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REPUBLIK INDONESIA REPUBLIC OF INDONESIA LEMBAGA ILMU PENGETAHUAN INDONESIA (L.LP.L) INDONESIAN INSTITUTE OF SCIENCES
FLORA MALESIANA
BEING AN ILLUSTRATED SYSTEMATIC ACCOUNT OF THE MALESIAN FLORA / INCLUDING KEYS FOR DETERMINATION / DIAGNOSTIC DESCRIPTIONS | REFERENCES TO THE LITERATURE|/|SYNONYMY]/AND DISTRIBUTION / AND NOTES ON THE ECOLOGY OF ITS WILD AND COMMONLY CULTIVATED PLANTS
PUBLISHED UNDER THE AUSPICES OF LEMBAGA BIOLOGI NASIONAL BOTANIC GARDENS OF INDONESIA / BOGOR / JAVA AND OF THE RIJKSHERBARIUM / LEYDEN / NETHERLANDS
PREPARED ON ANINTERNATIONAL 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 VOLUME 9
SPERMATOPHYTA
GENERAL EDITOR:
Dr C; G. G. J. VAN SITEENIS DIRECTOR OF THE FOUNDATION ‘FLORA MALESIANA’
PUBLISHED BY:
MARTINUS NIJHOFF / DR W. JUNK PUBLISHERS THE HAGUE / BOSTON / LONDON WONT — iN Oks)
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 Center
P.O. Box 322
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The Netherlands
Library of Congress Cataloging in Publication Data
(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, Cornelis Gijsbert Gerrit Jan van, ed.
II. Indonesia. Departemen Pertanian. III. Series.
QK366.F56 582 .09598 (2p le
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 © 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 NETHERLANDS
CONTENTS ERROR ene Se bea CNG ia «6 aa «1a oleae ale ante d sr BSE bas ar aie (3) UICTIINI Eo as ee a a wid ea C28OCE GE eek ss ee (5) Dedication by R.E.G. Pichi Sermolli & C.G.G.J. van Steenis ...................- (7) aia RISENTIS ORINIT SIDED ete aa. cee os 2 2's? Jas 3S P a eke ae ere Pee bee ee ee (45)
TAXONOMICAL REVISIONS
in alphabetical sequence
aeaemnimeene —T ENY WY, Be; ENDENENSOOEN eet sin ca ae as gs eta a i cre 1
eyperaceac—IP by: 3-H. Kern .&,H:-P..Nooteboom. 22825 2e> ene eee ss ee ee 107
Pcrocm pacene by PS. AShtonis... 33.9 2n;s ke ee ee oe eee 237
mehecne—F By: 1:P. Jessop). cia. 2 nsec ton oan. Ree eee eee 189 ADDENDA
to volumes 4—9
Addenda, corrigenda et emendanda by C.G.G.J. van Steenis ef al. ............... 553
INDEX
Index to scientific plant names by M.J. van Steenis-Kruseman & E.E. van Nieuwkoop 575
1906
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Dedicated to the memory of ODOARDO BECCARI
DEDICATION
A dedication to ODOARDO BECCARY, 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 Nelle foreste di Borneo, Viaggi e ricerche di un naturalista (1902), excellently translated into English (in a somewhat abbreviated form) by Prof. E. GicLiotiand revised and edited by F.H.H. GUILLEMARD as 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 BEccaRiin 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- Nuccl, from Radda in Chianti in Tuscany, died soon after his birth, and his father GruSEPPE 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 Minuccl. 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 Abbé IGNAzio MezzetTtT1! and by his Professor of Botany in the Lyceum of Lucca and Director of the Botanic Garden, CESARE Biccui. 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.
Beccarr’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 Biccut, Agg. Fl. Lucch. (1860) 21, nom. nud.; 1 Giardini 8 (1861) 50, t. 2.
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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 WeBp’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, BECCARIcommenced his studies in the Faculty of Natural Science at the University of Pisa. At first, perhaps under the influence of Biccut, he devoted his attention to the Cryp- togams and already in 1862 BECCARIs 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 PlETRO SAvi made 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, BEccaARthad 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 OpoarRpDo, 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 Doria 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 BEccARIand 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
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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 BEccaRt 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 May 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, BEccarr 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
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FLORA MALESIANA
to overcome some impassable rapids or to cross the 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, BEccaARI arrived at Kuching where he concluded this long, hard and risky enterprise.
In the first two years of his stay in Borneo BEccarr’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 BEccARIwas 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 BEccarti had prob- ably assembled his collections. The house soon became a meeting place for young lovers of natural history including LEvreR and Sommrer. 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 (CESATI 1876), mosses (HAMPE 1872), lichens (VON KREMPELHUBER 1875) and hepatics (DE NoTaris 1876), efc. (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 Giornale Botanico Italiano, which was intended as a replacement of the Giornale Botanico Italiano founded in 1844 by F. PARLATORE, but interrupted in 1852. BEccaARI edited three volumes of the new periodical (1869, 1870, 1871) with the help of his friend Marcuccl, 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
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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 (315 species of sper- matophytes and pteridophytes and 289 species of mosses, algae, fungi and lichens) enumerated and partly described in MARTELLI’ 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 Luici MarRIA 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. Kraatand his wife, the Italian lady AMALIA 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, Beccarti 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’ALBERTISs 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 BEccariable 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 difficulties and serious health troubles, he had assembled a col- lection of about 700 species of plants and a rich amount of zoologica!, 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 KraaLand 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
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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. BEccARtI 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, BEccaRIsailed 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. BEccarthad 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 Beccarr’s funds had been running out, but early in 1874 he had already written from Makassar to his friend G. Dortain 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, BEccartsoon 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, BEccarileft 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 ‘Deli’ with a crew of 10, and accompanied by 8 men and a young boy for collecting plants and animals.
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Dedication
He sailed on January 22, 1875, 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 18 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 Gurpo 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. GiGLioti, 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, BECCARIreceived 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 BEccaARr’s botanical and zoological in-
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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, Beccart 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 1878, 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 BEccarr’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 BECcCaARr’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.
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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 Beccarr’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. BEccaARI 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 Beccarr’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, BEccart 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 BEccaRr’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 BEccarr’s collections. The Royal Imperial Museum of Physics and Natural History in Florence was founded in 1775 by PreTRo 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 19th 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 Pump BARKER WEBB died in Paris and bequeathed his invaluable herbarium and library to
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the Florence Museum, together with an annual income of 6945.58 lira (derived from the sale of a palace in Paris) and known as Wesp’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.
Fitippo PARLATORE died on September 9, 1877. 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 BEccaRr’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 BEccaARI 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-
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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 Beccaritook 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 Beccarr’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. DortA, the Director of the Civic Museum of Natural History for an appraisal of BEccaRr’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. BEccaRt 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
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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 Beccaricame 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 Wess 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 BEccarthad 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
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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. BEccArt 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 Semplici. 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 Semplici and adjoining buildings near San Marco, on the op- posite side of the river Arno. However, BEccartdid not give in, and he continued to publish 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 1896, 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, BEccaARI 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
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ancient Giardino dei Semplici, was brought to a conclusion.
However, let us resume the course of BEccart’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 FLAMIN], from a noble family of Ca- sentino, in the high valley of the Arno. They had four sons: NELLo, Dino, 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. Picut 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. Ricasott, 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 of 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
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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 BEccarRt’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 Beccarthad 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 BECcaRI to overcome his scientific inactivity, but in the early days of May 1897 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, area 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 BECCARIon June 10, 1897.
The preparation of the book, Nelle foreste di Borneo, led BECcaRI to recover a certain interest for scientific work and it was not difficult for Prof. OrREsTE 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 publication of his book. However, BEccarthad 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
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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 GEorRGE 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 PHit1p 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 Beccarrand 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.
Beccariled 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 BEccarr’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 BEccaART’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 BEccARIon the expedi- tions to Borneo and the Red Sea. BEccaRI paid attention to special groups and assembled several
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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 BEccaArRI’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).
Beccari also 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 BEccarr’s books Nelle foreste 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. GiGLioL1, 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. MANTEGAZzZzA and E. REGALIA.
BEccaRr’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 Beccarthad 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’ 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 BEccarr’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.
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After about a decade of complete scientific inactivity, due to the sad vicissitudes of his academic life and the end of Malesia, BEccaripublished Nelle foreste 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 Beccarimanaged 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. BEccarRi 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-
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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. BECCARI was 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 Beccarireturned from Borneo to Florence he founded the 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. GesTroin 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, efc. 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 Beccarthimself, 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 Beccartever 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
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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 Diets (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, BEccaARI 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 HooKker’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 |’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 subfamily Arecoideae, was completed and published by PicHiI SERMOL- L1in 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 BEccarRI are described by LuIGI PAMPALONI, Apparecchio fotografico universale per laboratorio biologico ideato dal Dottor Beccari. Rend. Congr. Bot. Naz. Palermo (1903) 164—168, cum fig., and Gli apparecchi microfotografici del Dott. O. Becca- ri. Bull. Soc. Fotogr. Ital. 14 (1902) 129—145, fig. 1—7.
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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 BEccARIShowed 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. Gicuiot1and 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 BeccarRi 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 CorRNER. 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 (1876) 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 (1889) on the flowering of Amorphophallus titanum, the gigantic Ara- cea, and was summarized in his book, Nelle foreste di Borneo.
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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 variability 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 BEccARtalso 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 lively 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
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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.
BeEccar! 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. Picut SERMOLLI & C.G.G.J. VAN STEENIS
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Appendix 1 — Bibliography of Odoardo Beccari
A — Books and papers (excluding reviews of books)
1861. Escursione botanica. — L’Araldo Cattolico, Lucca, anno XVIII, 14 agosto 1861, n. 33: 264 (not seen).
1862. Illustrazione dell’ Arnoldia cyathodes Massal. — Comment. Soc. Crittog. Ital. 1: 128—130, tewle
1868. Descrizione di tre nuove specie di piante Bornensi. — Atti Soc. Ital. Sc. Nat. 11: 197—198.
— Cenno di un viaggio a Borneo. — Boll. Soc. Geogr. Ital. 1: 193-214.
1869. Illustrazione di nuove specie di piante Bornensi. (Balanophoreae, Rafflesiaceae). — Nuovo Giorn. Bot. Ital. 1: 65—91, t. 2—5S.
—— Varieta e notizie. — Ibid.: 158—160.
— Lamenti del Redattore. Rivista bibliografica. — Ibid.: 222—224.
1870. Illustrazione di nuove specie di piante Bornensi. (Aristolochiaceae). — Nuovo Giorn. Bot. Ital. 2: 5—8, t. 1.
— Nota di una nuova specie del genere Stenomeris. — Ibid.: 8—12, t. 2.
— Nota sul Trichopodium zeylanicum Thw. — Ibid.: 13-19, t. 3.
— Nota sull’embrione delle Dioscoreaceae. — Ibid.: 149-155, t. 4.
—— Disepalum coronatum nuova specie di Anonacea bornense. — Ibid.: 155—156, t. 5.
—— (O. Antinori & A. Issel). Relazione sommaria del viaggio nel Mar Rosso dei Signori Antinori, Beccari e Issel. — Boll. Soc. Geogr. Ital. 5(2): 43-60.
1871. Descrizione di due nuove specie di Hydnora d’ Abissinia. — Nuovo Giorn. Bot. Ital. 3: 5—7.
— Petrosavia. Nuovo genere di piante parassite della famiglia delle Melanthaceae. — Ibid.: (Sle ieeale — Note sopra alcune palme Bornensi. — Ibid.: 11—30.
— Le Hydrocotyle d’Europa. — Ibid.: 102.
—— Sui generi Bihania ed Eusideroxylon. — Ibid.: 102—103.
— Cattedra di Botanica nell’Universita di Torino. — Ibid.: 103-104.
—— Illustrazione di nuove 0 rare specie di piante Bornensi. (Anonaceae). — Ibid.: 177—193, t. 2-7.
1874. Descrizione di una nuova specie del genere Myrmecodia della famiglia delle Rubiaceae. — Nuovo Giorn. Bot. Ital. 6: 195—197, t. 6.
1875. Osservazioni supra alcune Rafflesiaceae. — Nuovo Giorn. Bot. Ital. 7: 70—75.
1876. Le Capanne ed i Giardini dell’ Amblyornis inornata. — Ann. Mus. Civ. St. Nat. Genova 9: 382—400, t. 8.
—— Lettera del Dr. A.B. Meyer al Mse. G. Doria (con nota di O. Beccari). — Ann. Mus. Civ. St. Nat. Genova 8: 333-334.
1877. Relazione intorno alla Baia di Assab. — Cosmos 4: 230—232.
— Della organogenia dei fiori feminei del Gnetum gnemon L. — Nuovo Giorn. Bot. Ital. 9: 91—100, t. 7 (reprinted in Amer. Journ. Sc. 13: 469—471).
—— Sulla Cardiopteris lobata Wall. — Nuovo Giorn. Bot. Ital. 9: 100—108, t. 8.
— Die Flora der Aru-Inseln. — Ausland 50: 759-760.
—— Sul nuovo genere Scorodocarpus e sul genere Ximenia L. della famiglia delle Olacineae. — Ibid: 273=279 ts 11.
