Genera Orchidacearum Volume 6: Epidendroideae (Part 3) [Illustrated] 0199646511, 9780199646517

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GEN ERA ORCHI DACEARUM Vol um e 6 Ep iden dr o ideae (Par t T h r ee)

G E N E R A O R C H I DA C E A R U M Volume 6 Epidendroideae (Part Three) Edited by Alec M. Pridgeon, Phillip J. Cribb, Mark W. Chase Royal Botanic Gardens, Kew

and

Finn N. Rasmussen Natural History Museum of Denmark

1

3 Great Clarendon Street, Oxford, OX2 6DP, United Kingdom Oxford University Press is a department of the University of Oxford. It furthers the University’s objective of excellence in research, scholarship, and education by publishing worldwide. Oxford is a registered trade mark of Oxford University Press in the UK and in certain other countries © Board of Trustees of the Royal Botanic Gardens, Kew, 2014 The moral rights of the authors have been asserted First Edition published in 2014 Impression: 1 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, without the prior permission in writing of Oxford University Press, or as expressly permitted by law, by licence or under terms agreed with the appropriate reprographics rights organization. Enquiries concerning reproduction outside the scope of the above should be sent to the Rights Department, Oxford University Press, at the address above You must not circulate this work in any other form and you must impose this same condition on any acquirer Published in the United States of America by Oxford University Press 198 Madison Avenue, New York, NY 10016, United States of America British Library Cataloguing in Publication Data Data available Library of Congress Control Number: 2001273293 ISBN 978–0–19–964651–7 Printed and bound by CPI Group (UK) Ltd, Croydon, CR0 4YY

PREFACE

What began as the germ of an idea by Phillip Cribb in the late 1990s blossomed into a 15-year orchid project that was produced chiefly at the Royal Botanic Gardens, Kew, but involved more than 200 experts throughout the world when everyone is taken into account – systematists, anatomists, palynologists, cytogeneticists, ecologists, artists, photographers, growers, and hybridizers. For all taxa in this final volume, 28 contributors provided up-to-date information on nomenclature, derivation of name, description, distribution (with maps), anatomy, palynology, cytogenetics, phytochemistry, phylogenetics, ecology, pollination, uses, and cultivation. Line drawings illustrating all genera appear here either for the first time or are reproduced with permission as noted in figure captions and acknowledgements. Numbers assigned to genera, continuous with the last volume, are based solely on the alphabetic order of their appearance in the text and bear no connection to phylogenetic position or level of evolutionary advancement. On the endpapers of this and other volumes is an alphabetic/numeric list of genera covered to date as an aid to finding generic treatments within and among volumes. Authorities for binomials are provided at the first occurrence in the text or in tabular form. Finn N. Rasmussen has provided a comprehensive glossary of terms covering all volumes in the end matter. This series was never intended to serve as the final classification of Orchidaceae, for that would be the antithesis of scientific integrity. Indeed, as long as there is even one orchid systematist working, as long as developing technologies improve our understanding of genetic relationships, as long as orchid species exist, their classification is forever destined to be a work in progress. Rather than a final monograph, then, the editors hope that Genera Orchidacearum becomes a benchmark for future scientists and identifies those gaps in our knowledge where further explanation and research are needed. ALEC M. PRIDGEON PHILLIP J. CRIBB MARK W. CHASE FINN N. RASMUSSEN

ACKNOWLEDGEMENTS

This final volume in the series would not have been possible without the dedicated work of so many talented individuals. We thank the many people who assisted our efforts in one way or another and in one place or another. Works of the following artists illustrated significant diagnostic elements of taxa: Juliet Beentje, Diane Bridson, Andrew Brown, Maria Helena Burian, Eleanor Catherine, C. L. Chan, Chen Li-jun, Maureen E. Church, G. C. K. Dunsterville, Mark Fothergill, Mary Grierson, Linda Gurr, N. Hallé, Cherry Ann Lavrih, Lucy F. L. Liew, Marco Antonio López Rosas, Loura Mason, Stella RossCraig, Gunnar Seidenfaden, Shim Phyau Soon, Judi Stone, Susanna Stuart-Smith, J. J. Vermeulen, Chin Wan Wai, Mrs. Wil Wessel-Brand, and Sue Wickison. We are especially indebted to Judi Stone, who has contributed hundreds of original drawings for this series, sometimes at a moment’s notice. We also acknowledge the photographers whose brilliant colour images enhance this volume: Peter Adams, Greg Allikas, Coen Arends, W. Bachmann, João Batista, Pierre Bonnet, Chen Li-jun, Mark Clements, Petr Efimov, Suranjan Fernando, Eberhard Fischer, Roger Hammer, Johan Hermans, Rudolf Jenny, Alex Kocyan, C. Satish Kumar, Anthony Lamb, Chuck McCartney, Piotr Markiewicz, Peter O’Byrne, Malcolm Perry, Imaduwa Priyadarshana, Dian Rabardja, Royal Botanic Garden, Edinburgh, Gerardo Salazar, André Schuiteman, Joyce Stewart, Milan Svanderlik, J. Tanner, Nguyen Thien Tich, Johan van Valkenburg, Jaap Vermeulen, Ed de Vogel, H. Vorster, Rogier van Vugt, Lubbert Westra, and Mark Whitten. We are grateful to the publishers of Blumea, Orchid Monographs, Nordic Journal of Botany, Orchids of Borneo, The Orchids of Bhutan, Orchids of Peninsular Malaysia and Singapore, Orchids of the Solomon Islands and Bougainville, Kew Bulletin, Flore de la Nouvelle Calédonie et dépendances. No. 8, Orchidacées, Flore du Cameroun, Orchidées, The Orchids of Mount Kinabalu, Flora of Tropical East Africa, Orchidaceae (Part 3), Flora of West Tropical Africa, Volume 3, Part 1-2, and Flora Zambesiaca, Volume 11, Part 2 for allowing us to reproduce drawings. Their participation in this project is deeply appreciated and added immeasurably to the final product. We thank Professor Stephen Hopper and Richard Deverell (past and present Director of the Royal Botanic Gardens, Kew, respectively) and Board of Trustees of the Royal Botanic Gardens, Kew, for continuing to support this international effort and also for providing laboratories and computer support. Professor David Mabberley (past Keeper of the Herbarium), Dr David Simpson (Acting Keeper of the Herbarium), and Professor Mark W. Chase (Keeper of the Jodrell Laboratory) offered facilities as well as encouragement. We and the other contributors must also acknowledge those who donated plant materials for study and/or DNA sequencing. The World Checklist of Selected Plant Families, compiled by Rafaël Govaerts and others, has been an invaluable resource for accepted plant names and standardization of author abbreviations, following the style established in Authors of Plant Names edited by Dick Brummitt and C. E. Powell. Directly or indirectly, every section and unit at the Royal Botanic Gardens, Kew, contributed to this project, and all warrant grateful recognition here. Diego Bogarín and Cassio van den Berg were kind enough to help prepare the distribution map for Campylocentrum, and Mario Blanco brought errors in previous volumes to our attention. Special thanks are reserved for André Schuiteman, who helped to secure artwork and resolved a multitude of problems, large and small, as they arose. This series would not have been possible without the hard work and patience of the various commissioning editors of Oxford University Press, particularly Ian Sherman, who has overseen the project since its inception. We also thank the many copy-editors and production editors who helped to ensure high standards throughout. It has been a great pleasure working with all of them.

A C K N OW L E D G E M E N T S

As for past volumes, we reserve our deepest gratitude for Lady Sainsbury, now 101 years old at this writing (1 April 2013), who has been a steadfast and generous supporter of Genera Orchidacearum and orchid conservation research. ALEC M. PRIDGEON PHILLIP J. CRIBB MARK W. CHASE FINN N. RASMUSSEN

CONTENTS

List of contributors  xiii List of colour plates  xv

I

 EPIDENDROIDEAE (Par t T hree)  1 A DENDROBIEAE 3 621. Bulbophyllum 4 622. Dendrobium 51 B VANDEAE 101 Adrorhizinae 104 623. Adrorhizon 104 624. Bromheadia 106 625. Sirhookera 109 Aeridinae 111 626. Acampe 129 627. Adenoncos 132 628. Aerides 133 629. Amesiella 137 630. Arachnis 139 631. Biermannia 142 632. Bogoria 142 633. Brachypeza 144 634. Calymmanthera 146 635. Ceratocentron 148 636. Chamaeanthus 150 637. Chiloschista 152 638. Chroniochilus 156 639. Cleisocentron 158 640. Cleisomeria 161 641. Cleisostoma 161 642. Cleisostomopsis 166 643. Cottonia 168 644. Cryptopylos 168 645. Deceptor 170 646. Dimorphorchis 172 647. Diplocentrum 175 648. Diploprora 177 649. Dryadorchis 180 650. Drymoanthus 182 651. Dyakia 183 652. Eclecticus 186 653. Gastrochilus 188 654. Grosourdya 191 655. Gunnarella 194 656. Holcoglossum 195

CONTENTS

657. Hymenorchis 201 658. Jejewoodia 203 659. Luisia 205 660. Macropodanthus 208 661. Micropera 210 662. Microsaccus 212 663. Mobilabium 212 664. Omoea 214 665. Ophioglossella 216 666. Papilionanthe 218 667. Papillilabium 220 668. Paraphalaenopsis 224 669. Pelatantheria 226 670. Pennilabium 229 671. Peristeranthus 231 672. Phalaenopsis 233 673. Phragmorchis 241 674. Plectorrhiza 242 675. Pomatocalpa 245 676. Porrorhachis 247 677. Pteroceras 249 678. Renanthera 252 679. Rhinerrhiza 253 680. Rhinerrhizopsis 256 681. Rhynchogyna 258 682. Rhynchostylis 260 683. Robiquetia 263 684. Saccolabiopsis 264 685. Saccolabium 266 686. Santotomasia 269 687. Sarcanthopsis 271 688. Sarcochilus 274 689. Sarcoglyphis 278 690. Sarcophyton 278 691. Schistotylus 280 692. Schoenorchis 282 693. Seidenfadenia 286 694. Seidenfadeniella 288 695. Singchia 290 696. Smithsonia 292 697. Smitinandia 295 698. Spongiola 295 699. Stereochilus 297 700. Taeniophyllum 299 701. Taprobanea 304 702. Thrixspermum 305 703. Trachoma 309 704. Trichoglottis 311 705. Tuberolabium 315 706. Uncifera 315 707. Vanda 317 708. Vandopsis 327 709. Xenikophyton 328 Agrostophyllinae 331 710. Agrostophyllum 331 711. Earina 335 x

CONTENTS

Angraecinae 338 712. Aerangis 344 713. Aeranthes 349 714. Ambrella 351 715. Ancistrorhynchus 353 716. Angraecopsis 355 717. Angraecum 358 718. Beclardia 363 719. Bolusiella 365 720. Calyptrochilum 367 721. Campylocentrum 370 722. Cardiochilos 372 723. Chauliodon 373 724. Cribbia 374 725. Cryptopus 376 726. Cyrtorchis 379 727. Dendrophylax 383 728. Diaphananthe 385 729. Dinklageella 389 730. Distylodon 390 731. Eggelingia 391 732. Erasanthe 393 733. Eurychone 395 734. Jumellea 397 735. Lemurella 399 736. Lemurorchis 402 737. Listrostachys 404 738. Margelliantha 405 739. Microcoelia 407 740. Mystacidium 411 741. Neobathiea 413 742. Nephrangis 415 743. Oeonia 417 744. Oeoniella 419 745. Ossiculum 421 746. Plectrelminthus 423 747. Podangis 424 748. Rangaeris 427 749. Rhaesteria 429 750. Rhipidoglossum 431 751. Sobennikoffia 433 752. Solenangis 435 753. Sphyrarhynchus 438 754. Summerhayesia 440 755. Taeniorrhiza 442 756. Triceratorhynchus 444 757. Tridactyle 445 758. Ypsilopus 448 Polystachyinae 450 759. Hederorkis 451 760. Polystachya 453 Addendum 463 Additional genera  463 761. Galeoglossum 463 762. Hemipiliopsis 466 xi

CONTENTS

763. Neolindleya 469 764. Pseuderia 471 765. Psychopsiella 473 Revised genera  476 122A. Nothostele 476 14A. Disa 479 458A. Cyanaeorchis 479 498A. Brassia 480 513A. Fernandezia 482 527A. Notyliopsis 483 569A. Gongora 485 References 487 Cumulative glossary  501 Corrigenda 513 Effective dates of publication  515 Generic synonyms  517 Series contributors  523 Index to scientific names  525 Subject index  535

xii

CONTRIBUTORS

DR PETER B. ADAMS (PA)

School of Botany, University of Melbourne, Victoria 3010, and Royal Botanic Gardens and National Herbarium of Victoria, Birdwood Avenue, South Yarra, Victoria 3141, Australia PROF. JOÃO A. N. BATISTA (JB)

Departamento Botânica, Universidade Federal de Minas Gerais, Av. Antônio Carlos 6627, Pampulha, Caixa Postal 486 31270-901 - Belo Horizonte, MG, Brasil LUCIANO B. BIANCHETTI (LB)

Embrapa, Recursos Genéticos e Biotecnologia, Parque Estação Biológica, P.O. Box 02372, Brasília, DF 70770-901, Brasil DR BARBARA S. CARLSWARD (BC)

Department of Biological Sciences, Eastern Illinois University, Charleston, Illinois 61920, USA PROF. MARK W. CHASE (MC)

Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, UK DR PHILLIP J. CRIBB (PC)

Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, UK R. H. S. Suranjan Fernando (SF)

Post Graduate Institute of Science, University of Peradeniya, Peradeniya, Sri Lanka DR GÜNTER FISCHER (GF)

Kadoorie Farm and Botanic Garden, Flora Conservation Department, Tai Po, Hong Kong, China DR LAUREN M. GARDINER (LG)

Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, UK DR BARBARA GRAVENDEEL (BG)

Naturalis Biodiversity Center, 2333 BE Leiden, The Netherlands DR RENÉE J. GRAYER (RG)

Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, UK JOHAN HERMANS (JH)

Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, UK DR ALEXANDER KOCYAN (AK)

University of Potsdam, Institute of Biochemistry and Biology, Biodiversity Research/Systematic Botany, Maulbeerallee 2a, D-14469 Potsdam, Germany Dr C. SatHish Kumar (CSK)

Tropical Botanic Garden & Research Institute, Palode, Thirivananthapuram, Kerala, India DR SHERYL D. LAWSON (SL)

Royal Botanic Gardens and National Herbarium of Victoria, Birdwood Avenue, South Yarra, Victoria 3141, Australia PROF. LUO YI-BO (LY-B)

State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China DR MARTIN MOTES (MM)

25000 Farmlife Road, Redland, Florida 33031, USA

CONTRIBUTORS

DR ALEC M. PRIDGEON (AP)

Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, UK EMERITUS PROF. FINN N. RASMUSSEN (FR)

University of Copenhagen, Botanical Museum and Garden, Gothersgade 130, DK-1123 Copenhagen K, Denmark DR GERARDO A. SALAZAR (GS)

Herbario Nacional de México (MEXU), Departamento de Botánica, Instituto de Biología, Universidad Nacional Autónoma de México, Apartado Postal 70-367, 04510 México, D.F., Mexico André Schuiteman (AS)

Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, UK ANTON SIEDER (TS)

Botanical Garden, University of Vienna, 1030 Vienna, Austria PROF ERIC DE CAMARGO SMIDT (ES)

Departamento de Botânica, sala 446, Setor de Ciências Biológicas, Centro Politécnico – UFPR, Caixa Postal 19031, Cep 81531-990, Curitiba, PR, Brasil PROF WILLIAM LOUIS STERN (WS)

Department of Biological Sciences,  Biscayne Bay Campus,  Florida International University,  North Miami, Florida 33181, USA DR NIGEL C. VEITCH (NV)

Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, UK DR JAAP JAN VERMEULEN (JJV)

Naturalis Biodiversity Center, 2300 RA Leiden, The Netherlands ROGIER RODERIK VAN VUGT (RV)

Hortus botanicus, Leiden University, 2300 RA Leiden, The Netherlands JEFFREY WOOD (JW)

Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, UK

xiv

COLOUR PLATES

Plate 1.

Bulbophyllum amplebracteatum Teijsm. & Binn. subsp. orthoglossum (H.Wendl. & Kränzl.) J.J. Verm. & P.O’Byrne, cult. (B. sect. Lepidorhiza) (Photo: Rogier van Vugt)

Plate 2.

Bulbophyllum bicolor Lindl. (B. sect. Cirrhopetaloides), cult. (Photo: Rogier van Vugt)

Plate 3.

Bulbophyllum cochleatum Lindl. var. bequaertii (De Wild.) J.J.Verm. (B. sect. Taurostalix), cult. (Photo: Rogier van Vugt)

Plate 4.

Bulbophyllum coloratum J.J.Sm. (B. sect. Codonosiphon), cult. (Photo: André Schuiteman)

Plate 5.

Bulbophyllum dayanum Rchb.f. (B. sect. Acrochaene), cult. (Photo: Jaap Vermeulen)

Plate 6.

Bulbophyllum delitescens Hance (B. sect. Macrostelydia), cult. (Photo: Rogier van Vugt)

Plate 7.

Bulbophyllum eberhardtii (Gagnep.) Seidenf. (B. sect. Cirrhopetalum), cult. (Photo: Rogier van Vugt)

Plate 8.

Bulbophyllum erythrosema J.J.Verm. (B. sect. Epibulbon), cult. (Photo: André Schuiteman)

Plate 9.

Bulbophyllum guttulatum (Hook.f.) N.P.Balakr. (B. sect. Brachyantha), cult. (Photo: André Schuiteman)

Plate 10. Bulbophyllum incisilabrum J.J.Verm. & P.O’Byrne (B. sect. Beccariana), cult. (Photo: André Schuiteman) Plate 11. Bulbophyllum lindleyanum Griff. (B. sect. Hirtula), cult. (Photo: Rogier van Vugt) Plate 12. Bulbophyllum longiscapum Rolfe (B. sect. Intervallatae), cult. (Photo: Rogier van Vugt) Plate 13. Bulbophyllum molossus Rchb.f. (B. sect. Pachychlamys), cult. (Photo: Malcolm Perry) Plate 14. Bulbophyllum monomeria J.J.Verm. (B. sect. Monomeria), cult. (Photo: Rogier van Vugt) Plate 15. Bulbophyllum nummularia (H.Wendl. & Kränzl.) Rolfe (B. sect. Trias), cult. (Photo: Piotr Markiewicz) Plate 16. Bulbophyllum nummularioides Schltr. (B. sect. Monanthes), cult. (Photo: Rogier van Vugt) Plate 17. Bulbophyllum patens King ex Hook.f. (B. sect. Stenochilus), cult. (Photo: Ed de Vogel) Plate 18. Bulbophyllum pedilochilus (Schltr.) J.J. Verm. (B. sect. Pedilochilus), cult. (Photo: Rogier van Vugt) Plate 19. Bulbophyllum piestobulbon Schltr. (B. sect. Piestobulbon), cult. (Photo: Dian Rabardja) Plate 20. Bulbophyllum plumatum Ames (B. sect. Plumata), cult. (Photo: André Schuiteman) Plate 21. Bulbophyllum psittacoglossum Rchb.f. (B. sect. Leopardinae), cult. (Photo: Piotr Markiewicz) Plate 22. Bulbophyllum saurocephalum Rchb.f. (B. sect. Saurocephalum), cult. (Photo: Jaap Vermeulen) Plate 23. Dendrobium angustipetalum J.J.Sm. (D. sect. Brevisaccata), cult. (Photo: André Schuiteman) Plate 24. Dendrobium archipelagense Howcroft & W.N.Takeuchi (D. sect. Spatulata), cult. (Photo: André Schuiteman) Plate 25. Dendrobium bigibbum Lindl. (D. sect. Phalaenanthe), Australia (Photo: Peter Adams)

C O L O U R P L AT E S

Plate 26. Dendrobium aff. bilobum Lindl. (D. sect. Biloba), cult. (Photo: André Schuiteman) Plate 27. Dendrobium bulbophylloides Schltr. (D. sect. Microphytanthe), cult. (Photo: Ed de Vogel) Plate 28. Dendrobium cacuminis Gagnep. (D. sect. Sarcopodium), cult. (Photo: André Schuiteman) Plate 29. Dendrobium caudiculatum (M.A.Clem. & D.L.Jones) Schuit. & de Vogel (D. sect. Rhizobium), cult. (Photo: André Schuiteman) Plate 30. Dendrobium densiflorum Wall. ex Lindl. (D. sect. Dendrobium), cult. (Photo: André Schuiteman) Plate 31. Dendrobium djamuense Schltr. (D. sect. Grastidium), cult. (Photo: André Schuiteman) Plate 32. Dendrobium ellipsophyllum Tang & F.T.Wang (D. sect. Distichophyllae), cult. (Photo: André Schuiteman) Plate 33. Dendrobium formosum Roxb. ex Lindl. (D. sect. Formosae), cult. (Photo: André Schuiteman) Plate 34. Dendrobium fytchianum Bateman (D. sect. Fytchianthe), cult. (Photo: André Schuiteman) Plate 35. Dendrobium gracilicaule F.Muell. (D. sect. Dendrocoryne), cult. (Photo: Peter Adams) Plate 36. Dendrobium hasseltii (Blume) Lindl. (D. sect. Pedilonum), cult. (Photo: André Schuiteman) Plate 37. Dendrobium lichenastrum (F.Muell.) Nicholls (D. sect. Lichenastrum), cult. (Photo: Peter Adams) Plate 38. Dendrobium lindleyi Steud. (D. sect. Dendrobium), cult. (Photo: Alec Pridgeon) Plate 39. Dendrobium minjemense Schltr. (D. sect. Diplocaulobium), cult. (Photo: André Schuiteman) Plate 40. Dendrobium aff. monticola P.F.Hunt & Summerh. (D. sect. Stachyobium), cult. (Photo: André Schuiteman) Plate 41. Dendrobium moschatum (Buch.-Ham.) Sw. (D. sect. Dendrobium), cult. (Photo: André Schuiteman) Plate 42. Dendrobium pinifolium Ridl. (D. sect. Conostalix), cult. (Photo: André Schuiteman) Plate 43. Dendrobium pleianthum Schltr. (D. sect. Pleianthe), cult. (Photo: Royal Botanic Garden, Edinburgh) Plate 44. Dendrobium polysema Schltr. (D. sect. Latouria), cult. (Photo: Alec Pridgeon) Plate 45. Dendrobium pseudoglomeratum T.M.Reeve & J.J.Wood (D. sect. Calyptrochilus), cult. (Photo: André Schuiteman) Plate 46. Dendrobium rumphiae Rchb.f. (D. sect. Cadetia), cult. (Photo: André Schuiteman) Plate 47. Dendrobium schneiderae F.M.Bailey (D. sect. Monophyllaea), cult. (Photo: Peter Adams) Plate 48. Dendrobium spatella Rchb.f. (D. sect. Aporum), cult. (Photo: André Schuiteman) Plate 49. Dendrobium spurium (Blume) J.J.Sm. (D. sect. Fugacia), cult. (Photo: André Schuiteman) Plate 50. Dendrobium squamiferum J.J.Sm. (D. sect. Amblyanthus), cult. (Photo: André Schuiteman) Plate 51. Dendrobium verruciferum Rchb.f. (D. sect. Macrocladium), cult. (Photo: Mark Clements) Plate 52. Dendrobium vonroemeri J.J.Sm. (D. sect. Herpethophytum), cult. (Photo: André Schuiteman) Plate 53. Dendrobium xantholeucum Rchb.f. (D. sect. Crinifera), cult. (Photo: André Schuiteman) Plate 54. Adrorhizon purpurascens (Thwaites) Hook.f., Sri Lanka (Photo: Suranjan Fernando) Plate 55. Bromheadia grandiflora Kruiz. & de Vogel, cult. (Photo: André Schuiteman) Plate 56. Sirhookera lanceolata (Wight) Kuntze, Sri Lanka (Photo: Suranjan Fernando) Plate 57. Acampe rigida (Buch.-Ham. ex Sm.) P.F.Hunt, cult. (Photo: André Schuiteman) Plate 58. Adenoncos sumatrana J.J.Sm., cult. (Photo: André Schuiteman)

xvi

C OLOUR PLATES

Plate 59. Aerides quinquevulnera Lindl., cult. (Photo: André Schuiteman) Plate 60. Amesiella philippinensis (Ames) Garay, cult. (Photo: Lubbert Westra) Plate 61. Arachnis grandisepala J.J.Wood, Sabah (Photo: Anthony Lamb) Plate 62. Brachypeza indusiata (Rchb.f.) Garay, cult. (Photo: André Schuiteman) Plate 63. Chiloschista exuperei (Guillaumin) Garay, cult. (Photo: Lubbert Westra) Plate 64. Cleisocentron merrillianum (Ames) Christenson, cult. (Photo: André Schuiteman) Plate 65. Cleisomeria pilosulum (Gagnep.) Seidenf. & Garay, cult. (Photo: Alexander Kocyan) Plate 66. Cleisostoma duplicilobum (J.J.Sm.) Garay, cult. (Photo: André Schuiteman) Plate 67. Cleisostoma simondii (Gagnep.) Seidenf., cult. (Photo: André Schuiteman) Plate 68. Cleisostomopsis eberhardtii (Finet) Seidenf., cult. (Photo: Nguyen Thien Tich) Plate 69. Cryptopylos clausus (J.J.Sm.) Garay, cult. (Photo: André Schuiteman) Plate 70. Dimorphorchis lowii (Lindl.) Rolfe, cult. (Photo: André Schuiteman) Plate 71. Diplocentrum recurvum Lindl., cult. (Photo: Alexander Kocyan) Plate 72. Diploprora championii (Lindl.) Hook.f., Sri Lanka (Photo: Imaduwa Priyadarshana) Plate 73. Dryadorchis dasystele Schuit. & de Vogel, cult. (Photo: André Schuiteman) Plate 74. Drymoanthus minutus Nicholls, cult. (Photo: Alexander Kocyan) Plate 75. Dyakia hendersoniana (Rchb.f.) Christenson, cult. (Photo © Greg Allikas, www.orchidworks. com) Plate 76. Gastrochilus retrocallus (Hayata) Hayata, cult. (Photo: André Schuiteman) Plate 77. Gastrochilus sororius Schltr., cult. (Photo: André Schuiteman) Plate 78. Grosourdya muscosa (Rolfe) Garay, cult. (Photo: André Schuiteman) Plate 79. Holcoglossum flavescens (Schltr.) Z.H.Tsi, cult. (Photo: André Schuiteman) Plate 80. Hymenorchis tanii Schuit. & de Vogel, cult. (Photo: André Schuiteman) Plate 81. Luisia morsei Rolfe, cult. (Photo: André Schuiteman) Plate 82. Luisia tristis (G.Forst.) Hook.f., Sri Lanka (Photo: Suranjan Fernando) Plate 83. Macropodanthus sabahensis J.J.Wood & A.L.Lamb, Malaysia (Photo: Alexander Kocyan) Plate 84. Micropera pallida (Roxb.) Lindl., cult. (Photo: André Schuiteman) Plate 85. Microsaccus truncatus Carr, cult. (Photo: André Schuiteman) Plate 86. Ophioglossella chrysostoma Schuit. & Ormerod, cult. (Photo: Johan van Valkenburg) Plate 87. Papilionanthe teres (Roxb.) Schltr., cult. (Photo: Lubbert Westra) Plate 88. Paraphalaenopsis labukensis Shim, A.L.Lamb & C.L.Chan, cult. (Photo: André Schuiteman) Plate 89. Pelatantheria rivesii (Guillaumin) Tang & F.T.Wang, cult. (Photo: André Schuiteman) Plate 90. Pennilabium struthio Carr, cult. (Photo: André Schuiteman) Plate 91. Phalaenopsis amabilis (L.) Blume, cult. (Photo © Greg Allikas, www.orchidworks.com) Plate 92. Phalaenopsis amboinensis J.J.Sm. ‘July Copper’, cult. (Photo © Greg Allikas, www.orchidworks.com) Plate 93. Phalaenopsis appendiculata Carr, cult. (Photo: André Schuiteman) Plate 94. Phalaenopsis bellina (Rchb.f.) Christenson ‘Krull’s Perfection’ FCC/AOS, cult. (Photo © Greg Allikas, www.orchidworks.com) xvii

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Plate 95. Phalaenopsis floresensis Fowlie, cult. (Photo: André Schuiteman) Plate 96. Phalaenopsis pulcherrima (Lindl.) J.J.Sm., cult. (Photo: André Schuiteman) Plate 97. Plectorrhiza tridentata (Lindl.) Dockrill, cult. (Photo: Peter Adams) Plate 98. Pomatocalpa maculosum (Lindl.) J.J.Sm., Sri Lanka (Photo: Suranjan Fernando) Plate 99. Pomatocalpa marsupiale (Kränzl.) J.J.Sm., cult. (Photo: André Schuiteman) Plate 100. Porrorhachis galbina (J.J.Sm.) Garay, cult. (Photo: André Schuiteman) Plate 101. Pteroceras teres (Blume) Holttum, cult. (Photo: André Schuiteman) Plate 102. Renanthera bella J.J.Wood, cult. (Photo: André Schuiteman) Plate 103. Rhinerrhiza divitiflora (F.Muell. ex Benth.) Rupp, cult. (Photo: Peter Adams) Plate 104. Rhinerrhizopsis moorei (Rchb.f.) Ormerod, cult. (Photo: André Schuiteman) Plate 105. Rhynchostylis coelestis (Rchb.f.) A.H.Kent ‘Frandys Delight’ AM/AOS, cult. (Photo © Greg Allikas, www.orchidworks.com) Plate 106. Robiquetia angustifolia Schltr., cult. (Photo: André Schuiteman) Plate 107. Robiquetia rosea (Lindl.) Garay, Sri Lanka (Photo: Suranjan Fernando) Plate 108. Robiquetia witteana (Rchb.f.) Kocyan & Schuit., cult. (Photo: André Schuiteman) Plate 109. Sarcochilus dilatatus F.Muell., cult. (Photo: Peter Adams) Plate 110. Sarcoglyphis smithianus (Kerr) Seidenf., cult. (Photo: André Schuiteman) Plate 111. Sarcophyton pachyphyllum (Ames) Garay, cult. (Photo: Lubbert Westra) Plate 112. Schoenorchis gemmata (Lindl.) J.J.Sm., cult. (Photo: Lubbert Westra) Plate 113. Seidenfadenia mitrata (Rchb.f.) Garay, Laos (Photo: Pierre Bonnet) Plate 114. Smithsonia saldanhae C.S.Kumar & Theuerkauf, cult. (Photo: C. Sathish Kumar) Plate 115. Smitinandia micrantha (Lindl.) Holttum, cult. (Photo: André Schuiteman) Plate 116. Stereochilus dalatensis (Guillaumin) Garay, cult. (Photo: Lubbert Westra) Plate 117. Taeniophyllum alwisii Lindl., Sri Lanka (Photo: Suranjan Fernando) Plate 118. Taeniophyllum montanum Carr, cult. (Photo: André Schuiteman) Plate 119. Thrixspermum pinocchio P.O’Byrne & J.J.Verm., cult. (Photo: André Schuiteman) Plate 120. Trachoma speciosum D.L.Jones, B.Gray, M.A.Clem. & J.J.Wood. cult. (Photo: André Schuiteman) Plate 121. Trichoglottis kinabaluensis Rolfe, cult. (Photo: André Schuiteman) Plate 122. Trichoglottis smithii Carr, cult. (Photo: André Schuiteman) Plate 123. Trichoglottis tenera (Lindl.) Rchb.f., Sri Lanka (Photo: Suranjan Fernando) Plate 124. Tuberolabium odoratissimum (J.J.Sm.) Garay, cult. (Photo: André Schuiteman) Plate 125. Uncifera thailandica Seidenf. & Smitinand, cult. (Photo: André Schuiteman) Plate 126. Vanda dives (Rchb.f.) L.M.Gardiner, cult. (Photo: André Schuiteman) Plate 127. Vanda falcata (Thunb.) Beer, cult. (Photo © Greg Allikas, www.orchidworks.com) Plate 128. Vanda garayi (Christenson) L.M.Gardiner, cult. (Photo: André Schuiteman) Plate 129. Vanda tessellata (Roxb.) Hook.f. ex G.Don, Sri Lanka (Photo: Imaduwa Priyadarshana) Plate 130. Vanda testacea (Lindl.) Rchb.f., Sri Lanka (Photo: Suranjan Fernando) Plate 131. Vandopsis gigantea (Lindl.) Pfitzer, cult. (Photo © Greg Allikas, www.orchidworks.com)

xviii

C OLOUR PLATES

Plate 132. Agrostophyllum majus Hook.f., cult. (Photo: André Schuiteman) Plate 133. Earina valida Rchb.f., cult. (Photo: André Schuiteman) Plate 134. Aerangis articulata (Rchb.f.) Schltr., cult. (Photo: Johan Hermans) Plate 135. Aerangis hariotiana (Kränzl.) P.J.Cribb & Carlsward, cult. (Photo: Johan Hermans) Plate 136. Aerangis monantha Schltr., cult. (Photo: Johan Hermans) Plate 137. Aeranthes filipes Schltr., cult. (Photo: Johan Hermans) Plate 138. Aeranthes schlechteri Bosser, cult. (Photo: Johan Hermans) Plate 139. Ambrella longituba H.Perrier, cult. (Photo: H. Vorster) Plate 140. Ancistrorhynchus cephalotes (Rchb.f.) Summerh., cult. (Photo: André Schuiteman) Plate 141. Angraecopsis parviflora (Thouars) Schltr., cult. (Photo: Johan Hermans) Plate 142. Angraecum eburneum Bory, cult. (Photo: Johan Hermans) Plate 143. Angraecum magdalenae Schltr. & H.Perrier, cult. (Photo: Johan Hermans) Plate 144. Angraecum sesquipedale Thouars, cult. (Photo: Johan Hermans) Plate 145. Angraecum viguieri Schltr., cult. (Photo: Johan Hermans) Plate 146. Beclardia macrostachya (Thouars) A.Rich., cult. (Photo: Johan Hermans) Plate 147. Bolusiella maudiae (Bolus) Schltr., Rwanda (Photo: Eberhard Fischer) Plate 148. Calyptrochilum christyanum (Rchb.f.) Summerh., cult. (Photo: Johan Hermans) Plate 149. Campylocentrum brenesii Schltr., cult. (Photo: Rudolf Jenny) Plate 150. Cribbia confusa P.J.Cribb, cult. (Photo: André Schuiteman) Plate 151. Cryptopus paniculatus H.Perrier, cult. (Photo: Johan Hermans) Plate 152. Cyrtorchis arcuata (Lindl.) Schltr., cult. (Photo: Johan Hermans) Plate 153. Cyrtorchis ringens (Rchb.f.) Summerh., cult. (Photo: Johan Hermans) Plate 154. Dendrophylax funalis (Sw.) Benth. ex Rolfe, cult. (Photo: Mark Whitten) Plate 155. Dendrophylax lindenii (Lindl.) Benth. ex Rolfe, USA (Photo: Roger Hammer) Plate 156. Diaphananthe pellucida (Lindl.) Schltr., cult. (Photo: Johan Hermans) Plate 157. Eggelingia ligulifolia Summerh., cult. (Photo: Eberhard Fischer) Plate 158. Erasanthe henrici (Schltr.) P.J.Cribb, Hermans & D.L.Roberts, cult. (Photo: Johan Hermans) Plate 159. Eurychone rothschildiana (O’Brien) Schltr., cult. (Photo: Johan Hermans) Plate 160. Jumellea rossii Senghas, cult. (Photo: Johan Hermans) Plate 161. Lemurella culicifera (Rchb.f.) H.Perrier, cult. (Photo: Johan Hermans) Plate 162. Lemurorchis madagascariensis Kränzl., cult. (Photo: Johan Hermans) Plate 163. Listrostachys pertusa (Lindl.) Rchb.f., cult. (Photo: Rudolf Jenny) Plate 164. Margelliantha leedalii P.J.Cribb, Tanzania (Photo: Phillip Cribb) Plate 165. Microcoelia koehleri (Schltr.) Summerh., cult. (Photo: Johan Hermans) Plate 166. Microcoelia perrieri (Finet) Summerh., cult. (Photo: Johan Hermans) Plate 167. Mystacidium venosum Harv. ex Rolfe, cult. (Photo: Johan Hermans) Plate 168. Neobathiea grandidieriana (Rchb.f.) Garay, cult. (Photo: Johan Hermans) Plate 169. Nephrangis filiformis (Kränzl.) Summerh., cult. (Photo: Eberhard Fischer) xix

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Plate 170. Oeonia rosea Ridl., cult. (Photo: Johan Hermans) Plate 171. Oeoniella polystachys (Thouars) Schltr., cult. (Photo: Johan Hermans) Plate 172. Plectrelminthus caudatus (Lindl.) Summerh., cult. (Photo: Johan Hermans) Plate 173. Podangis dactyloceras (Rchb.f.) Schltr., cult. (Photo: Johan Hermans) Plate 174. Podangis rhipsalisocia (Rchb.f.) P.J.Cribb & Carlsward, cult. (Photo: Johan Hermans) Plate 175. Rhaesteria eggelingii Summerh., Rwanda (Photo: Eberhard Fischer) Plate 176. Rhipidoglossum stolzii (Schltr.) Garay, cult. (Photo: J. Tanner) Plate 177. Sobennikoffia humbertiana H.Perrier, cult. (Photo: Johan Hermans) Plate 178. Solenangis wakefieldii (Rolfe) P.J.Cribb & J.Stewart, cult. (Photo: Joyce Stewart) Plate 179. Sphyrarrhynchus schliebenii Mansf., Tanzania (Photo: J. Tanner) Plate 180. Summerhayesia laurentii (De Wild.) P.J.Cribb, cult. (Photo: Coen Arends) Plate 181. Triceratorhynchus viridiflorus Summerh., Rwanda (Photo: Eberhard Fischer) Plate 182. Tridactyle bicaudata (Lindl.) Schltr., cult. (Photo: Johan Hermans) Plate 183. Tridactyle tanneri P.J.Cribb, Tanzania (Photo: Phillip Cribb) Plate 184. Ypsilopus longifolius (Kränzl.) Summerh., cult. (Photo: W. Bachmann) Plate 185. Polystachya clareae Hermans, cult. (Photo: Johan Hermans) Plate 186. Polystachya concreta (Jacq.) Garay & H.R.Sweet, Sri Lanka (Photo: Suranjan Fernando) Plate 187. Polystachya concreta (Jacq.) Garay & H.R.Sweet, Florida (USA) (Photo: Chuck McCartney) Plate 188. Polystachya goetzeana Kränzl., cult. (Photo: Milan Svanderlik) Plate 189. Polystachya porphyrochila J.Stewart, cult. (Photo: Joyce Stewart) Plate 190. Polystachya tsaratananae H.Perrier, cult. (Photo: Johan Hermans) Plate 191. Galeoglossum tubulosum (A.Rich. & Galeotti) Salazar & Soto Arenas, cult. (Photo: Gerardo Salazar) Plate 192. Nothostele acianthiformis (Rchb.f. & Warm.) Garay, cult. (Photo: João Batista) Plate 193. Hemipiliopsis purpureopunctata (K.Y.Lang) Y.B.Luo & S.C.Chen, China (Photo: Chen Li-jun) Plate 194. Hemipiliopsis purpureopunctata (K.Y.Lang) Y.B.Luo & S.C.Chen (close-up), China (Photo: Chen Li-jun) Plate 195. Neolindleya camtschatica (Cham.) Nevski, Russia (Photo: Petr Efimov) Plate 196. Neolindleya camtschatica (Cham.) Nevski, (close-up) Russia (Photo: Petr Efimov) Plate 197. Pseuderia floribunda Schltr., Papua New Guinea (Photo: Peter O’Byrne)

xx

I E P I DENDROI D E AE

(Part Three)

A . T RI BE DENDRO BIEAE

Dendrobieae Endl., Gen. Pl., 190 (1837). Type: Dendrobium Sw. Bulbophylleae Pfitzer, Entw. Nat. Anordn. Orchid., 104 (1887), as ‘Bolbophyllinae’. Type: Bulbophyllum Thouars Sunipieae Szlach., Folia Geobot. Phytotax., 26, 323 (1991). Type: Sunipia Buch.-Ham. ex Sm., syn. nov. Taxa at the rank of subtribe referable to Dendrobieae: Dendrobiinae Lindl., Gen. Sp. Orch. Pl., 45 (1830), as ‘sect. Dendrobieae’. Type: Dendrobium Sw. Bulbophyllinae Schltr., Orchideen, ed. 1, 319 (1914). Type: Bulbophyllum Thouars Genyorchidinae Schltr., Orchideen, ed. 1, 336 (1914). Type: Genyorchis Schltr. Sunipiinae Dressler, Selbyana, 5, 205 (1979). Type: Sunipia Buch.Ham. ex Sm. Epigeneiinae M.A.Clem., Telopea, 10, 278 (2003). Type: Epigeneium Gagnep. Grastidiinae M.A.Clem., Telopea, 10, 278 (2003). Type: Grastidium Blume

Description Epiphytic, lithophytic, or rarely terrestrial, sympodial herbs with one- to many-leaved, heteroblastic or homoblastic pseudobulbs or with few- to many-leaved, cane-like stems. Leaves distichous, conduplicate, with or without sheathing base, glabrous or rarely pubescent or papillose. Inflorescence basal, lateral or terminal, oneflowered or racemose, rarely branching. Flowers resupinate or not, minute to large, ephemeral to long-lasting; distichous, secund or helically arranged; fragrant or non-fragrant. Sepals entire, free or variously connate; the lateral sepals often forming a mentum together with the column foot. Petals entire or variously divided, free. Labellum free or adnate to the column foot, rigidly attached or hinged through a ligament and then highly mobile; entire or lobed, with or without keels and/or callosities, not spurred. Column usually straight, with or without lateral wings or apical stelidia; column foot usually present and well-developed, rarely reduced to almost absent; anther helmet-shaped, rarely hornlike and elongate, two-locular; pollinia usually four, sometimes two, usually oblong and laterally flattened, hard-waxy, if four then cohering in two pairs, those of a pair subequal to strongly unequal in size, without caudicles, rarely with a hamular stipe in some Bulbophyllum species; stigma a ventral concavity, rarely with a raised lower margin; rostellum simple, usually short, but sometimes beak-like and elongate; a well-defined, detachable viscidium usually absent, when present semiliquid or rarely solid. Ovary

and pedicel terete or winged. Capsule ellipsoid, oblong, obovoid to subglobose, unilocular, sometimes winged or triangular in cross-section, without endocarpic elaters. (AS)

Distribution (Fig. A.1) Dendrobieae are a pantropical tribe of two genera, Bulbophyllum and Dendrobium, with about 3650 species. Dendrobium is absent from Africa and America. Both genera are most diverse in Southeast Asia and Australasia, especially in New Guinea, where, for both genera, almost 30% of all species occur. (AS)

Phylogenetics Endlicher (1837) was the first to apply the rank of tribe to a group of genera for which Lindley (1830) had already used the name Dendrobieae, but as a ‘section’ of the tribe Malaxideae. If we disregard some misplaced genera, such as Diglyphosa and Microcoelia, then Lindley’s concept essentially covers the genera Bulbophyllum, Dendrobium, Polystachya, and Eria and their allies. Until the advent of molecular methods, Dendrobium was most often considered to be closely related to Eria, for example by Schlechter (1927) in his influential work Die Orchideen. DNA studies (e.g. Clements 2006) have shown, however, that Dendrobium is the sister group of Bulbophyllum, and that Eria and Polystachya are more distantly related. Van den Berg et al. (2005) performed a Bayesian analysis of four combined DNA regions of species in subfamily Epidendroideae and found that Dendrobieae are sister to a strongly supported clade that they called the vandoid orchids (in their analysis including Agrostophyllinae, Polystachyinae, Aeridinae, and other subtribes). However, the clade Dendrobieae + vandoid orchids has a low support, which renders this result inconclusive. More recently, Górniak et al. (2010), using the low-copy nuclear gene Xdh, confirmed the sister group relationship of Dendrobium (including Epigeneium) and Bulbophyllum but found Dendrobieae to be sister to Malaxideae with strong bootstrap support. However, the combined clade Dendrobieae + Malaxideae had no strongly supported sister group relationship with any other, much like all other major clades thus far identified in Epidendroideae. The morphological differences between Bulbophyllum and Dendrobium are few, as indicated in the key below. However, the two genera differ largely in their pollination biology: Bulbophyllum is mainly, with few known exceptions, fly-pollinated (but not syrphid flies, apparently), whereas Dendrobium is mainly bee- and (to 3

DENDROBIEAE

Fig. A.1.  Distribution map of Dendrobieae.

a lesser extent) bird-, wasp-, and syrphid fly-pollinated. Owing to the different pollination syndromes, Dendrobium species tend to have showier, more brightly coloured flowers than Bulbophyllum species, and it is usually easy to assign the species to the proper genus using either vegetative or floral characters. (AS)

Taxonomic literature Clements, M. A. (2006). Molecular phylogenetic systematics in Dendrobieae (Orchidaceae). Aliso, 22, 465–80. Endlicher, S. (1837). Genera plantarum secundum ordines naturales disposita. Fr. Beck, Vienna. Górniak, M., Paun, O., and Chase, M. W. (2010). Phylogenetic relationships within Orchidaceae based on a low-copy nuclear coding gene, Xdh: congruence with organellar and nuclear ribosomal DNA results. Molecular Phylogenetics and Evolution, 56, 784–95. Lindley, J. (1830). The genera and species of orchidaceous plants. Ridgways, London. Schlechter, R. (1927). Die Orchideen; ihre Beschreibung, Kultur und Züchtung. 2nd ed., Paul Parey, Berlin. Van den Berg, C., Goldman, D. H., Freudenstein, J. V., Pridgeon, A. M., Cameron, K. M., and Chase, M. W. (2005). An overview of the phylogenetic relationships within Epidendroideae inferred from multiple DNA regions and recircumscription of Epidendreae and Arethuseae (Orchidaceae). American Journal of Botany, 92, 613–24.

Artificial key to the genera of Dendrobieae (AS) 1. Plants with pseudobulbs (sometimes reduced) consisting of a single internode; pseudobulbs with one or two non●

4



●

sheathing leaves, rarely with a tuft of several needle-shaped leaves. Inflorescence arising from the base of the pseudobulb or the rhizome. Pollinia four, often distinctly unequal in size, sometimes two . . . . . . . . . . . . . 621. Bulbophyllum Plants with cane-like stems or with pseudobulbs (sometimes reduced) consisting of one to many internodes; stems or pseudobulbs one- to many-leaved, the leaves sheathing or not. Inflorescence arising above the base of the pseudobulb or stem, laterally or apically. Pollinia four, subequal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 622. Dendrobium

6 2 1 .  BU L B OPH YL LU M Bulbophyllum Thouars, Hist. Orchid., t. 3 (1822), nom. cons. Type species: Bulbophyllum nutans (Thouars) Thouars, typus cons. (basionym: Phyllorkis nutans Thouars), cons. pending. Phyllorkis Thouars, Nouv. Bull. Sci. Soc. Philom. Paris, 1, 319 (1809). Type species: Phyllorkis nutans Thouars Sunipia Buch.-Ham. ex Sm. in A.Rees, Cycl. 34, Art. Stelis, nos. 11 & 13 (1816). Lectotype (designated by Senghas 1991): Sunipia racemosa (Sm.) Tang & F.T.Wang (=Bulbophyllum reptans (Lindl.) Lindl. ex Wall.), syn. nov. Tribrachia Lindl., Bot. Reg., 10, t. 832 (1823) Type species: Tribrachia reptans Lindl. (=Bulbophyllum reptans (Lindl.) Lindl. ex Wall.) Osyricera Blume, Bijdr. Fl. Ned. Ind., 307 (1825). Type species: Osyricera crassifolia Blume (=Bulbophyllum osyricera J.J.Sm.) Epicranthes Blume, Bijdr. Fl. Ned. Ind., 306 (1825). (Epicranthes). Type species: Epicranthes javanica Blume (=Bulbophyllum epicrianthes (Blume) Hook.f.)

BULBOPHYLLUM

Ephippium Blume, Bijdr. Fl. Ned. Ind., 308 (1825). Lectotype (designated by Vermeulen 2008): Ephippium lepidum Blume (=Bulbophyllum lepidum (Blume) J.J.Sm.) Diphyes Blume, Bijdr. Fl. Ned. Ind., 310 (1825). Lectotype (designated by Szlachetko & Rutkowski 2008): Diphyes tortuosa Blume (=Bulbophyllum tortuosum (Blume) Lindl.) Cochlia Blume, Bijdr. Fl. Ned. Ind., 320 (1825). Type species: Cochlia violacea Blume (=Bulbophyllum salaccense Rchb.f.) Anisopetalon Hook., Exot. Fl., 2, t. 149 (1825). Type species: Anisopetalon careyanum Hook. (=Bulbophyllum careyanum (Hook.) Spreng.) Zygoglossum Reinw., Syll. Pl. Nov., 2, 5 (1825). Type species: Zygoglossum umbellatum Reinw. (=Bulbophyllum longiflorum Thouars) Megaclinium Lindl., Bot. Reg., 12, t. 989 (1826). Type species: Megaclinium falcatum Lindl. (=Bulbophyllum falcatum (Lindl.) Rchb.f. var. falcatum) Odontostylis Breda, Gen. Sp. Orchid. Asclep., t. 4 (1827), nom. illeg. Lectotype: Odontostylis triflora (Blume) Breda (=Bulbophyllum triflorum (Blume) Blume) Sestochilos Breda, Gen. Sp. Orchid. Asclep., 1, t. 3 (1827). Type species: Sestochilos uniflorum Breda (=Bulbophyllum lobbii Lindl.) Hippoglossum Breda, Gen. Sp. Orchid. Asclep., 3, t. 14 (1829). Type species: Hippoglossum umbellatum Breda (=Bulbophyllum lepidum (Blume) Lindl.) Cirrhopetalum Lindl., Gen. Sp. Orchid. Pl., 45 (1830), nom. cons. Type species: Cirrhopetalum thouarsii Lindl. (=Bulbophyllum longiflorum Thouars) Lyraea Lindl., Gen. Sp. Orchid. Pl., 46 (1830). Type species: Lyraea prismatica Lindl. Monomeria Lindl., Gen. Sp. Orchid. Pl., 61 (1830). Type species: Monomeria barbata Lindl. Lectotype (designated by Averyanov 1994): Sunipia scariosa Lindl., syn. nov. Trias Lindl., Gen. Sp. Orchid. Pl., 60 (1830). Lectotype (designated by Averyanov 1994): Trias oblonga Lindl., syn. nov. Macrolepis A.Rich. in J.S.C.Domont d’Urville, Voy. Astrolabe, 2, 25 (1833). Type species: Macrolepis longiscapa A.Rich. (=Bulbophyllum longiscapum (A.Rich.) Rolfe) Drymoda Lindl., Sert. Orchid., t. 8 C (1838). Type species: Drymoda picta Lindl., syn. nov. Malachadenia Lindl., Edwards’s Bot. Reg., 25 (Misc.), 67 (1839). Type species: Malachadenia clavata Lindl. (=Bulbophyllum malachadenia Cogn.) Oxysepala Wight, Icon. Pl. Ind. Orient., 5, 17 (1851). Type species: Oxysepala ovalifolium Wight (=Bulbophyllum clandestinum Lindl.) Bolbophyllaria Rchb.f., Bot. Zeitung (Berlin), 10, 934 (1852). Type species: Bolbophyllaria bracteolata Rchb.f. (=Bulbophyllum bracteolatum Lindl.) Bolbophyllopsis Rchb.f., Bot. Zeitung (Berlin), 10, 933 (1852). Type species: Bolbophyllopsis morphologorum Rchb.f. (=Bulbophyllum umbellatum Lindl.) Didactyle Lindl., Fol. Orchid., 1, 1 (1852). Type species: Didactyle exaltata Lindl. (=Bulbophyllum exaltatum Lindl.) Taurostalix Rchb.f., Bot. Zeitung (Berlin), 10, 933 (1852). Type species: Taurostalix herminiostachys Rchb.f. (=Bulbophyllum pumilum (Sw.) Lindl.)

Xiphizusa Rchb.f., Bot. Zeitung (Berlin), 10, 919 (1852). Type species: Xiphizusa chloroptera Rchb.f. (=Bulbophyllum chloropterum Rchb.f.) Acrochaene Lindl., Fol. Orchid., 2, 1 (1853). Type species: Acrochaene punctata Lindl. (=Bulbophyllum kingii Hook.f.), syn. nov. Ione Lindl., Fol. Orchid., 2, 1 (1853). Lectotype (here designated): Ione paleacea Lindl., syn. nov. Henosis Hook.f., Fl. Brit. India, 5, 771 (1890). Type species: Henosis longipes Rchb.f. Adelopetalum Fitzg., J. Bot., 29, 152 (1891). Type species: Adelopetalum bracteatum (F.M.Bailey) Fitzg. Genyorchis Schltr., Westafr. Kautschuk-Exped., 280 (1900). Type species: Genyorchis pumila Schltr. Pedilochilus Schltr. in K.M.Schumann & C.A.G.Lauterbach, Fl. Schutzgeb. Südsee, Nachtr., 218 (1905). Type species: Pedilochilus papuanum Schltr., syn. nov. Pelma Finet, Notul. Syst. (Paris), 1, 112 (1909). Syntype: Pelma absconditum J.J.Sm., Pelma neocaledonicum Schltr. (= B. absconditum J.J.Sm.) Saccoglossum Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 683 (1912). Lectotype: Saccoglossum papuanum Schltr., syn. nov. Codonosiphon Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 893 (1913). Type species: Codonosiphon codonanthum (Schltr.) Schltr. (basionym: Bulbophyllum codonanthum Schltr.) Dactylorhynchus Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 890 (1913). Type species: Dactylorhynchus flavescens Schltr. (=Bulbophyllum latipes J.J.Sm.) Monosepalum Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 682 (1913). Lectotype (designated by Van Royen 1979): Monosepalum muricatum (J.J.Sm.) Schltr. (basionym: Bulbophyllum muricatum J.J.Sm.) Tapeinoglossum Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 892 (1913). Lectotype: Bulbophyllum centrosemiflorum Schltr. Hyalosema (Schltr.) Rolfe, Orchid Rev., 27, 130 (1919). Type species: Hyalosema grandiflorum (Blume) Rolfe (basionym: Bulbophyllum grandiflorum Blume) Chaseella Summerh., Kirkia, 1, 88 (1961). Type species: Chaseella pseudohydra Summerh., syn. nov. Canacorchis Guillaumin, Bull. Mus. Natl. Hist. Nat., II, 35, 653 (1964). Type species: Canacorchis lophoglottis Guillaumin (=Bulbophyllum lophoglottis (Guillaumin) Hallé) Hapalochilus (Schltr.) Senghas, Orchidee (Hamburg), 29, 248 (1978). Lectotype (designated by Senghas 1978): Hapalochilus nitidus (Schltr.) Senghas. (=Bulbophyllum nitidum Schltr.) Jejosephia A.N.Rao & Mani, J. Econ. Taxon. Bot., 7, 217 (1985). Type species: Jejosephia pusilla (Joseph & H.Deka) A.N.Rao & K.J.Mani, syn. nov. Ferruminaria Garay, Hamer & Siegerist, Nordic J. Bot., 14, 635 (1994). Type species: Ferruminaria brastagiensis (Carr) Garay, Hamer & Siegerist Mastigion Garay, Hamer & Siegerist, Nordic J. Bot., 14, 635 (1994). Type species: Mastigion appendiculatum (Rolfe) Garay (=Bulbophyllum appendiculatum (Rolfe) J.J.Sm.) Rhytionanthos Garay, Hamer & Siegerist, Nordic J. Bot., 14, 637 (1994). Type species: Rhytionanthos cornutum (Lindl.) Garay, 5

DENDROBIEAE

Hamer & Siegerist (basionym: Cirrhopetalum cornutum Lindl. (=Bulbophyllum helenae (Kuntze) J.J.Sm.)) Synarmosepalum Garay, Hamer & Siegerist, Nordic J. Bot., 14, 639 (1994). Type species: Synarmosepalum kittredgei Garay, Hamer & Siegerist (=Bulbophyllum kittredgei (Garay, Hamer & Siegerist) J.J.Verm.) Vesicisepalum Garay, Hamer & Siegerist, Nordic J. Bot., 14, 641 (1994). Type species: Vesicisepalum folliculiferum (J.J. Sm.) Garay, Hamer & Siegerist (basionym: Bulbophyllum folliculiferum J.J.Sm.) Oncophyllum D.L.Jones & M.A.Clem., Orchadian, 13, 420 (2001). Type species: Oncophyllum minutissimum (F.Muell.) D.L.Jones & M.A.Clem. (basionym: Bulbophyllum minutissimum F.Muell.) Peltopus (Schltr.) Szlach. & Marg., Polish Bot. J., 46, 114 (2001). Type species: Peltopus greuterianus Szlach. & Marg. (=Bulbophyllum peltopus Schltr.) Blepharochilum M.A.Clem. & D.L.Jones, Orchadian, 13, 499 (2002). Type species: Bulbophyllum purpurascens F.M.Bailey (=Bulbophyllum macphersonii Rupp) Carparomorchis M.A.Clem. & D.L.Jones, Orchadian, 13, 499 (2002). Lectotype (designated by Seidenfaden 1979): Carparomorchis macrantha (Lindl.) M.A.Clem. & D.L.Jones: Bulbophyllum macranthum (Lindl.) Kaurorchis D.L.Jones & M.A.Clem., Orchadian, 13, 499 (2002). Type species: Kaurorchis evasa (T.E.Hunt & Rupp) D.L.Jones & M.A.Clem. (basionym: Bulbophyllum evasum T.E.Hunt & Rupp) Ichthyostomum D.L.Jones, M.A.Clem. & Molloy, Orchadian, 13, 499 (2002). Type species: Ichthyostomum pygmaeum (Sm.) D.L.Jones, M.A.Clem. & Molloy (basionym: Bulbophyllum pygmaeum (Sm.) Lindl.) Fruticicola (Schltr.) M.A.Clem. & D.L.Jones, Orchadian, 13, 499 (2002). Type species: Fruticicola albopunctata M.A.Clem. & D.L.Jones (=Bulbophyllum fruticicola Schltr.) Papulipetalum (Schltr.) M.A.Clem. & D.L.Jones, Orchadian, 13, 500 (2002). Type species: Papulipetalum angustifolium M.A.Clem. & D.L.Jones (=Bulbophyllum papulipetalum Schltr.) Serpenticaulis M.A.Clem. & D.L.Jones, Orchadian, 13, 500 (2002). Type species: Serpenticaulis bowkettiae (F.M.Bailey) M.A.Clem. & D.L.Jones (basionym: Bulbophyllum bowkettiae F.M.Bailey) Spilorchis D.L.Jones & M.A.Clem., Orchadian, 15, 37 (2005). Type species: Spilorchis weinthalii (R.S.Rogers) D.L.Jones & M.A.Clem. (basionym: Bulbophyllum weinthalii R.S.Rogers) Hamularia Aver. & Averyanova, Komarovia, 4, 18 (2006). Type species: Hamularia puluongensis Aver. & Averyanova (=Bulbophyllum griffithii Rchb.f.), syn. nov. Lepanthanthe (Schltr.) Szlach., Richardiana, 7, 82 (2007). Type species: Lepanthanthe lepanthiflora (Schltr.) Szlach., syn. nov. (basionym: Bulbophyllum lepanthiflorum Schltr.) Trachyrhachis (Schltr.) Szlach., Richardiana, 7, 85 (2007). Lectotype (designated by Vermeulen 1993): Bulbophyllum barbilabium Schltr. (=Bulbophyllum bulliferum J.J. Sm.), syn. nov.

6

Hordeanthos Szlach., Richardiana, 7, 88 (2007). Type species: Hordeanthos lemniscatus (C.S.P.Parish ex Hook.f.) Szlach. (basionym: Bulbophyllum lemniscatum C.S.P.Parish ex Hook.f.) Tripudianthes (Seidenf.) Szlach. & Kras, Richardiana, 7, 94 (2007). Type species: Tripudianthes tripudians (C.S.P.Parish & Rchb.f.) Szlach. & Kras. (basionym: Bulbophyllum tripudians C.S.P.Parish & Rchb.f.)

Derivation of name From the Greek bolbos, bulb, and phyllon, leaf, referring to the leaf-bearing pseudobulbs. (BG)

Description (Plates 1–22; Fig. 621.1–621.7) Epiphytic, occasionally lithophytic, or rarely terrestrial herbs. Sympodia arising from or (well) above one subterminal nodes from a previous sympodium. Basal part of each sympodium with three to many nodes forming a creeping or patent rhizome enveloped in cataphylls when young. Last node of each sympodium usually swollen into a pseudobulb (rarely with lower nodes swollen as well), sometimes not or hardly swollen, apex 1–3-leaved (6–12-leaved in B. sect. Chaseella). Leaves usually persistent, sometimes deciduous, duplicate, inarticulate, usually petiolate, thinly herbaceous to coriaceous, usually glabrous. Inflorescences usually heteranthous, consisting of modified sympodia sprouting from or (well) above nodes along rhizome, ofter close to pseudobulbs, each with several bract-bearing nodes, one- to many-flowered, apical or along distal part of rhizome. Rachis usually a peduncle, but sometimes swollen into a spindle-shaped body, or widened and bilaterally flattened with flowers either inserted on flat side, or along the edges; usually glabrous, or papillose, rarely pubescent; floral bracts appressed to reflexed, caducous or persistent. Flowers either distichous or spirally arranged, (partly) resupinate or not, opening either simultaneously or in succession, (almost) closed to widely open. Sepals free to adherent to connate (the dorsal to laterals, the laterals along their upper or lower margins, or all three), margins entire to erose to fimbriate, glabrous to papillose to ciliate, surface glabrous to papillose or hirsute; lateral sepals equal to dorsal sepal or distinctly different in length, shape, and surface ornamentation, and fused to column foot along basal part of their lower margins. Petals free, similar to sepals or not. Labellum usually with the base hinged to column foot by a flat strip of tissue allowing free movement of labellum parallel to bilateral plane of symmetry of flower, sometimes (partly) immobile because the strip of tissue is too short or too thick to allow movement, or because the labellum and column foot are fused; undivided to trilobed, margins entire to denticulate to fimbriate, glabrous to papillose to ciliate to vesiculose, adaxially with or without longitudinal ridges, more rarely with transverse ridges. Column usually ending in two stelidia; often winged along lower margins, column foot present; anther connected to apex of column by a thin strip of tissue, bilocular, pollinia two or four, the inner pair as large as the outer or (much) smaller, usually waxy, hard to soft, sometimes with stalks: caudicles, hamular or tegular

BULBOPHYLLUM

B

2 cm

A 4 mm

C G

2 mm

D

I 3 mm

H

E

F Fig. 621.1.  Bulbophyllum antheae (J.J.Verm. & A.L.Lamb) J.J.Verm. (B. sect. Trias). A. Habit; B. Flower; C. Flower analysis, from left to right: dorsal sepal, petal, lateral sepal, labellum; D. Labellum, upper surface; E. Labellum, lower surface; F. Column and labellum, side view; G. Anther, ventral view; H. Anther, dorsal view; I. Pollinia, one pair (left), two pairs (right). Drawn by J. J. Vermeulen from Tenom Orchid Centre TOC cult. 2600 (L).

stipes; stigma concave, transversely elliptic to a longitudinal slit, separated from anther by seam-like, thin or fleshy rostellum; viscidium usually inconspicuous, semi-liquid. Ovary and pedicel glabrous or covered in hairs; node at the base of pedicel level with attachment of subtending bract or (well) above it. Capsule obovoid to ellipsoid to subglobose, stalked or not, sometimes winged or triangular in cross-section. (BG, JJV)

to the Neotropics (about 100 species). The main centres of diversity are Madagascar (200 species) and New Guinea (600 species). (BG)

Distribution (Fig. 621.8)

Artificial key to the continental African sections of Bulbophyllum (BG, JJV)

Bulbophyllum comprises about 2200 species and is widely distributed from continental tropical Africa, the Comoros, Madagascar, the Seychelles, Reunion and Mauritius (about 200 species), India, Nepal, Bhutan, China, Taiwan, Thailand, Malaysia, Indonesia, the Philippines, New Guinea, Australia, New Zealand and the tropical Pacific islands as far east as Tahiti (about 1700 species)

Infrageneric treatments african sections:

 1. Pseudobulbs 6–12-leaved at apex . . . . B. sect. Chaseella Pseudobulbs 1–2-leaved at apex . . . . . . . . . . . . . . . . . . . . 2 ● ●

 2. Labellum attached to column foot by a thick strip of tissue and hence immobile . . . . . . . . 7. B. sect. Genyorchis ●

7

DENDROBIEAE

H

C

1 mm

I

6 mm

B

D

A

3 mm

F

2 mm

G

E

5 cm

Fig. 621.2.  Bulbophyllum calyptratum Kränzl. var. calyptratum. (B. sect. Megaclinium). A, B. Habit; C. Flower; D. Flower analysis, from left to right: dorsal sepal, petal, lateral sepal, labellum; E. Labellum, upper surface (left), lower surface (right); F. Column and labellum, side view; G. Column; H. Anther, ventral view (left), dorsal view (right); I. Pollinia, one pair (left), two pairs (right). Drawn by J. J. Vermeulen from FHI 42079 (K), Deighton 5026 (K), and Segerbaeck 1182 (K). Reproduced with permission from Orchid Monographs, 2 (1987).

8

BULBOPHYLLUM

G

H

B 3 mm

1 mm

D

3 mm

C

E

3 mm

A

F

3 cm

Fig. 621.3.  Bulbophyllum osyricera Schltr. (B. sect. Brachystachyae). A. Habit; B. Flower; C. Flower analysis, from left to right: dorsal sepal, petal, lateral sepal, labellum; D. Labellum, upper surface; E. Labellum, lower surface; F. Column and labellum, side view; G. Anther, ventral view (above) dorsal view (below); H. Pollinia. Drawn by J. J. Vermeulen from Lamb 2008/1295 (SAN).



Labellum attached to a thin, flexible strip of tissue, hence mobile, if not kept (partly) in position by other flower parts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3  3. Inflorescence a subumbellate raceme; pedicel plus ovary twice as long as rachis or longer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Cirrhopetalum Inflorescence an elongated raceme; pedicel plus ovary of the lowermost flower equal to length of the rachis or shorter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 ●

●

●

 4. Raceme with distichous flowers or all inflorescences with 1–3 flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 ●



Raceme with spirally arranged flowers; inflorescence with 4 flowers or more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5  5. Rachis slightly to distinctly swollen compared to peduncle, spindle-shaped, cylindrical or obovoid . . . . . . . . . . . 6 Rachis about as thick as peduncle . . . . . . . . . . . . . . . . . . . 7  6. Labellum adaxially hirsute; petals obovate to spatulate, adaxially papillose toward apex . . . . . . . . B. sect. Comata Labellum adaxially glabrous; petals ovate-oblong to narrowly triangular to linear, adaxially glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Lupulina  7. Pseudobulbs 1-leaved at apex . . . . . . . . . . . . . . . . . . . . . . 8 ●

●

● ●

●

●

9

DENDROBIEAE

B

10 mm

H

1 mm

C

D

4 mm

F

G

E

A 3 cm Fig. 621.4.  Bulbophyllum sannio J.J.Verm. (B. sect. Polymeres). A. Habit; B. Flower analysis, from left to right: dorsal sepal, petal, lateral sepal, labellum; C. Labellum, upper surface; D. Labellum, lower surface; E. Column and labellum, side view; F. Anther, ventral view; G. Anther, dorsal view; H. Pollinia, one pair (above), two pairs (below). Drawn by J. J. Vermeulen from Jongejan 1028 (L). Reproduced with permission from Nordic Journal of Botany, 26 (2008).

10

BULBOPHYLLUM

B

8 mm

F

G

1 mm

A C

5 mm

E 81

5 mm

5 mm

D

Fig. 621.5.  Bulbophyllum schinzianum Kränzl. var. phaeopogon (Schltr.) J.J.Verm. (B. sect. Ptiloglossum). A. Habit; B. Flower; C. Flower analysis, from left to right: dorsal sepal, petal, lateral sepal, labellum; D. Labellum, upper surface (left), lower surface (right); E. Column and labellum, side view; F. Anther, dorsal view (left), ventral view (right); G. Pollinia, one pair (left), two pairs (right). Drawn by J. J. Vermeulen from Le Testu 5178 (K). Reproduced with permission from Orchid Monographs, 2 (1987).



●

Pseudobulbs 2-leaved at apex . . . . . . B. sect. Denticulata

 8. Dorsal sepal distinctly wider than lateral sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Lupulina Dorsal sepal about as wide as lateral sepals or narrower . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Ptiloglossum ●

●

 9. Pseudobulbs 2-leaved at apex . . . . . . . . . . . . . . . . . . . . . 10 Pseudobulbs 1-leaved at apex . . . . . . . . . . . . . . . . . . . . . 14 ● ●

10. Lateral sepals reflexed at base and appressed against rachis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Bifaria ●

11

DENDROBIEAE

3 cm 5 mm

A

B

C

2 mm

E

5 mm

D

G

F 1 mm

Fig. 621.6.  Bulbophyllum tindemansianum J.J.Verm., de Vogel & A.Vogel. (B. sect. Epicranthes). A. Habit; B. Flower; C. Flower analysis, petal (above); from left to right (below): dorsal sepal, lateral sepal, labellum; D. Labellum, upper surface (left), lower surface (right); E. Column and labellum, side view; F. Anther, ventral view (left), dorsal view (right); G. Pollinia, two pairs (left), one pair (right). Drawn by J. J. Vermeulen from Leiden cult. 20081503 (L). Reproduced with permission from Blumea, 55 (2010).



Lateral sepals porrect to spreading, or lateral sepals reflexed well above base and not appressed against rachis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11. Anther abaxially with a dorsiventrally flattened, rounded, truncate to bifid protrusion anteriorly, with flat sides or with one (deeply) concave side that distinctly overtops anterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Anther abaxially without a protrusion, or with a protrusion that does not overtop anterior margin, or with a protrusion that overtops anterior margin but is not dorsiventrally flattened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 ●

●

●

12

12. Rachis 2- or 3-flowered; dorsal sepal 8.2–10.0 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Gilgiana Rachis 5–64-flowered; dorsal sepal 2.0–6.2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Oreonastes 13. Labellum margins ciliate or ciliolate, or glabrous and labellum thick and firm; rachis not thickened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Ptiloglossum Labellum margins glabrous; labellum usually thick, but soft and easy to compress, thin when dry; rachis often thickened and flattened, with the flowers inserted along midvein . . . . . . . . . . . . . . . . . . . . . . . B. sect. Megaclinium ●

●

●

●

BULBOPHYLLUM

A C

5 cm

5 mm

B G

1 mm

F

D

E

2 mm

Fig. 621.7.  Bulbophyllum vaginatum (Lindl.) Rchbf. (B. sect. Recurvae). A. Plant; B. Flower; C. Flower analysis, from left to right: dorsal sepal, petal, lateral sepal, labellum; D. Labellum, upper surface (left), lower surface (right); E. Column and labellum, side view; F. Anther, ventral view (above), dorsal view (below); G. Pollinia, one pair (above), two pairs (below). Drawn by J. J. Vermeulen from from Vermeulen 658 (L).

14. Rachis not or slightly thickened, orbicular in section, without edges, or rachis widened and flattened, with rounded edges; flowers arising from midvein of rachis . . . . . . . . 15 Rachis widened and flattened, sharply four-edged in section with two narrow, concave sides; flowers arising from narrow, concave sides of rachis . . . . . . . . . . . . . . . . . . . . 16 15. Rachis widened and flattened with rounded edges or not or slightly thickened, orbicular in section, without edges; labellum usually thick, but soft and easy to compress, thin when dry . . . . . . . . . . . . . . . . . . . . . . B. sect. Megaclinium ●

●



Rachis not thickened, orbicular in section, without edges; labellum thick and firm . . . . . . . . . . B. sect. Ptiloglossum 16. Stelidia 1.0–1.2 mm long, acuminate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Carnosisepala Stelidia 0.5 mm long or shorter, rounded to acute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Oreonastes ●

●

●

●

Bulbophyllum sect. Bifaria G.A.Fischer & J.J.Verm., in prep. Type species (proposed): Bulbophyllum bifarium Hook. 13

DENDROBIEAE

Epiphytes or lithophytes with creeping rhizomes. Pseudobulbs 2-leaved. Inflorescence a 6–30-flowered, elongate raceme with distichous flowers; rachis not or hardly thickened, sharply fouredged in section with two concave sides from which the flowers arise. Floral bracts as long as or longer than flowers. Flowers non-resupinate. Sepals free, glabrous abaxially, the lateral sepals reflexed at the base and appressed against the rachis. Labellum mobile, undivided, margins glabrous; thick and solid. Anther abaxially with a conical protrusion overtopping anterior margin; pollinia 4. Three species in montane forests up to 2050 m in Cameroon, Guinea, Kenya, and Liberia. (JJV)

Bulbophyllum sect. Comata G.A.Fischer & J.J.Verm., in prep. Type species (proposed): Bulbophyllum comatum Lindl. Epiphytes with creeping rhizomes. Pseudobulbs 1-leaved. Inflorescence a many-flowered, elongate raceme with spirally arranged flowers; rachis swollen, spindle-shaped, cylindrical or obovoid. Floral bracts about as long as the flowers. Flowers resupinate. Sepals free, with long hairs abaxially. Labellum mobile, auriculate near base, thick, solid, margins and adaxial surface hirsute. Anther abaxially without a protrusion; pollinia 4. A single species in forests up to 1300 m in Equatorial Guinea, Gabon, Ivory Coast, Liberia, Nigeria, Sierra Leone, and Rwanda. (JJV)

Bulbophyllum sect. Carnosisepala G.A.Fischer & J.J.Verm., in prep. Type species (proposed): Bulbophyllum carnosisepalum J.J.Verm. Epiphytes with creeping rhizomes. Pseudobulbs 1-leaved. Inflorescence a 3–12-flowered, elongate raceme with distichous flowers; rachis thickened, sharply four-edged in section with two concave sides from which flowers arise. Floral bracts shorter than flowers. Flowers non-resupinate. Sepals free, glabrous abaxially. Labellum mobile, undivided, margins glabrous; thick and solid. Stelidia 1.0–1.2 mm long, acuminate. Anther with an abaxial, dorsiventrally flattened protrusion overtopping the anterior margin; pollinia 4. One species in lowland and montane forests in Cameroon, Gabon, Ivory Coast, Uganda, and Zaire. (JJV)

Bulbophyllum sect. Denticulata G.A.Fischer & J.J.Verm., in prep. Type species (proposed): Bulbophyllum denticulatum Rolfe Epiphytes with creeping rhizomes. Pseudobulbs 2-leaved. Inflorescence a many-flowered, elongate raceme with spirally arranged flowers; rachis not thickened, round in section. Floral bracts shorter than flowers. Flowers resupinate. Sepals free, finely papillose abaxially. Labellum mobile, undivided, margins glabrous; thick, solid. Anther abaxially with a rounded protrusion overtopping the anterior margin; pollinia 4. A single species in montane forests up to 900 m in Ivory Coast, Liberia, and Sierra Leone. (JJV)

Bulbophyllum sect. Chaseella (Summerh.) G.A.Fischer & J.J.Verm., in prep. Type species (proposed): Bulbophyllum pseudohydra (Summerh.) G.A.Fischer & J.J.Verm. Epiphytes with creeping rhizomes. Pseudobulbs 6–12-leaved, needle-shaped. Inflorescence 1(–2)-flowered; rachis, if present, not thickened, round in section. Floral bracts shorter than the flowers. Flowers resupinate. Sepals free, glabrous abaxially. Labellum mobile, undivided, margins glabrous; thick and solid. Anther with an abaxial, rounded protrusion slightly overtopping the anterior margin; pollinia 2. A single species in montane forests up to 2000 m in Kenya, Zambia, and Zimbabwe. (JJV)

Bulbophyllum sect. Genyorchis (Lindl.) G.A.Fischer & J.J.Verm., in prep. Type species (proposed): Bulbophyllum apetalum Lindl. Epiphytes with creeping rhizomes. Pseudobulbs 1- or 2-leaved. Inflorescence a 5–15-flowered, elongate raceme with distichous flowers; rachis not thickened, round in section. Floral bracts shorter than the flowers. Flowers non-resupinate. Sepals free, glabrous abaxially. Labellum firmly attached to the column foot, undivided or distally auriculate, margins glabrous; thick, solid. Anther abaxially with a protrusion not overtopping the anterior margin; pollinia 4, stipitate. Seven species in mangrove and tropical dry forests up to 1400 m in Cameroon, Congo, Gabon, Ghana, Guinea Islands, Nigeria, Ivory Coast, Sierra Leone, and Zaire. (JJV)

Bulbophyllum sect. Cirrhopetalum (Lindl.) Rchb.f., Walpers Ann. Bot. Syst., 6, 259 (1861). Type species: Bulbophyllum longiflorum Thouars Epiphytes with creeping rhizomes. Pseudobulbs 1-leaved. Inflorescence a many-flowered, subumbellate raceme with flowers spirally arranged; rachis thickened, round in section. Floral bracts shorter than the flowers. Flowers resupinate. Dorsal sepal free; lateral sepals twice to 5.5 times as long as the dorsal, twisted near the base so that the upper margins turn inwards, adnate along their upper margins. Labellum mobile, undivided, margins glabrous; thick and solid. Anther abaxially with a small protrusion not overtopping the anterior margin; pollinia 4. One species in montane forests up to 1800 m in Zaire, Tanzania, Malawi, Zimbabwe, Madagascar and surrounding islands and Asia eastward to the Pacific. (JJV) 14

Bulbophyllum sect. Gilgiana G.A.Fischer & J.J.Verm., in prep. Type species (proposed): Bulbophyllum gilgianum Kränzl. Epiphytes with creeping rhizomes. Pseudobulbs 2-leaved. Inflorescence a 1–3-flowered, elongate raceme with spirally arranged flowers; rachis slightly swollen, spindle-shaped. Floral bracts shorter than flowers. Flowers non-resupinate. Sepals free, sparsely hirsute abaxially. Labellum mobile, undivided, margins ciliate; thick and solid. Anther abaxially with a dorsiventrally flattened protrusion overtopping the anterior margin; pollinia 4. One species in montane forests up to 1900 m in Tanzania. (JJV) Bulbophyllum sect. Lupulina G.A.Fischer, in prep. Type species (proposed): Bulbophyllum occultum Thouars

BULBOPHYLLUM

Epiphytes with creeping rhizomes. Pseudobulbs 1- or 2-leaved. Inflorescence a 4- to many-flowered, elongate raceme with ­spirally arranged flowers; rachis not thickened, round in section to spindle-shaped. Floral bracts shorter than or about as long as the flowers. Flowers non-resupinate. Sepals free, glabrous abaxially. Labellum mobile, undivided, margins entire to fimbriate; thick and solid. Anther with an abaxial, rounded protrusion that does not overtop the margin, to a dorsiventrally flattened and bifid one overtopping the anterior margin; pollinia 4. Three species in savanna woodlands and montane forests up to 1800 m in Burundi, Tanzania, Malawi, South Africa, Zaire, Zambia, Zimbabwe, and Madagascar. Note that B. lupulinum is not included here but rather in B. sect. Oreonastes. (JJV) Bulbophyllum sect. Megaclinium G.A.Fischer & J.J.Verm., in prep. Type species (proposed): Bulbophyllum falcatum (Lindl.) Rchb.f. Epiphytes or lithophytes with usually creeping rhizomes (patent in B. fayi J.J.Verm.). Pseudobulbs 1- or 2(–3)-leaved. Inflorescence a 2- to many-flowered, elongate raceme with distichous flowers; rachis usually thickened and flattened, with the flowers arranged along the midvein, sometimes not thickened, round in section. Floral bracts (slightly) shorter than flowers. Flowers resupinate or non-resupinate. Sepals free, glabrous to hirsute abaxially. Labellum mobile, undivided, margins entire to denticulate to lacerate, usually thick, soft and easy to compress. Anther without an abaxial protrusion or with a rounded protrusion that may overtop the anterior margin; pollinia 4. Nineteen species in mangrove forests, savanna woodland, plantations, lowland and montane forests up to 2300 m in Angola, Cameroon, Congo, Equatorial Guinea, Ethiopia, Gabon, Ghana, Guinea, Ivory Coast, Kenya, Liberia, Malawi, Mozambique, Nigeria, Sierra Leone, South Africa, Tanzania, Togo, Uganda, Zaire, Zambia, and Zimbabwe. (JJV)

Epiphytes, occasionally lithophytes or terrestrials, with creeping rhizomes. Pseudobulbs 1- or 2-leaved. Inflorescence 1-flowered or a 2- to many-flowered, elongate to contracted raceme with spirally or distichously arranged flowers; rachis not or hardly thickened, usually round in section, sometimes slightly angular. Floral bracts usually shorter than or as long as the flowers (longer than the flowers in B. pandanetorum Summerh.). Flowers resupinate or not. Sepals free, glabrous abaxially. Labellum mobile, undivided (trilobed in B. nigritianum Rendle); margins glabrous to long-ciliate; thick and solid at least near the base. Anther without an abaxial protrusion or with a rounded protrusion that may overtop the anterior margin; pollinia 2 or 4. Thirty species in savanna woodlands, mangrove, lowland and montane forests up to 2400 m in Angola, Burundi, Cameroon, Central African Republic, Congo, Equatorial Guinea, Ethiopia, Gabon, Ghana, Guinea, Ivory Coast, Kenya, Liberia, Malawi, Mozambique, Nigeria, Rwanda, Sierra Leone, Tanzania, Uganda, Zaire, Zambia, and Zimbabwe. (JJV) malagasy sections:

Artificial key to the Malagasy sections of Bulbophyllum (GF, JJV):  1. Pseudobulbs 1-leaved at apex . . . . . . . . . . . . . . . . . . . . . . 2 Pseudobulbs 2-leaved at apex . . . . . . . . . . . . . . . . . . . . . 11  2. Labellum margin partly or entirely ciliate, fimbriate or with mobile appendages . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Labellum margin entire, glabrous to papillose . . . . . . . . . 5  3. Labellum margin ciliate . . . . . . . B. sect. Pantoblepharon Labellum margin fimbriate or with mobile appendages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4  4. Flowers with labellum turned away from rachis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Kinethrix Flowers with labellum facing rachis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Lupulina  5. Lateral sepals connivent to connate along their lower margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Lateral sepals free, lower margins approximately diverging . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7  6. Lateral sepals connivent to connate along their lower margins; raceme elongate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Ploiarium Lateral sepals connivent to connate along their upper margins; raceme subumbellate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Cirrhopetalum (see African species)  7. Pseudobulbs bilaterally flattened . . . B. sect. Polyradices Pseudobulbs globose, ellipsoid, ovoid or dorsiventrally flattened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8  8. Apical part of labellum incurved, adaxial side deeply concave . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Elasmatopus Apical part of labellum porrect, or recurved, adaxial side flat or convex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9  9. Inflorescence 6–12-flowered; raceme contracted, globose . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Lyperocephalum Inflorescence 1–3-flowered; raceme elongate . . . . . . . . 10 ● ●

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Bulbophyllum sect. Oreonastes G.A.Fischer & J.J.Verm., in prep. Type species (proposed): Bulbophyllum oreonastes Rchb.f. Epiphytes or lithophytes with creeping rhizomes (patent in B. teretifolium Schltr.). Pseudobulbs 1- or 2-leaved. Inflorescence a many-flowered, elongate raceme with flowers arranged distichously; rachis thickened, sharply four-edged in section with 2 narrow, concave sides from which the flowers arise. Floral bracts shorter or longer than the flowers. Flowers resupinate or not. Sepals free, glabrous to papillose to sparsely hirsute abaxially. Labellum mobile, undivided or trilobed, margins entire to erose to denticulate; thick and solid. Stelidia 1.0–1.2 mm long, acuminate. Anther abaxially with a dorsiventrally flattened protrusion overtopping the anterior margin; pollinia 4. Ten species in lowland rain forests, plantations, semideciduous and montane forests up to 2300 m in Angola, Burundi, Cameroon, Ethiopia, Equatorial Guinea, Gabon, Ghana, Guinea, Ivory Coast, Kenya, Liberia, Malawi, Mozambique, Nigeria, Rwanda, Sierra Leone, South Africa, Tanzania, Uganda, Zaire, Zambia, and Zimbabwe. (JJV)

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Bulbophyllum sect. Ptiloglossum Lindl., J. Linn. Soc. Bot., 6, 125 (1862). Type species: Bulbophyllum barbigerum Lindl.

