EPIGENEIUM (ORCHIDACEAE) OF JAVA AND SUMATRA DESTRI

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1 EPIGENEIUM (ORCHIDACEAE) OF JAVA AND SUMATRA DESTRI THE GRADUATE SCHOOL BOGOR AGRICULTURAL UNIVERSITY 2011

2 STATEMENT OF RESEARCH ORIGINALITY AND INFORMATION SOURCE This is to verify that my thesis entitled: Epigeneium (Orchidaceae) of Java and Sumatra is my own work and never been submitted to any institution before. All the incorporated data and information are valid and stated clearly in the text and listed in the references. Bogor, February 2011 Destri NRP. G

3 ABSTRACT Study on Epigeneium (Orchidaceae) in Java and Sumatra was conducted based on herbarium specimens and living collections. Previously three species from Java and 11 species from Sumatra were reported, however present study found five species from Java and 14 species from Sumatra. One new record for Sumatra (E. geminatum) and two species proposed as new species (E. longibulbon sp. nov. and E. sikijangensis sp. nov.) and for Java E. triflorum var. elongatum were altered into E. elongatum. Two species from Java were endemic (E. arjunoense and E. triflorum), nine species endemic to Sumatra, two species were found in both islands (E. cymbidioides and E. geminatum). Identification key was provided for all species found from Java and Sumatra. Key words: Epigeneium, Orchidaceae, Java, Sumatra.

4 ABSTRAK Kajian tentang jenis-jenis Epigeneium (Orchidaceae) di Jawa dan Sumatera dilakukan berdasarkan spesimen herbarium dan koleksi hidup. Sebelumnya dilaporkan tiga jenis dari Jawa dan 11 jenis dari Sumatra, namun dari hasil kajian ini ditemukan lima jenis dari Jawa dan 14 jenis dari Sumatra. Terdapat satu catatan baru untuk Sumatra (E. geminatum) dan dua jenis diusulkan untuk jenis baru (E. longibulbon sp. nov. dan E. sikijangensis sp. nov.), dan untuk Jawa E. triflorum var. elongatum dinaikkan ke tingkat jenis menjadi E. elongatum. Dua jenis endemik di Jawa (E. triflorum dan E. arjunoense), sembilan jenis endemik di Sumatera, dua jenis terdapat di kedua kawasan (E. cymbidioides dan E. geminatum). Untuk melengkapi dibuat kunci identifikasi untuk seluruh jenis yang terdapat di kedua kawasan ini. Kata kunci: Epigeneium, Orchidaceae, Jawa, Sumatera

5 SUMMARY DESTRI. Epigeneium (Orchidaceae) of Java and Sumatra. Supervised by SRI SUDARMIYATI TJITROSOEDIRDJO and IRAWATI Study on Epigeneium (Orchidaceae) in Java and Sumatra was conducted based on herbarium and living collections. Specimens from Herbarium Bogoriense (BO) and Herbarium Singapore Botanic Garden (SING) were observed during this study. Photo images of specimens and types from Leiden (L), Oakes Ames Orchid Herbarium (AMES), and Kew (K) were have been incorporated. Other additional specimens were from Andalas University Herbarium (ANDA). Based on morphological characters, Epigeneium has been revised into five species of Epigeneium from Java: E. arjunoense, E. cymbidioides, E. elongatum, E. geminatum, and E. triflorum. Epigeneium elongatum previously was placed as one of the variety of E. triflorum, now it has been placed back to species level because there are some distinct characters from the other two varieties were observed. In Sumatra there was one new record, E. geminatum since this species was never been found in this island before. Two new species were proposed from Sumatra: E. longibulbon Destri sp. nov. and E. sikijangensis Destri sp. nov. Epigeneium longibulbon was named base on its long pseudobulb, and E. sikijangensis was named after the original habitat where this orchid was collected (Rawa Sikijang), located at Mt. Kerinci, Jambi province. Epigeneium spp. inhabits primary forest from lowland up to mountain forest. They mostly found from lower mountain forest to mountain zone. In Java and Sumatra these orchids prefer an area with high rain fall. The identification key of Epigeneium from both islands, description of each species and illustrations of the new species were provided. Key words: Epigeneium, Orchidaceae, Java, Sumatra.

6 RINGKASAN DESTRI. Epigeneium (Orchidaceae) of Java and Sumatra. Dibimbing oleh SRI SUDARMIYATI TJITROSOEDIRDJO dan IRAWATI Penelitian tentang jenis-jenis Epigeneium (Orchidaceae) yang terdapat di Jawa dan Sumatera dilakukan berdasarkan spesimen herbarium dan koleksi hidup. Spesimen herbarium Bogoriense (BO) dan herbarium Kebun Raya Singapura (SING), diamati dalam penelitian ini. Foto spesimen dan tipe dari beberapa herbarium yaitu Leiden (L), Herbarium Anggrek Oakes Ames (AMES), dan Kew (K). Tambahan data lainnya diperoleh dari herbarium Universitas Andalas (ANDA). Berdasarkan pengamatan terhadap karakter morfologi direvisi lima jenis Epigeneium yang terdapat di Jawa, yaitu E. arjunoense, E. cymbidioides, E. elongatum, E. geminatum, dan E. triflorum. Epigeneium elongatum sebelumnya merupakan varietas dari E. triflorum. Studi ini menunjukkan bahwa jenis ini dikembalikan ke status sebelumnya yaitu Epigeneium elongatum karena terdapat beberapa karakter yang sangat berbeda dari dua varietas lainnya sehingga layak dinaikan ke tingkat jenis. Di Sumatera terdapat satu catatan baru yaitu E. geminatum, sebelumnya belum ada catatan mengenai keberadaanya di Sumatera. Dua jenis baru diusulkan dari Sumatra yaitu E. longibulbon Destri sp.nov dan E. sikijangensis Destri sp. nov. E. longibulbon dinamakan berdasarkan panjang umbi semu yang jauh melebihi jenis lain dalam marga ini, sementara penamaan E. sikijangensis berdasarkan lokasi tempat jenis ini ditemukan, yaitu di Rawa Sikijang, gunung Kerinci, propinsi Jambi. Epigeneium spp. tumbuh mulai dari hutan dataran rendah hingga pegunungan, terbanyak dijumpai dari hutan pegunungan bawah hingga zona montana. Di Jawa maupun Sumatra anggrek ini lebih sering dijumpai di kawasan yang lebih basah atau dengan curah hujan yang lebih tinggi. Hasil ini juga dilengkapi dengan kunci identifikasi, deskripsi, dan untuk jenis baru juga dilengkapi dengan ilustrasi. Kata kunci: Epigeneium, Orchidaceae, Jawa, Sumatera.

7 Copyright 2010, Bogor Agricultural University Copyrights are protected by law 1. It is prohibited to cite all parts of this thesis without referring to and mentioning the source a. Citation only permitted for the sake of education, research, scientific writing, critical writing or reviewing scientific problems. b. Citation does not inflict the name and honor of Bogor Agricultural University. 2. It is prohibited to republish and reproduce all part of this thesis without the written permission from Bogor Agriculture University.

8 EPIGENEIUM (ORCHIDACEAE) OF JAVA AND SUMATRA DESTRI A thesis submitted to fulfill one of the requirements for Master Degree in Biology at Study Program of Plant Biology, The Graduate School, Bogor Agricultural University THE GRADUATE SCHOOL BOGOR AGRICULTURAL UNIVERSITY 2011

9 External Examiner: Dr. Nunik Sri Ariyanti, M.Si.

10 Title of Thesis : Epigeneium (Orchidaceae) of Java and Sumatra Name : Destri Registration Number : G Study Program : Biology Approved by Supervisor Committee Dr. Sri Sudarmiyati Tjitrosoedirdjo, M.Sc. Chairman Dr. Irawati Member Head of Plant Biology Study Program Dean of Graduate School Dr.Ir. Miftahudin, M.Si Prof.Dr.Ir. Khairil A. Notodiputro, M.S. Date of examination: Date of Graduation: December 23 rd, 2010 February 7, 2011

11 ACKNOWLEDGMENTS The great gratitude to Allah SWT for makes it happen. The author realized that this thesis will never possible without encouragement and assistance from colleagues and friends who have contributed in many ways. I would like to express my grateful to my supervisors Dr. Sri Sudarmiyati Tjitrosoedirdjo, M.Sc. and Dr. Irawati for their valuable advices and critical correction in completing this thesis; to Dr. Nunik Sri Ariyanti for giving important suggestions on the manuscript. I would like to acknowledge Dr. Didik Widyatmoko, M.Sc. the Head of Cibodas Botanic Garden LIPI, for granting me permission during my study; Prof. Dr. Eko Baroto Walujo, Head of Botanic Division, Center Research for Biology, LIPI for the permission using specimens of the Herbarium Bogoriense (BO); Prof. Dr. Ir. Khairil A. Notodiputro, M.S., the Dean of Post Graduate School, Bogor Agricultural University; Dr. Ir. Miftahudin, M.Si., the Head of Study Program of Plant Biology for their support during this study. I deeply thank to Herbarium Bogoriense for all facilities, to Dra. Diah Sulistyorini, M.Si., Arief Hidayat, S.Si. and staff for their assistance during my work in BO; to Herbarium of Singapore Botanic Garden, Dr. Serena Lee and staff, to Dr. Barbara Gravendeel and Dr. Jeffrey J. Wood for valuable publications; Dr. Titien Ng. Praptosuwiryo, S.Si, M.Si. for her advices and discussion. I also thanks to Amas, SE, MM. and Ir. Rismita Sari M.Sc. for permission in using facilities at Bogor Botanic Garden, and also to all researchers, staff, and technicians in Cibodas Botanic Garden for discussion and assistance; to all my friends at the Plant Biology Study Program 2008, the Graduate School, Bogor Agricultural University: Arief, Doel, Ina, Rina, Sukiman, Wina and Yesi for sharing and their support. Finally to my parents, my brother and sisters for their prays, keep believing and encouraging. Bogor, February 2011 Destri

12 CURRICULUM VITAE The author was born in Payakumbuh, West Sumatra on December , the youngest daughter of four children from Mr. Zulfahmi and Mrs. Dahniar Abbas. In 1992, she attended the Department of Biology, Faculty Mathematic and Natural Science, Andalas University, Padang and graduated in Since 2001, she was employed by Cibodas Botanic Garden, Technical Implementing Unit for Plant Conservation, Indonesia Institute of Sciences (LIPI) and promoted as junior researcher in In 2008 she continued her study at Bogor Agricultural University (IPB) for her Master degree majoring on Plant Biology.