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Dedication
— Della disseminazione delle palme. — Bull. R. Soc. Tosc. Ortic. 2: 167—173.
—— II sagu della Nuova Guinea, Metroxylon rumphii Mart. — Ibid.: 247—249.
— Catalogue of the plants of the Fly River (New Guinea) collected by De Albertis, 1877. — Journal of Melbourne Logbook 30 Jan. 1877 (not seen).
— Le specie di palme raccolte alla Nuova Guinea da O. Beccari e dal medesimo adesso descritte, con note sulle specie dei paesi circonvicini. — Malesia 1: 7—96, t. 1 & 2.
— Nuove osservazioni sulle palme della Nuova Guinea. — Ibid.: 97—102.
— Studio monografico sopra le piante della famiglia della Icacineae e delle Menispermaceae sin qui scoperte nella Malesia e nella Nuova Guinea. — Ibid.: 103—165, t. 3—8.
1877—78. Piante nuove o rare dell’Arcipelago Malese e della Nuova Guinea, raccolte, descritte ed illustrate da O. Beccari. — Leguminosae-Caesalpinieae, Palmae, Chailletiaceae, Gymno- spermae papuanae, Coniferae, Gnetaceae, Cycadaceae, Violaceae, Magnoliaceae, Monimia- ceae, Araliaceae, Ericaceae, Vacciniaceae, Nepenthaceae, Corsia, Burmanniaceae. — Ibid.: 1 (1877) 167—192; contd. (1878) 193—254, t. 9-15.
1878. Sulle piante raccolte alla Nuova Guinea dal Sig. L.M. D’Albertis durante l’anno 1877, con descrizione di tre nuove specie di Icacineae. — Ibid.: 1 (1878) 255—256; contd. (1883) 257—258 (incl., pp. 257—258: Su di un nuovo genere della famiglia delle Olacineae).
— II Conophallus titanum Beccari. — Bull. R. Soc. Tosc. Ortic. 3: 290—293, f. 32.
1879. Le Nepenthes. — Bull. R. Soc. Tosc. Ortic. 4: 13-19.
— La pit piccola delle Araceae: Microcasia pygmaea Becc. — Ibid.: 179-181, cum fig.
1880. Notes on the plants collected by Sig. L.M. D’Albertis in New Guinea. — In: L.M. D’Alber- tis, New Guinea: what I did and what I saw. 2: 391—395.
—— Catalogue of the plants of the Fly River, 1877. — Ibid.: 396—400.
— Sul viaggio di O. Beccari in Sumatra. Giugno—Ottobre 1878. — Boll. Soc. Geogr. Ital. ser. 2, 5 (1880) 300—302.
— Lettera di O. Beccari al Preside della Facolta di Scienze naturali del R. Istituto di Studi Supe- riori di Firenze. Dicembre 1880 (Published by U. Martelli, Webbia 5 (1921) 329—330).
1881. R. Museo di Fisica e Storia Naturale. L’Orto botanico. — Giornale ‘La Nazione’ 6 Gennaio 1881.
—— (& G. Doria) Viaggio ad Assab nel Mar Rosso, dei signori G. Doria ed O. Beccari con il R. Avviso ‘Esploratore’ dal 16 Novembre 1879 al 26 Febbraio 1880. — Ann. Mus. Civ. St. Nat. Genova 16: 523—524.
— A proposito del progetto di remozione delle collezioni botaniche dal Museo di Storia Natura- le di Firenze (not seen).
— Sull’abbandono del museo e del giardino botanico della Specola a Firenze. Protesta dei culto- ri della botanica in Firenze colle adesioni e le osservazioni dei botanici italiani ed esteri. — Firen- ze, Tip. Sborgi, in 8°, 72 pp.
— Cenni sopra la Flora di Assab. — Bull. R. Soc. Tosc. Ortic. 6: 108—110.
—— Beitrage zur Pflanzengeographie des malayischen Archipels. — Bot. Jahrb. 1: 25—40 (con- densed version in German of Malesia 1: 214—238).
1883. Su di un nuovo genere della famiglia delle Olacineae. — Malesia 1: 257—258 (included in: Sulle piante raccolte alla Nuova Guinea dal Sig. L.M. D’Albertis durante l’anno 1877, con de- scrizione di tre nuove specie di Icacineae. See 1878).
1884. Piante ospitatrici, ossia piante formicarie della Malesia e della Papuasia descritte ed illustra- te da O. Beccari. — Ibid.: 2: 5—128, f. 1—9, t. 1-25; contd. (1885) 129—212, f. 10—15, t. 26—54; contd. (1886) 213—284, f. 16—17, t. 55—65.
—— Veratronia malayana Miq. — Bull. R. Soc. Tosc. Ortic. 9: 70—72, t. 4.
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— II The in Italia. — Boll. Notiz. Agrar. Minist. Agric. 6: 279—281; also in L’ Agricolt. meridio- nale 7: 116-118, and La Natura, Milano, 1: 267—268.
— II colera e le mosche. — La Natura 1884(33): 81-83.
1885. Orto Botanico di Firenze. — Giornale il ‘Corriere’, 14 Maggio 1885.
— Reliquiae Schefferianae. Illustrazione di alcune palme viventi nel Giardino Botanico di Bui- tenzorg. — Ann. Jard. Bot. Buitenzorg 2: 77—171, t. 1—14.
—— Cyrtosperma (Alocasia Hort.) johnstonii Becc. — Bull. R. Soc. Tosc. Ortic. 10: 5—7.
—— Plantes a fourmis de l’Archipel Indo-Malais et de la Nouvelle Guinée. (Compte rendu de Mr. E. Levier). — Archiv. Ital. Biol. 6: 305—341.
1886. Rivista delle specie del genere Nepenthes. — Malesia 3: 1—15, t. 1—3.
—— Rivista delle felci e licopodiacee di Borneo e della Nuova Guinea, enumerate o descritte dal Barone V. Cesati nella memoria che porta il titolo: ‘‘Felci e specie nei gruppi affini raccolte a Borneo dal Sig. O. Beccari’’ e dell’altra: ‘‘Prospetto delle felci raccolte dal Sig. O.B. nella Poli- nesia’’. — Ibid.: 16—55, t. 4—S.
— Nota sopra alcune felci raccolte dal Sig. J.E. Teysmann all’isola di Sumba o Sandal-wood ed in Timor. — Ibid.: 56—57.
— Nuovi studi sulle Palme Asiatiche. — Ibid.: 58—149, t. 6—11.
— Sulla Quercus robur Linn. Nota di botanica legale. Perizia nell’interesse della causa Le Pen- nec e Monetti contro Vitali, Picard, Charles e Comp.i. — Firenze, Tip. Nicolai, in 4°, 44 pp.
1887. Turgescenza dei petali di Magnolia Yulan. — Malpighia 1: 420.
1887—88. Le palme incluse nel genere Cocos Linn.: Studio preliminare. — Malpighia 1 (1887) 343-354, 441-454, t. 9; 2 (1888) 85—95, 147-156.
1888. Nuove specie di palme recentemente scoperte alla Nuova Guinea. — Nuovo Giorn. Bot. Ital. 20: 177-180.
1889. Nuove palme asiatiche. — Malesia 3: 169—200.
— Le Bombaceae Malesi descritte ed illustrate da O. Beccari. — Ibid.: 201—280, f. 1—16, t. 12—36 (plates 12—35 were published before the text in vol. 3, no. 3, August 1887).
— Fioritura dell’Amorphophallus titanum Becc. — Bull. R. Soc. Tosc. Ortic. 14: 250—253, 266—278, t. 8 (Reprint with separate pagination: 17 pp., 3 plates).
— Palmae. — In: K. Schumann & M. Hollrung, Die Flora von Kaiser Wilhelms Land (Beiheft zu den Nachrichten tiber Kaiser Wilhelms Land und den Bismarck-Archipel): 15—17.
1890. Le palme del genere Pritchardia. — Malesia 3: 281—317, t. 37—38 (the plates were published before the text in vol. 3, no. 4, September 1889).
— Le Triuridaceae della Malesia. — Ibid.: 318—344, t. 39—42 (the plates were published before the text in vol. 3, no. 4, September 1889).
—— Rivista monografica delle specie del genere Phoenix Linn. — Ibid.: 345—416, f. 17, t. 43—44.
1891. Letter to Prof. G. Papasogli. — In: G. Papasogli, La nitrobenzina usata come insetticida. — Agric. Toscana 9: ?—? [repr. 1—6].
1892—93. (& J.D. Hooker) Palmae. — In: J.D. Hooker, The Flora of British India 6: 402—448, 449-483.
1893. Palmae. — In: G.F. Scott Elliot, New and little known Madagascar plants, collected and enumerated by G.F. Scott Elliot, M.A., B.Sc., F.L.S. — J. Linn. Soc. London, Bot. 29: 61—62.
1901. Lettera al Presidente della Societa Botanica Italiana. — Bull. Soc. Bot. Ital. (1901) 119—124 (Dated March 23, 1901).
—— Per il Giardino Botanico. — Giornale ‘La Nazione’, 12 Maggio 1901.
— Trasloco delle collezioni botaniche del Museo di Firenze. Lettera al Presidente della Societa Botanica Italiana. — Bull. Soc. Bot. Ital. (1901) 202—210.
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Dedication
— Sul trasloco delle collezioni botaniche della Malesia dal Museo di Storia Naturale in Via Ro- mana ne’ nuovi locali presso il Giardino de’ Semplici. — Corrispondenza del D. Odoardo Bec- cari coll’Istituto di Studi Superiori. — Firenze, Tipogr. S. Landi, 4 pp. (Dated July 2, 1901).
1902. Nelle Foreste di Borneo. Viaggi e ricerche di un naturalista. — Firenze, xvi+667 pp., 81 fig., 4 maps.
—— Systematic enumeration of the species of Calamus and Daemonorops with descriptions of the new ones. — Rec. Bot. Surv. India 2: 197—230.
1903. L’Istituto di Studi Superiori di Firenze, la chiusura del Museo Botanico e le sue peripezie. Osservazioni e Critiche. — Rocca San Casciano, Licinio Cappelli, 16 pp., 3 fig.
— Per un Istituto Agricolo Botanico Coloniale. — Giornale ‘Fieramosca’, 21 Marzo 1903 (not seen).
— L’avvenire agricolo dell’Eritrea. — ‘La Tribuna’, Roma, | Giugno 1903 (not seen).
1904. Palmae. — In: J. Perkins, Fragmenta Florae Philippinae 1: 45—48.
— Wanderings in the great forests of Borneo: travels and researches of a naturalist in Sarawak (transl. by E.H. Giglioli, rev. and ed. by F.H.H. Guillemard). — Constable, London, xxiv + 424 pp., 61 fig., 3 maps.
1905. Le palme del genere Trachycarpus. — Webbia 1: 41—68, f. 1—16.
— Note anatomiche sul frutto dei Trachycarpus. — Ibid.: 68—72, f. 17—18.
— Notizie sul Nannorhops ritchieana H. Wendl. — Ibid.: 72—73; also Bull. R. Soc. Tosc. Ortic. 30: 325—326.
— Palme nuove papuane. — Ibid.: 281—313, f. 1-8.
— Le palme delle Isole Filippine. — Ibid.: 315—359.
— Palmae. — In: K. Schumann & C. Lauterbach, Nachtrage zur Flora der Deutschen Schutzge- biete in der Stidsee: 60—61.
1906. Palmarum madagascariensium synopsis. — Bot. Jahrb. 38, Beibl. 87: 1—41, f. 1.
1907. Le Palme Americane della tribu delle Corypheae. — Webbia 2: 1—343.
— Notes on Philippine palms. I. — Philip. J. Sc. 2: Bot. 219—240.
1908. Asiatic palms — Lepidocaryeae. Part I: The species of Calamus. — Ann. R. Bot. Gard. Calc. 11 [Fol.]: Letter-press I-VI, 1—518, t. 1—2; Plates t. 1-236 (1—231+25A, 104A, 172A, 188A, 191A).
— Le palme ‘Dum’ od ‘Hyphaene’ piu specialmente quelle dell’ Affrica italiana. — Agric. Co- lon. 2: 137-183, f. 1—2, t. 1-3.
—— Palmae novae antillanae. — In: Fedde’s Repert. Bot. Syst. 6: 94—96.
— The palms of the Batanes and Babuyanes Islands. — Philip. J. Sc. 3: Bot. 339-342.
1909. New or little known Philippine palms. — Elmer, Leafl. Philip. Bot. 2: 639-650.
— Notes on Philippine palms. II. — Philip. J. Sc. 4: Bot. 601—639, t. 30—31.
—— Palmae. — Nova Guinea 8: 203—222, t. 50—S1.
1910. Glaziova treubiana, nouvelle espece de Cocoinée, avec observations sur le genre Cocos. — Ann. Jard. Bot. Buitenzorg ser. 2, suppl. 3: 791—806, 2 fig., t. 31.
— Le palme del genere Raphia. — Agric. Colon. 4: 137—170, f. 1—3, t. 1—6 (reprinted with the original pagination, as an issue of the series ‘Biblioteca Agraria Coloniale’).
