on August 7 after a long voyage, full of adventures. Beccari and d'Albertis established their home and headquarters for the exploration of the region in a large Papuan hut built on high palafittes near the Andai river surrounded by the forest, with a splendid view of both the sea and the Arfak Mts. While d'Albertis made a trip in the mountains, Beccari collected intensively in the vicinity of Andai, but on September 28 he moved to Putat on the lower slopes of the Arfak Mts with the intention of exploring the higher regions. Unfortunately on October 9, he was informed that d'Albertis had fallen seriously ill and he was compelled to return to Andai. The poor health of d'Albertis necessitated that the travellers return to Ambon, but only on November 2 was Beccari able to find an indigenous boat to reach Sorong and later Ambon. But in Sorong, unexpectedly, they found a schooner sent from Ambon in search of them. After recovering, safe and sound, the collections left there four months before, they sail- ed to Ambon, where they arrived on December 5, 1872. There Beccari and d'Albertis were greatly surprised to find the Italian Royal Corvette 'Vettor Pisani' on which d'Albertis obtained a passage, leaving his friend alone. Thus, Beccari concluded his first expedition to New Guinea, during which, despite all sorts of difficuhies and serious health troubles, he had assembled a col- lection of about 700 species of plants and a rich amount of zoological, ethnographical and mine- ralogical specimens. Beccari remained in Ambon for about two months to arrange and pack the collections. There, as a guest of Captain Kraal and his wife, he soon regained his health and prepared a trip to the Aru and Kei islands. Beccari departed from Ambon on February 8, 1873, having obtained a passage on a Dutch (11) Flora Malesiana Government steamer. Stricken with smallpox en route, Beccari nonetheless reached the Aru is- lands on February 22, and with his base on Wokam, he collected plants and animals and made a topographical survey of the islands (see Appendix 1 , C: Maps). On July 6, he moved to the Kei islands in a big local sailing boat, a Bughis prahu, on which Beccari was the guest of its Chinese master. But the boat suffered shipwreck on the east coast of Grand Kei. Fortunately he could save all his collections and collecting equipment. Beccari found the flora of these islands unexpectedly poor and after visiting Small Kei as well, he sailed on October 4 to Ambon with four men in a small indigenous sailing boat of only 4 tons, bought at Dulan. Despite the premonitions of the natives, the risky voyage of about 350 miles was successful and on October 23 he reached Ambon, where he stayed for two weeks, partly to arrange his collections, but chiefly to recover his strength, being again a guest of his friends, the Kraals. On November 5, 1873, Beccari sailed by the steamer 'Koning Willem III' towards the West Mo- luccas (Buru and Ternate), proceeding via North Celebes towards Southwest Celebes, where he disembarked at Makassar on November 18. He stayed in the region for nearly three months until February 6, 1874. From there, as a paying passenger on an old Chinese boat of about 40 tons, similar to a prahu, he went to the larger islands south and southeast of Celebes (Kabaena and Muna) and to Kendari on the southeast coast of Celebes where he arrived on February 23. He re- mained in this district for six months to collect and make topographical surveys, but the collec- tions did not increase very much because the flora was not particularly interesting and because the region was plagued with pirates on the sea, and head-hunters on land. He chiefly collected in- land at Lepo-Lepo. Here he was informed that a Dutch vessel was looking for him at Kendari. It was the Escort vessel 'Sumatra' of the Royal Dutch Navy, which had been sent from Makassar in search of him, since it was rumoured that he was in danger from the pirates which infested the sea of Kendari. Beccari had already decided to leave Kendari and accepted with pleasure the kind offer of the Captain of the vessel to take him and his men aboard to Makassar. Having packed his collections he sailed from Kendari on August 10 and arrived at Makassar after a voyage of five days. In these last months Beccari's funds had been running out, but early in 1874 he had already written from Makassar to his friend G. DoRiAin Genoa for financial help to carry out his project of a second expedition to New Guinea. When he was back at Makassar on August 29 he received the joyfull news that his friend had convinced the authorities of Genoa to contribute 15,000 lira towards a new, second expedition to the great island. Aware that the season was not suitable for sailing to New Guinea, Beccari soon left Makassar by the same steamer 'Koning Willem III' on which he had travelled from Ambon to Makassar some months before. He proceeded to Bali, Surabaja, Semarang, and through the interior of Java to Bogor, to recuperate and to sort out his collections. There, he also spent some days at Tjibodas and on Mt Pangerango collecting. Unwearied, Beccari left Jakarta on October 15, 1874, and via Surabaja, Makassar, the island of Bima and Timor in the Lesser Sunda Islands he arrived at Ter- nate island in the Moluccas on November 11 . He remained there about 20 days and assembled rich botanical and zoological collections in the primitive forest near the hut (named by him 'Para- disino') which the Dutch Resident had built for him on the slopes of the volcano. Beccari intended to organize his travel to New Guinea from Ternate, but soon he realized that this was impossible, and on December 4 he left the island by the mail-steamer arriving at Ambon three days later. There he prepared for his new expedition financed by the Province and the Municipality of Genoa to West New Guinea, his old hunting grounds of 1872. He hired for his voyage the brig-schooner 'Deh' with a crew of 10, and accompanied by 8 men and a young boy for collecting plants and animals. (12) Dedication He sailed on January 22, 1 875, and arrived at Sorong Island on February 1 , establishing his base in the schooner and making trips to Ramoi, Dorei Hum, Mt Morait and venturing inland from Has as far as a river, the War Samson, not then recorded on the maps. Together with plants he amassed a rich collection of birds. On March 5 he left Sorong and after a visit of some days to Waigeu Island chiefly hunting for birds, he proceeded to Dorei and soon to Warbusi and Momi on the west coast of Geelvink Bay mainly with the aim of obtaining some specimens of cassowa- ries. Later, in April, always in the 'Deli', he went to the islands of the Bay which he had not pre- viously visited, spending nearly twenty days on Japen, a week on the uninhabited Mios Num, twenty days on Schouten Island and a week on a small island, Pulo Manim, near Mafor. On June 2 he arrived at Dorei where he found the Italian Corvette 'Vettor Pisani', and where he received the warmest welcome. Having arranged his collections he prepared the expedition to the Arfak Mts; on June 16 he started from Andai towards the mountains establishing his exploration base at Hatam (1500 m) in the centre of the mountain group. On June 23 he climbed one of the summits of the Arfak Mts (about 2000 m). He had planned to remain two months in the mountains, but on July 12 he was forced to cut short his exploration and to come down to the coast because of beri-beri among the crew of the schooner: two men had already died and the others were seriously ill. Thus Beccari realized that there was no choice; he had to go back to Ternate: his second ex- pedition to New Guinea was nearing its end; his dream of climbing and exploring the highest re- gions of the Arfak Mts had vanished for a second time. On July 1 8 he left Mansinam, near Andai, where the 'Deli' was riding at anchor, and on August 4, 1875, he arrived at Ternate, but in the meantime the beri-beri had killed most of the crew. He remained at Ternate three months ar- ranging his collections, notes and observations. The scientific results of the second expedition to New Guinea were very important. Even if the botanical specimens were not particularly numerous, the zoological collections were very plenti- ful, especially the skins of birds which surpassed 2000 in number, and included a set of birds of paradise which still remains one of the best of its kind. No less abundant were the ethnological collections consisting of every sort of object in use by natives. Also an important set of approxi- mately 200 Papuan skulls enriched the anthropological collections. But the expedition was also very fruitful because of his untiring activity as a naturalist and explorer in making notes of every- thing that attracted his attention and in studying all the aspects of the regions he visited. During the expedition he had also made various topographic surveys which later allowed the geographer GuiDO Cora (see Appendix 1, C: Maps) to draw maps of some regions of New Guinea, and he had assembled a great wealth of botanical, zoological, ethnological and anthropological observa- tions which are profusely reported in his letters published by E.H. Giglioli, and in his book Nuova Guinea, Selebes e Molucche. Full of interest are the observations on the characteristics and origin of the Papuans, and on the life of birds, particularly those on the bower-birds of paradise, Amblyornis inornata, and its 'capanne e giardini', which are carefully and at the same time poeti- cally described by him in a paper full of interesting scientific and philosophical considerations, pervaded with a deep-rooted love and admiration of Nature. Learning that a Dutch expedition to New Guinea was being prepared, with the vessel 'Soeraba- ja', with the aim of performing a bathymetric survey, Beccari received permission to accompany this. It lasted from November 11, 1875 till January 29, 1876, visiting Dorei, the Bay of Wanda- men, the islands of Roon and Krudu, the Bay of Humboldt, the island Arimosa, Awek (Japen I.), Dorei, Waigeu, Misool, the Bay of MacCluer, the Bay of Gouns, the island of Geser (off Southeast Ceram), and Ambon. From there he returned on the mail steamer to Ternate. He stayed there about a month to arrange and pack his latest collections and to ship them to Italy. This third expedition to New Guinea had not yielded results as far as Beccari' s botanical and zoological in- (13) Flora Malesiana terests were concerned, but it had allowed him to assemble many ethnological and anthropological notes, and to improve his topographic surveys. On March 12, 1876, Beccari sailed from Ternate to Java, on the first stage of his homeward voyage. He arrived, unexpectedly, at Florence on June 18, 1876, after about four years of bold and glorious exploration. On his return Beccari was received with great honour. On July 14, 1876, the Municipality of Florence bestowed the freedom of the city on him; some scientific societies, such as the Zoological Society in London, and the Italian Anthropological Society, elected him an honorary member. Other scientific associations, such as the Italian Geographical Society and the Tuscan Society of Horticulture as well as the Faculty of Science of the Royal Institute of Advanced Studies of Flor- ence awarded him a gold medal. But he was not affected by these honours and devoted his time to his collections and to his friends in Florence and Genoa. However, the glamour of exploration and the call of the wild were too strongly in his nature and after a year Beccari made one further long voyage to the Malesian islands. He and Captain Count Enrico A. d'Albertis, a cousin of his former companion, set out from Genoa on October 14, 1877, on a trip, properly intended more for pleasure than for science, to Australia, en route travelling through India from Bombay, Lahore, Delhi, Benares, Lucknow, to Calcutta, touching Singapore and Kuching (December 1877), meeting in Australia Ferdinand VON Mueller, and proceeding to Tasmania and New Zealand. On the return voyage he parted from d'Albertis in Singapore and proceeded to Jakarta and then to Bogor where he spent two weeks, preparing a collecting trip in Central West Sumatra. Sailing from Jakarta on 28 May 1 878, he arrived in early June via Padang and Padang Pandjang at Mt Singalang, a primary-forest-clad, long-extinct volcano of nearly 2900 m height. Here he had a hut built, as before in Sarawak and Ternate, which he made his headquarters. The hut, named by him 'Bellavista', was placed above the limit of cultivation and on the lower fringe of the primi- tive forest, at an altitude of about 1700 m. He remained there from June 12 to early August, mak- ing rich collections on the flanks and on the top of the volcano. Later he set his base in a house in the village of Ajer Mantcior at the base of Mt Singalang till September 20. After a short stay in Padang to arrange his collections, he undertook a journey on October 4 in the provinces and on October 22 he sailed from Padang to Bangkok where he arrived on November 10. During his travels in West Sumatra (see map of his itineraries in Beccari 1930) he assembled large botanical and zoological collections; the largest were made on Mt Singalang, the harvest of plants running to a thousand numbers in all. Amongst them were the famous Rafflesia arnoldii and the then un- known, largest, erect aroid in the world, Amorphophallus titanum, a really colossal herbaceous plant, the tuber being up to 53 cm in diameter, the inflorescence more than 1 .5 m high, the lamina of leaf covering a surface of about 15 m in circumference and the petiole attaining about 29 cm diameter at the base. From Bangkok he began his homeward journey to Italy, arriving in Florence on December 28, 1878, thus concluding the last of his fascinating explorations in Malesia. Reviewing the results of his botanical activities in the six years exploration in the field through almost the whole of Malesia, it is evident that Beccari's exploration in Sarawak was the most fruitful and thorough, with the huge number of over 4000 collections in two and a half years. The great virtue and value of his collections can only be properly estimated if one takes into considera- tion that Beccari collected species rather than specimens, and that he seldom collected a species twice. Each species was studied, dissected and annotated on the spot and mostly carried flowers and fruit. The Sumatran collection again was rather large, about 1000 numbers in five months travel, especially when one considers that Beccari's interests were wide; in Sumatra he also dedi- cated time to the study of agriculture, forest products and fruit trees, as he had done in Borneo. (14) Dedication The amount of these collections clearly contrasts with less than 1000 numbers in the three years spent in the Moluccas, Celebes and New Guinea (see Appendix 4), although really there are more, as most collections from the Kei and Aru islands and Kendari in Southeast Celebes are unnum- bered and unlisted. As the flora of at least New Guinea is not less rich in proportion to that of Borneo or Sumatra, the reasons for this contrast can only be explained by Beccari's activity in the field. Firstly, we must consider that he was a thorough collector and disliked gathering occa- sional or incomplete specimens; he preferred always to stop some days to collect systematically in a place which he considered botanically interesting, rather than to gather here and there along his path, en route, when moving from one place to another in his long expeditions. In a word, he preferred to collect intensively rather than extensively. In Borneo and in Sumatra where he had huts for drying, labelling and drawing his specimens ('Vallombrosa' and 'Bellavista' respectively) his collections were more numerous than in New Guinea, Celebes and Moluccas when he frequent- ly moved his collecting base, sometimes being forced to do so because threatened by native head-hunters, or by pirates. Finally, we must also bear in mind that the second expedition to New Guinea was made thanks to the financial support of the Province and Municipality of Genoa, se- cured on the warmest recommendation of Giacomo Doria, his friend and zoologist of the expedi- tion to Borneo. Beccari knew that he longed to enlarge the zoological collections of the Civic Museum of Natural History which he had founded in 1867, and, in his profound honesty, he felt bound to assemble large zoological collections for the Genoa Museum and to put zoology before botany. Indeed, the zoological collections were very rich in quality and in quantity, while the bo- tanical collections were not particularly numerous. Another reason for this contrast can also be found in Beccari's health which was worse in eastern Malesia than in Borneo and Sumatra. In 1878 Beccari was still only 35 years old, but had accumulated an unrivalled, immense amount of material, great scientific-botanical experience and knowledge of the Malesian flora, in fact had proved himself the greatest explorer of his time. He would prove himself also to be the greatest botanist in the elaboration of his results, surpassing Blume in the width of the field he covered, including plant-geography, ecology and biology. Having concluded his explorations in Malesia, Beccari devoted the rest of his life to the study of his collections and of palms, except for an unhappy experience as Director of the Botanical Col- lections and Garden of the Royal Museum of Physics and Natural History of Florence in 1878- 1879, and a short journey to Ethiopia soon after. The vicissitudes of Beccari's life as Director of the Botanical Collections and Garden cannot be understood without knowledge of some of the events in the history of the Florence Museum and the sale of Beccari's collections. The Royal Imperial Museum of Physics and Natural History in Florence was founded in 1775 by Pietro Leopoldo di Lorena, Grand Duke of Tuscany, and was installed in a building in Via Romana, not far from the Palazzo Pitti, the palace of the Grand Duke, at the base of the great and famous Giardino di Boboli. Part of this was soon designated as the Botanical Garden of the Museum. Thanks to the great interest of the Grand Duke in Natural Science, the scientific collections were greatly increased in the years thereafter and the Museum was subdivided into various sections (Cabinets) and in 1789 an astronomical observatory was also installed there. From then on all the Museum complex was usually named 'La Specola' by the Flo- rentines. The botanical section consisted of the so-called Botanical Collections (herbaria, carpo- logical collections, vegetable products, wax models and fossil plants) and the Botanical Garden. After various events dominated by the historical course of Tuscany in the first half of the I9th century, Filippo Parlatore was appointed in 1842 director of the Collections and Garden. He greatly contributed to the growth of the herbaria and to the organization of the department. In 1854 Philip Barker Webb died in Paris and bequeathed his invaluable herbarium and library to (15) Flora Malesiana the Florence Museum, together with an annual income of 6945.58 lira (derived from the sale of a palace in Paris) and known as Webb's Legacy, for their maintenance and increase. In 1859 the Government of Tuscany established the Istituto di Studi Superiori Pratici e di Perfezionamento in Firenze, which had its centre near the Monastery of San Marco; and the Museum of Physics and Natural History with its collections and the botanical garden, although situated on the op- posite side of the river Arno, became part of that Institute as the seat of the Faculty of Science. However, the Museum continued to have a director of its own, and in 1868 Parlatore was ap- pointed to that office. Very unfortunately, the Institute of Advanced Studies, which only became the University of Florence in 1923, had no Rector responsible for the scientific and didactic activi- ty, but only an Administrative Board which determined the course of events in the Museum in the following years without an adequate knowledge of the problems and needs of scientific re- search. In 1860, after various vicissitudes, the Giardino dei Semplici, founded in 1545 and one of the most ancient in the world, became state property, and, in 1869, together with the adjoining buildings (originally the stables of the Grand Duke) was assigned to the Institute of the Advanced Studies being situated near the centre of the Institute at San Marco. In 1872 the Italian Govern- ment, the Province and the Commune of Florence signed a convention for the enlargement of the Institute of Advanced Studies, and the Board of the Institute decided to move some of the Cabi- nets of the Museum to the centre of the Institute. The latter also foresaw the removal of the Botan- ical Collections and Garden of the Museum from La Specola to the Giardino dei Semplici and pertinent buildings at San Marco, in order to have the botanical department nearer to the Institute, and to maintain only one garden, the Giardino dei Semplici, by far more famous than that of the Museum. Thus, during 1877 and 1880 the Cabinets of Chemistry, Physics, Geology and Mineralo- gy were shifted from the Museum to San Marco. Only in 1879 did the Commune of Florence ac- tually hand over the Giardino dei Semplici to the Institute of Advanced Studies. The latter decided that the Botanical Collections and Garden ought soon to be moved to San Marco. However, they were conveyed there only several years later owing to the opposition of several botanists, among whom Beccari, and other personalities, who were against the removal of the botanical collec- tions, as they considered it, for several reasons, to be a great error. FiLiPPO Parlatore died on September 9, 1 877. He was the last Director of the Museum of Phys- ics and Natural History in Florence as the Institute had decided that the Dean of the Faculty was to hold the directorship of the Museum. However, the office of Director of the Botanical Collec- tions and Garden was vacant, and according to general opinion Beccari was the best qualified and most worthy successor to the work of Parlatore, who had so greatly enlarged the herbarium and library and raised them to the level of the greatest museums in the world. But Beccari's ap- pointment was strongly opposed by the Dean of the Science Faculty and the Board of the Institute of Advanced Studies, particularly because Beccari was firmly convinced that the Director of the Botanical Collections and Garden should have no hand in teaching. On October 14, 1877, hardly more than a month after Parlatore's death, Beccari undertook his travels with E. d'Albertis, sketched above, and at that time no resolution had been taken. Only on March 26, 1878, while Beccari was journeying in Australia, was he, in spite of the opposition, appointed Director of the Botanical Collections and Garden of the Royal Museum of Physics and Natural History of Florence, with the duty of supervising the practical phytographic research of the students. Becca- ri returned to Florence from the exploration of Sumatra on December 28, 1878, and soon took up the office of Director. In the preceding years Beccari had organized, worked and studied on his own in the field of botany, and his brilliant achievements were naturally a one-man show. In the field he had to make his own decisions, and learned to do so immediately. He had no rivals and had always very subor- (16) Dedication dinate personnel whom he could command. That was very different from the situation in which he was now placed as a Director, with a graded staff accustomed to some privileges acquired dur- ing the long directorship of Parlatore. On the other hand Beccari was indefatigable and tidy in his work and he required everybody to be active and precise in carrying out his duty. He wanted to infuse new life into the operations of the botanical collections and garden, introducing methods that differed from the traditional ones. Very soon his reforms became unpopular among several of the staff, who felt that he lacked respect for the memory of his predecessor; this state of affairs acerbated the hostility towards him of the Institute of Advanced Studies. But such hostility was not something to scare him. It is clear, of course, that Beccari, with his enormous drive and ambition, proved by his unique exploration and study of the tropical floras, thinking big, botanically and otherwise, wanted to raise the Florentine centre into an institute which could compete with the leading world herbaria. For this there was excellent opportunity as the Florence botanical collections had already acquired the enormous and (still) most important herbarium of Philip Barker Webb, bristling with types of old collections, with funds attached for its maintenance, and further the important herbaria of Cesalpino, Micheli, Targioni, and Parlatore. To these could now be added his own numer- ous collections from the East, with its enormous mass of duplicates for the further enrichment of the Florence herbarium by exchange. There was, therefore, every reason and opportunity to fulfill his ambition. When Beccari took on the directorship, he was aware that the botanical collections were under threat of removal from the Museum to the buildings adjoining the Giardino dei Semplici near the centre of the Institute of Advanced Studies. At first he was rather in favour of the project since he thought that the Institute had large funds for the construction of new buildings and that they would be better and more suitable for the collections than those of the Museum. But when he dis- covered that the funds of the Institute were scarce, and the buildings were the old stables of the Grand Duke of Tuscany, very humid, unsuitable for both the herbarium and library, and far worse than those of the Museum, he became a most obstinate and relentless opponent to the removal of the collections. His hostility was the primary cause of a wide gulf between him and the Institute Board. On the other hand Beccari began to understand that his ambition of raising the Florence Botanical Collections to the level of the other great European herbaria would be dif- ficult to realize. But this controversy was not the reason for his resignation as Director, at least not the main one. Indeed Beccari's resignation was for a different reason. On one of his visits to Java during his travels he had been requested by the Dutch East Indies Government to sell his collections to the Bogor (Buitenzorg) Herbarium for the cash payment of a considerable amount of money and his appointment as botanical explorer in the Garden, or a life annuity of 5000 lira. The offer was alluring, but Beccari wished his collections to remain in Italy and to spend the rest of his life in Florence, attending to their study. However, his own estate was seriously compromised owing to the expenses for his long expeditions, and before his depar- ture for the last journey to Australia, New Zealand and Sumatra (1877-1878) Beccari undertook negotiations with the Florence Institute of Advanced Studies for the sale of his Malesian collec- tions. The Institute asked Marquis G. Doria, the Director of the Civic Museum of Natural History for an appraisal of Beccari's botanical collections, which were estimated at 75,065 lira. On the basis of this valuation and considering the offer of the Dutch East Indies Government, the Insti- tute of Advanced Studies offered to buy all his botanical collections from Malesia against an annu- ity of 5000 lira for the rest of his life. Beccari accepted this offer but on the condition that the collections were entrusted to the Museum of Physics and Natural History of Florence and that he was entitled to have them at his disposal during his lifetime; in exchange he would assume (17) Flora Malesiana responsibility for their study and conservation. During his journey, in May 1878, when he was in Batavia prior to his expedition to West Sumatra, Beccari was informed that the Board of the Institute of Advanced Studies had accepted his conditions and had officially approved the pur- chase of his collections. Consequently he refused the offer of the Dutch East Indies Govern- ment. When Beccari came back to Florence and took up the directorship of the Botanical Collections and Garden of the Museum, he ought to have accepted the contract but learnt that the Board of the Institute of Advanced Studies intended to use the money from the Webb legacy for the pay- ment of his life annuity. Beccari was greatly disappointed on hearing this decision, since he had thought that his life annuity would be paid by different Institute funds: he disliked the idea that the Webb collections were to be deprived of nearly all their endowment until his own death. Fur- thermore, he knew that the Webb legacy was the main source of income of the botanical depart- ment of the Museum and without it, his ambition to make the Florence herbarium one of the greatest in the world and a leading centre of tropical botany could not be accomplished. He under- stood too that under these conditions, the sale of his collections was incompatible with the duty of his office as Director. Indeed, he got a personal benefit from the sale, while as Director of the Collections and Garden it was his duty to avoid that these were deprived of a large amount of money necessary for their maintenance and increase during his lifetime. Thus he made every effort to persuade the Board of the Institute of Advanced Studies to use different funds to purchase his collections, but without result. He did not underestimate the hostility that the Institute had shown him since the beginning, and particularly recently, and when he was invited to sign the contract, he clearly understood that he was regarded by them as a troublemaker, and that the decision to pay the price of his collections with the Webb legacy was merely an expedient devised by the In- stitute in order to compel him either to lose his prestige as Director or to resign from his office. Indeed, if Beccari had sold his collections and kept his office he would have lost his prestige as a man and as Director, having put his personal interest before his duty. But the Board of the In- stitute knew that Beccari was a man of honour and that it would achieve its aim: his resigna- tion. Then, as a last attempt, he tried to find some way in which, without going back on his word, he could withdraw from the compromise of the sale, but without success. Thus, on July 26, 1879, Beccari resigned as Director of the Botanical Collections and Garden of the Florence Museum. Only later, on October 31, 1879, as a private citizen, did he sign the contract for the sale of his own collections. The fight had been lost, but his honour was saved! Soon after the end of this unhappy and painful experience, on November 16, 1879, Beccari left for Ethiopia to stay with his old friend and benefactor. Marquis Giacomo Doria, as members of an Italian expedition to the Assab Bay on the Red Sea which he had already visited in 1870. They also spent some days collecting in Aden and returned to Florence on February 26, 1880. After his return from Ethiopia, Beccari resumed the study of his collections, which were lo- cated in a few small rooms on the top floor of the Museum of Natural History. In those modest and secluded rooms, alone, like a hermit, without any assistant or help, but together with his rich collections, he worked until his death. There, he wrote his famous works on Malesian plants and on palm taxonomy, and made the splendid drawings and photographs which adorn his publications. The first months there, however, were unfortunately rather hard for him. After his resignation, in November 1880 Teodoro Caruel was appointed Director of the Botanical Collections and Garden of the Museum. He was soon instructed by the Institute of Advanced Studies to study the advisability of removing the Botanic Collections and Garden from the Museum to the Giardino (18) Dedication dei Semplici and adjoining buildings, and eventually to prepare a project for such a removal which had already been decided, but not realized, before Parlatore's death. Beccari was aware that Caruel between 1866 and 1871 had been Director of the Giardino dei Semplici and that he was in favour of the removal and was preparing the pertinent project. Beccari had already expressed his resolute opposition to the removal, chiefly because the buildings near the Giardino dei Semplici were unsuitable for the collections and library as they were very humid, smaller and worse than those of the Museum, but also because library, herbaria and garden were well settled in the Museum and there was no need to remove them, and in doing so waste a large amount of money which could have been used for their maintenance and growth. Besides, he was strongly convinced that the great botanical collections and library at the Museum, as a centre of taxonomic research, had to be kept distinct from the centre of teaching and research on anatomy and physiology at the Giardino dei SempHci. Thus, in 1880 and 1881 Beccari tried everything, with letters and ar- ticles in various Italian newspapers, to convince the Faculty of Science, the Board of the Institute of Advanced Studies, and public opinion that the removal of the botanical collections and garden would be a great and irreparable mistake. He carried out a referendum against the removal of the herbaria and library among botanists in Italy and abroad. This was spread far and wide and was discussed in many Italian and foreign publications. Numerous botanists from every part of the world, and among them the most eminent taxonomists of the time, declared themselves against the removal. In 1881 Beccari published a paper in which the reasons for the protest against the removal of the botanical department from the Museum and the result of the pertinent referendum were given. In the same year Caruel published his study for carrying into effect the project of the removal of the Botanical Collections and Garden. Despite the opinions of many and outstanding botanists expressed in the referendum, the Board of the Institute of Advanced Studies decided on the removal of the Botanical Collections, Library and Garden from the Museum of Natural History, in Via Romana, to the Giardino dei SempHci and adjoining buildings near San Marco, on the op- posite side of the river Arno. However, Beccari did not give in, and he continued to pubhsh other articles and papers against the removal until 1903. Even if his campaign did not gain its aim, it greatly contributed to further resolutions of the Institute of Advanced Studies which decided to enlarge and improve the buildings annexed to the Giardino dei Semplici and later to reserve for Botany the part of them originally intended for the Zoology department, which remained at the Museum, at La Specola, where it still is today. These deliberations greatly delayed the removal of the Botanical Collections, though the living plants of the Garden of the Museum were all moved to the Giardino dei Semplici during 1883. Caruel continued to give his botanical lectures at the Museum until his retirement, in 1 896, but the following year, his successor, O. Mattirolo, under- took his teaching in the building near the Giardino dei Semplici. When in 1900 he moved to Turin, P. Baccarini succeeded him as Director. At that time, the Library and the Botanical Collections were still located in the Museum. In 1901, when the removal was close at hand, Beccari was requested to inform the Institute of Advanced Studies in which rooms of the new botanical building at the Giardino dei Semplici he wished to have his Malesian collections deposited. He disdainfully replied that he wished his collections to remain at the Museum in Via Romana in agreement with the contract of their sale, adding that, if the Institute had decided to move them to the new buildings, he would not follow them and would give up their study. His Malesian collections remained in the same rooms at the Museum until the end of his life, but, in 1905, in spite of further protest and particularly after controversy with P. Baccarini, the removal of the Library and the Botanical Collections (includ- ing all the herbaria) from the Museum of Natural History to the new Botanical Institute near the (19) Flora Malesiana ancient Giardino dei Semplici, was brought to a conclusion. However, let us resume the course of Beccari's life after his sad experience as Director of the Botanical Collections and Garden of the Museum in 1878-1879, and his hard fight against their removal started in 1881. These regrettable events marked a turning-point in Beccari's career. He realized that he had lost the chance of making the Florence Herbarium one of the leading centres for research in plant taxonomy, and decided to retire to private life, devoting himself entirely to taxonomic research, chiefly to elaborate his own Malesian collections for which he had gathered a wealth of field observations and drawings. On January 23, 1882, Beccari married Nella Goretti de Flaminj, from a noble family of Ca- sentino, in the high valley of the Arno. They had four sons: Nello, Ding, Baccio and Renzo. The eldest, his devoted son Nello, became a professor of Comparative Anatomy at Florence Uni- versity and took great pains in editing some posthumous papers by the father, among them the book Nuova Guinea, Selebes e Molucche based on the original diaries of his father's explorations in eastern Malesia from 1871 to 1876. He also encouraged U. Martelli and R.E.G. Picm Ser- MOLLi to revise and edit some works on palm taxonomy which had been left unfinished by his father. The years immediately following Beccari's marriage, entirely devoted to his family and to study, were peaceful and fruitful. He set up his home in a villa inherited from his father, the me- diaeval Castello del Bisarno, near Ripoli in the immediate vicinity of Florence, and he lived there until his death. According to information obtained from his nephew and from letters to his friends, we know that he also had another house in the city of Florence where he and his family spent the week-days, particularly in winter. We also know that he used, as in the years before his marriage, to spend several weeks, particularly in the summer holidays and during the grape-har- vest at Radda in Chianti, on the old country estate of his mother's family. He was very fond of country life and, following the tradition of the old families of the region, he was particularly inter- ested in wine-making, in which he attained great experience. He was one of the first producers, together with Baron B. Ricasoli, of that typical wine, well-known in Italy and abroad as 'Chianti, Gallo nero'. Nevertheless, during his holidays he did not stop his research, even if he did not work so actively as in the Florence Museum, where his collections were housed. However, after a few years his life was troubled by another sad event. In 1877 he had under- taken the publication of a great work, Malesia, mainly with the intention of embodying in it the results of the studies dealing with his own collections from the Malesian Archipelago; in addition, other papers or abstracts of works published elsewhere on plants of that region were also to be included. The first two volumes were printed in Genoa and Beccari was greatly helped in editing them by his faithful friend R. Gestro, the Director of the Civic Museum of Natural History of Genoa, particularly during his last journey to the East. However, Beccari undoubtedly corrected the proofs of all the instalments of Malesia and also those of fascicle 3 of volume 1 , issued when he was in West Sumatra. This is proved by a letter to Gestro from Buitenzorg, now Bogor (dated May 2, 1878), which accompanied the corrected proofs of that fascicle. Beccari published the first two fascicles of volume 1 at his own expense, but later Malesia became a publication of the Florence Institute of Advanced Studies. However, despite its great interest, the sale of this work was obviously limited, the text being written entirely in Italian. Consequently the Institute of Ad- vanced Studies decided that it was not worth continuing its publication and suddenly, in 1887, stopped all contributions to it while fascicle 3 of volume 3 was not yet complete. Actually, in the cover of fascicle 3 we find a note which informs us that the publication oi Malesia is ended and explains the reasons for it. However, Beccari wished to publish at least the text pertinent to the drawings of the account on Bombacaceae already issued in fascicle 3, but as far as possible, also (20) Dedication other papers and drawings ready for the press. Hence, he was compelled to beg in Italy and abroad for funds necessary to publish the last two fascicles of volume 3 of Malesia. Fortunately, the Min- ister of Education, Paolo Boselli, and the Bentham Trust in England, where he was highly es- teemed, allowed him the necessary financial support for bringing volume 3 of Malesia to a close. The last issue appeared in March 1890. Great was Beccari's disappointment at the unhappy conclusion of the publication of Malesia, not only because the resolution of the Institute of Advanced Studies represented a slight to him and to his work, but also because he had lost a safe and certain means of publication for the results of the study of his collections. He was so much upset by this event that he even thought of visiting Malesia again. Actually, in the letter to GESTRO(April 4, 1890) which accompanied the last fascicle {'ultimo definitivo') of Malesia he asked his friend for information on the departures from Genoa to Batavia and about the liners. Anyhow, the end of Malesia was another turning-point in his life: it marked the beginning of a long period of inactivity, after which he never resumed the study of his own Malesian collections. Having concluded the studies already undertaken, he published no scientific papers from 1893 to 1902, except some articles and letters protesting against the removal of the Botanical Collec- tions of the Florence Museum, described above, and the temporary closing of the herbaria and library in connection with this removal. In these years he was on the point of giving up his botanical activity entirely and none of his colleagues and friends, not even his devoted pupil Ugo- LiNO Martelli, were able to induce him to resume his research. However, another person was to have the credit for reviving in him the enthusiasm for the country where he had spent the most fruitful period of his youth. After his explorations in Sarawak BECCARihad kept alive his friendship with the Rajah and the Ranee of Sarawak . They liked to spend part of the year in the surroundings of Genoa. It is difficult to say whether it was by chance or with the definite intention of helping BECCARito overcome his scientific inactivity, but in the early days of May 1 897 the Ranee visited Florence and met Beccari. A woman of great culture and sensibility, deeply fond of her kingdom of Sarawak, Lady Marga- ret Brooke succeeded in convincing Beccari to write a book on his fascinating explorations in Borneo. He soon began his work; the Ranee kindly assisted him in the preparation of the book, particularly in providing him with the illustrations. Several of them, in fact, are a selection from many fine photographs taken by the Ranee herself in Sarawak; these were assembled in a great album, still kept in the Florence Botanical Museum, which she presented to Beccari on June 10, 1897. The preparation of the book, Nelleforeste di Borneo, led Beccari to recover a certain interest for scientific work and it was not difficult for Prof. Oreste Mattirolo, the Director of the Botanical Department of the Museum, to complete the Ranee's enterprise and convince him to resume botanical research after the pubHcation of his book. However, Beccari had already real- ized that the removal of the library and the herbaria from the Museum to the Giardino dei Semplici was close at hand, and that without them the elaboration of his own Malesian collections would be rather difficult. Consequently, he did not resume their study, interrupted in 1890 after the pub- lication of Malesia was stopped, and decided to devote his entire botanical activity to a single group. This decision was neither easy to take nor satisfying for him, but represented the best solu- tion for carrying out his research without a big library and a great herbarium at his disposal at any moment of the day. The selection of the group was easy, as the Palms had intrigued him since his first visit to Malesia, and he had already done some research on them. On the other hand he knew that this group offered him a taxonomically nearly unexplored, big field. Thus, Beccari started again on his studies with renewed enthusiasm, spending the whole day at the Museum in his (21) Flora Malesiana small rooms where he had concentrated everything necessary for his work, including the great camera he had designed himself specially for making the marvellous photographs which are repro- duced in the plates of his truly monumental works on Palms. In the years that followed he had no difficulty in publishing his writings thanks to the great esteem he enjoyed abroad and the friendship of several Italian botanists. Actually, the publication of his chief work, Asiatic palms, was made possible by Sir George King, the Director of the Botanic Garden of Calcutta, and other important papers were published by his close friend, U. Martelli, in the periodical Webbia, which the latter had founded in honour of Philip Barker Webb. Some interesting works appeared also in L'Agricoltura Coloniale, a journal edited by the Istituto Agricolo Coloniale, the founda- tion of which was promoted by Beccari and other Italian personalities. Several other papers on palm specimens from all over the world entrusted by their collectors to him for determination, were issued in various periodicals and books. Beccari led this last period of his life completely secluded from Italian academic life and nearly forgotten by most Italian botanists, but he was always overwhelmed by the sympathy and esteem of foreign botanists. In this period, perhaps more than before, he enjoyed the affection of his old and devoted friends and particularly of Ugolino Martelli, his only pupil, a very keen botanist himself, well known for his basic works on the great family of Pandanaceae, whose study he had undertaken on Beccari's advice. This period, entirely devoted to his family and the palm studies, was serene and creative. In the last years of his active and eventful life he assembled the materials for a book on his explorations of eastern Malesia. He had already sorted out a final copy of his diaries and he had also begun to prepare the illustrations for his book, but unexpectedly death prevented him from accomplish- ing this last performance. He died peacefully in the evening of the 25th of October 1920, in Florence, at the age of 77. Odoardo Beccari was a great explorer but at the same time a very clever, many-sided, careful collector. Indeed he did incredible work in the field. In his long and lonely explorations in Malesia, Beccari was attracted by all aspects of nature and human life and assembled an enormous wealth of botanical, zoological, ethnological and anthropological collections. Not only the number of the specimens makes his collections really invaluable, but also the fact that these are often accom- panied by notes and descriptions and sometimes by splendid and detailed drawings made in the field. The botanical collections from Malesia are kept in the Herbarium and Museum of the Florence University. They amount to more than 21,000 sheets, about 2400 flasks of material in alcohol, about 800 carpological specimens and more than 200 wood samples with the pertinent voucher specimens (see further information in van Steenis's Thesaurus Beccarianus). Many collection numbers are represented in Beccari's herbarium by more than one sheet. To these specimens we must add many duplicates which were distributed to the most important herbaria, among which those of the British Museum, Kew, Paris, Geneva, Leningrad, Berlin, Leiden, Vienna, Munich, Stockholm, and Bogor. The zoological collections consist of several thousand specimens belonging to a very high num- ber of species, many of which were described as new. Those in the higher groups are represented by skins, but sometimes also by skeletons, skulls and even heads or other parts of the body pre- served in alcohol. They are kept in the Civic Museum of Natural History of Genoa, which bears the name of its founder and Maecenas, Giacomo Doria, the companion of Beccari on the expedi- tions to Borneo and the Red Sea. Beccari paid attention to special groups and assembled several (22) Dedication collections of particular importance such as that of the great ape, the orang-utan (48 specimens including a foetus), the set of birds of paradise, one of the best of its kind, and the collections of fishes, spiders, coleoptera, and ants. These collections have supplied a very rich harvest of study for many specialists. Indeed 205 works devoted to the study of Beccari's zoological collections had been published by 1920 in the Annali del Museo Civico di Storia Naturale of Genoa alone (see Gestro's biography of Beccari). BECCARialso collected rich and interesting ethnological collections of great value and beauty. They are kept in the Anthropological and Ethnological Museum of Florence and some of them are exhibited in a hall of that Museum. A set of ethnological collections was sent to the Italian Geographical Society in Rome. Some of the idols, weapons, implements, ornaments, clothing, etc. were described and illustrated in Beccari's books Nelleforeste di Borneo and Nuova Guinea, Selebes e Molucche for the purpose of trying to establish, with the aid of the anthropological fea- tures, the origin of some of the peoples of Malesia. The value of these collections was greatly in- creased by the detailed information on the customs of those peoples which he described in his books and in the letters to his friends published by E.H. Giglioli, G. Cora and the Italian Geo- graphical Society. The anthropological collections are kept in the Anthropological and Ethnological Museum of Florence. They consist of skulls and a few skeletons of natives of Malesia. The most important is undoubtedly the set of some 200 skulls assembled in Korido in the island of Schouten in north- western New Guinea. These collections were the base of the first craniological investigations on Papuans, by P. Mantegazza and E. Regalia. Beccari's activity in the field also extended to the geographical features of the districts he visited. During the preparation of his expeditions, he had trained himself in geodetics and topo- graphy and he had also invented a new instrument (Nuovo orizzonte artificiale. Rivista Marittima 6, 1873, 198-200, f. 1 -5) for topographic surveys. During his travels he also made a topographic survey of several territories, particularly of the northwestern parts of New Guinea. These surveys later allowed G. Cora to prepare the maps he published in Cosmos (see Appendix 1, C: Maps). He also discovered a great river. War Samson, in northwestern New Guinea, near Sorong. Even though Beccari had succeeded in assembling such enormous and invaluable collections, his fame is mainly due to his scientific work condensed into more than 150 publications, some of which consist of monographs of basic importance and those on palms especially still remain stan- dard works even today. Beccari's versatile mind allowed him to devote his attention to problems in different branches of natural science, but he carried out his activity chiefly in botany. In the first period, during which he made the great expeditions to Malesia, his botanical activity was essentially applied to the study of a part of his collections; the result was published mainly in the Nuovo Giornale Botanico Italia- no which he founded in 1869. Even if some of these papers were written here and there during his explorations, his scientific output in this period was necessarily small. With the end of his explorations in Malesia, the second period of Beccari's activity begins. Probably he was unsatisfied when he limited himself to the descriptions of new genera and species, and in this period he spread his field of research to the monographic or semi-monographic treat- ment of those families or genera from Malesia which had most attracted him. In this period Bec- cari also undertook, at his own expense, the publication of a collection of botanical writings, Malesia, in order to facilitate the printing of his papers dealing with his Malesian plants and the reproduction of his splendid drawings. For this purpose he selected the quarto size. In Malesia he published some of his most outstanding works. The end of its publication in 1890 marks the end of this second period of his scientific activity and also the end of the study of his own collec- tions from Malesia. (23) Flora Malesiana After about a decade of complete scientific inactivity, due to the sad vicissitudes of his academic life and the end of Malesia, Beccari published Nelleforeste di Borneo in 1902. This year coincides with the resumption of his scientific studies and marks the beginning of the third period of his botanical research in which he devoted himself entirely to the study of the family of palms, becom- ing the best specialist who ever existed. In the following pages we want to go into more detail about the subjects contained in the mas- sive oeuvre of the Maestro. Confronted with the multitude of his activities we hope to weave this into a readable account, with a distinct feeling that our ability for writing falls short of the way in which Beccari could express himself, as testified by his 'Wanderings', which is still a thrilling guide for exploration in the tropics. Let us start with this work on Sarawak, the core of his main work in Malesia. He worked under favourable conditions, having ample equipment and time at his disposal, and the support of the Tuan-muda, Charles Brooke. His big hut, called 'Vallombrosa' on Gunong Mattang, a hill west of Kuching, was used as a study centre. From there he made excursions and gradually familiarized himself with the very rich flora of the primary forest. He focussed attention on the big trees {Dipte- rocarpaceae, Bombacaceae, and others) as well as on the evasive tiny creatures of the saprophytic Triuridaceae and Burmanniaceae, the parasitic plants, the lianas and so forth, making beautiful and exemplary complete specimens in a skilled, professional way. This was, especially with un- wieldy plants such as palms and pandans, gingers and aroids, quite an effort, as every field botanist must be aware. Perseverance and patience fed by infinite interest must have induced him to take particular care with these groups. A special characteristic is that he knew his plants; hardly ever did he make two collections of the same species. As a scientific collector he was never equalled, and only approached by E.J.H. Corner and L.J. Brass. What a contrast with most other collectors who, even today, stick to the disgusting grab-as-grab-can way of collecting on hurried cross-country trips, causing heavy dupli- cation and absence of vital field notes. How Beccari managed all this at the age of 22, with only a few months training in tropical form knowledge at Kew, can only be understood if we imagine him as an extraordinarily gifted person with an intense interest in botany; botany in the widest sense, because he was not satisfied only with the taxonomy of flowering plants, but collected for example also wood samples and crypto- gams of all major phyla. His horizon widened to collecting minerals and all sorts of animals, ob- servations on vegetation types, on edible and horticultural plants, and the way of life of his com- panions, the Dayak people; in short, he possessed the integrated interest of a born all-round natu- ralist, whose scope went far beyond the mere plant collecting and description in which he excelled. Beccari assembled a great wealth of data on the geographical features of Borneo, on the matter of useful and horticultural plants, on fibres, rattan, bamboos, resins, camphor, getah percha yielding trees, and medicinal plants. He was aware of the primitive domestication of species of Durio (durian), Eugeissona (a palm), Artocarpus (breadfruit), bananas and species of Nephelium, which he learned from observation of Dayak life. Beccari briefly reported on these subjects in a short summary of his journey in Sarawak to the Italian Geographical Society (1868), and later he incorporated the complete data in some appendices to his book Nelle foreste di Borneo. Prob- ably, when gliding easily in canoes on the rivers or proceeding with difficulty on the mysterious peat of the swamp forests during his long journey in the interior of Sarawak, he ventured on ideas about the origin of coal in Borneo - in which he was correct. When hunting for orang-utan in Batang-Lupar or excavating in the limestone caves of Sarawak he made his first guesses about the origin of man, which he developed in Nelle foreste di Borneo. He surely must have had a very busy life in his 'Vallombrosa', because he could never have con- (24) Dedication ceived the 'Wanderings' thirty years later without having copious notes of his field observations and full diaries. He must also have started in the field to compose botanical papers and prelim- inary revisions; when still in Sarawak he published some small papers in Italy. BECCARiwas much enchanted with Sarawak, the flora, the country, and the Dayak people, and he even conceived a project that the Italian government should purchase it as a crown colony. When Beccari returned from Borneo to Florence he founded \.