13 CONTENT page LIST OF FIGURES... xiii LIST OF PLATES.. xiv INTRODUCTION.. 1 MATERIAL AND METHOD... 4 Material Method RESULT AND DISCUSSION... 5 Diversity of Epigeneium spp. in Java and Sumatra Habitat and Ecology... 6 Distribution The General Characters of Epigeneium Taxonomy Treatment Generic Description.. 17 Key to Species E. arjunoense J.J. Wood & J.B. Comber E. cymbidioides (Blume) Summerh E. dempoense (J.J. Sm.) Summerh E. elongatum (Blume) Summerh E. exilifolium (Ames & Schweinf.) Garay & G.A. Romero E. geminatum (Blume) Summerh E. gracilipes (Burkill) Garay & G.A. Romero E. laurifolium (Kraenzl.) Summerh E. longibulbon Destri sp. nov E. longipes (Hook.f.) Summerh E.macropodum (Hook.f.) Summerh E. pulchellum (Ridl.) Summerh E. quinquecallosum (J.J. Sm.) Summerh E. sikijangensis Destri sp. nov E. triflorum (Blume) Summerh

14 16. E. wichersii (Schltr.) Summerh E. zebrinum (J.J. Sm.) Summerh.. 38 CONCLUSION.. 40 REFERENCES.. 41 PLATES

15 LIST OF FIGURES No page 1. Distribution map of E. cymbidioides and E. elongatum in Java Distribution map of E. triflorum in Java Distribution map of E. arjunoense and E. geminatum in Java 8 4. Distribution map of Epigeneium spp. in Sumatra: E. dempoense, E. exilifolium, E. geminatum, E. gracilipes, E. laurifolium, E. longibulbon, E. longipes Distribution map of Epigeneium spp. in Sumatra: E. cymbidioides, E. macropodum, E. pulchellum 6. Distribution map of Epigeneium spp in Sumatra: E. quinquecallosum, E. sikijangensis, E. wichersii, E. zebrinum Epiphytic Epigeneium in their natural habitat (Mt. Gede, West Java) Epigeneium compare to its relatives Leaf shapes of Epigeneium spp The number of inflorescence on each pseudobulb Some variations of shapes and colors in flower of Epigeneium spp Some variation of the labellum ornament in Epigeneium spp. 16

16 LIST OF PLATES No page 1. Epigeneium cymbidioides (Irawati 910) Holotype of Epigeneium dempoense (E. Jacobson and Ajoeb 443) Epigeneium elongatum (M. Fleicher 38) Holotype of Epigeneium exilifolium (Walter.N. & Catherine M. Bangham 1124, AMES Orchid Herbarium) Epigeneium geminatum from Sumatra (J.J. Afriastini 2720) Holotype of Epigeneium longibulbon (H. Nagamasu 3414) Flower parts of E. longibulbon: A. dorsal sepal, B. lateral sepal, C. petal, D. labellum (H. Nagamasu 3414) Epigeneium longipes (R.E Holtum and M.R. Henderson 23398) Epigeneium macropodum (H.N Ridley 16134) Epigeneium pulchellum (C.G.G.J. Van Steenis 9234) Holotype of Epigeneium sikijangensis (Wiguna Rahman 107) Flower parts of Epigeneium sikijangensis: A. dorsal sepal, B. lateral sepal, C. petal, D. labellum (Wiguna Rahman 107) Holotype of Epigeneium triflorum var. orientale Epigeneium triflorum var. triflorum (J.J. Smith s.n) Epigeneium wichersii (C.G.G.J. van Steenis 9164) Holotype of Epigeneium zebrinum (Herbarium of Singapore Botanic Garden).. 58

17 INTRODUCTION Epigeneium (Orchidaceae) belongs to tribe Dendrobieae, subtribe Dendrobiinae together with Cadetia, Dendrobium, Diplocaulobium, Flickingeria, and Pseuderia (Dressler 1993). Previously it was placed in genus Dendrobium but then it was separated into Epigeneium because it has one internode and short mentum. Seidenfaden and Wood (1992) stated that this genus consists of about 35 species, in Index Kewensis 1997, 44 species were listed and according to Wood (2004), Epigeneium has 48 species. The member of this genus is a medium size orchid and under favorable environment may grow to a dense clump. It has beautiful inflorescence and long lasting flower and this genus is potential for creating new hybrids. Otherwise it can be propagated as ornamental plant since it has attractive flowers. Unfortunately the beauty of these orchids are almost overlooked by plant breeder and become rarely cultivated (Comber 1990). Epigeneium distributes from China, Indochina, Taiwan, Thailand, the Philippines, the largest number of this genus is found in eastern part of Malaysia and Indonesia (Seidenfaden & Wood 1993). From Java there are 3 species, 11 species from Sumatra (Comber 1990, 2001), 4 species from Mount Kinabalu (Wood et al. 1993), and 8 species from the whole Borneo. Three center of diversities are located in Indochina, Sumatra, and Borneo. Most Epigeneium is mountain forest orchid (Seidenfaden & Wood 1992), epiphytic on higher tree branches to reach enough sun light (Comber 1990). In Java and Sumatra Epigeneium usually found from 1,500 m above sea level, such as E. cymbidioides, E. pulchellum, E. triflorum (Comber 1990, 2001). In Borneo this orchid found from lower mountain forest up to 1,700 m asl such as E. kinabaluense, E. tricallosum, and E. longirepens (Wood et al. 1993). This genus formerly was known as Sarcopodium which was proposed by Lindley, and at present time some member of this genus are separated into Bulbophyllum and Dendrobium. According to Lindley (1853), between Dendrobiums and Bulbophyllums there exist a race having the large flower of the former, and the peculiar habit of the latter, and hence referred to the one or other

18 2 genus according to the fancy of the observer. They agree with Dendrobiums in having four pollen masses and a hornless column but they have coriaceous, not thin half-transparent flowers, and a tough leathery lip, enlarge not contracted at the base. If they had a caudicle and gland to their pollen masses, they would be Asiatic Maxillarias. They form neither horn nor spur, but are simply inflated and expanded at the base of the sepals. On the other hand, although they grow like Bulbophyllums, yet they have no horn to their column, but two pollen masses and their large leathery flowers afford a further difference. To these plants, consisting of the Dendrobium amplum of Wallich, and the Bulbophyllum lobbii, B. affine leopardiana, B. cheirii, and B. macranthum of Lindley, the name Sarcopodium may be applied. Gapnepain (1932) proposed genus Epigeneium based on Dendrobium fargesii Finet, and this species became type of this genus. Together with E. fargesii, Gagnepain also described 3 other species: E. clemensiae, E. dalacourii, and E. chapaense. Recent papers by Tang and Wang in 1951, they reduced Gagnepain s genus Epigeneium and including several other species to Sarcopodium, and unfortunately this publication overlooked by A.D. Hawkes (Summerhayes 1957). According to Hawkes (1956), Sarcopodium is the genus name for fungi and he proposed Katherinea to replaced Sarcopodium. Because Gagnepain already proposed before Hawkes, therefore Epigeneium became legitimate name for this genus (Summerhayes 1957). Previously Dressler and Dodson (1960) put Epigeneium within tribe Epidendreae, subtribe Dendrobiinae together with Bulbophyllum, Dendrobium, Pedilochillus, Pseuderia, and Saccoglossum. Later Dressler (1993) placed Epigeneium within the same subtribe Dendrobiinae together with Cadetia, Dendrobium, Diplocaulobium, Flickingeria, dan Pseuderia. Garay dan Romero (1999) by using the character differences between Epigeneium Gagnep. and Katherinae A.D. Hawkes to established infrageneric categories and divided Epigeneium into three sections: Epigeneium, Katherinea, and Racemosum based on number of the flower, the length of the column foot, and the vegetative appearance.

19 3 Based on molecular data Clement in Everet and Wilson (2003) proposed subtribe Epigeneiinae which cover Epigeneium, after he found that only weak support for its position on phyllogram as the part of Dendrobiinae. He also found that Epigeneium is monophyly. In Java, Comber (1990) described three species; E. cymbidioides, E. geminatum and E. triflorum with three varieties (E. triflorum var. elongatum, E. triflorum var. orientale and E. triflorum var. triflorum. However, he also mentioned that E. triflorum var. elongatum is quite distinct and perhaps should be elevated to species rank. Meanwhile, in Sumatra according to Comber (2001), around 10% of orchid family waits to be found. On the other hand, Sumatra as one of the diversity s centers of Epigeneium still has limited scientific knowledge due to under collection. This genus needs a complete monograph study (Wood and Comber 1996). Therefore, an attempt in order to uncover the knowledge on Epigeneium is urgently needed. The result below reveals new information about species diversity, the distribution of this genus, and provide identification key for species in Java and Sumatra. The result also discuss about Comber s opinion on taxonomy of Epigeneium in Java. It is hoped that this result will contribute valuable information for further research regarding to this genus in any field of biological science.

20 4 MATERIALS AND METHOD Materials This study was based on herbarium specimens of Epigeneium of Herbarium Bogoriense (BO), and specimens from Herbarium of Singapore Botanic Garden (SING) were observed to accomplish the material which does not available at BO. Image of type specimen from Leiden (L), Oakes Ames Orchid Herbarium (AMES), and Kew (K) were observed during this study and they are available online. As additional information, Herbarium collection of Andalas University Herbarium (ANDA) was confirmed. Living collection in Cibodas Botanic Garden was also observed during this study. Method Surveys were conducted at Mt. Gede Pangrango (West Java) and Mt. Singgalang (West Sumatra) to obtain additional material. Field collection method was based on Rugayah et al. (2004). Herbarium specimens were observed in Biosystematic Laboratory at BO following Rifai (1976) and De Vogel (1987), and for morphological character terminology following Harris (1994) and Seidenfaden and Wood (1992).

21 5 RESULT AND DISCUSSION Diversity of Epigeneium in Java and Sumatra Observation on herbarium specimen and living material recognized five species of Epigeneium in Java and 14 species in Sumatra. Previously, Comber (1990, 2001) only described three species in Java and 11 species in Sumatra. Epigeneium arjunoense is a new species proposed by Wood and Comber (1996). There are no new species found in Java but this study decide to put back one variety of E. triflorum (E. triflorum var. elongatum) to species rank and named by its previous name proposed by Summerhayes (1957) which is E. elongatum (Blume) Summerh. This study found that the labellum of E. elongatum has three calli in the middle, the center one is the longest, the mid-lobe is rounded and smaller compare to other varieties and the margin of the mid-lobe is thickened. The vegetative part also different in having oblongoid pseudobulb with four sharp angles, leaves elliptic and tough. Epigeneium geminatum is a new record from Sumatra, although the specimen was collected from Siberut Island outside of the mainland. Comber (2001) did not describe this species in Sumatra. this species is also found in Borneo and Peninsular Malaysia. Two species are proposed as new species and was named E. longibulbon Destri sp. nov. based on its longest pseudobulb compare to other species. The herbarium specimen was collected from mount Talang, West Sumatra. Another one was named E. sikijangensis Destri sp.nov., the specific epithet refers to the locality where this orchid was found, Rawa Sikijang, located on Mt. Kerinci. The complete result of this study is presented in Table 1.

22 6 Table 1. List of Epigeneium spp. in Java and Sumatra No. Species Distribution area Note Java Sumatra 1. E. arjunoense J.J. Wood & J.B. Comber E. cymbidioides (Blume) Summerh E. dempoense (J.J. Sm.) Summerh E. elongatum (Blume) Summerh E. exilifolium (Ames & Schweinf.) Garay & G.A. Romero E. geminatum (Blume) Summerh. + + New record for Sumatra 7. E. gracilipes (Burkill) Garay & G. A. Romero E. laurifolium (Kraenzl.) Summerh E. longibulbon Destri sp. nov. - + Proposed as new species 10. E. longipes (Hook.f.) Summerh E. macropodum (Hook.f.) Summerh E. pulchellum (Ridl.) Summerh E. quinquecallosum (J.J. Sm.) Summerh E. sikijangensis Destri sp. nov. - + Proposed as new species 15. E. triflorum (Blume) Summerh E. wichersii (Schltr.) Summerh E. zebrinum (J.J. Sm.) Summerh. - + Habitat and Ecology Epigeneium spp. inhabit primary forest from lowland to mountain zone. In the lowland this species can be found from 300 m above sea level, for example E. geminatum and E. zebrinum. Epigeneium geminatum in Halimun Salak National Park was collected from 700 1,100 m asl but in Siberut Island it was found at 300 m asl. In cultivation on about 1,300 m asl, this species adapt very well. Another species which was found on lower mountain forest are E. wichersii and E. sikijangensis from 800-1,000 m asl.