—— Studio monografico del genere Raphia. — Webbia 3: 37—130, f. 1—8, t. 1.
—— Palme australasiche nuove 0 poco note. — Ibid.: 131—165, f. 1—6.
—— Descrizione di una nuova specie di Trachycarpus. — Ibid.: 187—190.
— Palmae dell’Indo-China. — Ibid.: 191—245.
— Contributo alla conoscenza delle Lepidocaryeae affricane. — Ibid.: 247—294.
— La Copernicia cerifera in Riviera ed una nuova specie di Livistona. — Ibid.: 295—305, f. 1, thy. (33)
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1911. Asiatic palms — Lepidocaryeae. Part II: The species of Daemonorops. — Ann. R. Gard. Calc. 12(1) [Fol.]: Letter-press [I—II], 1—237, t. I—II; Plates t. 1—109.
— Classification des palmiers d’Indo-Chine. — Bull. Mus. Hist. Nat. Paris 17: 148—160.
— Le palme che producono fibre di Piassava nel Madagascar. — Agr. Colon. 5: 320—324, f. A-D.
— The palms of the island of Polillo. — Philip. J. Sc. 6: Bot. 229—230.
1912. Palmae. — In: H. Winkler, Beitrage zur Kenntnis der Flora und Pflanzengeographie von Borneo. II. — Bot. Jahrb. 48: 89—93.
— Palmae. — In: I. Urban, Symbolae Antillanae 7: 170-172.
1912—13. The palms indigenous to Cuba with appended remarks on the general structure of the trunk of a palm. — Pomona Coll. J. Econ. Bot. 2 (1912) 253—276, f. 109—118; 351—377, f. 144—153; 3 (1913) 391—417, f. 154-172.
1912—14. Palme del Madagascar. — Firenze, Istituto Micrografico Italiano, [Fol.], [i—iv] +62 pp., 50 fig., 50 tav.
1913. Contributi alla conoscenza delle palme. — Webbia 4: 143—240, f. 1—17.
— Manipolo di palme nuove Polinesiane conservate nell’erbario di Kew. — Firenze, M. Ricci, October 1913 (preprinted with the same pagination from Webbia 4: 253—291, f. 1—31, 1914).
— Studio sui ‘Borassus’ e descrizione di un genere nuovo Asiatico di ‘Borasseae’. — Firenze, M. Ricci, November 1913 (preprinted with the same pagination from Webbia 4: 293—385, f. 32-42, 1914).
1914. Asiatic palms — Lepidocaryeae. Supplement to Part I: The species of Calamus. — Ann. R. Bot. Gard. Calc. 11 (App.) [Fol.]: Letter-press i—viii, 1—142; Plates t. 1—83.
— Contributo alla conoscenza delle palme a olio (Elaeis guineensis). — Agric. Colon. 8: 5—37, t. 1—5; 108-118, t. 7-14; 201—212, t. 24-26; 255—270, t. 40—41 (reprinted with separate pagi- nation: 80 pp., 18 tav., as an issue of the series ‘Biblioteca Agraria Coloniale’).
— Neue Palmen Mikronesiens. — In: G. Volkens, Beitrage zur Flora von Mikronesien. — Bot. Jahrb. 52: 4.
— Neue Palmen Papuasiens. — In: C. Lauterbach, Beitrage zur Flora von Papuasien iv: 26B. — Bot. Jahrb. 52: 23—39.
— Palmae. — In: K. Rechinger, Botanische und zoologische Ergebnisse einer wissenschaftli- chen Forschungsreise nach den Samoainseln, dem Neuguinea-Archipel und den Salomonsinseln von Marz bis Dezember, 1905. V. Teil. — Denkschr. Kaiserl. Akad. Wiss. Math.-Naturwiss. Kl. 89 (‘1913’) 502—S09, f. 5—12.
1916. Palmae. — In: J.F. Rock, Palmyra Island with a description of its flora. — Coll. Hawaii Publ. Bull. 4: 44—48, t. 17—18 (reprinted with the title ‘Palmae. Cocos nucifera L. forma pal- myrensis. Characteristics of the Coconuts of the Palmyra Islands’ and different pagination: p. 1—5, t. 18-19).
—— Il genere Cocos e le palme affini. — Agric. Colon. 10: 435—471, t. 1—4; 489-532, t. 5—13; 585—623, t. 14—15 (reprinted with separate pagination: 128 pp., 15 tav., as issue of the series ‘Biblioteca Agraria Coloniale’).
— Palmae. — In: E. Chiovenda, Resultati scientifici della Missione Stefanini-Paoli nella Soma- lia Italiana. Volume I. Le collezioni botaniche. Appendice: Le raccolte di Mangano, Scassellati, Mazzocchi e Provenzale in Somalia. — Firenze, Pubbl. R. Ist. Stud. Super. Mus. ed Erb. Co- lon. p. 176—177, 230.
1917. The origin and dispersal of Cocos nucifera. — Philip. J. Sc. 12: Bot. 27—43 (reprinted in Principes 7: 57—69, 1963).
— A new species of Calamus from Amboina. — Ibid.: 81.
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Dedication
— Ona new South Polynesian palm, with notes on the genus Rhopalostylis Wendl. et Drude. — Trans. & Proc. New Zeal. Inst. Bot. 49: 47—50.
— Palmae. — In: L.S. Gibbs, Dutch N.W. New Guinea. A contribution to the phytogeography and flora of the Arfak Mountains, &c. p. 91—98, 200.
— Pelagodoxa henriana Becc. — In: D. Bois, Pelagodoxa henriana. Palmier nouveau des Isles Marquises. — Rev. Hort. 89: 302—304, f. 76-79.
1918—1921. Asiatic palms — Lepidocaryeae. Part III. The species of the genera Ceratolobus, Ca- lospatha, Plectocomia, Plectocomiopsis, Myrialepis, Zalacca, Pigafetta, Korthalsia, Metroxy- lon, Eugeissona. — Ann. R. Bot. Gard. Calc. 12(2) [Fol.]: Letter-press [I—II], 1-231, t. i—vi; Plates t. 1-120 (1—118+50A, 63A).
1919. Palms of the Philippine Islands, collected and distributed by A.D.E. Elmer. — Elmer, Leafl. Philip. Bot. 8: 2997—3067.
—— The palms of the Philippine Islands. — Philip. J. Sc. 14: 295—362, t. 1—3.
1920. Palmae novae antillanae. II. — In: Fedde, Repert. Spec. Nov. Regni Veg. 16: 436—437.
— Recensione delle palme del vecchio mondo appartenenti alla tribu delle Corypheae con de- scrizione delle specie e varieta nuove che vi appartengono. — Firenze, M. Ricci, 10 December 1920 (preprinted with the same pagination from Webbia 5: 5—70, 1921).
— Le palme della Nuova Caledonia. — Firenze, M. Ricci, 128 pp., 13 tav., 10 December 1920 (preprinted with separate pagination from Webbia 5: 71—197, t. 1—13, 1921).
1921. Nelle foreste di Borneo. Viaggi e ricerche di un naturalista. Ed. 2. — Firenze, xvi+ 469 pp., 72 tav., 11 fig., 4 maps, 1 portr. (published posthumously by Beccari’s son Nello Beccari). 1923. Neue Palmen Papuasiens II. — In: C. Lauterbach, Beitrage zur Flora von Papuasien. X.
— Bot. Jahrb. 58: 441—462.
— Note botaniche e botanico industriali bornensi. — Webbia 5: 451—581, f. 1—6 (reprint of Appendix, Nelle foreste di Borneo, 1902).
1924. Neue Palmen Mikronesiens. — In: L. Diels, Beitrage zur Flora von Mikronesien und Poly- nesien. III. — Bot. Jahrb. 59: 11—16.
— Nuova Guinea, Selebes e Molucche; diarii di viaggio ordinati dal figlio Nello Beccari, con introduzione e note del Prof. Luigi Buscalioni. — Firenze, Soc. Anon. Edit. ‘La Voce’, XXXViii +468 pp., 53 fig., 22 tav., 1 map (Diaries published by Beccari’s son Nello Beccari).
— Palme della trib’ Borasseae. Ed. U. Martelli, G. Passeri, Firenze, 56 pp., 18 fig., 45 tav.
1927. Odoardo Beccari nel Mar Rosso e tra i Bogos. Frammenti di diario inediti, trascritti e ordi- nati dal figlio prof. Nello Beccari. — Boll. R. Soc. Geogr. Ital. ser. 6, 4: 625—646.
1930. Odoardo Beccari in Sumatra e la scoperta dell’ ‘Amorphophallus Titanum’. (Frammenti di diario inediti ed ordinati). — Boll. R. Soc. Geogr. Ital. ser. 6, 7: 569—595, f. 1—6, 1 map. (The author of the paper is given as Nello Beccari).
1933. Asiatic palms — Corypheae. The species of the genera Corypha, Nannorhops, Sabal, Co- pernicia, Serenoa, Brahea, Acoelorhaphe, Washingtonia, Pritchardia, Erythea, Livistona, Li- cuala, Pritchardiopsis, Phlolidocarpus, Teysmannia, Rhaphis, Chamaerops, Trachycarpus, Rhaphidophyllum, Trithrinax, Acanthorhiza, Hemithrinax, Thrinax, Coccothrinax, Cryso- phyla. (Posthumous work by Dr. Odoardo Beccari. Revised and edited by Prof. Ugolino Mar- telli). — Ann. R. Bot. Gard. Calc. 13: Letter-press [Qu.] [i—x], 1—356; Plates [Fol.] t. 1-102 (1—99+ 59 bis, 68 bis, 89 bis), (1931’).
1934. Generi, specie e varieta nuove di palme gerontogee della trib Arecaceae lasciate inedite dal Dr. Od. Beccari ed ordinate a cura di U. Martelli. — Atti Soc. Tosc. Sc. Nat. Pisa. Mem. 44: 114-176. [repr. pp. 1—65].
1935. (By U. Martelli from Beccari manuscripts). I generi e le specie delle palme gerontogee della
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tribu delle Arecaceae. Esposizione geografica secondo la monografia inedita del Dr. O. Beccari. — Nuovo Giorn. Bot. Ital. ser. 2, 41: 693—723 (‘1934’).
—— (By U. Martelli from Beccari manuscripts). La sinonimia delle palme gerontogee della tribu delle Areceae. — Ibid. 42: 17-88.
1955. (& R.E.G. Pichi Sermolli) Subfamiliae Arecoidearum palmae gerontogeae. Tribuum et ge- nerum conspectus. — Webbia 11: 1—187, f. 1—47.
B — Letters by Odoardo Beccari
During his travels BECCARI wrote several letters to his friends in Italy, chiefly to G. Doria, E.H. GIGLIOLI, G. CorA, T. SALVADORI, R. GESTRO, and O. ANTINORI. They contain a wealth of very interesting observations and comments on the botanical, zoological, ethnological and other natu- ralistic aspects of Malesia. These letters or fragments of them were published in various Italian periodicals, usually accompanied by information and comments on BEccarr’s scientific discover- ies, and on the itineraries and the main events of his adventurous travels.
The bibliographic citations of the papers in which these letters are published are given below together with an indication of the name of the friend to whom the letter was addressed, and the date and place in which it was written. In order to facilitate and render more systematic the consul- tation of these letters, they are quoted according to the periodicals in which they were published.
Letters published in the Nuova Antologia
BEcCARI’s travels in Malesia, Assab and the country of Bogos between 1865 and 1876 were de- scribed by Enrico H. GIGLIOLI in various instalments published in the Nuova Antologia with the general title of ‘Odoardo Beccari ed i suoi viaggi’. They were also reprinted, with independent pagination, and assembled in a special book (Firenze, Le Monnier, 309 pp., 9 fig., 2 small maps, 1872—76) with the title ‘I viaggi del Dott. Beccari da Firenze tracciati e commentati’. There Gi- GLIOLI published several letters or fragments of letters by BECCARI to his friends and also some passages from his original diaries. All of them are quoted verbatim between GIGLIOLI’s descrip- tions and comments.
Borneo. 1865—1868. — Nuova Antologia 21 (1872) 119—160 (Passages of the original diaries).
Samhara e Bogor. 1870. — Ibid.: 22 (1873) 658—668.
Malesia, Molucche e Papuasia. 1871—72—73. — Ibid.: 22 (1873) 668—709. (Letters to G. Doria, from the Red Sea, December 10—11, 1871; to E.H. Giglioli, from Batavia, February 8; to G. Doria, from Makassar, February 23; Ceram, March 13; Ambon, March 21; Kapaor, April 21, with a small map; to E.H. Giglioli, from Sorong, May 3; to G. Doria, from Sorong, June 21, 1872).
Papuasia: Sorong-Mansinam-Andai (Monti Arfak). (Giugno 1872 al Gennaio 1873). — Ibid.: 23 (1873) 194—225, 2 fig. (Letters to G. Doria, from Andai, August 30; Andai, September 8—9, 1872; Ambon, January 2; Ambon, February 2—7, 1873).