\\q Nuovo Giornale Botanico Ita- liano, in which he published freely some papers on spectacular Bornean plants; but before leaving for the expedition to New Guinea he handed over the journal to T. Caruel, since it would be diffi- cult both to edit it and to explore in Malesia. When, loaded with further materials, manuscripts, field notes and drawings, he returned from New Guinea to Florence, he realized the difficulty of publishing the botanical results of his explorations in the Nuovo Giornale Botanico Italiano. Thus he decided to undertake a collection of writings he named Malesia, in which he concentrated near- ly all his works of that period. The first two instalments of it were published during his one year interval, spent in Florence, between the last expedition to New Guinea and the journey to Austra- lia, New Zealand and Sumatra, and the third one appeared during the last mentioned journey. He undoubtedly must have worked very hard during that period, but he was able to manage it thanks to his efficient organization en route and the help of his friend R. Gestro in Genoa, where Malesia was printed. Beccari published in Malesia several works which are important for the subject in itself, but also various original considerations about some particular subjects, such as evolutionary pro- cesses, dispersal of seeds, geographical distribution, etc. are dealt with in them. First should be mentioned the extensive work on the ant plants devoted to the study of the symbiosis between plants and ants, which occupies the entire second volume. In it, Beccari gives us his interesting views on the evolution and the common origin of plants and animals, discussing concepts still valid and topical today. Likewise of great interest are his considerations on the origin of the insectivor- ous plants and the distribution of plants in the Malesian archipelago, dealing especially with the Nepenthaceae. Other important works are the monograph on the genus Phoenix and the account of the palm genus Pritchardia in which he resumes his considerations on the dispersal of seeds and fruits and the origin of the flora of the Pacific islands. The three volumes of Malesia contain also a number of monographic or semi-monographic revisions of families and genera from Malesia, e.g., Icacinaceae, Menispermaceae, Nepenthes, Bombacaceae, Triuridaceae, Burmanniaceae, etc., and also a first survey of the palms of New Guinea. We can conclude that Malesia was intended to embody the botany of Malesia as a repository. It must have aroused great interest in the botanical world, containing novelties of fascinating plants with marvellous drawings made by Beccari himself, a 'must' for every botanical institute. The use of the Italian language, even for monographic contributions from non-Italian collabora- tors, e.g., on Araceae by Engler, was certainly an obstacle to a wide sale, and the edition was subvented from Italian sources and the third and last volume could only appear thanks to the aid of the Bentham Trustees. Whether Beccari ever intended or hoped to achieve an ultimate incorporation of all the mono- graphs of Malesian plant families, that is, an attempt towards a true Flora Malesiana, remains uncertain. The fact that he had certainly intentionally explored all areas of Malesia, except the Philippines, and that the first instalments of Malesia contained several monographic treatises, may support this idea. What is certain is that he fully realized that he could never dream of accom- plishing all this himself. Accordingly he freely entrusted many groups to colleagues in Italy and to his many correspondents abroad as appears from the bibliography in Appendix 4. Through the distributed duplicates Beccari's material went to various herbaria and later came (25) Flora Malesiana into the hands of specialists, but much of his original collection, embodied in the original Herba- rium Beccarianum, has not been examined by specialists. This original material is separately shelv- ed in Florence, 400 bundles in 33 cupboards; to it belong a card system with field data and a cover with drawings; there is a rather large number of unicates or collections of which no duplicates were distributed. In 1951 van Steenis very roughly sampled a number of families and found that Bec- CARi often had indicated and annotated genera in sched. as new, which were later based on other material, e.g., Koordersiodendron Engl. (1898), Clavistylus J.J.S. (1910), Neosepicaea Diels (1922), Octamyrtus Diels (1922), Haplolobus H.J. Lam (1931), Kjellbergiodendron Burret (1936), Macadamia hillebrandii Steen. (1952), Eriandra v. Royen«& Steen. (1952), Whiteoden- dron Steen. (1952). It is a pity that in the past five decades too little initiative has been taken by the curators of the Florence Herbarium to attract and induce specialists to study the original Beccari collections in Florence. We suppose that it is due to the understaffing of this great Herbarium. Especially the Herbarium Beccarianum is not a reliquiae, not a closed chapter, but truly a thesaurus, still con- taining unknown botanical treasures. As this is not always realized by specialists, we urge them to borrow material of their speciality from this century-old, inexhaustible source. We sincerely hope that a revival of interest in the Her- barium Beccarianum is welcome to the future curators of the Florence Herbarium. Its possession brings with it the scientific obligation of using it, not just in honour of the Maestro, but mainly for the benefit of scientific botany and as a contribution to the fame of the Florence centre. After publication of Nelle foreste di Borneo in 1902, Beccari decided to concentrate, for the rest of his life, on the study of one large family on which he possessed more field knowledge than anyone, before or since, namely the palms. His first contribution to their knowledge dates from 1871, with a provisional account of those of Borneo. Old love never dies! In 1877 he had ac- counted for the palms of New Guinea, in 1885 for those cultivated in the Botanic Gardens at Bo- gor, but in about 1890 he spread his wings towards those beyond Malesia, the Indian empire, Indo- China, and later to Madagascar, Africa, the Pacific islands, etc. Everybody entrusted him with palm material and from this emanated a massive knowledge of Asiatic palms, embodied partly in the Records of the Botanical Survey of India, in Hooker's Flora of India, partly later in Web- bia, founded by his old pupil and friend U. Martelli, but largely in the sumptuous volumes of the Annals of the Royal Botanic Garden, Calcutta, which also included those of Malesia. For the large folio plates of these massive plants Beccari designed a special large camera with suitable accessories in order to achieve excellent illustrations'. He devoted his attention mainly to the tax- onomy of palms, but he also studied the cultivated species in some works which appeared in L'Agricoltura Coloniale, edited by that Institute once named Istituto Agricolo Coloniale, now Istituto Agronomico per I'Oltremare of which he had solicited the foundation in 1903. Of this big work a large number of unpublished manuscripts appeared in print after his death, through the untiring devotion of his pupil, friend, and colleague Martelli, who must be given a tribute of honour for his singularly unselfish efforts. The last of the manuscripts on palms which Beccari left unfinished, that of the sub famWy A recoideae, was completed and published by Pichi Sermol- Liin 1955. We should also refer here to Moore's important and competent evaluation of Becca- Ri's massive contribution to the knowledge of the fascinating palm family. (1) The large camera and other microphotographic cameras designed by Beccari are described by Luigi Pampaloni, Apparecchio fotografico universale per iaboratorio biologico ideato dal Dottor Beccari. Rend. Congr. Bot. Naz. Palermo (1903) 164-168, cum fig., andGli apparecchi microfotografici del Dott. O. Becca- ri. Bull. Soc. Fotogr. Ital. 14 (1902) 129-145, fig. 1-7. (26) Dedication Dealing with his botanical activity we cannot silently pass over his descriptive work. Also in this Beccari excelled and showed that he had a remarkable insight into affinities. It appears that his new genera were always placed in the proper plant family and, moreover, that hardly ever were new species proposed by him reduced later, stamping him as a most accurate taxonomist. Indeed he was a taxonomist, but Beccari showed his sharp intelligence in other branches of botany often including his considerations in taxonomical papers. Describing the details of Gnetum led him to considerations about the ancestry of the flowering plants from the Gymnosperms. The plant geo- graphy of the palms led him to hypothetical ideas about former landbridges and sunken conti- nents. His gatherings in Sumatra led him to consider the affinities of its flora with those of South- east Asia and Java, concluding that the flora of volcanic ranges must be much younger than that of the more ancient and more stable Sunda lands. Other observations deal with the dissemination by earthworms; the double dispersal, anemochorous and zoochorous, of the plants of the periodic swamp forest provided with floating fruits and succulent seeds; pollination by pigeons; the various colours of flowers of the forest plants, and some others on physiology and ecology. Beccari also left traces of his versatile genius in various writings (papers and letters to his friends) which lie outside botany but must be mentioned briefly to understand how great he was as a naturalist. Particularly interesting are the letters to E.H. GiOLiOLiand G. Cora in which he disclosed his views on the origin of the peoples of Malesia, in particular of the Papua-Mafor which he regarded as derived from a crossing of aboriginals, perhaps descended from Negritos and Hin- du peoples. Other interesting observations are those on the connections between mosquitos and malaria which he was one of the first to suppose, those on the agent of the bee pest which he sus- pected to be due to a protozoon later discovered in America, those on the connections between flies and cholera and numerous other observations particularly on the customs of animals. It was in Borneo that Beccari perceived the true value of evolution and was primarily fascinat- ed by the importance of adaptation to environmental conditions. But only later did his views on the processes of evolution take shape in his mind. It is a fact that the prolonged stays of gifted naturalists in the tropical wilderness, when their minds are set free from daily minutiae and domes- tics and solely occupied with the bewildering structural wealth of tropical plants and animals, allows their minds to open to new, big ideas and syntheses, generating philosophical thought. For this, one has only to think of von Humboldt, Junghuhn, Wallace, Darwin, and Corner. To this, Beccari, with his eager mind and astute power of observation, was no exception. Beccari was used to going back from the facts to the causes, and his views on the evolutionary processes, which arose from the observations he made in nature, were consolidating in his mind in the course of time. Thus we find his views sketched in some papers and later resumed in others, whenever he had the chance to develop them on the basis of particular new observations. He did not supply us, or perhaps he did not want to supply us, with a synthesis of his views on evolution in an ad hoc publication, perhaps out of humility, since he disliked giving the impression that he was able to explain the laws of evolution, or perhaps out of honesty, because he perceived that his ideas had made their way into his mind by intuition and reasoning, without adequate investiga- tion. Beccari was undoubtedly an evolutionist, but he was one in a very original manner. The first foundation of his theory of 'plasmation ' was explained in his paper (1 876) on the huts and gardens of Amblyornis inornata, the small bower-bird of paradise which builds a pretty hut with, in front, a lovely garden of soft moss on which it scatters flowers in shining colours changing them when they wither. This theory was resumed in the introduction to his work on ant plants (1884) and was later developed in his paper (1 889) on the flowering oi Amorphophallus titanum, the gigantic Ara- cea, and was summarized in his book, Nelle foreste di Borneo. (27) Flora Malesiana According to this theory the evolutionary processes of living beings took place, beginning with the most ancient geological times, fundamentally in two different epochs: a first epoch of plasma- tion, and a second epoch of conservative heredity, displaying in the course of time the two funda- mental rules of variability and fixity. In the first epoch, the plasmative strength, unhindered by heredity, may have given free play to the variability and to the adaptation stimulated by environ- mental conditions. In that epoch, the organism may have been liable to yield to the stimulus of external factors and more subject to modelling itself to them, the modifications occurring with the greatest of ease and even quite suddenly without the offspring necessarily being like the par- ents. This epoch may have been a period of youth for living beings in which each individual was allowed to modify itself in conformity with its needs, or rather even according to its wishes, its vanities, its whims. This epoch of plasmation, with a maximum of variability and a minimum of fixity, may have been followed by an epoch of conservative heredity, characterized by that strength which aims at the conservation of the acquired characters and owing to which the individuals belonging to a spe- cies transmit to their descendants the characteristics they have inherited from their ancestors. Thus the plasmative epoch may have been replaced by an epoch with a minimum of variabihty and a maximum of fixity. The strength of the conservative heredity becoming stronger in the course of time may consequently have weakened the faculty to vary, perhaps even to cancel it entirely, thus impressing the stamp of fixity on all living beings. He recognized the great influence of environmental factors on the plasmation of living beings and pointed out several cases of correlation between the morphology of some apparatuses and the environmental factors. For instance, he was the first to correlate the life form of flood-resistant plants with the environment and was struck by their similarity in leaf-shape: his 'stenophyllous plants', now called 'rheophytes'. A still more important correlation amply studied by him was that of the symbiosis between plants and ants, the 'piante ospitatrici', or formicarian plants, to which he devoted a very large and detailed account (1884). However, he clearly and repeatedly recog- nized that plasmation could also be stimulated by an interior strength, by the wish of having some particular functions facilitated, such as defence, pollination, seed dispersal, etc. But BECCARialso supposed that plasmation was even influenced, particularly in the animal kingdom, by a psychic push stimulated by the beauty of the environment, as could have happened in the birds of paradise desirous of imitating with their feathers the glowing colours of the aurora and dusk of the tropics, which they greet from the highest trees of the forest with very Hvely dances. Though recognizing that the extant being cannot, as a rule, undergo modifications because of the environment, Beccari admitted however that even today some changes can take place in the species on account of a cross between individuals of different species or of the sudden appearance of hereditary modifications of various, even if unknown, origin. He admitted that these could be induced by new poisonous substances or by new enzymes arising in the substratum; thus he fore- saw the existence of mutations and mutagenic substances. Beccari was a man of great intelligence, versatility and intuition, who united an exceptional personality and liberality with uncommon integrity and strength of mind. He was an indefatigable worker, who devoted hours and hours to his research, without a moment of rest. But he did not work out of a wish to be praised; prizes and honours did not interest him. He loved his research studies since he was zealous of the beauty and perfection of nature and only happy when he could entirely devote himself to investigating its manifold and marvellous aspects. For the same reason he liked to draw plants and animals and very few excelled him in scientific drawing. Beccari had an austere and inflexible character, but he was neither obstinate nor autocratic. His temperament was based on a keen sense of duty. Also in private life, although deeply fond of (28) Dedication his wife, sons and friends, every action was characterized by a clear austerity, and he was greatly beloved but at the same time he always inspired a certain awe. But this austerity concealed a great goodness of heart. He had also a great sense of justice and was a very courageous and stalwart man. These gifts united with the wish to inspire respect and love rather than fear, were greatly esteemed by native people and on his dangerous expeditions he never suffered injury from anybody. Undoubtedly he must have encountered serious danger, but he faced them with resolution, and he recounts them with indifference and without boasting. Beccari is described as a proud, almost misanthropic spirit and perhaps he was not an easy character and we understand why some regarded him as a troublesome person. From his youth, moulded in his long and solitary explorations in Malesia, he was set apart, destined to travel and to work alone, and he was not afraid of solitude; on the contrary, in his ripe age he found refuge in it, the better to devote himself to his studies and serve his single purpose and sole end: the science of nature. R.E.G. PicHi Sermolli & C. G.G.J, van Steenis (29) Flora Malesiana Appendix 1 — Bibliography of Odoardo Beccari A — Books and papers (excluding reviews of bool =more than (in size, number, etc.) < =less than (size, number, etc.) X 2/5 = 2/5 of natural size X montana = means that the epithet montana is that of a hvbrid (47) ARALIACEAE— I (W. R. Philipson, Christchurch)i Trees, shrubs, lianas, woody epiphytes or (extra-Mai.) more rarely herbs. Branches usually stout with leaves clustered at their ends; armed or unarmed; glabrous or with a tomentum of stellate or simple hairs ; buds either covered by the stipular sheaths of leaves or by cataphylls. Leaves spiral or rarely opposite or in whorls; petiole usually clasping the stem; stipules either distinct or united into a ligule or absent (in Osmoxylon the petiole bears ± elaborate crests around its base) ; lamina digitately compound or pinnate, sometimes to the second or third degree, or simple, when either entire or pinnately or palmately lobed, margin entire or dentate. Inflorescence terminal or more rarely lateral ; either simple or compound racemes or spikes, or more commonly of umbels or capitula, either solitary or arranged in compound umbels or panicles; bracts usually small and caducous; pedicel either articulated with the flower or continuous with it. Flowers hermaphrodite or hetero- sexual, sometimes dioecious; actinomorphic. Calyx lobes small, or reduced to a rim, or rarely absent. Petals 3 to numerous, often 5, sometimes fused into a calyptra, or forming a tube with spreading lobes {Osmoxylon), valvate or imbricate in bud, usually with a broad base but rarely narrowed below. Stamens usually as many as the petals and alternating with them, or twice as many, or indefinite; filaments inserted at the edge of the disk ; anthers dorsifixed, introrse, pollen sacs 4 or rarely 8. Ovary inferior, half inferior, or very rarely {extra-MaX.) superior, 1- to many-celled, the top of the ovary usually a fleshy disk; styles and stigmas as many as the cells, either connate or wholly or partially free. Ovules solitary, pendulous, anatropous, with the raphe ventral. Fruit baccate or drupaceous, exocarp usually fleshy, endocarp forming cartilaginous or membranaceous pyrenes around the seeds. Seeds one per pyrene, with a small embryo within smooth or ruminate endosperm. Distribution. About 50 genera with a roughly estimated 11 50 species, ranging mainly in the warmer parts of both hemispheres (especially in montane zones), a small number in or extending to cool-temperate regions. With the exception of SE. Asia, the family and its centres of distribu- tion are largely found within the land masses derived from ancient Gondwanaland. In Malesia 17 genera with a total (excluding Schefflera) of 117 species in 16 genera. (The largest genus, Schefflera, with an estimated 250 species for the region, is omitted from this account.) Three genera are endemic to Malesia (or nearly so). One of these, Anakasia (related to Polyscias) is of very local distribution in West New Guinea; the two others, Harmsiopanax (Java, Lesser Sunda Is., Celebes, to New Guinea) and Aralidium (Malay Peninsula, Sumatra and Borneo, with an extension to Thailand) are more widely distributed. Five other genera have their main centres of distribution within Malesia. Two of these extend further eastwards: Mackinlaya ranges from the Philippines and Celebes through New Guinea to the Solomon Is. and NE. Australia; Osmoxylon occurs from Botel Tobago (Taiwan) and the Marianas to Borneo, the Philippines, Celebes, the Moluccas, New Guinea to the Solomon Is. and the New Hebrides. A third, Trevesia, is confined to western Malesia (Lesser Sunda Is., Java, Borneo, Sumatra, and Malaya), with an extension into the Asian mainland. Wider ranges are recorded for Gastonia (widespread in Malesia to the Solomon Is.) with an additional range in the Seychelles, Mascarenes, and Madagascar (but not East Africa as previously reported) and Arthrophyllum (all over Malesia with extensions to Thailand, Laos, the Nicobar Islands, and New Caledonia. A significant element in the Malesian representation of the family consists of six genera which occur mainly in SE.-E. Asia, two of which further extending to the Americas, viz Aralia (in (1) The genus Schefflera is omitted and will be treated separately by Dr D. G. Frodin (University of Papua New Guinea). I enjoyed his assistance in drawing the general chapters. (1) 2 Flora Malesiana [ser. I, vol. 9^ America south to Mexico) throughout Malesia, and Dendropanax (tropical America) in West Malesia (Sumatra, Malay Peninsula, Borneo). The remaining four genera range in Malesia as follows : Macropanax and Brassaiopsis in Malay Peninsula, Sumatra and Java, Acanthopanax in Malay Peninsula, Sumatra and the Philippines, and Pentapanax only in East Java. Only one genus, Delarbrea, has its main centre of distribution to the east of Malesia (Melanesia, mostly New Caledonia) and Queensland, and extends through New Guinea to the Moluccas and Lesser Sunda Is. as far west as Timor. Two other genera have wider distributions. Schefflera, including several segregate genera, is pantropical although with but few representatives (13) on the African mainland. However, the several sections of Schefflera as redefined by Frodin (1970, 1975) often show distinctive regional distributions paralleling those of several of the other genera. The type section ranges from the New Hebrides to Samoa and New Zealand, including Fiji and New Caledonia. Polyscias is widely dispersed in the Old World tropics from the African mainland eastwards to the Society Is. and Australia, but with only a few species in SE. Asia and western Malesia. As in Scheffiem, the diverse series of species comprising the genus show distinctive regional distribution patterns, with the type series being mainly Melanesian and East Malesian (in West Malesia it is only cultivated or adventive). As said above several Malesian genera extend into, or have their main centres in, mainland Asia, but only Polyscias (in Sri Lanka) and Schefflera are shared with Africa and the same genera (with Gastonia) with Madagascar. Only Polyscias and Gastonia occur in the Mascarene Islands and Gastonia and Schefflera in the Seychelles. Two other genera are restricted to mainland Africa. Hedera of temperate Eurasia extends to the Canary Islands. The Americas have two (or three) distinctive endemic genera with recognizable affinities, as well as disjunct groups of the Austral- asian Pseudopanax, the Asiatic Pentapanax and Dendropanax and (in North America) Oplopanax, Aralia and Panax, all very closely related to those in East Asia; there are also distinctive sections of Schefflera in the neotropics which are nearer those in Africa than in Asia. Many distinctive endemic genera (or parts of the larger genera) occur in Oceania, the New Zealand region and Australia, some of them taxonomically very isolated. The ratio of species : genus is here estimated as 23 : 1 , but if the very large genus Schefflera is not accounted for this reduces to 1 3 : 1 . Some 30 genera have five or fewer species ; in view of the considerable insular endemism at generic or infrageneric level, weak intercontinental links, and great distinctiveness of many genera (even though small), the family is surely of great antiquity, although much of the available palaeobotanical evidence requires re-evaluation (c/. Dilcher & DoLPH, 1970). Many fossils previously referred to Oreopanax must now be placed in Platanaceae (Doyle, pers. comm.). Dendropanax has reliably been reported from Tertiary deposits in both Europe and North America where it is now absent (Dilcher & Dolph, I.e.). References: Dilcher & Dolph, Amer. J. Bot. 57 (1970) 153-160; Frodin, The complex of Cephaloschefflera in Schefflera (Araliaceae), Thesis, Cambridge, U.K. (1970); J. Arn. Arb. 56 (1975) 427-448. Ecology. Malesian Araliaceae are usually small trees, shrubs, or lianas, with a number being sometimes or always epiphytic (especially in Schefflera), and where terrestrial usually in the under- growth or lower stories of rain-forest, seldom reaching over 20 m. A remarkable exception is provided by the two species of Gastonia; of these G. spectabilis (Harms) Philipson of Papuasia can attain the great height of 40 m with a stem of 1 .75 m ; it is the largest araliad known and is of very striking appearance. Fig. 27. Araliads are almost always found scattered in forest and other vegetation, at least in Malesia. However, a notable exception is provided by the tree Schefflera rugosa (Bl.) Harms in Java; Lam (1924) recorded its gregarious occurrence on the volcanic cone of Mt Slamet in Central Java where it is co-dominant with the pyrophilous Albizia lophantha Bth. in elfin forest between 2500- 3050 m, above which it gives way to the open, rocky, treeless slopes below the summit (3428 m). It is also gregarious on Mt Tjeremai in West Java, where Lam (1925) noted that above some 2650 m a low forest dominated by this species replaces the high forest of Dacrycarpus imbricatus; this low forest extends to c. 3000 m. On the Gedeh-Pangrango complex above Puntjak Pass in West Java Schefflera rugosa is frequent in forest borders. On Mt Ulu Kali in Malaya, east of Kuala Lumpur, S. nervosa (King) Vig. is common in young regrowth along the main road 1979] Araliaceae— I (Philipson) 3 below the Genting Highlands hotel/casino complex from 1300-1600 m. This pioneering tendency is shared by a number of other species in the genus, especially in montane parts of Papuasia. Many of these species are terrestrial as pioneers or in secondary formations, but epiphytic in closed forest (and then often much more scattered). Gregarious occurrence, usually in pioneering situations or in forest borders, has also been observed in the various species of Harmsiopanax ; Steup ( 1 938) observed H. aculeatus (Bl.) Warb. ex BoERL. as a characteristic pioneer in grass thickets on hills in SW. Celebes. //. harmsii K. ScH. behaves similarly around Wau, Bulolo, and Sogeri (Rouna) in Papua New Guinea, especially in narrow intermontane valleys, while H. ingens Philipson can be locally abundant in open situa- tions in the highlands of New Guinea. Gastonia spectabilis (Harms) Philipson can be locally frequent as a pioneer in hill areas, e.g. around Bulolo and on the Madang-Ramu Divide; while Polyscias elegans (C. Moore & F.v.M.) Harms is frequently seen in stable monsoon scrub and forest borders in the Port Moresby region and P. ledermannii Harms can be an exceedingly com- mon regrowth tree in cut-over montane forest subject to frequent cloudiness and rain. In the subalpine zone of New Guinea Brass (1941) found two species of Schefflera conspicuous in the stunted forest oi Xanthomyrtus-Vaccinium-Papuacedms-Phyllocladus between 3200-3800 m on Mt Wilhelmina. From Lake Habbema upwards, the characteristic overtopping tree is S. alti- gena Frodin {sect. Brassaia), with its large foliage contrasting sharply with the surrounding microphyllous vegetation; at higher altitudes it gives way to S. pagiophylla Harms (5. falcata Philipson), a species of uncertain affinities. Schefflera chimbuensis Frodin and S. straminea Frodin are likewise often seen in forest borders on Mt Wilhelm and Mt Giluwe respectively in Papua New Guinea. In contrast to the large and conspicuous species of Schefflera, many others are more or less shade-loving epiphytes or vines of lower storeys within the forest, although they may occur in surmier situations where clouding is frequent (5. singularis B. C. Stone on Mt Ulu Kali). One group of species in New Guinea is almost limited to perhumid moist or wet rain-forests and usually are small epiphytic shrubs or even herbs (5. gemma Frodin). A few species are con- spicuous rosette-trees of the forest understorey (5. stahliana (Warb.) Frodin). Climate. In Malesia Araliaceae for the most part shun regions subject to a seasonal climate; all species are evergreen. A few exceptions include Schefflera thaumasiantha Harms from open savanna in the hill zone in SE. New Guinea and S. actinophylla (Endl.) Harms as a gallery tree in seasonal parts of southern New Guinea, but especially Harmsiopanax aculeatus whose range is for the greater part subject to an annual drought period ; in addition to SW. Celebes, it is also frequent on old lava-streams on Mt Idjen in East Java together with Wightia, Casuarinajunghuhn- iana, Dodonaea, and Wendlandia, and is one of the few araliads in the Lesser Sunda Islands. With respect to altitude, most Araliaceae occur in the lowland, hill and montane zone below 2300 m. The only genera of which all Malesian representatives occur in the hills and mountains above 1(XX) m are Pentapanax and Dendropanax, but even these do not exceed 3000 m. Only certain species of Schefflera and Harmsiopanax continue upwards much higher, especially in New Guinea where the highest known record belongs to S. pagiophylla Harms on the Carstensz complex, where F. J, Wissel found it in 1936 at 39(X) m. Flower biology. Little has been recorded about the floral biology of the family in Malesia, but Beccari's account (1878) of the 'false fruits' of Osmoxylon (including Boerlagiodendron) serving to attract doves which are assumed to eff"ect pollination has become a classic description and example of ornithophily. Heterosexual flowers, usually involving combinations of perfect and male flowers, occur frequently but understanding will require intensive study in the living state. Though flowers are not generally show^ and often veritably inconspicuous, the disk produces abundant nectar. They also may spread a rather disagreeable scent, somewhat spermatic, that in Schefflera rugosa resembling that of Ligustrum. For this species Docters van Leeuwen (1933) observed on Mt Gedeh, West Java, only rare visits (notably by Diptera), but assumed that cross- pollination will be the rule. Flowers (at least those oiFatsia) will also be visited by Hymenoptera. In Schefflera stahliana (Warb.) Frodin, the thick fleshy flowers, numerous stamens, and position of the inflorescence below the rosette of leaves all point to bat pollination. Dispersal in the family takes place generally by fruit-fall; but as fruits are baccate or (more 4 Flora Malesiana [ser. I, vol. 9^ usually) drupaceous, they will also be eaten by birds (for the most part) and bats, as recorded by Ridley (1930) for Aralia, Hedera, and Schefflera. The black fruits of Schefflera sect. Brassaia in New Guinea and Australia are especially popular with birds. The hooked mericarps of Harmsio- panax are exceptional. Seed germination is most likely after the seed having passed the gut of a bird or after mastica- tion of the fruit by a bat. References: Beccari, Malesia 1 (1878) 193-198; Brass, J. Am. Arb. 22 (1941) 271-342, esp. 318, 320, 323, 327; Docters van Leeuwen, Verh. Kon. Ak. Wet. A'dam sect. 2, 31 (1933) 195; H. J. Lam, Trop. Natuur 13 (1924) 20; ibid. 14 (1925) 6; Ridley, Disp. (1930); Steup, Trop. Natuur 27 (1938) 142. Morphology. Most erect Araliaceae are sparingly branched or even unbranched; their limbs are upright and the often massive, easily broken twigs generally bear rosettes of big, long- stalked leaves which leave large scars when they fall. Few form a true crown {Arthrophyllum, Gastonia, some species of Schefflera and Polyscias), while the others often look like elongated and grotesque shrubs with a candelabrum-like framework. Branches in most Araliaceae are entirely orthotropic; generally speaking, the plants would fit into the so-called Holttum, Corner, Tomlin- son, Chamberlain and Leeuwenberg models of Halle c.s. (1970, 1978), or their intermediates. Aralia scandens (Merr.) Ha is a true climber. Epiphytic species only occur in the genus Schefflera; it is not yet recorded that any of them may appear to turn into a hemi-epiphytic habit. Hedera (not native in Malesia) is almost the only genus with differentiated shoots and marked vegetative dimorphism. A preliminary account of shoot-morphology in the family has been given by Philipson (1978), but much further field work is required in this area. Vegetative buds may be either proleptic or sylleptic; in the resting phase such buds may be covered by the clasping bases of foliage leaves {Osmoxylon, Schefflera) or (more rarely) specialized cataphylls {Acanthopanax). In both types of bud the primordia and young leaves may be covered by exudations of resinous slime. There is so far little recorded evidence of the changes in leaf shape and configuration during the somatic phase of the life cycle known in many araliads in othei parts of the world; however, recent observations in New Guinea made by Frodin suggest that heteroblastism does occur, although its manifestation is not constant for a given species. Distinctive juvenile and intermediate foliage has been found in Schefflera eriocephala Harms and (to a lesser extent) in S. stolleana Harms. Other examples are seen in Brassaiopsis, Trevesia and Schefflera subg. Agalma (5. aroma- tica (Bl.) Harms; S. nervosa (King) Vig.); juvenile leaves of some of these are preserved in Herbarium Bogoriense. In Harmsiopanax, the configuration of the leaves changes abruptly just below the inflorescence, a phenomenon paralleled in some other genera although less dramati- cally. In Mackinlaya celebica (Harms) Philipson and M. schlechteri (Harms) Philipson, leaf polymorphism is very marked with the result that in the past several 'paper species' have been described on too limited a range of material; in this revision many reductions have been made. References: Halle & Oldeman, Essai sur I'architecture et la dynamique de croissance des arbres tropicaux, Paris (1970); Halle, Oldeman & Tomlinson, Tropical trees and forests: an architectural analysis, Berlin (1978); Philipson in Tomhnson & Zimmermann (eds.). Tropical trees as living systems (1978) 269-284. Anatomy. General accounts of the vegetative anatomy of the ivy family are given by Gussow (1900), ViGuiER (1906, 1909) and Metcalfe & Chalk (1950). Secretory canals are characteristic of the stems and leaves, but are absent from Aralidium. A comparison of the wood anatomy of Araliaceae and Cornaceae is made by Philipson (1967), and an extensive account of vegetative anatomy in the context of woody Umbellales was provided by Rodriguez (1957, 1971). The xylem of the former family is characterized by fibres with small, simple pits, and the presence of scalariform and reticulate perforation plates in the vessel elements (as opposed to simple perforations) is thought to be a less advanced feature. Recent special reports bearing on taxonomy include : on stomatal development (Inamdar c.s., 1969); on sievetube plastids (Behnke, 1972), and on epidermal papillae (Bui, 1974). Floral anatomy is discussed by Baumann-Bodenheim (1955), Philipson (1967, 1970) and especially by Eyde «&. Tseng (1971). Embryological characteristics of the family have been re- 1979] Araliaceae— I (Philipson) 5 viewed by Davis (1966) as well as by Rao (1972). The single pendulous anatropous ovule has the funiculus and ovular vascular bundle axial (Philipson, 1970). The embryo is small in a mass of endosperm (Martin, 1946; Grushvitzky, 1967). References: Baumann-Bodenheim, Bull. Soc. Bot. Suisse 65 (1955) 481-510; Behnke, Bot. Rev. 38 (1972) 155-197; Bui Ngoc-Sanh, Bull. Mus. Hist. Nat. Paris III, Bot. 18 (1974) 85-91 (whole no 271); Davis, Systematic embryology of the angiosperms. New York (1966); Eyde & Tseng, J. Am. Arb. 52 (1971) 205-239; Grushvitzky, Proc. Int. Symp. Physiol. Ecol. & Bio- chem. of Germination (ed. H. Borriss) (1967); GiJssow, Beitrage zur vergleichende Anatomie der Araliaceae, Thesis, Breslau (Wroclaw) (1900) 67 pp., illus. ; Inamdar, Gopal & Chohan, Ann. Bot. n.s. 33 (1969) 67-73; Martin, Amer. Midi. Nat. 36 (1946) 513-660; Metcalfe &. Chalk Anatomy of the dicotyledons II, Oxford (1950); Philipson, New Zeal. J. Bot. 5 (1967) 134-165 in Robson, Cutler & Gregory (eds.), New research in plant anatomy, London (1970) 87-100 Rao, Phytomorphology 22 (1972) 75-87; Rodriguez, Univ. Calif. Publ. Bot. 29 (1957) 145-318 in Heywood (ed.). The biology and chemistry of the Umbelliferae, London (1971) 63-91 Viguier, Ann. Sci. Nat. Bot. IX, 4 (1906) 1-209; ibid. IX, 9 (1909) 305^05. Palynology. Palynological studies of Malesian Araliaceae include: on Gastonia (Tseng, 1971); on Tupidanthus and Plerandra (= Schefflera) (Tseng, 1973); on Osmoxylon (as Boer- lagiodendron) (Tseng, 1974) and on Schefflera (Tseng & Shoup, 1978). A detailed consideration of the relations of Araliaceous pollen to those in other orders is given by Hideaux & Ferguson (1976), and of the affinities of Klotzschia {Umbelliferae I Hydrocotyloideae) to Araliaceae by Shoup & Tseng (1977). To date, much useful new evidence has been made available, but better correla- tion with other classes of attributes is required. References: Hideaux & Ferguson in Ferguson & Muller (eds.). The evolutionary significance of the exine, London (1976); Shoup & Tseng, Amer. J. Bot. 64 (1977) 461-463; Tseng, Amer. J. Bot. 58 (1971) 505-516; Grana 13 (1973) 51-56; Amer. J. Bot. 61 (1974) 717-721; Tseng & Shoup, Amer. J. Bot. 65 (1978) 384-394. Chromosome numbers. Lists of chromosome numbers for members of Araliaceae are given by Darlington & Wylie (1955), Sharma & Chatterji (1964) and Bolkovskikh c.s. (1969). The family shows considerable constancy of the basic number of x = 12, although x = 1 1 has been recorded for one non-Malesian group of Schefflera. The implications of chromosome data on the wider relationships of the Araliaceae are discussed by Moore (1971). References: Bolkovskikh c.s.. Chromosome numbers of flowering plants, Leningrad (1969); Darlington & Wylie, Chromosome atlas of flowering plants, ed. 2, London (1955); Moore in Heywood (ed.). The biology and chemistry of the Umbelliferae, London (1971) 233-255; Sharma &. Chatterji, Cytologia 29 (1964) 1-12. Phytochemistry. Information on the chemistry of the Araliaceae should be sought in Hegnauer (1964, 1978) where references to original sources are given. The family is characterized by the occurrence of essential oils and resins in canals and by the presence of polyacetylenic compounds (especially falcarinone-type), triterpenic sapogenins of the oleanene-, ursene- and dammarene-types, seed oils with petroselinic acid, and by the absence of true tannins. The chemistry of the family fully confirms its close relationship with the Umbelliferae and also more distantly with the Pittosporaceae and the Compositae (Hegnauer, 1969, 1971; Bohlmann, 1971). The rareness of flavones and the predominance of flavonols in Araliaceae suggests closer relationship with two of the three subfamilies of the Umbelliferae, flavones having not yet been found in Hydrocotyloideae and Saniculoideae (Harborne, 1971). The absence of iridoid sub- stances and true tannins and the presence of polyenes, petroselinic acid and isoprenylated coumarins in the Umbellales contrasts with the Cornales (Hegnauer, 1969; Jensen c.s., 1975), and this led to a suggestion that the Umbellales and Cornales had to be separated (Bate-Smith c.s., 1975), an argument with increasing support from other lines of inquiry. However, insufficient evidence appears to be yet available for the detection of possible lines of relationship within the Araliaceae. References: Bate-Smith c.s. Biochem. Syst. Ecol. 3 (1975) 79-89 ; Bohlmann in Heywood (ed.), The biology and chemistry of the Umbelliferae (1971) 279-291; Harborne, I.e. 293-314; Hegnauer, Chemotaxonomie der Pflanzen 3 (1964); in Harborne & Swain (eds.). Perspectives in phytochemistry (1969) 121-138; in Heywood (ed.), The biology and chemistry of the Umbelli- 6 Flora Malesiana [ser. I, vol. 9^ ferae (1971) 267-277; in Cauwet-Marc & Carbonnier (eds.), Les Ombelliferes. Contributions pluridisciplinaires a la systematique. Perpignan (1978) 335-363; Jensen, Nielsen & Dahlgren, Bot. Notis. 128 (1975) 148-180. Taxonomy. The Araliaceae are on all grounds closely connected with the Umbelliferae, a very large but mainly temperate and tropical-montane group comprising mostly aromatic herbs with a restricted floral scheme. It seems likely that the ancestors of the ivy family were the woody tropical stock from which the herbaceous Umbelliferae evolved under the rigours of cooler climates (Corner, 1 940) with the arborescent Heteromorpha of upland Africa and Myrrhidendron in Central and South America, both in the subfamily Apioideae, perhaps representing relics of the transition on account of the presence of a number of attributes primitive for the family (Rodriguez, 1957); on the other hand, the umbellifers may have originated as megaherbs on tropical mountains (Philipson, 1978). A few Araliaceous genera exhibit some features charac- teristic of Umbelliferae, such as Harmsiopanax, Mackinlaya, and especially Myodocarpus (from New Caledonia) and Stilbocarpa (from southern New Zealand and associated 'subantarctic' islands). This led Harms (1898) to show three different lines leading from Araliaceae to Umbelliferae, suggesting that considerable overlap between the families existed; more recently Rodriguez (1971) again called attention to this phenomenon and Thorne (1968, 1973) has gone further by merging Umbelliferae into Araliaceae (a step also advocated by Hallier/. in 1905) and assuming the three subfamilies of the former to have arisen separately, perhaps in different parts of the world at different times, from proto-Araliaceous ancestors. However both he and Rodriguez have concluded in agreement with Corner (I.e.) and Baumann-Bodenheim (1946) that the Araliaceae sensu stricto, "because of their greater evolutionary breadth and their retention of many more primitive features, would seem closest to the proto-araliad stock" (Thorne, 1973). Nevertheless, the concept of Araliaceae as a separate natural family appears to serve a useful purpose and has been retained for this Flora. An isolated, doubtfully included genus is the West Malesian Aralidium; it is the only genus lacking resin ducts but it would be equally anomalous in the Corndceae to which it has also been referred. It shows some resemblance with the New Zealand genus Griselinia, usually relegated to the Cornaceae, be it as a marginal member (Philipson, 1967). Subdivision. Since the first significant family monograph by Seemann (1868), several systems have been proposed which, taken together, are notable for their lack of consistency. This results from a lack of agreement on the relative importance of the comparatively minor structural and gross anatomical features of reproductive parts traditionally used and by conflicting claims on the relative 'antiquity' of polymery versus pentamery. Until recently, there has been for a priori reasons (c/. Eyde, 1975) little recognition of the potential value of vegetative features and their acceptance as valid evidence for a system ; and information from wood anatomy, floral histology, palynology, karyology, phytochemistry, and other areas is only beginning to be utilized. While woodiness is generally accepted as a primitive feature in Araliaceae, the impact of the work of Corner and others on tree structure and growth rhythms (summarized in Halle, Oldeman & ToMLiNSON, 1978; see also Borchert, 1969; Hladik, 1970; Philipson, 1978) has still to be fully assimilated. Much more work is also required on inflorescences, although Frodin (1970), Philipson (1970b) and others have made a beginning. The monothetic interpretation of most attribute states usual in systems of the family was first challenged by Baumann-Bodenheim {I.e.) who considered that phyletic changes could have taken place in parallel ; and Eyde & Tseng (1969) showed that at least some supposedly unidirectional sequences were reversible. This has tended to reduce the supposed importance of many of the traditional attributes, with a conse- quent reduction in the number of genera. Sufficient evidence is not yet available, however, for the construction of a more balanced, polythetically based system of the family, and the long-standing subdivision proposed by Harms (1898) into three tribes, Schefflereae, Aralieae, and Mackinlayeae, based monothetically on the structure and aestivation of the perianth, is retained. In spite of its now recognized imperfections, it is more valid than the systems of Viguier (1906) and Hutchinson (1967) and remains the most widely accepted. Some steps towards the formulation of a new system on polythetic principles have been taken by Eyde «&. Tseng (1971 : 221) who make a fundamental distinction based on 1979] Araliaceae— I (Philipson) 7 basic leaf-organization (either pinnate or palmate) and recognize the heterogeneity of Harms' Aralieae; however, this system was deliberately not completely developed. Generic delimitation. Generic limits within the Araliaceae have long been unstable. As in the Umbelliferae, the flower conforms to a simple and relatively uniform pattern throughout most of the family and systematists have resorted to small technical diff"erences to delimit genera. Harms recognized 51 genera, while Viguier, little more than a decade later, recognized 80; Hutchinson distinguished 84 genera but with criteria very differently weighted as compared with Viguier. Nevertheless, in Malesia as elsewhere there are several distinctive and very natural genera, including Harmsiopanax, Aralidium, Osmoxylon, Anakasia, Mackinlaya, Arthrophyllum, Delarbrea, and Trevesia. Many other genera in the family, including those Malesian ones not noted above, are mutually less distinctive and delimitation is based on various combinations of a number of attributes which have been subject to many different standards of weighting and inter- pretation. These include: petals valvate or imbricate; pedicel articulated or not; leaves digitately compound, pinnate, or simple; stamen and locule number and the relationship of these numbers; style free or connate; endosperm ruminate or not; and thorns present or not. The present treatment is characterized by a number of generic reductions or exclusions ; only one genus described since 1900 has been retained (Anakasia). Hederopsis is united with Macro- panax, a genus with a very similar facies and (in part) overlapping range and which was separated merely on the number of cells in the ovary and variations in inflorescence structure. Wardenia has been united with Brassaiopsis because apart from its simple palmately veined leaves there are no differences; better material collected in recent years has provided evidence that the ovary is in fact 2-locular, but that one ovule aborts and the fruit is as a result 1 -seeded. Acanthophora differs from Aralia only in habit (van Steenis, 1948). Philipson (1951) already regarded Anomo- panax as insufficiently distinct from the older Mackinlaya, a union retained here. The same author (1973) combined Boerlagiodendron with the earlier described Osmoxylon as a number of species intermediate between the two genera had come to light. The Malesian species formerly included in Tetraplasandra as well as Peekeliopanax were likewise by Philipson (1970a) reduced to two species of Gastonia; but it should be noted that in Malesia as on the SW. Indian Ocean islands this genus is very close to Polyscias, particularly the very similarly ranging sect. Eupteron (P. nodosa, P. ledermannii, etc.). The two largest Malesian genera are herein both treated in a wider sense. Polyscias includes as its type section a distinctive Melanesian/Micronesian group of species which in the wild state extends into eastern Malesia and the Philippines, and in cultivation further west. The remainder of the genus as represented in Malesia comprises some rather distinctive species, several of which have been given generic rank (Eupteron, Kissodendron, and Palmervandenbroekia). If the name Polyscias were to be confined to the type-section of the genus it would be necessary to recognize a considerable number of small genera. Retention of a wide concept for the genus therefore appears to be preferable. Schefflera, by far the largest genus of the family in Malesia, is likewise more broadly conceived than in the past, although in our region only a small number of species from segregate genera are involved (i.e. those formerly in Brassaia, Plerandra, Scheffleropsis, and Tupidanthiis). On a world-wide basis, Frodin (1975) recommended the reduction of 12 segregate genera; improved knowledge of the genus required that, as in Polyscias, this step be taken or have the genus split into a number of diflFerently organized smaller genera with more serious nomenclatural conse- quences. A broad concept of Schefflera has therefore been adopted. It may be noted here that the flowers in both the former genera Plerandra and Tupidanthus, characterized by numerous stamens, are thought by Frodin to represent a secondary development related to bat pollination, and the large fruits for dispersal by bats or larger birds; this is supported by the position of the inflorescences, which are beneath the leafy rosettes as a result of retarded development although the axes remain sympodial and the shoot units mostly orthotropic. As divisions of the larger genus Schefflera, they are not at all closely related, conforming to the views of Tseng (1974) on the pollen morphology and contrasting with the views of Harms (1898) and Eyde & Tseng (1971), who regarded them (as genera) as of close affinity and relatively primitive within the family. In fact, rather different levels of specialization are represented overall by the two taxa, not just in the pollen morphology (Tseng, I.e.). 