23 7 The sub mountain and mountain zone is the most habitat of this genus. In mount Gede Pangrango National Park E. triflorum was found from 1,500 asl, similar to E. dempoense and E. quinquecallosum were found on 1,400 1,700 m asl on mount Dempo, South Sumatra. Meanwhile, some species found at mountain zone; E. cymbidioides found from 2,000 m asl and up near to the peak of Mt. Pangrango, on the branch of short tree, so it can be observed easily; Epigeneium pulchellum found on the lower part of mossy tree trunk on 1,900 2,200 m asl at mount Kerinci, and even on the higher elevation in North Sumatra and Aceh, up to 2,500 m asl; on mount Singgalang E. laurifolium was collected from 1,900 m asl. All habitats of these orchids in Sumatra are on the western slope of Barisan Range which has greater rainfall (Whitten et al. 1997). In Java this orchid mostly found in West Java, the wettest area in Java. It can be concluded that this genus grow better in the area where the rainfall is high. However these orchids also love full sunlight, as seen in E. triflorum which mostly occupies the high tree branch. In cultivation these orchid grow better without shades. The flowering season varies; in cultivation it is possible to find it flowering at any months of the year, sometime different species flowering at the same time. Distribution Of five species found in Java, four of which were found in West Java: E. cymbidioides, E. elongatum, E. geminatum, E. triflorum. Epigeneium cymbidioides is spread to Central Java but not found in East Java (Figure 1). It seems like E. cymbidiodes prefer the mountain forest with high rainfall such as in Mt. Gede - Pangrango. E. triflorum was also found in Central Java and one of the varieties was mostly found from East Java (E. triflorum var. orientale) (Figure 2). Epigeneium elongatum spreads in West Java at Mt. Halimun and Mt. Gede- Pangrango (Figure 1), there was no specimen collected from other location in West Java. Epigeneium geminatum only found in mount Halimun and Salak, and there was no record from other location in Java. Epigeneium arjunoense until now

24 8 only known from East Java. Distribution map of this two species was shown in Figure 3. Figure 1. Distribution map of E. cymbidioides ( ) and E. elongatum ( ) in Java Figure 2. Distribution map of E. triflorum ( ) in Java Figure 3. Distribution map of E. arjunoense ( ) and E. geminatum ( ) in Java In Sumatra the large numbers of these species were found in West Sumatra (Figure 4-6). Of 4 species, 8 species were distributed in West Sumatra from northern to southern part: Mt Talaumau (E. cymbidioides), Mt. Singgalang (E. laurifolium, E. wichersii), Mt. Sago (E. cymbidioides, E. longipes), Laras Talang (E. quinquecallosum), Air Sirah (E. cf. sikijangensis), Mt. Talang (E. longibulbon), and Siberut island (E. geminatum). Epigeneium pulchellum found from Tanah Gayo (Aceh), Sidikalang (North Sumatra), and Mt. Kerinci.

25 9 Figure 4. Distribution map of Epigeneium spp. in Sumatra: E. dempoense ( ), E. exilifolium ( ) E. geminatum ( ), E. gracilipes ( ) E. laurifolium ( ), E. longibulbon ( ), E. longipes ( ) Figure 5. Distribution map of Epigeneium spp. in Sumatra: E. cymbidioides ( ), E. macropodum ( ), E. pulchellum ( )

26 10 Figure.6. Distribution map of Epigeneium spp in Sumatra: E. quinquecallosum ( ), E. sikijangensis ( ), E. wichersii ( ), E. zebrinum ( ) Epigeneium quinquecallosum also found in Mt. Dempo in South Sumatra together with E. dempoense. Epigeneium gracilipes was found in Riau Archipelago, so far this is the only species which found on eastern part of Sumatra. Epigeneium pulchellum occur in three different areas, but another species mostly found only one area. There are unusual patterns on their distribution as shown in this result. Epigeneium geminatum is a species with widest area of distribution (Java, Sumatra, Borneo, and Peninsular Malaysia) but in Java this species only found in Mt. Halimun and Salak. There is a possibility that this species need a specific microclimate which only match in particular area, the other reason is lack of collection or overlooked by the explorer. So far for 14 species from Sumatra 9 of them are endemic or lack of collection in Sumatra as mention before. Here, it is also shown that West Sumatra is the richest area for this genus or because many explorations had been conducted in many areas in this province compare to

27 11 others. It is obviously needed to conduct more explorations through Sumatra to get better information on this genus. The General Characters of Epigeneium. Habit Epigeneium is known as epiphytic or sometimes lithophytic orchid. Rhizome and root help this plant to creep and thrive on branch of the tree meanwhile the pseudobulb save the water, so this orchid will survive during dry season. Since they love sunlight, this orchid mostly found up on the top of the tree branches. It can be easily recognized by shiny, yellow pseudobulbs. The habit and the vegetative part are shown in Figure 7. Root Figure 7. Epiphytic Epigeneium in their natural habitat (Mt. Gede, West Java) Root is fibrous, sometime flexuous, and branching, it is mostly born from the lower part of the pseudobulb and sometimes also found along the rhizome. Some species of this genus show different root thickness as in E. longipes and E. zebrinum which have thicker root from all specimens examined, up to 2.5 mm and the other species less than that. Even some authors mention that this character is

28 12 obvious in some species, but it is not a good character to distinguish among species in this genus since it depends on the environment. Rhizome Epigeneium has tough woody rhizome, cylindrical, branching, and creeping, and covered by brown cataphylls. Rhizome thickness is from 2-7 mm, varies for different species, and it may grow and form a big clump on tree branch. The large and stout species like E. dempoense and E. zebrinum have thicker rhizome compare to the small one as E. pulchellum, although this character can be changed due to the environment condition. The rhizome is covered with imbricate layered cataphylls, reddish when immature and gradually turns to brown. In some species, immature cataphylls is light green with purple spots which make longitudinal streak like in E. pulchellum and E. sikijangensis. This cataphylls mostly persistent but in some species like E. geminatum, it was peeled off. Pseudobulb Epigeneium has single internode pseudobulb and shared this character with other two relatives: Flickingeria and Diplocaulobium. It is different from Dendrobium which has many internodes (Figure 8). 5 cm 3 cm 3 cm 3 cm A B C D Figure 8. Epigeneium compare to its relatives. A. Epigeneium B. Dendrobium, C. Diplocaulobium, D. Flickingeria

29 13 The shape of the pseudobulb varies from almost globose, ovoid, long ovoid to ellipsoid. In most cases with distinct angle, almost circular or quadrangular in cross section, deep groove (sulcate) at all sides; yellow, olive green and sometime purple when the pseudobulb start to open and gradually became greenish purple and shiny. Their size also varies from very short and small like in E. pulchellum to large as in E. zebrinum and longer as in Epigeneium longibulbon. The latest has the longest pseudobulb, up to 14 cm and with no angle, so that the name which proposed for this species is Epigeneium longibulbon (Epigeneium with long pseudobulb). In Epigeneium geminatum the shape of pseudobulb on the lower part and the upper part of the rhizome are different; the pseudobulb at lower part is ovoid with distinct angle; on the upper part the pseudobulb tend to be ellipsoid or shortly cylindrical, sometimes rather flattened. Leaves Epigeneium has one, two, and three to four leaves on each pseudobulb. The species from Borneo E. longirepens has single leaf similar to Flickingeria spp. There is variation within species such as E. triflorum var. triflorum which sometimes has 3 leaves, and it is unusual since this feature is not appeared on each individual plant; most plant consist only 2 leaves and some consist of 2 and 3 leaves. In Epigeneium longibulbon, the leaf number is 2-4 on each pseudobulb, this is new character for the genus. Leaf shaped ovate, narrower ovate, ovate, lanceolate, elliptic, linear, strap shaped, and there are combination within individual plant (Figure 9). Leaves size is highly influenced by the environment condition where this orchid grows. Leaves of E. cymbidioides become longer in cultivation. Among all species within this genus E. pulchellum has smallest leaf size from cm long.

30 14 3 cm Figure 9. Leaf shapes of Epigeneium spp. a. narrower ovate (E. pulchellum) b. elliptic (E. elongatum) c. strap shaped (E. triflorum), d. linear (E. triflorum), e. long elliptic (E. cymbidioides) Inflorescence a b c d e Inflorescences are terminal or semi terminal. Mostly with single inflorescence appears between the two leaves (Figure 10), except in E. geminatum dan E. zebrinum, which has 2 inflorescences on each pseudobulb. A B Figure 10. The number of inflorescences on each pseudobulb A. Two inflorescences (E. geminatum), B. Single inflorescence (E. cymbidioides). The inflorescence is longer, shorter or equal compare to the leaf. In some species this character became very obvious such as in E. longipes, E. longibulbon,

31 15 E. macropodum which have inflorescence longer than the leaves, and E. exilifolium has shorter inflorescence. The number of flower from each inflorescences are 1 to 20, whereas E. pulchellum usually only produces one or two flowers. All species show simultaneous inflorescences (racemose), and last for two weeks. Flower The flower cm width, sepal and petal vary in color; white, cream, peach, dull yellow or greenish yellow and one species yellow with fine red spots (Figure 11). C A D B E Figure 11.Some variations of shapes and colors in flower of Epigeneium spp. A. greenish yellow, sepal and petal obtuse (E. cymbidioides,) B. white, sepal and petal acute (E. triflorum) C. peach, sepal and petal acute (E. geminatum) D. yellow with fine red spots, sepal and petal obtuse (E. sikijangensis), E. yellow, sepal dorsal and petal obtuse, lateral sepal acute (E. elongatum) Each flower is supported by a small bract, acute or acuminate, greenish translucent. Pedicel is short with long ovary, slightly twist, length vary between

32 16 species. Sepal and petal in some species fleshy like in E. cymbidioides. The shape of dorsal sepal elliptic or oblong, and the apex acute or obtuse, petal usually narrower, sometime spathulate, lateral sepal elliptic to lanceolate, oblique and wider at base, connected to the column foot which make short mentum (chin). Self compatibility was also found within this genus where fertilization occurred just after flower bloom, and in this case, flower faded earlier. Labellum or lip has three lobes, the side lobes erect sometimes covering the column. The disc or lamina (between side lobes) is ornamented with 3-5 longitudinal calli (keels) or knobs. The arrangement of the keels is not quite different in this genus; sometimes with several irregular calli on the base of the labelum, another three in the middle with variation in the length of the callus (Figure 12). The mid-lobe is rounded to rhomboid, acute to acuminate, sometime very warty on the disc and some species with thickened margin. 0.5 cm 0.5 cm A C 0.5 cm 0.5 cm B D Figure 12. Some variations of the labellum ornament in Epigeneium spp. A. three keels between side lobes, the center one is the longest (E. elongatum), B. several keels on the base and three keels between side lobes, almost equal (E. geminatum), C. several keels on the base of labelum (different arrangement with the Previous one), three keels between side lobes, almost equal (E. longibulbon), D. three keels between the sidelobes, the center ones is placed far behind the two (E. triflorum)