Papuasia — Le isole Aru e Kei (Febbraio—Settembre 1873). — Ibid.: 24 (1873) 835—866, 1 map; 25 (1874) 163—192, 1 fig. (Letters to C. Correnti, from Ambon, January 3; to O. Antinori, from Ambon, January 3; Ambon, February 5; to G. Doria, from Dobbo (Aru Is.), February 24; Wo- kan (Aru Is.), March 10; Wokan, July 3—4; Tual (Kei Is.), August 27, 1873).
I. Macassar — Kandari (Celebes). II. I Papua (Dicembre 1873, Giugno 1874). — Ibid.: 27 (1874) 420—463, 5 fig. (Letters to E.H. Giglioli, from Makassar, December 4; to R. Gestro, from Ma-
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kassar, December 1; to G. Doria, from Makassar, December 4; Makassar, December 26, 1873; Makassar, January 15; to T. Salvadori, from Makassar, January 14; to R. Gestro, from Makas- sar, January 15; to G. Doria, from Kandary, April 23, and May 2 and 11, 1874).
Celebes, Giava, Ternate, Amboina (Giugno 1874—Gennaio 1875). — Ibid.: ser. 2, 2 (1876) 802—822. (Letters to G. Doria, from Kandari, June 4; Makassar, August 30; to O. Antinori, from Makassar, August 28; to T. Salvadori, from Makassar, August 30; to Baron Podesta, the Mayor of Genoa, from Makassar, end August; to G. Doria, from Ambon, December 7, 1874; to E.H. Giglioli, from Ambon, January 7; to ??, from Ambon, January 15, 1875).
Seconda esplorazione della Nuova Guinea. La baia di Geelvink (Febbraio—Novembre 1875). — Ibid.: ser. 2, 3 (1876) 147-163. (Letters to G. Doria, from Dorei, June 5; Andai, June 15; Hatam, June 21; to T. Salvadori, from Ternate, August 4, 1875).
Terza esplorazione della Nuova Guinea. La baia di Humboldt (Novembre 1875—Marzo 1876). — Ibid.: ser. 2, 3 (1876) 333—363, 1 fig. (Letters to G. Cora, from Ternate, March 11; to E.H. Giglioli, from Ternate, March 6, 1876).
Letters published in the Bollettino della Societa Geografica Italiana
Estratti e frammenti di lettere dirette al marchese Giacomo Doria dal naturalista botanico Odoar- do Beccari, durante il suo viaggio alla Nuova Guinea. — Boll. Soc. Geogr. Ital. 8 (1872) 148—151. (Letters to G. Doria, from Wahaai, March 13; Ambon, March 21; Sorong, May 3, 1872).
Odoardo Beccari nella Nuova Guinea e nelle isole Aru. — Ibid.: 9 (1873) 145—158. (Letters to G. Doria, from Andai, August 30, 1872; to C. Correnti, from Ambon, January 3; to O. Antino- ri, from Ambon, January 4; to G. Doria, from Ambon, February 2, 5, 7; to O. Antinori, from Ambon, February 5; to G. Doria, from Dobbo, Aru Is., February 24; Wokan, Aru Is., March 10, 1873).
Lettera di O. Beccari al marchese Doria. — Ibid. 10: 4—5 (1873) 66—69. (Letter from Wokan, Aru Is., July 3, 1873).
Frammento di lettera di O. Beccari a Doria da Makassar, il 18 Novembre 1873. — Ibid.: 10: 6 (1873) 38.
Odoardo Beccari alle isole Key. — Ibid.: 87-89. (Letter to G. Doria, from Tual (Small Kei), Au- gust 27, 1873).
Ultime notizie di O. Beccari. — Ibid.: 11 (1874) 78-81. (Letters to R. Gestro, from Makassar, December 1; to G. Doria, from Makassar, December 4; to O. Antinori, from Makassar, Decem- ber 7; to G. Doria, from Makassar, December 26, 1873).
Corrispondenze del dott. Odoardo Beccari. — Ibid.: 276—282. (Letters to T. Salvadori, from Ma- kassar, January 14; to R. Gestro, from Makassar, January 15; to G. Doria, from Makassar, January 15, 1874).
Lettere di Odoardo Beccari. — Ibid.: 480—488, carta della parte sud-est di Celebes. (Letters to G. Doria, from Kandari, April 23 and June 4, 1874).
Nota sui Papua e sulla Nuova Guinea. — Ibid.: 652—659. (Letter to O. Antinori, from Makassar, August 28, 1874).
Lettere di Odoardo Beccari. — Ibid.: 660—667. (Letters to the Mayor of Genoa, from Makassar, without dates; to T. Salvadori, from Makassar, August 30; to G. Doria, from Kandari, June 4, 1874).
Lettera di O. Beccari. — Ibid.: 12 (1875) 117—122. (Letter to G. Doria, from Ambon, January 4, 1875).
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La Nuova Guinea Olandese. — Ibid.: ser. 2. 1 (1876) 5SO—557. (Letter to G. Cora, from Ternate, March 11, 1876).
Letters published in Cosmos
The editor of the journal Cosmos of Turin, Gurpo Cora, gave ample information on BECCARIs travels in Malesia. He published several letters by BEccarito his friends in Italy, which are listed below. Furthermore, Cora gave various reports on the different stages of BECCARI’s journeys. The references to the latter are given in Appendix 3 dealing with BEccarrs itineraries.
Recenti spedizioni alla Nuova Guinea. Spedizione di Beccari e D’Albertis:
1. Da Singapore ad Amboina. — Cosmos | (1873—74) 11—15. (Letters to G. Doria, from Ba- tavia, February 8; Makassar, February 23, Wahai(N. Ceram), March 13; Amboina, March 2 1872):
2. Da Amboina a Sorong. — Ibid.: 15—20. (Letters to G. Doria, from Kapaor, April 21; So- rong, May 3 and June 21, 1872).
Recenti spedizioni alla Nuova Guinea. Esplorazioni di Odoardo Beccari:
1. Ricerche geografiche nella Nuova Guinea. — Cosmos 2 (1874—75) 7—9. (Letter to G. Cora, from Makassar, December 1, 1873).
2. Note sulle Isole Kei. — Ibid.: 9-10. (Letter to G. Cora, from Makassar, December 1, 1873).
3. Da Makassar a Kandari. — Ibid.: 92—96. (Letters to G. Doria, from Kandari, May 2 and 11, 1874).
4. Escursioni intorno a Kandari. Ritorno a Makassar. — Ibid.: 203—207. (Letters to G. Doria, from Kandari, June 4; Makassar, August 30, 1874).
5. Notizie sull’ornitologia di Celebes. — Ibid.: 207—208. (Letter to T. Salvadori, from Makas- sar, August 30, 1874).
6. Appunti etnografici sui Papua. — Ibid.: 400—404. (Letter to O. Antinori, from Makassar, August 28, 1874).
7. Soggiorno a Ternate. Da Ternate ad Amboina. Preparativi pel terzo viaggio alla Nuova Guinea. — Cosmos 3 (1875—76) 83—88. (Letters to G. Doria, from Amboina, January 4, 8, 9, 1875).
8. Da Amboina a Dorei, per Soron e Wakkaré. Scoperta del fiume Wa Samson. Esplorazione della baia di Geelvink, determinazione della sua vera ampiezza. — Ibid.: 88—92. (Letter to G. Doria, from Dorei, June 5, 1875).
9. Esplorazione dei Monti Arfak. Ritorno a Ternate per la via di Salvatti, Batanta, Koffiao. — Ibid.: 92—95. (Letters to G. Doria, from Andai, June 15; Hatam, June 21; to T. Salva- dori, from Ternate, August 4, 1875).
10. Viaggio a bordo del trasporto olandese ‘Soerabaja’. Itinerario progettato. Da Ternate a Dorei, per Salvatti. Visita ad Ansus. Lavori idrografici. — Ibid.: 220—221. (Letters to G. Doria, from Ternate, November 7; Dorei, November 26, 28, 1875).
11. Viaggio a bordo del trasporto olandese ‘Soerabaja’. Da Dorei alla baia di Humboldt per la baia Vandamen, l’isola Run, il sud di Jobi, le foci dell’ Ambermo. — Ibid.: 349-352. (Letter to G. Cora, from Ternate, March 11, 1876).
12. Saggio statistico sulla Nuova Guinea Olandese. Popolazione, Commercio, Climatologia, Nomenclatura. — Ibid.: 352—360. (Letter to G. Cora, from Ternate, March 11, 1876).
13. Viaggio a bordo del trasporto olandese ‘Soerabaja’. La Baia di Humboldt edi suoi abitanti.
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La Baia del Disinganno, il Vulcano Ciclope e le isole Arimoa. — Ibid.: 364—372. (Letter to E.H. Giglioli, from Ternate, March 6, 1876).
14. Viaggio a bordo del trasporto olandese ‘Soerabaja’. Dalla Baia di Humboldt a Ternate pel nord di Jobi, Dorei, Waigheu, Misol, il golfo Mac Cluer, la baia Gouns, Ghesser, Amboi- na. — Ibid.: 372—374. (Letter to E.H. Giglioli, from Ternate, March 6, 1876).
15. Questioni etnologiche sui Papua. — Ibid.: 375—379. (Letter to E.H. Giglioli, from Ternate, March 6, 1876).
Letters published in various journals
Il viaggio di O. Beccari alla Nuova Guinea. — Nuov. Giorn. Bot. Ital. 4 (1872) 208—212. (Letters to a friend of Florence, from Wahai, N. Ceram, March 13; Ambon, March 21, 1872).
Il viaggio di O. Beccari alla Nuova Guinea. — Ibid.: 291—294. (Letters to G. Doria, from Kapaor, April 21; Sorong, June 21, 1872).
Lettera di O. Beccari dalle isole Aru. — Ibid. 5 (1873) 330. (Letter to [G. Doria], July 3—4, 1873).
Brano di lettera di O. Beccari da Makassar in data 1° Dicembre 1873. — Ibid. 6 (1874) 205—206.
Una pianta maravigliosa. — Bull. R. Soc. Tosc. Ortic. 3 (1878) 270—271. (Letter to R. Corsi Sal- viati, from Sumatra, without date. Comment by E.O. Fenzi).
Lettera ornitologica di O. Beccari intorno agli uccelli osservati durante un recente viaggio alla Nuova Guinea. — Ann. Mus. Civ. St. Nat. Genova 7 (1875) 704—720. (Letter to T. Salvadori, from Ternate, August 4, 1875. Introduction by T. Salvadori).
Lettera del Prof. Odoardo Beccari a Giacomo Doria. — Ibid. 13 (1878) 451—455. (From Kajt Ta- nam, Sumatra, September 8, 1878).
C — Maps
In the period of preparation for his travels BEccarRI carefully trained himself also in geodetics and topography. He also invented a new instrument (Nuovo orizzonte artificiale) for topographic survey.
During his explorations BEccarI devoted great attention to the topography of the places he vis- ited and his surveys allowed him to draw some maps which greatly contributed to the delimitation of the coasts of certain areas of the Malesian Archipelago.
Some of these maps were published by BEccartr himself, others were utilized by G. Cora, to- gether with surveys of other explorers, to elaborate some of the maps published in his periodical Cosmos.
BeEccarr’s itineraries are traced on all maps listed below.
Carta originale del viaggio di Beccari e d’Albertis nella Nuova Guinea ovest (Aprile 1872), costrut- ta e disegnata da Guido Cora. Scala 1:700.000. — Cosmos (G. Cora) 1 (1873-74) Tav. I. — Explanatory notes in Cora G. Recenti spedizioni alla Nuova Guinea. Memoria sulla Tavola I. — Ibid.: 22—24.
Carta originale del viaggio di O. Beccari nel Sud-est di Celebes (Maggio— Agosto 1874), costrutta e disegnata da Guido Cora. Scala 1:1.200.000. — Ibid. 2 (1874-75) Tav. V. — Explanatory notes in Cora G. Viaggio di O. Beccari nel Sud-est di Celebes. Note sulla Tavola V. — Ibid.: 201-202.
Carta originale della Nuova Guinea N.O. e delle isole Salvatti, Batanta, William, ecc. secondo i rilievi originali di Lovera, Cerruti, Beccari e le esplorazioni anteriori costrutta e disegnata da
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Guido Cora. Scala 1:1.000.000. — Cosmos (G. Cora) 3 (1875) Tav. II. — Explanatory notes in Cora G. Recenti spedizioni alla Nuova Guinea. Note sulle Tavole II e III. — Ibid.: 81—83.
Carta originale della Baia di Geelvink e del littorale N.O. della Nuova Guinea secondo i rilievi di Odoardo Beccari, 1875 del Geelvink, di Duperrey, Dumont D’Urville, ecc. costrutta e dise- gnata da Guido Cora. Scala 1:1.800.000. — Including an inset map: Tracciato comparativo della Baia di Geelvink secondo la carta dell’ Amm. Ingl. N° 2759a ed i rilievi di O. Beccari. Scala 1:4.800.000. — Ibid.: Tav. III. — Explanatory notes in Cora G. Recenti spedizioni alla Nuova Guinea. Note sulle Tavole II e III. — Ibid.: 81—83.