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DENDROBIEAE

10. Inflorescence 3- to many-flowered; or inflorescence 1- or 2-flowered and labellum glabrous adaxially . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Bulbophyllum Inflorescence 1- or 2-flowered; labellum papillose adaxially . . . . . . . . . . . B. sect. Pantoblepharon (B. muscicola) 11. Lateral sepals connivent to connate along their lower margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Lateral sepals free, lower margins approximately diverging . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 12. Labellum oblong, elliptic, (ob-)ovate to spatulate with a widened base . . . . . . . . . . . . . . . . . . . . . B. sect. Ploiarium Labellum spatulate, gradually narrowing towards base . . . . . . . . . . B. sect. Bulbophyllum (B. brachystachyum) 13. Inflorescence 1-flowered; vegetative shoots usually developing from basal node of pseudobulb, then fused to it for more than half the length of the latter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Lichenophylax Inflorescence 1- to many-flowered; if 1-flowered then vegetative shoots not fused to the pseudobulb above the node from which they arise . . . . . . . . . . . . . . . . . . . . . . . 14 14. Pseudobulbs distinctly laterally flattened, with two distinct edges (other edges inconspicuous only if present); or inflorescence developing sinultaneously with the young shoot from which it arises; or both characters present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 Pseudobulbs not or only slightly laterally flattened, with three or more distinct ridges; inflorescence developing from (almost) mature shoots . . . . . . . . . . . . . . . . . . . . . . 18 15. Inflorescence developing sinultaneously with the young shoot from which it arises . . . . . . . . . . . . . . . . . . . . . . . . 16 Inflorescence developing from (almost) mature shoots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 16. Rachis with lowermost bracts 12 mm or longer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Alcistachys Rachis with lowermost bracts 7 mm or shorter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Pachychlamys 17. Flowers with labellum turned away from rachis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Inversiflora Flowers with labellum facing rachis . . . B. sect. Lupulina 18. Labellum margins fimbriate . . . . . . . . . . . . . . . . . . . . . . . 19 Labellum margins entire to erose (glabrous to papillose or ciliate) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 19. Flowers with labellum turned away from rachis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Kinethrix Flowers with labellum facing rachis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Lupulina 20. Labellum adaxially with two distinct ridges with an erose edge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Moratii Labellum adaxially without ridges, or with ridges with an entire edge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 21. Inflorescence many-flowered; rachis distinctly thickened as compared to peduncle and spindle-shaped, with flowers inserted in three longitudinal rows . . . . . . . . . . . . . . . . . . . . B. sect. Lupulina (B. senghasii) ●



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Inflorescence 1-flowered; rachis not thickened or rachis thickened and with flowers distichously or spirally arranged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 Base of labellum with two acicular teeth (inflorescence with one or two flowers, or flowers distichous) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Bifalcula Base of labellum without acicular teeth (inflorescence with one or two flowers, or flowers spirally arranged) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 Adaxial side of labellum with a cavity near apex or apical part of labellum curved upwards, or both characters combined present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 Adaxial side of labellum flat or convex near the apex; apical part of the labellum porrect or curved downwards . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 Rostellum protruding beyond stelidia apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Ikongoense Rostellum receding in between stelidia apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Elasmatopus Rhizome scales thick, fibrous, the apices patent when pseudobulb reaches maturity . . . . . B. sect. Pachychlamys Rhizome scales thin, membranous, the remnants weathering to woolly hairs or clinging to pseudobulb at maturity . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Bulbophyllum

Bulbophyllum sect. Alcistachys Schltr., Feddes Repert. Spec. Nov. Regni Veg. Beih., 33, 224–6 (1925). Type species: Bulbophyllum occlusum Ridl. Epiphytes or lithophytes with creeping rhizomes. Pseudobulbs close together, distinctly laterally flattened or not, two-leaved. Inflorescences synanthous, a many-flowered, elongate raceme with spirally arranged flowers; rachis not thickened, round in section. Lower floral bracts 12 mm long or longer. Flowers resupinate. Sepals free, glabrous abaxially. Petal margins entire to distally erose, glabrous to distally papillose. Labellum mobile, undivided, thick, margins entire, glabrous or (partially) papillose to ciliate. Column with triangular, acute stelidia (obtuse in B. occlusum), without a tooth along the lower margin. Anther with a rounded to high, conical protrusion that overtops the anterior margin. Six species in lowland to high-elevation moist forests in Madagascar, the Mascarenes, and the Comoros. The flowers smell of rotting meat, faeces, or decaying fish. (GF, JJV)

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Bulbophyllum sect. Bifalcula Schltr. Feddes Repert. Spec. Nov. Regni Veg. Beih., 33, 215–16 (1925). Type species (proposed): Bulbophyllum implexum Jum. & H.Perrier Epiphytes or lithophytes with creeping rhizomes. Pseudobulbs clustered to distant, not laterally flattened, 2-leaved. Inflorescence heteranthous, 1-flowered or a 2- to many-flowered, elongate raceme with distichously arranged flowers; rachis not thickened, round in section. Flowers resupinate. Sepals free, glabrous abaxially. Petal margins entire, glabrous. Labellum mobile, with aciculate auricles near the base, thick, margins entire, glabrous, adaxial side without ridges, or with ridges with an entire edge. Column

BULBOPHYLLUM

with triangular, acute stelidia, without a tooth along the lower margin. Anther with a rounded protrusion that (slightly) overtops the anterior margin. Four species, with several undescribed, in dwarf forests on poor soils along the humid coast of northwestern and eastern Madagascar. (GF, JJV) Bulbophyllum sect. Bulbophyllum G.A.Fischer & J.J.Verm., in prep. Type species (proposed): Bulbophyllum nutans (Thouars) Thouars Epiphytes or lithophytes with creeping rhizomes. Pseudobulbs 1- or 2-leaved, green, red or yellow, often surrounded by cottony white fibres, densely clustered or well spaced on rhizome. Leaves thin, coriaceous, sometimes succulent, oblong or lanceolate. Inflorescence many-flowered, erect, terete but never thickened in some species, setaceous. Flowers resupinate, glabrous, often white or yellowish-brownish, rarely red. Sepals and petals never fimbriate, often transparent. Labellum glabrous, mostly white but also red or yellow in a few species, curved-geniculate, never fimbriate. Column foot short or free, part of column foot elongate. Stelidia rarely as long as the anther. Pollinia 4. At least 32 species, in humid forests at all elevations in Madagascar, the Mascarenes, and the Comoros. Bulbophyllum sect. Elasmotopus Schltr., Feddes Repert. Spec. Nov. Regni Veg. Beih., 33, 211–15 (1925). Type species (proposed): Bulbophyllum oxycalyx Schltr. Epiphytes or lithophytes with creeping rhizomes. Pseudobulbs distant, not laterally flattened, 1- or 2-leaved. Inflorescence heteranthous, 1- to many-flowered, elongate raceme with spirally arranged flowers; rachis not thickened, round in section. Flowers resupinate. Sepals free, glabrous abaxially. Petal margins entire, glabrous to distally minutely papillose. Labellum mobile, undivided, thick, or with a thick basal part and a thin apical part, apical part with an adaxial cavity near the apex or apical part of the labellum curved upwards or with both characters combined; margins entire, glabrous or partly ciliate, adaxial side without ridges, or with inconspicuous ridges with an entire edge. Column with deltoid to triangular, acute stelidia, with or without a tooth along the lower margin. Anther with a rounded protrusion that overtops the anterior margin. Ten species in humid forests at all elevations in Madagascar. Most (not all) species have 3-veined petals; B. sect. Pachychlamys always has 1-veined petals. (GF, JJV) Bulbophyllum sect. Ikongoense G.A.Fischer, in prep. Type species (proposed): Bulbophyllum ikongoense H.Perrier Epiphytes with creeping rhizomes. Pseudobulbs distant, not or only slightly laterally flattened, 2-leaved. Inflorescence heteranthous, a many-flowered, elongate raceme with spirally arranged flowers; rachis thickened, spindle-shaped. Flowers resupinate. Sepals free, glabrous abaxially. Petal margins entire, glabrous. Labellum mobile, undivided, thin, or apical part of the labellum curved upwards, concave; margins entire, glabrous, adaxial side without ridges. Column with stelidia, with the rostellum protruding in between, deltoid, obtuse, without a tooth along the lower

margin. Anther without an abaxial protrusion. A single species in montane forests (1000–1400 m) on the eastern edge of the central plateau of Madagascar. (GF, JJV) Bulbophyllum sect. Inversiflora G.A.Fischer & P.J.Cribb, in prep. Type species (proposed): B. cardiobulbum Bosser Epiphytes with creeping rhizomes. Pseudobulbs close together, distinctly laterally flattened, 2-leaved. Inflorescences heteranthous, a many-flowered, elongate raceme with spirally arranged flowers; rachis thickened, spindle-shaped. Flowers non-resupinate, reflexed, with the labellum turned away from the rachis. Sepals free, glabrous abaxially. Petal margins entire, glabrous. Labellum mobile, undivided, thick, margins entire, glabrous or partly ciliate. Column with triangular, acute stelidia, without a tooth along the lower margin. Anther with a rounded to conical protrusion that overtops the anterior margin. Two species in open or dwarf montane forests on poor soils (800–1400 m) in eastern Madagascar. Ants often live underneath the pseudobulbs, which are appressed to the substrate, not unlike the leaves of many Dischidia species (Apocynaceae). (GF, JJV) Bulbophyllum sect. Kinethrix Schltr., Feddes Repert. Spec. Nov. Regni Veg. Beih., 15, 329 (1910). Type species (proposed): Bulbophyllum mirificum Schltr. Epiphytes with creeping rhizomes. Pseudobulbs distant, not or hardly laterally flattened, 1- or 2-leaved. Inflorescences heteranthous, a many-flowered, elongate raceme with spirally arranged flowers; rachis thickened or not, round in section. Flowers resupinate, reflexed, with the labellum turned away from the rachis. Sepals free, glabrous abaxially. Petal margins entire, glabrous or slightly papillose distally. Labellum mobile, undivided, thick, margins fimbriate or with mobile appendages. Column with triangular to ligulate, acute stelidia, with a distinct tooth along the lower margin, close to the base. Anther with a rounded to conical protrusion that overtops the anterior margin. Eleven species in humid mid- to high elevation, moist montane forests in Madagascar and the Mascarenes. The petals are 1/3 as long as the dorsal sepal or shorter. All species are self-incompatible (Fischer, unpublished data). (GF, JJV) Bulbophyllum sect. Lichenophylax Schltr., Feddes Repert. Spec. Nov. Regni Veg. Beih., 33, 193–7 (1925). Type species: Bulbophyllum lichenophylax Schltr. Epiphytes or lithophytes, resembling a moss in size. Rhizome ramified and creeping, nodes slightly swollen; sheaths tubular, membranous, covering new growth and pseudobulb bases, disintegrating when old. Pseudobulbs green, 2-leaved, often laterally flattened. Leaves green, elliptic, acute, thin and soft. Inflorescence 1-flowered; peduncle erect, not swollen, setaceous, with sheaths, the flower terminal. Flowers resupinate, with sepals acute to longly caudate, free, 3-veined, margins entire to finely fimbriate. Petals margins entire or fimbriate, one-veined, surface occasionally finely papillose. Labellum fleshy, often swollen, 17

DENDROBIEAE

glabrous or papillose, margins entire or fimbriate. Column often with a tooth in its dorsal margin. Stelidia truncate, triangular or hook-shaped. Pollinia 4. At least 17 species in low- to high-­ elevation, moist forests in Madagascar. (GF, JJV) Bulbophyllum sect. Lupulina G.A.Fischer, in prep. Type species (proposed): Bulbophyllum occultum Thouars Epiphytes with pseudobulbs 1- or 2-leaved, densely clustered or well spaced, often laterally flattened. Leaves succulent, oblong or lanceolate. Inflorescence erect or pendent, racemose, terete; rachis as thick as the peduncle or swollen with flowers inserted into dimples. Flowers resupinate, in rows or alternately arranged. Sepals free, margins glabrous or fimbriate, surface glabrous or papillate. Petals oblong, falcate, hastate or narrowly triangular, margins entire, fimbriate or lacerate. Labellum thick, recurved in the middle, margins glabrous, lacerate or fimbriate, surface glabrous, papillate or papillose. Column fleshy. Stelidia narrowly triangular, falcate or acicular. Anther with a protrusion often as long or longer as the stelidia; pollinia 4. Eighteen species in Madagascar, the Mascarenes, and the Comoros. Bulbophyllum humblotii Rolfe and B. pusillum Thouars also occur in Africa. Bulbophyllum malawiense B.Morris occurs only in Africa. The species occur in dry forests, savanna woodlands, deciduous and semi-deciduous forests as well as in humid coastal and montane rain forests. (GF) Bulbophyllum sect. Lyperocephalum Schltr. Feddes Repert. Spec. Nov. Regni Veg. Beih., 33, 182 (1925). Type species: Bulbophyllum lyperocephalum Schltr. Epiphytes with creeping rhizomes. Pseudobulbs distant, not flattened, 1-leaved. Inflorescence heteranthous, a many-flowered, congested, capitate raceme with spirally arranged flowers; rachis not thickened, round in section. Flowers resupinate. Sepals free, glabrous abaxially. Petal margins entire, glabrous. Labellum mobile, undivided, thick, margins entire, glabrous to slightly papillose. Column with triangular, acute stelidia, with a tooth along the lower margin. Anther abaxially with a protrusion that does not overtop the anterior margin. A single species in humid lowland and montane forests in Madagascar. (GF, JJV) Bulbophyllum sect. Moratii G.A.Fischer, in prep. Type species (proposed): Bulbophyllum moratii Bosser Epiphytes with creeping rhizomes. Pseudobulbs distant, not laterally flattened, 2-leaved. Inflorescence heteranthous, a manyflowered, elongate raceme with spirally arranged flowers; rachis not thickened, round in section. Flowers resupinate. Sepals free, glabrous abaxially. Petals with entire margins, slightly erose towards the base, glabrous. Labellum mobile, undivided, thick, margins entire, glabrous to slightly papillose; adaxial side with two distinct ridges with an erose edge. Column with triangular, acute stelidia, without a tooth along the lower margin. Anther with an abaxial protrusion that overtops the anterior margin. A single species in wet lower montane forests (600–1000 m) in northeastern Madagascar. (GF, JJV) 18

Bulbophyllum sect. Pachychlamys Schltr., Feddes Repert. Spec. Nov. Regni Veg. Beih., 33, 207–9 (1925). Type species: Bulbophyllum pachypus Schltr. Epiphytes with creeping rhizomes, rhizome scales thick and fibrous, the apices patent when the pseudobulb reaches maturity. Pseudobulbs clustered to distant, not laterally flattened, 2-leaved. Inflorescence synanthous or heteranthous, a manyflowered, elongate raceme with spirally arranged flowers; rachis not or hardly thickened, round in section. Lower floral bracts 7 mm long or shorter. Flowers resupinate. Sepals free, glabrous abaxially. Petal margins entire to distally minutely erose, glabrous. Labellum mobile, undivided, thick, apical part porrect or recurved, adaxially flat or convex, margins glabrous, adaxial side without ridges, or with ridges with an entire edge. Column with triangular, acute stelidia, without a tooth along the lower margin (with a tooth in B. multivaginatum Jum. & H.Perrier). Anther with a rounded protrusion that slightly overtops the anterior margin. Ten species in humid lowland and montane forests in Madagascar and the Mascarenes. (GF, JJV) Bulbophyllum sect. Pantoblepharon Schltr., Feddes Repert. Spec. Nov. Regni Veg. Beih., 33, 179–81;184–7 (1925). Type species: Bulbophyllum pantoblepharon Schltr. Epiphytes or lithophytes with creeping rhizomes. Pseudobulbs clustered to distant, flattened or not, 1-leaved. Inflorescence heteranthous, 1- to many-flowered, on an elongate raceme with spirally arranged flowers; rachis not thickened, round in section. Flowers resupinate. Sepals free, glabrous abaxially. Petals with entire to distally erose margins, papillose to ciliate. Labellum mobile, undivided, margins entire, ciliate (papillose in B. muscicola Rchb.f.). Column with triangular, acute stelidia, with or without a tooth along the lower margin. Anther with or without a rounded protrusion that does not overtop the anterior margin. Fifteen species in humid montane forests in Madagascar and the Mascarenes. (GF, JJV) Bulbophyllum sect. Ploiarium Schltr., Feddes Repert. Spec. Nov. Regni Veg. Beih., 33, 226–48 (1925). Type species: Bulbophyllum coriophorum Ridl. Epiphytes or lithophytes with creeping rhizomes. Pseudobulbs clustered to distant, laterally flattened or not, 1- or 2-leaved. Inflorescence heteranthous, 1- to many-flowered, elongate or congested with spirally arranged flowers; rachis not thickened, or slightly to distinctly thickened, spindle-shaped. Flowers resupinate. Sepals glabrous abaxially; the dorsal sepal free, the lateral sepals connivent to connate along their lower margins. Petals with entire, glabrous to ciliate margins. Labellum mobile, undivided, thick, margins glabrous to papillose. Column with triangular, rounded to acute stelidia, with a tooth along the lower margin. Anther with a rounded protrusion that does not overtop the anterior margin. About 75 species (number uncertain because of several unresolved species complexes) in humid lowland and montane forests in Madagascar, the Mascarenes, and the Comoros. (GF, JJV)

BULBOPHYLLUM

Bulbophyllum sect. Polyradices G.A.Fischer, Sieder & P.J.Cribb, Adansonia, 29, 19–25 (2007). Type species: Bulbophyllum petrae G.A.Fischer, Sieder & P.J.Cribb Epiphytes with membranous, persistent rhizome scales. Pseudobulbs 1-leaved, densely clustered and bilaterally flattened. Leaves coriaceous. Inflorescence synanthous, few-flowered, scattered; peduncle pendent; floral bracts shorter than flowers. Flowers resupinate. Petals finely papillose on margins. Labellum finely papillose beneath. Anther with a protrusion overtopping the anterior margin. A single species in wet lower montane forests up to 1400 m in east-central Madagascar. (GF) asian sections:

Artificial key to the Asian sections of Bulbophyllum (JJV) 1.

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Some or all inflorescences racemose (inflorescence with two or more flowers, the raceme elongate, congested or subumbellate) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 All inflorescences 1-flowered (1-flowered inflorescences may develop in clusters sometimes resembling a raceme) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Basal node of pedicel well above (1/3 the diameter of the pedicel or more) the floral bract attachment in at least some flowers of the inflorescence; raceme congested or elongate, pedicellate ovary of lowermost flower shorter than twice as long as rachis . . . . . . . . . . . . . . . . . Group A Basal node of pedicel approximately level with floral bract attachment in all flowers; or raceme subumbellate, pedicellate ovary of lowermost flower twice as long as rachis or longer, and several to all flowers per inflorescence opening simultaneously; or both characters present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Group B Basal node of pedicel well above (1/3 the diameter of the pedicel or more) floral bract attachment . . . . . . Group C Basal node of pedicel approximately level with floral bract attachment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Group D

Group A 1. 2. 3.

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Petal margins ciliate or ciliolate . . . . . . . . . . . . . . . . . . . . . 2 Petal margins glabrous, erose(-papillose) or denticulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Apex median sepal thickened, globular to ellipsoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Lepanthanthe Apex median sepal obtuse to caudate . . . . . . . . . . . . . . . 3 Floral bracts not amplexicaul . . . . . . . . . . . . . . . . . . . . . . 4 Floral bracts amplexicaul and usually tubular proximally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Pseudobulbs wrinkling irregularly with age . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Hirtula Pseudobulbs wrinkling with longitudinal furrows with age . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Eublepharon Margins of labellum ciliate . . . . B. sect. Monanthaparva Margins of labellum entire or erose to dentate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Intervallatae

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Inflorescences 1- or 2-flowered (without more flowers present as developing buds) . . . . . . . . . . . . . . . . . . . . . . . 7 Inflorescences 3- to many-flowered (2-flowered inflorescences may also be present) . . . . . . . . . . . . . . . . . . . . . . . 17 Labellum 8.0–17.0 mm long (without stretching or spreading) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Labellum 0.8–6.0 mm long . . . . . . . . . . . . . . . . . . . . . . . 11 Labellum trilobed with antrorse lateral lobes attached to basal 1/3 of its length . . . . . . . . . . . B. sect. Balaenoidea Labellum undivided, or labellum inconspicuously trilobed with lobes attached close to its base . . . . . . . . . . . . . . . . . 9 Roots densely and coarsely papillose to rugulose to verrucate . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Lepidorhiza Roots glabrous, finely hirsute locally . . . . . . . . . . . . . . . 10 Petals more than half as long as dorsal sepal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Leopardinae Petals half as long as dorsal sepal or shorter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Intervallatae Stelidia distally along lower margin with a distinct, downwardly directed, generally rounded tooth (the tooth protruding well below level of rostellum) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Uncifera Stelidia distally along lower margin without a tooth (sometimes with an antrorse tooth lower down, or teeth may be present between which rostellum is attached) . . . . . . . . 12 Column in lateral view with proximal edge of stigma protruding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Column in lateral view with proximal edge of stigma not protruding (a slight swelling may be present lower down on column foot) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Pollinia 2 . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Macrouris Pollinia 4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Pelma Sepals 5–7-veined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 Sepals 3-veined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 Dorsal sepal 9–14 mm long; lateral sepals abaxially without keel . . . . . . . . . . . . . . . B. sect. Sestochilus (B. anceps) Dorsal sepal c. 5 mm long; lateral sepals abaxially with keel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Biseta Pseudobulbs minute compared to size of plant or seemingly absent . . . . . . . . . . . . . . . B. sect. Aeschynanthoides Pseudobulbs distinctly present . . . B. sect. Adelopetalum Raceme with 3 or more distichous flowers (although only 1 per raceme may be open, with more visible as buds or developing flower bracts) . . . . . . . . . . . . . . . . . . . . . . . . . 18 Raceme with spirally arranged flowers, or tristichous . . 21 Apex of dorsal sepal thickened, globular to ellipsoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Lepanthanthe Apex of dorsal sepal obtuse to caudate . . . . . . . . . . . . . 19 Rhizome erect, patent to pendulous, with roots growing over or alongside rhizome towards substrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Pelma Rhizome creeping or straggling, with roots spreading . 20 Roots glabrous or locally and finely hirsute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Intervallatae 19

DENDROBIEAE



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Roots densely and coarsely papillose to rugulose to verrucate . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Lepidorhiza Rhizome erect, patent to pendulous, with roots growing over or alongside rhizome towards substrate . . . . . . . . 22 Rhizome creeping or straggling, with roots spreading . 25 Stelidia distally along lower margin with a distinct, ­downwardly directed, generally rounded tooth (the tooth protruding well below level of rostellum) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Uncifera Stelidia distally along lower margin without a tooth (sometimes with an antrorse tooth lower down, or teeth may be present between which rostellum is attached) . . . . . . . . 23 Pollinia 4 . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Imitatores Pollinia 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 Racemes eventually two to many, fasciculate on a short sympodium; labellum with distinct, antrorse lateral lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Lepanthanthe (species with hanging rhizome) Racemes solitary, or sometimes two together; labellum without lateral lobes . . . . . . . . . . . . . . . B. sect. Macrouris Lateral sepals attached to apex of column foot only, leaving a gap between lower margin of petals and upper margins of lateral sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 Lateral sepals attached along most of length of column foot, with no gap between petals and lateral sepals . . . 27 Labellum hirsute; petals less than half as long as dorsal sepal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Henosis Labellum glabrous; petals more than half as long as dorsal sepal . . . . . . . . . . . . . B. sect. Altisceptrum (B. gymnopus) Lateral sepals either with a distinct, entire to deeply dentate keel abaxially, or apices of lateral sepals thickened, globular to ellipsoid to almost cylindrical (sometimes incurved and difficult to see), or both characters present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 Lateral sepals without keel abaxially; apices of lateral sepals obtuse to caudate, not thickened . . . . . . . . . . . . . 29 Pseudobulbs distinctly present . . . . . . . . . . B. sect. Biseta Pseudobulbs minute and inconspicuous compared to size of plant . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Lepanthanthe (species with creeping rhizome) Stelidia distally along lower margin with a distinct, ­downwardly directed, generally rounded tooth (the tooth protruding well below level of rostellum) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Uncifera Stelidia distally along lower margin without a tooth (sometimes with an antrorse tooth lower down, or teeth may be present between which rostellum is attached) . . . . . . . . 30 Either petals 1-veined; or petals 3-veined and dorsal sepal 3-veined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 Petals 3–5-veined; dorsal sepal 5–7-veined . . . . . . . . . . 35 Pollinia 2, or pollinia 4 and inner pair less than half as long as outer pair . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 Pollinia 4, inner pair more than half as long as outer pair . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

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Group B 1. 2. 3. 4. 5.

Pseudobulb of flowering shoots 2-leaved . . . . . . . . . . . . 2 Pseudobulb of flowering shoots 1-leaved . . . . . . . . . . . . 3 Leaves persistent, thick . . . . . . . . . . B. sect. Blepharistes Leaves deciduous, thin . . . . . . . . . . . . B. sect. Lemniscata Rhizome erect to patent to pendulous . . . . . . . . . . . . . . . 4 Rhizome creeping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Pseudobulbs minute compared to size of plant, entirely hidden in rhizome bracts . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Pseudobulbs distinctly present . . . . . . . . . . . . . . . . . . . . . 7 Pollinia 4; raceme c. 15-flowered, dense and globose . . . . . . . . . . . . . . . . . . . B. sect. Brachystachyae (B. salaccense) Pollinia 2; raceme 2- to many-flowered, lax and elongate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Basal node of pedicel well above (1/3 the diameter of pedicel or more) floral bract attachment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Imitatores (B. dichotomum) Basal node of pedicel approximately level with floral bract attachment . . . . . . . . . . . . . . . . . . . . B. sect. Stachysanthes Lateral sepals distinctly twisted proximally, so that upper margins turned inwards, towards each other; petal margins (partly) finely ciliolate or finely papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Emarginatae Lateral sepals not twisted proximally, or only slightly so distally; petal margins entire or erose . . . . . . . . . . . . . . . . 8 Basal node of pedicel approximately level with floral bract attachment . . . . . . . . . . . . . . . . . . . B. sect. Desmosanthes Basal node of pedicel well above (1/3 the diameter of pedicel or more) floral bract attachment . . . . . . . . . . . . . 9

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32. Column in lateral view with proximal edge of stigma protruding, or with a distinct tooth; roots densely and coarsely papillose to rugulose to verrucate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Macrouris Column in lateral view with proximal edge of stigma not protruding; roots glabrous, locally finely hirsute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Adelopetalum 33. Petals minutely erose-papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Eublepharon (B. levinei) Petals entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 34. Petals 1-veined; labellum margins glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Rhinanthera Either petals 3-veined, or petals 1- or 2-veined; labellum margins partly papillose–ciliate . . . B. sect. Adelopetalum 35. Dorsal sepal 2.5–3.6 mm wide; l/w 3.6–5.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Macrouris (B. dekockii) Dorsal sepal 5–9 mm wide; l/w 1.5–2.6 . . . . . . . . . . . . . 36 36. Labellum adaxially glabrous; labellum not auriculate . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Sestochilus (B. anceps) Labellum adaxially irregularly and coarsely verrucate, or labellum proximally auriculate with antrorse auricles, or both characters present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Leopardinae

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BULBOPHYLLUM

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Column in lateral view with proximal edge of stigma protruding, or with a distinct tooth; roots densely and coarsely papillose to rugulose . . . . . . . B. sect. Macrouris Column in lateral view with proximal edge of stigma not protruding; roots glabrous, locally finely hirsute . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Imitatores (B. dichotomum)

10. Raceme subumbellate (pedicellate ovary of lowermost flower twice as long as rachis or longer) . . . . . . . . . . . . . 11 Raceme elongate or congested (pedicellate ovary of lowermost flower shorter than twice as long as rachis) . . . 35 ●

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11. Petals (partly) ciliolate, ciliate, fimbriate or paleate . . . . 12 Petals entire, erose(–papillose) or irregularly denticulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 12. Stelidia with an antrorse tooth on adaxial surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Cirrhopetalum Stelidia without a tooth on adaxial surface (teeth may be present along upper and lower margins) . . . . . . . . . . . . 13 ● ●

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13. 14.

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Labellum 5 mm or shorter . . . . . . . . . . . . . . . . . . . . . . . . 14 Labellum longer than 5 mm . . . . . . . . . . . . . . . . . . . . . . 18 Petals paleate . . . . . . . . . . . . . . . . . . . . . . B. sect. Plumata Petals ciliolate, ciliate or fimbriate . . . . . . . . . . . . . . . . . . 15



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Labellum 1.8–3.6 mm long . . . . . . . B. sect. Eublepharon

24. Lateral sepals distinctly twisted proximally, so that upper margins are turned inwards, towards each other . . . . . . 25 Lateral sepals not twisted proximally, or only slightly so distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 ●

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25. Labellum trilobed with lateral lobes attached to basal 1/3 of its length . . . . . . . . . . . . . . . . . . . . B. sect. Balaenoidea Labellum undivided . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 ●

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26. Inflorescence 2-flowered; stem sections between floral bract attachment and basal node of pedicel of both flowers fused . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Biflorae Inflorescence 2- to many-flowered; stem sections between floral bract attachment and basal node of pedicel of both flowers free . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 ●

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27. Stelidia widened distally, apex rounded to truncate to bifid to erose, or stelidia semi-elliptic and rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Macrostelydia Stelidia tapering towards an acute apex (apart from the wing sometimes present along lower margin, lower down) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 ●

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28. Either lateral sepals coherent to connate along upper margins, or lateral sepals free and leaves thick, without visible venation when alive . . . . . . . . . . . . . B. sect. Brachyantha Lateral sepals free; leaves thin, with a dense, reticulate venation visible when alive . . . . . . . . B. sect. Beccariana

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29. Dorsal sepal 3-veined . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 Dorsal sepal 5–19-veined . . . . . . . . . . . . . . . . . . . . . . . . .31

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15. Lateral sepals distinctly twisted proximally, so that upper margins are turned inwards, towards each other . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Recurvae Lateral sepals not twisted proximally, or only slightly so distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 16. Pseudobulbs wrinkling in an irregular pattern with age . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Hirtula Pseudobulbs wrinkling with longitudinal furrows with age . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 ●

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17. Labellum distinctly auriculate proximally; auricles antrorse with minutely denticulate margin . . . B. sect. Acrochaene Labellum undivided . . . . . . . . . . . . . B. sect. Eublepharon 18. Labellum distinctly auriculate proximally; auricles antrorse with minutely denticulate margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Acrochaene Labellum undivided (labellum may be widened proximally) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 ●

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19. Lateral sepals 6.3–10.0 mm long . . . . . . . B. sect. Hirtula Lateral sepals 20.0–300.0 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Cirrhopetaloides 20. Either margins of sepals, or margins of petals, or both, denticulate, (erose–)papillose; or margins of dorsal sepal ciliolate, ciliate, fimbriate or paleate . . . . . . . . . . . . . . . . 21 Margins of sepals and petals entire and glabrous (petals may be truncate and caudate at apex) . . . . . . . . . . . . . . . 24 ● ●

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21. Lateral sepals distinctly twisted proximally, so that upper margins are turned inwards, towards each other . . . . . . 22 Lateral sepals not twisted proximally, or only slightly so distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 ●

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22. Labellum 5.0–15.0 mm long . . B. sect. Cirrhopetaloides Labellum 1.9–3.5 mm long . . . . . B. sect. Rhytionanthos 23. Labellum 5.5–16.0 mm long . . . . . . B. sect. Leopardinae ● ●

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30. Shoots with rhizome fused to pseudobulb from which they arise over most of length of the pseudobulb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Planibulbus Shoots growing away from pseudobulb from which they arise, not fused to pseudobulb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Desmosanthes ●

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31. Petals 5–7-veined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 Petals 1–3-veined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 ● ●

32. Pedicel with basal node 0.5–1.0 x diameter of pedicel above floral bract attachment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Leopardinae Pedicel with the basal node level with floral bract attachment . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Beccariana ●

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33. Lateral sepals distinctly keeled abaxially . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Biseta (B. bisetoides) Lateral sepals not or hardly keeled abaxially . . . . . . . . . 34 ●

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34. Labellum length/width 1.5–2.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Desmosanthes Labellum length/width approximately 1 . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Brachyantha (B. violaceolabellum) ●

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35. Labellum not mobile, attached to column foot by a thick strip of tissue . . . . . . . . . . . . . . . . . . . . . . B. sect. Sunipia Labellum mobile, attached to column foot by a thin strip of tissue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 36. Stelidia distally along lower margin with a patent and downwardly directed, semi-elliptic tooth . . . . . . . . . . . . 37 ●

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21

DENDROBIEAE



Stelidia without a tooth along the lower margin, or stelidia with a tooth along the lower margin in a more proximal position . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38

50. Pseudobulbs minute, inconspicuous compared to size of plant . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 Pseudobulbs distinctly present . . . . . . . . . . . . . . . . . . . . 54

37. Petals triangular . . . B. sect. Beccariana (B. anaclastum) Petals ovate to elliptic . . . . . . . . . . . . . . . . B. sect. Uncifera

51. Pollinia 2 . . . . . . . . . . . . . . . . . . . . . B. sect. Stachysanthes Pollinia 4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52

38. Stelidia distally abruptly bent downwards but with an antrorse tip . . . . . . . . . . . . . . . . . B. sect. Saurocephalum Stelidia porrect or gradually upwards or downwards falcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 39. Either labellum proximally auriculate, with antrorse auricles along margins; or margins of petals denticulate, papillose, ciliate or ciliolate; or both characters (on labellum and petals) present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 Labellum undivided, or labellum trilobed with semicircular or semi-elliptic, patent or retrorse lateral lobes; margins of petals entire, glabrous . . . . . . . . . . . . . . . . . . . . . . . . . 61

52. Petals widest at or near base and gradually tapering into an approximately strap-shaped top part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Altisceptrum Petals elliptic to (ob-)ovate to triangular . . . . . . . . . . . . 53

40. Labellum proximally auriculate . . . . . . . . . . . . . . . . . . . . 41 Labellum undivided . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48

54. Petals widest at base or in proximal half, and gradually tapering into an approximately linear top part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Altisceptrum Petals elliptic to oblong to (ob-)ovate . . . . . . . . . . . . . . . 55

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41. Petal margins ciliate or ciliolate . . . . . . . . . . . . . . . . . . . . 42 Petal margins glabrous, denticulate or papillose . . . . . . 44 42. Pseudobulbs wrinkling in a random pattern with age . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Hirtula Pseudobulbs wrinkling with longitudinal furrows with age . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 ● ●

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43. Petals 1-veined, less than half as long as dorsal sepal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Monomeria Petals 3-veined, half as long as dorsal sepal or longer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Acrochaene 44. Stelidia with a distinct tooth or wing along lower margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 Stelidia without a tooth or wing along lower margin (lower down, proximal edge of stigma may be protruding in lateral view) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 ●

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53. Flowers in a dense raceme, arranged in a regular pattern, or flowers in a lax raceme, with lateral sepals held appressed or parallel to rachis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Brachystachyae Flowers in a lax raceme, not arranged in a regular pattern, nor with lateral sepals held appressed or parallel to rachis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Stachysanthes ●

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55. Petals entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56 Petals finely erose–denticulate . . . . . . . . . . . . . . . . . . . . . 57 ● ●

56. Stelidia downwards sigmoid towards apex, with a large wing along upper margin that widely overtops stelidia themselves (check the vein!) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Saurocephalum Stelidia approximately straight, with or without a tooth along lower margin . . . . . . . . . . . B. sect. Brachystachyae ●

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57. Lateral sepals 27–37 mm long . . . . . . B. sect. Gongorodes Lateral sepals c. 5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Eublepharon (B. vietnamense) ● ●

58. Labellum margins ciliate . . . . . . . . . . . . . B. sect. Hirtula Labellum margins glabrous to papillose . . . . . . . . . . . . . 59 ● ●

45. Sepals abaxially distinctly hirsute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Beccariana (B. taeter) Sepals abaxially glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Phreatiopsis (B. viridescens) 46. Pedicel distinctly longer than ovary . . . . . . . . . . . . . . . . 47 Pedicel approximately as long as the ovary or shorter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Racemosae 47. Petals caudate, caudae with thickened apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Antennata Petals acute to acuminate, apices not thickened . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Beccariana (B. striatum)

59. Petals widest near base and gradually tapering into an approximately linear top part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Altisceptrum (B. farinulentum) Petals elliptic to (ob-)ovate . . . . . . . . . . . . . . . . . . . . . . . 60

48. Adaxial side of labellum with a median ridge near base, this ridge proximally widening into a triangular callus that forms a roof over furrows bordering the ridge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Phreatiopsis Adaxial side of the labellum without ridge that proximally widens into a callus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 49. Petal margins entire to denticulate, glabrous to papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50 Petal margins ciliate or ciliolate . . . . . . . . . . . . . . . . . . . . 58

62. Proximal edge of stigma with a protruding, triangular tooth . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Pseudopelma Proximal edge of stigma rounded to truncate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Beccariana

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60. Stelidia with a rounded wing or a deltoid tooth along lower margin . . . . . . . . . . . . . . . . B. sect. Brachystachyae Stelidia without a tooth or wing along lower margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Eublepharon ●

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61. Dorsal sepal 5–19-veined, free from lateral sepals . . . . 62 Dorsal sepal 3-veined, or dorsal sepal 5-veined and fused to lateral sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63 ● ●

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63. Pseudobulbs minute, inconspicuous compared to size of plant . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64 Pseudobulbs distinctly present . . . . . . . . . . . . . . . . . . . . 67 ●

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64. Pollinia 2 . . . . . . . . . . . . . . . . . . . . . B. sect. Stachysanthes Pollinia 4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65 ● ●

BULBOPHYLLUM

65. Petals widest at or near the base and gradually tapering into an approximately strap-shaped top part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Altisceptrum Petals elliptic to (ob-)ovate to triangular . . . . . . . . . . . . 66 ●

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66. Flowers in a dense raceme, arranged into a regular pattern, or flowers in a lax raceme, with lateral sepals held appressed or parallel to rachis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Brachystachyae Flowers in a lax raceme, not arranged into a regular pattern, nor with lateral sepals held appressed or parallel to rachis . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Stachysanthes ●

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67. Lateral sepals with lower margins running approximately parallel (at least in proximal half of sepals or adherent (but flowers not cleistogamous) . . . . . . . . . . . . . . . . . . . 68 Lateral sepals with lower margins spreading, or flowers cleistogamous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69 ●

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68. Pseudobulbs cylindrical, at most slightly widened proximally . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Beccariana Pseudobulbs ovoid to ellipsoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Brachystachyae ●

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69. Leaves deciduous . . . . . . . . . . . . . . . . B. sect. Lemniscata Leaves persistent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .70 ● ●

70. Shoots with rhizome fused to pseudobulb from which they arise over most of length of pseudobulb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Planibulbus Shoots growing away from pseudobulb from which they arise, not fused to pseudobulb . . . . . . . . . . . . . . . . . . . . 71 ●

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71. Petals more than half as long as dorsal sepal . . . . . . . . . 72 Petals half as long as dorsal sepal or shorter . . . . . . . . . 75 ● ●

72. Petals widest at or near base and gradually tapering into an approximately linear top part, or widest in proximal half, then abruptly narrowed into a drawn-out distal half, or petals strap-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73 Petals elliptic to (ob-)ovate . . . . . . . . . . . . . . . . . . . . . . . 74 ●

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73. Pseudobulbs obovoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Desmosanthes (B. compressum) Pseudobulbs ovoid or ellipsoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Altisceptrum ●

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74. Pseudobulbs cylindrical, at most slightly widened proximally . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Beccariana Pseudobulbs ovoid to globose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Adelopetalum (B. bracteatum) ●

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Group C 1. Labellum deeply saccate with a thickened, recurved apical part overhanging anterior margin of saccate part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Pedilochilus Labellum convex, partly concave, or labellum saccate without a thickened, recurved apical part overhanging anterior margin of saccate part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Petal margins with 1–24 appendages differing from petal itself in thickness and/or surface ornamentation . . . . . . . 3 Petal margins entire to denticulate, glabrous to papillose or ciliate, or with appendages similar to petals in thickness and/or surface ornamentation . . . . . . . . . . . . . . . . . . . . . . 5 3. Dorsal sepal connate to lateral sepals for half of its length or more; lateral sepals free along their lower margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Monosepalum Sepals free, or all connate proximally for a short distance, or the lateral sepals connate along their lower margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Sepals approximately equal in shape and size; dorsal sepal 5.0–18.5 mm long . . . . . . . . . . . . . . . . B. sect. Epicrianthes Lateral sepals dissimilar from the median; dorsal sepal 25.0–145.0 mm long . . . . . . . . . . . . . . . B. sect. Hyalosema 5. Adaxial side of labellum proximally with a transverse ridge without a notch halfway; proximal surface of this ridge touches column foot (column foot and base of labellum often fused together) . . . . . . . . . . . B. sect. Codonosiphon Adaxial side of the labellum proximally without a transverse ridge, or with a transverse ridge notched halfway and not touching column foot . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6. Pseudobulbs conspicuous compared to size of plant; rhizomes creeping, with shoots developing at basal node of pseudobulb and fused to the latter along 1/3–4/5 of its length, or with shoots developing a similar distance above its basal node, perforating the pseudobulb over the distance between its basal node and the point of emergence (rhizome will break off in front of the fused part) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Pseudobulbs minute compared to size of plant; or rhizomes patent; or rhizomes creeping with shoots developing from nodes below pseudobulbs and not fused to the latter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 7. Either pollinia 2, or pollinia 4 with the inner half as long as the outer or less . . . . . . . . . . . . . . . . . B. sect. Macrocaulia Pollinia 4, inner approximately as long as the outer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Leopardinae 8. Stelidia distally along lower margin with a distinct, downwardly directed, rounded to obtuse tooth (the tooth protruding well below level of rostellum) . . . . . . . . . . . . . . . . 9 Stelidia distally along lower margin without a tooth (sometimes with an antrorse tooth lower down, or teeth may be present between which rostellum is attached) . . . . . . . . . 10 9. Anther with a deeply lacerated anterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Monanthaparva (B. tothastes) Anther with an entire anterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Uncifera ●

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75. Inflorescences sprouting from basal nodes of a new shoot while the latter develops . . . . . . . . . . . B. sect. Reptantia Inflorescences sprouting from fully mature shoots, either below pseudobulbs, or elsewhere along rhizome . . . . . 76 ●

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76. Pollinia 2 . . . . . . . B. sect. Adelopetalum (B. bracteatum) Pollinia 4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77 77. Rhizome bracts densely and finely papillose; roots densely and coarsely papillose . . . . . . . . . . . B. sect. Pseudopelma Rhizome bracts glabrous to colliculate; roots glabrous, finely hirsute locally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Desmosanthes ● ●

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23

DENDROBIEAE

10. Pollinia 4 (the inner pair may be minute and may be best visible if the outer pair is split) . . . . . . . . . . . . . . . . . . . . 11 Pollinia 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 11. Petals 5–12-veined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Petals 1–3(–4)-veined . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 12. Dorsal sepal 4.0–4.2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Monanthaparva (B. tothastes) Dorsal sepal 7.0–95.0 mm long . . . . . . . . . . . . . . . . . . . . 13 13. Basal node of pedicel larger than diameter of pedicel above floral bract attachment; labellum adaxially glabrous to (partly) hirsute or minutely papillose . . . . . . . . . . . . . 14 Basal node of pedicel approximately equal to diameter of pedicel above floral bract attachment or less, or labellum adaxially (partly) coarsely papillose . . . . . . . . . . . . . . . . . 15 14. Flowers resupinate . . . . . . . . . . . . . . . . B. sect. Sestochilus Flowers non-resupinate . . . . . . . . . . . B. sect. Stenochilus 15. Lateral sepals more than three times as long as dorsal sepal . . . . . . . . . . B. sect. Cirrhopetaloides (B. fascinator) Lateral sepals approximately as long as dorsal sepal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Leopardinae 16. Rhizome patent; roots developing along basal part of rhizome only or growing backwards along rhizome towards substrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Rhizome creeping or straggling; roots developing along most of length of rhizome or below most pseudobulbs, spreading in all directions . . . . . . . . . . . . . . . . . . . . . . . . 22 17. Column foot distinctly widening towards apex, usually with two lateral teeth at tip . . . . . . . . . . . . . . . . . . . . . . . 18 Column foot not or hardly widening towards apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 18. Pseudobulbs well protruding from rhizome scales; shoots proximally fused to pseudobulbs, arising from the latter at or above halfway its length . . . . . . . . . B. sect. Epibulbon Pseudobulbs largely covered by rhizome scales or well protruding from rhizome scales; shoots proximally not fused to pseudobulbs . . . . . . . . . . . . . . B. sect. Polymeres 19. Petals shorter than 1/10 of length of dorsal sepal . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Peltopus (B. subapetalum) Petals longer than 1/10 of the length of dorsal sepal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 20. Pedicel and ovary together 35 mm or longer . . . . . . . . . . . . . . . . . . . . . . . B. sect. Papulipetalum (B. ellipticifolium) Pedicel and ovary together 10 mm or less . . . . . . . . . . . 21 21. Pedicel longer than ovary; petal margins finely but distinctly erose–denticulate or entire and ciliate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Piestobulbon Pedicel shorter than ovary; petal margins entire or erose distally, glabrous . . . . . . . . . . . . . . . . . . . . . . B. sect. Pelma 22. Stelidia vestigial, (partly) included in distinctly projecting rostellum, sometimes only visible as thickenings on its side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 Stelidia distinct although sometimes short; rostellum anterior surface receding in between stelidia apices or level with these, or only slightly projecting . . . . . . . . . . . . . . . 25 ●

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23. Petals shorter than 1/4 the length of dorsal sepal; or column foot distinctly swollen distally above labellum attachment, this callus fitting in a cavity on adaxial surface of labellum, near the base; or both characters present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Peltopus Petals 1/4 the length of dorsal sepal or longer; column foot not swollen distally; labellum without a cavity on adaxial surface near base . . . . . . . . . . . . . . . . . . . . . . . . . 24 24. Labellum margin coarsely papillose proximally . . . . . . . . . . . . . . . . . . B. sect. Monanthaparva (B. hemiprionotum) Labellum margin glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Minutissima 25. Column foot distinctly widening towards apex and usually with two lateral teeth at apex . . . . . . . . . . . . . . . . . . . . . . 26 Column foot tapering towards apex or only slightly widening towards apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 26. Petals proximally (partly) fused to column or stelidia narrowly oblong and obtuse, or both characters present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Brachypus Petals not fused to column; stelidia deltoid to triangular to subulate, approximately acute . . . . . . . B. sect. Polymeres 27. Petals proximally fused to column . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Brachypus Petals entirely free . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 28. Dorsal sepal 35.0–145.0 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Hyalosema Dorsal sepal 1.5–20.0 mm long . . . . . . . . . . . . . . . . . . . . 29 29. Pseudobulbs seemingly absent or minute, not or hardly thicker than rhizome . . . . . . . B. sect. Aeschynanthoides Pseudobulbs distinct when compared to size of plant, distinctly thicker than rhizome . . . . . . . . . . . . . . . . . . . . . . . 30 30. Stelidia apex approximately level with rostellum anterior surface or nearly so . . . . . . . . . . . . . . B. sect. Minutissima Stelidia apex projecting well beyond rostellum anterior surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 31. Pseudobulbs distant, at least some rhizome sections between pseudobulbs longer than adjacent pseudobulbs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 Pseudobulbs in clusters, rhizome sections between pseudobulbs shorter than adjacent pseudobulbs . . . . . . . . . 34 32. Dorsal sepal margins glabrous to minutely papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Monanthaparva Dorsal sepal margins ciliate . . . . . . . . . . . . . . . . . . . . . . . 33 33. Dorsal sepal acute to acuminate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Papulipetalum Dorsal sepal rounded to obtuse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Hirtula (B. rariflorum) 34. Petal margins papillose, glandular or distinctly, deeply and finely erose; or stelidia distally glandulose; or both characters present . . . . . . . . . . . . . . . . . . B. sect. Papulipetalum Petal margins approximately entire or slightly, shallowly and coarsely erose; stelidia not glandular . . . . . . . . . . . . 35 35. Sepals 3-veined . . . . . . . . . . . . . . B. sect. Monanthaparva Sepals 5–11-veined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 ●

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BULBOPHYLLUM

36. Lateral sepals approximately only as long as dorsal sepal . . . . . . . . . . . . . . . . . . . B. sect. Leopardinae (B. griffithii) Lateral sepals approximately twice as long as dorsal sepal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Balaenoidea



37. Column foot distinctly widening towards apex, usually with two lateral teeth at apex . . . . . . . . . . . . . . . . . . . . . . 38 Column foot not or hardly widening towards apex (sometimes widened more proximally and tapering towards apex) without teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40

48. Labellum margins glabrous . . . . . B. sect. Papulipetalum Labellum margins papillose–ciliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Monanthaparva (B. lipense)

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38. Column foot deeply concave distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Tapeinoglossum Column foot flat or convex distally . . . . . . . . . . . . . . . . 39

Petals without caudate appendages, (ob-)ovate to elliptic to spatulate to wedge-shaped, or petals triangular and labellum adaxially without ridges proximally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Monanthaparva

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39. Pseudobulbs protruding from rhizome scales; shoots proximally fused to pseudobulbs, arising from the latter at or above half its length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Epibulbon Pseudobulbs largely covered or not by rhizome scales; shoots not proximally fused to pseudobulbs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Polymeres ●

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40. Dorsal sepal distinctly and often sharply conduplicate towards apex . . . . . . . . . . . . . . . . . . . . B. sect. Hoplandra Dorsal sepal not or hardly conduplicate towards apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 ●

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41. Pseudobulbs seemingly absent or minute, not or hardly thicker than rhizome . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 Pseudobulbs distinct when compared to size of plant, distinctly thicker than rhizome . . . . . . . . . . . . . . . . . . . . . . . 44 ●

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43. Column in lateral view with proximal edge of stigma protruding, or with a distinct tooth; roots densely and coarsely papillose to rugulose to verrucate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Macrouris Column in lateral view with proximal edge of stigma not protruding, roots glabrous, locally finely hirsute . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Imitatores (B. glaucum) ●

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44. Stelidia apices approximately level with rostellum anterior surface, or slightly longer, or slightly shorter . . . . . . . . . 45 Stelidia apices projecting well beyond rostellum anterior surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46

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Floral bract distinctly tubular proximally . . . . . . . . . . . . . 2 Floral bract not tubular (but sometimes amplexicaul, with the side just connected at base) . . . . . . . . . . . . . . . . . . . . . 5 Leaves deciduous; column foot elongate so that a wide gap exists between lower margins of petals and upper margins of lateral sepals . . . . . . . . . . . . B. sect. Drymoda Leaves persistent; column foot not elongate with no gap between petals and lateral sepals . . . . . . . . . . . . . . . . . . . . 3 Lateral sepals 5–11-veined, free . . . . . . . . . B. sect. Trias Lateral sepals (1–3-veined) or 5–7-veined and connate along their lower margins . . . . . . . . . . . . . . . . . . . . . . . . . 4 Roots glabrous . . . . . . . . . . . . . . . . . . B. sect. Monanthes Roots densely and coarsely papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Pseudopelma (B. fossatum) Labellum not mobile, attached to column foot by a thick strip of tissue . . . . . . . . . . . . . . . . . . . . . . B. sect. Sunipia Labellum mobile, attached to column foot by a thin strip of tissue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Sepals 3- or 5-veined; dorsal sepal connate to lateral sepals . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Stachysanthes Sepals 5–19-veined; dorsal sepal not connate to lateral sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Petals entire, glabrous . . . . . . . . . . . . B. sect. Beccariana Petals (sparsely) ciliolate, ciliate or fimbriate . . . . . . . . . . 8 Labellum 2.5–2.7 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Recurvae (B. subbullatum) Labellum 8.0–15.0 mm long . . B. sect. Cirrhopetaloides

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45. Rhizomes creeping or straggling; roots spreading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Minutissima Rhizomes patent to pendulous; roots growing backwards along rhizome towards substrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Schistopetalum (B. barbasapientis) ●

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46. Petal margins approximately entire or slightly, shallowly and coarsely erose, glabrous or with 3–9 caudate appendages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47 Petal margins papillose, glandular or distinctly, deeply and finely erose; petals without appendages . . . . . . . . . . . . . 48 ●

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47. Petals with 3–9 caudate appendages, or triangular and labellum adaxially with ridges proximally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Schistopetalum ●

Bulbophyllum sect. Acrochaene (Lindl.) J.J.Verm., in prep. Type species (proposed): Bulbophyllum kingii Hook.f. Epiphytes with creeping rhizomes. Pseudobulbs wrinkling with age along longitudinal lines, 1-leaved. Leaves persistent. Inflorescences a subumbellate to elongate raceme with flowers spirally arranged. Pedicel with the basal node above the attachment of the floral bract. Flowers resupinate. Sepals free, approximately equal; margins glabrous to ciliate or fimbriate, c. 7-veined. Petal margins ciliate to fimbriate, surface glabrous; 3-veined. Labellum mobile on a thin ligament, auriculate. Column with stigma proximally not protruding. Stelidia shorter than ½ the length of the column. Anther anterior surface not 25

DENDROBIEAE

concave, anterior margin somewhat drawn out or not; pollinia 4, with bifid stipe. Three species in semideciduous forests up to 1300 m from India to Vietnam, southwards to Thailand. (JJV) Bulbophyllum sect. Adelopetalum (Fitzg.) J.J.Verm., Orch. Monog., 7, 66 (1993). Type species: Bulbophyllum bracteatum (Fitzg.) F.M.Bailey Epiphytes or lithophytes with creeping rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescences usually an elongate raceme with flowers spirally arranged. Floral bracts amplexicaul or not; pedicel with the basal node above the attachment of the floral bract. Flowers resupinate. Sepals free, usually 3-veined. Petals 1–3-veined. Labellum mobile on a thin ligament, undivided or auriculate. Column with stigma proximally not protruding, column foot distally widened. Stelidia ½ the length of the column or shorter, with or without a tooth along the upper and/or lower margin. Anther anterior surface concave, anterior margin drawn out or not; pollinia 2 or 4. Eight species in lower montane forests up to 1800 m in Australia, New Zealand, and New Caledonia. (JJV) Bulbophyllum sect. Aeschynanthoides Carr, Gard. Bull, Straits Settlem., 5, 137 (1930). Type species: Bulbophyllum dryas Carr Epiphytes with creeping or straggling rhizomes. Pseudobulbs seemingly absent or minute compared to the size of the plant, 1-leaved. Leaves persistent. Inflorescences 1- or 2-flowered. Floral bract amplexicaul, usually tubular; pedicel with the basal node above the attachment of the floral bract. Flowers resupinate. Sepals free, approximately equal, 3-veined. Petals 1-veined, margins entire or sometimes slightly erose. Labellum mobile on a thin ligament, undivided. Column with the rostellum anterior surface approximately level with the stelidia apices, stigma proximally not protruding. Stelidia distinctly shorter than ½ the length of the column, almost absent or inconspicuous, with a tooth along the lower margin. Anther anterior surface concave or not, anterior margin not drawn out; pollinia 2 or 4; with or without a stipe. Probably 4 species in montane forests up to 1600 m in Japan, Taiwan, China, Peninsular Malaysia, Sumatra, and Borneo. (JJV) Bulbophyllum sect. Altisceptrum J.J.Sm., Bull. Jard. Bot. Buitenzorg, ser. 2, 13, 34 (1914). Type species: Bulbophyllum elongatum (Blume) Hassk. Epiphytes with creeping rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence a contracted to elongate raceme with flowers spirally arranged. Flowers resupinate, pedicel usually with the basal node level with the attachment of the floral bract. Sepals free, approximately equal, 3-veined. Petals usually 1-veined with margins entire to erose, glabrous to papillose or ciliolate. Labellum mobile on a thin ligament, usually undivided. Column with stigma proximally not protruding, column foot with lateral teeth near the apex. Stelidia shorter than ½ the length of the column, with or without tooth along the upper and/or the 26

lower margin. Anther anterior surface concave or not, anterior margin not drawn out; pollinia 4. Nine species in dry kerangas forests, limestone forests and montane forests up to 3000 m in India, Bhutan, China, Laos, Thailand, Malaysia, Sumatra, Java, Borneo, the Philippines, Sulawesi, and Papua New Guinea. (JJV) Bulbophyllum sect. Antennata Pfitz. in Engler & Prantl, Die Natürlichen Pflanzenfamilien, 2, 6, 179 (1888). Type species: Bulbophyllum fuscopurpureum Wight Epiphytes with creeping rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence an elongate raceme with flowers spirally arranged. Pedicel with the basal node level with attachment of the floral bract. Flowers resupinate. Sepals free, ca. 5-veined, the lateral sepals c. 1.2 times as long as the dorsal sepal. Petals 1-veined. Labellum mobile on a thin ligament, distinctly auriculate. Column with stigma proximally not protruding. Stelidia shorter than ½ the length of the column. Anther anterior surface not concave, anterior margin not drawn out; pollinia 4. A single species in montane forests up to 2260 m in India. (JJV) Bulbophyllum sect. Balaenoidea Pfitz. in Engler & Prantl, Die Natürlichen Pflanzenfamilien, 2, 6, 179 (1888). Type species: Bulbophyllum balaeniceps Rchb.f. Epiphytes with creeping rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence 1- or 2-flowered. Pedicel with the basal node above the attachment of the floral bract. Flowers resupinate. Dorsal sepal free, 9-veined; lateral sepals as the dorsal but adherent along their lower margin, c. 2 times as long as the dorsal sepal. Petals 3-veined, margins entire or erose. Labellum mobile on a thin ligament, trilobed. Column with stigma proximally not protruding. Stelidia shorter than ½ the length of the column, denticulate along the upper margin or not. Anther anterior surface concave, anterior margin drawn out into a beak; pollinia 4. A single, poorly known species from India and Sri Lanka. (JJV) Bulbophyllum sect. Beccariana Pfitz. in Engler & Prantl, Die Natürlichen Pflanzenfamilien, 2, 6, 179 (1888). Type species: Bulbophyllum beccarii Rchb.f. Epiphytes with creeping rhizomes with shoots usually not fused to the pseudobulbs. Pseudobulbs small to distinct compared to the size of the plant, 1-leaved. Leaves persistent. Inflorescence a 2- to many-flowered, subumbellate, congested or elongate raceme, or one-flowered with an abortive second flower present; peduncle bracts 3–10; rachis, if present, not or hardly thickened. Floral bracts amplexicaul or not; pedicel with the basal node level with the attachment of the floral bract or above it. Flowers resupinate or non-resupinate. Dorsal sepal free, (3-)5–19-veined; lateral sepals as the dorsal but 1.0–1.4 times as long, usually free. Petals (1–3-)9-veined. Labellum mobile on a thin ligament, usually undivided. Column with stigma usually proximally not protruding, usually without lateral lobes near the apex. Stelidia approximately ½ the length of the column or shorter, entire to erose along the upper margin, with a tooth or wing along the

BULBOPHYLLUM

lower margin. Anther anterior surface not concave, anterior margin drawn out or not; pollinia 4, with or without a single stipe. Approximately 65 species in peat swamp forests up to 800 m in Bhutan, Borneo, Sulawesi, and Papua New Guinea. (JJV) Bulbophyllum sect. Biflorae Garay, Hamer & Siegerist, Nordic J. Bot., 14, 626 (1994). Type species: Bulbophyllum biflorum Teijsm. & Binn. Epiphytes with creeping rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence a 2-flowered, subumbellate raceme. Flowers resupinate; pedicel with the basal node above the floral bract attachment, the stem portions between pedicel node and floral bract attachment fused. Dorsal sepal free, 5-veined; lateral sepals as for the dorsal but 2.2–8.0 times as long, twisted near the base so that the upper margins turn inward, adherent along their upper margins. Petals 3-veined. Labellum mobile on a thin ligament, undivided. Column with stigma proximally not protruding and (slight) lateral lobes near the apex. Stelidia shorter than ½ the length of the column, with or without a wing along the lower margin. Anther anterior surface concave, anterior margin drawn out into a rounded beak; pollinia 4. Five species in montane forests up to 1200 m in Thailand, Peninsular Malaysia, Sumatra, Java, Lesser Sunda Islands, Borneo, and the Philippines. (JJV) Bulbophyllum sect. Biseta J.J.Verm. Edinburgh J. Bot., 58, 120 (2001). Type species: Bulbophyllum bisetum Lindl. Epiphytes with creeping rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence a 2- to many-flowered, contracted or elongate raceme with the rachis (distinctly) thickened or not. Flowers resupinate or not; pedicel with the basal node above the attachment of the floral bract; ovary with the jugae usually extending between the sepals as teeth. Dorsal sepal free, 3-veined; lateral sepals as for the dorsal sepal but 1.0–1.6 times as long, adherent along the lower margin, 5-veined, midvein abaxially keeled. Petals usually 1-veined. Labellum mobile on a thin ligament, undivided. Column with stigma proximally usually protruding and/or with a tooth, column foot often with lateral wings near the apex. Stelidia approximately ½ the length of the column, upper margin with or without tooth, lower margin with a tooth. Anther anterior surface concave, anterior margin drawn out into a beak; pollinia 4. Probably four species in montane forests up to 1900 m in India, Peninsular Malaysia, Sumatra, and Borneo. (JJV) Bulbophyllum sect. Blepharistes J.J.Verm., in prep. Type species (proposed): Bulbophyllum blepharistes Rchb.f. Epiphytes with creeping or straggling rhizomes. Pseudobulbs 2-leaved. Leaves persistent. Inflorescence a many-flowered, subumbellate raceme. Flowers resupinate; pedicel with the basal node slightly above the attachment of the floral bract. Dorsal sepal free, 7–9-veined; lateral sepals as for the dorsal sepal but 1.1–1.2 times as long, connate along their lower margins. Petals 5-veined, the distal margin fimbriate. Labellum mobile on a thin

ligament, undivided. Column with stigma proximally not protruding. Stelidia shorter than ½ the length of the column, lower margin with an inconspicuous wing. Anther anterior surface hardly concave, anterior margin not drawn out; pollinia 4. A single species in temperate forests up to 800 m in India, Burma, Thailand, Laos, Vietnam, and Peninsular Malaysia. (JJV) Bulbophyllum sect. Brachyantha Rchb.f., Ann. Bot. Syst., 6, 264 (1861). Type species: Bulbophyllum umbellatum Lindl. Epiphytes with creeping rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence usually a 2- to many-flowered, subumbellate raceme. Flowers resupinate; pedicel with the basal node above the floral bract attachment. Dorsal sepal free, usually 3–5-veined; lateral sepals as for the dorsal sepal but usually 2–6 times as long, usually twisted near the base so that the upper margins turn inward, free or adherent or adnate along their upper margins. Petals 3–5-veined. Labellum mobile on a thin ligament, undivided. Column with stigma proximally not protruding, column foot with or without lateral lobes near the apex. Stelidia approximately ½ the length of the column or shorter, with or without a tooth along the upper and/or lower margin. Anther anterior surface concave, anterior margin drawn out into a rounded beak or not; pollinia 4. Approximately 26 species in montane forests up to 2200 m from India to Japan, and to the Philippines, with 1 species in New Guinea. (JJV) Bulbophyllum sect. Brachypus Schltr., Repert. Spec. Nov. Regni Veg. Beih. 1: 700–765 (1911). Type species: Bulbophyllum maxillarioides Schltr. Epiphytes with creeping rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescences 1-flowered. Flowers resupinate. Floral bract tubular with basal node of above attachment of the floral bract. Sepals usually free; usually subequal, 3–9-veined. Petals 1-veined, usually proximally fused to the column over a short distance. Labellum mobile on a thin ligament, undivided. Column with stigma usually not protruding. Stelidia shorter to distinctly longer than ½ the length of the column; with or without a tooth along the upper margin, usually without one along the lower. Anther anterior surface not concave, anterior margin not drawn out; pollinia 4. Approximately 21 species in montane forests up to 1100 m in New Guinea and the western Pacific islands. (JJV) Bulbophyllum sect. Brachystachyae Benth. & Hook.f., Gen. Pl., 3, 504 (1880). Type species: Bulbophyllum repens Wall. Epiphytes with creeping rhizomes. Pseudobulbs minute compared to the size of the plant, 1-leaved. Leaves persistent. Inflorescence a many-flowered, contracted or elongated, lax to dense raceme. Flowers resupinate, pedicel with the basal node level with the floral bract attachment. Dorsal sepal free or fused to the laterals, 3-veined, margins glabrous to papillose to ciliate; lateral sepals 3-(–5-)veined, otherwise as for the dorsal sepal but usually adherent along their lower margins, 0.9–1.5 times as long as the dorsal sepal. Petals 1–3-veined, margins 27

DENDROBIEAE

entire to erose, glabrous to ciliate. Labellum mobile on a thin ligament, undivided. Column with stigma proximally not or only slightly protruding. Stelidia shorter to longer than ½ the length of the column, upper and lower margin usually with a tooth. Anther anterior surface usually concave, anterior margin usually drawn out into a beak; pollinia usually 4. Approximately 43 species in montane forests up to 2200 m in India and Sri Lanka to China, southwards to Java, eastwards to the western Pacific. In Sulawesi and eastwards represented by a few species only. (JJV) Bulbophyllum sect. Cirrhopetaloides Garay, Hamer & Siegerist, Nordic J. Bot., 14, 625 (1994). Type species: Bulbophyllum longissimum (Ridl.) Ridl. Epiphytes with creeping rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence 1- to many-flowered, subumbellate raceme. Flowers resupinate; pedicel with the basal node not to slightly above the floral bract attachment. Dorsal sepal free, 5–9-veined, margins entire to erose–dentate, fimbriate with or without paleate fimbriae; lateral sepals as for the dorsal sepal but 1.3–15.0 times as long, twisted near the base so that upper margins turn inward, free or adherent or adnate along their upper margins, margins entire, glabrous. Petals 3-(–5-)veined, margins denticulate and fimbriate with or without paleate fimbriae. Labellum mobile on a thin ligament, undivided. Column with stigma proximally not protruding. Stelidia usually shorter than ½ the length of the column, usually with a tooth or wing along the upper and lower margin. Anther anterior surface slightly concave or not, anterior margin not or hardly drawn out; pollinia 4, usually without stipes. Approximately 19 species in montane forests up to 1500 m in India, Sri Lanka, China, Taiwan, Japan, Burma, Thailand, Laos, Vietnam, Peninsular Malaysia, Sumatra, Borneo, and the Philippines. (JJV) Bulbophyllum sect. Cirrhopetalum (Lindl.) Rchb.f., Ann. Bot. Syst., 6, 259 (1861). Type species: Bulbophyllum longiflorum Thouars Epiphytes with creeping rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence a many-flowered, subumbellate raceme. Flowers resupinate; pedicel with the basal node above the floral bract attachment. Dorsal sepal free, (3–)5-veined, margins entire or distally erose; lateral sepals as for the dorsal sepal, 3–5-veined, but 2.0–5.5 times as long, twisted near the base so that the upper margins turn inward, connate along their upper margins margins, entire, glabrous. Petals 3-veined, margins fimbriate. Labellum mobile on a thin ligament, undivided or auriculate. Column with stigma proximally not protruding. Stelidia ½ the length of the column or shorter, with a tooth on the adaxial surface, with or without a tooth along the upper and/ or lower margin. Anther anterior surface slightly concave, anterior margin drawn out; pollinia 4. Approximately nine species in montane forests up to 1800 m in Africa, Madagascar and surrounding islands, India, Burma, Thailand, Vietnam, Peninsular Malaysia, Sumatra, Java, Borneo, the Philippines, and eastward in the Pacific. (JJV) 28

Bulbophyllum sect. Codonosiphon Schltr., Repert. Spec. Nov. Regni Veg., 10, 178 (1911). Type species: Bulbophyllum codonanthum Schltr. Epiphytes with creeping to patent rhizomes, shoots fused to the pseudobulbs or not. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence 1-flowered. Flowers resupinate; floral bracts tubular; pedicel with the basal node above the attachment of the floral bract. Sepals usually free, usually equal, 3-veined. Petals 1-veined, margins entire to erose, glabrous to papillose. Labellum usually immobile, fused to or firmly appressed to the column foot over a smaller or larger area, this area in front bordered by a transverse, straight ridge; often auriculate. Column with stigma proximally not protruding, column foot widening and thickening towards its apex. Stelidia usually shorter than ½ the length of the column, margins often erose or dentate. Anther anterior surface not concave, anterior margin drawn out or not; pollinia 4. Seventy-seven species in montane forests up to 2200 m in Borneo, Sulawesi, Maluku, New Guinea, and the Pacific. (JJV) Bulbophyllum sect. Desmosanthes (Blume) J.J.Sm., Bijdr. Flora Ned. Indië, 7, 315 (1825). Type species: Bulbophyllum croceum (Blume) Lindl. Epiphytes with creeping to patent rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence a 2- to many-­ flowered, elongate to subumbellate raceme. Flowers resupinate; pedicel with the basal node hardly above the floral bract attachment. Sepals free, 3–7-veined, the laterals 1–3 times as long as the dorsal, 3–7-veined. Petals 1–3-veined. Labellum mobile on a thin ligament, undivided. Column with stigma proximally usually not protruding. Stelidia shorter to longer than ½ the length of the column, lower margin often with a tooth. Anther anterior surface concave or not, anterior margin drawn out into a beak or not; pollinia 4, with or without stipes. Approximately 70 species in montane forests up to 2000 m in India, China, Taiwan, eastwards to the Philippines, Sulawesi, and the Lesser Sunda Islands. (JJV) Bulbophyllum sect. Drymoda (Lindl.) J.J.Verm., in prep. Type species (proposed): Bulbophyllum drymoda J.J.Verm., ined. Epiphytes with creeping rhizomes. Pseudobulbs 2-leaved. Leaves deciduous. Inflorescence 1-flowered. Flowers resupinate. Floral bract tubular; pedicel with the basal node level with the attachment of the floral bract. Sepals free, 3–5-veined, the lateral sepals 1.2–3.3 times as long as the dorsal sepal. Petals 1-veined, margins entire to irregularly denticulate. Labellum mobile but attached to the column foot by a thick strip of tissue, undivided or sub-trilobed. Column with the proximal margin of stigma protruding, column foot with the lateral sepals attached near the apex so that a gap exists between the lower margins of the petals and the upper margins of the lateral sepals. Stelidia approximately ½ the length of the column or shorter, lower margin with or without a tooth. Anther anterior surface not concave, anterior margin slightly drawn out; pollinia 4. Two species in montane forests up to 900 m in Burma, Laos, and Thailand. (JJV)

BULBOPHYLLUM

Bulbophyllum sect. Emarginatae Garay, Hamer & Siegerist, Nordic J. Bot., 14, 625. Type species: Bulbophyllum emarginatum (Finet) J.J.Sm. Epiphytes with patent rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence a 1-flowered, or a 2- to many-­flowered, subumbellate raceme. Flowers resupinate; pedicel with the basal node above the floral bract attachment. Dorsal sepal free, 5–7-veined, margins glabrous to ciliolate; lateral sepals 2–3 times as long as the dorsal, twisted near the base so that the upper margins turn inward, adherent or connate along their upper and lower margins. Petals 3-veined, ciliolate or papillose distally. Labellum mobile on a thin ligament, undivided. Column with stigma proximally not protruding, column foot with or without lateral lobes near the apex. Stelidia approximately ½ the length of the column or shorter, with a wing along the lower margin. Anther anterior surface not concave, anterior margin not drawn out; pollinia 4. Three species in montane forests up to 2600 m in India, Nepal, Bhutan, China, Burma, Thailand, and Vietnam. (JJV) Bulbophyllum sect. Epibulbon Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 703 (1912). Type species: Bulbophyllum epibulbon Schltr. Epiphytes with patent rhizomes, with roots sprouting near the base of the rhizome, growing backwards along it until reaching the substrate, shoots fused proximally to the pseudobulbs. Pseudobulbs distinct to small compared to the size of the plant, 1-leaved. Leaves persistent. Inflorescence 1-flowered. Flowers resupinate; floral bracts tubular; pedicel with the basal node above the attachment of the floral bract. Sepals usually free, 3-(4–9-) veined, usually approximately equal, margins glabrous to papillose or ciliate. Petals usually 1-veined, with margins entire to approximately fimbriate, glabrous to ciliate. Labellum mobile on a thin ligament, undivided or auriculate. Column with stigma often proximally protruding, column foot widened and often thickened and usually with two lateral teeth towards its apex. Stelidia with upper margin sometimes with a tooth. Anther anterior surface concave or not, anterior margin not drawn out; pollinia 2 or 4. Seven species in montane forests up to 1300 m in Papua New Guinea. (JJV) Bulbophyllum sect. Epicrianthes (Blume) Hook.f., Flora Brit. India, 5, 753 (1890). Type species: Bulbophyllum epicrianthes (Blume) Hook.f. Epiphytes with patent rhizomes. Shoots proximally fused to pseudobulbs. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence 1-flowered. Flowers resupinate; floral bracts tubular, pedicel with the basal node above the attachment of the floral bract. Sepals free, approximately equal, 3–9-veined. Petals 3–5-veined, distally with 3–24 appendages, appendages usually differing in texture from the petal itself. Labellum mobile on a thin ligament, auriculate or undivided. Column with stigma proximally not protruding, column foot with or without small, antrorse lobes near the apex. Stelidia shorter than ½ the length of the column, upper margin with or without a tooth, lower margin with 1–2 teeth. Anther anterior surface concave or not, anterior margin usually drawn out, emarginate; pollinia 2 or 4, sometimes with a

stipe. Forty species in montane forests up to 1300 m in China, Burma to Java and Borneo, eastward to New Guinea and New Caledonia. (JJV) Bulbophyllum sect. Eublepharon J.J.Verm., in prep. Type species (proposed): Bulbophyllum eublepharon Lindl. Epiphytes with creeping rhizomes. Pseudobulbs distinct, wrinkling with age along longitudinal lines, 1-leaved. Leaves persistent. Inflorescence a 2- to many-flowered, elongate to subumbellate raceme. Flowers resupinate; pedicel with the basal node not or slightly above the attachment of the floral bract. Dorsal sepal free, usually 3-veined, margins entire to denticulate; lateral sepals 1–2 times as long as the dorsal, usually free, 3–5-veined. Petals 1–3-veined, margins erose to fimbriate. Labellum mobile on a thin ligament, undivided. Column with stigma proximally not protruding, column foot with teeth near the apex. Stelidia ½ the length of the column or shorter; with or without teeth along the upper and lower margin. Anther anterior surface not concave, anterior margin drawn out or not; pollinia 4. Approximately 10 species in montane forests up to 3300 m in Sri Lanka, India, Nepal, Bhutan, China, Taiwan, Japan, Laos, and Vietnam. (JJV) Bulbophyllum sect. Gongorodes J.J.Sm., Bull. Jard. Bot. Buitenzorg, ser. 2, 13, 35. Type species: Bulbophyllum digitatum J.J.Sm. Epiphytes with creeping rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence a many-flowered raceme. Flowers resupinate; pedicel with the basal node level with the attachment of the floral bract. Sepals free; the dorsal sepal 3–7-veined, lateral sepals 1.6–1.9 times as long as the dorsal sepal. Petals 1–3-veined, apices either widened and tridentate, or sharply folded twice, margins erose. Labellum mobile on a thin ligament, undivided. Column with stigma proximally not protruding, column foot with the lateral sepals attached halfway along its length so that a gap exists between the lower margins of the petals and the upper margins of the lateral sepals, with 2 teeth at the apex. Stelidia shorter than ½ the length of the column, triangular, subacute, with or without a slight tooth along the upper margin, with 1–2 teeth along the lower. Anther anterior surface concave, anterior margin drawn out or not; pollinia 4. Two species in montane forests up to 1500 m in New Guinea. (JJV) Bulbophyllum sect. Hemisterrantha J.J.Verm., in prep. Type species (proposed): Bulbophyllum hemisterranthum J.J.Verm. & P.O’Byrne Epiphytes with creeping rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence 1-flowered. Flower resupinate; floral bract tubular; pedicel with the basal node above the attachment of the floral bract. Sepals usually free, approximately equal, usually 3-veined. Petals usually 1-veined, margins entire to erose, glabrous to papillose. Labellum mobile on a thin ligament, undivided, fleshy. Column with stigma proximally often protruding, column foot with or without a dorsal tooth near the apex. Stelidia with upper and/or lower margin often with a tooth. Anther anterior surface concave or not, anterior margin drawn out into a 29

DENDROBIEAE

beak or not; pollinia 2 or 4. Only two species in montane forests up to 1200 m in Sulawesi. (JJV)

to 1200 m from Peninsular Malaysia and Sumatra eastward to the Pacific islands. (JJV)

Bulbophyllum sect. Henosis (Hook.f.) Ormerod, Edinburgh J. Bot., 58, 120. Type species: Bulbophyllum longipes Rchb.f. Epiphytes with creeping rhizomes. Pseudobulbs distinct, 1-leaved. Leaves persistent. Inflorescence a many-flowered, elongate raceme. Flowers resupinate; pedicel with the basal node above the attachment of the floral bract. Sepals free, fiveveined; lateral sepals 1.6–1.8 times as long as the dorsal sepal. Petals 1-veined. Labellum mobile on a thin ligament, undivided. Column distinctly widened, stigma proximally not protruding, column foot with the lateral sepals attached halfway along its length so that a gap exists between the lower margins of the petals and the upper margins of the lateral sepals, at the apex with lateral wings. Stelidia approximately ½ the length of the column, with or without a tooth along the upper margin, lower margin much widened. Anther anterior surface not concave, anterior margin drawn out; pollinia 4. A single species in lowland forests up to 1500 m in Burma and Thailand. (JJV)

Bulbophyllum sect. Hyalosema Schltr., Repert. Spec. Nov. Regni Veg., 10, 92 (1911). Type species: Bulbophyllum grandiflorum Blume Epiphytes with creeping rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence 1-flowered. Flower resupinate; floral bract tubular; pedicel with the basal node well above the attachment of the floral bract. Dorsal sepal free, 5–9-veined, margins glabrous to papillose or hirsute; lateral sepals free or connate along the lower margins, 5–7-veined, as long as or shorter than the dorsal sepal. Petals 1–3-veined, apices often with a thickened appendage. Labellum mobile on a thin ligament, usually undivided. Column with stigma proximally not protruding, column foot slightly widened toward the apex or not. Stelidia less than ½ the length of the column, with or without a tooth along the upper and lower margin. Anther anterior surface inflated and deeply concave, anterior margin not drawn out; pollinia 4. Approximately 19 species in montane forests up to 1500 m from Thailand and Java eastward to Pacific islands. (JJV)

Bulbophyllum sect. Hirtula Ridl., Mat. Flora Malay. Peninsula, 1, 68 (1908). Type species: Bulbophyllum hirtulum Ridl. Epiphytes with creeping rhizomes. Pseudobulbs distinct to minute compared to the size of the plant, wrinkling with age in a random pattern, 1-leaved. Leaves persistent. Inflorescence usually a many-flowered, subumbellate to elongate raceme. Flowers resupinate; pedicel with the basal node level with or above the attachment of the floral bract. Sepals free, 4.5–32.0 mm long, equal in length, usually 3–5-veined, glabrous to ciliate; usually 3–5-veined. Petals 1-veined, margins ciliate. Labellum mobile on a thin ligament, auriculate. Column with stigma proximally protruding or not. Stelidia approximately ½ the length of the column or shorter, with or without a tooth along the upper and lower margin. Anther anterior surface concave or not, anterior margin drawn out or not; pollinia 4. Forty species in montane forests up to 2500 m from India to China, southward to Java, eastward to the Philippines and Sulawesi. (JJV) Bulbophyllum sect. Hoplandra J.J.Verm., Nordic J. Bot., 26, 147 (2008). Type species: Bulbophyllum restrepia Ridl. Epiphytes with creeping rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence 1-flowered. Flowers resupinate; floral bracts tubular; pedicel with the basal node above the attachment of the floral bract. Dorsal sepal free, 3–5-veined, distally distinctly and often sharply conduplicate; lateral sepals free or connate along the lower margin and/or with the caudate apices entwined, 5–8-veined, 1.6–6.5 times as long as the dorsal sepal. Petals 1–3-veined. Labellum mobile on a thin ligament, undivided. Column with stigma proximally not protruding. Stelidia approximately ½ the length of the column or shorter, often with a tooth along the upper or lower margin. Anther anterior surface curved backward, inflated and deeply concave, anterior margin not drawn out; pollinia 2. Approximately 10 species in forests up 30

Bulbophyllum sect. Hymenobractea Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 699-752 (1913). Type species: Bulbophyllum infundibuliforme J.J.Sm. Epiphytes with creeping rhizomes. Pseudobulbs minute compared to the size of the plant, 1-leaved. Leaves persistent. Inflorescence a 1–10-flowered, patent to erect raceme with distichously arranged flowers. Flowers non-resupinate; floral bracts amplexicaul, often proximally tubular; pedicel with the basal node above the attachment of the floral bract. Sepals 5–9-veined with broadly attached base, the lateral sepals 1.0–1.4 times as long as the dorsal sepal; lateral sepals united at base to column foot forming a mentum. Petals 3–5-veined with broadly attached base, margins entire. Labellum mobile on a thin ligament, distinctly recurved at 1/3 to ½ of its length, entire. Column with stigma proximally often protruding, or with two protruding teeth; column foot present. Stelidia ½ the length of the column or shorter, with or without a tooth along the upper and lower margin. Anther anterior surface not concave, anterior margin not or slightly drawn out; pollinia 4. Two species in forests up to 1500 m in Papua New Guinea. (JJV) Bulbophyllum sect. Imitatores J.J.Verm., in prep. Type species (proposed): Bulbophyllum imitator J.J.Verm. Epiphytes with patent rhizomes. Pseudobulbs distinct to minute compared to the size of the plant, 1-leaved. Leaves persistent. Inflorescences 1- to many-flowered, subumbellate to elongated raceme. Flowers resupinate; floral bracts usually not amplexicaul, pedicel with the basal node above the floral bract attachment. Sepals free, approximately equal, usually 3-veined. Petals 1–3-veined. Labellum mobile on a thin ligament, undivided. Column with the rostellum anterior surface approximately level with the stelidia apices or receding in between these; stigma

BULBOPHYLLUM

proximally not protruding. Stelidia shorter than the length of the column. Anther anterior surface not concave, anterior margin not drawn out (with a small, triangular beak in B. glaucum Schltr.); pollinia 2 or 4. Four species in montane forests up to 2100 m in New Guinea and the western Pacific islands. (JJV) Bulbophyllum sect. Intervallatae Ridl., J. Linn. Soc., Bot., 31, 276. Type species: Bulbophyllum attenuatum Rolfe Epiphytes with creeping rhizomes. Pseudobulbs distinct to minute compared to the size of the plant, 1-leaved. Leaves persistent. Inflorescence a 2- to many-flowered, dense to elongate raceme with distichously arranged flowers. Flowers resupinate, usually one open at a time per inflorescence; floral bracts amplexicaul, often proximally tubular; pedicel with the basal node above the attachment of the floral bract. Sepals free, 5–11-veined, the lateral sepals1.0–1.4 times as long as the dorsal sepal. Petals 1–5-veined, margins entire to distally denticulate. Labellum mobile on a thin ligament, undivided or auriculate. Column with stigma proximally often protruding, or with two protruding teeth. Stelidia ½ the length of the column or less, with or without a tooth along the upper and lower margin. Anther anterior surface not concave, anterior margin not or hardly drawn out; pollinia 4, with or without stipes. Approximately 44 species in forests up to 1400 m in Peninsular Malaysia, Sumatra, Borneo, Sulawesi, the Philippines, New Guinea, and the western Pacific islands. (JJV) Bulbophyllum sect. Lemniscata Pfitz. in Engler & Prantl, Die Natürlichen Pflanzenfamilien, 2, 6, 179 (1888-1889). Type species: Bulbophyllum lemniscatum C.S.P.Parish ex Hook.f. Epiphytes with creeping rhizomes. Pseudobulbs usually 2-leaved. Leaves deciduous. Inflorescence a 2- to many-flowered, subumbellate to elongate raceme. Flowers resupinate; pedicel with the basal node approximately level with the attachment of the floral bract. Dorsal sepal free, 1–5-veined, margins entire to erose to irregularly fimbriate or paleate; lateral sepals free to adnate, usually along the lower margin, 3–5-veined, 1.0–6.9 times as long as the dorsal. Petals usually 1-veined, margins entire to fimbriate or paleate. Labellum mobile on a thin ligament, undivided. Column with stigma proximally not protruding. Stelidia approximately ½ the length of the column or less, often with a wing along the lower margin. Anther anterior surface not concave, anterior margin drawn out into a beak or not; pollinia 4. Approximately 32 species in forests up to 400 m in India, Nepal, Bhutan, China, Burma, Laos, Vietnam, Thailand, Peninsular Malaysia, Sumatra, Java, and Borneo. (JJV) Bulbophyllum sect. Leopardinae Benth. & Hook.f., Gen. Pl., 3, 502 (1880-1883). Type species: Bulbophyllum leopardinum (Wall.) Lindl. Epiphytes with creeping rhizomes, shoots proximally fused to the pseudobulbs. Pseudobulbs one-leaved. Leaves persistent. Inflorescence a 1–4-flowered, congested to elongate raceme with the flowers spirally arranged. Flowers resupinate; floral bracts

tubular; pedicel with the basal node above the attachment of the floral bract. Sepals free, approximately equal, 5–11-veined. Petals 3–7-veined, margins entire to erose. Labellum mobile on a thin ligament, undivided or auriculate. Column with stigma proximally not protruding. Stelidia shorter than ½ the length of the column, deltoid to triangular to subulate, with or without the upper and lower margin. Anther anterior surface not concave, anterior margin usually not drawn out; pollinia 4, with or without stipes. Seven species in forests up to 3300 m in India, Nepal, China, Taiwan, Indochina, Burma, Thailand, and the Philippines. (JJV) Bulbophyllum sect. Lepanthanthe Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 704-875 (1911). Type species: Bulbophyllum lepanthiflorum Schltr. Epiphytes with creeping or pendent rhizomes. Shoots proximally fused to the pseudobulb. Roots verrucate. Pseudobulbs minute compared to the size of the plant, 1-leaved. Leaves persistent. Inflorescence a many-flowered raceme with flowers usually spirally arranged. Flowers resupinate; pedicel with the basal node above the attachment of the floral bract. Sepals equal, free, apex a fleshy, subulate projection, margins glabrous to papillose to ciliate; usually 3-veined, abaxially keeled or not, dorsal sepal free; lateral sepals sometimes connate along their lower margins. Petals 1-veined, margins entire to erose, glabrous to ciliate. Labellum mobile on a thin ligament, undivided or trilobed. Column with the rostellum anterior surface approximately level with the stelidia apices or receding in between these, stigma proximally usually protruding. Stelidia ½ or less the length of the column, often with a distinct tooth along the lower margin. Anther anterior surface not concave, anterior margin drawn out into a beak or not; pollinia 2. Seventeen species in forests up to 3300 m in New Guinea and the western Pacific islands. (JJV) Bulbophyllum sect. Lepidorhiza Schltr., Repert. Spec. Nov. Regni Veg., 10, 93 (1911). Type species: Bulbophyllum amplebracteatum Teijsm. & Binn. Epiphytes with creeping rhizomes. Roots verrucate. Pseudobulbs usually distinct, 1-leaved. Leaves persistent. Inflorescence 2to many-flowered, usually elongate raceme with distichously arranged flowers. Flowers often open one at a time, resupinate; floral bracts usually tubular proximally; pedicel with the basal node usually above the attachment of the floral bract. Sepals 5–13-veined; dorsal sepal free, lateral sepals 1.0–1.7 times as long as the dorsal sepal, often twisted near the base so that the upper margins turn inward, free or adherent along their upper margins. Petals usually 3–12-veined. Labellum mobile on a thin ligament, undivided, auriculate or sub-trilobed. Column with stigma proximally often protruding. Stelidia often with a tooth along the upper and/or lower margin. Anther anterior surface not concave, anterior margin not or hardly drawn out; pollinia 4. Approximately 28 species in forests up to 1900 m in Borneo, Sulawesi, the Philippines, Maluku, New Guinea, and the western Pacific islands. (JJV) 31

DENDROBIEAE

Bulbophyllum sect. Macrocaulia (Blume) Aver., Ident. Guide Viet. Orch., 279 (1994). Type species: Bulbophyllum ovalifolium (Blume) Lindl. Epiphytes with creeping rhizomes. Shoots proximally fused to the pseudobulbs. Pseudobulbs 1-leaved. Leaves usually persistent. Inflorescence 1-flowered. Flowers resupinate; floral bracts tubular; pedicel with the basal node above the attachment of the floral bract. Dorsal sepal usually free, 3–5-veined, margins entire to erose, glabrous to papillose; lateral sepals 1–5 times as long as the dorsal sepal. Petals 1–3-veined, margins entire to erose. Labellum mobile on a thin ligament, usually undivided. Column with stigma proximally protruding or not. Stelidia with or without a slight tooth along the upper and lower margin. Anther anterior surface slightly concave or not, anterior margin drawn out or not; pollinia 2 or 4, stipes absent or present. Approximately 68 species in forests up to 2500 m in India, China, Burma, Laos, Vietnam, Thailand, Peninsular Malaysia, Sumatra, Java, Bali, Borneo, Sulawesi, and the Philippines. (JJV) Bulbophyllum sect. Macrostelydia Garay, Hamer & Siegerist, Nordic J. Bot., 14, 626 (1994). Type species: Bulbophyllum macraei (Lindl.) Rchb.f. Epiphytes with creeping rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence 1-flowered, or a 2- to many-flowered, subumbellate raceme. Flowers resupinate; pedicel with the basal node above the floral bract attachment. Sepals 5-veined, dorsal sepal free; lateral sepals 1.3–4.5 times as long as the dorsal sepal, (slightly) twisted near the base so that the upper margins turn inwards, free to connate along their upper margins. Petals 3-veined. Labellum mobile on a thin ligament, undivided. Stelidia widened distally, apex rounded to truncate to bifid or erose, with a wing low down along the lower margin. Anther anterior surface not concave, anterior margin not or hardly drawn out; pollinia 4. Four species in forests up to 2000 m in India, Sri Lanka, China, Taiwan, Japan, and Vietnam. (JJV) Bulbophyllum sect. Macrouris Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 703-865 (1913). Type species: Bulbophyllum macrourum Schltr. Epiphytes with creeping, straggling or patent rhizomes. Roots usually verrucate. Pseudobulbs distinct to minute compared to the size of the plant, 1-leaved. Leaves persistent. Inflorescence 1- to many-flowered, contracted or elongate raceme. Flowers resupinate; floral bracts sometimes amplexicaul; pedicel with the basal node above the attachment of the floral bract. Sepals usually 1–3-veined, dorsal sepal free; lateral sepals sometimes connate along their lower margin, 1–3 times as long as the dorsal sepal. Petals usually 1-veined, margins entire to erose. Labellum mobile on a thin ligament, undivided. Column with stigma proximally protruding. Stelidia with upper and/or lower margins sometimes with a tooth. Anther anterior surface not concave, anterior margin drawn out into a beak or not; pollinia usually 2. Twenty-eight species in forests up to 3000 m in New Guinea, Maluku, and the western Pacific islands. (JJV) 32