8 Flora Malesiana [ser. I, vol. 9^ Specific delimitation. This has offered many problems, but fortunately the very abundant material now available from many areas has revealed that many of the species described up to 1951 are in fact conspecific. Especially was the great plasticity in vegetative and inflorescence development not sufficiently appreciated in the past, partly due to imperfect field knowledge of the plants. This has led to considerable reductions in Arthrophyllum, Aralia, Osmoxylon, Gastonia, and parts of Schejflera and Folyscias. Other species have been reduced through a regional approach to the genera. However, in Osmoxylon and Schefflera, many species from the Philippines, Celebes, western New Guinea, and to a lesser extent Borneo and Sumatra are still known only from very few or even only a single collection; this has meant a rather tentative treatment in many cases. The same applies in a more limited way in some of the other genera. Certain species are rather polymorphic, and at least in Schefflera several 'species-complexes' have been discerned. References: Baumann-Bodenheim, Bull. Soc. Bot. Suisse 56 (1946) 13-112; Borchert, Amer. J. Bot. 56 (1969) 1033-1041; Corner, Ways. Trees Malaya (1940) 153; Eyde, Amer. Sci. 63 (1975) 430-437; Hyde & Tseng, Science 166 (1969) 506-508; J. Arn. Arb. 52 (1971) 205-239; Frodin, The complex of Cephaloschefflera in Schefflera (Araliaceae), Thesis, Cambridge, U.K. (1970); J. Arn. Arb. 56 (1975) 427^148; Halle, Oldeman & Tomlinson, Tropical trees and forests: an architectural analysis, Berhn (1978); Hallier /. New Phytol. 4 (1905) 151-162; Harms in E. & P. Nat. Pfl. Earn. 3, 8 (1898) 1-62; Hladik, Adansonia 10 (1970) 383-407; Hutchinson, Gen. Fl. PI. 2 (1967) 52-81; Philipson, Bull. Brit. Mus. Nat. Hist. Bot. 1 (1951) 3-20; New Zeal. J. Bot. 5 (1967) 134-165; Blumea 18 (1970a) 490-495; ibid. 18 (1970b) 497-505; ibid. 21 (1973) 81-89; in Tomlinson & Zimmermann (eds.), Tropical trees as living systems (1978) 269-284; Rodriguez, Univ. Calif. Publ. Bot. 29 (1957) 145-318; in Heywood (ed.). The biology and chemistry of the Umbelliferae (1971) 63-91 ; Seemann, Revision of the natural order of Hederaceae, repr. from J. Bot. London (1868); van Steenis, Bull. Bot. Gard. Btzg III, 17 (1948) 390-391 ; Thorne, Aliso 6 (1968) 57-66; Notes R. Bot. Gard. Edinb. 32 (1973) 161-165; Tseng, Amer. J. Bot. 61 (1974) 717-721; Viguier, Ann. Sci. Nat. Bot. IX, 4 (1906) 1-209. Uses. A variety of minor local uses are reported by Burkill (1966), Heyne (1927), and OcHSE & Bakhuizen van den Brink (1931) as well as in the notes under individual species in this Flora. The only species that form articles of trade are the taxa of Polyscias sect. Polyscias popularly grown as foliage and hedge plants ; however, many other species in a variety of genera are of actual or potential ornamental worth, with Schefflera actinophylla (Endl.) Harms and S. longifolia (Bl.) Vig. being particularly widely used. In Papua New Guinea, Gastonia spectabilis is cut for timber and the wood used for light carpentry, boxes, etc. Monkeys are fond of the flush of some aromatic species of Schefflera, as observed in West Java and North Sumatra. References: Burkill, Diet. rev. ed. (1966); Heyne, Nutt. PI. (1927); Ochse &. Bakhuizen VAN DEN Brink, Veget. D.E.I. (1931). Notes. Nomina nuda and invalidly published names have only been quoted if they have been cited in Index Kewensis. Notes for collectors. Many araliads present problems to collectors because of the size of their leaves and inflorescences. It is often advisable to select leaves of medium size, but the maximum size of leaves should be recorded on the label. It is important to preserve the junction of leaf and stem and also sufficient of the leaf to allow reconstruction of the whole. Likewise, with inflores- cences the base, main axis and some primary branches should be preserved so that the whole can be visualized — ultimate branches alone are insufficient. Fruiting material is as useful as a flower- ing specimen. Collectors should be alert to note the existence of vegetative heteroblasty and floral dimorphism and document these with specimens and notes; the form in immature plants is important. Rapid drying is essential or all parts will disarticulate and very fragmentary specimens result. key to the GENERA 1 . Petals imbricate. Tribe Aralieae. 2. Leaves simple, palmately or pinnately lobed, or entire. 3. Leaves palmately lobed, tomentose 1. Harmsiopanax 3. Leaves pinnately lobed or entire, glabrous 2. Aralidium 1979] Araliaceae — I (Philipson) 9 2. Leaves pinnately compound (or bi- or tripinnate). 4. Leaves twice (or more) pinnate 3. Aralia 4. Leaves once pinnate. 5. Leaflets many 4. Delarbrea 5. Leaflets 5 or fewer 5. Pentapanax \. Petals valvate. 6. Petals with a narrow base, or claw. Tribe Mackinlayeae 6. Mackinlaya 6. Petals with a broad base. Tribe Schefflereae. 7. Inflorescence rays trifid : central branch shorter with 'false fruits', the two lateral longer with normal flowers 7. Osmoxylon 7. Inflorescence branches not as above. 8. Ovary with one cell 8. Arthrophyllum 8. Ovary with more than one cell. 9. Leaves pirmate. 10. Pedicel not articulated below the flower 9. Gastonia 10. Pedicel articulated below the flower 10. Polyscias 9. Leaves not pinnate. 1 1 . Pedicel articulated below the flower. 12. Leaves digitately compound (or rarely unifoliolate) (Malay Peninsula, Sumatra, Java) 11. Macropanax 12. Leaves simple, not articulated with the petiole (West New Guinea) .... 12. Anakasia 1 1 . Pedicel not articulated below the flower. 13. Leaf simple (or unifoliolate) or palmately lobed. 14. Leaf palmately lobed. 15. Ovary 2-celled 13. Brassaiopsis 15. Ovary 10- or more-celled 14. Trevesia 14. Leaf simple (or unifoliolate). 16. Articulation present between petiole and blade Schefflera 16. No articulation between petiole and blade. 17. Ovary 2-celled 13. Brassaiopsis 17. Ovary 4- or more-celled 15. Dendropanax 13. Leaf digitately compound. 18. Petiolules joined together by a web of tissue 14. Trevesia 18. No such web of tissue present. 19. Styles or stigmas 2. 20. Style bifid 16. Acantbopanax 20. Styles united into a column 13. Brassaiopsis 19. Styles or stigmas more than 2 Scbefl9era 1. HARMSIOPANAX Warb. in E. & P. Nat. Pfl. Fam. Nachtr. 1 (1897) 166; Harms, Bot. Jahrb. 56 (1921) 413; Hutch. Gen. Fl. PI. 2 (1967) 62; Philipson, Blumea 21 (1973) 81. — Schubertia Bl. Bijdr. (1826) 884, nom. illeg., non Mirb. 1812. — Horsfieldia Bl. ex DC. Prod. 4 (1830) 87, non Willd. 1805; Bth. in B. & H. Gen. PI. 1 (1865) 937; BOERL. Handl. 1 (1890) 633; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 60. — Fig. 1-3. Sparingly branched or single-trunked often monocarpic trees, up to 18 m. Stems stout, bearing terminal clusters of large, palmately lobed, often peltate, exstipulate leaves. Trunk, petioles, and sometimes the blades spiny. Flowers in very large, repeatedly branched, terminal panicles which develop after the leaves have fallen. Umbellules arranged racemosely on the ultimate branchlets, sessile or peduncled, each consisting of a few to many pedicelled flowers. Pedicels not jointed, subtended by a bract and bearing two subulate bracteoles. Flowers hermaphrodite or with hermaphrodite flowers on terminal and male flowers on basal branches. Calyx a 10 Flora Malesiana [ser. I, vol. 9^ Fig. 1. Harmsiopanax ingens Philipson ssp. ingens. a. Leaf, x "/j, b. part of inflorescence, x V3, c. umbel- lule, X 4, d. developing fruit, x 12 (a NGF 36901, b-d Philipson 3483). 1979] Araliaceae — I (Philipson) 11 minute rim. Petals 5, free, valvate with a broad base. Stamens 5, dorsifixed, versa- tile, introrse. Ovary inferior, narrowly obconic, densely bristly; cells 2; disk conical, deeply cleft between the two subulate styles. Fruit consisting of 2 dry mericarps, each 3-ribbed and bearing a persistent slightly hooked style. Distr. Malesia: 3 spp. from Java, the Lesser Sunda Is., Celebes, and New Guinea. Ecol. Montane and mossy forest and in regrowth on grassy hillsides. Notes. Harmsiopanax is a small structurally isolated genus confined to Malesia. The three species are uniform both in their vegetative and their reproductive features. It has long been recognized that some of the characters of this genus are anomalous within Araliaceae and a return to its earlier position within Umbelliferae would have something in its favour. The monocarpic habit is unknown elsewhere in Aralia- ceae, but is not uncommon in Umbelliferae. The character of the fruit, which splits into two dry mericarps, closely approaches the fruit structure of Umbelliferae, and the vascularization of the gynoecium is also characteristic of that family. However, the structure of the leaf-base, the woody habit, and the shape of the petals all incline towards Araliaceae. KEY TO THE SPECIES 1 . Upper surface of leaves uniformly setulose. 2. Umbellules sessile 1. H. aculeatus 2. Umbellules peduncled 2. H. harmsii 1 . Upper surface of leaves with many (or rarely few) larger spines among the setulose hairs 3. H. ingens 1. Harmsiopanax aculeatus (Bl.) Warb. ex Boerl. Handl. 3 (1900) 88; Koord. Exk. Fl. Java 2 (1912) 719; Atlas 4 (1916) f. 668 & 669; Bakh. /. & OosTSTR. in Back. Bekn. Fl. Java (em. ed.) 7 (1946) fam. 159, p. 19; Back. & Bakh./. Fl. Java 2 (1965) 171; Steen. Mt. Fl. Java (1972) pi. 3-2; Philipson, Blumea 21 (1973) 82. — Schubertia aculeata Bl. Bijdr. (1826) 885. — Horsfieldia aculeata (Bl.) DC. Prod. 4 (1830) 87; Benn. PI. Jav. Rar. (1840) 123, t. 26; K. & V. Bijdr. 7 (1900) 57; Boerl. Handl. 1 (1890) 647. — Horsfieldia peltata Bth. in B. & H. Gen. PI. 1 (1862) 937. — Fig. 2. Tree up to 4 m, with a slender spiny trunk. Young stems covered more or less densely with woolly hairs, bristles, and spines with bulbous bases, the spines enlarging on older stems. Leaves rounded, variable in size, often 60 cm or more in 0, deeply palmately lobed, usually peltate in mature leaves, sinuses between the lobes broad or narrow, lobes 7-10, usually sharply and irregularly incised and toothed, apex acute, upper surface rather sparsely covered with evenly-spaced, appressed, sometimes branched hairs (denser on the main veins), underside densely clothed with a soft, woolly tomentum, often with some bristles on the main veins; petiole c. 60 cm, 1 cm at base, terete with clasping base, densely covered with woolley hairs, bristles, and some spines. Inflores- cence up to c. 70 cm long, main branches rather sparsely covered with a short tomentum and, when young, bearing numerous bracts similar to the leaves but smaller, not peltate, and often 3-lobed or entire; ultimate branchlets slender and often woolly-tomentose, bearing minute linear bracts which subtend the sessile umbellules. Umbellules about 4 mm in flower, the broadly ovate outer bracts forming a more or less distinct involucre. Flowers hermaphrodite or male, either mixed in an inflorescence, or separate, c. 10-15 per umbellule, each subtended by a lanceolate receptacular bract c. 2 mm long. Pedicel c. V2 rnm long. Calyx rim fringed. Petals strap-shaped, c. IV2 nim long at anthesis. Filaments c. 2 mm; anthers c. 0.3 mm long, orbicular. Ovary covered with cilia which lengthen as the fruit ripens. Mericarps long-ciliate, crowned with the divergent styles. Distr. Malesia: Java, Lesser Sunda Is. (Bali, Lombok, Sumbawa, Flores, Timor), southern half of Celebes. There is a single Korthals sheet in L, ticketed from Central Sumatra, but this is presum- ably wrongly localized. Ecol. Usually in rather dry, open localities, but also in forest, in secondary forest, also pioneering on rocks, in grasslands and on lava-streams, 300-1800 m. Fl.fr. April-Nov. Schmutz found it in Flores flowering in October, but leaves had fallen. Vern. Java: djankurang, d. tjutjuk, djogloran- grang, S, gabus, garang, g. lanang, gungrang, udulan laki, J. Note. In this species lateral shoots usually appear below the infructescences so that the trees are not normally monocarpic. 2. Harmsiopanax harmsii K. Sen. in K. Sch. & Laut. Nachtr. (1905) 329; Harms, Bot. Jahrb. 56 (1921)413. Tree up to 7 m, with a slender trunk. Young stem covered with bristles, hairs, and spines, older stems with smooth bark with small rounded lenti- cels and numerous spines. Monocarpic. Leaves 12 Flora Malesiana [ser. I, vol, 9^ Fig. 2. Habit of Harmsiopanax aculeatus (Bl.) Warb. ex Boerl. Coarse shrub on old lava-streams in E. Java (Mt Idjen) at c. 900 m altitude (Photogr. van Steenis). rounded, up to 30 by 40 cm, deeply palmately lobed, cordate at base, lobes 5-9 with broad sinuses between them, margin unevenly and sharply den- tate, apex acute, upper surface densely covered with evenly spaced bristles of varying size (larger on the main veins), appressed and directed towards the leaf margin, often with woolly hairs inserted on their enlarged bases, the underside very densely woolly and with many bristles, usually bearing crisped hairs on their enlarged bases; petiole 50 cm, V2 cm at base, terete with clasping base, densely covered with bristles, woolly hairs, and spines. Panicle at first with numerous leaf-like bracts, the principal branches with some spines, rather sparsely covered with bristles and hairs, ultimate branches slender and tomentose, bearing linear bracts c. 4 mm long subtending peduncled umbellules; peduncles up to 5 mm, slender, tomen- tose, bearing 2 minute bracts. Umbellules spheri- cal, c. 4-5 mm in flower, outer bracts not form- 1979] Araliaceae — I (Philipson) 13 ing a distinct involucre. Flowers hermaphrodite, maturing in basipetal succession, the lower bracts of a branch either with sterile umbellules or lacking flowers; up to 60 in an umbellule, each subtended by a lanceolate ciliolate bract c. 1 mm long, and borne on a glabrous pedicel c. VU mm long. Calyx rim fringed with many lacerate filaments. Petals ovate, c. 1 mm long. Filaments c. 1 mm; anthers c. V2 nim long. Ovary covered with cilia which lengthen as the fruit ripens. Mericarps with rounded ribs, long-ciliate, crowned by the diver- gent styles. Distr. Malesia: Papua New Guinea (Madang Distr., Western Highlands, Morobe Distr. & Central Distr.). Ecol. Forested hills, grassy slopes, and road- sides, 100-1800 m. Vern. Opme, Ganja, Mt Hagen, mafiong, Sattelberg, Morobe Distr. Note. Information about the habit is inade- quate. The stalked spherical umbellules are very distinctive. 3. Hamisiopanax ingens Philipson, Blumea 21 (1973) 84. ssp. ingens. — Fig. 1, 3. Unbranched tree up to 1 8 m with a thick or some- times slender trunk densely covered, except towards the base of mature specimens, with long, sharp, upwardly directed spines and marked with leaf-scars. Monocarpic. Leaves usually peltate, rounded, up to 1 m 0, deeply palmately lobed, lobes usually with minor lobes and coarsely dentate, apex acute, upper surface bearing few to many long spines, especially on the midrib and principal veins between which the surface is often rugose and glabrous except for the remains of a tomentum of branched hairs, or with many bristles often with woolly hairs on their bases, the under- surface also with few to many long spines and usually clothed with a fawn or greyish woolly tomentum of branched hairs, or densely furnished with bristles usually with woolly hairs on their bases, or occasionally glabrous between the spines Fig. 3. Harmsiopanax ingens Philipson. Left: apex of leafy stem; right: the large inflorescence (Photogr. Frodin, Murmur Pass, 1971). 14 Flora Malesiana [ser. I, vol. 9^ except for a few bristles; petiole up to 1 m and 3 cm 0, terete with clasping base, covered with woolly hairs and bearing many spines. Panicle up to 5 m long and 5 m wide, leafless or with lobed bracts c. 10-20 cm long, principal branches spiny especially below, ultimate branches slender, tomentose, bearing linear bracts c. 1 cm long sub- tending peduncled or sessile umbellules; peduncles elongating as the fruit ripens, up to 4 mm, rather stout, tomentose, bearing 1 or 2 minute bracts. Umbellules bowl-shaped, c. 6-10 mm in flower, enlarging slightly in fruit, with an involucre of about 8 ovate bracts, 2-4 mm long and ciliolate distally. Flowers hermaphrodite, maturing in basipetal succession, terminal branches bearing maturing fruit while lower branches bear flowers or unopened buds; usually c. 12-16 (8-20) in an umbellule each subtended by an involucral bract or a narrower receptacular bract and borne on a glabrous pedicel 1-2 mm long. Calyx rim fringed with many lacerate filaments. Petals ovate, 1-2 mm long. Filaments 2-3V2 mm; anthers ^2-^/4 mm long. Ovary covered with cilia which lengthen as the fruit ripens. Mericarps with rounded ribs, long- ciliate, crowned by the divergent styles. Distr. Malesia: New Guinea (NW. Irian and extending along the central mountains from the Orion Mts to the Owen Stanley Range, Murray Pass). Ecol. Montane and mossy forest and second- growth forest, 2000-3600 m, occasionally rather lower. Vern. Papua: mauku, Huli; Mandated Terr.: Sepik Distr., kamul, Hindenburg Ra.; Western Highlands: murri, Hagen, tolsan, Minj, mauri, Melpa, mat, Mendi, kinogore, makua, makw, Enga; Eastern Highlands : kimu, Ka, ollu, Chimbu. Notes. A striking, single-trunked, monocarpic tree bearing immense inflorescences. The bark is described as grey brown and the wood white with a wide pith. The inflorescence has the appearance of bearing female flowers above and male flowers below, but this is evidently due to a basipetal sequence of anthesis. The terminal flowers have stamens when freshly opened and all those on lower branches bear styles. Apparently, the female organs of the lower branches are functional because branches from mature inflorescences bear fruit uniformly. Nevertheless, herbarium specimens cannot adequately represent such a large inflores- cence so that the possibility remains that some female-sterile flowers occur in this species. Variation occurs in both tomentum and in- florescence characters. For example, most speci- mens from West Irian have small umbellules and fewer leaf-spines. In the eastern part of the Eastern Highlands District a number of gatherings display a series of variations: the under-leaves give the appearance of being glabrous between bristles, the inflorescence branches bear small leafy bracts, the umbellules are sessile, with rather numerous tc. 18-21) small flowers subtended by rather broad bracts. Specimens from Mt Otto show all these features combined, but other specimens from this region diverge from the typical state in only some of these characters. No specimens of this subspecies are known from the Finisterre Range and only one from the Owen Stanley Range. ssp. moniliformis Philipson, Blumea 21 (1973) 86. Umbellules disposed irregularly along the branches, singly or in small groups, with bare spaces intervening, sessile; flowers usually c. 20-30 per umbellule, floral parts smaller than in ssp. ingens; fruiting heads rather small (c. 5 mm 0). Distr. Malesia: Papua New Guinea (districts bordering on the Huon Gulf). Vern. Morobe Distr.: mobian, Finschhafen. Note. This subspecies occurs at lower altitudes than is usual for ssp. ingens i\5QQ-2QQO m). No specimens of either subspecies have been collected from higher altitudes in the mountains north of the Markham River and the Huon Gulf. At higher altitudes in the Owen Stanley Range ssp. ingens is known from one gathering. The most south-easterly gathering at present known (Carr 13603) has a distinctive appearance due to the straight rigid in- florescence branches with small sessile umbellules. 2. ARALIDIUM MiQ. PI. Jungh. 3 (1855) 423; Fl. Ind. Bat. 1, 1 (1856) 762, t. 13; Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 25; Bth. in B. & H. Gen. PI. 1 (1865) 936; Hemsl. in Hook. Ic. PI. 16 (1886) t. 1549; Boerl. Handl. 1 (1890) 631; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 60; Hutch. Gen. Fl. PI. 2 (1967) 59. — Fig. 4. Unarmed tree or shrub with simple, exstipulate pinnately lobed, irregularly incised, or entire leaves. Inflorescence a large panicle, with cymules of small flowers arranged racemosely on the branches. Pedicels articulated below the ovary. Dioecious. Male flowers: calyx 5 ; petals 5, imbricate; stamens 5, anthers dorsifixed. Female flowers: calyx and corolla similar to male; staminodes 5; ovary with 3 1979] Araliaceae — I (Philipson) 15 Fig. 4. Aralidiumpinmtifidum (Jungh. & de Vriese) Miq. a. Habit, x ^/„ b. ' variable in size and complexity; petiole up to 52 cm, Vu cm wide, terete, striate, with a membranous base ensheathing the stem; leaflets 5 (rarely 3) or the central petiolule (or the central 3 petiolules) frequently disiding to bear three, or more rarely 5 leaflets: lateral petioles short (c. 1-2 cm), the three central longer (up to 12 cm); lamina elliptic or ovate, up to 48 by 22 cm, base abruptly attenuated into the petiolule or subcordate, apex acuminate or gradually narrowed, acute, margin entire, denticulate or coarsely serrate, especially towards the apex, membranous, lamina of the terminal leaflet (and less frequently of the central three leaflets) sometimes deeply 3-lobed or with 3-5 separate leaflets, of which the lateral are markedly oblique at the base. Inflorescence a terminal compound umbel, very variable in size, either entirely of male flowers or with male and hermaphrodite flowers, often overtopped by sym- podial growth; peduncle terete, striate, stout, up to 30(^5) cm, ^4 cm Z, bearing lanceolate bracts below the rays; primary rays c. 9-18, 10-20 cm, striate, with distal small linear bracts; secondary rays about 5-10, 3-6 cm, dividing again (often repeatedly) either umbellately or in an irregular cymose manner, the central ray frequently more strongly developed. Calyx lobes 5, triangular, c. 1 mm long. Petals 5, obovate c. Vjj mm long. Filaments c. P 2 mm, anthers small. Ovary obconic, 1-2 mm long, narrowly turbinate in male flowers, ovoid and quickly swelling in female flowers. Fruit up to 2' 2 by 3 cm, compressed, rotund, constricted in the mid-axis, the two halves frequently unevenly developed. Distr. Solomon Is.; in Malesia: New Guinea (incl. New Britain and Am Is.), Celebes, and Central & S. Philippines. Ecol. Rain-forest, open hill forest, and montane forest, also in secondary grov.ihs, from sea-level to 1450 m. Uses. Lant-Poole (I.e.) recorded that at Mt Obree leaves and flowers are cooked with coconut oil and put in armlets in dances. Vern. Philippines: binlaon, C.Bis., pararau, Bag., tagima. Sub., lumot-lumot, Mindanao; New- Guinea: bugini, wale, yam bonga, Sepik Distr., lak-lak, W. Highlands, po'undo, S. Highlands, Papua, nere. Central Distr., Papua, narona, New- Britain. Notes. The leaves have a strong parsley-like odour. The flowers are creamy white, and the fruits blue to purple with a glaucous bloom. Salt is said to be obtained from the ashes of the leaves. Although collected frequently in the Philippines and New Guinea, this species is unrecorded for the Moluccas. There is considerable variation in the size of the leaves and of the inflorescence. A few New Guinea specimens are intermediate in charac- ter between this and the equally common M. schlechteri, and are interpreted as hybrids. 2. Mackinlaya radiata F*hillpson, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 6. Slender shrub to 5 m. Petiole e. 20 cm, terete and finely striate, base ensheathing the stem, mem- branous. Leaflets 5, or the central petiolule bearing three leaflets, the two lateral petiolules short (1- P 2 cm), the three central longer (6-8 cm, or the midpetiolule to 11 cm); lamina of the lateral leaflets elliptic or ovate, up to 20 by 12 cm, base abruptly attenuated into the petiolule, apex gradually narrowed, acute, margin entire or minutely denticulate tow-ards the apex, mem- branous: lamina of the central leaflet similar or deeply 3-lobed or with 3 separate leaflets of which the lateral are strongly oblique at the base. In- florescence a terminal compound umbel; peduncle terete, striate, stout, from 20 cm to considerably longer, 4—6 mm Z, bearing lanceolate bracts below the rays; primary rays numerous (c. 30-50), 9-18 cm, slender, striate, with distal minute, linear, caducous bracts; secondary rays (pedicels) numerous (35-130), filiform, 1-2 cm; outer flowers male, central hermaphrodite. Calyx lobes 5, narrowly triangular, e. V2 mm long. Petals 5, obovate, e. 1 mm long. Ovary narrowly obconic in male flowers, ovoid in hermaphrodite flowers, c. 0.7 mm long. Fruit (immature) ovate, com- pressed. Distr. Malesia: New Guinea (NW. Irian; Sepik Distr.). Ecol. Montane rain-forest and mossy forest, 900-1200 m. Vern. Apiyetimber, Sepik Distr. Note. Flowers creamy white. 3. Mackinlaya schlechteri (Harms) Philipson, Bull.Br. Mus. Nat. Hist. Bot. 1 (1951)7. — Anomo- panax schlechteri Harms in K. Sch. & Laut. Nachtr. (1905) 332, t. 13. — Anomopanax versteegii Harms, Nova Guinea 8 (1910) 276. — M. versteegii (Harms) Philipson, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 7. — M. brass a Philipson, I.e. 6. — M. klossii Philipson, I.e. 6. — M. subulata Philipson, I.e. 7. Slender shrub to 6 m. Leaves l-3-{ rarely 4-)folio- late. Petiole usually less than 10 cm, but occasion- ally longer (to 20 cm), especially in compound leaves, terete and finely striate, base ensheathing the stem, membranous. Lamina elliptic obovate, or oblong, occasionally irregularly lobed, up to 28 by 12 cm but usually considerably smaller, base cuneate or truncate (of lateral leaflets often 30 Flora Malesiana [ser. I, vol. 9^ Fig. 11. Osmoxylon novoguineense (Scheff.) Becc. a. Leaf half, b. base of petiole, both x ^U, c. inflores- cence, X '•/, /flower and CS of ovary, x 4, e. false fruit and ditto in CS (Craven & Schodde 789). Drawn by W. R. I>hilipson. 1979] Araliaceae— I (Philipson) 31 oblique), apex shortly acuminate, acute, margin reduce fever and to relieve ^korima\ Pieces of leaf entire or dentate towards the apex, membranous or placed in a cavity relieve toothache. The leaves are chartaceous. Inflorescence a terminal compound wrapped around taro at planting to encourage umh>el, often overtopped by a lateral branch at its growth. base; peduncle terete, striate, c. 10-20 cm, bearing Vern. Dako, Wissel Lakes, kolobang, kulbang, small lanceolate bracts below the rays; primary Sepik Distr., aw/:^, ^e/uj/a, mwA:/o/o, E. Highlands, rays 6-15, c. 4-7 cm, striate, with minute distal narona. New Britain. caducous bracts; secondary rays (pedicels) c. 10-20, Notes. The flowers are white and the ripe fruit filiform or rather rigid, usually 5-12 mm; male mauve to purple with a glaucous bloom, flowers towards the outside of the umbellules. A large number of collections made in recent Calyx lobes 5, triangular to subulate, "j-l mm years throughout New Guinea all have regularly long. Petals 5, obovate. Ovary narrowly turbinate, compound umbels with the flowers borne on in male flowers obconic or ovoid, c. ^U nim long in branches of the third degree in the form of strict hermaphrodite. Fruit large, 15 by 22 mm, com- umbels. In two of the earliest gatherings (Schlech- pressed, rotund, constricted above and below on ter 14365 and Versteeg 1419) the third degree the central axis; styles persistent, recurved. branches frequently divide again either umbellately Distr. Malesia: New Guinea (along the Central or cymosely. These two specimens were described Ranges, from the Star Mts east to Meyamya), also as species by Harms. Philipson later (1951) kept in New Britain. the forms with regular umbellules separate (de- Ecol. Rain-forest and montane forest, 600- scribing three species). All these five entities are now 2300 m. considered conspecific, the Schlechter and Uses. The cut stem exudes a viscous sap which Versteeg specimens being regarded as rare is an irritant. The leaves are aromatic. The plant is anomalies in a widespread and abundant species, reported to be poisonous and to have a number of It is possible that the Versteeg plant is a hybrid medicinal uses. The boiled leaves are eaten to with M. celebica. 7. OSMOXYLON MiQ. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 5; Bth. in B. & H. Gen. Pi. 1 (1865) 944; Becc. Malesia 1 (1878) 193; Boerl. Ann. Jard. Bot. Btzg 6 (1887) 123; O. K. Rev. Gen. PI. 1 (1891) 645; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 32; Bot. Jahrb. 56 (1920) 384; Hutch. Gen. Fl. PI. 2 (1967) 73; Philipson, Blumea 23 (1976) 99. — Eschweileria Zipp. ex Boerl. Ann. Jard. Bot. Btzg 6 (1887) 112, non Eschweilera Mart. 1828; Handl. 1 (1890) 640. —Pseudosandalum O. K. Rev. Gen. PI. 1 (1891) 271 {'Pseudosantalurn'), nom. illeg. — Boerlagiodendron Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 31 ; in K. Sch. & Laut. Fl. Schutzgeb. (1900) 484; Bot. Jahrb. 56 (1920) 377; Hutch. Gen. Fl. PI. 2 (1967) 72. — Fig. 11, 13-16. Unarmed, glabrous or tomentose shrubs or trees. Leaves palmately lobed or simple, rarely digitately compound; stipules forming a ligule, and the base of the petiole furnished with one to several spiral or transversal crests or collars (very rarely absent). Inflorescence a terminal compound umbel; peduncle short; primary rays each terminating into three branches; the central branch bearing a head or umbellule of almost always sterile bacciform flowers ('pseudo-fruits'); the two lateral branches each bearing a head or umbellule of hermaphrodite flowers. Calyx an obsolete rim or 0. Corolla with few to many lobes above, tubular below. Stamens 4-30, filaments thick, anthers oblong, exserted. Ovary inferior, not articulated with the pedicel, cells 1-many; disk flat with a central raised boss bearing the pustulate stigmas. Fruit subglobose (ribbed when dry); exocarp fleshy, endocarp crustaceous. Seeds compressed, endosperm smooth or wrinkled. Distr. About 50 spp., of which 40 occur in Malesia, extending from Borneo and the Philippines east- wards through Celebes and Moluccas to New Guinea, the remainder lying further north and east in Taiwan, Micronesia, Melanesia and the New Hebrides. Fig. 12. 32 Flora Malesiana [ser. I, vol. 9^ Ecol. Mainly understorey trees in primary rain-forest, also in second growth forest, usually at low altitudes, especially in shaded situations and near rivers, 15. O. bomeense a characteristic rheophyte. Notes. The foliage, inflorescence and flowers of this genus are all unique within the family. The base of the petiole often bears a spiral crest, or this may form a simple collar; the blade varies from simple to elaborately compound ; the inflorescence is composed of trifid rays, the central branch bearing sterile bacciform flowers; the corolla is tubular. The central bacciform flowers (pseudo-fruits) are sterile, except in 12. O. yatesii, in which apparently mature seed was once found. By exception a specimen of 40. O. luzoniense had apparently fertile flowers on the central branches of the inflorescence. KEY TO THE SPECIES 1 . Leaves simple, without lobes. 2. Petiole base without crests (New Guinea) 1. O. miquelii 2. Petiole base with 1 or more crests. 3. Petiole base with several crests (PhiUppines) 8. O. oblongifolium 3. Petiole base with a single collar-like crest. 4. Fertile flowers pedicelled (Philippines). 5. Leaf obovate to oblanceolate, petiole 3 cm or less 2. O. dinagatense 5. Leaf elliptic, petiole 5 cm or more 3. O. simplicifolium 4. Fertile flowers sessile (or subsessile). 6. Leaf broadly obovate (Solomons) 4. O. spathipedunculatum 6. Leaf lanceolate or narrowly obovate. 7. Pseudo-fruits pedicelled (Moluccas) 5. O. articulatum 7. Pseudo-fruits sessile. 8. Inflorescence c. 60 cm (Moluccas) 6. O. umbelliferum 8. Inflorescence c. 30 cm (New Ireland) 7. O. lanceolatum 1 . Leaves lobed or digitately compound. 9. Leaves digitately compound. 10. Ovary 10-16-celled (New Guinea) 13. O. geelvinkianum 10. Ovary 4-5-celled. 11. Leaflets lobed (Philippines). 12. Primary rays of inflorescence c. 10 or fewer 9. O. catanduanense 12. Primary rays of inflorescence more than 20. 13. Inflorescence c. 16-20 cm 10. O. caudatum 13. Inflorescence c. 5-6 cm 11. O. heterophyllum 11. Leaflets entire. 14. Leaflets elliptic-oblong to ovate (Philippines) 12. O. yatesii 14. Leaflets linear-lanceolate. 15. Leaflets 4-7, petiole 4-6 cm (Philippines) 14. O. lineare 15. Leaflets 9 or more, petiole longer than 10 cm (Borneo) 15. O. borneense 9. Leaves lobed (sometimes smaller simple leaves below inflorescence). 16. Pseudo-fruits sessile. 17. Petiolar crests long-pectinate. Bracts of the peduncle heavily setose (Philippines) 16. O. pulcherrimum 17. Petiolar crests fimbriate or undulate. Bracts of the peduncle not setose (Moluccas). 18. Lateral inflorescence branches with opposite, persistent bracts close to the base 17. O. soelaense 18. Lateral inflorescence branches with scars of opposite caducous bracts near the middle 18. O. globulare 16. Pseudo-fruits pedicelled. 19. Lateral inflorescence branches without an articulation. 20. Petiolar base with a single collar-like crest (Borneo) 19. O. kostennansii 20. Petiolar base with several crests. 21. Petiolar crests long-pectinate. 22. Fertile flowers pedicelled. Stamens at least 7 (Key Is.) 20. O. barbatum 22. Fertile flowers sessile (or subsessile). Stamens 5 (Bismarck Arch.) 21. O. pfeilii 21. Petiolar crests fimbriate or undulate. 23. Stamens and ovary cells more than 5. Flowers usually pedicelled (Moluccas) 22. O. palmatum 23. Stamens and ovary cells 4. Flowers sessile or subsessile (Philippines). . . . 23. O. ramosii 19. Lateral inflorescence branches with an articulation (2 bracts or bract-scars). 1979] Araliaceae— I (Philipson) 33 24. Fertile flowers pedicelled. 25. Pedicels of fertile flowers longer than 7 mm (New Guinea) 24. O. novoguineense 25. Pedicels of fertile flowers 5 mm or shorter. 26. Ovary cells 7 or more. 27. Leaf lobes elliptic (Celebes) 25. O. teysmannii 27. Leaf lobes pinnatilobed (Batjan) 41. O. insigne 26. Ovary cells 4-5. 28. Petiolar crests ri entire (Philippines) 26. O. humile 28. Petiolar crests with long setae. 29. Leaves coriaceous, margin thickened, teeth obtuse (Philippines) . . . . 27. O. pectinatum 29. Leaves membranaceous, margin not thickened, teeth setulose (Celebes) 28. O. masarangense 24. Fertile flowers sessile (or if subsessile bracteoles longer than the very short pedicels). 30. Ovary cells 10 or more (New Guinea). 3L Flower buds large (c. 9-12 mm long). Primary rays of inflorescence very strong 29. O. insidiator 3L Flower buds smaller (c. 4-6 mm long). Primary rays of inflorescence more tenuous. 32. Heads of fertile flowers — discoid at anthesis 30. O. boerlagei 32. Heads of fertile flowers globose 31. O. sessiliflonun 30. Ovary cells 6 or fewer. 33. Leaf lobes lanceolate. 34. Leaf lobes 4-7 ; petiole 4-6 cm (Philippines) 14. O. lineare 34. Leaf lobes 9 or more; petiole longer than 10 cm (Borneo) 15. O. bomeense 33. Leaf lobes broader. 35. Leaf surfaces retaining some trace of setulose tomentum. Umbellules of pseudo-fruits small (8 mm or less) (New Guinea) 38. O. micranthum 35. Leaf surfaces glabrous at maturity. Umbellules of pseudo-fruits larger (1 cm or more). 36. Ovary cells 3 (Philippines). 37. Petiolar crests — entire 32. O. camiguinense 37. Petiolar crests long-pectinate 33. O. fenicis 36. Ovary cells 4 or more. 38. Umbellules of pseudo-fruits large (3-5 cm 0) (Philippines). 39. Primary rays of the inflorescence 9-12 cm long. Leaves with strong radiating veins, usually more than 11 34. O. eminens 39. Primar>' rays of the inflorescence 4-5 cm long. Leaf-veins less strongly developed, usually fewer than 11 35. O. serratifolium 38. Umbellules of pseudo-fruits smaller (2^2 cm or less). 40. Leaf with a small triangular lobe below the middle of each side of the blade (not strictly palmately lobed) (Philippines) 8. O. oblongifolium 40. Leaf palmately lobed. 41. Petiolar crests long-pectinate (Celebes) 36. O. celebicum 41. Petiolar crests fimbriate, entire, or undulate. 42. Articulation of lateral branches of umbels close to the base (Talaud Is.) 37. O. talaudense 42. Articulation of lateral branches of umbels near the middle. 43. Inflorescence over 20 cm (New Guinea) 31. O. sessiliflorum 43. Inflorescence under 20 cm (Philippines). 44. Leaf usually 3-lobed. Inflorescence rays delicate, indistinctly setose to glabrous 39. O. trilobatum 44. Leaf usually 5-7-lobed. Inflorescence rays sturdy, markedly setose 40. O. luzoniense 1. Osmoxylon miquelii Boerl. Arm. Jard. Bot. Sparsely branched tree, 15 m. Leaves glabrous, Btzg 6 (1887) 125, t. 16: Harms, Bot. Jahrb. 56 simple, subcoriaceous ; stipules small forming a (1920) 384; Philipson, Blumea 23 (1976) 103. — bicuspid ligule; petioles long (to 19 cm), swollen O. amboinense MiQ. Ann. Mus. Bot. Lugd.-Bat. 1 distally; blade oblong-elliptic, 22-36 by 9-12 cm, (1863) 6, p.p.; Becc. Malesia 1 (1878) \9A,p.p.— subrounded at base and apex or mucronulate, Gastonia simplicifolia Zipp. ex Seem. J. Bot. 3 midrib prominent below, secondary veins arched- (1865) 75, nomen in synon.: ex Boerl. Aim. Jard. ascending and uniting, c. 1-2 cm apart, margin Bot. Btzg 6 (1887) 125, mm. inval. in synon. — entire to undulate. Umbel terminal, sessile, with Pseudosandalum miquelii (Boerl.) O. K. Rev. Gen. many (28-32) radiating rigid, angular, trifid PL 1 (1891) 271. branches c. 7 cm long to first joint. Central 34 Flora Malesiana [ser. I, vol. 9^ Fig. 12. Species density of Osmoxylon Miq. in Malesia; above the hyphen the number of endemic species, below it the non-endemics. branches unknown. Lateral branches c. 5 cm long, articulate near the base. Flowers 20-30, sessile on the expanded ends of the inflorescence branches. Corolla and stamens unknown. Drupes crowded, subrotund, c. 4 mm (dry), c. 8-10 ribbed when dry, crowned by a semiglobose entire stigma, 8-10-celled. Seeds with slightly ruminate endo- sperm. Distr. Malesia: West New Guinea. Only known from the type (coll. Zippelius). 2. Osmoxylon dinagatense (Merr.) Philipson, Blumea 23 (1976) 103. — Boerlagiodendron dinagatense Merr. Philip. J. Sc. 17 (1920) 301; Merr. En. Philip. 3 (1923) 222. Glabrous shrub, c. 2 m. Leaves crowded at the ends of the branches; petiole 2-3 cm, channelled above, with a small triangular base, bearing a short stipular ligule (2-3 mm long) and extending around the base of the petiole as a single narrow collar; blade obovate to oblanceolate, to 23 by 8' ,'2 cm; base narrowed into the petiole, apex rounded with a very short apiculum, margin slightly revolute, entire to obscurely undulate with minute teeth, coriaceous. Inflorescence a terminal compound umbel; peduncle c. IV2 cm, with 1-few bracts (reduced leaf-bases) with small triangular bracts (3 mm long) among the primary rays ; primary rays about 15, 8-10 mm long, flattened, bearing opposite bracts (2 mm long) at the apex, each bearing three branches; central branch c. 4 mm long, bearing a head of sessile, bacciform flowers c. 2V2 mm ; lateral branches VI 2-2 cm long with two opposite small bracts about the middle and ending in an involucre of minute rounded bracts around a terminal umbellule of c. 7-10 flowers; pedicels P/2-2V2 inm long. Calyx a minute rim. Corolla and stamens unknown. Ovary 3-4-celled. Ripe fruit unknown. Distr. Malesia: Philippines (Dinagat L). Note. A species clearly demonstrating the con- generity of Osmoxylon and Boerlagiodendron. 3. Osmoxylon simplicifolium (Elmer) Philipson, Blumea 23 (1976) 103. — Boerlagiodendron simplicifolia Elmer, Leafl. Philip. Bot. 7 (1914) 2329; Merr. En. Philip. 3 (1923) 224. Glabrous shrub, to 5 m, with numerous leaves clustered near the ends of the branches; petiole 5-7 cm, 2 mm wide, terete, with a small clasping base, an inconspicuous stipular ligule, and a single broad disk-like crest around the lower part of the petiole; blade simple, elliptic, base broadly cuneate, apex acute to apiculate, to 20 by 6'/2 cm, coriaceous, margin thickened, coarsely dentate, midrib prominent, principal nerves c. 8-10 mm apart. Inflorescence a terminal compound umbel, spherical, c. 7 cm 0; peduncle IV2-2 cm; primary rays c. 25-30, c. 1 cm long with two small obtuse bracts at the apex, ending in three branches; the central branch c. 6 mm long, bearing a subglobose umbel of c. 10-12 sterile bacciform flowers (2 mm 0), 3-4 mm pedicelled ; lateral branches c. 2-2^2 cm long, articulated about the middle, terminating in an umbel of c. 10-20 flowers; pedicels c. 2V2 mm. Calyx rim obsolete. Corolla 3-4-lobed, tubular below, 2 mm long. Stamens 3-4, exserted, 3 mm long. Ovary subcylindric, 2-4-celled, 1 mm long. Drupe spherical, c. 5 mm (dry), 2-4-ribbed when dry; surface of endosperm shallowly wrinkled. Distr. Malesia: Philippines (Mindanao: Agusan Prov., Cabadbaran). Ecol. On wind-swept ridge at 1750 m, on moss- covered soil with stones. Vern. Bolauanon, Mbo. Note. A wide-spreading shrub. Bark thick, yellowish, becoming grey. Wood soft, yellowish. Twigs repeatedly branched, the leafy portion sub- erect, leaves mostly ascending, rigidly coriaceous. Inflorescence branches green. Flowers orange, odourless. Berries becoming purple-black. 4. Osmoxylon spathipedunculatimi (Philipson) Philipson, Blumea 23 (1976) 103. — Meryta spathipedunculata Philipson, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 12. Glabrous tree, to 20 m, with spreading branches. Leaves crowded at the ends of the branches ; petiole to 14 cm with a small clasping base bearing a stipular ligule and a rim-like collar around the base of the petiole; blade obovate 17-30 by 10-15 cm, attenuate at the base, apex obtuse, margin entire, midrib prominent, secondary veins arched and uniting, 1^2-2 cm apart. Inflorescence a terminal compound umbel; primary rays c. 12, stout, com- pressed 9-17 cm long, bearing three branches at the apex; central branches and sterile flowers un- known; the two lateral branches 9-14 cm long with an articulation c. 1-2 cm from base, bearing helmet-shaped bracts which fall to reveal the terminal head of c. 12 flowers sessile on an ex- panded receptacle with an involucral rim c. 1 cm 0. Calyx rim 0. Corolla split into 5 lobes above, tubular below. Stamens 5. Ovary subcylindr.c, ± 1979] Araliaceae — I (Philipson) 35 9-celled ; disk raised in the centre to the pustulate stigmas. Drupes in a spherical head, globose, c. 12 mm 0, c. 9-ribbed when dry, crowned by the prominent, persistent, confluent stigmas. Distr. Solomon Islands (Bougainville and Guadalcanal). Ecol. Rain-forest, 800-1200 m. 5. Osmoxylon articulatum Philipson, Blumea 23 (1976) 103. Tree with stout branches, glabrous. Leaves well spaced towards the ends of the branches; petiole 10-1 3 cm, 2 mm broad, narrowly channelled above, with a small triangular base, bearing a short stipu- lar ligule {c. 2 mm long) and extending around the base of the petiole as a collar; blade obovate, to 27 by 9 cm, base narrowly cuneate, apex rounded or acute and shortly apiculate, margin thickened, remotely dentate towards the apex, midrib promi- nent, principal lateral veins c. l'/4-lV2 cm apart. Inflorescence a terminal compound umbel, almost sessile, saucer-shaped bracts caducous; primary rays c. 24, 8-11 cm long, flattened, c. 4 mm broad, bearing three branches at the apex; central branch 4-6 V2 cm long, the apex expanded and bearing an umbel of c. 10 sterile bacciform flowers c. 6 by 6 mm (when dry) apparently 1-celled, c. 9-14 mm pedicelled ; the two lateral branches c. 1 cm long at anthesis with an articulation c. 8-10 mm above the base, bearing helmet-shaped bracts which fall to reveal the terminal head of c. 15-18 flowers, sessile on an expanded receptacle with an involucral rim c. 6 mm 0. Calyx rim 0. Corolla split into c. 4 lobes above, tubular below, c. 2V2 rnm long. Stamens 5, exserted. Ovary subcylindric, c. IV2 rnrn long, 7-8-celled, disk with a pustulate central stigmatic boss. Drupes in a spherical head c. 2 cm (when dry), strongly 7-8-ribbed (when dry), c. 8 mm ; stigmas persistent, prominent. Distr. Malesia: Moluccas (Halmaheira: Ake Mumar to upper reaches of the Kakatua-matawe). Fr. Sept. Note. This plant is evidently similar to 6. O. umbelliferum described by Rumphius. However, a number of differences make it unlikely that it is the same species. The diameter of the inflorescence of the Halmaheira plant is only about half that given by Rumphius; the lateral rays of the inflores- cence are distinctly articulated near the base, a feature now shown in Rumphius' figure; and the sterile bacciform flowers are long-pedicelled, where- as Rumphius described and figured his as borne in capitula. 6. Osmoxylon umbelliferum (Lamk) Merr. Int. Rumph. (1917) 406; Philipson, Blumea 23 (1976) 104. — Pseudo-Sandalum amboinense Rumph. Herb. Amb. 2: 54, t. 12. — Aralia umbellifera Lamk, Encycl. 1 (1783) 225. — Hedera umbelli- ferum (Lamk) DC. Prod. 4 (1830) 262. — Gilibertia saururoides DC. I.e. 256. — Gastonia saururoides Roxb. [Hort. Beng. (1814) 90, nomen;] Fl. Ind. ed. Carey 2 (1832) 408 Csasuroides"). — O. amboi- nense MiQ. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 6, pp. — Pseudosandalum umbelliferum (Lamk) O. K. Rev. Gen. PI. 1 (1891) 271. According to Rumphius: Tree with stout trunk, the branches marked with prominent round leaf- scars. Leaves clustered at the ends of the branches, glabrous; petioles long; blades simple, lanceolate (30-36 by 10-12 cm), base rounded, apex acute, margin dentate. Flowers in large spreading umbels, the radiating branches tripartite, c. 30 cm long, each ending in a capitulum. Distr. Malesia: Moluccas. Infrequent on Am- bon, but said to be more numerous in Ceram and the Sula Islands. Only known from Rumphius' excellent plate and description; not yet re- collected in Ambon. Ecol. Evidently in forest in the hills of Ambon, and also planted at the time of Rumphius. Vern. Sasuru, Leytimor, tonokuku, Hitu. Notes. Valued for the perfume of its wood and foliage. Since this plant is known only from a description and a figure, some uncertainties remain as to its specific characters. In the description it is stated that the young leaves possess a few small teeth of which some signs remain on the older leaves. It is not clear whether the teeth are best developed on distinctive juvenile foliage, or whether the newly expanded normal foliage is intended. The leaves in the figure have prominent teeth, but as they are not shown associated with the inflorescence, they may be from a juvenile shoot. The description of the size of the flower buds is confusing, and it seems likely that sterile pseudo-fruits were mistaken for flower buds. Nevertheless, most characters of the plant are adequately portrayed and there can be no doubt that this species is distinct from the other simple-leaved species with a single, collar-like, petiolar crest. 7. Osmoxylon lanceolatum Philipson, Blumea 23 (1976) 104. — Fig. 13. Small tree with few branches, up to 16 m, glabrous. Many leaves clustered towards the ends of the branches; petiole 8-15 cm, terete, with a small triangular base, bearing a short stipular ligule {c. 2 mm long) and extending around the base of the petiole as a collar; blade oblanceolate, to 33 by 7V2 cm, base narrowly cuneate, apex acute or slightly apiculate, margin entire, midrib promi- nent, lateral veins arched ascending, c. 2-3 cm apart. Inflorescence a terminal compound umbel, almost sessile, saucer-shaped; bracts caducous; primary rays c. 15, c. 10 cm long, flattened, c. 4-5 mm broad, bearing three branches at the apex; central branch 5-6 cm long, the apex expanded and bearing a spherical head of c. 8-12 sessile, sterile, bacciform flowers c. 5 by 5 mm (when dry), 1-2-celled; the two lateral branches 36 Flora Malesiana [ser. I, vol. 9^ Fig. 13. Osmoxylon lanceolatum Philipson. Above: habit of inflorescence and leaves; belovi', left: twig showing collar-like crests at the base of the petioles ; below, right : the trifid branches of the inflorescence (New Ireland, Sands 795). 1979] Araliaceae — I (Philipson) 37 c. 5 cm long at anthesis, with an articulation immediately above the base, bearing two helmet- shaped bracts which fall to reveal the terminal head of c. 8-10 flowers, sessile on an expanded receptacle with an involucral rim c. 7 mm 0. Calyx rim 0. Corolla known only in bud, c. l^l^ mm long. Stamens c. 5. Ovary gibbous, c. 1 V2 mm high, 4-celled. Fruit unknown. Distr. Malesia: New Ireland (Namatanai Subdistr., Danfu R. area, inland from Manga). Ecol. Understorey tree in ridge top forest on limestone, 750-850 m. Note. The bark is pale grey, ± smooth with fine cracks. The twigs and cut branches are strongly aromatic. The wood is soft and dark straw- coloured. The central branches of the inflorescence rays are held ± horizontally or depressed and come to maturity before the lateral branches which are held erect. 8. Osmoxylon oblongifolium Philipson, Blumea 23 (1976) 105. Shrub c. 2 m, glabrous when mature, setulose on young parts. Leaves clustered at the ends of the branches; petiole to 16 cm, channelled above, 4 mm broad, with a clasping base prolonged upwards as a stipular ligule IV2-2 cm long, and with entire or fimbriate crests encircling the lower part of the petiole; blade simple, oblong-ovate, occasionally with a small triangular lobe on each side below the middle, to 46 by 17(-24) cm, base rounded to truncate, apex shortly acuminate, margin serrate, midrib prominent, principal lateral veins c. 3-4 cm apart (at broadest part of leaf). Inflorescence a terminal compound umbel, hemispherical, c. 13 cm 0; peduncle 3-4 cm, c. 6 mm wide, bearing fimbriate, lanceolate bracts (to 2 cm long) along its length and around and among the primary rays; primary rays c. 20, rather short and stout (16-20 by 3-4 mm) with small opposite caducous bracts at apex, each ray ending in three branches; the central branch c. 4 mm long; sterile flowers un- known; the two lateral branches VI2-4 cm long, articulated below the middle, terminating in a head of c. 20-30 sessile or subsessile flowers. Flowers unknown. Fruits crowded in a spherical head, drupes c. 5 mm (when dry) on pedicels c. 1 mm, 4-ribbed, pyrenes 4, cartilaginous; endosperm with faint reticulate ridging. Distr. Malesia: Philippines (Samar). Ecol. In dipterocarp forest, along creek bank, at 200 m. 9. Osmoxylon catanduanense (Merr.) Philipson, Blumea 23 (1976) 105. — Boerlagiodendron catanduanense Merr. Philip. J. Sc. 13 (1918) Bot. 318; En. Philip. 3 (1923)222. Shrub c. 1 m, glabrous except for parts of the inflorescence. Leaves clustered at the ends of the branches; petiole c. 30 cm, terete, 3 mm wide, base with a short ligule, and inconspicuous recurved crests; blade digitately compound, leaflets 7, chartaceous to subcoriaceous; petiolules 4-5 cm, the lateral shorter; leaflets lanceolate, mid-leaflet to 26 cm long, base cuneate, apex ± caudate; irregularly lyrately lobed, the sinuses reaching to within c. 8 mm of the midrib, lobes 2-4 cm long, patent, margin slightly thickened, entire or obscurely dentate. Inflorescence a terminal compound umbel, c. 10 cm 0, peduncle stout, bearing ovate bracts c. 1 mm long; primary rays c. 7-10, 2-2 V2 cm long, to 2 mm wide, minutely pubescent, with 2 broadly ovate bracts 4-5 mm long at apex ; central branch 3 mm or less, bearing a head of sterile flowers; lateral branches 2^2 cm long, bearing 2 broad bracts near the middle, and ending in a spherical head of c. 15 sessile flowers. Calyx an obsolete rim. Petals and stamens un- known. Ovary 4-celled. Fruit ovoid, 4-ridged when dry, 7 by 5 mm. Distr. Malesia: Philippines (Catanduanes). Ecol. On forested slopes, at c. 350 m. Note. Together with 10. O. caudatum and 11. O. heterophyllum this species forms a small group with leaves composed of lyrate leaflets. Although the foliage is similar (though not identical), the inflorescences are distinctive. In O. heterophyllum the rays are short, resulting in a compact compound umbel; in O. caudatum there are few rays (10 or fewer); in O. catanduanense there are many, relatively long rays, resulting in a large, diff"use compound umbel. Since O. catanduanense and O. caudatum are known only from the type collec- tions, the range of variation of these species is not known, but the inflorescence differences justify the retention of all three species. 10. Osmoxylon caudatum (Merr.) Philipson, Blumea 23 (1976) 105. — Boerlagiodendron caudatum Merr. Philip. J. Sc. 14 (1919) 440; En. Philip. 3 (1923) 222. Erect shrub, c. 2 m, becoming glabrous. Leaves clustered towards the ends of the branches ; petiole c. 45 cm, terete, striate, 4-5 mm wide, base with a stipular ligule c. IV2 cm long, and 2-3 fimbriate or pectinate crests; blade digitately compound, leaflets 5-9, subcoriaceous ; petiolules of the central leaflets to 5 cm long, lateral leaflets ± sessile; leaflets elliptic-lanceolate, mid-leaflet to c. 25 cm long, base decurrent on the slightly winged petio- lule, apex caudate-acuminate, the larger leaflets with 1-3 pairs of lyrate lobes reaching almost to the midrib, lobes ascending, margin slightly revolute with often prominent usually incurved teeth. Inflorescence a terminal compound umbel, c. 18 cm 0; peduncle stout, c. 2 cm, bearing few ligulate bracts c. VI 2-2 cm long, rough, with short setae on the back; primary rays c. 25, c. 4-5 cm long, glabrous and striate, subtended by ligulate bracts, similar opposite bracts at the apex, c. 1 cm long; central branch I-IV2 cm long, glabrous, ending in a whorl of obtuse rough coriaceous bracts 38 Flora Malesiana [ser. I, vol. 91 c. 3 mm long and a head (c. 1 cm 0) of sterile bacciform flowers c. 3 mm 0, on pedicels 3-8 mm; lateral branches 4-5 cm long, with opposite fimbriate bracts near the middle, bearing an ellipsoid head of c. 20 sessile flowers, each flower subtended and ± enclosed by an ovate fimbriate bract 3-4 mm long. Calyx an obsolete rim. Petals and stamens unknown. Ovary 2-2 V2 mm long, turbinate, obscurely angled, 4-celled. Fruit unknown. Distr. Malesia: Philippines (Luzon: Ilocos Norte Prov., Mt Palimlin). Ecol. On forested slopes near the summit, at c. 1000 m. Note. Apparently never re-collected. For a dis- cussion of distinctive features see under 9. O. catanduanense. 11. Osmoxylon heterophyllum (Merr.) Philipson, Blumea 23 (1976) 106. — Boerlagiodendron heterophyllum Merr. Philip. J. Sc. 9 (1914) Bot. 329; En. Philip. 