33 17 Taxonomy Treatment Generic Description Epigeneium Gagnep Gagnep, Bull. Mus. Hist. Nat. Paris, ser. 2, 4, 595 (1932), in clavi. Type: Dendrobium fargesii Finet. Epiphytic, rhizome cylindrical, woody, creeping, branching, 2-7 mm in thickness, covered by several brown cataphylls. Root terete, fibrous, sometime flexuous, branching mostly bears from lower part of pseudobulb. Pseudobulb with single internode, globose, ovoid, elliptic to long or narrower ovoid, blunt or with sharp angle, circular or quadrangular in cross section, sometime sulcate, yellow, olive green or purple, shiny. Leaf 1-4 on each pseudobulb, varies in shape; ovate, narrower ovate, lanceolate, elliptic, oblong, linear, strap shaped, the apex acute or obtuse, obliquely slightly bilobed, shiny, coriaceous with cm petiole. Inflorescences terminal or semiterminal, single or two on each pseudobulb, racemose, 1-20 flowers, peduncle longer or not much different than the rachis, with 2-3 sterile bracts, each flower supported by small bract. Flower c cm width, white, yellow to greenish yellow, sometime rather peach; dorsal sepal elliptic, oblong or lanceolate, the apex acute or obtuse, with several nerves, x cm, lateral sepal rather lanceolate or elliptic, acute or obtuse, wider at base, oblique, connect to column foot and make a short mentum, x cm, petal usually narrower, linear, sphatulate or narrow elliptic, acute or obtuse, x Labellum trilobe, x , recurved, fleshy, with ridge (callus) at the base, sometime elongate to the middle part, mostly with knob or keels near the mid-lobe, with very short claw, the side lobes erect, triangular with red or purple streak on inside surface, the mid-lobe rounded, ovate or triangular, acute, acuminate or obtuse, the margin thin or thickened, sometime with projection on the disc. Column 4-9 mm, stelidia very short, white or red streak on surface, column foot c. 2-5 mm, pollinia 4.

34 18 Key to Species 1. a. Two inflorescences from each pseudobulb b. Single inflorescence from each psedobulb a. Labellum without calli on the base, three keels between the side lobes, the center one is the longest, mid-lobe acute E. zebrinum b. Labellum with several calli on the base, three keels between the side lobes and almost equal in size, the mid-lobe sub acute. 6. E. geminatum 3. a. Inflorescence shorter than the leaves E. exilifolium b. Inflorescence longer than the leaves a. Inflorescence with more than 3 flowers b. Inflorescence with 1-3 flowers a. Peduncle much longer than the rachis (3:1) E. longipes b. Peduncle slightly longer than the rachis sometimes equal, the ratio between peduncle and rachis less than 1: a. Leaves linear or strap shaped E. triflorum b. Leaves ovate, elliptic or oblong a. Pseudobulb cm long, swollen basally and cylindrical toward the apex, with 2 leaves or more E. longibulbon b. Pseudobulb cm long, globose, ovoid or oblongoid, with only 2 leaves a. Dorsal sepal lanceolate b. Dorsal sepal elliptic or oblong a. Five calli between the side lobes of the labellum 13. E. quinquecallosum b. Three calli between the side lobes of the labellum a. Leaves oblong b. Leaves elliptic or long elliptic or broader elliptic... 12

35 a. Labellum with three keels from the lip base to its apex, the two outer one broader than the center ones E. wichersii b. Labellum with three keels at the base, the side one much longer; another three keels in the middle, the center ones extending to the base of the labellum E. laurifolium 12. a. Pseudobulb with sharp angle b. Pseudobulb with blunt angle a. Pseudobulb olive green, sepal and petal yellowish green, column light green E. elongatum b. Pseudobulb, purplish to greenish purple, sepal and petal yellow with fine red spots, column yellow with red streaks E. sikijangensis 14. a. Sepal and petal white basally and yellow toward the apex E. cymbidioides b. Sepal and petal pale ochre, pale orange, or pale yellow, lighter towards the base E. arjunoense 15. a. The mid-lobe almost circular, the margin with branched nerved E. macropodum b. The mid-lobe ovate, the margin very warty E. dempoense 16. a. Pseudobulb x in size, leaves narrower ovate elliptic c x cm E. pulchellum b. Pseudobulb 3 x 1.4 in size, leaves narrowly elliptic, 16 x 2.5 cm E. gracilipes 1. Epigeneium arjunoense J.J. Wood & J.B Comber Epigeneium arjunoense J.J. Wood & J.B. Comber, Orch. Rev. 104 (1996) Type: East Java, Mt. Arjuno 2200 m asl, November 12, 1998, Comber 1539 (K, holo; herbarium and spirit, n.v) Rhizome creeping, cataphylls 3-4, ovate-elliptic, obtuse, pergamaceus, 1-5 cm long, brown. Pseudobulb ovoid to ellipsoid, six angled, somewhat bilaterally compressed, x 1.3 cm, bifoliate, yellow. Leaves oblong elliptic to elliptic,

36 20 cuneate below, apex equally or unequally obtusely bilobed, texture coriaceous, 6-11 x cm, dark green. Inflorescence terminal, 3-9 flowered, erect to porrect, flowers borne cm apart, all opening simultaneously; peduncle 6-7 cm long; rachis 4-5 cm long; floral bract ovate to elliptic, acute, cm long. Flowers with pale ochre, pale orange or pale yellow sepals and petals, lighter toward the base, labellum whitish or pale yellow with orange or crimson streaks and spots inside on the side lobe and some brown or yellow pigment on the mid-lobe, pedicel and ovary slender, 3-4 cm long; dorsal sepal oblong, obtuse, x cm, 7 nerved; lateral sepal obliquely oblong to oblong-ovate, obtuse, x cm, obscurely carinate on the reverse, nerved; petal narrowly oblong, ligulate, obtuse, 1.6 x cm, 6 nerved; labellum trilobed, cuneate at base, x 0.8 cm, side lobes erect, obliquely oblong, obtuse to truncate, margin undulate, often irregularly lobed, cm long; mid-lobe transversely elliptic, trilobed, thickened, fleshy, x 0.6 cm, side lobules oblong, truncate, incurved, mid-lobule rounded; disc with several irregular, rounded papillate calli between side lobes and a thickened, papillose area at base of mid-lobe; column porrect, straight to gently curving, shallowly concave, dilated distally, apical margin uneven, cm long, foot incurved, 0.3 x 0.2 cm; anther cap cucullate, conical above, truncate below, smooth, x mm. Distribution: Mt. Arjuno, East Java. Habitat: Mountain forest at c m asl. Note: This species was described in 1996 by J.J. Wood and J.B. Comber and it seems to be close related to E. cymbidioides. So far, this species is only known from Mt. Arjuno East Java. The specimen is not available and the type in Kew is inaccessible. This description was based on publication (Wood and Comber 1996). 2. Epigeneium cymbidioides (Blume) Summerh. (Plate 1). Desmotrichum cymbidioides Blume, Bijdr. (1825) 552. Dendrobium cymbidioides (Blume) Lindl., Gen. & Sp. Orch. Pl. (1830) 77. Sarcopodium cymbidioides (Blume) Rolfe, Orch. Rev. 18 (1910) 238. Katherinea cymbidioides (Blume) A.D. Hawkes, Lloydia 19 (1956) 95.

37 21 Epigeneium cymbidioides (Blume) Summerh comb.nov., Kew Bull. (1957) 261; Comber, Orchids of Java (1990) 247; Comber, Orchids of Sumatra (2001) 678. Type: Java, collector unknown, s.n. (L!, holo, photo image). Rhizome creeping, branching, c mm in thickness, with brown layered cataphylls. Root terete, fibrous, branching, surface smooth. Pseudobulb globose - ellipsoid, x cm, angular, yellow, shiny, and placed very closed to 3.5 cm apart on the rhizome. Leaves two on each of pseudobulb, elliptic long elliptic, x cm, obtuse, obliquely bilobe, coriaceous, green and shiny, petiole 4-15 mm. Inflorescence terminal, 1 on each pseudobulb, peduncle cm with 2 sterile bracts, rachis shorter cm, bring 5-7 flowers. Flower c.4 cm wide, sepal and petal white at very base and dull yellow toward the apex; dorsal sepal oblong, obtuse, 2.5 x 0.8 cm, 7 nerved; lateral sepal slightly lanceolate, oblique and wider at base, sub acute, 2.3 x 0.9 cm 7 nerved; petal narrower or strap shape, obtuse, 2.4 x 0.55 cm; labelum trilobes, with several irregular undulating calli at the base, side lobes erect, two knob near the mid-lobe, the mid-lobe rhomboid, acute, mostly yellow and white at the margin; column white, c. 9 mm, purple streak on inside surface, column foot 5 mm. Distribution: Mt. Gede-Pangrango, Ranca Cibodas, Mt. Salak, Mt. Ciremai (West Java), Wonosobo (Central Java), Mt. Talamau, Mt Sago Malintang (West Sumatra). Habitat: on higher mountain forest from up to 2,000 m asl, flowering time: May, July, August, and October. Note: this species show great plasticity in cultivation; the pseudobulb become longer and the angle disappear or blunt, leaves become much longer and soft. The flower at the beginning yellowish green and later turn to dull yellow, the size is larger compare to the native ones. Lip shape almost similar to E. triflorum, but different in the calli found on the base of the labellum. Specimen examined: Java, 2,200 m asl, 1919, L.J. van Bligs sn (BO); West Java, Ranca Cibodas, 1,900 m asl, July , J.H. de Haan 7 (BO); Mt. Salak 2,200 m asl, August 8, 1931, V.J. Lam (BO); Kandang Badak 2,400 m asl, July , C.G.G.J. van Steenis 1979 (BO); Mt. Salak 1,600-2,200 m asl, July , C.G.G.J. van Steenis (BO); Kandang badak 2,300 m asl, May

38 22 (BO); Central Java, Dieng, Wonosobo, 2,550 m asl, Irawati 910 (BO); West Sumatra, Mt. Talamau 2,100 m, May 1919, Bunnenmeijer 921a (BO). 3. Epigeneium dempoense (J.J. Sm.) Summerh. (Plate 2) Dendrobium dempoense J.J. Sm. in Bull. Jard. Bot. Buitenz. 3 (1920) Sarcopodium dempoense (J.J.Sm.) Carr in Journ. As. Soc. Malaya, 11 (1933) 87. Epigeneium dempoense (J.J. Sm.) Summerh. comb.nov., Kew Bull. (1957) 261; Comber, Orchid of Sumatra (2001) 679. Type : Mt. Dempo, South Sumatra, E. Jacobson and Ajoeb 443, August 9, 1916 (BO!, holo) Rhizome stout, creeping, 5 mm in thickness, covered by c. 10 imbricate brown cataphylls, c cm long. Pseudobulb placed cm apart along the rhizome, x cm, ovoid - oblongoid with 4 sharp angles, sulcate. Leaves 2 on each pseudobulb, x cm with 4 second major nerves, petiole cm. Inflorescence single, from top of pseudobulb; peduncle 17.8 cm with 2 sterile bracts; rachis 10.7 cm, bring 11 flowers, open at the same time, placed cm apart. Flower c. 4.3 x3.1 cm in size, yellow, tip of the labellum red; dorsal sepal lanceolate, obtuse, 2.25 x 0.43 cm; lateral sepal obliquely oblongoid, 2.3 x 0.8 cm; petal narrowly lanceolate-spatulate, acute, 2.35 x 0.54 cm; labellum recurved, trilobed, 1.55 x 1 cm when spread out, with four very short ridges from the base and three more before the mid-lobe, side lobes erect, triangular, obtuse, mid-lobe recurved, ovate and rounded, 6.5 x 8 mm, its broad margin very warty; column straight, c. 8.5 mm long. Habitat: Collected from Mt. Dempo, South Sumatra at 1,400 m asl, flowering time: August. Distribution: South Sumatra (Mt. Dempo). Note: According to Smith (1920), this species differs from E. laurifolium and E. macropodum by the larger pseudobulb and leaves, laxer and less flower inflorescence, much larger flower and the lip with different calli, and very warty mid-lobe. The description was based on single specimen with unopened flower. Flower description was partly based on publication (Smith 1920).

