REINWARDTIA A JOURNAL ON TAXONOMIC BOTANY, PLANT SOCIOLOGY AND ECOLOGY Vol. 14(1): 1 - 2 4 8 , December 23, 2014 Chief Editor Kartini Kramadibrata (Mycologist, Herbarium Bogoriense, Indonesia) Editors Dedy Darnaedi (Taxonomist, Herbarium Bogoriense, Indonesia) Tukirin Partomihardjo (Ecologist, Herbarium Bogoriense, Indonesia) Joeni Setijo Rahajoe (Ecologist, Herbarium Bogoriense, Indonesia) Marlina Ardiyani (Taxonomist, Herbarium Bogoriense, Indonesia) Topik Hidayat (Taxonomist, Indonesia University of Education, Indonesia) Eizi Suzuki (Ecologist, Kagoshima University, Japan) Jun Wen (Taxonomist, Smithsonian Natural History Museum, USA) Managing Editor Himmah Rustiami (Taxonomist, Herbarium Bogoriense, Indonesia) Lulut Dwi Sulistyaningsih (Taxonomist, Herbarium Bogoriense, Indonesia) Secretary Endang Tri Utami Layout Editor Deden Sumirat Hidayat Medi Sutiyatno Illustrators Subari Wahyudi Santoso Anne Kusumawaty Correspondence on editorial matters and subscriptions for Reinwardtia should be addressed to: HERBARIUM BOGORIENSE, BOTANY DIVISION, RESEARCH CENTER FOR BIOLOGY- INDONESIAN INSTITUTE OF SCIENCES CIBINONG SCIENCE CENTER, JLN. RAYA JAKARTA - BOGOR KM 46, CIBINONG 16911, P.O. Box 25 Cibinong INDONESIA PHONE (+62) 21 8765066; Fax (+62) 21 8765062 E-MAIL: reinwardtia@mail.lipi.go.id 1 2 3 4 1 3 4 4 Cover images: 1. Begonia holosericeoides (female flower and habit) (Begoniaceae; Ardi et al.); 2. Abaxial cuticles of Alseodaphne rhododendropsis (Lauraceae; Nishida & van der Werff); 3. Dipo- dium puspitae, Dipodium purpureum (Orchidaceae; O'Byrne); 4. Agalmyla exannulata, Cyrtandra coccinea var. celebica, Codonoboea kjellbergii (Gesneriaceae; Kartonegoro & Potter). The Editors would like to thanks all reviewers of volume 14(1): Abdulrokhman Kartonegoro - Herbarium Bogoriense, Bogor, Indonesia Altafhusain B. Nadaf - University of Pune, Pune, India Amy Y. Rossman - Systematic Mycology & Microbiology Laboratory USDA-ARS, Beltsville, USA Andre Schuiteman - Royal Botanic Gardens, Kew, UK Ary P. Keim - Herbarium Bogoriense, Bogor, Indonesia Barry Conn - Royal Botanic Gardens National Herbarium of New South Wales, Sydney, Australia Dato' Abdul Latiff Mohamad - Universiti Kebangsaan Malaysia, Bangi, Selangor, Malaysia Daniel Potter - Department of Plant Sciences, University of California, Davis, California, USA Deby Arifiani - Herbarium Bogoriense, Bogor, Indonesia Ferry J. W. Slik - University of Brunei Darussalam, Brunei Henti H. Rachmat - Conservation and Rehabilitation Research and Development Center, Bogor, Indonesia Ian M. Turner - Royal Botanic Gardens, Kew, UK Iskandar Z. Siregar - Bogor Agricultural University, Bogor, Indonesia Jay H. Bernstein - Kingsborough Community College, Brooklyn, New York, USA Jens G. Rohwer - University of Hamburg, Hamburg, Germany Joan Pereira - SAN Herbarium, Sabah Forestry Department, Sabah, Malaysia Kuswata Kartawinata - Herbarium Bogoriense, Bogor, Indonesia Lars H. Schmidt - University of Copenhagen, Copenhagen, Denmark Mark Hughes - Royal Botanic Gardens, Edinburgh, UK Masahiro Kato - Kyoto University, Kyoto, Japan Nuril Hidayati - Herbarium Bogoriense, Bogor, Indonesia Ong Poh Teck - Forest Research Institute Malaysia, Kepong, Malaysia Peter C. van Welzen - National Herbarium Netherlands, Leiden University Branch, Leiden, Netherlands Reuben Nilus - Sabah Forestry Department, Sabah, Malaysia Rugayah - Herbarium Bogoriense, Bogor, Indonesia Ruth Kiew - Forest Research Institute of Malaysia, Kepong, Malaysia Uwe Braun - Institut fur Biologie Bereich Geobotanik und Botanischer Garten, Halle (Saale), Germany Yasuaki Sato - Osaka-Sangyo University, Osaka, Japan REINWARDTIA Vol 14, No 1, pp: 123 − 132 123 ON THE EVOLUTION OF DIPODIUM R. BR. Received January 09, 2014; accepted July 15, 2014 PETER O’BYRNE Research Fellow at Forest Research Institute Malaysia (FRIM). Waikiki Condominium H10-11, Tanjung Aru, 88100 Kota Kinabalu, Sabah, Malaysia. E-mail: oberonia@gmail.com. ABSTRACT O’BYRNE, P. 2014. On the evolution of Dipodium R. Br. Reinwardtia 14(1): 123 – 132. — Dipodium R. Br. (Orchidaceae), a genus of ca. 38 species, should be divided into an Australasian clade and a Malesian clade, based on morphological and geographic evidences. Dipodium section Dipodium and section Leopardanthus (Blume) O. Kuntze are re-defined to accommodate this change. An evolutionary scenario that explains this division is proposed. The lack of diversity of floral structure