3 (1923) 223. Erect tree, c. 5 m, glabrous. Leaves clustered at the ends of the branches; petiole to 25 cm, base clasping, prolonged as a stipular ligule to 2V2 cm long, and bearing several pectinate crests (becom- ing recurved when the bristles may be obscured or shed); blade to 35 cm long, digitately com- pound (or some leaflets imperfectly separated); leaflets 3-7, unequal in size, oblong-ovate to broadly oblong-oblanceolate, acuminate, lyrately lobed and irregularly dentate, the base gradually narrowed to the petiolule; petiolule up to 7 cm. Inflorescence a dense terminal compound umbel; peduncle stout, c. 2 cm, bearing many lanceolate bracts 2-3 cm long; primary rays c. 30, 10-15 mm long, subtended by lanceolate c. 2^2 cm long bracts, sometimes with bristles on the back, and bearing similar opposite terminal bracts 8-10 mm long, each ending in three branches; central branch short (not seen fully developed) terminating in an umbellule of c. 15 sterile bacciform flowers (c. 3 mm long), pedicels 3-4 mm ; lateral branches 8-10 mm long (? fully developed), articulation present ending in heads (c. 1 cm 0) of numerous flowers. Calyx rim obsolete. Corolla lobes 4, 2 mm long. Stamens 4. Ovary 4-celled. Fruit unknown. Distr. Malesia: Philippines (Samar, Biliran and Mindanao). Ecol. Primary forest, under shade near creek, 100-550 m. Vern. Arams, Bis., kayuang, Mbo, magusayag, C.Bis. Note. The description is partially based on the original publication as I have seen only immature inflorescences. For a discussion of distinctive features, see under 9. O. catanduanense. 12. Osmoxylon yatesii (Merr.) Philipson, Blumea 23 (1976) 106. — Boerlagiodendron yatesii Merr. Philip. J. Sc. 13 (1918) Bot. 44; En. Philip. 3 (1923) 225. Shrub, 1 m, glabrous, except for the inflores- cence. Leaves clustered towards the ends of the stout branches; petiole to 38 cm, channelled above, 5 mm 0, base with a stipular ligule 1 cm long, and 1-2 inconspicuous non-fimbriate crests around the back of the petiole; blade digitately compound, leaflets 5-7; petiolule 2-7 cm (the lateral shorter); blade elliptic-oblong to ovate, mid-leaflet to 23 by 8 cm, base gradually tapered, apex acuminate- caudate, margin dentate or somewhat undulate. Inflorescence a terminal compound umbel, sub- sessile or peduncle stout 1-3 cm, bearing few ovate bracts 1 cm long; primary rays 5-10, tomentose, c. 3 cm long and 3 mm wide, subtended by ovate bracts 6 mm long, similar opposite bracts at apex ; central branch 2-3 mm long, pubescent, ending in a whorl of blunt bracts (3 mm long) and an umbellule of c. 10 sterile flowers 4 mm 0, 2-4- celled, pedicels 3-8 mm; lateral branches 2-2 V2 cm long, pubescent, with small opposite bracts about the middle, bearing a head of 10-15 sessile flowers, subtended by ovate ciliate bracts. Calyx rim obsolete, sometimes fimbriate. Corolla 4-5- lobed, tubular below, 1 V2 mm long in bud. Stamens 4-5. Ovary 4-celled. Young fruit (Merrill, I.e.) shortly pedicelled (2-3 mm). Distr. Malesia: Philippines (Luzon and Catan- duanes). Ecol. In rain-forest and mossy forest, from low altitude to 1250 m. Vern. Magalayag, Dinagat. Note. The leaves are unlike any other Osmoxy- lon, resembling those of Macropanax or Schefflera. The flowers are described as yellow, and the fruit black. The only instance of a pseudo-fruit con- taining apparently normal seeds occurred in this species. 13. Osmoxylon geelvinkianum Becc. Malesia 1 (1878) 196; Philipson, Blumea 23 (1976) 106. — Eschweileria geelvinkiana (Becc.) Boerl. Ann. Jard. Bot. Btzg 6 (1886) 120. — Trevesia geelvin- kiana (Becc.) O. K. Rev. Gen. PI. 1 (1891) 272. — Boerlagiodendron geelvinkianum (Becc.) Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 32. — Eschweiler{i)a elegans Ridl. Trans. Linn. Soc. II, 9 (1916) 63. — Boerlagiodendron elegans (Ridl.) Harms, Bot. Jahrb. 56 (1920) 380. — Boerlagioden- dron stenolobum Harms, I.e. 382, f. 1 k-t. Glabrous shrub with few slender branches, up to 3 m high. Leaves clustered near the ends of the branches; petiole up to 22 cm, usually shorter, 2-3 V2 mm wide, narrowly channelled above, with a sheathing base prolonged as a membranous stipular ligule up to 4 cm or longer and with fimbriate or ± entire crests encircling the lower part of the petiole; blade up to 30 cm (usually 20 cm or less) very deeply 5-11-lobed, or with distinct digitately arranged leaflets, the lobes or 1979] Araliaceae — I (Philipson) 39 leaflets linear-lanceolate to lanceolate-obovate, entire or irregularly pinnatifid with narrow finely- tapering lobes, base gradually narrowed, apex narrowly caudate, margin serrate ; leaves below the inflorescence sometimes reduced to a single leaflet. Inflorescence terminal hemispherical, c. 12-20 cm ; peduncle 1 cm or less, with caducous lanceolate bracts mostly clustered below the primary rays, 1-2 cm long; primary rays rather few, spaced, 2V2-6 cm long, slender, bearing two caducous lanceolate bracts at the apex, up to IV2 cm long, each ray ending in three branches; the central branch c. 4-6 mm long bearing a whorl of lanceo- late caducous bracts and an umbel of c. 7-12 sterile, globose or ovoid bacciform flowers (c. 7 mm when dry) with pedicels c. 5 mm long and 6-9- celled; the two lateral branches c. 3-4 cm long at anthesis, articulated about the middle, terminating in a small head of 10-20 sessile or subsessile flowers. Calyx rim obsolete; corolla splitting into c. 4 irregular lobes above, tubular below, c. 2V2 mm long. Stamens 10-14, exserted, 3 mm long, anthers small. Ovary cylindric, c. 2 mm long, 10-16-celled; disk with a central raised boss formed by the pustulate stigmas. Fruit globose, fleshy (ribbed when dry), c. 10 mm 0. Distr. Malesia: New Guinea (Irian Jaya, to Sepik and Fly R. areas). Ecol. Primary forest, along creeks and river banks, flood-resistant, from near sea-level to 850 m. Vern. Amamutapu, Kamora, korinki, Ome, ida'pforpforsami, Kutubu. Note. The narrow leaf segments, almost or quite separated at their bases, are characteristic, even though variable in outline. The flowers are described as orange or reddish, and the soft fleshy fruits as dark purple, dark blue, or black. 14. Osmoxylon lineare (Merr.) Philipson, Blumea 23 (1976) 106. — Boerlagiodendron lineare Merr. Philip. J. Sc. 3 (1908) Bot. 253; En. Philip. 3 (1923) 223. Glabrous, erect shrub, c. 3 m. Leaves crowded near the ends of the branches ; petiole 4-6 cm, the base with a small stipular ligule (5 mm long) and a few fimbriate crests; blade to 20 cm 0, digitately compound with 4-7 leaflets (or very deeply divided into as many lobes); leaflets linear- lanceolate, c. 1-1 V2 cm wide, the base decurrent on the winged petiolule, apex attenuated, margin thickened, denticulate especially above. Inflores- cence a terminal compound umbel ; peduncle short (c. 1 cm), bracteate; primary rays c. 10, 2-2^/4 cm long, each ending in three branches; the central branch 4-5 mm long bearing a spherical head of numerous fimbriate bracts (sterile flowers fallen); the lateral branches 3-3 V2 cm long with a pair of minute fimbriate bracts about the middle, ending in a capitulum of c. 20 sessile flowers subtended by small fimbriate bracts, c. 1 mm 0. Calyx rim minute. Corolla and stamens unknown. Ovary 5-celled. Fruit ovoid c. 3 mm long, 5-ridged (when dry). Distr. Malesia: Philippines (Luzon). Note. Apparently never re-collected. The original specimen is without field information. The foliage is similar to that of 15. O. borneense but with shorter petioles and fewer leaflets. 15. Osmoxylon borneense Seem. J. Bot. 6 (1868) 141; Philipson, Blumea 23 (1976) 107. — O. helleborinum Becc. Malesia 1 (1878) 198. — Eschweileria helleborina (Becc.) Boerl. Ann. Jard. Bot. Btzg 6 (1887) t. 13. — Trevesia helleborina (Becc.) O. K. Rev. Gen. PI. 1 (1891) 272. — Boerlagiodendron helleborinum (Becc.) Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 31 . — Boerlagio- dendron borneense (Seem.) Merr. En. Bom. (1921) 456. Glabrous, spreading shrub, up to 3 m. Leaves clustered near the ends of the branches ; petiole up to 24 cm, narrowly channelled above, 2-3 (or 5) mm wide, with a sheathing base prolonged as a mem- branous stipular ligule up to 2^2 cm long, and usually with fimbriate, ± entire, or more rarely long-setose crests encircling the lower part of the petiole ; blade up to 20 cm digitately compound (or the bases of the leaflets joined by a very short web of tissue); leaflets 9-13, linear-lanceolate to lanceolate, gradually narrowed to the base and apex, up to 20 by 3 cm, usually much narrower, margin serrate, principle veins numerous, c. 5- 10 mm apart. Inflorescence terminal, hemispherical, c. 6-13 cm 0; peduncle 1-2 cm or shorter, with caducous lanceolate-ovate entire or fimbriate bracts mostly clustered below the primary rays, 1-1 V2 cm long; primary rays rather few (5-12), 2V2-4 cm long, with 2 ovate bracts at the apex, c. 8 mm long, each ray ending in three branches; central branch c. 5-12 mm, bearing an umbel of c. 6-16 sterile, globose, bacciform flowers, c. 3- 5 mm (when dry) with pedicels 4-5 mm long and 5-celled; the two lateral branches 1 V2-3 cm long at anthesis, articulated about the middle, terminating in a small head of c. 20-25 sessile flowers with minute rounded bracts. Calyx rim obsolete ; corolla splitting into few irregular lobes above, tubular below, c. VI 2-2 mm long; stamens 5-6, slightly exserted; ovary turbinate, angled, c. 1 mm long, 5-celled, disk with a central raised boss formed by the pustulate stigmas. Fruit globose, fleshy (ribbed when dry) c. 5 mm 0. Distr. Malesia: Borneo (Sarawak, Sabah and Kalimantan). Ecol. Characteristic of rocky river banks, not beyond flood-level, often in deep shade, from near sea-level to 950 m. Vern. Medong, Kayan, empasia abor, Iban, kayan, Tamang, koung, Kinabalu, bungor, Murut Bokan, salimpangaya, Murut Kalabakai. Notes. The leaves of some specimens of 13. O. geelvinkianum (New Guinea) resemble this 40 Flora Malesiana [ser. I, vol. 9^ species closely. This species can be distinguished by the more compact and smaller fertile flowers, and by its leaflets being uniformly unlobed. Both species are characteristic of river banks, and O. borneense has a low spreading habit, with the branches often rooting, resulting in extensive patches of this low shrub. It is a characteristic rheophyte confined to below flood-level. The flowers are described as greenish white or cream and the inflorescence branches are frequently dark purple. 16. Osmoxylon pulcherrimum Vidal ex F.-Vill. Nov. App. (1880) 102; Sinopsis Atlas (1883) 28, t. 55, f. B; Philipson, Blumea 23 (1976) 107. — Eschweileria pulcherrima (Vidal) Boerl. Ann. Jard. Bot. Btzg 6 (1887) 123. — Trevesia pulcher- rima (Vidal) O. K. Rev. Gen. PI. 1 (1891) 272. — Boerlagiodendron pulcherrimum (Vidal) Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 32; Merr. PhUip. J. Sc. 3 (1908) Bot. 254; En. Philip. 3 (1923) 224. — Boerlagiodendron sibuyanense Elmer, Leafl. Philip. Bot. 7 (1914) 2328; Merr. En. Philip. 3 (1923) 224. Erect, sparsely branched tree, up to 10 m, glabrous when mature, except for the inflorescence. Leaves crowded at the ends of the branches ; petiole to 1 m, channelled above, clasping base heavily lenticellate, prolonged as -a broad stipular ligule c. 2 cm long, usually with strong bristles on the back, and with strong long-pectinate crests encircling the base of the petiole; blade coriaceous, fan-shaped, c. 40 cm long, base broadly cuneate to truncate, palmately 7-11-lobed, lobes extending to within c. 12 cm from the base, lanceolate, coarsely serrate, sometimes irregularly lobulate, slightly narrowed towards the base, apex acute, sinuses rounded. Inflorescence a terminal compound umbel, c. 18 cm 0; peduncle very short, bearing heavily setose bracts ; primary rays 15-20, c. 3-4 cm long, 3^ mm broad, setulose, at the apex bearing opposite, ovate-lanceolate bracts 10-15 mm long, each ending in three branches; central branch c. 15-20 mm long, terminating in a globular head (c. 12 mm 0) of c. 20-30, sessile, sterile, bacciform flowers (3-4 mm 0) 3-celled, subtended by small ovate-lanceolate bracts; lateral branches c. 5V2 cm long (71/2 cm in fruit), with opposite bracts (c. 3-4 mm long) near the middle, terminating in a globose head of c. 40-50 sessile flowers, bracts between the flowers very small, setulose. Calyx rim obsolete. Corolla 4-lobed, tubular below, 2 mm long in bud. Stamens 4. Ovary 4-celled. Fruit globose c. 6-8 mm long, 4-ribbed (dry). Distr. Malesia: Philippines (Luzon, Mindoro and Sibuyan), recorded also from Formosa and Micronesia (Palau), cf. Kanehira, En. Micron. PI. (1935) 384. Ecol. Damp primary forests, 225-8(X) m. Vern. Cf. Merrill: paladukai, Bik., salapak, Neg.; cf. Elmer: palad-amok. Vis. Notes. The fan-shaped leaves with several nar- row lobes and prominent main veins resemble those of 34. O. eminens but are less strikingly developed. The inflorescence is considerably smaller with the pseudo-fruits forming a compact head borne on a comparatively long peduncle. The heads of true flowers, and of the fruits, are considerably smaller than those of O. eminens. Although Vidal's material is no longer avail- able, the figure and description relate well to later collections. The specimens on which Elmer based his Boerlagiodendron sibuyanense have the lobes of the leaf rather simpler in outline than is usual, but the fragments of young inflorescence are quite typical of the taxon and the name is reduced to synonymy. 17. Osmoxylon soelaense I*hilipson, Blumea 23 (1976) 108. A glabrous shrub or small tree. Petiole c. 32 cm, broadly channelled above, clasping base prolonged as a stipular ligule I'/a cm long, and bearing c. 3 fimbriate crests; blade c. 40 cm long, broadly cuneate at the base, palmately 7-lobed to within c. 12 cm from the base, lobes narrowly ovate to oblong-elliptic, slightly narrowed towards the rounded sinuses, apiculate, margin denticulate. Inflorescence a terminal compound umbel; peduncle c. 2 cm, bearing lanceolate bracts c. 2 cm long (similar smaller bracts subtend the primary rays); primary rays c. IV2 cm long, 5 mm wide, flattened, bearing opposite, terminal, persistent bracts 10-12 mm long, with lenticels and branched bristles on the back, ending in three branches; central branch c. 10 mm long terminating in a head 1 cm of 10-15 sessile sterile bacciform flowers (4 by 3 mm when dry; 3-celled) surrounded by an involucre of ovate bracts (3 mm long) and with minute bracts interspersed; lateral branches 4V2-5 cm long, bearing opposite ovate bracts (4 mm long) c. 8 mm above the base, terminating in a dense head c. 1 cm of 25-30 sessile flowers interspersed with inconspicuous obtuse fimbriate bracts. Calyx rim obsolete. Corolla 5-lobed above, tubular below. Stamens 5. Ovary 4-5-celled. Fruit unknown. Distr. Malesia: Moluccas (Sula Is.: Taliabu and Sulabesi). Note. For a discussion of the distinctive features, see under 31. O. sessiliflorum. 18. Osmoxylon globulare Philipson, Blumea 23 (1976)108. Shrub to 4 m, furfuraceous on the young parts. Petiole to 55 cm long, broadly channelled above, c. 1 cm wide, clasping base heavily lenticellate, prolonged as a stipular ligule 4 cm long sometimes scaly on the back, and bearing numerous irregular undulate crests on the base of the petiole often continued up the petiole, as rough fascicles of bristles as far as the blade ; blade 45 cm long, base 1979] Araliaceae — I (Philipson) 41 cordate or emarginate, with some bristles under- neath, palmately 7-lobed to within c. 10-15 cm from the base, lobes narrowly ovate to oblong- elliptic, slightly narrowed towards the broadly rounded sinuses, apex acute, margin serrate. Inflorescence a terminal compound umbel, spheri- cal, c. 15 cm 0; peduncle stout, 2-3 cm, bracts together with those among the primar>' rays caducous; primary rays 30-40, rigid only slightly flattened (subterete), 2-4 cm long, 2-2' 2 mm wide, bearing opposite bract-scars at the apex, ending in three branches; central branch 8-10 mm long, terminating in a head c. 13 mm Z of c. 20 sub- sessile sterile bacciform flowers (5 by 4 mm when dry, 2-3-celled); pedicels to 2 mm interspersed with minute bracts; lateral branches 3-4' 2 cm long, with the scars of opposite bracts near the middle, terminating in a dense head 1-1' 2 cm Z (in bud) of 20-30 sessile flowers interspersed with inconspicuous bracts. Calyx rim obsolete. Corolla 6-8-lobed above, tubular below, 2' 2 rnm long (in bud). Stamens 6-8. Ovary turbinate, obscurely ribbed, 5-8-celled (varying on the same plant). Fruit unknowTi. Distr. Malesia: Moluccas (Halmaheira, Moro- tai). Ecol. In forest from sea-level to 800 m. Said to be rare in Halmaheira but common in Morotai. Vern. Bungan-gutu, saha-sasate, Djailolo. Note. For a discussion of the distinctive features see under 31. O. sessiliflorum. 19. Osmoxylon kostermansii Philipson, Blumea 23 (1976) 108. Glabrous, small tree, 8 m. Leaves clustered near the ends of the branches; petiole up to 35 cm, narrowly channelled above, c. 3 mm broad, with a sheathing base prolonged as a stipular ligule c. 2 cm long, continued around the back of the leaf-base as a single wide crest with an entire recur\ed margin; blade up to 30 cm 2r, base cordate, deeply 5-7-lobed, lobes elliptic, slightly narrowed towards the sinuses and with a short acute apiculum, margin minutely serrate and sometimes with small sub-lobes, sinuses rounded. Inflorescence a terminal compound umbel, hemispherical, c. 14 cm Z at anthesis; peduncle c. 1 cm long, bearing small lanceolate bracts (c. 3 mm long) below and among the numerous (c. 20-24) primary rays ; primarv- rays 4-5 cm long and 1 mm broad, with opposite bracts (2 mm long) at the apex, each ending in three branches ; the central branch 5-6 mm long, bearing a spherical umbel of c. 20 small, sterile, bacciform flowers (2 mm Z) on pedicels c. 5-7 mm long, 2-celled; the two lateral branches c. 3'- 2-4 cm long, with no articulation or bracts except for a minute involucre around the terminal umbellule of c. 10-14 flowers; pedicels c. 2-3 mm. Calyx rim obsolete; corolla 4-lobed, 2 mm long in bud; stamens 4; ovary subcylindric, angled, c. 1 mm long, 5-7-celled, disk with a central stylar boss. Fruit spherical, c. 6 mm long, strongly ribbed when dry. Distr. Malesia: Borneo (Kalimantan: Sang- kulirang Distr., Mt Medadam). Ecol. On limestone at 450 m. Note. The foliage is similar to that of 22. O.pal- matum, except for the distinctive petiolar crest. The inflorescence is also similar to O. palmatum, especially in the lack of an articulation on the rays below the umbellules. 20. Osmoxylon barbatiun Becc. Malesia 1 (1878) 197; Philipson, Blumea 23 (1976) 109. — Esch- weileria barbata (Becc.) Boerl. Ajm. Jard. Bot. Btzg 6 (1886) 117. —Trevesia barbata (Becc.) O. K. Rev. Gen. PI. 1 (1891) 272. — Boerlagio- dendron barbatum (Becc.) Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 31. Small, glabrous tree. Leaves clustered near the ends of the branches; petiole to 32 cm, channelled above, 4-5 mm broad, with a sheathing base prolonged as a stipular ligule 2-3 cm long, and with several long setose crests encircling the lower part of the petiole; blade up to 43 cm 0, base cordate to truncate: deeply 5-7-lobed; lobes elliptic-lanceolate, narrowed towards the sinuses and tapered to an acuminate apex, margin minutely serrate, sinuses broadly rounded. Inflorescence a terminal compound umbel, hemispherical, to 12 cm Z; peduncle 1-2 cm; primary rays c. 20, 3-5 cm long, with three branches at the apex; the central branch c. 1 cm long, bearing a subglobose umbel of c. 15-20, sterile, bacciform flowers (3^ mm Z) on pedicels 5-7 mm long, 2^^celled; the two lateral branches about 4 cm long with no clear articulation but 1 or 2 obsolescent bracts, terminating in an umbellule of c. 15-20 flowers; pedicels c. 3-4 mm. Calyx rim obsolete; corolla 4-lobed above, tubular below, c. 4 mm long; stamens 7 or more, rarely fewer, exserted; ovary subcylindric c. 3 mm long, angled, 7-celled; disk with a central stigmatic boss. Fruit unknown. Distr. Malesia: SE. Moluccas (Key Is.). Note. Distinguished from the closely related 22. O . palmatum by the long-setose petiolar crests. For discrimination from 21. O. pfeilii see that species. 21. Osmoxylon pfeilii (Ware.) Philipson, Blumea 23 (1976) 109. — EschH-eileria pfeilii W.mib. Bot. Jahrb. 13 (1891) 396. — Boerlagiodendron pfeilii (Ware.) Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 32. A tree developing a crown when mature, up to 16 m, glabrous when mature, young pans slightly setulose. Leaves in terminal clusters; petiole up to 60 cm, channelled above, c. 6 mm broad, with a sheathing base prolonged as a membranous stipular ligule 2-^ cm long, and with many pro- nounced, long-setose crests encircling the lower part of the petiole; blade up to 75 cm Z, deeply 42 Flora Malesiana [ser. I, vol. 9^ 7-ll-lobed, base cordate or emarginate; lobes lanceolate to narrowly elliptic-oblong, narrowed towards the sinuses and tapered to an attenuate apex, margin sharply and remotely serrate, sinuses broadly rounded. Inflorescence a terminal com- pound umbel, hemispherical, to 20 cm ; peduncle short (2-3 cm) ; primary rays numerous (c. 30-40), 4-5 cm long, glabrous, with obsolete bracts at the apex, each ending in three branches; central branch c. 10 mm long, bearing a subglobose umbel of c. 20, small, sterile, bacciform flowers (1-1 V2 rnm 0) on pedicels c. 3-4 mm long, 2-5-celled; the two lateral branches c. 3-4 cm long, with no clear articulation but 1 or 2 obsolescent bracts, termi- nating in a head of c. 12-16 subsessile flowers (pedicel c. 1 mm, becoming longer in fruit) sur- rounded by an inconspicuous involucral rim. Calyx rim obsolete; corolla 5-lobed, 1 V2 nim long in bud; stamens 5; ovary subcylindric, angled, IV2 mrri long at anthesis, 5-16-celled, disk with a central boss formed by the united pustulate stigmas. Fruit spherical, fleshy, c. 8 mm 0, ribbed when dry, the stigmatic boss persistent and prominent; pyrenes cartilaginous. Distr. Malesia: Bismarck Archipelago (New Britain, Duke of York Group and New Ireland). Ecol. Primary rain-forest, from near sea- level to 600 m. Vern. Sare, sasare, sare a lauvolau. New Britain, Pomio; a ibalur. New Ireland. Notes. The bark is grey-brown and pustular, the wood straw-coloured and soft. The flowers are orange, and the ripe fruit dark red-violet. In the original description the ovary is recorded to possess 10-14 cells. However, some other specimens have as few as 5 cells in the ovary, but in other respects agree with specimens with the large number of seeds. Since the inflorescence, leaf-shape, and especially the nature of the petiolar crests, as well as the distribution, are all highly distinctive within the genus, all the specimens can be accepted as examples of one species with a highly variable number of carpels. This species is very close to 20. O. barbatum of the Key Islands. The original diagnostic character of the number of cells in the ovary has been found to be unreliable. However, since the primary rays in the inflorescence are more numerous and the pedicels of the fertile and sterile flowers are shorter this geographically distinct species is maintained. 22. Osmoxyion palmatum (Lamk) Philipson, comb, nov. — Folium polypi mas {et femina?) Rumph. Herb. Amb. 4: 101, t. 43. — Aralia palmata Lamk, Encycl. 1 (1783) 224, type, non Lour. 1790, nee R. & S. 1820. — Trevesia moluccana MiQ. Fl. Ind. Bat. 1, 1 (1856) 748; Bonplandia 4 (1856) 137. — Trevesia zippeliana MiQ. Ann. Mus. Bot. Lugd.- Bat. 1 (1863) 11. — Unjala bifida Reinw. ex de Vriese, pi. Ind. Or. (1867) 83, nomen in synon.; ex BOERL. Ann. Jard. Bot. Btzg 6 (1887) 166, in synon. — O. moluccanum (MiQ.) Becc. Malesia 1 (1878) 195; Philipson, Blumea 23 (1976) 109. — O. zippelianum (MiQ.) Becc. Malesia 1 (1878) 195. — Eschweileria palmata Zipp. ex Boerl. Ann. Jard. Bot. Btzg 6 (1887) 116, t. 14. — Boerlagio- dendron palmatum (Zipp. ex Boerl.) Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 31 ; Merr. Int. Rumph. (1917) 407. — Boerlagiodendron moluc- canum (MiQ.) Bakh./. & Ooststr. in Back. Bekn. Fl. Java (em. ed.) 7 (1948) fam. 159, p. 3; Blumea 6 (1950) 367 ; Back. & Bakh./. Fl. Java 2 (1965) 163. Small, glabrous tree to 15 m. Leaves clustered near the ends of the branches ; petiole up to 40 cm, channelled above, 4-5 mm broad, with a sheathing base prolonged as a stipular ligule up to 5 cm long, and with several fimbriate crests encircling the lower part of the petiole ; blade up to 45 cm 0, base cordate, deeply 5-9-lobed, lobes elliptic, slightly narrower towards the sinuses, acuminate, margin serrate and sometimes with small sub-lobes, sinuses rounded. Inflorescence a terminal, compound umbel, subspherical, to c. 20 cm at anthesis; peduncle c. 1-2 cm, with small caducous bracts below and among the numerous (20-60) primary rays; primary rays 3-5 cm long, 2-3 mm broad, with two opposite caducous bracts at the apex, each ending in three branches; central branch 6-10 mm long, bearing a spherical umbel of c. 20-30 small, sterile, bacciform, flowers (2-4 mm 0) on pedicels 4-6 mm long, 4-7-celled; two lateral branches c. 2^2-5 cm long, rigid and straight, with no articulation (indistinct scars of bracts may occur below the involucre), bearing a minute involucre around the terminal umbellule of c. 12-20 flowers; pedicels to c. 5 mm (occasionally flowers subsessile). Calyx rim obsolete. Corolla irregularly 4-5-lobed above, tubular below, c. 5 mm long. Stamens 6-9, exserted, filaments stout. Ovary subcylindric, angled, c. 2 mm long, 6-9-celled, disk with a raised central stigmatic boss. Fruit globose, fleshy, c. 10 mm 0, strongly ribbed when dry. Distr. Malesia: Celebes (once, not localized) and Moluccas (Buru, Ceram, Ambon, Banda, Tenimber Is.). Also cultivated in the Bogor Botanic Garden. Ecol. An understorey tree in primary rain- forest. Uses. The leaves are used for culinary and medi- cinal purposes (against gonorrhoea). Vern. Daun gurita, pelenda darat, saha-saha, Moluccas, fumala-alas, Tenimber Is. Note. The spherical inflorescence is characteris- tic, having straight rigid rays with no articulation on the secondary branches, and the pseudo-fruits are well separated from the true flowers. Miquel distinguished Trevesia zippeliana because the collector noted that its ovary was 4-celled. Possibly this number related to the sterile flowers. In two gatherings from Ambon (Waai, Teysmann; G. Salhutu, Boerlage 179) the fertile flowers are 1979] Araliaceae — I (Philipson) 43 subsessile in heads, but otherwise conform to the characters of this species. The only record of this genus from the Tenimber Is. consists of leaves only, but their characters conform to this species. 23. Osmoxylon ramosii (Merr.) Philipson, Blumea 23 (1976) 110. — Boerlagiodendron ramosii Merr. Philip. J. Sc. 11 (1916) Bot. 27; Merr. En. Philip. 3 (1923) 224. Erect, unbranched, or sparingly branched, 4 m high shrub, becoming glabrous. Petiole to 40 cm, the clasping base prolonged as a stipular ligule c. 2 cm long, and with few to several prominent recurved, obscurely fimbriate, crests surrounding the lower part of the petiole; blade to 30 cm long, base emarginate, palmately 3-7-lobed, lobes extending to within 3-8 cm from the base, oblong- ovate, margin serrate, sometimes lyrately lobulate, apex acuminate, sinuses broadly rounded. Inflores- cence a terminal compound umbel, 10-15 cm 0; peduncle stout, with lanceolate bristle-bearing bracts; primary rays c. 15, 2-3 cm long, 2-3 mm wide, flattened, subtended by lanceolate bracts c. 2 cm long, with bristles on the back and bearing similar opposite terminal bracts c. 1 cm long, each terminating in three branches; central branch c. 4 mm long, slightly pubescent, terminating in an umbellule (c. IV2 cm 0) of c. 10-15 sterile, bacci- form flowers 4—5 mm 0, 2-celled, pedicels 5-8 mm long, subtended by caducous bracts; lateral branches 2^2 cm long (slightly longer as fruits form), without any articulation or bracts except for a caducous small involucre around the terminal head of c. 25-35 sessile or very short-pedicelled flowers, c. 1 cm 0, bracts among the flowers obscure. Calyx rim obsolete. Corolla 4-lobed above, tubular below. Stamens 4, exserted, filaments broad. Ovary subcylindric, obscurely angled, 4-celled. Fruit spherical 9 mm 0, 4-ribbed when dry. Distr. Malesia: Philippines (Luzon). Ecol. On low-lying, wet ground, in forest, or on forested slopes, 700-800 m. Notes. The flat-toppjed inflorescence is c. 15 cm with much-reduced leaves below it. The in- florescence rays are dark purplish and the flowers orange-yellow. The bark is grey and the wood soft. This is the only species in the Philippines without opposite bracts on the lateral branches of the inflorescence rays. In this respect it resembles 22. O. palmatum and a few other species. 24. Osmoxylon novoguineense (Scheff.) Becc. Malesia 1 (1878) 197; Philipson, Blumea 23 (1976) 110. — Trevesia novo-guineensis Scheff. Ann. Jard. Bot. Btzg 1 (1876) 26. — Eschweileria novoguineensis (Scheff.) Boerl. ibid. 6 (1886) 118. — Boerlagiodendron novoguineense (Scheff.) Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 31; Back. & Bakh. /. Fl. Java 2 (1965) 163. — Boerlagiodendron lauterbachii Harms in K. Sch. & Laut. Fl. Schutzgeb. (1900) 484. — Fig. 11, 14. Fig. 14. Osmoxylon novoguineense (Scheff.) Becc. Petiolar base with ligiile and collar-like crests (Photogr. Philipson, Wantoat, 1968). 44 Flora Malesiana [ser. I, vol. 9^ Tree or shrub, unbranched or sparingly branched, up to 16 m, the young parts rufous- furfuraceous, glabrescent. Large leaves forming terminal crowns; petiole up to 1 m, stout (1-2 cm broad), flattened above, with a sheathing base prolonged as a strong stipular ligule up to 7 cm long, and with fimbriate crests encircling the lower part of the petiole; blade up to 1.20 m 0, with 5-7 strong ribs radiating from the top of the petiole, deeply lobed almost to the base of these ribs, the lobes in turn deeply lobed and incised, the central lobes especially being strongly pinnatisect or digitately tripartite, apices acute, margins serrate; upper leaves associated with inflorescences may be smaller, more simply lobed, or entire. Inflorescence terminal, a large compound umbel, bowl-shaped, up to 35 cm ; peduncle up to 10 cm, stout, with lanceolate caducous bracts (c. 4 cm long) below and among the numerous (c. 50-70) primary rays ; primary rays c. 12-15 cm long at anthesis, c. 3 mm 0, bearing two caducous bracts (1 cm long) at the apex, each ray ending in three branches; central branch c. 2 cm long, bearing an umbel of c. 20-40, sterile, bacciform flowers (c. 6 mm when dry) on pedicels c. 10 mm, and 2-6-celled; the two lateral branches c. 4—6 cm, with two opposite or sub- opposite bracts about the middle, terminating in a subspherical umbel 2^2-3 cm of 30-50 flowers on pedicels c. 8-10 mm long. Calyx rim obsolete, undulate. Petals with irregular erect lobes, tubular below. Stamens 6-10 exserted. Ovary turbinate somewhat angled; glabrous, 6-14-celled; disk flat with a central double row of pustulate stigmas. Fruits on stiff" radiating pedicels, ovoid or spherical, fleshy, ribbed when dry. Distr. Solomon Is.; in Malesia: throughout New Guinea and in the Bismarck Archipelago. Ecol. Primary and second-growth forest, from sea-level to 1600 m. Vern. Lebe, Mooi, teresakui, Manikiong, akriek, Biak, hoppung, Hottam, uger, Wagu, faliifalii, Tifal, ap gan dandam, aimaini, Mamig, ida'pfopforsami, ¥^\iinhu,pulaka. Gazelle Peninsula. Notes. The foliage is similar to that of 30. O. boerlagei, but the pedicelled flowers of O. novo- guineense distinguish it readily from that species. The ripe fruits are usually ovoid, but in the Solomon Is. they are characteristically spheroidal, and this feature recurs in some specimens from the Bis- marck Archipelago and the adjacent coast of New Guinea. The fawn bark is pustulate with many lenticels. An orange exudate flows from the cut stems. The wood is soft and straw-coloured. The inflorescence branches are dark purple, the corolla and stamens usually deep red, and the ripe fruit shining purple or blue-black. 25. Osmoxylon teysmannii (Boerl.) Philipson, Blumea 23 (1976) 111. — Eschweileria teysmannii Boerl. Ann. Jard. Bot. Btzg 6 (1887) 119. — Trevesia teysmannii (Boerl.) O. K. Rev. Gen. Pl. 1 (1891) 272. — Boerlagiodendron teysmannii (Boerl.) Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894)31. A small, glabrous tree, 6 m. Leaves clustered at the ends of the branches; petiole to 40 cm, chan- nelled above, 4 mm broad, with a sheathing base prolonged as a stipular ligule 2-2 V2 cm long, and with several fimbriate or entire crests on the lower part of the petiole; blade c. 30 cm 0, membranous, cordate at the base, deeply 7-lobed, lobes elliptic, slightly narrowed to the broadly rounded sinuses, narrowed to a fine apiculum at the apex, margin finely serrulate. Inflorescence a terminal compound umbel, c. 10 cm ; peduncle c. 1 cm, bearing ovate bracts (ligules of reduced leaves) and terminating in a cluster of bracts (c. 10 mm long) below and among the primary rays; primary rays c. 12-15, c. 3-4 cm long, with a pair of lanceolate bracts at the apex (c. 1 cm long) ; central branch c. 1 cm long, terminating in an umbellule of c. 5-8 sterile bacci- form flowers (c. 4 mm when dry) on pedicels 6-9 mm long interspersed with linear bracts 5 mm long; two lateral branches c. 3V2 cm long, articu- lated about the middle, terminating in an umbellule surrounded by caducous linear bracts (leaving a rim-like involucre); flowers c. 8-12, pedicels c. 2-3 mm. Calyx rim obsolete. Corolla 3 mm long, with 7-8 lobes above, tubular below. Stamens 7-8, filaments stout, anthers exserted. Ovary sub- cylindric, angled, c. IV2 rnm long, 7-8-celied; disk flat, with a central stigmatic boss. Fruit unknown. Distr. Malesia: SW. Celebes (Tjamba, Kosali- Porema) and NW. Central Celebes (Palu-Parigi and Mt Nokilalaki). Ecol. In rain-forest, 800-1000 m.. 26. Osmoxylon humile (Elmer) Philipson, Blumea 23 (1976) 111. — Boerlagiodendron humilis Elmer, Leafl. Philip. Bot. 7 (1914) 2327; Merr. En. Philip. 3 (1923) 223. Erect, small, sparsely branched shrub, up to IV2 m. Leaver clustered at the ends of the branches; petiole to 25 cm, terete, base prolonged as a stipular ligule c. 1 cm long, and with c. 3 entire crests at the base; blade palmately 5-lobed, 24 cm long, base truncate or cordate, lobes reaching to within 3-6 cm from the base, elliptic, 4-6 cm wide, narrowed towards the broadly rounded sinuses, tapered to an acute apiculum, margin serrate in the upper part, the outer lobes with a lobule on the lower edge. Inflorescence a terminal compound umbel, 9 cm 0, subtended by a few foliaceous bracts ; peduncle stout, 2-3 cm, with furfuraceous, oblong bracts; primary branches crowded, numer- ous, 2V2 cm, furfuraceous, flattened, striate with opposite minute bracts at the apex ; central branch c. 3 mm, bearing an umbellule of sterile, bacciform flowers 3V2 nim 0, pedicels 4'/2 mm; lateral branches 3-3V2 cm, articulated about the middle, terminating in an umbellule of c. 15 flowers, bracts 1979] Araliaceae — I (Philipson) 45 inconspicuous, fimbriate, pedicels IV2-2 mm. Calyx an obsolete rim. Corolla and stamens not known. Ovary VI 2-2 mm, 4-celled, with a flat disk and a raised central stigmatic boss, 4-celled. Fruit 6 by 4 mm (dry) 4-ribbed; pyrenes crusta- ceous; endosperm rugose. Distr. Malesia: Philippines (Mindanao). Ecol. Damp fertile ground in dense forest, on south side of Baruring R., at 1000 m. Vern. Sardng-ka-mdno, Bag. 27. Osmoxylon pectinatum (Merr.) Philipson, Blumea 23 (1976) 111. — Boerlagiodendron pectinatum Merr. Philip. J. Sc. 3 (1908) Bot. 253, 424; En. Philip. 3 (1923) 224; Kanehira, Form. Trees rev. ed. (1936) 520, f. 480; Hui-LiN Li, Woody Fl. Taiwan (1963) 666, f. 273. Shrub or small glabrous tree up to 8 m. Leaves clustered at the ends of the branches; petiole to 18 cm, with a clasping base prolonged as a short acute stipular ligule, and with several basal crests fringed with 1-2 cm long bristles ; blade to 25 cm 0, base truncate to broadly cuneate, palmately 5-7- lobed, lobes reaching to about the middle of the lamina, sinuses narrow-rounded, lobes oblong- elliptic, usually slightly narrowed below, obtuse to acute, margin thickened, coarsely dentate, coria- ceous. Inflorescence a terminal compound umbel; peduncle c. 1 cm, with small bracts (3 mm) below and among the primary rays; primary rays c. 25- 35, c. 2-3 cm long, with opposite ovate caducous bracts at the apex, each terminating in three branches; central branch c. 8-11 mm long, ending in an involucre of minute bracts (1 mm) surround- ing an umbellule of c. 15-20 ovoid sterile flowers (c. 3 mm long, 3-celled), pedicels 5-6 mm long; lateral branches c. 2V2 cm long at anthesis, with an articulation about the middle, ending in an umbel- lule c. 1 cm with minute fimbriate bracts; flowers c. 30, pedicels c. IV2 mm (elongating slightly in fruit). Calyx rim obsolete. Corolla lobes 4-5, tubular below, 2 mm long. Stamens 4-5. Ovary turbinate, IV2 mm long, 4-6-celled. Fruits globose, 5 by 5 mm (dry), 4-6-ribbed when dry. Distr. Taiwan (Botel Tobago and Lutao I., east off Taiwan proper); in Malesia: N. Philippines (Batan I.). Ecol. Forested slopes at 650 m. Vern. Narapan, Iv. 28. Osmoxylon masarangense Philipson, Blumea 23(1976) 111. Small tree, 5 m, the young parts setulose, becoming ± glabrous. Leaves in terminal clusters; petiole c. 17 cm, rather narrow (IV2-2 mm 0), channelled above, with a small clasping base, prolonged as a stipular ligule, 1-1 V2 cm long, setulose on the back, and with a number of long- setulose crests encircling the lower part of the petiole; blade c. 18 by 22 cm, deeply 3-5-lobed (or below the inflorescence sometimes simple), the base truncate or emarginate, lobes oblong or elliptic, slightly narrowed to the broadly rounded sinuses, apex with a short apiculum, membranous, margin finely setulose-serrate, sinuses c. 6 cm frob- base of the blade. Inflorescence a terminal sum sessile compound umbel; primary rays c. 10, setulose, P/4-2 cm long, each ray ending in three branches; central branch 4-5 mm long, ending in an umbellule, pedicels 6 mm, sterile flowers not known; two lateral branches 2-2 V2 cm long, 1 mm broad, with two bract scars about the middle but usually not opposite, terminating in an umbellule with c. 10 pedicels 3-5 mm long (in fruit). Flowers unknown. Fruit (when dry) ovoid, 6 by 4 mm, 5-seeded. Distr. Malesia: N. Celebes (Minahasa, Tomo- hon, Mt Masarang). Ecol. Secondary forest at edge of crater lake, at 1200 m. Note. This species is similar in aspect to the Philippine 39. O. triiobatum, but the petiolar crests are distinctive. 29. Osmoxylon insidiator Becc. Malesia 1 (1878) 195; Philipson, Blumea 23 (1976) 112. — O. carpo- phagarum Becc. Malesia 1 (1878) 196. — Esch- weileria insidiatrix (Becc.) Boerl. Ann. Jard. Bot. Btzg 6 (1886) 120. — Eschweileria carpophagarum (Becc.) Boerl. I.e. 121, t. 15. — Trevesia insidiator (Becc.) O. K. Rev. Gen. PI. 1 (1891) 272. — Trevesia carpophagarum (Becc.) O. K. I.e. — Boerlagiodendron insidiator (Becc.) Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 32. — Boerlagiodendron carpophagarum (Becc.) Harms, I.e. — Boerlagio- dendron pachycephalum Harms, Nova Guinea 8 (1910)271. Small tree to 12 m, young parts with uniform scurfy tomentum. Large leaves forming terminal crowns; petiole up to 80 cm, stout (1-2 cm 0), broadly channelled above, with a sheathing base prolonged as a strong stipular ligule c. 9 cm long, and with moderately developed irregular (not fimbriate) crests encircling the lower part of the petiole; blade up to 85 cm 0, with 5-7 strong ribs radiating from the top of the petiole, deeply lobed almost to the base of these ribs, lobes in turn deeply lobed and incised, the median often digitately tripartite, apices long acuminate, margin irregu- larly and remotely serrate, subglabrous when mature or showing remnants of the tomentum. Inflorescence a terminal hemispherical compound umbel, c. 15 cm high by 30 cm wide; peduncle short, stout (IV2-2 cm 0), with lanceolate bracts 4-6 cm long below and among the numerous (15-20) primary rays; primary rays 6-10 cm long, c. 5-12 mm wide, rigid, bearing 2 lanceolate bracts (2-3 cm long) at the apex, each ray ending in three branches; central branch c. IV2 cm long, bearing an umbel of c. 30 sterile bacciform flowers (c. 5- 12 mm when dry), the pseudo-fruits and their pedicels ± rufous tomentose, pedicels 10-12 mm, 46 Flora Malesiana [ser. I, vol. 9^ and 6-celled, surrounded by an involucre of short ovate bracts (3-8 mm long); two lateral branches c. 6 cm at anthesis, rigid, slightly flattened, to 8 mm broad, bearing a pair of bracts (c. IV2 cm long) about the middle, terminating in a subglobose head 3V2-4 cm of c. 30-40 sessile flowers, and surrounded by an involucre of ovate bracts c. 10-14 mm long. Calyx rim fimbriate. Petals irregularly 4-5-lobed, 7-8 mm long, connate below to form a fleshy tube, pubescent on the outer surface. Stamens c. 15-26, filaments strap- like, projecting beyond the corolla, anthers c. 4 mm long. Ovary shortly turbinate, 2-4 mm long, angled, furfuraceous, c. 13-25-celled; disk flat, with a central boss formed of the pustulate stigmas. Fruits in a compact spherical head, the individual drupes angled by mutual pressure, and bearing the persistent stigmas on the exposed face, c. 10-14 mm long, the numerous pyrenes compressed and flat; cartilaginous. Distr. Malesia: throughout New Guinea, but local ; also Waigeo I. Ecol. Primary rain-forest and regrowths, frequently beside streams, from sea-level to 350 m. Vern. Angit, kangit, Waigeo, pennifogo, Orakawa, Papua. Notes. The bark is greyish brown, slightly fissured with many lenticels. The wood is soft and white. The flowers are reddish-brown or purple, with orange-red filaments and the fruit purple. Beccari provided a detailed description of the living plant, and noted that the fruits are eaten by various species of pigeon. Boerlagiodendron pachycephalum Harms has very strongly developed umbels and leaves, but apart from size, it does not differ from this species. Since a range in stature is shown by the several gatherings now available, the whole is best re- garded as a single species. Similarly, the very short pedicels of O. carpo- phagarum, which Beccari used to distinguish it from O. insidiator, can in fact be matched on several specimens of that species. 30. Osmoxylon boerlagei (Warb.) Phujpson, Blumea 23 (1976) 112. — Eschweileria boerlagei Warb. Bot. Jahrb. 13 (1891) 395. — Boerlagioden- dron warburgii Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 32, nom. illeg. superfl.; in K. Sch. & Laut. Fl. Schutzgeb. (1900) 484. — Boerlagiodendron boerlagei (Ware.) Harms, Bot. Jahrb. 56 (1920) 382. — Fig. 15. Small to fairly large tree, unbranched or sparingly branched, up to 24 m, glabrous, at least when mature. Large leaves forming terminal crowns; petiole up to 1 m, stout (1-2 cm 0), Fig. 15. Osmoxylon boerlagei (Ware.) Philipson. Showing that each ray ends into a central umbel of bacciform flowers and two lateral umbels with normal flowers (Photogr. Philipson, Kassam Pass, E. New Guinea, 1968). 1979] Araliaceae — I (Philipson) 47 broadly channelled above, with a sheathing, heavily lenticellate base prolonged as a strong stipular ligule up to 7 cm long, and with lacerate crests encircling the lower part of the petiole; blade up to 1 . 1 5 m , with 5-7 strong ribs radiating from the top of the petiole, deeply lobed almost to the base of these ribs, lobes in turn deeply lobed and incised, the central lobes especially being strongly pinnatisect or digitately tripartite, apices acute, margin undulate or indistinctly serrate. Inflores- cence terminal (or overtopped by a lateral leafy branch), a large compound umbel, bowl-shaped with a slightly convex top, up to 60 cm : peduncle c. 10 cm, stout, with lanceolate bracts below and among the very numerous radiating primary rays; outer primary rays c. 20 cm long at anthesis (elongating in fruit), inner rather shorter, woody, bearing two caducous bracts at the apex, each ray ending in three branches; central branch c. 4 cm, bearing an umbel of c. 20 sterile bacciform flowers {c. 8 mm when dry) with rigid pedicels c. VI2 cm long, and 5-6-celled; two lateral branches c. 9 cm long at anthesis, articulated about the middle, terminating in a button-like head of c. 20-30 sessile flowers and surrounded by ovate bracts which soon fall leaving a bowl-shaped involucre, c. IV2 cm 0. Calyx rim obsolete. Petals c. 13, bud flat-topped, angled, minutely pubescent, apparently falling as a calyptra. Stamens 8-13. Ovary shortly turbinate, angled, glabrous, 10-14-celled; disk flat, with a central double row (or ellipse) of pustulate stigmas. Fruits spreading to form a ± spherical head obscuring the involucre, each c. 9 by 7 mm (when dry) with prominent persistent stigmas. Distr. Malesia: throughout New Guinea. Ecol. Primary forest, and secondary growths on old cultivations, from near sea-level to 1800 m. Vern. Eunya, Gimi, apiatambay, Washkuk, ma-korr-korr, Jal, teresahui, Manikiong. Note. Bark yellow grey with shallow fissures and many pale corky pustules; wood straw- coloured, fibrous. The large terminal inflorescence is shallowly convex on top and is surrounded by several large leaves. Flowers reddish brown. Fruit purplish black and succulent. 31. Osmoxylon sessiliflonim (Laut.) Philipson, Blumea 23 (1976) 113. — Boerlagiodendron sessiliflorum Laut. Nova Guinea 8 (1910) 272. Small tree, up to 18 m, glabrous when mature, or tomentum persistent on the inflorescence. Leaves crowded at the ends of the branches; petiole to 60 cm, with the clasping base prolonged as a stipu- lar ligule up to 6 cm long, and with few to several strong or weak crests around the base of the petiole, margin undulate or fimbriate ; blade to 50 cm long, base cordate, palmately 5-9-lobed, lobes extending to near the base, elliptic, coarsely serrate, often irregularly lobulate, apex acute, sinuses rounded. Inflorescence a terminal compound umbel; peduncle to c. 4 cm, bearing lanceolate bracts to 2' '2 cm long, caducous or persistent, occasionally with some bristles on the back, primary rays c. 20-30, c. 9 cm long, - pubescent, bearing opposite caducous or rarely persistent bracts at the apex; central branch variable in length (2-18 mm), bearing an umbellule or head 1-2' 2 cm of sterile bacciform flowers (4-5 mm 0), 4-8-celled, subtended by minute bracts, pedicels variable in length (5-18 mm); lateral branches 3-7 cm, articu- lated near the middle, terminating in a globose head of c. 20-30 sessile flowers (or pedicels IV 2 mm long), bracts between the flowers very small. Calyx rim obsolete. Corolla few- to many-lobed, tubular below, 1','2^ mm long in bud. Stamens 6-17. Ovary 5-18-celled. Fruit a globose head of drupes; drupes c. 10 by 6 mm, obovoid, ribbed when dry. Distr. Malesia: throughout New Guinea. Ecol. Rain-forest, especially along the muddy banks of rivers, from sea-level to 100 m. Vern. Akriek, Biak, korinki, Orne, kwita-kwita, Milne Bay, sapi-ai, Jense, terrasahui, Manikiong. Notes. Unbranched or sparsely branched with crowns of large leaves. The bark is light brown and the wood cream. The inflorescence branches are purple, the flowers red, and the succulent ripe fruits black. The variation in the numbers of floral parts is considerable. Most specimens have more than 10 stamens and the same number of cells in the ovary, or more. Three gatherings have from 5-7 stamens and cells. These may possibly require to be segregated as a distinct species, but other evidence to support this course is lacking. An even more dis- tinctive gathering has central branches to 3 cm long with the pseudo-fruits on short pedicels (less than 5 mm) and flowers with 17 stamens and 25 cells in the ovary. These characters have not been included in the specific description as this specimen is only very tentatively referred to this species. This species forms an eastward extension of a complex of species, represented in the Moluccas by 37. O. talaudense, 17. O. soelaense and 18. O. glo- bulare. "Several similar species occur in the Philip- pines. They are characterized by dense spherical heads of flowers. Osmoxylon talaudense resembles some specimens of O. sessiliflorum rather closely, and the difficulty of preserving the characters of these large-leaved plants in an herbarium probably obscures several good diagnostic features. The most reliable charac- ter to distinguish these two species is the position of the articulation on the lateral branches of the inflorescence rays. In specimens from the Talaud Is. this is close to the base (below the apex of the central umbellule of pseudo-fruits) and the bracts are persistent, whereas in the New Guinea material it is near (or above) the middle, and is usually marked by two inconspicuous scars which fre- quently are not opposite. In both species the pseudo-fruits are pedicelled, whereas in the other 48 Flora Malesiana [ser. I, vol. 9^ two Moluccan species the pseudo-fruits are sessile or subsessile forming spherical heads. In O. globu- lare (from Morotai and Halmaheira) the lateral branches are rigid and only slightly flattened with the articulation near the middle and the bracts caducous. In O. soelaense the articulation is much nearer the base, the bracts are persistent, and the branches are broader and much flattened. 32. Osmoxylon camiguinense (Merr.) Philipson, Blumea 23 (1976) 113. — Boerlagiodendron camiguinense Merr. Philip. J. Sc. 3 (1908) Bot. 252; En. Philip. 3 (1923) 222. Shrub to 2 m, glabrous except for the inflores- cence. Leaves clustered at the ends of the branches; petiole to 18 cm, with a sheathing base prolonged as a stipular ligule 1 cm long, and with several entire crests encircling the lower part of the petiole ; blade 20 by 22 cm, base truncate, 3-5-lobed to about the middle, lobes oblong, scarcely nar- rowed towards the base, apex acuminate, sinuses broad, rounded, margin dentate, coriaceous. Inflorescence a terminal compound umbel ; primary rays c. 2-2^2 cm, pubescent with opposite lanceo- late bracts (c. 12 mm long) at the apex; central branch c. 4 mm long, bearing a globose umbellule (c. 12 mm 0) of sterile bacciform flowers (c. 2 mm 0), pedicels c. 3 mm, subtended by numerous ligulate bracts ; lateral branches c. 3 cm long, with opposite lanceolate bracts 4 mm long, ending in a head of c. 20-30 sessile flowers. Corolla and stamens not seen, described by Merrill as 3-merous. Fruit globose 6 mm (dry), 3-seeded. Distr. Malesia: Philippines (Babuyan Is.: Camiguin I.). Ecol. On slopes in forest, at 500 m. 33. Osmoxylon fenicis (Merr.) Philipson, Blumea 23 (1976) 114. — Boerlagiodendron fenicis Merr. Philip. J. Sc. 13 (1918) Bot. 44; En. Philip. 3 (1923) 223. — Boerlagiodendron tayabense Merr. Philip. J. Sc. 13 (1918) Bot. 45; En. Philip. 3 (1923) 224. Erect shrub or treelet a few m high, glabrous except for the inflorescence. Petiole to 45 cm, clasping base prolonged as a broad stipular ligule, IV2-3 cm long, several prominent long pectinate petiolar crests extending up the petiole as oblique groups of bristles; blade to 30 cm long, base cordate or truncate, deeply palmately 3-7-lobed, lobes extending to within c. 7 cm of the base, broadly elliptic to oblong, entire or with subsidiary lobes, somewhat narrowed towards the sinuses, apex abruptly apiculate, margin remotely denticu- late or serrate, sinuses very broadly rounded. Inflorescence a terminal, compound, furfuraceous umbel, hemispherical, c. 10-12 cm 0; peduncle 2-3 cm, densely enclosed in broadly ovate bracts c. 2 cm long, bearing dense fascicles of strong bristles on their blades; primary rays 15-30, c. 2^2-3 cm long, 2V2 rnm wide, pubescent, sub- tended by large bristly bracts c. P/2 cm long, bearing opposite terminal usually bristly bracts 6-15 mm long, each ending in 3 branches; central branch c. 4-10 mm, pubescent, bearing a terminal umbellule of c. 10-20 sterile bacciform flowers 2-3 mm 0, 2-celled, pedicels 3-4 mm tomentose sometimes with a ruff" of hairs around the pseudo- fruits, subtended by early-caducous small fimbriate bracts; lateral branches 2V2-3 cm long, with two opposite bracts (2-3 mm) about the middle, bearing a terminal head (8 mm without corollas) of c. 15-30 sessile flowers, subtended by incon- spicuous ovate tomentose bracts. Calyx an obsolete rim. Corolla in bud c. 1 mm long. Stamens 3. Ovary 3-celled. Fruiting head 10-12 mm 0; drupes c. 6 mm long, crowded, sessile, 3-angled; seeds 3. Distr. Malesia: Philippines (Luzon). Ecol. In primary dipterocarp forest, on rocky slopes near streams, 300-400 m. Note. Closely allied to 40. O. luzoniense and 27. O. pectinatum, but the combination of long- fringed petiolar crests, pubescent inflorescence branches, excessively bristly bracts and 3-merous flowers is distinctive. 34. Osmoxylon eminens (Bull) Philipson, Blumea 23 (1976) 114. — Trevesia eminens Bull, Cat. New Plants (1884) 17; Retail List (1885) 64, fig. — Boerlagiodendron mindanaense Merr. Philip. J. Sc. 3 (1908) Bot. 154. — Boerlagiodendron eminens (Bull) Merr. En. Philip. 3 (1923) 223. Small tree, up to 10 m, with few stout branches. Leaves large, forming terminal crowns, glabrous when mature, young inflorescence densely fur- furaceous; petiole to 1 m, stout (to 2 cm 0), flattened above, base clasping the stem, heavily lenticellate, prolonged as a stipular ligule 2 cm long, bicuspid, often with scales or bristles on the back, and bearing few to several entire, fimbriate or occasionally long setose crests; blade to 60 cm long, palmately 10-19-lobed, base cordate, lobes reaching to near the base, lanceolate to oblong, up to 15 cm wide, in outline either strap-shaped or irregularly pinnatisect, or the central lobe occa- sionally distinctly tripartite, margin coarsely and irregularly dentate, apex acuminate. Inflorescence a terminal compound umbel c. 40 cm ; peduncle stout c. 6 cm, IV2 cm wide, bearing many lanceo- late scaly bracts 3-6 cm long; primary rays numerous, rigid, flattened, 9-12 cm long, 6-10 mm broad, bearing opposite oblong scaly bracts (2-3 cm long) at the apex, each ending in three branches ; central branch '/2-I cm long, terminating in an umbellule (3-5 cm 0) of c. 20-40 sterile bacciform flowers, 7 mm 0, 2-3-celled, pedicels 1-2 cm long, surrounded by an involucre of small bracts (to 8 mm long); lateral branches c. 12 cm long, with opposite bracts (6-10 mm long) near the middle, terminating in a head of c. 50-60 sessile flowers, l'/2-2 cm (with corollas), heads spherical at anthesis, ovoid after corollas absciss; 1979] Araliaceae — I (Philipson) 49 bracts between the flowers very small. Calyx rim obsolete. Corolla 5-6 lobed, tubular below, 4-5 mm long. Stamens 4-6, filaments 7 mm long, anthers IV2 mm long. Ovary 2-3 mm long (at anthesis), 5-6-celled. Fruits crowded in dense ovoid heads 3-4 by 2V2-3 cm, drupes c. 9 by 5 mm, 5-6-angIed by mutual pressure, narrowed to the base, crowned by the persistent stigmatic boss; pyrenes crustaceous; endosperm wrinkled. Distr. Micronesia: Carolines; in Malesia: throughout the Philippines. Ecol. In primary forest from low altitude (100 m) to ridge forest and mossy forest at 950 m, often in shady ravines. Vern. Cf. Merrill: apalong or apulong. Bis., bunglui-babde, piria-pina, Sul., mangunpulun. Bag., palad-ulot, S.L.Bis., ulo-ulo, C.Bis.; in addition: lolobongan, Lan. Notes. This is the most widespread and most frequently collected species in the Philippines. It is also the most striking. Its large, many-lobed, fan- shaped leaves and the strong inflorescences, with globular flower-heads and large clusters of pseudo- fruits are distinctive. Only 16. O. pulchenimum resembles it somewhat in its leaf characters, but the central branches of the inflorescence rays of that species are much longer and its pseudo-fruits are sessile. The inflorescence branches are described as dull reddish brown, the flowers as light orange, and the fruits as indigo-black. 