39 23 Specimen examined: South Sumatra, Mt Dempo 1,400 m asl, August 9, 1916 E. Jacobson and Ajoeb 443 (BO). 4. Epigeneium elongatum (Blume) Summerh. (Plate 3) Desmotricum elongatum Blume, Bijdr. (1825) 552. Katherinea elongata (Blume) A.D. Hawkes, Lloydia, 19 (1956) 95. Epigeneium elongatum (Blume) Summerh. comb.nov., Kew Bull. (1957) 261. Epigeneium triflorum var. elongatum (Blume) J.B. Comber, Orchid of Java (1990) 245. Rhizome long, creeping, 3 6 mm in thickness, covered by 12 layered, imbricate, brown cataphylls, ovate to long ovate, acute to acuminate. Pseudobulb oblongoid, with four sharp angles, about cm apart on the rhizome, shiny olive green. Leaves two on each top of the pseudobulb, elliptic to long elliptic, obtuse, shallowly obliquely bilobe, petiole cm, coriaceous. Inflorescence from the top of pseudobulb between two leaves, 1 on each pseudobulb, peduncle cm long with 2 sterile bracts, rachis cm long and bring 2-8 flowers. Flower greenish yellow; dorsal sepal 1.25 x 0.5 cm, elliptic, obtuse; lateral sepal 1.3 x 0.65 cm, lanceolate, acute; petal 1.3 x 0.35 cm, long elliptic to near spathulate; labellum trilobed, 1.05 x 1 cm, with three calli on the disc, the center one is longest, the side lobes 0.7 x 0.5 cm, erect, the mid-lobe is rounded, with very short claw, the margin thickened. Distribution: Mt. Gede (West Java). Habitat: Primary mountain forest from 1,500 m asl, flowering time: September, October. Note: J.B Comber in Orchid of Java placed this species as variation of E. triflorum var. elongatum but he also suggested to elevated this variety into species rank. Base on Summerhayes (1957), Epigeneium elongatum were found in Java formerly known as Dendrobium elongatum. After observation it s agreed to separate this species from E. triflorum. There are some characters that need to be considered: vegetative parts which show great differences on the shape of pseudobulb which is ellipsoid with sharp angle, leaves mostly elliptic or long

40 24 elliptic. Flower smaller than Epigeneium triflorum var triflorum, dull yellow with dark red spot on the labellum. Apex of the mid-lobe is rounded and its margin thickened and this character very obvious on living specimen. Specimen examined: West Java, Mount Gede, M. Fleicher 38 (BO); Tjibeureum, Mt. Gede, 1,600 m asl, September 17, 1953, W. Meijer 1826 (BO). 5. Epigeneium exilifolium (Ames & Schweinf.) Garay & Romero (Plate 4) Dendrobium exilifolium Ames & Schweinf. In Contrib. Arnold Arb. 8 (1934) 32. Sarcopodium verruciferum (J.J. Sm.) Rolfe, Orch. Rev. 18 (1910) 239. var. pauciflorum Carr, J. Mal. Br. Roy. As. Soc. 11 (1933) 86. Epigeneium exilifolium (Ames & Schweinf.) Garay & Romero, comb.nov. Harv. Paper Bot. 4 (1999) 475. Epigeneium exilifolium (Ames & Schweinf.) J.B. Comber, comb. nov., Orchid of Sumatra (2001) 674. Type: North Sumatra, Walter N. & Catherine M. Bangham 1124 (AMES!, holo, photo image). Rhizome long, creeping, 2-3 mm in thickness, covered by brown cataphylls. Root fibrous, branching, flexuous, from lower part of pseudobulb. Pseudobulb ovoid-narrower ovoid, x cm, placed cm apart on rhizome. Leaves 2 on each pseudobulb, linear, acute, shallowly obliquely bilobes, cm x cm, 3 second major nerves on each side of the midrib. Inflorescence from top of pseudobulb, shorter than the leaf, racemose; peduncle thin, c cm, rachis c.2.1 cm, brings about 3 flowers. Flower with red green petal on outside and yellow inside, dorsal sepal elliptic to slightly lanceolate, 1.7 x 0.65 cm, obtuse, 7 nerved, lateral sepal ovate to lanceolate, but wider at obliquely base, 1.7 x 0.75 cm, obtuse, petal elliptic linear, 1.8 x 0.4 cm, obtuse; labellum trilobes, 1.38 x 1.12 cm when spread out, side lobes incurved, 1.13 cm long, mid-lobe smaller, with three short fleshy ribs; column 0.9 cm, with wing on both side, column foot about 0.59 cm. Distribution: Near Lake Toba, North Sumatra. Habitat: high elevation from 1,200-1,500 m asl, flowering time: September. Note: this species has linear leaf which seems similar to E. triflorum but acute and the inflorescence shorter. Observation based on single specimen and

41 25 photograph of type from Oakes Ames Herbarium. Previously this species known as D. exilifolium. Carr named this species as Epigeneium verruciferum var. pauciflorum. Comber probably missed the publication by Garay and Romero (1999) which has been transferred Dendrobium exilifolium to E. exilifolium and proposed the same new combination name in Flower description was partly based on publication. Specimen examined: North Sumatra, 4,100-4,500 ft.,walter N. and Catherine M. Bengham 1124 (AMES, Image); North West of Lake Toba c. 5,000 ft, September 1931, C.E. Carr 2025 (SING). 6. Epigeneium geminatum (Blume) Summerh. (Plate 5) Desmotrichum geminatum Blume, Bijdr. (1825) 552. Dendrobium geminatum (Blume) Lindl., Gen. & Sp. Orch. Pl. (1830) 77. Sarcopodium geminatum (Blume) Rolfe, Orch. Rev. 18 (1910) 238. Katherinea geminata (Blume) A.D. Hawkes, Lloydia, 19, (1956) 96. Epigeneium geminatum (Blume) Summerh. comb.nov., Kew Bull (1957) 262; Comber, Orchid of Java (1990) 244. Type: Java, collector unknown, s.n. (L!, holo, photo image). Rhizome long, creeping, 3-5 mm in thickness, covered by up to 7-9 imbricate brown cataphyll, subulate, acute, partly persistent. Pseudobulb ovoid at the lower part of the rhizome and cylindrical on the upper part, x 0.5 cm cm with distinct angle, sulcate, shiny, placed cm apart on rhizome. Leaves ovoid elliptic, sometimes lanceolate to rather linear, x cm, apex shallow obliquely bilobed, coriaceous, dark green sometimes with purplish line at the margin, petiole 2 6 mm. Inflorescence from new growth pseudobulb, mostly 2 inflorescences on each top of pseudobulb, rarely 1 or 3; peduncle thin, cm long; rachis cm, bring 5-9 flowers. Flower small compare to other species, not open wide, light yellow or slightly peach; dorsal sepal lanceolate, 1.3 x 0.3 cm, sub acute, 5 nerved; lateral sepal lanceolate, 1.4 x 4 cm, acute, wider at base, 5 nerved; petal linear, 1.2 x 0.2 cm, labellum c. 1 cm long, irregular calli at the base, another three between the side

42 26 lobes and almost equal in size, sub acute, ovary and pedicel c. 2 cm; column 4 mm with 2 mm column foot. Distribution: Mt. Salak and Mt. Halimun (West Java), Siberut Island (West Sumatra), Mt. Kinabalu, Sarawak, Central Kalimantan (Borneo), Pahang, Taiping Hills (Peninsular Malaysia). Habitat: Lower mountain forest from 700 1,100 m above sea level but sometime found in lower altitude. Flowering time: February to March, June to August, and November. Note: Comber (2001) did not include E. geminatum as species from Sumatra. Base on this study there was a specimen collected from Siberut island, outside of Sumatra mainland on a lower altitude and has the same character. Specimen examined: West Java, Mt. Salak 800 1,300 m asl, June 4, 1941, Bloemberger (BO); Halimun Salak National Park, Cikaniki 1,000 m asl, July , Asep Sadili 1030 (BO); Sumatera, Siberut Mentawai, Gunung Simapipit Sikabaluan residence 300 m asl, May , J.J. Afriastini 2720 (BO); Central Kalimantan, Samba 340 m asl, February , Jarvi, J.K Ruskandi A 5939 (BO); Sarawak, Karangas Secondary Forest 3,500 ft asl, November , Paul Chain (SING); Lubuk Antu District, 2 nd Div. 3,200 ft asl, March 17, 1974, Paul Chain (SING); Pahang, Mt. Tahan 3,000-4,000 ft asl, August 28, 1928, R.E. Holtum (BO); Pahang, Fraser Hill 4,000 ft asl, November 1930, C.E. Carr (SING); Malay Peninsula, Taiping Hills, 1902 (SING). 7. Epigeneium gracilipes (Burkill) Garay & G. Romero Dendrobium gracilipes Burkill, J. Str. Br. Asiat. Soc. 79 (1918) 45. Epigeneium gracilipes (Burkill) Garay & Romero, comb.nov., Harv. Paper Bot. 4 (1999) 475. Epigeneium gracilipes (Burkill) J.B. Comber, comb. nov. Orchid of Sumatra, p (2001). Type: Riau Archipelago, cultivated in Singapore, K. Saheb s.n. (SING, holo n.v). Rhizome long, quite thick, bearing pseudobulb c.4 cm apart. Pseudobulb obliquely ovoid, shiny green, initially with a sheath which drops early, c. 3 x 1.4 cm in size. Leaves two on each pseudobulb, narrowly elliptic, sessile, apex