in the genus is probably caused by pollinator-specificity, while the diversity of plant form may be due to long-term environmental factors. An evolutionary explanation is suggested for the complex between Dipodium fevrellii J. J. Sm. and the hybrid D. pandanum Bailey in New Guinea. The complex in Java involving D. pictum (Lindl.) Rchb. f. and D. scandens (Bl.) J. J. Sm. is discussed, and the presence of a hybrid noted. In Borneo, the complex taxonomic situation caused by repeated inter-breeding of populations is noted. Key words: Dipodium, evolution, inter-specific complex, hybrid, New Guinea, Orchidaceae, section Dipodium, section Leopardanthus. ABSTRAK O’BYRNE, P. 2014. Evolusi Dipodium R. Br. Reinwardtia 14(1): 123 – 132. — Marga Dipodium R. Br. (Orchidaceae) yang mempunyai kurang lebih 38 jenis, seharusnya dibagi kedalam klad Australasian dan Malesian, berdasarkan bukti morfologi dan geografis. Dipodium seksi Dipodium dan seksi Leopardanthus (Blume) O. Kuntze dipertelakan ulang untuk mengakomodasi perubahan yang ada. Sebuah skenario evolusi yang menjelaskan pembagian ini telah diusulkan. Rendahnya keanekaragaman struktur pembungaan pada marga ini kemungkinan disebabkan oleh kekhususan polinator, sementara keanekaragaman bentuk tumbuhan kemungkinan disebabkan oleh faktor lingkungan. Penjelasan evolusi dianjurkan untuk jenis kompleks Dipodium fevrellii J. J. Sm. dan jenis hibrid D. pandanum Bailey di New Guinea. Jenis kompleks di Jawa meliputi D. pictum (Lindl.) Rchb. f. dan D. scandens (Bl.) J. J. Sm. dibahas dalam makalah ini, dan keberadaan jenis hibrid tersebut telah dicatat. Di Borneo, kondisi taksonomi yang kompleks disebabkan oleh persilangan antar populasi yang berulang. Kata kunci: Dipodium, evolusi, inter-spesifik kompleks, hibrid, New Guinea, Orchidaceae, seksi Dipodium, seksi Leopardanthus. INTRODUCTION Dipodium was established by R. Brown in 1810 for the Australian species D. punctatum. It currently has about 39 taxa, distributed from southern Indochina and Peninsular Thailand through the Malesian region to Palau (Caroline Islands), Queensland (Australia), Solomon Islands and New Caledonia and Vanuatu. Dipodium species show remarkable consistency in floral morphology, making it very difficult to determine species. This resulted in the genus gaining a reputation for being “difficult”, and most taxono- mists avoided working on it. There have been no substantial taxonomic publications on the genus, other than O’Byrne (2013), which found 11 characters in the floral morphology that permit determination at species level. A full taxonomic revision of Dipodium section Leopardanthus (O’Byrne, in prep.) contains many new taxa, some of which are referred to in this paper. Traditional treatment of Dipodium The relationship between Dipodium and related genera is unclear. Both Cameron et al (1999) and Stern and Judd (2002) placed Dipodium in subtribe Cyrtopodiinae and both showed that the Cyrtopodiinae is not a monophyletic group, but neither managed to resolve the relationships within the group. Pridgeon et al. (2009) placed Dipodium in tribe Cymbidieae, but did not comment on inter- generic relationships within the tribe and did not mention the infrageneric division into two sections. The genus has traditionally been divided into section Dipodium and section Leopardanthus (Blume) O. Kuntze, which correspond roughly to the two geographic centres of diversity, Australasia and Malesia. Section Dipodium has eleven species; nine endemic to Australia, one endemic to New Guinea, and D. squamatum (G. Forst.) R. Br. in Australia, New Caledonia and Vanuatu. They are homoge- REINWARDTIA 124 [VOL.14 nous in habit; all are leafless plants with subterra- nean rhizomes and an erect flowering stem that has many persistent bracts (reduced leaves). They all have an annual growth cycle. It has long been assumed that they are holomycotrophic, but Dearnaley (2006) suggests that some may be indirect parasites on tree roots. Section Leopardanthus has about 28 taxa, dis- tributed from southern Indochina and Peninsular Thailand through the Malesian region to Palau (Caroline Islands), Queensland (Australia) and Solomon Islands. All are leafy plants, but other- wise they are non-homogenous in habit. Three species are sympodial terrestrials (Fig. 1), with a disjunct distribution (Fig. 2). The remainder have traditionally been described as monopodial climbing plants (Fig. 3) and have a continuous