35. Osmoxylon serratifolium (Elmer) Philipson, Blumea 23 (1976) 114. — Boerlagiodendron serratifolium Elmer, Leafl. Philip. Bot. 2 (1908) 505; Merr. En. Philip. 3 (1923) 224. Sparingly branched shrub to 5 m. Petiole to 50 cm long, channelled above, clasping base pro- longed as an obtuse stipular ligule, and with few narrow ± fimbriate crests around the base of the petiole; blade to 50 cm long, base cordate, pal- mately lobed (up to 11 lobes), lobes extending to within about Vs from the base, narrowly elliptic, margin serrate (or slightly lobulate), apex acumi- nate, sinuses narrowly rounded. Inflorescence a terminal compound umbel c. 30 cm 0; peduncle stout, bracteate; primary rays 20-30, 4-5 cm long, 5-6 mm wide, flattened, subtended by lanceolate bracts 3-5 cm long, sometimes with bristles on the back, and bearing similar opposite terminal bracts 2V2 cm long, each terminating in three branches; central branch 12-15 mm long, terminating in an umbellule (3-4 cm 0) of c. 20-25 sterile bacciform flowers 5-6 mm 0, 3-4-celled, pedicels to 10 mm, interspersed with persistent small bracts; lateral branches c. 9 cm long, with opposite bracts (c. 6 mm long) 2-3 cm from the base, terminating in a spherical head of c. 30 sessile flowers c. 1 V2 cm (in bud), bracts between the flowers very small, obtuse, fimbriate. Calyx rim obsolete. Corolla 5-7-lobed above, tubular below, 5 mm long. Stamens 5-6, exserted, filaments 7 mm long, anthers 2 mm long. Ovary 3 mm long, 5-celled. Fruit unknown. Distr. Malesia: Philippines (Leyte, Camiguin, Panay). Ecol. Elmer noted that this species was rare in the low hills of Leyte. Notes. The flowers are orange-yellow (salmon), the fruits dark purple. The inflorescence is very similar to that of 34. O. eminens, but the leaves lack the many strong fan-like ribs of that species. 36. Osmoxylon celebicum Philipson, Blumea 23 (1976) 115. — Boerlagiodendron celebicum Harms ex KooRD. Minah. (1898) 489, nomen. A small, sparsely branched tree, 6 m high. Large leaves forming terminal crowns; petiole 50 cm by 8 mm, flattened above, with a sheathing base prolonged as a strong stipular ligule 3 cm long with branched fibrous setae on the outer surface, and with several crests bearing similar setae on the lower part of the petiole; blade 50 cm 0, base emarginate, deeply 9-lobed, lobes narrowly elliptic to lanceolate, narrowed towards the sinuses, apex broadly cuneate, margin minutely and remotely serrate, sinuses broadly rounded. Inflorescence a terminal compound subspherical umbel c. 20 cm ; peduncle short, stout (15 mm wide) with large setose bracts (c. 4 cm long) below and among the primary rays; primary rays c. 15, c. 6 cm long, 5 mm broad, with a pair of large setose bracts (22 by 10 mm) at the apex, each ending in three branches; central branch c. 6 by 2 mm, terminating in an involucre of setose ovate bracts (c. 4 mm long) and an umbel of c. 20-30 sterile bacciform flowers (c. 3 mm when dry, 2-3-celled, on pedicels 6-10 mm long) interspersed with bracts covered with crisp reddish-brown setulae; the two lateral branches c. 2 cm long, with opposite setulose bracts (c. 1 cm long) below the middle, terminating in a spherical head of c. 30-40 sessile flowers each sub- tended by a reddish brown setulose cymbiform bract. Calyx rim obsolete. Corolla c. 2V2 mm long in bud (not seen in open condition). Stamens 5. Ovary subcylindric, c. 1 mm long in bud, 5-celled; disk with a central stigmatic boss. Fruit unknown. Distr. Malesia: Celebes (Minahasa, Manado). Ecol. On rich volcanic sand, at 10 m. Vern. Sinomaha. Note. The flower buds are orange and the fruits deep purple. 37. Osmoxylon talaudense Philipson, Blumea 23 (1976) 115. Shrub or small tree, to 6 m, glabrous. Leaves at the ends of the stout branches; petiole to 60 cm, broadly channelled above, clasping base prolonged as a stipular ligule c. 2 cm long, and bearing 2-3 fimbriate crests; blade c. 50 cm long truncate to cordate at the base, palmately 7-1 1 -lobed to within 50 Flora Malesiana [ser. I, vol. 91 Fig. 16. Osmoxylon micranthum (Harms) Philipson. a. Habit, x V^, b. flower, c. false fruit and ditto in CS, X 15, d. CS of fruit, x 6 {a-c Kanis 1384, d Pullen 428). Drawn by W. R. Philipson. 1979] Araliaceae — I (Philipson) 51 ± V4 of the base, lobes elliptic oblong slightly narrowed towards the rounded sinuses, apiculate, margin denticulate to undulate. Inflorescence a terminal compound umbel, peduncle 1-2 cm, bearing broad ovate bracts 2-3 cm long, with bristles on the back, (similar persistent bracts subtend the primary rays); primary rays c. 15, c. 3-4 cm long, 4 mm wide, flattened, bearing opposite terminal persistent bracts IV2-2 cm long, sometimes with a few bristles on the back, ending in three branches; central branch 8-10 mm long, terminating in an umbellule 2 cm of 15-20 sterile bacciform flowers (4 mm 0, 2-celled) surrounded by an involucre of obtuse bracts 1-2 mm lon^, pedicels 5-7 mm; lateral branches 4V2-5 cm long, bearing opposite ovate persistent bracts (3-7 mm long) c. 5-10 mm above the base, terminating in a dense head c. 1 cm of 30-40 sessile flowers inter- spersed with inconspicuous obtuse bracts. Calyx rim obsolete. Corolla 5-lobed. Stamens 5. Ovary turbinate, 1 V4 mm long, 5-celled. Fruit in spherical heads 2 cm (when dry); drupes c. 9 by 6 mm, obovoid, 5-ribbed. Distr. Malesia: N. Moluccas (Talaud Is.: Karekelong and Salebabu). Ecol. Common in forest, besides streams, from near sea-level to 100 m. Vern. Laripatu, Talaud. Note. The flower is yellow-orange and the fruit dark purple. For a discussion of the distinctive features, see under 31. O. sessiliflorum. 38. Osmoxylon micranthum (Harms) Philipson, Blumea 23 (1976) 115. — Boerlagiodendron micranthum Harms, Bot. Jahrb. 56 (1920) 379. — Boerlagiodendron sayeri Harms, I.e. 379, f. 1 a-j. — Eschweileria gawadensis Baker/. J. Bot. 61 (1923) 22. — Boerlagiodendron tricolor Philipson, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 11. — Fig. 16. A sparsely branched shrub to 8 m, sometimes trailing or semi-scandent, young parts uniformly setulose, buds without cataphylls. Leaves in terminal clusters; petiole up to 30 cm, rather narrow (2-4 mm 0), becoming sparsely setulose, channelled above, with a sheathing base prolonged as a membranous stipular ligule up to 3 cm long, and with a number of lacerate crests encircling the lower part of the petiole; blade deeply 3-5- or more rarely 7-lobed, or below the inflorescence sometimes simple, base cordate or emarginate, the central lobe up to 30 cm long, the lobes oblong, lanceolate or broadly elliptic, entire or irregularly lobed or incised, or with small sub-lobes, apices long cuspidate, acute, margin serrate, sinuses between the lobes broad and rounded, surfaces become sparsely setulose to subglabrous. Inflores- cence a terminal compound umbel, often appearing subterminal by growth of a leafy braiich at the base of the peduncle; peduncle short (1-2 cm), heavily setulose, occasionally with a flowering ray arising from the axils of bracts on or below the peduncle, bearing distally many lanceolate bracts 5-10 mm long; primary rays 12-18, 10-20 mm long, setulose, with two lanceolate bracts at the apex, each ray ending in three branches; the central branch very short (2-3 mm) bearing a subglobose umbel of many (c. 40) small sterile bacciform flowers (c. IV2 by 1 mm) with filamentous pedicels c. 2 mm long, and 1-2 cells each with 1 abortive ovule; the two lateral branches c. 2 cm long, with two minute bracts about their middle, setulose, terminating in a head of c. 20 sessile flowers sur- rounded by an involucre of small rounded bracts. Calyx rim obsolete. Corolla 4(-5)-lobed, united below, c. 2 mm long. Stamens 4(-5), filaments ribbon-like elongating beyond the corolla tube at anthesis, 3-4 mm, anthers small. Ovary shortly subcylindric, c. 1 mm high, faintly angled, glabrous, 1-5-, usually 4-celled, disk fleshy, rising slightly to the central stigmas. Fruit an ellipsoid drupe with 1-5 cartilaginous pjTenes; seeds with smooth endosperm. Distr. Malesia: New Guinea (Irian Jaya: Idenburg R. to Milne Bay Distr.). Ecol. In primary forest from the foothills to the montane mossy forest, often in swampy or deeply shaded situations, 700-2400 m. Vern. Diande, Chimbu, kenata, Okapa. Note. The inflorescence branches are often red or purple, and the flowers either orange or reddish with yellow anthers. The ripe fruits are deep purple or black. The shape and size of the leaf can vary greatly, even on the same plant. The ovary usually has 4 cells, but plants with 3, 2 and 1 occur. Since these are alike in other respects they have been treated as a single species. Although the type of Boerlagiodendron tricolor has an ovary with 5 cells and is from much further west than other gather- ings, it is not considered to be specifically distinct. 39. Osmoxylon trilobatiun (Merr.) Philipson, Blumea 23 (1976) 116. — O. cumingii Seem. J. Bot. 6 (1868) 141, nomen. — Boerlagiodendron trilo- batum Merr. Philip. J. Sc. 2 (1907) Bot. 289; En. Philip. 3 (1923) 224. Slender shrub or small tree up to 5 m, becoming glabrous except for slight pubescence on the inflorescence. Leaves clustered near the ends of the branches; petioles to 25 cm, c. 3 mm wide, clasping base prolonged as a broad stipular ligule c. 1 cm long, and with 2-3 entire, or obscurely fimbriate, often recurved crests surrounding the base of the petiole; blade 3- or occasionally 5-lobed (leaves below the inflorescence sometimes simple), to 30 by 28 cm, base broadly cuneate, rounded or trun- cate (emarginate in 5-lobed leaves), lobes about ^2-^/3 of the blade, narrowly or broadly oblong, often slightly narrowed below and sharply acumi- nate to caudate, margin serrate. Inflorescence a terminal compound spherical umbel, 7-15 cm 0, either rather compact or branches lax; peduncle 2-3 cm with broad ovate bracts; primary rays 8-20 52 Flora Malesiana [ser. I, vol. 9 or more, slightly pubescent, 1V2-4 cm long, sub- tended by ovate bracts 5-10 mm long, opposite ovate bracts at the apex, 1-3 mm long; central branch IV2-6 mm long, pubescent, bearing an umbel (1-1 V2 cm 0) of sterile bacciform flowers up to 5 mm 0, 1-4-celled, pedicels 2-6 mm long, subtended by ovate bracts 1-3 mm long; lateral branches 1^2-3 cm with opposite small bracts about the middle, bearing a terminal head, c. 1 cm of c. 8-20 flowers, surrounded by an involucre of small rounded pubescent bracts, pedicels c. 1 mm or less (up to 3 mm in fruit). Calyx rim obsolete. Corolla 4-5-lobed above, tubular below, 2-3 mm long. Stamens 4-5, exserted. Ovary subcylindric, 4-5-celled. Fruit a spherical drupe (when dry 4-5- ribbed, 7 by 5 mm). Distr. Malesia: widespread in the Philippines (Luzon to Mindanao). Ecol. In primary forest, frequently beside streams in damp ravines, from the lowland at 75 m to 750 m. Vern. Kamay-kamay, Tag., ayum, C.Bis. Note. Sparingly branched but wide-spreading slender shrub, with yellowish bark, at first heavily dotted with brown lenticels. The flowers are white and the ripe fruit smooth and purple. 40. Osmoxylon luzoniense (Merr.) Philipson, Blumea 23 (1976) 116. — Boerlagiodendron luzoniense Merr. Philip. J. Sc. 3 (1908) Bot. 252; En. Philip. 3 (1923) 223. — Boerlagiodendron dementis Merr. Philip. J. Sc. 3 (1908) Bot. 155; En. Philip. 3 (1923) 222. — Boerlagiodendron agusanense Elmer, Leafl. Philip. Bot. 7 (1914) 2330; Merr. En. Philip. 3 (1923) 222. — Boer- lagiodendron diversifolium Merr. Philip. J. Sc. 10 (1915) Bot. 333; En. Philip. 3 (1923) 223. Erect, unbranched or sparsely branched shrub or tree to 8 m, becoming glabrous except for the inflorescence. Leaves clustered at the ends of the branches; petiole to 40 cm, with a clasping base prolonged as a short stipular ligule, and with several basal entire or shortly fimbriate crests; blade to 33 cm 0, base truncate or cordate, deeply palmately 3-7-lobed, sinuses broad, rounded, lobes elliptic, usually narrowed below, entire or with subsidiary lobes (the central lobe especially often narrow below and strongly pinnately lobed), apex acute, margin coarsely serrate, coriaceous ; upper- most leaves often reduced and simple. Inflorescence a terminal compound, subsessile umbel 10-15 cm ; primary rays c. 20-30, 2-3 cm long, subtended by lanceolate furfuraceous and ± fimbriate bracts, furfuraceous villose or ± hirsute, with opposite rounded or lanceolate hirsute bracts at the apex, each ending in three branches; central branch c. 2-8 mm long, hirsute, ending in an involucre of minute bracts (1 mm) surrounding a globose (2 cm 0) umbellule of c. 15-20 sterile flowers (c. 6 by 6 mm, 2-3-celled), pedicels 2-3 mm, hirsute ; lateral branches c. 2V2-3V2 cm long at anthesis. with an articulation about the middle, i hirsute, ending in a globose head, 2 cm (with open corollas), with ovate tomentose obtuse bracts, c. 2 mm long; flowers 30-^0, ± sessile (rarely pedicels to 2 mm). Calyx rim obsolete. Corolla 4-5-lobed above, tubular below, 3V2-4 mm long. Stamens 4-5, exserted, filament stout, 5 mm, anther 1 mm long. Ovary 4-5-celled. Fruit 6 by 5 mm (dry), strongly 4-5-ribbed. Distr. Malesia: widespread in the Philippines (Luzon to Mindanao), also in N. Celebes. Ecol. In forests, often by streams and on ridge in mossy forest, 280-1650 m. Vern. Philippines: bolwang hi inalahan. If., iyangnok, Mbo, malakapdyas, S.L.Bis., molonpolon, Buk., tangan-tangan-batu, Buk., tachung, vangang, Ig. Notes. Inflorescence yellow to red (salmon), fruits blue-black or purple. Merrill did not liken his Boerlagiodendron diversifolium (from Mindanao) to this species, no doubt because he gave importance to the occur- rence of variable leaves and the 5-merous flowers. However, specimens from Luzon may possess simple leaves below the umbel, and both 4- and 5-merous flowers occur in both Luzon and Mindanao. The greater range of material now available establishes the identity of the two species. Similarly, no features seem to distinguish Boer- lagiodendron dementis, and B. agusanense though a greater range of collections would be desirable. The species is treated here in a broad sense. The inflorescence characters of most specimens are uniform, being hirsute and with the bracts at the apex of the primary rays obtuse and short. Some specimens (Curran 5088, Elmer 16762) have longer lanceolate bracts with some bristles on the back. The northernmost specimen, from Ilocos Norte, has finer and less hairy inflorescence rays (recalling 39. O. trilobatum), but the foliage agrees with this species. Leaf-shape is more variable, even on the same specimen. Other species which resemble O. luzoniense in some respects are: 32. O. camiguinense with broader, shallower lobing, a more delicate inflorescence, and tri-merous flowers ; 27. O. pectinatum with glabrous inflores- cence branches and long-pectinate petiolar crests; and 26. O. humile with pedicelled flowers forming less dense heads. A specimen from Surigao Province (BS 83562) has a most interesting abnormal structure. The central branches of the inflorescence rays bear heads of apparently fertile flowers, with lobed corollas and exserted stamens. 41. Osmoxylon insigne (MiQ.) Becc. Malesia 1 (1878) 195; Philipson, Blumea 23 (1976) 117. — Trevesia insignis MiQ. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 222. — Trevesia palmata var. insignis Clarke, F1. Br. Ind. 2 (1879) 732, pro nomen. — Eschweileria insignis (MiQ.) Boerl. Ann. Jard. Bot. 1979] Araliaceae — I (Philipson) 53 Btzg 6 (1887) 122. — Boerlagiodendron insigne (MiQ.) Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 31. A glabrous tree. Leaves large, palmately lobed; petiole to 45 cm, 8-10 mm wide, flattened above, with a sheathing base prolonged as a stipular ligule, numerous fimbriate crests around the base of the petiole, and irregular tufts of bristles along the whole length of the petiole; blade to 55 cm 0, deeply 5-7-lobed, the sinuses broadly rounded and c. 4-5 cm from the base of the blade, lobes pinnatilobed, with a narrow base and an attenuated apex, margins serrate. Inflorescence a terminal compound umbel c. 16 cm ; peduncle stout, with lanceolate bracts (IV2-2 cm long) subtending the primary rays; primary rays 25-30, c. 4 cm long, bearing 2 caducous lanceolate bracts (c. 1 cm long) at the apex, each ray ending in 3 branches; central branch c. 12-16 mm long, bearing an umbel (c. 2 cm 0) of c. 20 sterile ovoid bacciform flowers (c. 4 mm long when dry) with pedicels c. 4-6 mm long, and 3-celled; two lateral branches c. 3^2 cm long, with an articulation about the middle, termi- nating in an umbellule (c. 2 cm 0) of c. 10-15 flowers on short stout pedicels c. 2 mm long, umbellules surrounded by a receptacular rim after caducous bracts have abscissed. Calyx rim minute. Petals c. 3V4 mm long in bud (when dry), with 8-9 lobes above, tubular below. Stamens 8-9, with stout filaments. Ovary cylindric, 8-9-celled; disk with a central double row of pustulate stigmas. Fruit unknown. Distr. Malesia: Moluccas (Batjan). Note. Tufts of bristles along the entire length of the petiole together with the pinnatifid lobes of the leaf are distinctive. Seemann (J. Hot. 4, 1866, 353) referred to 5-flowered umbels with 5-angled drupes, but this probably relates to the New Guinea specimen which he included under this name. Insufficiently known Boerlagiodendron ledermannii Harms, Bot. Jahrb. 56 (1920) 383; Philipson, Blumea23 (1976) 117. — Type: Ledermann 12293. Harms compared this species with Boerlagio- dendron geelvinkianum. The size of the foliage and flowers prevents it from being included within that species. If it represents a local species, it has not been re-collected since the original gathering of Ledermann in 1912. The type specimen, which was incomplete, was destroyed during the war. Boerlagiodendron monticola Harms in K. Sch. & Laut. Fl. Schutzgeb. Nachtr. (1905) 330; Philip- son, Blumea 23 (1976) 117. — Type: Schlechter 14471. The incomplete type specimen, gathered by Schlechter, was destroyed during the war. This species was evidently similar to 38. Osmoxylon micranthum, but the ovary was possibly 10-celled. I have tentatively identified Robbins 1644 as this species: it is close to O. micranthum but its ovary, with 8 cells, is outside the range of variation of that species and the pedicels are longer (in fruit). Its distribution (foothills of Adalbert Range) is not dissimilar to that of Boerlagiodendron monticola (Torricelli Mts). 8. ARTHROPHYLLUM Bl. Bijdr. (1826) 878; DC. Prod. 4 (1830) 266; MiQ. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 27; Bth. in B. & H. Gen. PI. 1 (1865) 944; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 54; Koord. Atlas 4 (1916) f. 675 & 676; Hutch. Gen. Fl. PI. 2 (1967) 80; Stone, Gard. Bull. Sing. 30 (1977) 276; Philipson, I.e. 299, f. 1-16; Adansonia 17 (1978) 329. — Mormoraphis Jack ex Wall. Cat. (1831) n. 4931, nomen. — Eremopanax Baill. Adansonia 12 (1878) 158. — Fig. 17, 19-23. Unarmed, sparingly branched trees or shrubs. Leaves on vegetative shoots and lower leaves spirally arranged, imparipinnate, often crowded at the end of the branches, those on flowering branches often opposite, smaller, or reduced to a single leaflet; petiole terete; rachis articulated at the insertion of the pinnae and leaflets ; leaflets entire ; stipular sheath clasping, small, ligule a mere rim. Inflores- cence consisting of compound umbels, either solitary and terminal or more commonly borne on a cluster of specialized leafy branches arising from the axils of the uppermost leaves; pedicels not articulated. Flowers bisexual. Calyx an undulate rim, sometimes with indistinct teeth, persistent. Petals 4-6, valvate in bud. Stamens 4-6, anthers curved, basifixed. Ovary turbinate, 1 -celled; disk 54 Flora Malesiana [ser. I, vol. 91 Fig. 17. Habit of Arthrophyllum diversifolium Bl. (Photogr. Hoogland, Bogor). 1979] Araliaceae — I (Philipson) 55 fleshy, rising in the centre to the ± sessile capitate stigma. Fruit ovoid or spheroidal, often oblique; exocarp leathery; endocarp cartilaginous, ^'ee^ solitary, pendulous; endosperm deeply transversely ruminate. Distr. About 31 spp. (17 in Malesia) extending from the Nicobar Is. and Indo-China to the Philippines, New Guinea and New Caledonia. Fig. 18. Notes. The principal distinctive characters of this isolated genus are the single-celled ovary and the arrangement of the umbels on specialized lateral branches in the majority of the species. The present treatment, following that which I gave in 1977, I.e., remains tentative until widespread field studies can be undertaken. KEY TO THE SPECIES 1 . Inflorescence becoming paniculate by the successive development of branches below the umbellules. Fig. 19a 1. A. prolifenun 1. Inflorescence a compound umbel. 2. Inflorescence with four orders of branching. 3. Leaflets lanceolate. 4. Leaflets c. 4-7 cm long. Fig. 20f 2. A. ashtonii 4. Leaflets much longer 3. A. angustifolium 3. Leaflets broader. 5. Pedicels (at early anthesis) c. 10 mm long. 6. Peduncles of umbellules with bracts or their scars. Fig. 21a 4. A. ahernianum 6. Peduncles of umbellules without bracts or their scars. Fig. 21b 5. A. engganoense 5. Pedicels (at early anthesis) c. 5 mm long, or shorter. 7. Leaves associated with the umbels rotund. Fig. 21e 6. A. collinum 7. Leaves (or leaflets) associated with the umbels ovate or elliptic. 8. Leaves associated with the umbels ovate, ± fleshy, with the lower surface smooth (lateral veins obscure). Fig. 21f 7. A. crassum 8. Leaves (or leaflets) associated with the umbels ± elliptic, coriaceous or chartaceous, veins visible. 9. Young parts glabrous 8. A. pacificum 9. Young parts with rufous tomentum. 10. Umbels at anthesis with numerous filamentous pedicels. Fig. 21c. . . 9. A. diversifolium 10. Umbels at anthesis with fewer stout pedicels. Fig. 21d 10. A. macrantbum 2. Inflorescence with three (or fewer) degrees of branching. 1 1 . Leaflets membranaceous or chartaceous. 12. Mid-leaflets c. 16 cm long 11. A. papyraceum 12. Mid-leaflets c. 8 cm long or shorter. 13. Leaflets usually 5-7 (Fig. 20a). Primary inflorescence branches usually short (c. 3-6 cm) and without articulations 12. A. maingayi 13. Leaflets more numerous (Fig. 20d). Primary inflorescence branches longer (10-20 cm), with one or more leafy nodes 13. A. kjellbergii 11. Leaflets coriaceous. 14. Leaflets c. 5 14. A. cenabrei 14. Leaflets more numerous. 15. Petals and stamens 6 15. A. pulgarense 15. Petals and stamens 4 or 5. 16. Leaflets 6 cm long, or longer. Fig. 20b 16. A. montanum 16. Leaflets shorter. Fig. 20c 17. A. alternifolium 1. Artbrophyllum prolifenun Philipson, Gard. sometimes oblique, apex acute. Inflorescences Bull. Sing. 30 (1977) 302, f. 3-4. — Fig. 19a-b. terminating specialized plagiotropic shoots, bear- Medium-sized, glabrous tree. Leaver muhijugate, ing axillary flowering branches and ending in up to 90 by 24 cm, of the flowering branches smaller umbellules of a few flowers below which pairs or with fewer pinnae or usually simple; petioles up to whorls of branches continue the growth of the 28 cm, 5 mm 0; petiolules 1-1 V2 cm; leaflets inflorescence to produce an elongated panicle of obovate-oblong,c. 16 by 7 cm, chartaceous, margin umbellules; pedicels 7-8 mm (slightly longer in slightly revolute, base broadly cuneate to truncate, fruit). Calyx often with 5 indistinct teeth. Petals 5, 56 Flora Malesiana [ser. I, vol. 9^ Fig. 18. Species density of Arthrophyllum Bl. in Malesia ; above the hyphen the number of endemic species, below it the non-endemics. Complete range of genus encircled; 17 of the total of 31 spp. occur in Malesia. 3V2 mm long, narrowly triangular. Stamens 5, filaments c. 2 mm, anthers reniform, c. V2 mm long. Ovary obconical, c. 2 mm long; disk fleshy, cushion-like; stigma capitate ± sessile. Fruit ovoid, fleshy, capped by the calyx and the enlarged beak-like stylopodium, c. 10 by 5 mm when dry. Distr. Malesia: E. New Guinea (Morobe Distr.). Ecol. Mid-mountain rain-forest, reaching the canopy, on steep slopes, 300-1200 m. Notes. The thick outer bark is grey-brown, fissured, and peeling in small flakes. Wood straw- coloured. Cut stems exude brown latex. The thick petals are yellow-green. The flower and fruit are typical of this well- defined genus, but the branching of the inflores- cence is unlike that found in all other species. 2. Arthrophyllum ashtonii Philipson, Gard. Bull. Sing. 30 (1977) 303, f. 12. — Fig. 20f. Slender small tree, to 5 m, with the leaves dis- persed for some distance from the apex of the branches, young parts with brown scurfy tomen- tum which persists on the umbellules. Leaves multijugate, to c. 30 cm long ; of the flower-bearing branches smaller with fewer pinnae, or simple; petioles slender, c. 7-9 cm; petiolules c. 4-7 mm; leaflets lanceolate to broadly lanceolate, 3-7V2 by 1-2 cm, thinly coriaceous, margin revolute, base broadly cuneate, apex tapered to subcaudate, veins chaimelled above, visible beneath. Inflorescence a terminal cluster of specialized leafy branches ; main rays variable in length in the same inflorescence, the longest from c. 16-25 cm long with a pair of opposite simple leaves about the middle (with flowering branches in their axils) and ending in a whorl of secondary rays subtended by simple leaves; secondary rays 4-12 cm long, bearing simple leaves, usually in an opposite pair and terminating in compound umbels ; umbellules witji c. 8-10 flowers; pedicels 4-10 mm, furfuraceous. Flower buds c. 2 mm long, calyx a furfuraceous rim. Petals 5, broadly triangular. Stamens 5, anthers curved, basifixed. Ovary glabrous, obconical, c. 1 mm long; stigma on a raised stylopodium at the centre of a flat disk. Fruit spheroidal, with a persistent stylopodium, c. 6 mm long when dry. Distr. Malesia: Borneo (Sarawak and Brunei). Ecol. Mossy forest on sandstone ridge, and in kerangas forest, 1000-1550 m. Note. The small narrow leaflets are very dis- tinctive. No other species with small leaflets has inflorescences which branch to the fourth degree. 3. Arthrophyllum angustifolium Ridl. J. Fed. Mai. St. Mus. 10 (1920) 136; Fl. Mai. Pen. 1 (1922) 885; Philipson, Gard. Bull. Sing. 30 (1977) 304. Shrub or small tree, up to 5 m, young parts rufous-tomentose, glabrescent. Lower leaves multi- jugate, rachis dilated, c. 70-90 by 30-40 cm; petioles c. 15-22 cm, 3-5 mm wide, ligule a rim c. 2 mm long; petiolules c. 7-10 mm long; leaflets coriaceous, lanceolate, c. 15-22 by IV2-2V2 cm, tapering to an acute or obtuse apex, base cuneate, margin slightly revolute; upper leaves reduced, mostly unifoliolate, opposite, broader, with petioles 2-4 cm long. Inflorescence a terminal cluster of specialized leafy branches; main rays 30 cm (or more) long, bearing simple leaves in opposite pairs with small flowering branches in their axils, ending in a whorl of c. 10-12 secondary rays subtended by simple leaves; secondary rays 8-12 cm long, bearing pairs of simple leaves, and terminating in an umbel of 5-12 tertiary rays c. 2-3 cm long, with bract scars about the middle, and ending in an umbellule of c. 8-12 flowers, pedicels c. 5 mm. Petals 5, 2-3 mm long in bud. Stamens 4. Ovary turbinate, inconspicuous at anthesis; disk fleshy; stigma ± sessile. Fruit spheroidal, c. 5 by 5 mm when dry, calyx and stylopodium small. Distr. Malesia: Malay Peninsula (Perak) and Borneo (Brunei). Ecol. Forest and old regenerated forest on peat swamp at low altitude or on ridges. Note. The lanceolate leaflets are unlike those of any other species. The grey bark is minutely fissured and bears many small orange lenticels. The wood is soft and white. The Malayan and Bornean specimens are similar, except that the flower buds are larger in the Brunei plant. 4. Arthrophyllum ahemianum Merr. Philip. J. Sc. 1 (1906) Suppl. 109; En. Philip. 3 (1923) 235; Philipson, Gard. Bull. Sing. 30 (1977) 304, f. 13. — A. pinnatum {non Clarke) F.-Vill. Nov. App. (1880) 103; ViDAL, Sinopsis Atlas (1883) 28, t. 55 f. c. — Macropanax sp. Vidal, Rev. PI. Vase. Filip. (1886) 145. — A. sablanense Elmer, Leafl. Philip. Bot. 1 (1908) 331. — A. borneense Merr. PI. Elm. Born. (1929) 231, non Baker, 1896. — A. elmeri Merr. Webbia 7 (1950) 319. — A. merrilliana Furtado, Gard. Bull. Sing. 19 (1962) 185. — Fig. 21a. 1979] Araliaceae — I (Philipson) 57 Fig. 19. Arthrophyllum prolifenim Philipson. a. Part of inflorescence in fruiting state, x V2. t>. ditto, showing detail of ultimate branches in flowering stage, nat. size. — A. maingayi Philipson. c. Terminal inflorescence, x V2- — /I. mo«?a«MW Ridl. ^. Single flowering shoot, x V3 (Courtesy Gard. Bull. Sing. 30, 1977). Tree up to 15 m, young parts with rufous tomen- tum. Leaves clustered at the ends of the branches, multijugate, up to 200 by 60 cm; of the flower- bearing branches smaller with fewer pinnae, or simple; petiole stout, up to 35 cm; petiolules 5-20 mm; leaflets ovate-oblong, occasionally oblong-lanceolate, up to 35 by 12 cm, membrana- ceous or chartaceous, margin revolute, base cuneate to rounded, usually oblique, apex short acuminate. Inflorescence a whorl of specialized leafy branches forming a terminal crown ; main rays up to 150 cm (or more), bearing pinnate leaves usually in 1-2 opposite pairs, and with flowering branches in the upper axils, ending in a whorl of secondary rays subtended by pinnate or more rarely simple leaves; secondary rays up to 30 cm bearing simple or pinnate leaves in opposite pairs, and terminating in compound umbellules; um- bellules with c. 10-20 flowers c. 3 cm 0; peduncles with 1-2 pairs of small simple, often caducous leaves; pedicels c. 1 cm (at anthesis) subtended by minute caducous bracts. Petals 5, 4 mm long. Stamens 5, anthers curved. Ovary turbinate; disk fleshy; stigma — sessile. Fruit c. 10 by 7 mm, ellipsoidal, calyx and stylopodium forming a prominent beak. 58 Flora Malesiana [ser, I, vol. 9^ Fig. 20. Foliage leaves of Arthrophyllum spp. a. A. maingayi Philipson, b. A. montanum Ridl., c. A. pulga- rense Elmer, d. A. kjellbergii Philipson, e. A. alternifolium Maingay ex Ridl.,/. A. ashto/iii Philipson. All X Vs (Courtesy Card. Bull. Sing. 30, 1977). Distr. Malesia: N. Borneo and throughout the Philippines to the northern Moluccas (Talaud, Ternate). Ecol. Primary and second-growth forest, from the lowlands to 1000 m. Vern. Philippines, cf. Merrill: alabihig, dokloi, P.Bis., binaldyon, C.Bis., danipo, Ig., higin, Mang., malapapaya, puyga-puygahan. Tag., pama- latdngen-a-purau. Ilk. Additional names: Philip- pines: lulpo, Luzon, mayari, Mindoro, bungyo, Palawan; Moluccas: langator'a, Talaud. Notes. This species replaces the more westerly 9. A. diversifoUum which it closely resembles. It is characteristically larger in all its parts, particularly in the size of the individual flowers and the length of their pedicels. There are fewer flowers in an umbellule. The distinction between these two species is not always easy to make, especially when the material is fragmentary; Philipson, I.e. A few specimens from the Philippines appear very similar to A. diversifoUum, and it is possible that this species extends beyond Borneo. I have regarded them as part of the range of variation of A. aherniamtm. Similarly, at least one specimen from S. Borneo approaches A. ahemianum in appearance. 1979] Araliaceae — 1 (Philipson) 59 5. Arthrophyllum engganoense Philipson, Gard. Bull. Sing. 30 (1977) 305, f. 14. — Fig. 21b. Tree to 21 m high, becoming glabrous. Lower leaves imparipinnate, multijugate, 60 cm long or more; petiole 24 cm, 6 mm wide, rachis articulated at the insertion of the leaflets; petiolules 10-18 mm; leaflets broadly elliptic to elliptic-oblong, c. 12-15 by 6-7 cm, base rounded with a short asymmetrical cuneate centre, apex shortly apiculate, margin entire, often undulate chartaceous. Flowering branches c. 40 cm; leaves opposite, simple, or unifoliolate, petiolules c. 5-7 cm, leaflets ovate, c. 13 by 6 cm, with inflorescence branches in their axils; ending in a whorl of simple leaves surround- ing a compound umbel to 30 cm ; secondary rays c. 8, c. 10-15 cm long at flowering, slender (2 mm 0) and striate, each bearing a pair of small leaves about the middle (sometimes with inflorescences in their axils) and ending in an umbel; tertiary rays c. 8, slender, c. 20-40 mm long, without bracts; tertiary rays pedicels c. 5-10 per umbellule, c. 10-15 mm at anthesis. Calyx an undulate rim. Petals 5, c. 2 mm long in bud. Stamens 5, anthers curved. Ovary turbinate, obscurely ribbed, c. VI2 mm long at anthesis. Fruit ellipsoid, c. 10 by 7 mm, with a rather small persistent calyx and stylopodium. Distr. Malesia: S. Sumatra (Enggano I.), two collections. Ecol. Forest at low altitude, up to c. 100 m. Vern. Langkapii iitan kaauh, Enggano. Note. The two known collections of this species are very similar and contrast with the widespread 9. A. diversifolium because of the few-flowered umbellules with long, spreading pedicels. 6. Arthrophyllum collinum Philipson, Gard. Bull. Sing. 30 (1977) 305, f. 17. — Fig. 21e. Sparingly branched shrub or small tree up to 12 m, all young parts with dense, rufous tomentum which persists on flowers and inflorescences. Leaves multijugate, up to 60 (or more) by 32 cm; of the flowering branches smaller with fewer pinnae or more frequently unifoliolate, leaflets more rotund and with longer petioles; petiole up to 20 cm; petiolules c. 1 cm; leaflets oblong, broadly elliptic or rotund, up to 16 by 7 cm, coriaceous, margin slightly revolute, base truncate to rounded, unequal, apex rounded, obtuse, or shortly and bluntly apiculate, midrib prominent, lateral veins usually clearly visible below, upper surface fre- quently rugose. Inflorescence a cluster of specialized leafy branches forming a terminal crown ; main rays up to 60 cm long, bearing one or more, rarely two, opposite pairs of usually unifoliolate rotund leaves often with flowering branches in their axils, and ending in a whorl of secondary rays, subtended by usually unifoliolate leaves; secondary rays up to 17 cm, bearing a pair of unifoliolate leaves with flowering branches in their axils, and terminating in compound umbellules; tertiary rays c. 5-10, Fig. 21. Umbellules of some Arthrophyllum spp. a. A. ahernianum Merr., b. A. engganoense Philipson, c. A. diversifolium Bl., d. A. macranthum Philipson. All nat. size. — Bracts of two Arthro- phyllum spp. e. A. collinum Philipson,/. A. crassum Philipson. Both ■: \3 (Courtesy Gard. Bull. Sing. 30, 1977). c. 3-6 cm long, articulated near the middle; pedicels c. 10, c. 2-3 mm long, rufous tomentose. Petals 5, broadly triangular, c. 2 mm long. Stamens 5, anthers curved. Ovary turbinate, rufous-tomen- tose, c. 1 V 2 mm long at anthesis ; disk fleshy, stigma sessile. Fruit ellipsoidal, c. 9 by 5 mm (when dry), the calyx and stylopodium prominent. Distr. Malesia: Borneo (Sarawak and Sabah). Ecol. Forest and scrub, sometimes growing as an epiphyte in the crowns of trees, 600-2700 m. Notes. The bark is grey and smooth, the wood pale and soft, and the cut stems exude a yellowish or orange latex. This species is characterized by the rotund, coriaceous leaves on the flowering branches. 7. .\rthrophylluin crassum Philipson, Gard. Bull. Sing. 30 (1977) 305, f. 18. — Fig. 21f. Sparingly branched shrub or small tree up to 8 m, occasionally epiphytic, all young parts with dense, rufous tomentum which persists on the 60 Flora Malesiana [ser. I, vol. 9^ flowers and inflorescence. Leaves multijugate, up to 100 (or more) by 50 cm, occasionally with 4 leaflets inserted at some of the lower articulations ; of the flowering branches smaller with fewer pinnae or more frequently unifoliolate; petiole up to 25 cm; petiolules 10-20 mm; leaflets elliptic, oblong or lanceolate, up to 24 by 7'/2 cm, coria- ceous, margin strongly revolute, base cuneate to rounded, often oblique, apex narrowed to an acute often caudate apiculum, midrib prominent, lateral veins faint to obscure. Inflorescence a cluster of specialized leafy branches forming a terminal crown; main rays up to 70 cm, bearing an opposite pair (or rarely more pairs) of unifoliolate or (less frequently) pinnate leaves, often with flowering branches in their axils, and ending in a whorl of secondary rays, subtended by usually unifoliolate leaves; secondary rays up to 25 cm, bearing a pair of usually unifoliolate leaves with flowering branches in their axils, and terminating in com- pound umbellules; tertiary rays c. 10, c. 2-3 cm long, articulated near the middle; pedicels c. 12-14, c. 3-4 mm long, rufous-tomentose. Petals 5, broadly triangular, c. 2 mm long. Stamens 5, anthers curved. Ovary turbinate, rufous-tomentose, c. IV2 mrn long at anthesis; disk fleshy, stigma sessile. Fruit spheroidal, c. 6 by 5 mm (when dry), calyx and stylopodium prominent. Distr. Malesia: Borneo (Sarawak and Kaliman- tan). Ecol. Swampy peat forest and heath woodland, both primary and disturbed, from sea-level to c. 1 50 m. Some fragmentary collections from higher altitude (1000 m) further inland may belong to this species. Note. The leaves associated with the in- florescence are distinctively fleshy, have a strongly revolute margin and a smooth lower surface with indistinct lateral veins, and are ovate with rather acute apex. The leaflets of the pinnate leaves on the vegetative shoots are also rather leathery with indistinct lateral venation. 8. Arthrophyllum pacificum Philipson, Gard. Bull. Sing. 30 (1977) 306. A slender, glabrous tree to 14 m. Leaves multi- jugate, c. 60 by 24 cm; of the flower-bearing branches smaller with fewer pinnae or simple; petioles c. 22 cm, 3^ mm 0; petiolules c. 1 cm; leaflets elliptic, oblong or ovate, c. 8-10 by 4-5 cm, rather membranaceous, margin entire, very slightly revolute, base abruptly cuneate, often oblique, apex obtuse, acute, or slightly apiculate (in a specimen from Morotai attenuated). Inflorescence a whorl of specialized leafy branches forming a terminal crown to the vegetative shoots; main rays 35 cm, bearing small pinnate leaves in opposite pairs and with flowering branches in the axils, ending in a whorl of secondary rays, subtended by a whorl of simple leaves (bracts) ; secondary rays 10-24 cm long, bearing simple or trifoliolate leaves usually in opposite pairs, and terminating in umbellules; umbellules c. 7-12-flowered; pedicels c. 5 mm at anthesis, slightly elongating in fruit, minute bracts caducous. Petals 5, broadly triangu- lar, 2V2-3 mm long. Stamens 5, anthers reniform. Ovary obconical, IV2 mm long; disk fleshy, furrowed when dry; stigma ± sessile. Fruit ellipsoidal, fleshy, somewhat oblique, conical beak with calyx and stigma, c. 9 by 5 mm when dry. Distr. E. Malesia: Moluccas (Morotai) to the Bismarck Archipelago. Ecol. Primary forest on mountain slopes, attaining Nothofagus mossy forest, 500-2000 m. Reported as common in most localities. Vern. Kainsoka, Ambai, Japen I. Notes. All specimens from islands to the north of New Guinea are similar, in having more delicate foliage than \0. A. macranthum from the mainland of New Guinea. The specimens from Morotai are sterile, and have narrower more tapering leaflets. The bark is light brown and the cut branches exude a clear latex. The flowers are light green with yellow anthers, and the fruit is black. 9. Arthrophyllum diversifolium Bl. Bijdr. (1826) 879; DC. Prod. 3 (1830) 266; MiQ. Fl. Ind. Bat. 1, 1 (1856) 767; Sum. (1861) 340, ind. var. lanceolata MiQ.; Clarke, Fl. Br. Ind. 2 (1879) 733; K. & V. Bijdr. 7 (1900) 46; Koord. Exk. Fl. Java 2 (1912) 717; Atlas 4 (1916) f. 675 & 676; Back. & Bakh. /. Fl. Java 2 (1965) 169; Stone, Gard. Bull. Sing. 30 (1977) 135; Philipson, I.e. 306, f. 15. — A.javani- cum Bl. Bijdr. (1826) 879; DC. Prod. 4 (1830) 266; Back. & Bakh. /. Fl. Java 2 (1965) 169. — A. ellipticum Bl. Bijdr. (1826) 879; DC. Prod. 3 (1830) 266. — Mormoraphis sumatrana Jack ex Wall. Cat. (1831) «. 4931, «owe«. — A. blumeanum Z. & M. Syst. Verz. (1846) 41, nam. illeg.; MiQ. Fl. Ind. Bat. 1, 1 (1856) 768; Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 27, ind. var. oblongatum MiQ., var. ellipticum (Bl.) Miq. et var. ovalifolium (JuNGH. & DE Vriese) Miq. — A. ovatifoUum JuNGH. & DE Vriese, Ned. Kruidk. Arch. 1 (1846) 19; Ann. Sc. Nat. Paris III, 6 (1846) 117; Miq. Fl. Ind. Bat. 1, 1 (1856) 768, t. 14 {'ovalifolium'); Ridl. Fl. Mai. Pen. 1 (1922) 885; Stone, Gard. Bull. Sing. 30 (1977) 136, f. 2. — A. dilatatum Miq. Fl. Ind. Bat. 1, 1 (1856) 768. — A. borneense Baker, Kew Bufl. (1896) 23; Merr. En. Bom. (1921) 457. — A. congestum Ridl. J. Fed. Mai. St. Mus. 10 (1920) 137; Fl. Mai. Pen. 1 (1922) 887; Stone, Gard. Bull. Sing. 30 (1977) 136. — A. havi- landii Ridl. Kew Bull. (1933) 494. — A. rufosepa- lum Ridl. Kew Bull. (1946) 40. — A. rubiginosum Ridl. I.e. 41. — Fig. 17, 21c, 22, 23. Small tree, up to 14 m, young parts with rufous tomentum. Leaves clustered at the ends of the branches, imparipinnate or bipinnate (rarely tripinnate) with leaflets at the insertion of the lateral rachides, multijugate, 150 by 45 cm (wider in bipinnate leaves); usually in opposite pairs on 1979] Araliaceae — 1 (Philipson) 61 Fig. 22. Arthrophyllum diversifolium Bl. Diagrammatic sketch of habit, showing vegetative and flowering branches, x Vso (Courtesy Card. Bull. Sing. 30, 1977). the inflorescence-bearing branches and smaller with fewer pinnae or unifoliolate ; petiole up to 40 cm; petiolules V2-IV2 cm; leaflets ovate-oblong or elliptic, up to 24 by 11 cm (leaflets of bipinnate leaves usually c. 10 by 5 cm), r: coriaceous or (especially in bipinnate leaves) somewhat mem- branaceous, margin slightly revolute, base truncate, rounded, or cuneate, often oblique, apex shortly acuminate, veins usually 5-7 pairs. Inflorescence a cluster of specialized leafy branches forming a terminal crown which abscisses after fruiting; main rays up to 150 cm, bearing pinnate (or more rarely unifoliolate) leaves mostly in opposite pairs and usually with flowering branches in the axils of the upper leaves, ending in an umbel of secondary rays subtended by a whorl of pinnate or unifoliolate leaves; secondary rays up to c. 30 cm, bearing mainly simple leaves in opposite pairs with flowering branches in their axils and terminating in compound umbellules each subtended by a whorl of often caducous bracts; tertiary rays (peduncles) c. 5 cm, articulated about the middle; umbellules with c. 30-40 flowers, c. 17-20 mm 0; pedicels c. 20, c. 5 mm at anthesis (longer in fruit). 62 Flora Malesiana [ser. I, vol. 9^ Fig. 23. Arthrophyllum divers if olium Bl. Schematic drawing of a single flowering shoot, with four degrees of branching, / Vs (Courtesy Gard. Bull. Sing. 30, 1977). 1979 Araliaceae — I (Philipson) 63 with an involucre of minute caducous bracts. Petals 5, c. 2 mm long. Stamens 5, anthers curved. Ovary turbinate, often inconspicuous at anthesis; disk fleshy, rising in the centre to a sessile stigma. Fruit ellipsoidal, f. 9 by 7 mm; calyx and stylo- podium forming a conspicuous beak. Distr. Malesia: Sumatra, Malay Peninsula, Java, Borneo, Celebes. Ecol. In a wide variety of habitats, on dry sandy soil to swampy humus, in primary lowland and montane rain-forest and also in secondary forest, heath-forest and waste land, from sea-level to 1600 m. Uses. Concoctions of the root and bark are reported to have medicinal properties, including a remedy for syphilis, and the plant has stupifying and poisonous properties. Vern. Sumatra: antjaneudeimg uding, bidju, bolu bolii, bulu, (kayu) abang-abang, k. attu tunit, legung, Riouw Arch., mapang, miii, obang, potah, silanta, tenjam dakan, Banka, tocrah. Malay Peninsula: chindangan utan, jolok hantu, lupa dahan, (pokok) restong, segan budahan, susun kelapa, turn bong ninyor, tusum perpah. Java: dajo, delahan, deleg, delek, djangkoirang, gombong, {kayu) gompang, kedrja, kidjangkurang, kingom- pang, ki ompong, klentjang, krepang, langitj, malas berdahan, pongporang, putjangan. Borneo: merje- meh, Sarawak, karadjungjung, Kalimantan. Celebes: kambabah, susangkangan. Notes. The very widespread A. diversifolium is variable in many characters, and may comprise a number of geographic subspecies, but no basis for this is apparent at present. Most individuals have the lower leaves simply imparipinnate, whereas others have bipinnate, or rarely tripinnate, leaves. The flowers and inflorescences of these forms appear to be identical, though rapid changes in the umbellules after flowering produce a deceptively distinctive appearance in specimens at diff"erent stages of development. Field experience over the whole range of the species will be required to understand this interesting leaf-polymorphism. In treating all forms as one species I am partly influenced by the fact that most authors who have been familiar with the plants in Java (where both forms occur) have regarded the complex as a single species (the fact that some authors have recognized the variant from Mt Salak as a distinct species does not aff"ect the problem of leaf-polymorphism). Apart from the strikingly diff'erent leaf forms just discussed, certain local variants may eventually be shown to justify specific rank. A form growing on Mt Salak (near Bogor) has often been regarded as distinct (see, for example, Hochreutiner, CandoUea 2, 1925, 481, and Backer & Bakhuizen VAN DEN Brink/. F1. Java 2, 1965, 169). Indeed this form is the basis of the name A. diversifolium. I retain this name in preference to the other two names published simultaneously by Blume because it has been most consistently adopted since it was first used in this comprehensive sense by Clarke, I.e. On the evidence available I do not consider the Salak plants any more distinctive than many other local variants. It might be considered that 5. A. engganoense is also no more than another such variant, but its facies is so marked that specific rank appears justified. It is possible that Ridley was correct in distin- guishing A. eongestum, but the material is not good and appears inadequate to confirm specific status. Five collections from Brunei and a neighbouring district of Sarawak are all very alike and suflFiciently distinct from both A. diversifolium and 7. A. crass um to suggest that they represent a separate taxon, but for the present they are tenta- tively retained as a form of y4. diversifolium. Similarly, the two collections described by Ridley as A. rubiginosum and A. rufosepalum are based on collections which are not altogether typical of A. diversifolium, but which come closest to that species. In the absence of more supporting material, it is advisable not to retain them as species. The first of these names {A. rubiginosum) has been widely used in identifications of Bornean specimens, but the specimens concerned are either typical A. diversifolium or belong to the distinctive 1 . A. crass um. Specimens from Mt Kinabalu described by Ridley as A. havilandii have bipinnate leaves, and appear to conform well with A. diversifolium. This form was again collected on Mt Kinabalu by Clemens and is also known from Sarawak. The smooth bark is whitish to greyish brown with pustulate lenticels; the wood is cream, with a colourless aromatic exudate. The flowers are yellowish with a sickly sweet scent. Seedlings have simple and trifoliolate leaves. 10. Arthrophyllum macranthum Philipson, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 18; Gard. Bull. Sing. 30 (1977) 308, f. \6. — A. diversifolium (non Bl.) Harms, Bot. Jahrb. 56 (1920) 413. — Fig. 21d. Tree up to 25 m, sparsely branched with leaves crowded at the ends of the branches, all young parts with dense rufous tomentum which may persist on the flowers and inflorescence. Leaves muhijugate, up to 100 by 30 cm; of the flower- bearing branches smaller with fewer pinnae, or simple; petioles stout, up to 40 cm, clasping base heavily lenticellate; petiolules V2-2 cm; leaflets ovate to oblong, up to 16 by 8 cm, coriaceous, mar- gin revolute, base rounded, truncate, or cordate, very rarely cuneate, often oblique, apex obtuse or bluntly apiculate, principal veins arched-ascending, reticulations visible especially beneath (pinnae of leaves on the flowering branches usually elliptic with a cuneate base). Inflorescence a whorl of specialized leafy branches forming a terminal crown; main rays up to 60 cm, bearing pinnate leaves often in opposite pairs and with flowering 64 Flora Malesiana [ser. I, vol. 9^ branches in the upper axils, and ending in a whorl of secondary rays subtended by a whorl of usually simple leaves (bracts); the secondary rays c. \0- 20 cm, bearing simple leaves, usually in opposite pairs, and terminating in compound umbels; umbellules (c. 2 cm) with c. 10-15 flowers; pedicels stout, 3-5 mm to 10 mm or more in fruit, sub- tended by minute bracts. Flower buds 5 mm or more long; calyx a rim or with 5 indistinct teeth. Petals 5, triangular, c. 4 mm long, fleshy. Stamens 5. Ovary obconical, c. 2V2 rnm long; disk fleshy (hemispherical in living material, conical and furrowed when dry); stigma capitate, ± sessile. Fruit ellipsoid, sometimes slightly oblique, c. 12 by 8 mm, the stylopodium forming a conical beak with the persistent stigma; exocarp fleshy, endo- carp cartilaginous. Distr. Malesia: New Guinea (from Irian Jaya to Milne Bay Distr. and the Bismarck Archipelago). Ecol. Usually a sub-canopy tree of rain-forest ranging from the lower montane zone to mossy subalpine woodland and scrubland, occasionally in second growth, usually above 1000 m (up to 2700 m), but occurring also on the coastal scarps of the Astrolabe Range. Vern. Maguva, W. Sepik, agagwa, agare, agugwa, alolo, angga, engga,pooli, tipilan, W. High- lands, arua, hagegoa, wonkurumeh, E. Highlands, kolom, S. Highlands. Note. Trees become very different in appearance when in flower or fruit: the spiral pinnate foliage leaves are surmounted by tufts of branches which end in inflorescences and bear much smaller leaves. The foliage leaves are fleshy, leathery and glossy. The ripe fruit is purple and shining. The bark is grey, at first smooth with many leaf-scars and lenticels, but small longitudinal fissures develop. The cut branches exude a brown latex and a scent of celery. The soft wood is white or straw-coloured. 11. Arthrophyllum papyraceum Philipson, Gard. Bull. Sing. 30 (1977) 308. Shrub, rufous-tomentose on the young parts. Leaves alternate, imparipinnate; petiole 15-20 cm; leaflets c. 7, membranaceous, elliptic, c. 12-24 by 5-10 cm, base broadly cuneate, apex finely acuminate, margin slightly revolute. Inflorescence a terminal compound umbel; primary rays few (2), 2-3 cm long, 2 mm wide, without bracts (caducous), secondary rays few (3), c. 13-18 mm long, articu- lated about the middle, ending in an umbellule of c. 10-12 flowers; pedicels 2-3 mm, slightly fur- furaceous. Petals 5, c. 2 mm long in bud. Stamens 5, anthers curved. Ovary turbinate, glabrous, obscurely ribbed. Fruit unknown. Distr. Malesia: E. Sumatra (East Coast Res. near Aek Sordang), one collection. Ecol. Primary rain-forest. Note. Known from a single collection (with no duplicates) this species resembles 12. A. maingayi in its simple inflorescence, and the few pinnae of its foliage leaves. However, the large size of the leaflets precludes its inclusion in that species. 12. Arthrophyllum maingayi Philipson, Gard. Bull. Sing. 30 (1977) 309, f. 7. — A. pinnatum Clarke, F1. Br. Ind. 2 (1879) 734, excluding synonyms [see also Seem. J. Bot. 4 (1866) 294]; RiDL. Fl. Mai. Pen. 1 (1922) 886. — Fig. 19c, 20a. Low shrub or slender tree, rarely as high as 10 m, rufous-tomentose on the very young parts, soon becoming glabrous. Leaves tufted at the ends of the branches, usually with 5-7 leaflets (but up to 15), up to 30 by 18 cm; petiole to 12 cm (usually shorter), 2 mm 0; petiolules c. 0-10 mm; leaflets elliptic or elliptic-lanceolate (occasionally the lowermost pair of leaflets is replaced by pinnate leaf segments), c. 8 by 3V2 cm, rather thin, base cuneate or rounded, apex apiculate or caudate, margin slightly revolute, lateral veins faint and obscure; leaves below the flowering branches sometimes reduced to 3 or 1 leaflet(s). Inflorescence consisting of a number of primary branches radiating from the end of a leafy shoot (which forms a longer or shorter peduncle); primary branches, often rather few, usually 3-6 cm long and devoid of leaves except for a few terminal simple or trifoliolate leaves around the compound umbels, occasionally the branches bear pairs of opposite leaves when they may be up to 30 cm long; secondary rays c. 2-3 cm long; pedicels 4-10 cm. Petals 5, P/2-2 mm long in bud. Stamens 5. Ovary turbinate, disk fleshy. Fruit spheroidal, c. 8 by 6 mm, calyx and stylopodium rather inconspicuous. Distr. Malesia: Central W. Sumatra (Mt Kerintji), throughout the Malay Peninsula (incl. Penang) and Borneo. Vern. Poko minta anak, Kedah, karon baru, Sumatra. Notes. The name 'A. pinnatum', misapplied by Clarke, I.e., has been in general use for this species. However, this name was based on Panax pinnatum Lamk, which in turn was based on the Rumphian name 'Scutellaria secunda'; this is quite a distinct plant (see under 6. Polyscias cumingiana). The three species A. maingayi, 16. /i. montanum and 17. A. alternifolium are similar in having simpler inflorescences than 9. A. diver sifolium. A. montanum can be distinguished from A. main- gayi by its more leathery leaves with more promi- nent nervation, and by the more woody and leafy flower-bearing primary branches of the inflores- cence. A. alternifolium is distinguished from both these species by its small, coriaceous leaflets with incon- spicuous nervation. Most specimens can be readily distinguished, but a few can be assigned to a species only doubtfully, usually because the material is inadequate. This is usually due to the junction between the vegetative (spiral phyllotactic) shoots and the flower-bearing branches being omitted. 