43 27 bilobe, the lobe obtuse, to 16 x 2.5 cm. Inflorescence from top of pseudobulb, two flowered, longer than the leaves. Flowers mostly ivory white; dorsal sepal placed at right angle to the ovary, acute; lateral sepal 1.5 x 0.5 cm, decurrent on the column foot forming a rectangular mentum, 5 mm in length, curved above this, acute; petal linear, 1.5 x 0.15 cm, acute; labellum trilobes, side lobes erect, the upper margin straight, spotted purple on the inside and with purple upper margin and apices, the lower margin saffron yellow, the mid-lobe broad basally, more or less acuminate apically, its margins and apex ivory white with some yellow toward the top, with three keels; column ivory white flushed red near the base. Distribution: Riau Archipelago (Riau). Habitat: lowland forest from 300 m asl. Note: There was no specimen available for this species, the description based on the publication (Garay and Romero 1999, Comber 2001). Garay and Romero transferred this species to Epigeneium in 1999, but it seems this publication was overlooked by Comber and in 2001 he published the similar new combination with previous ones. 8. Epigeneium laurifolium (Kraenzl.) Summerh. Sarcopodium laurifolium Kranzl. In Engl., Pflanzenr. Orch. Mon. Dendr. 1: 328 (1910). Dendrobium laurifolium (Kranzl.) J.J. Sm., Enum. Orch. Sumatra (1933) 286. Katherinea laurifolia (Kranzl.) A.D. Hawkes, Lloydia, 19 (1956) 96. Epigeneium laurifolium (Kranzl.) Summerh., Kew Bull. (1957) 263; Comber, Orchid of Sumatra (2001) 675. Type: West Sumatra, Mt. Singgalang, July 1901, Odoardo Beccarii (FL, holo, n.v). Rhizome long, creeping, pseudobulb placed 4-6 cm apart. Pseudobulb c. 4 x 2 cm, ovoid, quadrangular, when dried orange and shiny. Leaves c. 8 x 3 cm, oblong, incurved apically, more or less bilobe. Inflorescence well surpassing the leaves, 25 cm or longer, the rachis with up to 25 flowers; bracts triangular, acute 4-5 mm long, ovary and pedicel 3 cm. Flower rather small for the genus; dorsal

44 28 sepal 1.3 cm long, oblong, obtuse; lateral sepal 1.3 x cm, triangular, oblong, forming a rectangular mentum with their bases; petal strap shaped, same length as sepals; labellum trilobes, side lobes oblong, rounded in front, 6-7 mm long, mid-lobe transversely oblong to rhomboid, acute, 3-4 mm long, three keels from the base of the labellum, the side one larger than the center ones, another three keels in the middle of the labellum, the center one extending to the base of the labellum; column slender. Distribution: Mt. Singgalang, West Sumatra. Habitat: mountain forest, found at 1,900 m asl. Note: There was no specimen of this species found during this study. The type is maintained in Florence Herbarium (Fl) and inaccessible through the website. The description was based on Kraenzlin (1910) and Comber (2001). 9. Epigeneium longibulbon Destri sp. nov. (Plate 6-7) Pseudobulbus longissimus, strumolusus inferiora et cylindrical versus apicem, sine angulus, folia 2-4, differt. Type: West Sumatra, Mt. Talang 2,230-2,500 m asl, November 14, 1988, H. Nagamasu 3414 (BO!, holo; L, iso). Rhizome branching, c. 3 4 mm in thickness, covered by brown imbricate cataphyll, cm long, ovate, acute to acuminate. Root fibrous, branching, wrinkled on surface, c mm in thickness, bears from lower part of pseudobulb and also on the rhizome but less dense. Pseudobulb long c x cm, swollen at the base and cylindrical toward the apex, blunt, shiny yellow, arrange cm apart along the rhizome. Leaves 2 4 on each pseudobulb, elliptic to rather lanceolate, x cm, acute, obliquely bilobe with 4-5 major secondary nerved on each side, petiole 4 8 mm, shiny, and coriaceous. Inflorescence long, peduncle c. 17 cm with 2 sterile bracts, 8.5 mm, tubular, acute, translucent brown; rachis is shorter c. 7.7 cm and bring up to 12 flowers. Flower white, labellum with yellow stripe and violet patches; dorsal sepal 2.3 x 0.55 cm, long oval, acuminate, 7 nerved; lateral sepal almost same size but wider near to the base 2.2 x 0.65 cm, lanceolate, 7 nerved; petal 2.2 x 0.55 cm, long oval, acuminate, 6 nerves; labellum trilobes, 1.3 x 1.2 cm, side

45 29 lobe erect, blunt, 9 x 4.5 mm, the disc of the labellum with calli arrange parallels, undulating from the base to near the middle of the labellum, the three other calli near the base of the mid-lobe, almost with the same length, apex rounded, acuminate, column 8 mm with very short stelidia on each side, column foot 4 mm. Distribution: Mt Talang (West Sumatra), Mt. Kerinci (Jambi). Habitat: on mountain forest up to 2,400 m asl, flowering time: March and November. Note: This species is only represented by two sheets herbarium specimen and some living specimen in Cibodas Botanic Garden. The major differences compare to other genus is the longest pseudobulb and the leaves number on each pseudobulb from 2-4 (observed in living specimen, Wiguna Rahman 353). Flower structure is similar with other species in this genus; having white flower similar to E. longipes but with up to 11 flowers. Ornament on the labellum also show dissimilarity; the calli at base of labellum similar to E. cymbidioides, the other three calli between side lobes has the same length. Specimen was collected from Mt. Talang, West Sumatra and Mt. Kerinci (Jambi). Specimen examined: West Sumatra, Mount Talang 2,230-2,500 m asl, November 14, 1988, H. Nagamasu 3414, flowering specimen (BO); Solok residence, along trail from Desa Air dingin to Mt. Rasam summit at 2,100-2,400 m asl, March 5, 1996, H. Okada, H. Nagamasu, T. Fukuhara, Y. Mori (BO). 10. Epigeneium longipes (Hook.f.) Summerh. (Plate 8) Dendrobium longipes Hook.f., Fl. Brit. Ind. 5 (1890) 713. Sarcopodium longipes (Hook.f.) Rolfe, Orch. Rev. 18 (1910) 238. Katherinea longipes (Hook.f.) A.D. Hawkes, Lloydia, 19 (1956) 96. Epigeneium longipes (Hook.f.) Summerh. comb.nov., Kew Bull. (1957) 263. Comber, Orchid of Sumatra (2001) 680. Rhizome creeping, 3-5 mm in thickness, covered by c. 5 layer brown cataphylls, ovate, acuminate. Root thicker compare to other species, mm, fibrous, terete, long, branching with wrinkle surface. Pseudobulb ovoid-narrower

46 30 ovoid, 1-4 cm x 2-10 mm, placed very close from 0.5 to 7.3 cm apart on rhizome. Leaves two on each pseudobulb, from almost orbicular or elliptic to long elliptic, rather obtuse, obliquely bilobes, petiole cm. Inflorescence long surpass the leaf, peduncle cm, much longer than rachis, with 2 sterile bracts, rachis cm, bears 2-6 flowers. Flower white, open widely; sepal lanceolate 2.3 x 0.45 cm; petal the same shape and length, but much narrower; labellum white and yellow, 1.6 cm long with several calli between side lobes and three larger one near to the apical end, side lobes erect, yellow with red stripes, obtuse, the lower margin curved, mid-lobe triangular, acute; column arching, white above, yellow with fine red stripes under, extending just as far as side lobes. Distribution: Mt. Sago (West Sumatra), Cameron Highland, Pahang, Mt. Tahan, (Peninsular Malaysia). Habitat: mountain forest orchid from 1,700 m asl, flowering time: February, July, and September. Note: the most conspicuous character of this species is the peduncle three times longer compare to the rachis. Leaves were also with wide variety in shape and size; from almost orbicular to elliptic and sometime longer. During this study there was no specimen examined from Sumatra but Comber (2001) described this species from Mt. Sago, West Sumatra. Specimen examined: Malay Peninsula, Pahang, H.N. Ridley (SING); Pahang, HN Ridley (SING); Mt. Tahan, July 1923, CB Kloss (SING); Mt Tahan, February 1921, E. Seimund 221 (SING); Cameron Highland, September 1956, H.M. Burkill 860 (SING); Cameron Highland, April 1930, 1,800 m asl, R.E Holtum and M.R. Henderson (SING); Batang Padang, L. Wray Jr (SING). 11. Epigeneium macropodum (Hook.f.) Summerh. (Plate 9) Dendrobium macropodum Hook.f., Fl. Brit. Ind. 5 (1890) 713. Sarcopodium macropodum (Hook.f.) Rolfe, Orch. Rev. 18 (1910) 238. Katherinea macropoda (Hook.f.) A.D. Hawkes, Lloydia, 19 (1956) 96. Epigeneium macropodum(hook.f.) Summerh., Kew Bull. (1957) 263; Comber, Orchid of Sumatra (2001) 681.

47 31 Rhizome long, creeping, branching mm in thickness, covered by brown cataphylls. Root terete, fibrous, branching, wrinkle on surface, come from lower part of the pseudobulb. The pseudobulb ovoid narrower ovoid cm x 8-17 mm, sharp angle, sulcate, shiny, cm in distances. Leaves two on each pseudobulb, long elliptic x cm, acute, obliquely bilobed, 3 second major nerved on each side, coriaceous, petiole 3-5 mm. Inflorescence single on top of the pseudobulb, much longer than the leaf; peduncle cm with 2-3 sterile bract ovate-triangular, acute; rachis cm, with up to 15 flowers, placed 5-19 mm apart, ovary pedicel cm, shorter toward the top. Flower reddish brown when first open and becoming greenish brown later, about 3 cm broad; sepal narrowly lanceolate, acute, 1.7 cm x 5 mm, the lateral one a little broader forming a short rounded mentum with their bases; petal somewhat shorter and narrower; labellum trilobes, the side lobes erect and rounded, the mid lobe almost circular at the apex, obtuse, some irregular thickened ridge (calli) from the base of the mid-lobe, and another 3 near the apex. Habitat: mountain forest from 1,500 m asl, flowering on April, July, and August. Distribution: Padang Panjang (West Sumatra), Cameron Highland, Pahang, Mt. Tahan (Peninsular Malaysia). Note: Summerhayes (1957) mentioned that this orchid also found in Java, but during this observation there was no specimen collected from this island. The specimen examined from Sumatra was unavailable but Comber (2001) described this species from around Padang Panjang, West Sumatra. The specimen from Sumatra also held in Leiden but locality is unknown. Specimen examined: Malay Peninsula, Pahang, Mt. Batu Brinchang, Cameron Highland 6,700 ft asl, April 17, 1930, M.R. Herderson (SING); Cameron Highland, W. Steele (SING); Pahang, Mt. Tahan, July 1911, H.N. Ridley (SING); Pahang, Mt. Tahan 5,000 ft asl, August 29, 1928 (SING).

