SYSTEMATICS AND EVOLUTION OF THE PALM GENUS ARECA CHARLIE DANNY HEATUBUN THE GRADUATE SCHOOL BOGOR AGRICULTURAL UNIVERSITY 2009

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1 SYSTEMATICS AND EVOLUTION OF THE PALM GENUS ARECA BY CHARLIE DANNY HEATUBUN THE GRADUATE SCHOOL BOGOR AGRICULTURAL UNIVERSITY 2009

2 I, Charlie Danny Heatubun, certify that this dissertation is my own original work and has not been submitted in a previous application for a higher degree. June, 25 th 2009 Charlie Danny Heatubun

3 ABSTRACT CHARLIE DANNY HEATUBUN, Systematics and Evolution of Palm Genus Areca L. supervised by SRI S. TJITROSOEDIRDJO, JOHANIS P.MOGEA, WILLIAM J. BAKER, and MIEN A. RIFAI. Areca L. (Arecoideae: Areceae: Arecinae) is treated in this study as a genus of South Asian West Pacific palm and best known for its type species Areca catechu, the betel nut palm. This genus comprises 41 species and included into two subgenera; subgenus Areca and subgenus Beccarioareca. Seven species are newly recognized (A. bakeri Heatubun, A. churchii Heatubun, A. dransfieldii Heatubun, A. gurita Heatubun, A. mogeana Heatubun, A. riparia Heatubun, and A. triginticollina Heatubun). Eight previously recognized species (A. jobiensis Becc., A. multifida Burret, A. nannospadix Burret, A. nigasolu Becc., A. rechingeriana Becc., A. rostrata Burret, A. torulo Becc. and A. warburgiana Becc.) are reduced to synonymy with A. macrocalyx Zipp. ex Blume; two species (A. guppyana Becc. and A. salomonensis Burret) are also reduced to A. novohibernica (Lauterb.) Becc. and one species (A. celebica Burret) to A. oxycarpa Miq.; and also one species A. macrocarpa Becc. is also synonym to A. catechu L. While, three species (A. chaiana J. Dransf., A. congesta Becc., and A. ledermanniana Becc.) are considered to species incertae sedis (doubtful or uncertain species). A determination key is presented to subgenera and all easts of Wallace s line taxa, including their detailed descriptions and for new species. A phylogenetic analysis of certain species in the genus was performed based on DNA sequences from two low-copy nuclear genes, phosphoribulokinase (PRK) and the second largest subunit of RNA polymerase II (RPB2). The monophyly of Areca as genus is reconfirmed together with its two lineages are also recovered. Subgenera and sections in the genus were assessed based on phylogenetic relationship together with their biogeography explanation. Natural history observations, including uses and conservation status are also presented in this study. Key words: palms, Arecaceae, Areca, systematics, morphology, DNA, phylogeny, classification, species.

4 ABSTRAK CHARLIE DANNY HEATUBUN. Sistematika dan Evolusi Marga Palem Areca L. Dibimbing oleh SRI S. TJITROSOEDIRDJO, JOHANIS P. MOGEA, WILLIAM J. BAKER dan MIEN A. RIFAI. Areca L. (Arecoideae: Areceae: Arecinae) adalah marga palem yang distribusinya dari Asia Selatan sampai Pasifik Barat dan sangat terkenal dengan jenis tipenya Areca catechu, pinang sirih. Marga ini terdiri dari 41 jenis yang tergabung dalam dua anak marga; anak marga Areca dan anak marga Beccarioareca. Tujuh jenis adalah jenis baru (A. bakeri Heatubun, A. churchii Heatubun, A. dransfieldii Heatubun, A. gurita Heatubun, A. mogeana Heatubun, A. riparia Heatubun, and A. triginticollina Heatubun). Delapan jenis yang sebelumnya dikenal (A. jobiensis Becc., A. multifida Burret, A. nannospadix Burret, A. nigasolu Becc., A. rechingeriana Becc., A. rostrata Burret, A. torulo Becc. and A. warburgiana Becc.) merupakan sinonim baru untuk A. macrocalyx Zipp. ex Blume; dua jenis (A. guppyana Becc. and A. salomonensis Burret) juga merupakan sinonim A. novohibernica (Lauterb.) Becc. dan satu jenis (A. celebica Burret) untuk A. oxycarpa Miq.; demikian pula A. macrocarpa Becc. merupakan sinonim untuk A. catechu L. Sementera tiga jenis (A. chaiana J. Dransf., A. congesta Becc., and A. ledermanniana Becc.) dipertimbangkan sebagai jenis incertae sedis (meragukan atau jenis yang tidak pasti). Kunci determinasi ditampilkan untuk anak marga dan jenis-jenis yang berasal dari sebelah Timur Garis Wallace, demikian pula deskripsi lengkapnya dan juga untuk jenis baru. Analisis filogenetika untuk beberapa jenis pada marga didasarkan pada sekuens DNA dari dua low-copy gen inti; phosphoribulokinase (PRK) dan the second largest subunit of RNA polymerase II (RPB2). Monofiletik Areca sebagai marga kembali dikonfirmasi bersama dengan penemuan dua jalur evolusi di dalamnya. Anak marga dan seksi-seksi pada marga ini dikaji berdasarkan hubungan kekerabatan (filogeni) dan juga penjelasan tentang biogeografinya. Pengamatan tentang peri kehidupan alam termasuk kegunaan dan status konservasi juga ditampilkan dalam penelitian ini. Kata kunci: palem, Arecaceae, Areca, sistematika, morfologi, DNA, filogeni, klasifikasi, jenis.

5 SUMMARY Areca L. was the first palm genus was described by Linneaus in Species Plantarum in 1753 based on Rumphius Pinanga published in Herbarium Amboinense in 1741 (see Corner, 1966; Moore & Dransfield, 1979). It is also the type genus of the family name Arecaceae Bromhead. This genus is best known for its type species, Areca catechu the widely cultivated betel nut palm, the seed of which is chewed with the leaf or inflorescence of Piper betle L. (Piperaceae), lime and sometimes tobacco and spices, principally as a mild stimulant. Betel nut is consumed by an estimated million people world-wide (Gupta & Warnakulasuriya, 2002; Loo et al., 2006; Dransfield et al., 2008) extensively in the Asia-Pacific region and expatriate Asian communities making betel nut the fourth most widely abused substance after nicotine, alcohol and caffeine (Norton, 1998; Loo, et al. 2006). The genus Areca has approximately 47 species (Dransfield et al., 2008), and is distributed from India and south China through Malesia to New Guinea and the Solomon Islands (Dransfield, 1984; Uhl & Dransfield, 1987; Dransfield et al., 2008). The objective of this research is to provide a modern taxonomic treatment of Areca including studies of its phylogenetic relationships, natural history, uses and conservation status. The work will be based on exhaustive studies of existing literature, a thorough examination of herbarium materials in international herbaria, extensive fieldwork and laboratory-based molecular systematic research. The project consists of four main subprojects: 1). Taxonomic revision of the genus Areca; 2). Species level phylogeny estimation; 3). Comparative morphological and molecular study of the genus Areca with aim to understand the morphological changes that occurred during their evolution; 4). and if possible to reconstruct the historical biogeography and origin of A. catechu L. Taxon sampling and observations for the morphology and distribution of the species were based on herbarium samples or specimens (dried and spirit-preserved materials) deposited at international herbaria, namely A, AAU, B, BH, BO, BRI, FI, K, KEP, L, LAE, MAN, PNH, SAN, SAR, and SING (herbarium acronyms follow Holmgren et al., 1990), as well as the newly established small herbarium in Balai Penelitian Kehutanan (Forestry Research Institute) in Manokwari, West Papua, Indonesia. Measurements were taken from spirit-preserved material and dried herbarium specimens and from living collections. Floral parts were measured from spirit-preserved material or dried specimens and rehydrated by boiling. Basic morphological characters such as habit, stem, leaves, inflorescence, staminate flower, pistillate flower, fruit, seed and their details were used to describe and recognize taxa; all morphological data was used for producing the descriptions of each taxon, while the key to species was constructed from the diagnostic characters only. The morphological species concept or taxonomic species concept was applied as a framework to define taxa, and assessed later with a phylogenetic species concept (Davies & Heywood, 1963; McDade, 1995; Gornall, 1997; Mayden, 1997; Dransfield, 1999), especially based on the result of molecular phylogenetics analysis of the genus Areca (Heatubun et al., in prep.). The conservation status of each species of the genus Areca in east of Wallace s line was assessed based on the IUCN red list categories and criteria version 3.1 (IUCN, 2001). DNA-materials were collected directly in the fields or obtained from botanical gardens. Also were using collections of DNA bank of Jodrell Laboratory, Royal Botanic Gardens Kew, UK. All specimens will be fully vouchered with specimens, they were deposited in appropriate herbaria, and their identities confirmed. All available sequences of species of the genus Areca were included in the taxon sampling, with species of Nenga

6 and Pinanga (Loo et al. 2006), Bentinckia condapanna (Arecoideae: Areceae: unplaced), Clinostigma savoryanum (Arecoideae: Areceae: unplaced), Cyrtostachys renda (Arecoideae: Areceae: unplaced), Cyphokentia macrostachya (Arecoideae: Areceae: Clinospermatinae) and Tectiphiala ferox (Arecoideae: Areceae: Oncospermatinae) as outgroup (Norup et al., 2006; Dransfield et al., 2008; Baker et al., 2009). For DNA extraction, total genomic DNA extracted from silica gel-dried leaf materials (Chase & Hills, 1991) using the 2 CTAB method of Doyle & Doyle (1987). DNA were precipitated with 100% ethanol at 20 C, purified by cesium chloride/ethidium bromide gradients (1.55 g/ml) followed by dialysis and removal of ethidium bromide using butanol. Primer sequences for PRK and RPB2 were obtained from published sources (Lewis & Martinez, 2000; Lewis & Doyle, 2002; Roncal et al. 2005; Loo et al. 2006; Norup et al. 2006; Trenel et al., 2007; Cuenca et al., 2007). Sequence alignment follow Loo et al. (2006) and Norup et al. (2006) which every base position in the reverse and forward sequences were check and assembled using Sequencher 4.1 (Gene Codes Corp, Ann Arbor, Michigan, USA) and will be deposited in GenBank. The sequences then enter and aligned manually into data matrices in PAUP* version 4b10 (Swofford, 2002) for Macintosh computer. Phylogenetic analyses in this study were using parsimony analysis and Bayesian analysis. the standard procedures as follow: congruence between the PRK and RPB2 datasets will evaluated using the incongruence length difference (ILD) test of Farris et al. (1994) as implemented in PAUP* (Swofford 2002) and Mr. Test in MrBayes version 3.0b4 (Huelsenbeck & Ronquist, 2001). In parsimony analyses, uninformative characters and ambiguously aligned regions were excluded. All included characters were unordered and equally weighted. Initial analyses employed 1000 heuristic searches, each with starting trees obtained by random taxon addition, tree-bisection-reconnection (TBR) swapping, and keeping multiple trees per step (MulTrees on). Only groups that were found in the strict consensus tree and 50% or more of the replicates were recorded. Bayesian analyses were carried out using the program MrBayes version 3.0b4 (Huelsenbeck & Ronquist, 2001). Models of sequence evolution that the best fit the individual datasets were determined using Mr. Test. The models were evaluated by the Akaike information criterion (Akaike, 1973) implemented in the program. Parameters based on patterns in the data matrices. For all three datasets, four incrementally heated Markov chains were used in an analysis that was run for 100,000 generations initially with trees saved at every 10 th generations. Trees produced prior to stationarity were discarded as the burn-in. From this study, Areca is interesting palm genus not only for the species diversity but also for its natural history. This genus comprises 41 species and included into two subgenera; subgenus Areca and subgenus Beccarioareca. Seven species are newly recognized (A. bakeri Heatubun, A. churchii Heatubun, A. dransfieldii Heatubun, A. gurita Heatubun, A. mogeana Heatubun, A. riparia Heatubun, and A. triginticollina Heatubun). Eight previously recognized species (A. jobiensis Becc., A. multifida Burret, A. nannospadix Burret, A. nigasolu Becc., A. rechingeriana Becc., A. rostrata Burret, A. torulo Becc. and A. warburgiana Becc.) are reduced to synonymy with A. macrocalyx Zipp. ex Blume; two species (A. guppyana Becc. and A. salomonensis Burret) are also reduced to A. novohibernica (Lauterb.) Becc. and one species (A. celebica Burret) to A. oxycarpa Miq.; and also one species A. macrocarpa Becc. is also synonym to A. catechu L. While, three species (A. chaiana J. Dransf., A. congesta Becc., and A. ledermanniana Becc.) are considered to species incertae sedis (doubtful or uncertain species). A determination key is presented to subgenera and all easts of Wallace s line taxa, including their detailed descriptions and for new species. A phylogenetic analysis of certain species in the genus was performed based on DNA sequences from two low-copy nuclear genes, phosphoribulokinase (PRK) and the

7 second largest subunit of RNA polymerase II (RPB2). The monophyly of Areca as genus is reconfirmed together with its two lineages are also discovered. Subgenera and sections in the genus were assessed based on phylogenetic relationship together with their biogeography explanation. The morphological characters are still relevant and applicable to apply species concept in the genus Areca, however those characters are independently each others and evolved several times in this palm genus, so is needed a depth phylogeny interpretation to understand relationship and changes during the evolution. Dispersal is the main biogeography factor to reflect the recent distribution pattern of the genus Areca. The origin of the cultivated wide-spread species Areca catechu L. is still unresolved from this study. Natural history observations, including uses and conservation status are also presented in this study.

8 Copyright 2009, Bogor Agricultural University Copyright are protected by law 1. It is prohibited to cite all or part of this dissertation without referring to and mentioning the source. a. Citation only permitted for the sake of education, research, scientific writing, report writing, critical writing or reviewing scientific problems. b. Citation does not inflict the name and honour of Bogor Agricultural University. 2. It is prohibited to republish and reproduce all or part of this dissertation without the written permission from Bogor Agricultural University.

9 SYSTEMATICS AND EVOLUTION OF THE PALM GENUS ARECA BY CHARLIE DANNY HEATUBUN G As partial requirement fulfilment for the Doctoral Degree in Plant Systematics DEPARTMENT OF BIOLOGY THE GRADUATE SCHOOL BOGOR AGRICULTURAL UNIVERSITY 2009

10 External Examiners Examination I : Dr. Dra. Rugayah, M.Sc. Herbarium Bogoriense, Puslitbang Biologi, Indonesia Institute of Sciences (LIPI), Cibinong, Bogor, Indonesia. Dr. Dedy Duryadi Solihin, DEA Head of Biology Study Programme, Faculty of Mathematics and Natural Science, Bogor Agricultural University, Darmaga, Bogor, Indonesia. Examination II : Dr. Timothy M. A. Utteridge South-East Asia Section, Herbarium, Royal Botanic Gardens Kew, Richmond, Surrey TW9 3AB, United Kingdom. Dr. Kuswata Kartawinata Herbarium Bogoriense, Puslitbang Biologi, Indonesia Institute of Sciences (LIPI), Bogor, Indonesia; UNESCO office Jakarta, Indonesia and Department of Botany, Field Museum, Chicago, Illinois , USA.

11 Dissertation Title Name of Student Register Number Study Programme Sub Study Programme : Systematics and Evolution of the Palm Genus Areca : Charlie Danny Heatubun : G : Biology : Plant Taxonomy Approved by 1. Advisory Committee (Dr. Sri S. Tjitrosoedirdjo, M.Sc) (Chairman) (Prof. Dr. Johanis P. Mogea) (Member) (William J. Baker, M.Sc, Ph.D, FLS) (Member) (Prof. Dr. Mien A. Rifai) (Member) 2. The Biology Study Programme 3. The Graduate Schools (Dr. Ir. Dedy Duryadi Solihin, DEA) (Head) (Prof. Dr. Ir. Khairil A. Notodiputro, MS) (Dean) Date of Public Examination: July, 15 th 2009 Date of Graduation:

12 ACKNOWLEDGEMENTS Many individuals and institutions contributed to the success of this work and I am grateful to them all. I would like to thank my advisors Dr. Sri S. Tjitrosoedirdjo, Professor Johanis P. Mogea, Dr. William J. Baker, Professor Mien A. Rifai and Dr. John Dransfield, whose always guided my work and gave me countless and useful recommendations to finished this dissertation. Professor Edwino S. Fernando is gratefully acknowledged for sharing his expertise on the Philippine s Areca. Drs. Felix Forest, James J. Clackson and Professor Mark W. Chase to let me in to the lab, introduced me to DNA and Molecular Systematics, and guided me with the palm molecular phylogeny in the Jodrell Laboratory of Royal Botanic Gardens Kew. Drs. Christine D. Bacon and Raymond Baker from the Lyon Arboretum provided me with DNA materials of Areca from their garden collections in Hawaii. Dr. Tom D. Evans sent his collection of a new species of Areca from Cambodia. Professors Anders Barfod, Finn Borschenius and Henrik Balsev to invite me, arranged my travel and allowed me working with Areca collections in University of Aarhus and Copenhagen. Drs. Piero Cuccuini and Chiara Nepi also arranged my visit to Florence and allowed me working with the type specimens from Beccari s historical collections. Ms. Julia Anak Sang and Mr. Shahbuddin Moh. Shabki are thanked for assistance with permits and all things regarding my fieldtrips to Sarawak. Dr. George Argent is responsible to my visiting in Edinburgh and Dr. Jef Veldkamp for my Leiden trip. Dr. Benito Tan and Ms. Serena Lee for accessing herbarium data base and type specimens of Areca kept in the Singapore Botanic Garden. I am grateful to the Keepers and/or curators of herbaria A, AAU, B, BH, BO, BRI, FI, K, KEP, L, LAE, MAN, PNH, SAN, SAR, SING and BPK Manokwari for access to their specimens, data bases, and loan materials for study. I also thank to Lucy T. Smith for preparing the wonderful plates for my new species. Drs. Rugayah, Dedy Duryadi Solihin, Timothy M. A. Utteridge and Kuswata Kartawinata are acknowledged for their willingness being examiners in my final examination. I am grateful to Professor Eko B. Waludjo, Drs. Dedy Duryadi Solihin, Drajat Martianto, Dedy Jusadi, Marimin, Nunik Sri Aryanti, Harry Wiriadinata, Yohanes Purwanto, Dedy Darnaedi, Rugayah, Elisabeth Widjaja, Teguh Triono, and Ary Keim, and also Himmah Rustiami, Rani Asmarayani, Arief Hidayat from BO and IPB; Hans Arwam, Rudi

13 Maturbongs, Marthinus Iwanggin, Elieser Sirami dan Pilep Mambor from UNIPA; Daud Leppe, Herman Remetwa, Thomas Nifinruri and Krisma Lekitoo from BPK Manokwari; Drs. Rogier de Kok, Tim Utteridge, Aaron Davis and Soraya Villalba, and Melinda Trudgen, Edith Kapinos, Lzaslo Cziba, Steve Graham and Anne Morley-Smith from K; Hermus Indou, Herkilaus Rumaikewi, Tobias Paiki, Maikel Simbiak and Piter Matani supported my research in various way. My colleagues, friends and my Papuan families in Bogor always offered good suggestion and words of hope during the hard times of my study. Rector and the Dean of Fakultas Kehutanan Universitas Papua (UNIPA) Manokwari, Rector and the Dean of Sekolah Pascasarjana Institut Pertanian Bogor (IPB), they allow me to do my PhD. Financial supports came from BPPS Dikti Depdiknas and Royal Botanic Gardens Kew, UK for the PhD scholarships. The Royal Botanic Gardens Kew and Royal Botanic Gardens Edinburgh were funded my fieldtrips to Tanah Papua and North Sulawesi. My fieldtrips to Sarawak was granted by International Palm Society (IPS Endowment Fund 2007); visit to Nationaal Herbarium of Nederland, Leiden Branch by Flora Malesiana Foundation (Kostermann Funds) and Beasiswa Unggulan Dikti Depdiknas; visit to Royal Botanic Gardens Edinburgh, Scotland by the Sibbald Trust; travel to herbarium of University Aarhus and Monocot IV Symposium in Copenhagen, Denmark by the university of Aarhus; visit to Xishuangbanna Tropical Botanical Garden, Yunnan, China by Centre for Tropical Forest Science, Harvard University and China Academy of Sciences; and visit to herbarium Sezione Botanico, Meseo di Storia Naturale, Università degli Studi di Firenze, Florence, Italy by Royal Botanic Gardens Kew. The plates were funded by Pacific Biological Foundation and the Royal Botanic Gardens Kew. Finally, to my wife Oktarina Simanjuntak Heatubun, my son Edward Glorious Excelsa Heatubun and my daughter Narcissa Elegantia Heatubun for continuous supports during my PhD studies, especially for their patience while I was away far from home.

14 CURRICULUM VITAE Charlie Danny Heatubun is a son of Clemens Yoseph Heatubun ( ) and Selfina Endemina Hursepuny ( ). He was born on December 6 th, 1974 in Manokwari, West Papua. In 1985 he was graduated from primary school, and secondary school in 1988 and also high school 1991, all in Manokwari, West Papua. He was continued his study at Department of Forestry, Faculty of Agriculture, Cenderawasih University in Manokwari and graduated in Since 1997, he has been enrolled as academic staff in Forestry Department, Faculty of Agriculture; Cenderawasih University (now is Faculty of Forestry, University of Papua). In 2006, at the same year after finished his Master degree (M.Si) on Plant Taxonomy in Biology Department, the Graduate Schools of Bogor Agricultural University (SPs IPB); he was continued his study to doctoral level in the same department of the same university with the scholarships from BPPS Dikti Depdiknas and the Royal Botanic Gardens Kew, UK. He was married Oktarina S. Simanjuntak and has a son Edward Glorious Excelsa Heatubun and a daughter Narcissa Elegantia Heatubun. He is Fellow of the Linnean Society of London (FLS) and joint the International Association of Plant Taxonomy (IAPT), the Systematics Association (SYSTASS), the International Palm Society (IPS), the Biological Society of New Guinea, Indonesian Association of Plant Taxonomy (PTTI), and the IUCN Species Survival Commission (SSC) Palm Specialist Group (PSG) as member. He is the founder of Herbarium Papuaense of Forestry Research Institute in Manokwari (Balai Penelitian Kehutanan Manokwari) and he was the visiting fellow on several herbaria and botanic gardens in ASEAN, China, Australia and Europe. He is also co-authors of the books about tree species of Gunung Meja in Manokwari, West Papua and the edible fruits from the same area. His articles related with plant taxonomy and palm systematics were appeared in the several journals, including peer review international journals such as Systematic Botany, Kew Bulletin, Blumea, Flora Malesiana Bulletin and Folia Malaysiana.

15 CONTENTS LIST OF TABLES... LIST OF FIGURES... LIST OF APPENDIXES... Page xvii xviii xix GENERAL INTRODUCTION Background... 1 Objectives... 2 Literature Cited... 2 PHYLOGENY OF ARECA (ARECACEAE) BASED ON DNA Abstract... 6 Introduction... 7 Materials and Methods... 8 Results PRK Analysis RPB2 Analysis Combined Analysis Discussion Phylogenetics Value of PRK and RPB Morphology Systematics Implications Biogeography Acknoledgements Literature Cited... 24

16 A CONSPECTUS OF GENUS ARECA L. (ARECACEAE) Summary Introduction Materials and Methods Taxonomic Treatment Genus Description Infrageneric Classification of the Genus Areca Key to Subgenus of Areca I. Subgenus Areca II. Subgenus Beccarioareca Excluded and Uncertain Names Acknowledgements Literature Cited A MONOGRAPH OF ARECA OF EAST OF WALLACE S LINE Summary Introduction Materials and Methods Results and Discussion Morphology Habit Stems Leaves Indumentum Inflorescences Flowers... 98

17 Fruits and Seeds Natural History and Conservation Habitat and Ecology Pollination, Seeds Predation and Seeds Dispersal Biogeography Conservation Status Uses Taxonomic Treatment Key to Species of Areca East of Wallace s Line Species Description of Areca East of Wallace s Line Areca catechu L Areca macrocalyx Zipp. ex Blume Areca mandacanii Heatubun Areca novohibernica (Lauterb.) Becc Areca oxycarpa Miq Areca vestiaria Giseke Doubful and Uncertain Species Acknowledgements References Appendix: List of Specimens Examined and Identified GENERAL DISCUSSION Taxonomic Revision of the Genus Areca Phylogeny of the Genus Areca Literature Cited GENERAL CONCLUSION

18

19 LIST OF TABLES Page 2.1 Statistics calculated from parsimony analyses of PRK, RPB2 and combined datasets Comparison of previously recognised taxa and infrageneric classification in Areca with taxa and infrageneric classification recognised in this conspectus...34

20 LIST OF FIGURES Page 1.1 The 50% majority rule consensus tree of the Bayesian inference analysis of PRK dataset The 50% majority rule consensus tree of the Bayesian inference analysis of RPB2 dataset The 50% majority rule consensus tree of the Bayesian inference analysis of combined (PRK and RPB2) dataset Areca bakeri Heatubun Areca dransfieldii Heatubun Areca riparia Heatubun Areca churchii Heatubun Areca gurita Heatubun Areca mogeana Heatubun Areca triginticollina Heatubun Distribustion map of the species Areca which native to east of Wallace s line The inflorescence (infructescence) and the pistillate flowers morphology of Areca macrocalyx Reproductive organ of Areca vestiaria and Areca oxycarpa Areca macrocalyx Zipp. Ex Blume Areca novohibernica (Lauterb.) Becc...126

21 LIST OF APPENDIXES I. Papers have been published during PhD study ( ) in the Graduate Schools of Institut Pertanian Bogor (part of the dissertation and/or related to the Palms Systematics in general): 1. Dransfieldia (Arecaceae) A New Palm Genus from Western New Guinea. Systematic Botany 31: (2006). 2. Apakah Jenis Itu? (Sebuah Pelajaran Penerapan Konsep Jenis Pada Suku Palem- Paleman Di New Guinea). Rampak Serantau 14: (2007). 3. Two Species of Licuala (Arecaceae; Coryphoideae) from Western New Guinea. Blumea 53: (2008). 4. A new Areca from Western New Guinea. Palms 52: (2008) 5. A Monograph of Cyrtostachys (Arecaceae). Kew Bulletin 64: (2009).

22 GENERAL INTRODUCTION Background Areca L. was the first palm genus was described by Linneaus in Species Plantarum in 1753 based on Rumphius Pinanga published in Herbarium Amboinense in 1741 (see Corner, 1966; Moore & Dransfield, 1979). It is also the type genus of the family name Arecaceae Bromhead. This genus is best known for its type species, Areca catechu the widely cultivated betel nut palm, the seed of which is chewed with the leaf or inflorescence of Piper betle L. (Piperaceae), lime and sometimes tobacco and spices, principally as a mild stimulant. Betel nut is consumed by an estimated million people world-wide (Gupta & Warnakulasuriya, 2002; Loo et al., 2006; Dransfield et al., 2008) extensively in the Asia- Pacific region and expatriate Asian communities making betel nut the fourth most widely abused substance after nicotine, alcohol and caffeine (Norton, 1998; Loo, et al. 2006). The genus Areca also contains species of horticultural and agro-forestry significance (Ray & Reddy, 2001) and recent work has revealed considerable potential for therapeutic drugs to be extracted from some species, including cholesterol reducing agents (Byun et al., 2001) and neurological and dermatological treatments (Lee & Choi, 1999; Sullivan et al., 2000). The genus Areca has approximately 47 species (Dransfield et al., 2008), and is distributed from India and south China through Malesia to New Guinea and the Solomon Islands (Dransfield, 1984; Uhl & Dransfield, 1987; Dransfield et al., 2008). Although varying in size from undergrowth palmlets to moderately tall tree palms, the genus is well defined in its traditional concept by the presence of a crownshaft, infrafoliar inflorescences, the single inflorescence bract (prophyll), complete floral triads confined to the basal part of the branched inflorescence or its main axis, the symmetrical fruits with basally attached seed, and the ruminate endosperm with basal embryo (Dransfield, 1984). However, Areca displays perplexing variation in inflorescence structure and flower presentation that is in need of further analysis, particularly in view of its wide distribution throughout Southeast Asia, straddling Wallace s line (Dransfield, 1995). In the recent phylogenetic classification of the palm family, the genus Areca is placed together with Nenga and Pinanga in subtribe Arecinae; tribe Areceae; subfamily Arecoideae (Dransfield et al., 2005, 2008; Loo et al., 2006); differing from the previous classification which also included Gronophyllum, Gulubia, Hydriastele, Siphokentia (these four now reduced to a single genus Hydriastele see Baker & Loo, 2004) and Loxococcus (Dransfield & Uhl, 1986, 1998; Uhl & Dransfield, 1987, 1999).

23 2 Current phylogenetic evidence indicates that Areca is sister to a clade comprising Nenga and Pinanga (Loo et al., 2006; Norup et al., 2006; Baker et al., 2009). The last infrageneric classification of the genus Areca was proposed by Furtado (1933) comprising two subgenera and five sections: subgenus Blumeoareca (sections Arecella, Oeotheanthe and Axonianthe); and subgenus Beccarioareca (sections Microareca and Mischophloeus). However, these relationships within the genus are based on morphological affinities alone (Furtado, 1933; Dransfield, 1984; Harley & Dransfield, 2003), and it has been suggested that they need to be reassessed using modern methods (Dransfield, 1984). Moreover, the geographic origin of economic species A. catechu is still uncertain, with the Philippines (Beccari, 1919; Furtado, 1933), Malaysia (Corner, 1966; Jones, 1995) or Celebes (Corner 1966) currently hypothesised to be the locality of the species origin. Objectives The general aim of this PhD project is to provide a modern taxonomic treatment of Areca including studies of its phylogenetic relationships, natural history, uses and conservation status. The work will be based on exhaustive studies of existing literature, a thorough examination of herbarium materials in international herbaria, extensive fieldwork and laboratory-based molecular systematic research. The project consists of four main subprojects: 1). Taxonomic revision of the genus Areca; 2). Species level phylogeny estimation; 3). Comparative morphological and molecular study of the genus Areca with aim to understand the morphological changes that occurred during their evolution; 4). And if possible to reconstruct the historical biogeography and origin of A. catechu L. The main subprojects would be accommodating in the next following papers or chapters. Literature cited Asmussen, C. B. and M. W. Chase Coding and non-coding plastid DNA in palm systematics. Amer. J. Bot. 88: Baker, W. J. and A. H. B. Loo A synopsis of the genus Hydriastele (Arecaceae). Kew Bull. 59: Baker, WJ, Asmussen CB, Chase MW, Dransfield J, Forest F, Harley MM, Savolainen V, Uhl NW Wilkinson M Complete Generic Level Phylogenetic Analyses of Palms (Arecaceae) with Comparisons of Supertree and Supermatrix Approaches. Syst. Biol. doi: /sysbio/syp021

24 3 Beccari, O The palms of Philippines Islands. Phillipp. J. Sci. 14: Byun, S. J., H. S. Kim, S. M. Jeon, Y. B. Park, and S. M. Choi Supplementation of Areca catechu L. extract alters triglyceride absorption and cholesterol metabolism in rats. Ann. Nutri. Metabol. 45: Corner, E. J. H The natural history of palms. London: Weidenfeld and Nicolson. Dransfield, J The genus Areca (Palmae: Arecoideae) in Borneo. Kew Bull. 39: Dransfield, J. and N.W. Uhl An outline of classification of palms. Principes 30: Dransfield, J. and N. W. Uhl Palmae. In: Kubitzki, K (Ed.), the families and genera of vascular plants, vol. IV. pp , Berlin: Springer. Dransfield, J., N. W. Uhl, C. B. Asmussen, W. J. Baker, M. M. Harley and C. E. Lewis A new phylogenetic classification of the palm family, Arecaceae. Kew Bull. 60: Dransfield, J., N. W. Uhl, C. B. Asmussen-Lange, W. J. Baker, M. M. Harley and C. E. Lewis Genera Palmarum: The Evolution and Classification of Palms. Kew: Royal Botanic Gardens Kew. Flynn, T Morphological variation and species limits in the genus Areca (Palmae) in New Guinea and the Solomon Islands. Unpublished Master thesis, University of Wales, Bangor. Furtado, F. X The limits of the genus Areca L. and its sections. Repert. Spec. Nov. Regni Veg. 33: Govaerts, R., and J. Dransfield World checklist of palms. Kew: Royal Botanic Gardens Kew. Gupta, P. C., and S. Warnakulasuriya Global epidemiology of Areca nut usage. Addict. Biol. 7: Hahn, W. J. 2002a. A molecular phylogenetic study of the Palmae (Arecaceae) based on atpb, rbcl and 18S nrdna sequences. Syst. Biol. 51: Hahn, W. J. 2002b. A phylogenetic analysis of the Arecoid line of palms based on plastid DNA sequence data. Mol. Phylogenet. Evol 23: Harley, M. M. and J. Dransfield Triporate pollen in the Arecaceae. Grana 41: Jones, D. L Palms throughout the world. Sidney: Reed Books. Lee, K. K., and J. D. Choi Areca catechu L. extract. I. effect on elastase and aging. Journal of Cosmetics Science 49:

25 4 Lewis, C. E., and J. J. Doyle A phylogenetic analysis of tribe Areceae (Arecaceae) using two low-copy nuclear genes. Plant Syst. Evol. 236: Loo, A. H. B., J. Dransfield, M. W. Chase and W. J. Baker Low copy nuclear DNA, phylogeny and the evolution of dichogamy in the betel nut palms and their relatives (Arecinae; Arecaceae). Mol. Phylogenet. Evol. 39: Moore H. E., Jr. and J. Dransfield Typification of Linnean palms. Taxon 28: Norton, S. A Betel: consumption and consequences. J. Am. Acad. Dermatol. 38: Ray, A. K., and D. V. S. Reddy Performance of Areca-based high density multi species cropping system under different level of fertilizer. Tropical Agriculture 78: Sullivan, R. J., J. S. Allen, C. Otto, J. Tiobech, K. Nero Effects of chewing betel nut (Areca catechu) on the symptoms of people with schizophrenia in Palau, Micronesia. British Journal of Psychiatry 177: Uhl, N. W. and J. Dransfield Genera Palmarum: A Classification of Palms based on the work of Harold E. Moore Jr. Lawrance: L. H. Bailey Hortorium and International Palm Society. Uhl, N. W. and J. Dransfield Genera Palmarum after ten years. Mem. New York Bot. Gard. 83:

26 PHYLOGENY OF ARECA (ARECACEAE) BASED ON DNA CHARLIE D. HEATUBUN 1, 2, 3, 4, WILLIAM J. BAKER 3, JOHN DRANSFIELD 3, JIM J. CLARKSON 3, AND FELIX FOREST 3 1 Fakultas Kehutanan, Universitas Papua, Jl. Gunung Salju, Amban, Manokwari 98314, Papua Barat, Indonesia; 2 Departemen Biologi, Sekolah Pascasarja Institut Pertanian Bogor, Kampus Darmaga, Bogor 16680, Jawa Barat, Indonesia; 3 Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, UK. 4 Author for correspondence (charlie_deheatboen@yahoo.com) ABSTRACT. Phylogenenetics analysis of the palm genus Areca was performed based on the sequences of two low-copy nuclear genes phosphoribukilonase (PRK) and RNApolymerase II subunit B (RPB2). Two lineages of the genus Areca are discovered and the monophyly of Areca as a genus is also reconfirmed. The previous infrageneric classification within Areca (subgenera and sections) were assess based on the phylogenetic tree and together with their biogeography explanation. Dispersal is more likely the main factor to reflect the present distribution of the genus Areca. And the origin of the wide spread cultivated species Areca catechu still unresolved from this study.

27 HEATUBUN ET AL.: PHYLOGENY OF ARECA INTRODUCTION Areca is the type genus of the palm family Arecaceae and it distribute in the old world tropics from India, Srilanka, south of China to Malesian region to Solomon Island in the Pacific (Dransfield et al. 2008). The genus is well known from its type species, Areca catechu L., the betel nut palm or pinang. The betel nut palm is important commodity and has been developed to the large-scaled plantations for supplying the nuts demanding. Traditionally, the seeds are chewing for a mild stimulant (Loo et al., 2006; Dransfield et al., 2008), but in the recent years, the betel nut palm and other species of Areca have been used widely in therapeutic drugs; including cholesterol reducing agents (Byun et al., 2001), and neurogical and dermalogical treatments (Lee & Choi, 1999; Sullivan et al., 2000); horticulture and agroforestry significance (Ray & Reddy, 2001); and even further to maintain the biodiversity and conservation (The Economist, 8 th November 2008: 100). The genus Areca has approximately before 47 species by Dransfield et al. (2008) and reduces to 41 species in the recent revision (Heatubun et al., in prep.). Morphologically, the development of generic concept of Areca has been discussed in details by previous authors (Furtado, 1933; Dransfield, 1984; Uhl & Dransfield, 1987; Dransfield et al., 2008), including its relationships with other palm genera from morphology point of view (Harley & Dransfield, 2003; Baker et al., 2009). In the recent phylogenetic classification of the palm family, the genus Areca is placed together with Nenga and Pinanga in subtribe Arecinae; tribe Areceae; subfamily Arecoideae (Dransfield et al., 2005, 2008; Loo et al., 2006); differing from the previous classification which also included Gronophyllum, Gulubia, Hydriastele, Siphokentia (these four now reduced to a single genus Hydriastele see Baker & Loo, 2004) and Loxococcus (Dransfield & Uhl, 1986, 1998; Uhl & Dransfield, 1987, 1999). While, the current phylogenetic evidence indicates that Areca is sister to a clade comprising Nenga and Pinanga (Loo et al., 2006; Norup et al., 2006; Baker et al., 2009). The last infrageneric classification of the genus Areca was proposed by Furtado (1933) comprising two subgenera and five sections: subgenus Blumeoareca (sections Arecella, Oeotheanthe and Axonianthe); and subgenus Beccarioareca (sections Microareca and Mischophloeus). However, these relationships within the genus are based on morphological affinities alone (Furtado, 1933; Dransfield, 1984; Harley & Dransfield, 2003), and it has been suggested that they need to be reassessed using modern methods (Dransfield, 1984). 7

28 HEATUBUN ET AL.: PHYLOGENY OF ARECA Moreover, the geographic origin of economic species A. catechu is still uncertain, with the Philippines (Beccari, 1919; Furtado, 1933), Malaysia (Corner, 1966; Jones, 1995) or Celebes (Corner, 1966) and western New Guinea (Heatubun, 2008) currently hypothesised to be the locality of the species origin. The application of phylogenetics analysis to construct classification system in the palm family is widely accepted, and the sequences of specific DNA region also proved as powerful tools to solved taxonomic problem in palms. From DNA data used in the palms systematics, data from nuclear regions more variable and appear to evolve more rapidly (Dransfield et al., 2008), and this very useful at species level phylogeny analysis, especially low-copy nuclear gene (Lewis & Doyle, 2002). Recently, sequences of two low-copy nuclear genes have been shown to provide satisfactory phylogenetics information at lower taxonomic levels in the palm family (Bayton 2005; Lewis & Martinez 2000; Lewis et al. unpublished; Norup 2004; Roncal et al. 2005; Thomas et al. 2006), especially in subtribe Arecinae (Loo et al. 2006): nrdna PRK (phosphoribolukinase, a Calvin cycle enzyme) intron 4 (Lewis & Doyle 2001, 2002) and RPB2 (RNA polymerase II) intron 23 (Roncal et al. 2005). And also have been successful in molecular dated phylogeny analysis to construct biogeography history and dispersal events in palms (Gunn, 2004; Norup et al., 2006; Trenel et al., 2007; Cuenca et al., 2007). Based on an existing preliminary dataset of subtribe Arecinae (Loo et al., 2006), it is already known that these regions are useful for resolving relationships among certain species in subtribe Arecinae, including Areca. Thus, sequences from PRK and RPB2 were used in this study; to evaluating monophyly and relationships within the genus Areca and testing subgeneric and sectional grouping proposed by Furtado (1933), and if possible to reconstruct biogeographic history and origin of the betel nut palm Areca catechu. MATERIALS AND METHODS DNA-materials were collected directly in the fields by first author in western New Guinea (the Indonesian provinces of Papua and West Papua), North Sulawesi and Sarawak or obtained from botanical gardens, especially provided by Christine Bacon and Raymond Baker from Lyon Arboretum, Hawaii and Carl Lewis from Fairchild Tropical Botanical Garden Miami, Florida, USA. Also were using collections of DNA bank of Jodrell Laboratory, Royal 8

29 HEATUBUN ET AL.: PHYLOGENY OF ARECA Botanic Gardens Kew. All specimens will be fully vouchered with specimens, they were deposited in appropriate herbaria, and their identities confirmed. All available sequences of species of the genus Areca were included in the taxon sampling, with species of Nenga and Pinanga (Loo et al. 2006), Bentinckia condapanna (Arecoideae: Areceae: unplaced), Clinostigma savoryanum (Arecoideae: Areceae: unplaced), Cyrtostachys renda (Arecoideae: Areceae: unplaced), Cyphokentia macrostachya (Arecoideae: Areceae: Clinospermatinae) and Tectiphiala ferox (Arecoideae: Areceae: Oncospermatinae) as outgroup (Norup et al., 2006; Dransfield et al., 2008; Baker et al., 2009). For DNA extraction, total genomic DNA extracted from silica gel-dried leaf materials (Chase & Hills, 1991) using the 2 CTAB method of Doyle & Doyle (1987). DNA were precipitated with 100% ethanol at 20 C, purified by cesium chloride/ethidium bromide gradients (1.55 g/ml) followed by dialysis and removal of ethidium bromide using butanol. Primer sequences for PRK and RPB2 were obtained from published sources (Lewis & Martinez, 2000; Lewis & Doyle, 2002; Roncal et al. 2005; Loo et al. 2006; Norup et al. 2006; Trenel et al., 2007; Cuenca et al., 2007). The primers for PRK (prk717f: 5 -GTG ATA TGG AAG AAC GTG G-3, prk969r: 5 -ATT CCA GGG TAT GAG CAG C-3 ) have been designed to be specific to the smaller of the two paralogues known in the palms (Loo et al., 2006) The primers for RPB2 (RPB2-PALM-INT23F: 5 -CAA CTT ATT GAG TGC ATC ATG G-3, RPB2-PALM-INT23R: 5 -CCA CGC ATC TGA TAT CCA C-3 ) are specific to palms. Each PCR product was amplified using a 25 µl reaction mix consisting of 1.1 ReddyMix PCR Master Mix (2.5mM MgCl 2, ABgene), 0.3 µl of each primer (10mM, final concentration), 1 5µl of template DNA, 0.9 µl bovine serum albumin (BSA) and optional dimethyl sulfoxide (DMSO) at 1% for recalcitrant amplifications of PRK and RPB2. Reaction mixtures will be subjected to the following temperature profile: initial denaturation at 95 C for 5 min, 38 cycles of 96 C at 1 min each, 50 C for 1 min and 72 C for 1 min, and a final extension at 72 C for 5 min. Amplification products were clean using the QIAquick PCR Purification Kit. The cleaned PCR products were cycle sequenced using the PRISM BigDye Terminator Cycle Sequencing Ready Reaction Kit (Applied Biosystems: ABI) following the manufacturer s protocols. Amplification primers will use as sequencing primers. Subsequent cycle sequencing protocols and purification of cycle sequencing 9