1979 Araliaceae — I (Philipson) 65 13. Arthrophyllum kjellbergii Philipson, Gard. Bull. Sing. 30 (1977) 309, f. 10. — Fig. 20d. Small tree, 10 m, branches c. 1 cm 0, young parts covered by rufous tomentum. Leaves clustered towards the ends of the branches, multijugate, 25-30 by 16 cm; petioles c. 8 cm, 2 mm wide; petiolules c. 3-8 mm; leaflets elliptic or ovate- oblong, up to 8 by 3', 4 cm, chartaceous, margin minutely revolute, base rounded to cuneate often oblique, apex tapered to a blunt apiculum, or rounded and mucronate. Inflorescence a terminal cluster of c. 5-10 specialized branches (primary rays); primary rays c. 15-20 cm, 1^1 2-2 mm wide, bearing near the middle an opposite pair of simple leaves or sometimes trifoliolate leaves with short flowering branches in their axils, and sometimes with a second pair higher up, and 2-3 similar leaves below the terminal cluster of secondary rays; secondary rays c. 12, c. l^li-Vli cm long, each subtended by a small bract and bearing opposite caducous bracts near the middle, terminating in an umbellule of c. 12 flowers surrounded by an involucre of caducous bracts (1 mm long). Flowers known only in young bud. Fruit spheroidal, c. 5 by 4 mm, calyx and stylopodium prominent; pedicel 5-6 mm. Distr. Malesia: SE. Celebes (Kendari). Ecol. Primary rain-forest, 50-150 m. Note. The small leaves and relatively simple inflorescences are distinctive. 14. Arthrophyllum cenabrei Merr. Philip. J. Sc. 20 (1922) 417; En. Philip. 3 (1923) 235; Philipson, Gard. Bull. Sing. 30 (1977) 309. Glabrous tree, c. 10 m, ultimate branches c. 5 mm 0. Upper leaves pinnate, up to 10 cm long, leaflets mostly 5, sometimes 3, or the uppermost reduced to simple leaflets, the rachis and petiole c. 4 cm; leaflets mostly elliptic, 4V'2-6 by 2'/2-3^'2 cm, chartaceous to subcoriaceous, very shortly and obtusely acuminate, base acute, brownish oliva- ceous and slightly shining when dry, nerves 3-4 pairs, slender; petiolules 5-10 mm. Peduncles c. 4 cm, umbellately arranged at the tops of the branchlets, usually however with solitary inflorescences in the axils of the uppermost leaves, thus forming a somewhat leafy inflorescence. Fruits 5-8 in each umbel, ovoid, c. 7 mm ; pedicels 8-10 mm. Distr. Malesia: Philippines (Cebu; FB 28343, type, not seen). Ecol. On slopes at 600 m. Vern. Bingliu, C.Bis. Note. No specimen of this species has been located. The above description is taken from Merrill's original account. In placing this species in the key, it has been assumed that the inflorescence branching is relatively simple. 15. Arthrophyllum pulgarense Elmer, Leafl. Philip. Bot. 7 (1915) 2551 ; Merr. En. Philip. 3 (1923) 235; Philipson, Gard. Bull. Sing. 30 (1977) 311, f. 9. — Fig. 20c. Small tree, branches c. 1 cm 0, young parts with red tomentum, becoming glabrous except on the ovaries. Leaves clustered towards the ends of the branches, leaflets c. 6 pairs, c. 22 by 8 cm; petioles c. 6 cm, 3 mm 0; petiolules 5-6 mm; leaflets elliptic to rotund, c. 4 by 2^ 4 cm, coriaceous, margin revolute, rounded to broadly cuneate, ap»ex rounded or abruptly tapered to a short obtuse apiculum. Inflorescence a terminal cluster of specialized branches (primary rays); primary rays c. 6-10 cm, 3-4 mm 0, bearing near the middle an opposite pair of simple rotund leaves, sometimes with flowering branches in their axils, and with a whorl of similar leaves below the terminal cluster of secondary rays; secondary rays c. 6-8, 2-4 cm long, articulated about the middle with scars of bracts or bearing a pair of small simple leaves terminating in an umbellule of c. 8-12 flowers; bracts caducous. Calyx with indistinct teeth. Petals 6, 2* '2 rnm long (in bud). Stamens 6, anthers curved. Ovary obconical, 2 mm long, furfuraceous. Fruit ellipsoid, c. 8 by 5 mm, the stylopodium forming a beak with stigma and calyx; exocarp fleshy. Distr. Malesia: Philippines (Palawan: Mt Pulgar). Ecol. Common in montane forest on Mt Pulgar. Note. The coriaceous, small, often rotund leaf- lets are characteristic. 16. Arthrophyllum montanum Ridl. J. Fed. Mai. St. Mus. 4 (1909) 24; Fl. Mai. Pen. 1 (1922) 886; Philipson, Gard. Bull. Sing. 30 (1977) 311, f. 8. — A. nitidum Ridl. J. Fed. Mai. St. Mus. 7 (1916) 42; Fl. Mai. Pen. 1 (1922) 886. — A. ovatum Ridl. J. Fed. Mai. St. Mus. 7 (1916) 42; Fl. Mai. Pen. 1 (1922) 886. — Fig. 19d, 20b. Shrub or small tree to 6 m, unbranched or sparingly branched, rufous-tomentose on the young parts, becoming glabrous. Leaves tufted at the ends of the branches, multijugate, c. 30-55 by 12-22 cm; petiole 9-21 cm, 3 mm 0; petiolules c. 10-15 mm; leaflets elliptic or oblong, 6-10 by 2\'2^ cm, coriaceous or chartaceous, base cuneate, apex with a short blunt apiculum, margin entire, revolute, the few principal lateral veins usually rather prominent; the leaves associated with the umbels usually unifoliolate, broadly ellip- tic to rotund, with a petiole to A^ij cm. Inflores- cences on specialized leafy branches either in terminal clusters or axillary in the upper leaves; branches 10-30 cm, leaves mostly simple in opposite pairs, usually without flowering branches in their axils, branches ending in a whorl of simple leaves surrounding a compound umbel; primary rays c. 5-15(-20), 2-6 cm, with scars of caducous bracts; pedicels 4-8 mm. Petals (4-)5, 2 mm long in bud. Stamens (4-)5. Ovary turbinate; disk fleshy, stigma ^ sessile. Fruit spheroidal, c. 8 by 66 Flora Malesiana [ser. I, vol. 91 Fig. 24. Gastonia senatifolia (Miq.) Philipson. a. Habit, /, V3, b. lower bud, c. flower, c/. fruit, both x 5 {a-c ScHMUTZ 3612, d van Royen 4090). Drawn by Helene Mulder. 1979 Araliaceae — I (Philipson) 67 5 mm, calyx and stylopodium rather prominent. Distr. Malesia: Malay Peninsula (Kedah to Selangor). Ecol. Primary forest ascending to the montane zone, and in second-growth, 250-1500 m. Note. Similar to 12. A. maingayi but dis- tinguished by the more leathery leaves and by the leafy inflorescence branches which are usually absent in A. maingayi. 17. Arthrophyllum alternifolium Maingay ex RiDL. Fl. Mai. Pen. 1 (1922) 886; Philipson, Card. Bull. Sing. 30 (1977) 311, f. 11. — /i. pinnatum Clarke, FI. Br. Ind. 2 (1879) 734, p.p., e.xcl. basionym; King, J. As. Soc. Beng. 67, ii (1898) 59, p.p. — A. alternifolium Maingay ex Clarke, Fl. Br. Ind. 2 (1879) 734, nomen in synon. — Fig. 20e. Slender, sparingly branched shrub to 2 m, rufous-tomentose on the young parts, becoming glabrous. Leaves tufted at the ends of the branches, multijugate, c. 20-25(-30) by 9-12(-15) cm; petiole terete, c. 3-6(-9) cm, 2-3 mm ; petiolules c. 2 mm; leaflets ovate, elliptic or lanceolate, 3V2-^(-6) by l-2(-2'/4) cm, coriaceous, base cuneate, apex acuminate to caudate, obtuse, margin revolute, veins obscure; leaves associated with the umbels (if any) reduced, with fewer leaflets or unifoliolate, sometimes broadly ovate. Inflorescence usually a terminal compound umbel, occasionally a whorl of leafy branches (5-14 cm long) (leaves usually simple in opposite pairs), each ending in a compound umbel; peduncle IV2- 4'/2 cm, with one or more usually caducous simple (or trifoliolate) leaves at the apex; primary rays c. 5, 3^'/2 cm, with scars of caducous leaves about the middle, each ending in an umbellule of c. 12-25 flowers, pedicels 5-8 mm. Calyx a rim or minutely 4-5-dentate. Petals 4-5, 2 mm long in bud. Stamens 4-5. Ovary turbinate; disk fleshy, stigma - sessile. Fruit spheroidal, c. 5 by 5 mm when dry, calyx and stylopodium small. Distr. Malesia: Malay Peninsula (Johore: Mt Ophir; Pahang, Selangor, and Malacca). Ecol. In shady montane forest, with Rhodo- dendron and Dacrydium, 900 m and above. Note. The small, coriaceous, often apiculate leaflets are characteristic. Although collected most frequently on Mt Ophir it occurs on other high ridges in southern Malaya. Excluded Arthrophyllum ceylanicum MiQ. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 27 (type in L), is according to VAN Steenis, Rec. Trav. Bot. Neerl. 24 (1927) 819 = Oroxylum indicum (L.) KuRZ (Bignoniaceae). Arthrophyllum reiiculatum Bl. ex Miq. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 27 (type in L), is according to Miquel {I.e. 318) and van Steenis {vide supra) = Oroxylum indicum (L.) KuRZ {Bignoniaceae). 9. GASTONIA CoMM. ex Lamk, Encycl. 2 (1786) 610; Miq. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 5; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 43; Hutch. Gen. Fl. PI. 2 (1967) 68; Philipson, Blumea 18 (1970) 491, 497, f. 1-10. — Tetraplasandra {non A. Gray) Miq. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 4; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 29, p.p. ; Nachtr. 2 (1900) 253. — Indokingia Hemsl. in Hook. Ic. PI. (1906) t. 2805. — Peekeliopanax Harms, Notizbl. Berl.-Dahl. 9 (1926) 478, fig. — Fig. 24, 26, 27. Trees unarmed with thick branches, glabrous or tomentose. Leaves large, im- paripinnate, exstipulate; rachis articulated; leaflets in pairs entire or crenate; petiole terete, with clasping base. Flowers in umbellules which are arranged race- mosely, or in verticils, on strong inflorescence branches; pedicels not articulated below the ovary. Calyx forming a continuous rim with an entire or indistinctly denticulate margin. Corolla of 5-13 free petals or calyptrate, fleshy, valvate. Stamens either equal in number to the petals or up to several times as many; filaments usually rather short and thick; anthers large, often irregularly lobed, dorsifixed. Ovary inferior, broadly obconic, cells 7-22; disk fleshy with stylar processes equal in number to the cells arising from its centre. Fruit a spherical berry (strongly ribbed when dry), with an indistinct calyx rim, a flattened disk, and a 68 Flora Malesiana [ser. I, vol. 9^ prominent stylopodium bearing a ring of radiating stigmatic arms ; exocarp fleshy, endocarp crustaceous. Endosperm with smooth surface. Distr. About 10 spp. in East Africa, Madagascar, the Seychelles and Mascarenes, Malesia, and the Solomon Is. Ecol. Primary and second-growth forest, or in open country, from sea-level to lower montane zone. Taxon. I have amply analyzed and discussed the affinities of Gastonia (Blumea 18, 1970, 497). I have come to the conclusion that within its alliances Gastonia is the only genus west of Samoa; in Polynesia there are three other closely allied genera. KEY TO THE SPECIES 1. Corolla with free petals. Ovary cells and style arms 6-9(-12). Whole plant glabrous. Leaflets usually entire or sparsely crenate. Main inflorescence branches borne along an elongated axis with caducous bracts; peduncles of the umbellules mostly aggregated into pseudo-whorls .... 1. G. serratifolia 1. Corolla calyptrate. Ovary cells and style arms 12-18(-22). Young parts with scurfy tomentum, per- sisting on the ovary and bracts. Leaflets strongly crenate. Main inflorescence branches borne sub- umbellately on a short axis with persistent bracts ; peduncles of the umbellules scattered 2. G. spectabilis 1. Gastonia serratifolia (Miq.) Philipson, comb, nov. — Arthrophyllum serratifolium Miq. Sum. (1861) 341, type from Sibolga, leg. Teysmann (in U). — G. papuana Miq. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 5; Harms, Bot. Jahrb. 56 (1921) 408; Philipson, Blumea 18 (1970) 492, 500 f. 3. — Tetraplasandra paucidens Miq. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 4; Koord. Minah. (1898) 488. — G. eupteronoides T. & B. Nat. Tijd. N. I. 25 (1863) 416. — Polyscias papuana (Miq.) Seem. J. Bot. 3 (1865) 181. — Tetraplasandra koordersii Harms, Ann. Jard. Bot. Btzg 19 (1904) 12; Ic. Bog. 2 (1906) t. 178. — Tetraplasandra philippinen- sis Merr. Philip. J. Sc. 1 (1906) Suppl. 219; En. Philip. 3 (1923) 222. — G. winkleri Harms in Fedde, Rep. 15 (1917) 20. — Tetraplasandra solomonensis Philipson, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 11.— Fig. 24. Shrub or small tree, sometimes epiphytic, up to 27 m, with clear bole to 15 m, dbh 34 cm, crown sparsely branched with leaves crowded at the ends of the branches, glabrous. Leaves up to 80 by 20 cm; petiole c. 13 cm; leaflets c. 10 pairs, petiolules up to 1 cm; blade oblong, ovate or lanceolate, middle leaflets 8-14(-18) by 2'/2- 3V2(-8) cm, chartaceous, entire and subrevolute, or with a few obscure, more rarely several promi- nent crenations, apex rounded and bluntly apicu- late or tapering and acute, base broadly cuneate, midrib prominent. Inflorescence terminal, glab- rous, with a stout primary axis 15-25 cm long, bearing scattered or clustered branches along its length and ending in an umbel of c. 10 branches; bracts caducous; secondary branches 30^0 cm, bearing numerous subverticillate peduncles along their length and ending in an umbel; peduncles c. 3 cm, somewhat longer in fruit; pedicels I-2V2 cm, forming umbellules of c. 10 flowers. Flower buds (when dry) c . 7 by 3 mm. Calyx rim undulate. Petals 5-9, slightly fleshy, fully separated. Stamens variable in number, (7-) 14-55; filaments short ; anthers broad and irregularly lobed, variable in size. Ovary glabrous, smoothly rounded below, slightly constricted below the calyx; cells 6-12, usually c. 9, disk with a prominent rim and, at anthesis, a central boss formed by closely appressed subulate style arms equal in number to the ovary cells. Fruit c. 9 by 7 mm (without stylopodium), the flattened stylopodium ending in a ring or double row of radiating subulate stigmatic arms, black when ripe, the fleshy exocarp enclosing com- pressed crustaceous pyrenes. Distr. Solomon Is.; in Malesia: Central W. Sumatra (Sibolga, Enggano I.), Malay Peninsula (Johore), Sunda Straits (islet Dwars in den Weg), West Java, Lesser Sunda Is. (Sumba, Timor, Wetar, Flores), W. Borneo (east of Pontianak, Karimata), N. Borneo (Sabah), Philippines Fig. 25. Distribution of Gastonia in Malesia. Known localities of G. serratifolia (Miq.) Philip- son: black dots; range of G. spectabilis (Harms) Philipson: broken line. 1979] Araliaceae— 1 (Philipson) 69 Fig. 26. Gastonia spectabilis (Harms) Philipson, two inflorescences visible below the leaves (Photogr. Philipson, Kassam Pass, E. New Guinea, 1968). 70 Flora Malesiana [ser. I, vol. 9^ (Palawan, Balabac), Celebes fLepo-Lepo, Luwuk, Minahassa), N. Moluccas (Talaud Is.), New Guinea (Vogelkop, NW. & SW. Irian, Schouten I., Waigeu I.). Fig. 25. Ecol. Primary and secondary forest, or in open country, usually at low altitude and often on the shore or sea-cliffs, but ascending to 1000 m. Eyma noted it to be a characteristic tree on Mt Tam- bunan, Luwuk, E. Celebes. Vern. Bajur talang ekoaho, M, Enggano,jarum, Sabah, raka, Sumba, kre, wangka, Flores, lampo pad, bungku, Celebes, bungio, Palawan, lantora, Talaud Is., mansnongoree, mantsenongor, Schouten I., raauwrack, ara-orach, Vogelkop, Maibrat. Note. The entire or serrate nature of the leaf margins, the number of ovary cells, and especially the number of stamens are variable characters but show no discernable geographical segregation. The outer bark is described as light brown, with small oblong brittle scales. Inner bark, leaves and inflorescences with copious sticky juice. Wood soft white. The petals are variously described as purple, light green, and white. 2. Gastonia spectabilis (Harms) Philipson, Blumea 18 (1970) 494, pi. 1. — Peekeliopanax spectabilis Harms, Notizbl. Berl.-Dahl. 9 (1926) 478, fig.; Philipson, Blumea 18 (1970) 500 f. 2. — G. bori- diana Harms, Bot. Jahrb. 69 (1938) 282. — Fig. 26, 27. Tree up to 40 m high, clear bole to 28 m, dbh 1 .75 m, crown sparsely branched with the branches whorled or regularly forked and the leaves crowded at the ends of the branches, all young parts with scurfy indumentum more evident in dried material. Leaves up to 80 by 30 cm; petiole c. 15 cm, with some tomentum remaining at the joints; leaflets c. 1 1 pairs on petiolules c. 2-8 mm, oblong or elliptic, middle leaflets 10-15 by 4-6 cm, charta- ceous when dry, prominently crenate, apex nar- rowed to a short blunt apiculus, base rounded or truncate. Inflorescence in forks well below the leaves, when in bud covered with large scurfy cataphylls, and when mature with radiating branches forming clusters c. 130 cm 0. Primary axis short (5-10 cm) with persistent bracts; secondary branches numerous, radiating, up to 65 cm, bearing small, persistent, scurfy bracts and peduncles scattered along their length and clustered in a terminal umbel; peduncles c. l''2-5 cm; pedicels V2-l'/2 cm, forming umbellules of c. 5-12 flowers. Flower buds c . 8 by 5 mm when dry (fresh c. 12 by 8 mm). Calyx rim straight. Petals 6-12, very fleshy, incompletely separated (often splitting into c. 5 lobes). Stamens 25-66, often c. 35, filaments short; anthers broad and irregularly lobed, variable in size. Ovary with a short dense indumentum, smoothly rounded below, cells usually c. 16, very rarely fewer than 10, as many as 22; disk at anthesis with a prominent rim and a central boss formed by closely appressed subulate Fig. 27. Gastonia spectabilis (Harms) Philipson. Young tree, free bole 19 m, 47 cm 0, crown 10 m, longest leaves 2 m, not yet flowering (Photogr. G. Peekel, New Ireland, Ugana, 1940). styles equal in number to the cells. Fruit c. 8 by 10 mm (dry and without stylopodium), the flattened disk with a prominent stylopodium ending in an elliptical ring of radiating subulate stigmatic arms; the fleshy exocarp enclosing compressed crusta- ceous pyrenes. Distr. Solomon Is.; in Malesia: New Guinea (Vogelkop, NW. Irian, Papua New Guinea), Bismarcks, and New Ireland. Fig. 25. Ecol. Primary and secondary rain-forest, and in cultivated areas, 200-2000 m. Vern. Amoriga, bekuak, djak, ntjier, tuju, Vogelkop, bohko, boinga, gabiel, jamwa, mestic, sikoto, tubat, tumbala, Madang Distr., waki, W. Highlands, aita, E. Highlands, kuhuh, Papua. Note. Possibly the largest araliad known. The regular habit of branching results from the vegeta- 1979 Araliaceae — I (Philipson) 71 Fig. 28. Polyscias nodosa (Bl.) Seem. a. Leaf and part of inflorescence, x Vs, b. leaflet x U, c- branch of inflorescence, x 'l„ d. umbellule, e. flower bud and ditto in LS, x 8,/. ovary and ditto in CS, x 8, g stamens (Versteegh BW 3868). Drawn by Helene Mulder. 72 Flora Malesiana [ser. I, vol. 9^ tive shoots springing in pairs or whorls from below Bole without buttresses or with buttresses 1 m high terminal inflorescence buds. Several flushes of and 2 m wide. Outer bark brown with prominent growth occur as an inflorescence bud matures, so pustular lenticels and small shallow fissures, that at anthesis the inflorescences are situated in Exudate from cuts abundant, clear and aromatic, forks well below the leafy crown, with a succession Wood soft. Flowers cream, stamens yellow. Ripe of younger inflorescence buds in higher forks. fruit dark red-brown. 10. POLYSCIAS J. R. & G. FORSTER, Char. Gen. (1776) 63, t. 32; DC. Prod. 4 (1830) 257; Seem. J. Bot. 3 (1865) 179; Bth. in B. & H. Gen. PI. 1 (1876) 941 ; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 43; Koord. Atlas 4 (1916) f. 677-680; Merr. Int. Rumph. (1917) 408; Harms, Bot. Jahrb. 56 (1921) 409; Merr. En. Philip. 3 (1923) 233; Hutch. Gen. Fl. PL 2 (1967) 75; Bernardi, Candollea 26 (1971) 13; Philipson, Blumea 24 (1978) 169. — Eupteron Miq. PI. Jungh. 3 (1855) 423; Fl. Ind. Bat. 1, 1 (1856) 762; Hutch. Gen. Fl. PI. 2 (1967) 68. — Nothopanax Miq. PI. Jungh. 3 (1855) 425; Bonplandia 4 (1856) 139; Fl. Ind. Bat. 1, 1 (1856) 765; Seem. Fl. Vit. (1866) 114; Merr. Int. Rumph. (1917) 409; En. Philip. 3 (1923) 233. — Irvingia F.v.M. Fragm. 5 (1865) 17, non Hook./. 1860. — Kissodendron Seem. J. Bot. 3 (1865) 201; ibid. 6 (1868) 129; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 45; Bot. Jahrb. 56 (1921) 412; Hutch. Gen. Fl. PI. 2 (1967) 75. — Palmervandenbroekia GiBBS, Arfak (1917) 162; Hutch. Gen. Fl. PI. 2 (1967) 75. — Gelibia Hutch. Gen. Fl. PI. 2 (1967) 57. — Fig. 28, 30. Unarmed shrubs or trees, glabrous or furfuraceous. Leaves imparipinnate or 2-3-pinnate (rarely unifoliolate) with an elongated or short sheathing base; rachis articulated; petiole terete; leaflets in pairs, entire, crenate or dentate. Inflorescence terminal, often large, a panicle, corymb, or compound rarely simple umbel. Flowers in umbellules, capitula, or racemose; pedicels articulated below the ovary. Calyx a rim with an undulate or dentate margin. Petals 4-5(-8 or more), valvate. Stamens equal in number to the petals; anthers dorsifixed. Ovary inferior, 4-5(-8 or more)-celled; disk fleshy; styles either free and recurved (at least in fruit) or joined to form a beak-like stylopodium. Fruit a spherical or ovoid drupe, crowned by the persistent calyx rim and the styles or stylopodium; exocarp fleshy, endocarp chartaceous. Endosperm with an uneven surface, fissured, or rarely smooth. Distr. About 100 spp. throughout the tropics of the Old World (incl. Australia), and the Pacific Islands; in Malesia 23 spp. of which 3 adventive: rare in West (none native in Sumatra and Malaya), more common in East, the majority in New Guinea. Fig. 29. Ecol. Primary or second-growth forest, from sea-level to 2650 m. Note. Araliads with pinnate leaves and an articulated pedicel are here regarded as forming one genus, divided into several sections. These sections have formerly been segregated as genera, but I follow Bernardi, I.e., in uniting them. The most distinctive is sect. Folyscias, characterized by an elongated leaf- sheath. Several of the species of this section are cultivated and have a number of cultivars. KEY TO THE SECTIONS 1. Leaf-sheath elongated, extending along the petiole for about ^U of its length (Spp. 1-7) 1. Sect. Folyscias 1 . Leaf-sheath short or obsolete, restricted to the base of the petiole. 2. Style arms spreading, at least in fruit. 1979] Araliaceae— I (Philipson) 73 3. Flowers arranged racemosely (Sp. 8) 2. Sect. Gelibia 3. Flowers arranged in umbellules or capitula (Spp. 9-14) 3. Sect. Eupteron 2. Style arms erect, fused, forming a beak in fruit. 4. Inflorescence large, ± as long as the leaves (ipp. 15-19) 4. 5fc/. Kissodendron 4. Inflorescence much shorter than the leaves (Spp. 20-23) 5. Sect. Palmervandenbroekia 1. Section Polyscias Philipson, Blumea 24 (1978) 169. Aromatic, glabrous shrubs or small trees, often cultivated. Leaf-sheath elon- gated along the petiole for V3-V4 of its length. Styles spreading, at least in the fruit. Distr. Polynesia, Queensland, Malesia, and SE. Asia. KEY TO THE SPECIES 1. Leaves 2-3-pinnate 1. P. fruticosa 1. Leaves imparipinnate, or unifoliolate. 2. Ovary 2-celled (or predominantly so). 3. Main inflorescence branches diff"usely branched 2. P. macgillivrayi 3. Main inflorescence branches with verticils of short branches 3. P. verticillata 2. Ovary variable, but many flowers with more than 2 cells. 4. Leaflets orbicular or reniform. Leaves unifoliolate or trifoliolate 4. P. Scutellaria 4. Leaflets ovate, oblong, or elliptic (cultivated forms often laciniate or lanceolate). Leaves with 3 or more pairs of leaflets. 5. Leaf margin sharply serrate (blade often rhomboidal and variegated with light yellow) 5. P. guilfoylei 5. Leaf margin entire, or obscurely dentate. 6. Peduncles of the ultimate umbellules bearing 1 or more pairs of small bracts (reduced flowering branches sometimes present in their axils) 6. P. cumingiana 6. Peduncles of the ultimate umbellules either without bracts, or with 1 or more small bracts inserted singly 7. P. javanica 1. Polyscias fruticosa (L.) Harms in E. & P. Nat. Merr.; Int. Rumph. (1917) 410; Sp. Blanc. (1918) Pfl. Fam. 3, 8 (1894) 45; K. & V. Bijdr. 7 (1900) 2; 295; Harms, Hot. Jahrb. 56 (1921) 412; Merr. En. Harms in K. Sch. & Laut. Fl. Schutzgeb. (1900) Philip. 3 (1923) 233; Heyne, Nutt. PI. (1927) 1209; 486; KooRD. Exk. Fl. Java 2 (1912) 716; Atlas 4 Ochse & Bakh. Veg. D.E.I. (1931) 64, f. 37; (1916) f. 680; Bailey, Rhodora 18 (1916) 153, incl. Corner, Ways. Trees (1940) 150. — Nothopanax var. plumata (Hort.) Bailey; Back. & Bakh. /. obtusum (Bl.) Miq. Fl. Ind. Bat. 1, 1 (1856) 766; Fl. Java 2 (1965) 168; Philipson, Blumea 24 (1978) Seem. Fl. Vit. (1866) 114. —P. obtusa (Bl.) Harms 169. — Scutellaria tertia Rumph. Herb. Amb. 4: in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 45, nom. illeg., 78, t. 33. — Panax fruticosum Linne, Sp. PI. non Blanco, 1837, quae est Schefflera odorata ed. 2 (1763) 1513; Bl. Bijdr. (1826) 830; DC. (Blanco) Merr. &Rolfe; Koord. Exk. Fl. Java 2 Prod. 4 (1830) 254; Blanco, Fl. Filip. ed. 2 (1845) (1912) 716; Atlas 4 (1916) t. 679 S; Back. & Bakh. 156; ed. 3, 1 (1877) 281, t. 78; Clarke, Fl. Br. Ind. /. Fl. Java 2 (1965) 168. 2 (1879) 725; F.-Vill. Nov. App. (1880) 101; Glabrous shrub or small tree, up to 5 m, with BoERL. Handl. 1 (1890) 647; F.v.M. Descr. Pap. branches bearing spirally arranged leaves towards PI. 9 (1890) 60; Ware. Bot. Jahrb. 13 (1891) 396; their ends. Leaves tripinnate, variable in size to Koord. Minah. (1898)488; Merr. Philip. J. Sc. 3 c. 75 cm long; petiole up to c. 25 cm, with a sheath- (1908) Bot. 84. — Panax obtusum Bl. Bijdr. (1826) ing base to 5 cm long; pinnae to c. 25 cm long; 880; DC. Prod. 4 (1830) 254. — Aralia tripinnata leaflets shortly petiolate, very variable in shape and Blanco, Fl. Filip. (1837) 223, cf. Merr. Sp. Blanc. size, oblong or linear-lanceolate, or 1-12 by V2- (1918) 295. — Nothopanax fruticosum (L.) Miq. 4 cm, more or less deeply serrate or lobed, or PI. Jungh. 3 (1855) 425; Fl. Ind. Bat. 1, 1 (1856) irregularly pinnatisect, apex attenuate, acuminate 765 ; Seem. Fl. Vit. (1866) 1 14, 1 15 ; J. Bot. 4 (1866) or rounded, base cuneate, truncate or subcordate, 363; Merr. Fl. Manila (1912) 358, incl. var. midrib and lateral veins evident. Inflorescence a plumatum (Hort.) Merr. et var. victoriae (Hort.) diffuse panicle; primary axis up to c. 60 cm with 74 Flora Malesiana [ser. I, vol. 9^ secondary branches mostly in verticils at intervals along its length; secondary branches up to 30 cm bearing umbellules in an irregularly branched system towards their extremities; umbellules with 12-20 flowers on pedicels c. 3 mm long. Calyx a minute rim. Petals 5, 2 mm long. Stamens 5. Ovary turbinate, c. 1 mm high, 2-3(-4)-celled; styles at first erect, later spreading. Fruit subglobose, fleshy, c. 5 mm when dry. Distr. Native country not properly known, cultivated throughout the region, and in other parts of the Indo-Pacific tropics. Ecol. Grown at low and moderate altitudes (c. 1000 m) as an ornamental or hedge shrub and for culinary use. Vern. (from various sources). Sumatra: orang aring, Medan, kedongdong mekka, Palembang; Malaya: daun girang, siku kluang, M; Java: kedongdong alus, k. batur, k. laut, M, imba, kedongdong laki, k. tjina, randa nunut, S, tjakar kutjung, t. tjikri, J, kadongdung, k. laut, k.petedhan, Md; Celebes: boka ula rise, kendem rintek, Minahasa, Alfur lang., bombu, Makassar: Philip- pines: papud. Tag., Bik., bani, makan, Bik.; Moluccas: daun papeda papua, pagar pagar, Ambon, guarbati, tampusong, Ternate. The name kedongdong belongs properly to species of the fruit tree gpnus Spondias, but is sometimes also applied to trees of other families with pinnate leaves; papua means 'curly'. Notes. The foliage of this plant is extremely variable in size and form. Typical plants are figured by KooRDERS (1916, I.e.) and by Ochse & Bak- HUIZEN VAN DEN BRINK (l.c). Thcsc may be readily identified by the intricately compound leaf. The individual leaflets are characteristically ovate-lanceolate and serrate to deeply pinnatifid. However, broader leaflets with simpler outlines are not uncommon. The more rotund, blunter leaflets of P. obtusa are considered here to be an extreme form of this species (the type is bipinnate). w#^oir?&-^:.j> Fig. 29. Species density of Polyscias J. R. & G. FoRSTER in Malesia; above the hyphen the number of endemic species, below it the non-endemics. Only the native species. Occasional specimens have leaves so much reduced that they are simply pinnate or unifoliolate when they approach forms of 6. P. cumingiana. A number of names have been applied to horti- cultural forms belonging to this section of the genus [cf. Bailey, Rhodora 18, 1916, 153), but the interrelations of these will be understood only after intensive biosystematic study of the many cultivars. 2. Polyscias macgillivrayi (See.m.) Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 45; Philipson, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 14; Blumea 24 (1978) 169. — Nothopanax macgillivrayi Seem. F1. Vit. (1866) 114, nomen. — Panax macgillivraei (Seem.) Bth. F1. Austr. 3 (1866) 382. — P. grandi- folia VoLKENS, Bot. Jahrb. 31 (1901) 471; Kanehira, En. Micron. PI. (1935) 385. — Tieghe- mopanax macgillivrayi Viguier, Bull. Soc. Bot. Fr. 52 (1905) 313. — Polyscias sp. C. T. White, J. Arn. Arb. 10 (1929) 255. Glabrous shrub or small tree, up to 15 m, with few branches bearing terminal clusters of large leaves. Leaves imparipinnate, up to 1 m (or more) long, uppermost leaves smaller; petiole c. 15 cm, with a sheathing base extending for 6-10 cm along the petiole; petiolules c. 10-15 mm.; lamina oblong often broader near the base (lower and the terminal pinnules more ovate) c. 20-25 by 8-10 cm, slightly succulent when fresh, margin entire, revolute or occasionally minutely dentate, apex rounded and shortly apiculate, base rounded, truncate or sub- cordate, midrib and widely spaced laterals promi- nent. Inflorescence a large panicle; primary axis stout, rather short {c. 3-10 cm), often bearing reduced leaves or cataphylls basally, and a few lateral inflorescence branches, terminating in an umbel of long diff"usely branched rays; lateral branches and rays c. 40-50 cm, with secondary branches c. 6-12 cm borne singly or in subverticils along their length; secondary branches with numer- ous tertiar>' branches bearing lateral and terminal umbellules (or the branching may be of a high order); umbellules with c. 6-12 flowers on delicate pedicels c. 3 mm long. Calyx a minute rim. Petals 5, 2^2 mrn long. Stamens 5, anthers oblong I'/j mm long, filaments delicate, IV 2 mm long. Ovary turbinate in bud, 1^2 rnm long, rapidly becoming rotund, compressed with prominent veins, 2-celled ; styles 2, divergent. Fruit fleshy, black, compressed, c. 5 by 6 mm; styles persistent, recurved. Distr. Micronesia, Solomon Is., Queensland; in Malesia: New Guinea (Papua, Eastern and Milne Bay Distr.; Territory of New Guinea, Morobe Distr., Musi I.), also in the D'Entrecasteaux and Trobriand Is., New Britain, and Louisiades. Ecol. Strand vegetation and littoral rain-forest, often behind mangrove. Vern. Aikove, Mimific lang., ane, D'Entre- casteaux Is., gambou, Musi I., Morobe Distr., nakaigwoo, Trobriand Is., raumonas, Onjob lang. 1979 Araliaceae — I (Philipson) 75 3. Polyscias verticillata Stone, J. Arn. Arb. 47 (1966) 272, f. 1 ; Philipson, Blumea 24 (1978) 169. Glabrous, small tree to 7 m with few branches bearing spirally arranged leaves towards their ends. Leaves imparipinnate, up to 1 m long; petiole c. 24-30 cm, terete with a sheathing base c. 6-7 cm long; petiole 5-20 mm, leaflets oblong often broader near the base, c. 16-27 by 5-13 cm, margin entire, slightly revolute or sparsely denticulate, apex acuminate, base subcordate or truncate, often oblique, midrib and lateral veins evident. Inflores- cence a large panicle; primary axis stout, rather short, with broad cataphylls, bearing several long radiating secondary branches in a sub-umbel; secondary branches c. 50 cm, with numerous tertiary branches borne in well defined verticils and in a terminal umbel, bracts triangular c. 8 mm long, caducous; tertiary branches c. A-1 cm with small bracts near the middle; umbellules with c. 10-15 flowers, on pedicels 1-4 mm long. Calyx a minute rim. Petals 5, 2'/2 mm long. Stamens 5, anthers oblong, c. 1-1 V2 mm long. Ovary turbinate, c. 1 mm long, 2-celled; styles 2, at first erect, later recurved. Fruit globose, fleshy, black, compressed, c. 4 by 7 mm when dry; styles persistent. Distr. Solomon Is.; in Malesia: New Guinea (New Britain, New Ireland, New Hanover, Admiralty Is., and Bagabag I.). Ecol. Usually near the beach or in lowland forest, to 140 m. Uses. The young foliage is eaten fresh or boiled. Vern. Babagula, valagur, Kuana dial., la bara kiku, Nakanai, tauwol, Gasmata, palu, Vairamana, vela vela, Pomio. Note. Cut surfaces produce a sweet scented sticky exudate. The petals are violet within, the fruits purple-black. 4. Polyscias Scutellaria (Burm. /.) Fosb. Un. Hawaii Occ. Pap. 46 (1948) 9; Stone, Taxon 14 (1965) 284; Philipson, Blumea 24 (1978) 169. — Scutellaria prima Rumph. Herb. Amb. 4: 75, t. 31. — Scutellaria secunda latifolia Rumph. I.e. 76. — Crassula Scutellaria Burm./. F1. Ind. (1768) 78. — Aralia cochleata Lamk, Encycl. I (1783) 224. — Panax scutellarioides Reinw. ex Bl. Bijdr. (1826) 880; Span. Linnaea 15 (1841) 208. — Panax cochleatum (Lamk) DC. Prod. 4 (1830) 253; BOERL. Handl. 1 (1890) 647. — Panax conchifolium RoxB. Fl. Ind. ed. Carey 2 (1832) 77. — Notho- panax cochleatum (Lamk) Miq. PI. Jungh. 3 (1855) 425; Fl. Ind. Bat. 1, 1 (1856) 766; Seem. J. Bot. 4 (1866) 296; Fl. Vit. (1866) 116; Koord. Minah. (1898) 490; Exk. Fl. Java 2 (1912) 717; Atlas 4 (1916) f. 697. — Nothopanax tricochleatum Miq. Fl. Ind. Bat. Suppl. (1860) 135, 340; Merr. Int. Rumph. (1917) 409; En. Philip. 3 (1923) 234; Ochse &BAKH. Veg. D.E.I. (1931) 69, f. 39A. — Panax rumphii Hassk. Abh. Naturf. Ges. Halle 9 (1866) 220. — Nothopanax scutellarium (Burm. /.) Merr. Int. Rumph. (1917) 409; En. Philip. 3 1923) 234; Heyne, Nutt. PI. (1927) 1209; Ochse & Bakh. Veg. D.E.I. (1931) 67, f. 39; Corner, Ways. Trees (1940) 156; Back. & Bakh./. Fl. Java 2 (1965) 169. — P. tricochleata (Miq.) Fosb. Phyto- logia 5 (1955)290. Glabrous shrub or small tree, up to 6 m, with branches bearing spirally arranged leaves towards their ends. Leaves simple or trifoliolate, variable in size ; petiole often c. 6 cm, but as long as 28 cm, with a sheathing base 1-6 cm long; lamina rotund or reniform, often c. 8 cm but as wide as 28 cm, margin usually serrate or becoming sub-lobed towards apex, in larger leaves the serrations often inconspicuous, apex rounded, base ± cuneate, midrib and lateral veins evident. Inflorescence a diff"use panicle; primary axis usually long (up to 1 m) with secondary branches (mostly in verticils, at intervals along its length; secondary branches 15-30 cm, bearing umbellules in an irregularly branched system towards their extremities; umbellules with c. 8-16 flowers, on pedicels c. 3 mm long. Calyx a minute rim. Petals 4-5, 2 mm long. Stamens 4-5, anthers oblong, 1 mm long. Ovary turbinate, c. 1 mm high, (2-)3^-celled, styles at first erect, later recurved. Fruit subglobose, fleshy, c. 5 mm when dry. Distr. Native country not properly known, possibly East Malesia; cultivated throughout the region, extending through the tropical Pacific. Ecol. Grown at low and moderate altitudes (800 m) as an ornamental or hedge shrub. Uses. The foliage is aromatic and is used to furnish perfume. Forms with variegated foliage are in cultivation. Used medicinally as a diuretic, against breast cancer, and to prevent baldness (Heyne, I.e.). Also for culinary purpose (Ochse & Bakh. I.e. 67). Vern. Cf. Heyne: memangkokan, (pohon) mangkok, Mai. (after the dish(= mangkok)-shaped leaves which are in the Moluccas sometimes used as dishes); Java: godong mangkokan, M, mama- mekan, S, puring mangkok, Md; Lesser Sunda Is.: lanido, ndalido, ndari, ramido, Roti; Celebes: (daun) mangko, tuwo mangku, M, bobohang, boku ula, kendem wewene, woworan, Manado, angko mangko, Bug. & Mak.; Philippines: salapiin, Mindanao; Moluccas: daun koin, d. papeda, M, Ambon, ai laun niwel, ai lohoi, Alf., Ambon, goma ma tari. Gal., Halmaheira, sawoko, Loda, Halmah., rau paroro, Ternate. Note. This species is usually readily distin- guished by its simple, orbicular, saucer-shaped leaves, but plants with some or all of their leaves trifoliolate occur. Stone (Micronesica 2, 1965, 51) advanced evidence for uniting these trifoliolate plants with P. pinnata (= 6. P. cumingiana) while retaining the unifoliolate plants as a distinct species. Possibly all are forms of one polymorphous species complex, but here it is considered convenient to adhere to the view which unites all forms having 76 Flora Malesiana [ser. I, vol. 9^ orbicular usually bowl-shaped leaves under the concept P. Scutellaria. 5. Polyscias guilfoylei (Cogn. & Marche) L. H. Bailey, Rhodora 18 (1916) 153; Stone, Micro- nesica 2 (1965) 57; Philipson, Blumea 24 (1978) 169. — Aralia guilfoylei [Bull, Cat. (1873)] Cogn. & Marche, PL Ornam. 2 (1874) t. 58. — Notho- panax guilfoylei (Cogn. & Marche) Merr. Philip. J. Sc. 7 (1912) Bot. 242; Fl. Manila (1912) 357; En. Philip. 3 (1923) 234. Glabrous shrub to 3 m high, with few branches, bearing spirally arranged leaves towards their ends. Leaves imparipinnate, with 3^ pairs of leaflets, c. 60 cm long; petiole c. 18 cm, terete, with a sheathing base 3-4 cm long; petiolules c. IV2- 2V2 cm; leaflets rotund, oblong or rhomboidal, c. 10-14 by 6-7 cm, rather thin and flaccid, some- times rugose, margin sharply serrate to broadly cuneate (often decurrent on the petiole), often oblique, midrib and lateral veins evident. Inflores- cence a diff"use panicle; primary axis short (r. 3- 4 cm) with one or few lateral flowering branches, terminating in an umbel of long much-branched rays; lateral branches and rays c. 40-50 cm, with secondary branches (c. 5-8 cm) singly or mainly in subverticils along their length and in a terminal umbel ; secondary branches with one or more pairs of small bracts and ending in umbellules with a few lateral tertiary branches also ending in umbel- lules; umbellules with c. 8-12 flowers on pedicels c. 8-10 mm long. Calyx a minute rim. Petals 5, VI2 mm long. Stamens 5, anthers oblong, 1 V2 rnm long; filaments 2 mm. Ovary turbinate, c. 1 mm high, usually 3-celled, styles at first erect, but soon elongating and recurved. Fruit fleshy, globose, f . 4 by 5 mm. Distr. Native country unknown, possibly from East Malesia. Cultivated throughout the region, but less commonly than other cultivated species of this section. General throughout the tropical Pacific. Ecol. Usually grown as a hedge plant. Flowers only when left untrimmed or when growing as an escape from cultivation. Note. Recognizable by the shape, texture and serrations of the leaflets, which are usually varie- gated with whitish or yellowish blotches near the margins. The flowers are brown in bud but yellow- green when open. 6. Polyscias cumingiana (Presl) F.-Vill. Nov. App. (1880) 102; Philipson, Blumea 24 (1978) 169. — Scutellaria secunda angustifolia Rumph. Herb. Amb. 4: 76, t. 32. — Panax pinnatum Lamk, Encycl. 2 (1788) 715, nan P. pinnata J. R. & G. Forster, 1776; DC. Prod. 4 (1830) 254; Span. Linnaea 15 (1841) 208; Miq. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 15; Boerl. Handl. 1 (1890) 647; Koord. Minah. (1898) 15. — Panax secundum Schult. Syst. 6 (1820) 215, nom. superfl. illeg. — Panax bandanense Zipp. ex Span. Linnaea 15 (1841) 208, nom. inval. in synon. — Paratropia cumingiana Presl, Epim. (1851) 250. — Notho- panax pinnatum (Lamk) Miq. Bonplandia 4 (1856) 139; Miq. Fl. Ind. Bat. 1, 1 (1856) 766; Merr. Int. Rumph. (1917) 409; En. Born. (1921) 458; Heyne, Nutt. PI. (1927) 1209; Ochse & Bakh. Veg. D.E.L (1931) 67, f. 38; Corner, Ways. Trees (1940) 156. — Notlwpanax cumingii (Presl) Seem. Fl. Vit. (1865) 114. — Aralia filicifolia C. Moore, 111. Hortic. 23 (1876) 72, t. 240. — Arthrophyllum pinnatum (Lamk) Clarke, Fl. Br. Ind. 2 (1879) 734, pro basionym. — Panax cumingiana (Presl) Rolfe, J. Linn. Soc. Bot. 21 (1884) 310; Vidal, Phan. Cuming. (1885) 117. — ? Aralia naumannii E. Marchal, Bot. Jahrb. 7 (1886) 469. — Panax crispatum Bull, Cat. (1888) 9. — Panax ornatum Bull, I.e. — P. cumingii (Presl) Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 45. — P. rumphiana Harms, I.e. 45; in K. Sch. & Laut. Fl. Schutzgeb. (1900) 485; Bakh./. & Ooststr. in Back. Bekn. Fl. Java (em. ed.) 7 (1948) fam. 159, p. 13; Back. & Bakh./. Fl. Java 2 (1965) 167. — Nothopanax crispatum (Bull) Merr. Philip. J. Sc. 7 (1912) Bot. 241. — Nothopanax ornatum (Bull) Merr. I.e. — P. filicifolia (C. Moore) Bailey, Rhodora 18 (1916) 153. — P. sorongensis Gibbs, Arfak (1917) 216. — Anomopanax cumingianus (Presl) Merr. Philip. J. Sc. 17 (1920) 300; En. Philip. 3 (1923) 236. Glabrous shrub or small tree, to c. 4 m, un- branched or with few branches bearing spirally arranged leaves towards their ends. Leaves impari- pinnate, up to 100 cm; petiole to 20 cm, with a sheathing base c. 5-6 cm long; petiolules to 3 cm; leaflets ovate-oblong or elliptic, 10-30 by 2-13 cm, apex attenuated or acuminate, base rotund, trun- cate or broadly cuneate, often oblique, margin entire or minutely and distantly dentate (sometimes pinnatilobed or almost pinnatipartite); midrib and lateral veins evident. Inflorescence a large terminal panicle, or with flowering branches also in the axils of the upper leaves; primary axis up to 140 cm with secondary branches mostly in verticils at intervals along its length; secondary branches up to 120 cm, bearing umbellules in an irregularly branched system towards their extremities, peduncles of the umbellules with one or more pairs of small bracts; umbellules with c. 10-20 flowers; pedicels 4-8 mm. Calyx a minute rim with 4-5(-6) teeth. Petals 4-5(-6), l^U-l^ii mm long. Stamens 4-5(-6), 2 mm long. Ovary turbinate, c. 2 mm long, 2-5-celled ; styles at first erect, later spreading. Fruit subglobose, fleshy, 3-5 mm when dry. Distr. Cultivated throughout the region, and also apparently growing as part of the indigenous vegetation. Ecol. Rain-forest and secondary growth, usually at low altitude but ascending to 1700 m, native range not very clear. Uses. A common ornamental shrub, especially 1979 Araliaceae — I (Philipson) 77 the cultivars with dissected foliage. Also for culinary use. Vern. (from various sources). Celebes: daim grisik, d. mangko, d. papeda pandang, M, Manado; Philippines: bani, Luzon, Albay; Moluccas: k^nd^m, Ternate, gurahati, M; papua, Jappen I.; ^Iew Britain: awalagu. Gazelle Pen. OcHSE & Bakhuizen van den Brink record that the vernacular names applied to P.fruticosum also include P. cumingianum. Notes. A complex of forms requiring intensive biosystematic study. I adhere to the view that the Indo-Malayan material is specifically distinct from the Polynesian Polyscias pinna ta J. R. & G. Forster; cf. Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 45; Philipson, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 9, but some cultivated forms included here may have been derived from the Pacific rather than the Indo-Malayan species. Existing herbarium material, though voluminous, is inadequate to determine whether the complex includes species with more limited ranges. As in 1. P. fruticosa, cultivated forms with dis- sected foliage occur. See also under 4. P. Scutellaria for a discussion of the relationship with that species. I have not seen the type of Aralia naumannii Marchal which, from the description, I assume to be a synonym. 7. Polyscias javanica K. & V. Bijdr. 7 (1900) 13; KooRD. Atlas 4 (1916) f. 679 A-R; Bakh. /. & OosTSTR. in Back. Bekn. Fl. Java (em. ed.) 7 (1948) fam. 159, p. 12; Back. & Bakh. /. Fl. Java 2 (1965) 167; Philipson, Blumea 24 (1978) 169. Glabrous shrub or sparsely branched small tree, bearing spirally arranged leaves towards the ends of the branches. Leaves imparipinnate, c. 70 cm long; petiole c. 17 cm, with a wide membranous sheath- ing base (f. 7 cm long); petiolules r. 1 cm; leaflets ovate to elliptic-oblong, up to 22 by 8 cm, mem- branous, apex attenuated, base rounded to broadly cuneate, often oblique, margin entire or minutely distantly dentate, slightly revolute, midrib and lateral veins prominent. Inflorescence a panicle; rachis 40 cm, with caducous bracts (reduced leaves or leaf-sheaths), bearing few secondary branches and terminating in a compound umbel; secondary branches with one or more verticils of tertiary branches; peduncles of the umbellules with one obscure bract or none; umbellule of c. 10-20 flowers, pedicels c. 5 mm. Calyx an undulate rim, indistinctly 5-dentate. Petals 5, 2V2-3 mm long. Stamens 5, with very short filaments. Ovary turbi- nate, IV2 mni high, 5-celled, styles at first erect, later recurved. Fruit a globose or obovoid fleshy drupe (prominently 5-ribbed when dry) c. 7 by 5 mm; persistent styles united below, strongly reflexed above. Distr. Malesia: East Java, Lesser Sunda Is. (Lombok, Sumbawa, Sumba). Ecol. In forest up to 1650 m. Uses. The root is said to be made into a face powder in Sumbawa. Vern. Lesser Sunda Is.: pulosari, Sumbawa, amdari, Sumba. Note. The vegetative parts are rather similar to those of 6. P. cumingiana, but the smaller and simpler inflorescence is distinctive. 2. Section Gelibia (Hutch.) Philipson, Blumea 24 (1978) 169. — Gelibia Hutch. Gen. Fl. PI. 2 (1967) 57. Tree with bipinnate leaves. Flowers arranged racemosely; style arms free and divergent after anthesis. Distr. In New South Wales, Queensland, and Malesia (New Guinea) 1 sp. 8. Polyscias elegans (C. Moore & F.v.M.) Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 45; J. Am. Arb. 20 (1939) 323; Philipson, Blumea 24 (1978) 169. — Panax elegans C. Moore & F.v.M. Trans. Phil. Inst. Vict. 2 (1858) 68; Bth. Fl. Austr. 3 (1866) 383; C. Moore, Handb. Fl. N.S.W. (1893) 215. — Nothopanax elegans (C. Moore & F.v.M.) Seem. Fl. Vit. (1866) 114; J. Bot. 4 (1866) 294. — Tieghemopanax elegans (C. Moore & F.v.M.) ViGuiER, Bull. Soc. Bot. Fr. 52 (1905) 308. — P. branderhorstii Harms, Nova Guinea 8 (1910) 274; Bot. Jahrb. 56 (1920) 42; J. Arn. Arb. 20 (1939) 323. — Gelibia branderhorstii (Harms) Hutch. Gen. Fl. PI. 2 (1967) 57. — Gelibia elegans (C. MooRE & F.v.M.) Hutch. I.e. 58 Tree to 20 m high, with a spreading crown of thick branches bearing terminal clusters of large leaves, young parts with fawn to grey scurfy tomentum. Leaves bipinnate, becoming glabrous, up to 110 by 50 cm, or larger; petiole c. 13 cm, with a slightly clasping base; petiolules up to c. V2 cm; leaflets ovate or elliptic, c. 6 by 3 cm, chartaceous to somewhat leathery, margin entire and slightly revolute, apex with an obtuse apiculum, base cuneate, midrib prominent, lateral veins rather obscure. Inflorescence a large panicle, 78 Flora Malesiana [ser. I, vol. 9^ tomentum persistent especially on the pedicels; primary axis stout, c. 30 cm, bearing secondary axes along its length and in a terminal subumbel- late cluster, bracts caducous; secondary axes c. 20-30 cm, bearing tertiary axes (c. 10 cm) along their length; flowers borne racemosely along the tertiary axes on pedicels 1-2 mm long. Calyx a minute rim. Petals 5, oblong, rather fleshy, spreading at anthesis and soon falling. Stamens 5, 2 mm long on short filaments, soon falling. Ovary scurfy, at first turbinate, globose at anthesis, c. 4 mm high, 2-celled, disk rising to the 2 styles; style arms at first short and pressed together, elongating and diverging after the anthers have fallen. Fruit leathery, globose, compressed, c. 5 by 5 mm and irregularly ribbed when dry. Distr. New South Wales and Queensland; in Malesia: New Guinea (West Irian, Southern Distr. ; Papua, Western & Central Distr.). Ecol. Gallery and second growth forest in (seasonal) monsoon regions, at low altitudes, up to 300 m. Notes. The racemose flower arrangement has sometimes been considered sufficiently distinctive to justify the accommodation of this species in a separate genus. The bark is described as grey or patched grey- brown; the wood as soft and pale; and the cut surfaces having a clear sticky exudate which has a pleasant fragrance. The inflorescence branches are purplish, the petals maroon, the anthers cream to yellow on reddish filaments, and the fruit purplish and smooth. 3. Section Eupteron (MiQ.) Philipson, Blumea 24 (1978) 170. — Eupteron MiQ. PI. Jungh. 3 (1855) 420; Fl. Ind. Bat. 1, 1 (1856) 762. Trees or shrubs with imparipinnate or bipinnate leaves. Flowers in umbellules or capitula; style arms divergent at least in fruit. Distr. In Malesia 6 spp., but the section probably includes the majority of the genus in other parts of the Old World tropics. Note. With the exception of 14. P. philipsonii, the Malesian spp. of sect. Eupteron have umbellules or capitula arranged racemosely along the main inflorescence branches. The type species of the section (P. nodosa) has its flowers in capitula, but this is not considered a sufficient difference to require a separate section. KEY TO THE SPECIES 1 . Flowers in capitula 9. P. nodosa 1 . Flowers in umbellules. 2. Umbellules arranged racemosely along the principal rays. 3. Leaf margins crenate. 4. Leaflets oblong (Luzon) 10. P. florosa 4. Leaflets lanceolate (New Guinea) 11. P. ledermannii 3. Leaf margins entire (rarely with few minute dentations). 5. Leaflets ovate (Sabah, Palawan) 12. P. borneensis 5. Leaflets elliptic (New Guinea) 13. P. belensis 2. Umbellules in a short corymbose compound umbel 14. P. philipsonij 9. Polyscias nodosa (Bl.) Seem. J. Bot. 3 (1865) 181; F.-ViLL. Nov. App. (1880) 102; Vidal, Phan. Cuming. (1885) 117; Rev. PI. Vase. Filip. (1886) 145; Boerl. Handl. 1 (1890) 647; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 44; Koord. Minah. (1898) 491; K. & V. Bijdr. 7 (1900) 11; Merr. Philip. J. Sc. 1 (1906) Suppl. 110; Koord. Exk. Fl. Java 2 (1912) 716; Atlas 4 (1916) f. 677 & 678; Merr. Int. Rumph. (1917) 408; Sp. Blanc. (1918) 294; En. Philip. 3 (1923) 233; Heyne, Nutt. PI. (1927) 1208; Bakh. /. & Ooststr. in Back. Bekn. Fl. Java (em. ed.) 7 (1948) fam. 159, p. 12; Back. & Bakh./. Fl. Java 2 (1965) 167; Philipson, Blumea 24 (1978) MO. — Papaya silvestris Rumph. Herb. Amb. 1: 149, t. 53 f. 1. — Aralia nodosa Bl. Bijdr. (1826) 872. — Paratropia nodosa (Bl.) DC. Prod. 4 (1830) 265; Presl, Epim. (1851) 250. — Aralia umbraculifera Roxb. [Hort. Beng. (1814) 22, nomen;] Fl. Ind. ed. Carey 2 (1832) 108. — Aralia pendula Blanco, Fl. Filip. (1837) 223; ed. 2 (1845) 157; ed. 3, 1 (1877) 283. — Hedera nodosa (Bl.) Hassk. Tijd. Nat. Gesch. Phys. 10 (1843) 131. — Eupteron nodosa (Bl.) Miq. PI. Jungh. 3 (1855) 420; Fl. Ind. Bat. 1, 1 (1856) 762; Ann. 1979 Araliaceae — I (Philipson) 79 Mus. Bot. Lugd.-Bat. 1 (1864) 220; Hutch. Gen. Fl. PI. 2 (1967) 68. — IP. acuminata Vidal, Sinopsis Atlas (1883) 28, t. 55 f. A, non Seem. 1865; F.-ViLL. Nov. App. (1880) 102. — P. floribunda Elmer, Leaf!. Philip. Bot. 10 (1939) 3819, nom. inval. in synon. — Fig. 28. Tree to 25 m, unbranched or with few thick branches, bearing tufts of long leaves at their ends, young parts scurfy-tomentose. Leaves imparipin- nate, multijugate, becoming glabrous or minutely villose on the nerves beneath, up to 2 m (or even 3 m); petiole c. 30 cm, to 2 cm 0, with a short sheathing base; leaflets sessile, ovate-oblong com- monly 15 by 4 cm, or larger, chartaceous, margin slightly crenate, apex ± apiculate, base truncate. Inflorescence a large panicle (sometimes with additional flowering branches in the axils of the upper leaves), tomentum ± persistent; primary axis stout, c. I'/a m, bearing secondary axes along its length, bracts triangular, c. 5 mm long; secon- dary axes c. 20-40 cm; capitula borne racemosely along the secondary branches on peduncles c. 6- 15 mm long. Flowers c. 8-12 in a capitulum. Calyx a minute rim. Petals 5, triangular, 2 mm long. Stamens 5, anthers broadly ovate on short fila- ments. Ovary turbinate, c. 2^J2 mm high, 5-celled; disk flat, rising to the 5 central erect styles. Fruit subglobose (5-ribbed when dry), styles spreading and reflexed. Distr. Solomon Is. (Bougainville); in Males ia: Sunda Straits (Krakatau), Java, Lesser Sunda Is. (Lombok), Celebes, throughout the Philippines and Moluccas (Talaud, Ternate, Sulu Is., Ceram, Ambon, Banda, Tenimber, Aru Is.), New Guinea (eastwards to Milne Bay Distr.). Recorded doubtfully, on sterile material, from the Andaman Is. by Kurz (Veg. Andam. Is. 1870, 39) but unlikely as it is absent from Sumatra, Malaya, and Borneo. Ecol. Open thickets and rain-forest, mostly at low altitudes and on small islands, but recorded to 1000 m. Uses. Used medicinally against purpuric fever and to delay pregnancy (Mindanao). The leaves are used to stupify fish and the wood makes durable fence posts and handles for rice-knives. Vern. (from various sources). Java: ki langit, S, deleg, djaranan, mangle, putengan, J, penang- penangan, Md, kaju djaran, k. lanang, rangit, tjaliru, tua kalap; Philippines: bias-bias, bingliu. Tag., Bis., bungloi. Bis., bon liu, bungliu, goyung- goyung. Tag., bungdieu, Ig., hagdan-anak, C.Bis., mano-mdno. Yak., malapapaya. Tag., Bis., Pang., tukod-ldngit. Tag., Pamp.; Celebes: kambowa, lalusuhan, pasusinggala, (pohon) mamalapa, tama- lola, tundu; Sulu Is.: lua; Moluccas: kobo-kobo, Morotai, bat at opus, Ceram, papaya utan, pat a tulan, p. tulong, Ambon, matanglolan, Tenimber; West New Guinea: amorigh, laulako, sonomdoro, totja. Note. A widespread, common and noticeable species. The flowers are described as yellow and as having a fragrant scent. The wood is soft and white. 10. Polyscias florosa Philipson, Blumea 24 (1978) 170. Small tree, up to 10 m, with thick branches bearing terminal clusters of large leaves, glabres- cent. Leaves imparipinnate, multijugate, up to 2 m; petiole c. 25 by 1 V2 cm, with a short sheathing base; leaflets sessile, oblong, c. 22 by 8 cm, chartaceous, margin crenate, apex slightly to long acuminate, base truncate or subcordate. Inflores- cence a large glabrescent panicle; principal rays over 1 m, up to 3 cm 0, each bearing secondary branches (20-40 cm) along its length; bracts broadly ovate, 1 cm long, caducous; umbellules borne racemosely along the secondary branches on peduncles 1-2 cm long (in fruit to c. 4 cm). Flowers c. 6-12 in an umbellule, pedicels 3-5 mm with 2 minute bracts near the middle. Calyx an undulate rim. Petals 5, oblong, 3 mm long. Stamens 5. Ovary turbinate, c. 2 mm long, 5-celled, disk flat, rising to the 5 central erect styles. Fruit subglobose (5-ribbed when dry), 8 by 6 mm, styles spreading and reflexed. Distr. Malesia: Philippines (Luzon; Mt Bulusan, Sorsogon Prov. and Mt Malinao, Albay Prov.). Ecol. Rarely collected in montane forest, c. 560 m. Note. The leaves are very similar to those of P. nodosa, with which this species has often been confused. However the pedicelled flowers are quite distinctive. 