Bulbophyllum sect. Minutissima Pfitz. in Engler & Prantl, Die Natürlichen Pflanzenfamilien, 2, 6, 180 (1889). Type species: Bulbophyllum minutissimum (F.Muell.) F.Muell. Epiphytes with creeping or straggling rhizomes. Pseudobulbs distinct to minute compared to the size of the plant, 1-leaved. Leaves usually persistent. Inflorescence 1-flowered. Flower resupinate; floral bracts usually tubular; pedicel with the basal node above the attachment of the floral bract. Sepals free, approximately equal, 1–3-veined. Petals usually 1-veined. Labellum mobile on a thin ligament, usually undivided. Column with the rostellum anterior surface projecting beyond the stelidia apices to slightly receding in between, stigma proximally protruding or not. Stelidia with upper and/or lower margin sometimes with a tooth. Anther anterior surface concave or not, anterior margin drawn out into a beak or not; pollinia 2 or 4, with or without stipes. Approximately 23 species in forests up to 2400 m in Thailand, Sumatra, and Java to Australia (B. minutissimum (F.Muell.) F.Muell., B. globuliforme Nicholls), New Zealand (B. pygmaeum (Sm.) Lindl.), New Caledonia (B. keekee N.Hallé), and Pacific islands. Most species are from New Guinea. (JJV) Bulbophyllum sect. Monanthaparva Ridl., J. Linn. Soc., Bot., 32, 269. Type species Bulbophyllum striatellum Ridl. Epiphytes with creeping or straggling rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence usually 1-flowered. Flower resupinate; floral bracts tubular; pedicel with the basal node above the attachment of the floral bract. Sepals usually free, approximately equal, usually 3-veined. Petals usually 1-veined, margins entire to erose, glabrous to papillose. Labellum mobile on a thin ligament, undivided or auriculate. Column with stigma proximally often protruding, column foot with or without a dorsal tooth near the apex. Stelidia with upper and/or lower margin often with a tooth. Anther anterior surface concave or not, anterior margin drawn out into a beak or not; pollinia 2 or 4. Approximately 36 species in forests up to 2500 m in China and Vietnam, Thailand, Sumatra, and Java to the Philippines and Sulawesi, with only B. membranaceum Teijsm. & Binn. in New Guinea and Pacific islands. (JJV) Bulbophyllum sect. Monanthes (Blume) Aver., Ident. Guide Viet. Orch., 279 (1994). Bulbophyllum tortuosum (Blume) Lindl. Epiphytes with creeping, straggling or patent rhizomes, shoots either fused proximally to the pseudobulbs or not. Pseudobulbs distinct to minute compared to the size of the plant, 1-leaved. Leaves persistent. Inflorescence 1-flowered. Flower resupinate; floral bracts tubular; pedicel with the basal node level with the attachment of the floral bract. Sepals usually approximately equal, usually 3-veined, margins glabrous to ciliate; the dorsal sepal free; the lateral sepals free or connate along their lower margins. Petals usually 1-veined, margins entire to erose, glabrous to ciliate. Labellum mobile on a thin ligament, undivided, auriculate or trilobed. Column with stigma proximally protruding or not, column foot with or without a dorsal callus at the apex. Stelidia upper and

BULBOPHYLLUM

lower margin with or without tooth. Anther anterior surface not concave, anterior margin usually drawn out into a small beak; pollinia 2 or 4. Approximately 150 species in forests up to 3000 m in India (Darjeeling), China, Indochina to Australia and the Pacific. Most species are from New Guinea. Range extension largely determined by B. clandestinum Lindl., Australia excepted. Endemic species in Maluku, New Guinea, and Australia. (JJV) Bulbophyllum sect. Monomeria (Lindl.) J.J.Verm., in prep. Type species (proposed): Bulbophyllum monomeria J.J.Verm., ined. Epiphytes with creeping rhizomes. Pseudobulbs wrinkling with age along longitudinal lines, one-leaved. Leaves persistent. Inflorescence a many-flowered, elongate raceme. Flowers resupinate; pedicel with the basal node above the floral bract attachment. Sepals free, dorsal sepal 5–7-veined; lateral sepals 1.5–1.8 times as long as the dorsal sepal, upper margins distally converging or adherent, adaxially puberulous to hispid. Petals 1-(–2-) veined, margins entire to fimbriate, glabrous to ciliate. Labellum mobile on a thin ligament, distinctly auriculate. Column foot with the lateral sepals attached approximately halfway along its length so that a gap exists between the lower margins of the petals and the upper margins of the lateral sepals. Stelidia with upper margin winged. Anther anterior surface concave, anterior margin drawn out; pollinia 4, stipe present. Three species in forests up to 1200 m in India, Nepal, Bhutan, China, Vietnam, Thailand, and Peninsular Malaysia. (JJV) Bulbophyllum sect. Monosepalum (Schltr.) J.J.Sm., Nova Guinea, 12, 3–4, 374 (1916). Type species: Bulbophyllum muricatum J.J.Sm. Epiphytes or terrestrials with creeping rhizomes. Pseudobulbs distinct to small compared to the size of the plant, 1-leaved. Leaves persistent. Inflorescence 1-flowered. Flower resupinate, pendent; floral bract tubular; pedicel with the basal node above the attachment of the floral bract. Dorsal sepal connate to the lateral sepals, the lateral sepals free along their lower margins, all approximately equal; 5-veined. Petals 3-veined, margins entire or lobed, with three appendages; appendages differing in texture from the petal itself. Labellum mobile on a thin ligament, undivided. Column with stigma proximally protruding or not, column foot widening and thickening toward the apex, with a dorsal tooth near the apex. Anther anterior surface curved backward, inflated and concave, anterior margin distinctly drawn out; pollinia 4. Three species in forests up to 2500 m in New Guinea. (JJV) Bulbophyllum sect. Papulipetalum Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 700-769 (1913). Type species: Bulbophyllum papulipetalum Schltr. Epiphytes with creeping rhizomes. Pseudobulbs distinct to small compared to the size of the plant, 1-leaved. Leaves persistent. Inflorescence 1-flowered. Flowers resupinate; floral bract tubular; pedicel with the basal node above the attachment of the floral bract. Sepal usually free, 3–5-veined; the lateral sepals 1.0–1.6

times as long as the dorsal sepal. Petals 1-veined, margins entire to finely erose, glabrous to ciliolate. Labellum mobile on a thin ligament, undivided. Column with stigma proximally protruding or not. Stelidia distally often glandular, upper margin with or without a tooth. Anther anterior surface not concave, anterior margin not drawn out; pollinia 2 or 4. Approximately 22 species in forests up to 2250 m in New Guinea, western Pacific islands, and Australia. (JJV) Bulbophyllum sect. Pedilochilus (Schltr.) J.J.Verm. & P.O’Byrne, Bulbophyllum Sulawesi, 199 (2011). Type species: Bulbophyllum papuanum Schltr. Epiphytes with creeping rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence 1-flowered. Flowers resupinate; floral bract tubular; pedicel with the basal node above the attachment of the floral bract. Sepals free, equal, 3–5-veined, margins glabrous to ciliolate. Petals 1-veined, midvein abaxially keeled, margins entire to lacerate, glabrous to ciliolate. Labellum immobile, fused to the column foot by a thick strip of tissue, auriculate, saccate with a small, apical part overhanging the anterior margin. Column with foot thickened toward the apex, ending in one or two calli around which the labellum is attached. Stelidia with upper margin with or without a small tooth. Anther anterior surface concave, anterior margin drawn out; pollinia 2 or 4. Approximately 33 species in forests up to 3550 m in Sulawesi, New Guinea, and the Pacific islands, but most species in New Guinea. (JJV) Bulbophyllum sect. Pelma (Finet) Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 703–855 (1913). Type species: Bulbophyllum absconditum J.J.Sm. Epiphytes with patent rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence 1- to many-flowered, an elongate raceme with distichously arranged flowers. Flowers resupinate; floral bracts tubular; pedicel with the basal node above the attachment of the floral bract. Sepals free, 1–7-veined, lateral sepals 0.9–1.5 times as long as the dorsal sepal. Petals 1–3-veined, margins entire to erose, glabrous to papillose. Labellum on a thin ligament, but usually not mobile, usually undivided. Column with rostellum anterior surface slightly receding in between stelidia apices to distinctly projecting beyond them, stigma proximally usually protruding, column foot usually with a dorsal tooth near the apex that fits into a basal cavity of the labellum. Stelidia often vestigial. Anther anterior surface concave or not, anterior margin not drawn out; pollinia 4, sometimes with a stipe. Twentysix species in forests up to 2800 m in Sumatra, Java, Borneo, Sulawesi, the Philippines, the Moluccas, and New Guinea, eastward to New Caledonia. Most species endemic to New Guinea and the western Pacific islands, the range extension largely determined by that of B. absconditum J.J.Sm. (JJV) Bulbophyllum sect. Peltopus Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 700–760 (1913). Type species: Bulbophyllum peltopus Schltr. 33

DENDROBIEAE

Epiphytes with creeping or straggling rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence 1-flowered. Flowers resupinate; floral bract tubular; pedicel with the basal node above the attachment of the floral bract. Sepals free, approximately equal, 3-(–5-)veined, margins usually entire, glabrous to ciliate. Petals usually vestigial or minute compared to the size of the flower, usually 1-veined, margins entire to erose, glabrous to ciliate. Labellum on a thin ligament but usually not mobile, undivided. Column with the rostellum anterior surface drawn out into a beak-like projection, stigma proximally protruding or not, column foot usually with a dorsal tooth near the apex that fits into a basal cavity of the labellum. Stelidia vestigial. Anther anterior surface concave or not, anterior margin usually not drawn out; pollinia 4, with stipe (easily lost). Thirty-three species in forests up to 2900 m in New Guinea, western Pacific islands, and New Caledonia. (JJV) Bulbophyllum sect. Phreatiopsis J.J.Verm. & P.O’Byrne, Bulbophyllum Sulawesi, 61 (2011). Type species: B. phreatiopse J.J.Verm. Epiphytes with creeping rhizomes. Pseudobulbs minute to small compared to the size of the plant, 1-leaved. Leaves persistent. Inflorescence a many-flowered, elongate raceme. Flowers resupinate; pedicel with the basal node level with attachment of the floral bract. Sepals free, equal, 3-veined, margins glabrous to papillose distally. Petals 1-veined, margins glabrous to papillose. Labellum mobile on a thin ligament, undivided or auriculate; adaxially near the base with a triangular callus. Column foot with or without small lateral teeth near the apex. Stelidia with upper margin with or without a tooth, lower with a wing. Anther anterior surface not concave, anterior margin not drawn out; pollinia 4. Two species in forests up to 1000 m in Peninsular Malaysia, Borneo, Sulawesi, and New Guinea. (JJV) Bulbophyllum sect. Piestobulbon Schltr., Bot. Jahrb. Syst., 58, 139 (1923). Type species: Bulbophyllum piestobulbon Schltr. Epiphytes with ascending to patent rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence 1-flowered. Flowers resupinate; floral bract tubular; pedicel with the basal node above the attachment of the floral bract. Sepals free, approximately equal, 3-veined, margins glabrous to ciliate. Petals 1-veined, margins entire to denticulate, glabrous to ciliate. Labellum mobile on a thin ligament, undivided; adaxially concave at the base, this cavity in front bordered by a transverse ridge notched halfway or with a concave crest line. Column with stigma proximally protruding or not, column foot with a dorsal tooth at the apex. Stelidia with upper margin often with a tooth, lower margin with a tooth. Anther anterior surface not concave, margin drawn out; pollinia 4. Five species in forests of unknown elevation in New Guinea and also the Pacific islands. (JJV) Bulbophyllum sect. Planibulbus J.J.Verm., in prep. Type species (proposed): Bulbophyllum planibulbe Ridl. 34

Epiphytes with creeping rhizomes. Shoots proximally fused to the pseudobulb. Pseudobulbs 1-leaved. Leaves persistent, thick. Inflorescence a 2- to many-flowered, congested or subumbellate raceme. Flowers resupinate; floral bracts amplexicaul or not, pedicel with the basal node level with the attachment of the floral bract. Sepals free, 3-veined; lateral sepals 1.5–2.2 times as long as the dorsal sepal. Petals 1–3-veined, margins entire or erose. Labellum mobile on a thin ligament, undivided. Stelidia with lower margin with a slight tooth. Anther anterior surface not concave, anterior margin not drawn out; pollinia 4. Two species in forests of unknown elevation in Thailand, Peninsular Malaysia, Sumatra, and Borneo. (JJV) Bulbophyllum sect. Plumata J.J.Verm., in prep. Type species (proposed): Bulbophyllum plumatum Ames Epiphytes with creeping rhizomes. Pseudobulbs 1-leaved. Leaves persistent. Inflorescence 1-flowered or a 2–4-flowered, subumbellate raceme. Flowers resupinate; pedicel with the basal node above the attachment of the floral bract. Dorsal sepal 3–17-veined, margins glabrous to minutely ciliolate; lateral sepals 3–16 times as long as the dorsal sepal, connate along upper and usually the lower margin; 5–11-veined. Petals 1–3-veined, apical margin with 12–32 appendages, appendages differing in texture from the petal itself. Labellum mobile on a thin ligament, undivided. Column face furrowed longitudinally. Stelidia with upper and lower margin usually with a tooth. Anther anterior surface deeply concave, anterior margin not drawn out; pollinia 4. Four species in forests up to 1460 m in Thailand, Peninsular Malaysia, Sumatra, Java, Borneo, and the Philippines. (JJV) Bulbophyllum sect. Polymeres (Blume) J.J.Verm. & P.O’Byrne, Gard. Bull. Singapore, 60, 126 (2008). Type species: Bulbophyllum tenuifolium (Blume) Lindl. Epiphytes with creeping to patent rhizomes; shoots free or fused proximally to the pseudobulbs. Pseudobulbs distinct to small compared to the size of the plant, 1-leaved. Leaves persistent. Inflorescence 1-flowered. Flower resupinate; floral bracts tubular; pedicel with the basal node above the attachment of the floral bract. Sepals usually free, usually approximately equal, 3-(4–9-)veined, margins glabrous to papillose or ciliate. Petals usually 1-veined, margins entire to fimbriate, glabrous to ciliate. Labellum mobile on a thin ligament, undivided or auriculate. Column with stigma often proximally protruding, column foot widened and often thickened and usually with 2 lateral teeth toward its apex. Stelidia with upper margin sometimes with a tooth. Anther anterior surface concave or not, anterior margin not drawn out; pollinia usually 4. Approximately 164 species in forests up to 3400 m in India (Andaman Islands), Thailand, Sumatra, Java, eastward to the western Pacific islands. Morphologically, B. sect. Polymeres grades into B. sect. Brachypus and B. sect. Papulipetalum. Uniting all three would leave a group without a distinguishing character (the latter two do not have a widened and winged column foot typical for B. sect. Polymeres), and keeping all three apart yields sectional boundaries that are partly arbitrary. The boundaries also divide a few species groups

BULBOPHYLLUM

that otherwise are similar. Here, the three groups are kept separate, with the note that further investigation is needed, which should include the use of characters now ignored, such as rostellum shape. (JJV) Bulbophyllum sect. Pseudopelma J.J.Verm. & P.O’Byrne. Gard. Bull. Singapore, 55, 130 (2003). Type species: Bulbophyllum pseudopelma J.J.Verm. Epiphytes with creeping or straggling rhizomes; shoots developing from nodes below the pseudobulbs, not fused to them; bracts finely and densely papillose, spiral vessels apparent when decaying. Roots spreading, densely and coarsely ­ papillose. Pseudobulbs 1-leaved. Leaves persistent, thin, veins inconspicuous. Inflorescences developing from nodes below the pseudobulbs, or from nodes scattered along the rhizome, solitary or few-fasciculate, 1–5-flowered, elongate racemes; peduncle bracts 2 or 3; rachis not thickened. Flowers resupinate, open simultaneously if racemose; floral bracts tubular; pedicel with the basal node level with the floral bract attachment, approximately as long as or longer than the ovary. Sepals free, approximately equal, 7–9 mm long, elliptic to triangular, acute to acuminate, glabrous, margins entire. The dorsal sepal 3–5-veined, the lateral sepals 3-veined. Petals 3.2–3.6 mm long, elliptic(–ovate), rounded to obtuse, 3-veined, margins entire, glabrous. Labellum mobile on a thin ligament, undivided, 1.3–2.0 mm long, elliptic to ovate or sub-orbicular, margins entire, glabrous, adaxially with two distinct ridges. Column with the rostellum anterior surface level with (or receding in between) the stelidia apices, stigma with the proximal edge truncated by a transverse ridge or a triangular tooth; column foot not widened toward its apex, with or without two deltoid, rounded lobes near the apex. Stelidia approximately ½ the length of the column or less, triangular, with a patent to antrorse, deltoid, rounded to obtuse wing along the lower margin. Anther abaxially with or without a crest, anterior surface not concave, anterior margin not drawn out; pollinia 4, the inner ones more than half as long as the outer ones. Two species in lowland forests in Sulawesi. (JJV) Bulbophyllum sect. Racemosae Benth. & Hook.f., Gen. Pl., 3, 502 (1883). Type species: Bulbophyllum careyanum (Hook.) Spreng. Epiphytes with creeping to occasionally straggling rhizomes; shoots developing from nodes below the pseudobulbs, not fused to them; bracts approximately glabrous, spiral vessels unknown. Roots spreading, approximately glabrous to finely hirsute locally. Pseudobulbs 1-leaved. Leaves persistent, thick, veins inconspicuous. Inflorescences developing from nodes below pseudobulbs, solitary, many-flowered, contracted to elongate racemes with flowers spirally arranged, opening simultaneously; peduncle bracts 3–7; rachis thickened or not. Flowers resupinate; floral bracts not amplexicaul; pedicel with the basal node level with the attachment of the floral bracts, or less than the diameter of the pedicel above it, approximately as long as the ovary or less. Dorsal sepal free, 3–8 mm long, (ob-)ovate to elliptic, rounded to acuminate, 3–7-veined, margins entire,

glabrous, surface glabrous, abaxially sometimes verrucate; lateral sepals like the dorsal sepal but 1.2–2.3 times longer, (loosely) adhering along the lower margin, 5.5–12.0 mm long, ovate, subacute to acuminate. Petals 1.5–4.5 mm long, ovate to triangular, approximately acute to caudate, glabrous, 1-veined (3-veined in B. allenkerrii Seidenf., B. bittnerianum Schltr.), margins entire to erose or denticulate. Labellum mobile on a thin ligament, auriculate, 2–5 mm long, oblong to ovate to triangular, margins entire, glabrous to papillose to ciliolate; adaxially with two ridges, sometimes also with a dorsal ridge, surface glabrous to papillose or hirsute, abaxial surface glabrous, auricles attached along the basal ¼–²∕₅ of the labellum, antrorse, margins approximately entire or erose to denticulate, glabrous. Column with the rostellum anterior surface receding in between the stelidia apices, stigma proximally not protruding (with a tooth in B. tricorne Seidenf. & Smitinand, B. tricornoides Seidenf.), column foot not widened toward the apex. Stelidia approximately ½ the length of the column or less, distinct, triangular, sometimes with a small tooth along the upper margin. Anther abaxially with a (low) rounded crest, anterior surface concave or not, anterior margin drawn out or not; pollinia 4, the inner ones more than half as long as the outer ones. Approximately 38 species in forests up to 1200 m in India, Nepal, Bhutan, Burma, China, Vietnam, Laos, Thailand, and Peninsular Malaysia. (JJV)

Bulbophyllum sect. Recurvae (Garay, Hamer & Siegerist) J.J.Verm., in prep. Type species (proposed): Bulbophyllum corolliferum J.J.Sm. Epiphytes with creeping rhizomes; shoots developing from nodes below the pseudobulbs, not fused to them. Roots spreading and glabrous to hirsute. Pseudobulbs small compared to the size of the plant in B. nanobulbon Seidenf., 1-leaved. Leaves persistent, veins inconspicuous. Inflorescences developing from nodes below the pseudobulbs, 2- to many-flowered, subumbellate racemes, with flowers spirally arranged (1-flowered in B. lineatum (Teijsm. & Binn.) J.J.Sm., B. subbullatum J.J.Verm.); peduncle bracts 4–6; rachis, if present, thickened or not. Flowers per inflorescence all open simultaneously if racemose, resupinate; floral bracts not amplexicaul; pedicel with the basal node equal to its diameter above the floral bract attachment or less, approximately as long as the ovary or shorter. Dorsal sepal free, 3.0–7.5 mm long, elliptic to ovate, rounded to caudate, 3–5-veined, margins entire to erose, fimbriate, glabrous or adaxially papillose to hirsute distally; lateral sepals as for dorsal but 2–10 times as long (approximately equal in length in B. subbullatum), twisted near the base so that the upper margins turn inward (not so in B. subbullatum), free or adherent or connate along their upper margins, sometimes also along the lower margins, 10–50 mm long, margins entire to (minutely) erose, upper margin slightly fimbriate to glabrous; surface glabrous to papillose or hirsute abaxially. Petals 1.5–7.5 mm long, obovate, elliptic to rhombiform to triangular, obtuse to caudate, 3-veined, margins entire to erose, fimbriate, adaxial surface glabrous to papillose to hirsute distally. Labellum mobile on a thin ligament, undivided or auriculate, 2.2–4.2 mm long, oblong (with a slightly widened apical part) to ovate with a widened base, margins entire, usually glabrous; 35

DENDROBIEAE

adaxially without ridges, or with two inconspicuous to distinct ridges (starting near the base of the labellum as conical knobs), often with one more inconspicuous to distinct ridge in between, adaxial surface glabrous to partly verrucose, rugose or hirsute, abaxial surface glabrous, auricles if present attached near the base, patent, margins entire, glabrous. Column with the rostellum anterior surface receding in between the stelidia apices, stigma proximally not protruding, column foot tapering toward its apex or not, with or without slight lateral lobes at the apex. Stelidia approximately ½ the length of the column or (much) shorter, porrect to downward-falcate, truncate to rounded, acute or long-subulate, with or without an (antrorse) tooth along the upper margin, with or without a patent, deltoid, rounded wing along the lower (with a triangular, obtuse wing with an erose anterior margin in B. serratotruncatum Seidenf.). Anther abaxially with rounded crest, anterior surface concave, anterior margin drawn out into a rounded beak; pollinia 4, the inner ones more than half as long as the outer ones. Approximately 36 species in forests up to 1700 m in India (Andaman Islands), Bangladesh, China, Taiwan, Burma, Thailand, Laos, Vietnam, Peninsular Malaysia, Sumatra, Java, Borneo, the Philippines, Sulawesi, and Maluku. (JJV) Bulbophyllum sect. Reptantia J.J.Verm., Edinburgh J. Bot., 58, 121 (2001). Type species: Bulbophyllum reptans Lindl. Epiphytes with creeping or straggling rhizomes; shoots developing from nodes below the pseudobulbs, not fused to them; bracts glabrous to colliculate, distally sometimes papillose, spiral vessels unknown. Roots spreading, glabrous to verrucate, hirsute locally. Pseudobulbs distinct, 1-leaved, Leaves persistent, thin, veins inconspicuous. Inflorescences developing at the base of developing shoots, solitary, two- to many-flowered, elongate racemes with flowers spirally arranged or imperfectly distichous; peduncle bracts 3–7; rachis not thickened. Flowers resupinate, opening simultaneously; floral bracts not amplexicaul; pedicel with the basal node level with the floral bract attachment or less than half the diameter of the pedicel above this, longer than the ovary. Sepals free, the dorsal sepal 4–11 mm long, elliptic to ovate, acute, 3–5-veined, margins entire, glabrous; lateral sepals as for the dorsal but approximately 1.0–1.5 times as long; 4–16 mm long, elliptic to triangular. Petals 2.0–3.5 mm long, elliptic to oblong, obovate to spathulate, rounded to obtuse, 1–3-veined, margins entire, glabrous. Labellum mobile on a thin ligament, undivided, 2–5 mm long, elliptic to oblong, margins entire, glabrous, adaxially without ridges or with two ridges, adaxial surface glabrous (papillose along the dorsal line in B. triviale Seidenf.), abaxial surface glabrous. Column with the rostellum anterior surface receding in between the stelidia apices, stigma proximally not protruding (a callus lower down on the column foot in B. reptans (Lindl.) Lindl. ex Wall.), column foot slightly widening toward its apex or not. Stelidia approximately ½ the length of the column or shorter, triangular to almost subulate, upper margin with or without a small, deltoid, obtuse tooth, lower margin with or without a deltoid, obtuse wing. Anther abaxially with a low rounded crest, anterior surface concave, anterior margin not drawn out; pollinia 4, the inner ones more than half as long as 36

the outer ones. Six species in montane forests up to 2800 m in India, Nepal, Bhutan, and China, southward to Thailand. (JJV) Bulbophyllum sect. Rhinanthera, J.J.Verm., in prep. Type species (proposed): Bulbophyllum turpe J.J.Verm. & P.O’Byrne Epiphytes with creeping rhizomes. Shoots developing from nodes below pseudobulbs, not fused to them; bracts minutely papillose. Roots spreading, glabrous to minutely hirsute locally. Pseudobulbs distinct, 1-leaved. Leaves persistent, thin to thick, veins inconspicuous. Inflorescences developing from nodes below the pseudobulb, solitary, elongate racemes with flowers spirally arranged; peduncle bracts 4–10; rachis not thickened. Flowers resupinate, opening simultaneously; floral bracts not amplexicaul; pedicel with the basal node 0.5–1.5 times its diameter above the floral bract attachment, distinctly longer than the ovary. Sepals free, the dorsal sepal 6–10 mm long, obovate to ovate-triangular, rounded to acuminate, 3-veined, margins entire, glabrous; lateral sepals equally long or shorter, 4.5–9.0 mm long, obliquely ovate to triangular. Petals 2.5–5.5 mm long, (sub-)spatulate to elliptic to oblong to triangular, rounded to subacute, 1-veined, margins entire, glabrous. Labellum mobile on a thin ligament, undivided or sub-trilobed, 1.8–3.0 mm long, oblong to ovate, or divided in a wide, ovate basal part and a drawn-out, narrow, oblong to ovate apical part, margins entire, glabrous, adaxially with two short, knob-like ridges close to the base or with two distinct ridges more distally positioned. Column with the rostellum anterior surface almost level with the stelidia apices or receding in between, stigma proximally with a protruding tooth, column foot not widened toward the apex but with a dorsal tooth at the apex. Stelidia approximately ½ the length of the column (slightly longer in B. wrayi Hook.f.), triangular to subulate, with or without a rounded to triangular wing along the upper margin, with or without a deltoid to triangular, obtuse wing along the lower margin. Anther abaxially with a low, rounded crest, anterior surface concave, anterior margin distinctly drawn out into an elliptic to triangular beak (curved backward in B. tahanense Carr); pollinia 4, the inner ones slightly shorter than the outer ones. Four species in montane forests up to 2100 m in Peninsular Malaysia, Sumatra, and Java. (JJV) Bulbophyllum sect. Rhytionanthos (Garay, Hamer & Siegerist) J.J.Verm., in prep. Type species (proposed): Bulbophyllum helenae (Kuntze) J.J.Sm. Epiphytes with creeping rhizomes. Shoots developing from nodes below the pseudobulbs, not fused to them; bracts glabrous to papillose. Roots spreading, glabrous to hirsute or papillose. Pseudobulbs distinct, 1-leaved. Leaves persistent, thick, veins inconspicuous. Inflorescences developing from nodes below the pseudobulb, solitary, 2- to many-flowered, subumbellate racemes, with flowers spirally arranged (also 1-flowered in B. paraemarginatum Aver., B. rheedei Manilal & C.S.Kumar); peduncle bracts 3–5; rachis thickened or not. Flowers resupinate, opening simultaneously if racemose; floral bracts not amplexicaul; pedicel with the basal node less than diameter of the pedicel above floral bract attachment, as long as or longer than ovary. Dorsal

BULBOPHYLLUM

sepal free, 4–9 mm long, elliptic to (ob-)ovate, rounded to caudate, 5-veined, surface glabrous or abaxially verrucose to papillose, margins entire to erose to papillose (coarsely and irregularly papillose in B. chondriophorum (Gagnep.) Seidenf.); lateral sepals free or adherent or connate along their upper margins, sometimes also along the lower margins, otherwise as for the dorsal sepal but 2–5 times as long (only slightly longer in B. bomiense Z.H.Tsi), twisted near the base so that the upper margins turn inward, 7–42 mm long, margins entire or (upper margin) slightly erose (proximally). Petals 2.5–5.0 mm long, obovate to elliptic to triangular, rounded to caudate, 3-veined, margins erose to denticulate, glabrous to papillose (coarsely and irregularly papillose in B. chondriophorum). Labellum mobile on a thin ligament, undivided, 1.9–3.5 mm long, oblong to ovate to triangular with a widened base, margins entire, adaxially without ridges, or with two ridges, surface glabrous or slightly verruculose distally, abaxial surface glabrous. Column with the rostellum anterior surface receding in between the stelidia apices, stigma proximally not protruding, column foot tapering toward its apex or not, with or without slight lateral lobes at the apex. Stelidia shorter than ½ the length of the column, porrect, triangular, acute to shortsubulate, sometimes with an (antrorse) tooth along the upper margin, often with an inconspicuous, patent, deltoid, obtuse wing along the lower. Anther abaxially with low to high, rounded crest, anterior surface concave, anterior margin drawn out into a rounded beak; pollinia 4, the inner ones more than half as long as the outer ones. Approximately 16 species in forests up to 1500 m in India, Sri Lanka, Nepal, China, Burma, Thailand, Laos, Vietnam, Peninsular Malaysia, Sumatra, Java, Borneo, and the Philippines. (JJV)

ovate to triangular (subhastate with minute, patent, acute teeth in B. sp. from the Philippines), acute to acuminate (approximately ligulate and subacute in B. fraternum J.J.Verm. & P.O’Byrne), 1-veined, margins entire to slightly erose, glabrous to papillose to ciliate, adaxial surface glabrous to papillose or hirsute distally. Labellum mobile on a thin ligament, undivided or sub-trilobed (auriculate in B. uviflorum P.O’Byrne), 1.7–6.0 mm long, panduriform, spatulate, or ovate to hastate with patent to retrorse lateral lobes, margins entire (coarsely denticulate in B. uviflorum), glabrous to papillose; adaxially sometimes with a narrow transverse ridge near the base, without ridges or with two inconspicuous ridges, adaxial surface glabrous to finely papillose, abaxial surface approximately glabrous, or with a papillose patch in the centre (near the apex in B. condylochilum J.J.Verm. & P.O’Byrne). Column with the rostellum anterior surface receding in between the stelidia, stigma proximally protruding, with a transverse ridge with a bifid crest (in some species the ridge itself may be absent except for two teeth or wings along the edges of the column face), column foot tapering toward its apex, with or without lateral teeth near the apex. Stelidia ½ the length of the column or shorter, elliptic, oblong to wedge-shaped, or sub-triangular, distinctly sigmoid near the apex, upper margin with an elliptic, oblong to (obliquely) wedge-shaped, or sub-triangular wing much larger than the stelidia itself (stelidia no more than a minute tooth along the lower margin of the wing) and widely overtopping this. Anther abaxially with a low to high and conical crest, anterior surface concave or not, anterior margin widely drawn out into a beak or not; pollinia 4, with the inner ones almost as long as the outer ones. Approximately 16 species in forests up to 1200 m in Sulawesi and the Philippines. (JJV)

Bulbophyllum sect. Saurocephalum J.J.Verm., in prep. Type species (proposed): Bulbophyllum saurocephalum Rchb.f. Epiphytes with creeping, erect to pendent rhizomes; shoots developing from nodes below the pseudobulbs, not fused to them; bracts glabrous to papillose. Roots spreading, developing only below the older pseudobulbs in species with an erect to pendent rhizome, growing over or alongside the rhizome toward the substrate, glabrous to finely hirsute locally. Pseudobulbs distinct, 1-leaved. Leaves persistent, thick, veins inconspicuous. Inflorescences developing from nodes below pseudobulbs, many-flowered, elongate racemes with flowers spirally arranged; peduncle bracts 3–5; rachis distinctly thickened or not. Flowers resupinate, opening simultaneously; floral bracts not amplexicaul; pedicel with the basal node level with the floral bract attachment, as long as the ovary or shorter. Sepals free, the dorsal, 2.8–7.0 mm long, (ob-)ovate to elliptic, deeply hooded toward the apex, rounded to obtuse (also apiculate in B. hastiferum Schltr.), 3-veined, margins entire, glabrous to papillose to ciliate, adaxially glabrous, abaxially glabrous to papillose or pubescent; lateral sepals as for the dorsal but usually (partly) adherent to connate along their lower margins (together often forming a dish- or bowl-shaped structure around the labellum), 0.8–1.4 times as long as the dorsal sepal, 2.8–7.5 mm long, ovate to triangular (sometimes with a drawn-out apical part), flat toward the apex, obtuse to acute, 3-(–4-)veined. Petals 1.5–7.0 mm long,

Bulbophyllum sect. Schistopetalum Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 700–63 (1913). Type species: Bulbophyllum schistopetalum Schltr. Epiphytes with creeping rhizomes, often spreading downward along the substrate; bracts glabrous to minutely colliculate. Shoots developing from nodes below pseudobulbs, not fused to them. Roots spreading, distinctly papillose. Pseudobulbs distinct to small compared to the size of the plant, 1-leaved. Leaves persistent, thick, veins inconspicuous. Inflorescences developing from nodes below the pseudobulbs, single to many-fasciculate on short lateral sympodia, 1-flowered; peduncle bracts two or three. Flowers resupinate, opening one at a time; floral bracts tubular; pedicel with the basal node the same size as the diameter of the pedicel above the floral bract attachment, longer than the ovary (as long as the ovary in B. fissipetalum Schltr.). Sepals free, equal, 4.5–23.0 mm long, elliptic to ovate-triangular, subacute to caudate, glabrous, 3–7-veined, margins entire; glabrous. Petals 0.3–3.0 mm long excluding appendages if present, a narrow seam alongside the column base, or elliptic to triangular, caudate (also acute in B. hassalii Kores), or distal margin with 3–9 tapering, caudate appendages, glabrous, 1–3-veined, margins otherwise entire; appendages immobile, not differing in texture from the petal itself, triangular, acute to caudate. Labellum mobile on a thin ligament, undivided, 1.2–13.0 mm long, elliptic to triangular, margins entire, (partly) ciliate to long-hairy (glabrous in 37

DENDROBIEAE

B. aristopetalum Kores, B. hassalii), projecting near the base of the labellum, then converging, usually until they almost meet, at that level usually clasping two ridges running parallel, close to the dorsal ridge, beyond that level margins spreading again, adaxial surface glabrous, or distally hirsute or long-hairy, abaxial surface glabrous to distally long-hairy. Column with rostellum anterior surface receding in between the stelidia, stigma proximally usually with a distinct, rounded callus, column foot tapering toward its apex. Stelidia ½ the length of column or longer; distinct, narrowly triangular to subulate, with or without a small deltoid to triangular, rounded to obtuse tooth along the upper margin, with a larger, patent to antrorse, rounded to obtuse wing or tooth along the lower. Anther abaxially with a (low) rounded crest, anterior surface concave or not, anterior margin usually drawn out, emarginate; pollinia 2. Ten species in forests of unknown elevation in New Guinea and eastward to Samoa. (JJV) Bulbophyllum sect. Sestochilus (Breda) Benth. & Hook.f., Gen. Pl., 3, 502 (1883). Type species: Bulbophyllum lobbii Lindl. Epiphytes with creeping, sometimes straggling rhizomes; shoots usually developing from nodes below the pseudobulbs, not fused to them; rhizome bracts glabrous. Roots developing from nodes along the rhizome, spreading, glabrous to coarsely papillose. Pseudobulbs distinct to minute compared to the size of the plant, 1-leaved. Leaves persistent, thin to thick, veins inconspicuous. Inflorescences usually developing from nodes below the pseudobulbs, also from nodes scattered along the rhizome, solitary, 1-flowered; peduncle bracts 2–6. Flowers resupinate; floral bracts amplexicaul, not tubular or tubular near the base only; pedicel with the basal node more than 1.5 times the diameter of the pedicel above the floral bract attachment, distinctly longer than the ovary. Sepals free, equal or lateral sepals shorter; 7–95 mm long, elliptic, (ob-)ovate to triangular, obtuse to acuminate, glabrous, 5–14-veined, margins entire, glabrous. Petals 6–77 mm, elliptic, (ob-)ovate to triangular, obtuse to acuminate, glabrous, 5–12-veined, margins entire, glabrous. Labellum mobile on a thin ligament, undivided, or with distinct, retrorse auricles near the ligament, (trilobed with distinct, patent, rounded lateral lobes halfway along the labellum in B. capillipes C.S.P.Parish & Rchb.f., B. monanthos Ridl., with rounded lateral lobes near the base in B. apheles J.J.Verm.), 3.5–35.0 mm long, elliptic or oblong to (ob-)ovate to triangular, or with a wide, triangular, hastate to cordate basal part and a narrow, drawn out apical part, adaxially without ridges, or with two ridges, adaxial surface glabrous, sometimes partly papillose to hirsute, abaxial surface glabrous, margins (including those of the lateral lobes, if present) entire, glabrous to papillose. Column with rostellum anterior surface level with the stelidia apices or receding in between the stelidia, stigma proximally not protruding, column foot tapering toward the apex or not. Stelidia less than ½ the length of column, often inconspicuous, deltoid to triangular, without a tooth along the upper margin, with or without a deltoid, rounded or obtuse wing along the lower margin. Anther abaxially with a low, inconspicuous and rounded crest to a distinct conical one, anterior surface not concave, anterior margin not drawn out; pollinia 4, usually equal, in some species with a hamular stipe. Approximately 28 species in forests up to 2800 m in India, 38

Nepal, Bhutan, Burma, China, Taiwan, Japan, Vietnam, Laos, Thailand, Peninsular Malaysia, Sumatra, Java, Borneo, and the Philippines, eastward to the Solomon Islands. (JJV) Bulbophyllum sect. Stachysanthes (Blume) Aver., Ident. Guide Viet. Orch., 276 (1994). Type species: Bulbophyllum gibbosum (Blume) Lindl. Epiphytes with creeping, erect or pendent rhizomes; bracts glabrous to colliculate. Shoots developing from the basal node of the pseudobulbs, not fused to them, or from or (well) above the node below basal node of pseudobulb. Roots spreading; in species with a patent to pendent rhizome below the older pseudobulbs only, growing along the rhizome toward the substrate, glabrous to finely hirsute locally (spreading in B. rhizomatosum Ames & C.Schweinf.). Pseudobulbs minute compared to the size of the plant, 1-leaved. Leaves persistent, thin to thick, veins inconspicuous. Inflorescences usually developing from nodes below pseudobulbs, also from nodes scattered along the rhizome, solitary or sometimes two together, 2- to many-flowered, elongate or contracted racemes with flowers spirally arranged, or sometimes few-fasciculate and 1-flowered; peduncle bracts 3–9; rachis, if present, not thickened (slightly thickened in B. succedaneum J.J.Sm.). Flowers resupinate, successively or simultaneously; floral bracts not amplexicaul (amplexicaul, not tubular in some species with 1- or few-flowered inflorescences); pedicel with the basal node level with the floral bract attachment, distinctly shorter to longer than the ovary. Sepals free (in some species all three adherent to connate with a lumen between the lateral sepals); 2–40 mm long, elliptic, (ob-)ovate to triangular, rounded to caudate, 3-veined (5-veined in B. succedaneum), margins entire; lateral sepals as the dorsal but 1.0–1.5 times as long. Petals 1–7 mm long, elliptic to (ob-)ovate, obtuse to acuminate, glabrous to papillose, adaxial surface glabrous to minutely papillose distally, 1-veined, margins entire to erose distally. Labellum mobile on a thin ligament, undivided, 1–9 mm long, elliptic to (ob-)ovate to oblong, , glabrous to papillose to ciliate, adaxially without ridges, or often with two ridges, and/or close to the base with 2 short, converging ridges, knobs or teeth, or with 2 more parallel ridges slightly farther above the base, sometimes with two (parallel) ridges on the distal part of the labellum (e.g. in B. obtusipetalum J.J.Sm., B. sigmoideum Ames & C.Schweinf.) (distally with a dorsal ridge in B. armeniacum J.J.Sm., B. odoratum (Blume) Lindl.), surface glabrous to papillose distally (glandular-hairy in B. armeniacum), abaxial surface glabrous, margins entire to erose. Column with the rostellum anterior surface receding in between the stelidia (level with the stelidia apices in B. brevicolumna J.J.Verm., B. devogelii J.J.Verm.), stigma proximally protruding or not, or with a distinct tooth, column foot tapering toward its apex, with or without a dorsal callus or lateral auricles at the apex. Stelidia shorter than length of the column; deltoid to triangular, upper margin without teeth (also with a tooth in B. odoratum), lower margin often with a deltoid to triangular, rounded to acute wing or tooth. Anther with a (low) rounded, abaxial crest, anterior surface concave or not, anterior margin drawn out into a beak or not; pollinia usually 2 (4 with the inner ones shorter than the outer ones in B. ionophyllum J.J.Verm. and B. sopoetanense Schltr.), without or with

BULBOPHYLLUM

a single stipe. Approximately 66 species in forests up to 2500 m in India, China, and Taiwan, eastward to the western Pacific islands. Numerous species in Peninsular Malaysia, Sumatra, Java, Borneo, the Philippines, and Sulawesi but elsewhere represented by one or a few species only. (JJV)

Bulbophyllum sect. Stenochilus J.J.Sm., Bull. Jard. Bot. Buitenzorg, ser. 2, 13, 33 (1914). Type species: Bulbophyllum macranthum Lindl. Epiphytes with creeping, sometimes straggling rhizomes; bracts glabrous. Shoots usually developing from nodes below pseudobulbs, not fused to them. Roots developing from nodes along rhizome, spreading, glabrous to coarsely papillose. Pseudobulbs distinct to minute compared to the size of plant, 1-leaved. Leaves persistent, thin to thick, veins inconspicuous. Inflorescences usually developing from nodes below the pseudobulbs, also from nodes scattered along the rhizome, solitary, 1-flowered (2–5-flowered racemes with flowers spirally arranged in B. anceps Rolfe); peduncle bracts 2–6. Flowers simultaneously opening in B. anceps, non-resupinate; floral bracts amplexicaul, not tubular or tubular near the base only; pedicel with the basal node more than 1.5 times the diameter of the pedicel above the floral bract attachment, distinctly longer than the ovary. Sepals free, equal or lateral sepals shorter; 7–95 mm long, elliptic, (ob-)ovate to triangular, obtuse to acuminate, glabrous, 5–14-veined, margins entire. Petals 6–77 mm, elliptic, (ob-)ovate to triangular, obtuse to acuminate, glabrous, 5–12-veined, margins entire. Labellum mobile on a thin ligament, undivided, or with distinct, retrorse auricles near the ligament, 3.5–35.0 mm long, elliptic or oblong to (ob-)ovate to triangular, or with a wide, triangular, hastate to cordate basal part and a narrow, drawn out apical part, glabrous to papillose, adaxially without ridges, or with two ridges, adaxial surface glabrous, sometimes partly papillose to hirsute, abaxial surface glabrous. Margins (including those of the lateral lobes, if present) entire. Column with the rostellum anterior surface level with the stelidia apices or receding in between the stelidia, stigma proximally not protruding, column foot tapering toward the apex or not. Stelidia less than ½ the length of the column, often inconspicuous, deltoid to triangular, without a tooth along the upper margin (upper margin also erose in B. affine Wall. ex Lindl., with an antrorse, deltoid tooth in B. anceps), with or without a deltoid, rounded or obtuse wing along the lower margin. Anther abaxially with a low, inconspicuous and rounded crest to a distinct conical one, anterior surface not concave, anterior margin not drawn out; pollinia 4, equal. Approximately 12 species in forests up to 2800 m in India, Nepal, Bhutan, Burma, China, Taiwan, Japan, Vietnam, Laos, Thailand, Peninsular Malaysia, Sumatra, Java, Borneo, and the Philippines, eastward to the Solomon Islands. (JJV) Bulbophyllum sect. Sunipia (Lindley) J.J.Verm., in prep. Type species (proposed): Bulbophyllum sunipia J.J.Verm., ined.

Epiphytes or lithophytes with creeping rhizomes; bracts glabrous to papillose; shoots pseudobulbs, spreading, glabrous to hirsute locally. Pseudobulbs 1-leaved. Leaves persistent, thin to thick, veins inconspicuous. Inflorescences developing from nodes below pseudobulbs, solitary, 1- to many-flowered, elongate racemes with distichously arranged flowers; peduncle bracts 4–8; rachis, if present, not thickened. Flowers resupinate, opening simultaneously; floral bracts amplexicaul, not tubular; pedicel with the basal node level with the floral bract attachment, shorter to distinctly longer than ovary. Dorsal sepal free, 4–25 mm long, elliptic to triangular, emarginate to rounded to acute-acuminate, glabrous, 3–7-veined, margins entire (rarely erose); lateral sepals as for the dorsal sepal but free or adherent along their lower margin, 1.0–1.3 times as long as the dorsal sepal, 5–27 mm long, rounded to acute-acuminate, margins entire. Petals 1.7–12.0 mm long, orbicular, elliptic(-oblong) to (broadly) ovate to triangular, rounded to acute, surface glabrous, 1–7-veined, margins entire to minutely erose to irregularly denticulate, glabrous to partly ciliate. Labellum not mobile, attached to column foot by a thick strip of tissue, undivided or sub-trilobed, 4–23 mm long, elliptic to (ob-)ovate to triangular, often with a drawn-out, linear apical part; adaxially without ridges, or with a dorsal ridge, or with 2 ridges that are highest and often knob-shaped near the base, which start close together and either run parallel close to the dorsal line or spread, surface glabrous (distally papillose in B. grandiflorum Blume), abaxial surface glabrous, margins entire to erose or denticulate, glabrous. Column with rostellum anterior surface protruding in between stelidia, stigma proximally not protruding, column foot not widened nor thickened toward the apex. Stelidia vestigial. Anther abaxially with a (low) rounded crest, anterior surface not concave, anterior margin not drawn out; pollinia 4, equal, with one v-shaped stipe, with two separate stipes with a single viscidium, or with two separate viscidia or one entire stipe. Approximately 22 species in forests up to 2700 m in India, Nepal, Bhutan, China, Taiwan, Burma, Laos, Thailand, and Vietnam. (JJV) Bulbophyllum sect. Tapeinoglossum (Schltr.) J.J.Sm., Nova Guinea, 12, 3–4: 406 (1916). Type species: Bulbophyllum centrosemiflorum Schltr. Epiphytes with creeping or straggling rhizomes; bracts glabrous. Shoots usually developing from nodes below the pseudobulbs, not fused to them. Roots spreading, glabrous, hirsute in patches. Pseudobulbs distinct, 1-leaved. Leaves persistent, thin, veins inconspicuous. Inflorescences developing from nodes scattered along the rhizome, solitary or few-fasciculate, 1-flowered; peduncle bracts 3. Flowers resupinate; floral bracts tubular; pedicel with the basal node greater than the diameter of the pedicel above the floral bract attachment, longer than the ovary. Dorsal sepal connate to the lateral sepals for a short distance near the base, 15–16 mm long, elliptic to obovate, obtuse, surface shortly papillose-hirsute toward the margins, 5-veined, margins entire, ciliolate; lateral sepals as for the dorsal but shorter, connate along the lower margins, c. 11 mm long, margins glabrous to papillose or hirsute, adaxial surface long-hirsute, shortly papillosehirsute abaxially. Petals c. 2 mm long, elliptic to ovate, rounded 39

DENDROBIEAE

to obtuse, glabrous, 3-veined, margins entire. Labellum mobile on a thin ligament, undivided, 2.0–2.5 mm long, ovate, adaxially with two parallel, hirsute ridges, surface otherwise glabrous, margins entire, glabrous. Column with the rostellum anterior surface clearly receding in between stelidia; stigma proximally not protruding, column foot deeply concave, halfway along its length constricted by the inward-folded margins, apex without teeth. Stelidia less than ½ the length of column, triangular, with or without a small tooth along the upper margin close to the apex, with an inconspicuous, rounded wing along the lower margin. Anther abaxially furrowed, anterior surface inflated and deeply concave, anterior margin not drawn out; pollinia 2. Two species in montane forests up to 900 m in New Guinea. (JJV) Bulbophyllum sect. Trias (Lindl.) J.J.Verm, in prep. Type species (proposed): Bulbophyllum trias J.J.Verm. & G.Fischer Epiphytes with creeping rhizomes; bracts glabrous or the upper ones colliculate-papillose, shoots developing from nodes below the pseudobulbs, not fused to them. Roots spreading, glabrous to locally hirsute. Pseudobulbs distinct, 1-leaved. Leaves persistent, thick, veins inconspicuous. Inflorescences developing from nodes below the pseudobulbs, solitary or few-fasciculate, 1-flowered; peduncle bracts 2 or 3. Floral bract tubular; pedicel with the basal node level with the floral bract attachment or not, longer than the ovary. Flowers resupinate. Sepals free, equal, dorsal sepal 4.2–25.0 mm long, elliptic to ovate, obtuse to shortacuminate, glabrous, 3–7-veined, margins entire; lateral sepals as for the dorsal but elliptic to triangular, 5–9-veined. Petals 2–4(­–8) mm long, obovate to triangular (a narrow seam alongside the column in B. crassifolium Thwaites ex Trimen), rounded to acuminate, glabrous, 1–3-veined. Labellum mobile on a thin ligament, undivided or auriculate, 2.2–18.0 mm long, elliptic to triangular, adaxially without ridges (with two ridges in B. nanum De Wild., B. roseum Ridl.), surface glabrous to papillose, abaxial surface glabrous, margins entire, glabrous, auricles if present attached near the base of the labellum, antrorse, small (distinct in B. nummularia (H.Wendl. & Kränzl.) Rolfe). Column with rostellum anterior surface receding in between stelidia, stigma proximally protruding or not, column foot thickened or not toward its apex. Stelidia ½ the length of the column or shorter, deltoid to triangular, lower margin with or without an antrorse, triangular, rounded to acute tooth or deltoid, obtuse wing. Anther with a long-conical, cylindrical or wedge-shaped crest with a rounded to deeply bifid apex either adaxially (anterior margin not drawn out), or with the anterior margin drawn out into a similar structure with the anterior surface concave; pollinia 4, the inner ones more than half as long to as long as the outer ones. Approximately 15 species in forests up to 650 m in India, Burma, and Thailand. (JJV)

Bulbophyllum sect. Uncifera Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 697–738 (1913). Type species: Bulbophyllum ochroleucum Schltr. Epiphytes with creeping, straggling, or erect to stiffly pendent rhizomes; bracts minutely colliculate to papillose or transversely 40

rugate. Shoots developing from nodes below pseudobulbs, not fused to them. Roots spreading, glabrous to minutely hirsute locally. Pseudobulbs distinct, 1-leaved, Leaves persistent, thick, veins inconspicuous. Inflorescences developing from nodes below the pseudobulbs or from nodes scattered along the rhizome, 1- to many-fasciculate on short lateral sympodia, 1- to many-flowered, contracted to elongate racemes with flowers spirally arranged; peduncle bracts 2–8; rachis, if present, not thickened. Floral bracts tubular; pedicel with the basal node greater than the diameter of the pedicel above the floral bract attachment, as long as the ovary or (distinctly) longer. Flowers resupinate, opening simultaneously. Sepals free, equal, 2–21 mm long, elliptic to ovate, obtuse to acute (acuminate to caudate in B. cavibulbum J.J.Sm.), glabrous, 3-veined (5-veined in B. posticum J.J.Sm.), margins entire. Petals 1–6 mm long, elliptic to ovate, rounded to acute, glabrous, 1-veined (3-veined in B. posticum), margins entire, sometimes erose (distally), sometimes finely papillose distally. Labellum mobile on a thin ligament, undivided, 0.6–5.0 mm long, elliptic to ovate or oblong, sometimes subpanduriform, glabrous, adaxially without ridges, or with two ridges, often with two short, converging ridges, knobs or teeth close to the base (also with a weak dorsal ridge in B. exiguum F.Muell.), margins entire. Column with rostellum anterior surface receding in between the stelidia apices, stigma proximally not or minimally protruding, column foot tapering toward its apex. Stelidia ½ the length of the column or less but usually not vestigial, semi-elliptic, deltoid to triangular, with a distinct, patent and downward-directed, straight or recurved, semi-elliptic, apical tooth along the lower margin. Anther with a low, rounded, abaxial crest, anterior surface not concave, anterior margin drawn out into a small to distinct, semi-circular to triangular beak; pollinia 2 or 4, with the inner ones shorter to as long as the outer (longer in B. bigibbum Schltr.). Eight species in forests up to 2800 m in New Guinea, the western Pacific islands, and Australia (B. exiguum). (JJV) neotropical sections:

Artificial key to the Neotropical sections of Bulbophyllum (ES) 1. 2. 3. 4. 5.

Pseudobulbs bifoliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Bulbophyllaria Pseudobulbs unifoliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Inflorescence spiciform, rachis more than 4 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Furvescens Inflorescence racemiform, rachis less than 3 mm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Flowers spirally arranged . . . . . . . . . . B. sect. Micranthae Flowers distichously arranged . . . . . . . . . . . . . . . . . . . . . . 4 Petals patent; column foot with bilobed apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Didactyle Petals erect; column foot with entire apices . . . . . . . . . . . 5 Column foot shorter than column; lateral sepals united to form a synsepal . . . . . . . . . . . . . . . . . . . B. sect. Xiphizusa Column foot longer than column; lateral sepals free . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Napellii

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BULBOPHYLLUM

Bulbophyllum sect. Bulbophyllaria (Rchb.f.) Griseb., Brit. W. Ind., 613 (1864). Type species: Bulbophyllum bracteolatum Lindl. Epiphytes. Rhizome bracts membranous, disintegrating. Pseudobulbs 2-leaved. Leaves coriaceous. Inflorescence synanthous, a raceme with spirally arranged flowers; peduncle pendent. Flowers resupinate. Lateral sepals free. Petals erect. Anther without a protrusion overtopping the anterior margin. Three species in forests up to 400 m in Cuba, Colombia, Costa Rica, Dominican Republic, El Salvador, Guatemala, Honduras, Mexico, Nicaragua, Panama, and Venezuela. (ES) Bulbophyllum sect. Didactyle (Lindl.) Cogn., Fl. Bras. (Mart.), 3(5), 595 (1902). Type species: Bulbophyllum exaltatum Lindl. Epiphytes. Rhizome bracts membranous, disintegrating. Pseudobulbs 1-leaved. Leaves coriaceous. Inflorescence synanthous, a raceme with distichously arranged flowers; peduncle erect to apically drooping up to 45 degrees. Flowers resupinate. Lateral sepals free. Petals patent. Anther without a protrusion overtopping the anterior margin. Seven species in forests up to 1600 m in Bolivia, Brazil, Colombia, Ecuador, Guyana, Paraguay, Peru, and Venezuela. (ES) Bulbophyllum sect. Furvescens E.C.Smidt, Borba and van den Berg, in prep. Type species: Bulbophyllum nagelii L.O.Williams Epiphytes. Rhizome bracts membranous, disintegrating. Pseudobulbs 1-leaved. Leaves coriaceous. Inflorescence synanthous to proteranthous, a raceme with spirally arranged flowers; peduncle erect to apically drooping up to 20 degrees. Flowers resupinate. Lateral sepals free. Petals erect. Anther without a protrusion overtopping the anterior margin. Five species in forests up to 1900 m in Brazil, Ecuador, French Guiana, Guyana, Mexico, Peru, Suriname, and Venezuela. (ES) Bulbophyllum sect. Micranthae Barb.Rodr., Gen. Spec. Orch. Nov., 2, 117 (1882). Type species: Bulbophyllum micranthum Barb.Rodr. Epiphytes. Rhizome bracts membranous, disintegrating. Pseudobulbs 1-leaved. Leaves coriaceous. Inflorescence synanthous, a raceme with spirally arranged flowers; peduncle erect to apically drooping up to 45 degrees. Flowers resupinate. Lateral sepals free. Petals erect. Anther without a protrusion overtopping the anterior margin. Ten species in Bolivia and Brazil. (ES) Bulbophyllum sect. Napellii Rchb.f., Walp. Ann. Bot., 6, 249 (1861). Type species: Bulbophyllum napellii Lindl.

Epiphytes. Rhizome bracts membranous, disintegrating. Pseudobulbs 1-leaved. Leaves coriaceous. Inflorescence synanthous, a raceme with flowers distichously arranged; peduncle erect to apically drooping up to 135 degrees. Flowers resupinate. Lateral sepals free. Petals erect. Anther without a protrusion overtopping the anterior margin. Twelve species in forests up to 612 m in Argentina, Brazil, and Venezuela. (ES)

Bulbophyllum sect. Xiphizusa Rchb.f., Bot. Zeit., 10, 919 (1852). Type species: Bulbophyllum chloropterum Rchb.f. Epiphytes, occasionally lithophytes. Rhizome bracts membranous, disintegrating. Pseudobulbs 2-leaved. Leaves coriaceous. Inflorescence synanthous, a raceme with distichously arranged flowers; peduncle erect to apically drooping up to 145 degrees. Flowers resupinate. Lateral sepals united, forming a synsepal. Petals erect. Stelidia bidentate. Anther without a protrusion overtopping the anterior margin. Twenty-three species in forests and secondary vegetation up to 200 m in Bolivia, Brazil, Colombia, Jamaica, Mexico, Paraguay, Peru, and Venezuela. (ES)

Anatomy Chatin (1856) examined the structure and function of aerial roots in orchids, including B. careyanum (Hook.) Spreng., and later (Chatin 1857) broadly characterized the anatomy of rhizome, stem, and leaf of an assortment of epiphytic orchids including B. careyanum. Oudemans (1861) described the position of the epidermis in orchid aerial roots with brief mentions of Bulbophyllum species throughout. Leitgeb (1864) stated that the root in B. muscicola Rchb.f. (reported as Cirrhopetalum wallichii Lindl.) has a single-layered velamen. There is a detailed outline of aerial root anatomy in which Bulbophyllum species are fleetingly noted. Möbius (1887) described the anatomy of the leaf in several species of Bulbophyllum (including some as Cirrhopetalum). Meinecke (1894) recorded a single-layered velamen in Bulbophyllinae. Weltz (1897) reviewed the literature on anatomy of Bulbophyllinae followed by original detailed descriptions of the anatomy of eight species. Sprenger (1904) made a comprehensive study of the anatomical structure of Bulbophyllum including Cirrhopetalum, Bolbophyllaria, and Megaclinium. Tominski (1905) provided a general description of the leaf in B. thwaitesii Rchb.f. (as C. thwaitesii (Rchb.f.) Hook.f.). Moreau (1913) reported on organization and structure of vascular bundles in the pseudobulb of B. occultum Thouars and their relationship to its two leaves. Curtis (1917) reported on anatomy of epiphytic New Zealand species of Bulbophyllum, specifically the tiny B. pygmaeum and B. tuberculatum Colenso. Metzler (1924) described the 2-layered hypodermis with spiral water-storage cells in ground tissue of B. ­coriaceum Ridl. ex Stapf.

41

DENDROBIEAE

Fig. 621.8.  Distribution map of Bulbophyllum.

Oliver (1930) published notes on the leaf anatomy of B. tuberculatum. Solereder and Meyer (1930) listed several references to anatomy of Bulbophyllum (some species as Cirrhopetalum) in their comprehensive analysis of literature on orchid anatomy up to 1930. Duruz (1960) studied stomata of B. lobbii Lindl. treated as a xerophyte. Guttenberg (1968) discussed the anatomy of B. penduliscapum J.J.Sm. and B. lobbii. Sanford and Adanlawo (1973) stated that a 1-layered velamen is rare and appeared only in Bulbophyllum (some described as Genyorchis) species examined in their study of velamen and exodermis of West African epiphytic orchids. Singh and Singh (1974) described the stomatal apparatus in several Orchidaceae, including B. odoratissimum (Sm.) Lindl. ex Wall. N. Williams (1979) found a tetracytic stomatal apparatus on the lower leaf surface in Bulbophyllum. Olatunji and Nengin (1980) included 13 species of Bulbophyllum in their survey of tracheoidal idioblasts, which are elongate and sac-like and characterized by spiral or annular thickenings. Singh (1981) showed that stomata in B. viridiflorum (Hook.f.) Schltr. (as C. viridiflorum Hook.f.) may be either tetracytic and anisocytic but that those of B. umbellatum Lindl. (as C. maculosum Lindl.) and an unidentified species of Bulbophyllum are exclusively tetracytic. In mature leaves of B. viridiflorum anticlinal walls of subsidiary cells are dissolved, giving the impression of ‘floating’ stomata. Pridgeon et al. (1983) provided a systematic review of tilosomes among orchids, including Bulbophyllum (and some species of Cirrhopetalum). Kaushik (1983) investigated anatomy of leaf blade, petiole, pseudobulb, rhizome, and root of B. reptans (Lindl.) Lindl. ex Wall., B. penicillium C.S.P.Parish & Rchb.f., B. triste Rchb.f., B. griffithii (Lindl.) Rchb.f., B. khasyanum Griff., B. scabratum Rchb.f. (as C. caespitosum Wall. ex Lindl.), and 42

B. vaginatum (Lindl.) Rchb.f. (as Cirrhopetalum caudatum Wight). Contrary to other studies, Kaushik described the epidermis of the root as multiseriate, 1–3-layered in B. reptans, B. triste, B. griffithii, and B. khasyanum, 4- or 5-layered in B. penicillium, and 1- or 2-layered in B. vaginatum and B. scabratum Rchb.f. Other researchers noted the velamen of Bulbophyllum to be 1-­layered. Møller and Rasmussen (1984) indicated lack of stegmata throughout Bulbophyllum even though there is abundant sclerenchyma normally associated with stegmata. Singh (1986) discussed root anatomy in some Orchidaceae including B. odoratissimum, B. umbellatum, and B. viridiflorum, all of which have a 1-layered velamen. Mohana Rao and Khasim (1987) described the anatomy, ecology, and taxonomy of B. dyerianum (King & Pantl.) Seidenf. (=B. rolfei (King & Pantl.) Seidenf.), B. andersonii (Hook.f.) J.J.Sm., and B. leopardinum (Wall.) Lindl. ex Wall. Porembski and Barthlott (1988) noted a 1-layered velamen with plain cell walls, tilosomes in some species, and strongly thickened exodermal cell walls in Bulbophyllum. Mohana Rao et al. (1991) studied the anatomy of leaf, stem, and root of B. fischeri Seidenf. Prychid et al. (2004) found that stegmata are entirely absent from Bulbophyllum.

Leaf surface

Hairs with sunken bases present on both surfaces, more numerous below than above. Hairs glandular in B. dasypetalum Rolfe ex Ames, B. makoyanum (Rchb.f.) Ridl., and B. gracile Thouars. Cuticle much thicker in some species than in others; considerably so in B. molossus Rchb.f., papillate on the lower surface in B. tuberculatum. Epidermis one-layered, cells irregularly polygonal,

BULBOPHYLLUM

walls thickened on radial and inner tangential sides, oil droplets in cells of both epidermal surfaces. Stomata tetracytic, abaxial only, ‘floating’ in B. viridiflorum owing to the dissolution of anticlinal walls of subsidiary cells. transverse section

Epidermal cells smaller, flat-surfaced; in B. hirtum (Sm.) Lindl. ex Wall., upper epidermal cells larger, reaching a sixth of the leaf thickness and serving in water-storage. Hypodermal cells generally polyhedral, spirally thickened in B. thouarsii, mostly 1-layered ab- and adaxially with thin walls; adaxially only in B. penicillium C.S.P.Parish & Rchb.f., B. griffithii and B. reptans, abaxially only in B. khasayanum. Mesophyll heterogeneous in B. retusiusculum Rchb.f., B. clandestinum Lindl., B. saltatorium Lindl. var. calamarium (Lindl.) J.J.Verm., B. makoyanum, B. vaginatum (Lindl.) Rchb.f., and other species; homogeneous in B. leopardinum and other species. Water-storage cells with spiral thickenings occur among mesophyll cells of B. gracile Thouars, B. retusiusculum, B. reptans, and additional species in which they are elongate, sac-like, and parallel to the midvein, and in some species isodiametric; some larger hypodermal cells also store water as in B. khasayanum. Mucilage cells, raphides, and fibres occur in the hypodermis of B. virescens J.J.Sm., B. gibbosum (Blume) Lindl., and B. multiflorum Ridl. Small druses occur in mesophyll cells of B. acutiflorum A.Rich. Vascular bundles usually uniseriate. Bundles surrounded or capped by a fibrous sheath containing cells in which the lumen is almost absent; fibres more numerous on phloem side than on xylem side. A sclerenchyma bridge of thick-walled cells occurs in B. virescens, B. ambrosia (Hance) Schltr., and other species. Stegmata absent.

Pseudobulb transverse section

Cuticle thin in B. leopardinum and other species but thick in some species. Epidermal cells periclinal, squarish, with weakly vaulted outer walls in B. ambrosia. Cells beneath the epidermis 3- or 4layered, colourless, parenchymatous, sometimes collenchymatously thickened and mucilaginous. Where there are spiral cells in the leaf, there are spiral cells in pseudobulb parenchyma. Cortex and ground tissue cells parenchymatous, polygonal; in B. penicillium, B. reptans, and B. triste, some larger cells and water-storage cells of cortex and ground tissue with spirals; some cortical cells mucilaginous. Vascular bundles typically scattered in the cortex and ground tissue, but in some species there is a central ring of small bundles. In B. gibbosum outside the sclerenchymatous ring there is a parenchymatous cortex of polyhedral cells with spirals and large parenchyma cells with spirals inside the ring. Larger vascular bundles may be surrounded by a complete sclerenchymatous sheath; in smaller bundles there are a few sclerenchyma cells at the xylem pole, whereas sclerenchyma at the phloem pole is more pronounced.

Rhizome transverse section

Cuticle generally thick, especially in B. clavatum, B. occultum, B. acutiflorum, and B. lobbii. Epidermal cells are elongated in the direction of the axis; radial and tangential walls even, inner and outer walls weakly vaulted; radial and tangential walls thick in

B. careyanum. Cortex consists of five cell layers; cells are thickwalled and narrow-lumened and form a compact central cylinder. Spiral cells occur only in the ground tissue of B. lobbii and B. gibbosum. All Bulbophyllum species examined, with some exceptions, have a 1-layered sclerenchyma cylinder at the border of the cortex and ground tissue, e.g. B. makoyanum, B. gibbosum, and B. clandestinum. Two layers of sclerenchyma are present in B. fischeri, four in B. acutiflorum, and six in B. virescens. Vascular bundles are scattered in the central parenchyma cylinder, the peripheral ones adjacent to the inner border of the sclerenchyma ring. All forms have a 1-layered xylem sheath, and most have 2- or 3-­layered sclerenchyma forming a complete phloem sheath. A parenchyma bridge occurs in vascular bundles of B. acutiflorum and B. clavatum.

Root transverse section

Velamen 1-layered, cells isodiametric lacking ornamentation. Tilosomes broadly lamellate in most Bulbophyllum species; spongy in B. micholitzianum Kränzl. and B. ornatissimum (Rchb.f.) J.J.Sm. Exodermal walls ○-thickened in most species, ∩-thickened in B. medusae (Lindl.) Rchb.f., and ∩- and ○-thickened in B. imbricatum Lindl. Cortex 4- or 5-layered with spirally thickened columnar or sac-like water-storage cells in B. penicillium, B. reptans, and B. triste. Endodermal cells barrel-shaped; cell walls ○-thickened. Pericycle cells sclerenchymatous with thin-walled passage cells opposite xylem sectors. Vascular cylinder polyarch; xylem and phloem components alternating: B. odoratissimum 20–66-arch, B. membranifolium Hook.f. 7–13-arch, B. viridiflorum 6–12-arch, B. griffithii 10-arch, B. reptans 9- and 10-arch, B. triste 6-arch. Xylem and phloem elements embedded in thick-walled cells. Root hairs unicellular. Pith sclerotic. (WS)

Palynology Schill and Pfeiffer (1977) surveyed pollen of nine species of Bulbophyllum, including six then considered as Cirrhopetalum. Tetrads are concave to convex depending on species with generally laevigate sculpturing but also microfoveolate to weakly verrucose. The sexine is rounded (but also angular in B. schinzianum Kränzl. and calymmate in B. maximum (Lindl.) Rchb.f.) and 0.5–2.5 μm thick. (AP)

Cytogenetics The most common diploid numbers found in Bulbophyllum are based on x = 19 (2x = 38; Lim and Jones 1982; Lim 1985) or x = 20 (2x = 40; a few exceptions with 2x = 57 or 95). (BG)

Phytochemistry A review of the chemistry and pharmacology of Bulbophyllum with literature references from 1985–2004 lists more than 60 constituents, including bibenzyls, flavonoids, phenanthrenes, phenylpropanoids, and volatiles (Chen et al. 2005). This report was compiled in order to assess the use of Bulbophyllum as a substitute for Dendrobium in traditional Chinese medicine. The local 43

DENDROBIEAE

Fig. 621.9.  Typical stilbenoid constituents of Bulbophyllum.

names, distribution, habitat, and properties of 15 species of Bulbophyllum used medicinally in China have been documented (Yi et al. 2005). One-third of these have been investigated phytochemically, notably B. kwangtungense Schltr. and B. odoratissimum, but also B. andersonii, B. inconspicuum Maxim., and B. reptans. A predominant feature of Bulbophyllum phytochemistry is the presence of stilbenoids, which have been recorded from most species investigated. These typically comprise phenanthrenes, dihydrophenanthrenes (including dimeric forms), and bibenzyls (Fig. 621.9). Although the same classes of compound are well represented in Dendrobium, many specific examples are unique to Bulbophyllum, including five stilbenoids isolated from methanol extracts of the whole plant of B. andersonii (cited as Cirrhopetalum andersonii Hook.f.), comprising the phenanthrene cirrhopetalin (Majumder and Basak 1990), the dihydrophenanthrene cirrhopetalanthridin (Majumder and Basak 1991b), and the bibenzyls cirrhopetalidin, cirrhopetalinin, and cirrhopetalidinin (Majumder and Basak 1991a,b). The known compounds, batatasin-III (3,3′-dihydroxy-5-methoxybibenzyl) and gigantol (4,3′-dihydroxy-3,5′-dimethoxybibenzyl) were also obtained (Majumder and Basak 1991a,b). Batatasin-III occurs in B. ­fuscopurpureum Wight and B. guttulatum (Hook.f.) N.P.Balakr., which also contain the 9,10-dihydrophenanthropyrans flavidin and imbricatin, and flaccidin, respectively (Veerraju et al. 1989a). The structure of gymnopusin, a constituent of B. gymnopus Hook.f. (Majumder and Banerjee 1988), was revised to that of 2,7-dihydroxy-3,4,9-trimethoxyphenanthrene (Fig. 621.9) on the basis of chemical synthesis and extensive spectroscopic analysis (Hughes and Sargent 1989; Majumder and Banerjee 1989). In work on B. inconspicuum Maxim., Tan et al. (2006a) described 44

2-methoxy-3,4,7-trihydroxyphenanthrene, a novel compound, together with 2,5-dihydroxy-4-methoxyphenanthrene and its 9,10-dihydro analogue, and the benzopyran-4-one derivative eugenitin. Extracts of leaves and stems of B. kwangtungense Schltr. afforded the bibenzyls cumulatin and densiflorol and the phenanthrene plicatol B (Wu et al. 2006). Subsequently, the unique example, 2′,3′-dimethoxy-2-methyl-4,5-methylenedioxybibenzyl, was reported from the same source (Wu et al. 2008). This species also contains a number of rare dihydrodibenzoxepin derivatives (Wu et al. 2006, 2008), including one characterized as a stress metabolite produced by leaves and stems in response to abiotic elicitation with copper (II) chloride (Chen et al. 2011). Interest in the structures of these compounds prompted the total synthesis of one example, bulbophylol-B (Fig. 621.9), which also occurs in B. odoratissimum (Lin et al. 2008). A novel phenanthrene derivative, bulbophyllanthrin, has been found in methanol extracts of the whole plant of B. leopardinum (Majumder et al. 1985). The chemical constituents of B. odoratissimum have been investigated in greater detail than those of any other species in the genus. Among the stilbenoids reported are the phenanthraquinone bulbophyllanthrone (Fig. 621.9), which was isolated from methanol extracts of the whole plant (Majumder and Sen 1991). This compound also occurs in the New Zealand endemic, Earina autumnalis (G.Forst.) Hook.f., and was found to have cytotoxic and antibacterial activity (Hinkley and Lorimer 1999). Seven methylenedioxy-substituted bibenzyl derivatives from B. odoratissimum have been synthesized because of interest in their cytotoxicity against human cancer cell lines (Lin et al. 2006; Zhang et al. 2007a,b). These include cirrhopetalin (Fig. 621.9), which was first obtained from B. andersonii (Majumder and Basak 1990).

BULBOPHYLLUM

A large-scale extraction (20 kg) of the whole plant of B. odoratissimum yielded 3,7-dihydroxy-2,4,6-trimethoxyphenanthrene, which showed cytotoxicity against five human cancer cell lines (Chen et al. 2007). Similarly, densiflorol B (7-hydroxy-2-­methoxy1,4-phenanthraquinone) was the most cytotoxic of seven known stilbenoids isolated from the same source (Chen et al. 2008d). Two novel dihydrophenanthrene dimers, bulbophythrins A and B, were also cytotoxic against the same panel of cell lines (Xu et al. 2009). In the most recent study, six further examples of known stilbenoids were isolated from B. odoratissimum (Chen et al. 2010). Bulbophyllin and bulbophyllidin are bibenzyl derivatives that occur only in B. protractum Hook.f. (Majumder et al. 1997). In addition, this species contains several known bibenzyls and the 9,10-dihydrophenanthropyran flavidin. Methanol extracts of the whole plant of B. reptans yielded the phenanthrene dimers reptanthrin (Fig. 621.9) and isoreptanthrin, which appear to be formed by the oxidative coupling of gymnopusin, one of nine known stilbenoids also found in this species (Majumder et al. 1999b). Similarly, rigidanthrin, a dimeric phenanthrene derivative from B. rigidum King & Pantl., is probably formed by oxidative coupling of nudol (2,7-dihydroxy-3,4-dimethoxyphenanthrene) and gymnopusin (Majumder et al. 2008a). Tristin (3-methoxy-4,3′,5′trihydroxybibenzyl), which was first isolated from B. triste Rchb.f. together with dihydroresveratrol (3,5,4′-trihydroxybibenzyl) and 3,4′-dihydroxy-5-methoxybibenzyl (Majumder and Pal 1993), also occurs in a number of Dendrobium species (Table 622.3, 91). The phenanthrene dimer characterized from B. umbellatum Lindl. (cited as Cirrhopetalum maculosum Lindl.) as cirrhopetalanthrin (Majumder et al. 1990), is identical in structure to blestriarene C from Bletilla striata (Thunb.) Rchb.f. (Yamaki et al. 1989). Harrison and coworkers have explored the rich stilbenoid chemistry of B. vaginatum (Lindl.) Rchb.f. in a series of publications (Leong et al. 1997, 1999; Leong and Harrison 2004). In addition to many known examples, several phenanthrenes and dihydrophenanthrenes unique to this species were described (Leong et al. 1997, 1999). Two minor components of hexane extracts of the whole plant were characterized as a phenanthrene dimer and an uncommon phenanthro(4,3-b)furan derivative. The latter is thought to be the result of radical coupling between sinapyl alcohol and 1,4,5-trihydroxy-7-methoxy-9,10-dihydrophenanthrene (Leong and Harrison 2004). The flavonoid chemistry of Bulbophyllum is poorly known. C. Williams (1979) detected kaempferol (3,5,7,4′-tetrahydroxyflavone) in acid-hydrolysed leaf extracts of B. cochleatum Lindl. var. brachyanthum (Summerh.) J.J.Verm. (cited as B. bequaertii De Wild. var. brachyanthum Summerh.) and flavone C-glycosides in the leaves of B. gibbosum (Blume) Lindl. Chrysin (5,7-­dihydroxyflavone) and pinobanksin (3,5,7-­trihydroxyflavanone) occur in methanol extracts of the whole plant of B. odoratissimum (Majumder and Sen 1991). Information on other phenolic and aromatic compounds is limited. The lignans (+)-lyoniresinol 9′-O-βglucopyranoside, (+)-pinoresinol, and (±)-syringaresinol have been found in B. odoratissimum (Liu et al. 2005), B. triste (Majumder et al. 1994), and B. vaginatum (Leong et al. 1999), respectively. Bate-Smith (1968) detected para-coumaric, ferulic, and sinapic acids in acid-hydrolyzed extracts of leaves of B. gibbosum.

Aqueous ethanol extracts of the whole plant of B. odoratissimum yielded the novel phenolic glycoside bulbophyllinoside, together with three known examples (Liu et al. 2005). Other phenolic compounds reported from this species include ethyl orsellinate, 4-hydroxy-3-methoxycinnamaldehyde, syringaldehyde, 3-(4-hydroxyphenyl)propanoic acid and its methyl ester, and vanillic acid (Liu et al. 2005; Chen et al. 2008d). Dihydroconiferyl alcohol and the ethyl ester of ferulic acid were obtained from extracts of whole plants of B. protractum Hook.f. (Majumder et al. 1997) and B. rigidum (Majumder et al. 2008a), respectively. Presence of dihydroferulic, p-coumaric, and 3,4-dihydroxybenzoic acids in extracts of B. vaginatum has been noted (Leong et al. 1999). Information on alkaloids of Bulbophyllum is limited to reports of their presence or absence determined during wider investigations of Orchidaceae. In an early paper, Boorsma (1902) mentioned that traces of alkaloids were found in B. helenae (Kuntze) J.J.Sm. (cited as Cirrhopetalum cornutum Lindl.). Other records are found in the surveys of Lüning (1964, 1967) and Lawler and Slaytor (1969, 1970), including two under Cirrhopetalum and Ione. According to a subsequent review (Lüning 1974), of 134 species of Bulbophyllum tested, only nine had an alkaloid content ≥ 0.1%. For a small number of species listed under Acrochaene (one), Cirrhopetalum (four), and Ione (four), none had an alkaloid content ≥ 0.1%. Extracts of whole plants of B. elatum (Hook.f.) J.J.Sm. (cited as Cirrhopetalum elatum Hook.f.) contain the (E)- and (Z)- isomers of the p-coumaroyl ester of 24-methylenecycloartan-3β-ol, a triterpenoid of the cycloartane series (Majumder and Pal 1985; Majumder et al. 1987). The (E)-isomer, known by the trivial name pholidotin, also occurs in Pholidota rubra Lindl. (Majumder et al. 1987). Reports of common triterpenoids include sitosterol and its 3-O-β-glucoside from B. odoratissimum (Chen et al. 2010), sitosterol and stigmasterol from B. protractum (Majumder et al. 1997), and friedelin from B. vaginatum (Leong et al. 1997). Floral fragrances of B. fascinator (Rolfe) Rolfe (as ‘Cirrhopetalum fascinor’ Rolfe), B. graveolens (F.M.Bailey) J.J.Sm. (as Cirrhopetalum robustum Rolfe), and B. lobbii were analysed using gas chromatography coupled to mass spectrometry (GC-MS) by Kaiser (1993). According to Kaiser (1993), there are two aspects to the fragrance of B. fascinator; butyric acid contributes to the carrion-like scent from the labellum, and caryophyllene epoxide to a pleasant woody scent associated with the sepals. Occurrence of heterogenous scent centres in Bulbophyllum was recorded previously by Vogel (1962) for B. ornatissimum (as Cirrhopetalum ornatissimum Rchb.f.). The GC-MS profile of the B. fascinator fragrance indicates that caryophyllene (55%) is by far the major component, followed by limonone (9%), acetic acid (5%), and caryophyllene oxide (5%) at lower levels, and other more minor components, including butyric acid (0.7%). Kaiser (1993) has described the stench produced by B. graveolens as ‘one of the most penetrating produced by the orchid family’, citing a combination of acetic, butyric, 2-methylbutyric, and valeric acids as the main olfactory components. An unexpected feature of this scent chemistry is the presence of the N-methyl and N,N-dimethyl derivatives of acetamide and formamide. In contrast to B. graveolens, the floral fragrance of B. lobbii is said to be pleasant, comprising mainly 45

DENDROBIEAE

linalool and (E)-ocimene, and their derivatives, which are among 28 components identified using GC-MS. Technical difficulties due to flower fragility allowed only preliminary data on the scent of B. gracillimum (Rolfe) Rolfe (cited as ‘Cirrhopetalum gracillium’ Rolfe) to be obtained (Kaiser 1993). Components identified comprise acetic acid, geranylacetone, (Z)-3-hexenol, (Z)-3-hexenyl tiglate, indole, isoleucine methyl ester, and nonanal. A solid sample injector device allowing direct analysis of flower material was developed in order to overcome problems experienced during investigation of some Bulbophyllum volatiles (Da Silva et al. 1999). Using this methodology, the chemical scent compositions of B. involutum Borba, Semir & F.Barros, B. ipanemense Hoehne, and B. weddellii (Lindl.) Rchb.f., three Brazilian species pollinated by female flies of the genus Pholeomyia, were obtained. Profiles of volatiles present in B. involutum and B. weddelii, which have the same pollinators, are similar but distinct from that of B. ipanamense, which is pollinated by a different species of Pholeomyia (Da Silva et al. 1999). Scent compositions of Bulbophyllum species that attract male fruit flies of the genus Bactrocera were examined in a series of papers (Tan and Nishida 2000, 2005, 2007; Tan et al. 2002, 2006b; Nishida et al. 2004). Principal volatile attractants are zingerone for B. patens King ex Hook.f. (Tan and Nishida 2000) and B. baileyi F.Muell. (Tan and Nishida 2007), methyl eugenol for B. cheiri Lindl. (Tan et al. 2002) and B. vinaceum Ames & C.Schweinf. (Tan et al. 2006b), and raspberry ketone for B. apertum Schltr. (Tan and Nishida 2005). Volatile phenylpropanoids with structures similar to that of methyl eugenol also occur as minor components in the scent of B. cheiri (Nishida et al. 2004). (NV, RG)

Phylogenetics Until recently, subtribe Bulbophyllinae was believed to include a varying number of genera, depending on the author. Strongly contrasting views were expressed by, for instance, Lindley (1824), Bentham and Hooker (1883), Pfitzer (1889), Schlechter (1913), Seidenfaden (1979), Garay et al. (1994), Vermeulen (1987), Szlachetko and Margonska (2002), Clements and Jones (2002), and Vermeulen & O’Byrne (2011). Based on combined nuclear and plastid DNA data and morphological analyses, we argue that all genera proposed in that subtribe should be included in Bulbophyllum (Gravendeel et al., in preparation). Diagnostic sets ascribed to those genera consist of characters that: 1) are highly polythetic (e.g. rachis morphology in the former genus Megaclinium is not present in all species of this group); 2) are not restricted to one genus but occur as well in morphologically similar species, related or not (e.g. crest on the anther in Trias that also occurs in B. sect. Leopardinae); 3) include various nonhomologous states (e.g. the crest on the anther in Trias, which is formed either by a protuberance on the abaxial side of the anther or by its anterior margin); and 4) are better regarded as apomorphies at the species level because they distinguish species among what we assume are its closest relatives (e.g. species of Ichthyostomum, which is regarded here as part of B. sect. Minutissima). 46

We also found new or resurrected genera for which no precise diagnostic sets were given; the authors apparently relied on loose or outdated circumscriptions provided by earlier authors. In our opinion, the formerly recognized genera cause an imbalance in the taxonomic structure of Bulbophyllinae. The genera are generally set off morphologically against ‘the rest of Bulbophyllum’, which is in phylogenetic terms a recipe for extracting small clades as separate genera and leaving large numbers of species in Bulbophyllum as paraphyletic groups. When diagnostic characters of these segregate genera are plotted on all molecular results obtained so far based on combined plastid (matK, trnL intron, trnL-trnF spacer, trnF-ndhJ spacer psbA-trnH spacer, trnE-trnD spacer, trnS-trnG spacer) and nuclear (nrITS) DNA sequences (Fischer et al. 2007a; Smidt et al. 2011; Hosseini et al. 2012; Gravendeel et al., in preparation), they prove to be highly homoplasious. In all phylogenetic analyses, Bulbophyllum s.l. as delimited above appears to be monophyletic. The phylogenetic tree consists of main clades that are all geographically restricted (Fig. 621.10). Each comprises species of continental Africa, America or Madagascar. The terminal main clade consists of Asiatic and Australasian species. It consists of two subclades. One contains predominantly western Malesian sections, the other predominantly eastern Malesian sections, but the division is not as clear as in the more basal nodes of the cladogram. Plotting morphological characters on the phylogenetic tree yields less clear results. In fact, it is only successful in the terminal branches, optimizing small diagnostic character sets to small clades. These typically include 20–50 species, but the species count ranges 1–165 species. At best, these diagnostic sets are monothetic. In many clades, however, suites of species are included that show only part of the diagnostic set of the section, rendering the latter polythetic. For major basal clades, we cannot identify any macromorphological diagnostic characters, in spite of the fact that the connoisseur can often recognize individual plants by their general appearance as belonging to one of those clades, even without flowers. However, we feel that the strong geographical signal lends the cladogram relevance as a phylogenetic hypothesis for Bulbophyllum, which also supports our conclusions here. Additionally, with representatives of almost all sections included, we claim that its coverage is sufficient to give it broad significance as the basis for our treatment of sections here. The phylogenetic tree offers different options for distinguishing taxonomic units at whatever rank within Bulbophyllum. Most meaningful from a phylogenetic point of view would be to recognize the geographically restricted clades attached to basal nodes as taxonomic units. Lack of macromorphological traits makes this impossible. Although individual groups within these clades may sometimes display unique characters or character sets, all include series of species without these outstanding features and, conversely, these are morphologically similar to species in other clades. As stated above, only clades in the terminal branches have diagnostic sets, monothetic or, more frequently, polythetic. Considering that taxonomic units should be monophyletic and have a diagnostic set of characters, we state the principles that underlie taxonomic hypothesis employed here. First of all,

BULBOPHYLLUM

we distinguish a single genus, Bulbophyllum, because neither the major clades near the base of the cladogram nor the smaller clades rooting towards the terminal branches are reflected in the distribution of morphological characters. Second, we distinguish supraspecific taxonomical units within Bulbophyllum based on clades rooting in the terminal branches. Third, we refrain from creating a hierarchical structure of supraspecific taxonomic units within Bulbophyllum for reasons already stated. Fourth, given that some of these units display a general morphological similarity, others appear to grade into each other, many units have a polythetic character set, and few sections, if any at all, are characterized by characters that do not occur elsewhere within the genus, we have chosen a taxonomic level not far above that of species, that of sections. In a few cases, the topology of the phylogenetic tree compels us to depart from the principles outlined above. First, a single clade has a character distribution that makes the distinction of a paraphyletic grade a taxonomic convenience: B. sect. Polymeres. This is a largely eastern Malesian (New Guinean) clade with a single, almost monothetic diagnostic character (a widened column foot) from which numerous groups have evolved, each with a larger diagnostic set, while in most cases losing the diagnostic character (the widened column foot) of their common ancestor. Choosing in this particular case for a hierarchical arrangement of the taxonomic units (e.g. a subgenus with sections) would have led to the necessary distinction of subgenera throughout Bulbophyllum. Lack of characters would have made this a hazardous enterprise. We accept the consequence that the distinction of a paraphyletic grade has led to some arbitrary decisions about the delimitation of the sections rooted in it. Taxonomic stability would not be served with raising all our sections to the generic level, i.e. splitting up Bulbophyllum into numerous small genera. The nature of the diagnostic sets, as explained above, does not allow this in our view. Any decision to raise some sections to generic level while retaining others at section level would be highly arbitrary. Characters supporting the phylogenetic tree are mostly those that earlier supported taxonomic divisions in Bulbophyllinae: rhizome morphology (either erect with roots growing alongside the rhizome towards the substrate or creeping with roots spreading); presence/absence of pseudobulbs; number and persistence of leaves of the pseudobulbs; inflorescence morphology (1-flowered or racemose); flower morphology including presence of 1- to few-celled excrescences on the flower parts (papillae, ciliae) or many-celled excrescences (fimbriae, teeth, but also appendages of more complicated morphology as on petals in B. sects. Epicrianthes and Hyalosema or labellum in B. sect. Kinethrix). The importance of distinguishing suites of other characters came to light recently. One in particular enabled completion of a key to the numerous Asiatic sections of the genus: node displacement at the base of the pedicel (node at the base of the pedicel level with the floral bract attachment, or well above it). Node displacement also occurs at rhizome nodes, where axillary buds may sprout so high above the attachment of the subtending bract that they are almost anti-axillary to the bract of the next node (in a few species of B. sect. Monanthes and in several of B. sect. Epibulbon). This character, in a less extreme state, helped

greatly to delimitate B. sect. Macrocaulia. In a few cases other previously unappreciated characters were found that helped formulate a diagnostic set for single sections (e.g. petals basally fused to the column in B. sect. Brachypus). (BG, JJV, TS, RV)

Ecology Members of Bulbophyllum occur mainly in tropical or subtropical primary forests, ranging from always wet rain forests and montane forests to dry deciduous forests, savanna woodlands, or rocky fields with shrubby vegetation and only a few wet months each year. Many species of B. sect. Beccariana are adapted to the undergrowth of tropical rain forests. Bulbophyllum uniflorum (Blume) Hassk., for instance, is a slender plant with cylindrical pseudobulbs and thin leaves. On the other hand, B. blepharistes Rchb.f. (B. sect. Blepharistes) is adapted to dry, exposed environments. It has thick, leathery pseudobulbs and leaves. Many Madagascan and South American species occur in similar, harsh environments. Generally, species grow as epiphytes and occur on the bases of tree trunks to the tiniest twigs. Some grow in moss pads, whereas others prefer bare bark. A few species are humuscollectors, either in cupped leaves (e.g. B. beccarii Rchb.f.) or in clusters of erect secondary roots (e.g. B. penduliscapum J.J.Sm.). Many species may also be found as lithophytes or terrestrials; a small number of species are predominantly lithophytic or terrestrial. Some species may persist after disturbance of the forest as long as the environmental changes are not too profound. A few species are more frequently found in secondary woodlands, plantations, or even ornamental trees; these include B. medusae (Lindl.) Rchb.f. and B. vaginatum (Lindl.) Rchb.f. Less common environments that may harbour Bulbophyllum species are mangrove forests, open and exposed vegetation on limestone or ultrabasic bedrock, and high-elevation scrub. (BG, GF, JJV)

Pollination In many species of Bulbophyllum, the labellum is moved by the wind, and this movement is important for attraction as it appeals to the aggregation instinct of Diptera, the main functional group of pollinators reported for the genus (Borba and Semir 1998; Tan and Nishida 2000; Tan et al. 2002; Teck and Hong 2011, 2012). In addition to visual attraction, a movable labellum in Bulbophyllum plays an essential additional role in the pollination mechanism. Ridley (1890a) described how flies land on the hinged labellum, causing it to move downward. When the flies crawl up the labellum and pass the balance point, it returns to its original position, pressing the flies against the column. This mechanism is widespread in the genus with some variation in the basic patterns (van der Pijl and Dodson 1966; Braga 1977; Christensen 1992; Verola 2002). Another pollination mechanism, in which wind is necessary to move the labellum and then presses the insect against the column, was observed by Sazima (1978) and Borba and Semir (1998) for species from B. section Didactyle found in open areas in southeastern Brazil. Borba and Semir (1998) hypothesized that because pollination is dependent on an unpredictable external factor (winds range from 1.0 to 1.5 m/s), nectar is also necessary to keep the insect in the flower for a long 47

DENDROBIEAE

period. In B. section Didactyle, Borba and Semir (1999) described another mechanism to prevent self-pollination, consisting of a temporal variation in pollinarium size after its removal; a freshly removed pollinarium has a diameter twice that of the entrance of the stigmatic cavity and therefore requires approximately two hours to shrink enough to allow pollination.