30 HEATUBUN ET AL.: PHYLOGENY OF ARECA products follow that of Asmussen and Chase (2001) and Salazar et al. (2003). The sequences were run on an automated sequencer (ABI). Sequence alignment follow Loo et al. (2006) and Norup et al. (2006) which every base position in the reverse and forward sequences were check and assembled using Sequencher 4.1 (Gene Codes Corp, Ann Arbor, Michigan, USA) and will be deposited in GenBank. The sequences then enter and aligned manually into data matrices in PAUP* version 4b10 (Swofford, 2002) for Macintosh computer. Variable positions in the alignment will verified against the raw data to ensure that these will not a result of base-calling errors. Phylogenetic analyses in this study were using parsimony analysis and Bayesian analysis. the standard procedures as follow: congruence between the PRK and RPB2 datasets will evaluated using the incongruence length difference (ILD) test of Farris et al. (1994) as implemented in PAUP* (Swofford 2002) and Mr. Test in MrBayes version 3.0b4 (Huelsenbeck & Ronquist, 2001). Character partitions were designated in the combined dataset and analyzed in replicates, each with a single heuristic search, saving trees with each replicate. The two dataset partitions were analyzed separately and simultaneously. Taxon sampling varied between the PRK and RPB2 datasets, and therefore the combined dataset included only taxa for which both gene sequences were available. For parsimony analyses, uninformative characters and ambiguously aligned regions were excluded. All included characters were unordered and equally weighted. Initial analyses employed 1000 heuristic searches, each with starting trees obtained by random taxon addition, tree-bisection-reconnection (TBR) swapping, and keeping multiple trees per step (MulTrees on). Only groups that were found in the strict consensus tree and 50% or more of the replicates were recorded. Bayesian analyses were carried out using the program MrBayes version 3.0b4 (Huelsenbeck & Ronquist, 2001). Models of sequence evolution that the best fit the individual datasets were determined using Mr. Test. The models were evaluated by the Akaike information criterion (Akaike, 1973) implemented in the program. Parameters based on patterns in the data matrices. For all three datasets, four incrementally heated Markov chains were used in an analysis that was run for 100,000 generations initially with trees saved at every 10 th generations. Trees produced prior to stationarity were discarded as the burn-in. 10

31 HEATUBUN ET AL.: PHYLOGENY OF ARECA Table 1. Statistics calculated from pasimony analyses of PRK, RPB2 and combined datasets. Analysis Number of taxa Alignment length Ambigously aligned characters Parsimony Informative Characters MPTs Tree length CI RI RC Nodes in strict consensus Nodes with >50% bootstrap support PRK RPB Nodes with >90% bootstrap support Combined (PRK: 55; RPB2: 72)

32 HEATUBUN ET AL.: PHYLOGENY OF ARECA RESULTS Results of the phylogenetics analysis of the genus Areca based on the molecular data nuclear DNA; PRK, RPB2 and combined as shown in the Table 1 and Figs 1 3. PRK Analysis In the PRK dataset, there were 54 sequences of 36 taxa, including 34 sequences from 20 species of Areca available. The average length of the genes is about 700 bp in the Areca and the final length from data matrices is 627 bp and consisting of 23 bp exon 4, all of intron 4 and 130 bp exon 5, with 102 ambiguous characters and 63 parsimonious informative characters (Table 1). In the 5 samples, including A. chaiana was unsuccessful amplified and/or recovered apparently the existence of non-orthologous copies or pseudo-gene, thus all these genes were excluded from the analyses. Overall, parsimony and Bayesian inference analysis of PRK produced well-resolved and strongly supported topologies (Fig. 1). The parsimony bootstrap analysis resulted 300 the most parsimonious trees (MPT) with a length of 91 trees (CI = , RI = ) and recovered 23 nodes over 50% bootstraps supports (one not shown in Fig. 1) including 4 nodes over 90% of bootstraps supported. As previously expected, that the Bayesian inference analysis will be generating more strong supported and more resolved in the PRK analysis than the bootstrap parsimony analysis. Of 25 resolved nodes in the Bayesian majority-rule tree, 21 nodes has value above 90% Bayesian posterior probability support and only 2 nodes below 75%. The Arecinae is strongly resolved (100 Bayesian posterior probability, BPS and 90 bootstraps percentage, BP), whereas the outgroups is unresolved with polytomies (Bentinckia condapanna, Clinostigma savoryanum and Cyphokentia macrostachya) and poorly resolved in Cyrtostachys renda and Tectiphiala ferox (55 BPS and less than 50 BP). The topology supports the monophyly of Arecinae. The monophyly of subtribe Arecinae which contain genera Areca, Nenga and Pinanga is reconfirmed and Areca as sister to Nenga and Pinanga is resolved with strongly supports (100 BPS and 90 BP). And also Nenga is sister to Pinanga resolved with strongly supports in Bayesian (95 BPS) and moderate support in parsimony analysis (70 BP). Areca is strongly supported (100 BPS, 91 BP) with two lineages to define two different clades. The discovery of two lineages within Areca with highly resolution (100 BPS in both, 89 and 55 BP) is indicate the infrageneric classification as proposed before by 12

33 HEATUBUN ET AL.: PHYLOGENY OF ARECA Furtado (1933), two subgenera (subgenus Areca and Beccarioareca). However, the species are included in these two subgenera still need to re-arrangements based on the result of this study. The subgenus Areca is strongly supports in Bayesian (100 BPS) and weak in parsimony analysis (55 BP). In general, the topology is low resolution with polytomies and the relationships among the species still unclear, except to A. triandra clade (A. triandra and A. concinna) with strongly supports (100 BPS, 97 BP), the Philippines congested inflorescence species clade (A. caliso, A. camarinensis and A. ipot) also with strongly supports (100 BPS, 96 BP) and A. oxycarpa (Heatubun 977 and Asmarayani 461). The resolution in subgenus Beccarioareca is better than subgenus Areca, the node is highly resolved (100 BPS, 89 BP) and the relationships among the species in the subgenus more resolved. In this clade is also revealed that Areca furcata is sister to all species in subgenus Beccarioareca with relatively moderate supports (75 BPS, 69 BP). The clade consist A. minuta, A. jugahpunya and A. ridleyana is resolved with high resolution in Bayesian analysis (100 BPS) and moderate in parsimony bootstrap analysis (71 BP). Also within the clade, the relationships between one accession A. minuta (Heatubun 892) and A. ridleyana recovered with strongly supports (100 BPS, 84 BP), whereas another accession of A. minuta (s.n.) and A. jugahpunya still unclear. While, two accession of A. minuta (Heatubun 887 and 888) are not clearly resolved as polytomies together with A. tunku. Areca guppyana (= A. novohibernica) and A. vestiaria forming the Mischophloeus clade (94 BPS, 62 BP) and indicates A. novohibernica is sister to A. vestiaria. This clade is similar to section Mischophloeus of subgenus Beccarioareca in Furtado s infrageneric classification (1933). The other node is resolved with low supports (50 BPS, less than 50 BP) and expressed the relationships between A. insignis var. moorei and A. subacaulis. And this splits from other accession of A. insignis. RPB2 Analysis In RPB2 dataset, 54 sequences were in total from 36 taxa, including 30 sequences of 18 species of Areca were available. The length of the genes around 850 bp in the Areca and the final length from data matrices is only 776 bp of intron 23 with 73 parsimony informative characters. Similarly, about 7 samples unsuccessful, thus those sequences excluded from 13

34 HEATUBUN ET AL.: PHYLOGENY OF ARECA RPB2 analysis and also in the combined analysis. The tree topologies in RPB2, nodes support and resolution performed by the parsimony bootstrap analysis lower than the results of Bayesian analysis. In the analysis produced 3021 most parsimonious trees (MPT) with a tree length 106 (CI = , RI = ). Of 19 nodes resolved in majority rule tree 11 nodes over 90% BP (Fig. 2). Whereas in the Bayesian inference analysis, from 21 nodes resolved in the majority rule tree, 18 nodes exceeding 90% BPS. The monophyly of Arecinae is reconfirmed with strongly supports in Bayesian analyisis (100 BPS) and moderate in parsimony bootstrap analysis (78 BP). From outgroups, only Clinostigma savoyarum and Cyrtostachys renda is resolved with strongly supports (100 BPS, 92 BP), while Bentinckia condapanna, Cyphokentia macrostachya and Tectiphiala ferox are not clearly resolved their relationships. Three lineages have been discovered from Arecinae in this analysis; two lineages (evolutionary lines) are also recovered in PRK analysis which contains Areca line and Nenga- Pinanga line and the third is Areca chaiana line. The topology is supports exclusion of Areca chaiana from Areca and as sister to all genera of Arecinae. Areca is monophyletic and sister to Nenga and Pinanga are reconfirmed with strongly supports (100 BPS, 100 BP). Nenga is sister to Pinanga also reconfirmed from this analysis (90 BPS, 60 BP) and supports the monophyly of each genus in subtribe Arecinae. Two lineages within Areca are also discovered with highly resolution in strongly supports in both analyses Bayesian and parsimony (100 BPS, 99 BP and 100 BPS, 100 BP). Two clades defined as clade of subgenus Areca and clade of Beccarioareca recovered with different resolution in tree topology. The relationships among species within subgenus in Areca are more resolved than Beccarioareca. Subgenus Areca is composed by nine resolved nodes with different resolutions. From those resolved nodes, three major clades have been identified as the congested inflorescence species clade, Areca catechu clade and Areca triandra clade. The congested inflorescence species clade is containing species accession of Areca from The Philippines, Sulawesi and New Guinea. This clade is strong supports in Bayesian analysis (95 BPS) and relatively weak supports in parsimony bootstrap analysis (62 BP), and from topology supports A. ipot as sister to all congested inflorescence species (A. camarinensis, A. caliso, A. macrocalyx and A. oxycarpa). Three accessions of Areca macrocalyx from New Guinea (Baker 1317, Heatubun 787 and 876) defined a resolved clade with strongly supports (100 BPS, 95 BP) and two of 14

35 HEATUBUN ET AL.: PHYLOGENY OF ARECA them (Baker 1317 and Heatubun 876) shown relationships with weak supports (58 BPS, less than 50 BP). While, one accession of A. macrocalyx (Heatubun 798) is resolved separately from A. macrocalyx clade and nested together with Areca oxycarpa (Asmarayani 461 and Heatubun 877) from north Sulawesi with strongly supports (100 BPS, 88 BP). The other clade is resolved with strongly supports (100 BPS, 87 BP) contain the Philippines species of A. caliso and A. camarinensis. Accessions of Areca catechu and A. mandacanii resolved in one clade with strong supports in Bayesian analysis (91 BPS) and unresolved in parsimony analysis (less than 50 BP). Although the relationships is not clear with polytomies especially between A. catechu and A. mandacanii, except to two accessions of A. catechu (Heatubun 751 and 870) resolved (98 BPS, 62 BP), but this result supports Heatubun (2008) that A. mandacanii close to A. catechu based on morphology. As in PRK analysis, A. triandra and A. concinna in RPB2 analysis also resolved in one clade with strongly supports (100 BPS, 96 BP). The relationships of among the accessions of A. triandra and A. concinna are not clearly resolved with polytomies. The clade of subgenus Beccarioareca is low resolution in general and only A. minuta (Heatubun 892) resolved as sister to all species in the clade with relatively moderate supports (76 BPS, 65 BP). Combined Analysis In the combined analyses, 41 sequences of 29 taxa, including 26 sequences of the 17 species of Areca with alignment 1403 bp long, including 102 ambiguous characters and 127 parsimony informative characters. In parsimony analysis, produced 972 most parsimonious tree (MPT) with a tree length 179 (CI = , RI = ) in Table 1. Of 25 resolved nodes in majority-rule tree, 12 nodes over 90% bootstraps support (Fig. 3), and from 27 nodes on Bayesian inference analysis 23 nodes over 90%. In general, the topology of tree in combined analysis is more resolved than previous analysis of each gene PRK and RPB2. Increasing the resolutions and supports in the tree topologies of the phylogenenetic tree can be carrying out with combine the PRK and RPB2. By combining these two genes in this analysis, is to maximize the informative characters (as phylogeny characters also) in both genes, so that the number of informative characters will increase and improve the topology of the phylogeny tree that produced. 15

36 HEATUBUN ET AL.: PHYLOGENY OF ARECA FIG. 1. The 50% majority rule consensus tree of the Bayesian inference analysis of PRK dataset. Bayesian probability support values are shown above the brach, Parsimony bootstraps support values are shown below the branch. An asterisk (*) indicates a branch recovered with less than 50% bootstraps support. Infrageneric classifications within Areca (sensu Furtado, 1933) are indicated to the right of the species names. 16

37 HEATUBUN ET AL.: PHYLOGENY OF ARECA FIG. 2. The 50% majority rule consensus tree of the Bayesian inference analysis of RPB2 dataset. Bayesian probability support values are shown above the brach, Parsimony bootstraps support values are shown below the branch. An asterisk (*) indicates a branch recovered with less than 50% bootstraps support. Infrageneric classifications within Areca (sensu Furtado, 1933) are indicated to the right of the species names. 17

38 HEATUBUN ET AL.: PHYLOGENY OF ARECA FIG. 3. The 50% majority rule consensus tree of the Bayesian inference analysis of combined (PRK and RPB2) dataset. Bayesian probability support values are shown above the brach, Parsimony bootstraps support values are shown below the branch. An asterisk (*) indicates a branch recovered with less than 50% bootstraps support. Geographical regions are indicated to the right of the species names and followed by Infrageneric classifications within Areca (sensu Furtado, 1933). 18

39 HEATUBUN ET AL.: PHYLOGENY OF ARECA The monophyly of subtribe Arecinae, the monophyly of Areca and it position as sister to Nenga and Pinanga, the monophyly of Nenga and Pinanga and also Nenga is sister to Pinanga are reconfirmed (Loo et al., 2006; Norup et al., 2006; Baker et al., 2009). Including resolved the relationships between two outgroups Clinostigma savoryanum and Cyrtostachys renda with relatively strong supports in Bayesian analysis (86 BPS) and moderate in parsimony analysis (65 BP), this result similar to Norup et al. (2006) and slightly different with Baker et al. (2009), which Cyrtostachys renda is sister to Clinostigma savoryanum and Bentinckia condapanna. While, Bentinckia condapanna not resolved in this analysis. Two lineages are discovered within Areca with strongly supports (100 BPS, 100 BP of each node) sequence of A. chaina excluded from the combined analysis. Topologies are more clearly explained the relationships among the taxa in the clades, especially after incorporate with subgenera and sections sensu Furtado (1933) and their regions. Subgenus Areca contains with three distinct clades and eight resolved nodes but A. dransfieldii and A. rheophytica still unresolved (less 50 BPS and BP). While, the clade of the congested inflorescence species of Areca from the Philippines, Sulawesi (Celebes), Moluccas, New Guinea and Solomon Islands resolved with strongly supports in Bayesian analysis (100 BPS) and moderate in parsimony analysis (76 BP). In this clade, is also discovered three evolutionary lines based on the geographic regions with highly resolution: the Philippines line (100 BPS, 81 BP) contains A. caliso, A. camarinensis and A. ipot; Moluccas-New Guinea- Solomon line (100 BPS, 98 BP) contains two accessions of A. macrocalyx (Baker 1317 and Heatubun 876); and Sulawesi line (100 BPS, 95 BP) contains two accessions of A. oxycarpa (Asmarayani 461 and Heatubun 877). And the Philippines line revealed that A. ipot is sister to A. caliso and A. camarinensis. Areca catechu clade is resolved in high supports in Bayesian (81 BPS) but unresolved in parsimony bootstraps analysis (less than 50 BP). Similar to RPB2 analysis, the topology is low resolution with polytomies, except to two accessions of A. catechu from New Guinea (Heatubun 751 and 870) resolved (100 BPS, 60 BP). The relationships between all accessions of A. catechu and A. mandacanii are not clearly resolved however still indicated these two species are closely related (Heatubun, 2008). Areca triandra clade is also resolved with strongly supports (100 BPS, 100 BP) including A. concinna in the clade. The relationships among the accession of A. triandra and A. concinna is still not clearly resolved with polytomies. 19

40 HEATUBUN ET AL.: PHYLOGENY OF ARECA Subgenus Beccarioareca is resolved with highly resolution (100 BPS, 100 BP) and A. furcata from Borneo is sister to all taxa in the clade. Two lineages also discovered (89 BPS, 65 BP); one nodes is resolved with relatively high supports in Bayesian (80 BPS) and collapsed in parsimony analysis (less than 50 BP), which forming a clade with contains A. guppyana (= A. novohibernica), A. subacaulis and accessions of A. vestiaria (Heatubun 879 and 885). The other node resolved with highly supports (100 BPS, 80 BP) and contains A. jugahpunya and two accessions of A. minuta (Heatubun 892 and s.n.). Areca subacaulis from Borneo is resolved (96 BPS, 66 BP) as sister to Mischopleous species of A. guppyana (= A. novohibernica) from off-shore islands of Papua New Guinea (Manus Island, Bismarck Archipelago and Solomon Islands) and A. vestiaria from Sulawesi and North Moluccas. This result is never expecting before. Areca guppyana (= A. novohibernica) is also resolved as sister to two accessions of A. vestiaria (Heatubun 879 and 885) with strong supports in Bayesian analysis (99 BP) and relatively moderate in parsimony bootstraps analysis (65 BP). The morphological characters also support this relationship (Furtado, 1933; Heatubun et al., inprep.). While, the relationships among Bornean species of A. jugahpunya and two accessions of A. minuta still not clearly resolved with polytomies, although the clade is strongly supports. DISCUSSION Phylogenetic value of PRK and RPB2 The phylogenetic utility of the low-copy nuclear gene PRK and RPB2 have been proved useful in palm phylogeny (Lewis & Martinez 2000; Gunn, 2004; Norup 2004; Bayton 2005; Roncal et al. 2005; Loo et al., 2006; Norup et al., 2006; Thomas et al. 2006; Trenel et al., 2007; Cuenca et al., 2007; Baker et al., 2009). The level of variation in PRK and RPB2 is higher than other plastid genes used so far already discussed by Loo et al. (2006) and Norup et al. (2006), including the comparison between number of parsimony informative characters obtained from the specific DNA regions, number of taxa sampled in the studies and the level of homoplasious characters gained from the analyses. Despite from the limited number of samples used in study, percentages of parsimonyinformative aligned positions for each partition in combined analysis (10.5% for PRK, 9.3% for RPB2) still exceeded from five plastid DNA regions used by (Hahn, 2002), which range 20

41 HEATUBUN ET AL.: PHYLOGENY OF ARECA 2.1 9% and still in the top position of the range 6 10% three plastid markers used by Asmussen & Chase, 2001). However, consistency and retention indices are high indicating low level of homoplasious characters in this study. And influence the resolution in phylogeny tree by providing the sufficient characters to resolve relationships among taxa of Areca and outgroups using on the study. In this study has identified several new relationships and reconfirmed relations in the genus Areca addressed by morphology before (Furtado, 1933; Dransfield, 1984; Harley & Dransfield, 2003; Heatubun, 2008). We also discovered the compatibility of each region PRK and RPB2 to specific lineages in this palm genus, and by combining them, PRK and RPB2 will resolving the relationships in the lower taxonomic level such as species. Morphology In morphologically, these two lineages within Areca are supports by differences of the sepals of staminate flower and arrangement of the staminate flowers on the rachillae; the sepals being free and the staminate flowers uniseriate and distichous in subgenus Areca versus the sepal being united or calyx cup shaped and all or small portion of the staminate flowers spiral in subgenus Blumeoareca. Whereas, in the sectional level is difficult to addressed, due to homoplasious morphological characters occurs. The morphological characters used to define sections in the previous classification (Furtado, 1933) have been evolved several times independently and separately in the genus Areca. For example, the small or even stem less habit was used to recognised the section Microareca in subgenus Beccarioareca (Furtado, 1933) also occurs in subgenus Areca A. ahmadii (Dransfield, 1984) and in our result A. subacaulis nested together with A. guppyana (= A. novohibernica) and A. vestiaria of Furtado s section Mischopleous. Areca chaiana in PRK analysis (unsuccessful sequence in RPB2, excluded in combined) is sister to all genera of subtribe Arecinae, this taxon from morphological point of view is very distinctive and different from the generic concept of the genus Areca. Dransfield (1984) said that this is aberrant species of Areca when he was described it, because the nature of it inflorescence being spicate. More over, incomplete zonosulcate pollen aperture and tectum perforate reticulate are unusual in Areca were found in this taxon (Harley & Dransfield, 2003). 21

42 HEATUBUN ET AL.: PHYLOGENY OF ARECA Systematics implications Species of Areca that has been include to the subgenus and section from previous classification (Furtado, 1933; Dransfield, 1984; Harley & Dransfield, 2003; Heatubun, 2008) and applied to the phylogeny tree in combined analysis (Fig. 1 3). In general, this result is support infrageneric classification sensu Furtado (1933); two subgenera, namely Areca and Beccarioareca, except to A. jugahpunya and A. tunku which were included in subgenus Areca (Blumeoareca) section Arecella (Dransfield, 1984), whereas, in this result joins the clade defined as the Beccarioareca clade. Similar situation also found in sectional level within subgenera. In subgenus Areca, A. oxycarpa was in section Areca (Furtado, 1933) but in our results nested together with A. caliso, A. camarinensis, A. ipot and A. macrocalyx from section Axonianthe (Furtado, 1933). In previous infrageneric classification (Furtado, 1933), A. concinna belongs to section Areca and nested together with A. triandra (section Arecella). Subgenus Beccarioareca has similar problem, which A. subacaulis (section Microareca, Dransfield, 1984) as sister to A. guppyana (= A. novohibernica) and A. vestiaria of section Mischopleous (Furtado, 1933). Although the limitation of samples used and unresolved branching nodes (polytomies) in topology of the phylogenetics tree, in fact, the clades grouping of our results are different from sectional grouping recognised in previous infrageneric classification (Furtado, 1933; Dransfield 1984; Harley & Dransfield, 2003), thus it will bring a taxonomic consequence, in which, we do not recommend to use the sectional classification within the genus Areca for the future application. We also strongly recommend excluding A. chaiana J. Dransf. from the genus Areca to proper palm genus. Biogeography From the topology of the phylogenetics tree (Fig. 3), suggests that the present distribution Areca is mainly caused by the long dispersal event rather than vicariance. And it might be, the dispersal events take place in several times dispersed out from centre of origin to recent area of distribution. Also from the tree topology, there is no indication of bimodal distribution pattern exist between west and east Wallace s line in this genus and supports previous results from subtribe Arecinae (Loo et al., 2006). In our results, suggests Areca is 22

43 HEATUBUN ET AL.: PHYLOGENY OF ARECA origin from Borneo, shown clearly in the subgenus Beccarioareca, which A. furcata from Borneo is sister to other species in the subgenus, including east of Wallace s line species A. vestiaria and A. guppyana (= A. novohibernica). Confirmation of Borneo as the centre of origin for Areca is straight forward, however, Norup et al. (2006) was included Areca in Indian Ocean clade, and Loo et al. (2006) in the Sunda shelf as possible centre of origin from previous study in tribe Areceae and subtribe Arecinae. Of 20 from 41 species in the recent revision of the genus Areca (Heatubun et al., in prep.) are origins from Borneo and supports the idea of the centre of origin. Loo et al. (2006) explained the diversification of the subtribe Arecinae (Areca, Nenga and Pinanga) is associated with the long, geologically stable history of the Sunda shelf and its ecological condition during Pliocene and Quaternary as proposed by Morley (1999). Furthermore, they also noted that palms were widespread in South-East Asia throughout much of Tertiary, and the subtribe Arecinae resembles the remnant of the early palm flora on that era. That evidence could be the explanation for distribution of A. triandra, this species distributed from India, Indochina, Thailand, Malay Peninsula, Sumatra, Java and Borneo. Biogeographycal relationships between the west and east Wallace s line taxa is clearly demonstrated in subgenus Beccarioareca. Explanations about how the species of Areca crossed the Wallace s line and reached islands of Sulawesi, the Philippines, New Guinea and the Solomon in the Pacific; as proposed by Hall (1998: 115); in the mid Oligocene, the southern-most promontory was the Sulawesi-Philippines-Halmahera arc which could have provided a pathway into the Pacific, via volcanic island stepping stone, for organisms that could cross sea water, and the other promontories terminated in the deep ocean area of the Pacific. Further molecular dated phylogeny analysis is needed to explain and/or reconstruct the dispersal events of Areca species in this region, including the diversification of the genus Areca and the species Diaspora from the centre of origin in Borneo to westward and eastward cross the Wallace s line, like similar study conducted by Gunn (2004) in Cocoeae with special emphasis on Cocos nucifera, Trenel et al. (2007) in the wax palm sub family Ceroxyloideae and Cuenca et al. (2007) in tribe Chamaedoreeae. Since the discovery of A. mandacanii (Heatubun, 2008), the country of origin of betel nut palm (A. catechu) become more and more blur this also indicates in our results. As previously suggested, the betel nut palm might be come from the Philippines (Beccari, 1919; Furtado, 1933) and Malaysia (Corner, 1966; Jones, 1995) or Celebes (Corner 1966) and in 23

44 HEATUBUN ET AL.: PHYLOGENY OF ARECA New Guinea by now. The suggestions above based observation on the wild species that morphologically close related to the betel nut. However, more samples are needed and must be include wild species from the Philippines in the analysis to try to solve the origin of betel nut. ACKNOWLEDGEMENTS Many individuals and institutions contributed to the completion of this paper. We would like to thank Drs Christine Bacon, Raymond Baker and Carl Lewis to provided DNA materials. The authority in Sarawak, especially the Sarawak Forestry Company (SFC) was allowed CDH to conduct his fieldtrip in Sarawak. Julia Sang and Shahabuddin M. Shabki are thanked for their helps with permit and all supports including organized CDH s fieldtrip in Sarawak. This paper is part of CDH PhD thesis which conducted at Institut Pertanian Bogor and Royal Botanical Gardens Kew. CDH would like to express his gratitude to Dr. Sri S. Tjitrosoedirdjo, Prof. Johanis P. Mogea, and Prof. Mien A. Rifai to their generous encouragements, supports, helps and guidance during the study. And the financial supports came from International Palm Society to funding CDH s field trip to Sarawak through IPS Endowment Fund The Royal Botanic Gardens Kew UK, BAT Biodiversity Partnerships and the Indonesian Ministry of Education (BPPS Dikti Diknas) to funding CDH s PhD, to them are greatly acknowledged. LITERATURE CITED Asmussen, C. B. and M. W. Chase Coding and non-coding plastid DNA in palm systematics. Amer. J. Bot. 88: Baker, W. J., C. B. Asmussen, M. W. Chase, J. Dransfield, F. Forest, M. M. Harley, V. Savolainen, N. W. Uhl, M. Wilkinson Complete generic level phylogenetic analyses of palms (Arecaceae) with comparisons of Supertree and Supermatrix approaches. Systematic Biology. doi: /sysbio/syp021 24

45 HEATUBUN ET AL.: PHYLOGENY OF ARECA Barrow, S A monograph of Phoenix L. (Palmae: Coryphoideae). Kew Bull. 53: Bayton, R. P Borassus L. and the Borassoid palms: systematics and evolution. Ph.D. Thesis, University of Reading. Beccari, O The palms of Philippines Islands. Phillipp. J. Sci. 14: Byun, S. J., H. S. Kim, S. M. Jeon, Y. B. Park, and S. M. Choi Supplementation of Areca catechu L. extract alters triglyceride absorption and cholesterol metabolism in rats. Annals of Nutrition and Metabolism 45: Chase, M. W. and H. H. Hills Silica gel: an ideal material for field preservation of leaf samples for DNA studies. Taxon 40: Cuenca A, Asmussen-Lange CB, Borchsenius FB A dated phylogeny of the palm tribe Chamaedoreeae supports Eocene dispersal between Africa, North and South America. Molecular Phylogenetics and Evolution 46: Doyle, J. D. and J. L. Doyle A rapid DNA isolation procedure from small quantities of fresh leaf tissue. Phytochem. Bull. Bot. Soc. Am. 19: Dransfield, J The genus Areca (Palmae: Arecoideae) in Borneo. Kew Bull. 39: Dransfield, J A guide to collecting palms. Annals of the Missouri Botanical Garden 73: Dransfield, J Species and species concepts in old world palms. Mem. New York Bot. Gard 83: Dransfield, J. and N.W. Uhl An outline of classification of palms. Principes 30: Dransfield, J. and N. W. Uhl Palmae. In: Kubitzki, K (Ed.), the families and genera of vascular plants, vol. IV. Springer, Berlin, pp Dransfield, J., N. W. Uhl, C. B. Asmussen, W. J. Baker, M. M. Harley and C. E. Lewis A new phylogenetic classification of the palm family, Arecaceae. Kew Bull. 60: Dransfield, J., N. W. Uhl, C. B. Asmussen, W. J. Baker, M. M. Harley and C. E. Lewis Genera Palmarum: The evolution and classification of palms. Royal Botanic Gardens, Kew. Farris, J. S., M. Kallersjo, A. G. Kluge, C. Bult Testing significance of congruence. Cladistics 10:

46 HEATUBUN ET AL.: PHYLOGENY OF ARECA Felsenstein, J Confidence limits on phylogenies: an approach using the bootstrap. Evolution 39: Flynn, T Morphological variation and species limits in the genus Areca (Palmae) in New Guinea and the Solomon Islands. Unpublished Master thesis, University of Wales, Bangor. Furtado, F. X The limits of the genus Areca L. and its sections. Repert. Spec. Nov. Regni Veg. 33: Govaerts, R., and J. Dransfield World checklist of palms. Royal Botanic Gardens, Kew. Gunn BF The phylogeny of Cocoeae (Arecaceae) with emphasis on Cocos nucifera. Annals Missouri Botanical Garden 91: Gupta, P. C., and S. Warnakulasuriya Global epidemiology of Areca nut usage. Addict. Biol. 7: Hahn, W. J. 2002a. A molecular phylogenetic study of the Palmae (Arecaceae) based on atpb, rbcl and 18S nrdna sequences. Syst. Biol. 51: Hahn, W. J. 2002b. A phylogenetic analysis of the Arecoid line of palms based on plastid DNA sequence data. Mol. Phylogenet. Evol 23: Harley, M. M. and J. Dransfield Triporate pollen in the Arecaceae. Grana 42: Heatubun, C. D A new species of Areca from Western New Guinea. Palms 52: Huelsenbeck, J. P. and F. Ronquist MrBayes: Bayesian inference of phylogenetic tree. Bioinformatics 17: IUCN. (2001). IUCN red list categories and criteria. Version 3.1. Gland & Cambridge: IUCN. Jones, D. L Palms throughout the world. Reed Books. Lee, K. K., and J. D. Choi Areca catechu L. extract. I. effect on elastase and aging. Journal of Cosmetics Science 49: Lewis, C. E. and N. Martinez Identity of the Hyophorbe palms at the botanical garden of Cienfugoes, Cuba. Palms 44: Lewis, C. E., and J. J. Doyle Phylogenetic utility of the nuclear gene malate synthase in the palm family (Arecaceae). Mol. Phylogenet. Evol. 19:

47 HEATUBUN ET AL.: PHYLOGENY OF ARECA Lewis, C. E., and J. J. Doyle A phylogenetic analysis of tribe Areceae (Arecaceae) using two low-copy nuclear genes. Plant Syst. Evol. 236: Loo, A. H. B., J. Dransfield, M. W. Chase and W. J. Baker Low copy nuclear DNA, phylogeny and the evolution of dichogamy in the betel nut palms and their relatives (Arecinae; Arecaceae). Mol. Phylogenet. Evol. 39: Mayden, R. L A hierarchy of species concept: the denouement in the saga of the species problem. In: Claridge, M. F., H. A. Dawah & M. R. Wilson (Eds.). Species: the unit of biodiversity. New York, pp Moore, H. E., Jr. and J. Dransfield Tipification of Linnean palms. Taxon 28: Morley, R. J Origin and evolution of tropical rainforest. Chicester: Wiley. Norton, S. A Betel: consumption and consequences. J. Am. Acad. Dermatol. 38: Norup, M.V A molecular systematic study of Heterospathe and Rhopaloblaste (Arecaceae, Areceae). Masters Thesis, University of Aarhus. Norup, M. V., J. Dransfield, M. W. Chase, A. Barfod, E. Fernando, and W. J. Baker Homoplasious character combinations and generic delimitation: A case study from the Indo-pacific Arecoid palms (Arecaceae; Arecoideae). Am. J. Bot. 93: Ray, A. K., and D. V. S. Reddy Performance of Areca-based high density multi species cropping system under different level of fertilizer. Tropical Agriculture 78: Roncal, J., J. Francisco-Ortega, C. B. Asmussen, and C. E. Lewis Molecular phylogenetics of the tribe Geonomeae (Arecaceae) using nuclear DNA sequences of phosphoribulokinase and RNA polymerase II. Syst. Bot. 30: Salazar, G.A., Chase, M.W., Soto, M.A., Ingrouille, M Phylogenetics of Cranichideae with emphasis on Spiranthinae (Orchidaceae, Orchidoideae): Evidence from plastid and nuclear DNA sequences. Am. J. Bot. 90: Sullivan, R. J., J. S. Allen, C. Otto, J. Tiobech, K. Nero Effects of chewing betel nut (Areca catechu) on the symptoms of people with schizophrenia in Palau, Micronesia. British Journal of Psychiatry 177: Swofford, D. L PAUP*. Phylogenetic Analysis Using Parsimony (* and Other Methods). Version 4b10. Sinuer Associates, Sunderland, MA. 27

48 HEATUBUN ET AL.: PHYLOGENY OF ARECA Thomas, M. M., N. C. Garwood, W. J. Baker, S. Henderson, S. J. Russell, D. R. Hodel, and R. M. Bateman Molecular phylogeny of the palm genus Chamaedorea, based on the low-copy nuclear genes PRK and RPB2. Mol. Phylogenet. Evol. 38: Trenel P, Gustafsson MH, Baker WJ, Asmussen-Lange CB, Dransfield J, Borchsenius F Mid-Tertiary dispersal, not vicariance explains Gondwanan distribution pattern in wax palm subfamily (Ceroxyloideae: Arecaceae). Molecular Phylogenetics and Evolution 45: Uhl, N. W. and J. Dransfield Genera Palmarum. Allen Press, Lawrence, Kansas. Uhl, N. W. and J. Dransfield Genera Palmarum after ten years. Mem. New York Bot. Gard. 83:

49 A Conspectus of the Genus Areca L. (Arecaceae) CHARLIE D. HEATUBUN FLS 1, 2, 3, 5, WILLIAM J. BAKER 3, EDWINO S. FERNANDO 4 and JOHN DRANSFIELD 3 1 Faculty of Forestry, Universitas Papua, Jl. Gunung Salju, Amban, Manokwari 98314, Papua Barat, Indonesia 2 Biology Department, Sekolah Pascasarjana Institut Pertanian Bogor, Darmaga, Bogor 16680, Jawa Barat, Indonesia 3 Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, United Kingdom 4 Department of Forest Biological Sciences, College of Forestry and Natural Resources, The University of The Philippines Los Banos College, 4031 Laguna, The Philippines 5 Author for correspondence (charlie_deheatboen@yahoo.com) Summary. This conspectus provides the latest taxonomic revision of the genus Areca. The genus comprises two subgeners (subgenus Areca and Beccarioareca) and 41 species, including seven newly described species, namely Areca bakeri Heatubun, Areca churchii Heatubun, Areca dransfieldii Heatubun, Areca gurita Heatubun, Areca mogeana Heatubun, Areca riparia Heatubun and Areca triginticollina Heatubun. The distinctive species, Areca chaiana J. Dransf. is considered as a species incertae sedis. A determination key is presented to subgenera and also detailed descriptions provided for the new species. Distribution, ecology and conservation status of each new species are also discussed. ADDITIONAL KEYWORDS: palm family Areca Arecaceae Arecoideae Areceae Arecinae taxonomy nomenclature infrageneric classification species.

50 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 31 INTRODUCTION Areca is interesting and important genus in the palm family Arecaceae, not only for its historical background as the first genus published for the palm family but also for its diversity, natural history and economic importance. Taxonomy of this genus has been dynamic, in accordance with the development of different generic concepts (Linneaus, 1753; Scheffer, 1876; Beccari, 1877; Furtado, 1933; Monteiro, 1976; Dransfield, 1984), the infrageneric classification (Furtado, 1933) and the species concepts (e.g. Dransfield 1974). Although the genus Areca is well circumscribed (Dransfield, 1984; Dransfield et al., 2008) and the relationship with other genera of palm is better understood (Loo et al., 2006; Norup et al., 2006; Baker et al., 2009), the infrageneric classification still needs reassessment (Dransfield, 1984) using modern methods and to accommodate the new taxa that have been discovered on recent fieldtrips (e.g. Heatubun, 2008). Furthermore, the phylogenetic concept has been widely used in plant systematics to construct the classification, in which the classification must reflect evolutionary processes and the relationships of plant taxa, such as the recent classification of the palm family (Dransfield et al., 2005, 2008). The last infrageneric classification of Areca was proposed by Furtado (1933), and he tried to establish the relationships among the species within the genus based on morphological similarities. Two subgenera (Subgenus Blumeoareca Furtado and Beccarioareca Furtado), five sections (Arecella Wendl. et Drude, Oeotheanthe (Scheff.) Furtado, Axonianthe Scheff., Microareca Furtado and Mischophloeus (Scheff.) Becc.) were published. Furtado (1933) also provided an enumeration of species under his subgenera and sections. The genus Areca is widely distributed from India, Sri Lanka, and south China through Malesia to New Guinea and the Solomon Islands (Dransfield et al., 2008) and several taxonomic monographs have been produced for each diversity hotspot (Beccari, 19 has been published (Govaerts & Dransfield, 2005). All these previous taxonomic works provide a basis to do a revision of the entire genus, incorporating the results of the molecular phylogenetic analysis (Heatubun et al., in prep.).

51 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 32 MATERIALS AND METHODS Taxon sampling and observations for morphology and distribution of the species were based on herbarium samples or specimens (dried and spirit-preserved materials) deposited at international herbaria, namely A, AAU, B, BH, BO, BRI, FI, K, KEP, L, LAE, MAN, PNH, SAN, SAR, and SING (herbarium acronyms follow Holmgren et al., 1990), as well as the newly established small herbarium in Balai Penelitian Kehutanan (Forestry Research Institute) in Manokwari, West Papua, Indonesia that will be appear in specimens citations as BPK Manokwari. Many new specimens have been collected recently by the author, particularly from western New Guinea (the Indonesian Province of Papua and West Papua), North Sulawesi and Sarawak by the first author in Specimens were made in the field using standard preparation guidelines to the palms (Dransfield, 1986; Baker & Dransfield, 2006). Measurements were taken from spirit-preserved material and dried herbarium specimens and from cultivated individuals. Floral parts were measured from spirit-preserved material or dried specimens and rehydrated by boiling. Basic morphological characters such as habit, stem, leaves, inflorescence, staminate flower, pistillate flower, fruit, seed and their details were used to describe and recognize taxa. All morphological data were used for producing the description of each new species of Areca, while the key to subgenus and species were constructed from the diagnostic characters only. The morphological species concept or taxonomic species concept was applied to recognise species in the genus Areca, while, the infrageneric classification was constructed based on the monophyletic concept on the phylogeny tree which generated from two lower copy nuclear genes PRK and RPB2 (Heatubun et al. in prep.).

52 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 33 Table 1. Comparison of previously recognised taxa and infrageneric classification in Areca with taxa and infrageneric classification recognised in this conspectus. Synonyms are shown in italics. Previous accepted infrageneric classification and taxa of Areca (Furtado, 1933; Dransfield, 1984; Harley & Dransfield, 2003; Govaerts & Dransfield, 2005) I. Subgenus Areca (Blumeoareca Furtado): A. Section Arecella Wendl. et Drude: 1. Areca ahmadii J. Dransf. 2. Areca hutchinsoniana Becc. 3. Areca jugahpunya J. Dransf. 4. Areca laosensis Becc. 5. Areca montana Becc. 6. Areca rheophytica J. Dransf. 7. Areca triandra Roxb. ex Buch. Ham. 8. Areca tunku J. Dransf. & C. K. Lim 9. Areca vidaliana Becc. B. Section Axonianthe Scheff.: 10. Areca caliso Becc. 11. Areca camarinensis Becc. 12. Areca congesta Becc. 13. Areca ipot Becc. 14. Areca jobiensis Becc. 15. Areca ledermanniana Becc. 16. Areca macrocalyx Zipp. ex Blume 17. Areca multifida Burret 18. Areca nannospadix Burret 19. Areca nigasolu Becc. 20. Areca rechingeriana Becc. 21. Areca rostrata Burret 22. Areca torulo Becc. 23. Areca warburgiana Becc. C. Section Oetheanthe Scheff.: 24. Areca catechu L. 25. Areca celebica Burret 26. Areca concinna Thwaites 27. Areca costulata Becc. 28. Areca kinabaluensis Furtado 29. Areca macrocarpa Becc. 30. Areca oxycarpa Miq. 31. Areca parens Becc. 32. Areca whitfordii Becc. II. Subgenus Beccarioareca Furtado: D. Section Microareca Furtado: 33. Areca abdulrahmanii J. Dransf. 34. Areca andersonii J. Dransf. 35. Areca arundinacea Becc. 36. Areca brachypoda J. Dransf. 37. Areca chaiana J. Dransf. 38. Areca dayung J. Dransf. 39. Areca furcata Becc. 40. Areca insignis (Becc.) J. Dransf. Infrageneric classification and taxa of Areca accepted in this conspectus I. Subgenus Areca: 1. Areca ahmadii J. Dransf. 2. Areca bakeri Heatubun sp. nov. 3. Areca caliso Becc. 4. Areca camarinensis Becc. 5. Areca catechu L. Areca macrocarpa Becc. 6. Areca concinna Thwaites 7. Areca costulata Becc. 8. Areca dransfieldii Heatubun sp. nov. 9. Areca hutchinsoniana Becc. 10. Areca ipot Becc. 11. Areca kinabaluensis Furtado 12. Areca laosensis Becc. 13. Areca macrocalyx Zipp. ex Blume Areca jobiensis Becc. Areca rechingeriana Becc. Areca nigasolu Becc. Areca torulo Becc. Areca warburgiana Becc. Areca nannospadix Burret Areca rostrata Burret Areca multifida Burret 14. Areca mandacanii Heatubun 15. Areca montana Ridl. 16. Areca oxycarpa Miq. Areca celebica Burret 17. Areca parens Becc. 18. Areca rheophytica J. Dransf. 19. Areca riparia Heatubun sp. nov. 20. Areca triandra Roxb. ex Buch.-Ham. 21. Areca vidaliana Becc. 22. Areca whitfordii Becc. II. Subgenus Beccarioareca Furtado: 23. Areca abdulrahmanii J. Dransf. 24. Areca andersonii J. Dransf. 25. Areca arundinacea Becc. 26. Areca brachypoda J. Dransf. 27. Areca churchii Heatubun sp. nov. 28. Areca dayung J. Dransf. 29. Areca fucata Becc. 30. Areca gurita Heatubun sp. nov. 31. Areca insignis (Becc.) J. Dransf. var. insignis var. moorei J. Dransf. 32. Areca jugahpunya J. Dransf. 33. Areca klingkangensis J. Dransf. 34. Areca minuta J. Dransf.

53 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 34 var. insignis var. moorei J. Dransf. 41. Areca klingkangensis J. Dransf. 42. Areca minuta Scheff. 43. Areca ridleyana Becc. ex Furtado 44. Areca subacaulis (Becc.) J. Dransf. E. Section Mischophloeus (Scheff.) Becc.: 45. Areca guppyana Becc. 46. Areca novohibernica (Lauterb.) Becc. 47. Areca salomonensis Burret 48. Areca vestiaria Giseke 35. Areca mogeana Heatubun sp. nov. 36. Areca novohibernica (Lauterb.) Becc. Areca guppyana Becc. Areca salomonensis Burret 37. Areca ridleyana Becc. ex Furtado 38. Areca subacaulis (Becc.) J. Dransf. 39. Areca triginticollina Heatubun sp. nov. 40. Areca tunku J. Dransf. & C. K. Lim 41. Areca vestiaria Giseke Species Incertae Sedis: Areca chaina J. Dransf.