11. Polyscias ledermannii Harms, Bot. Jahrb. 56 (1921) 409; Philipson, Blumea 24 (1978) 170. — Panax murrayi (non F.v.M.) F.v.M. Descr. Pap. PI. 7 (1886) 29; Boerl. Handl. 1 (1890) 647. — P.forbesii Baker/. J. Bot. 56 (1923) Suppl. 22. — P. clemensiana Harms, Bot. Jahrb. 69 (1938) 283. Glabrous tree, to 28 m, with few thick branches bearing terminal clusters of large leaves. Leaves imparipinnate, up to IV4 rn; petiole to c. 16 cm, base slightly dilated; petiolules c. 2-4 mm; leaflets lanceolate or oblong-lanceolate, c. 10-20 by 2-4 cm, papyraceous, margin crenulate, apex acuminate, base truncate or rounded, often oblique, midrib prominent, lateral veins numerous, faint. Inflorescence a panicle; primary axis stout, c. 15 cm, bearing crowded secondary axes along its length and in a subumbellate cluster at its apex, bracts triangular, c. 3 mm long, caducous; secondary axes 2O-3O(-40) cm; umbellules borne racemosely along the secondary axes (often in subverticils) on peduncles 2-5 cm long, usually bearing 2 minute bracts. Flowers c. 8-12 in an umbellule, pedicels c. 5 mm. Calyx a minute rim. Petals 2V2 rnm long. Stamens 5. Ovary turbinate, 2 mm high, 3-4- celled, disk rising to the central styles. Fruit 80 Flora Malesiana [ser. I, vol. 9^ globose, 4-5 mm high (when dry), 3-4-ribbed, styles 3-4, divergent, persistent. Distr. Malesia: New Guinea (Vogelkop to Milne Bay). Ecol. In rain-forest, gallery forest and secondary growths, usually above 1500 m (to 3000 m), but occasionally as low as 500 m. Vern. Agitgwa, Hagen, gapin, Morobe Distr., gowi, Efogi lang., panda panda, Mendi lang., puri, Kepilan. Notes. A common small tree. Carr estimated the height of one specimen as 28 m, but most are considerably smaller. The flowers are described as green. The wood is soft and pale. The plant has a scent resembling celery. The species closely resembles P. murrayi (F.v.M.) Harms from Queensland. 12. Polyscias borneensis Philipson, J. Bot. 78 (1940) 118; Blumea 24 (1978) 170. Glabrous shrub or small tree with few thick branches bearing terminal clusters of leaves. Leaves imparipinnate, c. 50 cm; petiole c. 15 cm, base slightly dilated; petiolules c. 1 cm; leaflets ovate, up to 10 by 4 cm, coriaceous, margin entire or slightly undulate, apex attenuate, base rounded or broadly cuneate, often asymmetrical, midrib prominent, lateral veins rather obscure. Inflores- cence a large panicle; primary axis stout, up to c. 40 cm, bearing secondary axes along its length and in a subumbellate cluster at its apex, bracts triangular c. 5 mm long; secondary axes c. 35 cm; umbellules borne racemosely along the upper part of the secondary branches on peduncles 2-5 cm long. Flowers about 10 in an umbellule, pedicels c. 10 mm long. Calyx a minute undulate rim. Petals oblong 3 mm long. Stamens 5, oblong c. 2V2 rnm long, filaments 2 mm. Ovary turbinate, 2 mm high, 5-celled, disk rising to the 5 central erect styles. Fruit unknown. Distr. Malesia: N. Borneo (Mt Kinabalu), Philippines (Palawan, Mt Mantalingahan). Ecol. Montane forest, c. 1500 m. Note. Specimens collected on Palawan by Edano were distributed with the specific epithet 'palawanensis\ but this name was never published. 13. Polyscias belensis Philipson, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 13; Blumea 24 (1978) 170. Glabrous tree, to 14 m. Branches bearing termi- nal clusters of large leaves. Leaves imparipinnate, up to 80 cm long; petiole c. 12-16 cm, base slightly dilated; leaflets subsessile or with a petiolule to 1 cm; leaflets elliptic to narrowly elliptic, c. 12 by 3V2-6 cm, subcoriaceous, margin entire or sparsely denticulate, slightly revolute, narrowed to an obtuse apex, base cuneate, midrib prominent, lateral veins c. 12. Inflorescence a panicle; primary axis stout, c. 18-22 cm, bearing many secondary branches along its length and in a subumbellate cluster at its apex, bracts caducous; secondary axes c. 22-35 cm, bearing umbellules racemosely along their length, bracts triangular c. 3 mm long; peduncles c. 1 V2-3 cm. Flowers c. 10 per umbellule, pedicels 5-6 mm. Calyx a minute rim. Petals 4, oblong. Stamens 4, anthers oblong, 2 mm long, filaments very short. Ovary turbinate, IV2-2 mm high (in anthesis), 4-5-ceIled, disk flat with 4-5 free styles, at first erect, later divergent. Fruit unknown. Distr. Malesia: New Guinea (West Irian, Bele R. near Lake Habbema; Papua, Morobe Distr., Mt Kaindi and Aseki Patrol area). Ecol. Infrequent tree in montane forest and regrowth, 1450-2650 m. Note. The inflorescence is similar to that of 11. P. ledermannii, but the shape of the leaflets is distinctive. In the type (from West Irian) the leaf margins are entire, but in the three gatherings from Morobe there are some dentations, and it is possible these are not conspecific. The corolla is maroon (Morobe). The plant has an odour like celery. 14. Polyscias philipsonii Bernardi, Ber. Schweiz. Bot. Ges. 74 (1966) 364; Philipson, Blumea 24 (1978) 170. — P. fraxinifolia Philipson, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 13, non Harms, 1894. A sympodial, unbranched, glabrous shrub or epiphyte to 2 m high, with leaves spirally arranged towards the apex, new shoots arising below the inflorescence. Leaves imparipinnate, or occa- sionally 3 leaflets inserted together, or the leaves bipinnate, upto35cmlong; petiole to 10 cm, with a very short sheathing base; petiolules to 7 mm, leaflets variable in size and shape, broadly or narrowly elliptic, up to 9 by 4^2 cm, but usually smaller, narrowed to the apex or apiculate, base cuneate or rounded, margin minutely setose- crenulate, midrib prominent, lateral veins few, arched-ascending. Inflorescence a corymb; peduncle short, 1-2 cm, sometimes bearing a reduced leaf or cataphyll distally; primary rays c. 5, subequal, c. 3-4^ 1 2 cm, bearing terminal com- pound umbels (occasionally with a lateral umbel) ; ultimate umbellules with 6-12 flowers on pedicels c. 4 mm long. Calyx a minute rim with indistinct teeth. Petals 5, triangular, 2 mm long. Stamens 5, anthers oblong, I mm long. Ovary turbinate, c. 2 mm high, 5-celled, the fleshy disk rising to the 5 central erect styles. Fruit fleshy, globose, 5-ribbed when dry, c. 4 by 6 mm, styles persistent, recurved. Distr. Malesia: West New Guinea (Idenburg R.). Ecol. In mossy forest at 1800 m, and on an exposed slope at 2150 m. Notes. The short corymbose inflorescence is similar to that in sect. Palmervandenbroekia, but the free recurved style arms exclude it from that sec- tion. The shrub is variable in aspect, and may grow as an epiphyte or as an undershrub. The panicle branches are described as purple and the flowers as green. 1979] Araliaceae — I (Philipson) 81 4. Section Kissodendron (Seem.) Philipson, Blumea 24 (1978) 170. — Kissodendron Seem. J. Bot. 3 (1865) 201. Trees or shrubs with pinnate or bipinnate leaves. Umbellules arranged in diffuse panicles. Style arms fused, forming a beak-like projection on the fruit. Distr. In Malesia (Moluccas and New Guinea) 5 spp., three of which extend to Queensland. KEY TO THE SPECIES 1 . Leaves bipinnate 15. P. bipinnata 1. Leaves simply imparipinnate. 2. Leaflets in 3-4(-5) pairs 16. P. zippeliana 2. Leaflets in more than 5 pairs. 3. Fruits rotund, c. 8 mm broad 17. P. royenii 3. Fruits ovoid, c. 4-5 mm broad. 4. Base of leaflets cuneate 18. P. schultzei 4. Base of leaflets truncate, subcordate, or rounded 19. P. australiana 15. Polyscias bipinnata (Gibbs) Philipson, Blumea 24 (1978) 170. — Kissodendron, bipinnatum Giees, Arfak (1917) 161; Harms, Bot. Jahrb. 56 (1921) 413. A small tree; branches with spirally arranged leaves towards their ends ; young parts brown fur- furaceous. Leaves bipinnate, 70 by 68 cm; petiole 26 cm, with a short sheathing base; rachis with a pair of leaflets at each articulation; petiolules to IV4 cm; leaflets narrowly oblong or ovate, c. 6-12 by 2V2-4(-5) cm, apex attenuated, base broadly cuneate or rounded, margin entire, irregularly undulate slightly revolute, midrib prominent. Inflorescence a diffuse panicle (sometimes also with flowering branches in the axils of the upper leaves) ; peduncle c. 6 cm, terminating in few (3) primary rays; primary rays 40-50 cm, bearing verticils of pairs of secondary branches along their length and ending in a compound umbel; umbellules of c. 10-15 flowers on pedicels c. 10 mm long. Calyx an undulate rim. Petals 5, 2V2 mni long. Stamens 5, filaments 1 mm, anthers IV4 mm long. Ovary turbinate 2-3-celled, disk with a central stylo- podium 1 mm long. Fruit ovoid, fleshy, 5 by 3 mm (when dry) crowned by the calyx rim and the beak- like stylopodium, stigmas minutely capitate. Distr. Malesia: W. New Guinea (Vogelkop: Angi Lakes). Ecol. Montane forest, at c. 2000 m. Vern. Louklouwko. Note. The individual leaflets are similar to those of 18. P. schultzei, but their bipinnate arrangement is distinctive. The flowers are white. 16. Polyscias zippeliana (MiQ.) Valeton, Bull. Dep. Agr. Ind. Need. 10 (1907) 42; Philipson, Blumea 24 (1978) 170. — Panax zippelianum MiQ. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 15; Boerl. Handl. 1 (1890) 645; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 46. — Nothopanax zippelianum (MiQ.) Seem. F1. Vit. (1866) 115. — Kissodendron australianum [non (F.v.M.) Seem.] Boerl. Handl. 1 (1890) 650. — P. caroli Harms, Bot. Jahrb. 56 (1920)411. Shrub, lV2(-5) m, with few branches, bearing leaves spirally arranged near their ends, glabrous except a slight rufous tomentum on young parts. Leaves imparipinnate, with 3-4 pairs of leaflets, up to c. 60 cm; petiole to 17 cm, with a very short sheathing base; petiolules V2-IV2 cm; leaflets oblong, ovate or elliptic, 6-13 by 3'/2-6 cm, papyraceous, apex attenuate or apiculate, base cuneate, often oblique, margin entire, slightly revolute, midrib and few lateral veins evident. Inflorescence a panicle, often appearing lateral by rapid growth of a bud at its base ; branches rather fine and wiry, rachis often angled at the nodes, up to c. 60 cm; secondary branches borne singly at wide intervals and in a few rayed terminal umbels, c. 20 cm long, bearing umbellules on side branches and terminally; umbellules with c. 10-15 flowers on pedicels V2-I cm long. Calyx a minute rim. Petals 5, broadly oblong, 2 mm long. Stamens 5, anthers oblong, 1 mm long, filaments 1 mm. Ovary turbi- nate, 2 mm high, 2-celled, the fleshy disk rising to the central fused stylar column. Fruit ovoid, crowned by the prominent stylopodium with a capitate stigma, c. 7 by 4 mm when dry. Distr. E. Malesia: SE. Moluccas (Aru Is.) and New Guinea (S. Irian and southern part of Western Distr. of Papua; also in the Sepik Distr.). Ecol. In the Aru Is. and S. New Guinea in obviously seasonal savannah country associated with Acacia and Melaleuca at low altitudes, in the Sepik Distr. in the mountains at c. 1000 m. Vern. Mirmur, uwah, perdi-perdi, W. Irian. 82 Flora Malesiana [ser. I, vol. 9^ Fig. 30. Polyscias schultzei Harms, a. Habit, x ^/j, b. flower and ditto in LS, x 10, c. petal, d. stamens, e. fruit and ditto in CS, x 5 {a-d Hoogland & Pullen 5841, e Kalkman 4200). Drawn by Helene Mulder. 1979 Araliaceae — I (Philipson) 83 Notes. The identity of P. zippeliana has been discussed by Boerlage, Valeton, and Harms, ll.cc. The short leaf-sheath and the connivent styles place it within sect. Kissodendron. A comparison of the type specimen with the species usually known as P. caroli Harms shows them to be conspecific. The type of P. caroli is no longer available but specimens collected by Brass can be considered authentic as they were identified by Harms. It should be noted, however, that the type of P. caroli is from the Sepik Distr. (probably in rain-forest) at 1000 m, while all other specimens are from the south of the island and from seasonal savannahs at much lower altitudes. The single gathering from the Aru Is. approaches 19. P. australiana having larger leaves with more pairs of leaflets than other gatherings of P. zippe- liana. 17. Polyscias royenii Philipson, Blumea 24 (1978) 170. Small tree up to 5(-15) m, monocaulous sym- podial, persistently rufous-furfuraceous, with leaves spaced along the upper part of the stem. Leaves imparipinnate, with c. 9 pairs of leaflets, up to 100 cm long or more; petiole to 30 cm, with a short sheathing base; petiolules of mid-leaflets c. 8-20 mm; leaflets normally oblong-ovate, broad- est near the base, up to 24 by 11 cm, coriaceous, apex attenuated or apiculate, base truncate to subcordate; often oblique, margin entire, irregu- larly undulate, slightly revolute; midrib prominent, reticulations depressed above (rugose) raised and furfuraceous below. Inflorescence a panicle, flowering branches also often present in the axils of the uppermost leaves; rachis up to 60 cm with verticils of secondary branches subtended by caducous unifoliolate bracts and a terminal com- pound umbel; lower secondary branches up to 50 cm, with verticillate tertiary branches and terminal compound umbels; umbellules with 10-20 flowers on furfuraceous pedicels c. 8 mm long. Caly.x a furfuraceous rim, with 5 small teeth. Petals 5, strap-shaped, c. 3 mm long, soon falling. Stamens 5. Ovary subcylindric, furfuraceous, c. 3 mm long, 2(-3)-celled, the fleshy disk forming a projecting conical stylopodium 2 mm long; stigmas 2, appressed at anthesis. Fruit rotund, compressed, with prominent ribs when dry 7-9 by 8-10 mm, crowned by the prominent persistent calyx and a short stout beak-like stylopodium; stigmas short, divergent. Distr. Males ia: New Guinea (Cyclops Mts to the E. Highlands Distr.). Ecol. Terrestrial or epiphytic in primary or secondary montane forest, 1200-2400 m. Vern. Irian: ato, Kapauku lang. ; Papua New Guinea: habia, S. Highlands, magabin, W. Sepik, momin, Mendi lang. Note. A species readily characterized by the sympodial monocaul habit with thick leathery leaflets, a large rigid furfuraceous inflorescence, and large, compressed, black fruits with a glaucous bloom. 18. Polyscias schultzei Harms, Bot. Jahrb. 56 (1921) 410; Philipson, Blumea 24 (1978) 171. — P. gjellerupii Harms, Bot. Jahrb. 56 (1921) 410. — Fig. 30. Shrub or small tree, often unbranched, 4-20 (-26) m high, young parts densely brown furfura- ceous, branches with spirally arranged leaves towards their ends. Leaves imparipinnate, to 80 cm long, with c. 8-11 pairs of leaflets; petiole up to 35 cm, with a short sheathing base; petiolules c. 8 mm; leaflets ovate-lanceolate, oblong or elliptic, up to 14(-17) by 4-5 cm, apex attenuated or acuminate, base broadly cuneate, oblique, margin entire, midrib prominent. Inflorescence a diff"use, repeatedly compound umbel, with reduced leaves — persistent at the nodes; peduncle usually rather short (c. 5 cm) ending in a group of several primary rays (and sometimes with one or more lateral rays); primary rays 50-60 cm, with verticils and a terminal umbel of secondary rays which in turn are branched; umbellules with c. 5-10 flowers on fine pedicels c. 5-8 mm long (elongating in fruit to 12-15 mm). Calyx a minute rim with 4-5 small teeth, usually glabrous but occasionally furfura- ceous. Petals 4-5, c. 3 mm long. Stamens 4-5, 2 mm long. Ovary — cylindric, glabrous or occasionally furfuraceous, c. 2 mm long, 2-celled, the fleshy disk forming a blunt stylopodium. Fruit ovoid, fleshy, c. 5 mm long, crowned by the incon- spicuous calyx and the projecting beak-like stylo- podium (c. 2 mm long); stigmas slightly divergent or capitate. Distr. Queensland; in Malesia: Moluccas (Morotai) and New Guinea (throughout most of the island, from the Vogelkop Peninsula and Japen I. to the Central and Morobe Distr.). Ecol. Usually in lower and mid-montane rain- forest (with Castanopsis, Nothofagus, Lithocarpus), also in regrowth, usually above 1200 m, ascending to 2400 m, but also descending to near sea-level. Vern. New Guinea: atok, Wissel Lakes, funim, Telefomin, houklouwko, Angi Lakes, paul, Wabag, agugna, Hagen. Notes. A widespread and frequently collected species recognized by the multijugate leaves with rather small, thin, attenuate leaflets with cuneate base. The flowers are white to yellowish and the fruits black. The grey bark exudes a small amount of gummy sap; the wood is white. Harms recognized that P. gjellerupii was very similar and the abundant material now available indicates the variability of this common species. 19. Polyscias australiana (F.v.M.) Philipson, Blumea 24 (1978) 171. — Hedera australiana F.v.M. Fragm. 4 (1864) 120; Bth. F1. Austr. 3 (1866) 384. — Kissodendron australianum (F.v.M.) 84 Flora Malesiana [ser. I, vol. 9^ Seem. J. Bot. 3 (1865) 201; ibid. 6 (1868) 129. — Irvingia australiana (F.v.M.) F.v.M. Fragm. 5 (1865) 18. var. disperma (F.v.M.) Philipson, Blumea 24 (1978) 171. — Kissodendron australianum (F.v.M.) Seem.' var. disperma F.v.M. Descr. Not. Pap. PI. 5 (1877) 88; Harms, Bot. Jahrb. 56 (1921) 412. Small tree, often unbranched, branches with leaves arranged spirally towards their ends, young parts brown furfuraceous. Leaves imparipinnate, multijugate, to 1 m long; petiole c. 25 cm, with a short sheathing base; petiolules to IV2 cm; leaflets broadly ovate or oblong, broadest near the base, up to 14 by 6 cm, ± coriaceous, apex attenu- ated to an obtuse apex, or with an obtuse apiculum, base rounded, truncate, or subcordate, oblique, margin entire, slightly revolute, midrib and lateral veins prominent. Inflorescence a diff"use panicle (sometimes also with flowering branches in the axis of the upper leaves); peduncle short, terminating in several primary rays, reduced leaves subtending the principal branches; primary rays c. 50 cm, bearing verticils of secondary branches along their length and ending in a compound umbel; umbel- lules of c. 10-15 flowers on pedicels c. 8-10 mm long. Calyx an undulate rim. Petals 5, c. IV2 mm long. Stamens 5, c. 1 mm long. Ovary turbinate, 2-celled, a fleshy disk rising to a low conical stylo- podium. Fruit ovoid, fleshy, 6 by 4 mm (when dry), crowned by the persistent calyx and beak-like stylopodium (2 mm long). Distr. Malesia: New Guinea (Vogelkop Penin- sula and Southern Distr. in Papua). Ecol. Primary rain-forest at low altitudes and also in lower montane forest at 1750 m. Note. Distinguished from 18. P. schultzei by the leathery leaflets with a truncate base, and from 17. P. royenii by the smaller fruits. 5. Section Palmervandenbroekia (GiBBs) Philipson, Blumea 24 (1978) 171. — Palmervandenbroekia Gibbs, Arfak (1917) 162. Trees or shrubs with imparipinnate leaves. Umbellules arranged in short corym- bose compound umbels. Style arms fused, forming a beak-like projection on the fruit. Distr. Malesia: New Guinea (Vogelkop Peninsula and Cyclops Mts in W. Irian, Western Distr. in Papua). KEY TO THE SPECIES 1 . Pedicels tomentose (at anthesis). 2. Inflorescence a compact regular compound umbel with rigid rufous branches (Vogelkop and Cyclops Mts) 20. P. sleumeri 2. Inflorescence more open, less regularly branched, with branches becoming ± glabrous (Mt Bosavi, Papua) 21. P. jacobsii 1. Pedicels glabrous (at anthesis). 3. Flower buds lanceolate, stylopodium at anthesis c. 2 mm long (Vogelkop, Arfak Mts) 22. P. palmervandenbroekii 3. Flower buds ovoid, stylopodium at anthesis c. 1 mm long (Vogelkop, Nettoti Range and Aifat R.) 23. P. vogelkopensis 20. Polyscias sleumeri Philipson, Blumea 24 (1978) 171. Monocaulous sympodial shrub, 1-1 V2 ni, with leaves spaced along the upper part of the stem; young parts red-furfuraceous. Leaves imparipin- nate, with 3-4 pairs of leaflets (leaflets occasionally in threes), c. 35 cm long; petiole to 9 cm, with a very short sheathing base; petiolules 8-15 mm; leaflets oblong, ovate or elliptic, up to 16 by 6 cm, chartaceous, apex attenuate or apiculate, base rounded or cuneate, margin entire, irregularly undulate, slightly revolute, midrib prominent. Inflorescence a compound umbel, red-furfuraceous (stem growth renewed by a bud t)etween the inflorescence and the uppermost leaf); peduncle 3-6 cm, bearing scars of caducous reduced leaves, occasionally with a lateral umbellule, and ending in an umbel of c. 1 primary rays; primary rays l-2'/2 cm, sometimes with a minute bract near the middle; umbellule with 10-20 flowers on pedicels 2-3 mm long, densely furfuraceous. Calyx a rim with 5 broad teeth. Petals 5, broadly oblong, 3-4 mm long. Stamens 5, anthers 1 mm long, filaments 2 mm. Ovary turbinate, 2 mm high, 2-celled, the fleshy disk forming a projecting coni- cal stylopodium; stigmas 2 appressed at anthesis. 1979 Araliaceae — I (Philipson) 85 Fruit rotund to ovoid compressed 5-6 by 5 mm crowned by the prominent persistent calyx and the beaic-like stylopodium (2-2' /2 rnm long); stigmas not prominent. Distr. Malesia: New Guinea (Vogelkop Penin- sula and Cyclops Mts). Ecol. Undergrowth in primary forest, 450- 1000 m. Note. The flowers are creamy white, the fruit dark red. 21. Polyscias jacobsii Philipson, Blumea 24 (1978) 171. A monocaulous sympodial shrub to 3 m high, with leaves spaced along the upper part of the stem, young parts furfuraceous. Leaves imparipinnate, 4-5 pairs of leaflets (leaves of juveniles smaller with fewer leaflets), c. 45 cm long; petiole to 12 cm, with a short sheathing base; petiolules I-IV2 cm; leaflets oblong or elliptic, c. 12 by 4 cm, charta- ceous, apex attenuate or caudate, base truncate or cuneate, margin entire, irregularly undulate, slightly revolute, midrib prominent. Inflorescence a compound umbel (stem growth renewed by a bud between the inflorescence and the uppermost leaf); peduncle 1-4 cm (sometimes bearing a uni- foliolate leaf about the middle with an axillary flowering branch), ending in an umbel of 2-4 primary rays; primary rays sometimes subtended by 1-2 unifoliolate leaves, c. 4 cm long, ending in 2-3 tertiary rays which may branch again before ending in umbellules; umbellules with 6-10 flowers on furfuraceous pedicels c. 8 mm long. Flower buds with an apical umbo. Calyx a rim with 5 acute teeth. Petals 5, attenuated. Stamens 3 mm long, anthers IV2 rnm long. Ovary subcylindric, furfuraceous, 4 mm high, 2-celled, disk forming a projecting conical stylopodium (IV2 rnrn long at anthesis), stigmas 2. Fruit rotund, compressed, 8 by 9 mm, crowned by the small calyx and the persistent stylopodium (2 mm long) with sub- capitate stigmas. Distr. Malesia: New Guinea (Papua: Mt Bosavi). Only known from the type. Ecol. On old well-drained volcanic soil in primary mixed forest, mostly on ridges and upper slopes, 1600-2100 m. Note. An aromatic, single-stemmed shrub. The flowers are white and the fruit dark purple. 22. Polyscias palmervandenbroekii Bernardi, Candollea 26 (1971) 16; Philipson, Blumea 24 (1978) 171. — Palmervandenbroekia papuana GiBBS, Arfak (1917) 162, f. 15, non P. papuana Seem. 1865. Glabrous shrub, c. l'/2 rn, sparsely branched, with leaves spaced along the upper parts of the stems. Leaves imparipinnate, with 3-5 pairs of leaflets, up to 13 cm long; petiole to 4 cm, slightly channelled above, with a very small sheathing base; petiolules 0-5 mm; leaflets lanceolate, obo- vate to elliptic, 1V4-4 by '/4-2 cm, coriaceous or chartaceous, apex attenuated (sometimes emargi- nate) to apiculate, base cuneate, margin entire, revolute, midrib prominent, lateral veins obscure. Inflorescence a simple or compound umbel; peduncle slender, c. 2 cm; primary rays (when present) few, c. 1 cm; umbellules of c. 10 flowers on glabrous pedicels 3-6 mm long. Flower buds lanceolate, acute, 3-6 mm. Calyx a rim with 5 broad teeth. Petals 5, attenuate, 5-8 mm long. Stamens 5, filaments c. 3 mm, anthers c. 1 mm. Ovary turbinate, striate, 2 mm high, 2-celled, disk projecting as a conical stylopodium (2 mm long). Fruit ovoid, compressed, 6 by 5 mm, crowned by the persistent calyx and the beak-like stylopodium; stigmas 2, small, divergent. Distr. Malesia: New Guinea (Vogelkop: Arfak Mts). Ecol. In summit heath vegetation (Tristania- Dacrydium scrub) and Nothofagus forest, 1900- 2400 m. Note. Collections made since the original de- scription show that this interesting plant is quite variable as regards leaf-shape, texture and amount of rolling of the margin, and also in the size of its flowers. The calyx is purplish brown, the petals reddish purple outside and white within. The ripe fruit is purplish black. The thin papery bark is light grey. 23. Polyscias vogelkopensis Philipson, Blumea 24 (1978) 171. An often monocaulous sympodial shrublet usually under 1 m high, with leaves spaced along the upper parts of the stem; young parts slightly furfuraceous, but soon becoming glabrous. Leaves imparipinnate, with 3^ pairs of leaflets (leaflets occasionally in threes) variable in size; petiole 2-7 cm, slightly channelled above with a short inconspicuous sheathing base; petiolules 2-8 mm ; leaflets elliptic or ovate, 3-1 2 by 1 ' j^-A cm, chartaceous, apex attenuated or apiculate, base broadly to narrowly cuneate, margin entire, irregularly undulate (occasionally with isolated dentations) slightly revolute, midrib prominent. Inflorescence a small compound umbel; peduncle up to 8 cm, occasionally with a lateral or basal umbellule, ending in c. 2-3 primary rays (some- times subtended by a reduced leaf); primary rays 1-2 cm; umbellules with c. 10-15 flowers on glabrous pedicels c. 3-6 mm long. Flower buds ovoid, obtuse. Calyx a rim with 4-5 broad undula- tions. Petals 4-5, broadly oblong, 4 mm long. Stamens 4-5, anthers 1 mm, filaments 3 mm. Ovary turbinate, 2 mm high, 2-celled, the fleshy disk rising to a conical stylopodium (1 mm long); stigmas 2, appressed at anthesis. Fruit ovoid, compressed, 5 by 4 mm, crowned by the persistent calyx and the prominent beak-like stylopodium; stigmas inconspicuous. 86 Flora Malesiana [sen I, vol. 9^ Distr. Malesia: New Guinea (Vogelkop and Merrill (1923, l.c) stated that this name was Wandammen Peninsula, Nettoti Range and Aifat excluded by Blanco from his 2nd edition, and that R.)- it is unrecognizable from the very short and im- Ecol. Primary submontane forest {Nothofagus, perfect description. Possibly a Rubiacea. Castanopsis, and conifers), heath vegetation, or in n i ■ n n ^ , i. t. t^ 1 1-iAA inr>r> Poiyscias roemeriana Harms, Bot. Jahrb. 56 open places, 1200-2000 m. /ir>-ii^.i'5c/a5705A:e/ L. S. Gibbs, J. Linn. Soc. Bot. 39 226; Merr. Sp. Blanc. (1918) 384; En. Philip. 3 (1909) 149. — This species was by error ascribed by (1923) 233. — Philippines. Index Kewensis to the Philippines; it is from Fiji. 11. MACROPANAX MiQ. Fl. Ind. Bat. 1, 1 (1856) 764; Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 13; Bth. in B. & H. Gen. PI. 1 (1865) 945; Boerl. Hand!. 1 (1890) 643; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 47; Hutch. Gen. Fl. PI. 2 (1967) 80. — Hederopsis Clarke, Fl. Br. Ind. 2 (1879) 739; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 56; King, J. As. Soc. Beng. 67, ii (1898) 62; Hutch. Gen. Fl. Pi. 2 (1967) 78; Stone, Gard. Bull. Sing. 30 (1977) 141. — Fig. 31. Small, unarmed trees. Leaves digitately compound or unifoliolate; petiole terete, with a sheathing base; stipules inconspicuous; leaflets entire or denticulate. Inflorescence a terminal panicle of umbellules. Pedicels articulated below the ovary. Flowers bisexual or on lower branches male. Calyx 5-6-dentate. Petals 5-6, valvate. Stamens 5-6, extrorse. Ovary inferior, 2-6-ceIled. Disk fleshy with the styles united below. Fruit ellipsoid or obovate, surmounted by an awl-shaped or conical stylopodium; exocarp fleshy, endocarp crustaceous. Endosperm ruminate. Distr. 4 spp. from India, Burma and southern China to West Malesia: 3 spp., Sumatra, Malaya and Java. Ecol. Lowland and montane rain-forest. Notes. Hederopsis is united with Macropanax because the species are very similar in appearance and in their technical characters. The sole diff"erence is the number of cells in the ovary, 2 (or sometimes 3) in Macropanax and 5-6 (sometimes 4) in Hederopsis. This distinction does not seem adequate to split a few very similar species into distinct genera. This genus is distinguished from Pseudopanax (China, New Caledonia, Tasmania, New Zealand, Chile) by the ruminate endosperm. Cf. Philipson, New Zeal. J. Bot. 3 (1965) 333. KEY TO THE SPECIES 1 . Ovary 2-celled. 2. Ovary broadly campanulate, not ribbed; 'epicalyx' below ovary distinct. Leaf margin usually dis- tinctly serrate. Inflorescence branches usually bearing lateral umbellules 1. M. dispermus 2. Ovary narrowly turbinate, strongly ribbed; 'epicalyx' below the ovary absent. Leaf margin entire or minutely dentate. Inflorescence branches usually without lateral umbellules . . 2. M. concinnus 1 . Ovary 5- or 6-celIed 3. M. maingayi 1, Macropanax dispermus (Bl.) O. K. Rev. Gen. Tjib. 2 (1923) 227; Bakh./. & Ooststr. in Back. PI. 1 (1891) 271; KooRD. Exk. Fl. Java 2 (1912) Bekn. Fl. Java (em. ed.) 7 (1948) fam. 159, p. 14; 716, 717; Atlas 4 (1916) f. 681 & 682 F-P; Fl. Back. & Bakh./. Fl. Java 2 (1965) 168. — Aralia 1979 Araliaceae — I (Philipson) 87 disperma Bl. Bijdr. (1826) 872. — Hedera disperma (Bl.) DC. Prod. 4 (1830) 265. — Aralia calyculata Z. & M. in Mor. Syst. Verz. (1846) 41. — M.flori- bunda MiQ. Fl. Ind. Bat. 1, 1 (1856) 764. — M. oreophilus MiQ. I.e. 764; Suppl. (1860) 135; Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 13; KuRZ, For. Fl. Burma 1 (1877) 541 ; Clarke, Fl. Br. Ind. 2 (1879) 738; BoERL. Hand). 1 (1890) 644; K. & V. Bijdr. 7 (1900) 16. Small tree, up to 18 m. Leaves dispersed along the shoots; petiole to 25 cm, striate, with a small basal sheath, the connate stipules forming a small ligule within the petiole; leaflets 5-7, petiolules up to c. 5 cm (the lateral shorter); blade elliptic or oblanceolate, usually c. 10-23 by 2' ,-9 cm, gradually tapered to an acute apex, base cuneate, rounded or oblique, margin coarsely dentate or sometimes only finely serrate, glabrous when mature. Inflorescence a panicle of umbellules, often stellate-furfuraceous, with a stout rachis to 40 cm and lateral (secondary) branches which terminate in umbellules. and which commonly bear tertiary branches. Pedicels c. 5-7 mm, with a distinct involucre ("epicalyx") around the base of the flower. Calyx rim undulate or indistinctly dentate, puberulous. Petals 2-3 mm long. Ovary subglobose, c. 2' 2 rnm high. Disk fleshy, surmounted by an awl-shaped stylar column, which bifurcates at apex. Fruit ovate, 1 by ^ , cm. crowned by the persistent stylar column. Distr. India, Burma, and southern China; in Malesia: Sumatra, Malay Peninsula, and through- out Java. Ecol. Mountain rain-forests, 1000-2300 m. in Java especially in Central and East Java. Vern. Java: panggang puju. p. serem, p. siju, ramo-gentjel, tjerem, S,pampung, sahang, tanganan, J, konjingal, Md. 2. Macropanax concinnus MiQ. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 220; Boerl. Handl. 1 (1890) 643; K. & V. Bijdr. 7 (1900) 18; Koord. Exk. Fl. Java 2 (1912) 717; Atlas 4 (1916) f. 682 Q-\V. — M. undulatus (W.\LL. ex G. Don) Seem. J. Bot. 2 (1864) 19 A, pro specim. malac.\ Bakh./. & Ooststr. in Back. Bekn. Fl. Java (em. ed.) 7 (1948) fam. 159. p. 14: B.ACK. & B.\KH./. Fl. Java 2 (1965) 168. Small tree, up to 15 m. Leaves dispersed along the shoots; petiole slender, to 20 cm, with a small basal sheath, the connate stipules forming a small ligule within the petiole; leaflets 5 or 3, petiolules slender, up to c. 5\ 2 cm (the lateral shorter) ; blade elliptic, usually c. 7-15 by 2' 2-5 cm, gradually tapered to a caudate-falcate acuminate apex, base narrowly cuneate, or rarely obtuse, often oblique, margin entire or sometimes a few very fine serra- tions, glabrous, membranous. Inflorescence a panicle of umbellules, minutely stellate-pubescent, with a stout rachis to 30 cm, and lateral (secondan.-) branches which terminate in umbellules; pedicels c. 6-7 mm, only slightly swollen at the articulation. Calyx rim undulate or indistinctly dentate. Petals c. 2 mm long. Ovary narrowly turbinate with distinct longitudinal ribs, c. 2 mm high. Disk fleshy, surmounted by an awl-shaped stylar column. Fruit ovate, 8 by 4 mm, crowned by the persistent stylar column. Distr. Malesia: widespread in Java, but local and uncommon. Ecol. Damp mountain forests, descending to lower altitudes (400 m) than the preceding sfjecies. Vern. Djampang tjerem, panggang puju, S. tanganan, J. Notes. This species has sometimes been identified with .\/. undulatus (Wall, ex G. Dos) See.m. from the Himalayan region, but that species differs in the more verticillate inflorescence branches, the large involucre around the umbellule, the hermaphrodite flowers more confined to the terminal umbellules, and the broader less promi- nently ribbed fruit. A considerable geographical gap separates the two species. A specimen collected by Grifhth labelled "Malacca" is perhaps M. undulatus, but the locality may be incorrect. 3. Macropanax maingayi (Clarke) Philipson, comb. nov. — Hederopsis maingayi Clarke, Fl. Br. Ind. 2 (1879) 739: Ridl. Fl. Mai. Pen. 1 (1922) 888 ; Stone, Card. Bull. Sing. 30 (1977) 287, f. 7. — Arthrophyllum trifoliatum Ridl. J. Fed. Mai. St. Mus. 7 (1916) 42. — Hederopsis major RiDL. Kew Bull. (1929) 124. — Fig. 31. Small tree, up to 15 m. Leaves dispersed along the shoots; petiole to 20 cm, striate, with a small basal sheath; leaflets 5, or (below the inflorescence) 3 or 1 , petiolule of the central leaflet to 6 cm, of the lateral leaflets much shorter {c. 1 cm); blade elliptic or ovate, up to 20 by 8 cm. base rounded or broadly cuneate, apex acuminate, margin denticulate or nearly entire, membranous or chartaceous. Inflorescence a terminal panicle of umbellules, with a stout rachis to 20 cm, termi- nating in an umbellule and bearing lateral secon- dary branches (c. 12 cm) which also terminate in umbellules and bear racemosely arranged tertiary branches (c. 4 cm) which terminate in umbellules; umbellules terminating the primary and secondary branches with hermaphrodite flowers, those terminating the tertiary branches smaller and apparently mostly functionally male. Flowers of small umbellules usually 5-merous, of larger umbellules usually 6-7-merous, pedicels c. 1 cm at anthesis, slightly elongating in fruit. Calyx rim obscurely dentate. Petals of terminal flowers c. 3 mm long, fleshy. Ovary subglobose, c. 2 mm high: disk fleshy surmounted by the style arms which become divergent in their upper part at anthesis. Fruit enlarging to a drupe 2 by 1 cm with a persistent calyx rim and a prominent conical stylopodium tapering into the stylar column bear- ing the recurved distal parts of the styles. Flora Malesiana [ser. I, vol. 9^ Fig. 31. Macropanax maingayi (Clarke) Philipson. a. Leaf and upper branches of inflorescence, x ^/j, b. flower bud and CS of ovary, x 2, c. flower, x 2, d. fruit, slightly enlarged (Phytochem. Surv. Mai. 2519). Drawn by W. R. Philipson. 1979] Araliaceae— 1 (Philipson) 89 Distr. Tonkin; in Malesia: Malay Peninsula Note. The trunk may have small buttresses, the (from Kedah southwards) and Central Sumatra bark is rugose with large Icnticels, and produces a (East Coast Res. and Mt Kerintji). watery exudate when cut. The flowers are greenish Ecol. Uncommon, in forest, up to 1050 m. yellow. 12. ANAKASIA W. R. Philipson, Blumea 21 (1973) 87, fig. on p. 88. — Fig. 32. Shrub with large, simple, exstipulate leaves. Inflorescence axillary; rachis bearing racemosely arranged umbellules. Pedicels very short with an articulation below the flower. Calyx a short rim with minute lobes. Petals 5 or 6, valvate, triangular. Stamens 5 or 6, filaments thick, anthers large, dorsifixed. Ovary inferior, broadly obconic, (4-)5-6-celled. Disk fleshy with (4^)5-6 stylar arms. Fruit broadly obo- void, strongly ribbed when dry; exocarp fleshy. Pyrenes compressed, crustaceous; endosperm smooth. Distr. Malesia: West New Guinea. Monotypic. Note. The large, simple, oblanceolate leaves clustered at the ends of the branches recall Meryta, but the flowers do not share the highly distinctive features of that genus. The floral and fruit characters are not unlike those of Polyscias {e.g. there is an articulation below the flower, the style arms are free, and the endosperm is smooth), but the general facies is unlike that genus, and this, together with the distinctive inflorescence and leaf, make the plant quite distinct from any species of Polyscias. 1. Anakasia simplicifolia W. R. Philipson, Blumea recurve and spread beyond the calyx. Fruit with 21 (1973) 87, fig. on p. 88. — Fig. 32. (4-)5-6 prominent ridges when dry, c. 2 by 1 V2 cm. Glabrous shrub, 5 m, branches marked with with the persistent style arms in a terminal prominent lenticels. Leaves crowded at the ends of depression. the branches, sessile or with a very short petiole, Distr. Malesia: West New Guinea (Vogelkop lanceolate, up to 135 by 18 cm, narrowed gradually Peninsula and near Babo). to the base, apex acuminate, margin entire or Ecol. Primary forest near sea-level, undulate, midrib prominent, lateral veins arching Note. Beccari described (in sched.) the in- upwards, reticulation rather indistinct. Inflores- florescence branches as erect in flower and cence rachis simple or forked, to 70 cm, at maturity reflexed in fruit. He noted that the outer flowers of 5 mm 0, bearing small lanceolate bracts. Peduncles the umbellules are probably male and those at the arising from the axils of all but the lowest bracts, centre female, though he could not be certain 3-4 mm long, elongating to c. 15 mm in fruit. of this. His observations cannot be confirmed from Umbellules with c. 1 1 minute bracts. Flower buds the material available. Evidently all flowers are when dry c. 3 mm long. Calyx rim with 5-6 minute structurally hermaphrodite, but all developing teeth. Petals slightly fleshy, 2 mm long. Stamens fruits seen are attached to central pedicels. The 5-6, filaments broad ; anthers 4-celled, ^U mm long. corolla is green and soon falls, the disk yellow, the Ovary glabrous. Disk with a central boss formed anthers cream, and the mature fruit blue (azure- by closely appressed subulate arms which soon violet) and aromatic. 13. BRASSAIOPSIS Decne & Planch. Rev. Hort. IV, 3 (1854) 106; Bth. in B. & H. Gen. PI. 1 (1865) 945; Clarke, F1. Br. Ind. 2 (1879) 735; Boerl. Handl. 1 (1890) 643; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 42; King, J. As. Soc. Beng. 67, ii (1898) 61; Ngoc-Sanh Bui, Adansonia 6 (1966) 437, pi. 1 (map); Hutch. Gen. Fl. PI. 2 (1967) 79; Stone, Gard. Bull. Sing. 30 (1977) 280. — Araliopsis Kurz, Rep. Andam. (1870) 39, nom. invai, in synon., non Engler, 1896. — Wardenia King, J. As. Soc. Beng. 67, ii (1898) 60; Harms in E. & P. Nat. Pfl. Fam. Nachtr. 2 90 Flora Malesiana [ser. I, vol. 9^ Fig. 32. Anakasia simplicifolia Philipson. a. Leaf, x V?, b. inflorescence, x Va, c. single umbellule with an immature fruit, d. flower bud, x \1, e. immature fruit, x 2 (Beccari PP 282). 1979] Araliaceae— I (Philipson) 91 (1900) 51; RiDL. Fl. Mai. Pen. 1 (1922) 887; Hutch. Gen. Fl. PI. 2 (1967) 81; cf. Frodin, Misc. Rec. Found, Fl. Males. 3 (1973) 8. — Euaraliopsis Hutch. Gen. Fl. PI. 2 (1967) 80, 624. — Pseudobrassaiopsis Banerjee, J. Bomb. Nat. Hist. Soc. 72(1975)71. — Fig. 33, 34. Shrubs or trees, usually prickly and tomentose. Leaves digitately compound, palmately lobed or simple, with a sheathing base and a usually bicuspid ligule; petiole terete. Inflorescence usually a terminal panicle of umbellules, often with persistent small bracts; pedicel not articulated below the ovary. Calyx rim 5-den- tate. Petals 5, valvate. Stamens 5, extrorse. Ovary inferior, 2-celled; disk shallow; styles united into a usually long column. Fruit globose, exocarp fleshy, endocarp chartaceous; seed 2, not compressed, endosperm smooth. Distr. More than 20 spp., extending from India, Tibet and southern China to West Malesia: Sumatra, Malay Peninsula and W. Java. In Malesia 6 spp., 1 endemic in Sumatra, 3 endemic in the Malay Penin- sula, 2 in Indo-Malaya. Ecol. Understorey of damp evergreen forest, mostly in mountainous districts, up to 2400 m. Notes. A genus characterized by the 2-celled ovary, columnar style, and non-articulated pedicel, com- bined with leaves which may be either entire, palmately lobed, or digitately compound. Possibly close to Trevesia, which is similar vegetatively, but which has more massive inflorescences and ovaries with more numerous cells. In dividing the species into two genera merely on the basis of leaf shape, Hutchinson I.e. fragmented what appears to be a coherent assemblage. KEY TO THE SPECIES 1 . Inflorescence shorter than the petioles (or at most about equal to them). Leaves palmately lobed, but variable in the variety 1. B. sumatrana 1 . Inflorescence much longer than the petioles. 2. Leaves simple, entire, minutely and remotely dentate. 3. Leaf base subcordate; blade to 35 cm or more long; petioles c. 15 cm, 5 mm . . 2. B. simplex 3. Leaf base cuneate or rounded; blade to 25 cm long; petioles c. 7 cm, 2 mm . . 3. B. minor 2. Leaves palmately lobed or digitately compound (occasionally some simple). 4. Leaves palmately lobed 4. B. polyacantha 4. Leaves digitately compound. 5. Branches of the inflorescence bearing several umbellules racemosely (as well as one at end); umbellules with c. 25-35 flowers 5. B. glomerulata 5. Branches of the inflorescence bearing a terminal umbellule only (or occasionally with separate flowers below it); umbellule with f. 10-13 flowers 6. B. elegans 1. Brassaiopsis sumatrana Ridl. J. Fed. Mai. St. lobes elliptic acuminate, midrib prominent and Mus. 8, 4 (1917) 43. — Kalopanax sumatranum lateral veins widely spaced and arched-ascending, MiQ. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 17, margin irregularly spinulose-serrate. Inflorescence heterotypic; Boerl. Handl. 1 (1890) 647; Harms a short, rufous-tomentose panicle, arising below in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 51. — the leaves (actually terminal but overtopped by ? Kalopanax resectum MiQ. Ann. Mus. Bot. Lugd.- sympodial innovations); rachis c. 4-6 cm, lateral Bat. 1 (1863) 18, see note below. — Euaraliopsis branches ascending, c. 2-4 cm, subtended by small sumatrana {Rioi..) Hutch. Gen. Fl. PI. 2 (1967) 624. linear bracts and sometimes branching to the third degree; umbellules with c. 8-20 flowers; pedicels var. sumatrana. c. 1 cm long, subtended by numerous minute Shrub or small tree, to 15 m, with stout prickly bracts. Calyx a minute rim with 5 indistinct teeth, branches, rufous-tomentose in its younger parts. Petals 5, triangular, 2^2 mm long. Stamens 2 mm Leov^i palmately lobed, clustered at the ends of the long, anthers oblong, 1 mm long. Ovary obconic, shoots; petiole c. 15-32 cm, striate, sometimes c. 2 mm high; disk fleshy, cushion-shaped, sur- with prickles, with an enlarged basal sheath mounted by a slender awl-shaped stylar column, and a small bicuspid ligule; blade rounded, cor- Fruit subglobose, c. 8 mm high, with a persistent date, c. 14-22 by 20-28 cm, palmately 5-7-lobed, calyx-rim and stylar column. 92 Flora Malesiana [ser. I, vol. 9^ Fig. 33. Brassaiopsis elegans Ridl. a. Habit, x ^/s, b. fruit and ditto in CS, x 4 (Kloss 1912). Drawn by P. Prendergast. 1979] Araliaceae — I (Philipson) 93 Distr. Malesia: West Sumatra (Mts Kerintji, Merapi, Singalang). Ecol. Montane, evergreen forest, 1500-2400 m. Vern. Semonloni,', Karo. Notes. The foliage is similar to that of 4. B. polyacantha and of Trevesia sundaica, but the small inflorescence is distinctive. The prickles persist on the older branches and the trunk. The petals and filaments are white, the anthers light purple, and the disk yellow. Kalopanax resectum MiQ. I.e., described on a KoRTHALS collection from Sumatra, possibly belongs here, but the inflorescence may be larger and most leaves are decidedly digitately compound. Possibly it represents a hybrid between B. suma- trana and 5. B. glomerulata. var. variaefolia Philipson, var. nov. — B. ovalifolia RiDL. J. Fed. Mai. St. Mus. 8, 4 (1917) 43. Frutex var. sumatrana humilior et tenen'or, foliis secus ramis dispersis, inermis. Folia minora (10- 20 cm longa) forma variantia etiam in eodem ramo, vel simplicia et ovata, vel profunde bi- vel trilobata, vel digitatim composita. — Typus: W. Meijer 7200 (L), Mt Sago near Pajakumbuh, W. Sumatra, 1000-1200 m, 8-VIII-1957. A smaller, more delicate and unarmed shrub than var. sumatrana, with the leaves scattered along the branches. Leaves smaller (10-20 cm long) and variable in shape, even on the same branch, being either simple and ovate, or deeply bi- or trilobed, or digitately compound. Distr. Malesia: Central West Sumatra. Ecol. Montane, evergreen forest, 1000-1200 m. Note. It is possible that Ridley was correct in regarding this as a distinct species, but the size and shape of the inflorescence links it with B. sumatrana, and the variable form of the leaf suggests that it may be no more than an extreme form of B. suma- trana. Nevertheless, typical forms of that species, with their stout prickly stems and rather coarse palmately lobed leaves, are very distinct. On the evidence available it is not possible to decide whether this variety consists of shade and/or juvenile forms of B. sumatrana, or whether it should constitute a distinct species. 2. Brassaiopsis simplex (King) Stone, Card. Bull. Sing. 30 (1976) 282, f. 3. — Wardenia simplex King, J. As. Soc. Beng. 67, ii (1898) 60; Harms in E. & P. Nat. Pfl. Fam. Nachtr. 2 (1900) 51; Ridl. F1. Mai. Pen. 1 (1922) 887; Hutch. Gen. Fl. PI. 2 (1967) 81; Philipson, J. Linn. Soc. Bot. 63, Suppl. (1970) 90. Shrub, to 5 m high, branches prickly, rufous- tomentose in its younger parts. Leaves large, simple, coriaceous, clustered towards the tips of the branches; petiole c. 14-20 cm, stout, striate, with a basal sheath and a short, bicuspid ligule; blade ovate, c. 35-40 by 12-18 cm, apex shortly acuminate, base slightly cordate, rufous-stellate hairs persistent beneath, midrib prominent, lateral veins very widely spaced, arched ascending, margin minutely dentate. Inflorescence a panicle of umbellules; rachis c. 10-20 cm; lateral branches reflexed, 5-12 cm, subtended by small lanceolate bracts and often bearing 1-2 bracts from which tertiary branches arise; umbellules with c. 10-20 flowers; pedicels \^U-1 cm, slender, subtended by numerous minute bracts. Calyx a narrow rim with 5 indistinct teeth. Petals 5, falling as a calyptra. Stamens 2 mm long, anthers oblong. Ovary ob- conic, c. 2 mm high, disk cushion-like, surmounted by an awl-shaped stylar column l'/2 mm long. Fruit globose, fleshy, c. 5 mm 0, with a persistent calyx rim and a stylar column c. 2 mm long. Distr. Malesia: Malay Peninsula (Perak, Selangor). Ecol. In open bamboo forest, 100-800 m. Note. Originally described as a distinct genus because of its supposedly 1 -celled ovary. Ridley (1922) did not include this feature in his descrip- tion. He distinguished Wardenia from Brassaiopsis because of its simple leaves, but several species of Brassaiopsis from further north also have undivided leaves. Both Harms (1900) and Hutchinson (1967) accepted the original statement that the ovary is 1 -celled at the time of flowering, but I found the ovary to be 2-celled even in the young state (1970, I.e.). 3. Brassaiopsis minor Stone, Gard. Bull. Sing. 30 (1976) 282, f. 5. Small shrub, c. 60 cm, with very few small prickles, rufous tomentose on its young parts. Leaves simple, spaced towards the ends of the branches, subcoriaceous; petiole 5-8 cm, c. 2 mm 0, with a basal sheath and a small ligule; blade elliptic or ovate 20-26 by 10-12 cm, apex shortly acuminate, base rounded to cuneate, rufous stellate hairs persistent beneath, midrib prominent, lateral veins widely spaced, arched ascending, margin minutely and remotely dentate. Inflores- cence: peduncle short, bearing lanceolate bracts; umbellules on short lateral branches, with 15-20 flowers on slender pedicels c. IV2 cm long, sub- tended by minute puberulent bracts. Calyx with 5 indistinct teeth. Ovary rotund, surmounted by a slender stylar column 1 mm long. Distr. Malesia: Malay Peninsula (Trengganu). Ecol. Humid situations in lowland to montane forest, to 1 100 m. 4. Brassaiopsis polyacantha (Wall.) Banerjee, Ind. For. 93 (1967) 341; Stone, Gard. Bull. Sing. 30 (1977) 282. — Hedera polyacantha Wall. PI. As. Rar. 2 (1831) 82, t. 190. — Panax palmatum Roxb. [Hort. Beng. (1814) 21, nomen;] Fl. Ind. ed. Carey 2 (1832) 74. — B. palmata (Roxb.) Kurz, J. As. Soc. Beng. 39, ii (1870) 77; Clarke, Fl. Br. Ind. 2 (1879) 735; King, J. As. Soc. Beng. 67, ii (1898) 61; Ridl. Fl. Mai. Pen. 1 (1922) 887. — 94 Flora Malesiana [ser. I, vol. 9^ Fig. 34. Brassaiopsis polyacantha (Wall.) Banerjee. In flower at Kuala Lumpur (Photogr. Philipson, 1975). Euaraliopsis palmata (Roxb.) Hutch. Gen. Fl. PI. 2 (1967) 80. — Pseudobrassaiopsis polyacantha (Wall.) Banerjee, J. Bomb. Nat. Hist. Soc. 72 (1975)72. — Fig. 34. A small sparsely branched tree with buttresses, to c. 13 m; branches prickly, rufous-tomentose on the younger parts and persisting on the leaves and inflorescence. Leaves palmately lobed, clustered at the ends of the shoots; petiole to c. 70 cm, longi- tudinally grooved, with an enlarged basal sheath and a pair of small stipular processes; blade rounded, cordate, c. 18-30 by 20-40 cm, deeply palmately 5-9-lobed, lobes oblong-elliptic, acumi- nate, midrib prominent and main lateral veins widely spaced and arched-ascending, margin inconspicuously serrate. Inflorescence a large panicle often overtopped by sympodial innova- tions; rachis c. 40-60 cm, lateral branches c. 8- 15 cm, subtended by very small caducous bracts and often bearing a pair of minute bracts near the middle, from which short tertiary branches may arise, branches terminating in spherical umbellules ; umbellules with c. 15-25 flowers, subtended by numerous minute bracts; pedicels ■'/4-IV4 cm. Calyx of 5 small teeth, rusty tomentose. Petals 5, triangular, 2'/2 mm long. Stamens 2 mm long, anthers oblong, 1 mm. Ovary obconic, c. 2 mm high; disk fleshy, cushion-shaped, surmounted by an awl-shaped stylar column IV2 rnrn long. Fruit globose, 6-8 mm 0, with a persistent calyx rim and stylar column. Distr. Widespread in SE. Asia (India, Nepal, Burma, Andamans); in Malesia: Malay Peninsula (from Kedah to Selangor and Pahang). Ecol. Montane rain-forest, to c. 1800 m. Note. The inflorescence is similar to that of 5. B. glomeriilata but the leaves of these two species cannot be confused. 5. Brassaiopsis glomerulata (Bl.) Regel, Gartenfl. 12 (1863) 275, t. 411; Koord. Exk. Fl. Java 2 (1912) 715; Atlas 4 (1916) f. 674; Fl. Tjib. 2 (1923) 226; Back. & Bakh. /. Fl. Java 2 (1965) 167; Ngoc-Sanh Bui, Adansonia 6 (1966) 437; Stone, Gard. Bull. Sing. 30 (1977) 282. — Aralia glomeru- lata Bl. Bijdr. (1826) 872. — Hedera glomerulata (Bl.) DC. Prod. 4 (1830) 265 ; Hook. Bot. Mag. 80 (1854) t. 4804. — B. speciosa Decne & Planch. 1979] Araliaceae — I (Philipson) 95 Rev. Hort. 4, 3 (1854) 106; Boerl. Handl. 1 (1890) 643; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 43, f. 1 B; K. & V. Bijdr. 7 (1900) 8; Bakh. /. & OosTSTR. in Baci<. Bekn. Fl. Java (em. ed.) 7 (1948) fam. 159, p. 11. — Macropanax glomerulatum (Bl.) MiQ. Fl. Ind. Bat. 1, 1 (1856) 764. — Macro- panax cyrtostyliim MiQ. Ann. Mus. Bot. Lugd.- Bat. 1 (1863) 13. — B. cyrtostyla (MiQ.) Seem. J. Bot. 2(1864)293. Tree up to 10 m, with stout often sinuous branches prickly in their upper parts, the young parts of the shoots and inflorescences rufous- tomentose. Leaves digitately compound, clustered at the ends of the shoots; petiole c. 25-60 cm, with an expanded basal sheath with two aculeate stipules; leaflets 5-9, petiolules c. 8-10 cm (the laterals shorter), blade elliptic-oblong, c. 15-25 by 6-10 cm, apex acuminate, base broadly cuneate to rounded, midrib prominent, lateral veins widely spaced, arched-ascending, margin serrulate. In- florescence a large panicle ; rachis 30-60 cm, lateral branches c. 20-30 cm, with umbellules arranged racemosely along them on peduncles c. 2-5 cm long, bracts numerous, lanceolate, c. V2 cm long, mostly persistent; umbellules with c. 25-35 flowers, pedicels 1-1 V4 cm, subtended by numerous minute ferruginous bracts. Calyx of 5 small teeth. Petals 5, triangular, li-A mm long. Filaments 2-2'/2 rnm, anthers oblong, 1 mm. Ovary obconic at anthesis, c. 2 mm high; disk fleshy, cushion-shaped, sur- mounted by an awl-shaped stylar column. Fruit globose, 6-7 mm high, with a persistent calyx rim and a stylar column usually 2 mm long but occa- sionally much shorter (even in the same inflores- cence). Distr. India to SW. China; in Malesia: Malay Peninsula (Pahang, Negri Sembilan), Sumatra, W. Java. Ecol. Humid, evergreen, montane rain-forest, 1200-2200 m. Vern. Sumatra: kayu aro, M (Kerintji); Java: panggang, p. pujut (or puju), p. rami (or ranto), S. Note. Juvenile plants bear leaves which are simple, irregularly lobed, or with 2, 3 or 5 leaflets. The flowers are pale yellowish cream. The cut stems exude a yellowish sap. 6. Brassaiopsis elegans Ridl. J. Linn. Soc. Bot. 41 (1913) 291; Fl. Mai. Pen. 1 (1922) 888; Stone, Card. Bull. Sing. 30 (1977) 282, f. 4. — Fig. 33. Shrub, stems with few to many prickles, young parts rufous-tomentose, becoming glabrous. Leaves digitately compound, or occasionally simple; petiole c. 14 cm, striate, with a small basal sheath and a short, bicuspid ligule; leaflets 3-5 (or 1), petiolules c. 3 cm (laterals shorter); blade elliptic, c. 12-16 by 4-5 cm, apex long acuminate, base cuneate, midrib prominent, lateral veins widely spaced, arched-ascending, margin minutely denti- culate. Inflorescence a lax gracefully pendent raceme of umbellules; rachis c. 23 cm, lateral branches c. 5 cm, subtended by small lanceolate bracts and bearing 1-2 minute bracts near the middle, terminating in spherical umbellules and occasionally with separate flowers arising below the apex; umbellules with c. 10-13 flowers, sub- tended by numerous minute bracts; pedicels at anthesis c. V2 cm elongating to c. IV4-IV2 cm in fruit. Calyx of 5 minute teeth. Petals 5, triangular, 2V2 rnm long. Stamens 2 mm long, anthers oblong. Ovary obconic, c. 2 mm high; disk fleshy, sur- mounted by an awl-shaped stylar column. Fruit globose, 6 mm high, with a persistent calyx rim and stylar column. Distr. Malesia: Malay Peninsula (Selangor, Pahang), very local. Ecol. Montane, evergreen rain-forest, 800 m. 14. TREVESIA VisiANi, Giorn. Tosc. Sc. Med. 1 (1840) 72; Mem. Accad. Torino II, 4 (1842) 262; MiQ. Fl. Ind. Bat. 1, 1 (1856) 747; Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 10; Bth. in B. & H. Gen. PI. 1 (1865) 942, p.p.; Boerl. Ann. Jard. Bot. Btzg 6 (1887) 107; Handl. 