There are many unrelated families of Diptera reported as pollinators of Bulbophyllum species: Milichiidae, Chloropidae, Sciridae, and Tachinidae in the Neotropics and Tephritidae in Indomalaysia. In the last family, male flies belonging to the genus Bactrocera are attracted to the flowers by zingerone, a phenanthrene that resembles fruit-fly attractants. Bactrocera males acquire

Fig. 621.10.  Schematic summary of molecular phylogeny of Bulbophyllinae based on nrITS sequences (BG, ES, GF, JJV).

48

BULBOPHYLLUM

Fig. 621.10.  Continued.

49

DENDROBIEAE

this floral essence and store it in a gland or convert zingerone to zingerol. The latter compound is stored in the rectal gland and subsequently released to attract conspecific females. This is a rare example of a mutualistic interaction between insects and plants via a floral synomone, in which both organisms obtain direct benefit to their reproductive efforts (Tan and Nishida 2000). Fischer (unpublished) reports pollination of Bulbophyllum ambrosia Schltr. by Apis cerana, indicating that shifts in pollination systems known from other orchid genera also occur in Bulbophyllum. (ES, GF)

Uses Several species of Bulbophyllum have artistic and medicinal uses in Africa and Asia (Lawler 1984). In Papua New Guinea, several species are used to provide a red color for plaited leg-, arm- and waistbands. In Nepal, B. careyanum and B. leopardinum are used to initiate abortions (Subedi 2011). In China, several species are used for treatment of pulmonary tuberculosis and stomach cancer. Many artificial hybrids have been created for horticultural purposes, often with species of B. sects. Beccariana, Cirrhopetaloides, Lepidorhiza, and Sestochilus as parents. These include, for example, Bulbophyllum Fascination, B. Elizabeth Ann, and B. Warren’s Wizardry, for which either B. lobbii or B. longissimum were used as one of the parents to create hybrids with large flowers with a pleasant fragrance. (BG, JJV)

Cultivation Species of the genus are easily grown in pots or attached to slabs of cork or bark. Preferred temperatures depend on the elevational range of the species in nature, but moderate conditions in cultivation will suit many (with minimum temperature of 15 °C at night). Appropriate relative humidity depends, again, on conditions under which a species grows in the wild. Generally, high humidity is important, even for species adapted to environments characterized by long, dry periods and high light intensity, such as B. blepharistes Rchb.f. from Asia, and many Madagascan and American species. High humidity should always accompany abundant air movement to prevent rot. If a dry season is needed, the plants are given less water. A short resting period with cooler temperatures and drier growing conditions is required by many species, but many small species should be kept perpetually damp. Shrivelling of pseudobulbs is always to be avoided. Larger species, e.g. those of B. section Beccariana, often need to produce a certain number of pseudobulbs before they flower. Artificial light in the winter season enhances growth significantly. Species with patent and hanging rhizomes and with roots that grow alongside the rhizome towards the substrate should be rejuvenated by removing the ends of the rhizomes and establishing them as new plants. Plants of species with patent rhizomes have a limited lifespan, depending on how long the plant clings to the substrate with its roots once the basal part of the rhizome has collapsed with age. Spontaneous rejuvenation by producing new shoots at the base of the plant may occur, particularly after disturbance. (GF, JJV, TS, RV)

50

Taxonomic literature Averyanov, L. V. (1994). Identification guide to Vietnamese orchids. Mir I. Semya Publishing House, St. Petersburg, Russia. Averyanov, L. V. and Averyanov, A. L. (2006). New orchids from Vietnam. Komarovia, 4, 19. Bentham, G. and Hooker, J. D. (1883). Bulbophyllinae. Genera plantarum, 3, 1–1258. L. Reeve, London. Blume, C. L. von (1825). Javaansche Orchideen. Bijdragen tot de Flora van Nederlandsch Indië, 1, 306–20. Breda, J. G. S. von (1827). Genera et species orchidacearum et asclepiadearum, t. 3–14. Gandavi, Ghent, Belgium. Buchanan-Hamilton, F. and Lindley, J. (1816). Sunipia. Rees’s Universal Dictionary of the Arts and Sciences, 11, 13. Longman, Hurst, Rees & Orms, London. Carr, C. E. (1930). Some Malayan orchids 2. Gardens’ Bulletin Straits Settlements, 5, 124–60. Carr, C. E. (1933). Some Malayan orchids 4. Journal of the Straits Branch of the Royal Asiatic Society, 11, 66–109. Clements, M. A. and Jones, D. L. (2002). Nomenclatural changes in the Australian and New Zealand Bulbophyllinae and Eriinae (Orchidaceae). Orchadian, 13, 485–500. De Witte, L. C. and Vermeulen, J. J. (2010). The Bulbophyllum species attributed to section Hymenobractea (Orchidaceae). Blumea, 55, 80–5. Finet, A. (1909). Pelma, orchidacearum genus novum. Notulae Systematicae, 1, 112–14. Fischer, G. A., Sieder, A., Cribb, P. J., and Kiehn, M. (2007b). Two new species and a new section of Bulbophyllum (Orchidaceae) from Madagascar. Adansonia, 29, 19–25. Fischer, G. A., Sieder, A., Hermans, J., Andriantiana, J., Kiehn, M., and Cribb, P. J. (2009). Description of four new species of Bulbophyllum (Orchidaceae) from Madagascar. Novon, 19, 344–52. Fitzgerald, R. D. (1891). Two new Australian orchids. Journal of Botany, 29, 152. Garay, L. A. and Kittredge, W. (1985). Notes from the Ames Orchid Herbarium. Botanical Museum Leaflets, Harvard University, 30, 47–59. Garay, L. A., Hamer, F., and Siegerist, E. S. (1994). The genus Cirrhopetalum and the genera of the Bulbophyllum alliance. Nordic Journal of Botany, 14, 609–46. Gaudichaud-Beaupré, C. (1826). Voyage in L’Uranie and La Physicienne, Botany, 27. Pillet, Paris. Guillaumin, A. (1929). Contribution à la flore des NouvellesHébrides 3. Supplément aux plantes recueillies par M. Levat. Bulletin de la Société Botanique de France s. 5, 5, 298–303. Guillaumin, A. (1964). Plantes nouvelles, rares ou critiques des serres du muséum. Bulletin du Muséum National d’Histoire Naturelle, 2(35), 653. Hooker, J. D. (1825). Anisopetalum. Exotic Flora, 2, t. 103, 149. Hooker, J. D. (1872). Bolbophyllum lemniscatum. Curtis’s Botanical Magazine s. 3, 28, 59–61. Hooker, J. D. (1890). Henosis. The flora of British India, 5, 771. L. Reeve, London. Hosseini, S., Go, R., Dadkhah, K., and Nuruddin, A. A. (2012). Studies on maturaseK sequences and systematic classification of Bulbophyllum in Peninsular Malaysia. Pakistan Journal of Botany, 44, 2047–54. Jones, D. L. and Clements, M. A. (2001). Oncophyllum, a new genus of Orchidaceae from Australia. Orchadian, 13, 421.

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Jones, D. L. and Clements, M. A. (2004). Miscellaneous new species, new genera, reinstated genera and new combinations in Australian Orchidaceae. Orchadian, 14, 15. Jones, D. L. and Clements, M. A. (2005). Miscellaneous nomenclatural changes in Australian, New Guinea and New Zealand Orchidaceae. Orchadian, 15, 37. Kuntze, C. E. O. (1891). Phyllorchis. Revisio generum plantarum, 2, 675. A. Felix, Leipzig. Lindley, J. (1824). Cirrhopetalum, Tribrachia. Botanical Register, 10, t. 832. Lindley, J. (1826). Megaclinium. Botanical Register, 12, t. 989. Lindley, J. (1829). Trias. Wallich’s numerical list, n. 1977. East India Company, London. Lindley, J. (1838). Sertum orchidaceum: a wreath of the most beautiful orchidaceous flowers. Ridgway and Sons, London. Lindley, J. (1840). The genera and species of orchidaceous plants. J. Ridgway, London. Lindley, J. (1845). Porpax. Botanical Register misc., 62. Lindley, J. (1852). An enumeration of the known species of orchids. Folia Orchidacea, 1, 1. J. Matthews, London. Lindley, J. (1853). Acrochaene. Folia orchidacea, 2, 1. J. Matthews, London. Lindley, J. and Paxton, J. (1850). Sarcopodium. Paxton’s Flower Garden, 1, 155. Luer, C. A. (1972). The native orchids of Florida. New York Botanical Garden, New York. Ormerod, P. (2002). Orchidaceae fragmentae 4. Oasis, 2, 4-7. Pearce, N., Cribb, P. J., and Renz, J. (2001). Notes relating to the flora of Bhutan 44. Taxonomic notes, new taxa and additions to the Orchidaceae of Bhutan and Sikkim (India). Edinburgh Journal of Botany, 58, 99–122. Pfitzer, E. H. H. (1889). Orchidaceae. In A. Engler and K. Prantl (eds.), Die Natürlichen Pflanzenfamilien, 2(6), 52–224. Rao, A. N. and Mani, K. J. (1985). Jejosephia Nageswara Rao et Mani (Orchidaceae) - a new genus from Meghalaya, India. Journal of Economic and Taxonomic Botany, 7, 216. Reichenbach, H. G. (1852). Bolbophyllaria. Botanische Zeitung, 10, 919–34. Reinwardt, C. G. C. (1828). Zygoglossum. Sylloge Plantarum Novarum, 2, 5. Ribeiro, P. L., Borba, E. L., Smidt, E. C., Lambert, S. M., SelbachSchnadelbach, A., and van den Berg, C. (2008). Genetic and morphological variation in the Bulbophyllum exaltatum (Orchidaceae) complex in the Brazilian “campo rupestres”: implications for taxonomy and biogeography. Plant Systematics and Evolution, 270, 109–37. Rolfe, R. A. (1919). The genus Hyalosema. Orchid Review, 27, 130. Schlechter, F. R. R. (1905). Pedilochilus. Nachtrage zur Flora der Deutschen Schutzgebiete in der Sudsee (mit Ausschluss Samoa’s und der Karolinen), 218. Borntraeger, Leipzig, Germany. Schlechter, F. R. R. (1907). Beitrage zur Flora von Afrika XXVIII. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie, 38, 11. Schlechter, F. R. R. (1913). Bulbophyllinae. Repertorium Specierum Novarum Regni Vegetabilis, Beihefte, 1, 683–893. Schlechter, F. R. R. (1915). Orchidaceae Stolzianae, ein Beitrag zur Orchideenkunde des Nyassa Landes. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie, 53, 564. Seidenfaden, G. (1973). Notes on Cirrhopetalum Lindl. Dansk Botanisk Arkiv, 29, 1–277. Seidenfaden, G. (1979). Orchid genera in Thailand VIII. Bulbophyllum Thou. Dansk Botanisk Arkiv, 33(3), 1–228. Senghas, K. (1978). Hapalochilus, eine alte, neue Orchideengattung aus Neu Guinea. Orchidee, 29, 245–48.

Smidt, E. C., Borba, E. L., Gravendeel, B., Fischer, G. A., and van den Berg, C. (2011). Molecular phylogeny of the Neotropical sections of Bulbophyllum (Orchidaceae) using nuclear and plastid spacers. Taxon, 60, 1050–64. Smidt, E. C., Silva-Pereira, V., Borba, E. L., and van den Berg, C. (2007). Richness, distribution and important areas to preserve Bulbophyllum in the Neotropics. Lankesteriana, 7, 107–13. Summerhayes, V. S. (1961). A new genus of orchids from southern Rhodesia. Kirkia, 1960–61, 88. Szlachetko, D. L. and Margonska, H. B. (2001). Genera et species orchidalium III. Polish Botanical Journal, 46, 114. Szlachetko, D. L. and Margonska, H. B. (2002). Gynostemia orchidalium II. Acta Botanica Fennica, 173, 123–47. Szlachetko, D. L. and Kulak, M. (2007). Contribution a la revision generique de Bulbophyllinae (Epidendroideae). Richardiana, 7, 82–94. Thouars, A. A. du Petit-. (1809). Extrait de trois Mémoires lus à la première classe de l’Institut, sur l’histoire des plantes orchidées des iles australes d’Afrique. Nouveau Bulletin de Sciences, publié par la Société Philomathique Paris, 1, 314–19. Thouars, A. A. du Petit-. (1822). Histoire particuliere des plantes orchidées recueillies sur les trois Iles Australes d’Afrique, de France, de Bourbon et de Madagascar, 90. Published by the author, Paris. Thouars, L. M. A. (1804-1819). Cahier des six plances. Published by the author, Paris. Vermeulen, J. J. (1991). Bulbophyllum. In Orchids of Borneo. Vol. 2. Bulbophyllum (ed. P. J. Cribb). Bentham-Moxon Trust, Royal Botanic Gardens, Kew, and Toihaan Publishing Company, Kota Kinabalu, Sabah, Malaysia. Vermeulen, J. J. (1993). A taxonomic revision of Bulbophyllum sections Adelopetalum, Lepanthanthe, Macrouris, Pelma, Peltopus, and Uncifera (Orchidaceae). Orchid Monographs, 7, 1–324. Vermeulen, J. J. (2002). A taxonomic revision of Bulbophyllum (Orchidaceae) 2. Sections Altisceptrum and Hirtula. Gardens’ Bulletin Singapore, 54, 1–151. Vermeulen, J. J. (2008). New species of Bulbophyllum from eastern Malesia (Orchidaceae). Nordic Journal of Botany, 26, 129–95. Vermeulen J. J. and O’Byrne, P. (2008). Thirty-two new species of Bulbophyllum (Orchidaceae) from Sulawesi. Gardens’ Bulletin Singapore, 60, 73–153. Vermeulen, J. J. and O’Byrne, P. (2011). Bulbophyllum of Sulawesi. Natural History Publications, Borneo. Sabah, Malaysia. Wight, R. (1851). Oxysepala. Icones plantarum Indiae orientalis, 5, 17. Pharoah, Madras.

6 2 2 .   DENDROB I U M Dendrobium Sw., Nova Acta Regiae Soc. Sci. Upsal., 6, 82 (1799), nom. cons. Conserved type species: Dendrobium moniliforme (L.) Sw. (basionym: Epidendrum moniliforme L.) Callista Lour., Fl. Cochinch., 2, 519 (1790). Type species: Callista amabilis Lour. Ceraia Lour., Fl. Cochinch., 2, 518 (1790). Type species: Ceraia simplicissima Lour. Dendrorkis Thouars, Nouv. Bull. Sci. Soc. Philom. Paris, 19, 314 (1809) nom. superfl. Aporum (Blume) Blume, Bijdr., 334 (1825). Type species: Aporum lobatum Blume Desmotrichum (Blume) Blume, Bijdr., 329 (1825), nom. rej., not Desmotrichum Kütz. Type species: Desmotrichum angulatum Blume 51

DENDROBIEAE

Grastidium (Blume) Blume, Bijdr., 333 (1825). Type species: Grastidium salaccense Blume Macrostomium (Blume) Blume, Bijdr., 335 (1825). Type species: Macrostomium aloifolium Blume Onychium (Blume) Blume, Bijdr., 323 (1825). Type species: Onychium japonicum Blume Pedilonum (Blume) Blume, Bijdr., 320 (1825). Type species: Pedilonum secundum Blume Cadetia Gaudich. in Freyc., Voy. Bol., 422, t. 33 (1826). Type species: Cadetia umbellata Gaudich. Gersinia Néraud in Freyc., Voy. Uranie, 27 (1826). Type species: Not designated (no species have as yet been assigned to Gersinia), syn. nov. Schismoceras C.Presl, Rel. Haenk., 1, 96, t. 13, fig. 2 (1827). Type species: Schismoceras disticha C.Presl Orthoglottis Breda, Gen. Sp. Orchid. Asclep., fasc. 4, s. n. (1830). Type species: Orthoglottis imbricata Breda Thelychiton Endl., Prodr. Fl. Norf., 32 (1833). Type species: Thelychiton macropus Endl. Endeisa Raf., Fl. Tellur., 2, 51 (1836, publ. 1837). Type species: Endeisa flava Raf., nom. illeg. (basionym: Dendrobium densiflorum Wall ex Lindl.) Tropilis Raf., Fl. Tellur., 2, 95 (1836, publ. 1837). Type species: Tropilis aemula (R.Br.) Raf., as ‘emulum’ (basionym: Dendrobium aemulum R.Br.) Froscula Raf., Fl. Tellur., 4, 44 (1836, publ. 1838). Type species: Froscula hispida Raf., nom. illeg. (basionym: Dendrobium longicornu Lindl.) Ormostema Raf., Fl. Tellur., 4, 38 (1836, publ. 1838). Type species: Ormostema purpurea Raf., nom. illeg. (basionym: Epidendrum moniliforme L.) Ditulima Raf., Fl. Tellur., 4, 41 (1836, publ. 1838). Type species: Ditulima anceps (Sw.) Raf. (basionym: Dendrobium anceps Sw.) Pierardia Raf., Fl. Tellur., 4, 41 (1836, publ. 1838). Type species: Pierardia bicolor Raf., nom. illeg. (basionym: Dendrobium pierardii Roxb.) Thicuania Raf., Fl. Tellur., 4, 47 (1836, publ. 1838). Type species: Thicuania moschata (Buch.-Ham.) Raf. (basionym: Epidendrum moschatum Buch.-Ham.) Latouria Blume, Rumphia, 4, 41, t. 195, 199 (1848), nom. illeg., non (Endl.) Lindl. Type species: Latouria spectabilis Blume Sarcopodium Lindl., Paxt. Flow. Gard., 1, 155 (1850), nom. illeg., non Ehrenb. ex Brogn. Type species: Sarcopodium amplum (Lindl.) Lindl. & Paxton (basionym: Dendrobium amplum Lindl.) Aclinia Griff., Notul. Pl. Asiat., 3, 320 (1851). Type species: Dendrobium aclinia Lindl., nom. illeg., non Rchb.f. Dichopus Blume, Mus. Bot. Lugd. Bat., 2, 176 (1856). Type species: Dichopus insignis Blume Coelandria Fitzg., Austral. Orchids, 1(7), t. 2 (1882). Type species: Coelandria smillieae (F.Muell.) Fitzg. (basionym: Dendrobium smillieae F.Muell.) Sayeria Kränzl., Oesterr. Bot. Zeitschr., 44, 298 (1894). Type species: Sayeria paradoxa Kränzl. Diplocaulobium (Rchb.f.) Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21, 331 (1910). Type species: Diplocaulobium nitidissimum (Rchb.f.) Kränzl. (basionym: Dendrobium nitidissimum Rchb.f.) 52

Inobulbum (Schltr.) Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21, 316 (1910), as ‘Inobulbon Schltr. & Kränzl.’; Schlechter (1912) did not accept Inobulbum as a genus and therefore should not be considered co-author. Type species: Inobulbum muricatum (Finet) Kränzl. (basionym: Dendrobium muricatum Finet) Goldschmidtia Dammer, Orchis, 4, 86 (1910), nom. invalid. Type species: Goldschmidtia gracilis Dammer Epigeneium Gagnep., Bull. Mus. Hist. Nat. Paris, sér. 2, 4, 593 (1932). Type species: Epigeneium fargesii (Finet) Gagnep. (basionym: Dendrobium fargesii Finet) Katharinea A.D.Hawkes, Lloydia, 19, 94 (1956). Type species: Katharinea ampla (Lindl.) A.D.Hawkes (basionym: Dendrobium amplum Lindl.) Flickingeria A.D.Hawkes, Orquidea, 27, 301 (1961). Type species: Flickingeria angulata (Blume) A.D.Hawkes (basionym: Desmotrichum angulatum Blume) Ephemerantha P.F.Hunt & Summerh., Taxon, 10, 102 (1961). Type species: Ephemerantha angulata (Blume) P.F.Hunt & Summerh. (basionym: Desmotrichum angulatum Blume) Amblyanthus (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 686 (1981), nom. illeg., non A.DC. Type species: Amblyanthus melanostictus (Schltr.) Brieger (basionym: Dendrobium melanostictum Schltr.) Australorchis Brieger in Schltr., Orchideen, ed. 3, 1, 741 (1981). Type species: Australorchis monophylla (F.Muell.) Brieger (basionym: Dendrobium monophyllum F.Muell.) Bolbodium (Lindl.) Brieger in Schltr., Orchideen, ed. 3, 1, 721 (1981), as ‘Bolbidium’ (not Bolbidium (Lindl.) Lindl.). Type species: Bolbodium pusillum (Blume) Rauschert (basionym: Desmotrichum pusillum Blume) Conostalix (Kränzl.) Brieger in Schltr., Orchideen, ed. 3, 1, 659 (1981). Type species: Conostalix calcarata Brieger (basionym: Dendrobium calcaratum Lindl., nom. illeg., non A.Rich.) Dockrillia Brieger in Schltr., Orchideen, ed. 3, 1, 745 (1981). Type species: Dockrillia linguiformis (Sw.) Brieger (basionym: Dendrobium linguiforme Sw.) Eriopexis (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 656 (1981). Type species: Eriopexis schlechteri Brieger (basionym: Dendrobium eriopexis Schltr.) Euphlebium (Kränzl.) Brieger in Schltr., Orchideen, ed. 3, 1, 722 (1981). Type species: Euphlebium spurium (Blume) Brieger (basionym: Dendrocolla spuria Blume) Herpethophytum (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 660 (1981). Type species: Herpethophytum schlechteri Rauschert (basionym: Dendrobium herpetophytum Schltr.) Kinetochilus (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 686 (1981). Type species: Kinetochilus pectinatus (Finet) Brieger (basionym: Dendrobium pectinatum Finet) Latourorchis Brieger in Schltr., Orchideen, ed. 3, 1, 727 (1981). Type species: Latourorchis spectabilis (Blume) Brieger (basionym: Latouria spectabilis Blume) Microphytanthe (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 742 (1981). Type species: Microphytanthe bulbophylloides (Schltr.) Brieger (basionym: Dendrobium bulbophylloides Schltr.)

DENDROBIUM

Monanthos (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 660 (1981), as ‘Monanthus’. Type species: Monanthos bilobus (Schltr.) Brieger (basionym: Dendrobium bilobum Lindl.) Trachyrhizum (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 687 (1981). Type species: Trachyrhizum schlechteri Brieger (basionym: Dendrobium trachyrhizum Schltr.) Amblyanthe Rauschert, Feddes Repert., 94, 436 (1983). Type species: Amblyanthe melanosticta (Schltr.) Rauschert (basionym: Dendrobium melanostictum Schltr.) Cannaeorchis M.A.Clem. & D.L.Jones, Lasianthera, 1, 132 (1997). Type species: Cannaeorchis fractiflexa (Finet) M.A.Clem. & D.L.Jones (basionym: Dendrobium fractiflexum Finet) Winika M.A.Clem., D.L.Jones & Molloy, Orchadian, 12, 214 (1997). Type species: Winika cunninghamii (Lindl.) M.A.Clem., D.L.Jones & Molloy (basionym: Dendrobium cunninghamii Lindl.) Tetrodon (Kränzl.) M.A.Clem. & D.L.Jones, Orchadian, 12, 310 (1998). Type species: Tetrodon oppositifolium (Kränzl.) M.A.Clem. & D.L.Jones (basionym: Eria oppositifolia Kränzl.) Abaxianthus M.A.Clem. & D.L.Jones, Orchadian, 13, 485 (2002). Type species: Abaxianthus convexus (Blume) M.A.Clem. & D.L.Jones (basionym: Desmotrichum convexum Blume) Aporopsis (Schltr.) M.A.Clem. & D.L.Jones, Orchadian, 13, 485 (2002). Type species: Aporopsis macfarlanei (F.Muell.) M.A.Clem. & D.L.Jones (basionym: Dendrobium macfarlanei F.Muell.) Bouletia M.A.Clem. & D.L.Jones, Orchadian, 13, 485 (2002). Type species: Bouletia finetiana (Schltr.) M.A.Clem. & D.L.Jones (basionym: Dendrobium finetianum Schltr.) Cepobaculum M.A.Clem. & D.L.Jones, Orchadian, 13, 486 (2002). Type species: Cepobaculum canaliculatum (R.Br.) M.A.Clem. & D.L.Jones (basionym: Dendrobium canaliculatum R.Br.) Ceratobium (Lindl.) M.A.Clem. & D.L.Jones, Orchadian, 13, 486 (2002). Type species: Ceratobium antennatum (Lindl.) M.A.Clem. & D.L.Jones (basionym: Dendrobium antennatum Lindl.) Chromatotriccum M.A.Clem. & D.L.Jones, Orchadian, 13, 493 (2002). Type species: Chromatotriccum lawesii (F.Muell.) M.A.Clem. & D.L.Jones (basionym: Dendrobium lawesii F.Muell.) Davejonesia M.A.Clem., Orchadian, 13, 487 (2002). Type species: Davejonesia lichenastrum (F.Muell.) M.A.Clem. (basionym: Bulbophyllum lichenastrum F.Muell.) Dendrobates M.A.Clem. & D.L.Jones, Orchadian, 13, 487 (2002). Type species: Dendrobates virotii (Guillaumin) M.A.Clem. & D.L.Jones (basionym: Dendrobium virotii Guillaumin) Distichorchis M.A.Clem. & D.L.Jones, Orchadian, 13, 487 (2002). Type species: Distichorchis revoluta (Lindl.) M.A.Clem. & D.L.Jones (basionym: Dendrobium revolutum Lindl.) Durabaculum M.A.Clem. & D.L.Jones, Orchadian, 13, 487 (2002), nom. illeg. Type species: Durabaculum undulatum (R.Br.) M.A.Clem. & D.L.Jones (basionym: Dendrobium undulatum R.Br.) Eleutheroglossum (Schltr.) M.A.Clem. & D.L.Jones, Orchadian, 13, 489 (2002). Type species: Dendrobium eleutheroglossum Schltr.

Eurycaulis M.A.Clem. & D.L.Jones, Orchadian, 13, 490 (2002). Type species: Eurycaulis lamellatus (Blume) M.A.Clem. & D.L.Jones (basionym: Onychium lamellatum (Blume) Lindl.) Exochanthus M.A.Clem. & D.L.Jones, Orchadian, 13, 496 (2002). Type species: Exochanthus pleianthum (Schltr.) M.A.Clem. & D.L.Jones (basionym: Dendrobium pleianthum Schltr.) Leioanthum M.A.Clem. & D.L.Jones, Orchadian, 13, 490 (2002). Type species: Leioanthum bifalce (Lindl.) M.A.Clem. & D.L.Jones (basionym: Dendrobium bifalce Lindl.) Maccraithea M.A.Clem. & D.L.Jones, Orchadian, 13, 496 (2002). Type species: Maccraithea cuthbertsonii (F.Muell.) M.A.Clem. & D.L.Jones (basionym: Dendrobium cuthbertsonii F.Muell.) Oxyglossellum M.A.Clem. & D.L.Jones, Orchadian, 13, 490 (2002). Type species: Oxyglossellum cyanocentrum (Schltr.) M.A.Clem. & D.L.Jones (basionym: Dendrobium cyanocentrum Schltr.) Sarcocadetia (Schltr.) M.A.Clem. & D.L.Jones, Orchadian, 13, 490 (2002). Type species: Sarcocadetia funiformis (Blume) M.A.Clem. & D.L.Jones (basionym: Dendrobium funiforme Blume) Stelbophyllum M.A.Clem. & D.L.Jones, Orchadian, 13, 490 (2002). Type species: Stelbophyllum toressae (F.M.Bailey) M.A.Clem. & D.L.Jones (basionym: Dendrobium toressae F.M.Bailey) Tetrabaculum M.A.Clem. & D.L.Jones, Orchadian, 13, 490 (2002). Type species: Tetrabaculum tetragonum (A.Cunn.) M.A.Clem. & D.L.Jones (basionym: Dendrobium tetragonum A.Cunn.) Vappodes M.A.Clem. & D.L.Jones, Orchadian, 13, 492 (2002). Type species: Vappodes bigibba (Lindl.) M.A.Clem. & D.L.Jones (basionym: Dendrobium bigibbum Lindl.) Anisopetala (Kränzl.) M.A.Clem., Telopea, 10(1), 283 (2003). Type species: Anisopetala mutabilis (Blume) M.A.Clem. (basionym: Dendrobium mutabile Blume)

Derivation of name From the Greek dendron, tree, and bios, life, referring to the epiphytic habit of most Dendrobium species. (AS)

Description (Plates 23–53; Fig. 622.1–622.6) Epiphytic, lithophytic, or rarely terrestrial herbs, sympodial but individual shoots sometimes with indeterminate apical growth (D. sections Grastidium, Herpethophytum, and others). Roots terete, glabrous or papillose-warty. Stems cane-like or having pseudobulbs, glabrous to hirsute. Cane-like stems fleshy or not, branching or not, long-lived, entirely or partially covered by leaf sheaths if present. Pseudobulbs consisting of one to many internodes, one- to many-leaved. Leaves conduplicate, usually articulate, with or without a tubular leaf sheath, distichous (rarely quaquaversal in D. section Grastidium), dorsiventrally flattened, bilaterally flattened or terete, margins entire. Inflorescence lateral or terminal, usually arising from a mature shoot, one-flowered or racemose, rarely paniculate, sometimes fasciculate. Flowers resupinate or not, ephemeral to long-lasting, distichous, secund or helically arranged, sometimes fragrant. Sepals entire, free or variously connate, lateral sepals similar or dissimilar to the dorsal sepal and usually forming a mentum together with column foot, mentum sometimes slender and spur-like. Petals entire, free, similar to 53

DENDROBIEAE

C

E

D

6 mm

G 2 mm

I

1 mm 5 cm

B A

H

6 mm

2 mm

F

Fig. 622.1.  Dendrobium brevicolle J.J.Sm. (D. sect. Diplocaulobium). A. habit; B. Flowering shoot; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum, front view; G. Labellum and column, side view; H. Anther cap, dorsal view; I. Pollinia. Drawn by C. L. Chan and Jaap Vermeulen from material cultivated at Tenom Orchid Centre, Sabah. Reproduced with permission from J. J. Wood (2003), Orchids of Borneo. Volume 4. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

the dorsal sepal or strongly dissimilar. Labellum adnate to sides of column foot, or only attached to apex of column foot and then rigidly attached (will break off when bent downwards), flexibly attached (can be bent downwards without breaking off), or rarely hinged and highly mobile, entire or lobed, with or without keels and/or callosities, not spurred but often forming a spur-like structure with column foot. Column usually straight, without lateral wings, rarely with apical stelidia; column foot well-developed, often with a cavity or a shallowly concave area at apex; anther helmet-shaped, two-locular, glabrous or papillosehairy, pollinia 4, oblong, triangular in cross-section, hard-waxy, strongly cohering in two subequal pairs, without caudicles, usually yellow, sometimes brown, purplish or grey; stigma a ventral concavity on the column, sometimes raised and callus-like; 54

rostellum simple or sometimes bilobed, usually swollen and forming a reservoir containing a sticky, often white and opaque, sometimes hyaline or purple liquid, which rapidly dries out to a soft, putty-like substance; viscidium usually absent but, when present, shapeless and formed instead of (or in addition to) rostellar liquid on the underside of the rostellum [‘diffuse viscidium’ as used by Rasmussen (1982) and Freudenstein and Rasmussen (1999)], often with a downwardly projecting, rigid, truncate or emarginate lamella between rostellum and stigma (the ‘scraper’). Ovary and pedicel terete or winged, sometimes papillose or hairy. Capsule ellipsoid, oblong, obovoid to subglobose, unilocular, sometimes winged or triangular in cross-section, usually opening along three longitudinal slits, sometimes with a single opening near the apex, without endocarpic elaters. (AS)

DENDROBIUM

1 cm

P

A

M H

E

1 mm

O

B C

F D

I G

N J

K

L 0.5 mm

Fig. 622.2.  Dendrobium bulbophylloides Schltr. (D. sect. Microphytanthe). A. Flower, face view; B. Dorsal sepal; C. Petal; D. Lateral sepal; E. Labellum, proximal region; F. labellum, distal region; G. Labellum, transverse section; H. Column and labellum, side view; I. Column, ventral view; J. Anther cap, ventral view; K. Anther cap, side view; L. Pollinia; M. Flower, side view; N. Ovary, transverse section; O. Habit; P. Ovary, surface spotting. Single bar = 1 mm, double bar = 1 cm. Drawn by Andrew Brown from Reeve 325 (K, including spirit material) and Sands 1825.

Distribution (Fig. 622.7) The genus of about 1450 species ranges from Sri Lanka throughout tropical Asia and the Pacific region, north to Japan, east to Tahiti, and south to New Zealand. (AS)

Artificial key to the sections of Dendrobium (AS) 1.

●



●

2.

●



●

Shoot 1-leaved; inflorescence 1-flowered (possibly in fascicles) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Shoot with 2 or more leaves or inflorescence with more than 1 flower . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Stems or pseudobulbs consisting of more than 1 internode . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Stems or pseudobulbs consisting of a single internode . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

3.

●



●

4.

●

5.

●

●



●

6.

●

Labellum without keels or callosities, adaxially finely pubescent all over . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Dendrobium (D. jenkinsii) Labellum with keels or callosities, adaxially glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Pseudobulbs distant along a creeping or erect rhizome . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Crinifera Pseudobulbs crowded, rhizome not creeping . . . . . . . . . 5 Pseudobulbs quadrangular in cross-section; labellum hinged with column foot, mobile; flowers ephemeral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Fugacia Pseudobulbs rounded in cross-section; labellum firmly affixed to column foot, not mobile; flowers longer-lasting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Latouria Labellum with 2 or more keels that extend from base to at least halfway . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 55

DENDROBIEAE

K

2 mm

I

J 2 mm

1 cm

G

F

H

5 cm

E

5 mm

B

D

1 cm

A

C Fig. 622.3.  Dendrobium kinabaluense Ridl. (D. sect. Sarcopodium). A. Habit; B. Pseudobulb, transverse section; C. Labellum, spread out; D. Column, dorsal view; E. Column and labellum, longitudinal section; F. Lateral sepal; G. Dorsal sepal; H. Petal; I. Ovary, transverse section; J. Pollinia; K. Anther. Drawn by Chan Chew Lun from Lamb AL 40/83. Reproduced with permission from Chan et al. (1994), Orchids of Borneo. Volume 1. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.



●

7.

●



●

8.

●



●

9.

56

Labellum without such basal keels, possibly with callosities near middle, or unadorned . . . . . . . . . . . . . . . . . . . . 10 Inflorescences arising from a (usually yellow) spathe at apex of pseudobulb . . . . . . . . . . D. sect. Diplocaulobium Inflorescences arising at or below apex of pseudobulb or stem, not from a spathe . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Leaves fleshy, terete, or if flat, without a distinct midvein . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Rhizobium Leaves not fleshy, not terete, with a distinct midvein . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Rhizome erect, or, if creeping, the individual pseudobulbs terminating short, erect branches of rhizome, or rhizome pendulous and pseudobulbs highly reduced;

●



flowers ephemeral, from apex of pseudobulb or laterally just below apex . . . . . . . . . . . . . . . . . . . . D. sect. Crinifera Rhizome creeping or horizontally spreading, with welldeveloped, sessile pseudobulbs; flowers longer-lasting, arising from apex of pseudobulb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Sarcopodium

●

10. Plant without pseudobulbs, with highly reduced stem, the fleshy leaves seemingly sessile on rhizome . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Lichenastrum Plant with distinct pseudobulbs or stems . . . . . . . . . . . . 11 ●

●

11. Labellum with basal part closely appressed and adnate to column foot; mentum often spur-like; column pubescent in front . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Cadetia ●

DENDROBIUM

F

H

E 5 mm

5 mm

G

5 cm

A B

D 1 cm

1 mm

I

K

C 1 mm

J 5 mm Fig. 622.4.  Dendrobium patentilobum Ames & C.Schweinf. (D. sect. Aporum). A. Habit; B. Flower, front view; C. Flower, side view; D. Ovary, transverse section; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum, flattened; I. Pedicel with ovary, column, and labellum, longitudinal section; J. Anther cap, dorsal view; K. Pollinia. Drawn by C. L. Chan and Lucy F. L. Liew from Lamb AL 1117/89. Reproduced with permission from J. J. Wood (1997), Orchids of Borneo. Volume 3. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.



Labellum with the basal part held away from and hinged with column foot; mentum not spur-like; column glabrous in front . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 12. Labellum with distinct, erect side lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Sarcopodium Labellum entire or indistinctly lobed, the lobes not erect . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Microphytanthe 13. Stem or pseudobulb 1- or 2-leaved at apex, the leaves not attached to a tubular sheath . . . . . . . . . . . . . . . . . . . . . . . 14 Stem or pseudobulb with more than two leaves, or leaves attached to a tubular sheath . . . . . . . . . . . . . . . . . . . . . . . 25

15. 16.

14. Leaves terete, or fleshy and bilaterally flattened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Rhizobium

18. Pseudobulbs oblong or ovoid to broadly conical, not with a narrow base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

●

●

●

●

●

●

Leaves dorsiventrally flattened . . . . . . . . . . . . . . . . . . . . 15 Rhizome elongate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 Rhizome short . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 Labellum linear, entire, the basal part adnate to column foot . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Calyptrochilus Labellum trilobed or wedge-shaped, the basal part not adnate to column foot . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 17. Column about half as long as column foot. Australia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Monophyllaea Column almost as long as or longer than column foot. Asia . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Sarcopodium ●

● ●

●

●

●

●

●

57

DENDROBIEAE

C

B

H

D

A F I 3 cm 5 cm 2 mm

E

G

Fig. 622.5.  Dendrobium spectatissimum Rchb.f. (D. sect. Formosae). A. Plant; B. Capsule; C. Capsule, transverse section; D. Labellum; E. Petal; F. Lateral sepal; G. Column and labellum, longitudinal section; H. Dorsal sepal; I. Anther. Drawn by Chan Chew Lun from Cribb in Bailes 818. Reproduced with permission from Chan et al. (1994), Orchids of Borneo. Volume 1. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.



Pseudobulbs narrowly cylindrical or with a narrow base (pseudobulbs spherical, or spindle-shaped to clubshaped) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 19. Inflorescence erect, branched or not; labellum trilobed, with a bilobed callus or three, horn-like projections at base of midlobe; no callus between lateral lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Macrocladium Inflorescence, if erect, not branched; labellum if trilobed with a callus that extends from base . . . . . . . . . . . . . . . . 20 20. Labellum suborbicular, thin-textured, ecallose . . . . . . . . . . . . . . . . D. sect. Dendrobium (D. lindleyi, D. jenkinsii) Labellum, if suborbicular, thick-textured, with a callus 21



21. Labellum with a basal callus; sepals fleshy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Latouria

24. Flowers with thick, rigid sepals and petals, sometimes with conspicuous sharp keels or thick hairs on sepals or

●

●

●

●

●

●

58

Labellum with an apical callus; sepals thin . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Monophyllaea (D. schneiderae) 22. Column foot much shorter than column; labellum mobile; flowers ephemeral . . . . . . . . . . . . . . . . . . D. sect. Fugacia Column foot about as long as or longer than column; labellum not mobile; flowers longer-lasting or ephemeral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 23. Labellum basally adnate to column foot; inflorescence 1-flowered; flowers ephemeral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Aporum Labellum only attached to apex of column foot; inflorescence 1- to many-flowered; flowers longer lasting . . . . 24 ●

●

●

●

●

●

DENDROBIUM

J C

I

O

H

D

F

B

A

K

M

E G

N

L

Fig. 622.6.  Dendrobium verruciferum Rchb.f. (D. sect. Macrocladium). A. Habit; B. Base of plant; C. Flower; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum; H. Column and labellum, side view; I. Column, anther cap removed, front view; J. Column apex, front view; K. Column, anther cap removed, side view; L. Anther cap, dorsal view; M. Anther cap, ventral view; N. Pollinia; O. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from MacKee 19, sheet 25288 (K); B from MacKee 19, sheet 28703 (K); others from Kew Spirit Collection no. 53131.

ovary, not with narrow sepals, long-lasting (more than two weeks); rostellum swollen, white, exuding a white liquid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Latouria Flowers with thin parts, without sharp keels or hairs on sepals or ovary, small, or, if large, with narrow sepals; not long-lasting (about two weeks); rostellum not swollen, hyaline, not exuding a white liquid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Dendrocoryne 25. Leaves laterally flattened . . . . . . . . . . . . . D. sect. Aporum Leaves dorsiventrally flattened or terete to needle-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 ●

● ●

26. Leaves terete to needle-shaped . . . . . . . . . . . . . . . . . . . . 27 Leaves flat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 27. Leaf sheaths covered with blackish or dark brown hairs, at least during development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Conostalix Leaf sheaths glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 28. Stems much shorter than leaves, fleshy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Pedilonum (D. hellwigianum) Stems elongate, much longer than leaves, not fleshy, or with a few abruptly swollen internodes basally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 ● ●

●

●

●

●

59

DENDROBIEAE

Fig. 622.7.  Distribution map of Dendrobium.

29. Inflorescence at base subtended by laterally flattened scales that persist for a long time after the flowers have fallen off, 1- or 2-flowered; stems never with swollen internodes; column foot much shorter than dorsal sepal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Grastidium Inflorescence at base with a few tight, tubular scales that wither away soon after flowering, 1-flowered; stem with or without swollen internodes basally; column foot as long as or longer than dorsal sepal . . . . . . . . . . . D. sect. Aporum 30. Stem slender, with a few of the lower, leafless internodes abruptly swollen above base . . . . . . . . . . D. sect. Aporum Stem either swollen throughout (sometimes with thinner branches) or slender throughout, or with upper internodes swollen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 31. Leaf sheaths densely covered with short, blackish or dark brown hairs, at least when developing . . . . . . . . . . . . . . 32 Leaf sheaths, if present, glabrous, or with white hairs, or long hairs along margin only, or scattered brownish scale hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 32. Flowers showy . . . . . . . . . . . . . . . . . . . D. sect. Formosae Flowers dull-coloured . . . . . . . . . . . . D. sect. Conostalix 33. Leaves not attached to a tubular sheath . . . . . . . . . . . . . 34 At least the lowermost leaf attached to a tubular sheath . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 ●

●

●

●

●

●

● ●

● ●

60

34. Column foot much shorter than column; labellum mobile; flowers ephemeral . . . . . . . . . . . . . . . . . . D. sect. Fugacia Column foot about as long as or longer than the column; labellum not mobile; flowers longer-lasting . . . . . . . . . . 35 ●

●

35. Labellum entire, adaxially finely pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Dendrobium Labellum entire or trilobed, adaxially entirely or mostly glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 ●

●

36. Flowers with thick, rigid sepals and petals, sometimes with conspicuous sharp keels or thick hairs on sepals or ovary, small to large, not with narrow sepals; long-lasting (more than two weeks); rostellum swollen, white, exuding a white liquid . . . . . . . . . . . . . . . . . . . . . . D. sect. Latouria Flowers with thin parts, without sharp keels or hairs on sepals or ovary, small, or, if large, with narrow sepals; not long-lasting (about two weeks); rostellum not swollen, hyaline, not exuding a white liquid . . . . . D. sect. Dendrocoryne ●

●

37. Stems fleshy or not, short or elongate; if not fleshy, each stem with limited apical growth (older stems have clearly ceased growing), often becoming leafless after one or two growing seasons; flowers mainly produced on stems of the current or previous growing season . . . . . . . . . . . . . 38 Stems not fleshy, of even thickness throughout, not becoming grooved with age, elongate, with seemingly ●

●

DENDROBIUM

unlimited apical growth, not becoming completely leafless after only a few growing seasons; flowers produced on both recent and old stems . . . . . . . . . . . . . . . . . . . . . 56 38. Mentum in lateral view bilobed; petals broader than sepals . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Phalaenanthe Mentum not bilobed in lateral view or, if bilobed; petals not broader than sepals . . . . . . . . . . . . . . . . . . . . . . . . . . 39 ●

●

39. Free part of labellum with two or more lamellate keels or a single longitudinal callus extending from base to halfway or beyond . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 Free part of labellum without keels or long basal callus, or with keels or calli starting well above base of free part of labellum, or with a short basal callus not reaching the middle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 ●

●

40. Inflorescences short, few-flowered, laterally on leafless stems of previous growing season; labellum with a single longitudinal keel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Dendrobium Inflorescences elongate, or, if not, on leafy stems; keels single or multiple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 ●

●

41. Inflorescences short, 1- or few-flowered; leaves not deciduous after a single growing season . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Distichophyllae Inflorescences elongate, or, if short and few-flowered, leaves deciduous after a single growing season . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 ●

●

42. Basal callus bulging backwards over claw of labellum; roots smooth or warty . . . . . . . . . . . D. sect. Brevisaccata Labellum not with a backward-projecting basal callus; roots smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 43. Labellum at base abruptly narrowed into a claw that is adnate to column foot, forming a conical spur; plant small to tall . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Spatulata Labellum not abruptly narrowed into a basal claw that is adnate to column foot; plant small . . . . . . . . . . . . . . . . . 44 ●

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●

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44. Labellum bilobed at apex. Australia and New Caledonia . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Eleutheroglossum Labellum entire or fringed at apex. Continental Asia and Malay Archipelago, east to Lesser Sunda Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Stachyobium 45. Inflorescences from apical part of stem, erect, elongate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 Inflorescences from apical part of stem or lower down, horizontal or pendulous, or, if erect, short . . . . . . . . . . 47 ●

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●

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46. Petals narrower than sepals; labellum bilobed at the apex . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Eleutheroglossum Petals broader than the sepals; labellum entire at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Fytchianthe 47. Leaf sheaths and usually also outer surfaces of sepals with scattered, appressed, brown scale-hairs; labellum with a thin, flat, usually lobed or fringed appendage at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Amblyanthus Leaf sheaths glabrous or with long hairs; outer surfaces of sepals glabrous, papillose or with long patent hairs; ●

●

●

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labellum without appendage or with a thick, callus-like appendage at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 48. Mentum shorter than dorsal sepal, or, if not, mentum broad and chin-like; labellum not adnate to column foot; inner surface of labellum often finely and densely pubescent; roots glabrous . . . . . . . . . . . . . D. sect. Dendrobium Mentum about as long as dorsal sepal or longer, not broad and chin-like but spur-like or conical, or, if mentum shorter than dorsal sepal, then labellum adnate to the column foot; inner surface of labellum not finely and densely pubescent; roots glabrous or warty . . . . . . . . . . . . . . . . . 49 ●

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49. Inflorescence sympodially branching (check remains of older inflorescences), producing one flower at a time . . . . . . . . . . . . . . . D. sect. Brevisaccata (D. viridiflorum) Inflorescence not branching, 1- to many-flowered . . . . 50 ●

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50. Roots rough or warty; stems sometimes laterally flattened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Pedilonum Roots smooth; stems not laterally flattened . . . . . . . . . . 51 ●

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51. Flowers 5–10 cm wide; petals much broader than sepals; inflorescence horizontal or pendent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Formosae Flower smaller; petals often narrower but not much broader than sepals; inflorescence horizontal, pendent or erect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52 ●

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52. Petals broader than dorsal sepal; labellum trilobed, with a large, flat midlobe pubescent at base; inflorescence racemose . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Fytchianthe Petals usually not broader than dorsal sepal; labellum entire or trilobed, not pubescent at base; inflorescence 1-flowered or racemose . . . . . . . . . . . . . . . . . . . . . . . . . . 53 ●

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53. Labellum with distinct lateral lobes or entire; if entire, midlobe warty and clearly more fleshy than sepals and petals; inflorescence short; flowers white, green or brown, not brightly coloured . . . . . . . . . D. sect. Distichophyllae Labellum entire, not warty, not more fleshy than sepals and petals; inflorescence short or elongate; flowers usually brightly coloured, less often white, green or ochre-brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54 ●

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54. Stems not fleshy, or, if fleshy, ovary with wings or papillae, or labellum apex inflexed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Calyptrochilus Stems distinctly fleshy; ovary terete and glabrous; labellum apex not inflexed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55 ●

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55. Species occurring west of Sulawesi, not including the Philippines; also D. secundum, D. amethystoglossum, and D. hasseltii from Sulawesi, the Philippines and farther east (see Taxonomic notes) . . . . . . . . . . . . . . . . D. sect. Pedilonum Species occurring in Sulawesi and farther east, including the Philippines, except D. secundum, D. amethystoglossum, and D. hasseltii (see Taxonomic notes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Calyptrochilus ●

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56. Inflorescence appearing as a cluster of many, almost sessile flowers that breaks through leaf sheath . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Pleianthe ●

61

DENDROBIEAE



Inflorescence 1- or few-flowered, or a raceme . . . . . . . 57

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57. Labellum with a long, narrow claw; mentum usually spurlike due to adnate lateral sepals, often partly closed in front . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58 Labellum not with a long, narrow claw; the basal part rectangular or cuneate; mentum broad, chin-like, open in front, with free lateral sepals . . . . . . . . . . . . . . . . . . . . . . 59 ●

species, from Peninsular Malaysia, Sumatra, Borneo, and New Guinea to the Solomon Islands. All but two species endemic in New Guinea; only one, aberrant species west of New Guinea (D. microglaphys Rchb.f.).

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58. Inflorescence 1-flowered; flower not resupinate; labellum with a callus or longitudinal keels in centre of blade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Herpethophytum Inflorescence 1- to many-flowered; flowers resupinate or not; labellum without a callus or keels on blade, sometimes with a short, V-shaped callus or a horizontal ridge on claw . . . . . . . . . . . . . . . . . . . . . D. sect. Calyptrochilus ●

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59. Inflorescence basally subtended by laterally flattened scales that persist for a long time after flowers have fallen, 1- or 2-flowered, if 2-flowered the flowers arising at same level on peduncle . . . . . . . . . . . . . . . . D. sect. Grastidium Inflorescence not subtended by persistent scales, 1- to many-flowered, if 2-flowered the flowers arising at different levels on peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . 60 ●

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60. Inflorescence 1-flowered, peduncle largely covered by scales; flowers non-resupinate . . . . . . . . . . D. sect. Biloba Inflorescence 1- to many-flowered, peduncle largely exposed; flowers resupinate . . . . . . . . . . . . . . . . . . . . . . . 61 ●

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61. Labellum with a prominent basal callus projecting backward . . . . . . . . . . . . . . . . . . . . . . . . . . D. sect. Brevisaccata Labellum without a basal callus, or with a callus that does not project backward . . . . . . . . . . D. sect. Macrocladium ●

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Infrageneric treatment (AS) Dendrobium sect. Amblyanthus Schltr. in K.Schum. & Lauterb., Nachtr. Fl. Deutsch. Schutzgeb. Südsee, 150 (1905). Type species: Dendrobium melanostictum Schltr. Taxa in other ranks with their type species in D. sect. Amblyanthus: Amblyanthus (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 686 (1981), nom. illeg., non A.DC. Type species: Amblyanthus melanostictus (Schltr.) Brieger (basionym: Dendrobium melanostictum Schltr.) Amblyanthe Rauschert, Feddes Repert., 94, 436 (1983). Type species: Amblyanthe melanosticta (Schltr.) Rauschert (basionym: Dendrobium melanostictum Schltr.) Stems clustered, cane-like, fleshy, usually many-leaved, not branching. Leaves with a tubular sheath at base. Sheaths and usually (except in D. microglaphys Rchb.f.) sepals covered abaxially with appressed brown scale-hairs. Inflorescence usually short, appearing laterally from leafy stems, sometimes with an elongate peduncle and then pendulous in distal part, several-flowered; floral bracts often large; mentum conical, parallel with the ovary. Labellum attached to apex of column foot, not adnate to it, entire or weakly lobed, fleshy, with a flat, appressed, backward-pointing, lobed or lacerate appendage at base, rarely the appendage reduced to a horizontal ridge; labellum often pubescent apically. About 17 62

Dendrobium sect. Aporum Blume, Tab. Pl. Jav. Orchid., in Clavis generum (1825). Type species: Dendrobium lobatum (Blume) Miq. (basionym: Aporum lobatum Blume) Dendrobium sect. Macrostomium Blume, Tab. Pl. Jav. Orchid., in Clavis generum (1825). Type species: Dendrobium aloifolium (Blume) Lindl. (basionym: Macrostomium aloifolium Blume) Dendrobium sect. Bolbodium Lindl., Paxton’s Fl. Gard., 1, 136 (1851). Type species: Dendrobium pusillum (Blume) Lindl., nom. illeg., non D.Don (basionym: Desmotrichum pusillum Blume) Dendrobium sect. Strongyle Lindl., Paxton’s Fl. Gard., 1, 134 (1851). Type species: Dendrobium subulatum (Blume) Lindl. (basionym: Onychium subulatum Blume) Dendrobium sect. Crumenata Pfitzer in Engl. & Prantl, Nat. Pflanzenfam., II, 6, 174 (1889). Type species: Dendrobium crumenatum Sw. Dendrobium sect. Ampullaria Pfitzer in Engl. & Prantl, Nat. Pflanzenfam., II, 6, 174 (1889). Type species: Dendrobium planibulbe Lindl. Dendrobium sect. Clavatae Ridl., J. Linn. Soc. Bot., 32, 232 (1896). Type species: Dendrobium clavator Ridl. Dendrobium sect. Virgatae Hook.f., Fl. Brit. Ind., 5, 711 (1890). Type species: Dendrobium crumenatum Sw. Dendrobium sect. Hemiphylla Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21, 201 (1910). Type species: Dendrobium aloifolium (Blume) Lindl. (basionym: Macrostomium aloifolium Blume) Dendrobium sect. Holophylla Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21, 201 (1910). Type species: Dendrobium lobatum (Blume) Miq. (basionym: Aporum lobatum Blume) Dendrobium sect. Rhopalanthe Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 449 (1912). Type species: Dendrobium crumenatum Sw. Aporum sect. Aporopsis (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 673 (1981) , nom. invalid. Type species: Aporum macfarlanei (F.Muell.) Rauschert (basionym: Dendrobium macfarlanei F.Muell.) Aporum sect. Crumenata (Pfitzer) Brieger in Schltr., Orchideen, ed. 3, 1, 671 (1981). Type species: Aporum crumenatum (Sw.) Brieger (basionym: Dendrobium crumenatum Sw.) Aporum sect. Linearifolia Brieger in Schltr., Orchideen, ed. 3, 1, 674 (1981). Type species: Aporum gracile (Blume) Brieger (basionym: Onychium gracile Blume) Aporum sect. Macrostomium (Blume) Brieger in Schltr., Orchideen, ed. 3, 1, 676 (1981). Type species: Aporum aloifolium (Blume) Brieger (basionym: Macrostomium aloifolium Blume) Aporum sect. Strongyle (Lindl.) Brieger in Schltr., Orchideen, ed. 3, 1, 674 (1981). Type species: Aporum subulatum (Blume) Rauschert (basionym: Onychium subulatum Blume) Taxa in other ranks with their type species in D. sect. Aporum: Ceraia Lour., Fl. Cochinch., 2, 518 (1790). Type species: Ceraia simplicissima Lour.

DENDROBIUM

Aporum (Blume) Blume, Bijdr., 334 (1825). Type species: Aporum lobatum Blume Macrostomium (Blume) Blume, Bijdr., 335 (1825). Type species: Macrostomium aloifolium Blume Schismoceras Presl, Rel. Haenk., 1, 96, t. 13, fig. 2 (1827). Type species: Schismoceras disticha Presl Orthoglottis Breda, Gen. Sp. Orchid. Asclep., fasc. 4, (1830) s. n. Type species: Orthoglottis imbricata Breda Ditulima Raf., Fl. Tellur., 4, 41 (1836, publ. 1838). Type species: Ditulima anceps (Sw.) Raf. (basionym: Dendrobium anceps Sw.) Dendrobium subgen. Aporum (Blume) Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21, 201 (1910). Type species: Dendrobium lobatum (Blume) Miq. (basionym: Aporum lobatum Blume) Dendrobium. subgen. Bolbodium (Lindl.) Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21, 26 (1910). Type species: Dendrobium pusillum (Blume) Lindl., nom. illeg., non D.Don (basionym: Desmotrichum pusillum Blume) Dendrobium subgen. Crumenata (Pfitzer) Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21, 224 (1910). Type species: Dendrobium crumenatum Sw. Dendrobium subgen. Strongyle (Lindl.) Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21, 220 (1910). Type species: Dendrobium subulatum (Blume) Lindl. (basionym: Onychium subulatum Blume) Dendrobium subgen. Xerobium Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 441 (1912). Type species: Dendrobium lobatum (Blume) Miq. (basionym: Aporum lobatum Blume) Dendrobium subgen. Rhopalobium Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 441 (1912). Type species: Dendrobium crumenatum Sw. Dendrobium sect. Rhopalanthe subsect. Aporopsis Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 565 (1912). Type species: Dendrobium macfarlanei F.Muell. Bolbodium (Lindl.) Brieger in Schltr., Orchideen, ed. 3, 1, 721 (1981), as ‘Bolbidium’ (not Bolbidium (Lindl.) Lindl.). Type species: Bolbodium pusillum (Blume) Rauschert (basionym: Desmotrichum pusillum Blume) Aporopsis (Schltr.) M.A.Clem. & D.L.Jones, Orchadian, 13, 485 (2002). Type species: Dendrobium macfarlanei F.Muell. Stems clustered or on short rhizomes, of three main types: 1. Cane-like, with many internodes of which a few of the lower ones abruptly swollen and leafless, the remainder wiry and leafy, or with a leafless top part (Crumenata group); 2. As type 1, but without swollen internodes (Aporum group); 3. Short, clavate, with only one prominent internode and two, non-sheathing leaves at apex (Bolbodium group). Stems of type 2 may continue to grow for several years. Leaves long-lived, attached to a tubular sheath, except in the Bolbodium group; laterally flattened or terete to needle-shaped in the Aporum and part of the Crumenata group, dorsiventrally flattened in the Bolbodium and part of the Crumenata group. Inflorescences one-flowered, appearing laterally or apically from leafy stems, sometimes on a leafless, apical part of stem, which may resemble a racemose inflorescence. Flowers often thin-textured and ephemeral (in all three groups), or more fleshy and longer-lived (in part of the Aporum group); mentum usually long, sometimes spur-like. Labellum attached to the apex of column foot, often adnate to it apically; entire or distinctly

lobed. Rostellum usually with a soft but discrete viscidium. About 190 species, distributed largely as for the genus, but not in Japan or New Zealand.

Dendrobium sect. Biloba J.J.Sm., Orchid. Ambon, 67 (1905). Type species: Dendrobium bilobum Lindl. Dendrobium sect. Monanthos Schltr. in K.Schum. & Lauterb., Nachtr. Fl. Deutsch. Schutzgeb. Südsee, 150 (1905). Type species: Dendrobium bilobum Lindl. Taxa in other ranks with their type species in D. sect. Biloba: Monanthos (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 660 (1981), as ‘Monanthus’. Type species: Monanthos bilobus (Schltr.) Brieger (basionym: Dendrobium bilobum Lindl.) Stems clustered, cane-like, numerous, with many internodes, wiry and leafy, not branching, with apical growth continuing for several seasons. Leaves long-lived, attached to a tubular sheath. Inflorescences one-flowered, appearing laterally from leafy stems. Flowers non-resupinate, not ephemeral, usually dull-coloured; mentum usually short, chin-like. Labellum flexibly attached to apex of column foot, entire or rather weakly trilobed, often with a single low, bar-like, glossy callus in basal part; midlobe often fleshy. About 35 species, from Sulawesi, the Moluccas, New Guinea, the Solomon Islands, Fiji, and Australia, with most species in New Guinea.

Dendrobium sect. Brevisaccata Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21, 158 (1910). Type species: Dendrobium agrostophyllum F.Muell. Dendrobium sect. Trachyrhizum Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 447 (1912). Type species: Dendrobium trachyrhizum Schltr. Taxa in other ranks with their type species in D. sect. Brevisaccata: Trachyrhizum (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 687 (1981). Type species: Trachyrhizum schlechteri Brieger (basionym: Dendrobium trachyrhizum Schltr.). Roots usually verrucose. Stems clustered, cane-like, fleshy, manyleaved, not branching, apical growth continuing for several seasons. Leaves with a basal tubular sheath. Inflorescences usually elongate, horizontal or arching, few-flowered, sometimes forming a small sympodium of successive, one-flowered branches (D. viridiflorum F.M.Bailey); mentum large. Labellum attached to apex of column foot, or sometimes adnate to it, subentire or distinctly trilobed, fleshy, with a prominent, backward-pointing callus at the base, which is usually connected to a pair of high lamellate keels basally, sometimes labellum without keels but with a swollen median band. About ten species, from the Moluccas to New Guinea, the Solomon Islands, and Fiji, with one species in Australia. New Guinea is clearly the centre of diversity for this section.

Dendrobium sect. Cadetia (Gaudich.) Benth. & Hook.f., Gen. Pl., 3, 499 (1883). Type species: Dendrobium umbellatum 63

DENDROBIEAE

(Gaudich.) Rchb.f. (basionym: Cadetia umbellata Gaudich.) Dendrobium sect. Pseudocadetia Rchb.f. in Ann. Bot. Syst. (Walpers), 3, 533 (1853), nom. invalid. Type species: Dendrobium funiforme Blume Cadetia sect. Eu-Cadetia Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 425 (1912), nom. illeg. Type species: Cadetia umbellata Gaudich. Cadetia sect. Pterocadetia Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 424 (1912). Type species: Cadetia trigonocarpa Schltr. Cadetia sect. Sarcocadetia Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 424 (1912). Type species: Cadetia funiformis (Blume) Schltr. (basionym: Dendrobium funiforme Blume) Taxa in other ranks with their type species in D. sect. Cadetia: Cadetia Gaudich. in Freyc., Voy. Bol., 422, t. 33 (1826). Type species: Cadetia umbellata Gaudich. Dendrobium subgen. Cadetia (Gaudich.) P.Royen, Alp. Fl. New Guinea, 2, 285 (1979). Type species: Dendrobium umbellatum (Gaudich.) Rchb.f. (basionym: Cadetia umbellata Gaudich.) Sarcocadetia (Schltr.) M.A.Clem. & D.L.Jones, Orchadian, 13, 490 (2002). Type species: Sarcocadetia funiformis (Blume) M.A.Clem. & D.L.Jones (basionym: Dendrobium funiforme Blume) Stems or pseudobulbs usually clustered, or along an elongate creeping or erect rhizome, terete or four-angled, of one internode, one-leaved at apex. Leaves not sheathing at base. Inflorescences one-flowered, from apex of pseudobulb, or laterally just below apex, or both, sometimes from a whitish spathe. Flowers usually white; mentum often spur-like. Labellum usually adnate to column foot, entire or trilobed, without keels basally but sometimes with transverse calli near middle, or with keels or calli on midlobe; glabrous or pubescent. Column pubescent in front. Ovary glabrous or with hair-like papillae, terete or three-winged. About 60 species, from Java, Borneo, Sulawesi, the Philippines, New Guinea, Australia, the Solomon Islands, and Vanuatu, with a clear centre of diversity in New Guinea. Dendrobium sect. Calyptrochilus Schltr. in K.Schum. & Lauterb., Nachtr. Fl. Deutsch. Schutzgeb. Südsee, 150 (1905). Type species: Dendrobium lawesii F.Muell. Dendrobium sect. Capitata Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21, 130 (1910). Type species: Dendrobium purpureum Roxb. Dendrobium sect. Cuthbertsonia Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 445 (1912). Type species: Dendrobium cuthbertsonii F.Muell. Dendrobium sect. Dolichocentrum Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 451 (1912). Type species: Dendrobium furcatum Schltr. Dendrobium sect. Glomerata Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21, 101 (1910). Type species: Dendrobium glomeratum Rolfe Dendrobium sect. Oxyglossum Schltr. in K.Schum. & Lauterb., Nachtr. Fl. Deutsch. Schutzgeb. Südsee, 149 (1905). Type species: Dendrobium cyanocentrum Schltr. 64

Pedilonum sect. Calyptrochilus (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 681 (1981). Type species: Pedilonum lawesii (F.Muell.) Rauschert (basionym: Dendrobium lawesii F.Muell.) Pedilonum sect. Cuthbertsonia (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 682 (1981). Type species: Pedilonum cuthbertsonii (F.Muell.) Brieger (basionym: Dendrobium cuthbertsonii F.Muell.) Pedilonum sect. Oxyglossum (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 684 (1981). Type species: Pedilonum cyanocentrum (Schltr.) Rauschert (basionym: Dendrobium cyanocentrum Schltr.) Taxa in other ranks with their type species in D. sect. Calyptrochilus: Coelandria Fitzg., Austral. Orchids, 1, 7, t. 2 (1882). Type species: Coelandria smillieae (F.Muell.) Fitzg. (basionym: Dendrobium smillieae F.Muell.) Dendrobium sect. Eudendrobium subsect. Pycnostachyae Benth. & Hook.f., Gen. Pl., 3, 500 (1883). Type species: Dendrobium purpureum Roxb. Dendrobium sect. Glomerata subsect. Mesocentra Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21, 102 (1910). Type species: Dendrobium glomeratum Rolfe Dolichocentrum (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 659 (1981). Type species: Dolichocentrum furcatum (Schltr.) Brieger (basionym: Dendrobium furcatum Schltr.) Chromatotriccum M.A.Clem. & D.L.Jones, Orchadian, 13, 493 (2002). Type species: Chromatotriccum lawesii (F.Muell.) M.A.Clem. & D.L.Jones (basionym: Dendrobium lawesii F.Muell.) Maccraithea M.A.Clem. & D.L.Jones, Orchadian, 13, 496 (2002). Type species: Maccraithea cuthbertsonii (F.Muell.) M.A.Clem. & D.L.Jones (basionym: Dendrobium cuthbertsonii F.Muell.) Oxyglossellum M.A.Clem. & D.L.Jones, Orchadian, 13, 490 (2002). Type species: Oxyglossellum cyanocentrum (Schltr.) M.A.Clem. & D.L.Jones (basionym: Dendrobium cyanocentrum Schltr.) Roots glabrous. Stems or pseudobulbs highly variable: often clustered, cane-like, fleshy or wiry, usually many-leaved, branching or not; less often with distinct globose to fusiform, one- to fewleaved pseudobulbs. Leaves with a tubular sheath at base (except in some species with distinct pseudobulbs), short-lived or less often longer-lived. Inflorescences one- to many-flowered, appearing laterally or apically from leafy or leafless stems. Flowers often brilliantly coloured; mentum usually elongate, often spur-like. Labellum attached to apex of column foot or usually adnate to it, entire or weakly lobed, often linear and with a V-shaped callus basally, otherwise without keels or calli; apex of labellum sometimes inflexed and with denticulate margin. Rostellum usually with a soft but discrete viscidium. About 140 species, from Thailand, through the Malay Archipelago and the Philippines to Fiji, one species in Australia. Few species occur west of Sulawesi; by far the greatest diversity is in New Guinea. Dendrobium sect. Conostalix Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21, 160 (1910). Type species: Dendrobium conostalix Rchb.f. (basionym: Dendrobium calcaratum Lindl., nom. illeg., non A.Rich.)

DENDROBIUM

Taxa in other ranks with their type species in D. sect. Conostalix: Conostalix (Kränzl.) Brieger in Schltr., Orchideen, ed. 3, 1, 659 (1981). Type species: Dendrobium conostalix Rchb.f. (basionym: Dendrobium calcaratum Lindl., nom. illeg., non A.Rich.) Stems clustered, cane-like, with many internodes, wiry or fleshy, leafy, branching or not, with or without apical growth continuing for several seasons. Leaves long-lived, attached to a tubular sheath. Leaf sheaths, when immature, covered with dark brown or blackish hairs, although these are apparently lacking in certain Bornean species. Inflorescences one- to few-flowered, appearing laterally from leafy stems. Flowers usually non-resupinate, not ephemeral, dull-coloured; mentum long. Labellum adnate to column foot, trilobed, fleshy. About 20 species from Burma, Thailand, Laos, Vietnam, Malaysia, Indonesia, the Philippines, New Guinea, Australia, and the Solomon Islands. Dendrobium sect. Crinifera Pfitzer in Engl. & Prantl, Nat. Pflanzenfam., II, 6, 175 (1889). Type species: Dendrobium criniferum Lindl. Dendrobium sect. Desmotrichum Blume, Tab. Pl. Jav. Orchid., in Clavis generum (1825), nom. rej. Type species: Dendrobium angulatum (Blume) Lindl., nom. illeg., non Lindl. (basionym: Desmotrichum angulatum Blume) Flickingeria sect. Plicatiles Seidenf., Dansk Bot. Ark., 34, 1, 16 (1980). Type species: Flickingeria fimbriata (Blume) A.D.Hawkes (basionym: Desmotrichum fimbriatum Blume) Flickingeria sect. Bilobulatae Seidenf., Dansk Bot. Ark., 34, 1, 17 (1980). Type species: Flickingeria angustifolia (Blume) A.D.Hawkes (basionym: Desmotrichum angustifolium Blume) Taxa in other ranks with their type species in D. sect. Crinifera: Desmotrichum (Blume) Blume, Bijdr., 329 (1825), nom. rej., not Desmotrichum Kütz. Type species: Desmotrichum angulatum Blume Flickingeria A.D.Hawkes, Orquidea, 27, 301 (1961). Type species: Flickingeria angulata (Blume) A.D.Hawkes (basionym: Desmotrichum angulatum Blume) Ephemerantha P.F.Hunt & Summerh., Taxon, 10, 102 (1961). Type species: Ephemerantha angulata (Blume) P.F.Hunt & Summerh. (basionym: Desmotrichum angulatum Blume) Abaxianthus M.A.Clem. & D.L.Jones, Orchadian, 13, 485 (2002). Type species: Abaxianthus convexus (Blume) M.A.Clem. & D.L.Jones (basionym: Desmotrichum convexum Blume) Rhizome elongate, creeping and rooting along its length or more frequently erect and shrubby and rooting mainly in the basal part, sometimes thin and pendulous. Pseudobulbs distant, usually of a few internodes of which only the upper one is swollen, usually laterally flattened, sometimes highly reduced and slightly swollen, one-leaved at apex. Leaves not sheathing at base. Inflorescences one-flowered, from apex of pseudobulb, or laterally just below apex, or both. Flowers ephemeral; mentum well-developed but not spur-like. Labellum flexibly attached to apex of column foot, sometimes adnate to column foot, usually distinctly trilobed, usually with lamellate and often wavy keels. About 70 species, distributed as for the genus, east to Samoa, absent from Japan

and New Zealand, with centres of diversity in Sumatra, Borneo, and Sulawesi. Dendrobium sect. Dendrobium Dendrobium sect. Onychium Blume, Tab. Pl. Jav. Orchid., in Clavis generum (1825). Type species: Dendrobium japonicum (Blume) Lindl. (basionym: Onychium japonicum Blume) Dendrobium sect. Eudendrobium Lindl., Paxton’s Fl. Gard., 1, 134 (1851), nom. illeg. Type species: Dendrobium moniliforme (L.) Sw. (basionym: Epidendrum moniliforme L.) Dendrobium sect. Holochrysa Lindl., J. Linn. Soc. Bot., 3, 7 (1859). Type species: Dendrobium moschatum (Buch.-Ham.) Sw. (basionym: Epidendrum moschatum Buch.-Ham.) Dendrobium sect. Chrysantha Rchb.f. in Ann. Bot. Syst. (Walpers), 6, 287 (1861), nom. invalid. Type species: Dendrobium chrysanthum Lindl. Dendrobium sect. Planifolia Rchb.f. in Ann. Bot. Syst. (Walpers), 6, 282 (1861). Type species: Dendrobium nobile Lindl. Dendrobium sect. Breviflores Hook.f., Fl. Brit. Ind., 5, 711 (1890). Type species: Dendrobium breviflorum Lindl. Dendrobium sect. Densiflora Finet, Bull. Mus. Hist. Nat. (Paris), 9, 205 (1903). Type species: Dendrobium densiflorum Lindl. Dendrobium sect. Aquea Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21, 51 (1910). Type species: Dendrobium aqueum Lindl. Dendrobium sect. Aurea Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21, 40 (1910). Type species: Dendrobium aureum Lindl. Dendrobium sect. Chrysotoxa Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21, 62 (1910). Type species: Dendrobium chrysotoxum Lindl. Dendrobium sect. Herbacea Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21, 71 (1910). Type species: Dendrobium herbaceum Lindl. Dendrobium sect. Macrostachya Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21, 57 (1910). Type species: Dendrobium macrostachyum Lindl. Dendrobium sect. Nobilia (Pfitzer) Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21, 27 (1910). Type species: Dendrobium nobile Lindl. Dendrobium sect. Stuposa Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21, 55 (1910). Type species: Dendrobium stuposum Lindl. Dendrobium sect. Goldschmidtia Dammer, Orchis, 4, 86 (1910), nom. illeg. Type species: Dendrobium vexans Dammer Dendrobium sect. Brachyanthe Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 446 (1912). Type species: Dendrobium bicameratum Lindl. Dendrobium sect. Callista (Lour.) Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 444 (1912). Type species: Dendrobium amabile (Lour.) O’Brien (basionym: Callista amabilis Lour.) Dendrobium sect. Eugenanthe Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 445 (1912). Type species: Dendrobium nobile Lindl. Dendrobium sect. Actinomorpha Pradhan, Indian Orchids: Guide Ident. Cult., 2, 301 (1979). Type species: Dendrobium lawanum Lindl. Taxa in other ranks with their type species in D. sect. Dendrobium: Callista Lour., Fl. Cochinch., 2, 519 (1790). Type species: Callista amabilis Lour. 65

DENDROBIEAE

Onychium (Blume) Blume, Bijdr., 323 (1825). Type species: Onychium japonicum Blume Endeisa Raf., Fl. Tellur., 2, 51 (1836, publ. 1837). Type species: Endeisa flava Raf., nom. illeg. (basionym: Dendrobium densiflorum Wall. ex Lindl.) Ormostema Raf., Fl. Tellur., 4, 38 (1836, publ. 1838). Type species: Ormostema purpurea Raf., nom. illeg. (basionym: Epidendrum moniliforme L.) Pierardia Raf., Fl. Tellur., 4, 41 (1836, publ. 1838). Type species: Pierardia bicolor Raf., nom. illeg. (basionym: Dendrobium pierardii Roxb.) Dendrobium sect. Eudendrobium subsect. Chrysantha Rchb.f. in Ann. Bot. Syst. (Walpers), 3, 533 (1853), nom. invalid. Type species: Dendrobium chrysanthum Lindl. Dendrobium sect. Eudendrobium subsect. Grandia Rchb.f. in Ann. Bot. Syst. (Walpers), 3, 532 (1853), nom. invalid. Type species: Dendrobium anosmum Lindl. Dendrobium sect. Eudendrobium subsect. Transparentia Rchb.f. in Ann. Bot. Syst. (Walpers), 3, 533 (1853), nom. invalid. Type species: Dendrobium transparens Lindl. Dendrobium sect. Eudendrobium subsect. Calvae Lindl., J. Proc. Linn. Soc. Bot., 3, 14 (1859). Type species: Dendrobium breviflorum Lindl. Dendrobium sect. Stachyobium subsect. Aureum Rchb.f., Gard. Chron., n.s., 10, 300 (1878), nom. illeg., as ‘Stachyobium aureum’. Type species: Dendrobium binoculare Rchb.f. Dendrobium sect. Eudendrobium subsect. Calostachyae Benth. & Hook.f., Gen. Pl., 3, 500 (1883). Type species: Dendrobium pulchellum Roxb. ex Lindl. Dendrobium sect. Eudendrobium subsect. Fasciculata Benth. & Hook.f., Gen. Pl., 3, 501 (1883). Type species: Dendrobium moniliforme (L.) Sw. (basionym: Epidendrum moniliforme L.) Dendrobium sect. Stachyobium subsect. Ramosissimae Benth. & Hook.f., Gen. Pl., 3, 500 (1883). Type species: Dendrobium ramosissimum Wight Dendrobium. sect. Bolbodium subsect. Indivisa Pfitzer in Engl. & Prantl, Nat. Pflanzenfam., II, 6, 175 (1889). Type species: Dendrobium jenkinsii Lindl. Dendrobium sect. Eudendrobium subsect. Calceolaria Pfitzer in Engl. & Prantl, Nat. Pflanzenfam., II, 6, 173 (1889). Type species: Dendrobium moschatum (Buch.-Ham.) Sw. (basionym: Epidendrum moschatum Buch.-Ham.) Dendrobium sect. Eudendrobium subsect. Chrysostachya Pfitzer in Engl. & Prantl, Nat. Pflanzenfam., II, 6, 173 (1889). Type species: Dendrobium fimbriatum Hook. Dendrobium sect. Eudendrobium subsect. Chrysantha Rchb.f ex Pfitzer in Engl. & Prantl, Nat. Pflanzenfam., II, 6, 173 (1889). Type species: Dendrobium chrysanthum Lindl. Dendrobium sect. Eudendrobium subsect. Nobilia Pfitzer in Engl. & Prantl, Nat. Pflanzenfam., II, 6, 174 (1889). Type species: Dendrobium nobile Lindl. Dendrobium sect. Eudendrobium subsect. Rhodostachya Pfitzer in Engl. & Prantl, Nat. Pflanzenfam., II, 6, 173 (1889). Type species: Dendrobium dalhousieanum Wall. Dendrobium sect. Eudendrobium subsect. Trilobata Pfitzer in Engl. & Prantl, Nat. Pflanzenfam., II, 6, 174 (1889). Type species: Dendrobium maccarthiae Thwaites 66

Dendrobium subgen. Eu-Dendrobium Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 220, nom. illeg. Type species: Dendrobium nobile Lindl. Dendrobium sect. Aurea subsect. Binocularia Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 40. Type species: Dendrobium binoculare Rchb.f. Dendrobium sect. Aurea subsect. Moniliformia Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 41, nom. invalid. Type species: Dendrobium moniliforme (L.) Sw. (basionym: Epidendrum moniliforme L.) Dendrobium sect. Aurea subsect. Percnochila Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 40. Type species: Dendrobium heterocarpum Wall. ex Lindl. Dendrobium sect. Aurea subsect. Subcylindracea Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 41. Type species: Dendrobium chrysanthum Lindl. Dendrobium sect. Glomerata subsect. Brachycentra Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 101. Type species: Dendrobium minahassae Kränzl. Dendrobium sect. Nobilia subsect. Fimbrilabia Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 27. Type species: Dendrobium devonianum Paxton Dendrobium sect. Nobilia subsect. Integrilabia Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 27. Type species: Dendrobium nobile Lindl. Goldschmidtia Dammer, Orchis, 4, 86 (1910), nom. invalid. Type species: Goldschmidtia gracilis Dammer Dendrobium subgen. Eugenanthe P.Royen, Alp. Fl. New Guinea, 2, 325 (1979). Type species: Dendrobium nobile Lindl. Roots glabrous. Stems or pseudobulbs usually cane-like, fleshy or rarely wiry, usually many-leaved, branching or not, less often with distinct, ovoid to fusiform, one- to few-leaved pseudobulbs. Leaves usually short-lived, with or without a tubular sheath at the base. Inflorescences usually short, pendulous when elongate, one- to many-flowered, appearing laterally or apically from leafy or leafless stems, most often from the leafless stem of the previous year. Flowers often showy; mentum short, chin-like or less often narrowly conical. Labellum (usually rigidly) attached to apex of column foot, not adnate to it, entire or weakly lobed, adaxially often darker coloured in the centre, often densely and finely pubescent, often with finely fringed margins often with a linear, basal claw, without multiple keels but sometimes with a single callus or with a single longitudinal keel. About 100 species, almost throughout the range of the genus, including Australia, but not in the Pacific Islands east of New Guinea, and with a clear centre of diversity in continental Asia. Dendrobium sect. Dendrocoryne Lindl., Edwards’s Bot. Reg., 30, sub t. 53 (1844). Type species: Dendrobium tetragonum A.Cunn. ex Lindl. Dendrobium sect. Speciosa Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 269. Type species: Dendrobium speciosum Sm. Taxa in other ranks with their type species in D. sect. Dendrocoryne: Thelychiton Endl., Prodr. Fl. Norf., 32 (1833). Type species: Thelychiton macropus Endl.