Carta originale della Nuova Guinea Nord dai Monti Arfak alla Baia d’ Humboldt secondo i rilievi di Odoardo Beccari, 1875—76, della nave oland. ‘Soerabaja’, cap. Swaan e le esplorazioni ante- riori, costrutta e disegnata da Guido Cora. Scala 1:2.000.000. — Including an inset map: I Mon- ti Arfak. Scala 1:800.000. — Ibid. 3 (1876) Tav. X. — Explanatory notes in Cora G. Recenti spedizioni alla Nuova Guinea. Note sulla Tavola X. — Ibid.: 347-349.
Piano della Baia d’ Humboldt (Telokh Lintciu) secondo i rilievi della nave oland.e ‘Etna’ 1858 e le ricerche di O. Beccari, 1875, di Guido Cora. Scala 1:70.000. — Ibid.: Tav. XI.
Carta originale della Provincia di Sarawak, compilata sopra vari documenti e secondo le osserva- zioni dell’autore. — In O. Beccari, Nelle foreste di Borneo (1902) fig. 36 (p. 187).
Abbozzo di carta (originale) del fiume Bintulu e suoi affluenti. — Ibid.: fig. 58 (p. 351).
Carta del sistema idrografico e delle attuali divisioni politiche di Borneo, compilata sopra i docu- menti pili recenti e le osservazioni dell’autore. — Ibid.: fig. 66 (p. 407).
Carta orginale degli itinerari dell’autore in Sarawak. — Ibid.: fig. 75 (facing page 504).
Carta speciale della Nuova Guinea Ovest cogli itinerari di O. Beccari e L.M. D’Albertis (1872—1876) costrutta e disegnata da Guido Cora. — Including an inset map: Carta originale del viaggio di O. Beccari nel Sud Est di Celebes. 1874. Disegnata da G. Cora. — In O. Beccari, Nuova Guinea, Selebes e Molucche (1924) at the end of the book.
Carta delle Isole Aru secondo i rilievi di Odoardo Beccari. Luglio 1873. — In O. Beccari, Lettera di O. Beccari al Marchese Doria. — Boll. Soc. Geogr. Ital. 10: 4—5 (1873) 64—65, at the end of the paper.
Carta della Parte Sud-est delle Celebes secondo la relazione di O. Beccari e traccia del suo viaggio da Makassar a Kandari, Febbraio 1874. — Ibid.: 11 (1874) 480—488, at the end of the paper.
Appendix 2 — Biographies of Odoardo Beccari
BALDASSERONI, V. & D. CaRAzzi. L’opera biologica di Odoardo Beccari. Rassegna Sci. Biol. 3 (1921) 84-88.
BARGAGLI PETRUCCI, G. L’ opera biologica di Odoardo Beccari. Pubbl. Ist. Stud. Sup. Firenze, Sez. Sci. Fis. Nat. In memoria di Odoardo Beccari, pp. 5—16. 1921.
BeccarI, N. Enciclopedia Italiana di Scienze, Lettere ed Arti. Milano, Ist. G. Treccani, 6 (1930) 462.
—— Brief Obituary, Itineraries and Bibliography — Manuscript compiled by Beccari’s son Nello for Flora Malesiana, Oct. 1947 (in Library Rijksherbarium, Leiden).
BeEGuINoT, A. Boll. R. Soc. Geogr. Ital. ser. 5, 12 (1923) 194—209, portr.
— Rivista Biol. 3 (1921) 118—121, portr.
BERTACCHI, C. Geografi ed esploratori italiani contemporanei. Milano, De Agostini Ed. 1929, pp. 368—372, portr.
BurkiLL, I.H. & J.C. MouLton. Journ. Str. Br. R. Asiat. Soc. 83 (1921) 166-173.
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Dedication
BUSCALIONI, L. La vita e l’opera di Odoardo Beccari. In: O. Beccari, Nuova Guinea, Selebes e Molucche. Firenze, 1924, pp. xxiii—Xxxviii.
CavaRA, F. Bull. Soc. Afr. Ital. 40 (1921) 46—S51.
CHIOVENDA, E. Notizie biografiche. Odoardo Beccari. Annuario R. Ist. Stud. Sup. Firenze 1920-21 (1921) 155—156.
— Nuovo Giorn. Bot. Ital. ser. 2, 28 (1921) 5—35, portr., bibl. (in historical sequence).
Cora, G. Recenti spedizioni alla Nuova Guinea. I. Spedizione di Beccari e d’Albertis. (Studi e viaggi di Beccari dalla sua adolescenza al 1872). Cosmos (G. Cora) 1 (1873-74) 8-11.
Dori, G. I naturalisti italiani alla Nuova Guinea e specialmente delle loro scoperte zoologiche. Boll. Soc. Geogr. Ital. ser. 2, 3 (1878) 154-169.
Gestro, R. Ann. Mus. Civ. St. Nat. Genova, ser. 3, 9 (1921) 242—297, 11 fig. (10 portr.).
GUBERNATIS, A. DE. Dizionario biografico degli scrittori contemporanei. Firenze, 1879, 1: 117-118, portr.
— Dictionnaire international des écrivains du jour. Florence, 1888, 1: 220—221.
— Dictionnaire international des écrivains du monde latin. Suppl. et Ind. Florence, 1906, 30*.
Jackson, B.D. (‘B.D.J.’). Proc. Linn. Soc. London 134 (1922) 37—40.
LEONE E. DE & M. A.ippI CAPPELLETTI. Dizionario biografico degli Italiani. Roma, 1965, 7: 440-444.
Lo Priore, G. Ber. Deut. Bot. Ges. 39 (1922). Gen. Vers. Heft (56)—(87), portr. bibl. (alphabeti- cal according to titles).
MarRTELLI, U. Webbia 5 (1921) 295-353, portr., 3 maps, bibl. (alphabetical according to titles).
Moore Jr., H.E. Odoardo Beccari (1843—1920). Principes 25 (1981) 29—35, portr., bibl. (on palms).
NissEN, CL. Die botanische Buchillustration. 2 (1951) 11 (on his drawings).
PALADINI, C. La morte di Odoardo Beccari. ‘Il Nuovo Giornale’ 27 Ottobre 1920.
PAMPANINI, R. Agricoltura Coloniale 14 (1920) 449—453.
Po.iaccl, G. Atti Ist. Bot. Univ. Pavia. ser. 6, 6 (1935), i—xiii, bibl. (in historical sequence).
Puccion1, N. L’opera etnografica ed antropologica di Odoardo Beccari. Pubbl. Ist. Stud. Sup. Firenze, Sez. Sci. Fis. Nat. In memoria di Odoardo Beccari. pp. 17—26, 1921.
Ropotico, F. Naturalisti esploratori dell’Ottocento Italiano. Firenze, Le Monnier, 1967, pp. 191—222, t. 4 (with reprint of original passages).
Roster, G. Bull. R. Soc. Tosc. Ortic. 46 (1921) 33—36, portr.
Saccarbo, P.A. La Botanica in Italia. Materiali per la storia di questa scienza. Mem. Ist. Veneto Sci. Lett. Arti. 25:4 (1895) 25; 26:6 (1901) 16.
S.A.S. Kew Bull. (1920) 369—370.
STEENIS, C.G.G.J. vAN. Thesaurus Beccarianus. Webbia 8 (1952) 427—436.
STEENIS-KRUSEMAN, M.J. vAN. Flora Malesiana I, 1 (1950) 43—46, portr.
—— Flora Malesiana I, 8 (1974) xiii (brief addition).
Wittrock, V.B. Catalogus illustratus Iconothecae Botanicae Horti Bergiani Stockholmiensis. Acta Horti Berg. 3:2 (1903) 160, t. 31 (portr.); 3:3 (1905) 175.
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Appendix 3 — Accounts of Odoardo Beccari’s itineraries in Malesia and Ethiopia (see also Letters and Maps)
Beccarl, N. Brief Obituary, Itineraries and Bibliography. — Manuscript compiled by Beccari’s son Nello for Flora Malesiana, Oct. 1947. (in Library Rijksherbarium, Leiden).
Burtt, B.L. Beccari’s ascent of ‘Mount Poi’. Flora Males. Bull. 19 (1964) 1131—1132. (Pointing out that Mt Poi (Poe, Pueh) in Southwest Sarawak, mentioned in the ‘Wanderings’, a famous type locality, is not what is nowadays on maps indicated as Mt Poi, but a more southeasterly peak in the Poi Range, now known as Gunung Berumput, or Gunung Rumput).
Cora, G. Spedizione italiana alla Nuova Guinea. Roma, Stab. Civelli, 1872, 39 pp.
— Recenti spedizioni alla Nuova Guinea. — Cosmos (G. Cora) 1 (1873-74) 7-8.
— Recenti spedizioni alla Nuova Guinea. Spedizione di Beccari e D’Albertis. — Ibid.: 8-11, 141-143.
— Recenti spedizioni alla Nuova Guinea. Odoardo Beccari. — Ibid.: 215-218, 265; 2 (1874-75) 2—4, 86; 3 (1875-76) 75—76.
— Viaggio di O. Beccari nel Sud-est di Celebes. — Ibid.: 2 (1874—75) 200—202, t. V (map).
— Recenti spedizioni alla Nuova Guinea. Secondo viaggio della ‘Vettor Pisani’. — Ibid.: 3 (1875—76) 77-78.
— Recenti spedizioni alla Nuova Guinea. Quarto viaggio di O. Beccari alla Nuova Guinea (1875—1876) — Ibid.: 217.
— Recenti spedizioni alla Nuova Guinea. I Monti Arfak. — Ibid.: 217-218.
— Carta speciale della Nuova Guinea Ovest cogl’itinerari di O. Beccari e L.M. D’Albertis (1872—1876) costrutta e disegnata da Guido Cora. — Ann. Mus. Civ. St. Nat. Genova 12 (1878).
— (see also Appendix 1 — Letters by O. Beccari). Recenti spedizioni alla Nuova Guinea. — Cosmos 1 (1873—74) 11—15, 15—20; 2 (1874-75) 7-9, 9-10, 92—96, 203—207, 207—208, 400—404; 3 (1875-76) 83-88, 88-92, 92-95, 220-221, 349-352, 352-360, 364-372, 372—374, 375-379.
Gestro, R. Ricordo biografico di Giacomo Doria. Ann. Mus. Civ. St. Nat. Genova, ser. 3, 10 (1921) 1—78, portr. (information on Beccari’s travels).
GiGuio1!, E.H. (see also Appendix 1 — Letters by O. Beccari). Odoardo Beccari ed i suoi viaggi. — Nuova Antologia 21 (1872) 119-160; 22 (1873) 658—668; 668—709; 23 (1873) 194—225; 24 (1873) 835—866; 25 (1874) 163-192; 27 (1874) 420—463; ser. 2, 2 (1876) 802—822; 3 (1876) 147-163, 333-363.
Maaistris, L.F. DE. Biografie di Geografi e di Esploratori contemporanei. IV. Giacomo Doria. Novara, Ist. Geogr. De Agostini, 1917, 18 pp., portr. (information on Beccari’s travels).
MaRTELLI, U. Odoardo Beccari. Webbia 5 (1921) 295—353, 3 maps.
STEENIS-KRUSEMAN, M.J. vAN. Flora Malesiana I, 1 (1950) 43—45.
Vink, W. Nova Guinea, Bot. n. 22 (1965) 479—481, f. 6. (on itinerary in New Guinea).
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Dedication
Appendix 4 — Studies based on Odoardo Beccari’s botanical collections (incomplete)
ARCANGELI, G. L’Amorphophallus titanum Beccari, illustrato da G. Arcangeli. — Nuovo Giorn. Bot. Ital. 11 (1879) 217—223, cum fig.
ASCHERSON, P. Plantae phanerogamae marinae, quas Cl. Eduardus Beccari in Archipelago Indico annis 1866 et 1867, et in Mari Rubro anno 1870 collegit, enumeratae. — Nuovo Giorn. Bot. Ital. 3 (1871) 299-302.
BAGLIETTO, F. Lichenes in regione Bogos Abissiniae septentrionalis lecti ab O. Beccari; illustravit F. Baglietto. — Nuovo Giorn. Bot. Ital. 7 (1875) 239-254.
BaILLon, H. Stirpes exoticae novae. — Adansonia 11 (1873—76) and 12 (1876—79). — Referring to several Bornean plants collected by Beccari.
BAKER, J.G. On a collection of ferns made by Dr. Beccari in western Sumatra. — Journ. Bot. 18 (1880) 209-217.
BARGAGLI-PETRUCCI, G. Sulla struttura dei legnami raccolti in Borneo dal dott. O. Beccari. — Malpighia 17 (1903) 280—371, t. 4—15.
CESATI, V. Felci e specie nei gruppi affini raccolte a Borneo dal Signor Odoardo Beccari. — Atti Accad. Sci. Fis. Mat. Napoli 7:8 (1876) 1—37, t. 1-4.
—— Prospetto delle Felci raccolte dal Sign. O. Beccari nella Polinesia, durante il suo secondo viaggio d’esplorazione in quei mari. — Rend. Accad. Sci. Fis. Mat. Napoli 16 (1877) 23—31.
— Mycetum in itinere Borneensi lectorum a cl. Od. Beccari enumeratio. — Atti Accad. Sci. Fis. Mat. Napoli 8:3 (1879) 1—28, t. 1—4.