1 (1890) 639; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 32; King, J. As. Soc. Beng. 67, ii (1898) 57; Koord. Atlas 4 (1916) f. 664-666; Hutch. Gen. Fl. PI. 2 (1967) 71. — Petasula Noronha, Verh. Bat. Gen. 5 (1790) ed. 1, art. 4: 3, p.p., cf. Steen. Regn. Veget. 71 (1970) 376. — Fig. 35, 36. Shrubs or trees, unarmed or prickly, tomentose at least on young parts; buds enclosed in cataphylls. Leaves large, palmately lobed or digitately compound, stipules forming a bicuspid ligule within the terete petiole. Flowers in umbellules which are arranged racemosely along a rachis or in panicles; pedicels not articu- lated below the ovary. Calyx a small rim. Petals 7-12, valvate, often falling at 96 Flora Malesiana [ser. I, vol. 9^ Fig. 35. Trevesia burckii Boerl. a. Habit, x '/s, b. flower, x 2, c. fruit and ditto in CS, x 1 V2 (a Rahmat SI ToROES 5048, b Yapp 99, c Rahmat si Toroes 5049). Drawn by P. Prendergast. 1979 Araliaceae — I (Philipson) 97 anthesis as a calyptra. Stamens as many as the petals; anthers large ovoid, dorsi- fixed, introrse, filament strap-like. Ovary inferior, broadly obconic or subglobose, 7-12-celled; disk rising gradually to a central boss formed by the united styles. Fruit globose or ovoid, crowned by the disk and prominent stylar column; pyrenes coriaceous, compressed, endosperm smooth. Distr. 6 spp. ranging from eastern India and SW. China to Malesia; 5 of the species occur in W. Malesia: Sumatra, Malay Peninsula, Java, Borneo, Lesser Sunda Is. (Lombok). Formerly erroneously credited to the Philippines, cf. Merrill, En. Philip. 3 (1923) 237. Ecol. Primary rain-forest and second growth, especially in humid, shady localities. Note. The genus comprises three well known species, each occupying distinct areas, except that the ranges of two overlap in Sumatra. In addition there are three very local and imperfectly known species. KEY TO THE SPECIES 1. Leaves digitately compound. 2. Petiolules connected by a web of tissue for most of their length 1. T. burckii 2. Petioles free. Continental SE. Asia T. palmata (DC.) Vis. 1. Leaves palmately lobed. 3. Inflorescence branches and petioles densely covered with long red setae .... 2. T. rufo-setosa 3. Inflorescence branches not as above (but sometimes setulose, furfuraceous, pubescent or glabrous). 4. Flowers sessile or subsessile 3. T. beccarii 4. Flowers pedicelled. 5. Ovary c. 14-celled, pedicels stout, c. 4 mm 4. T. snndaica 5. Ovary c. 8-10-celled, pedicels more slender, 1-2 mm 5. T. arborea 1. Trevesia burckii Boerl. Ann. Jard. Bot. Btzg 6 (1887) 110, pi. 12(-14):Handl. 1 (1890) 649: Merr. En. Born. (1921) 456; Masa.mune, En. Phan. Born. (1942) 566. — T. palmata (DC.) Vis. \ar. cheirantha Clarke, F1. Br. Ind. 2 (1879) 732; King, J. As. Soc. Beng. 67, ii (1898) 58. — T. cheirantha (Clarke) O. K. Rev. Gen. PI. 1 (1891) 272; Ridl. Fl. Mai. Pen. 1 (1922) 882. — Fig. 35. Shrub or small, sparsely branched tree up to 10 m, branches stout with small prickles, young parts rufous-furfuraceous, cataphylls prickly. Leaves crowded at the ends of the shoots, digitately compound; leaflets usually 7-9; petioles c. 60 cm, striate, sometimes with small prickles or bristles, dilated into a clasping base, ligule w ith two lanceo- late lobes; petiolules united for all or most of their length by a foliaceous web; leaflets oblong- lanceolate or elliptic, up to 30 by 10 cm, apex acuminate, base cuneate or rounded, margin finely serrate especially in the upper part, principal veins arched-ascending, prominent. Inflorescence a large terminal panicle, often overtopped by lateral shoots; rachis to 60 cm, slightly prickly or not, bearing branches along its length and usually end- ing an umbel of branches, bracts caducous or persistent; secondary branches (peduncles) c. 10- 18 cm, sometimes with 1 or few bracts along their length and terminating in spherical umbellules of 40-50 flowers; pedicels c. 20-35 mm, slender. Calyx an irregular obsolete rim. Corolla hemi- spherical, c. 6 mm high, falling as a calyptra, petals c. 7-10. Stamens c. 7-10, filament flattened c. 2^2 mrn, anthers broadly triangular, c. 3 mm long. Ovary at anthesis broadly obconic, usually rufous-furfuraceous, c. 7-10-celled; disk broadly conical surmounted by the connate styles, stigmas slightly swollen. Fruit subglobose V ^ by 1\4 cm, surmounted by the stylopodium and connate styles. Distr. Malesia: Sumatra, Malay Peninsula (from Kedah southwards), Borneo (Sarawak). Ecol. Primary rain-forest, mostly below 500 m, rarely up to 1000 m. Vern. Sumatra: tapa arimaii, tada lada, M; Malaya: kakabit, tapak itek, t. rimau, M. Note. The stamens are described as cream on an orange disk. Seedling plants have leaves which are palmately lobed or entire. 2. Trevesia rufo-setosa Rjdl. J. Str. Br. R. As. Soc. n. 86 (1922) 294; Fl. Mai. Pen. 1 (1922) 883. A shrub with stout prickly stems, young parts, petioles and inflorescence densely covered with red-brown flat setae. Leaves palmately lobed, clustered at the ends of the shoots: petioles c. 60 cm, dilated into a clasping base with a bicus- pid ligule: blade c. 30 by 30 cm, with 7 lobes; lobes c. 20 by 9 cm, elliptic, shortly cuspidate, mid- veins and widely spaced arched-ascending laterals prominent, margin with many upwardly directed spinulous teeth. Inflorescence terminal; rachis flexuous, c. 20 cm, bearing lateral umbellules, bracts linear c. 12 mm; pedicels short. Flowers heterosexual. Calyx rim obscure. Petals 5, triangu- lar, 6 mm long, setose on the outside, spreading. Stamens 5. Ovary in the fertile flowers obconic. 98 Flora Malesiana [ser. I, vol. 9^ Fig. 36. Trevesia sundaica MiQ. in mixed evergreen mountain forest in E. Java at Sarangan, r. 1200 m altitude, with Quercus (Photogr. Jeswiet, 1925). 1979 Araliaceae — I (Philipson) 99 setose, 5-celled; disk with cylindrical projecting stylar column. Fruit unknown. Distr. Malesia: Malay Peninsula (Selangor: Semangkok Pass). Ecol. Montane forest, at c. 800 m. 3. Trevesia beccarii Boerl. Ann. Jard. Bot. Btzg 6 (1887) 110, pi. 11. Shrub, c. 3 m, with stout prickled stems. Leaves palmately lobed, clustered at the ends of the branches; petiole c. 50 cm, hirsute with short patent bristly hairs when young, some persisting, dilated into a clasping spinulose base with a bicus- pid ligule; blade c. 30 by 40 cm, with 7-9 lobes, base cordate, lobes c. 15 by 7 cm, broadly elliptic- oblong, apex rather blunt, rusty stellate-pubescent when young, sometimes persisting, mid-veins and rather numerous arched-ascending lateral veins prominent, margin irregularly serrate. Inflorescence when young hirsute like the petioles, sometimes persistent; rachis c. 90 cm, bearing many lateral branches (peduncles) along its length, bracts lanceolate, caducous; peduncles c. 5-15 cm with some caducous, linear bracts along their length and around the base of the umbellules (capitula). Capitula of c. 10-20 sessile (or subsessile) flowers. Calyx rim short, undulate. Corolla conical in bud, falling as a calyptra. Stamens 10-12, anthers oblong, filaments stout. Ovary obconic, angular from contact with neighbouring flowers, 10-12- celled; disk broadly conical; surmounted by the massive connate styles, stigmas slightly swollen. Fruit obconic angular from mutual pressure, c. 10 by 7 mm. Distr. Malesia: West Central Sumatra. Ecol. Forest, ascending to 1200 m. Vern. Likabau, M. 4. Trevesia sundaica Miq. PI. Jungh. 3 (1855) 420; Fl. Ind. Bat. 1, 1 (1856) 747; de Vriese, PI. Ind. Or. (1857) 81; MiQ. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 11; Boerl. Ann. Jard. Bot. Btzg 6 (1887) 111, pi. 12 (16); Handl. 1 (1890) 649; K. & V. Bijdr. 7 (1900) 4, ind. var. glomerata K. & V. I.e. 5; KooRD. Exk. Fl. Java 2 (1912) 710; Atlas 4 (1916) f. 664-666; Fl. Tjib. 2 (1923) 222; Bakh./. & OosTSTR. in Back. Bekn. Fl. Java (em. ed.) 7 (1948) fam. 159, p. 4; Back. & Bakh./. Fl. Java 2 (1965) 163. — Aralia palmata Reinw. ex Bl. Cat. Btzg (1823) 43, nomen; ex de Vriese, PI. Ind. Or. (1857) 82, nomen in synon.; Boerl. Ann. Jard. Bot. Btzg 6 (1887) 111, in synon., non Lamk, 1783, nee Lour. 1790. — Sciadophyllum palmatum Bl. Bijdr. (1826) 875, non T. palmata (Roxb.) Vis. 1842. — Aralia reinwardtiana Steud. Nomencl. 1 (1840) 118, nomen. — Gastonia sundaica (Miq.) Baill. Hist. PI. 7 (1880) 161, f. 202. — Actinophyllum palmatum Bl. ex Boerl. Ajin. Jard. Bot. Btzg 6 (1887) 111, />z synon. — Fig. 36. Shrub or small, sparsely branched tree, up to 8 m; branches stout, with small prickles, young parts stellate-tomentose, becoming glabrous, buds with cataphylls. Leaves palmately lobed, clustered at the ends of the branches; petiole striate, c. 50- 60 cm, unarmed or with a few prickles near the base, dilated with a clasping, heavily lenticellate base, ligule bicuspid; blade c. 60 by 60 cm, with 7-1 1 lobes, base cordate, lobes c. 30 by 40 cm, elliptic-oblong, apex acute to acuminate, separated by wide sinuses, mid-veins and widely spaced arched-ascending lateral veins prominent, margin serrate, occasionally irregularly incised. Inflores- cence arising among or below the leaves, a raceme of umbellules, at first rusty pubescent, glabrescent; rachis up to 60 cm, often much shorter, bearing branches along its length and ending in an umbel, bracts lanceolate, caducous; secondary branches (peduncles) c. 8-16 cm, bearing small bracts along their length and around the umbellules ; umbellules of c. 20-35 flowers, pedicels c. 2-5 cm, lateral umbellules sometimes bearing small sterile (male) flowers. Calyx rim very short, undulate. Corolla hemispherical in bud, c. 6 mm high, falling as a calyptra, petals 8-12. Stamens 8-12, filaments stout, anthers ovate, c. 3 mm long. Ovary at anthesis broadly obconic, rufous-furfuraceous, c. 8-10-celled; disk broadly conical passing into the stylar column. Fruit semiglobose, IV4-IV2 cm 0, surrounded by the stylopodium and connate styles. Distr. Malesia: Sumatra, Java, Lesser Sunda Is. (Lombok). Ecol. Evergreen rain-forest, especially in ravines, ascending to 1500 m. Vern. Sumatra: ahab, tapa arimau, M; Java: borang, djemporang, dorang, gabus, gorang, lontanglanting, panggang, p. lembur, p. puju, p. tjutjuk, papanggangan, S. 5. Trevesia arborea Merr. Contr. Am. Arb. 8 (1934) 116. Tree up to 15 m, young parts shortly reddish stellate-tomentose, prickly. Leaves palmately lobed, clustered at the ends of the branches; petioles 17-50 cm, shortly rusty tomentose, glabrescent, unarmed, dilated into a sheathing base, ligule bi- cuspid; blade rotund, 25-40 cm, base cordate, glabrous above, shortly red-pubescent below, coriaceous, usually 9-lobed, sinuses narrow, lobes oblong-elliptic or oblong-oblanceolate, acuminate, margin distantly serrate distally. Inflorescence a raceme of umbellules, at first reddish pubescent, glabrescent, sometimes with a few spines below; rachis stout (c. 1 cm 0), c. 35 cm, bearing branches along its length, and ending in an umbel, bracts oblong-ovate, acuminate 1-2 cm long; primary branches (peduncles) few, spreading, stout, without spines, to c. 15 cm; umbellules of c. 8-13 flowers, pedicels IV2-2V2 cm, stout (3-4 mm 0), reddish tomentose, basal bracts triangular, c. 5 mm long. Flowers hermaphrodite. Calyx an irregular short 100 Flora Malesiana [ser. I, vol. 9^ Fig. 37. Dendropanax borneensis (Philipson) Merr. a. Habit, x V2, b. flower, x 5, c. fruit and ditto in CS, X 5, d. bifid leaf with area enlarged to x 100 to show glands {a-b Nooteboom 2258, c Clemens 51039, d Clemens 28927). Drawn by P. Prendergast. 1979] Araliaceae— I (Philipson) 101 rim. Corolla hemispherical in bud, c. 6 mm high, hemispherical, shortly furfuraceous, 14-celled. 12 mm diam., densely rusty pubescent outside. Fruit unknown. falling as a calyptra. Stamens c. 13, filaments stout, Distr. Malesia: N. Sumatra (Atjeh). 4 mm, anthers ovate, c. 4 mm long. Ovary broadly Ecol. Primary, evergreen forest, 1 100 m. 15. DENDROPANAX Decne & Planch. Rev. Hort. IV, 3 (1854) 107; Bth. in B. & H. Gen. PI. 1 (1865) 943; Clarke, FI. Br. Ind. 2 (1879) 733; Merr. Brittonia 4 (1941) 129; Hui-Lin Li, Sargentia 2 (1942) 38; Philipson, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 18; Hutch. Gen. Fl. PI. 2 (1967) 71; Stone, Gard. Bull. Sing. 30 (1977) 148. — Gilibertia Ruiz & Pav. Prod. Fl. Peruv. (1794) 50, non J. F. Gmel. 1791 ; Bth. in B. & H. Gen. PI. 1 (1865) 944; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 40 & Nachtr. 2 (1900) 254; Philipson, J. Bot. 78 (1940) 1 16. — Textoria Miq. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 12. — Fig. 37. Unarmed, usually glabrous small trees, or erect or subscandent shrubs. Leaves simple, entire, or sometimes palmately 3-5-lobed, often with pellucid glands, with- out articulation between blade and petiole; stipular sheath inconspicuous. Umbels solitary or compound, or on racemosely arranged branches or paniculate; pedicels not articulated below the ovary. Flowers sexually dimorphic. Calyx on entire rim or 5-dentate. Petals 5, valvate. Stamens 5-8, extrorse. Ovary inferior, 4-8-celled; styles united throughout all or part of their length, or free. Disk fleshy, conical. Fruit globose or ellipsoid, usually strongly ribbed when dry; exocarp fleshy; endocarp cartilaginous. Endosperm smooth or rarely weakly ruminate. Distr. About 30 spp. in tropical America and eastern Asia south to Malesia (3 spp.): N. Sumatra, Malay Peninsula, Borneo. Fig. 38. Ecol. Rain-forest in lowland and montane regions, 300-3100 m. Note. Distinguished from Hedera by the normally erect habit and the smooth (or only weakly ruminate) endosperm. KEY TO THE SPECIES 1. Leaves without pellucid gland-dots. 2. Inflorescence a compound umbel 1. D. lancifolius 2. Inflorescence a simple umbel 2. D. maingayi 1 . Leaves with pellucid gland-dots 3. D. borneensis 1. Dendropanax lancifolius (Ridl.) Ridl. Fl. Mai. Distr. Malesia: Malay Peninsula (Perak) and Pen. 1 (1922) 884; Stone, Gard. Bull. Sing. 30 N. Sumatra (Toba Lands). (1977) 148, f. 6 C. — Arthrophyllum lancifolium Ecol. Montane rain-forest, c. 300-1500 m. Ridl. J. Str. Br. R. As. Sec. «. 75 (1917) 38. Vern. Sumatra: modang simarla siak, Batak Tree to c. 14 m. Leaves dispersed along the lang. shoots; petiole c. 5 cm, slightly dilated at the base and with an obscure ligular rim; blade to c. 15 by 2. Dendropanax maingayi King, J. As. Soc. Beng. 3^/4 cm, chartaceous, lanceolate, base narrowly 67, ii (1898) 48; Ridl. Fl. Mai. Pen. 1 (1922) 884; cuneate, apex gradually tapered to an acute point. Stone, Gard. Bull. Sing. 30 (1977) 48, f. 6 B. — margin entire but undulate, midrib well defined, D. parviflorus [non (Champ.) Bth.] Clarke, secondary and tertiary veins delicate. Umbel Fl. Br. Ind. 2 (1879) 733. — Gilibertia maingayi terminal, compound; peduncle c. 2 cm; secondary Philipson, J. Bot. 78 (1940) 117. rays few, 1-2 cm; pedicels 5-10, c. 1-2 cm. Calyx a Slender shrub to c. IV2 m, with few branches, rim with indistinct teeth. Petals triangular, falling Buds enclosed in small cataphylls. Leaves dis- as a cap or separating. Ovary 4-5-celled, styles persed along the shoots; petiole Vz-S cm, chan- forming an erect column. nelled above, slightly dilated at the base and with 102 Flora Malesiana [ser. I, vol, 9^ Fig. 38. Range of Dendropanax Decne & Planch. a small to obscure ligule within the petiole; blade 5-15 by 2-5^:4 cm, thinly coriaceous to membran- ous, ovate, elliptic, oblong or lanceolate, base rounded to narrowly cuneate, apex acute, margin entire, often undulate, midrib prominent, some- times with three distinct principal veins. Umbel terminal, simple, peduncle c. "1-2 cm; pedicels 1-2 cm, slightly elongating in fruit, bracts cadu- cous. Flowers sexually dimorphic, males pre- dominate in lateral umbellules. Calyx margin minute. Petals triangular, c. 2^ '2 mm long. Stamens 5. Ovary 5-6-celled, reduced in male flowers, disk fleshy, conical; styles 5-6, at first erect, but the free upper half spreading in fruit. Fruit globose or ellipsoid, succulent, 5-6-ribbed when dry, c. 10 by 8 mm, crowned by the persistent spreading styles. Distr. Malesia: Malay Peninsula (from Kedah southwards). Ecol. Montane rain-forest, 1000-2000 m. Note. Very like the next species in the general appearance of leaf and inflorescence. However, all specimens from the Malay Peninsula lack pellucid glands in the lamina of the leaf, whereas these are present in all specimens from Borneo. 3. Dendropanax borneensis (Philipson) Merr. Brittonia 4 (1941) 132. — Gilibertia borneensis Philipson, J. Bot. 78 (1940) 116, fig. — Fig. 37. Slender shrub or small tree, with few or no branches, occasionally prostrate or subscandent, sometimes reaching 5-6 m. Buds enclosed in few broadly ovate cataphylls. Leaves dispersed along the shoots; petiole 4-10 cm, channelled above, slightly dilated at base and with a small to obscure ligule within the petiole; blade 7-15 by 3-7 cm, coriaceous, with pellucid glands, usually ovate, elliptic or oblong, occasionally irregularly lobed, base rounded to broadly cuneate, apex acute, margin entire, often undulate, midrib and lateral veins prominent, frequently with 3 distinct princi- pal veins. Umbel terminal, simple, or occasionally compound, single or two borne side by side, with c. 20 flowers; peduncle and secondary rays (if present) usually rather short {V 12-^11 cm), but occasionally much longer (to 13 cm); pedicels c- l\/4-l\2 cm (occasionally to 3 cm), elongating slightly in fruit, bracts ovate caducous. Flowers sexually dimorphic, males predominate in lateral umbellules. Calyx margin minute. Petals triangular, c. 2',2 mm long. Stamens 5. Ovary 4-6-celled, reduced in male flowers; disk fleshy, conical with 4-6 styles, at first erect, but the free upper half spreading in fruit. Fruit globose or ellipsoid, succu- lent, 4-6-ribbed when dry, 5-10 by 6-12 mm, crowned by the persistent spreading styles. Distr. Malesia: N. Sumatra (Mts Goh Lembuh and Pinto) and northern Borneo (Sabah: Mt Kinabalu; Brunei: Pagon Ridge; Sarawak: Mt Murut and Kalabit Highlands). Ecol. Local in montane, mossy forest, and alpine thickets, 1700-3100 m. Vern. Borneo: merit, Kalabit Highlands. Notes. Flower greenish or cream, fruit black. This species exhibits considerable variation in size and shape of leaf, inflorescence and fruit, and 1979] Araliaceae— I (Philipson) 103 also in venation pattern, but these variations are in Borneo. The presence of pellucid glands between not correlated with geographical distribution. the reticulations of the veins of the lamina dis- Specimens from the two Sumatran localities have tinguishes this species from the other two species small fruits and leaves, but similar specimens occur in the area. 16. ACANTHOPANAX (Decne & Planch.) H. Witte, Ann. Hort. Bot. 4 (1861) 89; Miq. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 10; Bth. in B. & H. Gen. PI. 1 (1865) 938; Seem. J. Bot. 5 (1867) 238; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 49; Merr. Philip. J. Sc. 1 (1906) Suppl. 217; Hutch. Gen. Fl. PI. 2 (1967) 69. — Panax subg. Acantho- panax Decne & Planch. Rev. Hort. IV, 3 (1854) 105. — Fig. 39. Small trees or scandent shrubs, prickly or unarmed. Leaves digitately compound, stipules absent, glabrous or with bristles. Inflorescence terminal, with umbels either solitary or more usually umbellately or racemosely arranged; pedicels not articulated below the flower. Flowers hermaphrodite or sexually dimorphic. Calyx margin minutely dentate. Petals 4 or 5, valvate. Stamens as many as the petals, anthers dorsifixed, introrse. Ovary 2-4(-5)-celled; disk fleshy; styles 2, connate to about the middle. Fruit a subglobose drupe, crowned by the persistent bifid style, 2-4(-5)-seeded ; exocarp fleshy, endocarp chartaceous or cartilaginous. Endosperm smooth or undulate. Distr. About 30 spp. in eastern Asia and the Himalayan region, south to Malesia (2 spp.): Malay Peninsula, N. Sumatra (Gajo Lands), Philippines (N. Luzon). Ecol. Usually in montane forest or in secondary growth. Note. A sterile specimen from Gunong Iran, Cameron Highlands (Symington CF 36300) in the Kepong Herbarium, probably represents an undescribed species. KEY TO THE SPECIES 1. Twigs with prickles. Leaf margins without bristles 1. A. trifoliatus 1. Twigs without prickles. Leaf margins with bristles 2. A. malayanus 1. Acanthopanax trifoliatus (L.) Merr. Philip. J. rays c. 4-5, or rarely solitary, slender or the central Sc. 1 (1906) Suppl. 217. — Zanthoxylum trifoliatum ray stronger (the laterals sometimes apparently LiNNE, Sp. PI. (1753) 270. — Panax aculeatus An. male), with minute furfuraceous bracts at their Hort. Kew. ed. 1, 3 (1789) 448. — A. aculeatus bases, occasionally with a few prickles, glabrous, H. Witte, Ann. Hort. Bot. 4 (1861) 89, nom. illeg.\ c. 3-6 cm long; secondary rays (pedicels) numerous, Seem. J. Bot. 5 (1867) 238; Harms in E. & P. Nat. slender, c. \-Vu cm. Calyx a rim with 5 minute Pfl. Fam. 3, 8 (1894) 50. teeth. Petals 5, ligulate, c. 2 mm long. Stamens 5, Scandent shrub, c. 2-8 m, with broadly based filaments c. 2 mm. Ovary turbinate, c. l'/2 mm high, recurved prickles sparsely disposed on the branches, 2-celled; styles 2, connate to about the middle, usually below the nodes; buds with small brown Fruit a spheroidal drupe, c. 5 mm when dry, cataphylls. Leaves disposed along the branchlets; crowned by the persistent bifid style; endocarp petiole up to c. 5 cm, flattened above, with a chartaceous. Endosperm surface slightly undulate, slightly dilated base, glabrous, usually with a few Distr. From the Himalayas through S. China prickles; leaflets (l-)3(-5); petiolules up to 8 mm, to Japan and Formosa; in Malesia: Philippines articulated with the petiole, channelled above; (N. Luzon: Benguet; Bontoc). blade up to 3V2 by 2^4 cm, broadly ovate to subro- Ecol. In montane forest and thickets, 1100- tund, apex acute, base cuneate, margins serrate, 1400 m. chartaceous, glabrous, lateral veins conspicuous on both surfaces, pinnate, reticulation inconspicuous. 2. Acanthopanax malayanus M. R. Henderson, Inflorescence terminating the main or lateral Card. Bull. S. S. 7 (1933) 105, pi. 22. — Fig. 39, branches, a sessile compound umbel; primary Unarmed tree to 17 m, trunk to 1 m 0; branch- 104 Flora Malesiana [ser. I, vol. 9^ "Ss- Fig. 39. Acanthopanax malayanus Henderson, a. Habit, x '/a, b. flower, x 7, c. fruit, x 4 (a, 6 de Wilde c.s. 16725, c DE Wilde c.s. 15925). Drawn by W. R. Philipson. 1979] Araliaceae — I (Philipson) 105 lets rather stout (c. 5 mm 0) ; bark greyish brown, glabrous, smooth; buds with small brown cata- phylls. Leaves crowded at the ends of the branch- lets; petiole terete, striate, glabrous, with a slightly dilated base, up to I2V2 cm; leaflets 3-5; petiolules up to 7 mm, articulated with the petiole, channelled above; blade up to 17 by 5'/2 cm, ovate-lanceolate to narrowly elliptic, apex acute, base cuneate or one side truncate, chartaceous, the principal veins pinnately arranged and conspicuous, the minor veins forming a dense reticulation, margin with numerous spinulose teeth. Inflorescence a terminal, sessile, compound umbel, often on short lateral shoots, apparently dioecious; primary rays c. 3-7, slender, terete, glabrous, striate, c. 4-8 cm; secondary rays (pedicels) of male flowers numer- ous, slender, IV2-2 cm, of female flowers stouter. Calyx a rim with 4-5 minute teeth. Petals 4-5, ligulate, c. 2V2 by IV2 rnm long. Stamens 4-5, filaments c. 2^U mm long. Ovary turbinate, 3^- celled ; styles 2, at first connate, but becoming free to about the middle. Fruit an oblate spheroidal drupe, c. 10 by 8 mm when dry, with a small stylo- podium bearing a persistent bifid style; endocarp cartilagineous. Endosperm surface strongly undu- late. Distr. Malesia: Malay Peninsula (Pahang; Cameron Highlands), N. Sumatra (Gajo Lands). Ecol. Montane rain-forest and mossy forest, 1400-2600 m. Vern. Berlaki, Sakai lang. Notes. The interpretation of this species as having sexually dimorphic inflorescences cannot be proved with the available material. This appears to consist of twigs bearing either inflorescences of male flowers, or infructescences. Appearances suggest that the putative male flowers have rudi- mentary ovaries. There is no evidence whether the fruiting flowers had produced pollen. The tree is said to be conspicuous when flowering by reason of its delicate feathery foliage, which is pale green with a reddish tinge, the petioles and inflorescence also being reddish. Excluded Meryta colorata F. M. Bailey, Queensl. Agric. J. 3 (1898) 283; Harms, Bot. Jahrb. 56 (1920) 384. The type specimen of this species cannot be located, but the original description does not seem to relate to a member of the Araliaceae, and certainly not to a Meryta. Panax ? anisum DC. Prod. 4 (1830) 254. — Anisum moluccamm Rumph. Herb. Amb. 2: 131, t. 42. — Nothopanax ? anisum MiQ. Fl. Ind. Bat. 1, 1 (1856) 766; Seem. F1. Vit. (1866) 114. — Polyscias anisum Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 45. According to Merrill (Int. Rumph. 1917, 289) these names are all based on the Rumphian description and plate and must refer to some species of Fagara, possibly F. avicennae Lamk = Zanthoxylum avicennae (Lamk) DC. {Rutaceae). CYPERACEAE— ir (J. H. Kernf & H. P. Nooteboom, Leyden) 28. CAREX LiNNE, Gen. PI. ed. 5 (1754) 420; Sp. PI. (1753) 972; Endl. Gen. (1836) 110; KuNTH, En. 2 (1837) 368; Steud. Syn. 2 (1855) 182; Miq. F1. Ind. Bat. 3 (1856) 346; BooTT, Illustr. Genus Carex I-IV (1858-67); Boeck. Linnaea 39 (1875) 14; ibid. 40 (1876) 327; ibid. 41 (1877) 145; B. & H. Gen. PI. 3 (1883) 1073; Pax in E. & P. Nat. Pfl. Earn. 2, 2 (1887) 122; Clarke, F1. Br. Ind. 6 (1894) 699; J. Linn. Soc. Bot. 37 (1904) 1 ; Philip. J. Sc. 2 (1907) Bot. 107; Kuk. Pfl. R. Heft 38 (1909) 67; Philip. J. Sc. 6 (1911) Bot. 57; Ohwi, Mem. Coll. Sc. Kyoto Imp. Un. Bll (1936) 232; Nelmes, Kew Bull. (1946) 5; Reinwardtia 1 (1951) 221; ibid. 2 (1954) 373; Raym. Mem. Jard. Bot. Montreal n. 53 (1959) 17; Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1962) 149; Kern in Back. & Bakh. /. Fl. Java 3 (1968) 487. — Fig. 118-135. Perennial herbs with tufted or creeping rhizome, monoecious, rarely dioecious. Stems arising centrally or laterally, erect or obliquely erect, mostly triquetrous or trigonous, rarely subterete, solid or sometimes hollow, often clothed at the base by persistent leaf-sheaths or their fibrous remains. Leaves tristichous, usually narrowly linear, sheathing at the base, with a ligule at the junction of blade and sheath, rarely lanceolate or elliptic with a more or less distinct petiole and eligulate, mostly basal and subbasal, 0-several higher on the stem, the lower ones often reduced to bladeless sheaths; sheaths of the stem-leaves and bracts closed. Inflorescence paniculiform, racemiform or spiciform, more rarely reduced to a single spikelet. Spikelets 1-very numerous, terete, sessile or peduncled, few- to many-flowered, wholly male, wholly female, or bisexual (androgynous when male flowers above, gynaecandrous when female flowers above). Bracts foliaceous or glume-like, often sheathing, sometimes wanting. Base of the branches of the inflorescence usually with a utriculiform or ocreiform bracteole {cladoprophyllum) surrounding it. Flowers unisexual, naked, solitary in the axils of the spirally arranged glumes; male flowers consisting of 3 free or rarely more or less connate stamens; anthers linear; female flowers consisting of a single pistil enclosed in a bottle-shaped prophyll {utricle, perigynium). Style either continuous with the ovary and persistent, or articulated with it and deciduous, straight or flexuous, often incrassate at the base; stigmas 2 or 3, protruding through the small terminal orifice of the utricle. Vestigial rachilla (see Uncinid) rarely present. Utricles membranous, chartaceous, or coriaceous, bicarinate, sometimes winged, sessile or stipitate, beakless to strongly beaked, nerveless, nerved, or ribbed, glabrous, or pubescent or hispid, papillose or puncticulate or smooth, sometimes spongy at the base; beak truncate, obliquely cleft, bidentate, or bifurcate at the top. Nut trigonous (when stigmas 3), or lenticu- lar (plano-convex or biconvex; when stigmas 2), enclosed within the utricle. Distr. A large genus with 600 to 1000 spp., the majority of them outside the tropics. However, the most primitive section, Vigneastra with a compound, paniculate inflorescence and andro- gynous spikelets, occurs mainly in the tropics of the Old World, from sea-level up to 3000 m. This section is represented in Malesia with 1 1 spp., and is there by far the largest section. Subg. Carex is, besides with the 1 1 spp. of sect. Vigneastra, represented in Malesia by 25 sections and 49 spp., subg. Vignea with 6 sections and 6 spp. (1) Continued from volume 7, part 3 (1974) p. 753. (107) 108 Flora Malesiana [ser. I, vol. 9^ Of the 66 spp. in all, 12 spp. are endemic and mostly confined to one island. Of the other 54 spp. the majority occur also in the northern hemisphere, either widely or only in SE. and/or E. Asia. Of these, 25 spp. extend without noticeable disjunctions into Malesia, either to its western part or further eastward. Another 4 of the E. Asian spp. occur in Formosa and in Malesia only in the Philippines (mostly only in Luzon), viz 12. C satziimensis, 30. C rhynchachaenium, 32. C. dolichostachya, and 33. C. formosensis. Among the Asian spp. 4 show a distinct disjunction between SE. Asia and Java, viz 11. C vesi- culosa, 14. C. helferi, 63. C. nubigena, and 58. C. longipes (which is also once found in Celebes and once in New Guinea); all are mountain species. A similar disjunction is found in the lowland species 10. C. stramentitia and 38. C. tricephala; this disjunction is understandable as both are bound to a distinct seasonal climate which shows a similar pattern (see van Steenis, Reinwardtia 5, 1961, 420-429). Still larger disjunctions are shown by 5 E. Asian spp. which are in Malesia almost only found in New Guinea, Formosa being mostly the nearest station, viz 46. C. brachyathera, 47. C.finitima (also in N. Sumatra), 48. C. mich.auxiana, 56. C. bilateralis, and 61 . C. duriuscula; all are mountain plants. Not a few Carex spp. occur in both the northern and southern hemisphere (Australia, often Tasmania and New Zealand) and are also found in the intervening tropical zone in Malesia. There are 1 1 of such species in all. Among these the following 7 are found in more than one island, viz 6. C indica, 31. C. breviculmis, 39. C maculata, 40. C capillacea, 57. C brunnea, 5. C. hors- fieldii, and 64. C echinata, although the latter two are very rare. Four others are also transtropical but are in Malesia only found in New Guinea, and thus show almost the same disjunction between Formosa and New Guinea as mentioned above; these are 43. C. brownii, 49. C. pseudo- cyperus, 55. C gaudichaudiana, and 66. C. curta. They occur in Malesia only in bogs and marshes on (high) mountains. The southern hemisphere has contributed very little to the Malesian Carices, as there is only one southern species from Australia, New Zealand, and New Caledonia, viz 62. C. appressa, which is found in Malesia, and then again only in New Guinea at very high altitude. Summarizing, one cannot withdraw oneself from concluding that only the presence of moun- tain bogs, marshes and grasslands gives the opportunity for the maintenance of Carex in the Malesian tropics. This would explain why the big mountainous island of New Guinea — where this habitat is most abundantly represented, much more than in any other island — is so rich in species. It is then also understandable that this is the place where relict areas could maintain their last stand in the tropical mountains of Malesia. Ecol. Most species prefer moist or wet localities, some are found in rain-forest, others in open habitats like in mountain marshes, grassland, heaths, etc. Only 22 spp. grow below 1000 m, 8 of them belonging to sect. Vigneastra, 13 are found below 500 m, 4 of them of sect. Vigneastra; 14 spp. occur exclusively above 2000 m. 57. C. brunnea, which is found from 80 m up to 3400 m, has the largest altitudina! range. The majority of the Carex spp. is growing on more or less acid soils. Three species of the lowland tropics occupy a different habitat, and are bound to a seasonal climate, viz 10. C stra- mentitia, 38. C. tricephala, and 37. C. speciosa. The first two occur only in Java or Madura I., amongst others with the teak-forest; the third of these drought species occurs also in some local dry spots in E. Sumatra and SE. Borneo, and is obviously adapted to a slightly less severe dry season. Of the three mentioned species, C. tricephala and C. speciosa are confined to limestone, like one other species, 35. C. malaccensis, an endemic of N. Malaya (the Langkawi Is.). One species, 59. C. teinogyna, seems in Malesia to be confined to banks of swift streams and occupies a rheophytic habitat. Notes. The distinction of the sections is not always unanimously agreed on, and several species were put into different sections by different authors. Some sections are closely allied, the only difference being sometimes 2 versus 3 carpels. As this has great influence on the shape of the nut, it often is considered a very important character. In my opinion the difference is due to a minor genetic difference, and is taxonomically -not very important. Besides, in some collections 2-carpelled and 3-carpelled fruits are found in the same inflorescence, e.g. in species of sect. 1979] Cyperaceae— II (Kern & Nooteboom) 109 Graciles, which has normally 2 carpels, and of sect. Oligostachyae (Decorae) which has normally 3 carpels (Noot.). Acknowledgement. The manuscript of Carex was for the greater part elaborated by Dr. J. H. Kfrn before he died in 1974. The obstacle for finishing it in time was his worry about the status cq. evaluation of a number of names in sect. Vigneastra, which also prohibited the finishing of the practical key to the species. Dr. H. P. Nooteboom has done this evaluation and he made the final draft of the key, for which he is responsible. He is also responsible for the treatment offered here for several species, viz 3. C. cniciata, 18. C. perakensis, 19. C. tunita, 20. C. verticil- lala, and 31. C breviculmis, and for some critical notes which have been marked "Noot." between brackets. The division in subgenera and sections is largely from Dr. Kern. The revision of Uncinia is entirely by Dr. Nooteboom. (Ed.) Explanation of some terms. The term 'stems' refers to the flower-bearing stems, except when stated otherwise, whereas 'inflorescence' means the entire flower-bearing stem including all the spikelets. The terms used for the inflorescence ('panicle', 'raceme') indicate the disposition of the spikelets, not of the individual flowers. 'Stems central' means that the leaves are arranged around the flower-bearing stem (usually towards the base). 'Stems lateral' means that the leaves are placed on a vegetative shoot, and that the flower-bearing stems arise separately from the rhizome and bear few short-bladed leaves. The flowering stems can also be situated in the axils of the leaves. This situation is sometimes difficult to distinguish from 'stems central', because the leaves are often crowded in a basal rosette. The bracts are inserted on the stem and they are sheathing or not; they bear in their axils a spikelet or a partial inflorescence, the peduncles of which are often basally surrounded by a cladoprophyllum; this is a bracteole which is utriculiform or ocreiform. If the cladoprophyllum is ocreiform, it usually is hidden entirely in the sheath of the bract. Each flower is solitary in the axil of a glume; several flowers together form a spikelet which can be S, ?, or bisexual; in the latter case the S flowers are usually placed only at the base or at the apex of a spikelet. Gynaecan- drous spikelets have ? flowers above, 3 below, androgynous spikelets have S flowers above and $ below. The figures 118-123, depicting utricles and nuts of nearly all the species, and some glumes (fig. 132), are given to facilitate use of the keys. KEY to subgenera AND SECTIONS as represented in Malesia 1 . Stigmas 3 and nut trigonous, or, when stigmas 2 and nut lenticular (plano-convex or biconvex) some or all of the spikelets peduncled or terminal spikelet c?- Spikelets either dissimilar in appearance (the terminal one S and some or all the lateral ones ?), or similar in appearance (all bisexual). Base of the peduncles of the lateral spikelets or of the branches of the inflorescence surrounded by a utriculiform or ocreiform bracteole {cladoprophyllum); inflorescence sometimes reduced to a single terminal spikelet I. SuBO. Carex 2. Stigmas 3; nut trigonous. 3. Inflorescence consisting of more than one spikelet. 4. Lateral spikelets arising from a gaping, utriculiform, fertile {i.e. nut-bearing) cladoprophyll. Spikelets androgynous, up to 15 mm long. Bracts not sheathing. Stems 3-20 cm tall. Sp. 12 2. Sect. Japonicae 4. Lateral spikelets arising from a sterile cladoprophyll, or sessile without cladoprophyll. 5. Stems scapiform, surrounded at the base by spadiceous sheaths in place of leaves. Leaves wide, conduplicate-petioled below. Secondary panicles contracted, very dense. Utricles with very long linear beak and very oblique mouth. Sp. 14 4. Sect. Mapaniifoliae 5. Otherwise. 6. Spikelets peduncled, in fascicles in the axils of at least some of the sheathing bracts, all androgy- nous or some wholly (^ or ?. Utricles long-beaked. tSpp. 17-20 6. 5ecr. Oligostachyae 6. The peduncled or sessile spikelets or secondary panicles single or binate in the axils of the bracts. 7. Spikelets similar in appearance, androgynous (upper flowers S, lower ones ?). 8. Lower bracts not or only shortly (some millimeters) sheathing. 9. Apex of nut truncate, ^U-^U mm 0, hollowed out. Style centred in the apical hollow of the nut. Utricles lageniform or rhomboid-lageniform. Spp. 27-30. . . 10. Sect. Lageniformes 110 Flora Malesiana [ser. I, vol. 9^ 9. Apex of nut not truncate. Style base pyramidally thickened, often broader than apex of nut. Spp. 35-38 12. Sect. Radicales 8. Lower bracts long sheathing. 10. Flowering stems arising from the axils of the leaves which are crowded on a short stem and much longer than the flowering stems. Style flexuous. Nut with irregularly excavated sides, seated on a spongy, disk-like stipe. 5p. 26 9. Sect. Cryptostachyae 10. Flowering stems arising either from the centre of the leaves, or from lateral shoots bearing a few short-bladed leaves. 11. Spikelets small, up to 15(-20) mm long. 12. Stems central. Spikelets numerous, ovoid or oblong-ovoid, arranged in often diffuse, compound panicles. Bracts long. 5p/7. 1-11 1. 5ec?. Vigneastra 12. Stems lateral. Spikelets few, globose-ovoid, in a simple panicle. Bracts shorter than 8 cm, the sheaths not included. Sp.l\ 7. Sect. Surculosae 1 1 . Spikelets longer. 13. Utricles densely many-nerved. Spikelets l-8(-14). 14. Utricles ellipsoid, short-beaked, more or less ciliate on the margins, spongy-thickened at the base. Spikelets l-3(-4). Spp. 35-38 12. Sect. Radicales 14. Utricles narrowly ellipsoid, long-beaked, more or less hispidulous on the faces, not spongy-thickened at the base. Spikelets 2-8(-14). Spp. 17-20 . 6. Sect. Oligostachyae 13. Utricles distinctly but not densely nerved. Leaves with reddish sheaths splitting into fibres. Spikelets 6- numerous. 5pp. 15-16 5. 5ecr. Polystachyae 7. Spikelets dissimilar in appearance, unisexual (or rarely gynaecandrous), the terminal one or some upper ones c?, the remainder 5 (occasionally the 5 spikelets may have a few (^ flowers, and the cJ spikelets a few 2 flowers). Or most of the spikelets 2, shortly cylindrical, the S spikelets lateral, 1-5 just below some of the terminal $ spikelets. 15. Leaves not crowded in a basal rosette but inserted all over the stem, the upper ones merging into leafy bracts, the lower ones gradually decreasing in length and merging into bladeless sheaths. 16. Inflorescence a compound panicle consisting of 5-8 secondary panicles each containing numerous 2 spikelets and 1-5 (^ spikelets. Spikelets small, 4-8 mm long. Utricles glabrous, with short, V4 mm long beak. Leaves sparsely hairy beneath., 5p. 13 3. Sect. Hypolytroldes 16. Inflorescence simple, racemose, consisting of 4-9 spikelets; terminal spikelet (^ (sometimes a second, smaller one added), remaining spikelets $, IV2-4 cm long. Utricles densely hispid, with distinct, c. 1 mm long beak. Leaves glabrous. Sp. 50 21. Sect. Occlusae 15. Leaves crowded at the base of the stem, often also some higher on the stem. 17. Nut excavated in the middle, the apex abruptly narrowed into a conspicuous, hastiform, i.e. cylindric, V2-V3 mm long beak expanding into the annulate apex. Spp. 22-25 8. Sect. Rhomboidales 17. Nut without hastiform beak. 18. Apex of nut truncate, ^U-\ mm wide, hollowed out. Style centred in the apical hollow of the nut. Utricles lageniform or rhomboid-lageniform. Spp. ll-ZQ 10. Sect. Lageniformes 18. Apex of nut not truncate, at most V2 mm wide. 19. Nut mitrate, i.e. contracted at the apex and then expanded into a discoid annulus. Utricles ovoid or ellipsoid. Spp. 31-34 11. Sect. Mitratae 19. Nut not mitrate. 20. Utricles densely papillose, erostrate or but shortly beaked. Sp. 39 13. Sect. Trachychlaenae 20. Utricles not papillose, distinctly beaked. 21. $ Spikelets broadly ovoid or subglobose, with few narrowly lanceolate, c. 1 cm or more long utricles. Bracts sheathing. Rhizome short. Sp.A%. . . . 19. Sect. Folliculatae 21. ? Spikelets not broadly ovoid-subglobose. Utricles smaller. 22. Spikelets linear-cylindric, loosely flowered. Utricles nerveless (except for 2 submar- ginal nerves) or obscurely nerved, with long beak and oblique mouth. Bracts sheathing. 23. Utricles hispidulous, brownish. 5>. 46 17. Sect. Fernigineae 23. Utricles glabrous, light green. Sp. Al 18. Sect. Sylvaticae 22. Spikelets oblong or cylindric. Utricles many-nerved. 24. Spikelets short-cylindric, few-flowered. Utricles fusiform-ellipsoid or rhomboid, at least towards the base many-nerved. Flowering stems lateral or central. Spp. 22-25 8. Sect. Rhomboidales 24. Spikelets cylindric, densely many-flowered. 1979] Cyperaceae— II (Kern «fe Nooteboom) 111 25. Utricles firm, of thick texture, short-beaked. Rhizome emitting strong stolons. Leaves conspicuously septate-nodulose. Spikelets erect. Bracts not or scarcely sheathing. Sp. 60 26. Sect. Paludosae 25. Utricles membranous, with long beak (except in C. brownii ssp. brownii). 26. Spikelets erect. Utricles patulous, subinflated, many-nerved, with shortly bidentate beak. Style straight. Rhizome often creeping. Spp. 42-45 . 16. Sect. Anomalae 26. Lower spikelets cernuous. Utricles widely patent to refiexed, closely many-ribbed, with bifurcate beak. Style flexuous. Rhizome cespitose. Sp. 49 20. Sect. Pseudocypereae 3. Inflorescence consisting of a single terminal spikelet. 27. Utricles with whitish pubescent margins or hispidulous, densely and strongly many-nerved, 4-10 mm long. Spp. 35-38 12. Sect. Radicales 27. Utricles glabrous, l'/2-5 mm long. 28. Utricles rather large (4V2-5 mm long), with slender, linear, bidentate beak. Sp. 41 15. Sect. Rhizopodae 28. Utricles small (1V2-4 mm long), with short, subentire or slightly emarginate beak. Sp. 40 14. Sect. Capitellatae 2. Stigmas 2; nut lenticular (plano-convex or biconvex). 29. Bracts sheathing. Spikelets often in fascicles. Utricles often hispidulous, membranous, nerved, usually long-beaked. Stigmas often very long. Spp. 56-59 25. Sect. Graciles 29. Bracts not sheathing (or very shortly). 30. Lateral spikelets sessile or the lowest shortly peduncled, erect. Terminal spikelet o. Glumes muticous. 5;?. 55 24. Sect. Carex 30. Spikelets peduncled, cernuous. Glumes often awned. 31. Spikelets 3-8, single at the nodes, cylindric, short peduncled. Terminal spikelet o, or (often with exception df basal spikelets) spikelets gynaecandrous. Spp. 52-54. ... 23. Sect. Praelongae 31 . Spikelets 6-50, single or binate at the nodes, the lower ones long-peduncled, all androgynous or some 0, linear-cylindric. Sp. 51 22. Sect. Longispicae 1. Stigmas 2 and nut lenticular. Spikelets similar in appearance, bisexual, sessile. Cladoprophyila usually absent IL Subg. Vignea 32. Spikelets androgynous (upper flowers o, lower ones 5). 33. Rhizome long-creeping. Spikelets 4-7. Utricles coriaceous, plano-convex, sharp-edged but not winged. 5/7. 61 27. Sect. Divisae 33. Rhizome short, forming dense tufts. Spikelets numerous or very numerous. 34. Utricles coriaceous, with rounded margins. Bracts inconspicuous. Sp. 62 28. Sect. Paniculatae 34. Utricles membranous, with acute or winged margins. Lower bracts foliaceous. Sp. 63 29. Sect. Multifiorae 32. Spikelets gynaecandrous (upper flowers, 1, lower ones o)- Utricles membranous. 35. Utricles with rounded margins, densely whitish-puncticulate, beakless or very shortly beaked. Spikelets ebracteate (bracts glumiform). Sp. 66 32. Sect. Heleonastes 35. Utricles with acute or winged margins, not puncticulate. 36. Utricles obliquely erect, also when mature. Spikelets more or less distinctly bracteate. Sp. 65. 31. Sect. Elongatae 36. Utricles widely spreading or reflexed when mature. Spikelets not or inconspicuously bracteate (bracts glumiform). Sp. (A 30. Sect. Stellulatae KEY TO THE SPECIES Only complete specimens with nearly or fully mature utricles are identifiable. As the distribution of sexes is variable, especially in the species with a terminal male spikelet, it is advisable to examine several specimens if possible. In the measurements of the length of the utricles the beak is included. Unless stated otherwise 'glumes' refers to the glumes subtending Xht female flowers. 1. Inflorescence consisting of a single (androgynous) spikelet terminating the stem. Stigmas 3; nut trigonous. 2. Utricles with whitish pubescent margins, or hispidulous, or winged and scabrid margined, 4-8 mm long. Glumes (many-)nerved. Style pyramidally thickened towards the base, persistent on the nut. Leaves (2-)3-10 mm wide. 3. Spikelets 2-8 cm long. Utricles with whitish pubescent margins 37. C. speciosa 3. Spikelets 6-12 mm. Utricles hispidulous or glabrous with scabrid margins. 112 Flora Malesiana [ser. I, vol. 9^ 4. Utricles hispidulous in the upper ^/j, 4-6 mm, with conic-cyhndric, I-2V2 mm long beak. 38. C. tricephala 4. Utricles glabrous with scabrid, distinctly winged margins, 6-8 mm, gradually narrowed into the 3 mm long beak 36. C. ramosii 2. Utricles glabrous, membranous and otherwise different, 2-5 mm long. Glumes 3-nerved in the centre, otherwise nerveless. Style not or but slightly thickened towards the base. Leaves '/2-3 mm wide. 5. Utricles (I V2-)2','2-3'/2(-4) mm long, rather gradually tapering into a very short, subentire or slightly emarginate beak. Leaves ' ,-2 mm wide. Glumes F/2-3 mm long ... 40. C. capillacea 5. Utricles 4V2-5 mm long, abruptly narrowed into a slender, linear, bidentate, c. I'/j mm long beak. Leaves 2-3 mm wide. Glumes 3-4 mm long 41. C. eremostachya 1. Inflorescence consisting of 2 or more spikelets. 6. Terminal and most of the other spikelets 2, shortly cylindrical, the ^ spikelets lateral, 1-5, just below some of the terminal 2 spikelets. Inflorescence a compound panicle. Leaves all over the stem 13. C. hjTX)lytroides 6. Terminal spikelet bisexual or wholly £. 7. Terminal spikelet wholly o (exceptionally in some specimens with a few 2 flowers), or when spikelets in fascicles, 1-2 spikelets of the upper fascicle wholly 3. 8. Stigmas 3. 9. Leaves inserted all over the stem, the upper ones merging into leafy bracts, the lower ones gradu- ally decreasing in length and merging into bladeless sheaths. Inflorescence simple, racemose, consisting of 4-9 spikelets. Spikelets P ,-4 cm long. Utricles densely hispid, with distinct, c. 1 mm long beak. Leaves glabrous 50. C. maubertiana 9. Leaves basal and often also some higher on the stem. 10. Nut excavated in the middle, the apex abruptly narrowed into a conspicuous, cylindric, V2-IV4 mm long beak expanding into the annular apex 22. C. anomocarya 10. Nut different. 11. Apex of nut truncate, ^,4-1 mm wide, hollowed out. Style centred in the apical hollow of the nut. Utricles lageniform or rhomboid-lageniform. 12. Utricles 5-7 mm long. Apex of nut contracted into a cylindric, truncate, ^/j-l mm long and wide neck. 13. Leaves 5-10 mm wide. Beak of utricles 1*2-2 mm long. Stems arising from basal leaf axils 28. C. gracilispica 13. Leaves 1-3 mm wide. Beak of utricles^ 4-I mm long. Stems central 30. C. rhynchachaenlum 12. Utricles VU-^^li mm long. Apex of nut hardly or not contracted into a cylindric neck 27. C. breviscapa 11. Apex of nut not truncate, at most '2 mm wide. 14. Utricles densely papillose, subabruptly contracted into a very short, up to ' 2 mm long beak 39. C. maculata 14. Utricles not papillose, nut either mitrate or distinctly beaked. 15. Nut mitrate, i.e. contracted at the apex and then expanded into a discoid annulus. 16. Glumes of spikelets cup-shaped, the margins more than halfway connate in front. Stamens monadelphous (not always so in the lowermost flowers). Utricles many-nerved 34. C. tristachya 16. Glumes of o spikelets with free margins. Stamens free. 17. Nut rhomboid with deeply concave faces 33. C. formosensis 17. Nut (long-)ellipsoid to ovoid or obovoid. 18. Leaves 3-10 mm wide. Stems arising from basal leaf-axils. Utricles strongly many-nerved 32. C. dolichostachya 18. Leaves l-4(-6) mm wide. Stems central. Utricles nerveless to multinerved. 31. C. breviculmis 15. Nut not mitrate. 19. Spikelets in fascicles of 3-20 from the axils of at least one of the bracts. 20. Utricles narrowly ellipsoid, 4-6^/2 mm long, gradually tapering below into a stipe-like, ^U-\ mm long base, above into a 1-3 mm long beak 20. C. verticillata 20. Utricles ovoid or ellipsoid, 2V3-3S'2 mm long, scarcely stipitate; beak ^^-l mm long. 17. C. celebica 19. Spikelets solitary (exceptionally binate) from the axils of the bracts, or crowded at the af)ex of the stem. 21. Utricles (8-)12-13 mm, many-nerved. ? Spikelets broadly ovoid to subglobose, 15-25 mm long and wide, with few, divergent, narrowly lanceolate utricles. Bracts long-sheathing 48. C. michauxiana 1979] Cyperaceae— JI (Kern & Nooteboom) 113 21. Utricles at most T'/j mm long. Other characters not so combined. 22. Broader leaves 1-2 cm wide. Spikelets 5V2-16cm long. Glumes vinaceous to dark red with a green, central stripe. StoloniferoLis 45. C. olivacea 22. Broader leaves at most 10 mm wide. Spikelets usually much shorter. 23. Stems lateral. Lateral spikelets 6-12 by 4-7 mm, V or androgynous. Utricles at least below many-nerved, 5-7 mm long. Glumes 2V2-4 mm. Nut 3-4 mm. 24. Leaves 3-7 mm wide. Glumes 2V2-3'/2 mm, with a l'/2-6 mm long awn. Lateral spikelets ? 24. C. lateralis 24. Leaves l'/2-3 rnm wide. Glumes c. 4 mm long, muticous or mucronulate. Lateral spikelets androgynous 25. C. loheri 23. Stems central. Lateral spikelets more than 10 mm long, 9 or rarely androgynous, when shorter than 12 mm nut 2'/2-3 mm. 25. Utricles 5-7'/2 nim long; nut at least 2'/4 mm long. 26. Glumes 2-2V2 mm, with an antrorsely scabrous, 2-4 mm long awn. Utricles with deeply furcate l'/2-2 mm long beak, strongly and densely many-nerved, when ripe widely spreading or reflexed. Lower bracts much overtopping the inflorescence, scarcely sheathing 49. C. pseudocyperus 26. Glumes 3-6 mm long, whether or not awned. 27. Utricles nerveless or obscurely few-nerved, except for the 2 marginal veins, 5-7V2 mm long. $ Spikelets linear-cylindric, lax-flowered, 2-9 cm by 3-7 mm. Glumes 3'/2- 6 mm, oblong to ovate, acute to very obtuse, translucent, with broad white margins and greenish midrib, the latter not reaching the apex, but sometimes excurrent into a short awn up to l(-2) mm long 47. C. finitima 27. Utricles (strongly) many-nerved. 28. Utricles 5 '/2-7V2 mm, fusiform. Glumes oblong-ovate, acute, very thin, dirty white with 3-nerved greenish central stripe, 3-572 mm long, muticous or mucronulate, rarely with a short awn 23. C. jackiana 28. Utricles 5-5V2 mm. Glumes ovate, deeply emarginate at the top, pale with purplish margins, c. 3 mm long, the strong midrib excurrent into a firm, smooth or scabrid awn as long as the glume 60. C sp. (^ Palndosae) 25. Utricles 3-5(-7) mm long; nut at most 2'/2 mm long, but mostly shorter; if the utricles are longer than 5 mm, the nut is at most P/3 mm long. 29. Utricles sparsely to rather densely hispidulous, nerveless except for 2 submarginal nerves, 3-4V2 mm long, with stout, l-l'^mmlong beak. ? Spikelets linear-cylindric, 1 V2-6 cm by 2V2-3V2 mm. Glumes 3-4'/2 mm, with an hispidulous awn up to 1 mm long 46. C. brachyathera 29. Utricles glabrous, many-nerved or -ribbed. 30. Utricles 4-5(-7) mm long, strongly many-nerved. Glumes 2-2V2 mm, with a 2-4 mm long awn. ? Spikelets 2V2-5 cm by 7-15 mm. Leaves 6-10 mm wide. 49. C. pseudocyperus 30. Utricles 3-4 mm (up to 6 mm in C. brownii ssp. transversa), strongly many-ribbed or plurinerved. Glumes IV2-2V2 mm, excurrent into a V2-3V2 mm long awn. ? Spikelets IV2-I2 cm by 4-6 mm. Leaves 2-10 mm wide. 31. Utricles strongly many-ribbed. ? Spikelets short-cylindric to cylindric, 1V2-3 cm by 5-6 m.m. Glumes 1-2 mm with a V2-3V2 mm long awn. Nut 2V4-2V2 mm. Leaves 3-4 mm wide 43. C. brownii 31 . Utricles plurinerved. ? Spikelets (narrowly) cylindric, (1 V2-)2V2-12 cm by 4-6 mm. Nut IV2-2V4 mm. Leaves 2-10 mm wide. 32. Plant densely cespitose, without stolons. Lower bracts long sheathing. Utricles olive-brown to fuscous. ? Spikelets 2V2-I2 cm long 44. C. oedorrhampha 32. Plant stoloniferous. Bracts not sheathing. Utricles pale greenish to stramineous. ? Spikelets (lV2-)2-5 cm 42. C. alopecuroides 8. Stigmas 2. 33. Utricles densely covered with raised glandular papillae, slenderly nerved or nerveless. At least the lower spikelets subcernuous to pendulous. Lowest bract much overtopping the inflorescence. 34. Glumes acute to obtusish, muticous or only the lower ones excurrent in a short, up to 1 mm long awn. Utricles slenderly nerved, 3-4 mm long 53. C. pruinosa 34. Glumes truncate or bilobed-emarginate, distinctly awned (awn V2-2 mm long). Utricles mostly nerveless, 2V4-3V2 mm 52. C. phacota 33. Utricles not papillate, whether minutely punctulate or not. 114 Flora Malesiana [ser. I, vol. 9^ 35. Stems 3^0(-75) cm by I-l V2 mm. Utricles distinctly 8-14-nerved, 2-3 V2 mm. Spikelets erect or suberect, terminal <^, remainder ? or with a few