54 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 35 TAXONOMIC TREATMENT ARECA L. SP. PL.: 1189 (1753). TYPE: ARECA CATECHU L. Mischophloeus Scheff., Ann. Jard. Bot. Boitenzorg 1: 115, 134 (1876). Gigliolia Becc., Malesia 1: 171 (1877). Pichisermollia H.C. Monteiro, Rodriguesia 28: 195 (1976). Description: Solitary or clustering, very small to moderate, acaulescent to erect, unarmed, pleonanthic, monoecious palms. STEM slender to moderate, occasionally stilt-rooted, internodes very short to elongate, leaf scars often conspicuous. LEAVES undivided and pinnately ribbed, with or without an apical notch, or pinnate; sheaths forming a well defined crownshaft with leaves neatly abscising, or rarely crownshaft not well developed when leaves marcescent or sheaths partly open; petiole present or absent, adaxially channelled or rounded, abaxially rounded, glabrous or variously indumentose; leaflets regularly or irregularly arranged, 1 several fold, acute, acuminate or lobed, the lobes corresponding to folds, the apical pair almost always lobed, held in one plane except to one species (A. mandacanii) in different planes, very rarely (A. insignis) with basal auricles reflexed across the rachis, blade variously scaly or hairly, transverse veinlets obscure. INFLORESCENCE erect or pendulous, mostly infrafoliar, rarely interfoliar in acaulescence species with marcescent leaf sheaths (e.g. A. jugahpunya), branched to 3 orders basally, very rarely spicate, protandrous (or very rarely recorded as protogynous); peduncle very short to long; prophyll thin, membranous, enclosing the inflorescence in the bud, quickly splitting and falling, other bracts very inconspicuous; rachis shorter or longer than the peduncle; rachillae glabrous or variously indumentose; rachilla bracts minute; triads confined to the proximal part of the main axis, or to the proximal part of each order of branching, or rarely to a subdistal part of main axis only; rachillae otherwise bearing solitary or paired staminate flowers arranged spirally, distichously, or in 2 approximate rows on one side of the rachilla, the rachilla tips sometimes devoid of the flowers. STAMINATE FLOWERS frequently minute, sessile, or with a stalk formed from the receptacle; calyx with 3 distinct, slightly imbricate, triangular sepals or cupular with triangular lobes, corolla with 3 triangular, valvate petals, rarely briefly connate at the base, much longer than the sepals; stamens free or briefly epipetalous, 3, 6, 9, or up to 30 or more, filaments short to elongate, anthers linear or sinuous, sometimes very irregular,

55 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 36 lartrorse or rarely opening by apical pores; pollen elliptic or circular, monosulcate with the sulcus short to extensive, trichotomosulcate, monoporate (ulcerate), or triporate, with perforate, foveolate, or finely to coarsely reticulate tectate or intectate exine; pistilode present and conspicuous as trifid coloum as long as the stamens, or minute, or often absent. PISTILLATE FLOWERS sessile, usually much larger than the staminate, ± globular; sepals 3, distinct, imbricate; petals similar to sepals, 3, distinct, sometimes valvate at the very tip, otherwise imbricate; staminodes 3 9 or absent; gynoecium unilocular, uniovulate, globose to ovoid, stigma 3, fleshy, triangular, ± reflexed at anthesis, ovule anatropous, basally attached. RACHILLA distal to pistillate flowers drying after anthesis, portions bearing fruits sometimes becoming brightly coloured. FRUITS globose, ovoid, or spindle-shaped, often brightly coloured, rarely dull brown or green, stigmatic remains apical; epicarp smooth, shiny or dull, mesocarp thin to moderately thick, fleshy or fibrous, endocarp composed of robust longitudinal fibers, usually closely appressed to the seed, becoming free at the basal end or not. SEED conforming to the fruit shape or slightly hollowed at the base, with basal hilum and raphe branches anastomosing, endosperm deeply ruminate; embryo basal. GERMINATION adjacent ligular; eophyll bifid or rarely entire with minute apical cleft. CITOLOGY 2n = 32. Distribution: India and south China, through Malesia to New Guinea and the Solomon Islands. Number of species: 41 species. INFRAGENERIC CLASSIFICATION OF THE GENUS ARECA KEY TO SUBGENUS OF ARECA 1. Staminate flowers uniseriate, or distichous and/or alternate; sepals free or sometimes connected; stamens Subgenus Areca 1. Staminate flowers all or small portions spirally arranged; sepals united; stamens 6 more.subgenus Beccarioareca

56 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 37 The infrageneric classification is mainly based on the result of the molecular phylogenetic analysis of the genus Areca using two low-copy nuclear genes PRK and RPB2 (Heatubun et al. in prep.). The consensus tree from the combined analysis showed that the genus Areca is monophyletic and comprises two lineages with strong support in both Bayesian and Parsimony analysis. Two lineages represent the two subgenera, but the groups (clades) do not correspond with the previous sections proposed by Furtado (1933). This result has taxonomic consequences; we recognise only two subgenera to accommodate the two lineages (evolutionary lines) within the genus and without any sections. After optimization the morphological characters to the phylogenetic tree, only two characters staminate flower arrangement (spiral and secund or distichous), and sepals of staminate flowers (free and united) support the topology of the tree. These characters are used here as diagnostic characters to define two subgenera in Areca subgenus Areca and Beccarioareca. I. Subgenus Areca Blumeoareca Furtado, Repert. Spec. Nov. Regni. Veg. 36: 225 (1933). Type species: A. catechu L. 1. ARECA AHMADII J. DRANSF. Areca ahmadii J. Dransf., Kew Bull. 39: 4 (1984). Type: BORNEO, Sarawak, 1 st Division, Semengoh Forest Reserve, i.1980, Lai Shak Teck S (holotype: K; isotypes: BH, BO, KEP, L, SAR). Distribution: Borneo (Sarawak). 2. ARECA BAKERI HEATUBUN SP. NOV. (FIG.1). Type: BORNEO, Sarawak, cultivated in the Semengoh Forest Arboretum, 42 m, 22.iv.1996, Baker et al. 716 (holotype: K; isotypes: KEP, SING, SAR). Diagnosis: Ad subgenerum Arecam, a ceteris speciebus Borneensibus habitu acaule caespitoso, petiolis longis cm, inflorescentia erecta rigida, floribus staminatis distichiss, floribus foemineis singulariter dispositis distincta.

57 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 38 Description: Undergrowth palmlet densely clustered with c. 20 stems in a clump to 1 m high tall, unarmed. STEM subterranean to very short ( 1 cm), c. 2 diam.; internodes very close (0.5 1 cm long), not conspicuous, covered by marcescent leaf sheaths. LEAVES 4 6 in crown, litter trapping, pinnate, cm long (excluding petiole); sheath tubular, cm long and 3 4 cm wide, smooth, not fibrous, yellowish green; crownshaft well defined, up to 40 cm long and up to 4 cm diam.; petiole cm long, channelled adaxially, rounded abaxially; rachis ascending but not arching, with adaxial longitudinal ridge, rounded abaxially; blade with slightly irregularly arranged leaflets, 7 8 broad on each side; leaflets near petiole c cm, with 7 9 folds, sigmoid, the middle leaflets cm, with 5 9 folds, slightly sigmoid and the terminal leaflets c cm, with up to folds, slightly sigmoid, tips pointed except for the terminal slightly oblique-lobed, papery, darker adaxially than abaxially. INFLORESCENCE infrafoliar, erect, bursting out among marcescent sheaths, cm long and cm wide, protandrous, branching to 1 order; peduncle 6 10 cm long, pale yellowish green; prophyll not available; rachis yellowish green; rachis bracts caducous; rachillae 11 13, cm long and mm wide, very stiff and stout, flattened, pale green, elongate. STAMINATE FLOWERS small, triangular, c mm, asymmetric; sepals 3, low; petals 3, triangular, small, striate; stamens 6, small, anthers shorter than the filaments; filaments slender, elongate; pistilode c mm, pointed. PISTILLATE FLOWERS larger than the staminate, triangular, borne on the enlarged rachillae, only one flower on each rachillae, buds varying greatly in size depending on stage of development, c. 17 mm long and 8 mm wide in late anthesis; sepals 3, strongly imbricate, c. 7 7 mm, triangular, asymmetrical, striate; petals 3, imbricate, triangular, c mm, striate; gynoecium c mm long and 4 mm wide at the base; stigma c. 8 mm long, pointed with 3 lobes, split 8 mm to the base; style c. 1.5 mm long; staminodes c. 6, irregularly dentiform, mm. FRUITS elongate, sickled-shaped, cm (in young fruits), beak mm; epicarp smooth, shiny, dark green (young), mature fruits and seed not known; endosperm sparsely ruminate. Distribution and Ecology: Only known from very limited collections from 4 th Division of Sarawak in Northern Borneo and from the plant cultivated in Semengoh Forest Arboretum

58 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 39 near Kuching. This species grows in primary forest in mixed Dipterocarp forest in river valleys. This palm was successfully planted and fruits in Semengoh Forest Arboretum in Kuching, Sarawak. Local Name and Uses: Not recorded. Conservation Status: Data Deficient (IUCN, 2001). There is no population data record of this palm in the wild. However, the species is successfully grown in Semengoh Forest Arboretum as part of the ex situ conservation program run by Sarawak Forestry Department Specimens Studied: BORNEO. MALAYSIA: Sarawak, 4 th Division, Tatau, Ulu Anap, 22.vi.1982, Mokhtar &Othman S (K!, L!, KEP, SAN, SAR!). CULTIVATED. MALAYSIA: Sarawak, Semengoh Forest Arboretum, 42 m, 22.iv.1996, Baker et al. 716 (holotype: K!; isotypes: KEP, SING, SAR!); 31.iii.2008, Heatubun & Kuda 903 (K!, SAR!). Notes: Areca bakeri is similar to A. jugahpunya (subgenus Beccarioareca) in its acaulescent-clustering habit and broad leaflets, but can immediately be distinguished by the densely clustering habit with c. 20 stems in the clump, very long and slender petioles, leaflets more papery and the tips of terminal leaflets slightly obliquely lobed. The inflorescence of A. bakeri is more similar to A. dransfieldii (subgenus Areca) rather than A. tunku and A. jugahpunya, especially with its swollen-flattened rachillae and only one pistillate flower on each rachilla. The differences between A. bakeri and A. dransfieldii and the relationships with other species of Areca with swollen-flattened rachilla are discussed below (see notes under A. dransfieldii). The specific epithet of this new Areca is named after the collector of the type specimen and co-author of this paper, Dr. William J. Baker of Royal Botanic Gardens Kew.

59 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 40 Figure 1. Areca bakeri Heatubun. A, Apical portion of leaf. B, Middle portion of leaf. C, Basal portion of leaf and petiole attached to crownshaft. D, Inflorescence attached to stem just below the crownshaft and showing acaulescence habit. E, F, Staminate flower whole and in section. G, Pistillate flower. Scale bar: A, B = 8 cm; C, D = 3 cm; E, F = 2.2 mm; G = 7 mm. A D from Baker et al. 716; E G from Mokhtar & Othman S Drawn by Lucy T. Smith.

60 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA ARECA CALISO BECC. Areca caliso Becc., Leafl. Philipp. Bot. 8: 2998 (1919). Type: PHILIPPINES, Mindanao, Davao, Mt. Apo, Todaya, May 1909, Elmer (holotype: FI; isotype: K). Distribution: The Philippines. Notes: In the herbarium at Kew, there is a specimen (B.S ) collected by McGregor from Biliran, the Philippines and it had been designated by Beccari as a type of Areca biliranensis Becc., but this species was never published. Later one of us, ESF, determined that specimen in 1986 as A. caliso and also mentioned about the invalid name of A. biliranensis Nomen nudum. 4. ARECA CAMARINENSIS BECC. Areca camarinensis Becc. Philipp. J. Sci. 14: 309 (1919). Type: THE PHILIPPINES, Luzon, Camarines Province, Goa, Mt. Isarog, Apr. 1909, Aquilar (holotype: FI; isotype: ). Distribution: The Philippines. 5. ARECA CATECHU L. Areca catechu L., Sp. Pl.: 1189 (1753). Type: Pinanga Rumphius, Herb. Amboin. 1: t. IV (Lectotype). Areca cathechu Burm.f., Fl. Indica: 241 (1768). Type: Pinanga Rumphius, Herb. Amboin. 1: t. IV Areca faufel Gaertn., Fruct. Sem. Pl. 1: 19 (1788). Type: Pinanga Rumphius, Herb. Amboin. 1: t. IV Areca catechu var. nigra Giseke, Prael. Ord. Nat. Pl.: 73 (1792). ( Rumphius Pinang Itam ). Areca hortensis Lour., Fl. Cochinch.: 568 (1790). Type: Pinanga Rumphius, Herb. Amboin. 1: t. IV Areca catechu var. alba Blume, Rumphia 2: 68 (1839). ( Rumphius pinang poetih ).

61 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 42 Areca catechu L. var. batanensis Becc., Philipp. J. Sci. 3: 304 (1908). Type: the Philippines, Batanes Island, 6.vi.1907, Fenix 3834 (holotype: FI). Areca macrocarpa Becc., Phillip. J. Sci. C4: 601 (1909). Type: the Philippines, Mindanao, Zamboanga District, Port Banga, i.1908, Whitford & Hutchinson 9103 (holotype: FI). Areca catechu L. var. longicarpa Becc., Philipp. J. Sci. 6: 229 (1911). Type: the Philippines, Polillo Island, 10.xi.1909, Mc Gregor (holotype: FI). Areca catechu L. f. communis Becc., Philipp. J. Sci. 14: 304 (1919). Type: the Philippines, Mindanao, Misamis Province, Katajan, Mt. Malindang, Mearns & Hutchinson 4717 (holotype: FI). Areca catechu L. var. silvatica Becc., Becc., Philipp. J. Sci. 14: 304 (1919). Type: the Philippines, Palawan, Lake Manguao, iv.1913, Merrill 9447 (holotype: FI; isotypes: K, L). Distribution: Cultivated old world tropics. 6. ARECA CONCINNA THWAITES Areca concinna Thwaites, Enum. Pl. Zeyl.: 328 (1864). Type: SRI LANKA (Ceylon), C. P. 620 (holotype: not traced; isotypes: FI, K). Distribution: Sri Lanka. Notes: Areca concinna is poorly known and is only represented by its type specimen. Although there are a few specimens of Areca collected from Sri Lanka they are all Areca triandra. 7. ARECA COSTULATA BECC. Areca costulata Becc., Philipp. J. Sci. 14: 310 (1919). Type: THE PHILIPPINES, Leyte, Dagami, viii.1912, Ramos (holotype: FI; isotype: K). Distribution: The Philippines.

62 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA ARECA DRANSFIELDII HEATUBUN SP. NOV. (FIG. 2). Type: BORNEO, Sarawak, Miri, Lambir National Park, Path from waterfall, 17 m, 1.iv.2008, Heatubun & Rebi 901 (holotype: K; isotype: SAR). Diagnosis: Ad subgenerum Arecam inflorescentia structura A. tunku J. Dransf. & Lim Chong Keat affinis, sed caule caespitoso, inflorescentia cremea vel viride, floribus staminatis uniseriatim vel distichis, staminibus antheris dorsifixis, floribus foemineis solitariis in rachillis differt. Description: Small, slender, unarmed, monoecious, pleonanthic undergrowth palmlet, clustering (rarely solitary) with aerial branching, sometimes decumbent, stilt roots up to 40 cm. STEM 2 3 m tall, cm diam.; internodes 1 4 cm long, green near tip and greyish at the base, nodal scars conspicuous, whitish brown. LEAVES 5 7 in crown, pinnate, cm long (including petiole); sheath tubular, cm long, smooth, pale green to mid green; crownshaft well defined, cm long and up to cm diam.; petiole cm long, channelled adaxially, rounded abaxially; rachis ascending but not arching, with adaxial longitudinal ridge, rounded abaxially; blade with regularly arranged leaflets, 9 23 leaflets on each side; leaflets somewhat descending, 1 2 main veins running parallel from the base to the tip, leaflets near petiole small, c cm, slightly sigmoid, the middle leaflets about cm and the terminal cm, linear, pointed tip except for the terminal slightly oblique-lobed, papery, green adaxially and pale green abaxially. INFLORESCENCE infrafoliar, erect, cm long at anthesis, protandrous, branching to 1 order; peduncle cm long, cream when young and turning green when mature; prophyll green, c cm, elongate-triangular with pointed tip; rachis cream to green; rachis bracts not persistent; rachillae 5 7 and 7 11 cm long, very stiff and stout, flattened, elongate. STAMINATE FLOWERS small, triangular, mm, asymmetric, cream to pale green; sepals 3, low, about mm; petals 3, triangular, mm, striate; stamens 6, c. 2.5 mm 1.9 mm,; anthers dorsifixed, mm, cream-coloured, elongate, longer than the filaments; filaments mm, dark brown; pistillode low, c mm, rounded. PISTILLATE FLOWERS larger than the staminate, triangular, borne on the enlarged rachillae, buds varying greatly in size depending on stage of development, just before anthesis mm long and 10 11

63 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 44 mm wide, pale green to green; sepals 3, strongly imbricate, c mm, triangular, asymmetrical; petals 3, imbricate, triangular, mm; staminodes 3 4, irregular dentiform, 0.8 mm high. FRUITS elongate, sickle-shaped to ovoid, immature c cm, apical stigmatic remains, perianth sometimes persistent; epicarp smooth, shiny, very thin, dark green when young; mesocarp fibrous, c. 2.6 mm thick in immature fruit; endocarp very thin, adhering closely to seed. SEEDS elongated with pointed tip, c mm in immature fruit; endosperm ruminate; embryo basal. EOPHYLL bifid. Distribution and Ecology: Central and Northern parts of Borneo (in Central Kalimantan, Sarawak and Brunei Darussalam). This palm grows in primary forest in mixed Dipterocarp forest. Other palms that occupied the same habitat are Areca insignis var. moorei, Licuala sp., and Pinanga spp. Local Name and Uses: Pinang Nyaring and/or Pinang Bandang (local dialect in Central Kalimantan, Indonesia). There is no record of uses of this palm. Conservation Status: Data Deficient (IUCN, 2001). This species has been reported from a very restricted area in three different localities in Central Kalimantan (Indonesia), Lambir National Park in Sarawak (Malaysia) and Ladan Hills Forest Reserve in Lamunin, Tutong (Brunei Darussalam). Further population studies would be needed to assess the conservation status of this palm. Specimens Studied: BORNEO. MALAYSIA: Sarawak, Miri, Lambir National Park, Path from water fall, 17 m, 1.iv.2008, Heatubun & Rebi 901 (holotype: K!, isotype: SAR!). BRUNEI DARUSSALAM: Tutong, Lamunin, Ladan Hills Forest Reserve, 50 m, 20.v.1995, Dransfield JD 7531 (K!, BRUN). INDONESIA: Central Kalimantan Province, Upper Katingan River, c km West of North West Tumbang Samba, Tumbang Merak, 200 m, 20.xi.1982, Mogea & de Wilde 3452 (K!, L!, BO!).

64 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 45 Figure 2. Areca dransfieldii Heatubun. A, Stem, stiltroots and clustering habit. B, Middle portion of leaf. C, Apical portion of leaf. D, Inflorescence attached to stem just below the crownshaft. E, Detail portion of rachilla showing staminate and pistillate flowers in bud. F, G, Staminate flower whole and in section. H, Pistillate flower. I, Fruit still attached to rachilla. Scale bar: A C = 6 cm; D = 3 cm; E = 1.5 cm; F, G = 2.5 mm; H = 7 mm; I = 2 cm. A E, H from Mogea 3452; F G from Dransfield JD 7531; I from Heatubun 901. Drawn by Lucy T. Smith.

65 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 46 Notes: Areca dransfieldii is similar to A. tunku in the inflorescence structure, but it can be distinguished immediately by the clustering habit with aerial branching and sometimes decumbent stems. The inflorescence of A. dransfieldii is also cream when young and turning green when mature, the staminate flowers are mostly distichously arranged on the rachilla, with sepals free and stamens with dorsifixed anthers, and also only one pistillate flower for each rachilla rather than purple inflorescence, with staminate flowers uniseriate and spiral near the rachilla tip, sepals united to form a cup shaped calyx, sagittate anthers and more than one pistillate flower in A. tunku. Moreover, this new palm is restricted to Borneo species of Areca, whereas A. tunku is a Sumatran and Malay Peninsular species. The swollen-flattened rachillae in the inflorescence structure suggest that A. dransfieldii has close relationships with A. bakeri and A. tunku. Dransfield & Lim (1992) mentioned that A. tunku has relationships with A. ahmadii and A. jugahpunya in its swollen rachillae, but actually these two acaulescent species have long-cylindrical rachillae rather than really flattened as in A. tunku. The distinction between A. dransfieldii and A. tunku has already described above, while, A. bakeri is easily distinguished from A. dransfieldii by its acaulescent litter-trapping habit, leaves with very long petioles and broad leaflets,and infrafoliar inflorescences among marcescent leaf sheath swith slightly long peduncle. The distichous staminate flowers with free sepals free indicate that A. bakeri and dransfieldii belong to subgenus Areca, while Areca jugahpunya and A. tunku belong to subgenus Beccarioareca; they have united sepals (calyx) in the staminate flowers, and the spirally-arranged staminate flowers also observed in their inflorescence, even for small portions near the tip of the rachillae. This separation has been confirmed in the recent molecular phylogenetic analysis (Heatubun et al. in prep.). The specific epithet to honour Dr. John Dransfield from Royal Botanic Gardens at Kew, co-author of this paper in recognition his great achievements in palm botany in general and to palm genus Areca in Borneo in particular. 9. ARECA HUTCHINSONIANA BECC. Areca hutchinsoniana Becc., Philipp. J. Sci. 14: 312 (1919). Areca mammillata var. mindanaoensis Becc., Philipp. J. Sci. C 4: 62 (1909).

66 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 47 Type: THE PHILIPPINES, Mindanao Island, Bangka Province, Zamboanga, Port Banga, 20 m, xii.1907, Whitford & Hutchinson 9141 (holotype: FI). Distribution: The Philippines. 10. ARECA IPOT BECC. Areca ipot Becc., Leafl. Philipp. Bot. 2: 639 (1909). Areca catechu var. humilis Blanco (1845: 495). Type: THE PHILIPPINES, Luzon, Laguna, Nagcarlan, ii.1915, Elmer 9292 (lectotype: FI; isolectotypes: BM, BO, E, K, L, NSW, NY). Areca ipot var. polillensis Becc., Philipp. J. Sci. 6: 229 (1911). Type: THE PHILIPPINES, Island of Polillo, Robinson B.S (holotype: FI, isotypes: BM, K). Distribution: The Philippines. 11. ARECA KINABALUENSIS FURTADO Areca kinabaluensis Furtado, Repert. Spec. Nov. Regni Veg. 33: 228 (1933). Type: BORNEO, Sabah, Mt. Kinabalu, Lumu-Lumu, 1.iv.1932, Clemens J & M. S (holotype: SING; isotypes: B, K). Areca hallieriana Becc. ex Marteli, Atti Soc. Tosc. Sci. Nat. Pisa Processi Verbali 44: 114 (1934). Type: BORNEO, Kalimantan, at the foot of G. Demus, , Hallier 454 (holotype: BO; isotype: FI). Distribution: Borneo. 12. ARECA LAOSENSIS BECC. Areca laosensis Becc., Webbia 3: 191 (1910). Type: INDOCHINA, Laos, Phulet Phay, Attopeu, iv.1877, Harmand 1201 (holotype: P; isotype: FI). Distribution: Indochina (Lao PDR).

67 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA ARECA MACROCALYX ZIPP. EX BLUME Areca macrocalyx Zipp. ex Blume, Rumphia 2: 75 (1839). Type: NEW GUINEA, SW coast, Zippel s.n. (holotype: L). Areca jobiensis Becc., Malesia 1: 21 (1877). Type: New Guinea, Gelvink Bay, Japen Island, Ansus, iv.1875, Beccari s.n. (holotype: FI; isotypes: K). Areca macrocalyx var. zippelliana Becc., Malesia 1: 19 (1877). Type: NEW GUINEA, SW coast, Zippel s.n. (holotype: L). Areca macrocalyx var. aruensis Becc., Malesia 1: 20 (1877). Type: MOLUCCAS, Aru Island, Vokan (Wokam), iii.1875, Beccari s.n. (holotype: FI; isotype: K). Areca macrocalyx var. conophyla Becc., Malesia 1: 20 (1877). Type: NEW GUINEA, NW coast, Ramoi and Amberbaken, 1872, Beccari s.n. (holotype: FI; isotype: K). Areca macrocalyx var. waigheuensis Becc., Malesia 1: 20 (1877). Type: NEW GUINEA, Waigeo Island, Wakkere (Wakre), iii.1875, Beccari s.n. (holotype: FI; isotype: K). Areca rechingeriana Becc., Webbia 3: 163 (1910). Type: PAPUA NEW GUINEA, Bougenville Island, Kieta, Rechinger 3992 (holotype: B ; isotype: FI). Areca macrocalyx var. intermedia Becc., in R. Rechinger, Teil. Denkschr. Kaiserl. Akad. Wiss. Wien Math.-Naturwiss. Kl. 89: 506 (1913). Type: PAPUA NEW GUINEA, Bougenville Island, Kieta, Rechinger 4182 (holotype: B, isotype: FI). Areca nigasolu Becc., Webbia 4: 256 (1914). Type: SOLOMON ISLANDS, Treasury Island, Guppy 95 (holotype: K, photo FI). Areca torulo Becc., Webbia 4: 253 (1914). Type: SOLOMON ISLANDS, Treasury Island, Guppy 94 (holotype: K). Areca warburgiana Becc., Bot. Jarhrb. Syst. 52: 24 (1914). Type: NEW GUINEA, Sigar, Warburg 20 (holotype: B, photo FI!; isotype: FI). Areca nanospadix Burret, J. Arnold Arbor. 12: 265 (1931). Type: PAPUA NEW GUINEA, Ihu, Vailala River, rain forest, 9.iii.1926, Brass 921 (holotype: A). Areca rostrata Burret, Notizbl. Bot. Gart. Berlin-Dahlem 12: 322 (1935). Type: PAPUA NEW GUINEA, Diemi, Onange road, Central Division, 6.v.1933, Brass 3971 (holotype: A). Areca multifida Burret, Notizbl. Bot. Gart. Berlin-Dahlem 13: 331 (1936). Type: PAPUA NEW GUINEA, Veiya, 11.iii.1935, Carr (holotype: B, isotype: A, K). Distribution: Moluccas, New Guinea and Solomon Islands.

68 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA ARECA MANDACANII HEATUBUN Areca mandacanii Heatubun, Palms 52: 199 (2008). Type: INDONESIA, West Papua Province, Sorong Selatan, Teminabuan, Sayal, Maampou Forest, ii.2003, Heatubun 423 (holotype: BO; isotypes: K, MAN). Distribution: New Guinea (Bird s head peninsula in western New Guinea). 15. ARECA MONTANA RIDL. Areca montana Ridl., Mat. Fl. Malay. Penins. 2: 136 (1907). Type: MALAY PENINSULA, Selangor, Semangkok pass, 1.iii.1905, Burn-Murdoch s.n. (holotype: SING). Areca latiloba Ridl., J. Straits Branch Roy. Asiat. Soc. 86: 310 (1922). Type: MALAY PENINSULA, Negeri Sembilan, Bukit Tangga, Mountain forest, 800 m, Ridley s.n. (holotype: K). Areca recurvata Hodel, Palm J. 134: 28 (1997). Type: THAILAND, SW of Narathiwat, 800 m, Hodel & Vatcharakorn 1634 (holotype: BK). Distribution: Thailand, Malay Peninsula, Sumatra and Java. 16. ARECA OXYCARPA MIQ. Areca oxycarpa Miq., Verh. Kon. Ned. Akad. Wetensch. Afd. Natuurk. 11: 1 (1868). Type: CELEBES, Menado, Mt. Pisah, Riedel s.n. (holotype: L). Areca celebica Burret, Repert. Spec. Nov. Regni Veg. 32: 115 (1933). Type: CELEBES, Tomohon, Sarasin s.n. (holotype: B). Distribution: Celebes (Sulawesi). 17. ARECA PARENS BECC. Areca parens Becc., Philipp. J. Sci. 14: 307 (1919). Type: THE PHILIPPHINES, Luzon, Camarines, iv.1914, Penas, Soriano & Abellanosa F.B (holotype: FI). Distribution: The Philippines.

69 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA ARECA RHEOPHYTICA J. DRANSF. Areca rheophytica J. Dransf., Kew Bull. 39: 18 (1984). Type: BORNEO, Sabah, Telupid, 23.x.1979, Dransfield et al. JD 5780 (holotype: K; isotypes: L, SAN). Distribution: Borneo. 19. ARECA RIPARIA HEATUBUN SP. NOV. (FIG. 3) Type: CAMBODIA, Province Koh Kong, Tatai Chveng waterfall, less than 20 m, 27.ix.2008, Evans 174 (holotype: K). Diagnosis: Ad subgenerum Arecam pertinens, a ceteris speciebus Cambodianis distinctissima; A. rheophytica J. Dransf. Borneensi similis, sed palma caespitosa et staminibus 3 differt. Description: Clustering, unarmed, monoecious, pleonanthic, small tree palm. STEM to 2.5 m tall, cm diam.; internodes cm long, green near the crown, shiny, brown near the base, nodal scars conspicuous, white to whitish brown. LEAVES about 7 in the crown, pinnate, to 80 cm long (including petiole); sheath tubular, c cm long, smooth, greenish cream; crownshaft well defined, cm long and 1 2 cm diam.; petiole short, cm long, 5 5 mm at the base, channelled adaxially, rounded abaxially; rachis somewhat arching, very slender with adaxial longitudinal ridge, rounded abaxially; blade with regularly arranged leaflets, about 25 leaflets on each side; leaflets somewhat arching, single fold, linear, cm long, acuminate, with pointed tips, papery, green adaxially and light green abaxially, slightly discolorous when dried; continuous white woolly ramenta present along the mid-ribs on abaxial surface. INFLORESCENCE infrafoliar, erect, small and slender, cm long and 6 8 cm wide, protandrous, branching to 1 order and sometimes the basalmost rachilla branches to 2 order; peduncle cm long; prophyll c cm, lanceolate, 2-keeled, papery, cream coloured, entirely enclosing the inflorescence, then splitting longitudinally and falling before staminate flower anthesis;

70 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 51 rachis cream to greenish; rachillae numerous, divaricate, elongate, yellowish, usually with a basal triad, rarely with several triads, otherwise bearing paired staminate flowers in a single row on one side. STAMINATE FLOWERS small, triangular, c mm (young stage), asymmetric, cream to pale brown; calyx low, triangular cup-shaped, c mm wide and 0.5 mm high (at young stage), 3-lobed; petals 3, strongly keeled, valvate, c mm (at young stage); stamens 3, about 1 mm long (at young stage), basifixed; anthers c. 0.8 mm long (at young stage), white to cream-coloured, longer than the filaments; filaments c. 0.2 mm (at young stage), orange to brown; pistillode lacking. PISTILLATE FLOWERS larger than the staminate, triangular, c. 6.2 mm long and 4 mm wide (at young stage); sepals 3, strongly imbricate, c. 4 3 mm (at young stage), triangular; petals 3, imbricate, triangular, c. 5 3 mm at anthesis; gynoecium about 3 mm (at young stage), lacking staminodes. FRUITS cm, elongate, fusiform, apical stigmatic remains, epicarp smooth, mesocarp fleshy, endocarp fibrous, bright crimson when ripe. SEEDS c cm, obovoid, endosperm ruminate, embryo basal. Distribution and Ecology: Distribution of this palm is restricted and only known from one collection (and few photographs) taken by Tom Evans in Tatai Chveng waterfall in Koh Kong Province, Cambodia. Based on the field notes, this palm grows as a rheophyte in a wet, well vegetated jumble of rocks along the banks of a forest river, just downstream of the falls out of the splash zone but below the peak flood level. Local Name and Uses: Not recorded. Conservation Status: Data Deficient (IUCN, 2001). Further population study is needed to assess the conservation status of this palm, although the collector had stated that this palm is common on the river system near the collection area. Specimens Studied: CAMBODIA. Province Koh Kong, Tatai Chveng waterfall, less than 20 m, 27.ix.2008, Evans 174 (holotype K!).

71 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 52 Figure 3. Areca riparia Heatubun. A, Stem, leaf and inflorescence. B, Detail of rachilla showing staminate and pistillate flowers in bud. C, Staminate flower in section. D, Pistilate flower. E, F, Fruit whole and in section. Scale bar: A = 4 cm; B = 7 mm; C = 1 mm; D = 3 mm; E, F = 1 cm. All from Evans 174. Drawn by Lucy T. Smith.

72 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 53 Notes: Areca riparia is the only rheophytic species of Areca from Cambodia (Indochina). This species is similar to the Bornean rheophyte A. rheophytica in general appearance, except its clustering habit, pistillate flowers more broad with slightly rounded tips, and stamens 3 in A. riparia, rather than habit solitary, pistillate flowers narrow with pointed tips and stamens 6 in A rheophytica. Areca riparia is the third species of Areca with three stamens; the other two are A. montana and A. triandra. However, Areca montana is solitary, with very few broad leaflets, inforescences always branched to 1 order and the triad is only seated at the very base of the rachilla and almost a half way of rachilla length from the dyads of staminate flowers start to assembly (there is a disjunction or space between triad and dyads of staminate flowers), and also protogynous. While, A. triandra is medium palm, with inflorescence branched to 3 oders and more than one triad in one rachilla. Moreover, A. montana and A. triandra are ecologically very different; they are not rheophytic. The specific ephitet reflects the specific habitat where this species grows. 20. Areca triandra Roxb. ex Buch.-Ham., Areca triandra Roxb. ex Buch.-Ham., Mem. Wern. Nat. Hist. Soc. 5: 310 (1826). Type: ANDAMAN ISLAND, Buchanan-Hamilton s.n. (holotype: not traced) Areca laxa Buch.-Ham., Mem. Wern. Nat. Hist. Soc. 5(2): 309 (1826). Type: ANDAMAN ISLAND, Buchanan-Hamilton s.n. (holotype: not traced) Areca nagensis Griff., Calcutta J. Nat. Hist. 5: 453 (1845). Nenga nagensis (Griff.) Scheff., Ann. Jard. Bot. Buitenzorg 1: 120 (1876). Type: INDIA, Assam, Naga hills, 1842, Griffith s.n. (holotype: BR; K (photo)). Ptychosperma polystachyum Miq., Fl. Ned. Ind., Eerste Bijv.: 590 (1861). Areca polystachya (Miq.) H. Wendl. in O. C. E. de Kerchove de Denterghem, Palmiers: 232 (1878). Type: SUMATRA, Bangka Island, Jebus, Teijsmann s.n. (holotype: BO). Areca triandra var. bancana Scheff., Natuurk. Tijdschr. Ned.-Indië 32: 165 (1873). Type: SUMATRA, Bangka Island, Jebus, Teijsmann s.n. (holotype: BO). Areca borneensis Becc., Malesia 1: 22 (1877). Type: BORNEO, Sarawak, Rejang, Beccari P.B (holotype: FI; isotype: K).

73 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 54 Areca aliceae W. Hill ex F. Muell., Gartenflora 28: 199 (1879). Type: AUSTRALIA, cultivated in Brisbane Botanic Garden, from plant Trinity bay, (holotype: K). Distribution: India, Sri Lanka, Andaman Island, Indochina, Thailand, Malay Peninsula, Sumatra and Borneo. 21. ARECA VIDALIANA BECC. Areca vidaliana Becc., Philipp. J. Sci., C 2: 222 (1907). Type: THE PHILIPPINES, Palawan, iv.1887, Vidal 3955 (holotype: FI; isotypes: K). Areca mammilata Becc., Philipp. J. Sci., C 2: 220 (1907). Type: THE PHILIPPINES, Palawan, ii.1906, Curran F.B (holotype: FI). Distribution: The Philippines. 22. ARECA WHITFORDII BECC. Areca whitfordii Becc., Philipp. J. Sci., C 2: 219 (1907). Type: THE PHILIPPINES, Mindoro Island, Bongabon River, i.1906, Whitford 1372 (holotype: FI). Areca whitfordii Becc. var. luzonensis Becc., Philipp. J. Sci. 14: 306 (1919). Type: THE PHILIPPINES, Luzon Island, Nueva Vizcaya, Dupax, Mc Gregor (holotype: FI; isotype: BM). Distribution: The Philippines. II. Subgenus Beccarioareca Furtado 23. ARECA ABDULRAHMANII J. DRANSF. Areca abdulrahmanii J. Dransf., Bot. J. Linn. Soc. 80: 33 (1980). Type: BORNEO, Sarawak, 4 th Division, G. Mulu National Park, west of Sungai Berar Camp, iii.1978, Jermy (holotype: SAR; isotype K). Distribution: Borneo.

74 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA ARECA ANDERSONII J. DRANSF. Areca andersonii J. Dransf., Kew Bull. 39: 6 (1984). Type: BORNEO, Sarawak, 4 th Division, Miri, Niah, Mt. Subis, Sungai Sekaloh, 60 m, viii.1972, Anderson S (holotype: K; isotype: SAR). Distribution: Borneo. 25. ARECA ARUNDINACEA BECC. Areca arundinacea Becc., Malesia 1: 23 (1877). Type: BORNEO, Sarawak, Sakarang, ix.1867, Beccari P.B (holotype: FI; isotype: K). Distribution: Borneo. 26. ARECA BRACHYPODA J. DRANSF. Areca brachypoda J. Dransf., Kew Bull. 39: 8 (1984). Type: BORNEO, Sarawak, 1 st Division, Sematan, Lundu, Mt. Pueh Forest Reserve, v.1981, Dransfield et al. JD 6011 (holotype: K; isotypes: BH, BO, L, SAN, SAR). Distribution: Borneo. 27. ARECA CHURCHII HEATUBUN SP. NOV. (FIG. 4) Type: BORNEO, West Kalimantan, Serawai, Sungai Merah, 2 km to W of camp along Gaharu trail towards Sungai Labang, surrounding ridges and valleys, 600 m, ii.1995, Church et al (holotype: K; isotypes: BO, A). Diagnosis: ad subgenerum Beccarioarecam Furtado pertinens, palma solitaria, caude gracillimo, inflorescentia fortiter effusa, floribus staminatis spiraliter dispositis, sepalis elongatis, staminibus 11 12, filamentis interdum tortilis et circinatis, floribus foemineis semper solitaria in rachillis, 1 2 cm longioribus in rachillis basaliter dispositis, differt. Description: Solitary, slender, unarmed, monoecious, pleonanthic, undergrowth palmlet. STEM up to 2 m tall, c. 2 cm diam.; internodes 1 2 cm long. LEAVES c. 8 leaves in crown,

75 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 56 pinnate, cm long; sheath tubular, not fibrous, up to 20 cm long, striate; crownshaft well defined, cm long, 3 cm diam.; petiole up to 65 cm long, channelled adaxially, rounded abaxially, 5 6 mm wide and 6 7 mm thick; rachis with adaxial longitudinal ridge, rounded abaxially; blade with irregularly arranged leaflets, 5 7 leaflets on each side, 3 11 cm spacing between leaflets, leaflets with 2 7 folds, leaflets near petiole c cm, slightly sigmoid and tip oblique-lobed, middle leaflets cm, and terminal leaflets about 33 7 cm, tips bifid with lobes cm depth, papery, dark adaxially and pale abaxially when dried. INFLORESCENCE infrafoliar, upright, cm at anthesis, protandrous, branching to 1 order; peduncle cm long, prophyll not known; rachis pale yellow at the base and peach-coloured towards the apex and light brown when dried; rachis bracts caducous; rachillae 13 17, cm long, slender, spreading, covered by minute light brown to chocolate-brown indumentum, elongate, sinuous to slightly zigzag in appearance. STAMINATE FLOWERS small, triangular, elongate, mm, asymmetric, spirally arranged on the rachillae; calyx cupshaped, mm wide and 2 mm high, 3-lobed, the lobes c mm; petals 3, valvate, elliptic to slightly spatulate, c. 7.7 mm long and 1.8 mm wide at anthesis, connate at the base, cream-coloured; pistillode lacking; stamens 11 12, mm long, elongate, basifixed; anthers mm long and 0.3 mm wide after anthesis, cream-coloured; filaments mm long and 0.2 mm wide, dark brown, sometimes twisted and coiled, free near corolla and connate at the centre. PISTILLATE FLOWERS larger than the staminate, triangular, mm, only one flower on each rachilla and never on the rachis or main axis of the inflorescence, positioned 1 2 cm from the base of rachilla; sepals 3, imbricate, triangular, striate, 7 9 mm long and 6 mm wide at anthesis; petals 3, imbricate, triangular, striate, c mm; gynoecium 13 4 mm at anthesis, tubular shaped; stigma trifid, 4 4 mm; style 9 mm long, 4 mm wide; staminodes lacking. FRUITS very young, dull green and yellow at the base, mature fruits not known. Distribution and Ecology: Only known from the type of locality in Sungai Merah, Serewai, West Kalimantan. This species grows on hill slopes above the perennial stream of Sungai Labang in Dipterocarp forest in association with Shorea, Dipterocarpus, Lauraceae and Sapotaceae.

76 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 57 Figure 4. Areca churchii Heatubun. A, Apical portion of leaf. B, Middle and basal portion of leaf. C, Petiole, leaf sheath and crownshaft. D, Inflorescence with pistillate flowers still attached. E, Detail portion of rachilla showing staminate and pistillate flowers in bud. F, G, Staminate flower whole and in section. H, I, Pistillate flower whole and in section. Scale bar: A C = 6 cm; D = 4 cm; E = 2 cm; F, G = 3 mm; H, I = 5 mm. A E, H from Mogea 3452; F G from Dransfield JD 7531; I from Heatubun 901. Drawn by Lucy T. Smith.