distribution (Fig. 4). O’Byrne (2013) states that the division into terrestrial and climbing plants is an over-simplification, as climbing plants go through terrestrial and scrambling phases when immature. Horticultural experience supports this; Fig. 1. Dipodium paludosum is one of the three leafy sympodial species tradi- tionally described as “terrestrial”. Fig. 2. Map showing the disjunct distribution of the three leafy sympo- dial Dipodium species. Fig. 4. Map showing the continuous distribution of climbing Dipodium species. Fig. 3. The majority of section Leopardanthus species have traditionally been de- scribed as monopodial climbing plants. Note the copious climbing roots on this Dipodium fragrans. Photo by P.T.Ong. 2014] 125 O’BYRNE: On the evolution of Dipodium R. Br. so-called “terrestrial” plants can be cultivated as epiphytes, and climbing phase specimens can be cultivated as terrestrials. Section Leopardanthus can have a seasonal growth cycle or can grow continuously; in some taxa these life-forms are interchangeable. The origin and evolution of Dipodium (Fig. 5). If we consider Dipodium to be divided into Australasian and Malesian clades, then it is analogous to Dendrobium, where the division into two geographic clades is now well supported (Clements (2003); Schuiteman (in preparation); Wood (2006). It is reasonable to assume that both genera had similar origins and each developed into two clades through similar causes. This could mean, as suggested by Wood (2006) for Dendrobium, that Dipodium has an ancient origin and that the ancestral species was present on both the Australian and the Indian continental plates when (or shortly after) the ancient supercontinent of Pangea broke up. The ancestral species was probably not dissimilar to the modern-day Dipodium ensifolium F. Muell., a leafy sympodial terrestrial with a seasonal growth cycle. As the two continental plates drifted apart, the Dipodium populations on each plate developed separately, in isolation from each other. Consider the following scenario. As Australia drifted slowly north-east, it became drier and more arid, and Dipodium responded by becoming subterranean and developing holomycotrophy or indirect parasitism as their energy source. This enabled them to reduce water-loss by reducing the leaf area; in most modern day species, the leaves are vestigial, present only as non-photosynthetic scales at the stem base. Since their stems are short-lived (their only function is to support the inflorescence), these species never developed the ability to produce adventitious roots along the stems. The Indian continental plate drifted northwards until it collided with Eurasia. The Dipodium popu- lation travelled across the collision zone and then spread south-east through what is now Myanmar into Thailand and Malaysia. Somewhere on this journey, they encountered a barrier of rainforest and responded by developing the ability to climb up tree trunks. In order to do this, they need adventitious roots along the stems and these are present in copious quantities in all members of the Malesian clade. Once they had adapted to living in rainforest, the Malesian clade were able to spread south-east through the island chains, eventually reaching Caroline Islands, Australia and the Solo- mon Islands, the outer limits of their present-day distribution. The main binary morphological difference between the two Dipodium clades is therefore not the presence or absence of leaves (section Dipodium has reduced leaves on the stem), but the presence or absence of adventitious roots along the stem. Section Dipodium lacks adventitious roots at stem nodes; section Leopardanthus possesses them. This enables us to resolve the sectional position of the two of the “leafy terrestrial” taxa traditionally placed in Leopardanthus. The Austra- lian Dipodium ensifolium F.Muell. does not form adventitious roots at stem nodes, and therefore belongs in the Australasian clade, despite the presence of green leaves. Dipodium gracile Schltr. from Sulawesi is known only from the type (destroyed), but the protologue does not mention adventitious roots and Schlechter stated it was closer to D. ensifolium than D. paludosum (which is in section Leopardanthus). Dipodium gracile is there- fore tentatively placed in Section Dipodium, becoming its northernmost member and the only species outside Australasia. Dipodium Sections redefined; summary Section Dipodium Australasian Clade. 