CHRISTENSEN, C. Revision of the Bornean and New Guinean ferns collected by O. Beccari and described by V. Cesati & J.G. Baker. — Dansk Bot. Arkiv 9 (1937) 33-52.
DuBarpD, M. Description de quelques types nouveaux ou peu connus de Sapotacées (Illipées) d’aprés les documents de L. Pierre. — Bull. Mus. Hist. Nat. Paris 14 (1908) 405—409.
ENGLER, A. Araceae, specialmente Bornensi e Papuane raccolte da O. Beccari. — Bull. R. Soc. Tosc. Ortic. 4 (1879) 265—271; 295—302. (with an introduction, pp. 265—266, by O. Beccari).
—— Araceae della Malesia e della Papuasia raccolte da Beccari, determinate ed illustrate. — Malesia 1 (1883) 259—304, t. 16—28.
GEHEEB, A. Weitere Beitrage zur Moosflora von Neu Guinea. I. Uber die Laubmoose, welche Dr. O. Beccari in den Jahren 1872—73 und 1875 auf Neu Guinea, besonders dem Arfak-Gebirge, sammelte. II. Uber einige Moose von westlichen Borneo. — Bibl. Bot. Heft 44 (1898) 29 pp., Qetave
— Musci frondosi in monte Pangerango insulae Javae a Dr. O. Beccari annis 1872 et 1874 lecti. — Rev. Bryol. 21 (1894) 81-85.
Hampg, E. Musci frondosi in insulis Ceylon et Borneo a Dr. Od. Beccari lecti. — Nuovo Giorn. Bot. Ital. 4 (1872) 273-291.
Hem, F. Diptérocarpacées de Borneo. — Bull. Mens. Soc. Linn. Paris 120 (1891) 954—958; 122 (1891) 970—976.
— Deux Richetia nouveaux. — Bull. Mens. Soc. Linn. Paris 123 (1891) 979-981.
IRMSCHER, E. Neue Begoniaceen von O. Beccari in Malesien gesammelt. — Webbia 9 (1954) 469—509, f. 1-8.
KRANZLIN, F. On orchids collected by Beccari, in Pflanzenreich; corrections on it by R. Schlechter in Fedde, Repert. 9 (1911) 286—287.
KREMPELHUBER, A. VON. Lichenes foliicoli quos legit O. Beccari annis 1866—1867 in insula Bor- neo. — Miinchen, 1874.
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FLORA MALESIANA
—— Lichenes quos legit O. Beccari in insulis Borneo et Singapore annis 1866 et 1867. — Nuovo Giorn. Bot. Ital. 7 (1875) 5—67, t. 1-2.
MaRTELLI, U. Le Composte raccolte dal Dottore O. Beccari nell’ Arcipelago Malese e nella Papua- sia. — Nuovo Giorn. Bot. Ital. 15 (1883) 281—305.
— Le Dilleniaceae Malesi e Papuane delle collezioni Beccari. — Malesia 3 (1886) 150—167.
— Florula Bogosensis. Enumerazione delle piante dei Bogos raccolte dal Dott. O. Beccari nell’anno 1870, con descrizione delle specie nuovo 0 poco note. — Firenze, 1886. vii+ 169 pp., 1 tav.
— Contribuzione alla flora di Massaua. — Nuovo Giorn. Bot. Ital. 20 (1888) 359—371.
MERRILL, E.D. A brief survey of the present status of Bornean botany. — Webbia 7 (1950) 309-324.
Notaris, C. DE. Epatiche di Borneo raccolte dal Dre O. Beccari nel Ragiato di Sarawak durante gli anni 1865—66—67. — Torino, Stamperia Reale Paravia, 1874, 44 pp., 35 tav. (preprinted from Mem. Acad. Sci. Torino 18 (1876) 267—308, t. 1—35; reprinted also in Nuovo Giorn. Bot. Ital. 8 (1876) 217-251).
PASSERINI, G. Funghi raccolti in Abissinia dal Signor O. Beccari. — Nuovo Giorn. Bot. Ital. 7 (1875) 180—192, t. 4—S.
PicHi SERMOLLI, R. Rapporti tra parassita ed ospite nella Rafflesia tuan-mudae Becc. e Cissus sp. — Nuovo Giorn. Bot. Ital. ser. 2, 44 (1937) 385—421, t. 8—10, f. 1-16.
REICHENBACH, H.G. Odoardo Beccari novitiae orchidaceae papuanae describuntur. — Bot. Centr. Bl. 28 (1886) 343—346.
SCHLECHTER, R. Orchidaceae novae Beccarianae. — Notizbl. Berl. Dahl. 8 (1921) 14—20.
SCHUMANN, K. Sterculiaceae Beccarianae. — Bot. Jahrb. 24 (1897), Beibl. 58: 14—21.
Soitms-LAUBACH, H. Uber die von Beccari auf seiner Reise nach Celebes und Neu Guinea gesam- melten Pandanaceae. — Ann. Jard. Bot. Buitenzorg 3 (1883) 89—104, t. 16.
STEENIS, C.G.G.J. vAN. Thesaurus Beccarianus (including ‘Malaysian sea-grasses collected by Beccari’ and ‘Some records of Malaysian plants’). Webbia 8 (1952) 427—436.
— & M.J. vAN STEENIS-KRUSEMAN. Numberlists of Beccari: Sumatra, Borneo, New Guinea. 1951—1953. Typed copies in Leiden, Bogor, Florence and the Arnold Arboretum, the Leiden copy kept up to date by the first author. Piante Sumatrane, PS 1—979, Piante Borneensi, PB 1—4069, Piante Papuane, PP 1—986. These are his collecting numbers; they do not correspond with the herbarium ‘sheet numbers’ as given in the ‘Herbarium Beccarianum’ at Florence.
STEENIS-KRUSEMAN, M.J. VAN. Flora Malesiana I, 5 (1958) cclvii. — Explaining that Beccari later renumbered his collections originally named ‘Plantae Beccarianae’ in three series, each starting with number 1, viz the PS series (Piante Sumatrane), PB (Piante Borneensi) and PP (Piante Pa- puane).
VENTURI, G. DE. Muschi raccolti dal Signor Odoardo Beccari nella terra dei Bogos in Abissinia. — Nuovo Giorn. Bot. Ital. 4 (1872) 7—22.
ZANARDINI, J. Phycearum indicarum pugillus, a cl. Eduardo Beccari ad Borneum, Sincapore et Ceylanum annis MDCCCLXV—VI-—VII collectarum. — Mem. R. Ist. Veneto Sci. Lett. Arti 17 (1872) 109—170, t. 1-12.
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ABBREVIATIONS AND SIGNS
acc. = according
Ak. Bis. = Aklan Bisaya (Philip. language) Alf. Cel. = Alfurese Celebes (language) alt. = altitude
Anat. = Anatomy
Ap. = Apayao (Philip. language)
app. =appendix, appendices
appr. = approximate
Apr. = April Arch. = Archipelago atl. = atlas
auct. div. =auctores diversi; various authors
auct(t). mal. = auctores malayenses; authors dealing with Malesian flora
auct(t). plur. = auctores plures; several authors
Aug. = August
Bag. = Bagobo (Philip. language)
basionym = original name of the type specimen; its epithet remains permanently attached to the taxon which is typified by it provided it is of the same rank.
Bg. = Buginese (language)
Bik. = Bikol (Philip. language)
Bil. = Bila-an (Philip. language)
Bill. = Billiton
Bis. = Bisaya (Philip. language)
Bon. = Bontok (Philip. language)
Born. = Borneo
Bt= Bukit; mountain
Bug. = Buginese (language)
Buk. = Bukidnon (Philip. language)
c. =circiter; about
C. Bis. = Cebu Bisaya (Philip. language)
cf. =confer; compare
Chab. = Chabecano (Philip. language)
citations = see references
cm = centimetre
c.n. =see comb. nov.
comb. nov. =combinatio nova; new combination
CS = cross-section or transversal section of an organ
c.s. =cum suis; with collaboration
cum fig. =including the figure
cur. = curante; edited by
D (after a vernacular name) = Dutch
Daj. = Dyak (language)
d.b.h. =diameter at breast height
D.E.I. = Dutch East Indies
descr. added behind a reference=means that this contains a valid description
diam. = diameter
Distr. (as an item) = Distribution
Distr. (with a geographical name) = District
ditto =the same, see do
Div. = Division, or Divide
div. = diversus (masc.); various
do = ditto (Ital.); the same
Dum. = Dumagat (Philip. language)
dupl. = duplicate
E=east (after degrees: eastern longitude)
E (after a vernacular name) = English
Ecol. = Ecology
ed. = edited; edition; editor
e.g. =exempli gratia; for example
elab. = elaboravit; revised
em(end). = emendavit; emended
em(erg). ed. =emergency edition
Engl. = English
etc., &c. =et cetera; and (the) other things
ex auctt. =ex auctores; according to authors
excl. = exclusus (masc.); excluding, exclusive of
ex descr. =known to the author only from the de- scription
f. (before a plant name) = forma; form
f. (after a personal name) = filius; the son
f. (in citations) = figure
fam. = family
Feb(r). = February
fide =according to
fig. = figure
fl. =flore, floret (floruit); (with) flower, flowering
For. Serv. = Forest Service
fr. =fructu, fructescit; (with) fruit, fruiting
Fr. (after a vernacular name) = French
G.= Gunung (Malay); mountain
Gad. = Gaddang (Philip. language)
gen. = genus; genus
genus delendum = genus to be rejected
Germ. = German
geront. =Old World
haud = not, not at all
holotype=the specimen on which the original de- scription was actually based or so designated by the original author
homonym =a name which duplicates the name of an earlier described taxon (of the same rank) but which is based on a different type species or type specimen; all later homonyms are nomenclaturally illegitimate, unless conserved
I. = Island
ib(id). =ibidem; the same, in the same place
Ibn. = Ibanag (Philip. language)
ic. =icon, icones; plate, plates
ic. inedit.=icon ineditum, icones inedita; inedited plate(s)
id. =idem; the same
i.e. =id est; that is
If. =Ifugao (Philip. language)
Ig. =Igorot (Philip. language)
Ilg. =Ilongot (Philip. language)
Ilk. = Iloko (Philip. language)
in adnot. =in adnotatione; in note, in annotation
incl. = inclusus (masc.); including, inclusive(ly)
indet. = indetermined
Indr. = Indragiri (in Central Sumatra)
inedit. =ineditus (masc.); inedited
in herb. =in herbario; in the herbarium
in litt. =in litteris; communicated by letter
in sched. =in schedula; on a herbarium sheet
in sicc. =in sicco; in a dried state
in syn. =in synonymis; in synonymy
Is. = Islands
Is. (after a vernacular name) =Isinai (Philip. lan- guage)
Ism. = Isamal (Philip. language)
isotype=a duplicate of the holotype; in arboreous plants isotypes have often been collected from a single tree, shrub, or liana from which the holotype was also derived
Iv. =Ivatan (Philip. language)
J(av). = Javanese (language)
Jan. = January
Jr = Junior
Klg. = Kalinga (Philip. language)
Kul. =Kulaman (Philip. language)
Kuy. = Kuyonon (Philip. language)
Lamp. = Lampong Districts (in S. Sumatra)
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FLORA MALESIANA
Lan. = Lanao (Philip. language)
lang. = language
l.c. =loco citato; compare reference
lectotype=the specimen selected a posteriori from the authentic elements on which the taxon was based when no holotype was designated or when the holotype is lost
livr. =livraison, part
IKeex= ex (plurs)
LS = longitudinal or lengthwise section of an organ
m= metre
M = Malay (language)
Mag. = Magindando (Philip. language)
Mak. = Makassar, Macassar (in SW. Celebes)
Mal. = Malay(an)
Mal. Pen. = Malay Peninsula
Mand. = Mandaya (Philip. language)
Mang. = Mangyan (Philip. language)
Mar. = March
Mbo = Manobo (Philip. language)
Md. = Madurese (language)
Minangk. = Minangkabau (a Sumatran language)
min. part. =pro minore parte; for the smaller part
mm = milimetre
Mng. = Mangguangan (Philip. language)
Morph. = Morphology
ms(c), MS(S) = manuscript(s)
Mt(s) = Mount(ains)
n. =numero; number
N=North (after degrees: northern latitude); or New (e.g. in N. Guinea)
NE. = northeast
nec=not
neerl. = Netherlands, Netherlands edition
Neg. = Negrito (Philip. language)
N.E.I. = Netherlands East Indies
neotype=the specimen designated to serve as no- menclatural type when no authentic specimens have existed or when they have been lost; a neotype retains its status as the new type as long as no auth- entic elements are recovered and as long as it can be shown to be satisfactory in accordance with the original description or figure of the taxon
N.G. = New Guinea
N.I. = Netherlands Indies
no =numero; number
nom. = nomen; name (only)=nomen nudum
nom. al.=nomen aliorum; name used by other authors
nom. alt(ern).=nomen_ alternativum; name
nom. cons(erv).=nomen conservandum, nomina conservanda; generic name(s) conserved by the In- ternational Rules of Botanical Nomenclature
nom. fam. cons.=nomen familiarum conservan- dum; conserved family name
nom. gen. cons. =see nomen conservandum
nom. gen. cons. prop.=nomen genericum conser- vandum propositum; generic name proposed for conservation
nom. _ illeg(it). =nomen name
nom. leg(it). =nomen legitimum; legitimate name
nom. nov. =nomen novum; new name
nom, nud. = nomen nudum; name published without description and without reference to previous pub- lications
alternative
illegitimum; illegitimate
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nom. rej(ic.) = nomen rejiciendum; name rejected by the International Rules of Botanical Nomenclature
nom. seminudum=a name which is provided with some unessential notes or details which cannot be considered to represent a sufficient description which is, according to the International Rules of Botanical Nomenclature, compulsory for valid publication of the name of a taxon
nom. subnudum = nomen seminudum
nom. superfl. =a name superfluous when it was pub- lished; in most cases it is a name based on the same type as an other earlier specific name
non followed by author’s name and year, not placed in parentheses, and put at the end of a citation = means that this author has published the same name mentioned in the citation independently. These names (combinations) are therefore homo- nyms.