DENDROBIUM

Tropilis Raf., Fl. Tellur., 2, 95 (1836, publ. 1837). Type species: Tropilis aemula (R.Br.) Raf., as ‘emulum’ (basionym: Dendrobium aemulum R.Br.) Dendrobium sect. Stachyobium subsect. Speciosae Benth. & Hook.f., Gen. Pl., 3, 500 (1883). Type species: Dendrobium speciosum Sm. Dendrobium subgen. Dendrocoryne (Lindl.) Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 26. Type species: Dendrobium tetragonum A.Cunn. ex Lindl. Tetrabaculum M.A.Clem. & D.L.Jones, Orchadian, 13, 490 (2002). Type species: Tetrabaculum tetragonum (A.Cunn.) M.A.Clem. & D.L.Jones (basionym: Dendrobium tetragonum A.Cunn.) Stems or pseudobulbs clustered, cane-like or fusiform to clavate, fleshy, with 2­–7 leaves apically, not branching (sometimes forming keikis), with limited apical growth. Leaves without a tubular sheath at base. Inflorescences elongate, erect or arching, few- to many-flowered. Flowers variable in size, if large then with narrow sepals and petals, thin-textured, lasting from several days to a few weeks, often fragrant; mentum large but not spur-like. Labellum flexibly attached to apex of column foot, distinctly trilobed, usually with 1–3 low and narrow keels, sometimes with a raised and ribbed callus. Rostellum not exuding a white liquid when damaged. About 12 species from Australia, Lord Howe Island, Norfolk Island, Vanuatu, New Caledonia, Fiji, and Samoa. Only three species occur outside mainland Australia. Dendrobium sect. Diplocaulobium Rchb.f., J. Linn. Soc., 15, 112 (1876). Type species: Dendrobium nitidissimum Rchb.f. Dendrobium sect. Goniobulbon Schltr. in K.Schum. & Lauterb., Nachtr. Fl. Deutsch. Schutzgeb. Südsee, 149 (1905). Type species: Dendrobium chrysotropis Schltr. Dendrobium sect. Mekynosepalum Schltr. in K.Schum. & Lauterb., Nachtr. Fl. Deutsch. Schutzgeb. Südsee, 149, (1905). Type species: Dendrobium mekynosepalum Schltr. Dendrobium sect. Longicollia J.J.Sm., Orchid. Ambon, 39 (1905), nom. invalid. Type species: Dendrobium longicolle Lindl. Diplocaulobium sect. Goniobulbon (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 735 (1981), nom. illeg. Type species: Diplocaulobium chrysotropis (Schltr.) A.D.Hawkes (basionym: Dendrobium chrysotropis Schltr.) Taxa in other ranks with their type species in D. sect. Diplocaulobium: Diplocaulobium (Rchb.f.) Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21, 331 (1910). Type species: Diplocaulobium nitidissimum (Rchb.f.) Kränzl. (basionym: Dendrobium nitidissimum Rchb.f.) Pseudobulbs usually clustered or spaced along an elongate, creeping or pendulous rhizome, of one internode, often attenuated apically, one-leaved at apex. Leaves not sheathing at base. Inflorescences one-flowered, often in fascicles, from apex of pseudobulb, arising from a bilaterally flattened, long-persistent, yellow spathe. Flowers often white, ephemeral or rarely lasting a few days, mentum chin-like. Labellum flexibly attached to apex of column foot, entire or trilobed, with lamellate keels at least basally. About 85 species, from Peninsular Malaysia, Singapore, Sumatra, Borneo,

Sulawesi, Moluccas, New Guinea, the Philippines, Australia, the Solomon Islands, the Carolines, New Caledonia, Fiji and Samoa, with a clear centre of diversity in New Guinea and only two species west of Sulawesi. Dendrobium sect. Distichophyllae Hook.f., Fl. Brit. Ind., 5, 711 (1890). Type species: Dendrobium revolutum Lindl. Dendrobium sect. Distichophyllum Schltr., Bot. Jahrb. Syst., 39, 68 (1906). Type species: Dendrobium revolutum Lindl. Dendrobium sect. Revoluta Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 176. Type species: Dendrobium revolutum Lindl. Taxa in other ranks with their type species in D. sect. Distichophyllae: Distichorchis M.A.Clem. & D.L.Jones, Orchadian, 13, 487 (2002). Type species: Distichorchis revoluta (Lindl.) M.A.Clem. & D.L.Jones (basionym: Dendrobium revolutum Lindl.) Stems clustered, cane-like, with many internodes, fleshy, often longitudinally grooved, leafy, usually with many equal-sized and regularly arranged leaves, not branching, with or without apical growth continuing for several seasons. Leaves usually long-lived, attached to a tubular sheath; leaf sheaths glabrous. Inflorescences one- to few-flowered, appearing laterally from stems. Flowers resupinate, often white or with combinations of green and brown, mentum often spur-like. Labellum adnate to column foot, usually trilobed, fleshy, with low longitudinal ridges, or with prominent papillae or callosities. About 35 species from Burma, Thailand, Cambodia, Laos, Vietnam, Peninsular Malaysia, Sumatra, Java, Borneo, New Guinea, the Philippines, the Solomon Islands, Vanuatu, and New Caledonia, with a centre of diversity in Borneo. Dendrobium sect. Eleutheroglossum Schltr., Bot. Jahrb. Syst., 39, 67 (1906). Type species: Dendrobium eleutheroglossum Schltr. Taxa in other ranks with their type species in D. sect. Eleutheroglossum: Eleutheroglossum (Schltr.) M.A.Clem. & D.L.Jones, Orchadian, 13, 489 (2002). Type species: Dendrobium eleutheroglossum Schltr. Stems or pseudobulbs clustered, cane-like or clavate, fleshy, with 2–5 leaves apically, not branching, with limited apical growth. Leaves with a tubular sheath at base, which may be short on the upper leaves. Inflorescences elongate, erect, few-flowered. Flowers with the mentum relatively large but not spur-like. Lateral sepals usually reflexed. Labellum flexibly attached to apex of column foot, distinctly trilobed, often with two lamellate keels; midlobe bilobulate. Four species, one in Australia and three in New Caledonia. Dendrobium sect. Formosae (Benth. & Hook.f.) Hook.f., Fl. Brit. Ind., 5, 710 (1890). Type species: Dendrobium formosum Lindl. Dendrobium sect. Eudendrobium subsect. Formosae Benth. & Hook.f., Gen. Pl., 3, 500 (1883). Type species: Dendrobium formosum Lindl. 67

DENDROBIEAE

Dendrobium sect. Nigrohirsutae (Lindl.) Brieger in Schltr., Orchideen, ed. 3, 1, 681 (1981), nom. illeg., as ‘Nigrohirsuta’. Type species: Dendrobium formosum Lindl. Dendrobium sect. Oxygenianthe Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 448 (1912). Type species: Dendrobium formosum Lindl. Taxa in other ranks with their type species in D. sect. Formosae: Froscula Raf., Fl. Tellur., 4, 44 (1836, publ. 1838). Type species: Froscula hispida Raf., nom. illeg. (basionym: Dendrobium longicornu Lindl.) Dendrobium sect. Eudendrobium subsect. Nigrohirsutae Lindl., J. Proc. Linn. Soc., Bot., 3, 16 (1859). Type species: Dendrobium formosum Lindl. Dendrobium subgen. Nigrohirsutae (Lindl.) Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 26, as ’Nigro-hirsuta’. Type species: Dendrobium formosum Lindl. Stems clustered, cane-like or less often spindle-shaped, with several to many internodes, fleshy, with few to many leaves usually mainly apically, not branching, of limited apical growth. Leaves usually long-lived, attached to a tubular sheath; leaf sheaths densely covered with short, blackish hairs at least when developing, glabrous in a few species of uncertain position. Inflorescences one- to few-flowered, appearing in the upper part from stems. Flowers showy, usually white with orange or red markings on the lip, rarely non-resupinate; mentum often spur-like. Labellum adnate to column foot, usually indistinctly lobed, fleshy, with low longitudinal ridges, irregular callosities or with prominent hair-like outgrowths. About 50 species from India, Nepal, Bhutan, Bangladesh, Burma, Thailand, China, Laos, Cambodia, Vietnam, Peninsular Malaysia, Sumatra, Borneo, Sulawesi, and the Philippines, with centres of diversity in continental southeast Asia and Borneo. Dendrobium sect. Fugacia J.J.Sm., Orchid. Java, 343, (1905). Type species: Dendrobium spurium (Blume) J.J.Sm. (basionym: Dendrocolla spuria Blume) Dendrobium sect. Euphlebium Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 242, as ‘Euphlebia’, 263, as ‘Euphlebium’. Type species: Dendrobium spurium (Blume) J.J.Sm. (basionym: Dendrocolla spuria Blume) Taxa in other ranks with their type species in D. sect. Fugacia: Gersinia Néraud in Freyc., Voy. Uranie, 27 (1826). Type species: Not designated (no species have as yet been assigned to Gersinia) Euphlebium (Kränzl.) Brieger in Schltr., Orchideen, ed. 3, 1, 722 (1981). Type species: Euphlebium spurium (Blume) Brieger (basionym: Dendrocolla spuria Blume) Pseudobulbs clustered, clavate, often quadrangular in cross-section, fleshy, with two or three leaves apically, not branching, with limited apical growth. Leaves without a tubular sheath at base. Inflorescences one- to few-flowered, arising from a concavity in the stem. Flowers ephemeral, mentum short. Labellum hinged with apex of column foot, mobile, entire, usually with a few projections adaxially. About 12 species from Peninsular Malaysia, 68

Singapore, Sumatra, Java, Lesser Sunda Islands, Borneo, perhaps Sulawesi, Moluccas, New Guinea, and the Philippines, with the greatest diversity in the three last-mentioned areas. The synonymous genus Gersinia has until now wrongly been referred to Bulbophyllum.

Dendrobium sect. Fytchianthe Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 446 (1912). Type species: Dendrobium barbatulum Batem. Taxa in other ranks with their type species in D. sect. Fytchianthe:

Dendrobium sect. Stachyobium subsect. Elatiores Benth. & Hook.f., Gen. Pl., 3, 500 (1883). Type species: Dendrobium barbatulum auct. non Batem.: Wight Dendrobium sect. Eudendrobium subsect. Perulacea Pfitzer in Engl. & Prantl, Nat. Pflanzenfam., II, 6, 174 (1889). Type species: Dendrobium barbatulum Batem. Stems clustered, cane-like, fleshy, leafy throughout, not branching, with limited apical growth. Leaves with a tubular sheath at base, short-lived. Inflorescences elongate, erect or patent, few- to manyflowered, arising from upper part of leafless stems. Flowers with petals broader than sepals; mentum well-developed, narrowly conical or spur-like. Labellum adnate to column foot, trilobed, the midlobe much larger than lateral lobes, with a pubescent area at base of midlobe. Three or four species from India and Burma.

Dendrobium sect. Grastidium Blume, Tab. Pl. Jav. Orchid., in Clavis generum (1825). Type species: Dendrobium salaccense (Blume) Lindl. (basionym: Grastidium salaccense Blume) Dendrobium sect. Bambusifoliae Ridl., Mat. Fl. Malay Penins., 1, 31 (1907). (‘Bambusaefoliae’). Type species: Dendrobium gemellum auct. non Lindl.: Ridl. Dendrobium sect. Dianthe Schltr. in K.Schum. & Lauterb., Nachtr. Fl. Deutsch. Schutzgeb. Südsee, 150, (1905). Type species: Dendrobium gemellum auct. non Lindl.: Ridl. Dendrobium sect. Gemella Ridl. ex Schltr. in K.Schum. & Lauterb., Nachtr. Fl. Deutsch. Schutzgeb. Südsee, 168, (1905), nom. invalid. Type species: Dendrobium gemellum auct. non Lindl.: Ridl. Dendrobium sect. Angustifolia Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 165. Type species: Dendrobium salaccense (Blume) Lindl. (basionym: Grastidium salaccense Blume) Dendrobium sect. Bambusacea Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 165. Type species: Dendrobium salaccense (Blume) Lindl. (basionym: Grastidium salaccense Blume). Dendrobium sect. Eugrastidium Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 188, nom. illeg. Type species: Dendrobium rugosum (Blume) Lindl. (basionym: Grastidium rugosum Blume). Dendrobium sect. Dichopus (Blume) Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 450 (1912). Type species: Dendrobium insigne (Blume) Rchb.f. ex Miq. (basionym: Dichopus insignis Blume).

DENDROBIUM

Dendrobium sect. Eriopexis Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 450 (1912). Type species: Dendrobium eriopexis Schltr. Taxa in other ranks with their type species in D. sect. Grastidium: Dichopus Blume, Mus. Bot. Lugd. Bat., 2, 176 (1856). Type species: Dichopus insignis Blume Grastidium (Blume) Blume, Bijdr., 333, (1825). Type species: Grastidium salaccense Blume Dendrobium sect. Eudendrobium subsect. Foliosae Benth. & Hook.f., Gen. Pl., 3, 501 (1883). Type species: Dendrobium salaccense (Blume) Lindl. (basionym: Grastidium salaccense Blume) Dendrobium subgen. Grastidium (Blume) Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 159. Type species: Dendrobium salaccense (Blume) Lindl. (basionym: Grastidium salaccense Blume) Dendrobium sect. Grastidium subsect. Tridentifera Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 604 (1912). Type species: Dendrobium tridentiferum Lindl. Eriopexis (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 656 (1981). Type species: Eriopexis schlechteri Brieger (basionym: Dendrobium eriopexis Schtr.) Grastidium subgen. Dichopus (Blume) M.A.Clem. & D.L.Jones, Lasianthera, 1, 116 (1997). Type species: Grastidium insigne (Blume) M.A.Clem. & D.L.Jones (basionym: Dichopus insignis Blume) Stems clustered, cane-like, often numerous, with many internodes, wiry and leafy, usually not branching, with apical growth continuing for several seasons, rarely twining. Leaves long-lived, attached to a tubular sheath, flat or rarely terete. Inflorescences one- or usually two-flowered, appearing laterally from stems, at the base subtended by a few laterally flattened, long-persistent bracts. Flowers ephemeral; mentum chin-like. Labellum flexibly attached to apex of column foot, entire or trilobed, usually with a single, low, longitudinal ridge basally, often with warts or transverse ridges adaxially, sometimes with a hairy midlobe. About 180 species, distributed as for the genus, but absent from Japan and New Zealand, with the greatest diversity in New Guinea.

Dendrobium sect. Herpethophytum Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 451 (1912). Type species: Dendrobium herpetophytum Schltr. (Note: the spelling ‘Herpethophytum’ is linguistically incorrect but must be followed.) Taxa in other ranks with their type species in D. sect. Herpethophytum: Herpethophytum (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 660 (1981). Type species: Herpethophytum schlechteri Rauschert (basionym: Dendrobium herpetophytum Schltr.) Stems clustered, with many internodes, wiry and leafy, branched, with apical growth continuing for several seasons. Leaves longlived, attached to a tubular sheath. Inflorescences one-flowered, appearing laterally from stems. Flowers non-resupinate, ephemeral, usually white with purplish markings on the labellum; mentum spur-like. Labellum flexibly attached to apex of column foot,

entire or usually trilobed, usually with a pair of fleshy ridges or a U-shaped callus in the centre. About 18 species, from Sulawesi (one species) and New Guinea. Dendrobium sect. Latouria Miq., Fl. Ned. Indië, 3, 645 (1859). Type species: Dendrobium spectabile (Blume) Miq. (basionym: Latouria spectabilis Blume) Dendrobium sect. Leiotheca Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 250. Type species: Dendrobium spectabile (Blume) Miq. (basionym: Latouria spectabilis Blume) Dendrobium sect. Superbientia Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 257. Type species: Dendrobium johnsoniae F.Muell. Dendrobium sect. Trachytheca Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 243. Type species: Dendrobium macrophyllum A.Rich. Dendrobium sect. Athecebium P.Royen, Alp. Fl. New Guinea, 2, 299 (1979), nom. illeg. Type species: Dendrobium spectabile (Blume) Miq. (basionym: Latouria spectabilis Blume) Taxa in other ranks with their type species in D. sect. Latouria: Latouria Blume, Rumphia, 4, 41, t. 195, 199 (1848), nom. illeg., non (Endl.) Lindl. Type species: Latouria spectabilis Blume Dendrobium sect. Dendrocoryne subsect. Hirsuta Pfitzer in Engl. & Prantl, Nat. Pflanzenfam., II, 6, 174 (1889). Type species: Dendrobium macrophyllum A.Rich. Sayeria Kränzl., Oesterr. Bot. Zeitschr., 44, 298 (1894). Type species: Sayeria paradoxa Kränzl. Dendrobium. subgen. Athecebium Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 441 (1912). Type species: Dendrobium spectabile (Blume) Miq. (basionym: Latouria spectabilis Blume) Latourorchis Brieger in Schltr., Orchideen, ed. 3, 1, 727 (1981). Type species: Latourorchis spectabilis (Blume) Brieger (basionym: Latouria spectabilis Blume) Leioanthum M.A.Clem. & D.L.Jones, Orchadian, 13, 490 (2002). Type species: Leioanthum bifalce (Lindl.) M.A.Clem. & D.L.Jones (basionym: Dendrobium bifalce Lindl.) Stems or pseudobulbs clustered, cane-like or clavate to fusiform or almost spherical, fleshy, with 1–10 leaves apically, not branching, with limited apical growth. Leaves without a tubular sheath at base. Inflorescences elongate, erect or arching, mainly from apical part of stem, one- to many-flowered. Flowers often with broad petals; fleshy, long-lasting, sometimes fragrant; mentum not spur-like. Labellum flexibly attached to apex of column foot, entire or trilobed, usually with high fleshy callosities or ribs. Rostellum exuding a white liquid when damaged. About 58 species from Java, Borneo, Lesser Sunda Islands, Moluccas, New Guinea, Philippines, Australia, Palau Islands, Solomon Islands, Vanuatu, New Caledonia, Fiji, and Samoa; relatively few species occur outside New Guinea, where some 40 species are endemic. Dendrobium sect. Lichenastrum (Brieger) Dockrill, Austral. Indig. Orchids, 2, 681 (1992). Type species: Dendrobium lichenastrum (F.Muell.) Nicholls (basionym: Bulbophyllum lichenastrum F.Muell.) 69

DENDROBIEAE

Dockrillia sect. Lichenastrum Brieger in Schltr., Orchideen, ed. 3, 1, 746 (1981). Type species: Dockrillia lichenastrum (F.Muell.) Brieger (basionym: Bulbophyllum lichenastrum F.Muell.) Taxa in other ranks with their type species in D. sect. Lichenastrum: Davejonesia M.A.Clem., Orchadian, 13, 487 (2002). Type species: Davejonesia lichenastrum (F.Muell.) M.A.Clem. (basionym: Bulbophyllum lichenastrum F.Muell.) Stelbophyllum M.A.Clem. & D.L.Jones, Orchadian, 13, 490 (2002). Type species: Stelbophyllum toressae (F.M.Bailey) M.A.Clem. & D.L.Jones (basionym: Dendrobium toressae F.M.Bailey) Rhizome creeping, mat-forming. Stems one-leaved, highly reduced, not swollen. Leaves not sheathing at base, fleshy, terete or laterally flattened or with a broad furrow. Inflorescences one-flowered. Flower resupinate or not; mentum not spur-like. Labellum flexibly attached to column foot, entire, fleshy, without keels or other ornamentation. Three species, all endemic to Australia. Dendrobium sect. Macrocladium Schltr., Bot. Jahrb. Syst., 39, 68 (1906). Type species: Dendrobium sarcochilus Finet Dendrobium sect. Inobulbum Schltr., Bot. Jahrb. Syst., 39, 67 (1906). Type species: Dendrobium muricatum Finet Dendrobium sect. Kinetochilus Schltr., Bot. Jahrb. Syst., 39, 68 (1906). Type species: Dendrobium pectinatum Finet Eria sect. Tetrodon Kränzl., Vierteljahresschr. Naturf. Ges. Zürich, 74, 89 (1929). Type species: Eria oppositifolia Kränzl. Grastidium sect. Macrocladium (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 653 (1981), nom. invalid. Type species: Grastidium sarcochilus (Finet) Brieger (basionym: Dendrobium sarcochilus Finet) Dendrobium sect. Tetrodon (Kränzl.) Ormerod, Austral. Orchid Rev., 61, 4, 39 (1996). Type species: Dendrobium oppositifolium (Kränzl.) N.Hallé (basionym: Eria oppositifolia Kränzl.) Dendrobium sect. Finetianthe Ormerod, Oasis, 2, 2, 5 (2002). Type species: Dendrobium finetianum Schltr. Dendrobium sect. Dendrobates (M.A.Clem. & D.L.Jones) H.P.Wood, Dendrobiums, 578, (2006). Type species: Dendrobium virotii Guillaumin Dendrobium sect. Winika (M.A.Clem., D.L.Jones & Molloy) H.P.Wood, Dendrobiums, 578, (2006). Type species: Dendrobium cunninghamii Lindl. Taxa in other ranks with their type species in D. sect. Macrocladium: Inobulbum (Schltr.) Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 316, as ‘Inobulbon Schltr. & Kränzl.’; Schlechter (1912) did not accept Inobulbum as a genus and therefore should not be considered co-author. Type species: Inobulbum muricatum (Finet) Kränzl. (basionym: Dendrobium muricatum Finet) Kinetochilus (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 686 (1981). Type species: Kinetochilus pectinatus (Finet) Brieger (basionym: Dendrobium pectinatum Finet) Dendrobium subgen. Inobulbum (Schltr.) N.Hallé, Adansonia n.s., 20, 353 (1981), as ‘Inobulbon’. Type species: Dendrobium muricatum Finet 70

Cannaeorchis M.A.Clem. & D.L.Jones, Lasianthera, 1, 132 (1997). Type species: Cannaeorchis fractiflexa (Finet) M.A.Clem. & D.L.Jones (basionym: Dendrobium fractiflexum Finet) Winika M.A.Clem., D.L.Jones & Molloy, Orchadian, 12, 214 (1997). Type species: Winika cunninghamii (Lindl.) M.A.Clem., D.L.Jones & Molloy (basionym: Dendrobium cunninghamii Lindl.) Tetrodon (Kränzl.) M.A. Clem. & D.L.Jones, Orchadian, 12, 310 (1998). Type species: Tetrodon oppositifolium (Kränzl.) M.A.Clem. & D.L.Jones (basionym: Eria oppositifolia Kränzl.) Bouletia M.A.Clem. & D.L.Jones, Orchadian, 13, 485 (2002). Type species: Bouletia finetiana (Schltr.) M.A.Clem. & D.L.Jones (basionym: Dendrobium finetianum Schltr.) Dendrobates M.A.Clem. & D.L.Jones, Orchadian, 13, 487 (2002). Type species: Dendrobates virotii (Guillaumin) M.A.Clem. & D.L.Jones (basionym: Dendrobium virotii Guillaumin) Roots sometimes verrucose. Stems either clustered, cane-like, with many internodes, leafy, usually not branching, with apical growth continuing for several seasons, or swollen into pseudobulbs, with two leaves at apex. Leaves long-lived, attached to a tubular sheath in species with cane-like stems, without sheaths in species with pseudobulbs. Inflorescences one- to many-flowered, appearing laterally from stems or pseudobulbs, sometimes branching. Flowers with a chin-like mentum. Labellum flexibly attached to apex of column foot, sometimes hinged and mobile, entire or trilobed, usually with three or more basal keels or with callosities at base of midlobe. Fifteen species, with one endemic to New Zealand and all others endemic to New Caledonia. Dendrobium sect. Microphytanthe Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 442 (1912). Type species: Dendrobium bulbophylloides Schltr. Taxa in other ranks with their type species in D. sect. Microphytanthe: Microphytanthe (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 742 (1981). Type species: Microphytanthe bulbophylloides (Schltr.) Brieger (basionym: Dendrobium bulbophylloides Schltr.) Rhizome elongate, creeping. Pseudobulbs well-spaced along rhizome, of one internode, one-leaved. Leaves not sheathing at base. Inflorescences one-flowered, from apex of pseudobulb or laterally just below apex. Flower with mentum not spur-like. Labellum flexibly attached to apex of column foot, entire or weakly lobed, fleshy, without keels but sometimes with transverse calli near middle, glabrous. Three or four species, endemic to New Guinea. Dendrobium sect. Monophyllaea Benth., Fl. Austral., 6, 276 (1873). Type species: Dendrobium monophyllum F.Muell. Dendrobium sect. Australorchis (Brieger) Dockr., Austral. Indig. Orchids, 1, 439 (1982). Type species: Dendrobium monophyllum F.Muell. Taxa in other ranks with their type species in D. sect. Monophyllaea: Australorchis Brieger in Schltr., Orchideen, ed. 3, 1, 741 (1981). Type species: Australorchis monophylla (F.Muell.) Brieger (basionym: Dendrobium monophyllum F.Muell.)

DENDROBIUM

Dendrobium sect. Dendrocoryne subsect. Monophylla Pfitzer in Engl. & Prantl, Nat. Pflanzenfam., II, 6, 174 (1889). Type species: Dendrobium monophyllum F.Muell. Rhizome creeping, with well-spaced pseudobulbs, or with pseudobulbs clustered. Pseudobulbs of one internode, one- or twoleaved at apex. Leaves without a tubular sheath at base. Inflorescences elongate, erect, many-flowered. Flowers thin-textured; mentum not spur-like. Labellum flexibly attached to apex of column foot, distinctly trilobed, with 1–3 low and narrow keels or without keels but with a flat callus apically. Three species, endemic to Australia. Dendrobium sect. Pedilonum Blume, Tab. Pl. Jav. Orchid., in Clavis generum (1825). Type species: Dendrobium secundum (Blume) Lindl. (basionym: Pedilonum secundum Blume) Dendrobium sect. Dendrocoryne Lindl., Edwards’s Bot. Reg., 28, Misc. 76 (1842), nom. invalid., not Lindl. 1844. Type species: Dendrobium compressum Lindl. Dendrobium sect. Platycaulon Schltr. in K.Schum. & Lauterb., Nachtr. Fl. Deutsch. Schutzgeb. Südsee, 150 (1905) (‘Platybulbon’). Type species: Dendrobium lamellatum (Blume) Lindl. (basionym: Onychium lamellatum Blume) Dendrobium sect. Calcarifera J.J.Sm., Bull. Dép. Agric. Indes Néerl., 15, 14 (1908). Type species: Dendrobium pedicellatum J.J.Sm. Dendrobium sect. Anisopetala Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 68. Type species: Dendrobium mutabile (Blume) Lindl. Dendrobium sect. Platycaula Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 266. Type species: Dendrobium platycaulon Rolfe Dendrobium sect. Secunda Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 97. Type species: Dendrobium secundum (Blume) Lindl. (basionym: Pedilonum secundum Blume) Pedilonum sect. Platycaulon (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 685 (1981). Type species: Pedilonum lamellatum (Blume) Brieger (basionym: Onychium lamellatum Blume) Pedilonum sect. Sanguinolenta Brieger in Schltr., Orchideen, ed. 3, 1, 681 (1981). Type species: Pedilonum sanguinolentum (Lindl.) Brieger (basionym: Dendrobium sanguinolentum Lindl.) Taxa in other ranks with their type species in D. sect. Pedilonum: Pedilonum (Blume) Blume, Bijdr., 320 (1825). Type species: Pedilonum secundum Blume Dendrobium sect. Eudendrobium subsect. Pedilonum (Blume) Pfitzer in Engl. & Prantl, Nat. Pflanzenfam., II, 6, 174 (1889). Type species: Dendrobium secundum (Blume) Lindl. (basionym: Pedilonum secundum Blume) Dendrobium subgen. Pedilonum (Blume) Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 96. Type species: Dendrobium secundum (Blume) Lindl. (basionym: Pedilonum secundum Blume) Dendrobium sect. Glomerata subsect. Camptocentra Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 101. Type species: Dendrobium epidendropsis Kränzl. Dendrobium sect. Glomerata subsect. Macrocentra Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 103. Type species: Dendrobium ionopus Rchb.f.

Eurycaulis M.A.Clem. & D.L.Jones, Orchadian, 13, 490 (2002). Type species: Eurycaulis lamellatus (Blume) M.A.Clem. & D.L.Jones (basionym: Onychium lamellatum (Blume) Lindl.) Eurycaulis subgen. Calcariferus (J.J.Sm.) M.A.Clem., Telopea, 10, 285 (2003). Type species: Eurycaulis pedicellatum (J.J.Sm.) M.A.Clem. (basionym: Dendrobium pedicellatum J.J.Sm.) Anisopetala (Kränzl.) M.A.Clem., Telopea, 10, 283 (2003). Type species: Anisopetala mutabilis (Blume) M.A.Clem. (basionym: Dendrobium mutabile Blume) Roots glabrous or verrucose. Stems clustered, cane-like, fleshy, sometimes laterally flattened, usually many-leaved, branching or not. Leaves short-lived, with a tubular sheath at base. Inflorescences one- to many-flowered, appearing laterally or apically from the leafy or leafless stems. Flowers with mentum elongate, often spurlike. Labellum attached to apex of column foot or (less often) adnate to it, entire or weakly lobed, the basal part often linear and with a V-shaped callus, sometimes with indistinct and irregular ridges on midlobe or with a laterally flattened callus at base of midlobe, otherwise without keels or calli. Viscidium soft, discrete. About 125 species, from Sri Lanka through India, Bhutan, Burma, Thailand, Laos, Cambodia, Vietnam, the Malay Archipelago, and the Philippines to Fiji, but not in Australia. The greatest diversity is in Sumatra, Java, the Philippines, and especially in Borneo.

Dendrobium sect. Phalaenanthe Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 447 (1912). Type species: Dendrobium bigibbum Lindl. Taxa in other ranks with their type species in D. sect. Phalaenanthe: Dendrobium sect. Dendrocoryne subsect. Bigibba Pfitzer in Engl. & Prantl, Nat. Pflanzenfam., II, 6, 174 (1889). Type species: Dendrobium bigibbum Lindl. Dendrobium sect. Ceratobium subsect. Platypetala Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 139. Type species: Dendrobium affine (Decne.) Steud. (basionym: Onychium affine Decne.) Vappodes M.A.Clem. & D.L.Jones, Orchadian, 13, 492 (2002). Type species: Vappodes bigibba (Lindl.) M.A.Clem. & D.L.Jones (basionym: Dendrobium bigibbum Lindl.) Stems clustered, cane-like, fleshy, with 3–9 leaves apically, not branching, with limited apical growth. Leaves with a tubular sheath at base. Inflorescences elongate, erect, few- to many-flowered. Flowers showy, white-mauve-purple, infrequently bicoloured, with broad petals, mentum bilobed in lateral view, with an abruptly narrowed, spur-like apical part. Labellum adnate to column foot, forming a narrow conical spur, abruptly widened above spur, trilobed, with low ridges and hair-like papillae basally. Two species from the Lesser Sunda Islands, Moluccas, New Guinea, and Australia.

Dendrobium sect. Pleianthe Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 450 (1912). Type species: Dendrobium pleianthum Schltr. 71

DENDROBIEAE

Grastidium sect. Pleianthe (Schltr.) Brieger in Schltr., Orchideen, ed. 3, 1, 653 (1981), nom. invalid. Type species: Grastidium pleianthum (Schltr.) Brieger (basionym: Dendrobium pleianthum Schltr.) Taxa in other ranks with their type species in D. sect. Pleianthe: Exochanthus M.A.Clem. & D.L.Jones, Orchadian, 13, 496 (2002). Type species: Exochanthus pleianthum (Schltr.) M.A.Clem. & D.L.Jones (basionym: Dendrobium pleianthum Schltr.) Stems clustered, cane-like, with many internodes, not fleshy, leafy, not branching, with apical growth continuing for several seasons. Leaves long-lived, attached to a tubular sheath. Inflorescences manyflowered, appearing laterally from the stems as a dense cluster of flowers piercing leaf-sheath. Flowers with mentum chin-like. Labellum flexibly attached to apex of column foot, trilobed, with a warty callus on midlobe continuing as a low, longitudinal band toward base. Probably only one species, from Sulawesi, New Guinea, and the Solomon Islands. Dendrobium sect. Rhizobium Lindl., Paxton’s Fl. Gard., 1, 136 (1851). Type species: Dendrobium linguiforme Sw. Dendrobium sect. Carnifolia Rchb.f. in Ann. Bot. Syst. (Walpers), 6, 282 (1861). Type species: Dendrobium linguiforme Sw. Dendrobium sect. Strongylophyllum Schltr., Bot. Jahrb. Syst., 39, 67 (1906). Type species: Dendrobium teretifolium R.Br. (here designated) Taxa in other ranks with their type species in D. sect. Rhizobium: Dendrobium subgen. Rhizobium (Lindl.) Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 289. Type species: Dendrobium linguiforme Sw. Dockrillia Brieger in Schltr., Orchideen, ed. 3, 1, 745 (1981). Type species: Dockrillia linguiformis (Sw.) Brieger (basionym: Dendrobium linguiforme Sw.) Rhizome elongate, creeping or pendulous. Stems appearing as branches of rhizome, not swollen, one-leaved. Leaves not sheathing at base, fleshy, terete, laterally flattened or rarely gherkin-shaped or dorsiventrally flattened and less fleshy, without midvein. Inflorescence one-flowered or a few- to many-flowered raceme, arising near apex of upper internode of stem, or sometimes from lower internodes. Flowers non-resupinate, mentum prominent but not spur-like. Labellum flexibly attached to column foot, usually trilobed, with three lamellate keels. About 27 species, from the Aru Islands, New Guinea, Australia, Solomon Islands, possibly Vanuatu, New Caledonia, Fiji, Samoa, and Tahiti, with centres of diversity in New Guinea and Australia.

Dendrobium sect. Sarcopodium Benth. & Hook.f., Gen. Pl., 3, 499 (1883). Type species: Dendrobium amplum Lindl. Epigeneium sect. Katherinea (A.D.Hawkes) Garay & G.Romero, Harvard Pap. Bot., 4, 479 (1999). Type species: Epigeneium amplum (Lindl.) Summerh. (basionym: Sacropodium amplum Lindl.) Epigeneium sect. Racemosum Garay & G.Romero, Harvard Pap. Bot., 4, 479 (1999). Type species: Epigeneium acuminatum 72

Summerh. (basionym: Dendrobium acuminatum Rolfe ex Ames, nom. illeg., non Kunth) Taxa in other ranks with their type species in D. sect. Sarcopodium: Sarcopodium Lindl., Paxt. Flow. Gard., 1, 155 (1850), nom. illeg., non Ehrenb. ex Brogn. Type species: Sarcopodium amplum (Lindl.) Lindl. & Paxton (basionym: Dendrobium amplum Lindl.) Dendrobium sect. Stachyobium subsect. Geminatae Benth. & Hook.f., Gen. Pl., 3, 500 (1883). Type species: Dendrobium geminatum (Blume) Lindl. (basionym: Desmotrichum geminatum Blume) Dendrobium sect. Bolbodium subsect. Triloba Pfitzer in Engl. & Prantl, Nat. Pflanzenfam., II, 6, 175 (1889). Type species: Dendrobium cymbidioides (Blume) Lindl. (basionym: Desmotrichum cymbidioides Blume) Epigeneium Gagnep., Bull. Mus. Hist. Nat. Paris, sér. 2, 4, 593 (1932). Type species: Epigeneium fargesii (Finet) Gagnep. (basionym: Dendrobium fargesii Finet) Katharinea A.D.Hawkes, Lloydia, 19, 94 (1956). Type species: Katharinea ampla (Lindl.) A.D.Hawkes (basionym: Dendrobium amplum Lindl.) Roots glabrous or verrucose. Rhizome elongate, creeping or patent, with well-spaced pseudobulbs. Pseudobulbs of one internode, one- or two-leaved at apex. Leaves without a tubular sheath at base. Inflorescences erect, one- to many-flowered, arising from apex of pseudobulb. Flowers with mentum not spur-like. Labellum flexibly attached to apex of column foot, distinctly trilobed, usually with a few calli or interrupted ridges basally. About 38 species, from India, Nepal, Bhutan, Burma, Thailand, China, Laos, Vietnam, Taiwan, Peninsular Malaysia, Sumatra, Java, Borneo, and the Philippines, with no obvious centre of diversity.

Dendrobium sect. Spatulata Lindl., Hook. London J. Bot., 2, 235 (1843). Type species: Dendrobium antennatum Lindl. Dendrobium sect. Ceratobium Lindl., Paxton’s Fl. Gard., 1, 134 (1851). Type species: Dendrobium veratrifolium Lindl., nom. illeg., non Roxb. Dendrobium sect. Antennata Rchb.f., Gard. Chron., n.s., 25, 266 (1886), nom. illeg. Type species: Dendrobium antennatum Lindl. Dendrobium sect. Strebloceras Schltr. in K.Schum. & Lauterb., Nachtr. Fl. Deutsch. Schutzgeb. Südsee, 149 (1905). Type species: Dendrobium antennatum Lindl. Dendrobium sect. Tokai Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 242. Type species: Dendrobium tokai Rchb.f. Taxa in other ranks with their type species in D. sect. Spatulata: Dendrobium sect. Stachyobium subsect. Antennata Rchb.f., Gard. Chron., n.s., 10, 300 (1878), nom. illeg., as ‘Stachyobium Antennata’. Type species: Dendrobium antennatum Lindl. Dendrobium sect. Eudendrobium subsect. Antennata Rchb.f. ex Pfitzer in Engl. & Prantl, Nat. Pflanzenfam., II, 6, 174 (1889). Type species: Dendrobium antennatum Lindl. Dendrobium sect. Ceratobium subsect. Minacia Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 138. Type species: Dendrobium minax Rchb.f.

DENDROBIUM

Dendrobium sect. Ceratobium subsect. Mirbeliana Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 140. Type species: Dendrobium mirbelianum Gaudich. Dendrobium sect. Ceratobium subsect. Taurina Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 139. Type species: Dendrobium taurinum Lindl. Dendrobium sect. Ceratobium subsect. Undulata Kränzl. in Engl., Pflanzenr., IV. 50. II. B., 21 (1910) 140. Type species: Dendrobium undulatum R.Br. Cepobaculum M.A.Clem. & D.L.Jones, Orchadian, 13, 486 (2002). Type species: Cepobaculum canaliculatum (R.Br.) M.A.Clem. & D.L.Jones (basionym: Dendrobium canaliculatum R.Br.) Ceratobium (Lindl.) M.A.Clem. & D.L.Jones, Orchadian, 13, 486 (2002). Type species: Ceratobium antennatum (Lindl.) M.A.Clem. & D.L.Jones (basionym: Dendrobium antennatum Lindl.) Durabaculum M.A.Clem. & D.L.Jones, Orchadian, 13, 487 (2002), nom.illeg. Type species: Durabaculum undulatum (R.Br.) M.A.Clem. & D.L.Jones (basionym: Dendrobium undulatum R.Br.) Stems clustered, usually cane-like, fleshy, many-leaved, not branching, with limited apical growth, sometimes with clustered pseudobulbs having 2–7 narrow, fleshy leaves apically. Leaves with a tubular sheath at base. Inflorescences elongate, erect or arching, few- to many-flowered, arising from apical part of stem. Flowers often with twisted petals longer than dorsal sepal; mentum with an abruptly narrowed, spur-like apex. Labellum adnate to column foot, forming a narrow conical spur, abruptly widened above spur, trilobed, with lamellate, basal keels often abruptly raised on midlobe. About 68 species from Java, Lesser Sunda Islands, Sulawesi, Moluccas, Philippines, New Guinea, Australia, Solomon Islands, Vanuatu, New Caledonia, Samoa, Fiji, and Tonga, with a clear centre of diversity in New Guinea. Dendrobium sect. Stachyobium Lindl., Edwards’s Bot. Reg., 30, 55 (1844). Type species: Dendrobium denudans D.Don Dendrobium sect. Aclinia (Griff.) Lindl., J. Linn. Soc., Bot., 3, 9 (1859). Type species: Dendrobium aclinia Lindl. Taxa in other ranks with their type species in D. sect. Stachyobium: Aclinia Griff., Notul. Pl. Asiat., 3, 320 (1851). Type species: Dendrobium aclinia Lindl. Dendrobium sect. Stachyobium subsect. Eriaeflorae Benth. & Hook.f., Gen. Pl., 3, 500 (1883). Type species: Dendrobium eriiflorum Griff. Dendrobium sect. Stachyobium subsect. Humiles Benth. & Hook.f., Gen. Pl., 3, 499 (1883). Type species: Dendrobium humile Wight Dendrobium subgen. Stachyobium (Lindl.) Brieger in Schltr., Orchideen, ed. 3, 1, 697 (1981), nom. illeg. Type species: Dendrobium denudans D.Don Pseudobulbs clustered, fleshy, variable in shape, sometimes hardly swollen and cane-like, leafy throughout or with leaves in apical part only, not branching, with limited apical growth. Leaves with a tubular sheath at base, short-lived. Inflorescences elongate,

usually with wiry peduncle and rachis, erect or patent, sometimes pendulous, few- to many-flowered, arising apically and laterally from leafy shoots of current growing season, often while shoot is not fully grown, or sometimes when shoot has barely developed; often with three or more inflorescences arising from the same shoot simultaneously. Flowers often whitish with green or purplish markings on labellum; mentum well-developed, conical. Labellum flexibly hinged with column foot, usually trilobed, with lamellate keels basally, or with a flattened, rod-shaped callus often lobed at apex, the midlobe often with undulate margins, sometimes with numerous clavate appendages. About 44 species from Sri Lanka, India, Nepal, Bhutan, Burma, China, Thailand, Laos, Cambodia, Vietnam, Peninsular Malaysia, Java, and the Lesser Sunda Islands, with centres of diversity in India and Thailand. (AS, PA)

Anatomy Solereder and Meyer (1930) provided anatomical data for Orchidaceae, including many species of Dendrobium, to that time. Mejstrik (1970) studied root anatomy in Dendrobium cunninghamii. Khasim and Rao (1989) investigated the anatomy of four Indian species. Yukawa et al. (1990) reported leaf surface structure in some species, and Yukawa et al. (1992) discussed two stomatal shapes and their systematic importance. Stern et al. (1994) presented an anatomical analysis of thick-leaved species in D. section Rhizobium. Morris et al. (1996) prepared an anatomical and systematic analysis of vegetative parts of plants in the genus. Carlsward et al. (1997) made a comparative anatomical study of thick-leaved species in two sections of Dendrobium. The discovery of Dendrobium winikaphyllum with abaxial brachyparacytic stomata together with Earina fouldenensis (Agrostophyllinae) from Early Miocene deposits of New Zealand represents the earliest unequivocal vegetative orchid fossil (Conran et al. 2009). The following description represents results from original anatomical studies on species of Dendrobium including those formerly treated as Cadetia.

Leaf surface

Hairs ambifacial, uniseriate, uni- to tricellular in D. angustifolium (Blume) Lindl. and D. pallens Ridl., glandular, base sunken in epidermal crypt walls. Epidermis: cells polygonal, anticlinal walls straight-sided and curvilinear, sinuous in a few species. Stomata abaxial, ab- and adaxial in D. canaliculatum R.Br.; paracytic in some species, mostly tetracytic. transverse section

Cuticle: thin to moderately thick, smooth to irregular. Epidermis: cells elliptical, square, rectangular, and hexagonal. Stomata superficial. Hypodermis present, or absent in D. amethystoglossum Rchb.f. and D. salaccense (Blume) Lindl., of one or more layers, cells with helical wall thickenings in D. sect. Cadetia, parenchyma cells sometimes alternating with fibre bundles. Fibre bundles: present, or absent in D. comatum (Blume) Lindl. and D. chrysotropis Schltr.; with stegmata in D. malbrownei Dockrill or without stegmata in D. amplum Lindl., bearing spherical, rough-surfaced silica bodies. Mesophyll: homogeneous in D. salaccense and D. 73

DENDROBIEAE

cunninghamii Lindl., or heterogeneous in D. chrysotropis and D. jonesii Rendle; water-storage cells with helical thickenings present in D. munificum (Finet) Schltr. and D. sect. Cadetia. Vascular bundles collateral in one row, phloic and xylic sclerenchyma present; stegmata with spherical, rough-surfaced silica bodies present opposite vascular sclerenchyma, or absent in D. senile C.S.P.Parish & Rchb.f. Leaves in species of D. sections Rhizobium and Aporum are unique in Dendrobium because exposed surfaces are for the most part abaxial. Species in D. section Rhizobium stand out because of their fleshy, usually terete or subterete leaves and lack of fleshy stems. Plants in D. section Aporum have coriaceous, unifacial, and laterally flattened leaves borne in fascicles on upright or pendent stems arising from a short rhizome. Collateral vascular bundles occur in two facing rows between which is a suture or an incipient suture in species of D. section Aporum and a lacuna or an incipient lacuna in species of D. section Rhizobium. There are no fibre bundles in leaves of D. section Rhizobium, but these occur abaxially in leaves of D. section Aporum. Stomata abaxial only; paracytic in species of D. section Aporum and tetracytic in D. section Rhizobium. Hairs uniseriate, bicellular, and glandular in both sections. An adaxial epidermis is exposed in the foliar groove of leaves in D. rigidum R.Br. and D. torresae (F.M.Bailey) Dockrill. In most species of D. section Rhizobium the adaxial epidermis is internal completely surrounded by cells of the mesophyll. It forms an epithelium enclosing a canal represented as a lacuna in transverse section, the cells of which are covered by a cuticle and subtended by a hypodermis.

Stem transverse section

Hairs present in D. senile or absent. Cuticle moderately thick, smooth. Stomata absent. Ground tissue: parenchymatous, bior trizonate, or non-zonate; water-storage cells with helical cell wall thickenings in inner region of zonate ground tissue present in D. munificum; water-storage cells absent in D. cunninghamii. Vascular bundles collateral, concentrated in inner regions of stems with zonate parenchyma and sometimes forming a ring between outer zones; randomly dispersed in non-zonate ground tissue. Phloic and xylic sclerenchyma present; stegmata with spherical rough-surfaced silica bodies present, absent in D. senile and D. rigidum.

Root transverse section

Velamen: 2–17-layered with variably shaped cells and fine or relatively wide, helical, anastomosing thickening bands in D. pachyphyllum (Kuntze) Bakh.f. Tilosomes absent. Exodermis: dead cells ○-, ∪-, or ∩-thickened; anticlinal walls with tenuous scalariform bars present in D. pachyphyllum. Cortex: cells polygonal, thin-walled, parenchymatous. Endodermis uniseriate, cells isodiametric or anticlinal, cell walls ○-thickened opposite phloem sectors, thin-walled opposite xylem sectors. Pericycle configured like endodermis. Vascular cylinder 5–27-arch; conductive cells embedded in sclerenchyma; xylem and phloem elements alternate around the periphery of the vascular cylinder. Pith sclerenchymatous or parenchymatous. (WS) 74

Palynology Schill and Pfeiffer (1977) examined pollen of three species of Cadetia, two of Diplocaulobium, and 28 of Dendrobium. Tetrads of Dendrobium sect. Cadetia (studied as the genus Cadetia) were level (convex in one species) and had laevigate sculpturing with a calymmate sexine about 3.3 μm thick. Those of Dendrobium sect. Diplocaulobium (formerly the genus Diplocaulobium) were concave and had microfoveolate sculpturing and a rounded sexine. Tetrads of the species studied as Dendrobium were concave, convex, or even depending on the species, and sculpturing was essentially laevigate (sometimes weakly microfoveolate or verrucose). The sexine of Dendrobium species was mostly rounded except angular in D. anceps Sw. In D. cariniferum Rchb.f., D. bracteosum Rchb.f., and D. capituliflorum Rolfe the sexine was calymmate and 1–11 μm thick (3–4 μm thick in the middle). Pollen data from the former species of Cadetia and Diplocaulobium fall within the observed range for Dendrobium. (AP)

Cytogenetics Reported chromosome numbers for Dendrobium are 2n = 18, 30, 32–35, 36, 36 + Bs, 38, 38 + Bs, 39, 40, 40 + Bs, 41, 42, 43, 57, 76, and 80 (Jones et al. 1982; Brandham 1999). Brandham (1999) suggested a main basic number of x = 19, so that 2x = 38, 3x = 57, and 4x = 76; another basic number, x = 20 or 10, would explain 2n = 40, 80. It is possible that x = 10 gave rise to x = 19 by doubling followed by dysploid reduction (Brandham 1999). Chromosome numbers reported for genera now included in Dendrobium are either 2n = 38 (Diplocaulobium, Flickingeria) or 2n = 40 (Epigeneium, Ephemerantha), the most common numbers for Dendrobium (Brandham 1999). (AP)

Phytochemistry More has been published on the phytochemistry of Dendrobium than on any other orchid genus. The most important contributing factor in this regard is the use of some species in traditional Chinese medicine, with a documented history covering almost two millennia. Hew et al. (1997) gave an excellent introduction to the subject in which several of the species concerned are described together with a brief overview of conditions treated and typical formulations of the crude drugs. No comprehensive review of Dendrobium phytochemistry is available in the literature. Existing accounts are mainly those found within general surveys of Orchidaceae chemistry (Hegnauer 1963, 1986) or specific classes of natural product, such as Orchidaceae alkaloids (Lüning 1974; Slaytor 1977). Some updates on progress in the characterization of chemical constituents from Dendrobium have appeared subsequently in the Chinese literature (Chen and Guo 2001; Zhang et al. 2003) but are not widely accessible. The large number of species for which phytochemical data are available makes individual accounts impractical; instead the following discussion is organized by classes of compound. Alkaloids, sesquiterpenoids, and stilbenoids are described first, as these are the most frequently reported constituents of Dendrobium. The subsequent commentary is concerned with other significant groups of natural product found in the genus, including

DENDROBIUM

anthraquinones, coumarins, di- and triterpenoids, flavonoids and anthocyanins, lignans, phenolics, polysaccharides, and scent constituents. Constituents that do not fall within the above categories but are nevertheless worthy of record are the glucosylated lactones, cis- and trans-crassinodine, isolated from D. pendulum Roxb. (cited as D. crassinode Benson & Rchb.f.), and 3,6,9-trihydroxy3,4-dihydroanthracen-1(2H)-one, a novel tetrahydroanthracene derivative found in stem extracts of D. polyanthum Wall. ex Lindl. (Dahmén et al. 1976; Hu et al. 2009).

Alkaloids Presence and localization of alkaloids in Dendrobium were investigated at the end of the nineteenth century using microchemical techniques based on staining and microscopy. De Wildeman (1892) observed that alkaloids were present in D. ×ainsworthii T.Moore and D. nobile Lindl., noting that the latter contained them in all tissues. These findings were confirmed in 1896 by De Droog, who also listed eight species in which alkaloids were not detected (reprinted as De Droog 1906). Other early surveys for alkaloid content that include results for Dendrobium are those of Boorsma (1902) and Wester (1921). Extensive sampling of Orchidaceae for alkaloid content using chemical methods has created more than 200 records for named species of Dendrobium (including a few citations as Cadetia, Diplocaulobium, and Ephemerantha), which appear in the surveys of Lüning (1964, 1967) and Lawler and Slaytor (1969, 1970). In a subsequent evaluation, based partly on the last but also on unpublished material, 32 of 391 species of Dendrobium (including Cadetia and Diplocaulobium) tested were reported to have an alkaloid content ≥ 0.1% (Lüning 1974). Table 622.1 gives details of the identity and sources of more than 30 alkaloids reported from Dendrobium. The best known of these is dendrobine (Fig. 622.8, 1), which Suzuki et al. isolated as a crystalline substance from stem extracts of D. nobile in 1932, although more than three decades elapsed before its structure was determined (Inubushi et al. 1964a; Onaka et al. 1964; Yamamura and Hirata 1964) and later confirmed by synthesis (Inubushi et al. 1972; Yamada et al. 1972). Good accounts of the history of alkaloid research in Dendrobium are available that summarize important contributions made by Swedish and Japanese research groups in the 1960s and early 1970s (Lüning 1974; Slaytor 1977). Dendrobine alkaloids (Fig. 622.8) are unique to Dendrobium and for convenience can be classified into four structural classes represented by dendrobine (type A), nobilonine (type B), dendroxine (type C), and dendrowardine (type D). Biosynthesis of dendrobine appears to proceed from a rearranged sesquiterpenoid that has its origin in the mevalonate pathway (Yamazaki et al. 1966; Corbella et al. 1973). Type A dendrobine alkaloids are the most numerous of the four classes. Of the 12 examples currently recorded, eight occur in D. nobile, with dendrobine the major alkaloid of this species. Nobilonine (type B), a ring-opened form of dendrobine that may be a biosynthetic precursor, is the second most abundant alkaloid of D. nobile but is also found in three other species. Dendroxine (type C) is an elaborated form of dendrobine characterized by an additional

heterocyclic ring, and four derivatives are known. The structure of dendrowardine (type D), a constituent of D. wardianum R.Warner, is unique insofar as an O-heterocyclic ring replaces the analogous N-heterocyclic ring of dendrobine, and a choline moiety is attached as a functional group. A comprehensive overview and critique of the numerous syntheses of dendrobine alkaloids was given by Gössinger (2010). Relatively little has been published on the biological activity of these compounds, especially in recent literature, although some information is available in general reviews (Gutiérrez 2010; Hossain 2011). Dendrobine itself was reported to have mild analgesic and antipyretic properties to produce moderate hyperglycemia and reduce cardiac activity; at higher doses it lowers blood pressure, suppresses respiration, and causes convulsions (Chen and Chen 1935a). Lethal doses for small animals are in the range 17–22 mg/kg, although sodium amytal (sodium isoamyl ethyl barbiturate) can be effective as an antidote (Chen and Chen 1935b; Chen and Rose 1936). A few representatives of the imidazole, indolizidine, phthalide, pyrrolidine, and quinolizidine classes of alkaloid are also found in Dendrobium (Table 622.1 and Fig. 622.9, 22–31). The structure of anosmine (22), a unique imidazolinium salt from D. anosmum Lindl. and D. parishii Rchb.f., was confirmed using X-ray crystallography and synthesis (Leander and Lüning 1968; Söderberg and Kierkegaard 1970). This alkaloid is derived from two molecules of the amino acid lysine, which are incorporated via cadaverine and pipecolic acid units (Hemscheidt and Spenser 1991, 1993). The phthalide alkaloid pierardine (26) is related structurally to the phthalide-pyrrolidine alkaloid shihunine (30). In solution, the latter isomerizes to a betaine, which is probably the naturally occurring form in the plant (Elander et al. 1971). Studies on the biosynthesis of shihunine (30) in D. aphyllum (Roxb.) C.E.C.Fisch. (as D. pierardii Roxb. ex Hook.) indicated that o-succinylbenzoic acid is a direct precursor (Leete and Bodem 1976). The unusual spiro structure of shihunine (30) has attracted the interest of synthetic chemists, and several total syntheses are available (Gore et al. 1990).

Sesquiterpenoids More than 50 sesquiterpenes representing seven structural classes have been isolated from Dendrobium (Table 622.2, Fig. 622.10 and 622.11). The most abundant sources of compounds are extracts prepared from stems of D. moniliforme (L.) Sw. and D. nobile. Picrotoxanes, which are the largest group of Dendrobium sesquiterpenes (53–83), are of special interest because of their structural similarity to dendrobine alkaloids. As such, they are generally considered to be precursors in the biosynthesis of the latter (Gössinger 2010). Nobilomethylene (81) is probably an artefact generated from nobilonine N-oxide (Okamoto et al. 1972). Some Dendrobium sesquiterpenes possess immunomodulatory activity. According to the results of in vitro assays using murine spleen cells, dendroside A (34), dendronobiloside A (73), and dendrosides D–G (37, 51, 75, 76) stimulate the proliferation of B and T lymphocytes, whereas dendronobiloside B (74) shows inhibitory activity (Zhao et al. 2001; Ye et al. 2002). Similarly, dendromonilisides A (44) and C (63) stimulate proliferation of B lymphocytes but inhibit proliferation of T lymphocytes (Zhao et al. 2003a). 75

DENDROBIEAE

Table 622.1.  Alkaloids reported from species of Dendrobium. No.

Alkaloid class and name

1

DENDROBINE-TYPE A Dendrobine

2

Dendramine (6-hydroxydendrobine)

3 4 5 6 7 8 9

10-Hydroxydendrobine 2-Oxodendrobine (mubironine A) 3-Hydroxy-2-oxodendrobine N-Demethyldendrobine (mubironine B) Seco-dendrobine; methyl ester (mubironine C) Dendrobine N-oxide N-Methyldendrobinium iodide

D. findlayanum D. Snowflake ‘Red Star’ D. nobile D. Snowflake ‘Red Star’ D. Snowflake ‘Red Star’ D. nobile D. nobile

10 11

N-Isopentenyldendrobinium bromide Dendrine

D. nobile D. nobile

12

D. nobile

13

Dendronobiline A DENDROBINE-TYPE B Nobilonine (nobiline)

14

6-Hydroxynobilonine

15

Moniline DENDROBINE-TYPE C Dendroxine 6-Hydroxydendroxine 8-Hydroxydendroxine N-Isopentenyldendroxinium chloride

D. friedericksianum D. moniliforme (L.) Sw. D. signatum (as D. hildebrandtii) D. moniliforme

Granelli et al. (1970), Suzuki et al. (1975) Hedman et al. (1971) Yamamura and Hirata (1964), Onaka et al. (1965) Elander and Leander (1971) Hedman et al. (1971) Liu et al. (2007a) Elander and Leander (1971) Liu et al. (2007a)

D. nobile D. nobile D. nobile D. friedericksianum D. nobile D. signatum (as D. hildebrandtii) D. friedericksianum D. nobile D. signatum (as D. hildebrandtii)

Okamoto et al. (1966a) Okamoto et al. (1966b) Okamoto et al. (1972) Hedman et al. (1971) Hedman and Leander (1972) Elander and Leander (1971) Hedman et al. (1971) Hedman and Leander (1972) Elander and Leander (1971)

D. wardianum R.Warner

Blomqvist et al. (1973)

D. anosmum Lindl., D. parishii Rchb.f.

Leander and Lüning (1968), Söderberg and Kierkegaard (1970), Hemscheidt and Spenser (1991, 1993)

D. crepidatum Lindl. ex Paxton D. crepidatum D. polyanthum Wall. ex Lindl. (as D. primulinum Lindl.)

Elander et al. (1973) Elander et al. (1973) Lüning and Leander (1965), Blomqvist et al. (1972)

D. aphyllum (Roxb.) C.E.C.Fisch (as D. pierardii Roxb. ex Hook.)

Elander et al. (1969, 1971)

D. chrysanthum Wall. ex Lindl. D. polyanthum (as D. primulinum) D. chrysanthum D. chrysanthum D. aphyllum (as D. pierardii) D. lohohense Tang & F.T.Wang

Lüning and Leander (1965) Lüning and Leander (1965) Yang et al. (2005) Ekevåg et al. (1973) Elander et al. (1971) Inubushi et al. (1964b, 1968)

D. crepidatum

Kierkegaard et al. (1970), Elander et al. (1973)

16 17 18 19 20

N-Isopentenyl-6-hydroxydendroxinium chloride

22

DENDROBINE-TYPE D Dendrowardine IMIDAZOLE Anosmine (as bromide salt)

23 24 25

INDOLIZIDINE Crepidamine Dendrocrepine Dendroprimine

26

PHTHALIDE Pierardine

27

PYRROLIDINE Hygrine

28 29 30

Dendrochrysanine: (E)- and (Z)-isomers Dendrochrysine: (E)- and (Z)-isomers Shihunine

31

QUINOLIZIDINE Crepidine

21

76

Species

Reference

D. findlayanum C.S.P.Parish & Rchb.f. D. linawianum Rchb.f. D. nobile Lindl. as ‘Chin-Shih-Hu’ D. friedericksianum Rchb.f. D. nobile D. signatum Rchb.f. (as D. hildebrandtii Rolfe)

Granelli et al. (1970) Suzuki et al. (1934) Suzuki et al. (1932) Hedman et al. (1971) Inubushi et al. (1964a, 1966), Okamoto et al. (1966b) Elander and Leander (1971) Granelli et al. (1970), Suzuki et al. (1975) Morita et al. (2000) Wang et al. (1985) Morita et al. (2000) Morita et al. (2000) Hedman and Leander (1972) Inubushi et al. (1964a), Hedman and Leander (1972) Hedman and Leander (1972) Inubushi and Nakano (1965), Granelli and Leander (1970) Liu and Zhao (2003)

D. findlayanum D. friedericksianum D. nobile D. signatum (as D. hildebrandtii)

DENDROBIUM

Fig. 622.8.  Dendrobine-type alkaloids (1–21) of Dendrobium (see Table 622.1).

Stilbenoids This is the best-represented class of natural products in Dendrobium, with more than 120 examples described to the end of 2011. For convenience these have been divided into three subclasses: bibenzyls (84–137, Table 622.3), phenanthrenes (138–198, Table 622.4), and fluorenones (199–207, Table 622.5). Selected examples are illustrated (Fig. 622.12– 622.14). Some compounds listed in Tables 622.3–622.5 occur in many species of Dendrobium, whereas others are known only from a single source. Batatasin III (87), gigantol (96),

moscatilin (109), and chrysotobibenzyl (113) are the most commonly reported bibenzyls (Veerraju et al. 1989a; Yang et al. 2006b), and among the phenanthrenes moscatin (139) is the most widespread (Yang et al. 2006b). Although the majority of Dendrobium bibenzyls are simple derivatives with only hydroxy and methoxy substituents (84–119), other more complex structures have been described, including dimers and various conjugated forms (120–137). Similarly, most phenanthrenes and 9,10-dihydrophenanthrenes are also characterized by various combinations of hydroxy and methoxy substituents (138–178), 77

DENDROBIEAE

Fig. 622.9.  Imidazole (22), indolizidine (23–25), phthalide (26), pyrrolidine (27–30), and quinolizidine (31) alkaloids of Dendrobium (see Table 622.1). Ph = phenyl.

9,10-­dihydrophenanthrenes isolated from whole plant extracts of D. loddigesii inhibited nitric oxide production, whereas ­bibenzyls from the same source did not. The most active compound was loddigesiinol A (155). Evaluation of inhibition of nitric oxide production by stilbenoids isolated from stem extracts of D. nobile revealed that 9,10-dihydrophenanthrenes were more active than phenanthrenes and bibenzyls (Hwang et al. 2010). The anti-fibrotic and antioxidant properties of stilbenoids ­present in stem extracts of D. nobile have also been examined (Yang et al. 2007a; Zhang et al. 2007e, 2008d). Similarly, Sritularak et al. (2011) reported antioxidant activity for stilbenoids present in a tea prepared from the stems of D. draconis, which contained batatasin III (86), gigantol (96), hircinol (140), 149, and 184. Among bibenzyls isolated from stem extracts of D. gratiosissimum, gigantol (96), moscatilin (109) and the dimers dengraols A (131) and B (132) showed cytotoxicity to human leukemia HL-60 cells (Zhang et al. 2008a). Trigonopol A (136), a bibenzyl conjugate from stems of D. trigonopus, exhibited antiplatelet aggregation activity (Hu et al. 2008b). The phenanthrenes denthyrsinin (172), denthyrsinol (193), and denthyrsinone (194), which occur uniquely in stem extracts of D. thyrsiflorum, showed cytotoxicity in several human cancer cell lines (Zhang et al. 2005b). Eight fluorenones (199–206) and one fluoren-9-ol (207) have been recorded as constituents of Dendrobium (Table 622.5, Fig. 622.14). Among these, the most frequently reported examples are dengibsin (199), dendroflorin (202), and denchrysan A (203). The first naturally occurring fluorenones to be discovered were dendroflorin from D. densiflorum (Talapatra et al. 1984), and dengibsin and dengibsinin from D. gibsonii (Talapatra et al. 1985), although the structures proposed for the compounds were later revised (Fan et al. 2001; Sargent 1987). Some fluorenones isolated from Dendrobium have been tested in bioassays. In studies on the stem constituents of D. nobile, both nobilone (200) and dendroflorin (202) showed higher antioxidant activity than vitamin C in an oxygen radical absorbance capacity (ORAC) assay, and inhibited NO production without cytotoxicity (Zhang et al. 2007e). Three fluorenones (201–203) isolated from whole plant extracts of D. chrysotoxum showed selective cytotoxicity to human hepatoma BEL-7402 cells when tested against a panel of human cancer cell lines (Chen et al. 2008b). Denchrysan B (207), the sole example of a fluoren-9-ol, occurs in three species of Dendrobium (Table 622.5). The absolute configuration of a sample of this compound obtained from stem extracts of D. chrysotoxum was determined to be (9R) using Mosher’s method (Yang et al. 2004a).

Other natural products anthraquinones

with one example of an unusual glycosylated form afforded by denchrysan A (173). The remaining phenanthrenes comprise a phenanthrenone (179), a phenanthropyran (180), phenanthropyranones (181, 182), phenanthraquinones (183–189), a phenanthradiquinone (190), and dimers and other conjugates (191–198). Biological activity of some Dendrobium phenanthrenes has been reviewed (Kovács et al. 2008). In more recent work, Ito et al. (2010) demonstrated that phenanthrenes and 78

Reports of anthraquinones as chemical constituents of Dendrobium have emerged only from large-scale extractions of stem material. The species concerned are D. chrysanthum, which yielded chrysophanol, emodin, and physcion (Yang et al. 2004c), D. fimbriatum, from which aloe-emodin, chrysophanol, emodin, physcion, and rhein were isolated (Bi et al. 2001a,b, 2003), and D. thyrsiflorum, which contains chrysophanein, the 8-O-glucoside of chrysophanol (Zhang et al. 2005a).

DENDROBIUM

Table 622.2.  Sesquiterpenoids reported from species of Dendrobium. No.

Sesquiterpenoid class and name

Species

Reference

32

AROMADENDRANES Dendrobiumane A

Dendroside D 10β,12,14-Trihydroxyalloaromadendrane 10β,13,14-Trihydroxyalloaromadendrane

D. moniliforme (L.) Sw. D. nobile D. nobile D. moniliforme D. nobile D. nobile D. moniliforme D. nobile D. nobile D. nobile D. moniliforme

Zhao et al. (2003b) Zhang et al. (2007c) Zhang et al. (2007d) Zhao et al. (2003a) Zhao et al. (2001) Ye and Zhao (2002) Zhao et al. (2003a) Ye and Zhao (2002) Ye et al. (2002) Ye and Zhao (2002) Zhao et al. (2003b)

33 34

Dendronobilin H Dendroside A

35 36

Dendroside B Dendroside C

37 38 39 40 41 42

CADINANES Dendronobilin G Dendronobiloside C Dendronobiloside D

D. nobile D. nobile D. nobile

Zhang et al. (2007d) Ye and Zhao (2002) Ye and Zhao (2002)

D. nobile D. moniliforme D. nobile D. nobile D. nobile D. nobile

Ye and Zhao (2002) Zhao et al. (2003a) Zhang et al. (2007d) Zhang et al. (2007d) Zhang et al. (2008c) Ye and Zhao (2002)

50

COPACAMPHANES Dendrobane A Dendromoniliside A Dendronobilin A Dendronobilin I Dendronobilins K and N Dendronobiloside E CYCLOEUDESMANES Dehydroshizukanolide

D. fimbriatum Hook.

Qing et al. (2009)

51

EMMOTIN-GROUP Dendroside E

D. nobile

Ye et al. (2002)

52

OPPOSITANES Bullatantriol

D. nobile

Zhang et al. (2007c)

53 54 55 56 57–60 61

PICROTOXANES Aduncin Amoenin Amotin Crystallinin Dendrobiumanes B–E Dendrodensiflorol

62–64

Dendromonilosides B–D

65–69

Dendronobilins B–F

70 71, 72 73, 74 75, 76

Dendronobilin J Dendronobilins L and M Dendronobilosides A and B Dendrosides F and G

77 78 79, 80 81 82 83

α-Dihydropicrotoxinin 10,12-Dihydroxypicrotoxane Flakinins A and B Nobilomethylene Picrotin 6α,10,12-Trihydroxypicrotoxane

D. aduncum Lindl. D. amoenum Wall. ex Lindl. D. amoenum D. crystallinum Rchb.f. D. moniliforme D. densiflorum Lindl. D. nobile D. moniliforme (B–D) D. nobile (D only) D. crystallinum (B only) D. nobile (B–F) D. nobile D. nobile D. nobile D. moniliforme (F only) D. nobile (F and G) D. moniliforme D. nobile D. Snowflake ‘Red Star’ D. nobile D. moniliforme D. nobile

Gawell and Leander (1976) Dahmèn and Leander (1978) Dahmèn and Leander (1978) Wang et al. (2009) Zhao et al. (2003b) Tang et al. (2004) Zhang et al. (2007c) Zhao et al. (2003a) Shu et al. (2004) Wang et al. (2009) Zhang et al. (2007d) Zhang et al. (2007c) Zhang et al. (2008c) Zhao et al. (2001) Zhao et al. (2003a) Ye et al. (2002) Zhao et al. (2003b) Zhang et al. (2007c) Morita et al. (2000) Okamoto et al. (1972) Zhao et al. (2003b) Zhang et al. (2007c)

43 44 45 46 47, 48 49

coumarins

Ayapin (6,7-methylenedioxycoumarin), scoparone (6,7-dimethoxycoumarin), and scopoletin (7-hydroxy-6-methoxycoumarin) have been detected in acid-hydrolysed extracts of leaves of D. thyrsiflorum (Wrigley 1960; Bate-Smith 1968), and coumarin itself in similarly treated leaf material of D. moschatum (Buch.-Ham.)

Sw. (Bate-Smith 1968). Later work on stem extracts of D. thyrsiflorum revealed the presence of ayapin, coumarin, scopoletin, and two glycosides of the latter (scopolin and xeroboside), as well as denthyrsin, a coumarin-benzofuran dimer (Zhang et al. 2005a,b). Stem extracts of this species also contain (E)- and (Z )-2-glucosyloxycinnamic acid derivatives that are presumed to 79

DENDROBIEAE

Fig. 622.10.  Sesquiterpenes of Dendrobium from the aromadendrane (32–39), cadinane (40–42), copacamphane (43–49), cycloeudesmane (50), emmotingroup (51), and oppositane (52) classes (see Table 622.2).

be biogenetic precursors of ayapin, scoparone, and scopoletin, and which are responsible for their formation under conditions of acid hydrolysis (Zhang et al. 2006a). Levels of these metabolites in different tissues, and in stem material of D. thyrsiflorum collected from various locations in Yunnan province, China, have been evaluated for quality purposes using high performance liquid chromatography coupled to diode-array detection (Zhang et al. 2006a). Additional records of the occurrence of coumarin have been published for D. chryseum Rolfe (Zheng et al. 2000a) and D. denneanum (as D. aurantiacum var. denneanum; Yang et al. 2006a; Liu et al. 2009b). Stem extracts of D. densiflorum contain 3,4-­dihydroayapin, which has not been reported from any other 80

source, together with ayapin, scoparone, and scopolin (Zheng et al. 2000b). Scoparone is also present in D. catenatum Lindl. (as D. tosaense Makino; Lo et al. 2004), ayapin and scoparone in D. farmeri (Majumder and Chakraborti 1989), and scopolin methyl ether in D. fimbriatum (Bi et al. 2003). diterpenoids and triterpenoids

Compared with sesquiterpenoids, diterpenoids are rarely reported in Dendrobium, and only a few examples are known. Ephemeric acid and ephemeroside are uncommon cis-clerodanetype diterpene lactones isolated from extracts of the whole plant of D. comatum Blume (published as Ephemerantha comata (Blume) P.F.Hunt & Summerh.) (Niwa et al. 1987). Ephemeranthoside is a glucoside of a pimarane-type diterpene obtained by Tezuka

DENDROBIUM

Fig. 622.11.  Picrotoxane-type sesquiterpenes (53–83) of Dendrobium (see Table 622.2). Glcp = β-glucopyranosyl. The structures are drawn in relation to dendrobine (1, Fig. 622.8), for which the absolute configuration is known (for the majority of Dendrobium picrotoxanes, the relative configuration was determined).

81

DENDROBIEAE

Table 622.3.  Bibenzyls reported from species of Dendrobium. No.

Bibenzyl type and trivial name

84 85 86

Tri-O-substituted 3,4,3′-TriOH 3,5,3′-TriOH 3,5,4′-TriOH

87 88

3,3′-DiOH-5-OMe (batatasin III) 3,4′-DiOH-5-OMe

89 90

3-OH-5,3′-diOMe 3,5,3′-TriOMe

91 92 93

Tetra-O-substituted 4,3′,5′-TriOH-3-OMe (tristin) 3,4-DiOH-5,3′-diOMe (dendrobin A) 3,4-DiOH-5,4′-diOMe (moniliformine)

95 96 97

3,4-DiOH-5,4′-diOMe; 3,4-di-O-β-Glcp (dendromoniliside E) 4,3′-DiOH-3,5-diOMe (aloifol I) 4,3′-DiOH-3,5′-diOMe (gigantol) 4,4′-DiOH-3,5-diOMe

98

3′-OH-3,4,5′-diOMe

99 100 101 102

4-OH-3,5,3′-triOMe (dendrobin B) 4-OH-3,5,4′-triOMe (amoenylin) 3′-OH-3,4,5-triOMe (isoamoenylin) 3′-OH-5′-OMe-3,4-OCH2O- (densiflorol A)

103 104 105 106 107 108 109 110

Penta-O-substituted 3,4,3′,4′-TetraOH-5-OMe (dendrocandin E) 3,4,4′-TriOH-5,α(S )-diOMe (dendrocandin C) 3,4,4′-TriOH-5-OMe-α(S )-OEt (dendrocandin D) 3,4-DiOH-5,4′,α(R)-TriOMe (dendrocandin A) 4,5-DiOH-3,3′,α-triOMe (nobilin A) 4,3′-DiOH-3,5,α-triOMe 4,4′-DiOH-3,5,3′-triOMe (moscatilin) 4-OH-3,5,3′,4′-tetraOMe (chrysotoxin)

111

3′-OH-3,4,5,4′-tetraOMe (erianin)

112

4′-OH-3,4,5,3′-tetraOMe (crepidatin)

113 114 115

94

82

Species

Reference

D. longicornu Lindl. D. cariniferum Rchb.f. D. aphyllum D. gratiosissimum Rchb.f. Many recordsa D. amoenum D. gratiosissimum D. moniliforme D. amplum Lindl. D. amplum

Hu et al. (2008a) Liu et al. (2009a) Zhang et al. (2008b) Wang et al. (2007), Zhang et al. (2008a)

Many recordsb D. nobile D. gratiosissimum D. moniliforme D. moniliforme

Majumder et al. (1999a) Wang et al. (2007), Zhang et al. (2008a) Zhao et al. (2003b) Majumder et al. (2008b) Majumder et al. (2008b)

Ye and Zhao (2002) Wang et al. (2007), Zhang et al. (2008a) Bi et al. (2002, 2004) Zhao and Zhao (2003)

D. longicornu Many recordsc D. aphyllum D. crystallinum D. moniliforme (as D. candidum Wall. ex Lindl.) D. moniliforme (as D. candidum) D. plicatile Lindl. D. xantholeucum Rchb.f. (as Ephemerantha lonchophylla (Rchb.f.) P.F.Hunt & Summerh.) D. nobile D. amoenum D. amoenum D. densiflorum

Hu et al. (2008a) Yang et al. (2006b) Zhang et al. (2008b) Wang et al. (2008) Li et al. (2008)

Li et al. (2009a) Li et al. (2009a) Li et al. (2009a) Li et al. (2008) Zhang et al. (2006b) Hu et al. (2008a)

3,4,5,3′,4′-pentaOMe (chrysotobibenzyl)

D. moniliforme (as D. candidum) D. moniliforme (as D. candidum) D. moniliforme (as D. candidum) D. moniliforme (as D. candidum) D. nobile D. longicornu Many recordsc D. chrysanthum D. denneanum Kerr (as D. aurantiacum var. denneanum (Kerr) Z.H.Tsi) D. nobile D. chrysotoxum Lindl. D. moniliforme D. chrysanthum D. crepidatum D. denneanum (as D. aurantiacum var. denneanum) D. nobile Many recordsc

Hexa-O-substituted 4,4′,α-TriOH-3,5,3′-triOMe (nobilin D) 3,3′-DiOH-4,5,4′,5′-tetraOMe (cumulatin)

D. nobile D. cumulatum Lindl.

Zhang et al. (2007e) Majumder and Pal (1993)

Li et al. (2008) Yamaki and Honda (1996) Tezuka et al. (1991) Ye and Zhao (2002) Majumder et al. (1999a) Majumder et al. (1999a) Fan et al. (2000, 2001)

Yang et al. (2006c) Yang et al. (2007d) Zhang et al. (2007e) Yang et al. (2006b) Yang et al. (2006b) Yang et al. (2004c) Majumder and Chatterjee (1989) Liu et al. (2009b) Zhang et al. (2007e)

DENDROBIUM

Table 622.3.  (Continued ). No.

Bibenzyl type and trivial name

Species

Reference

116 117 118

4,4′-DiOH-3,5,3′,α(S)-tetraOMe (nobilin B) 4,4′-DiOH-3,5,3′,α(R)-tetraOMe (loddigesiinol C) 4-OH-3,5,3′,4′,α-pentaOMe (nobilin C)

D. nobile D. loddigesii Rolfe D. nobile

Zhang et al. (2006b) Ito et al. (2010) Zhang et al. (2006b)

119

Hepta-O-substituted 4,4′-DiOH-3,5,3′-triOMe-α,α′-dioxo (loddigesiinol D)

D. loddigesii

Ito et al. (2010)

D. crystallinum D. moniliforme (as D. candidum) D. moniliforme (as D. candidum) D. falconeri Hook. D. nobile D. gratiosissimum D. gratiosissimum D. longicornu D. nobile D. trigonopus Rchb.f.

Wang et al. (2009) Li et al. (2008) Li et al. (2009b) Sritularak and Likhitwitayawuid (2009) Liu et al. (2007b) Zhang et al. (2008a) Zhang et al. (2007f) Hu et al. (2008a) Zhang et al. (2007e) Hu et al. (2008b)

120, 121 122 123-126 127, 128 129, 130 131, 132 133 134 135 136, 137

Complex structures Dencryols A and B Dendrocandin B Dendrocandins F–I Dendrofalconerols A and B Dendronophenols A and B Dengraols A and B Isomoniliformine Longicornuol A Nobilin E Trigonopols A and B

a

Some species records for batatasin III are given by Veerraju et al. (1989a). D. aphyllum (Zhang et al. 2008b); D. denneanum (as D. aurantiacum var. denneanum) (Yang et al. 2007d); D. densiflorum (Fan et al. 2001); D. gratiosissimum (Zhang et al. 2008a); D. longicornu (Hu et al. 2008a); D. nobile (Liu et al. 2007b); D. thyrsiflorum Rchb.f. ex André (Zhang et al. 2004); D. trigonopus (Zhang et al. 2005c). c Many species records for gigantol, moscatlin, and chrysotobibenzyl are given by Yang et al. (2006b). b

et al. (1991) from stem extracts of D. xantholeucum (as Ephemerantha lonchophylla). Later work on this species revealed two additional pimarane diterpenes, lonchophylloids A and B, which were evaluated in cell-based assays for their ability to modulate multidrug resistance to the anticancer drug, doxorubicin (Ma et al. 1998). Large-scale extraction of plant material sourced from Dendrobium typically leads to reports of commonly occurring sterols such as daucosterol, β-sitosterol, and stigmasterol, which are of marginal interest. In some species, however, the terpenoid chemistry is novel. Examples include D. ochreatum Lindl., which yielded the steroidal glycosides dendrosteroside, ochreasteroside, and epi-ochreasteroside (Behr et al. 1975; Behr and Leander 1976), and D. fimbriatum, from which denfigenin was isolated (Talapatra et al. 1992). The last has the structure, (25R)-22α-Ospirost-5-ene-3β,16α,17α-triol, and is a derivative of diosgenin, which also occurs in this species. flavonoids and anthocyanins

Flavonoid constituents of Dendrobium comprise mainly simple flavones, flavonols, flavanones, and, in some cases, flavone C-glycosides or flavonol O-glycosides, although only a few species have been studied in detail. Bate-Smith (1968) detected the flavonol kaempferol in acid-hydrolysed extracts of D. thyrsiflorum leaves; in later work on this species the flavones chrysoeriol and tricin, and flavanones naringenin and homoeriodictyol, were found (Zhang et al. 2004). A wider survey of Orchidaceae leaf flavonoids based on analysis of acid-hydrolysed extracts reported kaempferol in D. chrysanthum, D. densiflorum, D. farmeri, D. fimbriatum (as D. fimbriatum var. oculatum Hook.), D. gracilicaule var. howeanum Maiden, and D. kingianum Bidwill ex Lindl., and quercetin in D. densiflorum, D. farmeri, D. kingianum, and D. ×superbiens (Williams 1979). Quercetin also occurs in D. catenatum (as D. tosaense; Lo et al. 2004), and kaempferol in D. denneanum (as

D. aurantiacum var. denneanum), together with naringenin (Yang et al. 2006a). Additional records of flavones comprise tricin, its 4′-methyl ether, and tricetin 7,3′,5′-trimethyl ether from D. aphyllum (Shao et al. 2008), luteolin from D. denneanum (as D. aurantiacum var. denneanum; Liu et al. 2009b), and apigenin from D. crystallinum (Wang et al. 2009) and D. gratiosissimum (Wang et al. 2007). The flavanone naringenin has also been reported from D. crystallinum (Li et al. 2007; Wang et al. 2008), D. longicornu (Hu et al. 2008a), and D. trigonopus (Zhang et al. 2005c; Hu et al. 2008b). Flavone C-glycosides are present in leaves of D. chrysotoxum, D. moschatum, D. plicatile (as Ephemerantha fimbriata), and D. triflorum (Blume) Lindl. (as Epigeneium triflorum (Blume) Summerh.), according to Williams (1979). Isoviolanthin (6-C-rhamnosyl-8C-glucosylapigenin) occurs in the stems of D. crystallinum, but the identity of a second flavone-C-glycoside reported from this species as ‘6″′-glucosyl-vitexin’ is uncertain (Wang et al. 2009). Four new 6,8-di-C-glycosylflavones have been isolated from leaf and stem extracts of D. catenatum (as D. huoshanense Z.Z.Tang & S.J.Cheng) together with the known compound, isoschaftoside (Chang et al. 2010). Flavonol O-glycosides are less widely reported, although quercetin 3-O-rutinoside (rutin) was detected in the white flowers of D. crumenatum Sw. (Lowry and Keong 1973), and 3-O-glycosides, 7-O-glycosides, and 3,7-diO-glycosides of kaempferol and quercetin in flower tissue of D. ‘Pompadour’, together with trace amounts of the flavone C-glycoside, vicenin-2 (Williams et al. 2002). Early observations on anthocyanins of Dendrobium were concerned with localization rather than chemical constitution of pigments (Gertz 1906). Although the presence of anthocyanins in the flowers of D. nobile was noted by Shibata et al. in 1919, the first study in which characterization of specific pigments was addressed was that of Lowry and Keong (1973), who 83

DENDROBIEAE

Table 622.4.  Phenanthrenes reported from species of Dendrobium. No.

Phenanthrene type and trivial name

138

Tri-O-substituted 2,4,7-TriOH-9,10-dihydro

139 140 141 142

2,5-DiOH-4-OMe (moscatin) 2,5-DiOH-4-OMe-9,10-dihydro (hircinol) 2,7-DiOH-4-OMe (flavanthrinin) 2,7-DiOH-4-OMe-9,10-dihydro (coelonin)

143 144

4,5-DiOH-2-OMe-9,10-dihydro 4,7-DiOH-2-OMe-9,10-dihydro (lusianthridin)

145 146

2-OH-4,7-diOMe-9,10-dihydro 5-OH-2,4-diOMe

147

Tetra-O-substituted 2,3,5-TriOH-4-OMe (fimbriol B)

148

2,3,5-TriOH-4-OMe-9,10-dihydro (ephemeranthol C)

149

2,5,7-TriOH-4-OMe-9,10-dihydro

150

2,5,9-TriOH-4-OMe-9,10-dihydro (rotundatin)c

151 152

2,5-DiOH-3,4-diOMe 2,5-DiOH-4,9-diOMe

153

2,7-DiOH-3,4-diOMe (nudol)

154

2,7-DiOH-3,4-diOMe-9,10-dihydro (erianthridin)

155 156 157

2,9-DiOH-4,5-diOMe (loddigesiinol A) 3,5-DiOH-2,4-diOMe (bulbophyllanthrin) 3,7-DiOH-2,4-diOMe

158 159 160 161

3,7-DiOH-2,4-diOMe-9,10-dihydro (ephemeranthol B, flavanthridin) 4,5-DiOH-2,6-diOMe-9,10-dihydro 4,7-DiOH-2,3-diOMe 4,7-DiOH-2,3-diOMe-9,10-dihydro (ephemeranthol A)

162 163 164 165 166 167 168 169 170

84

Species

Reference

D. aphyllum D. chrysotoxum D. polyanthum Many recordsa Many recordsb D. nobile D. amplum D. nobile D. nobile D. aphyllum D. densiflorum D. loddigesii D. nobile D. plicatile (as Ephemerantha fimbriata (Blume) P.F.Hunt & Summerh.) D. nobile D. loddigesii

Zhang et al. (2008b) Yang et al. (2002) Hu et al. (2009)

Yang et al. (2007a) Tezuka et al. (1993) Hwang et al. (2010) Tezuka et al. (1993) Sritularak et al. (2011) Hu et al. (2008a) Ito et al. (2010) Honda and Yamaki (2000) Majumder and Pal (1992) Yang et al. (2007a) Yang et al. (2006c) Li et al. (2009d) Zhang et al. (2008d) Yang et al. (2007a) Majumder and Pal (1992) Yang et al. (2007a) Yamaki and Honda (1996) Tezuka et al. (1991), Chen et al. (2000)

2-OH-3,4,7-triOMe-9,10-dihydro 3-OH-2,4,7-triOMe 3-OH-2,4,7-triOMe-9,10-dihydro

D. nobile D. plicatile (as E. fimbriata) D. nobile D. plicatile (as E. fimbriata) D. draconis Rchb.f. D. longicornu D. loddigesii D. plicatile D. rotundatum (Lindl.) Hook.f. D. nobile D. chrysanthum D. chrysotoxum D. nobile D. nobile D. rotundatum D. nobile D. plicatile D. xantholeucum (as Ephemerantha lonchophylla) D. loddigesii D. nobile D. chrysotoxum D. nobile D. plicatile D. xantholeucum (as E. lonchophylla) D. nobile D. xantholeucum (as E. lonchophylla) D. nobile D. xantholeucum (as E. lonchophylla) D. nobile D. xantholeucum (as E. lonchophylla) D. nobile D. nobile D. nobile

Penta-O-substituted 2,3,5-TriOH-4,9-diOMe 2,3,7-TriOH-4,6-diOMe 2,3,7-TriOH-4,6-diOMe-9,10-dihydro 1,7-DiOH-2,5,6-triOMe 1,7-DiOH-2,5,6-triOMe-9,10-dihydro 2,5-DiOH-3,4,9-triOMe

D. nobile D. amplum D. amplum D. nobile D. nobile D. plicatile (as E. fimbriata)

Yang et al. (2007a) Majumder et al. (2008b) Majumder et al. (2008b) Yang et al. (2007a) Yang et al. (2007a) Tezuka et al. (1993)

Zhang et al. (2008d) Majumder et al. (2008b) Yang et al. (2007a) Yang et al. (2007a) Chen et al. (2008a) Fan et al. (2001) Ito et al. (2010) Lee et al. (1995), Yang et al. (2007a) Tezuka et al. (1993), Yamaki and Honda (1996) Yang et al. (2007a) Ito et al. (2010)

Ito et al. (2010) Yang et al. (2007a) Li et al. (2009d) Zhang et al. (2008d) Yamaki and Honda (1996) Chen et al. (2000) Hwang et al. (2010) Tezuka et al. (1991) Ye and Zhao (2002), Liu et al. (2007b) Tezuka et al. (1991) Yang et al. (2007a) Tezuka et al. (1991) Yang et al. (2007a) Yang et al. (2007a) Yang et al. (2007a)

DENDROBIUM

Table 622.4.  (Continued ). No.