77 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 58 Local Name and Uses: There is no record of local name and uses of this palm. Conservation Status: Data Deficient (IUCN, 2001). Only known from the type of locality in Sungai Merah area in Serawai, Indonesian Province of West Kalimantan, Borneo. Specimens Studied: BORNEO. INDONESIA: West Kalimantan Province, Serawai, Sungai Merah, 2 km to west of camp along Gaharu trail towards Sungai Labang, surrounding ridges and valleys, 600 m, ii.1995, Church et al (K!, BO!, Harvard). Notes: Areca churchii is very striking with its relatively large staminate flowers borne on slender-spreading rachillae. The spiral arrangement of staminate flowers on the rachilla, the sepals united and 12 stamens indicate this species belongs to subgenus Beccarioareca. The elongate-elliptical petals of the staminate flower are rather similar to those of A. costulata, A. jugahpunya, A. novohibernica and A. vestiaria in the same subgenus. This character was reported also to occur in A. ahmadii from different subgenus Areca (Dransfield 1984). The fused, twisted and coiled filaments are never reported in the genus Areca or even in the subtribe Arecinae; in the tribe Areceae this character has only been spotted in the monotypic and endemic genus Tectiphiala from Mauritius and also perhaps in the genus Calyptrocalyx and Chambeyronia (Uhl & Dransfield 1987, Dransfield et al. 2008). Areca churchii is similar to others miniature Areca species A. abdulrahmanii, A. andersonii and A. klingkangensis in its slender-solitary habit and pinnate leaves in general, although there are still some distinctions in leaf segmentation and texture, but basically they differ in the inflorescence and flower structure, including number of stamens. Areca churchii has an inflorescence with long-slender rachillae and 12 stamens, while A. abdulrahmanii has an inflorescence with 3 divaricate-sinous rachillae and 16 stamens, A. andersonii has an inflorescence with 3 4 very slender rachillae and 6 stamens and A. klingkangensis has an inflorescence with only 3 rachillae and 9 stamens. The divaricate structure of the inflorescence and the sinuous to slightly zigzag rachillae of A. churchii are similar to those of A. abdulrahmanii, A. arundinacea, A. furcata, A. minuta, and A. ridleyana. However, except to the important characters of staminate flowers already mentioned above, this new species can be easily distinguished by the presence of the only

78 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 59 one pistillate flower on each rachilla, and also this pistillate flowers is positioned 1 2 cm from the base of the rachilla. The specific epithet of this species homours the collector of the type specimen, A. C. Church from Harvard University, USA. 28. ARECA DAYUNG J. DRANSF. Areca dayung J. Dransf., Bot. J. Linn. Soc. 81: 30 (1980). Type: BORNEO, Sarawak, 4 th Division, Bintulu, Bukit Nyabau, 13.iv.1968, Dransfield JD 785 (holotype: K; isotype: BH). Distribution: Borneo. 29. ARECA FURCATA BECC. Areca furcata Becc., Malesia 1: 23 (1877). Type: Borneo, Sarawak, 7 th Division, on hill of Belaga, ix.1867, Beccari P.B (holotype: FI). Distribution: Borneo. 30. ARECA GURITA HEATUBUN SP. NOV. (FIG. 5). Type: CULTIVATED, Borneo, Sarawak, Semengoh Forest Arboretum, Palm Collection, 42 m, 31.iii.2008, Heatubun & Kuda 896 (holotype: K; isotype: SAR). Diagnosis: Ad subgenerum Beccarioarecam, a ceteris speciebus Borneensibus elegantissima, palmae caespitosae, A. minuta Scheff. habitu similis, inflorescentia octopiformis, rhachillis recurvatis, floribus secundis ab imum usque ad apicem et spiraliter ad apicem, floribus staminatis alternatis, antheris connectivo liniare, differt. Description: Clustering with many suckers, unarmed, monoecious, pleonanthic, small undergrowth-palmlet. STEM to 1.5 m high, 1 cm in diam.; internodes 3 cm long, smooth and green near the crown, light brown to whitish near the base. LEAVES c. 11 in crown,

79 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 60 pinnate, c. 32 cm long (including petiole); sheath tubular, c. 20 cm long and 1 2 cm wide, light brown to brown, and light brown when dried; crownshaft well defined, up to 30 cm long and 1 2 cm diam.; petiole c. 3 cm long, channelled adaxially, rounded abaxially; rachis very slender with adaxial longitudinal ridge, rounded abaxially; blade with irregularly arranged leaflets, 2 4 leaflets on each side, spacing between leaflets cm; leaflets from narrow to broad, leaflets near petiole c cm, slightly sigmoid, tip obligue acuminate, the middle leaflets about c cm, linear with 1 vein, the terminal leaflets broad, flabellate, c cm, margin toothed, papery, concolorous. INFLORESCENCE infrafoliar, erect, small and dense, to 10 cm long and to 10 cm wide, protandrous, always branching to 1 order; peduncle short to 3 cm, covered in thick brown indumentum; prophyll persistent; rachis also covered in rusty brown indumentum; rachillae 3 8 (including rachis), divaricate, to 10 cm long and c. 5 mm wide near base, recurved; flowers secund in general appearance, spiral at the tip of rachilla, triads alternate. STAMINATE FLOWERS small, triangular to rounded, mm, asymmetric, white to cream; calyx fused, triangular cup-shaped, c mm before anthesis, 3-lobed, light brown to brown; petals 3, fused near base, obovate, mm at anthesis, white to cream-coloured; stamens 6, mm 0.5 mm, basifixed; anthers c mm, cream-coloured, sagittate, longer than filaments; filaments c mm, dark brown, connective linear; pistillode absent. PISTILLATE FLOWERS larger than the staminate, triangular, borne at the rachillae base (1 3 on each rachilla) and distributed to half of rachis (main axis) length, 5 15 flowers in the rachis, c. 5.5 mm long and 3.5 mm wide before anthesis, greenish; sepals 3, strongly imbricate, mm before anthesis, somewhat triangular, asymmetrical; petals 3, imbricate, triangular, c mm before anthesis; lacking staminodes; Gynoecium c. 4 mm (including stigma). FRUITS (young) white with green at the tip, obovoid with beak, c. 2 cm long and 1 cm wide (young fruit still in development), apical stigmatic remains, mature fruits not known. Distribution and Ecology: This species is only known from several collections in Bintulu Division of Sarawak. It grows in heath forest to montane forest from an altitude m above sea level. Local Name and Uses: Not recorded.

80 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 61 Figure 5. Areca gurita Heatubun. A, Habit. B, Leaf. C, Inflorescence showing staminate and pistillate flowers in bud and their arrangement on rachillae. D, E, Staminate flower whole and in section. F, Fruit still attached to rachilla. Scale bar: A = 31.5 cm; B = 3 cm; C = 3.6 cm; D, E = 1.2 mm; F = 1.3 cm. All from Heatubun 896. Drawn by Lucy T. Smith.

81 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 62 Conservation Status: Data Deficient (IUCN, 2001). Further population study is needed to assess the conservation status of this palm. However, this species has been cultivated in Semengoh Forest Arboretum near Kuching and the palm has been flowering and fruiting. Specimens Studied: BORNEO. Sarawak, Kuching, Semengoh Forest Arboretum, Palms Collection, 42 m, 31.iii.2008, Heatubun & Kuda 896 (holotype: K!; isotype: SAR!); Bintulu Division, Bukit Kana, Ulu sg. Sanggan, Tatau, 680 m, 25.iii.1995, Yii & Jugah S (K!, KEP, SAR!); Lumut Range, 750 m, 22.ix.1992, Mohtar & Yii S (SAR!); Sabal, Logging road, Sabal sisi, 440 m, 18.ii.1995, Runi et al. S (K!, KEP, L!, SAR!); Miri Division, Lavang Forest Protected, 80 m, 24.iv.1994, Yii et al. S (KEP, SAR!). Notes: Areca gurita is similar to A. minuta in habit, leaves and/or in general appearance. However, they can easily be differentiated by inflorescence structure and flower arrangement. The rachillae of A. gurita are recurved and the flowers are secund; they seem like octopus tentacles. The flower arrangement is alternate and the anther connectives are also linear, rather than the spiral flower arrangement and furcate connectives in A. minuta. The flower arrangement in A. gurita is actually a modification from spiral to secund or alternate; the spiral arrangement still occurs in the tip of rachillae. The rachillae are expanded on one side and display all flowers in the one direction. Although the pistillate flowers appear secund, but the fused calyx (or sepals united) still indicate this species belong to subgenus Beccarioareca. The specific epithet is gurita, the Indonesian word for octopus, to express the special appearance of the inflorescence of this species. 31. ARECA INSIGNIS (BECC.) J. DRANSF. Areca insignis (Becc.) J. Dransf., Kew Bull. 39: 13 (1984). Gigliolia insignis Becc., Malesia 1: 172 (1877). Pichisermollia insignis (Becc.) H. Montiero, Rodriguésia 28: 198 (1976).

82 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 63 Type: BORNEO, Sarawak, Bintulu, , Beccari P.B (holotype: FI; isotype: K). var. insignis Distribution: Borneo. var. moorei (J. Dransf.) J. Dransf., Kew Bull. 39: 13 (1984). Pichisermollia insignis var. moorei J. Dransf., Bot. J. Linn. Soc. 81: 40 (1980). Type: BORNEO, Sarawak, 1st Division, Bako National Park, Lintang path between North entrance and Bukit Tambi, i Moore & Chai 9109 (holotype: K!; isotypes: BH, L, SAR). Distribution: Borneo. 32. ARECA JUGAHPUNYA J. DRANSF. Areca jugahpunya J. Dransf., Kew Bull. 39: 13 (1984). Type: BORNEO, Sarawak, 7 th Division, Ulu Sungai Kapit, 25.ii.1975, Chai et al. S (holotype: K; isotypes: BH, L, SAR). Distribution: Borneo. 33. ARECA KLINGKANGENSIS J. DRANSF. Areca klingkangensis J. Dransf., Kew Bull. 39: 13 (1984). Type: BORNEO, Sarawak, 1 st Division, Serian, Sabal Tapang Forest Reserve, G. Gaharu, v.1981, Dransfield et al. JD 6103 (holotype: K; isotypes: BH, SAR). Distribution: Borneo. 34. ARECA MINUTA SCHEFF. Areca minuta Scheff., Ann. Jard. Bot. Buitenzorg 1: 146 (1876). Type: BORNEO, West Kalimantan, Landak, Teijsmann s.n. (holotype: BO). Areca tenella Becc., Malesia 1: 22 (1877). Type: BORNEO, Sarawak, , Beccari P.B (lectotype: FI; isolectotype: K).

83 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 64 Areca amdjahi Furtado, Repert. Spec. Nov. Reg. Veg. 33: 235 (1933). Type: BORNEO, East Kalimantan, Samenggaris, Amdjah 1019 (holotype: B ; isotype: BO, SING (photo)). Areca bongayensis Becc. ex Furtado, Repert. Spec. Nov. Reg. Veg. 33: 234 (1933). Type: BORNEO, Sabah, Bongaya, xii.1897, Ridley 9090 (holotype: SING; isotype: K). Areca hewittii Furtado, Repert. Spec. Nov. Reg. Veg. 33: 234 (1933). Type: BORNEO, Sarawak, Lingga Mountain, Hewitt 25 (holotype: SING). Areca hullettii Furtado, Repert. Spec. Nov. Reg. Veg. 33: 235 (1933). Type: BORNEO, Sarawak, G. Matang, vii.1890, Hullett s.n. (holotype: SING). Distribution: Borneo. 35. ARECA MOGEANA HEATUBUN SP. NOV. (FIG. 6). Type: Borneo, Indonesia, Kalimantan Barat Province, Kabupaten Hulu Kapuas, Bentuang Karimun National Park, Mogea ITTO/BA 0605 (Holotype: SAR). Diagnosis: Ad subgenerum Beccarioarecam, a ceteris speciebus Borneensibus, A. abdulrahmanii J. Dransf. habitu et inflorescentia structura similis, sed staminibus 6, floribus foemineis plus quam uno ( 10) semper in rhachis sitis ab imum usque ad apicem, differt. Description: Solitary, unarmed, monoecious, pleonanthic, small undergrowth-palmlet. STEM to 2 m tall, 5 8 mm diam.; internodes cm long. LEAVES 5 7 in crown, pinnate, to 50 cm long (including petiole); sheath tubular, c. 10 cm long and cm wide, striate, dark green to tinged red, tomentose, and brown to dark brown when dried, covered by punctiform scales, sparse or very dense near the petiole; crownshaft well defined, up to 17 cm long and 1 cm diam.; petiole 4 10 cm long, channelled adaxially, rounded abaxially, covered with thick brown indumentum; rachis very slender with adaxial longitudinal ridge, rounded abaxially; blade with irregularly arranged leaflets, 5 6 leaflets on each side, spacing between leaflets 3 6 cm; leaflets from narrow to slightly broad, leaflets near petiole cm, linear, tip obligue acuminate, the middle leaflets about cm, linear with 1 3 veins, the terminal leaflets broad, sometimes less broad than middle leaflets or sometimes flabellate, cm, somewhat

84 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 65 lanceolate with small cleft, papery, discolorous when dried, slightly pale adaxially and dark abaxially, with sparse brown ramenta along the mid-ribs in abaxial surface. INFLORESCENCE infrafoliar, erect, small and slender, cm long and cm wide, protandrous, always branching to 1 order; peduncle 2 5 mm long, covered by thick brown indumentum; prophyll not available; rachis also covered by brown indumentum; rachis bracts persistent, low, triangular; rachillae 5 8 (including main axis), divaricate, 7 10 cm long and c. 2 mm wide near base, elongate, very slender, sinuous. STAMINATE FLOWERS small, triangular, sickle-shaped when young to elongate at anthesis, mm, asymmetric, cream to pale brown, spirally arranged on the rachilla; calyx fused, triangular cup-shaped, c mm at anthesis, 3-lobed; petals 3, fused near base, obovate, mm at anthesis, striate; stamens 6, mm 0.5 mm, basifixed; anthers c mm, cream-coloured, elongate, sagitatte, shorter than the filaments; filaments c. 0.5 mm long, dark brown, connective furcate; pistillode absent. PISTILLATE FLOWERS larger than the staminate, triangular, borne on the enlarged rachillae and distributed to half the rachis length, only one on each rachilla and usually abortive, more than one ( 10) on the rachis (main axis), c mm (young stage); sepals 3, strongly imbricate, c mm (young stage), triangular, asymmetrical; petals 3, imbricate, triangular, c mm (young stage); staminodes absent. FRUITS (young) light orange at the base becoming red and darker red at the tip, elongate, sickle-shaped, c cm long and 0.3 cm wide, apical stigmatic remains, mature fruits not known. Distribution and Ecology: This species is only known from several collections collected in Bentuang Karimun National Park in Kabupaten Hulu Kapuas, Kalimantan Barat Province, Indonesia, Borneo. This palm was recorded growing in submontane forest at an altitude 1100 m above sea level. Local Name and Uses: Not recorded. Conservation Status: Data Deficient (IUCN, 2001). Further population study is needed to assess the conservation status of this palm.

85 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 66 Figure 6. Areca mogeana Heatubun. A, Stem, leaf and inflorescence. B, Inflorescence showing staminate and pistillate flowers in bud and their arrangement on rachillae. C, D, Staminate flower whole and in section. Scale bar: A = 4 cm; B = 2 cm; C, D = 1.5 mm. A, C, D from Mogea ITTO/BA 0595; B from Mogea ITTO/BA Drawn by Lucy T. Smith.

86 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 67 Specimens Studied: BORNEO. INDONESIA: Kalimantan Barat Province, Kabupaten Hulu Kapuas, Bentuang Karimun National Park, trail to Bukit Condong, 1100 m, 17ix.1997, Chai et al. ITTO/BA 0368 (SAR!); Mogea ITTO/BA 0585 (BO, SAR!); Mogea ITTO/BA 0595 (BO, SAR!); Mogea ITTO/BA 0605 (holotype SAR!). Notes: Areca mogeana is similar to A. abdulrahmanii in its solitary habit, dissected-leaf blade with several leaflets and inflorescence structure with elongated-sinuous rachillae. The two differ from each other in triad arrangement and staminate flowers, including stamens number triads more than one ( 10) and mainly confined in the rachis rather than the rachillae in A. mogeana, and if there any triads on the rachillae they are usually abortive and in A. abdulrahmanii, the triads borne at the base of the rachilla. Moreover, ther are six stamens in A. mogeana versus 16 in A. abdulrahmanii. The specific epithet honours the collector of the type specimen Dr. Johanis P. Mogea, palm botanist from Herbarium Bogoriense LIPI (Indonesian Institute of Science) in recognition of his contribution to our understanding of the palm flora of Indonesia. 36. ARECA NOVOHIBERNICA (LAUTERB.) BECC. Areca novohibernica (Lauterb.) Becc., Bot. Jahrb. Syst. 52: 24 (1914). Nenga novohibernica Lauterb., Bot. Jarhb. Syst. 45: 357 (1911). Type: PAPUA NEW GUINEA, Bismarck Archipelago, New Ireland, Nabumai, Urwald, Peekel 110 (holotype: FI; isotype: K, B ). Areca guppyana Becc., Webbia 4: 258 (1914). Type: SOLOMON ISLANDS, Shorland Islands, Alu Island, 1 to 2 miles from coast, Guppy 107 (holotype: K). Areca salomonensis Burret, Notizbl. Bot. Gart. Berlin-Dahlem 13: 70 (1936) Areca novohibernica var. salomonensis Burret, Notizbl. Bot. Gart. Berlin-Dahlem 13: 69 (1936). Type: PAPUA NEW GUINEA, Bougenville Island, Kugumaru, Buin District, 2.vii.1930, Kajewski 1908 (holotype: B ; isotype: A). Distribution: Bismarck Archipelago and the Solomon Islands.

87 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA ARECA RIDLEYANA BECC. EX FURTADO Areca ridleyana Becc. ex Furtado, Repert. Spec. Nov. Reg. Veg. 33: 236 (1933). Type: MALAY PENINSULA, Johor, G. Janeng, 1892, Kelsall s.n. (holotype: SING). Distribution: Malay Peninsula. 38. ARECA SUBACAULIS (BECC.) J. DRANSF. Areca subacaulis (Becc.) J. Dransf., Kew Bull. 39: 20 (1984). Gigliolia subacaulis Becc., Malesia 1: 174 (1877). Pichisermolia subacaulis (Becc.) H. C. Monteiro, Rodriguésia 28: 196 (1976). Type: BORNEO, Sarawak, Gunung Matang, near Kuching, Beccari P.B (holotype: FI; isotypes: K). Distribution: Borneo. 39. ARECA TRIGINTICOLLINA HEATUBUN SP. NOV. (FIG. 7). Type: INDONESIA, Sumatra, Riau Province, Indragiri Hulu Regency, Seberida District, Bukit Tiga Puluh National Park, 10 km North of Jambi border, 250 m, 5.v.1991, Gundersen et al. 15 (holotype: K; isotype: BO). Diagnosis: a ceteris speciebus sumatranis Arecae ad subgenerum Arecam pertinens, superficialiter A. tunku J. Dransf. & Lim Chong Keat in habitu et A. jugahpunya J. Dransf. inflorescentiae structura similis, floribus staminatis spiraliter dispositis, differt. Description: Solitary, unarmed, monoecious, pleonanthic palm. STEM 2 5 m tall, 3 4 cm diam.; internodes 3 6 cm long. LEAVES in crown, pinnate, cm long (including petiole); sheath tubular, cm long and 6 10 cm wide, smooth, yellowishgreen to grey brown, covered in punctiform brown indumenta, sparse or very frequent near the petiole; crownshaft well defined, up to 50 cm long and 15 cm diam., sometimes swollen; petiole very close or lacking, channelled adaxially, rounded abaxially; rachis with adaxial longitudinal ridge, rounded abaxially; blade with irregularly arranged leaflets, 6 12 leaflets on each side, spacing between leaflets 6 7 cm; leaflets variable from narrow to

88 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 69 broad, leaflets near petiole shorter than others, c cm, slightly sigmoid, the middle leaflets about cm, with up to 4 folds, the terminal leaflets broad, cm, linear, tips pointed except for the terminal leaflets with notched tips, papery, slightly dark adaxially and pale abaxially. INFLORESCENCE infrafoliar, upright, cm long and 5 6 cm wide, protandrous, branching to 1 (sometimes 2) order; peduncle 2 5 cm long, cream to warm yellow; prophyll cm, borne about 1/2 way up the peduncle, lanceolate, 2-keeled, boat shaped, papery, cream to light brown, entirely enclosing the inflorescence, then splitting longitudinally and disintegrating into numerous fibres, still persistent after staminate anthesis; rachis cream-coloured; rachis bracts not persistent; rachillae c. 13, ( 15) cm long, very stiff, flattened, wide or swollen near base, zigzag, slightly congested. STAMINATE FLOWERS small, triangular, sickle-shaped when young to elongate at anthesis, mm, asymmetric, cream to pale brown; calyx fused, cup-shaped, c mm at anthesis, 3-lobed, the lobes triangular about mm; petals 3, fused near base, lanceolate, mm at anthesis, striate; stamens 6, c. 6.7 mm 0.6 mm, elongated, basifixed; anthers c mm, creamy, elongated, sagittate, shorter than filaments; filaments c mm, joined to anther from base to tip, dark brown; pistillode absent. PISTILLATE FLOWERS larger than the staminate, triangular, borne on the enlarged rachillae and distributed to half of rachillae length, 3 6 flowers on each rachilla, buds varying greatly in size depending on stage of development, mm long and 5 6 mm wide at anthesis; calyx with 3 sepals; sepals free, strongly imbricate, c mm at anthesis, triangular, asymmetrical; petals 3, imbricate, triangular, c mm at anthesis; staminodes absent. FRUITS (young) green, elongate, apical stigmatic remains, mature fruits not known. Distribution and Ecology: This species is only known so far from Bukit Tiga Puluh National Park in Riau Province in South Western of Sumatra, Indonesia. It was recorded growing on a ridge crest on primary forest at m above sea level. Local Name and Uses: Not recorded. Conservation Status: Data Deficient (IUCN, 2001). Further population study is needed to assess the conservation status of this palm.

89 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 70 Figure 7. Areca triginticollina Heatubun. A, Apical portion of leaf. B, Middle and basal portion of leaf. C, Basal portion of leaf and leaf sheath showing no petiole. D, Prophyll with disintegrating fibrous mass. E, Inflorescence. F, Details of rachilla showing staminate and pistilate flowers in bud and their arrangements. G, Pistillate flowers at anthesis on rachillae. H, I, Staminate flower whole and in section. J, Pistillate flower in section. Scale bar: A C = 6 cm; D, E = 3 cm; F = 1.5 cm; G = 2.5 cm; H J = 4 mm. A F, H J from Witono et al. 150; G from Gundersen et al. 15. Drawn by Lucy T. Smith.

90 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 71 Specimens Studied: INDONESIA. SUMATRA: Riau Province, Indragiri Hulu Regency, Seberida District, Bukit Tiga Puluh National Park, 10 km North of Jambi border, 250 m, 5.v.1991, Gundersen et al. 15 (holotype: K!, isotype: BO); 5 km West of Kelesa, 150 m, 15.v.1991, Gundersen et al. 32 (K!, BO); Talang Langkat village, 120 m, 9.viii.1991, Witono et al. 146 (K!, BO!); 150 m, 10.viii.1991, Witono et al. 150 (K!, BO!). Notes: Areca triginticollina is similar to A. tunku in many respects, including the littertrapping crown and the inflorescence sometimes bursting out from the persistent prophyll. However, they can be distinguished by inflorescence structure; A. triginticollina has swollen-flattened rachillae, which are folded in a zigzag fashion, this corresponding to the position of the pistillate flowers. Also the triads are more than one (3 6) for each rachilla and they are distributed along half of the rachilla length similar to A. jugahpunya in the same subgenus, whereas, A. tunku has erect, stiff and rigid rachillae and triads borne at the base of the rachillae only. The specific epithet refers to the type locality of this new palm in Taman Nasional Bukit Tiga Puluh in Riau, Sumatra, Indonesia. Triginticollina in latin mean thirty hills or bukit tiga puluh in bahasa Indonesia. 40. ARECA TUNKU J. DRANSF. & C. K. LIM. Areca tunku J. Dransf. & C. K. Lim, Principes 36: 81 (1992). Type: MALAY PENINSULA, Terengganu, Besut, Ulu Setiu Forest Reserve, viii.1977, Dransfield et al. JD 5178 (holotype: K; isotype: KEP). Areca bifaria Hodel, Palm J. 136: 7 (1997). Type: THAILAND, SW of Narathiwat, Hodel & Vatcharakorn 1753 (holotype: BK). Distribution: Thailand, Malay Peninsula and Sumatra. 41. ARECA VESTIARIA GISEKE Areca vestiaria Giseke, Prael. Ord. Nat. Pl.: 78 (1792). Pinanga vestiaria (Giseke) Blume, Rumphia 2: 77 (1839). Seaforthia vestiaria (Giseke) Mart., Hist. Nat. Palm. 3: 313 (1849).

91 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 72 Ptychosperma vestiarium (Giseke) Miq., Fl. Ned. Ind. 3: 31 (1855). Mischophloeus vestiarius (Giseke) Merr., Interpr. Herb. Amboin.: 121 (1917). Type: Pinanga sylvestris e Buro Rumphius, Herb. Amboin. 1: (Lectotype Dransfield, 1974). Ptychosperma paniculatum Miq., Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk. 11(5): 3 (1868). Areca paniculata (Miq.) Scheff., Tijdschr. Ned.-Indië 32: 179 (1873). Mischophloeus paniculatus (Miq.) Scheff., Ann. Jard. Buitenzorg 2: 152 (1876). Type: NORTH MOLUCCAS, Bacan Island, Teysmann & De Vriese (isotype: L). Areca leptopetala Burret, Notizbl. Bot. Gart. Berlin-Dahlem 13: 199 (1936). Type: SOUTH CELEBES, Porema, 10.ix.1929, Kjellberg 2324 (holotype: B, isotype: BO). Areca langloisiana Potztal, Willdenowia 2: 628 (1960). Type: A. C. Langlois s.n. (holotype: B). Distribution: Celebes and Moluccas. EXCLUDED AND UNCERTAIN NAMES Areca alba Bory, Voy. Îles Afrique 1: 306 (1804) = Dictyosperma album (Bory) Scheff., Ann. Jard. Bot. Buitenzorg 1: 157 (1876). Areca angulosa Giseke, Prael. Ord. Nat. Pl.: 80 (1792) = uncertain. Areca appendiculata F. M. Bailey, Dept. Agric. Bot. Div. Bull. 4: 18 (1891) = Oraniopsis appendiculata (F. M. Bailey) J. Dransf., A. K. Irvine & N. W. Uhl., Principes 29: 61 (1985). Areca angusta Kurz, J. Bot. 13: 331 (1875) = Rhopaloblaste angusta (Kurz) H. E. Moore, Principes 14: 79 (1970). Areca aurea Van Houtte, Ann. Gèn. Hort. 7: 43 (1867) = Dictyosperma album var. aureum Balf. f., Fl. Mauritius: 384 (1877).

92 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 73 Areca bacaba Arruda, in H. Koster., Trav. Brazil: 490 (1816), nom. nud. = Oenocarpus bacaba Mart. Hist. Nat. Palm. 2: 24 (1823). Areca banaensis (Magalon) Burret, Notizbl. Bot. Gart. Berlin-Dahlem 13: 198 (1936) = Nenga banaensis (Magalon) Burret, Notizbl. Bot. Gart. Berlin-Dahlem 13: 347 (1936). Areca banksii A. Cunn. ex Kunth., Enum. Pl. 3: 185 ( 1841), pro syn = Rhopalostylis sapida (Sol. ex G. Forst.) H. Wendl. & Drude, in O. C. E. Kerchove de Denterghem, Palmiers: 255 (1878). Areca baueri Hook.f. ex Lem., Ill. Hort. 15: 575 (1868) = Rhopalostylis baueri (Hook.f. ex Lem.) H. Wendl. & Drude, Bot. Zeitung (Berlin) 35: 638 (1877). Areca borbonica Kunth, Enum. Pl. 3: 186 (1841) = Dictyosperma album (Bory) Scheff., Ann. Jard. Bot. Buitenzorg 1: 157 (1876). Areca calapparia Blume, Rumphia 2: t. 100 (1843) = Actinorhytis calapparia (Blume) H. Wendl. & Drude ex Scheff., Ann. Jard. Bot. Buitenzorg 1: 156 (1876). Areca chaiana J. Dransf., Kew Bull. 39: 10 (1984) = uncertain. Based on molecular phylogenetic analysis in one gen RPB2 (Heatubun et al. in prep.), this species is a sister to all members of subtribe Arecinae (Areca, Nenga and Pinanga). Dransfield (1984) published this species and noted that this is an aberrant species of Areca, the only one with a truly spicate inflorescence. Harley & Dransfield (2003) also found that this species has unique pollen morphology in the palm family. Areca cincta Walp., Ann. Bot. Syst. 5: 808 (1860) = Acanthophoenix rubra (Bory) H. Wendl., Fl. Serres Jard. Paris 2(6): 181 (1866). Areca congesta Becc., Bot. Jahrb. Syst. 58: 441 (1923) = uncertain (see Heatubun et al. in prep.).

93 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 74 Areca cocoides Griff., Calcutta J. Nat. Hist. 4: 454 (1845) = Actinorhytis calapparia (Blume) H. Wendl. & Drude ex Scheff., Ann. Jard. Bot. Buitenzorg 1: 156 (1876). Areca communis Zipp. ex Blume, Rumphia 2: 73 (1839) = Drymophloeus litigiosus (Becc.) H. E. Moore, Principes 13: 76 (1969). Areca cornuta Giseke, Prael. Ord. Nat. Pl.: 81 (1792) = uncertain. Areca coronata Blume ex Mart., Hist. Nat. Palm. 3: 179 (1838) = Pinanga coronata (Blume ex Mart.) Blume, Rumphia 2: 83 (1839). Areca costata (Blume) Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 43(2): 200 (1874) = Pinanga coronata (Blume ex Mart.) Blume, Rumphia 2: 83 (1839). Areca crinita Bory, Voy. Îles Afrique 1: 307 (1804) = Acanthophoenix rubra (Bory) H. Wendl., Fl. Serres Jard. Paris 2(6): 181 (1866). Areca cuneifolia Stokes, (Bot. Mag. Med. 2: 318 (1812) = uncertain. Areca curvata Griff., Ic. Pl.Asiat. 3: t. 248 (1851) = Pinanga disticha (Roxb.) H. Wendl., in O. C. E. de Kerchove de Denterghem, Palmiers: 253 (1878). Areca curvata Griff., Not. Pl. Asiat. 3: 164 (1851) = Pinanga paradoxa (Griff.) Scheff., Tijdschr. Ned.-Indië 32: 179 (1871). Areca dicksonii Roxb., Fl. Ind. ed. 1832, 3: 616 (1832) = Pinanga dicksonii (Roxb.) Blume, Rumphia 2: 77 (1839). Areca disticha Roxb., Fl. Ind. Ed. 1832, 3: 620 (1832) = Pinanga disticha (Roxb.) H. Wendl., in O. C. E. de Kerchove de Denterghem, Palmiers: 253 (1878).

94 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 75 Areca elaeocarpa Reinw. ex Kunth, Enum. Pl. 3: 195 (1841) = Drymophloeus oliviformis (Giseke) Mart., Hist. Nat. Palm. 3: 314 (1849). Areca erythrocarpa H. Wendl., in O. C. E. de Kerchove de Denterghem, Palmiers: 231 (1878) = Cyrtostachys renda Blume, Bull. Sci. Phys. Nat. Nèerl. 1: 66 (1838). Areca erythropoda Miq., J. Bot. Nèerl. 1: 6 (1861) = Cyrtostachys renda Blume, Bull. Sci. Phys. Nat. Nèerl. 1: 66 (1838). Areca flavescens Voss, Vilm. Blumengärtn. Ed. 3, 1: 1153 (1895) = Dypsis lutescens (H. Wendl.) Beentje & J. Dransf., Palms Madagascar: 212 (1995). Areca furfuracea H. Wendl., in O. C. E. de Kerchove de Denterghem, Palmiers: 231 (1878) = Dictyosperma album (Bory) Scheff., Ann. Jard. Bot. Buitenzorg 1: 157 (1876). Areca gigantea H. Wendl., in O. C. E. de Kerchove de Denterghem, Palmiers: 254 (1878) = Pinanga rumphiana (Mart.) J. Dransf. & Govaerts, World Checklist Palms: 178 (2005). Areca glandiformis Lam., Encycl. 1: 241 (1783) = uncertain. Areca globulifera Lam., Encycl. 1: 895 (1783) = Pinanga globulifera (Lam.) Blume, Bull. Sci. Phys. Nat. Nèerl. 1: 65 (1838). Areca gracilis Thouars ex Kunth, Enum. Pl. 3: 188 (1841), pro syn. = Dypsis pinnatifrons Mart., Hist. Nat. Palm. 3: 180 (1838). Areca gracilis Roxb., Fl. Ind. ed. 1832, 3: 619 (1832), nom. illeg. = Pinanga gracilis Blume, Rumphia 2: 77 (1839).

95 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 76 Areca gracilis Buch.-Ham., Mem. Wern. Nat. Hist. Soc. 5(2): 310 (1826) =? Pinanga gracilis Blume, Rumphia 2: 77 (1839). Areca haematocarpon Griff., Not. Pl. Asiat. 3: 184 (1851) = Pinanga malaiana (Mart.) Scheff., Tijdschr. Ned.-Indië 32: 175 (1871). Areca henrici Furtado, Repert. Spec. Nov. Regni Veg. 33: 232 (1933) = uncertain. See notes Dransfield (1974) and Heatubun et al. (in prep.). Areca herbstii W. Watson, Gard. Chron., n. s., 22: 426 (1884) = Acanthophoenix rubra (Bory) H. Wendl., Fl. Serres Jard. Paris 2(6): 181 (1866). Areca hexasticha Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 43(2): 201 (1874) = Pinanga hexasticha (Kurz) Scheff., Ann. Jard. Bot. Buitenzorg 1: 148 (1876). Areca himalayana Griff. ex H. Wendl. in O.C.E.de Kerchove de Denterghem, Palmiers: 231 (1878). Nom. nud. Areca horrida Griff., Calcutta J. Nat. Hist. 5: 465 (1845) = Oncosperma horridum (Griff.) Scheff., Tijdschr. Ned.-Indië 32: 191 (1871). Areca humilis Roxb. ex H. Wendl., in O. C. E. de Kerchove de Denterghem, Palmiers: 231 (1878) = Pinanga disticha (Roxb.) H. Wendl., in O. C. E. de Kerchove de Denterghem, Palmiers: 253 (1878). Areca humilis Blanco ex H. Wendl. in O. C. E. de Kerchove de Denterghem, Palmiers: 231 (1878). = uncertain. Areca humilis Willd. = Areca oryziformis var. saxatilis = = uncertain basionim. Areca ilsemannii Andrè, Rev. Hort. 70: 261 (1898) = uncertain.

96 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 77 Areca lactea Miq., Fl. Ned. Ind. 3: 10 (1855) = Dictyosperma album (Bory) Scheff., Ann. Jard. Bot. Buitenzorg 1: 157 (1876). Areca lansiformis Giseke, Prael. Ord. Nat. Pl.: 81 (1792) = uncertain. Areca latisecta (Blume) Scheff., Tijdschr. Ned.-Indië 32: 168 (1871) = Pinanga latisecta Blume, Rumphia 2: 79 (1839). Areca litoralis Blume, Rumphia 2: 123 (1843), nom. inval. = Drymophloeus litigiosus (Becc.) H. E. Moore, Principes 13: 76 (1969). Areca lutescens Bory, Voy. îles Afrique 2: 296 (1804) = Hyophorbe indica Gaertn., Fruct. Sem. Pl. 2: 186 (1791). Areca madagascariensis Mart., Hist. Nat. Palm. 3: 179 (1838) = uncertain. Areca malaiana (Mart.) Griff., Calcutta J. Nat. Hist. 5: 457 (1845) = Pinanga malaina (Mart.) Scheff., Tijdschr. Ned.-Indië 32: 175 (1871). Areca micholitzii Sander, Cat. 1895: 46 (1895) = uncertain. Areca microspadix Burret, Notizbl. Bot. Gart. Berlin-Dahlem 13: 198 (1936) = Nenga banaensis (Magalon) Burret, Notizbl. Bot. Gart. Berlin-Dahlem 13: 347 (1936). Areca minor W. Hill, Rep. Brisbane Bot. Gard.: 6 (1874) = Linospadix minor (F. Muell.) F. Muell., Fragm. 11: 58 (1878). Areca monostachya Mart., Hist. Nat. Palm. 3: 178 (1838) = Linospadix monostachya (Mart.) H. Wendl. & Drude, Linnaea 39: 199 (1875). Areca nenga Blume ex Mart., Hist. Nat. Palm. 3: 179 (1838) = Nenga pumila (Blume) H. Wendl., in O. C. E. de Kerchove de Denterghem, Palmiers: 251 (1878).

97 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 78 Areca nibung Mart., Hist. Nat. Palm. 3: 173 (1838) = Oncosperma tigillarium (Jack) Ridl., J. Straits Branch Roy. Asiat. Soc. 33: 173 (1864). Areca nigra Giseke ex H. Wendl. in O.C.E.de Kerchove de Denterghem, Palmiers: 231 (1878). Nom. nud. Areca nobilis auct., Gard. Chron. 1868: 349 (1868), pro. syn. = Nephrosperma vanhoutteanum (H. Wendl. ex Van Houtt.) Balf.f., in J. G. Baker, Fl. Mauritius: 386 (1877). Areca normanbyi F. Muell., Fragm. 8: 235 (1874) = Normanbya normanbyi (F. Muell.) L. H. Bailey, Gentes Herb. 2: 188 (1930). Areca oleracea Jacq., Select. Stirp. Amer. Hist.: 278 (1763) = Roystonea oleracea (Jacq.) O. F. Cook, Bull. Torrey Bot. Club 28: 554 (1901). Areca oliviformis Giseke, Ord. Nat. Pl. 79 (1830) = Drymophloeus oliviformis (Giseke) Mart., Hist. Nat. Palm. 3: 314 (1849). Areca oliviformis var. gracilis Giseke, Prael. Ord. Nat. Pl.: 80 (1792) = Drymophloeus oliviformis (Giseke) Mart., Hist. Nat. Palm. 3: 314 (1849). Areca oriziformis Gaertn., Fruct. Sem. Pl. 1: 20 (1788) = Pinanga globulifera (Lam.) Blume, Bull. Sci. Phys. Nat. Nèerl. 1: 65 (1838). Areca oriziformis var. gracilis Giseke, Prael. Ord. Nat. Pl.: 80 (1792) = Pinanga coronata (Blume ex Mart.) Blume, Rumphia 2: 83 (1839). Areca oriziformis var. saxatilis Burm.f. ex Giseke = uncertain. Areca oviformis Giseke, Prael. Ord. Nat. Pl.: 81 (1792) = uncertain.

98 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 79 Areca paradoxa Griff., Calcutta J. Nat. Hist. 5: 463 (1845) = Pinanga paradoxa (Griff.) Scheff., Tijdschr. Ned.-Indië 32: 179 (1871). Areca passalacquae Kunth, Ann. Sci. Nat. (Paris) 8: 420 (1826) = Medemia argun (Mart.) Wurttenb. ex Wendl., Bot. Zeitung (Berlin) 39: 93 (1881). Areca pisifera Lodd. ex Hook.f. Rep. Progr. Condition Roy. Bot. Gard. Kew 1882: 54 (1884) = Dictyosperma album (Bory) Scheff., Ann. Jard. Bot. Buitenzorg 1: 157 (1876). Areca procera Zipp. ex Blume, Rumphia 2: 45 (1843) = Hydriestele procera (Blume) W. J. Baker & Loo, Kew Bull. 59: 67 (2004). Areca propia Miq., Fl. Ned. Ind. 3: 10 (1855) = Dictyosperma album (Bory) Scheff., Ann. Jard. Bot. Buitenzorg 1: 157 (1876). Areca pumila Blume, Rumphia 2: 71 (1839) = Nenga pumila (Blume) H. Wendl., in O. C. E. de Kerchove de Denterghem, Palmiers: 251 (1878). Areca punicea Zipp. ex Blume, Rumphia 2: 79 (1839) = Pinanga rumphiana (Mart.) J. Dransf. & Govaerts, World Checklist Palms: 178 (2005). Areca purpurea auct., Ill. Hort. 24: t. 298 (1877) = uncertain. Areca rubra H. Wendl., in O. C. E. de Kerchove de Denterghem, Palmiers: 232 (1878) = Dictyosperma album (Bory) Scheff., Ann. Jard. Bot. Buitenzorg 1: 157 (1876). Areca rubra Bory, Voy. Îles Afrique 1: 306 (1804) = Acanthophoenix rubra (Bory) H. Wendl., Fl. Serres Jard. Paris 2(6): 181 (1866).

99 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 80 Areca sanguinea Zipp. ex Blume, Rumphia 2: 72 (1839) = Pinanga rumphiana (Mart.) J. Dransf. & Govaerts, World Checklist Palms: 178 (2005). Areca sapida Sol. ex G. Forst., Pl. Esc.: 66 (1786) = Rhopalostylis sapida (Sol. ex G. Forst.) H. Wendl. & Drude, in O. C. E. Kerchove de Denterghem, Palmiers: 255 (1878). Areca sechellarum (H. Wendl.) Baill., Hist. Pl. 13: 348 (1895) = Phoenicophorium borsigianum (K. Koch) Stuntz, U.S.D.A. Bur. Pl. Industr. Invent. Seeds 31: 88 (1914). Areca speciosa Lem., Ill. Hort. 13: t. 462 (1866) = Hyophorbe amaricaulis Mart., Hist. Nat. Palm. 3: 309 (1849). Areca spicata Lam., Encycl. 1: 241 (1783) = Calyptrocalyx spicatus (Lam.) Blume, Rumphia 2: 103 (1843). Areca spinosa Hasselt & Kunth, Enum. Pl. 3: 185 (1841) = Oncosperma tigillarium (Jack) Ridl., J. Straits Branch Roy. Asiat. Soc. 33: 173 (1864). Areca sylvestris Lour., Fl. Cochinch. 2: 568 (1790) = Pinanga sylvestris (Lour.) Hodel, Palm J. 139: 55 (1998). Areca tigillaria Jack, Malayan Misc. 2(7): 88 (1820) = Oncosperma tigillarium (Jack) Ridl., J. Straits Branch Roy. Asiat. Soc. 33: 173 (1864). Areca vaginata Giseke, Prael. Ord. Nat. Pl.: 78 (1792) = Drymophloeus oliviformis (Giseke) Mart., Hist. Nat. Palm. 3: 314 (1849). Areca verschaffeltii Lem., Ill. Hort. 13: t. 462 (1866) = Hyophorbe verschaffeltii H. Wendl., Ill. Hort. 13(1): t. 462 (1866). Areca wallichiana Mart., Hist. Nat. Palm. 3: 178 (1838) = Iguanura wallichiana (Mart.) Becc., Malesia 3: 100 (1886).

100 HEATUBUN ET AL.: A CONSPECTUS OF THE GENUS ARECA 81 Areca wendlandiana (Scheff.) H. Wendl., in O. C. E. de Kerchove de Denterghem, Palmiers: 332 (1878) = Nenga pumila (Blume) H. Wendl., in O. C. E. de Kerchove de Denterghem, Palmiers: 251 (1878). Drymophloeus vestiarius Miq., Palm. Archip. Ind.: 24 (1868). Nom. inval. ACKNOWLEDGEMENTS Many individuals and institutions contributed to the completion of this paper. We would like to thank the Keepers and staff of herbaria AAU, BO, E, FI, K, L, MAN and SAR for access to their specimens, databases and for providing loans in many cases. The authorities in Sarawak, especially the Sarawak Forestry Company (SFC) allowed CDH to conduct his fieldtrip in Sarawak. Julia Sang and Shahabuddin M. Shabki are thanked for their helps with permits and all support including the organisation of CDH s fieldtrip in Sarawak. Tom D. Evans sent his collection of a new Areca from Cambodia. Lucy T. Smith rendered beautiful illustrations for the new species funded by Royal Botanic Gardens Kew. This paper is part of CDH PhD thesis, conducted at Institut Pertanian Bogor and Royal Botanic Gardens Kew. CDH would like to express his gratitude to Dr. Sri S. Tjitrosoedirdjo, Prof. Johanis P. Mogea, and Prof. Mien A. Rifai to their generous encouragement, support, help and guidance during the study. Financial support came from International Palm Society in funding the field trip to Sarawak through IPS Endowment Fund The Royal Botanic Gardens Kew, British American Tobacco Biodiversity Partnership and the Indonesian Minister of High Education (BPPS Dikti Diknas) funded CDH s PhD; to them are greatly acknowledged.

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106 A Monograph of Areca of East of Wallace s Line CHARLIE D. HEATUBUN FLS 1, 2, 3, 5, WILLIAM J. BAKER 3, SRI S. TJITROSOEDIRDJO 2, JOHANIS P. MOGEA 4, and JOHN DRANSFIELD 3 1 Faculty of Forestry, Universitas Papua, Jl. Gunung Salju, Amban, Manokwari 98314, Papua Barat, Indonesia 2 Biology Department, Sekolah Pascasarjana Institut Pertanian Bogor, Darmaga, Bogor 16680, Jawa Barat, Indonesia 3 Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, United Kingdom 4 Herbarium Bogoriense, Puslitbang Biologi LIPI, Jl. Ir. H. Juanda No. 22, Bogor, Jawa Barat, Indonesia 5 Author for correspondence (charlie_deheatboen@yahoo.com) Summary. This monograph provides a review of the current taxonomic status of the palm genus Areca of east of Wallace s line together with ecological information, uses and conservation status. Six species of Areca are recognised in this treatment, including the widespread cultivated species A. catechu L. the betel nut palm. Five species are native to areas east of Wallace s line, namely A. macrocalyx Zipp. ex Blume, A. mandacanii Heatubun, A. novohibernica (Lauterb.) Becc., A. oxycarpa Miq. and A. vestiaria Giseke. Of the twelve previously accepted species, five varieties and one form are treated as new synonyms, with two species considered as doubtful taxa. ADDITIONAL KEYWORDS: Areca palms taxonomy Wallace s line conservation natural history.