48 Epigeneium pulchellum (Ridl.) Summerh. (Plate 10) Sarcopodium pulchellum Ridl., Journ. Fed. Malay States Mus. 8 (1917) 93. Dendrobium brevibulbum J.J. Sm., Bull. Jard.Bot.Buitenz. 3 (1928) 62. Katherinea pulchella (Ridl.) A.D. Hawkes, Lloydia, 19 (1956) 97. Epigeneium pulchellum (Ridl.) Summerh. comb.nov., Kew Bull. (1957) 264; Comber, Orchid of Sumatra (2001) 675. Rhizome long, creeping c mm in thickness, covered by c.10 layered cataphylls, partially persistent, triangular, and acute. Root mostly found on the lower part of the pseudobulbs along the rhizome, fibrous, branches. Pseudobulb small, globose to ovoid, sometimes longest, c x cm, placed cm apart on the rhizome. Leaves 2 on each top of pseudobulb, narrower ovate to elliptic, c x cm, apex shallowly lobed, unequal, obtuse, petiole mm. Inflorescence short, peduncle and rachis c. 1 cm, with 1-2 flower. Flower white, quite large in proportion to the plant size, 3 x 2.7 cm; dorsal sepal 1.55 cm x 6 mm, elliptic, acute; lateral sepal 1.4 cm x 7.5 mm, elliptic but wider at base, acute; petal 1.5 cm x 6 mm, elliptic, acute; labellum trilobed, 1.3x1.25 cm, side lobes erect, mid-lobe with three longitudinal keels, the middle ones thinner and elongate to the base, the mid-lobe ovate, obtuse; column 9 mm, column foot 4 mm, with short wings. Distribution: Tanah Gayo (Aceh), Mt. Kerinci (Jambi). Habitat: Mountain forest, in mount Kerinci it was collected from 1,900 m asl. Some herbarium specimen were collected from Aceh and North Sumatra from above 2,000 m asl. Flowering time: January, February, August, and December. Note: So far Epigeneium pulchellum is the smallest Epigeneium found in Sumatra. In absent of flower, this species can be distinguished from other species is from its small size. Another difference is, it has only single flower on each inflorescence. At a glance E. pulchellum seem to be similar with E. longirepens but the last one only has single leaf on each pseudobulb and distribution area is restricted to Borneo (Endemic), meanwhile E. pulchellum endemic to Sumatra. Specimen examined: Aceh, Gajolanden 2,300-2,700 m asl, January 1937, C.G.G.J. van Steenis 8344 (BO); Gajolanden 2,500-3,000 m asl, C.G.G.J. van Steenis 9026 (BO); Gajolanden 3,300-3,500 m asl, C.G.G.J. van Steenis 8582

49 33 (BO); Mt. Leuser 3,200-3,400 m asl, W.J.J.O de Wilde and B.E.E. de Wilde- Duyfjes (BO). 13. Epigeneium quinquecallosum (J.J. Sm.) Summerh. Dendrobium quinquecallosum J.J. Sm., Bull. Jard. Bot. Buitenz. 3 (1920) 68. Epigeneium quinquecallosum (J.J. Sm.) Summerh. comb. nov., Kew Bull. (1957) 264; Comber, Orchid of Sumatra (2001) 674. Rhizome creeping, branching, mm in thickness, covered by c. 5 layered cataphylls, imbricate, elliptic, and acute. Root terete, fibrous, branching. Pseudobulb ovoid, x 0.8 cm, with deep groove on the side, angular, placed apart on rhizome. Leaves two on each pseudobulb, elliptic, x cm obtuse, shallowly obliquely bilobe, with 4 second major vein, coriaceous. Inflorescence between two leaves on top of pseudobulb, peduncle with 2 sterile bracts, tubular, acute, rachis c cm, bring about 4 flowers. Flower white, dorsal sepal lanceolate, narrowly obtuse, apiculate, 2.75 x 0.77 cm; lateral sepal obliquely, lanceolate, acute, 2.8 x 8 cm; petal obliquely lanceolate, acute 2.9 x 0.65 cm; labellum trilobe, curved, 1.7 cm long, with four small calli near the base and five larger one placed between the side lobe just below the mid-lobe, the middle one of this five is longer, side lobes erect, reaching over the column, concave, triangular obtuse, the mid-lobe recurved in the middle, ovate and acuminate with seven wavy nerves, 8.5 x 0.7 cm; column slightly curved, concave below, 9 mm long. Habitat: mountain forest at around 1,700 m asl, flowering on November. Distribution: Laras Talang (West Sumatra), Mt. Dempo (South Sumatra). Note: five calli between the side lobes is the key character to distinguish this species from others within this genus. It is only represented by single photo of the specimen deposit in BO. The specimen also keeps in Leiden but the image is unavailable. Flower description base on publication (Smith 1920). Specimen examined: Sumatra, Laras Talang 1,750 m asl, November 11, 1918, H.A.B. Bunnemeijer 5437 (BO).

50 Epigeneium sikijangensis Destri sp. nov. (Plate 11-12) Sepalis et petala flaveolus cum rubigunosus puncticulosus, differt. Type: Sumatra, Jambi, Mt. Kerinci, 1000 m asl, June 2008, Wiguna Rahman 107 (BO!, holo). Rhizome long, creeping, 5-6 mm in thickness, covered by layered imbricate cataphylls. Cataphylls consist of c. 15 layers, from c.0.5 to 6 cm long, light green with longitudinal purple streak when immature and gradually turn to reddish brown. Root from the base of pseudobulb, c.1.5 mm in thickness, branching, wrinkle on surface. Pseudobulb ovoid to long ovoid, x cm in size with 3 or 4 sharp angle, purplish sometime greenish, shiny. Leaves two on each pseudobulb, conduplicate, oblong, x cm in size, obtuse, shallowly obliquely bilobe, not shiny, 3-4 second major nerves on each side of the mid-rib. Inflorescence longer than the leaves; peduncle 11.4 cm long with 2 sterile bracts, tubular or triangular, acute, mm, purplish translucent; rachis 9.8 cm long, bears up to 5 flowers which placed cm apart. Flower c. 3 cm in wide, sepal and petal greenish yellow with fine red spot except a little part at the base; dorsal sepal elliptic, acute, 1.85 x 0.65 cm; lateral sepal elliptic rather lanceolate, oblique and wider at base, 1.8 x 0.7 cm; petal narrower elliptic, acute, 1.9 x 0.5 cm; labellum trilobed 1.25 cm long, side lobes erect white with red streaks on inside surface, three keels between the side lobes, the middle one elongate from the base to near the apex, mid-lobe deltoid, obtuse; column yellow with dark red streak. Distribution: Barisan Range (West Sumatra), Mt. Kerinci (Jambi). Habitat: lower mountain forest on 1,000 m asl, flowering on August October. Note: among the species observed this is the only one has purplish pseudobulb and flower with fine red spots. This species represented by two living plant specimens cultivated in Cibodas Botanic Garden. This species was collected from mount Kerinci, Sumatra and it was quite exclusive since there was none of their relative found in that location so that this species could be new one instead of natural hybrid. There was a sterile specimen in BO, quite similar to this living collection, collected from West Sumatra on 1,000 m asl, sepal and petal with fine brownish spots.

51 35 Specimen examined: West Sumatra, Barisan Range, Air Sirah, above Padang, 1,000 m asl, March 6, 1974, Ed F. de Vogel 2835, sterile (BO); Jambi, Mt. Kerinci 1,000 m asl, June 2008, Wiguna Rahman 107 (BO). 15. Epigeneium triflorum (Blume.) Summerh. (Plate 13-14) Desmotrichum triflorum Blume, Bijdr. (1825) 551. Dendrobium triflorum (Blume) Lindl., Gen. Spec. Orch. Pl. (1830) 77. Sarcopodium triflorum (Blume) Rolfe, Orch. Rev. 18 (1910) 239. Epigeneium triflorum (J.J. Sm.) Summerh. comb. nov., Kew Bull. (1957) 265; Comber, Orchid of Java (1990) Rhizome terete, creeping, branching, tough, mm in thickness, covered by brown cataphylls, about 12 layers, elliptic, acute, lower ones is the smallest. Root terete, fibrous, mm in thickness, branching, mostly at lower part of rhizome. Pseudobulb almost globose ovoid or long ovoid, cm x cm, yellow, shiny, bears 2 or 3 leaves. Leaves linear to strap shape, obtuse, obliquely bilobed, petiole cm. Inflorescence single come out between two leaves, long peduncle, 2 bracts sterile, tubular, acute, rachis almost the same, bears 3 many flowers. Flower white flushing with light yellow; dorsal sepal long elliptic, acute; lateral sepal lanceolate, acute, broader at the base; petal rather spathulate, acute; labellum trilobe, side lobe erect, apex of midlobe rhomboid, acute, the margin curved backward; column erect, with short stelidia. Two varieties within this species: Epigeneium triflorum var. orientale (J.J. Sm.) J.B. Comber, Orchid of Java (1990) 246. Dendrobium elongatum Lindl. var. orientale J.J. Sm., Bull. Dep. Agr. Indes. Neerl. 43 (1910) 52. Type: East Java, Lumajang, culta. (BO!, holo; L, iso n.v) Leaves linear to strap shaped, x cm, obtuse, obliquely bilobed, petiole 5-7 mm, shiny, coriaceous. Inflorescence single between two leaves, peduncle c cm, 2 bracts sterile, tubular, acute, rachis c. 9.3, bring more than 5 flowers. Flower white flushing with light yellow; dorsal sepal long

52 36 elliptic, 2.7 x 0.6 cm, acute, 7 nerved; lateral sepal long elliptic, 2.6 x 0.7 cm, acute, broader at oblique base; petal spatulate, 2.8 x 0.4 cm, acute, labellum trilobes, 1.45x1.2 cm, side lobes erect 9 x 5 mm, mid-lobe with 2 calli near the apex, apex rhomboid, acute, the margin curved backward; column erect, 9 mm, column foot 4.5 mm. Epigeneium triflorum var. triflorum (J.J. Sm.) J.B. Comber, Orchid of Java (1990) 245. Leaves 2 or 3 on each top of pseudobulb, linear, slightly obtuse, obliquely bilobe, shiny. Inflorescence from between leaves, peduncle 10.3 cm, thin, with 2 sterile bracts, rachis 4.9 cm, brings more than 3 flowers. Flower white; dorsal sepal lanceolate, acute, 2.05 x 0.6 cm, 5 nerved; lateral sepal lanceolate, acute, wider at base, 2.15 x 0.7 cm, 5 nerved; petal rather spathulate, 2.1 x 0.35 cm, acute; labellum trilobe, 1.65 x 1.15 cm, with erect side lobe 8.5 x 4 mm, triangular, with purple streaks on inner surface; mid-lobe with two ridge from the base of toward the center, three knobs near the apex, the middle one is the longest and placed nearer to the base, apex of mid-lobe rhomboid, acute, yellow in the middle, white margin, and gradually curved backward which make this part seem narrow; column 9 mm, white, inner suface (front), light yellow with short red streak, column foot c. 5 mm, stelidia very short. Distribution: Mt. Gede-Pangrango, Mt. Halimun, Mt. Salak, Mt. Tangkuban Prahu (West Java); Mt Slamet (Central Java); Lumajang and Pasuruan (East Java). Habitat: Mountain forest at Montana zone from 1,500 m asl, on higher branches which is exposed to sunlight. Flowering time: March, April, and June to October. Note: some variation was found on the vegetative parts of this species. Leaves vary from linear to strap shape, color from light green to dark green, surface waving or flat, and the leaf number 2 to 3 on each pseudobulb. Specimens also show the consistency of combination between 2 and 3 leaves in certain individual plant. Flower size sometimes differs among individual plant, usually it has the same size with E. cymbidioides but some are smaller. Another variation is observed for specimens with deep curved tepal backward so that its appearance slightly different compared to E. cymbidioides, this species has white tepal,