13 species. Distribution: Australia (11 species, 10 endemic), 1 extending to New Caledonia and Vanuatu, New Guinea (1 endemic), Sulawesi (1 endemic). Terrestrial. Sym- podial. Stems with green leaves (2 species) or leaves reduced to scales/bracts. Stems without adventitious roots at nodes. Column apex not yellow. Section Leopardanthus (Blume) O. Kuntze in Post & Kuntze, Lexicon Gen. Phanerog.: 179 (1904). Malesian Clade. ca. 26 taxa. Distribution: Indo- china, Malaysia, Indonesia, Philippines, Caroline Islands (1 taxon), New Guinea (3 taxa, 1 endemic), 1 taxon extending to Australia and Solomon Islands, Louisiade Archipelago (1 endemic). Adult phase either sympodial terrestrial (1 species) or monopodial climbing. Leafy. Stems with adventitious roots at nodes. Column apex yellow (3 exceptions). Pollinators, conservation of floral stucture, and the significance of flower color The flowers of all Dipodium species are unusually consistent in size, shape and structure, especially when compared to the great diversity of floral form found in genera of similar evolutionary REINWARDTIA 126 [VOL.14 Fig. 5. Diagram summarising the proposed evolution of Dipodium, the development of different plant habits, and the infrageneric division into sections. age, such as Dendrobium. Despite the long period of geographically-enforced separate development, there is hardly any difference in the flowers of the Australasian and Malesian clades. The simplest explanation for such highly conserved floral morphology is that from an early stage in its evolu- tionary history, the ancestral Dipodium was producing flowers that were a perfect match with its pollinator and neither flower nor insect have undergone major external changes since then. This does not mean that there has been no evo- lution of floral form since the time of the ancestral Dipodium, just that the evolutionary changes have been minor. The only visible difference between flowers of the Australasian and Malesian clades is the colour of the column apex. No species in the Australian clade have a yellow column apex, while most species in the Malesian clade (and all the successful ones) have a bright yellow column tip (Fig. 6). The evolutionary paths of the pollinating insects in Australasia and Malesia have diverged, but only slightly. The Dipodium species in the two areas have co-evolved along with the pollinators, resulting in a minor floral difference in flower colour between the two clades. The highly- conserved yellow column apex in the Malesian clade must be a pollinator-attractant, while the pollinators of the Australasian clade do not require yellow on the column to attract them to the flowers. The pollinators in Dipodium are not known, but there are two relevant records, both from Australia. Bernhardt & Burns-Balogh (1983) reported the removal of pollinia from Dipodium punctatum (Sm.) R. Br. by a female leafcutter bee (Chalicodoma derelicta) and Jones (1988) ob- served wasps removing pollinia from D. stenochi- lum O. Schwarz. Note that removal of pollinia does not automatically equate with being a polli- nator; Ling (2011) and Ong (2011) both report stingless bees (Trigona sarawakensis) collecting pollinia from orchid flowers without causing pol- lination. 2014] 127 O’BYRNE: On the evolution of Dipodium R. Br. It is possible to write a list of characters that the pollinators of Dipodium must possess: A medium to large size flying insect that is long-lived and travels long distances. Belongs to an old insect group that currently occurs throughout Malesia and Australasia. Possesses some degree of colour vision. Solitary bees (eg. leafcutter bees) and solitary wasps (eg. parasitoid wasps of subfamily Eumeninae) fulfill the criteria for being pollinators in Dipodium. Honeybees (Apis sp.) and bumble- bees (Bombus sp.) are excluded because they are not native to Australia. Unusually in Orchidaceae, flower colour is a conserved character in all Dipodium species except for D. pandanum (O’Byrne, 2011). Furthermore, most species share the same colour palette; white to cream tepals with pink to purple spots. Several species have yellow tepals with red spots. Only a small number of species deviate from these colour palettes and they are all rare, i.e. not very successful. This level of colour- conservation must be linked to pollinator- attraction. It also implies that the pollinators are likely to be attracted to and capable of