107 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 89 INTRODUCTION The centre of diversity of the genus Areca is in Malesia, and within this area there are three regional hotspots for the genus that also reflect palm diversity in general in Southeast Asia: west of Wallace s line is primary centre of diversity; the second centre of diversity is east of Wallace s line (EWL); and the third in the Philippines. Wallace s line is the imaginary biogeographical line which first recognised by English naturalist A. R. Wallace in 1863 and named later by Thomas Henry Huxley (Johns, 1995). This biogeographical boundary is separating Asian and Australian biota and running between the island of Bali and Lombok and northwards through the Makassar straits separating the island of Sulawesi and Borneo. Although this demarcation is very significant in fauna, but in some study in vascular plants also revealed that the distributions of flora in this region follow this line (Van Steenis, 1950, 1979; Van Balgooy, 1984; Baker et al., 1998), especially in palms (Dransfield, 1981, 1987). A thorough revision of the entire genus is needed. As a step towards this goal, we present here an account of the genus in its EWL hotspot, the area in which the genus is perhaps most poorly understood. Thus, we focus here on species of Areca from Sulawesi, the Moluccas, New Guinea and the Solomon Islands. The taxonomic history of Areca was described in detail in Furtado (1933), especially the limits of the genus and infrageneric classification, including the amalgamation of Mischophloeus Scheff. into Areca. Dransfield (1984) included the genus Pichisermollia H.C.Monteiro within Areca. The historical understanding of Areca as a palm genus is closely connected to the EWL hotspot and the Moluccas Islands in particular. Dating back to Rumphius time, this genus first appeared as Pinanga in Herbarium Amboinense in 1741, and was used by Linnaeus as basis of the genus Areca, together with its species A. catechu L. in his Species Plantarum in 1753 based on Rumphius s plate IV; this plate was subsequently selected by Moore & Dransfield (1979) as the lectotype for A. catechu The second species from the region, A. vestiaria Giseke was published by Giseke (1792) also based on Rumphius s Pinanga (Dransfield, 1974) and followed by 16 other species which accepted in the World Checklist of Palms (Govaerts & Dransfield, 2005and updated in the online World Checklist of Monocotyledons : The other sixteen accepted species of the genus Areca based on Govaerts & Dransfield (2005) are A.

108 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 90 macrocalyx Zipp. ex Blume (Blume, 1839); A. oxycarpa Miq. (Miquel, 1868); A. jobiensis Becc. (Beccari, 1877); A. rechingeriana Becc. (Beccari, 1910); A. nigasolu Becc., A. guppyana Becc., A. torulo Becc., A. warburgiana Becc, A. novohibernica (Lauterb.) Becc. (Beccari, 1914a, b); A. congesta Becc., A. ledermanniana Becc. (Beccari, 1923); A. nannospadix Burret (Burret, 1931); A. celebica Burret (Burret, 1933); A. rostrata Burret (Burret, 1935); A. multifida Burret and A. salomonensis (Burret, 1936). Furtado (1933) divided Areca into two subgenera (subgenus Beccarioareca Furtado and Blumeoareca Furtado) based on the arrangement of the pairs of the staminate flowers; being spiral in Beccarioareca and unilateral or distichous in Blumeoareca. In addition, he established two sections in Beccarioareca based on the number of stamens, the number of branching orders of the inflorescence, and dimensions of the fruits: section Microareca Furtado has six or more stamens, always a simple-branching inflorescence, fruits 2 3 longer than wide and with the inflorescence and the staminate flowers very small in general; section Mischophloeus (Scheff.) Becc. always has six stamens, the inflorescence is simple or branched to 2 3 orders, the fruits are ovoid to ellipsoid, with the inflorescences and staminate flowers generally large. Subgenus Blumeoareca has three sections, namely Arecella Wendl. & Drude, Oeotheanthe (Scheff.) Furtado and Axonianthe Scheff. This separation is mainly based on the inflorescence branching order, the arrangement of staminate flowers, the number of stamens and the position of pistillate flowers on rachilla or triads. Section Arecella differs from the two other sections in the alternate-arrangement of the staminate flowers on the rachilla and 3 to 6 stamens rather than a distichous-arrangement of the staminate flowers always with six stamens.section Oeotheanthe differs from section Axonianthe in the inflorescence being branched to 2 3 orders, divaricate branches, and the pistillate flowers or triads at the base near the branches. Section Axonianthe has the inflorescence branched to 1 order (simple), branching near to primary axis, the pistillate flowers or triads borne on the main axis or at the base of the branches and congested around the main axis. In addition, Furtado (1933) also gave an enumeration of the species of Areca in each section on his synopsis; 15 of the 17 species recognised by Furtado of east of Wallace s line are still accepted as valid in the World Checklist of Palms (Govaerts & Dransfield, 2005; For nomenclatural reasons, Harley & Dransfield (2003) corrected Furtado s subgenus Blumeoareca and the section Oeotheanthe Scheff. to subgenus Areca and section Areca to follow nomenclatural rules.

109 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 91 Figure 1. Distribustion map of the species Areca which native to east of Wallace s line. Areca mandacanii (solid square), Areca macrocalyx (rounded dots), Areca novohibernica (dash lines), Areca oxycarpa (square dots) and Areca vestiaria (solid line). There was no important work on Areca in the east of Wallace s line for almost 70 years, although Eva Potztal (1960) described A. langlosiana Potztal from North Celebes (Sulawesi) but this was regarded as superfluous and treated as synonym of A. vestiaria by Dransfield (1974). Subsequently, Flynn (2004), in an unpublished Master thesis, followed Furtado s infrageneric classification and reassessed species delimitation of Areca in New Guinea and the Solomon Islands using a morphometric approach, proposing a reduction of seven species of section Axonianthe to one species (A. macrocalyx Zipp. ex Blume), and two species in section Mischophloeus to one (A. guppyana Becc.). Despite this broad species concept and its delimitation of Areca in New Guinea, Heatubun (2008) discovered a new species from the Bird Head Peninsula in Western New Guinea Areca mandacanii Heatubun, notable in being the first New Guinean species of Areca to have a close relationship with the widely cultivated betel nut palm A. catechu.

110 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 92 MATERIALS AND METHODS Taxon sampling and observations for the morphology and distribution of the species were based on herbarium samples or specimens (dried and spirit-preserved materials) deposited at international herbaria, namely A, AAU, B, BH, BO, BRI, FI, K, KEP, L, LAE, MAN, PNH, SAN, SAR, and SING (herbarium acronyms follow Holmgren et al., 1990), as well as the newly established small herbarium in Balai Penelitian Kehutanan (Forestry Research Institute) in Manokwari, West Papua, Indonesia that will appear in specimens citations as BPK Manokwari. Many new specimens have been collected recently by the authors, particularly from western New Guinea (the Indonesian Provinces of Papua and West Papua) and North Sulawesi by the first author in Specimens were made in the field using standard preparation guidelines to the palms (Dransfield, 1986; Baker & Dransfield, 2006). Measurements were taken from spirit-preserved material and dried herbarium specimens and from living collections. Floral parts were measured from spirit-preserved material or dried specimens and rehydrated by boiling. Basic morphological characters such as habit, stem, leaves, inflorescence, staminate flower, pistillate flower, fruit, seed and their details were used to describe and recognize taxa; all morphological data was used for producing the descriptions of each taxon, while the key to species was constructed from the diagnostic characters only. The morphological species concept or taxonomic species concept was applied as a framework to define taxa, and assessed later with a phylogenetic species concept (Davies & Heywood, 1963; McDade, 1995; Gornall, 1997; Mayden, 1997; Dransfield, 1999), especially based on the result of molecular phylogenetics analysis of the genus Areca (Heatubun et al., in prep.). The conservation status of each species of the genus Areca in east of Wallace s line was assessed based on the IUCN red list categories and criteria version 3.1 (IUCN, 2001).

111 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 93 RESULTS AND DISCUSSION MORPHOLOGY Habit This character, together with the certain floral characters, was used by Furtado (1933) to define sections in his infrageneric classification, specifically sections Microareca and Mischophloeus. Section Microareca was created to accommodate the small or miniature taxa of Areca west of Wallace s line, while section Mischophloeus included those species of Areca from east of Wallace s line with a moderately sized habit, and even relatively big palms. However, these sections are not recognisable in the new conspectus of the genus based on molecular phylogenetics (Heatubun et al., in prep.). In general, the habit of Areca species can be very variable, from acaulescent undergrowth-palmlets to robust-emergent tree palms, and from solitary to clustering (Dransfield, 1984; Uhl & Dransfield, 1987; Dransfield et al., 2008). In the EWL hotspot, taxa of Areca can be slender small palms (e.g., A. macrocalyx and A. oxycarpa) to robust tree palms (e.g., A. macrocalyx and A. catechu), but never have an acaulescent habit, as in several species in Borneo (Dransfield, 1980, 1984; Heatubun et al., in prep.). All species in this region are solitary; except for A. vestiaria (although in some population of A. vestiaria is found also solitary). Two species of Areca in the region have prop roots or stilt-roots, namely A. novohibernica and A. vestiaria. Although the ecological significance of stilt roots has been debated (Dransfield et al., 2008), this phenomenon is probably related to the habitat where the palm grows, usually on slopes and/or in habitats which are temporarily or permanently inundated by water. These roots are essential to mechanical and vascular support (Frangi & Ponce, 1985; Tomlison, 1990), but never for walking as thought by Leopold (2000). According to Tomlinson (1990), stilt-roots usually occur in solitary palms (like A. novohibernica) to support the trunk, but in some populations of A. vestiaria can be clustering with stilt-roots this is also found in a new species of Areca from Borneo (Heatubun et al., in prep.). Clustering or solitary and with or without stilt roots (prop roots) are important and relatively constant characters that are very useful for species level taxonomy of Areca in

112 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 94 this region. These two characters are very practical and significant, especially when recognizing species in the field. Areca macrocalyx is an exceptional species in the east of Wallace s line hotspot, with a wide range in size and habit from being small and slender to robust-canopy palms, but is consistent in having a solitary habit without stilt roots. Stems The stem size varies from very slender (1 1.5 cm in diam.) in A. oxycarpa and A. macrocalyx to moderate (15 20 cm in diam.) in A. catechu and A. macrocalyx. The internodes or leaf scars are prominent, obvious and sometimes contrast strongly with the shiny green stem. Internode length is variable, and is related to the physiology of the palm and the habitat where the individual palm grows. Stem colour is shiny green (or yellowish green) near the stem tip and becomes brown to whitish near the base. Elongated internodes with a shiny black stem have been observed in the populations of A. macrocalyx from the heath forest in the centre of Japen Island, New Guinea. The stems of Areca species lack the black fibres as found in other genera of palm in the east of Wallace s line hotspot, especially in New Guinea, such as Actinorhytis, Dransfieldia, Drymophleous, Ptychococos or Rhopaloblaste. Without the fibres, the stems of Areca are softer than the stems of those palms with fibres; this is why the stem of Areca has limited properties for local use when compared to the palm genera mentioned above. The stem anatomy of the genus has been studied in detail from the cultivated widespread species A. catechu by Tomlison (1961), and previously by Schoute (1912) from the west of Wallace s line species of A. triandra. Leaves All species of Areca in the EWL hotspot have pinnately-ribbed leaves, contrasting strongly with the simple-bifid leaves of several species of Areca in west of Wallace s line in Borneo (Dransfield, 1980, 1984; Heatubun et al. in prep.). The number of leaves in the crown is ranges from 5 to 11 in almost all species in this region. There is a correlation between the number of leaves in the crown and the crown-shape in palms. Three general shapes of crown have been recognized in palms (Tomlinson, 1990) - especially tree palms: the spherical crown-shape, the hemispherical crown-shape and the shuttle-cock shape; all occur in Areca. The shuttle-cock crown shape is a result of very few leaves in the crown (Tomlinson, 1990) and is very useful at species level as in the genus Cyrtostachys

113 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 95 (Heatubun et al., 2009). However, it appears not to work well in Areca, especially in A. catechu as all three types of crown-shape have been observed in the cultivars of the betel nut in this region, as well as in the highly variable A. macrocalyx. The petiole ranges from lacking or very short (up to 5 cm long) to long (then about c. 50 cm long). From six species of Areca of EWL, only three species (A. catechu, A. macrocalyx and A. mandacanii) have a very short petiole and three species (A. novohibernica, A. oxycarpa and A. vestiaria) have a relatively long petiole. The arrangement of leaflets is regular to irregular in Areca, but all species in the east of Wallace s line are irregular and located in the same plane as the leaf rachis, except A. mandacanii (Heatubun, 2008). This unique species has plumose leaves, with leaflets are irregular with a single-folde and located in a different plane or several ranked in the leaf rachis. The plumose leaf has previously never be reported in the genus Areca or even in the subtribe Arecinae and is very rare in tribe Areceae. Only the Australian monotypic genera Normanbya and Wodyetia of tribe Areceae and several species of South American genera Dictyocaryum, Iriartea, Socratea and Wettinia from tribe Iriarteeae have plumose leaves; this division of leaflets is unusual in palms in general (Dransfield et al., 2008). Structurally, the leaves become plumose by the longitudinal splitting of a single leaflet (Dransfield et al., 2008), but this process is slightly modified in A. mandacanii, in which the broad leaflets splits longitudinally to produce a single folded-leaflet with a prominent midrib, and the veins running parallel to midrib from the base to the acuminate-notch tip. In contrast to other plumose-leaved palms, the splitting comes from the relatively similar sized of leaflets and the veins divergent from base to praemorse tip, and with the midrib not prominent. The total number of leaflets on one side of the leaf ranges from 5 to 75 in east of Wallace s line Areca, with the lowest number of leaflets in A. novohibernica and A. oxycarpa, and the highest number in A. macrocalyx. The number of leaflets is very variable within the genus and shows a high degree of phenoplasticity, and sometimes the variation is gradual and continues in the leaf plication, like in A. macrocalyx, from a few numbers of broad-leaflets to many single-folded leaflets. However, Areca mandacanii always has single-folded leaflets and is therefore striking among the EWL hotspot species of Areca. The crownshaft is formed by long tubular leaf sheaths and in Areca; the crownshaft is well defined and is one of the main characters of the genus. The crownshaft is dark brown to black only in A. oxycarpa, yellow to orange in A. vestiaria (sometimes reddish), and

114 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 96 green in all other species. Although a red crownshaft (bright red to reddish brown) has been reported in A. macrocalyx,this species has a green crownshaft in general. The leaf sheath varys from thin in small-undergrowth palms to thick in emergent tree palms, and sometimes has disintegrating fibres at the margins just below the petiole. The leaf anatomy of Areca has been discussed by Tomlinson (1961) based on his observation on betel nut A. catechu and the Sri Lankan species A. concinna. Indumentum Indumentum plays a minor contribution to the taxonomy of Areca in this region, although it occurs on leaf sheaths, leaves, inflorescences and staminate flowers. There are several types of indumentum present: fine purple scales, filamentous brown scales and black thick scales occur on leaf sheaths and petioles; dense white, greyish and brown scales occur on the leaf rachis; the inflorescences, especially on peduncle and rachilla of A. mandacanii, carry minute red to brown dots and a light brown to chocolate brown thick indumentum. The velvet indumentum which occurs on the staminate flowers is the one of the important characters to differentiate A. novohibernica from A. vestiaria. Inflorescences The inflorescence of all species of Areca of east of Wallace s line is infrafoliar, although interfoliar inflorescences are found in several species of Areca in Borneo (Beccari, 1877; Dransfield, 1984). Only A. vestiaria is always branched to two orders, three other species are usually branched to one order (A. macrocalyx, A. novohibernica and A. oxycarpa), and two species arebranched from two to three orders (A. catechu and A. mandacanii). Occasionally, the basal most rachillae of the first order branched-species branch again. Mostly species in this region have erect inflorescences in the early stages of development but will become pendulous when the fruits have been developed; however, A. oxycarpa always has erect inflorescences even when bearing ripe fruits. Areca macrocalyx has some populations with an always erect inflorescence, other populations are always pendulous, and combinations of both in other populations, but generally, the inflorescence of A. macrocalyx is erect in the early stages of development and becomes pendulous later. This observation indicates that the erect or pendulous inflorescence is not a good taxonomic character to use especially in complex species, like A. macrocalyx.

115 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 97 Figure 2. The inflorescence (infructescence) and the pistillate flowers morphology of Areca macrocalyx. A, Laxly and pendulous inflorescence. B and C, Crowded and erect inflorescence. D, Long pendulous, laxly and paired fruits. E, Pendulous with more than one triad on basalmost-rachilae. F, Turning from erect to pendulous. G, Pistillate flowers at anthesis, position of corolla lower than stigma (St). Scale bar A = 10 cm, B, C and F = 5 cm, D = 20 cm, E = 25 cm, G = 1 cm. Photos by: A (William J. Baker); B (John Dransfield); C G (Charlie D. Heatubun). The prophyll is thin, membranous and encloses the inflorescence in bud, quickly splitting and falling, other bract very inconspicuously and appearing no additional bract. A single large bract in the inflorescence is an important character that defines the genus Areca (Dransfield et al., 2008). The prophyll, or bracts, varies in colour, and is generally similar to the crownshaft or leaf sheath colour; a bright red prophyll has been observed in A. macrocalyx populations although the crown shaft is brown. The peduncle is shorter than the rachis in all species in this region. The rachillae are glabrous, and the rachillae bracts are tiny. The rachillae bear solitary or paired staminate flowers arranged spirally, distichously, or in two approximate rows on one side of rachilla the rachilla tips are sometimes devoid of flowers (Dransfield et al., 2008). Only A. macrocalyx has less number of triads which born at the base of rachillae and congested to the rachis, then the rachillae will dry and drop off when the fruits have been developed.

116 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 98 Therefore, the rachis bearing congested and compacted fruits is peculiar and gives the appearance of a spicate-like inflorescence (Fig. 2). Flowers In common with almost all Arecoideae, the flowers are arranged in triads with a central pistillate flower and two lateral staminate flowers (Heatubun et al., 2009), borne usually at the base of the rachilla (Furtado, 1933; Dransfield, 1984; Uhl & Dransfield, 1987; Dransfield et al., 2008). According to Dransfield et al. (2008), triads in Areca are confined to the proximal part of the main axis, or to the proximal part of each order of branching, and only rarely on a subdistal part of the main axis. The dispositions of triads in the rachilla are important taxonomically when defining species in Areca, especially in the east of Wallace s line hotspot. Numerous triads occur along the half length of the rachillae are found in A. novohibernica and A. vestiaria; few triads occurring along a 1/3 of the length of the rachillae are found only in A. oxycarpa; and only one triad seated in each node of rachillae branches is found in A. catechu and A. mandacanii. Areca macrocalyx has more than one triad (up to 5) on the basal most rachilla, and only one triad along the rest of rachillae. In common with many palms, the flowers in Areca are sexually dimorphic. The staminate or male flowers are smaller than the pistillate or female flowers. In species in the east of Wallace s line hotspot, the staminate flowers are cream-coloured in the bud and white to yellowish-white at anthesis. The pistillate flowers are cream-coloured to white or greenish-white with green at the tip in bud, and cream-coloured to green with creamcoloured near the tip at the anthesis. The staminate flowers are an important character in Areca, not only for its taxonomic information to distinguish taxa but also to reveal the evolutionary lines within the genus. The arrangement of staminate flowers was used by Furtado (1933) to develop his infrageneric classification in Areca as subgeneric character staminate flowers arranged spirally versus a uniseriate and/or distichous arrangement. And this morphology character supports separation of two lineages that have recently been recovered in the recent molecular phylogenetic study of the genus Areca (Heatubun et al., in prep.). The arrangement is spiral in A. novohibernica and A. vestiaria, while in the other species of Areca in the east of Wallace s line hotspot are distichous. The staminate flowers have three, distinct, slightly imbricate, triangular sepals or a cupular calyx with three triangular lobes; the corolla has three triangular to elongated

117 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 99 triangular or slightly spathulate, valvate petals much longer than the sepals. All species in this region have six free stamens, except for the epipetalous stamens of A. novohibernica. In male flowers the filaments are short to elongated with linear to sinuous anthers, sometimes very irregular, latrorse; the pistillode is present and conspicuous as a trifid column as long as the stamens, but can be and minute or absent. Two distinct groups can be revealed easily based on the nature of the staminate flowers among species of Areca in the east of Wallace s line hotspot: the first with staminate flowers with a cupular calyx and three, elongated triangular to slightly spathulate petals (containing A. novohibernica and A. vestiaria), and the other with the calyx with three, slightly imbricate, triangular sepals and three-triangular petals in the other group (including A. catechu, A. macrocalyx, A. mandacanii and A. oxycarpa). The group of A. novohibrnica and A. vestiaria actually resembles Furtado s section of Mischopleous, and they form a very isolated group within the genus Areca. Areca is diverse in pollen morphology. In the recent studies of palm pollen conducted by Harley (1990, 1996, 1999), Harley and Baker (2001), and the genera specific study by Harley and Dransfield (2003), the pollen of Areca is given special attention, especially with the presence of triporate and zonosulcate apertures. Although the east of Wallace s line hotspot taxa were not included in these studies, in general pollen of Areca is usually ellipsoidal, symmetric or slightly asymmetric, less frequent oblate triangular or oblate spheroidal; the aperture a distal sulcus, in some species an extended sulcus, trichotomosulcus, or incomplete, presumed equatorial zonosulcus, rarely brevi or monoporate, or triporate; ectexine tectate or semi-tectate, finely to coarsely perforate, foveolate or finely reticulate, occasionally perforate-rugulate, aperture margin similar or slightly finer; infratectum columellate; longest axis µm; post-meiotic tetrads tetrahedral, rarely tetragonal or rhomboidal (Dransfield et al., 2008). The pistillate flowers are sessile, globular to triangular and usually much larger than the staminate flowers. Sepals are three, distinct, imbricate, and united near the base in A. novohibernica and A. vestiaria; the petals are similar to the sepals, sometimes valvate at the very tip, otherwise imbricate; staminodes are 4 6 or absent, triangular, ovate or circular with 4 6 teeth, membranous to fleshy; the gynoecium is unilocular, uniovulate, globose to ovoid, stigma three, and reflexed at anthesis.

118 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 100 Figure 3. Reproductive organ of Areca vestiaria and Areca oxycarpa: A, The triad on Areca vestiaria; two lateral staminate flowers and a central pistillate flower. B, Different stage at anthesis of the staminate flowers of Areca vestiaria, the flesh fly (Sarcophagidae) visiting the flowers. C, The inflorescence of Areca oxycarpa, slightly congested and protogynous. D, Ripen and young fruits of Areca oxycarpa. A B from Heatubun et al. 885; C from Heatubun et al. 883; D from Heatubun et al Scale bar A = 2 mm, B = 10 mm, C and D = 7 cm. Photos by Charlie D. Heatubun. Fruits and Seeds The fruits of the species of Areca from the east of Wallace s line hotspot are small to slightly moderate (to mm), varying in shape from obovate to ellipsoid, ellipsoid to slightly ovoid, ellipsoid to elongated with a beak to sickle-shaped, green to yellow or yellow to red when ripe, and one-seeded. The epicarp is thin and smooth with a mesocarp that can be thin to moderately thick and fibrous or fleshy and juicy; and with a fibrous endocarp. The seeds are small (to mm), sub globose, slightly globose to ellipsoid, ovate, and rounded apical and flatted or slightly hollowed at the base, with a basal hilum and anastomosing raphe branches. The size and shape of fruits and seeds varies greatly within individuals and populations, especially in the widespread native or cultivated species A. catechu, A. macrocalyx and A. vestiaria. Although fruit dimensions

119 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 101 were used by Beccari (1919) as the main character to distinguish the Philippine species of Areca and their varieties, and later used by Furtado (1933) to recognise sections in subgenus Blumeoareca, but there is no fundamental taxonomic character in the fruits and seeds useful to separate species in this region, except the mesocarp condition. A fleshy and juicy mesocarp is found only in A. novohibernica and A. vestiaria, while the other species are fibrous. Essig and Young (1979) studied the histology of the fruits of Areca together with other genera in subtribe Arecinae, and found that the systematic information obtained was only useful at higher taxonomic levels rather than for species level relationships in the genus. NATURAL HISTORY AND CONSERVATION Habitat and Ecology Almost all species of Areca in the EWL hotspot grow in lowland areas varying in soil types and habitat. Generally, Areca in this region occupies Entisols, sometimes Ultisols and very rarely Inceptisols (Soil Survey Staff, 1998; Hope & Hartemink, 2007). Tthere is a correlation between species of Areca and soil types especially with the parent materials which composed that soil. The alluvial and the limestone-based soils are the best substrates for Areca. Areca novohibernica and A. vestiaria have been collected from the volcanic soils in the Solomon Islands and near the volcano in the north of Sulawesi and the islands in the north Moluccas. Although A. macrocalyx found in the island of Waigeo and Gag, in the group of Raja Ampat Islands, but it never grows on ultrabasic soils, this palm grows on the limestone parts of those islands. Areca mandacanii grows in forest transitional between swampy and lowland rain forest, where the soils are temporarily inundated by water (Heatubun, 2008). Areca macrocalyx is also observed and collected from kerangas (heath forest) below Mt. Jaya, in southern part of New Guinea (Dransfield et al., 2000) and around Ambaidiru, in the central of Japen Island. This species is also one of the important species in the monsoon forest and savannah in Merauke area, in the southern part of New Guinea. High rainfall, humidity and temperatures are characteristic requirements for palms in the tropics, including Areca. Indeed, the high rate of precipitation is crucial for species growing in extreme habitats in this region, such as A. macrocalyx grow on limestone outcrops in the small off-shore islands of New Guinea, where the limestone s substrate almost without soil layer.

120 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 102 The litter-trapping grow forms, sometimes found in A. macrocalyx with the small undergrowth habits. Leaves of that palm are porrect, forming the shuttlecock-shaped crown that collects forest debris and provides a habitat for micro- and macro-decomposers responsible for breaking down the litter, such as fungi, arthropods (like the millipedes: Diplopoda, and the field cricket: Gryllidae) and earthworms. As observed in Sommieria (Heatubun, 2002), during rainfall, the crown also traps water and organic matter that is quickly channelled down the trunk to the ground where roots can take up nutrients and water. Unlike any other litter-trapping species of Areca from Borneo, the petiole of A. macrocalyx is very short or lacking and the stem is well developed. Whereas, the stemless and elongated-petiole are typical for the Bornean litter-trapping Areca. In the tropical forest communities, Areca as palm will occupy stratum C in tropical forest stratification (Richard, 1952), however, this position is depend on the stem habits of each species of Areca. The small-undergrowth form of A. macrocalyx, A. novohibernica and A. oxycarpa are in stratum D with stem tall 1 5 m high, together with tall herbs, large ferns and shrubs of various forms. Areca macrocalyx and A. mandacanii, each both sometimes tall to 20 m and 15 m high, they occupy stratum B. Sometimes, these two species will reaching up above the forest canopy in their habitat, in the kerangas forest, and the transitional between swamp forest and lowland rain forest where the forest only has four strata. Areca vestiaria is in stratum C, with its tall up to 10 m high. And all seedlings of Areca of east of Wallace s line will be in stratum E. Meanwhile, the planted A. catechu makes pure stands in the rural area through this region, and also cultivated in the garden near the forest margin. Sometimes, this species can found in the old and abandoned gardens or in the secondary forest and it become wild. Areca also plays important roles and supports other life forms in tropical forest ecosystem in this region. The stem is used as a mechanical support by liana, mostly Piper spp. (Piperaceae) and aroids (Araceae), like few species of Pothos, Rhaphidophora and Scindapsus to climbing up to the forest canopy for the light-demanding. The bark also is ideal substrates for the epiphyte-ferns and mosses or even orchids to grow. Further study is needed to understand these relationships. Pollination, Seeds Predation and Seeds Dispersal Despite the importance of understanding pollination (Corlett, 2004; Appanah, 1990), there have been no detailed studies on pollination of wild palms in the Indomalayan region comparable with those in the Neotropics (e.g. Silberbauer-Gottsberger, 1990; Listabarth,

121 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA , 1993, 1994, 1996). However, some observations (without went to detailed) have been carried out and increased our knowledge about palm-pollination in Malesian region (Dransfield, 1972, 1975, 1979, 1982; Essig, 1973; Mogea 1978; Madulid, 1980; Ferguson et al., 1983; Utami & Kahono 1989). Most species in the genus Areca are protandrous rather than protogynous (Loo et al., 2006, Dransfield et al., 2008). Of six species of Areca occurs in the east of Wallace s line, two species are protogynous (A. macrocalyx and A. oxycarpa) and four others are protandrous (A. catechu, A. mandacanii, A. novohibernica and A. vestiaria). The dichogamy is thought to be correlated with pistillate flowers morphology (Loo et al., 2006), pollen structure and pollination syndromes (Dransfield et al., 2008). In protogynous species which are thought to be beetle pollinated (Dransfield et al., 2008), the inflorescence is rather condensed and bears crowded pistillate flowers, and the pollen has more diverse exine sculpturing such as in Pinanga (Ferguson et al., 1987; Dransfield et al. 2008). In contrast, the putatively bee-pollinated protandrous species have laxly-branched inflorescences with the pistillate flowers rather distant, and the pollen is more uniform with low relief exine sculpturing (Essig, 1977; Dransfield et al., 2008). Areca macrocalyx and A. oxycarpa have congested inflorescence and the pistillate flowers crowded at base of rachillae near the rachis. Further investigation still needed to confirm the pollen morphology, the pollinator and other aspect of pollination biology on those palms. Based observation in the natural habitat of A. vestiaria in the north Sulawesi, the flowers of this palm have been visited by flesh flies (Sarcophagidae) (Fig. 3B) and it might be responsible for pollination. Utami & Kahono (1989) were also reported self-compatibility in A. vestiaria based on the plants cultivated in Bogor Botanic Garden. Self-compatibility also observed in A. macrocalyx, in which the prophyll still enclosed the inflorescence during the anthesis and apparently impenetrable to insect, but fruit set is abundant. Seed predation and dispersal is crucial in terms of their relevance to conservation. However, not much information about seeds predation and dispersal of Areca is available. The fruits (seeds still covered by the fibrous mesocarp) of A. macrocalyx have been observed in cassowary dung together with fruits of Orania regalis Zipp. ex Blume (Arecaceae: Arecoideae: Oranieae) and others forest fruits, such as Campnosperma sp. (Anacardiaceae), Eleaocarpus sp. (Eleaocarpaceae) Lepiniopsis sp. and Cerbera sp. (Apocynaceae) in Onin Peninsula, Western New Guinea (pers. obs.). The fruits of this palm also found floating in the streams, small rivers or even traps in dry gullies after being washed out by run-off.

122 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 104 Areca vestiaria has been reported in many occasions, that the fruit eaten by the black macaque (Macaca nigra) or yaki (local dialect in north Sulawesi) and that is why A. vestiaria in local dialect called as Pinang yaki. Perhaps, there is correlation between the distribution of A. vestiaria and the macaques (Macaca spp.), because almost all areas in Sulawesi and North Moluccas where the A. vestiaria occurs also found the macaques. But this still need proper study to proof and understand this relationships. Biogeography Of all six species of Areca in east of Wallace s line, five species are native to the region and one species is widely cultivated across the tropics, the betel nut palm (A. catechu). And from five native species; two species (A. macrocalyx and A. vestiaria) have relatively wide distribution and three other species (A. mandacanii, A. novohibernica and A. oxycarpa) have limited distribution or restricted to the certain areas in this region. Areca mandacanii just confined to the specific habitat in the lowland area in the Bird s Head Peninsula in western New Guinea (Heatubun, 2008). Areca oxycarpa also has similar distribution pattern to A. mandacanii, but this species is restricted to Minahasa and Gorontalo area in the north of Sulawesi or Celebes. While, A. novohibernica is the offshore islands Areca species in the north to east Papua New Guinea and it has been collected from Manus Island, New Britain and New Ireland in the Bismarck Archipelago and the Solomon Islands. Whereas A. macrocalyx distributed from Moluccas Islands in the west (including Aru and Tanimbar in the south-east, and Ternate and Halmahera in the north), through New Guinea and ended in the Solomon Islands in the east. Areca vestiaria is mainly distributed in the island of Sulawesi (Celebes) and eastward via Sula Islands to the Moluccas Islands (Buru, Ternate, Bacan and Halmahera). It is difficult to explain the pattern that showed by each species of Areca in east of Wallace s line. However, these distribution patterns are more likely reflect the dispersal events rather than vicariances as suggested in the phylogenetic analysis of the genus Areca based on DNA (Heatubun et al., in prep.). The roles of specific dispersal agents may also contribute to this curious distribution pattern. There is an indication that the distribution pattern of two closely related species A. novohibernica and A. vestiaria ( and also A. costulata in the Philippines) have been followed the soil preferences, the distribution of volcanic soil and it might be correlated with geological history of the islands in the region.

123 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 105 Conservation Status Three species of Areca in this region meets the criteria of Least Concern (LC IUCN, 2001), namely A. catechu, A. macrocalyx and A. vestiaria. These all three species have wide areas of distribution and are also widely cultivated as nut-produced and/or ornamental palms especially A. catechu and A. vestiaria. Two species (A. mandacanii and A. oxycarpa) meet the criteria of Near Threaten (NT IUCN, 2001), they both have very restricted areas of distribution whereas the environmental pressure around their natural habitat are so high, from land-clearings, re-settlements, coffee and cacao plantations, and paddy fields mainly in north Sulawesi. Before, the conservation status of A. mandacanii was regarded as Data Deficient (Heatubun, 2008), but it is important now to increase the category of this endemic palm having recognised more potential threats to its area of distribution. Oil palm plantation has been planned in the area of Sorong Selatan as a result of greater autonomy in districts of the Bird s Head Peninsula of Western New Guinea. Moreover, the local people usually chop down the tree when they want to harvest the fruits or nuts as a betel nut substitute. Areca novohibernica meets criterion Data Deficient (DD IUCN, 2001), more populations and distribution data are required to assess the conservation status of this species. Uses The enormous scale of betel nut palm use (A. catechu) is beyond doubt (Norton, 1998; Lee & Choi, 1999; Sullivan et al., 2000; Byun et al., 2001; Ray & Reddy, 2001) and it has become one of the most important stimulant products in the world, used by around million people (Norton, 1998; Gupta & Warnakulasuriya, 2002; Zumbroich, 2008). The origin and disperal of the betel nut palm and the chewing habit have been reported and discussed for many years, but more comprehensive study from various backgrounds and details is provided by Zumbroich (2008). The betel nut palm plantation also has been showed playing an important role to maintain the biodiversity in Western Ghats, India (The Economist, 8 th November 2008: 100). Whereas the uses of the native species of Areca of east of Wallace s line, in general, the fruits of all species used as the betel nut substitution. While, there are some minor utilization of A. macrocalyx in New Guinea, such as stems and leaves for building materials (flooring and thatching for hut or temporary house) and fruits for medicine. The fruits of Areca vestiaria are traditionally used for tonic and male contraception in Dumoga Nani Warta Bone National Park in north Sulawesi.

124 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 106 TAXONOMIC TREATMENT OF ARECA EAST OF WALLACE S LINE ARECA L. SP. PL.: 1189 (1753). TYPE: ARECA CATECHU L. Mischophloeus Scheff., Ann. Jard. Bot. Boitenzorg 1: 115, 134 (1876). Gigliolia Becc., Malesia 1: 171 (1877). Pichisermollia H.C. Monteiro, Rodriguesia 28: 195 (1976). Distribution: India and south China, through Malesia to New Guinea and the Solomon Islands. Number of species: 41 species in total and 6 species in east of Wallace s line. KEY TO SPECIES OF ARECA EAST OF WALLACE S LINE 1. Palm with stilt roots (prop roots); staminate flowers spirally arranged on rachilla, sepals united or calyx tubular, petals elongated and spathulate; pistillate flowers spirally arranged and distributed to the half length of rachilla; fruits with fleshy and juicy mesocarp Palm without stilt roots (prop roots); staminate flowers uniseriate or distichously (or sub distichously) arranged on rachilla, sepals triangular and free, petals triangular; pistillate flowers not spirally arranged and distributed only at the base of rachillae; fruits with fibrous mesocarp Small palm to 4 m ( 5 m) high and to 5 cm in diameter; always solitary; crown shaft green; inflorescence branched to 1 order (rare 2 orders); staminate flowers covered by velvet-like indumentum, stamens epipetalous, anther irregular and sometimes twisted; Manus Island, the Bismarck Archipelago and the Solomon Islands A. novohibernica 2. Moderate to big palm to 10 m ( 15 m) high and 7 15 cm in diameter; solitary or clustering; crown shaft orange to reddish; inflorescence always branched to 2 orders; staminate flowers glabrous, not epipetalous stamens, anther arrowheadshaped and elongated; Sulawesi to Moluccas. 6. A. vestiaria 3. Small-undergrowth palm to robust-emergent tree palm; inflorescence congested, branched to one order (rare 2 orders); protogynous.... 4

125 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA Moderate tree palm to robust-emergent tree palm; inflorescence divaricate, branched from 2 to 3 orders; protandrous Small-undergrowth palm to robust emergent tree palm ( 20 m high); stem 3 20 cm diam.; leaf sheath and crownshaft mostly green in colour (sometimes reddish to bright red); inflorescence strongly congested and bottle brush-shaped; triads just borne at the base of rachilla; Moluccas to Solomon Island A. macrocalyx 4. Small undergrowth palm ( 3 m high); stem 1 3 cm in diam.; leaf sheath and crownshaft always dark brown to black in colour; inflorescence slightly congested but not bottle-brush shaped inflorescence; triads occur to 1/3 length of rachilla from base to tip; North Sulawesi A. oxycarpa 5. Multi-fold leaflets with several folds at the same plane of the leaf rachis; cultivated across the Old Wold tropics.. 1. A. catechu 5. Single-folded leaflets at different plane of the leaf rachis (plumose); native to Bird s head peninsula in western New Guinea A. mandacanii SPECIES DESCRIPTION OF ARECA EAST OF WALLACE S LINE 1. Areca catechu L. Sp. Pl.: 1189 (1753). Type: Pinanga Rumphius, Herb. Amboin. 1: t. IV (Lectotype Moore & Dransfield, 1979). Areca cathechu Burm.f., Fl. Indica: 241 (1768). Type: Pinanga Rumphius, Herb. Amboin. 1: t. IV (Lectotype: designated here). Areca faufel Gaertn., Fruct. Sem. Pl. 1: 19 (1788). Type: Pinanga Rumphius, Herb. Amboin. 1: t. IV (Lectotype: designated here). Areca catechu var. nigra Giseke, Prael. Ord. Nat. Pl.: 73 (1792). ( Rumphius Pinang Itam ). Synon. Nov. Areca hortensis Lour., Fl. Cochinch.: 568 (1790). Type: Pinanga Rumphius, Herb. Amboin. 1: t. IV (Lectotype: designated here). Areca catechu var. alba Blume, Rumphia 2: 68 (1839). ( Rumphius pinang poetih ). Synon. Nov. Areca catechu L. var. batanensis Becc., Philipp. J. Sci. 3: 304 (1908). Type: Philippines, Batanes Island, 6.vi.1907, Fenix 3834 (holotype: FI). Synon. Nov.