Phenanthrene type and trivial name

Species

Reference

171

2,5-DiOH-4,9,10-triOMe (plicatol A)

D. nobile D. plicatile

Yang et al. (2007a) Honda and Yamaki (2000)

172

2,6-DiOH-1,5,7-triOMe (denthyrsinin)

173 174 175

2,6-DiOH-1,5,7-triOMe-2-O-rutinoside (denchryside A) 2,7-DiOH-1,5,6-triOMe (confusarin) 2,7-DiOH-3,4,6-triOMe

Fan et al. (2001) Zhang et al. (2005b) Ye et al. 2003

176

2,7-DiOH-3,4,6-triOMe-9,10-dihydro

177 178

4,7-DiOH-1,5,6-triOMe 2-OH-1,5,6,7-tetraOMe (chrysotoxene)

D. densiflorum D. thyrsiflorum D. chrysanthum Many recordsd D. amplum D. chrysotoxum D. amplum D. rotundatum D. nobile D. denneanum (as D. aurantiacum var. denneanum)

179

Phenanthrenone Ephemeranthone

D. xantholeucum (as E. lonchophylla)

Chen et al. (1999)

180

Phenanthro[4,5-bcd]pyran 2,6-DiOH-7-OMe-9,10-dihydro (flaccidin)

D. amoenum

Veerraju et al. (1989b)

181

Phenanthro[4,5-bcd]pyranone 2,7-DiOH-8-OMe (fimbriatone)

182

1-OH-2-OEt-7-OMe (crystalltone)

D. chrysotoxum D. fimbriatum D. nobile D. crystallinum

Yang et al. (2004a) Bi et al. (2003) Zhang et al. (2008d) Wang et al. (2009)

183

1,4-Phenanthraquinones 5-OH-7-OMe-9,10-dihydro (dendronone)

184 185

5-OMe-7-OH-9,10-dihydro 7-OH-2-OMe (densiflorol B)

186

7-OH-2-OMe-9,10-dihydro (ephemeranthoquinone)

187

5-OH-3,7-diOMe (denbinobin)

188 189

7-OH-2,8-diOMe (cypripedin) 7-OH-5,6-diOMe

D. cariniferum D. longicornu D. draconis D. chrysotoxum D. densiflorum D. plicatile D. xantholeucum (as E. lonchophylla) D. moniliforme D. nobile D. plicatile (as E. fimbriata) D. xantholeucum (as E. lonchophylla) D. densiflorum D. moniliforme

Chen et al. (2008c) Hu et al. (2008a) Sritularak et al. (2011) Li et al. (2009d) Fan et al. (2000, 2001) Yamaki and Honda (1996) Tezuka et al. (1991) Lin et al. (2001) Yang et al. (2007a) Tezuka et al. (1993) Tezuka et al. (1991) Fan et al. (2001) Bae et al. (2004)

190

1,4,5,8-Phenanthradiquinones 2,6-DiOMe (moniliquinone)

D. moniliforme

Lin et al. (2001)

191 192 193 194 195

Phenanthrenoid dimers and conjugates Amplumthrin Dendrochrysanene Denthyrsinol Denthyrsinone Flavanthrin

196 197 198

Loddigesiinol B [4,7-diOH-2-OMe-9,10-dihydro]-dimer [2-OH-3,4,7-triOMe-9,10-dihydro]-dimer

D. amplum D. chrysanthum D. thyrsiflorum D. thyrsiflorum D. amplum D. aphyllum D. loddigesii D. plicatile D. nobile

Majumder et al. (2008b) Yang et al. (2006c) Zhang et al. (2005b) Zhang et al. (2005b) Majumder et al. (2008b) Chen et al. (2008a) Ito et al. (2010) Yamaki and Honda (1996) Yang et al. (2007a)

Majumder et al. (2008b) Yang et al. (2002) Majumder et al. (2008b) Majumder and Pal (1992) Hwang et al. (2010) Yang et al. (2007d)

a

Many species records for moscatin are given by Yang et al. (2006b). D. aphyllum (Zhang et al. 2008b); D. draconis (Sritularak et al. 2011); D. loddigesii (Ito et al. 2010); D. nobile (Hwang et al. 2010); D. thyrsiflorum (Zhang et al. 2004); D. trigonopus (Hu et al. 2008b). c Mistakenly assigned trivial name of plicatol C when already published as rotundatin. d D. chrysotoxum, D. lohohense, D. plicatile (as Flickingeria fimbriata) (Yang et al. 2006b); D. denneanum (as D. aurantiacum var. denneanum) (Yang et al. 2007d); D. fimbriatum (Bi et al. 2003); D. nobile (Zhang et al. 2008d). b

reported cyanidin 3-glucoside in the red column of D. crocatum Hook.f., and a cyanidin glycoside in the purple flower of D. hasseltii (Blume) Lindl. (as D. cornutum Hook.f.). In more recent investigations, structures of a number of acylated cyanidin

3,7,3′-­glycosides have been described from the flowers of species and cultivars of Dendrobium, including D. bigibbum var. bigibbum Lindl. & Paxton, D. bigibbum var. superbum Rchb.f., D. williamsianum Day & Rchb.f., and D. ×superbiens (Tatsuzawa et al. 2005, 2006), D. 85

DENDROBIEAE

Table 622.5.  Fluorenones reported from species of Dendrobium. No.

Fluorenone type and trivial name

Species

Reference

199 200

Tri-O-substituted 2,5-DiOH-4-OMe (dengibsin) 2,7-DiOH-4-OMe (nobilone)

Many recordsa D. nobile

Zhang et al. (2007e)

201 202 203 204 205

Tetra-O-substituted 1,4,5-TriOH-7-OMe 1,4,7-TriOH-5-OMe (dendroflorin) 2,4,7-TriOH-4-OMe (denchrysan A) 2,4,7-TriOH-5-OMe 3,5-DiOH-2,4-diOMe (dengibsinin)

D. chrysotoxum Many recordsb Many recordsc D. chrysotoxum D. gibsonii Paxton

Chen et al. (2008b)

206

Penta-O-substituted 2,4,7-TriOH-1,5-diOMe

D. chrysotoxum

Yang et al. (2004b)

207

Fluoren-9-ol 2,5,9-TriOH-4-OMe (denchrysan B)

D. chrysanthum D. chrysotoxum D. thrysiflorum

Ye et al. (2003) Yang et al. (2004a) Zhang et al. (2004)

a

Yang et al. (2004b) Talapatra et al. (1985), Sargent (1987)

D. brymerianum Rchb.f., D. chrysanthum, D. chrysotoxum, D. densiflorum, D. fimbriatum, D. hancockii Rolfe, D. lohohense, D. nobile, D. thyrsiflorum (Yang et al. 2006b); D. denneanum (as D. aurantiacum var. denneanum) (Yang et al. 2006a); D. farmeri Paxton (Majumder and Chakraborti 1989); D. gibsonii (Talapatra et al. 1985). b D. brymerianum, D. chrysanthum, D. chrysotoxum, D. densiflorum, D. thyrsiflorum (Yang et al. 2006b); D. denneanum (as D. aurantiacum var. denneanum) (Yang et al. 2006a); D. nobile (Zhang et al. 2007e); D. trigonopus (Zhang et al. 2005c). c D. brymerianum, D. chrysotoxum, D. hancockii, D. lohohense, D. nobile (Yang et al. 2006b); D. chrysanthum (Ye et al. 2003); D. thyrsiflorum (Zhang et al. 2004).

Fig. 622.12.  Typical bibenzyl constituents of Dendrobium (see Table 622.3).

86

DENDROBIUM

Fig. 622.13.  Typical phenanthrene constituents of Dendrobium (see Table 622.4).

‘Pompadour’ (Williams et al. 2002), and D. ‘Pramot’ (Saito et al. 1994). This pattern of glycosylation is a characteristic feature of anthocyanins in Epidendroideae (Andersen and Jordheim 2006). lignans

Nine species of Dendrobium are reported to contain lignans. Most are well-known compounds, but the glycoside denchryside B from D. chrysanthum (Ye et al. 2004) and the bibenzyl-lignan conjugates longicornuol A (134) and trigonopols A (136) and B (137) from D. longicornu and D. trigonopus, respectively, are novel (Hu et al. 2008a,b). Other recorded occurrences are (+)-syringaresinol from D. aphyllum (Shao et al. 2008), (+)-lyoniresinol, dehydrodiconiferyl alcohol-4-Oβ-glucoside, and 7,7′-bis(4-hydroxy-3,5-dimethoxyphenyl)8,8′-dihydroxymethyltetrahydrofuran-4-O-β-glucoside from D. chrysanthum (Ye et al. 2004), (+)-syringaresinol from D. chrysotoxum (Gong et al. 2006), (–)-medioresinol and (–)-pinoresinol from D. loddigesii (Ito et al. 2010), episyringaresinol and its 4′-O-βglucoside from D. longicornu (Hu et al. 2008a), (+)-pinoresinol, (+)-syringaresinol, and acanthoside B from D. moniliforme (Zhao et al. 2003a,b), lirioresinol A, medioresinol, pinoresinol, and

syringaresinol from D. nobile (Zhang et al. 2008d), (+)-pinoresinol and (+)-syringaresinol from D. plicatile (as Ephemerantha fimbriata) (Tezuka et al. 1993), and (+) and (–)-syringaresinol from D. trigonopus (Zhang et al. 2005c; Hu et al. 2008b). phenolics

In addition to stilbenoids, other types of phenolic compounds present in Dendrobium include phenolic acids, phenylethanoids, phenylpropanoids, and phenolic esters. Recorded occurrences of simple phenolic compounds comprise syringic acid, p-hydroxybenzylacetic acid, and p-hydroxyphenylpropionic acid methyl ester from D. aphyllum (Chen et al. 2008a; Shao et al. 2008; Zhang et al. 2008b), methyl 2,4-dihydroxy-3,6-dimethylbenzoate from D. chryseum (Zheng et al. 2000a), 3,4-dihydroxybenzoic acid, vanillin, 3,4-dimethoxybenzoic acid and its methyl ester from D. chrysotoxum (Yang et al. 2002; Gong et al. 2006), p-hydroxybenzaldehyde from D. denneanum (as D. aurantiacum var. denneanum) (Liu et al. 2009b), p-hydroxybenzoic, 3-hydroxy-2-methoxy-5,6-­ dimethylbenzoic, and syringic acids from D. crystallinum (Wang et al. 2008, 2009), p-hydroxybenzaldehyde and p-hydroxybenzoic acid from D. falconeri (Sritularak and Likhitwitayawuid 2009), 87

DENDROBIEAE

Fig. 622.14.  Selected fluorenone constituents of Dendrobium (see Table 622.5).

p-hydroxybenzaldehyde from D. gratiosissimum (Wang et al. 2007), methyl 2,4-dihydroxy-3,6-dimethylbenzoate, ethyl 3-formyl2,4-dihydroxy-6-methylbenzoate, and gallic acid from D. longicornu (Hu et al. 2008a; Li et al. 2009c), p-methoxybenzaldehyde, vanillin, and vanilloloside from D. moniliforme (Lin et al. 2000; Zhao et al. 2003a), acetovanillone, α-hydroxypropiosyringone, protocatechuic acid, syringaldehyde, syringylethanone, syringic acid, and vanillin from D. nobile (Ye and Zhao 2002; Zhang et al. 2005d), and 3,4,5-trimethoxybenzoic acid from D. plicatile (as Ephemerantha fimbriata; Tezuka et al. 1993). Methyl 2,4-dihydroxy3,6-dimethylbenzoate (methyl β-orsellinate; D. chryseum, D. longicornu) and ethyl 3-formyl-2,4-dihydroxy-6-­methylbenzoate (ethyl haematommate; D. longicornu) are typically found as constituents of lichens, and may be artefacts (Hylands and Ingolfsdottir 1985). The phenylethanoid derivatives icariside D2 and salidroside occur in D. aphyllum (Shao et al. 2008; Zhang et al. 2008b) and 3-hydroxy-4-methoxyphenylethanol in D. nobile (Zhang et al. 2005d). Reports of simple phenylpropanoid derivatives include dihydro-p-coumaric acid from D. farmeri (Majumder and Chakraborti 1989), dihydroconiferyl alcohol, coniferyl aldehyde, and eugenyl O-β-glucoside from D. longicornu (Hu et al. 2008a; Li et al. 2009c), coniferyl aldehyde, dihydroconiferyl alcohol, dihydroconiferyl dihydro-p-coumarate, dihydro-p-coumaric acid, and 2-hydroxyphenylpropanol from D. nobile (Zhang et al. 2005d), dihydroconiferyl dihydro-p-coumarate and dihydro-pcoumaric acid from D. plicatile (as Ephemerantha fimbriata; Tezuka et al. 1993), and coniferyl alcohol from D. trigonopus (Hu et al. 2008b). 88

Presence of 2-glucosyloxycinnamic acids in Dendrobium was first described by Dahmén et al. (1975), who isolated the (E)- and (Z)- forms of densifloroside from methanol extracts of D. densiflorum. Other sources of 2-glucosyloxycinnamic acids are D. chrysotoxum, D. denneanum (as D. aurantiacum var. denneanum), and D. thyrsiflorum (Zhang et al. 2005a, 2006a; Yang et al. 2007b,c). Under conditions of mild acid-hydrolysis, these compounds afford the corresponding coumarins. Methods for qualitative and quantitative analyses of 2-glucosyloxycinnamic acids in Dendrobium have been developed (Yang et al. 2007c; Zhang et al. 2006a). Phenolic esters derived from cinnamic acids and long-chain alcohols have been isolated from more than 10 species of Dendrobium. The first of these to be characterized was defuscin (n-triacontyl p-coumarate), which Talapatra et al. (1989) obtained from whole-plant extracts of D. defuscescens Griff. The same compound was later reported from D. chryseum (together with n-­octacosyl ferulate; Zheng et al. 2000a), D. chrysotoxum (Gong et al. 2006), D. densiflorum (Zheng et al. 2000b), D. fimbriatum (Talapatra et al. 1992; Bi et al. 2001a), D. gratiosissimum, also with n-octacosyl ferulate (Wang et al. 2007), and D. thyrsiflorum (Zhang et al. 2005a). Sources of other alkyl cinnamates include D. crystallinum (Wang et al. 2008), D. falconeri (Sritularak and Likhitwitayawuid 2009), D. fimbriatum (Bi et al. 2003), D. longicornu (Li et al. 2009c), and D. moniliforme (Lin et al. 2000; Bi et al. 2004). polysaccharides

Investigation of beneficial effects ascribed to some Dendrobium species used in traditional Chinese medicine has prompted studies of macromolecules as well as smaller constituents. The former include polysaccharides, which are typically obtained from stem material either by hot water extraction, or by Soxhlet extraction in organic solvents followed by hot water extraction of the resulting residues. Further purification of polysaccharide-containing extracts can be achieved using column chromatography on various media. Crude, semi-purified, and purified polysaccharides obtained from stem extracts of Dendrobium have been evaluated in bioassays, mainly for immunostimulant and antioxidant activity. Partial characterization of polysaccharide structures requires a combination of chemical and spectroscopic methods. Most Dendrobium polysaccharides are high molecular-weight molecules comprising a backbone of sugar residues arranged in repeating units, and branches that contain additional sugar residues, some of which may be acetylated. Three polysaccharides (AP-1, AP-2, and AP-3) obtained from stems of D. aphyllum are O-acetylglucomannans that showed immunostimulant activity in bioassays (Zhao et al. 1994). Studies on the polysaccharides of D. catenatum Lindl., which are also O-acetylglucomannans, were published under the synonyms D. huoshanense Z.Z.Tang & S.J.Cheng (Zha et al. 2007; Hsieh et al. 2008) and D. officinale Kimura & Migo (Hua et al. 2004). Hsieh et al. (2008) demonstrated that the polysaccharide fraction obtained from stem mucilage of this species stimulated production of cytokines and growth factors in murine splenocytes, whereas a deacetylated preparation of this fraction failed to induce cytokine production. The polysaccharide (HPS-1B23) obtained by Zha et al. (2007) from D. catenatum (as D. huoshanense) had high immunostimulant activity. A high molecular weight

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(150 kDa) polysaccharide (DCL) obtained from stems of D. chrysotoxum showed antioxidant, anti-hyperglycemic, and immunostimulant activities (Zhao et al. 2007). Antioxidant activity of polysaccharides extracted from D. denneanum (Fan et al. 2009; Luo et al. 2011) and D. fimbriatum (as D. fimbriatum var oculatum) has also been reported (Luo and Fan, 2011). In an early paper in this field, Wang et al. (1988) described three O-acetylglucomannan polysaccharides from D. moniliforme (as D. candidum Wall ex Lindl.). Polysaccharides obtained from stem extracts of D. nobile have attracted considerable attention, with several studies reporting on their immunostimulant (Wang et al. 2010a,b,c) and antioxidant activity (Luo et al. 2009, 2010). Extensive structural analysis of immunostimulant polysaccharides (DNP-W2, DNPW3, and DNP-W5) has been carried out, revealing that DNPW2 and DNP-W3 are a 2-O-acetylgalactomannoglucan and a rhamnoarabinogalactan, respectively (Wang et al. 2010b,c). DNPW5 is a complex pectic polysaccharide with a backbone structure of galacturonosyl and rhamnosyl residues (Wang et al. 2010a). Polysaccharides present in different species of Dendrobium or in accessions of the same species from different locations can be compared using saccharide mapping (Xu et al. 2011). scent constituents

Chemical composition data for the floral fragrances of numerous species of Dendrobium are available, largely as a result of analytical work carried out by Kaiser (1993, 2011) using GC-MS (gas chromatography coupled to mass spectrometry). Species included are D. anosmum, D. antennatum Lindl., D. brymerianum, D. cariniferum, D. chrysotoxum, D. delacourii Guillaumin, D. lichenastrum (F.Muell.) Rolfe, D. moniliforme (white- and purple-flowered forms), D. monophyllum F.Muell., D. nobile, D. pugioniforme A.Cunn. ex Lindl., D. schoeninum Lindl. (as D. beckleri F.Muell.), D. spectabile (Blume) Miq., D. treacherianum Rchb.f. ex Hook.f. (as Epigeneium lyonii (Ames) Summerh.) D. trigonopus, D. unicum Seidenf., D. virgineum Rchb.f., and D. williamsonii. Scent composition of D. anosmum (also cited using the synonym D. superbum Rchb.f.) is of particular interest because of the presence of the so-called ‘raspberry ketone’, 4-(4-hydroxyphenyl)butan-2-one, which is a specific attractant for the male melon fly, Dacus cucurbitae. Detecting this component proved elusive; neither Flath and Ohinata (1982) nor Kaiser (1993) were able to confirm its presence, although subsequent analysis of a specimen sourced from Mount Kinabalu found it at levels of 0.1% (Kaiser 2011). Nishida et al. (1993) detected it as a volatile component in an extract of D. anosmum petals (as D. superbum), and as a sequestered compound in the rectal glands of Dacus cucurbitae males. Although D. anosmum is sometimes known as the ‘unscented dendrobium’, Kaiser (2011) deemed this term inappropriate, owing to the ‘strong and distinct fruityfloral and somewhat herbal scent reminiscent of raspberry and rhubarb’. The main scent component is pentadecan-2-one, which accounts for up to 50% of the volatiles (Kaiser 1993, 2011). Flowers of D. sinense Tang & F.T.Wang produce (Z)-11eicosen-1-ol as a volatile component. This compound is a major constituent of the alarm pheromones of Asian and European honey bees and appears to attract the pollinator, the hornet Vespa bicolor, by chemical mimicry (Brodmann et al. 2009). Other volatiles detected in the floral scent of D. sinense were benzyl

acetate, benzyl alcohol, octadecan-1-ol, and eicosan-1-ol. Major fragrance components from scents of several members of D. section Dendrocoryne, comprising D. gracilicaule F.Muell., D. kingianum, D. speciosum Sm., and the natural hybrids D. ×delicatum (F.M.Bailey) F.M.Bailey (D. kingianum × D. tarberi), and D. ×suffusum Cady (D. gracilicaule × D. kingianum), were identified using GC-MS and discussed in relation to bee pollination syndromes (Adams 1988). The compounds detected, which were present in a non-specific pattern in these taxa, included methyl esters of butanoic and octanoic acids, and simple aromatics (1,4-dimethoxybenzene, ethoxybenzene, and 3-methoxybenzaldehyde). (NV, RG)

Phylogenetics In recent years, several authors have published molecular phylogenetic studies of Dendrobium (Yukawa 2001; Yukawa et al. 1993, 1996, 2000; Clements 2003, 2006; Wongsawad et al. 2005; Burke et al. 2008; Sathapattayanon 2008; Schuiteman 2011), reviewed by Adams (2011). No single, well-resolved result has emerged from these studies, but they agree in many respects. It is now wellestablished, as first observed by Yukawa et al. (1993) employing plastid DNA restriction site analysis, that Dendrobium consists of three main clades: 1. The first and smallest comprises only D. sect. Sarcopodium (formerly the genus Epigeneium), which is sister to two much larger clades, which are sometimes called the northern and the southern clade or alternatively the Asian and Australasian clade. 2. The northern/Asian clade here comprises D. sects. Amblyanthus, Aporum, Calyptrochilus, Conostalix, Dendrobium, Distichophyllae, Formosae, Fytchianthe, Pedilonum, and Stachyobium. 3. The southern/Australasian clade here comprises D. sect. Biloba, Brevisaccata, Cadetia, Crinifera, Dendrocoryne, Diplocaulobium, Eleutheroglossum, Fugacia, Grastidium, Herpethophytum, Latouria, Lichenastrum, Macrocladium, Microphytanthe, Monophyllaea, Phalaenanthe, Pleianthe, Rhizobium, and Spatulata. Yukawa et al. (2000), using plastid matK sequences, obtained trees that largely supported the existing morphology-based sectional classification of Dendrobium, but their sampling was limited. They also found that Dendrobium secundum (Blume) Lindl., the type species of D. sect. Pedilonum, was sister to a combined clade of two species (in the present classification), one each in D. sects. Calyptrochilus and Aporum, which in turn formed two strongly supported subclades. This result foreshadowed later DNA studies (especially Clements 2003), which demonstrated that the Pedilonum alliance, comprising the traditionally circumscribed D. sects. Calcarifera, Calyptrochilus, Cuthbertsonia, Dolichocaulon, Oxyglossum, Pedilonum, and Platycaulon, which were generally considered to be closely related, is not monophyletic. Yukawa (2001) presented a tree based on plastid matK and ITS nuclear ribosomal (nr) DNA data and including 93 species. He noted that matK and ITS when analysed separately produced similar results. The combined tree shows several morphologically plausible and often well-supported clades: 1) Conostalix with Distichophyllae and Formosae; 2) Biloba (as Monanthos) with Grastidium, Herpethophytum, and Pleianthe; 3) Spatulata with Eleutheroglossum 89

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and Phalaenanthe; 4) Dendrocoryne with Rhizobium, Lichenastrum, and Monophyllaea (as Australorchis), and 5) Diplocaulobium with Crinifera (formerly genus Flickingeria) and Microphytanthe; the last two are more disparate morphologically than the others. The strongly supported sister-group relationship displayed between D. sects. Calyptrochilus (as here circumscribed) and Aporum (ibid.) is difficult to explain in terms of morphology. Dendrobium sects. Densiflora (as Callista), Pedilonum, and Dendrobium appear to be polyphyletic (but D. leptocladum Hayata is mistakenly listed as a species of D. sect. Grastidium rather than D. sect. Dendrobium). Dendrobium microglaphys Rchb.f. (D. sect. Amblyanthus) is shown as being nested in a strongly supported clade comprising species allied to D. densiflorum Lindl., whereas another species of D. sect. Amblyanthus, D. melanostictum Schltr., is sister to this clade with weak support. Morphologically and biogeographically, this association of D. sect. Amblyanthus with part of D. sect. Densiflora is hard to explain. Clements (2003) used ITS nrDNA data and a sample of 66 species (not counting outgroup taxa) to produce an analysis in which he focused on species of the Pedilonum alliance as defined above. He confirmed the strongly supported sister-group relation between D. sect. Calyptrochilus (as here circumscribed) and D. sect. Aporum (ibid.), and found that D. sects. Pedilonum and Calcarifera, as previously delimited, are polyphyletic. Clements also noted that D. sect. Bolbodium (as Bolbidium) was nested within D. sect. Crumenata (as Rhopalanthe). Wongsawad et al. (2005) concentrated on D. sects. Dendrobium and Densiflora (as Callista). Monophyly could not be established for these two sections. In the case of D. sect. Dendrobium, this is mainly due to a small group of species not being part of the clade that contains by far the greater number of species of this section (the main clade). These outlying species, which include D. capillipes Rchb.f. and D. albosanguineum Lindl., do not display any obvious morphological characters that would support their exclusion from the main clade. Dendrobium senile C.S.P.Parish ex Rchb.f., which is traditionally included in D. sect. Dendrobium, does not appear as part of the main clade either, but this species is morphologically unusual, for example in the long white hairs on the leaves. Wongsawad et al. found that species of D. sects. Stuposa and Breviflores were nested in D. sect. Dendrobium. Dendrobium sect. Densiflora as previously circumscribed appears to include three clades: 1) D. chrysotoxum Lindl., which is nested in D. sect. Dendrobium; 2) D. lindleyi Steud. and D. jenkinsii Wall. ex Lindl., two similar species that form a strongly supported clade; 3) species related to D. densiflorum Wall. ex Lindl., which form a well-supported clade sister to D. sulcatum Lindl., usually included in this section. These three clades are unresolved. Clements (2006), using ITS nrDNA data with a well-balanced sampling of 118 taxa of Dendrobium s.l., produced a tree that largely confirmed the results by Yukawa (2001). He included a broader sample of species from New Caledonia, which showed little support for the several endemic sections split off from D. sect. Macrocladium. An unusual, although weakly supported, result is the inclusion of D. herpethophytum Schltr. in the New Caledonian clade rather than in the Grastidium alliance. Clements included two species of D. sect. Fugacia (D. coxii F.M.Bailey [= D. 90

lacteum Kränzl.] and D. aff. spurium (Blume) J.J.Sm.) in his analysis. This is one of the most distinctive and seemingly natural sections in the genus, but in Clements’s tree, although with weak bootstrap support, the two species do not form a single clade but are nested in separate parts of the Australasian clade. Burke et al. (2008) sampled the Australian sections, including all species previously included in D. sect. Dendrocoryne, and species representing the main vegetative types of D. sect. Rhizobium. In a consensus tree, bootstrap support for the northern and southern clades was 100%. Dendrobium section Macrocladium was sister to all other sections in the southern clade as found for the Australasian sections sampled by Yukawa (2001) and Clements (2006). Dendrobium sects. Cadetia, Diplocaulobium, Spatulata, and Phalaenanthe formed an internally unresolved sister group to the remaining sections. Two Dendrocoryne species, D. aemulum and D. callitrophyllum, nest in D. sect. Crinifera (Flickingeria), and these are sister to D. sect. Australorchis, which is in turn sister to D. sect. Rhizobium and the majority of D. sect. Dendrocoryne species. Together with morphology, this provides evidence that D. sect. Lichenastrum, sometimes regarded as part of D. sect. Rhizobium, is a distinct section. A phylogram based on morphological characters was well resolved, showing geographical patterns (Adams et al. 2006). Natural hybridization is well documented in D. sect. Dendrocoryne and Rhizobium and may contribute to the lack of monophyly (Adams 2011). Burke et al. (2008) identified pseudogenes among the ITS sequences but suggested that these did not strongly influence the tree produced. Satthapatayanon (2008) provided a comprehensive phylogenetic analysis of D. sect. Formosae, utilizing both plastid matK and ITS nrDNA sequence data. Separate analyses indicated that D. sect. Formosae is biphyletic. Although the ITS tree shows strong support for a sister group relation between D. sections Densiflora (p.p., as Callista) and Amblyanthus, as earlier found by Yukawa (2001), the matK tree has D. sect. Amblyanthus sister to D. sect. Stachyobium, but with weak support, and D. sect. Densiflora (p.p.) sister to D. sects. Distichophyllae + Conostalix, also with weak support. In summary, although much of the section-based classification by Schlechter (1911–1914) has been supported by DNAbased analyses, relationships between and within the sections have not often been investigated. The cladogram shown here (Fig. 622.15, 622.16) displays most of the features of the studies discussed above. The inferred grouping of D. sects. Biloba, Grastidium, and Herpethophytum, although with weak bootstrap support, is morphologically plausible. All have slender, thin stems with multi-annual apical growth, and short, one- or twoflowered inflorescences. The cladogram indicates that Grastidium may not be monophyletic. The grouping of D. sects. Spatulata, Eleutheroglossum, and Phalaenanthe is in agreement with their vegetative and floral morphology as well. There is some evidence that D. sect. Spatulata is paraphyletic when D. sects. Phalaenanthe and Eleutheroglossum are not included in it. The position of the strongly supported D. sects. Diplocaulobium and Crinifera within a clade also comprising D. sects. Dendrocoryne, Lichenastrum, Rhizobium, and Monophyllaea is morphologically unexpected, and, given the weak support at the basal nodes, may not be reliable. As also

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found by Burke et al. (2008), there are indications that D. sects. Dendrocoryne and Rhizobium are not monophyletic, and that D. sect. Rhizobium is nested within D. sect. Dendrocoryne. Moreover, D. sect. Lichenastrum does not group with the species of D. sect. Rhizobium included here. Dendrobium sect. Macrocladium, which is endemic to New Caledonia and New Zealand, is sister to the remainder of the southern clade, with strong support for the sister clade. The northern clade as a whole is well supported. As also found by Yukawa (2001) and Clements (2003, 2006), three clades, here (Fig. 622.16) named Calyptrochilus, Aporum, and Pedilonum, have strong support, but they conflict with traditional circumscriptions of the sections bearing these names. Dendrobium sect. Calyptrochilus includes all species of the morphologically circumscribed D. sects. Calyptrochilus (s.s.), Oxyglossum, Cuthbertsonia, Dolichocentrum, Calcarifera (p.p.), and Pedilonum (p.p.). These former sections do not all form clades, although there is strong support for certain subsets of species. The previously circumscribed D. sects. Pedilonum and Calcarifera in particular are polyphyletic. Dendrobium sect. Aporum comprises four morphologically circumscribed former sections: Aporum (s.s.), Strongyle, Bolbodium, and Crumenata. Two strongly supported subclades can be seen within D. sect. Aporum, one containing D. crumenatum Sw. among other species, one containing D. reflexitepalum J.J.Sm. among other species. The first subclade contains species with partly swollen stems, traditionally included in D. sects. Crumenata and Bolbodium (D. hymenanthum Rchb.f.), but also two species lacking swollen stems (D. pseudocalceolum J.J.Sm. and D. reginanivis P.O’Byrne & J.J.Verm.), which were traditionally included in D. sect. Aporum. In the second subclade we find species lacking swollen stem internodes, traditionally included in D. sects. Aporum (s.s.) and Strongyle. Species with laterally flattened leaves are found in both subclades. The Pedilonum clade (Fig. 622.16) includes D. sects. Pedilonum (s.s.), Calcarifera (p.p.), and Platycaulon. Even within this clade, species of D. sects. Pedilonum and Calcarifera as previously circumscribed do not form clades. For example, D. hasseltii (Blume) Lindl. (previously in D. sect. Pedilonum) forms a strongly supported clade with D. derryi Ridl. (D. sect. Calcarifera) and D. platygastrium Rchb.f. (D. sect. Platycaulon), and not with the subclade comprising D. secundum (Blume) Lindl. and D. amethystoglossum Rchb.f. (both D. sect. Pedilonum). Similarly, D. mutabile (Blume) Lindl. and D. sanguinolentum Lindl. (both D. sect. Calcarifera) do not group with D. derryi. It would seem that there has been much convergent evolution in the set of species previously ascribed to D. sects. Pedilonum and Calcarifera. Dendrobium sects. Conostalix and Distichophyllae form a strongly supported clade, but this is not, as in most other studies, sister to D. sect. Formosae (Fig. 622.16). This may be caused by the small sample of species of these sections in the present cladogram. Morphology indicates that these sections are closely related (Sathapattayanon 2008). Dendrobium sect. Stachyobium forms a strongly supported clade with high sequence divergence, but perhaps for that reason does not have a well-supported sister group relationship. It is shown (Fig. 622.16) as sister to the remainder of the northern clade, but in other published cladograms it is usually shown nested within

the northern clade (but always with weak support for any particular sister-group relationship). In terms of floral morphology, this clade is unusual within the northern clade in having a labellum with a distinct, elongate, basal callus or with lamellate keels, characters otherwise only seen in the southern clade. A number of species, in particular D. trigonopus Rchb.f., D. jerdonianum Wight, and D. senile, are not readily assigned to any of the traditional sections on morphological grounds, and are here considered ‘unplaced’ due to the weak support for their position in the cladogram. Analyses based on different genome regions may bring more clarity, but it is also possible that ancient and recent (Adams 2011) hybridization events have produced a reticulate pattern, which cannot adequately be captured in a tree-like structure. In that case, it is equally impossible to produce a classification that corresponds to a simple nested hierarchy. Relationships of Dendrobium subgroups may well be more complex than any standard method of classification is capable of representing; see Knapp et al. (2004) for a phylogenetic classification of Nicotiana L. (Solanaceae), which includes many species of demonstrated hybrid origin. (AS, PA)

Biogeography There is a clear biogeographical signal in the main clades of Dendrobium. All sections in the northern clade are poorly or not represented in Australia and New Caledonia. All sections that are well represented there belong to the southern clade. Six sections are (semi-)endemic to Australia + New Caledonia: Dendrocoryne, Eleutheroglossum, Lichenastrum, Macrocladium, Monophyllaea, and Rhizobium. Some southern sections are clearly centred on New Guinea. Dendrobium sect. Microphytanthe is endemic to New Guinea, and D. sects. Biloba, Brevisaccata, Cadetia, Diplocaulobium, Grastidium, Herpethophytum, Latouria, Pleianthe, and Spatulata have distributions extending to the north or south. Finally, a number of southern clade sections are neither largely confined to Australia + New Caledonia nor centred on New Guinea: D. sects. Crinifera, Fugacia, and Phalaenanthe. Dendrobium sect. Crinifera is much more species-rich in northern and western areas and as such would be more appropriate as part of the northern clade. Within the northern clade a fairly clear distinction can be made between a set of western and eastern sections, where ‘western’ means the Philippines and all areas to the west of Sulawesi. Western sections of Dendrobium include Aporum, Conostalix, Dendrobium, Distichophyllae, Formosae, Fytchianthe, Pedilonum, and Stachyobium. Eastern sections of Dendrobium are Amblyanthus and Calyptrochilus. However, only D. sects. Fytchianthe and Stachyobium are exclusively western, and none is exclusively eastern. Dendrobium sect. Sarcopodium, the sister group of the combined northern and southern clades, is endemic in the western area just defined. The origins of Dendrobium are uncertain. Brieger and others attempted to explain some of these patterns by postulating a Gondwana origin. Wood (2006) favoured a bicentric origin involving the northward movement of India and tectonic changes in the Australian and Philippine plates. In this view, the earliest Dendrobieae may have diversified in the early Tertiary or more 91

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Fig. 622.15.  Phylogenetic relationships in Dendrobium based on parsimony analysis of ITS sequence data (in part provided by NCB Naturalis DNA barcoding lab). Bootstrap support values of 1000 replicates shown. I: Sarcopodium clade; II: southern clade; III: northern clade (shown in Fig. 622.16). (Schuiteman, unpublished).

92

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Fig. 622.16.  Phylogenetic relationships in Dendrobium, northern clade. Letters refer to traditional sectional classification: B. D. sect. Brevisaccata; C. D. sect. Calcarifera; D. D. sect. Densiflora; P. D. sect. Pedilonum; S. D. sect. Stuposa.

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recently, in a part of the present distribution of the Asian clade, where as yet no fossils have been found. The only date assigned to Dendrobium is a minimum of about 23 mya (Early Miocene), based on the first vegetative orchid fossil, the New Zealand D. winikaphyllum Conran, Bannister & Lee (Conran et al. 2009). This consists of leaves that resemble many present-day dendrobiums and may indicate a longer history. Gustaffson et al. (2010) used D. winikaphyllum to estimate that Dendrobium is about 32 million years old with a Bayesian confidence interval of 25 to 40 mya and a 90% probability. Development of all of Dendrobium almost certainly postdates the Gondwana breakup c. 80 million years ago. A vicariance explanation has been used for many major floral and faunal groups of eastern Australia, New Caledonia, and New Zealand, which is supported by endemicity, geological models, and molecular clock dating (Ladiges and Cantrill 2007); however, more evidence is needed to explain convincingly the distribution of Dendrobium and other taxa with a similar distribution pattern. In the midEocene to late Oligocene, land persisted in a complex geological history, with connections between these areas (Schellart et al. 2006; Ladiges and Cantrill 2007) allowing old biota to persist in an admixture with more recently evolved endemics. New Caledonia and New Zealand were finally isolated by the late Oligocene (Grandcolas et al. 2008; Trewick et al. 2007). New Caledonia itself was wiped clean of its vegetation until about 32 million years ago because it was totally submerged. The New Caledonian and eastern Australian orchid flora share terrestrial genera and species, including species of several Dendrobium sections, whereas D. sect. Macrocladium (except the New Zealand species D. cunninghamii Lindl.) and other Dendrobium species are endemic to New Caledonia. Speciation in eastern Australia since the mid- to late Tertiary is associated with contraction of rain forest and a progressively drier climate (Greenwood 1996). New Guinea, with many habitats created by orogenic activity, has numerous endemics, mainly in the southern/Australasian clade, but also in D. sect. Calyptrochilus from the northern clade, which indicates at least two separate colonization events. It has a complex geological history of land connections in the Middle Eocene, 40–45 mya, continuing until 10,000 years ago, allowing dispersal in several directions to and from Asia in the north and Australia and the Pacific to the south and east. (AS, PA)

Ecology Dendrobium species are predominantly epiphytes, less often lithophytes, and are rarely exclusively terrestrial. Throughout their large area of occurrence they may be encountered wherever epiphytic orchids occur. The greatest concentration of species is found in primary forests between 800 and 1500 m above sea level, with few species occurring above 2200 m anywhere, except in New Guinea. On Mount Kinabalu in Sabah, North Borneo, only six out of 70 locally occurring species of Dendrobium are found above 1800 m with none above 2400 m (Wood et al. 2011). In the Himalayas, D. fimbriatum Hook. has been reported from as high as 3660 m (Pearce and Cribb 2002), but this is exceptional, as only a handful of Dendrobium species occur there above 94

2500 m. In New Guinea, around 30 species of Dendrobium occur at or above 3000 m (Schuiteman, unpublished). At lower elevations, species are common in rain forest, for example in New Guinea, whereas in tropical areas with a strongly seasonal climate relatively few Dendrobium species occur in the lowlands and then usually in localities with a more humid microclimate, such as river valleys, gorges, and hill tops. In general, Dendrobium species are not found in deep shade. Even in the relatively dry, semi-deciduous forests of continental Asia, most species occur high up in the trees in relatively exposed positions. They mainly grow on the larger branches and the upper part of tree trunks. However, in areas with a pronounced dry climate combined with low humidity, as in much of Australia, they are more often found in shaded locations. Specialized twig epiphytes are uncommon in the genus and are almost limited to species of D. section Stachyobium in continental Asia and of D. sects. Calyptrochilus (e.g. D. masarangense Schltr.) and Cadetia in New Guinea and neighbouring areas. The latter may even occur as epiphytes on other orchids. Open grassy forests and woodlands support few species, with frequent fires limiting populations. Some lithophytic species (e.g. D. speciosum Sm.) in fire-prone, drier areas of eastern Australia have adapted to fire with the capacity to regrow from basal shoots after being burned to ground level. Epiphytes tend to be more fire-sensitive, with populations re-establishing from surviving pockets of vegetation. Few Dendrobium species are subjected to frost. The few exceptions are found at the fringe of the distribution, for example D. falcorostrum Fitzg. in Nothofagus Blume forests of eastern Australia, D. cunninghamii Lindl. in New Zealand, and D. moniliforme (L.) Sw. in Japan. In D. sects. Dendrocoryne and Rhizobium, D. speciosum, D. kingianum Bidwill ex Lindl., and D. striolatum Rchb.f. are sometimes exposed to cold harsh winters and summer temperatures of over 40 °C, close to the lethal limit in hot summers. In the mountains of New Guinea, a few species of Dendrobium regularly experience brief spells of frost, such as D. dekockii J.J.Sm. and D. brevicaule Rolfe. The latter occurs up to 4000 m above sea level and holds the elevational record in the genus. Among the more exceptional habitats where species may occur are: mangrove forests (to which in New Guinea the inconspicuous D. viridiflorum F.M.Bailey is almost restricted); roadside trees in big cities (D. crumenatum Sw. in Singapore and Palembang); subalpine shrubland between 3000 and 4000 m above sea level (D. brevicaule in New Guinea); boggy areas with sphagnum moss (D. lobbii Teijsm. & Binn.); coconut palms along the coast (D. bifalce Lindl. in New Guinea); ultrabasic, rocky soil among shrubs (D. verruciferum Rchb.f. in New Caledonia); bare, vertical sides of large boulders (D. venustum Teijsm. & Binn. in Laos and D. sect. Dendrocoryne species in Australia); and tree-fern trunks in montane grassland (D. cuthbertsonii in New Guinea). (AS, PA)

Pollination The main pollination syndromes in Dendrobium involve both deception and rewards. Information on pollination in the genus is incomplete and uneven (Cingel 2001), with a paucity of

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observations on Asian species and more comprehensive information on Australian ones. Much reporting of pollination in orchids generally contains incomplete descriptions of pollination events and speculation based on structural features of flowers and potential pollinators. Pollination is often confused with visitation of flowers. Criteria and definitions for confirmed, probable, and suggested pollinator status recommended and used to assess pollination reports (Adams and Lawson 1993) have been applied here. The minimum criteria for confirmed pollinator status include observation of removal of pollinia by the vector, travel to a flower of the same species, deposition of pollinia on the stigma, and accurate identification of orchid and vector. Australian species have been the most studied because they flower near ground level and also because of relative ease of access along the more populated eastern coast. In contrast to the highly specific pollinators described for many terrestrial Australian orchids, the major pollination syndromes for epiphytes are nonspecific and involve bees of the genera Trigona, Lasioglossum, Hylaeus, and Euryglossina (Adams and Lawson 1993). Eusocial Trigona bees are by far the most important confirmed or probable pollinators for D. sects. Dendrocoryne, Lichenastrum, and Monophyllaea. Trigona is confirmed for D. kingianum Bidwill ex Lindl., several D. speciosum Sw. varieties, and D. adae F.M.Bailey, and has been suggested for varieties of D. teretifolium R.Br. For D. sect. Dendrocoryne, the pollination syndrome involves mass flowering in spring and heavy emission of fragrance collected by Trigona under particular weather conditions of warm days (temperatures above 24 °C) with light breezes. Homalictus bees are confirmed pollinators for D. jonesii Rendle. Some species may be pollinated by more than one species. For example, the introduced honey-bee, Apis mellifera, is also confirmed for larger-flowered D. kingianum plants and other species with labellar dimensions that allow access to the pollen and stigma. Most members of D. sect. Dendrocoryne are nectarless, but D. kingianum may produce nectar as the flowers age. In D. sect. Rhizobium, most species are fragrant, but only D. linguiforme Sw. has confirmed pollinators, two species of Euryglossina. In D. sect. Grastidium, Trigona bees have been suggested as pollinators of D. luteocilium Rupp from northern Australia. Many species in D. sect. Spatulata have flowers that are fragrant and have bright colours on different parts. Dendrobium canaliculatum R.Br. in northern Australia is pollinated by an undescribed Hylaeus species. In D. sect. Monophyllaea, most species emit a fragrance, and bee pollination is likely, but there are no confirmed or probable pollinators reported. Insect visitors have been observed in D. sect. Phalaenanthe, but there are no descriptions of pollination events despite many years of field observations. In D. sects. Calyptrochilus and Pedilonum, the brightly coloured flowers are usually open for weeks to several months. Many display characteristics of ornithophily, such as tubular, firm-textured, red or orange (less often purple, white, yellow or green) flowers that produce copious nectar and lack fragrance. Honeyeaters (Meliphagidae) have been observed visiting the flowers of Dendrobium lawesii F.Muell., D. pseudoglomeratum T.M.Reeve & J.J.Wood, and D. smillieae F.Muell., all in D.

sect. Calyptrochilus, but a full pollination sequence has not been described. Dendrobium cuthbertsonii F.Muell. in the same section has colour polymorphism of the flowers. Plants growing in proximity may have different combinations of colours. It has been suggested that this species imitates Rhododendron flowers and that the colour polymorphism prevents the pollinators, most likely honeyeaters, from learning to avoid this species, as it is presumably unrewarding. Dendrobium cuthbertsonii is also remarkable for its extremely long-lived flowers, which may last for up to nine months. Honeyeaters do not occur west of Bali and Sulawesi. Sunbirds (Nectariniidae) have been observed as visitors of D. hasseltii (Blume) Lindl. in Java, D. secundum (Blume) Lindl. in Singapore, and D. lampongense J.J.Sm. in Borneo (Wood et al. 2011, with photograph), all in D. sect. Pedilonum, but no full pollination sequence has been reported. Pollination reports are scarce for the Asian species. Kjellson et al. (1987) detailed Bombus eximius (Apidae) pollination of D. infundibulum Lindl. (D. sect. Formosae), interpreted as a case of floral mimicry with Rhododendron lyi H.Lév. in a mountainous region of Thailand. This may represent a reasonably common pollination syndrome for similar orchids, as many species of D. sect. Dendrobium appear to lack nectar. Brodmann et al. (2009) reported a study on the apparently rewardless orchid D. sinense (D. sect. Formosae, endemic to Hainan, China), with the hornet Vespa bicolor (Hymenoptera: Vespoidea) recorded as a confirmed pollinator. These social wasps are known to collect honeybees for feeding their brood, and the authors identified compounds mimicking the scent of the bee prey. This indicates that the hornet is attracted in a syndrome of deceptive pollination by chemical mimicry. In the fragrant but nectarless D. unicum Seidenf. (D. sect. Dendrobium, from Thailand and Laos), Davies and Turner (2004) investigated labellar structures identified previously by Kjellsson and Rasmussen (1985) and described labellar hairs acting as pseudopollen and containing starch, unlike the protein described in pseudopollen from other orchid species. They suggested that the unknown pollinators may be using them as a food source, a possible new pollination syndrome. Dendrobium crumenatum Sw. (D. sect. Aporum), with ephemeral flowers, is pollinated by the Asiatic honeybee Apis cerana indica, as reported by Ong (2010), confirming previous observations. Moth- or butterfly-pollination has not been observed in Dendrobium, but some species have pale-coloured flowers with a long and narrow, spur-like mentum, such as D. furcatum Reinw. ex Lindl. and D. citrinum Ridl. Hoverflies are likely natural pollinators of Dendrobium species and frequently pollinate Australian Dendrobium species in cultivation (S.D.Lawson, personal communication), but there is no confirmed pollination in nature. It has been suggested that D. macrostachyum, D. rarum, D. mirbelianum, and D. affine, and some species with flowers that never open fully, are self-pollinating. These reports require confirmation. A few species, or varieties of species, are self-pollinating, for example D. womersleyi T.M.Reeve var. autophilum Dauncey. An especially unusual case is D. pandaneti Ridl., which has a pair of extra, lateral anthers (actually half-anthers) on the column. According to Carr (1928), this species pollinates itself, but only from the two pollinia produced in each of the lateral anthers; the 95

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normal anther, with four pollinia, is not involved. It is not known if this species is ever pollinated by insects. There is usually high fertility between most Dendrobium species, including members of different sections, as indicated in the vast array of registered artificial hybrids. There are hybrids between species from the northern and the southern clades (for example between D. sects. Phalaenanthe and Formosae), although these tend to be more difficult to produce (Kamemoto et al. 1999). Most species tested are fertile in all types of pollination. Some individual plants are self-infertile. Within the Australasian D. sects. Dendrocoryne, Rhizobium, and Spatulata, natural hybrids are relatively common and in many cases have been shown to result from cross-pollination by Trigona bees (Adams 2011). In 1905, Schlechter recorded the first natural hybrid in D. sect. Spatulata, D. ×schumannianum Schltr. (D. antennatum Lindl. × D. lineale Rolfe). Within D. sect. Calyptrochilus, D. ×yengiliense T.M.Reeve is a natural hybrid between D. alaticaulinum P.Royen and D. leucocyanum T.M.Reeve from New Guinea. Natural hybrids between sections, such as D. sects. Phalaenanthe and Spatulata, occur occasionally, and many artificial intersectional hybrids have been made, for example, Dendrocoryne and Brevisaccata, Dendrocoryne and Rhizobium, Dendrocoryne and Phalaenanthe, Crinifera and Dendrocoryne, Eleutheroglossum and Phalaenanthe, Formosae and Dendrobium, Formosae and Distichophyllae, Formosae and Phalaenanthe, Latouria and Formosae, Phalaenanthe and Dendrobium, Spatulata and Dendrocoryne, Spatulata and Latouria, Spatulata and Rhizobium, and others. (PA, SL, AS)

Uses Dendrobium species have been used by indigenous people for decorative, practical, medicinal, and spiritual purposes throughout Asia and Australia (Lawler 1984; Lavarack et al. 2000). The long, fibrous stems of some species, especially in D. sect. Diplocaulobium, turn yellow with drying, often increased by heating from the sun or fires. They can then be cut into strips for weaving and decoration. In Papua New Guinea and the Moluccas, the dried yellow stems of Diplocaulobium are used to weave belts, girdles and armbands, and specimens are often grown in villages. In Bougainville and the Solomon Islands, the yellow stems of D. solomonense (Carr) Schuit. & P.B.Adams (D. sect. Diplocaulobium) are plaited into amulets. Dendrobium polysema Schltr. (D. sect. Latouria) is also planted in New Guinea villages, where the dried stems are used in making armbands. In northern Australia, D. johannis Rchb.f. (D. sect. Spatulata) pseudobulbs were used to weave decorative dresses and bands. Dendrobium gouldii Rchb.f. (D. sect. Spatulata) is used similarly in the Solomon Islands. Flowers, leaves and stems of various species have been applied in ceremonial purposes and for body decorations. In Papua New Guinea, plants have been cultivated for these purposes. Dendrobium leaves and flowers are picked for their fragrance throughout Asia. Ancient Chinese medical herbals describe extensive use of Dendrobium species. The medicine chin-shi-hu consists of dried stems of around 20 Dendrobium species and is recommended for longevity and as an aphrodisiac. Dendrobium nobile Lindl., a major component, contains several alkaloids, the pharmacological properties of which are being studied. Large-scale collecting 96

of these Dendrobium species from the wild in countries such as Laos for export to China is a serious threat to these species. Populations of the more sought-after species, such as D. devonianum Paxton, have declined greatly as a result in many areas (Schuiteman, personal communication). Dendrobium species are also used extensively as medicines in India and Sri Lanka. An extract from D. amoenum Wall. ex Lindl. reportedly shows some antioxidative and antibacterial properties. Plants of D. sect. Biloba are used to treat intestinal haemorrhage in Papua New Guinea. Species of D. sect. Spatulata have been used in Australia and the Pacific as antibiotics, oral contraceptives, and abortifacients, and for anorexia and alopecia. Other Dendrobium species have application as ointments, antiparasitics, painkillers, and aphrodisiacs. In Australia, pseudobulbs of D. sect. Dendrocoryne species have been dried for decoration, as a source of yellow ochre, and also for food. Pseudobulbs of D. canaliculatum R.Br. were baked and eaten in northern Australia, and cooked stems of D. speciosum Sm. are also edible. The juice of D. affine (Decne.) Steud. stems is a fixative for rock and bark paintings in northern Australia and Groote Eylandt. Dendrobium bigibbum var. superbum Hort. ex Rchb.f. is the state emblem of Queensland, Australia, and D. engae T.M.Reeve adorns the official flag of the Enga Province in Papua New Guinea. Dendrobium plants are among the most popular orchids for growing and hybridizing. The sections most used in producing pot plants and cut flowers are Phalaenanthe, Spatulata, Dendrobium, Formosae, and Dendrocoryne, followed by Pedilonum, Calyptrochilus, Latouria, Rhizobium, and Sarcopodium. Sections Amblyanthus, Aporum, Biloba, Brevisaccata, Cadetia, Conostalix, Crinifera, Diplocaulobium, Distichophyllae, Eleutheroglossum, Fugacia, Fytchianthe, Grastidium, Herpethophytum, Lichenastrum, Macrocladium, Microphytanthe, Monophyllaea, Pleianthe, and Stachyobium are at present hardly grown commercially and are mainly found in species collections. The most significant use of Dendrobium is the ongoing international cut-flower trade and pot-plant industry, based primarily on the Australian species D. bigibbum (often referred to in the industry as D. phalaenopsis Fitzg.), D. nobile, and related species. Hybridization commenced during the Second World War and continued for four decades at the University of Hawaii under the direction of Haruyuki Kamemoto, who pioneered research on chromosome numbers and polyploidy. This was applied to develop superior seed-propagated, amphidiploid cultivars. Species of D. sect. Phalaenanthe, predominantly forms of D. bigibbum, were repeatedly crossed with species of D. sects. Spatulata, Latouria, Dendrobium, and Formosae. Since 1951, Hawaiians have registered about 40 new hybrids annually, with peaks of 80–110 per year (Kamemoto et al. 1999). The export industry in countries such as Thailand has developed from this base, particularly by expansion in Singapore in the early 1960s. Dendrobium moniliforme (L.) Sw. cultivars have been grown for centuries in Japan, and the ‘soft cane’ D. nobile has been in cultivation in temperate countries for over a century, with many generations of selections and hybrids. The first artificial Dendrobium hybrid, listed in 1864, was probably D. Dominianum (D. linawianum Rchb.f. × D. nobile) made by John Dominy of Messrs. Veitch and Sons. Species and hybrids of D. sects. Dendrobium, Formosae, and Pedilonum were introduced to the stove houses of

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the United Kingdom and Europe in the 19th century and are now widely grown in collections. Species in D. sects. Phalaenanthe and Spatulata and the natural hybrid between two of them, D. ×superbiens, all imported from northern Australia, were grown for cut-flowers in the United Kingdom and Europe. They were used in hybridization by Vacherot & Lecoufle in France and by Stuart Low in England. In temperate regions, especially Australia and the west coast of the USA, the primarily Australian D. sect. Dendrocoryne is widely grown either as true species, line-bred species or hybrids that exceed six generations. The hybrids are based mainly on D. kingianum Bidwill ex Lindl., D. speciosum, and D. tetragonum A.Cunn., providing mass-flowering pot-plants in a wide range of colours for much of the year. Intersectional hybrids between Dendrocoryne and Phalaenanthe (known as hot/cold hybrids) have extended the flowering period to include autumn and winter. Line-breeding of the most commonly grown D. sect. Dendrocoryne species, D. kingianum, has resulted in significant increases in overall flower size and segments, as well as creating more intense colour and unusual colour forms such as deep purple, cream, and yellow on compact plants (Adams and Lawson 1995). There is also extensive breeding within and between varieties of D. speciosum, with hybridizers focusing on high flower count and reduced plant size. In recent years, there has been increasing interest in hybridizing and line-breeding within D. sect. Rhizobium, particularly crossing Australian and New Guinea species to produce heavily flowering, pendulous plants (Adams 2008). The flowers are often larger and more heavily textured than those of most species, with interesting colour combinations and spotting. A range of cool-growing, vigorous plants produce flowers throughout the year with peaks in winter and spring. (PA, SL)

Cultivation Dendrobium species occur in diverse habitats under a wide range of climatic conditions; therefore it is difficult to make general comments on their cultivation. In nature, most species grow in fairly exposed positions, usually high up in trees or on rocks. Almost all species demand perfect drainage and cannot tolerate soggy conditions at the roots, but at the same time they require ample watering during the growing season. They are best grown in light shade in slab or pot culture using an open medium, often based on bark, which provides support and excellent drainage. Stone, scoria, perlite, polystyrene, coconut fibre, and other components are often added to the mix, but single-component and simple mixes are usually satisfactory. Some epiphytes, especially large, pendulous species, require mounts or hanging baskets. Fertilizing should generally be in the form of weak solutions and used sparingly, except for robust plants growing rapidly in the summer that can tolerate fertilizer without root damage. In most regions, housing is required to control climatic conditions and protect against excessive sun, rain, cold, hail, snow, and insect damage, and to provide heating when ambient temperature falls below minimum requirements. The main problems with enclosed housing are maintaining humidity, adequate air movement, and ventilation. These problems may be overcome

with fans and evaporative coolers, misting systems, venting, shade-cloth sides and ends, large doors, and roof components that open to the exterior. Plants require high humidity, which can be difficult to provide constantly in outdoor conditions. Particular species may grow on trees and mounts and in pots outdoors in favourable climates but are at greater risk from damage, especially to flowers. Detailed cultural information for Dendrobium can be found in Baker and Baker (1996), Upton (1989), and Adams and Lawson (1995). The sections can be divided into five broad cultural groups as follows, but there are many exceptions. 1) Dendrobium sects. Dendrobium, Formosae, Fytchianthe, continental Asian Sarcopodium, and Stachyobium. The first two contain some of the most commonly grown and showy species, such as D. nobile Lindl. and its relatives, and the species allied to D. formosum Roxb. ex Lindl. They can grow to form massive specimens with hundreds of flowers. Commercial growers refer to them collectively as the ‘soft cane dendrobiums’, in contrast to the ‘hard cane dendrobiums’ in D. sects. Phalaenanthe and Spatulata. They are mainly continental Asian in origin, and as such are usually subjected to a distinct alternation between a cooler dry season lasting a few to several months and a warmer and wet rainy season. Many species in D. sects. Dendrobium, Fytchianthe, and Stachyobium drop their leaves during the dry season. Most require intermediate conditions, with much light, lower temperatures (down to about 10 °C at night), and reduced watering during the winter months. They grow rapidly in summer, when they require frequent watering and benefit from fertilizing. Most species in D. sects. Formosae and Sarcopodium should not be kept dry for long even during the resting period. The often small plants in D. sect. Stachyobium are best grown on slabs so that the roots do not remain wet. 2) Dendrobium sects. Amblyanthus, Aporum, Biloba, Brevisaccata, Cadetia, Calyptrochilus, Conostalix, Crinifera, Diplocaulobium, Distichophyllae, Fugacia, Grastidium, Herpethophytum, Latouria, Macrocladium, Microphytanthe, Pedilonum, Pleianthe, and Malesian Sarcopodium. These are cool- to warm-growing, depending on the species. Many are from New Guinea or the mountains of Java, Sumatra, Borneo, and the Philippines, where they do not experience a distinct dry season. They generally require wet conditions throughout the year but always with excellent drainage. Most of these sections are of limited horticultural interest, except Calyptrochilus, Latouria, and Pedilonum. The majority are easily cultivated under (warm-) intermediate conditions, such as D. (Calyptrochilus) lawesii F.Muell., D. (Latouria) forbesii Ridl., and D. (Pedilonum) secundum (Blume) Lindl. Many species of D. sects. Aporum and Diplocaulobium are especially easy; the latter often form large clumps rapidly. More difficult in cultivation are the small, colourful highland species of D. sect. Calyptrochilus, such as D. cuthbertsonii F.Muell. and D. vexillarius J.J.Sm. These are intolerant of disturbance, over-­ fertilization, and pesticides and should never be allowed to dry out completely. They require cool and humid conditions without stagnant moisture. Little, if any, success has been achieved with some highly desirable subalpine species from New Guinea, such as D. brevicaule Rolfe, D. dekockii J.J.Sm., and D. vannouhuysii J.J.Sm. These need a combination of high light, high humidity, and low temperatures that is almost impossible to simulate in cultivation. 97

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Dendrobium cinnabarinum, a beautiful red-flowered species from the mountains of Borneo, grows in moss cushions under intermediate-cool conditions and is more intolerant of dryness than most of the less showy allied species in D. sect. Aporum. Species of D. sect. Macrocladium are grown in pots of welldrained sandy mixtures with plenty of organic material and may become large. They respond to filtered sunlight with yearround watering. The flowers last a few weeks. Some species may flower throughout the year, whereas others peak in summer and autumn. 3) Dendrobium sects. Phalaenanthe and Spatulata. These are mostly warm-growers, commonly seen commercially and in hobbyist collections. They are the most widely used Dendrobium species in hybridization. Most require a distinct seasonal climate, with hot, humid, and wet summers for the growth phase and a dry resting phase in winter. Heavy watering and high humidity are essential in summer. Small pots with a coarse, well-drained medium, mounts of hardwood or cork slabs or hanging baskets are used. Growth is assisted by frequent applications of weak fertilizers to promote rapid stem development. Species and hybrids of D. sect. Phalaenanthe, such as D. bigibbum, are warm-growers, preferring a range of 18–38 °C, with a dry rest period in winter and spring. New growths must mature and initiate racemes by late summer to flower in autumn and early to mid-winter. Bright light and high air movement are required, with lower humidity during flowering to prevent fungal infestation. In northern Australia, the Dendrobium beetle will attack all plant parts if kept outdoors. Species and hybrids of D. sect. Spatulata can be grown outdoors in tropical regions. They require glass-houses or heated houses in cooler climates. Watering should be infrequent in winter, just sufficient to prevent shrivelling of pseudobulbs. If not kept dry in winter, they are likely to drop all their leaves and develop root and stem rot. 4) Dendrobium sects. Dendrocoryne, Rhizobium, and Eleutheroglos­ sum. These are cool-growing, with a growing season over spring– summer and periods of relative dryness in winter when temperatures can be close to freezing. A few species prefer intermediate conditions. Most are easy and tolerate a range of temperatures and humidity, preferring frequent watering in summer and some fertilizing with weak solutions, pelleted fowl manure, or, best of all, annual repotting in fresh medium. Dendrobium section Dendrocoryne is the most commonly grown and includes such popular species as the fragrant D. kingianum. Most species are easily cultivated in temperate conditions and can develop into large specimens. For example, D. speciosum forms huge specimen plants with mass displays of over a thousand flowers (Adams 2008). Flowers can last up to two weeks under cool conditions. They are grown predominantly in pots with well-drained medium based on bark, stone, and coconut fibre. Plants can be mounted but will then require more frequent watering. Flowering is usually in spring to summer. Dendrobium tetragonum A.Cunn. may flower twice a year, in autumn and spring. Most species of D. sect. Rhizobium, such as D. teretifolium R.Br., grow well in pots and baskets or on mounts. They require vigorous air movement, bright light, well-drained media, high humidity, and frequent watering in summer. They can be established 98

successfully on tree branches in suitable climates. Roots are best left undisturbed by remounting on bigger slabs or sinking smaller pots into larger ones as plants advance in size. The terete-leaved and pendulous species can grow into specimens several feet long, bearing hundreds of flowers. Flowering is usually between winter and early summer, but a few species flower in later summer and autumn. Species of D. sect. Eleutheroglossum are predominantly cultivated on slabs or branches with 60–70% shade and flower in spring and summer. The Australian D. fellowsii F.Muell. is difficult to grow. 5) Dendrobium sects. Lichenastrum and Monophyllaea. These are cultivated in specialized collections and are of little horticultural value. They require cool to occasionally intermediate conditions, with year-round watering and high humidity, with more frequent watering in summer. Species of D. sect. Lichenastrum are small plants with a creeping habit suited to slab culture on tree fern, cork or hardwood in fairly low light. They can be grown on top of an inverted small pot or on plastic mesh (e.g. gutter guard) placed in a moss bed. These small, mat-forming orchids usually grow slowly and flower throughout the year. Species of D. sect. Monophyllaea, such as D. monophyllum F.Muell., are grown only occasionally. They have a creeping habit, requiring a mount on weathered wood or culture in shallow pots and may be difficult to grow in conditions other than intermediate. They are larger plants than are found in D. sect. Lichenastrum, tolerating more light and less humidity as their tough pseudobulbs retain moisture. (PA, SL, AS)

Taxonomic notes A sound infrageneric classification is invaluable in representing the diversity of such a huge genus as Dendrobium. Proposals to recognize sections, subsections, and subgenera in this genus have been numerous from the early nineteenth century onwards, starting with Blume and Lindley, as summarized by Wood (2006). Many early infrageneric groups were mixtures of species with highly diverse morphology that were probably only distantly related. As late as 1910, in Kränzlin’s monograph of Dendrobium, there are a number of sections comprised of species that might as well have been chosen randomly. This treatment was severely criticized by Schlechter (1912), who at the same time published what was to prove the definitive infrageneric classification of Dendrobium for a long time. Although Schlechter’s subgenera were as artificial as those of Kränzlin, his classification at the sectional level displayed for the first time a firm grasp of the natural affinities of the species across the whole genus. Indeed, most of the sections recognized here can already be found in Schlechter’s work, although sometimes under different names, and often with a different delimitation. The three main clades of Dendrobium (D. sect. Sarcopodium, northern clade, and southern clade) could possibly be classified as subgenera, but we have not used this rank, mainly because there are no consistent morphological differences between the northern and the southern clades, and bootstrap support is variable depending on the sampling used in different analyses. As

DENDROBIUM

DNA sequences of more species are analyzed, our insight into the phylogeny within sections may improve, which may lead to proposals to split up certain of the sections recognized here or establish subsections within them. We have refrained here from using the rank of subsection because our knowledge of many clades is still insufficient. It is clear, however, that in sections such as Cadetia, Crinifera, Dendrobium, Diplocaulobium, Grastidium, Macrocladium, Pedilonum, and others, distinct subclades can and probably should be formally recognized. Excessive splitting, on the other hand, defeats the purpose of having a formal classification. Dendrobium sect. Macrocladium, for example, has been proposed as comprising seven distinct sections by Wood (2006): Dendrobates, Finetianthe, Inobulbum, Kinetochilus, Macrocladium, Tetrodon, and Winika. Most of these contain only one or two species. This level of splitting, and that of Clements (2003, 2006) and others at generic level, appears impractical, as there are no satisfactory morphological correlates, too many name changes, and adverse consequences for horticultural and other users (Adams 2011; Schuiteman and Adams 2010; Schuiteman 2011). In the Australian clade, 19 sections are here recognized, which are generally well distinguished by morphological characters. There is a lack of bootstrap support and some evidence for lack of monophyly, for example in D. sects. Dendrocoryne and Grastidium, but also no strong support for polyphyly. In the Asian clade, ten sections are here provisionally recognized, some seemingly polyphyletic, and others likely to be divided into subsections in future classifications. Where support and resolution are low, morphology has been used to make decisions on sectional delimitation, even when in a few cases this has led to groupings that may eventually turn out to be para- or even polyphyletic. A problematic group is formed by D. sects. Conostalix, Distichophyllae, and Formosae. Sathapattayanon (2008) has shown that certain species previously included in D. sect. Formosae, such as the Philippine D. sanderae Rolfe and its relatives, are probably not closely related to species with superficially similar flowers, such as D. formosum Roxb. ex Lindl. Her cladograms also show that the large-flowered members of D. sect. Distichophyllae may be more closely related to D. sect. Conostalix than to other species in D. sect. Distichophyllae. Perhaps more sections should be recognized in D. sect. Formosae, or, conversely, it may be preferable to distinguish only one broadly circumscribed section comprising Distichophyllae and Conostalix. The infrageneric key toward the beginning of this treatment of Dendrobium is unsatisfactory for some members of these three sections. Dendrobium sects. Pedilonum (which here includes the former sections Calcarifera p.p., Pedilonum p.p., and Platycaulon) and Calyptrochilus (which here includes the former sections Calcarifera p.p., Calyptrochilus s.s., Cuthbertsonia, Dolichocentron, Oxyglossum, and Pedilonum p.p.) seem well resolved by DNA sequences, but morphology provides few clues that help assign the species to one or the other section. Both sections contain members of the D. sects. Calcarifera and Pedilonum as recognized by most previous authors (see species marked with P and C in Fig. 622.16). Within this alliance, species with rough or even warty roots are only found in D. sect. Pedilonum (but are also present in certain members of the southern clade and D. sect.

Sarcopodium), whereas species with papillose or winged ovaries, an inflexed labellar apex or non-fleshy stems are found only in D. sect. Calyptrochilus. This leaves unclassified a considerable number of ‘uncharacteristic’ species with smooth roots, glabrous and terete ovaries, and swollen stems. These are difficult to place in either D. sect. Pedilonum or D. sect. Calyptrochilus on the basis of morphology alone. Molecular data indicate that all ‘uncharacteristic’ species west of Sulawesi (which include a few that in addition occur farther east) belong in D. sect. Pedilonum (for example, D. hasseltii), whereas all ‘uncharacteristic’ species endemic to the Philippines, Sulawesi, and farther east, which form the great majority, belong in D. sect. Calyptrochilus (for example, D. bracteosum Rchb.f.). The only exception known so far is D. amethystoglossum Rchb.f., a Philippine endemic in D. sect. Pedilonum. Dendrobium sect. Dendrobium here includes D. sects. Densiflora (syn. D. sect. Callista), Stuposa, and Breviflores (species indicated in Fig. 622.16 with D, S, and B, respectively). The oligotypic D. sect. Herbacea is here tentatively included in D. sect. Dendrobium on morphological grounds, but could also be considered unplaced. Monophyly has not been established for D. sect. Dendrobium. However, overall morphological similarity and weak bootstrap support would indicate that the polyphyly shown in almost all published cladograms for the species concerned may not reflect their true evolutionary history. (AS, PA)

Taxonomic literature Adams, P. B. (2011). Systematics of Dendrobiinae (Orchidaceae), with special reference to Australian taxa. Botanical Journal of the Linnean Society, 166, 105–26. Adams, P. B., Burke, J. M., and Lawson, S. D. (2006). Systematic analysis of Dendrobium Swartz section Dendrocoryne in the Australian region. Plant Systematics and Evolution, 260, 65–80. Brieger, F. G. (1981). Subtribus Dendrobiinae. In R. Schlechter’s Die Orchideen, 3rd edition (ed. F. G. Brieger, R. Maatsch, and K. Senghas), 1 (11–12), pp. 636–752. Verlag Paul Parey, Berlin. Burke, J. M., Bayly, M. J., Adams, P. B., and Ladiges, P. Y. (2008). Molecular phylogenetic analysis of Dendrobium (Orchidaceae), with emphasis on the Australian section Dendrocoryne, and implications for generic classification. Australian Systematic Botany, 21, 1–14. Clements, M. A. (2003). Molecular phylogenetic systematics in the Dendrobiinae (Orchidaceae), with emphasis on Dendrobium section Pedilonum. Telopea, 10, 247–98. Clements, M. A. (2006). Molecular phylogenetic systematics in Dendrobieae (Orchidaceae). Aliso, 22, 465–80. Clements, M. A. and Jones, D. L. (2002). Nomenclatural changes in the Dendrobieae (Orchidaceae). 1: The Australasian region. Orchadian, 13, 485–97. Cribb, P. J. (1983). A revision of Dendrobium section Latouria (Orchidaceae). Kew Bulletin, 38, 229–306. Cribb, P. J. (1986). A revision of Dendrobium section Spatulata (Orchidaceae). Kew Bulletin, 41, 615–92. Dauncey, E. A. (2003). A taxonomic revision of Dendrobium section Pedilonum. Harvard Papers in Botany, 7, 151–320. Freudenstein, J. V. and Rasmussen, F. N. (1999). What does morphology tell us about orchid relationships? – A cladistic analysis. American Journal of Botany, 86, 225–48.

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Kränzlin, F. (1910). Orchidaceae–Monandrae–Dendrobiinae I. In: A. Engler, Das Pflanzenreich, Heft 45. Lavarack, P. S., Harris, W., and Stocker, G. (2000). Dendrobium and its relatives. Timber Press, Portland, Oregon. Morris, M. W., Stern, W. L., and Judd, W. S. (1996). Vegetative anatomy and systematics of subtribe Dendrobiinae (Orchidaceae). Botanical Journal of the Linnean Society, 120, 89–144. Reeve, T. M. (1983). A revision of Dendrobium section Microphytanthe. Orchadian, 7, 203–6. Reeve, T. M. and Woods, P. J. B. (1989, publ. 1990). A revision of Dendrobium section Oxyglossum (Orchidaceae). Notes from the Royal Botanic Garden Edinburgh, 46, 161–305. Sathapattayanon, A. (2008). Taxonomic revision of orchids in the genus Dendrobium Sw. section Formosae (Benth. & Hook.f.) Hook.f. in Thailand and adjacent areas. Thesis, Chulalongkorn University, Bangkok. Schlechter, R. (1911–1914). Die Orchidaceen von Deutsch-NeuGuinea. Repertorium Specierum Novarum Regni Vegetabilis. Beihefte, 1, 1–1079. Schuiteman, A. (2011). Dendrobium (Orchidaceae): To split or not to split? Gardens’ Bulletin. Singapore, 63, 245–57. Schuiteman, A. (2012). Typification of infrageneric taxa in Dendrobium. Muelleria, 30, 3–7. Schuiteman, A. (2013). A guide to Dendrobium of New Guinea. Natural History Publications (Borneo), Kota Kinabalu. Schuiteman, A. and Adams, P. B. (2010). A broad look at Dendrobium. Orchadian, 16, 447–59. Schuiteman, A. and Adams, P. B. (2011). New combinations in Dendrobium (Orchidaceae). Muelleria, 29, 62–8. Schuiteman, A. and de Vogel, E. F. (2002). Flora Malesiana: Orchids of New Guinea Vol. II; Dendrobium and allied genera. CD-ROM. ETI, Amsterdam.

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Seidenfaden, G. (1985). Orchid genera in Thailand. XII. Dendrobium Sw. Opera Botanica, 83, 1–295. Wongsawad, P., Handa, T., and Yukawa, T. (2005). Molecular phylogeny of Dendrobium Callista-Dendrobium complex. In Proceedings of the 17th World Orchid Conference (ed. H. Nair and J. Arditti), pp. 131–3. Natural History Publications (Borneo), Kota Kinabalu, in association with Shah Alam City Council, Shah Alam. Wood, H. P. (2006). The dendrobiums. A.R.G. Gantner Verlag, Ruggell. Yukawa, T. (2001). Molecular phylogeny of Dendrobium. In Proceedings of the Seventh Asia Pacific Orchid Conference (ed. H. Nagata and S. Ichihashi), pp. 69–71. APOC7, Nagoya, Japan. Yukawa, T., Kita, K., and Handa, T. (2000). DNA phylogeny and morphological diversification of Australian Dendrobium (Orchidaceae). In Monocots: systematics and evolution (ed. K. L. Wilson and D. A. Morrison), pp. 465–71. CSIRO Publishing, Melbourne. Yukawa, T., Kurita, S., Nishida, M., and Hasebe, M. (1993). Phylogenetic implications of chloroplast DNA restriction site variation in subtribe Dendrobiinae (Orchidacaeae). Lindleyana, 8, 211–21. Yukawa, T., Ohba, H., Cameron, K. M., and Chase, M. W. (1996). Chloroplast DNA phylogeny of subtribe Dendrobiinae (Orchidaceae): insights from a combined analysis based on rbcL sequences and restriction site variation. Journal of Plant Research, 109, 169–76. Yukawa, T. and Uehara, K. (1996). Vegetative diversification and radiation in subtribe Dendrobiinae (Orchidaceae): evidence from chloroplast DNA phylogeny and anatomical characters. Plant Systematics and Evolution, 201, 1–14.

B. TRI BE VANDE AE

Vandeae Lindl., Coll. Bot., App. (1821). Type: Vanda R.Br. Polystachyeae Pfitzer, Entw. Nat. Anordn. Orch., 100 (1887). Type: Polystachya Hook. Sarcantheae Pfitzer, Entw. Nat. Anordn. Orch., 100 (1887), as ‘Sarcanthinae’. Type: Sarcanthus Lindl. Polyrrhizeae Small, Man. Southeast. Fl., 274 (1933). Type: Polyrrhiza Pfitzer Adrorhizeae (Schltr.) Szlach., Fragm. Florist. Geobot. Suppl., 3, 67 (1995). Type: Adrorhizon Hook.f.

Description Epiphytic, lithophytic, or rarely terrestrial, monopodial or sympodial herbs. Stems one- to many-leaved; in sympodial taxa: stems congested or on a rhizome, often swollen into hetero- or homoblastic pseudobulbs. Leaves usually distichous, rarely secund or quaquaversal, conduplicate, with sheathing base, glabrous. Inflorescence lateral or terminal, racemose or paniculate, sometimes one-flowered. Flowers resupinate or not, ephemeral to long-lasting, distichous, secund, or quaquaversal. Sepals entire, free or rarely connate. Petals entire, free or rarely adnate to sepals. Labellum free, rigidly attached or hinged to the column or column foot, entire or lobed, with or without keels and/or callosities, with or without a spur. Column usually without apical stelidia; column foot present or absent; anther helmet-shaped, two- or fourlocular, pollinia two (sometimes porate or cleft), four (sometimes unequal-sized) or eight, hard-waxy, with or without caudicles, with or without a stipe, viscidium present or rarely absent, sometimes of complex shape; stigma a ventral concavity; rostellum short or beak-like elongated, entire or bi- or tridentate. Ovary and pedicel terete or six-ribbed, glabrous or rarely pubescent, rarely winged. Capsule subspherical to oblong to narrowly cylindrical, unilocular, smooth or six-ribbed, rarely winged. (AS)

Distribution (Fig. B.1) Vandeae comprise five subtribes with 139 genera and about 2600 species distributed in tropical America, tropical and southern Africa, Madagascar and neighbouring islands, tropical and subtropical Asia, eastern Australia and Tasmania, and much of the tropical Pacific south to New Zealand and east to Tahiti. (AS)

Phylogenetics Morphologically, the tribe Vandeae is remarkably heterogeneous, comprising as it does taxa with both monopodial and sympodial

architecture, as well as those with lateral or terminal inflorescences, two, four, or eight pollinia with or without caudicles/ stipe/viscidium, with or without pseudobulbs. There are no obvious morphological or anatomical traits characterizing Vandeae as here understood. Aeridinae and Angraecinae share stegmata with spherical silica bodies (Møller and Rasmussen 1984). Dressler (1993) suggested that this could be a possible synapomorphy for Vandeae, Podochileae, and Dendrobieae and placed them in one clade. Gorniak et al. (2010) pointed out that spherical silica bodies found in Podochileae and Vandeae could have been inherited from a common ancestor in their phylogenetic tree. In analyses based on multiple DNA regions (ITS, trnLF, matK, and rbcL) by van den Berg et al. (2005), a moderately supported clade (bootstrap 85%) comprising Polystachyinae + Aeridinae + Angraecinae (‘core Vandeae’) is sister to tribe Cymbidieae (bootstrap less than 50), and these two are together sister to Agrostophyllinae, but this last node was also weakly supported. The authors denoted the assemblage of the three clades as the vandoid orchids. In a Bayesian analysis based on the Xdh gene region, Górniak et al. (2010) recovered similar relationships, except that some Agrostophyllinae are here situated far removed from the core Vandeae, being sister to Epidendreae + Calypsoeae, and Vandeae do not appear as sister to Cymbidieae, which are far removed as well, but to the genus Sirhookera (incorrectly labelled as Agrostophyllinae, in reality belonging to Adrorhizinae). The position obtained for Earina with Xdh is peculiar, and the authors noted that this copy of the gene could be a paralogue, and the position of Earina thus would be spurious. It certainly contrasts with all previous analyses. Xdh could not be amplified from Agrostophyllum, so it was absent from this analysis. Bromheadia, which had previously been suggested to be a member of Cymbidieae (Dressler 1993; although with doubts about its exact placement), is sister to core Vandeae, and to this pair Polystachya is sister. Although Bayesian methods are considered to be the most modern, it has been our experience that the posterior probabilities they produce are inflated and unreliable, particularly with data matrices of combined data. Bootstrap percentages are preferred and more conservative. Neither Sirhookera nor Bromheadia was included in the analysis of van den Berg et al. (2005). Since the van den Berg et al. (2005) analysis, material of several genera of problematic placement has been obtained, so we added them to the matrix of van den Berg et al. and show the results here (Fig. B.2). This analysis is the basis for the subtribal classification used here, which should be seen as tentative. 101

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Seychelles Mascarene Islands

Fig. B.1.  Distribution map of Vandeae.

The additional taxa include 1) Imerinaea and Hederorkis, which Dressler (1993) included in Polystachyinae, 2) Claderia, which was considered of uncertain affinities by Dressler (1993), and 3) Adrorhizon (Adrorhizinae). With this set of taxa, Imerinaea falls with Eulophia in Cymbidieae (bootstrap 87%). Claderia is weakly supported as a member of Cymbidieae. Adrorhizon and Sirhookera are a well supported pair, to which Bromheadia is strongly supported (bootstrap 89%) as sister. Hederokis is weakly supported (66%) as sister to this clade, rather than sister to Polystachya. This larger set of taxa is then moderately supported (81%) as sister to core Vandeae, and Agrostophyllinae (Earina plus Agrostophyllum) are sister to the larger clade, which we considered to be Vandeae s.l. As noted above, further work on this set of taxa is required to confirm relationships. (AS, MC, FR)

Taxonomic literature Dressler, R. L. (1993). Phylogeny and classification in the orchid family. Dioscorides Press, Portland, Oregon. Górniak, M., Paun, O., and Chase, M. W. (2010). Phylogenetic relationships within Orchidaceae based on a low-copy nuclear coding gene, Xdh: congruence with organellar and nuclear ribosomal DNA results. Molecular Phylogenetics and Evolution, 56, 784–95. Møller, J. D. and Rasmussen, H. (1984). Stegmata in Orchidales: character state distribution and polarity. Botanical Journal of the Linnean Society, 89, 53–76. Senghas, K. (1986–1990). Tribus: Vandeae. In Rudolf Schlechter Die Orchideen 3rd ed. (ed. F. G. Brieger, R. Maatsch, and K. Senghas), pp. 973–1422. Paul Parey, Berlin.

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Senghas, K. (1990–1991). Tribus: Polystachyeae. In Rudolf Schlechter Die Orchideen 3rd ed. (ed. F. G. Brieger, R. Maatsch, and K. Senghas), pp. 1422–57. Paul Parey, Berlin. Van den Berg, C., Goldman, D. H., Freudenstein, J. V., Pridgeon, A. M., Cameron, K. M., and Chase, M. W. (2005). An overview of the phylogenetic relationships within Epidendroideae inferred from multiple DNA regions and recircumscription of Epidendreae and Arethuseae (Orchidaceae). American Journal of Botany, 92, 613–24.

Artificial key to the subtribes of Vandeae (AS) 1. Plants monopodial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Plants sympodial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2. Species from the Americas, Africa, Madagascar and neighbouring islands, with two outliers in Sri Lanka (Angraecum zeylanicum Lindl. and Aerangis hologlottis (Schltr.) Schtr.); pollinia 2, with 1 or 2 stipes and 1 or 2 viscidia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Angraecinae Species from Asia, with two outliers in Africa and Madagascar (Acampe pachyglossa Rchb.f. and Taeniophyllum coxii (Summerh.) Summerh.); pollinia 2 or 4, with a single stipe, rarely with 2 stipes (Bogoria taeniorhiza (Schltr.) Schltr.), with one viscidium . . . . . . . . . . . . . . . . . . . . . . . . Aeridinae 3. Pollinia 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Pollinia 4 or 8 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4. Stems without pseudobulbs; flowers distichous, ephemeral, opening in succession at long intervals; column ­without a foot . . . . . . . . . . . . . . . . . . . . Adrorhizinae (Bromheadia) ● ●

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●

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VANDEAE

Fig. B.2.  Strict consensus tree with bootstrap percentages (>50%) produced by adding additional taxa to the matrix of van den Berg et al. (2005). Branches without a number received bootstrap percentages less than 50. Subtribes and tribes for most groups indicated on the right; Coeliinae and Bletiinae were assigned to Epidendreae in Genera Orchidacearum, volume 4, but this is not indicated here because they do not fall in this analysis with the other members of that tribe. Five subtribes are assigned to Vandeae in the broad sense.

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Stems with or without pseudobulbs; flowers quaquaversal, longer lived, not opening in succession at long intervals; column with a distinct foot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polystachyinae (Polystachya) 5. Stems 3- to many-leaved; pollinia 4 or 8 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Agrostophyllinae Stems 1- or 2-leaved; pollinia 4 . . . . . . . . . . . . . . . . . . . . . . 6 6. Rhizome elongate; pollinia unequal (2 large, 2 small), attached by a tegular stipe to a viscidium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polystachyinae (Hederorkis) Rhizome abbreviated; pollinia equal, without stipe or viscidium, sometimes with caudicles . . . . . . . . . Adrorhizinae ●

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●

S UB TR IBE A DRO R H I Z I N AE Adrorhizinae Schltr., Die Orchideen, 132 (1914). Type: Adrorhizon Hook.f. Bromheadiinae Dressler, Lindleyana, 5, 124 (1990). Type: Bromheadia Lindl., syn. nov.