pollinating, different Dipodium species. This means that hybrids between Dipodium species should occur; the remaining part of this paper deals with this issue. New Guinea and the Dipodium pandanum com- plex The island of New Guinea has 5 Dipodium species. Dipodium elatum J. J. Sm. is in section Dipodium; the other 4 species are in section Leopardanthus and arrived in New Guinea from Malesia. One of these, Dipodium brevilabium Metusala & O’Byrne, occupies an isolated position in the section, while the other three form Fig. 6. Most species in the Malesian clade have a bright yellow column tip. Left to right: Dipodium bicallosum J.J.Sm., Dipodium purpureum, Dipodium conduplicatum J. J. Sm. the Dipodium pandanum complex. The species involved are D. fevrellii J. J. Sm., D. pandanum Bail. and D. brassii O’Byrne (in prep.). Their distributions are shown in Fig. 7. Dipodium section Leopardanthus would have reached New Guinea by island-hopping to the south-east (from modern-day Maluku) while the island was being pushed in the opposite direction. New Guinea arrived at its present position only recently in geological time, having been pushed to the north-west by the northward drift of Australia. The Louisiade Archipelago, which was originally part of mainland New Guinea, got left behind and became a chain of isolated islands. Dipodium brassii is the only species found in the Louisiade Archipelago and is endemic there. Although it is possible that it arose in-situ by reversion, its glabrous midlobe flanks are caused by a fundamentally different bud-development to the other New Guinea Leopardanthus taxa, all of which have hairy midlobe flanks. It is more likely that the ancestor of D. brassii island-hopped to New Guinea from the north-east and occupied the whole island before the Louisiade Archipelago became isolated. Dipodium brassii (or its immediate ancestor) was probably the first Leopardanthus to reach New Guinea and although it has been replaced by more vigorous competitors on the mainland, it has survived on isolated islands that the competitors never reached. O’Byrne (in prep.) shows that Dipodium fevrellii is widely distributed across the Malesian islands (Fig. 7). It is one of the most easily recognised species in the section, with red-spotted bright yellow sepals and petals (Fig. 8) and a red-marked white midlobe which has stiff bristle- like hairs on its flanks. (Fig. 9). In New Guinea, it is found only in the west of the island, where it is often hard to distinguish from the rather variable REINWARDTIA 128 [VOL.14 Dipodium pandanum. Dipodium pandanum is the most common section Leopardanthus in New Guinea. It is unique in the genus because of the variation in tepal colour (Fig. 10). Since flower colour is a conserved character in all other Dipo- dium species, this level of colour variation points to a hybrid origin for D. pandanum. The species also shows variation in sidelobe length and an ab- normally high degree of variation in the quantity and nature of hairs on the midlobe flanks and the simplest explanation for this is hybridization between a species with glabrous flanks and another with hairy flanks. There are only two possible parents of this hybrid; D. brassii (purple-spotted creamy-white tepals, short side lobes, glabrous Fig. 7. Map showing the distribution of the three species in the Dipodium pandanum complex. midlobe flanks) and D. fevrellii (red-spotted yellow tepals, long sidelobes, hairy midlobe flanks). A hypothetical scenario that accounts for these observations is that D. brassii occupied New Guinea before the Louisiade Archipelago became isolated and before the relatively young volcanic island chain of New Britain, Bougainville and the Solomon Islands emerged. After the Louisiade Archipelago became isolated, D. fevrellii (or its immediate ancestor) invaded New Guinea from the west. Since the two species have the same pollinators, they hybridized, forming D. panda- num. The hybrid displaced both parent species and occupied New Guinea, later spreading through a newly emerged New Britain to Bou- gainville and the Solomon Islands and (only re- cently) crossing the Torres Straits to Australia. Dipodium fevrellii has probably invaded New Guinea several times, each time hybridizing with Fig. 8. Dipodium fevrellii. Photo by R. Kusumawati. Fig. 9. The lip of Dipodium fevrellii. Arrows point to the rows of bristle-like hairs. Photo by R. Kusumawati. Fig. 10. Variation in tepal colour in Dipodium panda- num. A: Solomon Islands, photo by P. J. Cribb. B: alba-form, Indonesian Papua, photo by F. Handoyo. C: Indonesian Papua, showing the influence of D. fevrellii, photo by E. F. de Vogel. D: Papua New Guinea, photo by P. O’Byrne. 