126 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 108 Areca macrocarpa Becc., Phillip. J. Sci. C4: 601 (1909). Type: Philippines, Mindanao, Zamboanga District, Port Banga, i.1908, Whitford & Hutchinson 9103 (holotype: FI). Synon. Nov. Areca catechu L. var. longicarpa Becc., Philipp. J. Sci. 6: 229 (1911). Type: Philippines, Polillo Island, 10.xi.1909, Mc Gregor (holotype: FI). Synon. Nov. Areca catechu L. f. communis Becc., Philipp. J. Sci. 14: 304 (1919). Type: Philippine, Mindanao, Misamis Province, Katajan, Mt. Malindang, Mearns & Hutchinson 4717 (holotype: FI). Syn. Nov. Areca catechu L. var. silvatica Becc., Becc., Philipp. J. Sci. 14: 304 (1919). Type: Philippine, Palawan, Lake Manguao, iv.1913, Merrill 9447 (holotype: FI; isotypes: K, L). Synon. Nov. Invalid names: Areca himalayana Griff. ex H. Wendl. in O.C.E.de Kerchove de Denterghem, Palmiers: 231 (1878). Nom. nud. Areca nigra Giseke ex H. Wendl. in O.C.E.de Kerchove de Denterghem, Palmiers: 231 (1878). Nom. nud. Sublimia areca Comm. ex Mart., Hist.Nat. Palm. 3: 169 (1838). Nom. inval. Description: Moderate to large, solitary tree palm m ( 30 m). STEM cm in diam.; internodes closely to elongated ( 20 cm long). LEAVES 8 10 in crown, crown appearing shuttle-cock shaped to arching, cm long (including petiole); sheath tubular, cm long; crownshaft cm long, cm in diam., light green to green; petiole almost missing or very short to 15 cm long, channeled adaxially, rounded abaxially; leaflets distributed evenly along the rachis in one plane, leaflets on each side, basal leaflets c , middle leaflets c cm, apical leaflets c cm, briefly pointed and sometimes notched at apices, green, concolorous when dried. INFLORESCENCE infrafoliar, divaricate, cm long, branched 2 to 3 orders, rachillae numerous; prophyll caducous, cm, leathery; peduncle short, c. 8 cm wide at the base; rachillae c cm long, rachilla bract not conspicuous, sometimes calyx persistent on rachillae after fruits fallen. FLOWERS dimorphic, borne in triads at the base of rachillae and/or on the proximal part of each order of branching, staminate flowers arrange one-sided to distichous. STAMINATE FLOWERS mm, asymmetrical; calyx cupular, low; petals 3, mm, triangular, cream coloured; stamens 12; filaments

127 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA mm long; anthers mm; pistillode to 3 mm long, trifid. PISTILLATE FLOWERS mm, triangular; sepals 3, to mm, imbricate, strongly keeled with cream to green coloured at anthesis; petals 3, mm, cream coloured; gynoecium c mm (including stigma 5 mm), stigma trifid; staminodes circular, membranous. FRUITS cm, various shapes from ovoid, ovoid-ellipsoid, narrowly ellipsoid, elongate-ellipsoid, rather ventricose, green and yellow to orange or reddish orange when ripen; epicarp smooth and thin, sometimes with scars or ramenta; mesocarp fibrous and thick; endocarp fibrous and thin. SEEDS cm, various shapes from subglobose, subglobose-depressed ovoid, ovoid conical, with more less flattish base to the base flat, hard; endosperm ruminate. Distribution and Ecology: This is the most widely cultivated species in the genus Areca and it has been distributed throughout tropics. Due to domestication of this palm, the country of origin is not known with certainty. However, several locations have been suggested based on the occurrence of the wild close relative species, such as the Philippines, Malaysia, Celebes (Sulawesi) and New Guinea (Beccari, 1919; Furtado, 1933; Corner, 1966; Jones, 1995; Heatubun, 2008). Specimen studied: EUROPE UK. Great Britain, England, Kew, cultivated at Royal Botanic Gardens, no. x.1.776, s.n. (K!). AFRICA TANZANIA. Dar Es Salaam, University Agriculture and Forestry campus, Morogoro, T6, 535 m, 11.xi.1976, Wingfield 3708 (K!). INDIA. Bengal, Cari Bangar, Baharchara, 11.x.1943, Sinclair 3272 (E!). East Bengal, , Griffith s.n. (K!). Assam, Masters s.n. (BO!). Calcutta, Botanic Garden, cultivated at Botanic Garden, s.n. (E!, L!). SRILANKA. 1.iv.1860, Dubuc 1869 (E!). Paradineya, Botanic Garden, lawn Q 306, 27.iii.1986, Rutherford & Bandara 124 (K!); 23.ii.1909, Baker 136 (BO!, K!). Sinharaja, Barfod s.n. (AAU photo!). CHINA. Yunnan, Yunnan Institute of Tropical Botany, 600 m, 2.vii.1978, Chen (K!). TAIWAN. Kagi Province, Kagi, 24.ii.1918, Wilson 9891 (K!); Orchid Island, Ten-tzu, 31.viii.1969, Huang & Kao 5234 (L!). Hainan, 16.vii.1933, Liang (K!); xi.1889, Henry 8406 (K!); Central eastern Hainan, between Kachek and Ling Mon, 5.vi.1922, the Canton Christian College 9808 (E!, K!); Taam Chau District, Na Lin Sham, 30.v.1928, Tsang 146 (K!); Ngai District, Naam Shan Leng, 16.vi.1932, Lau 266 (E!, K!). THAILAND. Bangkok, 4.vi.1920, Graff 134 (K!). Phangnga Province, Laemson National Park, Kampuan Substation, Ton Lan, hill Dipterocarp forest, m,

128 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA ii.1994, Barfod et al (AAU!, BKF, PSU). SINGAPORE. Botanic garden, iv.1920, s.n. (K!); lawn X, 19.vi.1929, s.n. (K!); lawn J, 26.vii.1929, s.n. (K!). The National University Hospital, Kent Ridge wing, 13.vii.2001, Ching & Tan TC 01 (K!). THE PHILIPPINES. Luzon, Manila, Isaac Peral Street, 1935, Bartlett (K!). Rizal Province, Antipolo, xi.1914, Merrill Species Blancoanae 213 (BO!, K!, L!, PNH). Laguna Province, Los Banos, Mt. Makiling, vi vii.1917, Elmer (BO!). Tayabas Province, Lucban, v.1907, Elmer 7795 (BO!, E!). Isabela Province, Jones, St. Domingo, 120 m, 22.iv.1987, Fernando 665 (K!). Bataan Province, Mt. Mariveles, Lamao River, 3.xii.1903, Williams 330 (K!); vii.1904, Borden (Forestry Bureau 1272) (K!). Batanes Province, Batan Island, Mt. Iraya, viii.1930, Ramos (Bureau of Science) (K!). Camarines Norte Province, Labo, Bo Fondado, Coconut grove, 14.vii.1985, Fernando 546 (K!). Visayas Western Samar Province, Mt. Malingon, Brgy Lokilokon, Paranas, 13.x.1992, Reynoso et al (K!). Mindanao, Zamboanga Peninsula, La Paz, Camp Susana, 500 m, 4.vii.1986, Fernando 600 (K!). Mindoro, Mt. Yagaw, SE Slope. 5.viii.1953, Conklin (L!); 300 m, 6.ii.1958, Conklin 1055 (PNH 37593) (L!). BORNEO. Sarawak, , Beccari PB 3112 (FI!, K!); Triboh scheme, Serian, 14.iii.1988, Othman & Munting S (K!, SAR!); Batu Bedanan, Ngarai Talong, Sungai Engkari, Batang Ai, Lubok Antu, 270 m, 31.vii.1994, Lai S (K!, L!, SAR!, KEP); Kampung Melayu, Ulu Layar, Betong, 4.vii.1988, Lee S (K!, L!, KEP, SAN, SAR!); Kapit, Upper Rejang River, 1929, Clemens (K!). Sabah, Ranau District, Bundu Tuhan village, Siba, Himbaan village, 13.v.1994, Soibeh 778 (K!); Poring village, at hill slopes, 31.x.1993, Sambuling 8 (K!); 6.ii.1994, Sambuling 76 (K!); Sandakan, miles 12, 10.iii.1949, Acuadian 202 (K!); Lahad Datu, Masuri, Ulu Segama, 20.xi.1949, Acuadian 254 (K!); Sorinsim village, Hutan muda, 13.iii.1993, Sibil 153 (K!); Melangkap Tomis village, 10 m from village hall, 29.v.1995, Lugas 442 (K!); Tongod District, 22 km logging road to upstream Milian River, 31.x.1984, Mansus & Aban SAN (K!, L!); Kiaunuluh village, Nuluhon, 4.ii.1993, Duaneh 255 (K!); Kota Marudu District, Serinsim village, 9.iii.1995, Bakia 416 (K!). INDONESIA. Riau Islands Province, Bengkalis, Singoro, 4 m, 17.ix.1919, Beguin 330 (BO!). West Sumatera Province, Siberut, 9.ix.1924, Boden-Kloss (BO!); Mentawai Islands, Sipora Island, 18.x.1924, Iboet 440 (BO!). Banten Province, Panaitan Island, Ciharahas, upstream, 5.ix.1951, Borssum- Waalkes 350 (BO!, K!); Peucang Island, Ujung Kulon, 4 m, 13.iv.1971, Dransfield 1403 (BO!); track to Cibunar, 5 m, 17.iv.1971, Dransfield 1467 (BO!); Dransfield 1471 (BO!); Ujung Kulon National Park, c. 1 km NW of Cibunar security post, along the bank of

129 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 111 Cibunar River, 40 m, 2.x.1998, Noblick et al (BO!, K!, MBC). DKI Jakarta Province, Bidara Cina, xi.1863, Edeling s.n. (BO!). West Java Province, Bogor, Botanic Garden, ex cultivated, lawn II.F.9, s.n. (BO!); lawn VII.B.78, s.n. (BO!); lawn VII.B.79, s.n. (BO!, L!); lawn VII.B.80, s.n. (BO!); lawn VII.B.81, s.n. (BO!); lawn VII.B.84, s.n. (BO!, L!, G); lawn VII.B.85, s.n. (BO!, L!); lawn VII.B.87, s.n. (B, BO!, K!, L!); lawn VII.B.88, s.n. (BO!, L!); lawn VII.B.89, s.n. (BO!, L!); lawn VII.B.91, s.n. (BO!); lawn XI.B(XX)6, s.n. (BO!); lawn XIII.A.6, s.n. (BO!); Ciherang, 700 m, 8.i.1928, van Steenis 212 (BO!); W of Bogor, Leuwiliang, Pasir Honje, 300 m, 5.vi.1927, Bakhuizen van den Brink 6724 (BO!); Bakhuizen van den Brink 6782 (BO!); Bakhuizen van den Brink 6788 (BO!); Bakhuizen van den Brink 6789 (BO!); 7.vi.1927, Bakhuizen van den Brink 6807 (BO!, L!); 14.viii.1927, Bakhuizen van den Brink 6866 (BO!); Tasikmalaya, Nusa Gede, Penjalu, 720 m, 30.vii.1917, Koorders 396 (BO!). Central Java Province, Banyumas, Jagadanda River, 3 m, 10.v.1921, Backer (BO!). Gorontalo Province, 250 km W of Gorontalo, 75 km inland from Papayuto, on tributary of Papayuto River, 150 m, 30.iii.1990, Burley et al (A, BO!, K!). South Sulawesi Province, c. halfway South road Soroaku Wasupoda, 700 m, 1.vi.1979, de Vogel et al (BO!, L!). North Sulawesi Province, Bolaang Mongondow, Pindool District, Lolak, 50 m, 19.x.1973, Dransfield & Mogea 3808 (BO!, L!). Southeast Sulawesi Province, Kolaka area, Mt. Watuwila foothills, above Sangguna, Mokuwu camp, 200 m, 30.x.1989, Coode 6074 (BO!, K!). East Nusa Tenggara Province, Alor Island, Atimelang, 750 m, 24.i.1939, Du Bois 24 (BO!); Timor Island, Baumata village, 50 m, 21.iii.1939, Bloembergen 3555 (BO!); Bloembergen 3556 (BO!). North Moluccas Province, Halmahera Island, Ekor, near the village, 20 m, 1.x.1974, de Vogel 3266 (BO!, K!, L!); Tidore, 17.v.1921, Beguin 1613 (BO!, B); Bacan Island, NE Bacan, 2.ix.1985, Sidiyasa et al.tcw 3580 (K!). Moluccas Province, Seram Island, Masohi, Waipia, Jerili village, 10 m, 18.xi.1981, Mogea 3140 (BO!). West Papua Province, Raja Ampat Islands District, Gag Island, 10 m, 29.vii.2006, Heatubun et al. 751 (BPK Manokwari!); Heatubun et al. 752 (BPK Manokwari!); Heatubun et al. 753 (BPK Manokwari!). Papua Province, Japen Island, Konti Nuai village, 100 m, 25.ii.2008, Heatubun et al. 870 (BPK Manokwari!, BO!, K!). Notes: Areca catechu is the most widespread cultivated species in the genus Areca and it has been a subject of selection by humans for a long time. This species has been planted throughout tropical regions and also been developed to the large-scale plantations these days. This palm is mainly planted for the nuts production, thus the fruits and seeds are the

130 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 112 priority or main target for the selection or breeding, although cultivation for ornamental purpose has increased in recent years. Anthropogenic selection pressure makes this species very variabele in cultivation, especially the habit, fruits and seeds in the sizes and shapes, colours or even tastes, including other characters within the fruits and seeds, such as epicarp, mesocarp, endocarp, raphe, endosperm rumination and embryo. In this treatment, we apply the more wide species concept to A. catechu and do not recognise any varieties within this species as proposed before by the earlier authors (Giseke, 1792; Blume, 1839; Beccari 1908, 1911, 1919). And if any distinct individuals, populations or groups which found within the morphological range of A. catechu, they must be treated as cultivar(s) because they are the product of cultivation. There is no record of the wild population of A. catechu so far, although Merrill collected his specimen (Merrill 9447) in primeval forest in Palawan, the Philippines, but he also noted in his letter to Beccari (Beccari, 1919) that the trees he found and collected originated from seeds accidentally left there by natives. The two varieties, var. nigra (Giseke, 1792) and var. alba (Blume, 1839) based on interpretation of Rumphius s pinang itam and pinang poetih in Herbarium Amboinense, and these two varieties erected on the basis of different colours in the plant and reproductive organs, including fruits more dark and white or yellow from the common variety. Further, Beccari (1919) keyed out three varieties and one form of A. catechu in the Philippines; var. batanensis (Beccari, 1908), var. longicarpa (Beccari, 1911), var. silvatica (Beccari, 1919) and forma communis based on the stem thick and short, inflorescence dense and rachillae bearing flowers short, the fruits and seeds shapes or dimension. Areca macrocarpa Becc. synonymous with A. catechu, as already mentioned by Merrill (1923: 26). Areca macrocarpa is merely a form of Areca catechu L. with much larger fruits. This has been followed by Fernando (in prep.) in his provisional annotated checklist of the palms of the Philippines who could find no other specimen of A. macrocarpa, except the type specimen kept in Florence. Beccari (1919) in his conspectus of the Philippines species of Areca, differentiated A. macrocarpa from A. catechu by seed having vascular bundles of the integument arising straight, almost erect, from raphal side and slightly branching, and fruit elongate-ellipsoid, twice as long as broad (7 cm long and 3.2 to 3.5 cm broad), and also seed ovoid-conical with a blunt apex rather than seed having a vascular bundles of the integument arching on the sides, and strongly anastomising immediately from its base, even on the raphal side, and fruit globose-ovoid or ovoid-ellipsoid, not more than one-third or one-fourth longer than broad (4 to 5 cm long and 3 cm or little less

131 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 113 broad), and seed subglobose with a more or less flattish base. After examining many collections of A. catechu from throughout its range, including type materials of A. macrocarpa and all varieties of A. catechu, we find no reason to maintain more than one species Areca catechu is similar to the EWL species A. mandacanii in all parts, except the leaves plication and leaflets arrangement on the leaf rachis. See discussion under the leaves and notes under A. mandacanii for more explanation. 2. Areca macrocalyx Zipp. ex Blume, Rumphia 2: 75 (1839). Type: New Guinea, SW coast, Zippel s.n. (holotype: L). Areca jobiensis Becc., Malesia 1: 21 (1877). Type: New Guinea, Geelvink Bay, Japen Island, Ansus, iv.1875, Beccari s.n. (holotype: FI; isotypes: K). Synon. Nov. Areca macrocalyx var. zippelliana Becc., Malesia 1: 19 (1877). Type: New Guinea, SW coast, Zippel s.n. (holotype: L). Synon. Nov. Areca macrocalyx var. aruensis Becc., Malesia 1: 20 (1877). Type: Aru Island, Vokan (Wokam), iii.1875, Beccari s.n. (holotype: FI; isotype: K). Synon. Nov. Areca macrocalyx var. conophyla Becc., Malesia 1: 20 (1877). Type: New Guinea, NW coast, Ramoi and Amberbaken, 1872, Beccari s.n. (holotype: FI; isotype: K). Synon. Nov. Areca macrocalyx var. waigheuensis Becc., Malesia 1: 20 (1877). Type: New Guinea, Waigeo Island, Wakkere (Wakre), iii.1875, Beccari s.n. (holotype: FI; isotype: K). Synon. Nov. Areca rechingeriana Becc., Webbia 3: 163 (1910). Type: Papua New Guinea, Bougainville Island, Kieta, Rechinger 3992 (holotype: B ; isotype: FI). Synon. Nov. Areca macrocalyx var. intermedia Becc., in R. Rechinger, V. Teil. Denkschr. Kaiserl. Akad. Wiss. Wien Math.-Naturwiss. Kl. 89: 506 (1913). Type: Papua New Guinea, Bougainville Island, Kieta, Rechinger 4182 (holotype: B, isotype: FI). Synon. Nov. Areca nigasolu Becc., Webbia 4: 256 (1914). Type: Solomon Islands, Treasury Island, Guppy 95 (holotype: K, photo FI). Synon. Nov. Areca torulo Becc., Webbia 4: 253 (1914). Type: Solomon Islands, Treasury Island, Guppy 94 (holotype: K). Synon. Nov. Areca warburgiana Becc., Bot. Jarhrb. Syst. 52: 24 (1914). Type: New Guinea, Sigar, Warburg 20 (holotype: B, photo FI; isotype: FI). Synon. Nov.

132 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 114 Areca nannospadix Burret, J. Arnold Arbor. 12: 265 (1931). Type: Papua New Guinea, Ihu, Vailala River, rain forest, 9.ii.1926, Brass 921 (holotype: A). Synon. Nov. Areca rostrata Burret, Notizbl. Bot. Gart. Berlin-Dahlem 12: 322 (1935). Type: Papua New Guinea, Diemi, Onange road, Central Division, 6.v.1933, Brass 3971 (holotype: A). Synon. Nov. Areca multifida Burret, Notizbl. Bot. Gart. Berlin-Dahlem 13: 331 (1936). Type: New Guinea, Papua New Guinea, Veiya, 11.iii.1935, Carr (holotype: B, isotypes: A, K). Synon. Nov. Invalid names: Areca glandiformis Lam., Encycl. 1: 241 (1783). Nom. nud. Areca macrocalyx var. keyensis Becc. in Martelli, Nov. Giornale Bot. Italiano 42: 24 (1935). Nom. nud. Description: Solitary, slender to robust palm, tall m. STEM cm in diam.; internodes closely to elongated, 2 20 cm long, light brown to whitish with conspicuous leaf scars, and shiny green near the crown. LEAVES 6 10 in crown, lamina glabrous, to 250 cm long (including petiole); sheath tubular, to 92 cm long; crownshaft to 150 cm long, green to dark green with numerous black dot scales and reddish green to bright red in some population; petiole from almost missing to 10 cm long, channelled adaxially, rounded abaxially; leaflets in regularly, irregularly or clustered arranged, papery, 6 75 leaflets on each side, from broad leaflet with several main veins to single folded leaflet, slightly sigmoid in basal most leaflets and oblique-acuminate tip except to terminal leaflets, slightly flabellate to linier with notch tips, green, discolorous when dried, dark-coloured adaxially, and paler abaxially. INFLORESCENCE infrafoliar, elongate, bottle-brush like, to 65 cm long, branched to 1 order (sometimes basal most rachillae with 2 branches), erect to pendulous; prophyll to cm, caducous, cream to reddish; peduncle short to 10 cm long, elongate; rachillae numerous (12 600), to 41 cm long, cream to green, sometimes perianth persistent on rachillae after fruits fall. STAMINATE FLOWERS numerous, arranged distichously along rachillae from immediately above pistillate flowers to rachilla tip, c mm, triangular, asymmetrical, creamy; sepals 3, united at the base, low, 3 lobes, c. 7 4 mm; petals 3, thick and fleshly, c mm, triangular, striata; stamens 6, 7 9 mm long; filaments c. 3 4 mm long; anthers 6 7 mm long; pistillode dimunitive. PISTILLATE FLOWERS to mm, triangular, congested to the main rachis; sepals 3, imbricate, 10

133 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA mm; petals 3, imbricate, mm; gynoecium mm (including stigma 5 mm); staminodes circular, membranous. FRUITS typically obovate, but somewhat variable from connate to spindle shaped to almost globose, to 5 3 cm, with beak mm long, green to bright yellow or orange when ripen. SEEDS to 3 2 cm, ovate, rounded apically and flatted basally, hard; endosperm ruminate. Distribution and ecology: This is the most widespread species of Areca in the EWL hotspot with distribution from the Moluccas Islands in the west through New Guinea to the Solomon Islands in the east. This species is also occupying the widest ecological spectrum, growing in the lowlands to the highland from sea level to 1500 m. Specimen studied: INDONESIA. Moluccas Province, Seram Island, Masohi, Waipia, Jerili village, 10 m, 19.xi.1981, Mogea 3139 (BO!, K!); Aru Islands, Kobroor Island, 10 m, 6.xi.1994, van Balgooy 6864 (L!, BO!); Vokan (Wokam), iii.1875, Beccari s.n. (holotype FI!; isotype K!). West Papua Province, Kepulauan Raja Ampat District, Misool Island, Motlol, 10 m, 22.i.2002, Heatubun et al. 360 (K!, MAN!); c. 10 km SW of Limalas village, around Wavari camp, 5 m, 20.i.2002, Wanggai et al. 03 (AAU!, BO!, K!, LAE); Gag Island, Kaplebet River, 10 m, 28.vii.2006, Heatubun et al. 747 (BO! BPK Manokwari!, K!); Waigeo Island, Omrab, 21.viii.1956, Manuputty BW 958 (BO!, CANB!, L!, MAN!); Sorong District, road between Bayangkate village to Makbon, 72 m, 9.iii.2008, Heatubun et al. 876 (BO!, BPK Manokwari!, K!); Manokwari District, Manokwari sub district, Amban, Pantai Anggori, 5 m, 9.viii.1995, Dransfield et al. JD 7533 (BO!, K!, MAN); Mupi, c. 40 km S of Manokwari, along coast of Arfak Nature Reserve, 50 m, 11.iv.1994, Mogea 6204 (BO!, K!, MAN!); Ransiki subdistrict, Siwi village, 28.i.1999, Heatubun 292 (AAU!, BO!, FTG!, MAN!); Mountains S of the Arfak Plains, step ridge between the Arfak Plains and Mt. Itsiwei, 900 m, 27.iv.1994, Sands et al (BO!, K!); Teluk Bintuni District (before as Bintuni subdistrict of Manokwari District), near Saengga village, 10 m, Maturbongs et al. 728 (BO!, K!); Kaimana District (before as Kaimana subdistrict of Fakfak District), Etna bay, Urie Base camp, km-36 road of PT. Kaltim Hutama, 70 m, 1.ii.2001, Heatubun et al. 331 (AAU!, K!, MAN); SW coast New Guinea, Kaimana District, Etna bay subdistrict, Zippel 169 (holotype L!).

134 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 116 Figure 4. Areca macrocalyx Zipp. ex Blume. A, Apical portion of leaf. B, Middle portion of leaf. C, Petiole. D, Inflorescence. E, Portion of rachilla with staminate and pistillate flowers. F, G, Staminate flower whole and in section. H, I, Pistillate flower whole and in section. J, K, Fruit whole and in section. Scale bar: A C = 8 cm; D, E = 6 cm; F I = 7 mm; J, K = 1.5 cm. All from Baker Drawn by Lucy T. Smith.

135 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 117 Papua Province, Japen Island, Ambaidiru village, 800 m, 23.xi.1998, Maturbongs et al. 605 (BO!, K!, MAN!); Japen Island, Ansus, iv.1875, Beccari s.n. (holotype FI!; isotypes K!); Mamberamo Raya District, Idenburg River, 4 km SW of Bernhard Camp, 850 m, iii.1939, Brass (A!, L!); Brass 13437A (A!, L!); Versteeg 1592 (BO!, L!); Bivak Eiland, 6.x.1907, Versteeg 1782 (L!); Jayapura District, North Cyclops Mts., m, 30.i.2001, Desianto 04 (AAU!, K!, MAN!); Keerom District, Tami River, 91 m, 16.ii.2008, Heatubun et al. 796 (BO!, BPK Manokwari!, K!); Heatubun et al. 798 (BO!, BPK Manokwari!, K!); Heatubun et al. 799 (BO!, BPK Manokwari!, K!); Arso, Tami River, Yawu, m, 16.iii.2002, Gusbager et al. 20 (K!, MAN!). PAPUA NEW GUINEA. Sandaun Province, Bewani District, N of Bewani Patrol Post, 150 m, 28.viii.1985, Karenga LAE (L!, LAE!); Bewani, 0 m, iii.2000, Barfod et al. 488 (AAU!, BRI!, CANB!, K!, LAE!); Round house village, m, 27.xi.1996, Barfod 416 (AAU!, K!); Vanimo, Warastron Forestry Station, 1 2 m, 9.ix.1982, Karenga LAE (L!, LAE!); Sepik District, Aitape subdistrict, near Sumo village, Rhainbrum River, 15 m, 7.vii.1961, Darbyshire & Hoogland 8089 (CANB!, L!, LAE!); Telefomin District, Mt. Entaldam, immediately S of Busilmin airstrip, 1500 m, 28.iii.1975, Vinas LAE (L!, LAE!); Hak valley, head of Bal Creek, tributary of Sek River, Donner Mts., S of airstrip, 1050 m, 15.x.1993, Morren 3049 (K!). Western Province, Palmer River, 2 miles below junction, Black River, 100 m, vi.1936, Brass 7001 (A!, L!); vii.1936, Brass 7170 (A!); Brass 7188 (A!); Brass 7386 (A!); lower Fly River, Sturt Island, x.1936, Brass 8189 (A!, A photo!); near ingembit village, 144 m, 16.vi.1967, Henty et al. NGF (LAE!); Henty et al. NGF (L!, LAE); Oriomo River, 2 7.vii.1968, Reeve 899 (CANB!); upper Fly River, near N glei village, c. 10 miles N of Kiunga, 60 m, 10.vii.1967, Pullen 7299 (CANB!); Kiunga, 25 m, 12.ix.1972, Streimann & Lelean NGF (LAE!); Streimann & Lelean NGF (BRI, L!, LAE); Lake Daviumbu, middle Fly River, ix.1936, Brass 7901 (A!, L!). East Sepik Province, Wewak-Anggoram area, c. 5 miles N of Timbunke on Kwoiwut track, right hand side of Minjim River, 30 m, 12.ix.1959, Pullen 1712 (CANB!, L!, LAE!); Sepik District, Ambunti, Eastern ridge of Sumset (Mt. Hunstein), 1140 m, 18.viii.1966, Hoogland & Craven (CANB!, K!, L!, LAE); Sepik River, vicinity of Langu village, 40 m, 18.viii.1994, Takeuchi (A!, LAE). Southern Highlands Province, Mt. Bosavi, northern side, m, 25.ix.1973, Jacobs 8761 (L!); m, 4.x.1973, Jacobs 9002 (L!); m, 26.x.1973, Jacobs s.n. (L!); near Bosavi mission, Dudessa or Ludessa village, 750 m, 5.ii.1996, Baker et al. 631 (K!, LAE); Tari, Mt. Bosavi, head of Kuru Creek, 1300 m,

136 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA viii.1986, Gideon LAE (L!, LAE!); E of Mendi, upper Agimo River, Mt. Gilure, Southern slopes, (?) m, 27.vi.1961, Pullen 2638 (CANB!). Madang Province, Bogia, mouth of Ramu River, Bogia-Bosmun road, near to Bosmun 2 village, 50 m, 20.i.1996, Baker & Utteridge 591 (K!, LAE); Josephstaal FMA area, near Kumamdeber, along streambed flowing to SW of Expedition camp 1, 160 m, 31.vii.1999, Takeuchi et al (A!, LAE). Eastern Highlands Province, Kainantu District, between Ayura and Akuna, 1800 m, 12.x.1957, Pullen 725 (CANB!, L!, LAE). Gulf Province, Kikori District, Victory junction (confluence of Sirebi River and Kuru River), 34 km N of Kikori, 50 m, 20.xi.2000, Baker et al (AAU!, K!, LAE, NY); Kopi-Kikori road, 10 km NW of Kikori, 40 m, 21.xi.2000, Baker et al (AAU!, BRI, K!, LAE, NY); Panini Creek, 13.xi.1959, White NGF (LAE!); Baimuru, Purari River, 240 m, 27.iii.1974, Croft et al. LAE (BRI, L!, LAE). Morobe Province, Finschhafen District, Jivewaneng village, 14 km NW of Finschhafen, 500 m, 5.xii.2000, Banka et al (AAU!, K!, LAE, NY); Wantot (Wantoat), c m, 11.iv.1940, Clemens (K!); Lae, Eriku, cultivated in Bulae International Prmary School, 20.iii.1996, Gideon s.n. (K!); Houn Peninsula, Masba Creek, 3 miles S of Pindiu, 600 m, 22.v.1964, Hoogland 9026 (CANB!, LAE, NY); NW of Waria River, along the streambed Wara Eya, near Yai village, m, 15.vi.1999, Takeuchi et al (A!, LAE). Central Province, Gulf District, Kikori River, 13.ii.1959, White NGF (K!, LAE); Karuku District, 2 miles N of Maipa village, 97.5 m, 10.ix.1962, Darbyshire 921 (CANB!); Abau District, Cape Rodney, Mori River, 60 m, 19,vi.1968, Henty NGF (L!, LAE); Dieni, Ononge road, 500 m, ivv.1933, Brass 3971 (A!, NY); Nunumai, c. 12 km N of Amazon bay, 30 m, 19.vi Pullen 7641 (CANB!, LAE). Milne Bay Province, Alotau District, 2 km NE of Kapurika village, 200 m, 11.v.1978, Essig & Young LAE (LAE!); SE of Kaporika village, road to Baraga and Cumuni villages, 70 m, 24.xi.1975, Larivita & Katik LAE (LAE!); Sagarai, Pini Range, Southern slopes, 200 m, 2.iii.1984, Gideon LAE (CANB, K!, L!, LAE, USF); Gideon LAE (L!, LAE); Raba-Raba District, junction Ugat and Mayu River, near Mayu I, 350 m, 29.vi.1972, Streimann & Katik NGF (LAE!); 1100 m, 19.vii.1972, Streimann & Katik NGF (L!, LAE); Gwariu River, Biniguni camp, 200 m, iii-iv.1938, Brass (A!). West New Britain Province, West Wakanau, near cape Hoskins, Glilo village, 3.viii.1954, Floyd 6422 (A!, LAE); Gasmata, m, 29.v.1987, Karenga & Obedi LAE (L!, LAE, USF). East New Britain Province, Pomio District, Aiwit River, E of Fulleborn harbour, 50 m, 11.v.1973, Croft & Katik NGF (L!, LAE); Gaselle Peninsula, Warangoi valley, 60 m, ix.1955, Kazakoff

137 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 119 NGF 7057 (A!, LAE). New Ireland Province, Feni group, Ambitle Island, E side of Nanum Caldera, 50 m, 13.xi.2003, Takeuchi (A!, LAE). North Solomon Province, Bougainville Island, , Waterhouse s.n. (K!); vicinity of Aku village, c. 10 miles, W of Buin Station, 30 m, 14.ix.1964, Schodde & Craven 4029 (A!, BRI, CANB!, K!, LAE). THE SOLOMON ISLANDS. Eastern District, Santa Cruz Island, Towuto Noi Island, Ngambwani area, 4.iv.1972, Powell BSIP (CANB!, BSIP); St. Ysabel Island, Bogutu Peninsula, Longuhutu River, SE corner of Tanegoba harbour, 0 30 m, 21.iii.1964, Moore & Whitemore 9303 (BSIP 4051) (BSIP, K!); Guadacanal, Honiara, behind Lengakiki ridge, 2.xii.1963, Whitmore BSIP 1268 (K!, BSIP); New Georgia, 15.vii.1929, Waterhouse 185 (K!); San Cristobal, Wairaha River, 5 miles from North Coast, 300 m, 11.v.1964, Whitmore BSIP 4273 (K!). CULTIVATED. Sri Lanka, Paradineya Botanic Garden, Q 451, 23.vii.1986, Rutherford & Bandara 154 (K!). Thailand, Bangkok, Sumawong s.n. (K!). Singapore, Botanic Gardens Singapore, Lawn X, 9.x.1929, Furtado s.n. (K!, SING). Notes: Areca macrocalyx is easy to distinguish from other species of Areca in east of Wallace s line by the nature of inflorescence being congested and club-like infructescence. The male only portions of the rachillae are very thin compared to female part and that they dry and fall off as the fruits mature leaving a club-like infructescence. The authors have the privilege to observe this species directly in its natural habitat, to understand its variations and to collect adequate representative herbarium specimens and other materials to allow us to complete this study. Flynn (2004) analysed Areca in New Guinea and the Solomon Islands for his master thesis project using morphometrics approach. Although he was faced the missing data and incomplete information in some materials, he reached the conclusion to include seven species (A. congesta, A. jobiensis, A. ledermanniana, A. multifida, A. nannospadix, A. rostrata and A. warburgiana) under A. macrocalyx. We concur with this decision. The morphology of A. macrocalyx is quite complex, perhaps as a response to different types of habitat and distribution. This species occupies a wide range of ecological conditions from littoral and swampy areas in lowlands to heath forest in lower montane vegetation, from evergreen rain forest to dry areas in savannah lands and from the main island of New Guinea to small off-shore islands, the Moluccas and the Solomon Islands. The adaptation to various habitats is reflected in the very variable appearance, a plasticity that occurs in size and shape. If two specimens from different localities are compared in

138 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 120 isolation, the impression may be gained that more than one taxon exists. However, with careful examination of the type specimens and protologues of the species listed in the synonymy above and a large number of other herbarium specimens, we found that morphological variation among them is continuous and overlapping, as Flynn discovered. No disjunctions in variation occur that would allow the consistent separation of multiple species in this complex as recognised by previous authors. The narrow species concept used in the past reflects limited information obtained from single collections. While our treatment is aligned with that of Flynn (2004), we have included three species (A. rechingeriana, A. nigasolu and A. torulo) and five varieties of A. macrocalyx were not covered by him and considered two other species (A. ledermanniana and A. congesta) as doubtful taxa rather than as synonyms of A. macrocalyx. Areca nigasolu and A. torulo were described by Beccari (1914) from Guppy collections in the Solomon Islands. The materials are very inadequate and consists a fragment of leaf and a few fruits, but the fruits are typical fruits of A. macrocalyx. The varieties of A. macrocalyx (var. zippeliana, var. aruensis, var. conophylla, var. waigheuensis and var. intermedia) were described by Beccari (1877, 1913) based on different fruit characters and collections localities. The types of Areca rechingeriana and A. warburgiana along with several specimens from recent collections (e.g. Heatubun et al. 796, 798, 799, Tami River, Keerom Indonesian Province of Papua) possess inflorescences that are elongated and slender with a very thin rachis very thin and laxly arranged pistillate flowers. The arrangement of two pistillate flowers in triads seated very close to each other at the base of the rachilla, and with the stalk formed by the rachilla gives the distinct impression as the fruit develop of fruit borne in pairs. However, the typical inflorescences of A. macrocalyx with condense and crowded pistillate flowers and/or fruits also been observed on that populations. The morphological plasticity that occurs in A. macrocalyx displays gradual and often overlapping variation and this has influenced the broad species concept used in this revision. Two specimens in Florence Herbarium (Jaheri n. 245 ex Bogor and n. 279 culta in Bogor Botanic Garden, lawn V. K ) are annotated by Beccari with his hand writings as A. macrocalyx var. keyensis but this name is never published. In the synonym list of palm tribe Areceae provided by Martelli (1935), this variety was cited as having been published in Malesia (1: 20). This is not the case and the name is in fact a nomen nudum. Similar to A. glandiformis (Lamarck, 1783) was published without designated a type and proper description.

139 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 121 Although in several collections of A. macrocalyx from main island of New Guinea have striking appearance with thick-brown indumentum distributes on the leaf sheaths, the bright red crownshaft (from Finschhafen), the leaves and the inflorescences but these may reflects to ecological conditions, such as the heath forest. We treat A. congesta and A. ledermanniana as doubtful taxa in this monograph because the type specimens of A. congesta (Ledermann & 7250) and A. ledermanniana (Ledermann 9766) were destroyed during the Second World War in Berlin and we have been unable to locate another duplicate. Beccari s descriptions (Beccari, 1923) indicate that these taxa could fit within the range of variation of A. macrocalyx, but in the absence of authentic materials it is not possible to synonymise these two species. 3. Areca mandacanii Heatubun, Palms 52: 199 (2008). Type: Indonesia, West Papua Province, Sorong Selatan, Teminabuan, Sayal, Maampou Forest, ii.2003, Heatubun 423 (holotype: BO; isotypes: K, MAN). Description: Solitary, moderate tree palm. STEM up to 15 m tall, 8 10 cm diam.; internodes cm long, dark green, shiny, nodal scars conspicuous, white. LEAVES 8 in crown, pinnate, appearing plumose, cm long (including petiole); sheath tubular, c. 92 cm long, smooth, light green; crownshaft well defined, up to 152 cm long and up to 15 cm diam.; petiole short to 6 cm long, channelled adaxially, rounded abaxially; rachis somewhat arching, with adaxial longitudinal ridge, rounded abaxially; blade with irregularly arranged leaflets, divided into groups, in several planes, c. 60 leaflets on each side, near petiole small and gradually becoming larger, terminal leaflets regularly arranged (c leaflets); leaflets somewhat arching, single-fold, linear, cm long, 2 cm wide, tip acuminate, notched, split to 3 cm long and pointed in a few leaflets near the petiole, papery, green adaxially and light green abaxially. INFLORESCENCE infrafoliar, c. 60 cm long at anthesis, protandrous, branching to 2 3 orders; peduncle 5 cm long, green with numerous minute red-brown dots; prophyll 64 6 cm, borne about 1/3 way up the peduncle, lanceolate, 2-keeled, papery, cream to light brown, entirely enclosing the inflorescence, then splitting longitudinally and falling before staminate anthesis; rachis green to whitish green; rachis bracts not persistent; rachillae numerous, 37 cm long, covered by light brown to chocolate-brown thick indument, highly contrasting with the rachis, first branching rachillae c. 50 cm long, elongate. STAMINATE FLOWERS small,

140 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 122 triangular, mm, asymmetric; sepals 3, low, about 2 1 mm; petals 3, strongly keeled, 4.5 mm long, c mm wide at basal; pistillode mm, trifid, dark brown; stamens 6, 1.5 mm long, sagittate; anthers 1.5 mm long, arrow head shape, creamy, longer than the filaments, twisted basally; filaments 0.5 mm long, dark brown. PISTILLATE FLOWERS larger than the staminate, triangular, mm diam., sitting on the branchnodes; sepals 3, imbricate, triangular, mm long, mm wide, 1 3 mm thick, asymmetrical, thicker at base, cream to light green; petals 3, imbricate, triangular, mm, 0.5 mm thick, cream; staminodes ovate, 10 mm high, 9 mm wide, pointed, brown coloured at the tip, fleshy. FRUITS ellipsoidal, mm, with conspicuous woody discoid depression at apex, shallowly concave, cm diam., stigmatic remains persistent in centre of depression, perianth persistent; epicarp smooth, shiny, c. 0.5 mm thick, dark green when young, turning to golden yellow or light orange when mature; mesocarp fibrous, c. 10 mm thick, but much thicker at the base of fruit (below the seed), where c. 20 mm thick; endocarp very thin, adhering closely to seed. SEEDS c mm, subglobose; endosperm ruminate; embryo basal. Distribution and ecology: Bird s Head Peninsula of Western New Guinea. This palm grows in the transition between swamp forest and lowland rain forest, where the soils are temporarily inundated by water. Specimen examined: INDONESIA. West Papua Province, Sorong Selatan District, Teminabuan subdistrict, Sayal village, Maampow forest, 10 m, 21.ii.2003, Heatubun et al. 423 (holotype BO!, isotypes K!, MAN!); Heatubun et al. 413 (K! spirit collection, MAN!); Heatubun et al. 424 (MAN!); West Papua Province, exact locality unknown, Maturbongs s.n. (K!, MAN). CULTIVATED. Indonesia, West Papua Province, Manokwari District, Reremi, 75 m, 10.iv.2008, Heatubun & Iwanggin 902 (MAN!). Notes: The differences between A. mandacanii and New Guinean Areca including A. catechu already discussed in detail by Heatubun (2008). However, the plumose leaves are the crucial taxonomic character not only for the genus Areca but also in high level classification, in subtribe Arecinae and tribe Areceae see discussion under the leaves morphology.

141 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA Areca novohibernica (Lauterb.) Becc., Bot. Jahrb. Syst. 52: 24 (1914). Nenga novohibernica Lauterb., Bot. Jarhb. Syst. 45: 357 (1911). Type: Papua New Guinea, Bismarck Archipelago, New Ireland, Nabumai, Urwald, Peekel 110 (holotype: B, photo FI; isotypes: FI, K). Areca guppyana Becc., Webbia 4: 258 (1914). Type: Solomon Islands, Shorland Islands, Alu Island, 1 to 2 miles from coast, Guppy 107 (holotype: K). Syn. Nov. Areca salomonensis Burret, Notizbl. Bot. Gart. Berlin-Dahlem 13: 70 (1936) Areca novohibernica var. salomonensis Burret, Notizbl. Bot. Gart. Berlin-Dahlem 13: 69 (1936). Type: Papua New Guinea, Bougainville Island, Kugumaru, Buin District, 2.vii.1930, Kajewski 1908 (holotype: B ; isotype: A). Syn. Nov. Description: Solitary, small and slender, undergrowth palm, tall 2 4 m ( 5 m), with stilt roots or prop roots. STEM 5 10 cm in diam.; internodes 3 4 cm long, light brown to whitish with conspicuous leaf scars (± 1 cm), and shiny green near the crown. LEAVES 5 8 in crown, lamina cm long (including petiole); sheath tubular, cm; crownshaft cm long, light green to green; petiole cm long, c. 1 cm wide at the base, channeled adaxially, rounded abaxially, cover by thick brown indumenta and continue to leaf rachis; leaflets slightly regularly arranged, papery, c. 5 leaflets on each side, the basal most leaflets cm, 3 6 folds and slightly sigmoid with oblique-acuminate tip, the middle leaflets cm, 5 7 folds, slightly sigmoid with oblique-notch tip, splits between folds 3 5 cm depth, terminal leaflets more broad, cm, with 9 15 folds, flabellate with truncate tip, green, slightly discolorous when dried, dark-coloured adaxially, and paler abaxially. INFLORESCENCE infrafoliar, cm, branched to 1 order (sometimes basal most rachillae with 2 branches), erect and will pendulous when fruits in development; prophyll papery, cm, caducous; peduncle 3 6 cm long and 1 2 cm wide at the base; rachilla (including main axis), divaricate, 7 16 cm long and 2 4 mm wide, greenish cream to green, sometimes perianth persistent on rachillae after fruits fallen-off. FLOWERS are borne in triads from the base to tip along the half way length of the rachilla, and continue with only pairs of staminate flowers to the tip. STAMINATE FLOWERS small, elongated, triangular, mm, asymmetrical, white to creamy green, glaucous, spirally arranged on rachilla; calyx fused, low, cup-shaped with margin lobes, mm; petals 3, thick and fleshly, striate, mm, vary in shapes; stamens 6, alternate, mm long, fused with petals up to the half of petal height (epipetalous);

142 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 124 filaments shorter than anther, mm long; anthers mm long, sometimes twisted; pistillode low, about 0.5 mm high, various shapes. PISTILLATE FLOWERS bigger than staminate flowers, mm; sepals 3, imbricate, united 2 4 mm from the base, mm, obovate, thick; petals 3, imbricate, mm; gynoecium mm (including stigma mm), stigma trifid; staminodes circular with 4 6 irregular toothed, membranous. FRUITS small, cm, ellipsoid to slightly ovoid, green yellow orange red when ripen; epicarp thin and smooth; mesocarp fleshy and juicy; endocarp fibrous. SEEDS small, cm, slightly globose to ellipsoid, rounded apical and flatted basally, hard; endosperm ruminate; embryo basal. Distribution and ecology: Areca novohibernica grows on the soil from volcanic ash and limestone at m above sea level, fron the island of Manus, New Britain and New Ireland in the Bismarck Archipelago, and the Solomon Islands. Specimen studied: PAPUA NEW GUINEA. Manus Province, Manus Island, Western Manus, South Coast, 1 km from Kabuli village, 25 m, 24.xi.1975, Sands et al (K!; LAE). West New Britain Province, Kapiura River, near Lavege village, 30 m, 16.iv.1968, Henty & Lelean NGF (BRI, L!, LAE!); Hoskins District, near Dami, Kavui logging area, 15 m, 25.iv.1972, Essig LAE (L!, LAE); Nuau, 210 m, 19.ii.1971, Lelean & Stevens LAE (A!, BRI!, CANB!, K!, L!, LAE); Talasea, Mt. Tangis, ridge below rim of crater, 1350 m, 30.v.1966, Frodin NGF (BRI!, L!, LAE); Kandrian District, Pulie River, 10 miles from the mouth, west side, 30 m, 14.iii.1966, Henty NGF (BRI, L!, LAE). East New Britain Province, Rabaul District, Powell harbour, 30 m, 19.vi.1972, Foreman LAE (LAE!); Awung village, 600 m, 23.iii.1968, Ridsdale & Katik NGF (L!, LAE). New Ireland Province, Namatanai District, Hans Meyer Range, Danfu River valley, c. 8 km W and upstream of Danfu bridge, near Manga, upper terrace, behind base camp, to the north, 220 m, 27.i.1970, Sands 726 (L!, LAE); close to Mindih Lake, east coast, 6 km W of NW Toron, 650 m, 6.x.1975, Sands et al (K!, LAE); Northern Hans Meyer Range, 70 km SE of Namatanai, 850 m, 31.x.1984, Gideon LAE (L!, LAE); ridge adjacent to Weitin River, 1175 m, 27.i.1994, Takeuchi & Wiakabu 9592 (A!, LAE); 31.i.1994, Takeuchi & Wiakabu 9733 (A!, LAE); Feni group, Ambitle Island, Nanum caldera, thermal spring area, 125 m, 16.xi.2003, Takeuchi (A!, LAE); New Hanover, west Lavongai, 2 km N of

143 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 125 Matemulai village, 8.x.1974, Croft & Lelean LAE (L!, LAE). North Solomon Province, Bougainville Island, 15 miles N of Buim, Lake Loloru Crater, lower South slopes, 750 m, Craven & Schodde 202 (CANB!, L!); Kugumaru, Buin, 150m, 2.vii.1930, Kajewski 1980 (isotype A!). THE SOLOMON ISLANDS. Shortland Islands, Alu Island, 1 2 miles from the coast, ii.1885, Guppy 107 (holotype K!). Fauro Island, Kauriki village, ridge leading to peak, 24.iv.1964, Whitmore BSIP 4132 (BSIP, K!, L!). Malaita, Mt. Alasa, 21.x.1965, Corner RSS 223 (K!, BSIP). Notes: Areca novohibernica is similar to Areca vestiaria in its stilt roots, the staminate flowers spirally arranged on the rachilla, the sepals united or calyx tubular, the petals elongated and spathulate, the pistillate flowers also spirally arranged and distributed to the half length of the rachilla, the fruits with fleshy and juicy mesocarps. However, it differs from the latter by its slender solitary habit, the green crown shaft and inflorescence branched to 1 order (rarely 2 orders). It differs from all other EWL species in presence of the velvet-like indumentum on the staminate flowers, epipetalous stamens, anthers irregular and sometimes twisted, and in its distribution at the very limits of subtribe Arecinae in Manus Island, the Bismarck Archipelago and the Solomon Islands. On re-examining the type specimens of A. guppyana (Guppy 107), A. novohibernica (Peekel 110) and A. salomonensis (Kajewski 1908) alongside more recent collections we see no reason to maintain these distinct taxa. All three species (A. guppyana, A. novohibernica and A. solomonensis) were described from very inadequate or single specimen, thus very narrow species concept was applied to those species by earlier authors. The fruits characters used by previous authors (Beccari, 1914a, b; Burret, 1936) to separate the three species, show particularly high phenoplasticity within populations, individuals or even inflorescences.

144 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 126 Figure 5. Areca novohibernica (Lauterb.) Becc. A, Apical portion of leaf. B, Middle portion of leaf. C, Portion of rachilla with staminate and pistillate flowers. D, E, Staminate flower whole and in section. F, G, Pistillate flower whole and in section. H, Inflorescence. I, J, Fruit whole and in section. Scale bar: A, B = 8 cm; C, I, J = 1.5 cm; D, E = 3 mm; F, G = 7 mm; H = 6 cm. A B from Sands 726; C G, I J from Sands 2124; H from Takeuchi Drawn by Lucy T. Smith.