Compare 56b line 5—4 from bottom. The same can happen with generic names.
(non followed by abbreviation of author’s name) be- fore a reference (citation) headed by an other author’s name = means that the second author has misinterpreted the taxon of the first author. Compare p. 419a under species 47 the synonym H. celebica. DIELS misapplied the name H. celebica as earlier described by BURCK.
non al. =non aliorum; not of other authors
non vidi=not seen by the author
nov. =nova (femin.); new (species, variety, efc.)
Nov. = November
n.s. =new series
n. sp. =nova species; new species
n. (sp.) prov.=nomen (specificum) provisorium; provisional new (specific) name
n.v. =non vidi; not seen
NW. =northwest
Oct. = October
op.cit. = opere citato; in the work cited
Pp. =pagina; page
P.=Pulau, Pulu (in Malay); Island
Pal(emb.) = Palembang
Pamp. = Pampangan (Philip. language)
Pang. = Pangasinan (Philip. language)
paratype = a specimen cited with the original descrip- tion other than the holotype
part. alt. =for the other part
P. Bis. = Panay Bisaya (Philip. language)
P.I. = Philippine Islands
pl. =plate
plurim., = plurimus; most
D.p. = pro parte; partly
pr. max. p. = pro maxima parte; for the greater part
pro=as far as is concerned
prob. = probabiliter; probably
prop. = propositus; proposed
Prov. = Province
pr.p. = pro parte; partly
pt = part
quae est =which is
quoad basionym, syn., specimina, efc. =as far as the basionym, synonym(s), specimen(s), efc. are con- cerned
references =see for abbreviations the list in vol. 5, pp. cxlv—clxv
Res. = Residency or Reserve
resp. = respective(ly)
Abbreviations and signs
S=south (after degrees: southern latitude)
S (after a vernacular name) = Sundanese (language)
Sbl. =Sambali (Philip. language)
SE. = southeast
sec. =secus; according to
sect. =sectio; section
sens. ampl. (ampliss.)=sensu amplo (amplissimo); in a wider sense, in the widest sense
sens. lat. =sensu lato; in a wide sense
sens. str. (Sstrictiss.)=sensu stricto (strictissimo); in the narrow sense, in the narrowest sense
Sept. = September
seq., seqq. =sequens, sequentia; the following
ser. = series
s.1. =sensu lato; in a wide sense
S.-L. Bis. =Samar-Leyte Bisaya (Philip. language)
Sml. = Samal (Philip. language)
s.n.=sine numero; (specimen) without the collec- tor’s number
Sp. =Spanish (language)
sp(ec). = species; species
specim. = specimen(s)
sphalm. = sphalmate; by error, erroneous
Spp. = species; species (plural)
Sr = Senior
S.S. =see sens. Str.
ssp. = subspecies; subspecies
s.str. =see sens. Str.
stat. nov. =Status nova; proposed in a new rank
Sub. = Subanum (Philip. language)
subg(en). =subgenus; subgenus
subsect. = subsectio; subsection
subsp. = subspecies; subspecies
Sul. = Sulu (Philip. language)
Sum. E.C. = Sumatra East Coast
Sum. W.C.=Sumatra West Coast
Suppl. = Supplement
SW. =southwest
syn. =synonymum; synonym
synonyms =the names of taxa which have been re- ferred to an earlier described taxon of the same rank and with which they have been united on taxonomical grounds or which are bound together nomenclaturally
syntypes = the specimens used by the original author when no holotype was designed or more specimens were simultaneously designated as type
t. = tabula; plate
Tag. = Tagalog (Philip. language)
Tagb. = Tagbanua (Philip. language)
Tagk. = Tagaka-olo (Philip. language)
Tapan. = Tapanuli (in NW. Sumatra)
taxon=each entity throughout the hierarchic ranks of the plant kingdom which can be described and discriminated from other taxa of the same rank
Taxon. = Taxonomy
Tg =Tandjung (Malay); cape
Ting. = Tinggian (Philip. language)
Tir. = Tirurai (Philip. language)
transl. = translated
type = each taxon above the rank of a species is typi- fied by a type belonging to a lower rank, for in- stance a family by a genus, a genus in its turn by a species; a species or infraspecific taxon is typified by aspecimen. The name of a taxon is nomenclatu- rally permanently attached to its type; from this it cannot be inferred that the type always represents botanically the most typical or average structure found in the circumscription of the taxon.
type specimen=the specimen or other element to which the name of a species or infraspecific taxon is (nomenclaturally) permanently attached; botan- ically a type specimen is a random specimen on which the name was based by description. There- fore, it does not need to represent the average or most typical representative of a population. See holotype, isotype, lectotype, syntype, paratype, and neotype
typ. excl. = typo excluso; type excluded
typ. incl. =typo incluso; type included
typus =see type and type specimen
var. = varietas; variety
var. nov. =varietas nova; new variety
Vern. = Vernacular
vide = see
viz. = videlicet; namely
vol. = volume
W =west (after degrees: western longitude)
Yak. = Yakan (Philip. language)
+ =about
) = male (flower, efc.) 2 =female (flower, efc.)
(
)=monoecious with unisexual flowers
)= polygamous
) = polygamous
c’= many
> =more than (in size, number, efc.)
< =less than (size, number, efc.)
xX 2/5=2/5 of natural size
Xx montana= means that the epithet montana is that of a hybrid
(47)
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vrs
ARALIACEAE—I (W. R. Philipson, Christchurch)!
Trees, shrubs, lianas, woody epiphytes or (extra-Mal.) 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-Mal.) 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 1150 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 Scheffiera 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)
9) 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 Schefflera, 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 Scheffiera 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 nere estimated as 23 : 1, but if the very large genus Schefflera is not accounted for this reduces to 13 : 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 (cf. DILCHER & Do pn, 1970). Many fossils previously referred to Oreopanax must now be placed in Platanaceae (DoyLeE, pers. comm.). Dendropanax has reliably been reported from Tertiary deposits in both Europe and North America where it is now absent (DILCHER & DOLPH, /.c.).
References: DiLCHER & DoLpH, Amer. J. Bot. 57 (1970) 153-160; FRopIN, 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 Brn. 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 (1938) observed H. aculeatus (BL.) WARB. ex BOERL. as a characteristic pioneer in grass thickets on hills in SW. Celebes. H. harmsii K. Scu. 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. MOorRE & F.v.M.) HArMs is frequently seen in stable monsoon scrub and forest borders in the Port Moresby region and P. /Jedermannii 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 of Xanthomyrtus-Vaccinium-Papuacedrus-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 (S. falcata PHILIPSON), a species of uncertain affinities. Scheffiera 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 Scheffiera, many others are more or less shade-loving epiphytes or vines of lower storeys within the forest, although they may occur in sunnier situations where clouding is frequent (S. 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 (S. gemma FRODIN). A few species are con- spicuous rosette-trees of the forest understorey (S. 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, Casuarina junghuhn- 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 1000 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 3900 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 effect 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 showy 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 of Fatsia) 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. 91
usually) drupaceous, they will also be eaten by birds (for the most part) and bats, as recorded by RiD.Ley (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. Arn. 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; RipLey, 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 (4rthrophyllum, 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 othe: 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 (S. 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 l’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 Tomlinson & 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 (Bul, 1974).
Floral anatomy is discussed by BAUMANN-BODENHEIM (1955), PHILIPSON (1967, 1970) and especially by EypE & 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; But NGoc-SAnu, Bull. Mus. Hist. Nat. Paris III, Bot. 18 (1974) 85-91 (whole no 271); Davis, Systematic embryology of the angiosperms, New York (1966); EypE & TsENG, J. Arn. Arb. 52 (1971) 205-239; GRUSHVITZKY, Proc. Int. Symp. Physiol. Ecol. & Bio- chem. of Germination (ed. H. Borriss) (1967); GUssow, 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. Midl. 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; VicuiER, Ann. Sci. Nat. Bot. IX, 4 (1906) 1-209; ibid. TX, 9 (1909) 305-405.
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/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); SHourp & 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.5S. (1969). The family shows considerable constancy of the basic number of x = 12, although x = 11 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, /.c. 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. 91
ferae (1971) 267-277; in Cauwet-Marc & Carbonnier (eds.), Les Ombelliféres. Contributions pluridisciplinaires a la systématique. 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, 1940) 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 f. 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 (/.c.) 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 (cf. 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 dnd 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 (l.c.) who considered that phyletic changes could have taken place in parallel; and EypE & 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) |
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 differences 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 VicuiER. 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 Tupidanthus). 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 differently 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 FRopDIN 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, /.c.).
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 Schefflera and Polyscias. 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; Eypge, Amer. Sci. 63 (1975) 430-437; EYDE & 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) 427-448; HALLE, OLDEMAN & TOMLINSON, Tropical trees and forests: an architectural analysis, Berlin (1978); HALLIER f. New Phytol. 4 (1905) 151-162; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1898) 1-62; HLapik, Adansonia 10 (1970) 383-407; HUTCHINSON, Gen. FI. Pl. 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; RopRIGUEZ, 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; Vicuter, 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.) Vic. 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, Dict. rev. ed. (1966); HEYNE, Nutt. Pl. (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. Ss EeaVvesspalmatelvalObeGaitOMment OSC mm uriiCinrsince cnn mnie a on 1. Harmsiopanax 3, Leaves pinnarciy lobed on entire, glabrous’ =). 3 << 2) eet Renee ee 2. Aralidium
1979] ARALIACEAE—I (Philipson) 9
2. Leaves pinnately compound (or bi- or tripinnate).
Amibeavesmwice (Or more) pinnate (eo ee ac sk ke sk we tw ee 3. Aralia 4. Leaves once pinnate. Seebenheroumany csi: ©. seen Lae ROE ew. Stes. sla ao oe aes 4. Delarbrea Sra HerNE OTC WOR 8, oe Re eM ed can Ban 055) odd ios outa 5. Pentapanax 1. 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
TOG: 9: Sahara iment Meme ys Bent ee er en A 7. Osmoxylon 7. Inflorescence branches not as above. fo. LO EER Sod a NVoy 1 ite] | Oe aan SP ee De et ee a 8. Arthrophyllum
8. Ovary with more than one cell. 9. Leaves pinnate. lOfPedicel not articulated below the flower.5..> 44 ne eee 9. Gastonia OM Pedicellarticulatedibelow;themowel cme ae en eee een oe ae: 10. Polyscias 9. Leaves not pinnate. 11. 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 11. Pedicel not articulated below the flower. 13. Leaf simple (or unifoliolate) or palmately lobed. 14. Leaf palmately lobed.
BO VARY 2-COMEG: (yes a4 oc si o Sehst a os Ste S o 13. Brassaiopsis IS Ovary,.10=orsmore-celledvose-s.... oachaaet wud) Bile Soo Gickbs OR Seoue 14. Trevesia 14. Leaf simple (or unifoliolate). 16. Articulation present between petiole and blade. ........2.2.2.2.. Schefflera 16. No articulation between petiole and blade. AV ROVATYP2-COLCA) graye cers. oy Bn sk She Te Rs By GED tsuki, Sieies dates ess 13. Brassaiopsis ieeOvary’4-.or;more-celled o..) 0.546" ¢ os EA ee thes Cl NS at db es 15. Dendropanax 13. Leaf digitately compound. 18. Petiolules joined together by a web of tissue. . .........2.2.4.-. 14. Trevesia
18. No such web of tissue present. 19. Styles or stigmas 2.
Boe Otyle DING teri Woes 6 We sb earane 6 Feb. oxen? ecaeey WA Moa aee” 16. Acanthopanax
20 estylestunited intola columnysure-ges eee lee) eRe ee nome) oe lee 13. Brassaiopsis
ios Styles or stigmas more: than.2. oi Spill nls Shen sin eine Nelsen iene Schefflera 1. HARMSIOPANAX
Wars. in E. & P. Nat. Pfl. Fam. Nachtr. 1 (1897) 166; HArRms, Bot. Jahrb. 56 (1921) 413; Hutcu. Gen. FI. Pl. 2 (1967) 62; PHiLipson, Blumea 21 (1973) 81. — Schubertia BL. Bijdr. (1826) 884, nom. illeg., non MirRB. 1812. — Horsfieldia BL. ex DC. Prod. 4 (1830) 87, non WILLD. 1805; Bru. in B. & H. Gen. Pl. 1 (1865) 937; BoerL. Hand. 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
, Rh SN a SS SH litte: y YY a ea a a Sana 2) REREeeah es Seen +) ,—,, NAN, . = ON * 4 r. i, o ayiNatal BP une > NY See Nea pa era SRO SN AS RY ACO Ke -—. Brea Ne vo Ss r ‘a rl (3 AR Sy a p = > =, SORUN AALS OS. Ueraaaia SS | aay, Sg AN oa, ie ete TA ba
TR RS eS mame ie i
MS
[ser. I, vol. 91
FLORA MALESIANA
10
Seep Nd als ‘ Bs a
lule, x 4, d. developing fruit, x 12 (a NGF 36901, b-d Puiipson 3483).