2014] 129 O’BYRNE: On the evolution of Dipodium R. Br. the native Dipodium population. The existence of pure-form D. fevrellii in the west of the island is an unhybridized population from the most recent invasion. The cline of D. pandanum forms that is found from west to east is due to the percentage of D. fevrellii in the population; plants in the west have the highest percentage and thus the most bristles amongst the hairs on the midlobe; plants in the Solomon Islands have the lowest percent- age, and thus the midlobe flanks are covered in long soft hairs, not bristles. Fig. 11 shows some of the lip variation in D. pandanum and the lips of both parents. Although D. pandanum may be of hybrid origin, it has been in existence for a long time, has a widespread population and is stable in the eastern part of its range. Therefore, this paper does not propose any taxonomic change to its specific status. In the western part of the range, the population seems to contain many F2, F3 (etc.) hybrids, which cannot be readily distinguished from each other, or from the puta- tive parents. It is beyond the scope of this paper to resolve their status, so they are all treated as D. pandanum, but their existence is noted. For similar reasons, this paper does not propose to change the specific status of D. fevrellii. The species is widespread and easily-recognised across the whole of its range except in western New Guinea, where repeated hybridization events apparently caused it to merge with D. pandanum. If it were reduced to synonomy under D. panda- num, it would become impossible to identify these hybrids; it makes no taxonomic sense to make such a change. Java & the Dipodium scandens – D. pictum complex Dipodium scandens (Bl.) J. J. Sm. and D. pictum (Lindl.) Rchb. f. are the two earliest names in section Leopardanthus. Both species are confined to West Java, where D. pictum is known only from the type and D. scandens is still found in scattered populations (O’Byrne, in prep). Both species have been widely misinterpreted, partly because of poor protologues (D. pictum is a mix- tum), but also because a hybrid between the two species is more common than either parent. In books and photos of Javanese orchids, this hybrid has invariably been referred to as “D. pictum”. The confusion probably originated with J. J. Smith (1924), who illustrated 2 different taxa under D. pictum, but chose the wrong one for publication. The published illustration shows the hybrid; the unpublished one is D. pictum. Dipodium scandens and D. pictum have totally different flowers, different lip shapes and different stipes. The hybrid, Dipodium javanicum (O’Byrne, in prep), is intermediate between the parents in each of these (Fig. 12). It is being described at specific status because it has formed a stable popu- lation that has been in existence for at least one hundred years. There are other, unknown taxa in the Javanese complex. The photograph in Comber (1990) labelled “D. scandens, West Java, 820 m.” shows a taxon from Gunung Halimun that is neither D. scandens nor D. javanicum. It probably represents another hybrid; the influence of D. scan- dens on the lip shape is clear. I have seen no herbarium material that matches this photo. Borneo The Australasian section Dipodium never reached Borneo; here, all taxa belong to section Leopardanthus. Apart from a few distinct, uncommon (i.e. not very successful) species, the situation in Borneo is extremely complicated. The best-known species is Dipodium purpureum J. J. Sm. (Fig. 13), but D. fragrans O’Byrne & Vermeulen (Fig. 14), D. paludosum (Griff.) Rchb. f. (Fig. 15) and D. puspitae O’Byrne (in prep) (Fig. 16) are also widespread across the island, while pockets of D. fevrellii can be found in the east. All these taxa probably share the same pollinators and should be capable of interbreeding. It is hardly surprising that many Borneo Dipodium specimens cannot be placed under any of these existing names and appear to be intermediates between two or more of these species. With the few exceptions mentioned above, Dipodium in Borneo is in a state of population-flux, with new populations constantly emerging, then