145 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA Areca oxycarpa Miq., Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk. 15: 1 (1868). Type: Celebes, Menado, Mt. Pisah, Riedel s.n. (holotype: L). Areca celebica Burret, Repert. Spec. Nov. Regni Veg. 32: 115 (1933). Type: Celebes, Tomohon, Sarasin s.n. (holotype: B). Syn. Nov. Description: Solitary, small and slender, undergrowth palm, tall m. STEM cm in diam.; internodes cm long, light brown to whitish with conspicuous leaf scars, and shiny green near the crown. LEAVES 6 8 in crown, lamina glabrous, cm long (including petiole); sheath tubular, cm; crownshaft cm long and cm in diam., dark brown to blackish purple with numerous black dot scales; petiole cm long, mm wide and 4 6 mm thick at the base, channeled adaxially, rounded abaxially; leaflets irregularly arranged, papery, 4 8 leaflets on each side, the basal most leaflets cm, ribbed 2 8 ribs, the middle leaflets cm, ribbed 5 7 ribs, sometimes with 1-folded leaflet between two broad leaflets, slightly sigmoid and oblique-acuminate tip except to terminal leaflets, terminal leaflets cm, flabellate, 7 8 ribs, and truncate tip, green, slightly discolorous when dried, dark-coloured adaxially, and paler abaxially. INFLORESCENCE infrafoliar, 8 15 cm long, branched to 1 order (sometimes basal most rachillae with 2 branches), erect; prophyll and bract caducous, not seen; peduncle 1 2 cm long and 5 7 mm wide at the base; rachillae divergent, 8 16, 5 8 cm long, greenish cream to green, sometimes perianth persistent on rachillae after fruits fallen-off. STAMINATE FLOWERS mm, triangular, asymmetrical, creamy to greenish yellow, distichous arranged on rachilla; calyx fused, low, cup-shaped with margin lobes, c mm; petals 3, thick and fleshly, c. 6 3 mm, triangular, striata; stamens 6, mm long; filaments c mm long; anthers mm long; pistillode dimunitive. PISTILLATE FLOWERS to 8 5 mm; calyx c. 7 5 mm, imbricate, greenish; petals 3, c. 8 4 mm, imbricate, cream; gynoecium 8 3 mm (including stigma 1 mm), stigma trifid; staminodes circular, membranous. FRUITS small, cm, elongated to ellipsoid, with 3 4 mm long beak, green to bright yellow or orange when ripen; epicarp smooth and thin; mesocarp fibrous; endocarp thin and fibrous. SEEDS small, cm, ovate, rounded apical and flatted basally, hard; endosperm ruminate; embryo basal.

146 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 128 Distribution and ecology: This species is endemic to the northern part of Sulawesi in the Indonesian provinces of North Sulawesi and Gorontalo. Areca oxycarpa grows at an altitude m above sea level, on volcanic soil in primary forest near streams on slopes. Specimen studied: INDONESIA. North Sulawesi Province, Bolaang Mongondow District, Dumoga Nani Warta Bone National Park, Tapak Kulintang, 280 m, 7.iii.1984, Mogea 5070 (BO!, K!); Dumoga Nani Warta Bone National Park, West Dumoga subdistrict, Matayangan village, Tumokang Kasinggolan forest, 225 m, 16.iii.2008, Heatubun et al. 877 (BO!, BPK Manokwari!, K!); 272 m, 16.iii.2008, Asmarayani et al. 461 (BO!, BPK Manokwari!); 284 m, 17.iii.2008, Heatubun et al. 883 (BO!, BPK Manokwari!, K!); Kasinggolan, 200 m, 28.ii.1994, Mogea 4961 (BO!, K!); Gunung Mogogonipa, 300 m, 1.iv.1985, de Vogel & Vermeulen 6950 (BO, L!, K!); Duloduo, 250 m, 16.ix.1984, Whitmore & Sidiyasa 3407 (BO!, FRI Bogor, K!); Minahasa District, Tomohon subdistrict, 450 m, iv.1974, Kaseger 210 (BO!, K!, L!); Tomohon, Sarasin s.n. (holotype B!). CULTIVATED. USA, Hawaii, Honolulu, the Harold L. Lyon Arboretum, accession no. L , i.2008, Bacon 115 (K!). Notes: The inflorescence of A. oxycarpa is slightly congested and similar to A. macrocalyx in branching to two orders and being protogynous. However, they can be differentiated easily. Areca oxycarpa is always a small and slender-undergrowth palm, with erect and rather stiff inflorescences. The pistillate flowers (in triads) are confined to the basal 1/3 length of rachilla. Moreover, A. oxycarpa has a dark brown to black leaf sheath. Burret (1933) published A. celebica from inadequate material collected by Sarasin in Tomohon, North Sulawesi. He separated his species, A. celebica from A. oxycarpa just based on fruit characters. However, the fruits characters in the genus Areca are very variable and there is no justification for maintaining these taxa on these grounds. Furtado (1933) expressed his suspicion about A. celebica in his monograph. However, he was just mention that A. celebica may be a form or variety of A. oxycarpa without makes any formal transferred. Our observations in the field and herbariuim support the reduction to a single species.

147 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA Areca vestiaria Giseke, Prael. Ord. Nat. Pl.: 78 (1792). Pinanga vestiaria (Giseke) Blume, Rumphia 2: 77 (1839). Seaforthia vestiaria (Giseke) Mart., Hist. Nat. Palm. 3: 313 (1849). Ptychosperma vestiarium (Giseke) Miq., Fl. Ned. Ind. 3: 31 (1855). Mischophloeus vestiarius (Giseke) Merr., Interpr. Herb. Amboin.: 121 (1917). Type: Pinanga sylvestris e Buro Rumphius, Herb. Amboin. 1: (Lectotype: Dransfield, 1974). Ptychosperma paniculatum Miq., Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk. 11(5): 3 (1868). Areca paniculata (Miq.) Scheff., Tijdschr. Ned.-Indië 32: 179 (1873). Mischophloeus paniculatus (Miq.) Scheff., Ann. Jard. Buitenzorg 2: 152 (1876). Type: North Moluccas, Bacan Island, Teysmann & De Vriese (isotype: L). Areca leptopetala Burret, Notizbl. Bot. Gart. Berlin-Dahlem 13: 199 (1936). Type: South Celebes, Porema, 10.ix.1929, Kjellberg 2324 (holotype: B ; isotype: BO). Areca langloisiana Potztal, Willdenowia 2: 628 (1960). Type: A. C. Langlois s.n. (holotype: B). Description: Solitary to clustering, moderate palm, tall to 10 m ( 15 m) high, with stiltroots or prop roots up. STEM 7 10 cm in diam.; internodes cm long, greenish yellow with conspicuous leaf scars. LEAVES about 11 in crown, cm long (including petiole); sheath tubular, cm; crown shaft cm long and cm in diam., orange to reddish with numerous dot brown scales; petiole cm long, 2 4 cm wide and 1 2 mm thick at the base, channeled adaxially, rounded abaxially, yellowish green to orange; leaflets somewhat irregularly arranged, papery to leathery, leaflets on each side, the basal-most leaflets cm, 2 3 folds, lanceolate to sigmoid with oblique tip, the mid-leaflets cm, 3 4 folds, slightly sigmoid with oblique-notch tips, terminal leaflets cm, flabellate, 4 11 folds, notch at tips with splits 5 11 cm long between two folds, green, slightly discolorous when dried, darkcoloured adaxially, and paler abaxially, with fine sparse ramenta on mid-vein on abaxial surface. INFLORESCENCE infrafoliar, cm long and cm wide, branched to 2 orders, erect to re-curved and later pendulous in fruit development; prophyll cm, thin, papery, caducous, orange to reddish; peduncle 3 5 cm long and cm wide at the base; rachillae numerous, up to 20 cm long, yellowish cream, orange to red, sometimes perianth persistent on rachillae after fruits fallen. FLOWERS are borne in triads from the base to tip along the half way length of the rachilla, and continue with only pairs of staminate flowers to the tip. STAMINATE FLOWERS mm at anthesis, triangular, asymmetrical, elongated, creamy to yellowish white, spiral arranged on rachilla; calyx

148 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 130 fused, low, cup-shaped with shallow margin lobes, 2 mm wide and mm high; petals 3, thick and fleshly, mm, elongated, ovate; stamens 6, sagittate basally, white to yellowish cream, mm; filaments shorter than anther, mm, white and becoming dark brown after anthesis; anthers mm long, arrow-head shaped; pistillode low, various shapes. PISTILLATE FLOWERS mm, triangular, asymmetrical; sepal 3, mm, triangular, strongly imbricate; petal 3, almost similar size with sepal, triangular, strongly imbricate; gynoecium mm (including stigma), tubular; staminodes 4 6, triangular (wedge shaped), low, membranous. FRUITS cm, obovate to ellipsoid, yellowish orange to red when ripen, costate when dried; epicarp thin and smooth; mesocarp fleshly and juicy; endocarp thin and fibrous. SEEDS cm, ovate, rounded apical and flatted basally, hard; endosperm deeply ruminate; embryo basal. Distribution and ecology: This species has relatively wide distribution in the EWL hotspot, occurring throughout Sulawesi and the Moluccas except for Ambon and the islands in southeast (Kei, Aru and Tanimbar). Areca vestiaria grows on volcanic soil from the lowland area to the highland (below 2000 m above sea level). Two collections (Mogea 1303, 1305) have been made on the summit of Mt. Maleno in Central Sulawesi, at an altitude 2260 m above sea level, from the populations with the palm tall to 30 m high and these quite exceptional to A. vestiaria. Specimen studied: INDONESIA. North Sulawesi Province, Minahasa District, Mt. Soputan, Langoan, 900 m, 11.x.1973, Dransfield & Mogea JD 3755 (BO!, K!, L!, BH); Tomohon subdistrict, Mt. Lokon, 950 m, 3.x.1973, Dransfield & Mogea 3714 (BO!, K!, L! BH); 1000 m, 3.x.1973, Dransfield & Mogea 3716 (BO!, L!); Bolaang Mongondow District, Dumoga Nani Warta Bone National Park, West Dumoga subdistrict, Matayangan/Kasinggolan village, 225 m, 17.iii.2008, Heatubun et al. 879 (BO!, BPK Manokwari!, K!); 257 m, 17.iii.2008, Heatubun et al. 885 (BO!, BPK Manokwari!, K!); Edwards camp, 750 m, 27.iii.1985, de Vogel & Vermeulen 6768 (BO!, K!, L!); vicinity of Mt. Sinombayuga, 1750 m, 29.ix.1991, Milliken & Bonde 1115 (BO!, K!); Dumoga Nani Warta Bone National Park, 250 m, 16.ix.1984, Whitmore & Sidiyasa TCW 3411 (K!); Mt. Mogogonipa, 650 m, 10.iv.1985, de Vogel & Vermeulen 7070 (BO!, K!, L!); 220 km W of Manado, 50 km inland from Pangi, on tributary of Ilanga River, m, 13.iii.1990, Burley et al (A, BO!, K!). Central Sulawesi Province, Kulawi subdistrict, Moa, on

149 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 131 the summit of Mt. Maleno, 2260 m, 17.x.1977, Mogea 1303 (BO!, K!, L!); Mogea 1305 (BO!, K!, L!); Luwuk area, inland from Batui and Saseba, on Batui River, at Totup camp 2, 2 hours upriver from Sinsing, Mt. Sohean, 170 m, 21.x.1989, Coode 6045 (BO! K!); Palu, road Palu-Sopu valley, c. 40 km SSE of Palu, 600 m, 21.iv.1979, de Vogel 5006 (BO!, K!, L!); Palu, Sopu valley, c. 80 km SSE of Palu, 1000 m, 27.v.1979, de Vogel 5065 (BO!, K!, L!); Donggala District, Pangi-Binangga Nature Reserve, Puncak Beringin study area, 560 m, 14.x.1991, Bynum 8201 (A, BO!, K!); Mt. Roroka, Timbu, west slopes, 2000 m, 1979, van Balgooy 3267 (BO!, K!, L!); by the river S of Tongoa, 650 m, 3.iii.1981, Johansson et al. 151 (BO!, K!, L!); 5.iii.1981, Johansson et al. 179 (BO!, L!, K!); area of Mt. Nokilalaki, of lake Lindu to Sidaunta, 1000 m, 3.v.1975, Meijer 9993 (A, BO!, L!, MO, LEX). West Sulawesi Province, Mamuju District, Kaluku subdistrict, Popangatalu village, Kona-Kona River, c. 200 m, 4.ii.1993, Afriastini 2060 (BO!, K!). North Moluccas Province, Bacan Island, Mt. Sibela, near Waiaua, 1050 m, 25.x.1974, de Vogel 3655 (BO!, K!, L!); 250 m, 28.x.1974, de Vogel 3725 (BO!, K!, L!); near Amasing River, 5 m, 6.xi.1974, de Vogel 3926 (BO!, K!, L!); de Vogel 3927 (BO!, K!, L!); de Vogel 3929 (BO!, K!, L!); Halmahera, 20 km SE of Dodinga, Darco/Modul logging camp, Tapayo, 600 m, 9.ix.1985, Sidiyasa et al. TCW 3611 (K!); 630 m, 13.ix.1985, Sidiyasa et al. TCW 3651 (K!); Mt. Sahu, near Susupu, 500 m, 5.x.1974, de Vogel 3272 (BO!, K!, L!); 6.x.1974, de Vogel 3285 (BO!, K!, L!). Moluccas Province, Buru Island, west Buru, Wae Nibe, wood industry base camp 2, 20 km, Wae Ili, 9.xi.1984, Mogea & Ismail 5202 (K!, BO!). CULTIVATED. Malaysia, Malay Peninsula, Kepong, FRIM office, Barfod s.n. (AAU photo!). Indonesia, Bogor Botanic Garden, II. J. 14, iv-v.1936, Furtado 155 (K!). Notes: Areca vestiaria is a unique species in the genus Areca with a distinct combination of morphological characters: moderate, clustering or solitary palm with stilt roots, the brilliant orange to bright red crownshaft, the inflorescence always branched to 2 orders, the flowers (staminate and pistillate) spirally arranged on rachillae and the fruits with fleshy and juicy mesocarps. See discussion above for comparison with A. novohibernica. The arrangement of staminate flowers was used by Scheffer (1876) as the basis for Mischophloeus, a genus distinct from Areca. Subsequently, Furtado (1933) synonymized it with Areca, reducing it to a section under subgenus Beccarioareca. This palm has a relatively wide range of distribution in the EWL hotspot and is morphologically variable, thus this taxon has been a subject of taxonomic confusion and nomenclature problems (Giseke, 1792; Martius, 1849; Miquel, 1868; Scheffer, 1873, 1876; Merrill, 1917; Furtado,

150 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA ; Burret, 1936; Potztal, 1960; Dransfield, 1974) since it first appeared in Herbarium Amboinense (Rumphius, 1741). DOUBTFUL OR UNCERTAIN NAMES Areca congesta Becc., Engl. Bot. Jahrb. 58: 441 (1923). Type: Papua New Guinea, Sepik, Ledermann (holotype: B ). See notes under Areca macrocalyx for discussion. Areca ledermanniana Becc., Engl. Bot. Jahrb. 58: 441 (1923). Type: Papua New Guinea, April River, Ledermann 9766 (holotype: B ). See notes under Areca macrocalyx for discussion. ACKNOWLEDGEMENTS Many individuals and institutions contributed to the completion of this work. We would like to thank the Keepers and staff of herbaria AAU, BO, E, FI, K, L, MAN, SAR and BPK Manokwari for access to their specimens, databases and for providing loans in many cases. The authorities in the Provinces of Papua, West Papua, North Sulawesi, and Gorontalo were allowed CDH to conduct his fieldtrips in those areas. Maikel Simbiak, Herkilaus Rumaikewi, Lukas Manua, Krisma Lekitoo, Tobias Paiki, and Dr. Herni Simbala are thanked for their helps with the permits and all supports including organized CDH s fieldtrips on their area. Special thank to Balai Penelitian dan Kehutanan Manokwari for their supports and helps in many ways. Lucy T. Smith was rendered beautiful illustrations and using in this monograph. Drs. Christine D. Bacon and Raymond Baker from Lyon Arboretum, Hawaii which have kindly provided herbarium vouchers and DNA materials of Areca from their garden collections. Professor Edwino E. Fernando sharing his expertise on the Philippine s Areca. And Dr. Tom D. Evans to his kindly was sending his collection of a new Areca from Cambodia. This paper is part of CDH s PhD thesis which was conducted at Institut Pertanian Bogor and Royal Botanical Gardens Kew. And he would like to express his gratitude to funding supports provided by the Royal Botanic Gardens Kew, UK (BAT Biodiversity Partnership and Pacific Biological Foundation) and BPPS Dikti Depdiknas for his PhD scholarships.

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154 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 136 Harley MM Occurrence of simple, tectate, monosulcate or trichotomosulcate pollen grains whitin the Palmae. Review of Palaeobotany and Palinology 64: Harley MM Palm pollen and the fossil record. Ph.D. thesis, University of East London. Harley MM Palm pollen: overview and examples of taxonomic value at species level. Memoirs of the New York Botanical Garden 83: Harley MM, Baker WJ Pollen aperture morphology in Arecaceae: application within phylogenetics analyses, and summary of the fossil record of palm-like pollen. Grana 40: Harley MM, Dransfield J Triporate pollen in Arecaceae. Grana 42: Heatubun CD A monograph of Sommieria (Arecaceae). Kew Bulletin 57: Heatubun CD A new species of Areca from Western New Guinea. Palms 52: Heatubun CD, Baker WJ, Mogea JP, Tjitrosoedirdjo SS, Dransfield J A monograph of Cyrtostchys (Arecaceae). Kew Bulletin 64: Holmgren PK, Holmgren NH, Barnett LC Index herbariorum, Eight editions. New York: New York Botanical Garden. Hope GS, Hartemink AE Soils of Papua. In: Marshall AJ, Beehler BM, Eds. The ecology of Papua. Singapore: Periplus Editions, Johns RJ Malesia an introduction. Curtis s Botanical Magazine 12: Lee KK, Choi JD Areca catechu L. extract. I. effect on elastase and aging. Journal of Cosmetics Science 49: Leopold AC Many modes of movement. Science 288: Lewis CE, Doyle JJ A phylogenetic analysis of tribe Areceae (Arecaceae) using two low-copy nuclear genes. Plant Systematics and Evolution 236: Linneaus C Species Plantarum: p Listabarth C A survey of pollination strategies in the Bactridinae. Bulletin de l'institut franáais d'etudes andines 21: Listabarth C Pollination in Geonoma macrostachys and three congeners, G. acaulis, G. gracilis, and G. interupta. Acta Botanica 106: Listabarth C Pollination and pollinator breeding in Desmoncus. Principes 38: Listabarth C Pollination of Bactris by Phyllotrox and Epurea. Implications of the palm breeding beetles on pollination at the community level. Biotropica 28:

155 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 137 Listabarth C Pollination studies of palm populations: A step toward the application of a biological species concept. Memoirs of the New York Botanical Garden 83: Loo AHB, Dransfield J, Chase MW, Baker WJ Low copy nuclear DNA, phylogeny and the evolution of dichogamy in the betel nut palms and their relatives (Arecinae; Arecaceae). Molecular Phylogenetics and Evolution 39: Martelli U La sinonimia delle Palme gerontogee della tribù delle Areceae. Nuovo Giornale Botanico Italiano 42: Martius CFP von Historia Naturalis Palmarum, vol. 3. Munich. Merrill ED An interpretation of Rumphius s Herbarium Amboinense. Manila: Bureau of Science. Merrill ED An Enumeration of Philippine Flowering Plants, vol. 1. Manila: Bureau of Printing. Miquel FAG Flora Indiae Batavae, vol. 3. Amsterdam. Miquel FAG De palmis Archipelagi Indici observationes novae. Verhandelingen der Koninklijke Nederlandsche Akademie van Wetenschappen, afdeeling Natuurkunde 11: McDade LA Species concepts and problems in practice: Insight from botanical monograph. Systematic Botany 20: Moore HE Jr., Dransfield J Typification of Linnean palms. Taxon 28: Norton SA Betel: consumption and consequences. Journal of American Academy of Dermatology 38: Potztal E Eine neue Palme von Celebes. Willdenowia 2: Ray AK, Reddy DVS Performance of Areca-based high density multi species cropping system under different level of fertilizer. Tropical Agriculture 78: Read RW New chromosome counts in palmae. Principes 10: Richard PW The tropical rain forest. Cambridge: Cambridge University Press. Rumphius GE Herbarium Amboinense, vol. 1. Amsterdam. Sato D Karyotype alteration and phylogeny, VI. Karyotype analysis in Palmae. Cytologia 14: Scheffer RHCC Sur quelques palmiers du groupe des Arécinées 1. Natuurkundig Tijdschrift voor Nederlandsch Indië 32: Scheffer RHCC Sur quelques palmiers du groupe des Arécinées 2. Annales du Jardin Botanique de Buitenzorg 1:

156 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 138 Schoute JC Ueber das dickenwachstum der palmen. Annales du Jardin Botanique de Buitenzorg 2: Sharma AK, Sarkar SK Cytology of different species of palms and its bearing on the solution of the problems of phylogeny and speciation. Genetica 28: Silberbauer-Gottsberger I Pollination and evolution in palms. Python 30: Sullivan RJ, Allen JS, Otto C, Tiobech J, Nero K Effects of chewing betel nut (Areca catechu) on the symptoms of people with schizophrenia in Palau, Micronesia. British Journal of Psychiatry 177: Soil Survey Staff Soil taxonomy: a basic system of soil classification for making and interpreting soil surveys. Natural Resources Conservation Service, Agricultural Handbook 436. U.S. Department of Agriculture, Washington, D.C. Tomlinson PB Palmae. In: Metcalfe CR, Ed. Anatomy of the monocotyledones II. Oxford: Clarendon Press. Tomlinson PB The structural biology of palms. Oxford: Clarendon Press. Trenel P, Gustafsson MH, Baker WJ, Asmussen-Lange CB, Dransfield J, Borchsenius F Mid-Tertiary dispersal, not vicariance explains Gondwanan distribution pattern in wax palm subfamily (Ceroxyloideae: Arecaceae). Molecular Phylogenetics and Evolution 45: Uhl NW, Dransfield J Genera Palmarum. Kansas: Lawrence, Allen Press. Uhl NW, Dransfield J Genera Palmarum after ten years. Memoirs of the New York Botanic Garden 83: Van Balgooy MMJ The phytogeographical position of Sulawesi (Celebes). In: Hovenkamp P, Ed. Systematics and Evolution: a matter of diversity. Utrecht: University of Utrecht, Van Steenis CGGJ The delimitation of Malaysia and its main plant geographical regions. Flora Malesiana 1, 1: 1xx 1xxv. Van Steenis CGGJ Plant-geography of east Malesia. Botanical Journal of the Linnean Society 79: Wallace AR On the physical geography of the Malay Archipelago. Journal of the Royal Geographical Society 33: Zumbroich TJ The origin and diffusion of the betel chewing: a synthesis of evidence from South Asia, Southeast Asia and beyond. Electronic Journal of Indian Medicine 1:

157 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 139 APPENDIX LIST OF SPECIMENS EXAMINED AND IDENTIFIED Species numbers are given in the brackets and in bold after collection numbers. Key: (1) A. catechu, (2) A. macrocalyx, (3) A. mandacanii, (4) A. novohibernica, (5) A. oxycarpa, (6) A. vestiaria. Afriastini: 2060 (6); Anonymous: (Cult. Kew) s.n. (1); (Cult. Calcutta) s.n. (1); (Cult. Singapore) s.n. (1); (Cult. Singapore, lawn X) s.n. (1); (Cult. Singapore, lawn J) s.n. (1); (Cult. Bogor, lawn II.F.9) s.n. (1); (Cult. Bogor, lawn VII.B.78) s.n. (1); (Cult. Bogor, lawn VII.B.79) s.n. (1); (Cult. Bogor, lawn VII.B.80) s.n. (1); (Cult. Bogor, lawn VII.B.81) s.n. (1); (Cult. Bogor, lawn VII.B.84) s.n. (1); (Cult. Bogor, lawn VII.B.85) s.n. (1); (Cult. Bogor, lawn VII.B.87) s.n. (1); (Cult. Bogor, lawn VII.B.88) s.n. (1); (Cult. Bogor, lawn VII.B.89) s.n. (1); (Cult. Bogor, lawn VII.B.91) s.n. (1); (Cult. Bogor, lawn XI.B.(XX)6) s.n. (1); (Cult. Bogor, lawn XIII.A.6) s.n. (1); Asmarayani et al.: 461 (5); Backer: (1); Baker: 136 (1); Bacon: 115 (5); Baker & Utteridge: 591 (2); Baker et al.: 631 (2); 1098 (2); 1100 (2); Bakhuizen van den Brink: 6724 (1); 6782 (1); 6788 (1); 6789 (1); 6807 (1); 6866 (1); Bakia: 416 (1); Banka et al.: 2001 (2); Barfod: s.n. (1); Barfod et al.: 416 (2); 488 (2); (1); Bartlett: (1); Beccari: PB 3112 (1); s.n. (2); Beguin: 330 (1); 1613 (1); Bloembergen: 3555 (1); 3556 (1); Boden-Kloss: (1); Borden: (Forest Bureau 1272) (1); Borssum-Waalkes: 350 (1); Brass: 3971 (2); 7001 (2); 7170 (2); 7188 (2); 7386 (2); 7901 (2); 8189 (2); (2); 13437A (2); (2); Burley et al.: 3856 (6); 4215 (1); Bynum: 8201 (6); Canton Christian College: 9808 (1); Ching & Tan: TC 01 (1); Chen: (1); Clemens: (2); (1); Conklin: 1055 (1); (1); Coode: 6045 (6); 6074 (1); Corner: RSS 223 (4); Craven & Schodde: 202 (4); Croft & Katik: NGF (2); Croft & Lelean: LAE (4); Croft et al.: LAE (2); Darbyshire: 921 (2); Darbyshire & Hoogland 8089 (2); Desianto: 4 (2); de Vogel: 3266 (1); 3272 (6); 3285 (6); 3655 (6); 3725 (6); 3926 (6); 3927 (6); 3929 (6); 5006 (6); 5065 (6); de Vogel & Vermeulen: 6768 (6); 6950 (5); 7070 (6); de Vogel et al.: 6048 (1); Dransfield: JD 1403 (1); JD 1467 (1); JD 1471 (1); Dransfield & Mogea: 3714 (6); 3716 (6); JD 3755 (6); JD 3808 (1); Dransfield et al.: JD 7533 (2);Duaneh: 255 (1); Dubuc: 1869 (1); Du Bois: 24 (1);

158 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 140 Edeling: s.n. (1); Elmer: 7795(1); (1); Essig: LAE (4); Essig & Young: LAE (2); Fernando: 546 (1); 600 (1); 665 (1); Floyd: 6422 (2); Foreman: LAE (4); Frodin: NGF (4); Furtado: 155 (6); s.n. (2); Gideon: LAE (2); LAE (2); LAE (2); LAE (4); s.n. (2); Graff: 134 (1); Griffith: s.n. (1); Guppy: 107 (4); Gusbager et al.: 20 (2); Heatubun et al.: 292 (2); 331 (2); 360 (2); 413 (3); 423 (3); 424 (3); 747 (2); 751 (1); 752 (1); 753 (1); 796 (2); 798 (2); 799 (2); 870 (1); 876 (2); 877 (5); 879 (6); 883 (5); 885 (6); Heatubun & Iwanggin: 902 (3); Henry: 8406 (1); Henty: NGF (4); NGF (2); Henty & Lelean: NGF (4); Henty et al.: NGF (2); NGF (2); Hoogland: 9026 (2); Hoogland & Craven: (2); Huang & Kao: 5234 (1); Iboet: 440 (1); Jacobs: 8761 (2); 9002 (2); s.n. (2); Johansson et al.: 151 (6); 179 (6); Kajewski: 1980 (4); Karenga: LAE (2); LAE (2); Karenga & Obedi: LAE (2); Kaseger 210 (5); Kazakoff: NGF 7057 (2); Koorders: 396 (1); Lai: S (1); Larivita & Katik: LAE (2); Lau: 226 (1); Lee: S (1); Lelean & Stevens: LAE (4); Liang: (1); Lugas: 442 (1); Mansus & Aban: SAN (1); Manuputty: BW 958 (2); Masters: s.n. (1); Maturbongs et al.: 605 (2); 728 (2); s.n. (3); Meijer: 9993 (6); Merrill Species Blancoanae: 213 (1); Milliken & Bonde: 1115 (6); Mogea: 1303 (6); 1305 (6); 3139 (2); 3140 (1); 4961 (5); 5070 (5); 6204 (2); Mogea & Ismail: 5202 (6); Moore & Whitemore: 9303 (BSIP 4051) (2); Morren: 3049 (2); Noblick et al.: 5180 (1); Othman & Munting: S (1); Powell: BSIP (2); Pullen: 725 (2); 1712 (2); 2638 (2); 7299 (2); 7641 (2); Ramos: Bureau of Science (1); Reeve: 889 (2); Reynoso et al.: 7363 (1); Ridsdale & Katik: NGF (4); Rutherford & Bandara: 124 (1); 154 (2); Sambuling: 8 (1); 76 (1); Sands: 726 (4); Sands et al.: 2124 (4); 2756 (4); 6365 (2); Sarasin: s.n. (5); Schodde & Craven: 4029 (2); Sibil: 153 (1); Sidiyasa et al.: TCW 3580 (1); TCW 3611 (6); TCW 3651 (6); Sinclair: 3272 (1); Soibeh: 778 (1); Streimann & Katik: NGF (2); NGF (2); Streimann & Lelean: NGF (2); NGF (2); Sumawong: s.n. (2); Takeuchi: (2); (2); (4); Takeuchi & Wiakabu: 9592 (4); 9733 (4); Takeuchi et al.: (2); (2); Tsang: 146 (1);

159 C. D. HEATUBUN ET AL.: EAST OF WALLACE S LINE ARECA 141 van Balgooy: 3267 (6); 6864 (2); van Steenis: 212 (1); Veersteg: 1592 (2); 1782 (2); Vinas: LAE (2); Wanggai et al.: 3 (2); Waterhouse: 185 (2); s.n. (2); White: NGF (2); NGF (2); Whitmore: BSIP 1268 (2); BSIP 4132 (4); BSIP 4273 (2); Whitmore & Sidiyasa: 3407 (5); TCW 3411 (6); Williams: 330 (1); Wilson: 9891 (1); Wingfield: 3708 (1); Zippel: s.n. (2);

160 GENERAL DISCUSSION Taxonomic Revision of Genus Areca Areca as palm genus is defined in its traditional concept by the presence of a crownshaft, infrafoliar inflorescences, the single inflorescence bract (prophyll), complete floral triads confined to the basal part of the branched inflorescence or its main axis, the symmetrical fruits with basally attached seed, and the ruminate endosperm with basal embryo (Dransfield, 1984; Uhl & Dransfield, 1987; Dransfield et al., 2008). And this concept has been verified in modern phylogenetics analysis, in which the genus Areca is monophyletic (Loo et al., 2006; Norup et al., 2006; Baker et al., 2009) and reconfirmed in this study. This palm genus has wide range of morphological variations, as similar to its distribution extend from Asia to west Pacific. In general, the variation on morphological characters: from the stemless habit to robust palm trees; solitary to clustering; stilt roots presence and absence; from simple-bifid leaves to pinnate leaves with many leaflets; petiole very short or almost missing to long petiole; leaflets regularly arranged to irregularly arranged or plumose; inflorescence interfoliar to infra foliar; the inflorescence few to many with flat, stiff and thick or tubular and rounded rachillae; staminate flowers uniseriate, distichous to spirally arranged on rachillae; only one triads at rachillae base to some and on the rachis; fruits and seeds varying in the shapes and sizes, including pollen morphology. And to understand these variations within the genus Areca is need to carry out the morphological comparative study using large numbers of specimens and spending some time in the field observed variations in populations of Areca in their natural habitat. In related to revise the whole genus of Areca, our understanding about the variations is very important, especially if we work with morphological species concept as the frame work, with combining the field experience and the ability to access all source of data makes our decisions to delimit species more actual. There are differences between our results and previous workers to recognised taxa in the genus Areca is mainly due to different perspective in application of morphological species concept broader in ours, and the ability to access new data and modern methods, including using molecular technique and new herbarium materials from the recent expeditions. Our results, we recognised 41 species of Areca from 47 species as previously thought (Dransfield et al., 2008), including 8 new species (A. mandacanii Heatubun, 2008) and 12 names treated as new synonymy, 2 treated as doubtful species (species incertae sedis) and A. chaiana J. Dransf. must be excluded from the genus Areca to proper palm genus (species sedis mutabilis). We also recognised two subgenera

161 143 (Areca and Beccarioareca) within genus Areca based on the molecular phylogenetics analysis. Phylogeny of Genus Areca Phylogeny based classification for all living organisms are really urgently needed, with no exceptional for plants, palms and including this genus Areca. The last classification has been applied to the genus Areca since Furtado (1933) published his synopsis. The Furtado s infrageneric classification (1933) comprising two subgenera and five sections: subgenus Blumeoareca (sections Arecella, Oeotheanthe and Axonianthe); and subgenus Beccarioareca (sections Microareca and Mischophloeus). However, these relationships within the genus are based on morphological affinities alone (Furtado, 1933; Dransfield, 1984; Harley & Dransfield, 2003), and it has been suggested that they need to be reassessed using modern methods (Dransfield, 1984), such as the molecular phylogenetics analysis using DNA as source of taxonomic evidence. In our study, we used two regions of nuclear DNA (low-copy nuclear genes PRK and RPB2), in which these both genes have been proved satisfied from previous study (Bayton 2005; Cuenca et al., 2007; Gunn, 2004; Lewis & Martinez 2000; Lewis et al. unpublished; Loo et al. 2006; Norup 2004; Norup et al., 2006; Roncal et al. 2005; Thomas et al. 2006; Trenel et al., 2007) and also in our results to resolved the relationships among close related species in palms. We discovered two distinctive evolutionary lines within genus Areca and reconfirmed the relationships between Areca and other member of subtribe Arecinae with highly resolution in our phylogeny tree. Although the relationships in species level are not clearly resolved due to limitation of samples used in this study, but we have description of the relationships among the certain species used in the study, and let us to make prediction to the rest species of Areca, including compare to morphological affinities. Incongruence between morphological data and molecular data also faced in our study, however, a few morphological characters supports our results, especially in subgeneric level. From biogeography point of view, the long dispersal event is suggested as the factor to reflect the recent distributional pattern of the genus Areca, except to the widely cultivated species A. catechu L. the betel nut palm. Biogeographical barrier, the Wallace s line is not affected for the distribution of Areca species and this fact refused the existence of bimodal distribution pattern, west of Wallace s line or Sunda shelf and east of Wallace s line or Sahul shelf. These findings are also found in subtribe Arecinae (Loo et al., 2006).

162 144 Literature cited Asmussen, C. B. and M. W. Chase Coding and non-coding plastid DNA in palm systematics. Amer. J. Bot. 88: Baker, W. J., C. B. Asmussen, M. W. Chase, J. Dransfield, F. Forest, M. M. Harley, V. Savolainen, N. W. Uhl, M. Wilkinson Complete generic level phylogenetic analyses of palms (Arecaceae) with comparisons of Supertree and Supermatrix approaches. Systematic Biology. doi: /sysbio/syp021 Bayton, R. P Borassus L. and the Borassoid palms: systematics and evolution. Ph.D. Thesis, University of Reading. Cuenca A, Asmussen-Lange CB, Borchsenius FB A dated phylogeny of the palm tribe Chamaedoreeae supports Eocene dispersal between Africa, North and South America. Molecular Phylogenetics and Evolution 46: Dransfield, J The genus Areca (Palmae: Arecoideae) in Borneo. Kew Bull. 39: Dransfield, J. and N.W. Uhl An outline of classification of palms. Principes 30: Dransfield, J. and N. W. Uhl Palmae. In: Kubitzki, K (Ed.), the families and genera of vascular plants, vol. IV. pp , Berlin: Springer. Dransfield, J., N. W. Uhl, C. B. Asmussen, W. J. Baker, M. M. Harley and C. E. Lewis A new phylogenetic classification of the palm family, Arecaceae. Kew Bull. 60: Dransfield, J., N. W. Uhl, C. B. Asmussen-Lange, W. J. Baker, M. M. Harley and C. E. Lewis Genera Palmarum: The Evolution and Classification of Palms. Kew: Royal Botanic Gardens Kew. Flynn, T Morphological variation and species limits in the genus Areca (Palmae) in New Guinea and the Solomon Islands. Unpublished Master thesis, University of Wales, Bangor. Furtado, F. X The limits of the genus Areca L. and its sections. Repert. Spec. Nov. Regni Veg. 33: Govaerts, R., and J. Dransfield World checklist of palms. Kew: Royal Botanic Gardens Kew. Gunn BF The phylogeny of Cocoeae (Arecaceae) with emphasis on Cocos nucifera. Annals Missouri Botanical Garden 91:

163 145 Hahn, W. J. 2002a. A molecular phylogenetic study of the Palmae (Arecaceae) based on atpb, rbcl and 18S nrdna sequences. Syst. Biol. 51: Hahn, W. J. 2002b. A phylogenetic analysis of the Arecoid line of palms based on plastid DNA sequence data. Mol. Phylogenet. Evol 23: Harley, M. M. and J. Dransfield Triporate pollen in the Arecaceae. Grana 41: Heatubun, C. D A new species of Areca from Western New Guinea. Palms 52: Lewis, C. E., and J. J. Doyle A phylogenetic analysis of tribe Areceae (Arecaceae) using two low-copy nuclear genes. Plant Syst. Evol. 236: Lewis, C. E. and N. Martinez Identity of the Hyophorbe palms at the botanical garden of Cienfugoes, Cuba. Palms 44: Loo, A. H. B., J. Dransfield, M. W. Chase and W. J. Baker Low copy nuclear DNA, phylogeny and the evolution of dichogamy in the betel nut palms and their relatives (Arecinae; Arecaceae). Mol. Phylogenet. Evol. 39: Norup, M.V A molecular systematic study of Heterospathe and Rhopaloblaste (Arecaceae, Areceae). Masters Thesis, University of Aarhus. Norup, M. V., J. Dransfield, M. W. Chase, A. Barfod, E. Fernando, and W. J. Baker Homoplasious character combinations and generic delimitation: A case study from the Indo-pacific Arecoid palms (Arecaceae; Arecoideae). Am. J. Bot. 93: Roncal, J., J. Francisco-Ortega, C. B. Asmussen, and C. E. Lewis Molecular phylogenetics of the tribe Geonomeae (Arecaceae) using nuclear DNA sequences of phosphoribulokinase and RNA polymerase II. Syst. Bot. 30: Thomas, M. M., N. C. Garwood, W. J. Baker, S. Henderson, S. J. Russell, D. R. Hodel, and R. M. Bateman Molecular phylogeny of the palm genus Chamaedorea, based on the low-copy nuclear genes PRK and RPB2. Mol. Phylogenet. Evol. 38: Trenel P, Gustafsson MH, Baker WJ, Asmussen-Lange CB, Dransfield J, Borchsenius F Mid-Tertiary dispersal, not vicariance explains Gondwanan distribution pattern in wax palm subfamily (Ceroxyloideae: Arecaceae). Molecular Phylogenetics and Evolution 45:

164 146 Uhl, N. W. and J. Dransfield Genera Palmarum: A Classification of Palms based on the work of Harold E. Moore Jr. Lawrance: L. H. Bailey Hortorium and International Palm Society. Uhl, N. W. and J. Dransfield Genera Palmarum after ten years. Mem. New York Bot. Gard. 83:

165 GENERAL CONCLUSION Areca as a palm genus is interesting not only from its species diversity but also from all aspects of its natural history. Taxonomy of this genus is being revised with acceptance of 41 species, and two subgenera as the result of the discovery of two evolutionary lines in molecular phylogenetics analysis. The morphological characters are still relevant and applicable to apply the species concept in the genus Areca, however those characters are independently each others and evolved several times in this palm genus, thus is needed a depth phylogeny interpretation to understand relationships and changes during the evolution. Dispersal is the main biogeography factor to affect the recent distribution pattern of the genus Areca. Borneo is the centre of origin for palm genus Areca, however the country of origin for the cultivated wide-spread species Areca catechu L the betel nut palm is still unresolved from this study.

166 APPENDIXES Published Papers

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186 BLUMEA 53: Published on 29 October 2008 TWO NEW SPECIES OF LICUALA (ARECACEAE; CORYPHOIDEAE) FROM WESTERN NEW GUINEA Charlie D. Heatubun 1 & Anders S. Barfod 2 Summary Fieldwork in Western New Guinea has revealed a number of new palm species. Here we describe two small-sized species of Licuala with unbranched partial inflorescences. Key words: Arecaceae, Coryphoideae, Licuala, palms, New Guinea. Introduction Recent field work, conducted under the auspices of the Palms of New Guinea Project, has led to the discovery of a number of new palm species in the Indonesian provinces of Papua and West Papua. With an estimated number of 32 species Licuala is the second largest genus in New Guinea only surpassed by the genus Calamus which comprises about 60 species (Barfod et al. 2001, Baker & Dransfield 2006). Many of the New Guinean species of Licuala display character combinations that deviate markedly from species that are not present on the island. This applies particularly to the fruiting morphology. A number of species such as Licuala beccariana and the recently described L. crassiflora Barfod and L. longispadix Banka & Barfod have large-sized fruits with grooved endocarps (Barfod 2000, Banka & Barfod 2004). Here two new species of Licuala are described. They are both small palms with unbranched partial inflorescences. One species has leaves that are divided into narrow segments reminiscent of grass leaves, whereas the other has entire, bifid leaves. The two species are known from relatively few and mostly recent collections. Materials and methods Herbarium specimens from the following herbaria were available to us: AAU, BM, BO, BRI, CAN, K, L, LAE, MAN, MO. All measurements were derived from dried voucher specimens. The flowers were boiled prior to longitudinal sectioning and measured in a stereoscope using a stage micrometer. Indication of colour is based on field observations and label information. The line drawings were rendered from voucher specimens 1) Departemen Biologi, Sekolah Pascasarjana Institut Pertanian Bogor, Darmaga, Bogor 16680, Jawa Barat, Indonesia. (Permanent address: Fakultas Kehutanan Universitas Papua, Amban, Manokwari 98314, Papua Barat, Indonesia); corresponding author: charlie_deheatboen@yahoo.com. 2) Department of Biological Sciences, University of Aarhus, Ny Munkegade, bygn. 1540, DK-8000 Aarhus C, Denmark Nationaal Herbarium Nederland, Leiden University branch

187 430 BLUMEA Vol. 53, No. 2, 2008 d e a c b Fig. 1. Licuala bifida Heatubun & Barfod. a. Habit; b. leaf; c. infructescence; d. detail of infructescence; e. fruit. Scale bar: a = 20 cm, b = 8 cm, c = 4 cm, d = 1 mm, e = 7 mm (a: from photograph taken by Charlie Heatubun; b e: Wally 839, K).