Description Epiphytic or terrestrial, occasionally lithophytic, herbs. Roots glabrous. Rhizome creeping. Stem erect, simple, terete to laterally compressed, not swollen or with pseudobulbs. Leaves solitary to several, distichous, alternate, articulate, often stiffly herbaceous to hard-coriaceous, base tapering to petiole (Adrorhizon and Sirhookera); blade sometimes purple-spotted when young (Sirhookera), linear-oblong to oblanceolate, acute to subacute (Adrorhizon), oval, oblong or oblong-lanceolate (Sirhookera) or dorsiventally flattened, articulation perpendicular to stem, obovate to linear with a bilobed apex (Bromheadia sect. Bromheadia) or laterally flattened or terete, articulation parallel to stem, obliquely triangular to linear with an acute apex (Bromheadia sect. Aporodes); sheaths tubular, clasping the stem, imbricate. Inflorescence accrescent, arising from base of stem (Adrorhizon), terminal, lateral or both, racemose or cymose, sometimes with one or several raceme-like branches or paniculate (Sirhookera and Bromheadia divaricata Ames & C.Schweinf.); floral bracts imbricate, alternating in two rows, stiff, rarely membranous, many-veined, midrib carinate; in Bromheadia sect. Aporodes with a few membranous, sterile bracts. Flowers resupinate, opening in succession or a few open simultaneously, sometimes gregarious, short-lived to ephemeral. Sepals and petals free or connivent, base sometimes subsaccate (Sirhookera). Sepals oblong or elliptic to linear-lanceolate, thin-textured to fleshy, 1–9-veined. Lateral sepals sometimes asymmetric, carinate or not. Petals elliptic, narrowly spatulate, or oblong to linear-lanceolate or lanceolate, often slightly asymmetric, usually thinner-textured than sepals, not carinate, narrowed at base, three-veined. Labellum cymbiform or spatulate, emarginate and crenulate to trilobed, spur absent, sometimes saccate, disc sometimes with a transverse membrane between side lobes and midlobe (Sirhookera); hypochile (Bromheadia) thin-textured, mostly median swollen, glabrous or with scattered hairs; lateral 104

lobes variable in size, pronounced in front; callus, when present, ligulate, more pronounced at top, often densely hairy or papillose; epichile (Bromheadia) triangular to suborbicular or orbicular, or ligulate, flat to recurved, with upturned margins (a few species of B. sect. Bromheadia), with a pronounced, narrow to broad, ruminate, glabrous to papillose or hairy callus. Column curved, narrowly winged, with or without a semi-orbicular auricle at the base; foot absent; hood small or dilated, covering at least base of anther-cap, margin sometimes toothed; anther cap terminal, bilocular or imperfectly four-chambered, obovate to elliptic in outline from above, pollinia two or four, free, sometimes porate, flat, lanceolate-ovoid to triangular-ovoid, or pyriform in unequal pairs; stigma cupulate, top margin (Bromheadia) laterally on either side with a triangular to semi-orbicular flap bent to cover median part of top rim (lateral flaps absent in Bromheadia pendek de Vogel, B. sect. Bromheadia); viscidium detachable, semi-domeshaped, covering most of stigma. Capsule with six longitudinal ribs, perianth often persistent. (JW)

Distribution Adrorhizinae comprise three genera distributed from India and Sri Lanka eastward through Burma, Thailand and Indochina, Malaysia and Indonesia, north to the Philippines, south to New Guinea and northern Australia (Queensland). (JW)

Artificial key to the genera of Adrorhizinae ( JW) 1. Pollinia 2; leaves sometimes laterally flattened or terete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 624. Bromheadia Pollinia 4; leaves never laterally flattened or terete . . . . . . 2 2. Inflorescence a 1–3-flowered cyme; labellum entire, spatulate; disc without a transverse membrane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 623. Adrorhizon Inflorescence paniculate, many-flowered; labellum trilobed; disc with a transverse membrane between side lobes and midlobe . . . . . . . . . . . . . . . . . . . . . . . . 625. Sirhookera ●

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●

6 2 3 .  A DROR H I ZON Adrorhizon Hook.f. in Trimen, Handb. Fl. Ceylon, 4, 161 (1898). Type species: Adrorhizon purpurascens (Thwaites) Hook.f. (basionym: Dendrobium purpurascens Thwaites)

Derivation of name From the Greek (h)adros, stout, and rhiza, root, referring to the thickened roots arising from the rhizome (Gunawardena 1968). (SF)

Description (Plate 54; Fig. 623.1) Epiphytic herbs. Rhizome creeping, producing many long, stout, vermiform roots. Stem terete, unifoliate. Leaf linear-oblong to oblanceolate, acute to subacute, base tapering to petiole, midrib

ADRORHIZON

B

H

G

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K C

I E

L

D

F A

Fig. 623.1.  Adrorhizon purpurascens (Thwaites) Hook.f. A. Habit; B. Flower; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum; G. Column and labellum, side view; H. Column, from view; I. Column apex, anther cap removed; J. Anther cap, dorsal view; K. Anther cap with pollinia, ventral view; L. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from holotype (K) and Kew Spirit Collection no. 24813.

prominent, coriaceous, often purplish beneath. Inflorescence a 1–3-flowered cyme arising from base of stem, erect, glabrous, with two or three sterile, sheathing bracts. Flowers resupinate, 1.2–2.0 cm across, white, margins of column and anther cap purple. Sepals linear-oblong, obtuse to oblanceolate, acute, three-veined. Petals narrowly spatulate, subacute, one-veined. Labellum sessile on base of column, spatulate, emarginate, apex crenulate, base saccate, five-veined, disc smooth. Column erect, narrowly winged, club-shaped, top dilated with toothed margins, foot absent; anther terminal, imperfectly four-­ chambered, pollinia four, free, waxy, narrowly pyriform in

unequal pairs. Capsule narrowly oblong or pyriform, strongly six-ribbed. (SF)

Distribution (Fig. 623.2) Adrorhizon is a monospecific genus endemic to Sri Lanka. The main distribution lies in the central mountainous region of the island in Kandy, Nuwara Eliya, and Badulla districts. It also occurs at higher elevations of the Knuckles Mountains and Deniyaya-Rakwana range in the north and southwest of the ­central highlands, respectively. This species has not been 105

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japonica Thunb. (Pentaphylaceae) according to Jayaweera (1981). Flowering has been recorded from September to January. (SF)

Pollination Pollination by flies and ants has been observed in the Knuckles region in Sri Lanka (Suranjan Fernando, unpublished). (SF)

Uses There are no known uses for this species, and it is not in cultivation. (SF)

Taxonomic literature

Fig. 623.2.  Distribution map of Adrorhizon.

recorded in the northern and eastern dry and intermediate zones of Sri Lanka. (SF)

Anatomy Tominski (1905) investigated the leaves of this species. In transection, the cuticle is more strongly developed adaxially than abaxially. Stomata are confined to the abaxial surface. Mesophyll is differentiated into palisade and spongy tissues. Vascular bundles lie in one row. Xylem bundle-sheath fibres are one or two layers thick. Porembski and Barthott (1988) examined the roots of this species and designated the velamen as the Calanthe-type with three cell layers. Tilosomes are absent. (AP)

Palynology There are no reports on the pollen of this species. (AP)

Cytogenetics There are no published chromosome counts for this species. (AP)

Phytochemistry No reports have been found on the phytochemistry of Adrorhizon. (NV, RG)

Ecology Adrorhizon occurs in submontane and montane forests above 720 m but is most commonly found above 1000 m. It is found on tree trunks and large branches with moss cover, rarely on moss-covered rocks. In low-elevation locations, the species was observed in canopies of tall trees. The species is commonly found on Calophyllum trapezifolium Thwaites (Callophyllaceae), Bhesa ceylanica (Arn. ex Thwaites) Ding Hou (Centroplacaceae), Eugenia spp. (Myrtaceae), Prunus ceylanica Miq. (Rosaceae), Scolopia sp. (Salicaceae), Memecylon sp. (Melastomataceae), and Eurya 106

Gunawardena, D. C. (1968). Genera et species plantarum zeylaniae. Lake House Investments Ltd., Colombo. Jayaweera, D. M. A. (1981). Orchidaceae. In A revised handbook to the flora of Ceylon. Vol. 2. (ed. M. D. Dassanayake and F. R. Fosberg), pp. 4–320. Oxford and IBH Publishing Co., India.

6 2 4 .  B ROM H EA DI A Bromheadia Lindl. in Edwards’s Bot. Reg., 27: 89, misc. 184 (1841). Type species: Bromheadia finlaysoniana (Lindl.) Miq. (basionym: Grammatophyllum finlaysonianum Lindl.)

Derivation of name The name commemorates Sir Edward French Bromhead, F.R.S., a keen student of botany and friend of John Lindley ‘whose investigations of the natural affinities of plants are well known to systematical Botanists’. (JW)

Description (Plate 55; Fig. 624.1) Epiphytic or terrestrial, occasionally lithophytic, herbs with creeping rhizomes. Roots glabrous. Stem erect, terete to laterally compressed, not swollen. Leaves stiffly herbaceous to hard-coriaceous, blade dorsiventally flattened, articulation perpendicular to stem, obovate to linear with a bilobed apex (B. sect. Bromheadia), or blade laterally flattened or terete, articulation parallel to stem, obliquely triangular to linear with an acute apex (B. sect. Aporodes); sheaths tubular, clasping the stem, imbricate. Inflorescence racemose, sometimes with one or several raceme-like branches, or a panicle (B. divaricata); floral bracts imbricate, alternating in two rows, stiff, rarely membranous, many-veined, midrib carinate, with a few, membranous sterile bracts in B. sect. Aporodes. Flowers resupinate, opening in succession or a few open simultaneously. Sepals 7–9-veined. Lateral sepals slightly asymmetric. Petals elliptic or oblong to linear-lanceolate or lanceolate, slightly asymmetric. Labellum cymbiform, trilobed, hypochile thin-textured, epichile triangular to suborbicular or orbicular, or ligulate, flat to recurved, with upturned margins in a few species of B. sect. Bromheadia, callus ruminate, glabrous to papillose or hairy. Column curved, narrowly winged; anther cap bilocular, obovate to elliptic in outline from above, pollinia two, porate; stigma cupulate, top margin laterally on either side with a triangular to semi-orbicular flap bent to cover median part of top rim (lateral flaps absent in B. pendek, B. sect. Bromheadia); viscidium detachable, semi-dome-shaped, covering most of the stigma. (JW)

BROMHEADIA

E

G

J

F K H N

L

M

I

O B A

D

C

Fig. 624.1.  Bromheadia borneensis J.J.Sm. A, B. Habit; C. Example of a narrower leaf; D. Inflorescence; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum, flattened, front view; I. Pedicel with ovary, labellum, and column; J. Upper portion of pedicel with ovary and column, anther cap detached; K. Column with anther cap detached, front view; L. Anther cap, dorsal view; M. Anther cap, ventral view; N. Pollinarium, pollinia detached; O. Viscidium. A and B redrawn from Vermeulen 816 and C-O from Lamb SAN 93455 by Susanna Stuart-Smith. Single bar = 1 mm, double bar = 1 cm. Reproduced with permission from J. J. Wood (2003), Orchids of Borneo. Volume 4. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

Distribution (Fig. 624.2)

Infrageneric treatment (JW)

Bromheadia is a genus of 27 species (7 in B. sect. Bromheadia, 20 in B. sect. Aporodes) distributed from Sri Lanka eastward through Indochina, Malaysia, and Indonesia, north to the Philippines, south to New Guinea and northern Australia (Queensland). (JW)

Kruizinga et al. (1997) revised the genus and recognized two sections distinguished readily by their vegetative morphology as outlined below.

107

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Fig. 624.2.  Distribution map of Bromheadia.

Bromheadia Lindl. sect. Bromheadia. Type species: Bromheadia finlaysoniana (Lindl.) Miq. Bromheadia Lindl. sect. Aporodes Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 366 (1914). Type species: Bromheadia aporoides Rchb.f.

Artificial key to sections of Bromheadia ( JW)

●



●

Leaf blade dorsiventrally flattened, narrowed at base, apex bilobed; articulation with the sheath perpendicular to stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Bromheadia Leaf blade laterally flattened or terete, not narrowed at base, apex acute, articulation with the sheath parallel with stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. sect. Aporodes

finlaysoniana. Furthermore, whereas B. alticola has long root hairs (especially on the ventral surface attached to the substrate), B. finlaysoniana has only cuticularized papillae that do not protrude beyond other epidermal cells. Porembski and Barthlott (1988) confirmed the bilayered velamen of B. finlaysoniana and characterized it and the exodermis as of the Cymbidium-type. They also reported spirally thickened idioblasts in the cortex, as did Groom (1893) in B. palustris. Finally, Groom (1893) observed ‘a loosely coherent black-brown substance’ in B. alticola at the interface between exodermal passage cells and innermost layer of velamen, presumably tilosomes in modern parlance. (AP)

Palynology There are no published reports on pollen of Bromheadia. (AP)

Cytogenetics Anatomy Møller and Rasmussen (1984) reported conical silica bodies in the stem and leaf of B. aporoides Rchb.f. Groom (1893) studied roots of the epiphytic B. alticola Ridl. and the terrestrial B. finlaysoniana (as B. palustris Lindl.). He noted that there is a bilayered velamen in both species but that it breaks down in B. 108

Nothing is known about the chromosome numbers of these species. (AP)

Phytochemistry A preliminary survey of anthocyanin pigments present in the young leaves and flowers of Bromheadia finlaysoniana revealed

SIRHOOKERA

the presence of a cyanidin glucoside together with the common flavonol glycoside, rutin (quercetin 3-O-rutinoside; Lowry and Keong 1973). Genes encoding two enzymes involved in the biosynthesis of flavonoids and anthocyanins, chalcone synthase and dihydroflavonol 4-reductase, have been characterized in B. finlaysoniana by screening cDNA libraries constructed from the flowers (Liew et al. 1998a,b). A survey of alkaloid content in Orchidaceae gave positive results for B. alticola and B. finlaysoniana (Lüning 1967). (NV, RG)

Ecology Species are epiphytic or terrestrial in lowland mixed and hill forests, moss forests, and terrestrial in peat swamp forests and along roadsides. The widespread B. finlaysoniana is a terrestrial on loamy to sandy soils in disturbed and open habitats, where plants receive direct sunshine. It is also recorded as epiphytic low on tree trunks, high up on major branches, often in their forks, and on lianas. Species range in elevation from sea level to 3000 m. ( JW)

Pollination Ridley (1890b) discussed pollination of B. alticola, B. aporoides, and B. palustris by carpenter bees (Xylocopa spp.). Rolfe (1908) reported that the flowers of B. finlaysoniana are also pollinated by a Xylocopa species. ( JW)

Uses Bromheadia finlaysoniana is used by the Iban (vernacular name: wi buntak) and Kelabit (vernacular name: busak paya) in Sarawak to cure toothache; the latex is applied to the painful area (Christensen 2002). ( JW)

Cultivation Bromheadia finlaysoniana is sometimes cultivated. The epiphytic species can be grown successfully in partially shaded conditions in pots with a loose, well-drained medium hardwood bark or charcoal. The pot medium can be allowed to dry out but for short periods only. The terrestrial species, e.g. B. finlaysoniana, require a more retentive combination, such as 40% fibrous peat, 40% bark, and 20% perlite. Regular watering is necessary and a high relative humidity maintained. The epiphytic species of section Bromheadia and all species of section Aporodes can also be successfully grown on slabs. A minimum winter temperature of 12–15 °C should be provided. ( JW)

Taxonomic literature Kruizinga, J., van Scheindelen, H. J., and de Vogel, E. F. (1997). Revision of the genus Bromheadia (Orchidaceae). Orchid Monographs, 8, 79–118, figs. 29–55, pl. 4b–5b. Repetur, C. P., Welzen, P. C., and de Vogel, E. F. (1997). Phylogeny and historical biogeography of the genus Bromheadia (Orchidaceae). Systematic Botany, 22, 465–77. Rolfe, R. A. (1908). Bromheadia finlaysoniana. Orchid Review, 16, 200.

6 2 5 .   S I R H OOK ER A Sirhookera Kuntze Rev. Gen. Pl., 2, 681(1891). Type species: not designated. Josephia Wight, Icon. Plant. Ind. Orch., 5, 19, t. 1742 (1851). Type species: not designated.

Derivation of name The name commemorates British botanist Sir Joseph Dalton Hooker (1817–1911), Director of the Royal Botanic Gardens, Kew (1865–1885), traveller in India and other countries, collector of specimens of about 3000 species of plant, and co-author of Genera plantarum (Wight 1852; Kuntze 1891). (SF)

Description (Plate 56; Fig. 625.1) Epiphytic herbs. Roots vermiform. Pseudobulb tufted. Leaves one or two, surrounded by four green, purple-veined sheaths when young, base continuing into the petiolar sheath, later becoming papery and matted, blade oval, oblong, or oblong-lanceolate, coriaceous, lower surfaces of young leaves purple-spotted. Inflorescence persistent, paniculate, longer than leaves, continuously flowering. Flowers terminal or subterminal, bracts subulate, persistent, shorter than pedicels. Sepals and petals subequal, oblong, connivent, concave, base subsaccate. Labellum base recurved, concave, side lobes rounded, incurved, midlobe rounded, disc with a transverse membrane between side lobes and midlobe. Column erect, as long as the sepals, broadened above; anther terminal, two-loculed, pollinia four, waxy, pyriform, with caudicles. Capsule sessile. (SF)

Distribution (Fig. 625.2) Sirhookera is a genus with two species, S. lanceolata (Wight) Kuntze and S. latifolia (Wight) Kuntze. In Sri Lanka, the genus is distributed in the submontane and mountain area in the central region and southwestern lowlands at 190–2000 m. In India, it occurs in Western Ghats from Nilgiris (Abraham and Vatsala 1981) and Palni Hills (Seidenfaden 1999) in Tamil Nadu; Trivandrum, Idukki, Ponmudi, Wayand (Sathish Kumar and Manilal 2004) in Kerala; and Hassan (Saldanha 1976) and Kodagu (Rao 1998) districts in Karnataka. (SF)

Anatomy Porembski and Barthlott (1988) investigated roots of S. lanceolata and designated the velamen as the Calanthe-type with five cell layers. Tilosomes are absent. (AP)

Palynology There are no reports on the pollen of Sirhookera. (AP)

Cytogenetics There are no published chromosome counts for these species. (AP) 109

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N

B

L

O H M I

J

C E

K D A

G

F

Fig. 625.1.  Sirhookera lanceolata (Wight) Kuntze. A. Habit; B. Flower; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum, from above; G. Labellum, longitudinal section; H. Column and labellum, side view; I. Column, front view; J. Column apex, anther cap removed; K. Column, side view; L. Anther cap, dorsal view; M. Anther cap with pollinia, ventral view; N. Anther cap, ventral view; O. Pollinia. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from holotype (K) and Kew Spirit Collection no. 42679.

Phytochemistry No reports have been found on the phytochemistry of Sirhookera. (NV, RG)

Ecology Sirhookera species grow on trunks and large branches of trees as small colonies, usually in open places with about 60% sunlight. They also grow on moss-covered rocks. In submontane and montane forests they are abundant in frequently mist-covered areas with moss. In the lowlands they are commonly found 110

on trunks and branches of trees near water sources. In India, Saldanha (1976) reported them in semi-evergreen forests of Upper Ghats. Flowering occurs in May–December in Sri Lanka and July–September in Karnataka, India (Saldanha 1976). The curious character of the genus is simultaneous flowering each year on the same inflorescence. In Sri Lanka some of its host species are Calophyllum trapezifolium Thwaites (Calophyllaceae), Syzygium fergusonii Gamble (Myrtaceae), Palaquium grande (Thwaites) Engl. (Sapotaceae), Symplocos sp. (Symplocaceae), and Gordonia speciosa (Gardner) Choisy (Theaceae) according to Jayaweera (1981). (SF)

AERIDINAE

Seidenfaden, G. (1999). Orchidaceae. In The flora of the Palni Hills, south India, part 3 (ed. K. M. Matthew), pp. 1197–272. Rapinat Herbarium, Tiruchirapalli, India. Thwaites, G. H. K. (1864). Enumeratio plantarum Zeylaniae. Dulau & Co., London. Wight, R. (1852). Icones plantarum Indiae Orientalis. Vol. 5. Madras, India.

SU BTRIBE AERID IN AE

Fig. 625.2.  Distribution map of Sirhookera.

Pollination Pollination by ants has been observed in Adam’s Peak wilderness area in Sri Lanka (Suranjan Fernando, unpublished). (SF)

Uses There are no uses reported for this genus, and it is not in cultivation. (SF)

Taxonomic notes The genus was originally named Josephia by Robert Wight (1851), which was a later homonym of Josephia Salisb. (1809), a genus in Proteaceae. Kuntze (1891) renamed it Sirhookera. (SF)

Taxonomic literature Abraham, A. and Vatsala, P. (1981). An introduction to orchids. Tropical Botanic Garden and Research Institute. Trivandrum, India. Jayaweera, D. M. A. (1981). Orchidaceae. In A revised handbook to the flora of Ceylon. Vol. 2. (ed. M. D. Dassanayake and F.R. Fosberg), pp. 4–320. Oxford and IBH Publishing Co., India. Kuntze, O. (1891) Reviso generum plantarum. Vol. 2. A. Felix, Leipzig. Rao, T. A. (1998). Conservation of wild orchids of Kodagu in the Western Ghats. Karnataka Association for the Advancement of Science, Bangalore, India. Saldanha, C. J. (1976). Orchidaceae. In Flora of Hassan district, Karnataka, India (ed. C. J. Saldanha and D. H. Nicolson), pp. 809–855. New Delhi, India. Sathish Kumar, C. and Manilal, K. S. (2004). Orchids of Kerala, India. In Orchid memories: a tribute to Gunnar Seidenfaden (ed. K. S. Manilal and C. Sathish Kumar), pp. 155–254. Mentor Books, for Indian Association for Angiosperm Taxonomy, Calcutta, India.

Aeridinae Pfitzer, Entw. Nat. Anord. Orchid., 106 (1887), as ‘Aerideae’. Type: Aerides Lour. Vandinae Rchb.f., Ic. Fl. Germ. Helv., 13 & 14, IX (1851), as ‘Euvandeae’, nom. invalid. Sarcanthinae Benth., J. Linn. Soc., Bot., 18, 288 (1881), as ‘Sarcantheae’, nom. illeg. Type: Sarcanthus Lindl. (1826) Deceptorinae Szlach., Fragm. Florist. Geobot. Suppl., 3, 88 (1995). Type: Deceptor Seidenf., syn. nov. Diplocentrinae Szlach., Fragm. Florist. Geobot. Suppl., 3, 90 (1995). Type: Diplocentrum Lindl., syn. nov. Gastrochilinae Szlach., Fragm. Florist. Geobot. Suppl., 3, 89 (1995). Type: Gastrochilus D.Don, syn. nov. Pelatantheriinae Szlach., Fragm. Florist. Geobot. Suppl., 3, 90 (1995). Type: Pelatantheria Ridl., syn. nov. Phalaenopsidinae Szlach., Fragm. Florist. Geobot. Suppl., 3, 89 (1995). Type: Phalaenopsis Blume, syn. nov. Taeniophyllinae Szlach., Fragm. Florist. Geobot. Suppl., 3, 91 (1995). Type: Taeniophyllum Blume, syn. nov.

Description Epiphytic, lithophytic, or rarely terrestrial, monopodial herbs. Stems few- to many-leaved. Leaves usually distichous, rarely secund or quaquaversal, conduplicate, with sheathing base, glabrous. Inflorescence lateral, racemose or paniculate, sometimes 1-flowered. Flowers resupinate or not, ephemeral to long-lasting, distichous, secund, or quaquaversal. Sepals entire, free or rarely connate. Petals entire, free or rarely adnate to sepals. Labellum free, rigidly attached or hinged to the column or column foot, entire or lobed, with or without keels and/or callosities, with or without a spur. Column usually without apical stelidia; column foot present or not; anther helmet-shaped, two-locular, pollinia two or four, hard-waxy, when 2 then often partially slit or porate, when 4 often arranged in two bodies and those of a pair often unequal, with or without caudicles, attached to a tegular stipe, viscidium present, sometimes of complex shape; stigma a ventral concavity, rarely with raised margins; rostellum beak-like or not, bidentate. Ovary and pedicel terete or six-ribbed, glabrous or rarely pubescent. Capsule subspherical to oblong to narrowly cylindrical, unilocular, smooth or six-ribbed, rarely winged, with endocarpic elaters. (AS)

Distribution Subtribe Aeridinae comprises 83 genera with about 1550 species, distributed from India and Sri Lanka throughout tropical Asia and the Pacific region, north to Japan, east to Tahiti, south to New Zealand, with two species (in Acampe and Taeniophyllum) in tropical and southeastern Africa, and Acampe also found in Madagascar. (AS) 111

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Anatomy Members of Aeridinae are by far the most common vandaceous taxa included in orchid anatomical surveys. The anatomy of Luisia filiformis Hook.f. was examined during a survey of the ecological anatomy of several Sikkim Himalayan orchids (Mohana Rao et al. 1989). They found that the leaves possessed paracytic stomatal complexes, no trichomes, no hypodermis, an undifferentiated mesophyll, leaf idioblasts with ‘multispiral cellulosic thickenings’, and that the root had a ∪-thickened endodermis and lacked mycorrhizae. Kaushik (1983) published an extensive survey of the vegetative anatomy for Himalayan Orchidaceae. He broke the subtribe into the following groups, based on water-storage idioblasts in leaves, roots, and stems: A) Kingidium (= Phalaenopsis) + Camarotis (= Micropera), idioblasts absent; B) Vandopsis + Gastrochilus, idioblasts large with smooth cell walls; C) Cleisostoma + Aerides + Rhynchostylis + Vanda + Luisia, idioblasts club- or barrel-shaped with variously banded cell-wall thickenings. In a survey of developmental root anatomy for 10 Taiwanese orchid species, Chiang and Chou (1971) examined five vandaceous taxa: Haraella odorata Kudô (= Gastrochilus retrocallus (Hayata) Hayata), Phalaenopsis amabilis (L.) Blume, Sarcanthus fuscomaculatus (Ker Gawl.) Lindl. (= Cleisostoma paniculatum (Ker Gawl.) Garay), Saccolabium formosanum Hayata (= Gastrochilus formosanus (Hayata) Hayata), and Thrixspermum saruwatarii (Hayata) Schltr. Roots possessed 2–4 velamen layers with endophytic fungi and algae in all specimens examined. Singh (1986) examined the root anatomy of four species of Aeridinae: Aerides odorata Lour., Rhynchostylis retusa (L.) Blume, Vanda cristata Wall. ex Lindl., and V. roxburghii R.Br. (= V. tessellata (Roxb.) Hook. ex G.Don). Velamen layers ranged from two to eight; exodermal walls were either ∪- or ○-thickened; endodermal cells were either thin-walled or ○-thickened; and the number of vascular groups (xylem + phloem) ranged from 11–46. In their survey of leaf anatomy in nine epiphytic orchids, Oliveira and Sajo (1999) examined Vanda tricolor Lindl. and found lignified cells in the mesophyll as well as in the subepidermal layer but no water-storage tissue. In his monograph of aerial orchid roots, Leitgeb (1864) examined several genera of Vandeae, mainly Aeridinae. In this extensive treatment, Leitgeb was first to describe Deckzellen, or cover cells, in the velamen of many vandaceous genera. Cover cells form a tiered mantle or covering of 2–4 wedge-shaped cells that develop from the inner velamen above the passage (short) cells of the exodermis. Cover cells may be associated with tilosomes in other orchid tribes, and several other anatomists mentioned their occurrence within Vandeae (Solereder and Meyer 1930; Mulay and Panikkar 1956). Although Curtis (1917) did not observe cover cells (his ‘pyramidal cells’) in Sarcochilus adversus Hook.f. (= Drymoanthus adversus (Hook.f.) Dockrill), he did mention these structures in Dendrobium cunninghamii Lindl. (which lacks tilosomes). Areas of the velamen that remain dry after wetting were first observed by Leitgeb (1864) and subsequently termed ‘pneumathodes’ by Jost (1887). Haberlandt (1914) also described the 112

structure of pneumathodes in Taeniophyllum zollingeri Rchb.f. (= T. pusillum (Willd.) Seidenf. & Ormerod). According to Haberlandt (1914), the pneumathode was composed of three parts—a wedge-shaped area of air-filled velamen cells, one (usually) to several thin-walled exodermal cells, and several specialized cortical cells he called ‘complementary cells’ (a term typically used for lenticel tissue in the bark of trees). Dycus and Knudson (1957), in their study of the role of the velamen in aerial roots, described pneumathode cells as more heavily lignified than surrounding velamen cells, but they did not detect any specialized cortical complexes below those specialized velamen areas. In a scanning electron microscope (SEM) survey of plant epidermal surfaces, Barthlott (1981) described the ‘spotted’ appearance of the root surface for Taeniophyllum sp. Although Barthlott did not describe them as such, these spots probably correspond to pneumathodes in the velamen, which are able to retain air when wetted with water and are thus different from surrounding areas. Benzing et al. (1983) later examined pneumathodes in several leafless and leafy vandaceous taxa and compared them to nonvandaceous taxa. Their definition of pneumathode was more restrictive than Haberlandt’s (1914) and included only the airfilled cells of the velamen, whereas the adjacent cortical layers (exodermal cell + cells of the assimilatory cortical region) were defined as ‘aeration units’. These aeration units potentially act as cortical stomatal complexes, regulating gas exchange between the photosynthetic cortex and air-filled velamen cells, a function of pneumathodes that had been hypothesized by Schimper in 1888 (Pridgeon 1987). Pridgeon et al. (1983) examined tilosomes (rod-bodies or Stabkörper), speculated on their systematic occurrence in Orchidaceae, and created a classification scheme of tilosome types. Of the 31 vandaceous genera examined, only one unidentified species of Saccolabium possessed broadly lamellate tilosomes. In their extensive study of velamen types in Orchidaceae, Porembski and Barthlott (1988) did not find any tilosomes in Vandeae. The root multiple epidermis itself has been characterized in an extensive survey of velamen micromorphology assembled by Porembski and Barthlott (1988). They found a consistent and unique type of velamen, the ‘Vanda type’, for the 46 vandaceous genera surveyed. Velamina consisted of 2–5 cell layers with a distinct epivelamen, the cells of which are typically thinner-walled than those of the endovelamen. Cell wall thickenings were large, helical, and anastomosing; unthickened walls may show small pores. Sanford and Adanlawo (1973) previously examined the velamen and exodermis of several West African epiphytic taxa and related these characteristics to habitat tolerance. They found that velamen with more cell layers typically characterized taxa that grew in drier, harsher habitats than those with fewer cell layers. They also classified the velamen into three categories based on its wall striations: type I with broad, parallel, unbranched, and uncrossing striations; type II with broad, branched, and occasionally crossing striations; and type III with fine striations that usually crossed one another. Vandeae generally possessed two velamen layers with type I or type II thickenings.

AERIDINAE

Olatunji and Nengim (1980) examined the occurrence and distribution of ‘tracheoidal’ idioblasts among Orchidaceae. They speculated on the function of these cortical idioblasts and concluded that they most likely provide mechanical support to surrounding thin-walled parenchyma via their wall thickenings. Solereder and Meyer (1930) described these idioblasts as waterstorage cells with various thickenings (spiral, reticulate, ring- or ridge-like). The term ‘tracheoidal idioblast’ was originally coined by Foster (1956) to describe idioblasts resembling tracheids because of their wall thickenings. Although these idioblasts may serve in water storage and/or mechanical support, their function is in no way associated with water or mineral conduction (as the term ‘tracheoidal’ might suggest). Burr and Barthlott (1991) further confused the issue by collectively calling these tracheoidal cells a ‘pseudovelamen’ because their thickenings were similar to those of velamen cells. The development of these cortical cells is independent of velamen development. Perhaps because of the term ‘pseudovelamen’, Porembski and Barthlott (1988) included the presence of tracheoidal idioblasts in their survey of velamen in Orchidaceae. Among the 46 vandaceous taxa examined, they found idioblasts in the roots of: Aerides multiflora Roxb., Armodorum siamense Schltr., Holcoglossum quasipinifolium (Hayata) Schltr., and Smitinandia micrantha (Lindl.) Holttum. Cheadle and Kosakai (1982) surveyed occurrence of vessels in roots, stems, inflorescence axes, and leaves of Orchidaceae and included Phalaenopsis sp., Renanthera matutina (Poir.) Lindl., Sarcochilus falcatus R.Br., and Vanda tricolor. They found vessels with scalariform perforation plates in the roots, stems, inflorescence axes, and leaves of Vandeae sensu Dressler (1993). Tracheids were restricted to stems, inflorescence axes, and leaves. The most comprehensive surveys of stegmata in Orchidaceae were completed by Møller and Rasmussen (1984) and more recently by Prychid et al. (2003). The three conditions that exist among Orchidaceae are stegmata present with conical silica bodies, stegmata present with spherical silica bodies, and stegmata absent. Spherical silica bodies were consistently found in Vandeae as well as in the genera Apostasia, Eria, and Dendrobium. According to Rasmussen (1986), conical silica bodies are probably the ancestral condition for the family, with spherical silica bodies secondarily derived from either the conical or the absent condition. Prychid et al. (2003) provided alternatives to this hypothesis and suggested that the presence of both conical and spherical silica bodies may be ancestral or that both independently originated in Apostasia and the epidendroid groups. Studies of leaf anatomy that include Vandeae are less common than those of the root. N. Williams (1979) examined subsidiary cell development and distribution in several groups of Orchidaceae. Although the emphasis of that study was on Oncidieae, 24 species of Vandeae were included. Most Vandeae possessed stomata distributed along the abaxial surface only; Doritis pulcherrima Lindl. (= Phalaenopsis pulcherrima (Lindl.) J.J.Sm.), Saccolabium hendersonianum Rchb.f. (= Dyakia hendersoniana (Rchb.f.) Christenson), and Vanda tricolor exhibited an amphistomatal pattern of distribution. Subsidiary cells ranged from 2–4, and their cell wall divisions were consistently oblique. There was no attempt to derive systematic implications for Vandeae.

Das and Paria (1992) also examined the stomatal structure of several Aeridinae in their survey of Orchidaceae from India. Most taxa were hypostomatic, except for Aerides multiflora Roxb., Rhynchostylis retusa, and Vanda tessellata, which were amphistomatic. They found three categories of stomata: type I consisted of four similar subsidiary cells forming a circle around the guard cells; type II consisted of two elongate and parallel + two short and perpendicular subsidiary cells surrounding the guard cells; type III consisted of undifferentiated subsidiary cells. Among the seven species of Aeridinae studied, type I stomatal complexes were most common, type II stomatal complexes were less common, and type III stomatal complexes were absent in Vandeae. Detailed observations from Carlsward (2004) are provided below for Acampe, Amesiella, Chiloschista, Phalaenopsis, Taeniophyllum (including Microtatorchis), Trichoglottis, and Vanda (including Neofinetia). (BC)

Palynology Schill and Pfeiffer (1977) examined pollen of Aerides, Chiloschista, Cleistostoma (as Sarcanthus), Gastrochilus, Luisia, Phalaenopsis (including Doritis), Rhychostylis, Sarcochilus, Schoenorchis, and Vanda (including Ascocentrum). Their results are organized under the respective genera below. (AP)

Phylogenetics The nomenclatural history of Aeridinae has been characterized by major taxonomic shifts (e.g. Garay 1974, Christenson 1986), and the current set of genera is the result of using ever less obvious floral characters as the bases for recognizing genera. It has long been recognized that generic circumscription in Vandeae is ‘very finely split’ (Dressler 1993), especially compared to Dendrobium and Bulbophyllum as treated in this volume of Genera Orchidacearum. The generic scheme in Vandeae is similar to the situation in some subtribes of Cymbidieae, such as Maxillariinae, Oncidiinae, and Zygopetalinae. Floral features are complicated, and number of pollinia, a column foot, and details of the rostellum and pollinaria provide suites of floral characters for use in delimiting genera, but this has long been recognized to produce a highly artificial classification. A clear parallel existed within the genera related to Comparettia (Oncidiinae), and more and more genera were being carved out of Comparettia and Scelochilus as their floral morphology became more fully understood. If those who wished to delimit more morphologically homogeneous and much smaller genera want to see what the logical outcome of the process they have started (for example, carving out 15 genera from Gomesa s.l.), then subtribe Aeridinae is a good example of an extremely complicated taxonomic scheme that no one can fully understand and use. One could argue that the whole subtribe should be made into a single genus, but even then there are no clearly synapomorphic morphological characters uniting all these genera. Often a combination of monopodial architecture, incumbent anther orientation, and presence of a stipe was used to characterize this group, but all of these are also found in other orchid 113

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groups. In any event, now that their phylogenetic relationships are becoming better understood, many genera can be combined with others (e.g. as in Phalaenopsis and Vanda as circumscribed here), even though in the process more florally heterogeneous genera are being created. Smith (1934) was the first who proposed features of pollinia (numbers, degree of fusion) as the main characters to be used for generic delimitation. Holttum (1958) proposed an improved system on the basis of Smith’s (1934) ideas, which was adopted by several later authors and culminated in systems of Aeridinae based primarily on pollinia number/fusion and secondarily on column foot and labellum characters published by Seidenfaden (1988) and Senghas (1988–1990). Later, Dressler (1993) adopted a slightly modified Seidenfaden–Senghas system and accepted 102 genera (Table B.1). These authors recognized that their classifications were artificial but practical and provided a ‘system to work with, even if further study must change it’ (Dressler 1993). For example, Dressler (1993) placed Phalaenopsis in his series III (two cleft pollinia), whereas Doritis was in series IV (four unequal pollinia) and Nothodoritis in series V (four equal pollinia); here, these are all treated as Phalaenopsis. Clearly, the set of genera as treated here is a ‘work in progress’; it is undisputed that the Aeridinae scheme used previously is highly artificial and does not help us to circumscribe monophyletic generic concepts, but what should be done for each of the major clades (see below) remains to be sorted out in detail. From 1988 until 2005, only Christenson (1994) and Pedersen (1993) published papers about relationships within Aeridinae; Christenson proposed relationships in the Aerides-Vanda alliance and Pedersen for genera related to Pteroceras. With the first molecular phylogenetic treatment of Aeridinae by Topik et al. (2005, 2006) using matK (plastid) and ITS (nuclear ribosomal) markers, it became clear that characters related to pollinia and the column foot could no longer be supported, and the authors identified the stipe and viscidium as useful features in inferring phylogenetic relationships. The Topik et al. (2005) study contained information on 75 species representing 62 genera (most of which are represented by only one species) in 14 clades (which they termed ‘alliances’). Furthermore, it was clear despite the limited sampling of this analysis that at least three genera were not monophyletic (Cleisostoma, Phalaenopsis, and Sarcochilus). Similarly, Carlsward et al. (2006b) found in a study on the evolution of leafless monopodial orchids that Cleisostoma, Vanda, and Tuberolabium are not monophyletic, and Kocyan et al. (2008) showed that Vanda was problematic in a study on the genus Aerides. Interrelationships among the major clades remained unresolved in Topik et al. (2005) as the deeper nodes were only weakly supported. On the other hand, some relationships were supported, such as those between Aerides, Vanda, Christensonia, Trudelia, Neofinetia, Papilionanthe, Ascocentrum, Holcoglossum, and Seidenfadenia (e.g. Christenson 1994). Generally, there is a lack of molecular systematic treatments for most Aeridinae genera, although Phalaenopsis (with Kingidium and Doritis) has been a major exception (e.g. Padolina et al. 2005; Tsai et al. 2006). Phylogenetic treatments for Aerides (Kocyan et al. 2008) and Holcoglossum (Fan et al. 2009; Liu et al. 2011) have also been published, and studies are in various stages of completion 114

for Luisia, Renanthera, and Pteroceras. A more detailed study of Vanda has recently been published (Gardiner et al. 2013). The phylogenetic tree of Aeridinae illustrated here (Fig. B.3) is part of an as yet unpublished phylogenetic study by Alex Kocyan and uses sequence information from four plastid regions (matK exon, trnL intron, trnL-F intergenic spacer, and trnS-G intergenic spacer) and the nuclear ribosomal ITS for more than 200 species and all currently accepted genera except Calymmanthera, Phragmorchis, Saccolabium, Smithsonia, and Xenikophyton. Both parsimony and Bayesian analyses were performed on these data, and we illustrate only the parsimony results; the Bayesian trees are highly similar but mostly have much better support (posterior probabilities, PP), which is reported below in addition to bootstrap percentages (BP). Techniques for the parsimony analysis follow the standard protocols: 1000 replicates of random taxonentry order, extensive swapping (TBR) but saving only five trees per replicate, after which swapping on the trees saved until 10,000 equally parsimonious trees were collected; bootstrapping used the same settings except with no random replicates per bootstrap replicate. Bayesian analyses were started with two different starting trees but otherwise followed standard protocols. Aeridinae as recently circumscribed (Dressler 1993; Chase et al. 2003) is monophyletic (BP 100, PP 1.0; Fig. B.3). Within the subtribe, there are four main subclades with varying levels of bootstrap support but all with PP 1.0: 1) Chiloschista alone, 2) Phalaenopsis alone, 3) clade C, Australia/New Guinea clades, D and E and 4) Thrixspermum/Dimorphorchis plus all the other lettered clades and grade K. No clear support exists for their interrelationships (i.e., they form a polytomy in the strict consensus and Bayesian trees). We show only the parsimony tree, but the Bayesian tree differs in how the members of clade A are arranged (in the Bayesian tree, clade A does not exist). Chiloschista is monophyletic (BP 100, PP 1.0; clade A), but in the Bayesian tree is part of the large polytomy with the four major clades, whereas in the parsimony tree it is sister to just one species of Dimorphorchis, D. beccarii (Rchb.f.) Kocyan & Schuit. (which previously was considered to be in Arachnis), to which Dimorphorchis rossii Fowlie is then sister (i.e. Dimorphorchis is paraphyletic to Chiloschista). These relationships received BP 40-flowered; petals spatulate; pollinia 4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 680. Rhinerrhizopsis Inflorescence up to c. 25-flowered; petals variously shaped, sometimes spatulate; pollinia 2 (partly cleft) or 4; if pollinia 4, then petals narrowly oblong . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 677. Pteroceras Labellum consisting almost entirely of the cylindricclavate, laterally flattened spur, with highly reduced lateral lobes and a minute midlobe; flower bright pink to rosepurple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 651. Dyakia Labellum different; flower whitish, greenish or yellowish, with or without brown or purple markings . . . . . . . . . . 68 Inflorescence with flowers opening in succession, one or a few at a time . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69 Inflorescence with many flowers opening simultaneously . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 Stipe spatulate, much widened near pollinia; rostellum proboscis-like; spur usually much longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 670. Pennilabium Stipe linear to oblong, not notably widened near pollinia; rostellum tooth-like; spur not much longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 703. Trachoma

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70. Column with a distinct foot . . . . . . . . . 705. Tuberolabium Column without a foot . . . . . . . . . . . . . . . 698. Spongiola 71. Column with a distinct foot at least half as long as column, or labellum mobile . . . . . . . . . . . . . . . . . . . . . . . . . 72 Column foot absent or less than half as long as column, labellum rigidly attached . . . . . . . . . . . . . . . . . . . . . . . . . 81 72. Inflorescence producing flowers in succession, one or a few at a time . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73 Inflorescence with all or most flowers opening simultaneously . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77 73. Labellum mobile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74 Labellum rigidly attached . . . . . . . . . . . . . . . . . . . . . . . . . 75 74. Inflorescence finely muricate . . . . . . . . . 654. Grosourdya Inflorescence glabrous . . . . . . . . . . . . . . . 633. Brachypeza 75. Labellum with two parallel keels extending from base of midlobe into short, subglobose spur . . . . . 652. Eclecticus Labellum without keels extending from midlobe into spur . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76 76. Inflorescence with floral bracts arranged in two ranks or quaquaversal; column just longer than wide, without a distinct lateral constriction below stigma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 702. Thrixspermum Inflorescence with quaquaversal floral bracts; column longer than wide, or, if shorter, with a distinct constriction below stigma (column pandurate in front view) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 672. Phalaenopsis 77. Sepals and petals almost filiform; roots warty . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 679. Rhinerrhiza Sepals and petals not filiform; roots smooth . . . . . . . . . 78 78. Labellum articulate with column foot . . . . . . . . . . . . . . 79 Labellum not articulate with column foot . . . . . . . . . . . 80 79. Inflorescence finely pubescent or glabrous; anther with two wiry lateral appendages . . . . . . . . . . 637. Chiloschista Inflorescence glabrous; anther without wiry lateral appendages . . . . . . . . . . . . . . . . . . . . . . . . 688. Sarcochilus 80. Midlobe of labellum slightly projecting beyond lateral lobes; labellum ecallose, densely papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 667. Papillilabium Midlobe of labellum projecting well beyond lateral lobes, or labellum subentire; labellum with one or more calli, not papillose . . . . . . . . . . . . . . . . . . . . . . . . . 672. Phalaenopsis 81. Labellum with two spurs . . . . . . . . . . . 647. Diplocentrum Labellum with a single spur or sac . . . . . . . . . . . . . . . . . 82 82. Floral bracts almost extending to tip of dorsal sepal, finely pubescent . . . . . . . . . . . . . . . . . . . . . . . . . 640. Cleisomeria Floral bracts not more than half the length of dorsal sepal, glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83 83. Mouth of spur or sac mostly closed by one or two calli . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84 Mouth of spur or sac unobstructed . . . . . . . . . . . . . . . . 91 84. Rostellum raised and crest-like, extending across clinandrium . . . . . . . . . . . . . . . . . . . . . . . . . . . . 689. Sarcoglyphis Rostellum not raised and tooth-like or beak-like, not extending across clinandrium . . . . . . . . . . . . . . . . . . . . . 85 ● ●

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85. Interior of back wall of spur with an obliquely erect lamella or an uncinate callus . . . . . . . . . . . . . . . . . . . . . . 86 Interior of back wall of spur without ornaments or with a thick, cushion-shaped callus . . . . . . . . . . . . . . . . . . . . . . 87 ●

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86. Interior of back wall of spur with an obliquely erect lamella inserted well inside the spur . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 675. Pomatocalpa Interior of back wall of spur with an uncinate callus inserted near the mouth . . . . . . . . . . . . . . 683. Robiquetia ●

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87. Interior of back wall of spur without a callus . . . . . . . . 88 Spur on the back wall with a thick, cushion-shaped callus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90 ● ●

88. Midlobe of labellum not projecting beyond lateral lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 691. Schistotylus Midlobe of labellum projecting much wider than lateral lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89 ●

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89. Leaves linear; column foot absent or indistinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 707. Vanda Leaves elliptic; column foot well-developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 672. Phalaenopsis

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91. Sepals and petals connate at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 700. Taeniophyllum Sepals and petals free . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92 ●

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92. Labellum (excluding spur) longer than dorsal sepal; rostellum bifurcate, abruptly curved upwards, often surpassing anther; pollinia 4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93 Labellum (excluding spur) as long as or shorter than dorsal sepal; rostellum entire or bidentate, not abruptly curved upwards, not surpassing anther; pollinia 2 or 4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94

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93. Roots glabrous; pollinia unequal, arranged in two pairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 692. Schoenorchis Roots densely long-papillose; pollinia equal, separate, not arranged in pairs . . . . . . . . . . . . . . . . . 709. Xenikophyton ●

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94. Mouth of spur situated near middle of labellum . . . . . . . . . . . . . . . . 672. Phalaenopsis (subgenus Ornithochilus) Mouth of spur at base of labellum . . . . . . . . . . . . . . . . . 95 ●

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95. Interior of back wall of spur with an obliquely erect lamella . . . . . . . . . . . . . . . . . . . . . . . . . . . 675. Pomatocalpa Interior of back wall of spur without lamellae . . . . . . . 96 ●

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96. Labellum clearly divided into a blade-like epichile and a cup- or pitcher-like hypochile; front rim of hypochile sometimes with two horn-like lobes . . . . . . . . . . . . . . . . 97 Labellum otherwise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98 ●

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97. 98.

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Lobes of epichile absent or broad . . . . 653. Gastrochilus Lobes of epichile almost horn-like . . . . . 696. Smithsonia Labellum bowl-shaped, unlobed . . . . . 650. Drymoanthus Labellum not bowl-shaped, distinctly lobed . . . . . . . . . 99

Rachis of inflorescence much thicker than peduncle; peduncle usually finely muricate........................................... .........................654. Grosourdya (Ascochilopsis alliance) Rachis of inflorescence not much thicker than peduncle; peduncle glabrous............................................................ 100 Base of midlobe extended backwards, partly blocking mouth of spur; pollinia 4, equal............. 691. Schistotylus Base of midlobe not extended backwards over mouth of spur; pollinia 2 or 4 in two unequal pairs..................... 101 Leaves linear, strongly V-shaped in cross-section, not twisted at base through 90°............................................ 102 Leaves linear to elliptic, flat, usually twisted at base through 90°...................................................................... 103 Leaf tip bilobed or praemorse........................ 707. Vanda Leaf tip narrowly obtuse to subacute.................................. .................................................................656. Holcoglossum Pollinia 2, partly cleft or entire, or, if 4, rostellum with two cordate projections............................ 683. Robiquetia Pollinia 4; rostellum projections not cordate...................... .................................................................684. Saccolabiopsis Leaves bilaterally flattened............................................. 105 Leaves dorsiventrally flattened....................................... 107 Inflorescence 1-flowered; labellum clearly divided into hypochile and epichile................................658. Jejewoodia Inflorescence 1- or 2-flowered; labellum entire.......... 106 Inflorescence 1-flowered; flower c. 2 cm wide................... ...................................704. Trichoglottis (T. biglandulosa) Inflorescence 2-flowered; flower < 2 cm wide................... .....................................................................662. Microsaccus

107. Margins of petals serrate; leaf margins smooth or serrate...........................................................657. Hymenorchis Petals with smooth or irregular-erose margins; leaf margins smooth...................................................................... 108 ●

●

108. Pedicel arising from a longitudinal concavity delimited by sharp edges on rachis................................................. 109 Rachis with smooth or not sharply delimited concavities...................................................................................... 116 ●

●

109. Labellum consisting almost entirely of the cylindricclavate, laterally flattened spur; flower bright pink to rose-purple........................................................ 651. Dyakia Labellum not as above; flower whitish, greenish or yellowish, with or without brown or purple markings........... ............................................................................................ 110 ●

●

110. Column with foot about as long as column or longer; labellum mobile; pollinia 2, partly cleft, or 4............... 111 Column foot absent or not as long as column; labellum rigidly attached; pollinia 2, entire................................... 112 ●

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111. Labellum continuing the line of column foot; rostellum tooth-like.......................................................677. Pteroceras Labellum bent upwards relative to column foot; rostellum beak-like....................................660. Macropodanthus ●

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112. Inflorescence with flowers opening in succession, one or a few at a time.................................................................. 113 ●

127

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113.

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114.

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115.

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116.

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117.

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118. 119.

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120.

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121.

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122.

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123. 124. 125. 126.

128

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Inflorescence with many flowers opening simultaneously................................................................................... 115 Stipe strongly spatulate; rostellum proboscis-like; spur usually much longer than wide.............670. Pennilabium Stipe suborbicular, linear or oblong; rostellum tooth-like; spur not much longer than wide................................... 114 Labellum without lateral lobes..............676. Porrorhachis Labellum with well-developed lateral lobes........................ ........................................................................703. Trachoma Column with a foot; labellum midlobe narrower than lateral lobes............................................... 705. Tuberolabium Column without a foot; labellum midlobe wider than lateral lobes....................................................... 698. Spongiola Floral bracts almost extending to tip of dorsal sepal, finely pubescent........................................640. Cleisomeria Floral bracts not reaching more than half the length of dorsal sepal, glabrous...................................................... 117 Interior of spur mouth with one or two horizontal calli that largely obstruct entrance........................................ 118 Interior of spur mouth without calli that largely obstruct entrance, but possibly with a thin lamella on back wall or with calli lower down...................................................... 128 Back wall of spur without callus................................... 119 Back wall of spur with a callus...................................... 123 Column bent downwards, with rostellum projecting into mouth of spur; stipe with a hook-like extension that carries the pollinia................................................706. Uncifera Column not with rostellum projecting into mouth of spur; stipe without a hook-like extension.................... 120 Callus blocking spur entrance pubescent............................ ....................................................................674. Plectorrhiza Callus at spur entrance glabrous.................................... 121 Plant often well over 1 m tall, sometimes shorter; midlobe of labellum tooth-like; pollinia 4......661. Micropera Plant < 1 m tall; midlobe of labellum not tooth-like, suborbicular to flabellate, sometimes linear; pollinia 2 or 4... ............................................................................................ 122 Flowers c. 5 mm diam.; column without a foot; pollinia 4..................................................................697. Smitinandia Flowers > 1 cm diam.; column with a (sometimes indistinct) foot; pollinia 2, incompletely cleft.......628. Aerides Rostellum beak-like......................................................... 124 Rostellum tooth-like........................................................ 125 Anther beak-like, covering rostellum.....699. Stereochilus Anther not beak-like, leaving rostellum exposed............... ........................................................................661. Micropera Inflorescence much shorter than leaves; column often with a pair of stelidia....................................................... 126 Inflorescence elongate, as long as or longer than leaves; column without stelidia.................................................. 127 Interior of spur with a longitudinal septum; stipe broader than long.................................................669. Pelatantheria Interior of spur without septum; stipe strap-shaped, longer than broad................................... 704. Trichoglottis

127. Rostellum gibbose...................................690. Sarcophyton Rostellum flat........................................... 641. Cleisostoma ● ●

128. Inflorescence with rachis shorter than pedicellate ovary, one- to many-flowered.................................................... 129 Inflorescence with rachis longer than pedicellate ovary, 2–many-flowered............................................................. 140 ●

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129. Interior of back wall of spur with a callus or lamella....... ............................................................................................ 130 Interior of back wall of spur without ornaments....... 131 ●

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130. Interior of back wall of spur with a protruding lamella... ................................................................... 704. Trichoglottis Interior of back wall of spur with a low callus not extending to mouth...............................639. Cleisocentron ●

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131. Labellum divided into a flat epichile and cup-shaped hypochile without lateral lobes............. 653. Gastrochilus Labellum not with a cup-shaped hypochile, or hypochile with well-developed lateral lobes................................... 132 ●

●

132. Column with a foot......................................................... 133 Column without a foot................................................... 137 ● ●

133. Labellum mobile........................................688. Sarcochilus Labellum rigidly attached............................................... 134 ● ●

134. Inflorescence 1-flowered; labellum without a callus in centre; pollinia 2, entire..........................685. Saccolabium Inflorescence 1–many-flowered; labellum with or without a callus in centre; pollinia 2, porate or cleft, or 4........ ............................................................................................ 135 135. Flowers opening in succession, one or a few at a time; pollinia 4...............................................702. Thrixspermum Flowers largely opening simultaneously; pollinia 2, cleft but not entirely so............................................................ 136 ●

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136. Midlobe of labellum not longer than lateral lobes............ ...........................................................................695. Singchia Midlobe of labellum longer than lateral lobes................... .............................................................................. 707. Vanda 137. Inflorescence < 1 cm long............................................. 138 Inflorescence > 1 cm long............................................. 139 ●

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138. Leaves usually < 3 cm long; pollinia 2, entire..................... ............................................................................. 664. Omoea Leaves > 5 cm long; pollinia 4.............. 704. Trichoglottis ●

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139. Midlobe of labellum with densely papillose or verrucose surface; mouth of spur pubescent or glabrous; pollinia 4..........................................................................626. Acampe Midlobe of labellum with smooth surface; mouth of spur glabrous; pollinia 2.................................... 707. Vanda 140. Floral bracts bilaterally flattened, distichous...................... ...............................................................702. Thrixspermum Floral bracts dorsiventrally flattened, distichous or quaquaversal..................................................................... 141 ●

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141. Spur with a protruding lamella on interior of back wall... ............................................................................................ 142 Spur without a lamella on interior of back wall.......... 143 ●

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142. Lamella inserted deep inside spur, glabrous....................... ...................................................................675. Pomatocalpa ●

ACAMPE



●

Lamella inserted at mouth of spur, pubescent or glabrous......................................................... 704. Trichoglottis

143. Labellum mobile.............................................................. 144 Labellum rigidly attached............................................... 146



●

● ●

144. Column without a foot.................................630. Arachnis Column with a distinct foot........................................... 145 ● ●

Labellum (excluding spur) as long as or shorter or slightly longer than dorsal sepal; rostellum entire or bidentate, not abruptly curved upwards, not surpassing anther........ ............................................................................................ 159

158. Roots glabrous; pollinia in two unequal pairs..................... ................................................................... 692. Schoenorchis Roots densely papillose-hairy; pollinia equal, not in pairs....................................................... 709. Xenikophyton ●

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145. Column bent knee-like; labellum with a short, finger-like callus in centre...................................... 671. Peristeranthus Column straight; labellum with a bilobed callus in centre...........................................................688. Sarcochilus ●

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146. Pollinia 2, possibly partly but not entirely cleft........... 147 Pollinia 4........................................................................... 156 ● ●

147. Leaves almost overlapping at base, linear, thick and rigid, not twisted at base........................................................... 148 Leaves distant, variously shaped, flexible, often twisted at base through 90°......................................................... 151 ●

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148. Column with a foot......................................................... 149 Column without a foot................................................... 150 ● ●

149. Pollinia deeply cleft into two highly unequal parts............ ...........................................................................695. Singchia Pollinia shallowly cleft, or parts not highly unequal.......... .............................................................................628. Aerides ●

159. Labellum (excluding spur) not shorter than dorsal sepal; midlobe much larger than lateral lobes........626. Acampe Labellum (excluding spur) shorter than dorsal sepal, or midlobe not much larger than lateral lobes................. 160 ●

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160. Spur without a ring of hairs inside.........678. Renanthera Spur with a ring of hairs inside................... 645. Deceptor ● ●

6 2 6 .   AC A M P E Acampe Lindl., Fol. Orchid. Acampe, 1 (1853). Type species: Acampe multiflora (Lindl.) Lindl. (basionym: Vanda multiflora Lindl.), nom. conserv. Sarcanthus Lindl., Bot. Reg., 10, t. 817 (1824), non Lindl. (1826). Type species: Epidendrum praemorsum Roxb.

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●

Derivation of name

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From the Greek akampes , stiff, rigid, probably in reference to the rigid, brittle flowers. ( JW)

150. Margin of stigma not crater-like, raised.............................. ................................................................. 682. Rhynchostylis Margin of stigma crater-like, raised..................................... .............................................................................. 707. Vanda 151. Column with a foot to which lateral sepals are attached.. .............................................................................628. Aerides Column without a foot................................................... 152 ●

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152. Climbing plant > 1 m tall; flowers c. 3 cm wide................. .................................................................... 701. Taprobanea Plant < 1 m tall, not climbing (plant habit not known for Santotomasia); flowers < 1.5 cm wide............................. 153 ●

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153. Pollinia entire; inflorescence with 3 or more branches..... .................... 678. Renanthera (Porphyrodesme alliance) Pollinia cleft or, if entire, inflorescence simple or with 1 or 2 branches only........................................................... 154 ●

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154. Column bent downwards, with rostellum projecting into mouth of spur; stipe with a hook-like extension that carries pollinia.......................................................706. Uncifera Column not with rostellum projecting into mouth of spur; stipe without a hook-like extension.................... 155 ●

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155. Anther with upcurved, peltate rostrum; stipe ligulate....... ..................................................................686. Santotomasia Anther without or with a short, straight rostrum; stipe spatulate...................................................... 683. Robiquetia ●

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156. Labellum with a callus on back wall inside spur................. ..................................................................639. Cleisocentron Labellum without callus inside spur.............................. 157 ●

Description (Plate 57; Fig. 626.1) Coarse epiphytic or lithophytic herbs. Stems simple or branched, stout, leafy. Leaves distichous, conduplicate, coriaceous, sometimes fleshy, apex emarginate, bilobed or praemorse. Inflorescence axillary, racemose, corymbose, or paniculate, shorter than leaves; floral bracts inconspicuous, scale-like, persistent. Flowers with the labellum nearest the inflorescence axis, rigid and fleshy, fragile. Sepals and petals similar, flat to curved; lateral sepals adnate to the spur (when present). Labellum saccate or with a spur, adnate to column, immobile, rigid, variously lobed, fleshy, papillose, tuberculate or warty, sometimes dentate; spur lacking any internal tongue or median septum. Column stout and fleshy, glabrous or papillose; stelidia two, toothlike; foot absent; anther cap ovate, apiculate; pollinia four, globose, unequal, waxy, connected in two pairs, caudicle linear, viscidium ovoid; stigma transverse, concave, rostellum emarginate. (JW)

Distribution (Fig. 626.2) Acampe includes about 10 species distributed from tropical Himalayan regions to China and Indochina, and Southeast Asia, tropical and subtropical Africa, Madagascar, and the islands of the western Indian Ocean. ( JW)

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157. Labellum (excluding spur) slightly to much longer than dorsal sepal; rostellum bifurcate, abruptly curved upwards, often surpassing anther.................................. 158 ●

Anatomy Carlsward et al. (2006a) examined A. papillosa (Lindl.) Lindl. Their results are summarized below. 129

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D

B

F

E

C

A

H

G

J I

K Fig. 626.1.  Acampe praemorsa (Roxb.) Blatt. & McCann. A. Habit; B. Flower, side view; C. Labellum and column, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum and spur; H. Column apex, oblique view; I. Column apex, side view; J. Anther cap, dorsal and ventral views; K. Pollinarium. Single bar = 1 mm; double bar = 1 cm. Drawn by Susanna Stuart-Smith as A. papillosa (Lindl.) Lindl., A from Pantling 243 (K), B-K from cult. Kew accession no. 175–1924. Reproduced with permission from N. R. Pearce and P. J. Cribb (2002), The Orchids of Bhutan. Royal Botanic Garden, Edinburgh and Royal Government of Bhutan.

Leaf Cuticle smooth to ridged along the contours of the epidermal cells. Adaxial cuticle 11.25 μm thick; abaxial cuticle 8.75 μm thick. Hairs multicellular, glandular. Stomata abaxial, superficial to sunken; outer ledges thin, inner ledges thick. Substomatal chambers irregularly shaped. Epidermal cells 130

periclinally oriented to isodiametric. Hypodermis absent. Fibre bundles absent. Mesophyll homogeneous, 16 cells thick. Water-storage cells with birefringent bands. Cells above midrib distinctly modified. Vascular bundles collateral, in one row. Sclerenchyma associated with both xylem and phloem poles. Bundle sheath distinct. Stegmata contain spherical,

ACAMPE

Fig. 626.2.  Distribution map of Acampe.

rough-surfaced silica bodies found in association with phloem and xylem sclerenchyma.

Root Velamen two to three cells thick. Epivelamen cells isodiametric to radially elongate. Endovelamen cells angular, isodiametric to radially elongate; cells of outer layer thicker-walled than those of the inner layer. Distinct endovelamen thickenings absent. Cover cells present over short cells of exodermis. Hairs present. Exodermal cells radially elongate to isodiametric. Long cell walls primarily ○-thickened. Cortex 17 cells wide. Water-storage cells with birefringent bands. Aeration units absent. Endodermal cells ○-thickened. Pericyclic cells thin-walled opposite xylem and thick-walled opposite phloem. Vascular cylinder 12–14-arch. Vascular tissue embedded in sclerenchyma. Pith sclerenchymatous. (BC)

Cytogenetics Chen Xinqi and Wood (2009) recorded a chromosome count of 2n = 38 for A. ochracea (Lindl.) Hochr. and A. rigida (Buch.-Ham. ex Sm.) P.F.Hunt, and 2n = 36, 38, 72 for A. praemorsa (Roxb.) Blatt. & McCann (as A. papillosa). Felix and Guerra (2010) listed 2n = 36, 38 for various species. ( JW)

Phytochemistry Little is known about the phytochemistry of this genus. Evaluation of one species for alkaloid content indicated that these compounds were not present at levels ≥ 0.1% (Lüning 1974). (NV, RG)

Phylogenetics According to Topik et al. (2005), who used a much sparser sampling of genera, Acampe is related to Adenoncos and Micropera, but this relationship found no support in the results published here. Acampe is well supported as sister to monospecific Cottonia, and then this pair is sister to Trichoglottis s.l. (clade J). (AK, MC)

Ecology Species of Acampe are epiphytes or lithophytes from 0 to1300 m. Acampe ochracea is recorded from streamside trees in tropical wet evergreen forest between 300 and 900 m in Sri Lanka. Acampe pachyglossa Rchb.f. is recorded as epiphytic in woodlands, usually in hot, low-lying areas, and in riverine forest in Africa. Acampe praemorsa (Roxb.) Blatt. & McCann is recorded as epiphytic on Anacardium L. (Anacardiaceae) at 260 m in Nepal, on branches of trees in tropical wet evergreen forest and tropical savannas up to 900 m in Sri 131

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Lanka, and on mango (Mangifera indica Wall.; Anacardiaceae) trees in southern India. Acampe rigida is recorded as lithophytic on rocks in China and on limestone in Laos and as epiphytic on trees in tropical wet evergreen forests between 300 and 900 m in Sri Lanka. (JW)

Pollination There is no information available on the pollination of Acampe. ( JW)

Uses Subedi et al. (2011) reported the use in Nepal of a powder made from A. praemorsa as a medicine for rheumatism. Acampe rigida (syn. A. multiflora (Lindl.) Lindl.) was listed as a medicinal plant in Hong Kong (Lawler 1984). Species of Acampe are sometimes cultivated. (AS)

Cultivation Group 2 (see subtribal treatment). (MM)

Taxonomic notes Acampe was proposed in 1853 by John Lindley in his Folia Orchidacea. Although Garay (1972) pointed out that Lindley’s earlier name Sarcanthus (Lindl. 1824, non 1826, typified by Epidendrum praemorsum Roxb.) had priority, the name Acampe has now been conserved. The complex and confusing history of Acampe was reviewed by Seidenfaden (1977). ( JW)

Taxonomic literature Garay, L. A. (1972). On the systematics of the monopodial orchids I. Botanical Museum Leaflets, Harvard University, 23, 149–212. Seidenfaden, G. (1977). Thalia Maravara and the rigid air-blossom. Botanical Museum Leaflets, Harvard University, 25, 49–69. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

or sparsely branching in the basal part. Leaves conduplicate, articulate, distichous, oblong to linear, often semi-terete, apex obtuse to acute, mucronate; leaf sheaths finely transversely wrinkled. Inflorescences lateral, solitary or fascicled, racemose, 1–5-flowered. Flowers inconspicuous, resupinate or not, c. 0.3–1.0 cm wide, opening a few at a time, distichous, green or greenish yellow, becoming more yellow with age, sometimes scented like vanilla. Sepals entire, free, ovate-oblong, often sharply keeled externally. Petals narrower than sepals. Labellum not spurred, concave, entire or trilobed, rigidly attached to column, with an oblong, papillose callus basally; labellum apex usually apiculate. Column without a foot; anther cucullate, rostrate; pollinia four, subspherical; stipe about as long as pollinia. Capsule oblong, subsessile. (AS)

Distribution (Fig. 627.2) Adenoncos is a genus of about 17 species from Thailand, Vietnam, Peninsular Malaysia, Sumatra, Java, Borneo, Sulawesi, Moluccas, and New Guinea. (AS)

Cytogenetics There are no published chromosome counts for the genus. (AP)

Phytochemistry No reports have been found on the phytochemistry of Adenoncos. (NV, RG)

Phylogenetics Adenoncos (clade I) is well supported in the results published here as related to Jejewoodia and Microsaccus. Topik et al. (2005) found Adenoncos closely related to Acampe and Micropera, and Carlsward et al. (2006b) found it to be related to Aerides and Vandopsis. However, both these studies had a much sparser sampling at the generic level, which is likely to have affected the topologies obtained. (AK, MC)

Ecology Species are epiphytes in evergreen forest, from 0 to 1700 m. (AS)

627.  AD E N O N C O S

Pollination

Adenoncos Blume, Bijdr., 381 (1825). Type species: Adenoncos virens Blume Podochilopsis Guillaumin, Bull. Mus. Hist. Nat. (Paris), sér. 2, 34, 478 (1963). Type species: Podochilopsis dalatensis Guillaumin

Little is known about pollination in Adenoncos. Carr (1928) reported that the basal callus of the lip is often eaten away and that its absence is usually correlated with removal of the pollinia. He noted that fruit-set is about 70% in A. major Ridl. (AS)

Derivation of name

No uses have been reported for Adenoncos; it is uncommon in cultivation. (AS)

From the Greek aden, gland, and onkos, mass, referring to the callus at the base of the labellum. (AS)

Description (Plate 58; Fig. 627.1) Epiphytic herbs. Roots terete, not branching, glabrous, smooth. Stem patent-ascending to erect, leafy, rooting at the base, not branching 132

Uses

Cultivation Group 4 (see subtribal treatment). Most species are warm-to intermediate-growing epiphytes that may be cultivated tied to a slab of cork and misted frequently. The roots should be allowed to dry out between waterings. (MM, AS)

AERIDES

B

C

D

O P N M

K R F L H

Q

G

E A I

J Fig. 627.1.  Adenoncos parviflora Ridl. A. Habit; B. Flower, close-up; C. Flower, front view; D. Flower, side view; E. Leaf, transverse section; F. Dorsal sepal; G. Lateral sepal; H. Petal; I. Labellum, dorsal view; J. Labellum, longitudinal section; K. Column and labellum, side view; L. Column, anther cap removed, front view; M. Column, side view; N. Column, back view; O. Anther cap, dorsal view; P. Anther cap with pollinia, ventral view; Q. Pollinarium; R. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Wood 665 (K), others from Kew Spirit coll. no. 54310.

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogeny of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

6 2 8 .   A ER I DES Aerides Lour., Fl. Cochinch., 2, 525 (1790). Type species: Aerides odorata Lour. Aeridium Salisb., Trans. Hort. Soc., 1, 295 (1812). Type species: Aeridium odorum Salisb. Orxera Raf., Fl. Tellur., 4, 37 (1836, publ. 1838). Type species: Orxera cornuta (Roxb.) Raf. (basionym: Aerides cornuta Roxb.) 133

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Fig. 627.2.  Distribution map of Adenoncos.

Polytoma Lour. ex B.A.Gomes, Mem. Acad. Sc. Lisb. Cl. Sc. Pol. Mor. Bel.-Let., n. s. 4, 1, 30 (1868). Type species: Polytoma odorifera Lour. ex B.A.Gomes

Derivation of name From the Latin aer-, air, and the suffix -idus, having the nature of, in reference to the epiphytic habit. ( JW)

Description (Plate 59; Fig. 628.1) Coarse, epiphytic herbs. Stems ascending, enclosed by leafsheaths, with many nodes. Leaves linear-oblong to terete, coriaceous, distichous, apex bilobed, jointed and sheathing at base. Inflorescence racemose or paniculate, lateral, pendent, usually densely many-flowered. Flowers showy. Sepals and petals similar, broad, spreading; lateral sepals decurrent on column foot. Petals smaller than sepals. Labellum continuous with column foot, usually immobile, trilobed, spurred, side lobes decurrent on column, erect, midlobe often erose; spur narrowly conical or horn-shaped, bent forward, usually with swellings or calli inside. Column elongate, often broadened at apex, foot distinct; anther cap two-celled, beaked; pollinia waxy, two, subglobular, unequally cleft, attached by a long, slender stipe to a single, semicircular viscidium; rostellum well developed, bifid after removal of pollinia, porrect. ( JW) 134

Distribution (Fig. 628.2) There are about 20 species of Aerides distributed from Sri Lanka, India, Nepal, Bhutan, Burma, China, Thailand, Indochina, and Malaysia to the Philippines and Indonesia. ( JW)

Palynology Schill and Pfeiffer (1977) examined pollen of A. fieldingii Lodd. ex E.Morren (=A. rosea Lodd. ex Lindl. & Paxton) and described tetrads as convex with laevigate sculpturing and a rounded sexine. (AP)

Cytogenetics Chen Xinqi and Wood (2009) recorded a chromosome number of 2n = 38 for A. falcata Lindl. and A. rosea, and 2n = 36, 38, 40 for A. odorata. Felix and Guerra (2010) listed 2n = 38, 40, 76. Brandham (1999) suggested a basic number of x = 19, which would account for the numerous records of 2x = 38 as well as 4x = 76 along with dysploid variation. ( JW)

Phytochemistry Alkaloids are present at low levels (up to 0.01%) in 13 of 14 species surveyed (Lüning 1964, 1967), although four of the latter are now placed in other genera (Papilionanthe, Seidenfadenia, and

AERIDES

C K

A

D

L

F

E

B

I

H

J

G

Fig. 628.1.  Aerides multiflora Roxb. A. Habit; B. Flower, oblique view; C. Ovary, labellum and column, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum, front view; H. Labellum, longitudinal section; I. Column with anther cap, side view; J. Column, anther cap removed, side view; K. Anther cap, ventral view; L. Pollinarium. Single bar = 1 mm; double bar = 1 cm. Drawn by Susanna Stuart-Smith, A from Gamble 9432 (K), B-L from Trudel 714 (K). Reproduced with permission from N. R. Pearce and P. J. Cribb (2002), The Orchids of  Bhutan. Royal Botanic Garden, Edinburgh and Royal Government of Bhutan.

Vanda). Leaves of A. rosea (cited as A. fieldingii) contain flavonoid glycosides, including the 3-O-glucosides of kaempferol, quercetin, and dihydroquercetin; flavonol O-glycosides (kaempferol and quercetin) are also found in leaves of A. odorata Lour., but both species lack flavone C-glycosides (C. Williams 1979).

A novel stilbenoid derivative, aeridin (2,7-dihydroxy-1,3-dimethoxyphenanthropyran), has been isolated from extracts of the whole plant of A. crispa Lindl. (Anuradha and Rao 1998c). Chemical composition data are available for the floral fragrances of A. crassifolia C.S.P.Parish ex Burb., A. lawrenceae 135

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Fig. 628.2.  Distribution map of Aerides.

Rchb.f., A. odorata, and A. rosea (Kaiser 1993, 2011). The main component of the scent of A. crassifolia is benzyl acetate (75.0%); among the minor components are benzyl alcohol (2.4%), methyl benzoate (5.9%), nonadecane (4.7%), phenylethyl acetate (1.4%), phenylethyl alcohol (2.3%), and others at lower levels (Kaiser 1993). In contrast, the scent of A. lawrenceae is dominated by the monoterpenes, geraniol (35.6%) and nerol (41.9%). Other contributions to the character of this fragrance are made by (E,E)-farnesal (4.5%), geranial (4.0%), methyl phenylacetate (2.7%), neral (1.8%), and the trace components, coumarin, indole and methyl anthranilate (Kaiser 1993, 2011). Methyl ketones such as undecan-2-one (22.4%) and tridecan2-one (41.0%) are among the major components of the scent of A. odorata, together with nerol (25.8%); other methyl ketones are present as minor components (Kaiser 2011). Descriptions of the scent of A. rosea suggest a resemblance to the fragrances of lily of the valley (Convallaria; Asparagaceae) and cyclamen (Cyclamen; Primulaceae). The major component is (E)-nerolidol (40.1%), although several minor components contribute to the quality of the fragrance, including (E)-2,3-dihydrofarnesal, (E)-2,3-dihydrofarnesol (E,E)-farnesal, (E,E)-farnesol, linalool, and 2-phenylethyl acetate (Kaiser 1993, 2011). (NV, RG)

Phylogenetics In the results published here, Aerides is moderately to well supported as related to Renanthera s.l. and Arachnis s.s. (clade 136

G). Aerides has a long history of variable circumscription, and many species have been transferred to other genera and then returned subsequently to Aerides. Recently, Kocyan et al. (2008) found support for Aerides s.s., but the sections of the genus as previously recognized (based on floral characters) were not supported; some new sections were proposed. (AK, MC)

Ecology Aerides species are epiphytes or sometimes lithophytes, from 0 to 2000 m. Aerides krabiensis Seidenf. is found growing in full sun on dry limestone coastal rocks in Thailand. Aerides multiflora Roxb. is recorded as epiphytic on species of Dillenia L. (Dilleniaceae) and Fraxinus L. (Oleaceae) on river banks in Bhutan, on Mangifera indica Wall. (Anacardiaceae) trees in India, and in deciduous forest in Thailand. Aerides odorata often grows high in the branches of trees where it receives ample light; it is recorded from secondary subtropical forests in Bhutan, lithophytic in deciduous forests in Laos, epiphytic on Terminalia catappa L. (Combretaceae) in littoral vegetation in Kalimantan, and from lowland forests, hill forests, sometimes on ultramafic substrate, and lower montane forests in Sabah. Aerides ringens (Lindl.) C.E.C.Fisch. is known from moist semi-evergreen forest between the wet and dry zone and extending into tropical savannas in Sri Lanka. ( JW).

AMESIELLA

Pollination

Description (Plate 60; Fig. 629.1)

Carr (1928) reported that Aerides odorata is pollinated by carpenter bees (Xylocopa sp.), and Kocyan et al. (2008) reproduced a photo taken by Abishkar Subedi of the same species being visited by a Xylocopa species with several pollinia on its head. (AS)

Erect, epiphytic herbs. Stem leafy. Leaves distichously arranged, linear to narrowly elliptic or obovate, with a sheathing leaf base. Inflorescence axillary, simple, racemose, one- to few-flowered; peduncle and rachis glabrous; bracts triangular. Flowers white, sometimes with yellow in throat of labellum. Sepals erect, elliptic, obtuse, lateral sepals spreading. Petals spreading, elliptic, similar to sepals. Labellum concave, simple or obscurely trilobed, obovate or cordate, ecallose, with an elongate, incurved, cylindrical, nectariferous spur at base. Column lacking a foot; pollinia two, porate, stipe oblong-­linear, viscidium suborbicular; stigma concave. Ovary and pedicel glabrous. (PC)

Uses Aerides multiflora is used for ‘vahic’ disorders according to data label on material at Kew collected on Earthwatch Expedition 1983; Subedi et al. (2011) reported the use in Nepal of a powder made from A. multiflora as a tonic and of a paste made from the leaves of A. odorata as a poultice for wounds. In Indochina, fruits (seeds?) of A. odorata and A. falcata have been used to treat wounds and skin diseases (Lawler 1984). Species of Aerides are commonly cultivated. ( JW, AS)

Cultivation Group 2 (see subtribal treatment). Cultivate in containers or baskets filled with a well-draining mixture allowed to dry out between waterings. Most species grow under warm to intermediate conditions. All need good light and air movement. Those from continental Asia should be kept cooler and drier and as brightly lit as possible during the winter. (MM, AS)

Taxonomic notes The gender of Aerides has been the subject of some confusion, species variously being given feminine or neuter epithets. Roxburgh’s original designation of Aerides as feminine is followed here. The genus has been most recently considered by Christenson (1987) and by Kocyan et al. (2008). (PC)

Taxonomic literature Christenson, E. A. (1987). The taxonomy of Aerides and related genera. In Proceedings of the 12th World Orchid Conference (ed. K. Saito and R. Tanaka), pp. 35–40. 12th World Orchid Conference Organizing Committee, Tokyo. Garay, L. A. (1972). On the systematics of the monopodial orchids I. Botanical Museum Leaflets, Harvard University, 23, 149–212. Kocyan, A., de Vogel, E. F., Conti, E., and Gravendeel, B. (2008). Molecular phylogeny of Aerides (Orchidaceae) based on one nuclear and two plastid markers: a step forward in understanding the evolution of the Aeridinae. Molecular Phylogenetics and Evolution, 48, 422–43.

629.   A ME SIE LLA Amesiella Schltr. ex Garay, in Bot. Mus. Leafl., Harvard Univ., 23, 159 (1972). Type: Amesiella philippinensis (Ames) Garay (basionym: Angraecum philippinense Ames)

Derivation of name The name commemorates Oakes Ames (1874–1950), Professor of Botany at Harvard University and well-known orchid taxonomist. (PC)

Distribution (Fig. 629.2) Amesiella is a genus of three species endemic to the Philippines (Luzon and Mindoro). (PC)

Anatomy Carlsward et al. (2006a) examined A. philippinensis. Their results are summarized below.

Leaf Cuticle smooth to ridged along contours of epidermal cells. Adaxial cuticle 7.5 μm thick; abaxial cuticle 5.0 μm thick. Hairs multicellular, glandular. Stomata ad- and abaxial; inner ledges thicker than outer ledges. Substomatal chamber irregularly shaped. Epidermal cells periclinally oriented to isodiametric, infrequently conical abaxially. Hypodermis absent. Fibre bundles absent. Mesophyll 20 cells wide, isobilateral with columnar anticlinal cells surrounding central core of isodiametric cells. Waterstorage cells with finely banded, birefringent wall thickenings. Cells above midrib unmodified. Vascular bundles collateral, in one row. Sclerenchyma associated with both xylem and phloem poles. Bundle sheath distinct. Stegmata absent.

Root Velamen three cells thick. Endovelamen cells angular, isodiametric to radially elongate. Distinct endovelamen thickenings absent. Cover cells present over short cells of exodermis. Hairs present. Exodermal cells radially elongate to isodiametric. Long cell walls ○-thickened to infrequently ∩-thickened. Proliferations present. Cortex 17 cells wide. Water-storage cells with finely banded, birefringent wall thickenings. Aeration units present. Endodermal cells ○-thickened. Pericyclic cells thin-walled opposite xylem and thick-walled opposite phloem. Vascular cylinder 11-arch. Vascular tissue embedded in sclerenchyma. Pith sclerenchymatous. (BC)

Phytochemistry No reports have been found on the phytochemistry of Amesiella. (NV, RG)

Phylogenetics Amesiella belongs to clade E, all members of which are characterized by prominent spurs, but Amesiella is the most extreme, 137

VA N D E A E

K

J

B

L

M I

C H

E A

D N

G

F

Fig. 629.1.  Amesiella philippinensis (Ames) Garay. A. Habit; B. Flower and bract; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum, dorsal view; G. Bract; H. Column and labellum, side view; I. Column, front view; J. Column, side view; K. Anther cap, dorsal view; L. Anther cap, ventral view; M. Pollinarium; N. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone. Habit redrawn after drawing by Blanche Ames from Merrill 5698; others from Kew Spirit Collection no. 49660.

with a several centimetre-long spur and white flowers. Topik et al. (2005) found a similar topology; Amesiella is well supported as related to a set of genera for which we anticipate future changes to circumscription: Ceratocentron, Cryptopus, Hymenorchis, Macropodanthus, Porrorhachis, Trachoma, and Tuberolabium. A great deal more study is required before the taxonomy of this clade can be altered. (AK, MC)

Pollination

Ecology

Cultivation

Species of Amesiella occur from 0 to 2200 m. (PC)

Group 1 (see subtribal treatment). (MM)

138

Nothing is known about pollination in Amesiella, but the white flowers with the long, tubular spur would indicate moth pollination. (AS)

Uses No uses have been reported for Amesiella. Amesiella philippinensis is frequently cultivated. (AS)

ARACHNIS

Fig. 629.2.  Distribution map of Amesiella.

Taxonomic notes Garay (1972) established Amesiella based on Angraecum philippinense Ames, a species with a flower that is remarkably similar to that of Angraecum or Aerangis species. Oakes Ames had already realized the distinctive nature of this genus and had intended to describe it as a new genus but died before he was able to do so. Both recognized that its relationship lay with Asiatic members of Vandeae rather than with the Afro-Madagascan genera. Although distinctive, Amesiella belongs to a problematic group of genera closely allied to Pteroceras, which needs further study. (PC, AS)

Taxonomic literature Garay, L. A. (1972). On the systematics of the monopodial orchids. Botanical Museum Leaflets, Harvard University, 23, 159 (1972). Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

630.   A R AC H N IS Arachnis Blume, Bijdr., 365 (1825). Type species: Arachnis flosaeris (L.) Rchb. f. (basionym: Epidendrum flos-aeris L.) Armodorum Breda, Gen. Sp. Orchid. Asclep., t. 6 (1827). Type species: Armodorum distichum Breda

Arachnanthe Blume, Rumphia, 4, 55 (1849). Type species: Arachnanthe moschifera (Blume) Blume (basionym: Arachnis moschifera Blume) Arrhynchium Lindl., Paxton’s Fl. Gard., 1, 142 (1851). Type species: Arrhynchium labrosum Lindl. & Paxton Esmeralda Rchb.f., Xenia Orchid., 2, 38 (1874), syn. nov. Type species: Esmeralda cathcartii (Lindl.) Rchb.f. (basionym: Vanda cathcartii Lindl.)

Derivation of name From the Greek arachne, spider, in reference to the spider-like flower, with its usually narrow, striped sepals. ( JW)

Description (Plate 61; Fig. 630.1) Epiphytic or lithophytic herbs. Stems branched, often to several metres long, leafy, scrambling. Leaves distichous, oblong or gradually narrowing toward apex, apex bilobed, coriaceous, base sheathing. Inflorescence lateral, often branched, ascending or pendent, few- to many-flowered. Flowers resupinate, often showy, usually widely opening, often spider-like, often fragrant. Sepals and petals similar, free, ovate or (narrowly) oblong to linear, usually widened toward apex, spreading. Lateral sepals and petals often falcate-curved. Labellum trilobed, motile, attached to the column foot by a strap of tissue, base saccate or spurred, midlobe with a raised, central ridge or callus. Column with an 139

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B

A

H

C

F

E

J L

D

K

I

G

Fig. 630.1.  Arachnis longisepala (J.J.Wood) Shim & A.L.Lamb. A. Habit; B. Flower, oblique view; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Apex of dorsal sepal, reverse. G. Pedicel-with-ovary, labellum and column, side view. H. Labellum, front view; I. Entrance to spur, close-up; J. Column, front view; K. Anther cap with pollinarium, dorsal and ventral views; L. Pollinarium. Single bar = 5 mm; double bar = 1 cm. Drawn by Susanna Stuart-Smith from Cockburn in SAN 84949, holotype (A) and Bailes & Cribb 654 (B–L). Reproduced with permission from J. J. Wood (1997), Orchids of Borneo. Volume 3. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

inconspicuous or absent foot; pollinia four, appearing as two masses, those in each pair unequal, waxy, stipe broad, viscidium broadly ovate or saddle-shaped. Capsule cylindrical. ( JW)

Distribution (Fig. 630.2) Arachnis comprises about 14 species distributed from northeastern India and mainland Asia to Taiwan, the Ryukyu Islands, Indonesia (west of Sulawesi), and the Philippines. ( JW)

Cytogenetics Chen Xinqi and Wood (2009) cited a chromosome number of 2n = 38 for A. labrosa (Lindl. & Paxton) Rchb.f. Brandham (1999) also reported 2n = 42. ( JW) 140

Phytochemistry According to Lüning (1964), two species contain low levels of alkaloids (< 0.01%), A. flos-aeris (as A. moschifera), and A. siamense (Schltr.) Tang & F.T.Wang (as Armodorum siamense Schltr.). (NV, RG)

Phylogenetics Arachnis (clade G) was described by Blume in 1825 and more broadly interpreted in the past. Species once under this generic concept were moved to Armodorum, Dimorphorchis, Esmeralda, Renanthera, Stauropsis, Trichoglottis, Vanda, and Vandopsis, and some were moved back again (Tan 1975). With different sampling of genera than used here, Topik et al. (2005) found a relationship

ARACHNIS

Fig. 630.2.  Distribution map of Arachnis.

for Arachnis with Hygrochilus, Sarcoglyphis, Cleisocentron, Cleisostoma, Chiloschista, and Ornithochilus. However, the molecular analysis here places a narrowly defined Arachnis (including Esmeralda and Armodorum) in a well supported clade with Renanthera s.l. and Aerides. In the parsimony analysis (Fig. B.3), A. calcaratum falls outside the rest of Arachnis, but there is only weak support for this position, whereas in the Bayesian analysis (results not shown), this species falls with the rest of the genus. Although sometimes considered a member of Arachnis, A. beccarii falls with Dimorphorchis (clade A) and is considered a member of that genus here. (AK, MC)

Ecology Species of Arachnis are epiphytes or lithophytes from 0 to 2000 m. Arachnis calcarata Holttum is an epiphyte on branches in lower montane forests, sometimes on ultramafic substrate, between 1100 and 2000 m in Borneo. Arachnis cathcartii (Lindl.) J.J.Sm. grows in large clusters on trees in tropical broad-leaved forests and by streams in deep cool shade between 600 and 2000 m in Bhutan. Arachnis clarkei (Rchb.f.) J.J.Sm. is lithophytic on rocks along valleys or epiphytic on tree trunks in open forests between 500 and 2100 m in China. Arachnis flos-aeris occurs in forests, often on limestone, between 300 and 900 m in Borneo. Arachnis grandisepala J.J.Wood occurs in forests at 900 m in Borneo. Arachnis hookeriana (Rchb.f.) Rchb.f. occurs in coastal scrub on sandy soil in Singapore and Malaysia. Arachnis longisepala (J.J.Wood) Shim &

A.L.Lamb is found in forests on ultramafic substrates between 800 and 900 m in Borneo. ( JW)

Pollination Nothing is known about pollination in Arachnis. ( JW)

Uses Christensen (2002) reported that A. flos-aeris is used by the Iban people of Sarawak to cure toothache. Members of this genus are frequently cultivated and have been used in hybridization. ( JW)

Cultivation Group 3 (see subtribal treatment). (MM)

Taxonomic literature Tan, K. W. (1975). Taxonomy of Arachnis, Armodorum, Esmeralda and Dimorphorchis (Orchidaceae). Part I. Selbyana, 1, 1–15. Tan, K. W. (1976). Taxonomy of Arachnis, Armodorum, Esmeralda and Dimorphorchis. Part II. Selbyana, 1, 365–73. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

141

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631.  BIE R MA N N IA

Pollination

Biermannia King & Pantl., J. Asiat. Soc. Bengal, 66, 591 (1897). Type species: Biermannia quinquecallosa King & Pantl.

There is no information on pollination in Biermannia. ( JW)

Derivation of name

No uses have been reported for Biermannia; it is rare in cultivation. ( JW)

Named after Adolph Biermann (-1880), one-time Curator of the Calcutta Botanic Garden. ( JW)

Uses

Cultivation

Description (Fig. 631.1)

Group 1 (see subtribal treatment). (MM)

Epiphytic herbs. Stems enclosed in leaf-sheaths. Leaves fleshy, linear, unequally bilobed, lobules acute, jointed and sheathing at base. Inflorescence racemose, lateral, several-flowered. Flowers resupinate, usually opening successively, short-lived. Sepals and petals free, subequal, lateral sepals attached to column base, usually broader. Petals shorter than sepals. Labellum sessile, narrowly but firmly adnate to column foot at a right angle, sides enveloping or parallel with the column, base with a slit-like opening leading to a hidden pouch, trilobed, lateral lobes parallel to or embracing the column, midlobe linear to narrowly ovate; spur absent or short-saccate (B. calcarata Aver.). Column with an abbreviated foot; pollinia two, waxy, ovoid, slightly grooved or with a small cavity, attached by a linear-oblong stipe to a viscidium. ( JW)

Taxonomic notes

Distribution (Fig. 631.2) Biermannia has about 10 species and occurs in China, India, Thailand, Vietnam, Peninsular Malaysia, Sumatra, Java, and Bali. ( JW)

Delimitation of the genera closely allied to Pteroceras, including Biermannia, needs further study, and Biermannia is tentatively accepted here. Circumscription of this genus is unclear, and the relatively few species currently assigned to it are probably not closely related. Of two species analyzed by Kocyan (Fig. B.3) for his analysis of Aeridinae, one, B. ciliata (Ridl.) Garay, is nested within Grosourdya; the other, B. bimaculata (King & Pantl.) King & Pantl., falls in a clade with Pennilabium and Chroniochilus. (AS)

6 3 2 .  B OGOR I A Bogoria J.J.Sm., Orchid. Java, 566 (1905). Type species: Bogoria raciborskii J.J.Sm.