Fig. 1. Harmsiopanax ingens PHILIPSON ssp. ingens. a. Leaf, x 4/., b. part of inflorescence, x 1/3, c. umbel-
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. PemUuMIDEIUIESISESSIIC.. 5 6 es He 2 Wmbellules peduncled. . ..... .4 4.
1. Upper surface of leaves with many (or rarely few) larger spines among the setulose hairs
1. Harmsiopanax aculeatus (BL.) WARB. ex BOERL. Hand. 3 (1900) 88; Koorp. Exk. Fl. Java 2 (1912) 719; Atlas 4 (1916) f. 668 & 669; BakH. f. & OoststR. in Back. Bekn. Fl. Java (em. ed.) 7 (1946) fam. 159, p. 19; Back. & BAKH. f. Fl. Java 2 (1965) 171; STEEN. Mt. FI. Java (1972) pl. 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 BTuH. in B. & H. Gen. Pl. 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 @, 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
si Yous 46: JeMve® “eLie se, im. a5 ea) Pw) ie, Se
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. !/, mm long. Calyx rim fringed. Petals strap-shaped, c. 11/, mm 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. FI. fr. April-Nov. ScHMutTz 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. Scu. 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, !/, 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) 1)
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. 1'/, mm long. Calyx rim fringed with many lacerate filaments. Petals ovate, c. 1 mm long. Filaments c. 1 mm; anthers c. '/, mm 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 . 4 3 ® .
3. Harmsiopanax ingens PHILIPSON, Blumea 21 (1973) 84.
ssp. ingens. — Fig. 1, 3.
Unbranched tree up to 18 m witha 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 @, 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. 91
except for a few bristles; petiole up to 1 m and 3 cm @, 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 2-31/, mm; anthers 4/,—3/, 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., kKamul, Hindenburg Ra.; Western Highlands: murri, Hagen, tolsan, Minj, mauri, Melpa, mai, 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 witha 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 {c. 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 @).
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 {1500-2000 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
Mia. Pl. Jungh. 3 (1855) 423; Fl. Ind. Bat. 1, 1 (1856) 762, t. 13; Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 25; Bru. in B. & H. Gen. Pl. 1 (1865) 936; HEmsL. in Hook. Ic. Pl. 16 (1886) t. 1549; Bogert. Handl. 1 (1890) 631; Harms in E. & P. Nat. Pf. Fam. 3, 8 (1894) 60; Hutcu. Gen. FI. Pl. 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. Aralidium pinnatifidum (JUNGH. & DE VriesE) Mia. a. Habit, x 2/;, b. 3 flower and bud, c. 2 flower and bud, both x 8, d. fruit, seed, and CS, slightly enlarged (a VAN BALGOOY 2185, b CocKBURN FRI 8376, c SINCLAIR 9884, d fresh material). Drawn by W. R. PHILIPSON.
16 FLORA MALESIANA [ser. I, vol. 91
locules (2 abortive) and 1 ovule, styles 3-4, tapering from broad bases, stigmas terminal. Fruit drupe-like. Seed solitary, pendulous from a thickened funicle, 4-5-grooved; endosperm deeply ruminate.
Distr. Monotypic. Peninsular Thailand and Malesia: Malay Peninsula, Sumatra, and Borneo.
The record from Java by Mique (Ann. Mus. Bot. Lugd.-Bat. 1, 1863, 25) is erroneous (see K. & V. Bijdr. 7, 1900, 2).
Ecol. Primary and secondary forest, from sea-level to c. 1250 m.
Notes. The single species forms a genus with several unique features, namely the large, simple, charac- teristically pinnately lobed leaves, the absence of resin ducts, the diffuse panicles of male or female flowers, the 3-carpellate ovary with a single surviving loculus resulting in a single-seeded fruit, the dorsal raphe, the deeply ruminate endosperm, and the enlarged funicle.
The genus is treated here as a member of the Araliaceae mainly as a matter of convenience. Sometimes it has been placed in the Cornaceae (e.g. RiDL. Fl. Mal. Pen. 1, 1922, 894, and Vicuter, Ann. Sc. Nat. Bot. 4, 1906, 171), and on full investigation it may well prove to be better placed in that family. The absence of resin ducts and the dorsal raphe strongly support a relationship with the Cornaceae and its immediate allies, though the absence of borders to the pits of the xylary fibres is characteristic of Araliaceae.
Several genera formerly placed in the Cornaceae have now been elevated to the rank of family. If this course is followed then Aralidium should also be segregated. Many of the features of Aralidium approach those of Griselinia (segregated as Griseliniaceae) and possibly these two genera should be united as a
single family.
1. Aralidium pinnatifidum (JUNGH. & DE VRIESE Mia. FI. Ind. Bat. 1, 1 (1856) 763; HEMsL. in Hook. Ic. Pl. 16 (1886) t. 1549; BoerL. Handl. 1 (1890) 631; Ripv. Fl. Mal. Pen. 1 (1922) 895; Puiieson, J. Bot. 78 (1940) 118. — Aralia pinnatifida JUNGH. & DE VRIESE, Ned. Kruidk. Arch. 1 (1846) 15; Ann. Sc. Nat. III, 6 (1846) 115. — A. dentatum Mia. Sum. (1861) 340. — A. integrifolium Herne in Fedde, Rep. 54 (1951) 245. — Fig. 4.
Shrub or small tree up to c. 10 m, rarely reaching 20 m and 25 cm @, glabrous in its vegetative parts; buds enclosed in long sheathing leaf-bases. Leaves spaced with distinct internodes, usually c. 30 by 22 cm or more, + regularly pinnately incised, frequently as deep as the midrib, lobes oblong- acuminate and decurrent on the midrib, c. 2!/, cm wide or more, the lobing sometimes irregular, and occasionally the blade entire and broadly ovate (up to 25 by 20 cm) or rarely lanceolate, leaf mar- gin either entire or coarsely dentate, especially on the terminal lobe; petioles 5-12 cm, broadly channelled above, clasping the stem with a slightly dilated base, exstipulate. Panicles terminal, or occasionally in the upper axils, to 50 cm long, pendulous, puberulous; main bracts caducous, but the minute bracteoles often persisting until anthesis. Flowers numerous, small (buds c. 2!/, mm long), fragrant, creamy or red-tinged, ovary, calyx lobes and petals densely covered in a minute but coarse puberulence. Male flowers with the corolla persistent during anthesis, petals c. 1!/, mm long, strap-shaped, spreading, stamens c. 1 mm with flattened filaments and round anthers; stylopodium a succulent disk with a concave centre; styles absent, the ovary 11/, mm long, narrowly turbinate,
without a loculus. Female flowers with the corolla caducous at anthesis together with the staminodes, styles divergent from their gibbous bases, ovary ovate with a single loculus (two abort early); ovule pendulous. Fruit usually obliquely ellipsoid, tapering to the apex and c. 3-4'/, cm long, but rarely subspherical, white when immature, ripening to purplish or black, juicy; exocarp fleshy, endocarp chartaceous. Seed broadly ellipsoid, 2—2'/, cm long, with the surface patterned with deep ruminations.
Distr. Peninsular Thailand; in Malesia: Malay Peninsula (from Kedah southwards common; Singapore), throughout Sumatra (also in Simalur I.), Anambas Is. (Siantan) and throughout Borneo.
Ecol. Frequent in evergreen primary rain- forest, also in open bamboo forest and secondary growths, from sea-level to c. 1250 m, in Borneo up to 1500-1800 m. FI. fr. Jan.—Dec.
Uses. The only use, once mentioned, is from Brunei, as “leaves make good ghost medicine’’.
Vern. Malay Peninsula: /émpédu buaya, (poko) balai, pungar, sahalat, sébalai tingal, sibilai, tébalai, M; Sumatra: (kayu) attarodan, Asahan, Batak lang., ségéntut, Gajo, médung, M, maneél silai, mannel dotan, sukun dotan, M, Simalur; Anambas Is.: ballok, M, Siantan; Borneo: daun tutchol antu, Brunei, Iban lang.
Note. Entire leaves are not infrequent through- out the range of the species, so that the recognition of a second species using this character is not justified. Coarsely dentate leaf-margins were also employed as a specific character but are merely a minor variation. Some specimens from Mt Kina- balu have rather small globose fruits, but the typical form of fruit also occurs on that mountain.
1979] ARALIACEAE—I (Philipson) 17
3. ARALIA
LinnE, Gen. Pl. ed. 5 (1754) 134; Sp. Pl. (1753) 273; DC. Prod. 4 (1830) 257; Mia. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 6; BTH. in B. & H. Gen. Pl. 1 (1865) 936; BoeRL. Handl. 1 (1890) 629; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 56; STEEN. Bull. Bot. Gard. Btzg III, 17 (1948) 391; Hutcu. Gen. FI. Pl. 2 (1967) 63; STONE, Gard. Bull. Sing. 30 (1977) 134; PHiLipson, /.c. 97. — Acanthophora MERR. Philip. J. Sc. 13 (1918) Bot. 316, non LAMOUREUX, 1813 (Algae); STEEN. Bull. Bot. Gard. Btzg III, 17 (1948) 390. — Fig. 5, 6.
Sparingly branched shrubs or small trees, or climbing, rarely (extra-Mal.) herbaceous, glabrous or hairy, often prickly. Leaves pinnate to tripinnate, usually with leaflets at the insertion of the lateral pinnae; leaflets serrate; petiole with a sheathing base. /nflorescence a terminal panicle; flowers sessile or pedicelled, with an articulation below the flower; calyx with 5—6 teeth; petals 5—6, imbricate; ovary 2-6-celled; styles 2-6 free or shortly connate below. Fruit a fleshy drupe; pyrenes cartilaginous, compressed; endosperm uniform.
Distr. More than 30 spp. in North America (S. to Mexico) and East Asia, 6 spp. in Malesia: Sumatra, Malay Peninsula, Java, Lesser Sunda Is. (Sumba), Borneo, Celebes, Philippines, and West New Guinea.
Ecol. Usually on scrubby hillsides and in secondary growth, often in ravines or near streams, or in thickets near or above the limit of tree-growth, at low altitude (100 m), but usually in the montane zone, up to 3000 m.
Note. For a discussion of specific distinctions see VAN STEENIS, /.c. 391. Hut-Liw Li in Sargentia 2 (1942) 101, treated some species that extend into Malesia. MERRILL considered that the climbing habit and recurved spines of Acanthophora justified its separation as a distinct genus, but more recent authors have not agreed.
KEY TO THE SPECIES
1. Flowers sessile (capitate) or very shortly pedicelled. 2. Flowers sessile, underside of leaf + densely tomentose, hairs of the branches and inflorescence + appressed and felted, bracts around the capitula enveloped in hairs 1. A. dasyphylla 2. Flowers short-pedicelled, underside of leaf sparsely tomentose, hairs of the branches and inflorescence + patent, bracts around the capitula less densely tomentose 2. A. javanica 1. Flowers +/,-11/, cm pedicelled (umbellate). 3. Climbing or scrambling liana, spines curved 3. Erect shrubs or small trees, spines straight. 4. Leaflets glaucous beneath, margins with few crenations. Fruit small (c. 3 mm long) 4. A. bipinnata 4. Leaflets green (or with fawn pubescence) beneath, margins serrate. Fruit rather larger (4-6 mm long). 5. Young parts and undersurface of leaves glabrous (but with small spines) 5. A. ferox 5. Young parts and undersurface of leaves pubescent
Pe oR ee 2 eS ee te ce ee eA
1. Aralia dasyphylla Mia. FI. Ind. Bat. 1, 1 (1856) 751; Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 9, incl. var. strigosa Mia. et var. latifolia MiQ.; BOERL. Handl. 1 (1890) 646; K. & V. Bijdr. 7 (1900) 53; Koorp. Exk. Fl. Java 2 (1912) 718; Atlas 4 (1916) f. 673 A-K; Hut-Lin Lt, Sargentia 2 (1942) 20; BAKH. f. Blumea 6 (1947) 367, incl. var. urticifolia (BL. ex Mia.) BAKH. f.; BAKH. f. & OosTsTR. in Back. Bekn. Fl. Java (em. ed.) 7 (1948) fam. 159,