188 C.D. Heatubun & A.S. Barfod: Two new species of Licuala 431 except for the habit drawing on Fig. 1, which was based on a colour photograph from the type locality. Description of species 1. Licuala bifida Heatubun & Barfod, spec. nov. Fig. 1 Palma solitaria, foliis bifidis et inflorescentia partiali spicata. Typus: Wally 839 (holo MAN!; iso BO, K!), Indonesia, West Papua province, Teluk Bintuni regency (previously Manokwari regency), Merdey. Solitary, small to medium-sized palm. Stem to 2 m high, to 3 cm diam., internodes very short. Leaves c. 18 in crown; petiole cm long, 3 6 mm wide at the base to 3 6 mm across towards the apex, adaxial face flattened at the apex and shallowly channelled at the base, abaxial face rounded, greenish yellow in colour, armed; blade small, cm wide, simple-bifid leaves, adaxial light green, abaxial whitish, c. 112 cm long, truncate at apex and 5 10 mm wide, lobe split cm from apex to rachis, rachis c. 40 cm long. Inflorescences 1 or 2 in a single palm, shorter than leaves, cm long, branched to only 2 first-order branches; peduncle cm long; prophyll by mm; peduncular bracts lacking; rachis elongate, c. 16 cm long; rachis bract tubular, flattened, 2-keeled, by 1 mm, covered with brown and black indument, mouth loosely sheathing, splitting on one side, apex pointed; first-order branches close to or slightly away from mouth of rachis bract (c. 12 cm); rachillae 8 12 cm long, 2 3 mm wide. Flowers unknown. Fruits globose, 7 10 mm diam., orange when mature; seed globose 5 8 mm diam.; endocarp smooth; endosperm homogeneous. Distribution This has been reported from a restricted area in the Merdey valley (now Moskona Utara district of Teluk Bintuni regency), in South-Central Bird s Head Peninsula in Western New Guinea. Habitat This palm grows in lowland forest to 500 m asl. Conservation status Data deficient but probably under threat due to its restricted distribution. Vernacular name and uses Not recorded. Note The only New Guinea Licuala recorded so far with bifid leaves. The inflorescence structure is reminiscent of L. graminifolia from which it differs by normally having two partial inflorescences. Flowering material of this species has not yet been collected. Etymology The name refers to the entire, bifid leaves. Additional specimen studied: Indonesia. West Papua: Heatubun s.n. (MAN!), Manokwari, Merdey district (now is Moskona Utara district of Teluk Bintuni regency), forest behind Merdey village. 2. Licuala graminifolia Heatubun & Barfod, spec. nov. Fig. 2 Palma parva solitaria, foliis subtiliter divisis et inflorescentia non ramosa. Ab L. bifida, et ceteris speciebus inflorescentiam spicam praebens, folia in segmentis augustis divisis statim diagnoscenda. Typus: Van Royen & Sleumer 7345 (holo L!; iso K!), Indonesia, West Papua province (Vogelkop Peninsula), Kebar Valley, opposite Andjai, path to Hatai, above Aremi village, 800 m, 20 November 1961.

189 432 BLUMEA Vol. 53, No. 2, 2008 a g b c d Fig. 2. Licuala graminifolia Heatubun & Barfod. a. Leaf; b. stem apex with leaf sheaths and inflorescences; c. inflorescence; d. detail of the partial inflorescence; e. flower; f. longitudinal section of flower; g. staminal ring. Scale bar: a = 6 cm, b = 2.5 cm, c = 2 cm, d = 3 mm, e, f = 1.2 mm, g = 1 mm (all: Van Royen & Sleumer 7345, K). e f

190 C.D. Heatubun & A.S. Barfod: Two new species of Licuala 433 Solitary, small palm. Stem acaulescent to about 1 m high, 1.5 cm diam., internodes short. Leaves c. 9 in crown, sheath disintegrating into a loose mesh of fine fibres; petiole cm long, 2 4 mm wide at the base to mm across towards the apex, adaxially flattened to shallowly channelled towards the base, abaxially rounded, unarmed; lamina fan-shaped in outline, cm wide, divided into 9 11 segments, narrowly wedge-shaped or with slightly curved lateral margins, all segments with 1 3 adaxial ribs, more or less equal in length cm, indentations leading to adaxial ribs 2 3 mm long, those leading to abaxial splits considerably shorter, mid-segment truncate, 6 9 mm wide at apex, basal segment obliquely truncate at apex, usually wider than the mid-segment. Inflorescences 1 or 2 in one palm individual, shorter than leaves, cm long, carrying one spicate, partial inflorescence; prophyll cm long and up to 10 mm wide, loosely fitting, splitting on one side; peduncle as measured to insertion of rachis bract 8 20 cm long; peduncular bracts lacking; rachis bract tubular, 5 9 cm long and up to 7 mm wide, covered with scattered ferruginous ramenta basally, loosely fitting to weakly inflated towards the apex, with 2 or 3 splits, 5 12 mm long; peduncle of partial inflorescence equal to or slightly exceeding rachis in length; rachilla 6 8 cm long, 1 2 mm wide. Flowers solitary, hermaphroditic, bullet-shaped mm long; flowers borne on elevated points, subtending bract c. 1 mm long, bracteole more or less conspicuous at a lateral position, pedicel very short, calyx mm long, hyaline distally, with three mm long pointed lobes; corolla mm long, splitting deeply to c. 5 mm above the base; androecium fused to corolla for mm, staminal ring c. 0.5 mm high; anthers inconspicuously inserted in 2 levels, mm long; ovary mm long, attenuate apically; style mm long; locules in lower half. Fruits globose, 8 10 mm diam., green; seed 6 8 mm diam.; endocarp smooth; endosperm homogeneous. Distribution Known from several collections in Western New Guinea. Habitat This palm is distributed in lowland rain forest ranging up to 800 m asl. It is often found in association with Vatica rassak Blume. Conservation status No immediate concern since it is quite common locally but further research is required. Vernacular name and uses Pupuru (Jamur dialect) is the collective term for species of Licuala in the Lake Jamur area. No specific use of this palm has been recorded. Note Licuala graminifolia is similar to L. bifida in the inflorescence structure; but in contrast to this, the inflorescence is unbranched with only one, spicate, partial inflorescence. The leaf is easily distinguished by the fine segmentation of the blade. Etymology The name refers to the grass-like leaf segments. Additional specimens studied: Indonesia. West Papua: Heatubun 332 (K!, MAN!), Kaimana regency (previously Fakfak regency), Teluk Etna district, km 45 logging road to Lake Jamur; Heatubun 334 (MAN!), idem. Acknowledgements The authors are grateful to the keepers of the herbaria BM, BO, BRI, CAN, K, L, LAE, MAN and MO who sent material of Licuala on loan to AAU. The fieldwork to Lake Jamur and Sorong Selatan was funded by Royal Botanic Gardens, Kew through the Palms of New Guinea Project. The plates were skilfully rendered by Lucy T. Smith for the Palms of New Guinea Project. Dr. William Baker

191 434 BLUMEA Vol. 53, No. 2, 2008 kindly granted us permission to use them in this paper. CDH would like to thank the staff of AAU, K, and L for ground support and hospitality during his visits in June-July References Baker, W. & J. Dransfield Field guide to palms of New Guinea. Royal Botanic Gardens, Kew. Banka, R. & A.S. Barfod A new spectacular species of Licuala (Arecaceae, Coryphoideae) from New Guinea. Kew Bulletin 59: Barfod, A.S A new species of Licuala from New Guinea. Palms 44: Barfod, A.S., R. Banka & J.L. Dowe Field guide to palms in Papua New Guinea. AAU reports 40: 1 77.

192 PALMS Heatubun: New Areca Vol. 52(4) 2008 A New Areca from Western New Guinea CHARLIE DANNY HEATUBUN Departemen Biologi, Sekolah Pascasarjana Institut Pertanian Bogor, Bogor, Jawa Barat, Indonesia (Permanent address: Fakultas Kehutanan Universitas Papua, Amban, Manokwari 98314, Papua Barat, Indonesia) charlie_deheatboen@yahoo.com 1. Leaves of Areca mandacanii showing irregular leaflet arrangement; Martinus Iwanggin provides scale. A new species of Areca related to A. catechu has been discovered in swampy areas in western New Guinea. This remarkable palm has future potential not only as a nut-crop species, but also for its ornamental horticulture value. As part of the Palms of New Guinea project (Baker 2000, 2002), the author conducted a field trip to the area of Teminabuan (south of Sorong) in the Bird s Head Peninsula of Western New Guinea in During the field trip, many interesting palms were encountered and collected, such as the newly described genus Dransfieldia (Baker et al. 2006) and the 198 PALMS 52(4):

193 PALMS Heatubun: New Areca Vol. 52(4) 2008 rediscovery of Hydriastele flabellata (Becc.) W. J. Baker & Loo (Heatubun 2005) this latter beautiful undergrowth palm was originally collected by Beccari almost 132 years ago and the type locality has already vanished within the town center of Sorong. Among the other species found was a new species of Areca described in detail here. Areca mandacanii Heatubun sp. nov., habitu A. catechu L. similis, sed plica foliolorum solitaria, lineari, irregulariter disposita incisura acuminata, fructibus grandibus, ellipsoideis, seminibus globosis, differt. Typus: Indonesia, West Papua Province, Sorong Selatan Regency, Teminabuan District, Sayal Village, Maampou forest, Feb. 2003, Heatubun 423 (Holotypus BO!; isotypi K!, MAN!). Solitary, moderate pleonanthic, tree palm. Stem up to 15 m tall, 8 10 cm diam.; internodes cm long, dark green, shiny, nodal scars conspicuous, white. Leaves 8 in crown, pinnate, appearing plumose, cm long (including petiole); sheath tubular, ca. 92 cm long, smooth, light green; crownshaft well defined, up to 152 cm long and up to 15 cm diam.; petiole short to 6 cm long, channelled adaxially, rounded abaxially; rachis somewhat arching, with adaxial longitudinal ridge, rounded abaxially; blade with irregularly arranged leaflets, divided into groups, ca. 60 leaflets on each side, held in several planes, near petiole small and gradually becoming larger, terminal leaflets regularly arranged (ca. 13 or 14 leaflets); leaflets somewhat arching, single-fold, linear, cm long, 2 cm wide, tip acuminate, notched, split to 3 cm long and pointed in a few leaflets near the petiole, papery, green adaxially and light green abaxially. Inflorescence infrafoliar, ca. 60 cm long at anthesis, protandrous, branching to 2 or 3 orders; peduncle 5 cm long, green with numerous minute red-brown dots; prophyll 64 6 cm, borne about 1/3 way up the peduncle, lanceolate, 2-keeled, papery, cream to light brown, entirely enclosing the inflorescence, then splitting longitudinally and falling before staminate anthesis; rachis green to whitish green; rachis bracts not persistent; rachillae numerous, 37 cm long, covered by light brown to chocolate-brown thick indument, highly contrasting with the rachis, first branching rachillae ca. 50 cm long, elongate. Staminate flowers small, triangular, mm, asymmetric; sepals 3, low, about 2 1 mm; petals 3, strongly keeled, 4.5 mm long, ca mm wide at basal; pistillode mm, trifid, dark brown; stamens 6, Reproductive organs of Areca mandacanii: young inflorescence with staminate and pistillate flowers (left); young fruits and rachilla covered with brown indumentum (right). 199

194 PALMS Heatubun: New Areca Vol. 52(4) 2008 B F E G C H D I 3. Areca mandacanii: A. leaf; B. apical portion of leaf; C. mid portion of leaf; D. infructescence with immature fruits; E. portion rachillae showing position of staminate and pistilate flowers; F. staminate flowers on rachilae; G. staminate flower in section; H. pistillate flower whole and in section; I. fruit in section. Scale bar A = 62.5 cm; B, C, D = 4 cm; E, I = 3 cm; F= 7 mm; G =1.6 mm; H = 1 cm. A from photograph, B E & H from Heatubun 423, F, G & I from Heatubun 413. Drawn by Lucy T. Smith. A mm long, sagittate; anthers 1.5 mm long, arrow head shape, creamy, longer than the filaments, twisted basally; filaments 0.5 mm long, dark brown. Pistillate flowers larger than the staminate, triangular, mm diam., sitting on the branch-nodes; sepals 3, 200

195 PALMS Heatubun: New Areca Vol. 52(4) Fruits of Areca mandacanii: immature fruits (far left) and mature fruits (near left). imbricate, triangular, mm long, mm wide, 1 3 mm thick, asymmetrical, thicker at base, cream to light green; petals 3, imbricate, triangular, mm, 0.5 mm thick, cream; staminodes ovate, 10 mm high, 9 mm wide, pointed, brown coloured at the tip, fleshy. Fruits ellipsoidal, mm, with conspicuous woody discoid depression at apex, shallowly concave, cm diam., stigmatic remains persistent in centre of depression, perianth persistent; epicarp smooth, shiny, ca. 0.5 mm thick, dark green when young, turning to golden yellow or light orange when mature; mesocarp fibrous, ca. 10 mm thick, but much thicker at the base of fruit (below the seed), where ca. 20 mm thick; endocarp very thin, adhering closely to seed; seed subglobose, ca mm; endosperm deeply ruminate; embryo basal. Distribution. Bird s Head Peninsula of Western New Guinea. Specimens Examined. INDONESIA: West Papua Province, Sorong Selatan Regency, Teminabuan District, Sayal Village, 5. Fruit of Areca mandacanii (left) compared with fruits of two different varieties of A. catechu (center and right). Maampouw forest, 10 m asl., 21 Feb 2003, Heatubun et al. 423 (Holotype BO!, isotypes K!, MAN!); Heatubun et al. 413 (K! (spirit collection), MAN!); Heatubun et al. 424 (MAN); West Papua province, exact locality unknown, Maturbongs s.n (K!, MAN). CULTIVATED: Indonesia, West Papua Province, Manokwari, Reremi, 75 m asl., 10 Apr 2008, Heatubun & Iwanggin 902 (MAN!). Habitat. This palm grows in forest transitional between swampy areas and lowland rain forest, where the soils are temporarily inundated by water. Other palms that occupy the same habitat are Areca macrocalyx, Calyptrocalyx sp., Drymophloeus litigiosus, Licuala beccariana and Sommieria leucophylla. Local Name. Nggafa (Sayal dialect) and/or Pinang Hutan (Indonesian for forest betel nut). Uses. The stem of this palm is used for flooring by local people for their houses and the seed is chewed as a substitute for betel nut (A. catechu L.). Conservation Status. Data Deficient. This species has been reported from a very restricted area in the Bird s Head Peninsula of Western New Guinea. Further population studies would be needed to assess the conservation status of this palm. Notes. Areca mandacanii is similar to A. catechu in habit and inflorescence structure, but can immediately be distinguished by the leaf with single-fold leaflets, irregularly arranged in groups in several ranks, with tips acuminate and notched rather than the broad leaflets with several folds, arranged in a single plane and more or less regularly arranged, characteristic of A. catechu. The structure of the inflorescence and flowers indicate that A. mandacanii belongs to subgenus Areca (subgenus Blumeoareca) section Oeotheanthe (sensu Furtado)) together with A. catechu. In this infrageneric classification 201

196 PALMS Heatubun: New Areca Vol. 52(4) 2008 (Furtado 1933), subgenus Blumeoareca (correctly subgenus Areca) is characterized by the inflorescence branched to two or three orders, with many rachillae, staminate flowers unilateral or alternate-distichous, sepals free or sometimes united, stamens six, the pistillate flowers seated on the axis of the basal branches, subdistichous or subopposite or spiraled on the rachilla. Despite the variation and complexity of morphological characters in New Guinean Areca species, the new species is quite different from the widespread A. macrocalyx Zipp. ex Blume and its relatives especially in inflorescence form, which in the A. macrocalyx complex is congested. The attractive leaves and fruits make this species instantly identifiable from other New Guinean Areca. The irregular arrangement of leaflets (several planes) is reminiscent of Woodyetia bifurcata, the Australian foxtail palm. The fruit size, aroma and colour (yellowish golden) also make this palm of great interest to palm growers. This palm is named for Mr. Dominggus Mandacan, the regent of Manokwari and the chief of the Arfak tribes, who has paid so much attention to the development of scientific activities in the Vogelkop (Bird s Head Peninsula) area, including botanical research, and who also supported our expedition in many ways. Acknowledgments The author thanks Marthinus Iwanggin, Arius Sesa, Tobias Paiki, the peoples of Sayal village and the keeper and staff of AAU, BO, K, L, and MAN. Drs. John Dransfield and William J. Baker from K provided useful comments on the manuscript including help with the Latin diagnosis. The Bakers, Dransfields, Barfods and Veldkamps are much appreciated for their hospitality and ground support during the author s visits to AAU, K and L from June July, The plate was prepared by Lucy T. Smith with funding by the Pacific Biological Foundation. Fieldwork in 2003 was funded by British American Tobacco (BAT) to Royal Botanic Gardens Kew through The Palms of New Guinea Project. This paper is part of the author PhD project on genus Areca at Institut Pertanian Bogor, Indonesia, under the auspices of the Royal Botanic Gardens Kew, UK. LITERATURE CITED BAKER, W.J The Palms of New Guinea Project. Palms 44: 160, 165. BAKER, W.J The Palms of New Guinea Project. Fl. Males. Bull. 13: BAKER, W.J., S. ZONA, C.D. HEATUBUN, C.E. LEWIS, R.A. MATURBONGS AND M.V. NORUP Dransfieldia (Arecaceae) A new palm genus from western New Guinea. Syst. Bot. 31: DRANSFIELD, J The genus Areca (Palmae: Arecoideae) in Borneo. Kew Bull. 39: FURTADO, C.X The limits of the genus Areca L. and its sections. Feddes Repertorium 33: GOVAERTS, R. & DRANSFIELD, J World Palms Checklist. Royal Botanic Gardens, Kew. HEATUBUN, C.D The rediscovery of Beccari s Nenggela flabellata in Irian Jaya Barat. Folia Malaysiana 6:

197 KEW BULLETIN VOL. 64: (2009) A monograph of Cyrtostachys (Arecaceae) Charlie D. Heatubun 1,4, William J. Baker 2, Johanis P. Mogea 3, Madeline M. Harley 2, Sri S. Tjitrosoedirdjo 4 & John Dransfield 2 Summary. Cyrtostachys Blume (Areceae: Arecaceae) is treated in this study as a genus of tree palms with a disjunct distribution pattern across Malesia and consisting of seven species. Three species are newly recognised (C. bakeri Heatubun, C. barbata Heatubun and C. excelsa Heatubun). Five previously accepted species (C. brassii Burret, C. kisu Becc., C. microcarpa Burret, C. peekeliana Becc. and C. phanerolepis Burret) are reduced to synonymy with C. loriae Becc. and one species (C. compsoclada Burret) is removed to Heterospathe as Heterospathe compsoclada (Burret) Heatubun, while C. ledermanniana Becc. is considered as a doubtful taxon. A determination key is presented and detailed descriptions provided for all taxa. A phylogenetic analysis of all species in the genus was performed based on morphological data. Despite the poorly resolved tree topologies, Cyrtostachys is resolved as monophyletic, with C. glauca H. E. Moore as sister to all other species, and the west Malesian species C. renda Blume probably representing a dispersal from within a Papuasian clade into the Sunda shelf. Natural history observations, including uses and conservation status are also presented in this monograph. Key Words. Arecaceae, Areceae, Arecoideae, Cyrtostachys, Malesia, morphology, taxonomy. Introduction Cyrtostachys Blume is a genus of palms described almost 170 years ago by Blume (1838), based on the remarkable species, C. renda Blume, the sealing wax palm, a species well-known for its brilliant red leaf sheaths. The genus is highly desirable from a horticultural point of view and it is also used locally for a variety of purposes; the extraordinary disjunct distribution poses interesting biogeographic questions that can only be addressed when its interspecific relationships are better understood. Cyrtostachys has a disjunct distribution pattern (Map 1), with one species (C. renda) occurring to the west of Wallace s line in south Thailand, Malay Peninsula, Sumatra and Borneo, and the remaining six species on the island of New Guinea to the Solomon Islands (Moore 1973; Dransfield 1981, 1987; Baker et al. 1998). Cyrtostachys belongs to subfamily Arecoideae and tribe Areceae based on its pinnate, reduplicate leaf, monoecy with flowers borne in triads of a central pistillate and two lateral staminate flowers. It can be distinguished from other genera of arecoid palm by the strongly divaricate inflorescence branches, flowers borne in pits along the rachillae, connate petals and more than six stamens in staminate flowers (Uhl & Dransfield 1987). Dransfield & Uhl (1986) placed Cyrtostachys in its own subtribe, Cyrtostachydinae, as an isolated genus with no clear relatives. Moreover, in most studies (Asmussen & Chase 2001; Lewis& Doyle 2002; Hahn 2002; Essig& Litten 2004) no clear affinities with other palm genera have been found, apart from establishing that Cyrtostachys belongs to the large group of Indo-Pacific pseudomonomerous arecoid palm genera tribe Areceae (sensu Dransfield et al. 2005, 2008). Recently, a phylogenetic analysis of all genera of Areceae based on low-copy nuclear DNA provided moderate support for a relationship between Cyrtostachys and Clinostigma (Norup et al. 2006). However, further evidence is required to substantiate this position. Thus, in the new phylogenetic classification of the palm family (Arecaceae), Cyrtostachys remainsasanunplacedmember of Areceae (Dransfield et al. 2005, 2008). Although the genus Cyrtostachys is well circumscribed, the species, with the exception of C. renda, are as yet Accepted for publication July Faculty of Forestry, Universitas Papua, Jl. Gunung Salju, Amban, Manokwari 98314( Papua Barat, Indonesia. charlie_deheatboen@yahoo.com 2 Herbarium, Royal Botanic Gardens, Kew, Richmond( Surrey, TW9 3AB, UK. 3 Herbarium Bogoriense, Puslitbang Biologi LIPI, Jl. Ir. H. Juanda No. 22, Bogor( Jawa Barat, Indonesia. 4 Biology Department, Sekolah Pascasarjana Institut Pertanian Bogor, Baranangsiang, Bogor( 16680, Jawa Barat( Indonesia. The Board of Trustees of the Royal Botanic Gardens, Kew, 2009

198 68 KEW BULLETIN VOL. 64(1) Map 1. Distribution map of the genus Cyrtostachys Blume. rather poorly known. Most of these Papuasian species were described based on single specimens (Beccari 1905, 1914b, 1923; Burret1936, 1937, 1939; Moore1966). No monographic account has ever been published (Uhl & Dransfield 1987; Barfod et al. 2001). Infraspecific taxa have been described from cultivation (Tucker 1992; Ellison & Ellison 2001; Waddel 2002), and the number of these could increase in parallel to horticultural demand. The genus is thus in need of reassessment. In the 40 years since the last species, Cyrtostachys glauca H. E. Moore, was published (Moore 1966) the number of herbarium specimens of Cyrtostachys has increased greatly, particularly due to activities related to the Palms of New Guinea Project (Baker 2000, 2002); the palm collecting density in the western part of New Guinea, in particular, has increased dramatically. Now we have sufficient materials in the herbarium, field experience, and access to new data (anatomy and pollen morphology) to study the genus and appreciate the wide range of variation within it. Furthermore, there has also been success in tracing isotypes of Burret s species, the holotypes of which were destroyed during the Second World War in Berlin. This monograph aims to provide a modern taxonomic treatment of Cyrtostachys and to compile and reevaluate published information on its systematics, natural history, uses and conservation status. Taxonomic History The genus Cyrtostachys was described for the first time by Carl Ludwig von Blume in 1838 based on C. renda Blume (Blume 1838). Blume had intended to publish the genus in Rumphia, but publication of his great work was delayed until 1843 (Stafleu & Cowan 1976). In notes with the protologue, he wrote: in Rumphia l.c. behind the genus name and he provided only a brief description of the genus and species while in Rumphia he provided a full description of the genus and species accompanied by a plate with details of C. renda from a specimen collected by Korthals near Indrapura in Sumatra. The word Cyrtostachys comes from the Greek word cyrtos (arched) and stachys (a spike) in reference to the recurved inflorescence (Whitmore 1973; Jones 1995). Odoardo Beccari (1905) published the second species of the genus, Cyrtostachys loriae Becc., based on a Hartmann collection from the Owen Stanley Range, Papua New Guinea, and this was followed by C. kisu Becc. (Beccari 1914a), based on a specimen from the Solomon Islands made by Guppy and C. peekeliana Becc. (Beccari 1914b) from New Ireland (Papua New Guinea) based on a collection made by Peekel. In 1923 another species was published by Beccari, C. ledermanniana Becc. to accommodate Ledermann s specimen from Ettapenberg, Papua New Guinea (Beccari 1923). Max Burret published C. brassii Burret (Burret 1935) based on a specimen collected by L. J. Brass from Kubuna, Central Division, Papua New Guinea. Two more species of Cyrtostachys (C. phanerolepis Burret and C. compsoclada Burret) were published by Burret (1936). The last of Burret s species, C. microcarpa Burret, was published in 1939 and based on a L. J. Brass specimen from Lake Daviambu, Fly River, Papua New Guinea (Burret 1939). Harold E. Moore Jr discovered and published C. glauca H. E. Moore from Morobe district, Papua New Guinea (Moore 1966). The Board of Trustees of the Royal Botanic Gardens, Kew, 2009

199 A MONOGRAPH OF CYRTOSTACHYS (ARECACEAE) 69 Materials and Methods Morphology Morphology and distribution of the species were studied based on specimens (dried and spirit-preserved materials) deposited at A, AAU, BH, BO, BRI, FI, K, KEP, L, LAE, MAN, PNH, SAN, and SING (Holmgren et al. 1990). Many new specimens have been collected recently by the authors. In particular, new specimens were collected in western New Guinea (Indonesian province of Papua and Irian Jaya Barat) by the first author in Specimens were made in the field using standard preparation guidelines proposed by Dransfield (1986). Collections of leaf samples, inflorescence (rachillae with flowers in bud and/or at anthesis), and mature fruits were preserved in 70% alcohol for further studies. Measurements were taken from spirit-preserved material and dried herbarium specimens and from cultivated individuals. Floral parts were measured from spirit-preserved material or dried specimens rehydrated by boiling. Pollen Morphology Pollen material was obtained from dried herbarium specimens and spirit-preserved material. Seventeen samples were chosen to represent the wide range of taxa in the genus Cyrtostachys. Details of pollen preparation and methods for herbarium material follow Harley (1990, 1996) and terminology follows Punt et al. (1994), Harley (1999) and Harley & Baker (2001). Microphotographs were taken using a Nikon D100 Optiphot light microscope (LM) and a Hitachi S-2400 scanning electron microscope (SEM). Phylogenetic Analysis A preliminary phylogenetic analysis of all species of Cyrtostachys was conducted. In recent molecular phylogenetic studies, Cyrtostachys is resolved in a position sister to a clade including Nenga and Areca (Asmussen & Chase 2001), or Areca (Hahn 2002), or Iguanura (Lewis & Doyle 2002). Based on histological studies of the fruit (Essig & Litten 2004) the genus has some similarities with Hydriastele. Therefore, in our analysis we have included Areca catechu L., Hydriastele costata F. M. Bailey, Pinanga rumphiana (Mart.) J. Dransf. & Govaerts and Rhopaloblaste ledermanniana Becc. as outgroups. Seventeen morphological and palynological characters were included in the data matrix (Table 1), which was constructed in MacClade 4.0 (Maddison & Maddison 2000). A parsimony analysis was performed using PAUP 4.0b10 (Swofford 1998). All characters were treated as unordered and were equally weighted. Support for the clades was calculated by conducting a 10,000 replicate bootstrap analysis using the branch and bound algorithm. Character List General Habit Characters 1. Habit: solitary (0), clustering (1). This character has been discussed in detail by Holttum (1955), Dransfield (1978), Uhl & Dransfield (1987), Tomlinson (1990) and Fisher & Maidman (1999) not only for descriptive terminology, but also for developmental physiology and its value for systematics. 2. Crown shape: spherical (0), hemispherical (1), shuttle-cock shaped (2). The crown shape is an obvious character and very helpful in recognising species in the field. The crown itself is composed of many leaves and forms a distinctive shape in outline. A spherical shape is formed when leaves spread out in all directions, forming a circular outline when viewed from a distance. A hemispherical shape is formed when the lowest leaves make angles 15 above or below the horizontal or leaf inclination is ± 30, taking into account the angles of the lowest leaves either side of the horizontal. The shuttle-cock crown arises when the inclination of the lowest leaves is more than 30 above horizontal. 3. Leaflet arrangement: regular (0), irregular (1). 4. Leaflet thickness: papery (0), leathery (1). 5. Ramenta on abaxial surface of leaflet: absent (0), present (1). Table 1. Data matrix for phylogenetic analysis of morphological and palynological data of species of Cyrtostachys and outgroups. Inapplicable characters ( ) are indicated. For polymorphic characters, both states are shown. Characters Terminal taxa Cyrtostachys bakeri Cyrtostachys barbata Cyrtostachys elegans Cyrtostachys excelsa Cyrtostachys glauca Cyrtostachys loriae Cyrtostachys renda = Areca catechu Hydriastele costata Pinanga rumphiana Rhopaloblaste ledermanniana The Board of Trustees of the Royal Botanic Gardens, Kew, 2009

200 70 KEW BULLETIN VOL. 64(1) 6. Arrangement of ramenta: separate (0), continuous/ connected (1). 7. Leaf sheath surface: not glaucous (0), glaucous (1). 8. Crownshaft colour: green (0), red (1). General Reproductive Characters 9. Inflorescence basal branches: not strongly divaricate (0), strongly divaricate (1). The strongly divaricate inflorescence is defined based on the angle formed between the first primary branch and the rachis (main axis) being more than Inflorescence branching orders: spicate (0), 1 branching order (1), 2 branching orders (2), 3 branching orders (3), 4 branching orders (4). 11. Rachilla pits: superficial pits (0), deep pits (1). 12. Pistillate calyx persistent on rachilla: absent (0), present (1). 13. Petals in male flowers: free (0), connate (1). 14. Filaments in male flowers: free (0), connate (1). Pollen Characters 15. Pollen tectum surface: reticulate (0), perforate (1), rugulate (2), verrucate-gemmate (3). 16. Trichotomosulcate grains: absent (0), present (1). Seed Character 17. Seed endosperm: homogenous (0), ruminate (1). Conservation Status The conservation status of each species of the genus Cyrtostachys was assessed based on the IUCN red list categories and criteria version 3.1 (IUCN 2001). Results and Discussion Species Delimitation A morphological species concept (Davies & Heywood 1963; McDade 1995; Dransfield 1999) is used in this monograph. Like Barrow (1998), we also looked at the constancy of morphological character states both within and between populations and recognised as species only those smallest units which can be diagnosed by constant character states. Seven taxa of Cyrtostachys are recognised in this monograph (Table 2); three species are new (C. bakeri Heatubun, C. excelsa Heatubun and C. barbata Heatubun); five taxa (C. brassii Burret, C. kisu Becc., C. microcarpa Burret, C. peekeliana Becc. and C. phanerolepis Burret) are treated as new synonyms of C. loriae Becc. and one taxon (C. compsoclada Burret) is incorrectly described in Cyrtostachys and is transferred to Heterospathe. One species, C. ledermanniana Becc., is regarded as an imperfectly known taxon. For full discussion see Taxonomic Treatment. Morphology Habit Of the seven species of Cyrtostachys, three species are solitary (C. excelsa, C. loriae and C. barbata) and four others are clustering (C. bakeri, C. elegans, C. glauca, and C. renda). However, although generally solitary, some populations of C. loriae in Indonesia, Province of Papua, are clustering. Furthermore, C. renda also displays variation in habit character, especially in individuals in cultivation. Table 2. Comparison of previously recognised taxa in Cyrtostachys with taxa recognised in this monograph. Synonyms names are shown in italics. Species of Cyrtostachys accepted in World checklist (Govaerts & Dransfield 2005) Species of Cyrtostachys accepted in this monograph Cyrtostachys bakeri Heatubun sp. nov. Cyrtostachys barbata Heatubun sp. nov. Cyrtostachys elegans Burret Cyrtostachys elegans Burret Cyrtostachys excelsa Heatubun sp. nov. Cyrtostachys glauca H. E. Moore Cyrtostachys glauca H. E. Moore Cyrtostachys loriae Becc. Cyrtostachys loriae Becc. Cyrtostachys brassii Burret Cyrtostachys brassii Burret synon. nov. Cyrtostachys kisu Becc. Cyrtostachys kisu Becc. synon. nov. Cyrtostachys microcarpa Burret Cyrtostachys microcarpa Burret synon. nov. Cyrtostachys peekeliana Becc. Cyrtostachys peekeliana Becc. synon. nov. Cyrtostachys phanerolepis Burret Cyrtostachys phanerolepis Burret synon. nov. Cyrtostachys renda Blume Cyrtostachys renda Blume Areca erythropoda Miq. Areca erythropoda Miq. Cyrtostachys lakka Becc. Cyrtostachys lakka Becc. Cyrtostachys lakka var. singaporensis Becc. Cyrtostachys ledermanniana Becc. Cyrtostachys compsoclada Burret Imperfectly known (doubtful) taxon: Cyrtostachys ledermanniana Becc. Excluded Taxon: Heterospathe compsoclada (Burret) Heatubun comb. nov. The Board of Trustees of the Royal Botanic Gardens, Kew, 2009

201 A MONOGRAPH OF CYRTOSTACHYS (ARECACEAE) 71 Species of Cyrtostachys are moderate to robust palms with stems from 10 to 30 m high, they are entirely unarmed, pleonanthic and monoecious palms and grow in various conditions from tropical lowland rainforest to montane forest (900 m a.s.l.); sometimes they emerge above the forest canopy. The palm architecture of this genus conforms to Corner s model for solitary species and Tomlinson s model for clustering species (Hallé et al. 1976; Dransfield 1978; Tomlinson 1990; Uhl & Dransfield 1987). Stem The stem of Cyrtostachys varies from moderate ( cm in diameter) to robust (up to 30 or 40 cm in diameter). Leaf scars are prominent and demarcate conspicuous internodes that are mostly green near stem tip (yellowish green, yellow, orange to red purple with stripes in C. renda and its cultivars) and become brown to greyish near the base. The stem in cross section displays an outer cortex consisting of a thin layer of about 4 mm and an inner cortex with vascular bundles. As in Actinorhytis, Ptychococcus or Rhopaloblaste (Banka & Baker 2004) the outer stem is usually very hard while the inner part is softer with sparser fibres. However, wood of Cyrtostachys has a lower number of black fibres and is only used for certain construction purposes. It cannot be used for traditional weapons such as bows, arrow-heads, and spears (Heatubun et al. in prep.). Leaves The crown consists of leaves ranging from about 6 7 per stem in Cyrtostachys glauca to 14 in C. loriae. There are three types of crown outline in the genus: spherical (C. barbata, C. glauca and C. loriae), hemispherical (C. elegans) and shuttle-cock shaped (C. bakeri, C. excelsa and C. renda). According to Tomlinson (1990) the shuttle-cock or feather duster crown shape results from relatively few leaves in the crown, with the leaf axis held at a high angle and abscission of the leaf taking place as a unit before the axis droops. In contrast, C. glauca has the lowest number of leaves in the genus and has a spherical crown shape. All species of Cyrtostachys have regularly pinnate leaves, except for C. excelsa, in which leaflets are irregularly arranged in 1s or 2s. The inclination of leaflets, whether porrect or pendulous, is associated with the crown outline. Pendulous leaflets occur in spherical and hemispherical crowns and the porrect leaflets in shuttle-cock shaped crowns, except in C. bakeri which has a shuttlecock shaped crown with pendulous leaflets (Fig. 5A), and C. excelsa which has a shuttle-cock shaped crown and slightly curved-porrect leaflets. The thickness and coloration of leaflets when dried are leathery and discolorous in general in Cyrtostachys, except in C. glauca that has papery leaflets, and C. excelsa that has concolorous leaflets. The petiole is almost missing to short (to 10 cm long) in a few species, and elongate to 50 cm in C. renda and more than 50 cm (to 100 cm) in C. glauca. The long tubular leaf sheaths form a well-defined crownshaft. The crownshaft is glaucous only in Cyrtostachys glauca. In C. renda the crownshaft is bright red to orange, and in other species is green. The leaf sheath is thick and sometimes has disintegrating fibres at the margins just below the petiole. The leaf anatomy of Cyrtostachys is discussed briefly by Tomlinson (1961) based on his observation on C. lakka (= C. renda), and he noted that Cyrtostachys is readily distinguished from other Arecoid palms in having markedly sinuous epidermal cell-walls and having the hypodermal layer made up mostly of fibres instead of colourless cells. Indumentum Indumentum in Cyrtostachys occurs on leaf sheaths, leaves and inflorescences. A number of types of indumentum occur on the leaves. Fine purple scales, filamentous brown scales and brown to black thick scales occur on leaf sheaths; thin to thick, caducous, white, scale-like indumentum, with scattered, filamentous, brown to dark brown scales occurs on the petiole; dense white or brown scales occur on the leaf rachis, and inflexed, filamentous brown ramenta occur scattered and/or continuous along the abaxial surface of the mid-veins of the leaflets. The inflorescences carry less indumentum than the leaves, although brown to rusty brown woolly indumentum is present on the rachillae. The presence of beard-like filamentous brown ramenta along the mid-vein on the abaxial leaflet surface is a crucial character in recognising C. barbata from other species (Fig. 6C). Inflorescence All Cyrtostachys species have infrafoliar inflorescences and are branched from two to four orders. The 2- keeled prophyll encloses the inflorescence until leaf fall, borne just above the winged base of the peduncle and is tubular and lanceolate with winged margins, splitting and soon caducous. The single peduncular bract is similar to the prophyll and it too is soon caducous. Inflorescences are strongly divaricate (Fig. 1) with most basal branches rather diffuse and spreading. The peduncle is usually very short, oval in cross section. First order branches are robust, almost as thick as the main axis (Barfod et al. 2001), spreading, and with a short bare portion at the base, then branching to produce diverging rachillae or second-order branches; second order branches, when not bearing flowers, also have a short bare portion and then branch to produce rachillae. The rachillae are elongate, cylindrical, rather robust, glabrous, papillose, minutely roughened or indumentose, often brightly coloured and expanding long before anthesis. The The Board of Trustees of the Royal Botanic Gardens, Kew, 2009

202 72 KEW BULLETIN VOL. 64(1) Fig. 1. Cyrtostachys loriae Becc. in Japen Island (Heatubun et al. 546) strongly divaricate inflorescence with young fruits. PHOTO: CHARLIE D. HEATUBUN. The Board of Trustees of the Royal Botanic Gardens, Kew, 2009

203 A MONOGRAPH OF CYRTOSTACHYS (ARECACEAE) 73 rachilla bracts are low, triangular, spirally arranged, rather crowded, each partially enclosing a shallow pit bearing a triad of flowers. The numbers of orders of branches of the inflorescence and the nature of pits have important taxonomic value to define species in Cyrtostachys. An inflorescence branched to two orders is only found in C. renda. Four orders of branching are found in C. elegans and C. excelsa, while the other species have inflorescences branched to three orders. There are variations in the abundance and depth of pits from species to species, from superficial pits in C. glauca (Figs. 7D E) to deep pits in C. loriae (Figs. 8D E). Inflorescences in Cyrtostachys change in colour as they develop, from cream-coloured at the beginning after the prophyll has fallen, followed by greenish to green at flowering stage (anthesis of male flowers) to fruiting (young fruits), then yellowish-brown to dark brown at ripening (ripe fruits). All species follows this sequence, except C. excelsa and C. renda. In C. excelsa, pink to reddish-purple colour occurs after the prophyll has fallen until the phase before anthesis of male flowers, while C. renda has reddish-purple to blackish rachillae at fruiting (young fruits) to ripening stage (ripe fruits). Flowers In common with almost all Arecoideae, the flowers are arranged in triads of a central pistillate flower and two lateral staminate flowers, borne in pits along the entire length of the rachilla (e.g. Fig. 7D). The staminate flowers are cream-coloured to greenish-yellow in bud and white to cream-coloured at anthesis. The staminate flowers are about the same size as or slightly smaller than the pistillate. At the beginning of rachilla emergence the pistillate flowers are not visible, being obscured by the staminate buds. Once the staminate flowers have fallen, the pistillate flowers become visible and enlarge. Staminate flowers have three, distinct, imbricate, broad, strongly keeled sepals with minutely toothed margins; the petals are about twice as long as the sepals, united at the base to about one third their length distally with triangular, valvate tips. Stamens number 8 to 15, with awl-shaped filamentsthatare connate basally and inflexed apically in bud, the anthers being dorsifixed and latrorse. The connate petals and stamens are distinguishing key characters for separating Cyrtostachys from other genera in the Indo-pacific pseudomonomerous Arecoid palm clade. Pistillate flowers are about the same size as, or slightly larger than, the staminate flowers. Sepals are three, distinct, rounded, imbricate, the margins minutely toothed; petals are also three, distinct, imbricate proximally, asymmetrical, rounded with short triangular valvate tips. There is a very low membranous staminodal ring, bearing short truncate or irregular triangular teeth. The gynoecium is unilocular, ellipsoidal with three short recurved stigmas (e.g. Fig. 8H, J). Fruits and Seeds The fruits are small (to 16 6 mm), varying in shape from ellipsoid, ovoid to obovoid with a beak to sickleshaped, green to black when ripe, and one-seeded. The seeds are small (to mm), globose to ellipsoidal, apically attached, with an orbicular hilum, and homogeneous endosperm with basal embryo (e.g. Figs. 7K, 8K). There is no important taxonomic character in the fruits and seeds useful to separate species. The shape and size of fruits and seeds vary greatly within individuals and populations. Fruit histology of palms has given major insights into palm systematics (Chapin et. al. 2001; Essig 1977, 1982, 1999; Essig & Hernandes 2002; Essig & Young 1979, 1985; Essig et al. 1999, 2001). Variations of fruit pericarp among the representative species of Cyrtostachys have recently been studied in detail by Fred Essig and co-workers. The genus can be characterised by a combination of a papillate epidermis, a heavy layer of tanniniferous/pigmented cells below the epidermis, a system of vascular bundles with thick fibrous sheaths with purely fibrous bundles frequently above and below, the absence of brachysclereids, and a very thin sclerified locular epidermis (Essig & Litten 2004). The authors found that one species, C. kisu (= C. loriae), appears to be the most distinctive, with relatively few, large bundles and tanniniferous tissue throughout the pericarp. Pollen Morphology Pollen morphology was observed using both light microscopy (LM) and scanning electron microscopy (SEM) (Figs. 2 3). Using SEM the following details were obtained: the tectum surface (topology) is perforate, rugulate or in some species, verrucate and/or gemmate. Sometimes, in LM or SEM preparations, a few still intact tetrahedral post meiotic tetrads were observed. Pollen morphology for individual species is included in the species descriptions. Results from pollen morphological data provide limited variation in characters to aid species delimitation, especially in the Papuasian taxa. This reflects the findings of Harley (1999), who conducted a general survey of pollen morphology throughout the genera of the palm family and found that pollen data are not often useful in helping to support delimitation at the species level. However, the data give a clear distinction between the west Malesian Cyrtostachys renda and the Papuasian taxa, in which the topology of the tectum surface of C. renda is microfossulate-rugulate (Fig. 3J M), in contrast to the verrucate-gemmate tectum topology in the Papuasian The Board of Trustees of the Royal Botanic Gardens, Kew, 2009

204 74 KEW BULLETIN VOL. 64(1) Fig. 2. Some of the pollen morphological diversity in Cyrtostachys. A C C. bakeri (from Baker 1138): A whole grain, proximal face; B whole grain, distal face; C group of grains to show general appearance of pollen. D, E C. barbata (from Brass 13707): D whole grain, proximal face; E two rather collapsed grains, apparently conjoined. F H C. elegans (from Heatubun 341): F whole grain, proximal face; G whole grain, distal face showing trichotomosulcate aperture; H close up of verrucate surface, proximal face. J L C. excelsa (from Heatubun 330): J three grains adhered, showing proximal faces; K whole grain, proximal face; L close up of verrucate surface, proximal face. Scale bar: 5 m. The Board of Trustees of the Royal Botanic Gardens, Kew, 2009

205 A MONOGRAPH OF CYRTOSTACHYS (ARECACEAE) 75 Fig. 3. Some of the pollen morphological diversity in Cyrtostachys (continued). A C C. glauca (from Barfod 454): A whole grain, proximal face; B close up of verrucate surface, proximal face; C group of grains to show general appearance of pollen. D H C. loriae (D, G from Barfod 463; E, F, H from Whitmore 3945): D group of grains; E whole grain, distal face showing monosulcate aperture; F whole grain, distal face showing trichotomosulcate aperture; G two grains, showing proximal faces; H close up of gemmate surface, proximal face. J M C. renda (from Keith 2491): J whole grain, proximal face; K whole grain, distal face showing monosulcate aperture with membrane semi intact; L whole grain, distal face showing trichotomosulcate aperture; M close up of micro-fossulate surface, proximal face. Scale bar: 5 m. The Board of Trustees of the Royal Botanic Gardens, Kew, 2009