inter-breeding with neighbouring populations to produce new hybrids. This has probably been going on for tens or hundreds of millennia, with frequent invasions of species from neighbouring territories preventing the emergence of a stable population. O’Byrne (in prep) attempted to produce a specific level taxonomy using traditional morphological methods, but the results are unsatisfactory; it will require an inspired DNA analysis to fully resolve the relationships between these constantly changing populations. ACKNOWLEDGEMENTS The author would like to thank Phillip Cribb, Ed de Vogel, Frankie Handoyo, Richa Kusumawati and Ong Poh Teck, for allowing me to use their photos. Thanks to Forest Research Institute Malaysia (FRIM), Kebun Raya Bogor, Linus Gosuking at Kipandi Butterfly Park (Sabah), REINWARDTIA 130 [VOL.14 Fig. 11. Lip variation in D. pandanum, and the lips of D. fevrellii and D. brassii. Fig. 12. Lip and column of Dipodium javanicum, lip of D. pictum, lip and column of D. scandens. Four of the drawings after Smith (1905). Photos by F. Handoyo. 2014] 131 O’BYRNE: On the evolution of Dipodium R. Br. Fig. 13. Dipodium purpureum from Sabah, Borneo. Fig. 14. Dipodium fragrans from Central Kalimantan, Borneo. Photo by F. Handoyo. Fig. 15. Dipodium paludosum from Pahang, Peninsular Malaysia. Fig. 16. Dipodium puspitae holotype from Jambi, Suma- tra. The flowers of this species are hard to dis- tinguish from D. paludosum; the most obvious differences are in plant habit. REINWARDTIA 132 [VOL.14 Singapore Botanic Gardens and Tenom Orchid Centre (Sabah) for allowing me access to their collections of living plants. Thanks to Herbarium Bogoriense, Kepong Herbarium in FRIM, Singapore Botanic Gardens, the Royal Botanic Gardens at Kew, and the National Her-barium Nederland at Leiden for allowing me to examine their herbarium collections. Finally, my thanks to Andre Schuiteman and Ong Poh Teck for commenting on (although not necessarily agreeing with) the manuscript. REFERENCES BERNHARDT, P. & BURNS-BALOGH. 1983. 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Legends or illustration must be submitted separately at the end of the manuscript. References. Bibliography, list of literature cited or references follow the Harvard system as the following examples. Journal : KRAENZLIN, F. 1913. Cyrtandraceae novae Philippinenses I. Philipp. J. Sci. 8: 163-179. MAYER, V., MOLLER, ML, PERRET, M. & WEBER, A. 2003. Phylogenetic position and generic differentiation of Epithemateae (Gesneriaceae) inferred from plastid DNA sequence data. American J. Bot. 90: 321-329. Proceedings :TEMU, S. T. 1995. Peranan tumbuhan dan ternak dalam upacara adat "Djoka Dju" pada suku Lio, Ende, Flores, Nusa Tenggara Timur. In: NASUTION, E. (Ed.). Presiding Seminar dan Lokakarya Nasional Etnobotani II. LIP1 & Perpustakaan Nasional: 263-268. (In Indonesian). SIMBOLON, H. & MIRMANTO, E. 2000. Checklist of plant species in the peat swamp forests of Central Kalimantan, Indonesia. In: IWAKUMA et al. (Eds.) Proceedings of the International Symposium on: Tropical Peatlands. Pp. 179-190. Book : RIDLEY, H. N. 1923. Flora of the Malay Peninsula 2. L. Reeve & Co. Ltd, London. Part of Book : BENTHAM, G. 1876. Gesneriaceae. In: BENTHAM, G. & HOOKER, J. D. Genera plantarum 2. Lovell Reeve & Co., London. Pp. 990-1025. Thesis : BAIRD, L. 2002. A Grammar of Keo: An Austronesian language of East Nusantara. Australian National University, Canberra. [PhD. Thesis]. Website : http://www.nationaalherbarium.n1/fmcollectors/k/Kostermans AJGH.htm). Accessed 15 February 2012. Reinwardtia Published by Herbarium Bogoriense, Botany Division, Research Center for Biology, Indonesian Institute of Sciences Address: Jin. Raya Jakarta-Bogor Km. 46 Cibinong 16911, P.O. Box 25 Cibinong Telp. (+ 62) 21 8765066; Fax (+62) 21 8765062 E-mail: reinwardtia@mail.lipi.go.id REINWARDTIA Author Agreement Form Title of article Name of Author(s) : I/We hereby declare that: • My/Our manuscript was based on my/our original work. • It was not published or submitted to other journal for publication. • I/we agree to publish my/our manuscript and the copyright of this article is owned by Reinwardtia. • We have obtained written permission from copyright owners for any excerpts from copyrighted works that are included and have credited the sources in our article. Author signature (s) Date Name MUHAMMAD EFFENDI, TATIK CHIKMAWATI & DEDY DARNAEDI. New cytotypes of Pteris