53 37 mostly acute and less fleshy while E. cymbidioides has dull yellow, obtuse and fleshy tepal. According to Comber (1990) this species endemic to Java but there are some living collection in Bali Botanic Garden but locality is unknown. Specimen examined: East Java, Lumajang, culta (BO, holotype); Lumajang, August 1954, Connell E. s.n (BO); Pasuruan, June 1946, C.G.G.J. van Steenis 7385 (BO); West Java, Preanger, 1,500 m asl, June 1905, R.C. Backer v/d Brink 182 (BO); Mt. Gede, 1,450 m asl, March 1972, Ed de Vogel 1349 (BO); Mt. Kamojang 1,500 m asl, Van Leuween Rejnvaran 2425 (BO); Central Java, Baturraden, 800 m asl, C (BO). 16. Epigeneium wichersii (Schltr.) Summerh. (Plate 15). Dendrobium wichersii Schltr., Fedde, Repert. Spec. Nov. 8 (1910) 506. Katherinea wichersii (Schltr.) A.D. Hawkes, Lloydia, 19 (1956) 98. Epigeneium wichersii (Schltr.) Summerh., comb. nov., Kew Bull. (1957) 266; Comber, Orchid of Sumatra (2001) 678. Rhizome creeping, branching, mm, with brown cataphylls. Root fibrous, c. 1 mm, branching, mostly from lower part of pseudobulb but sometime also spread along the rhizome. Pseudobulb ovoid long ovoid, sometime elliptic, x cm, quadrangular, placed cm apart. Leaves two on each pseudobulb, oblong, 6-16 x cm, obtuse, shortly obliquely bilobe, petiole 2-3 mm, shiny, coriaceous. Inflorescence single on top of pseudobulb, peduncle 6 cm, to sterile bract; rachis c. 9.7 cm, bring c.12 flowers. Flower medium size; dorsal sepal oblong, obtuse 1.4 cm long; lateral sepal obliquely ovate, obtuse, form a short mentun at the base, petal obliquely strap shape, almost the same length with sepal; labellum trilobe, 1.5 cm long, side lobes erect, triangular, obtuse, mid-lobe smaller, semi circular, three ridge from the base of mid-lobe toward the apex, the middle one is the smallest. Distribution: Tanah Gayo (Aceh), Mt Singgalang (West Sumatra), Mt. Kerinci (Jambi). Habitat: Lower mountain forest 800 to 1,700 m asl, flowering time: March.

54 38 Note: There was no specimen available and also incomplete material for this species which make it difficult to observed more detail on its character. Flower description base on publication. Specimen examined: Sumatra, Aceh, Gajolanden 1,100-1,250 m asl, March 1937, C.G.G.J. van Steenis 9940 (BO); Mt. Kerinci 1,700 m asl, March 1920, H.A.B. Bunnenmeijer 8903 (BO). 17. Epigeneium zebrinum (J.J. Sm.) Summerh. (Plate 16) Dendrobium zebrinum J.J. Sm., Icon. Bogor. 2 (1903) 72. D. sulphuratum Ridl., Journ. Roy. As. Soc. Straits, 49 (1908) 28. Sarcopodium sulphuratum (Ridl.) Rolfe, Orch. Rev. 18, 239 (1910). S. zebrinum (J.J. Sm.) Kraenzl., Engl. Pflanzenr. Orch. Dendrob. 1 (1910) 324. Dendrobium citrino-castaneum Burkill, Gard. Bull. Straits Settlem. 2 (1923) 12. Sarcopodiuum citrino-castaneum (Burkill) Ridl., Fl. Malay Penin. 4 (1924) 29. Katherinea citrino-castaneum (Burkill) A.D. Hawkes, Lloydia, 19 (1956) 95. Epigeneium zebrinum (J.J. Sm.) Summerh. comb. nov., Kew Bull. (1957) 266; Comber, Orchid of Sumatra (2001) 682. Type: Malay Peninsula, September 1921 (SING!, holo, photo image, L, iso). Rhizome stout, creeping, 4-5 mm in thickness, covered by c layered cataphylls, acute, ovate, imbricate. Root fibrous, terete mostly from lower part of pseudobulb. Pseudobulb ovoid, sulcate, 4 sharp angles, x cm, placed cm on rhizome. Leaves two on each top of psedobulb, oblong, x cm, acute, petiole 5-6 mm, coriaceous. Inflorescence two from between two leaves of immature pseudobulb, peduncle c. 9 cm long, rachis c.2.9 cm with 4-9 flowers. Flower about 3.1 cm broad with c. 2.7 cm length of ovary and pedicels, sepals and petals yellow; sepal lanceolate, x 0.65 cm; petal acute, 1.7 x 0.5 cm; labellum trilobes 1.25 x 0.8 cm when spread out, side lobes erect, obtuse, the margin little erose, the mid-lobe long triangular, acute, more or less concave, thickly fleshy, yellow on lower surface and dark reddish brown above with yellow margin, three parelel ribs toward the apex of mid-lobe, the middle one is longest; column yellow, 6.5 x 3.5 mm. Distribution: Bayung Lencir (South Sumatra), Nanga Tekungai (West Borneo), and Pahang (Peninsular Malaysia).

55 39 Habitat: lowland forest at about 350 m asl, flowering time: July. Note: This species share the same character with E. geminatum which mostly has 2 inflorescences on each pseudobulb but the appearance quite different. Epigeneium zebrinum is much larger with stout rhizome while E. geminatum is smaller with smaller flower. Habitat range is also different, E. zebrinum only found on lower altitude meanwhile Epigeneium geminatum has wider habitat range from 350 1,000 m asl. There was no specimen from Sumatra but Comber described this species from Bayung Lencir, South Sumatra. Isotype which held in Leiden was cultivated in Bogor Botanic Garden but the locality is unknown. Flower description base on publications. Specimen examined: Malay Peninsula, Ulu Brang 350 m asl, L. Moysey (SING); Pahang, 500 m asl, November 1929, C.E. Carr 259 (SING); West Borneo, Nanga Tekungai, 80 m asl, Hans Winkler 1266 (BO).

56 40 CONCLUSION The study indicated that there are five species Epigeneium in Java and 14 species in Sumatra. Epigeneium geminatum is a new record from Sumatra and two species were proposed as new spesies: Epigeneium longibulbon and E. sikijangensis. Based on distribution area analysis, E. geminatum has the widest distribution from Java to Peninsular Malaysia but in Java this species only found in two location (Mt. Salak and Mt. Halimun), meanwhile E. triflorum, E. arjunoense so far only found in Java. In Sumatra E. dempoense, E. exilifolium, E. gracilipes, E. laurifolium, E. longibulbon, E. pulchellum, E. quinquecallosum, E. sikijangensis, and E. wichersii, were endemic. More comprehensive exploration and taxonomic study on this genus is needed to find new species especially from Sumatra.

57 41 REFERENCES Comber JB Orchid of Java. London. Bentham-Moxon Trust. The Royal Botanic Gardens Kew. Comber JB Orchid of Sumatra. London. The Royal Botanic Gardens, Kew. de Vogel EF in de Vogel EF, editor. Guidelines for the Preparation of Revision. Manual of Herbarium Taxonomy Theory and Practice. Jakarta. UNESCO.p. 76. Dressler RL Phylogeny and Classification of the Orchids Family. Portland, Oregon. Dioscorides Press. Dressler RL, Dodson CH Classification and Phylogeny in the Orchidaceae. Annals of The Missouri Botanical Garden 47: Clement MA Molecular Phylognetic systematic in the Dendrobiinae (Orchidaceae), with emphasis on Dendrobium section Pedilonum in Everett J, Wilson PG, editors Proceeding of the 5 th International Flora Malesiana Symposium; May Telopea 10 (1): Gagnepain F Huit genres nouveaux d Orchidees Indochinoises. Bull. du Mus. Nat. Paris 2 (4): Garay LA, Romero GA Schedulae Orchidum II. Harv. Paper Bot. 4: Harris JG, Haris MW Plant Identification Terminology, an Illustrated Glossary. Utah, Spring Lake Publishing. Hawkes AD Katherinea, a New Name for the Orchid Genus Sarcopodium. Lloyidia 19: Kraenzlin F Orchidaceae-Monandrae-Dendrobininae, part 1 in Engler, A, editor. Das Pflanzenr. 4: Lindley J Sarcopodium lobbii (alias Bulbophyllum lobbii Lindley). Paxton s Flower Garden 1: Rifai MA Sendi-sendi Botani Sistematika. Lembaga Ilmu Pengetahuan Indonesia (LIPI). Rugayah, Wijaya EA, Pratiwi, editor Pedoman Pengumpulan Data Keanekaragaman Flora. Bogor. Puslit Biologi LIPI. Seidenfaden G, Wood JJ The Orchids of Peninsular Malaysia and Singapore (A Revision of R.E. Holtum: Orchids of Malaya.). Fredensborg, Denmark. Olsen & Olsen. Smith JJ Dendrobium (sec. Sarcopodium) dempoense J.J. S. n. sp. Bull. Jard. Bot. Buitenz. 3: Smith JJ Dendrobium (sec. Sarcopodium) quinquecallosum J.J. S. n. sp. Bull. Jard. Bot. Buitenz 3: Summerhayes VS Notes on Asiatic Orchids: II. Kew Bull. :

58 42 Whitten T, Damanik SJ, Jazanul A Nazarudin S The Ecology of Sumatra. Periplus Edition. Wood JJ Epigeneium cacuminis Orchidaceae. Curtis Botanical Magazine 21: Wood JJ Epigeneium triflorum var. orientale Orchidaceae. Curtis Botanical Magazine 13: Wood JJ, Beaman RS, Beaman JH The Plants of Mount Kinabalu, Vol. 2: Orchids. London. Royal Botanic Gardens Kew. Wood JJ, Comber JB A new species of Epigeneium from East Java. Orchid Review 104: Index Kewensis. London. Royal Botanic Garden Kew.

59 Plate 1. Epigeneium cymbidioides (Irawati 910). 43

60 Plate 2. Holotype of Epigeneium dempoense (E. Jacobson and Ajoeb 443) 44

61 Plate 3. Epigeneium elongatum (M. Fleicher 38) 45

62 Plate 4.Holotype of Epigeneium exilifolium (Walter.N. & Catherine M. Bangham 1124, from AMES Orchid Herbarium). 46

63 Plate 5. Epigeneium geminatum from Sumatra (J.J. Afriastini 2720) 47

64 Plate 6. Holotype of Epigeneium longibulbon (H. Nagamasu 3414) 48

65 49 1 cm 1 cm A A C C 1 cm 0.5 cm B D Plate 7. Flower parts of E. longibulbon: A. dorsal sepal, B. lateral sepal, C. petal, D. labellum (H. Nagamasu 3414)

66 Plate 8. Epigeneium longipes (R.E Holtum and M.R. Henderson 23398) 50

67 Plate 9. Epigeneium macropodum (H.N Ridley 16134) 51

68 Plate 10. Epigeneium pulchellum (C.G.G.J. Van Steenis 9234) 52

69 Plate 11. Holotype of Epigeneium sikijangensis (Wiguna Rahman 107) 53

70 cm 0.5 cm A C 0.5 cm 0.5 cm B D Plate 12. Flower parts of Epigeneium sikijangensis: A. dorsal sepal, B. lateral sepal, C. petal, D. labellum (Wiguna Rahman 107)

71 Plate 13. Holotype of Epigeneium triflorum var. orientale 55

72 Plate 14. Epigeneium triflorum var. triflorum, J.J. Smith s.n 56

73 Plate 15. Epigeneium wichersii (C.G.G.J. van Steenis 9164) 57

74 Plate 16. Holotype of Epigeneium zebrinum (from Herbarium of Singapore Botanic Garden) 58

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