ensiformis var. victoria from Indonesia 133 SUZANA SABRAN, REUBEN NILUS, JOAN T. PEREIRA & JOHN BAPTIST SUGAU. Contribution of the heart of Borneo (HoB) initiative towards botanical exploration in Sabah, Malaysia 137 WENNI SETYO LESTARI, BAYU ADJIE, TASSANAI JARUWATANAPHAN, YASUYUKI WATANO & MADE PHAR- MAWATI. Molecular phylogeny of maidenhair fern genus Adiantum (Pteridaceae) from Lesser Sunda Islands, Indonesia based on Rbcl and Trnl-f 143 ELIZABETH A. WIDJAJA & DANIEL POTTER. Floristic study of Mekongga Protected Forest: towards establishment of the Mekongga National Park 157 YESSI SANTIKA, EKA FATMAWATI TIHURUA & TEGUH TRIONO. Comparative leaves anatomy of Pandanus, Freycinetia and Sararanga (Pandanaceae) and their diagnostic value 163 SUHARDJONO PRAWIROATMODJO & KUSWATA KARTAWINATA. Floristic diversity and structural characteristics of mangrove forest of Raj a Ampat, West Papua, Indonesia 171 IAN M. TURNER. A new combination in Orophea (Annonaceae) for Uvaria nitida Roxb. ex G. Don 181 IVAN S AVINOV. Taxonomic revision of Asian genus Glyptopetalum Thwaites (Celastraceae R. Br.) 183 YUSI ROSALINA, NISYAWATL ERWIN NURDIN, JATNA SUPRIATNA & KUSWATA KARTAWINATA. Floristic compo- sition and structure of a peat swamp forest in the conservation area of the PT National Sago Prima, Selat Panjang, Riau, Indone- sia 193 IMAN HID AY AT & JAMJAN MEEBOON. Cercospora brunfelsiicola (Fungi, Mycosphaerellaceae), a new tropical Cercosporoid fungus on Brunfelsia uniflora 211 MAX VAN BALGOOY & ELIZABETH A. WIDJAJA. Flora of Bali: a provisional checklist 219 EKA FATMAWATI TIHURUA & INA ERLINAWATI. Leaf anatomy of Pandanus spp. (Pandanceae) from Sebangau and Bukit Baka-Bukit Raya National Park, Kalimantan, Indonesia 223 JULIA SANG & RUTH KIEW. Diversity of Begonia (Begoniaceae) in Borneo - How many species are there? 23 3 DIAN LATIFAH, ROBERT A. CONGDON & JOSEPH A. HOLTUM. A Physiological approach to conservation of four palm species: Arenga australasica, Calamus australis, Hydriastele wendlandiana saALicuala ramsayi 237 REINWARDTIA Vol. 14. No. 1.2014 CONTENTS Page ABDULROKHMAN KARTONEGORO & DANIEL POTTER. The Gesneriaceae of Sulawesi VI: the species from Mekongga Mts. with a new species of Cyrtandra described 1 LIM CHUNG LU & RUTH KIEW. Codonoboea (Gesneriaceae) sections in Peninsular Malaysia 13 WISNU H. ARDI, YAYAN W. C. KUSUMA, CARL E. LEWIS, ROSNIATI A. RISNA, HARRY WIRIADINATA, MELISSA E. ABDO & DANIEL C. THOMAS. Studies on Begonia (Begoniaceae) of the Molucca Islands I: Two new species from Halmahera, Indonesia, and an updated description of Begonia holosericea 19 YUZAMMI, JOKO R. WITONO & WILBERT L. A. HETTERSCHEID. Conservation status of Amorphophallus discophorus Backer & Alderw. (Araceae) in Java, Indonesia 27 MOHAMMAD F. ROYYANI & JOENI S. RAHAJOE. Behind the sacred tree: local people and their natural resources sustainabil- ity 35 FIFI GUS DWIYANTI, KOICHI KAMIYA & KO HARADA. Phylogeographic structure of the commercially important tropical tree species, Dryobalanops aromatica Gaertn. F. (Dipterocarpaceae) revealed by microsatellite markers 43 SACHIKO NISHIDA & HENK VAN DER WERFF. Do cuticle characters support the recognition of Alseodaphne, Nothaphoebe and Dehaasia as distinct genera? 53 NURUL AMAL LATIFF, RAHAYU SUKMARIA SUKRI & FAIZAH METALI. Nepenthes diversity and abundance in five habi- tats in Brunei Damssalam 67 NURUL HAZLINA ZATNI & RAHAYU SUKMARIA SUKRI. The diversity and abundance of ground herbs in lowland mixed Dipterocarp forest and heath forest in Brunei Darussalam 73 MUHAMMAD AMIRUL AIMAN AHMAD JUHARI, NORATNI TALIP, CHE NURUL ATNI CHE AMRI & MOHAMAD RUZI ABDUL RAHMAN. Trichomes morphology of petals in some species of Acanthaceae 79 DIAN ROSLEINE, EIZI SUZUKI, ATIH SUNDAWIATI, WARDI SEPTIANA & DESY EKAWATI. The effect of land use history on natural forest rehabilitation at corridor area of Gunung Halimun Salak National Park, West Java, Indonesia 85 JULIUS KULIP. The Ethnobotany of the Dusun people in Tikolod village, Tambunan district, Sabah, Malaysia 101 PETER O'BYRNE. On the evolution of Dipodium R. Br 123 Reinwardtia is a LIPI accredited Journal (517/AU2/P2MI-LIPI/04/2013) Herbarium Bogoriense Botany Division Research Center for Biology - Indonesian Institute of Sciences Cibinong Science Center Jln. Raya Jakarta - Bogor, Km 46 Cibinong 16911, P.O. Box 25 Cibinong Indonesia barudepan 402-581-2-PB belakangbaru img577_Page_1 img577_Page_2 img577_Page_3 img577_Page_4