Derivation of name

Cytogenetics

The genus was named after the city of Bogor in western Java, near which the type species was first collected. Bogor is home of Indonesia’s National Botanic Garden (Kebun Raya). (AS)

Brandham (1999) listed a chromosome number of 2n = 36 for Biermannia. Similarly, Felix and Guerra (2010) reported n = 18. (AP)

Description (Fig. 632.1)

Phytochemistry No reports have been found on phytochemistry of Biermannia. (NV, RG)

Phylogenetics Two species of Biermannia were sequenced for this study, and they fall in two different clades. One is well supported as a member of clade C and is here treated as a member of Grosourdya, and the other species is well supported as a member of a clade with the small genera Chroniochilus, Pennilabium, and Spongiola (in clade E). Nomenclatural unification is for the moment unfeasible until type species of the respective genera are sequenced. All of these have pendent inflorescences with flowers that face downward. (AK, MC)

Ecology Biermannia bigibba (Schltr.) Garay is recorded from forests at 600 m in Sumatra. Biermannia ciliata (Ridl.) Garay is recorded as epiphytic on roadside trees in Pahang, Peninsular Malaysia, and in riverine forests in Peninsular Malaysia and Thailand at elevations of 500–600 m. ( JW) 142

Epiphytic herbs. Roots terete or flattened, not branching, glabrous, smooth. Stem pendent, rooting at base. Leaves 3–5, conduplicate, articulate, distichous, oblong to linear, subfalcate, twisted at base so as to lie in one plane with stem, margins entire, apex unequally bilobed. Inflorescence lateral below lowest leaf, pendent, solitary, racemose, many-flowered; rachis continuing the line of the peduncle, flexuose, slightly thicker than peduncle. Flowers resupinate on pendent inflorescence, about 0.5 cm wide, probably short-lived, opening a few at a time, quaquaversal. Sepals and petals entire, free, elliptic to obovate, obtuse. Labellum rigidly attached to apex of column foot, saccate at base, trilobed, side lobes falcate. Column with foot as long as the column or longer; anther cucullate; pollinia four, in two masses, stipe about as long as pollinia (B. taeniorhiza (Schltr.) Garay with two stipes), narrowly linear; viscidium reduced; rostellum bidentate. (AS)

Distribution (Fig. 632.2) Bogoria is a genus of four species from Sumatra, Java, Borneo, the Philippines, Sulawesi, and New Guinea. (AS)

Phytochemistry No reports have been found on the phytochemistry of Bogoria. (NV, RG)

BOGORIA

B

C H

E G

D

F

A

K I

L J

Fig. 631.1.  Biermannia bimaculata (King & Pantl.) King & Pantl. A. Habit; B. Inflorescence; C. Flower, side view; D. Column and labellum, side view; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum, front view; I. Column with anther cap, front view; J. Pollinarium; K. Capsule; L. Capsule, transverse section. Double bar = 1 cm. Drawn by Susanna Stuart-Smith after King & Pantling, The Orchids of the Sikkim Himalaya, t. 267 (1898). Reproduced with permission from N. R. Pearce and P. J. Cribb (2002), The Orchids of Bhutan. Royal Botanic Garden, Edinburgh and Royal Government of Bhutan.

Phylogenetics Bogoria is part of the Australia clade. Topik et al. (2005) and Carlsward et al. (2006b) produced a similar set of relationships. The position of Bogoria in this clade is somewhat anomalous as the genus occurs mostly from Malesia to the Philippines, but one species reaches New Guinea. The single species sampled so far

is sister to the rest of the Australia clade in the Bayesian analysis, but its position in both analyses is weakly supported. (AK, MC)

Ecology All species of Bogoria are epiphytes in evergreen forests from 500 to 1500 m. (AS) 143

VA N D E A E

Fig. 631.2.  Distribution map of Biermannia.

Pollination

Derivation of name

Nothing is known about pollination in Bogoria. (AS)

From the Greek brachys, short, and peza, foot, in reference to the characteristically short column foot. ( JW)

Uses No uses have been reported for Bogoria; it is rare in cultivation. (AS)

Description (Plate 62; Fig. 633.1)

Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogeny of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

Epiphytic herbs. Stems with 1–6 leaves dorsiventrally flattened or semi-terete, fleshy, broad, to 15–25 × 5 cm. Inflorescences 1–3, arching to pendent, as long as leaves or shorter, rachis clavate, not furrowed above flower nodes, 1–25-flowered. Flowers resupinate, fugacious. Sepals and petals free, spreading, narrowly oblong, to c. 1 cm long. Dorsal sepal erect, concave. Lateral sepals oblique. Labellum tubular, spurred, distally trilobed, articulated to column foot, side lobes erect, auriculate, midlobe recurved; spur bag-like. Column arcuate, with an inconspicuous foot; pollinia two, each split to the middle from the base, stipe cuneate-triangular or oblong, as long as pollinia; stigma terminal; rostellar projection triangular, bidentate. ( JW)

633.  BR AC H Y P E Z A

Distribution (Fig. 633.2)

Brachypeza Garay, Bot. Mus. Leafl., Harvard Univ., 23, 163 (1972). Type species: Brachypeza archytas (Ridl.) Garay (basionym: Saccolabium archytas Ridl.)

About ten species of Brachypeza are distributed from Thailand and Indochina through Malaysia and Indonesia north to the Philippines and east to New Guinea. ( JW)

Cultivation Group 1 (see subtribal treatment). (MM)

Taxonomic literature

144

BRACHYPEZA

L C

J

B N K M

O

I

P D F Q A E

G

H

Fig. 632.1.  Bogoria raciborskii J.J.Sm. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum, dorsal view; H. Labellum, longitudinal section; I. Column and labellum, side view; J. Column, front view; K. Column apex, anther cap removed; L. Column, side view; M. Anther cap, dorsal view; N. Anther cap with pollinarium, ventral view; O. Anther cap, ventral view; P. Pollinarium; Q. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Kew Spirit Collection no. 48150.

Cytogenetics Brandham (1999) reported chromosome 2n = 36–38 for the genus. (AP)

Ecology numbers

of

Phytochemistry No reports have been found on the phytochemistry of Brachypeza. (NV, RG)

Phylogenetics Brachypeza is a member of clade C, and in the circumscription used here includes some species that were formerly in Pteroceras. It is well supported as sister to Grosourdya. (AK, MC)

Brachypeza archytas (Ridl.) Garay is recorded from tall, closed forests on phosphate–pebble limestone and clay loam at 200 m on Christmas Island, Indian Ocean. Brachypeza indusiata (Rchb.f.) Garay occurs as an epiphyte in deep shade in lowland forests from sea level to 300 m in Peninsular Malaysia, in forests on limestone and in mixed dipterocarp forest at 200–300 m in Sarawak, and also forests in West Papua and Sabah at 200–300 m. Brachypeza zamboangensis (Ames) Garay grows as an epiphyte low on tree trunks and on lianas in forests on limestone at 500–600 m in Sabah and the Philippines. ( JW)

145

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Fig. 632.2.  Distribution map of Bogoria.

Pollination

Derivation of name

There is no information available on the pollination of Brachypeza. ( JW)

From the Greek kalymma, hood, and the Latin anthera, anther, referring to the hood of the anther, which covers the apex of the column. (AS)

Uses No uses have been reported for Brachypeza; it is uncommon in cultivation. ( JW)

Cultivation Group 1 (see subtribal treatment). (MM)

Taxonomic notes Garay (1972) noted that Brachypeza is distinguished from Pteroceras by the elongate column with a short foot, pendent labellum, and distinctly cleft pollinia. However, in Pteroceras (q.v.) the pollinia are also cleft. As a result of the phylogenetic analyses by Kocyan (Fig. B.3), several species with a long column foot, formerly included in Pteroceras, will be included in Brachypeza. See also the taxonomic notes under Pteroceras. ( JW, AS)

Description (Fig. 634.1) Epiphytic herbs. Roots terete, not or sparsely branching, glabrous, smooth. Stem patent to pendent, rooting at the base or along most of its length. Leaves conduplicate, articulate, distichous, narrowly oblong to linear; margins entire or crenulate near apex; apex asymmetric or bidentate; leaf-sheaths when young with fimbriate, membranous margin. Inflorescence lateral, solitary or fascicled, racemose or more frequently paniculate, with up to three levels of branching, 10–25-flowered. Flowers resupinate, probably ephemeral, many opening at the same time, quaquaversal, whitish or yellowish. Sepals and petals entire, free, narrow, acuminate. Labellum hinged with the apex of the column foot, not spurred, trilobed, midlobe longer than the lateral lobes, with a callus or pair of calli between lateral lobes. Column terete; column foot shorter than column; anther helmet-shaped, with a hood-like extension over the apex of the column; pollinia four, unequal, separate, subglobose, stipe about as long as the pollinia, oblong to linear, viscidium disc-shaped. (AS)

634.  C ALY MMA N TH E R A

Distribution (Fig. 634.2)

Calymmanthera Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 955 (1913). Type species: Calymmanthera tenuis Schltr.

Calymmanthera is a genus of five (possibly only three) species from the Moluccas, New Guinea, the Solomon Islands, and Fiji. (AS)

146

C ALYMMANTHERA

1 mm

J A 5 mm

E

G 1 cm 5 mm

F

C

B

H

5 cm

1 cm

D 5 mm

I

Fig. 633.1.  Brachypeza zamboangensis (Ames) Garay. A. Habit; B. Flower, oblique view; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum and column, side view; G. Labellum and column, longitudinal section; H. Anther cap, dorsal view; I. Pollinarium; J. Ovary, transverse section. Drawn by C. L. Chan and Chin Wan Wai from a plant cultivated at Tenom Orchid Centre, Sabah. Reproduced with permission from J. J. Wood (1997), Orchids of Borneo. Volume 3. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

Cytogenetics

Pollination

Felix and Guerra (2010) reported a chromosome number of 2n = 38 for the genus. (AP)

Nothing is known about pollination in Calymmanthera. (AS)

Phytochemistry No reports have been found on phytochemistry of Calymmanthera. (NV, RG)

Phylogenetics There is no phylogenetic information is available for this genus. (AK)

Ecology Species are epiphytes in forests from 0 to 1700 m. (AS)

Uses No uses have been reported for Calymmanthera; it is rare in cultivation. (AS)

Cultivation Group 2 (see subtribal treatment). (MM)

Taxonomic notes Calymmanthera has not yet been included in phylogenetic studies. Based on morphology and biogeography, it may be expected that Gunnarella is among its nearest relatives. (AS) 147

VA N D E A E

Fig. 633.2.  Distribution map of Brachypeza.

635.  C E R ATO C E N TRO N

Distribution (Fig. 635.2)

Ceratocentron Senghas, Orchidee (Hamburg), 40, 89 (1989). Type species: Ceratocentron fesselii Senghas

Ceratocentron is a monospecific genus endemic to the Philippines (Luzon). ( JW)

Derivation of name From the Greek keras, horn, and kentron, spur, in reference to the long spur adorned with a solid, upward-curving, horn-like, ventral protuberance. ( JW)

Description (Fig. 635.1) Epiphytic herb. Stem 3–5-leaved. Leaves broadly lanceolate to narrowly ovate, acuminate, coriaceeous. Inflorescences borne below the leaves, c. 1.0–3.5 cm long, 2–5-flowered, flowers opening simultaneously. Flowers resupinate, 1.8–2.0 cm wide, sepals, petals, and spur flame-red to orange, labellum pink, horn-like protuberance on spur pink. Sepals and petals free. Labellum immobile, entire, ovate, apiculate, margin minutely erose, 4 mm wide; spur cylindrical, 6 mm long, adorned with a solid, upward-curving, horn-like ventral protuberance, internal walls without callosities. Column with lateral auriculate wings; foot absent; pollinia two, entire, stipe linear, viscidium rounded to rhomboid; stigma broad; rostellum acutely bidentate. ( JW) 148

Cytogenetics No chromosome numbers have been recorded for Ceratocentron. ( JW)

Phytochemistry No reports have been found on phytochemistry of Ceratocentron. (NV, RG)

Phylogenetics Ceratocentron is a member of clade E, and the single species included in the analyses here is well supported as sister to the single sampled species of Tuberolabium. This pair is then sister to Amesiella/Cryptopylos/Dyakia/Macropodanthus. The previous molecular analysis that included Ceratocentron and Tuberolabium agreed upon their relationship (Topik et al. 2005). (AK, MC)

Ecology Ceratocentron fesselii occurs in exposed conditions in native and secondary forests at 700–1000 m. ( JW)

C ERATOCENTRON

B

C

E

D

F

G

H

I

J

A

Fig. 634.1.  Calymmanthera major Schltr. A. Habit, ×0.66; B. Flower, ×8; C. Anther cap, dorsal and ventral views, ×14; D. Column and labellum, ×14; E. Pollinia, ×14; F. Dorsal sepal, ×8; G. Petal, ×8; H. Lateral sepal, ×8; I. Column, ×14; J. Labellum, ×14. All magnifications as originally published. Drawn by Sue Wickison from Wickison 137. Reproduced with permission from B. A. Lewis and P. J. Cribb (1991), Orchids of the Solomon Islands and Bougainville. Royal Botanic Gardens, Kew.

149

VA N D E A E

Fig. 634.2.  Distribution map of Calymmanthera.

Pollination

6 3 6 .  C H A M A EA NT H U S

There is no information available on the pollination of Ceratocentron. The bright orange-red colour of the flowers and their morphology would indicate bird-pollination. (AS, JW)

Chamaeanthus Schltr. in J.J.Sm., Orch. Jav., 552 (1905). Type species: Chamaeanthus brachystachys Schltr.

Derivation of name Uses No uses have been reported for Ceratocentron; it is occasionally cultivated. ( JW)

Cultivation Group 2 (see subtribal treatment). (MM)

Taxonomic notes The generic delimitation of the genera closely allied to Pteroceras, including Ceratocentron, needs further study. Ceratocentron is only tentatively accepted here. It is easily recognized by the bright orangered flowers and the horn-like process on the labellum. (AS)

Taxonomic literature Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

150

From the Greek chamai, dwarf, and anthos, flower, referring to the insignificant flowers. ( JW)

Description (Fig. 636.1) Epiphytic herbs. Stems only a few centimetres long. Leaves few, fleshy, up to 7 × 1 cm, jointed and sheathing at base. Inflorescence lateral, simple, with up to about 20 flowers; rachis usually clavate. Flowers ephemeral, only a few open at a time, greenish yellow, superficially resembling Bulbophyllum. Sepals and petals similar, often linear and acuminate, to 4–6 × 3 mm; lateral sepals adnate to column foot. Labellum motile, trilobed, without a spur, sometimes saccate, lateral lobes auriculate, margins fimbriate, midlobe conical. Column with a foot 3 mm long, rostellar projection present; pollinia two, entire (although some specimens have a tiny notch), stipe ligulate, about twice the diameter of pollinia, viscidium obovate. ( JW)

Distribution (Fig. 636.2) Chamaeanthus consists of three species distributed in southern Thailand, Java, and Borneo, north to Taiwan and the Philippines. ( JW)

C H AMAEANTHUS

D

C

E

K J M

N

G

L I

H P O F A

B

Fig. 635.1.  Ceratocentron fesselii Senghas. A. Habit; B. Inflorescence; C. Flower, front view; D. Flower, side view; E. Flower, from above; F. Bract; G. Dorsal sepal; H. Lateral sepal; I. Petal; J. Column and labellum, from above; K. Column and labellum, side view; L. Labellum, longitudinal section; M. Column, anther cap removed, side view; N. Column, anther cap removed, side view; O. Anther cap; P. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Loher 2140 (K); flowers from photographs in H.H. Fessel & P. Balzer, Native Philippine Orchids, 121 (2000); others redrawn after K. Senghas in Orchidee (Hamburg) 40, 85–90 (1989).

Cytogenetics

Ecology

Brandham (1999) cited a chromosome count of 2n = 38 for the genus. (AP)

All three species are epiphytes from 0 to 900 m. Chamaeanthus brachystachys is recorded as epiphytic in hill forests on ultramafic substrate in Sabah and in swampy podsol forest in southern Kalimantan, Borneo. Comber (1990) recorded C. brachystachys growing in dry areas on isolated trees in Java. Chamaeanthus wenzelii Ames occurs on trees along seashores in the Philippines. ( JW)

Phytochemistry No reports have been found on the phytochemistry of Chamaeanthus. (NV, RG)

Phylogenetics There is no phylogenetic information available for this genus. (AK)

Pollination No information on the pollination of Chamaeanthus is available. ( JW) 151

VA N D E A E

Fig. 635.2.  Distribution map of Ceratocentron.

Uses

From the Greek cheilos, lip, and chistos, cleft, in reference to the divided labellum. ( JW)

clustered at stem apex at anthesis). Roots flattened, ribbon-like or subcylindric, scabrid, green, photosynthetic. Leaves (when present) oblong-elliptic to ligulate, obliquely subacute or unequally bilobed, sometimes slightly falcate, articulate. Inflorescence lateral, racemose, rarely paniculate (C. ramifera Seidenf. and C. trudelii Seidenf.), erect or pendent, many-flowered, pilose or glabrous; floral bracts ovate-lanceolate. Flowers resupinate, ephemeral, fragrant, white to yellow, with or without reddish or purple spots. Sepals and petals free, subequal, oblong-ovate or oblong-elliptic, lateral sepals and petals usually laterally adnate to column foot by a broad base. Labellum clawed, articulate to apex of column foot, motile, trilobed, basal portion concave or saccate, with an erect or ascending, entire or weakly bilobed, pilose callus on inner surface of posterior wall, side lobes erect, midlobe commonly abbreviated, transverse, entire or rarely bilobed, disc often pilose. Column semi-terete, foot perpendicular to column, broad, at least twice as long as column; clinandrium shallowly excavate, with a membranous posterior margin; anther cap provided on either side with a long-filiform, glandular-tipped appendage, or appendages reduced to a small tooth; pollinia four, waxy, stipe sublinear, viscidium small; stigma transverse, deeply set; rostellum deflexed, shortly bifid after removal of pollinarium. ( JW)

Description (Plate 63; Fig. 637.1)

Distribution (Fig. 637.2)

Epiphytic or lithophytic, usually stemless and commonly leafless herbs (C. godefroyana (Rchb.f.) Schltr. may have persistent leaves

There are about 10 (possibly more) species of Chiloschista collectively distributed from China and the Indian subcontinent

No uses have been reported for Chamaeanthus; it is rare in cultivation. ( JW)

Cultivation Group 4 (see subtribal treatment). (MM)

Taxonomic literature Comber, J. (1990). Orchids of Java. Bentham-Moxon Trust, Royal Botanic Gardens, Kew.

637.  C H ILO SC H ISTA Chiloschista Lindl., Edwards’s Bot. Reg., 18, sub. t. 1522 (1832). Type species: Chiloschista usneoides (D.Don) Lindl. (basionym: Epidendrum usneoides D.Don)

Derivation of name

152

C HILOSCHISTA

C N O P

B

L

I

J

M

E

G

K

A

F

D

H

Fig. 636.1.  Chamaeanthus brachystachys Schltr. A. Habit; B. Inflorescence; C. Flower; D. Bract; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum; I. Column and labellum, side view; J. Column, front view; K. Column, anther cap removed, front view; L. Column, side view; M. Column, anther cap removed, side view; N. Anther cap, dorsal view; O. Anther cap, ventral view; P. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Kew Spirit Collection no. 56439.

through Southeast Asia to Australia, the Palau Islands, and Fiji. ( JW)

Anatomy Carlsward et al. (2006a) examined three species of Chiloschista. Their results are summarized below.

Leaf Cuticle smooth to ridged along the contours of epidermal cells. Ad- and abaxial cuticle less than 1.25 μm thick. Hairs simple, eglandular. Epidermal cells raised around base of hairs. Stomata

ad- and abaxial; outer ledges thin to moderate, inner ledges moderate to thick; cuticular horns inconspicuous. Substomatal chambers small, irregularly shaped. Epidermal cells periclinally oriented to isodiametric. Hypodermis absent. Fibre bundles absent. Mesophyll homogeneous, 10–14 cells wide. Water-storage cells with birefringent walls absent. Cells above midrib unmodified. Vascular bundles collateral, in one row. Thin-walled sclerenchyma associated with phloem and xylem poles. Stegmata absent.

Root Velamen absent to three cells thick. If present, epivelamen cells isodiametric in C. lunifera; isodiametric to radially elongate in 153

VA N D E A E

Fig. 636.2.  Distribution map of Chamaeanthus.

C. parishii; radially elongate in all other specimens examined. Endovelamen cells angular, isodiametric to radially elongate. Distinct endovelamen thickenings absent. Cover cells present over short cells of exodermis. Hairs present in C. lunifera and C. parishii. Hyphae present in C. lunifera and C. parishii. Exodermal cells radially elongate to isodiametric. Long cell walls primarily ○-thickened. Proliferations present in C. lunifera, C. parishii, and C. usneoides (D.Don) Lindl. Cortex 10–13 cells wide. Hyphae present in C. lunifera and C. parishii. Water-storage cells with birefringent walls absent. Aeration units present. Endodermal cells ○-thickened. Pericyclic cells thin-walled opposite xylem and thick-walled opposite phloem. Vascular cylinder 5–8-arch. Vascular tissue embedded in sclerenchyma. Pith sclerenchymatous. (BC)

(Lüning 1967; Lawler and Slaytor 1969). The floral fragrance of C. lunifera is due mainly to methyl (E)-cinnamate (25.0%) and (E)-geranylacetone (10.4%), although (Z)-heptadec-8-ene (22.2%) and pentadecane (26.0%) are also present as major components (Kaiser 2011). (NV, RG)

Phylogenetics

Palynology

In the Bayesian analysis, Chiloschista is an isolated genus, but the parsimony tree puts it in a weakly supported position as sister to Dimorphorchis plus Thrixspermum in clade A. Topik et al. (2005) and Carlsward et al. (2006b) found the genus at different positions in their trees. One reason for the different placements may be that matK sequences used are pseudogenes for Chiloschista. Additional study is needed. (AK, MC)

Schill and Pfeiffer (1977) studied pollen tetrads of C. lunifera (Rchb.f.) J.J.Sm. and described them as convex with laevigate sculpturing and a rounded sexine. (AP)

Ecology

Cytogenetics Brandham (1999) reported chromosome counts of 2n = 38, confirmed by Felix and Guerra (2010). (AP)

Phytochemistry Relatively high levels of alkaloids (> 0.1%) are found in C. lunifera, whereas C. phyllorhiza (F.Muell.) Schltr. contains them at < 0.01% 154

Species are epiphytes from 0 to 2000 m. Kores (1991) recorded Chiloschista godefroyana as epiphytic on trees on limestone and in coastal areas in Fiji. Comber (1990) reported C. javanica Schltr. as epiphytic on tree trunks where the light level is high, e.g., on roadside trees, on poorly grown teak, etc. Chiloschista lunifera is epiphytic on trees in deciduous forests at 200 m and on trees overhanging streams at 600 m in Thailand. Pearce and Cribb (2002) recorded C. parishii as epiphytic on the branches of Panax pseudoginseng Wall. (Araliaceae) between 600 and 1800 m in

C HILOSCHISTA

D A

C I

B E

J F M

L

K

H G

Fig. 637.1.  Chiloschista usneoides (D.Don) Lindl. A. Habit; B. Inflorescence; C. Infructescence; D. Flower, front and back view; E. Ovary, labellum, and column, from above; F. Dorsal sepal; G. Lateral sepal; H. Petal; I. Labellum, longitudinal section showing disc; J. Column, side view; K. Column apex, two views; L. Anther cap, dorsal and ventral views; M. Pollinarium. Single bar = 1 mm; double bar = 1 cm. Drawn by Susanna Stuart-Smith from Ludlow et al. 20544 (A & B, D–M) and Hooker 192 (C). Reproduced with permission from N. R. Pearce and P. J. Cribb (2002), The Orchids of Bhutan. Royal Botanic Garden, Edinburgh and Royal Government of Bhutan.

Bhutan. Chiloschista phyllorhiza occurs epiphytically in mangrove and Melaleuca L. (Myrtaceae) swamps in Australia. Chiloschista segawae (Masam.) Masam. & Fukuy. is epiphytic on tree trunks in forests between 700 and 1000 m in Taiwan. Pearce and Cribb (2002) recorded C. usneoides as epiphytic on oaks and in

evergreen forests between 700 and 1900 m in Bhutan; it also occurs as a lithophyte on rocks in oak forests at about 1700 m in Bhutan. Chiloschista yunnanensis Schltr. is epiphytic on tree trunks at forest margins and in open forests between 700 and 2000 m in China. ( JW) 155

VA N D E A E

Fig. 637.2.  Distribution map of Chiloschista.

Pollination No information on the pollination of Chiloschista is available. ( JW)

Uses No uses have been reported for Chiloschista; it is sometimes cultivated. ( JW)

Cultivation

Pearce, N. and Cribb, P. J. (2002). Flora of Bhutan, including a record of plants from Sikkim and Darjeeling. Volume 3. Part 3. The orchids of Bhutan. Royal Botanic Garden Edinburgh and Royal Government of Bhutan. Seidenfaden, G. (1988). Orchid genera in Thailand XIV. Fifty-nine vandoid genera. Opera Botanica, 95, 168–81. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

Group 1 (see subtribal treatment). Species may be grown on slabs of cork with some moss under warm to cool conditions (depending on the species) with high humidity and fairly high light levels. Plants should be misted frequently but allowed to dry out between waterings. (MM, AS)

6 3 8 .  C H RONI OC H I LU S

Taxonomic literature

From the Greek chronios, lasting, persistent, and cheilos, lip, referring to the unusually long-lasting flowers of the type species. ( JW)

Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogeny of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Comber, J. (1990). Orchids of Java. Bentham-Moxon Trust, Royal Botanic Gardens, Kew. Kores, P. (1991). Orchidaceae. In Flora vitiensis nova: a new flora of Fiji (spermatophytes only). Volume 5, Angiospermae: dicotyledons, families 170–186, monocotyledons, family 32 (ed. A. C. Smith). National Tropical Botanical Garden, Lawai, Kauai, Hawaii.

156

Chroniochilus J.J.Sm., Bull. Jard. Bot. Buitenzorg, sér. 2, 26, 81 (1918). Type species: Chroniochilus tjidadapensis J.J.Sm.

Derivation of name

Description (Fig. 638.1) Epiphytic herbs. Stems erect to pendent. Leaves few, flat, unequally bilobed. Inflorescence simple, few-flowered; rachis fleshy or sometimes flattened. Flowers resupinate. Sepals and petals free, spreading. Labellum motile, sessile, sagittate, conical and solid in front,

C H RONIOCHILUS

3 cm

1 mm

M

A

I 4 mm

B G

E

C

D

6 mm

1 cm

4 mm

L

2 mm

H 6 mm

J

6 mm

K

F

Fig. 638.1.  Chroniochilus virescens (Ridl.) Holttum. A. Habit; B. Flower, oblique view; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum, front view; G. Labellum and column, front view; H. Ovary, labellum, and column, side view; I. Ovary, labellum, and column, longitudinal section; J. Anther cap, ventral view; K. Anther cap, dorsal view; L. Pollinarium; M. Ovary, transverse section. Drawn by Chan Chew Lun from a plant cultivated at Tenom Orchid Centre, Sabah. Reproduced with permission from J. J. Wood et al. (2011), The Orchids of Mount Kinabalu, Volume 2. Natural History Publications (Borneo), Kota Kinabalu, Sabah.

with large auriculate side lobes; spur absent. Column with a distinct foot to which the labellum is elastically attached; pollinia two, entire, stipe linear-oblong, viscidium oval, at least half as long as the stipe; rostellar projection prominent. ( JW)

Phytochemistry

Distribution (Fig. 638.2)

Phylogenetics

The four species of Chroniochilus are collectively distributed from southern Thailand through Malaysia to Indonesia. ( JW)

Cytogenetics No chromosome numbers have been reported for Chroniochilus. ( JW)

No reports have been found on the phytochemistry of Chroniochilus. (NV, RG)

Chroniochilus is well supported as a member of a clade with the small genera Biermannia (at least one species), Pennilabium, and Spongiola (in clade E). Type species of the respective genera need to be sequenced before altering generic limits. All of these have pendent inflorescences with flowers that face downward. (AK, MC) 157

VA N D E A E

Fig. 638.2.  Distribution map of Chroniochilus.

Ecology These species are epiphytes from 0 to 1000 m. Chroniochilus minimus (Blume) J.J.Sm. and C. virescens (Ridl.) Holttum are recorded from forests on ultramafic substrates between 700 and 1000 m in Borneo, Chroniochilus minimus grows on tea bushes between 800 and 1000 m in Java. Chroniochilus thrixspermoides (Schltr.) Garay has been recorded from a coffee plantation at 300 m in Sumatra. Chroniochilus virescens grows on small trees in grasslands at 100 m in Thailand. ( JW)

tentatively accepted here. The flowers are similar to those of Pteroceras, but the spur-like distal part of the labellum is solid, not hollow, and the pollinia are entire, not cleft. (AS)

6 3 9 .  C L EI S OC ENT RON Cleisocentron Brühl, Guide Orchids Sikkim, 136 (1926). Type species: Cleisocentron trichromum (Rchb.f.) Brühl (basionym: Saccolabium trichromum Rchb.f.)

Pollination

Derivation of name

There is no information available on the pollination of Chroniochilus. ( JW)

From the Greek kleiso-, hook or tongue, and kentron, a sharp point, in reference to the projection within the spur. ( JW)

Uses

Description (Plate 64; Fig. 639.1)

No uses have been reported for the genus; it is rare in cultivation. ( JW)

Epiphytic herbs. Stems pendent or erect, simple or branching, up to 1 m in length. Leaves oblong, ligulate or terete, unequally bilobed or acute. Inflorescence supra-axillary, racemose. Flowers resupinate, pinkish white, various shades of blue, deep magenta or dark purple. Sepals and petals spreading. Labellum firmly adnate to column, urceolate, trilobed above; spur cylindrical, gently curving, interior containing either an upward-pointing, central protuberance on the back wall or a decurved shelf-like back wall callus; front wall callus flap-like; median septum absent. Column erect, cylindrical, foot decurrent on the back wall of labellum as

Cultivation Group 2 (see subtribal treatment). (MM)

Taxonomic notes The generic delimitation of the genera closely allied to Pteroceras, including Chroniochilus, needs further study. Chroniochilus is only

158

C LEISOCENTRON

I

F

D

K

L

J

G

P

H

C M

O E B

A

N

Fig. 639.1.  Cleisocentron merrillianum (Ames) Christenson. A, B. Habits; C. Portion of leaf, close-up; D. Leaf apex variation; E. Leaf, two transverse sections; F. Flower, oblique view; G. Pedicel with ovary, labellum, and column with dorsal sepal, longitudinal section; H. Upper portion of ovary, labellum, and column with anther cap, side view; I. Dorsal sepal; J. Lateral sepal; K. Petal; L. Column with anther cap (left), without anther cap (right), front view; M. Anther cap, dorsal view; N. Anther cap, flattened, ventral view; O. Pollinarium; P. Stipe and viscidium, side view. Single bar = 1 mm; double bar = 1 cm. Drawn by Susanna Stuart-Smith, A and C from Lamb in SAN 89677, B from Lamb in SAN 89678, D from Collenette 756 (left), Lamb in San 89678 (centre), and Lamb in SAN 89677 (right), E–P from Collenette 1, cult. R.B.G. Kew EN. 657–60. Reproduced with permission from J. J. Wood (1997), Orchids of Borneo. Volume 3. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

a thickened cushion, or free; pollinia four, appearing as two, each completely divided into free halves, stipe long, linear, viscidium subquadrate; stigma oval, vertical; rostellum distinct, porrect or stiffly erect, bilobed, lobes broad. ( JW)

Cytogenetics

Distribution (Fig. 639.2)

Phytochemistry

Cleisocentron includes six species collectively distributed in India (Sikkim), Bhutan, Burma, Vietnam, and Borneo. ( JW)

No reports have been found on the phytochemistry of Cleisocentron. (NV, RG)

Brandham (1999) reported a chromosome number of 2n = 38 for the genus. (AP)

159

VA N D E A E

Fig. 639.2.  Distribution map of Cleisocentron.

Phylogenetics

Uses

The three species of Cleisocentron that have been sequenced form an exclusive subclade in clade L, which is dominated by species of Cleisostoma. Topik et al. (2005) found a similar set of relatives (e.g. species of polyphyletic Cleisostoma, Sarcoglyphis, and Sterochilus). Further phylogenetic work on Cleisostoma and relatives is needed, but it may well be that Cleisocentron should be included in a larger generic concept. None of this can be sorted out until the type species of these genera are sequenced. (AK, MC)

No uses have been reported for Cleisocentron; it is uncommon in cultivation. ( JW)

Ecology Species of Cleisocentron are epiphytes, ranging in elevation from 330 to 3000 m. Cleisocentron abasii Cavestro occurs in forests on ultramafic substrate at 1200–1500 m in Borneo. Cleisocentron gokusingii J.J.Wood & A.L.Lamb occurs in mossy forests at 1800 m in Borneo. Cleisocentron kinabaluense D.Metusala & J.J.Wood occurs on mossy trees in shade in moss forest on ridges at 2400–3000 m in Borneo. Cleisocentron merrillianum (Ames) Christenson occurs in forests, sometimes on ultramafic substrates, at 1100–3000 m in Borneo. Cleisocentron pallens (Cathcart ex Lindl.) N.Pearce & P.J.Cribb occurs in forests at 330–2000 m in Sikkim. ( JW)

Pollination There is no information available on the pollination of the genus. It is worth noting that Cleisocentron merrillianum, C. kinabaluense, and C. gokusingii J.J.Wood and A.L.Lamb have blue flowers, an unusual colour in orchid flowers. ( JW) 160

Cultivation Group 2 (see subtribal treatment). Plants may be grown in containers with a well-draining mixture that should be allowed to dry out between waterings, requiring high humidity and moderate to high light levels. Most species prefer cool to intermediate conditions. (MM, AS)

Taxonomic notes Cleisocentron merrillianum from Borneo exhibits leaf polymorphy. The holotype (Clemens s.n., AMES) comprises flowering stems displaying two leaf types, the lower terete and acute, the upper broader, ligulate, and acute. The isotype at K comprises only two detached leaves, one ligulate, one terete, and some loose flowers. Ames seems to have overlooked or perhaps ignored these odd leaf types when drawing up his description of Sarcanthus merrillianus Ames, believing them to be abnormal. An illustration by Blanche Ames included in Ames and Schweinfurth’s 1920 account of the orchids of Mount Kinabalu shows a specimen with ligulate, acute leaves. Christenson (1992), however, commented that the young shoots produce leaves with ligulate blades similar to C. trichromum (Rchb.f.) Brühl (= C. pallens) but which grade to terete ones on older shoots. ( JW)

C LEISOSTOMA

Taxonomic literature

Phylogenetics

Ames, O. and Schweinfurth, C. (1920). The orchids of Mount Kinabalu, British North Borneo. Orchidaceae, fascicle 6. Merrymount Press, Boston. Christenson, E. A. (1992). An enigmatic blue. American Orchid Society Bulletin, 61, 242–7. Garay, L. A. (1972). On the systematics of the monopodial orchids I. Botanical Museum Leaflets, Harvard University, 23, 149–212. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

Cleisomeria is well supported as a member of a subclade in clade L, in which it is related to Pelatantheria, some species of Cleisostoma, and Rhynchogyna, a topology that was not supported by Topik et al. (2005). Cleisostoma is grossly polyphyletic, and a decision about how to handle Cleisomeria and related genera awaits resolution of the status of this problematic genus. (AK, MC)

Ecology

640.   CLE IS O ME R IA

Species occur on trees and rocks in deciduous and evergreen forests from 200 to 1100 m. (AS)

Cleisomeria Lindl. ex D.Don in Loud., Encycl., ed. 4, Suppl., 2, 1447 (1855). Type species: Cleisomeria lanatum (Lindl.) Lindl. ex G.Don (basionym: Cleisostoma lanatum Lindl.)

Pollination

Derivation of name From the Greek kleistos, closed, and meris, part, referring to the spur, the entrance of which is almost closed by a pair of calli. (AS)

Nothing is known about pollination in Cleisomeria. (AS)

Uses No uses have been reported for Cleisomeria; it is uncommon in cultivation. (AS)

Description (Plate 65; Fig. 640.1)

Cultivation

Epiphytic herbs. Roots terete, sparsely branching, glabrous. Stem well developed, patent to erect, rooting. Leaves conduplicate, articulate, distichous, linear, thick-coriaceous, apex unequally bilobed. Inflorescence lateral, solitary, usually branching, manyflowered; peduncle elongate, pendent; rachis continuing the line of the peduncle, straight, finely pubescent; floral bracts conspicuous (unusual in this subtribe), covering the flowers, greenish yellow, often marked with maroon, finely pubescent. Flowers resupinate on a pendent inflorescence, about 0.5 cm wide, opening in succession, several at a time over a long period, quaquaversal, greenish yellow with maroon and purple markings. Sepals entire, free, obtuse, finely pubescent abaxially. Petals shorter than sepals, glabrous. Labellum rigidly attached, spurred, trilobed, with a single callus at the mouth of the spur just below the midlobe; spur saccate, parallel with ovary, without callosities inside. Column without column foot; pollinia four in two slightly unequal pairs, stipe longer than the pollinia, proximally forked, with one branch under each set of pollinia, longer than twice diameter of pollinia. Ovary densely and finely pubescent. (AS, FR)

Group 2 (see subtribal treatment). (MM)

Distribution (Fig. 640.2) Cleisomeria is a genus of two species—C. lanatum and C. pilosulum (Gagnep.) Seidenf. & Garay—from Bangladesh, Burma, Thailand, Laos, Cambodia, Vietnam, Peninsular Malaysia, and Borneo. (AS)

Cytogenetics There are no published chromosome counts for the genus. (AP)

Phytochemistry No reports have been found on phytochemistry of Cleisomeria. (NV, RG)

Taxonomic literature Seidenfaden, G. (1988). Orchid genera in Thailand XIV. Fifty-nine vandoid genera. Opera Botanica, 95, 131–2. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

6 4 1 .   C L EI S OS T OM A Cleisostoma Blume, Bijdr., 362 (1825). Type species: Cleisostoma sagittatum Blume Echioglossum Blume, Bijdr., 364 (1825). Type species: Echioglossum javanicum Blume Sarcanthus Lindl., Collect. Bot., pl. 39, B (1826), nom. illeg. Type species: Sarcanthus rostratrus Lindl. Carteretia A.Rich., Sert. Astrol., 10. t. 4 (1834). Type species: Carteretia paniculata A.Rich. Garayanthus Szlach., Fragm. Flor. Geobot. Supp., 3, 136 (1995). Type species: Garayanthus duplicilobus (J.J.Sm.) Szlach. (Basionym: Sarcanthus duplicilobus J.J.Sm.). Raciborskanthos Szlach., Fragm. Flor. Geobot. Supp., 3, 135 (1995). Type species: Raciborskanthos capricornis (Ridl.) Szlach. (basionym: Ascochilus capricornis Ridl.) Blumeorchis Szlach., Ann. Bot. Fennici, 40, 68 (2003). Type species: Blumeorchis crochetii (Guillaumin) Szlach. (basionym: Sarcanthus crochetii Guillaumin) Ormerodia Szlach., Ann. Bot. Fennici, 40, 68 (2003). Type species: Ormerodia belophora (Rchb.f.) Szlach. (basionym: Sarcanthus belophorus Rchb.f.) 161

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Fig. 640.1.  Cleisomeria pilosulum (Gagnep.) Seidenf. & Garay. A. Habit; B. Leaf; C. Flower, front view; D. Flower, side view; E. Bract; F. Dorsal sepal; G. Lateral sepal; H. Petal; I. Labellum, from above; J. Labellum, longitudinal section; K. Column, labellum, and ovary, side view; L. Column, front view; M. Column, anther cap removed, front view; N. Column, side view; O. Anther cap and pollinarium, dorsal view; P. Anther cap, ventral view; Q. Pollinarium; R. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Kew Spirit Collection no. 48115.

Derivation of name From the Greek kleistos, closed, and stoma, mouth, referring to the mouth of the spur narrowed by callosities. ( JW)

Description (Plates 66, 67; Fig. 641.1) Epiphytic or rarely lithophytic or terrestrial herbs. Stems rigid, erect or pendent, rarely creeping, usually simple, many-noded. Leaves few to many, distichous, dorsiventrally flattened and conduplicate or terete, apex bilobed or acute, coriaceous, articulate to a tubular sheathing base. Inflorescence lateral, axillary or opposite 162

leaves, perforating the leaf-sheath, racemose or paniculate, erect, horizontal or pendent, few- to many-flowered; floral bracts much shorter than pedicel and ovary. Flowers fleshy, resupinate. Sepals free, usually spreading, slightly unequal. Petals free, usually spreading, similar to sepals or smaller and narrower. Labellum adnate to column at its base or posterior margins of side lobes, immobile, trilobed, spurred at base; side lobes ascending, subdeltoid; midlobe porrect or erect, usually deltoid or sagittate, often with a prominent callus or thickening near base; disc often with longitudinal lamellae or ridges; spur conical, cylindrical or rarely shortly saccate, often longitudinally septate inside, rarely

C LEISOSTOMA

Fig. 640.2.  Distribution map of Cleisomeria.

septum reduced to a narrow keel, usually with a prominent callus on back wall near mouth and often with outgrowths on front wall closing the entrance, the callus grooved, either alone or in conjunction with an opposing callus on the midlobe, completely obscuring opening to spur. Column with foot inconspicuous or absent; clinandrium shallowly excavate; anther cap terminal, operculate, frequently rostrate, pollinia four, arranged into two appressed, unequal masses, waxy, semi-globular, with a common stipe, stipe usually narrow, linear or broadened near apex, rarely broadly rectangular, viscidium single, subglobose to broad and horseshoe-shaped; stigma transverse, deeply recessed. ( JW)

Distribution (Fig. 641.2) Cleisostoma comprises about 100 species known from Sri Lanka, India, China (including Taiwan), and mainland Southeast Asia, the Ryukyu Islands, Malaysia, and Indonesia to the Philippines, New Guinea, and northern Australia east to Fiji. ( JW)

Infrageneric treatment Seidenfaden (1975) recognized seven sections in Cleisostoma as follows: Cleisostoma sect. Cleisostoma. Type species: Cleisostoma sagittatum Blume. Leaves dorsiventral, not terete, bilobed; stipe simple, usually linear tapering or clavate; viscidium usually simple and subglobose.

Cleisostoma sect. Complicatum Seidenf. Type species: Vanda teretifolia Lindl. (= Cleisostoma simondii (Gagnep.) Seidenf.). Leaves terete; stipe of a complicated shape; viscidium with two arms turned down and backwards like a saddle or horseshoe, the arms most often protected by a pair of callosities, ridges or flanges on front of column. Cleisostoma sect. Echioglossum (Blume) Seidenf. Type species: Echioglossum javanicum Blume (= Cleisostoma javanicum (Blume) Garay). Leaves dorsiventral, not terete; stipe and viscidium complicated in shape; labellum apex with a bifurcate appendage. Cleisostoma sect. Mitriformes Seidenf. Type species: Sarcanthus arietinus Rchb.f. (= Cleisostoma arietinum (Rchb.f.) Garay). Leaves terete; stipe mitre-shaped, less than 0.5 mm high, viscidium relatively large. Cleisostoma sect. Paniculatum Seidenf. Type species: Aerides paniculata Ker. Gawl. (= Cleisotoma paniculatum (Ker. Gawl.) Garay). Leaves dorsiventral, not terete; stipe and viscidium complicated in shape; labellum apex without a bifurcate appendage. Cleisostoma sect. Pilearia (Lindl.) Seidenf. Type species: Aerides appendiculata Lindl. (= Cleisostoma appendiculatum (Lindl.) Benth. & Hook.f. ex B.D. Jacks.). Leaves terete; stipe narrowly triangular in front view, usually narrowing from a broader base on account of the sides being 163

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Fig. 641.1.  Cleisostoma discolor Lindl. A. Plant; B. Flower; C. Callus at spur entrance; D. Anther; E. Pollinarium; F. Column and labellum, longitudinal section; G. Ovary, transverse section; H. Floral bract. Drawn by Shim Phyau Soon from Lamb AL 75–83. Reproduced with permission from Chan et al. (1994), Orchids of Borneo. Volume 1. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

recurved, the recurved sides most often broadening upwards, viscidium a simple disc. Cleisostoma sect. Subulatum Seidenf. Type species: Cleisostoma subulatum Blume. Leaves dorsiventral, not terete, acute, often caudate or with a mucronate apex; stipe simple, usually linear, tapering or clavate; viscidium usually simple and subglobose. ( JW)

Cytogenetics

Palynology

Phytochemistry

Schill and Pfeiffer (1977) examined Cleisostoma filiforme (Lindl.) Garay (as Sarcanthus filiformis Lindl.) and two unidentified species listed under Sarcanthus. Tetrads were convex with laevigate– microfoveolate sculpturing and a rounded sexine about 1 μm thick. (AP)

The alkaloid content of several species has been evaluated (Lüning 1967). Higher levels (0.01–0.1%) are found in C. discolor Lindl. (as Sarcanthus termissus Rchb.f.) and C. subulatum (as Sarcanthus subulatus (Blume) Rchb.f.) than in C. appendiculatum (Lindl.) Benth. & Hook.f. ex B.D.Jacks (as Sarcanthus appendiculatus

164

Brandham (1999) reported chromosome numbers of 2n = 26, 38, 40, 42, 72, with 38 as the most common. He saw no clear relationship among the numbers, perhaps reflecting the fact that as presently understood the genus is not monophyletic. Felix and Guerra (2010) listed the numbers 2n = 38 and n = 19, 20, 36. (AP)

C LEISOSTOMA

Fig. 641.2.  Distribution map of Cleisostoma.

(Lindl.) Hook.f.) and C. racemiferum (Lindl.) Garay (as Sarcanthus pallidus Lindl.), which contain them at 0.001–0.01%. No alkaloids were detected in C. striatum (Rchb.f.) N.E.Br. (as C. brevipes Hook.f.). (NV, RG)

Pollination

Phylogenetics

Uses

Cleisostoma is highly problematic and falls into at least four independent positions within clade L. It is probably the most difficult genus in Aeridinae. At present it is still unclear whether different subclades of Cleisostoma in the tree correspond to some of Seidenfaden’s (1975) sections or whether Cleisostoma and allied genera (e.g. Cleisomeria, Pelatantheria, Rhynchogyna, Sarcoglyphis, Sarcophyton, Smitinandia, and Stereochilus) should all be included in one large genus. However, new generic combinations must wait until the type species of Cleisostoma, C. sagittatum, and those of the allied genera are sequenced. Earlier studies that reported non-monophyly for Cleisostoma were Topik et al. (2005) and Carlsward et al. (2006b). (AK, MC)

Ecology Species of Cleisostoma are epiphytes or sometimes lithophytes, rarely terrestrial, from sea level to 3050 m. They are known from dipterocarp forests, mangrove and kerangas forests, broad-leaved evergreen forests, Terai forests (Nepal), teak plantations, swamp forests, and scrub. Cleisostoma gjellerupii (J.J.Sm.) Garay occurs as a terrestrial in wet peaty glades at 3000 m in New Guinea. ( JW)

There is no information available on pollination of Cleisostoma. ( JW)

No uses have been reported for Cleisostoma; it is occasionally cultivated. ( JW)

Cultivation Group 2 (see subtribal treatment). Species with softer and thinner leaves may need lower light levels. (MM, AS)

Taxonomic notes As currently delimited, Cleisostoma is one of the more diverse genera in Aeridinae, comprising species with terete and strapshaped leaves, long or short stems, branched or unbranched inflorescences. The flowers are always small, with a spurred, rigidly attached labellum, the spur entrance of which is almost blocked by two opposing callosities. The taxonomy of the genera closely allied to Cleisostoma, such as Pelatantheria, Rhynchogyna, Sarcoglyphis, Sarcophyton, Stereochilus, and others, all with similar flowers, is still unresolved. As their phylogenetic relationships become better understood, at least some of these genera may have to be included in Cleisostoma. (AS) 165

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Taxonomic literature

6 4 2 .  C L EI S OS T OM OPS I S

Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogeny of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. O’Byrne, P. (2009). Notes on Malesian Cleisostoma: Part 1. Malesian Orchid Journal, 4, 77–98. O’Byrne, P. (2010). Notes on Malesian Cleisostoma: Part 2. Malesian Orchid Journal, 5, 89–114. Seidenfaden, G. (1975). Orchid genera in Thailand 2. Cleisostoma Bl. Dansk Botanisk Arkiv, 29(3), 1–80. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

Cleisostomopsis Seidenf., Opera Bot., 114, 370 (1992). Type species: Cleisostomopsis eberhardtii (Finet) Seidenf. (basionym: Saccolabium eberhardtii Finet)

Derivation of name From the genus Cleisostoma Blume, and the Greek opsis, appearance, in reference to its superficial resemblance to Cleisostoma. ( JW)

Description (Plate 68; Fig. 642.1) Epiphytic herbs. Stems spreading to pendent, enclosed in leafsheaths, many-leaved. Leaves terete, fleshy, obtuse, 5–15 cm long,

B

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A Fig. 642.1.  Cleisostomopsis eberhardtii (Finet) Seidenf. A. Habit; B. Flower; C. Column and labellum, with labellum cut open to show back wall callus; D. Labellum, from above; E. Column, front view; F. Column, anther cap removed, front view; G. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Redrawn by Judi Stone after G. Seidenfaden, Opera Botanica, 114, Fig. 249 (1992).

166

C LEISOSTOMOPSIS

2 mm in diameter, articulate to a sheathing base. Inflorescence lateral, up to 10 cm long, piercing leaf-sheaths, racemose, many-flowered; floral bracts 2 mm long, triangular shorter than ovary. Flowers white, labellum midlobe lilac. Sepals free, 2.2 × 1.6 mm, lateral sepals slightly larger than dorsal sepal, ovate-elliptic, obtuse. Petals 2 × 1 mm, ovate, rounded. Labellum adnate to column base, spurred, trilobed, side lobes 0.3 mm long, broadly ovate, erect, rounded, midlobe 1 mm long, ovate, cymbiform, acute and incurved; spur 3.0–3.5 mm long, cylindrical-clavate, dilated toward apex, interior with a Y-shaped callus on back wall but without septum. Column without a foot; pollinia four in two pairs, waxy, semi-globular, each pair with a caudicle, attached by a common lanceolate stipe to an oblong viscidium about 0.5 × 0.4 mm; stigma concave. ( JW)

Phylogenetics

Distribution (Fig. 642.2)

Uses

Cleisostomopsis includes two species, C. eberhardtii from China (southwestern Guangxi) to Vietnam and C. elytrigera (Seidenf.) Szlach. from Thailand. ( JW)

No uses have been reported for Cleisostomopsis; it is not known to be in cultivation. ( JW)

Cytogenetics Felix and Guerra (2010) reported chromosome numbers for the genus as n = 19, 20. ( JW)

Phytochemistry No reports have been found on phytochemistry of Cleisostomopsis. (NV, RG)

There is no phylogenetic information available for this small genus. (AK)

Ecology Cleisostomopsis eberhardtii is epiphytic on tree trunks at 600 m in China. ( JW)

Pollination No information on pollination of Cleisostomopsis is available. ( JW)

Cultivation Group 4 (see subtribal treatment). (MM)

Taxonomic notes Garay (personal communication) pointed out to Seidenfaden (1992) that the relationship of Cleisostomopsis with Cleisocentron needs investigation. In his key to the Thai Aeridinae, Seidenfaden

Fig. 642.2.  Distribution map of Cleisostomopsis.

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placed Cleisocentron among the genera having a distinct column foot following Garay’s (1972) description, which was based on a King and Pantling plate (1898, t. 306) of Saccolabium trichromum Rchb.f. According to Seidenfaden, Garay pointed out that King and Pantling’s plate shows a distinct column foot, which is lacking in Reichenbach’s plate in Xenia Orchidacea 2: t. 139 (1865) which should therefore represent the true Saccolabium trichromum. Seidenfaden noted that ‘if this is incorrect because this plant is something else, Cleisocentron would in my key move up beside Cleisostomopsis, and the validity of my new genus be questionable. It is accordingly necessary to study the authentic material that may be made available and decide on the fate of the new genus at the same time as studying a series of plants found to belong in this complex.’ This genus is, therefore, only tentatively accepted until such detailed studies can be completed. ( JW)

Taxonomic literature Garay, L. A. (1972). On the systematics of the monopodial orchids I. Botanical Museum Leaflets, Harvard University, 23, 149–212. King, G. and Pantling, R. (1898). The orchids of the Sikkim-Himalaya. Annals of the Royal Botanical Garden of Calcutta 8, parts 1-4. Bengal Secretariat Press, Calcutta. Reichenbach, H. G. (1865). Xenia orchidacea. Brockhaus, Leipzig. Seidenfaden, G. (1992). The orchids of Indochina. Opera Botanica, 114, 1–502.

Cytogenetics Brandham (1999) and Felix and Guerra (2010) reported chromosome counts of 2n = 38, 40 for the genus. (AP)

Phytochemistry No reports have been found on phytochemistry of Cottonia. (NV, RG)

Phylogenetics Monospecific Cottonia is sister to Acampe in clade J, and the pair are sister to Trichoglottis s.l. Sinking of the two genera in Trichoglottis would increase the heterogeneity of the enlarged genus considerably. In particular, the flowers of Cottonia are different from all others in the subtribe. Similar arguments against unification of Cottonia and Acampe apply, but given the close relationship of these two genera it is conceivable that the former could be embedded in the latter when more species are included. (AK, MC)

Ecology Cottonia peduncularis occurs on trees by streams and on rocks in tropical dry, mixed evergreen forests and in tropical savannas at elevations of 350–800 m. (PC)

643.  C O TTO N IA

Pollination

Cottonia Wight, Icon. Pl. Ind. Orient., 5 (I), 21, t. 1755 (1852). Type: Cottonia macrostachya Wight (= C. peduncularis (Lindl.) Rchb.f.)

The Ophrys-like floral morphology would indicate pollination by sexual deception, but this has not been demonstrated as yet. (AS)

Derivation of name

Uses

The name commemorates Major-General Cotton, an English officer of the 19th century who collected plants in southwestern India. (PC)

No uses have been reported for Cottonia; it is rare in cultivation. (AS)

Description (Fig. 643.1) Erect, epiphytic herbs with leafy stems. Leaves alternate, coriaceous, elongate, narrow, unequally bilobed at tip, articulated to a sheathing base. Inflorescence two- or three-branched, with fewflowered racemes; bracts minute, ovate. Flowers fleshy. Sepals spreading, subequal, obovate. Petals spreading, subsimilar to sepals. Labellum sessile at base of column, porrect, much longer than sepals, fleshy, flat, spreading, bee-like, obscurely five-lobed, purple, villose, basal lobes auricular, side lobes shallowly semielliptic, villose on edges, midlobe subpandurate, retuse; calli two, erect, auricular at base of lip, fringed at tip. Column lacking a foot, winged toward tip, puberulous; anther two-celled, pollinia two, pear-shaped, cleft, stipe linear with recurved sides, viscidium minute; stigma U-shaped. Capsule fusiform-oblong. (PC)

Distribution (Fig. 643.2) Cottonia is monospecific and confined to Sri Lanka and India (southwestern Ghats, Deccan). (PC)

168

Cultivation Group 2 (see subtribal treatment). (MM)

Taxonomic notes Although closely related to the genus Acampe, Cottonia is easily distinguished by the tall, erect inflorescences, which are much longer than the leaves, and especially by the insect-like labellum. (AS)

6 4 4 .  C RYP T OPYL OS Cryptopylos Garay, Bot. Mus. Leafl., Harvard Univ., 23, 176 (1972). Type species: Cryptopylos clausus (J.J.Sm.) Garay (basionym: Sarcochilus clausus J.J.Sm.)

Derivation of name From the Greek kryptos, hidden, and pyle, a gateway, referring to the small, concealed spur-entrance. (AS)

C RYPTOPYLOS

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Fig. 643.1.  Cottonia peduncularis (Lindl.) Rchb.f. A. Habit; B. Flower, front view; C. Flower, side view; D. Bract; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum; I. Column and labellum, side view; J. Column, front view; K. Column, side view; L. Anther cap, dorsal view; M. Anther cap, ventral view; N. Pollinarium; O. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Jayasuriya 1299 (K); others from Kew Spirit Collection no. 71026.

Description (Plate 69; Fig. 644.1) Epiphytic herbs. Roots terete, not branching, glabrous, smooth. Stem patent, rooting at the base. Leaves conduplicate, articulate, distichous, oblong to linear, subfalcate, twisted at the base so as to lie in one plane with the stem, margins entire, apex unequally bilobed. Inflorescence lateral below the lowest leaf, pendent, racemose, many-flowered; peduncle elongate, rachis continuing the line of the peduncle, flexuose, only slightly thicker than the peduncle. Flowers non-resupinate, about 2 cm wide, probably short-lived, opening in succession a few at a time, quaquaversal, glabrous, ochre-yellow, suffused or spotted with brown, labellum white marked with violet. Sepals and petals entire, free, obtuse.

Labellum articulated with the apex of the column foot, spurred, trilobed; spur with a narrow opening, cylindrical, curved, continuing the line of the column foot. Column long and slender, column foot shorter than column; anther cucullate, pollinia two, porate, stipe longer than the pollinia, linear-subspatulate, viscidium small. Ovary and pedicel curved. (AS)

Distribution (Fig. 644.2) The only known species, Cryptopylos clausus, has been recorded from Thailand, Laos, Cambodia, Vietnam, and Indonesia (AS) 169

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Fig. 643.2.  Distribution map of Cottonia.

Cytogenetics

Cultivation

Chromosome numbers for C. clausus have not been reported. (AP)

Group 1 (see subtribal treatment). (MM)

Phytochemistry No reports have been found on phytochemistry of Cryptopylos. (NV, RG)

Phylogenetics Cryptopylos is a member of clade E and well supported as related to Dyakia and Macropodanthus. Topik et al. (2005) found a similar relationship. (AK, MC)

Ecology Cryptopylos clausus is epiphytic in evergreen forests from 300 to 1500 m. (AS)

Taxonomic notes Cryptopylos is morphologically similar to Macropodanthus and Pteroceras. It may be recognized by the narrow, partly concealed opening of the spur and the non-resupinate flowers, resembling a Gongora species. The generic delimitation of the genera allied to Pteroceras needs further study; Cryptopylos is only tentatively accepted here. (AS)

Taxonomic literature Seidenfaden, G. (1988). Orchid genera in Thailand XIV. Fifty-nine vandoid genera. Opera Botanica, 95, 131–2. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

Pollination Nothing is known about pollination of the species. (AS)

Uses No uses have been reported for Cryptopylos; it is rare in cultivation. (AS) 170

6 4 5 .  DEC EPT OR Deceptor Seidenf., Opera Bot., 114, 363 (1992). Type species: Deceptor bidoupensis (Tixier & Guillaumin) Seidenf. (basionym: Saccolabium bidoupense Tixier & Guillaumin)

DECEPTOR

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D G I F J Fig. 644.1.  Cryptopylos clausus (J.J.Sm.) Garay. A. Habit; B. Flower, side view, with labellum lowermost (flowers normally non-resupinate); C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum, from above; G. Labellum, longitudinal section; H. Column and labellum, side view; I. Column, anther cap removed, front view; J. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from a photograph by J.B. Comber and Kew Spirit Collection no. 29877.

Derivation of name From the Latin deceptor, deceiver, in reference to Frederick Sander’s subsequent disappointment with the small flowers of the plant, which are without commercial potential. ( JW)

Description (Fig. 645.1) Epiphytic herb. Stem elongate, leafy. Leaves ligulate, obliquely obtusely bilobulate, coriaceous, up to 25.0 × 3.1 cm; sheaths imbricate. Inflorescence paniculate (up to 1000 or more flowers), up to 25 cm or more long, branches to 15 cm long, subtended by acute bracts; floral bracts less than 1/8 length of pedicel. Flowers non-resupinate, minute, sepals and petals

greenish yellow spotted purple below, labellum white, column white with two purple, basal blotches. Sepals and petals free, acute. Sepals 2 mm long, petals slightly shorter. Labellum broadly attached to column foot and so not motile, obscurely trilobed, saccate, 2 mm long, vertical, with a transverse ridge at base; sac obtuse, globose, ecallose, 2 mm deep, with a ring of hairs around the entrance, with a long tuft of hairs spreading downwards on inside of transverse ridge. Column less than 1 mm high, widening on either side of rostellum, stelidia absent; pollinia four, in two unequal, semi-globular pairs, stipe ligulate, slightly widening below pollinia, length 1.5 times diameter of pollinia, viscidium ovate; rostellum ovate. ( JW) 171

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Fig. 644.2.  Distribution map of Cryptopylos.

Distribution (Fig. 645.2)

Cultivation

Deceptor is monospecific and endemic to Vietnam. ( JW)

Group 2 (see subtribal treatment). (MM)

Cytogenetics

Taxonomic notes

No chromosome number has been recorded for Deceptor. ( JW)

Seidenfaden (1992) believed Deceptor to be most probably closely allied to Smitinandia, from which it is distinguished by several characters including the vertical labellum, hairy entrance to the sac, and long-hairy tuft on the inside of the transverse ridge at the base of the midlobe. There are as yet no DNA sequences for the genus. ( JW)

Phytochemistry No reports have been found on phytochemistry of Deceptor. (NV, RG)

Phylogenetics There is no phylogenetic information available for this genus. (AK)

Ecology

Taxonomic literature Seidenfaden, G. (1992). The orchids of Indochina. Opera Botanica, 114, 1–502.

6 4 6 .  DI M OR P H ORC H I S

There is no information available on the pollination of Deceptor. ( JW)

Dimorphorchis Rolfe, Orchid. Rev., 27, 149 (1919), nom. cons. prop. Type species: Dimorphorchis lowii (Lindl.) Rolfe (basionym: Vanda lowii Lindl.) Lowianthus Becc., Nelle Foreste di Borneo, 531 (1902), nom. rej. prop. Type species: Lowianthus borneensis Becc., nom. illeg. (basionym: Vanda lowii Lindl.)

Uses

Derivation of name

No uses have been reported for Deceptor; it is not known to be in cultivation. ( JW)

From the Greek di-, two, morphe, shape, and orchis, orchid, referring to the flowers that are of two different shapes on

This species is epiphytic in forests at 2000 m. ( JW)

Pollination

172

DIMORPHORCHIS

B C

O

J

A

E

K G

L

I

N

F

M D

H

Fig. 645.1.  Deceptor bidoupensis (Tixier & Guillaumin) Seidenf. A. Habit; B. Flower, front view; C. Flower, side view; D. Bract; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum, from above; I. Labellum, longitudinal section; J. Column and labellum, side view; K. Column, front view; L. Anther cap, dorsal view; M. Anther cap, ventral view; N. Pollinarium; O. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Micholitz s.n. (1903) (K) and Micholitz s.n. (1905) (K).

the same inflorescence in most of the species in this genus. (AS)

Description (Plate 70; Fig. 646.1) Epiphytic herbs. Roots from basal part of the stem opposite the leaves. Stem spreading to pendent, sometimes erect, leafy. Leaves distichous, conduplicate, linear, apex unequally bilobed or sometimes acute. Inflorescence pendent, racemose, sometimes erect and paniculate, axillary, 1–4; peduncle and rachis glabrous or covered by stellate hairs, few- to many-flowered. Flowers dimorphic or not; basal flowers, when different from the others, 2–4, yellow, highly scented, unspotted or lightly spotted with red-purple dots; upper flowers (or all flowers in non-dimorphic taxa) blotched or heavily spotted

with red-purple or maroon, labellum white to pale yellow, marked with red-purple or maroon. Labellum fleshy, hinged with base of column, trilobed, side lobes erect, oblong to subquadrate, midlobe with a raised, laterally flattened callus and an erect callus at base of larger callus. Column papillose-pubescent or glabrous, with a column foot; anther papillose or not, pollinia four in two appressed pairs, transversely oblong to rounded-obdeltoid, stipe oblong to pandurate, about as long as the pollinia, viscidium elliptic. (PC, AS)

Distribution (Fig. 646.2) Dimorphorchis is a genus of seven species from Borneo, the Philippines, Sulawesi, New Guinea, and the Solomon Islands. It probably also occurs in the Moluccas. Dimorphorchis s.s., 173

VA N D E A E

Fig. 645.2.  Distribution map of Deceptor.

comprising taxa with dimorphic flowers, is endemic to Borneo, with five species and one variety. (PC, AS)

Cytogenetics Chromosome numbers for the genus have not been published. (AP)

In the former it is sister to Chiloschista in clade A, with this pair then sister to Thrixspermum. In the Bayesian tree, it is sister to Thrixspermum, with Chiloschista distantly placed. A few former species of Arachnis has been added to Dimorphorchis. A close relationship between Dimorphorchis and Thrixspermum was found by Topik et al. (2005). (AK, MC)

Phytochemistry

Ecology

Several species of Dimorphorchis have been investigated for their alkaloid content, but under former generic names (Lüning 1964; Lawler and Slaytor 1970). Those giving positive results were D. breviscapa ( J.J.Sm.) Kocyan & Schuit. (studied as Vandopsis longicaulis Schltr.), whereas D. beccarii (Rchb.f.) Kocyan & Schuit. (studied as Vandopsis muelleri (Kränzl.) Schltr.) did not contain them. Large-scale extraction of fresh plant material of D. breviscapa afforded the alkaloids N-methylpiperidine N-oxide and N-methylpyridinium chloride, which were obtained in crystalline form as their hydrobromide and iodide salts, respectively (Brandänge and Lüning 1970). (NV, RG)

Pollination

Phylogenetics Circumscription of Dimorphorchis has been highly variable depending on the author, but in the analyses presented here it is inconsistently placed in the parsimony and Bayesian trees.

174

Species are epiphytes in forests from 0 to 1300 m. Dimorphorchis lowii and D. rossii Fowlie are mainly found in riverine forests overhanging the water, whereas D. beccarii usually grows as a lithophyte (or semi-terrestrial) on exposed rocky outcrops. (AS)

Nothing is known about pollination in Dimorphorchis. As in Grammatophyllum (Cymbidieae), the lowermost flowers in the inflorescence of several species are markedly different from the others. This phenomenon is as yet unexplained but probably plays a role in the pollination of these spectacular orchids. (AS)

Uses No uses have been reported for Dimorphorchis; it is occasionally cultivated. (AS)

DIPLOCENTRUM

A F

1 cm

1 cm

J

1 cm

G

I

H

B 1 cm

E 4 cm

C

6 mm

D

K

4 mm

L

Fig. 646.1.  Dimorphorchis graciliscapa (A.L.Lamb & Shim) P.J.Cribb. A. Plant; B. Flower, lower; C. Flower, upper; D. Labellum; E. Column, pedicel, and labellum, side view; F. Column and labellum, longitudinal section; G. Lateral sepal; H. Dorsal sepal; I. Petal; J. Column, ventral view; K. Pollinarium; L. Anther. Drawn as D. rossii Fowlie var. graciliscapa A.L.Lamb & Shim by Chan Chew Lun from cult. Hepburn in Lamb AL 598/86. Reproduced with permission from Chan et al. (1994), Orchids of Borneo. Volume 1. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

Cultivation

Taxonomic literature

Group 2 (see subtribal treatment). Plants of Dimorphorchis may be grown in large containers or baskets with a well-draining medium. These plants require high humidity, regular watering throughout the year, and moderate light levels, except for D. beccarii, which requires a more exposed position. (MM, AS)

Cribb, P. and Bell, A. (2008). The Bornean endemic genus Dimorphorchis (Vandeae: Aeridinae). Malesian Orchid Journal, 2, 77–92. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

Taxonomic notes The earliest valid name for this genus is Lowianthus, but it has never been adopted. A proposal is in preparation to conserve Dimorphorchis over Lowianthus (Schuiteman and Cribb, unpublished).

6 4 7 .   DI P L OC ENT RU M Diplocentrum Lindl., Edwards’s Bot. Reg., t. 1522 (1832). Type species: Diplocentrum recurvum Lindl.

175

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Fig. 646.2.  Distribution map of Dimorphorchis.

Derivation of name

Cytogenetics

From the Greek diploos, double, and kentron, spur, referring to the two spurs on the labellum. (AS)

Brandham (1999) and Felix and Guerra (2010) reported chromosome counts of 2n = 38 for the genus. (AP)

Description (Plate 71; Fig. 647.1)

Phytochemistry

Epiphytic herbs. Roots terete, unbranched, glabrous, smooth. Stem patent to erect, rooting at the base. Leaves conduplicate, articulate, distichous, linear to subterete, coriaceous; apex bilobed. Inflorescence lateral, racemose or paniculate, erect or pendent, many-flowered; rachis continuing the line of the peduncle, straight, glabrous. Flowers resupinate, about 0.5 cm wide, many opening simultaneously, quaquaversal, greenish or pinkish with pink to purple labellum. Sepals entire, free, obtuse. Petals shorter than sepals. Labellum rigidly attached, with two short spurs at base, entire, ligulate, longer than sepals; spurs conical, at right angles to ovary, without callosities inside. Column without column foot, with a transverse papillose lamella at base; pollinia four, compressed in two unequal pairs, stipe obspatulate, longer than pollinia, viscidium large; rostellar projection not elongate. Capsule obovoid. (AS)

No reports have been found on phytochemistry of Diplocentrum. (NV, RG)

Distribution (Fig. 647.2) Diplocentrum has two species—D. recurvum and D. congestum Wight—from southern India and possibly Sri Lanka, the latter known only from a single 19th-century record. (AS) 176

Phylogenetics Diplocentrum is sister to Diploprora and the pair sister to Sarcanthopsis. They are part of grade K. (AK, MC)

Ecology Both species are epiphytes in deciduous forests from 1200 to 1600 m. (AS)

Pollination Nothing is known about pollination in Diplocentrum. (AS)

Uses No uses have been reported for Diplocentrum; it is rare in cultivation. (AS)

DIPLOPRORA

C

I

B

F

D

J K

E

M N

G

A H P

O L

Fig. 647.1.  Diplocentrum congestum Wight. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum, from above; H. Labellum, from below; I. Column and labellum, side view; J. Column and labellum, front view; K. Column, side view; L. Column, anther cap removed, front view; M. Anther cap, dorsal view; N. Anther cap with pollinarium, ventral view; O. Pollinarium; P. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone after drawing 2687 in Kew Orchid Drawing Collection and from Kew Spirit Collection no. 12048.

Cultivation

Derivation of name

Group 2 (see subtribal treatment). Plants should be grown mounted on cork or similar substrate under intermediate conditions, with a cool and dry resting period in winter. (MM, AS)

From the Greek diploos, double, and prora, prow, in reference to the spreading bifurcation of the labellum apex. ( JW)

648.   DIPLO P RO R A

Epiphytic herbs. Stems pendent, terete or slightly flattened, sometimes branched, with many nodes. Leaves distichous, blade narrowly ovate to falcate-lanceolate, articulate to an amplexicaul sheath. Inflorescence lateral, racemose, few-flowered. Flowers appearing a few at a time, widely opening. Sepals and

Diploprora Hook.f., Fl. Brit. India, 6, 26 (1890). Type species: Diploprora championii (Lindl. ex Benth.) Hook.f. (basionym: Cottonia championii Lindl. ex Benth.)

Description (Plate 72; Fig. 648.1)

177

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Fig. 647.2.  Distribution map of Diplocentrum.

petals similar, free, spreading, abaxial surface of sepals carinate. Petals narrower than sepals. Labellum superior, spurless, broadly attached to column base, hypochile boat-shaped, with raised side lobes, epichile narrowed, apex forked into two tails, or truncate. Column lacking a foot; pollinia four, unequal, globose, stipe narrower at apex than at base, viscidium ovate. ( JW)

Distribution (Fig. 648.2) Two species—D. championii and D. truncata Rolfe ex Downie— are collectively distributed in Sri Lanka, Andaman Islands, India, Burma, Thailand, Vietnam, China, Taiwan, and the Ryukyu Islands. ( JW)

Cytogenetics Chen Xinqi and Wood (2009) recorded chromosome numbers of 2n = 16, 38 for D. championii. ( JW)

Phytochemistry Although a survey of alkaloid content in Orchidaceae revealed that one species of Diploprora did not contain these compounds at levels ≥ 0.1% (Lüning 1974), nothing further is known about the phytochemistry of this genus. (NV, RG)

was postulated by Topik et al. (2005), but with the greater sampling included here, that relationship appears to be unfounded. (AK, MC)

Ecology Both species are epiphytes from 200 to 1700 m. Diploprora championii is epiphytic on tree trunks or lithophytic in valleys between 200 and 1500 m in China. It is reported as an epiphyte on branches overhanging watercourses and in wet evergreen forests up to 700 m in Sri Lanka. Diploprora truncata is recorded as an epiphyte in evergreen forests at 1650 m in Thailand. ( JW)

Pollination No information on the pollination of Diploprora is available. ( JW)

Uses No uses have been reported for Diploprora; it is uncommon in cultivation. ( JW)

Cultivation Group 2 (see subtribal treatment). (MM)

Phylogenetics

Taxonomic literature

The two species of Diploprora fall together and then are successively sister to Diplocentrum and Sarcanthopsis. A relation to Malleola and Omoea

Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK

178

DIPLOPRORA

B

A

E

G

K F

M

H N L

J

D C I

Fig. 648.1.  Diploprora championii (Lindl. ex Bentham) Hook.f. A. Habit; B. Inflorescence; C. Floral bract, ovary, labellum, and column, side view; D. Ovary, labellum, and column, side view; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum, front view; I. Labellum, side view; J. Labellum, unfolded; K. Column apex, anther cap removed, front view; L. Anther cap, dorsal and ventral views; M. Pollinia; N. Stipe and viscidium, two views. Single bar = 1 mm; double bar = 1 cm. Drawn by Susanna Stuart-Smith from Hooker s.n. (K). Reproduced with permission from N. R. Pearce and P. J. Cribb (2002), The Orchids of Bhutan. Royal Botanic Garden, Edinburgh and Royal Government of Bhutan.

179

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Fig. 648.2.  Distribution map of Diploprora.

and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

649.  D RYA D O RC H IS Dryadorchis Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 976 (1913). Type species: Dryadorchis barbellata Schltr.

foot, not spurred, trilobed, often with a callus between lateral lobes, midlobe spur-like but not hollow. Column semi-terete, ciliate or pubescent, sometimes with glandular hairs, column foot hardly discernible, shorter than column; anther helmet-shaped, pollinia four, hemispherical, in two subglobose masses, stipe longer than pollinia, oblong; rostellum bidentate after removal of pollinarium. Capsule oblong, angular and slightly curved. (AS)

Derivation of name From Dryas, a tree-nymph in Greek mythology, and orchis, orchid, probably referring to the pretty flowers of these epiphytic orchids. (AS)

Description (Plate 73; Fig. 649.1) Epiphytic herbs. Roots terete, not or sparsely branching, glabrous, smooth. Stem rooting at the base, patent. Leaves 3–5, conduplicate, articulate, distichous, subfalcate, oblong, twisted at base so as to lie in one plane with stem, surface appearing velvety-opaque, margins finely crenulate, apex asymmetric. Inflorescence lateral, solitary, racemose, 5–12-flowered; peduncle elongated, patent, thinner than the swollen, angular, geotropic rachis. Flowers resupinate on pendent rachis, 1–3 cm wide, ephemeral, opening in succession or in successive groups, 1 to 12 at a time, quaquaversal, colourful, sometimes fragrant. Sepals and petals entire, free, obovate, concave, rounded to obtuse. Labellum hinged with apex of column 180

Distribution (Fig. 649.2) Dryadorchis comprises five species endemic to New Guinea. (AS)

Cytogenetics Chromosome numbers for Dryadorchis have not been published. (AP)

Phytochemistry No reports have been found on phytochemistry of Dryadorchis. (NV, RG)

Phylogenetics Dryadorchis forms a clade with one species of Gunnarella (another falls in clade E) and Rhinerrhizopsis in the New Guinea clade. (AK, MC)

DRYADORCHIS

K

J

B C M

L N

I

O

D

F

A H

E

G Fig. 649.1.  Dryadorchis huliorum (Schuit.) Christenson & Schuit. A. Leaves and inflorescence; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum, from above; H. Labellum, longitudinal section; I. Column and labellum, side view; J. Column, front view; K. Column apex, anther cap removed; L. Column, side view; M. Anther cap, dorsal view; N. Anther cap, ventral view; O. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Reeve 624 (K) and Kew Spirit Collection no. 59937.

Ecology

Cultivation

All species of Dryadorchis are epiphytes in coastal and other forests from 0 to 1700 m. (AS)

Nothing is known about pollination in Dryadorchis. (AS)

Group 1 (see subtribal treatment). Species are best cultivated on a slab of bark covered with moss and humus and kept in light shade under conditions of high humidity. The plants should not be allowed to dry out for any considerable length of time. Depending on the species, a warm or intermediate-cool greenhouse is required. (MM, AS)

Uses

Taxonomic literature

No uses have been reported for Dryadorchis; it is rare in cultivation. (AS)

Schuiteman, A. and Christenson, E. A. (1995). A synopsis of Dryadorchis (Orchidaceae). Blumea, 40, 421–4.

Pollination

181

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Fig. 649.2.  Distribution map of Dryadorchis.

Schuiteman, A. and de Vogel, E. F. (2004). A new species of Dryadorchis from Papua New Guinea. Journal für den Orchideenfreund, 11, 222–30.

Distribution (Fig. 650.2)

650.  D RY MOA N TH US

Cytogenetics

Drymoanthus Nicholls, Victorian Naturalist, 59, 173 (1943). Type species: Drymoanthus minutus Nicholls

Felix and Guerra (2010) reported chromosome numbers of 2n = 38, 76 for the genus. (AP)

Derivation of name

Phytochemistry

From the Greek drymos, forest, and anthos, flower, referring to the habitat. ( JW)

Description (Plate 74; Fig. 650.1) Epiphytic or lithophytic herbs. Stem up to 4 cm long. Leaves distichous, oblong or elliptic, sometimes falcate, acute. Inflorescence racemose, few-flowered. Flowers greenish, greenish white or yellow, often marked with red or purple, lasting at least two days. Sepals and petals subequal, not opening widely. Labellum attached broadly at an acute angle to base of column, cymbiform, or deeply concave, fleshy and thick-walled, entire, ecalcarate, ecallose. Column with tooth-like apical appendages, foot absent; anther cap with a porrect beak, pollinia four, in two closely appressed unequal pairs, stipe at least as long as a pollinium, narrow; rostellum porrect. ( JW) 182

The four species of Drymoanthus are collectively distributed in Australia, New Caledonia, and New Zealand. ( JW)

No reports have been found on phytochemistry of Drymoanthus. (NV, RG)

Phylogenetics Drymoanthus is strongly supported as sister to monospecific Mobilabium in the Australia clade. (AK, MC)

Ecology All species are epiphytes at 0 to 800 m. Drymoanthus adversus (Hook.f.) Dockrill is epiphytic on tree trunks and on shaded rocks in moist gullies in New Zealand. Drymonanthus minimus (Schltr.) Garay is epiphytic on trees in open forest and scrub at 0–800 m in New Caledonia. Drymoanthus minutus occurs in northern Queensland, Australia, as an epiphyte on trees and shrubs in rain forest, especially overhanging watercourses, at 100–880 m. (JW, PA)

DYAKIA

C

B

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F

H E G

I J A Fig. 650.1.  Drymoanthus adversus (Hook.f.) Dockrill. A. Habit; B. Inflorescence; C. Flower, front view; D. Flower, side view; E. Bract; F. Dorsal sepal; G. Lateral sepal; H. Petal; I. Labellum, from above; J. Labellum, longitudinal section; K. Column and labellum, side view; L. Column, front view; M. Column, side view; N. Anther cap with pollinia; O. Anther cap, ventral view; P. Pollinarium; Q. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Kew Spirit Collection no. 47476.

Pollination

6 5 1 .   DYA K I A

There is no information available on pollination of Drymoanthus. ( JW)

Dyakia Christenson, Orchid Dig., 50, 63 (1986). Type species: Dyakia hendersoniana (Rchb.f.) Christenson (basionym: Saccolabium hendersonianum Rchb.f.)

Uses No uses have been reported for Drymoanthus; it is rare in cultivation. ( JW)

Derivation of name From the Malay Dyak, a generic name for the aboriginal peoples of Borneo. ( JW)

Cultivation

Description (Plate 75; Fig. 651.1)

Group 1(see subtribal treatment). Plants may be cultivated as for Sarcochilus twig epiphytes. (MM, PA)

Epiphytic herbs. Stem leafy. Leaves conduplicate, oblong-ligulate, apex obtusely bilobed. Inflorescence simple, racemose, erect, 183

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Fig. 650.2.  Distribution map of Drymoanthus.

many-flowered; rachis with pedicels emerging from the base of crateriform depressions. Flowers resupinate, c. 1 cm across, bright pinkish red with a white labellum. Sepals and petals spreading. Lateral sepals cuneate-oblong, obtusely angled on outer side, lowermost portion connate with labellum at base. Labellum obscurely trilobed, solidly attached to column, side lobes auriculate, midlobe flat, triangular, appearing in the flat, uppermost front of spur; spur laterally compressed, oblongligulate, obtuse, anterior third solid, remainder hollow, with a pair of crimson pollinator guides at each side of entrance; back wall callus singular, prominent. Column foot absent, rostellar projection elongate, attenuate, sigmoid, longer than remainder of column; pollinia two, porate, stipe with two lateral apical appendages. ( JW)

Distribution (Fig. 651.2) Dyakia is monospecific and endemic to Borneo. ( JW)

Cytogenetics No chromosome numbers have been recorded for D. hendersoniana. ( JW)

Phytochemistry No reports have been found on the phytochemistry of Dyakia. (NV, RG) 184

Phylogenetics Although Dyakia has floral similarities to Vanda (especially the former species of Ascocentrum now included in Vanda), this monospecific genus is a member of clade E, which is only distantly related to Vanda. It is well supported as related to Cryptopylos and Macropodanthus. Topik et al. (2005) and Carlsward et al. (2006b) found a similar position for Dyakia. (AK, MC)

Ecology This species is epiphytic in lowland and hill forests. ( JW)

Pollination There is no information available on the pollination of Dyakia. ( JW)

Uses No uses have been reported for Dyakia; it is frequently cultivated. ( JW)

Cultivation Group 1 (see subtribal treatment). Dyakia hendersoniana is easily grown in an intermediate to warm greenhouse with a minimum night temperature of 16–18° C (60–65° F). Plants will thrive if

DYAKIA

B

C

Q L

K

M

N

O

P

I

E J

D

G F A

H R Fig. 651.1.  Dyakia hendersoniana (Rchb.f.) Christenson. A. Habit; B. Flower, front view; C. Flower, back view; D. Dorsal sepal, front view; E. Dorsal sepal, back view; F. Lateral sepal; G. Petal; H. Labellum; I. Labellum, longitudinal section; J. Column and labellum, side view; K. Column, front view; L. Column, anther cap removed, front view; M. Column, side view; N. Anther cap, dorsal view; O. Anther cap, ventral view; P. Pollinarium; Q. Viscidium, side view; R. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Lamb 270/86 (K) and Kew Spirit Collection no. 33851.

given abundant water, provided the foliage and roots dry quickly. An open, well-drained medium is essential. Christenson (1986) reported success using medium-grade tree bark in small clay pots suspended beneath an upper tier of plants with good air circulation and bright, diffused light. ( JW, MM)

Taxonomic notes Dyakia may be distinguished from species of Vanda formerly included in Ascocentrum by the flat, obscurely bilobed leaves, rudimentary labellar side lobes, pair of pollinator guides at the base of a bilaterally compressed spur, prominent back wall callus, appendaged stipe, and elongate, attenuate, sigmoid rostellum.

From Robiquetia it can be distinguished by the pollinator guides, bilaterally compressed spur, appendaged stipe, and erect inflorescence. It shares the long rostellum of Micropera but differs by its symmetrical, resupinate flowers and appendaged stipe. The recessed depressions on the fleshy rachis are similar to those found in Trachoma and Tuberolabium, genera which according to DNA sequence data are closely related (see Phylogenetics under subtribal heading). ( JW)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogeny of Vandeae (Orchidaceae) and

185

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Fig. 651.2.  Distribution map of Dyakia.

the evolution of leaflessness. American Journal of Botany, 93, 770–86. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

652.  E C LE C TIC US Eclecticus P.O’Byrne, Orchideenj., 3, 127 (2009). Type species: Eclecticus chungii P.O’Byrne

Derivation of name From the Greek eklektikos, choosing or eclectic, referring to features of different vandaceous genera found in Eclecticus. ( JW)

Description (Fig. 652.1) Epiphytic herbs. Roots slightly flattened. Stems inconspicuous. Leaves crowded, 2.5–7.0 × 0.9–1.9 cm, oblong to elliptic, unequally obtusely bilobed, often heavily suffused purple, especially on reverse and base. Inflorescences borne among roots, suberect to descending; peduncle not compressed; rachis 7–20-flowered, flowers opening in succession. Flowers resupinate, 7–8 mm long, 9–10 mm wide, sepals and petals greenish white to pale brownish cream, labellum white, side lobes with red margins and several broad, longitudinal, red stripes, spur externally gold, internally 186

yellow, column white with a red V below stigma. Sepals free, dorsal sepal erect with a concave, decurved apex, lateral sepals horizontal, the lower margin at base recurved and resting against spur. Petals with erose distal margins. Labellum trilobed, solidly attached to column foot, spurred, side lobes obliquely erect, quadrate, recurved at front, midlobe porrect, ovate, margins erose, basal third occupied by a papillose transverse thickening, a low median ridge in proximal half, hypochile with two parallel, spiculate, ridge-like keels that extend backward from midlobe onto floor of spur where a third, teardrop-shaped, spiculate keel lies between them; spur inverted subglobose, distinctly compressed dorsally at base. Column porrect, foot a downward-projecting plate that lies across spur aperture and partly closes it, isosceles-trapezoid; stelidia absent; anther cap cucullate, with a drawn-out linear base; pollinia two, each deeply cleft, stipe 2.5 mm long, linear, distal quarter expanded, obovate, concave, viscidium canaliculate; stigmatic cavity large; rostellum projection 2 mm long, linear, extending backward across stigmatic cavity. ( JW)

Distribution (Fig. 652.2) Eclecticus is monospecific and found in Thailand, Laos, and Cambodia. ( JW)

Cytogenetics No chromosome counts for Eclecticus have been recorded. ( JW)

ECLECTICUS

B

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J

K

G

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C E

H

D

A F

Fig. 652.1.  Eclecticus chungii P.O’Byrne. A. Habit; B. Flower, side view; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum; G. Column and labellum, side view; H. Column and labellum, longitudinal section; I. Column, front view; J. Columnn apex with pollinia; K. Anther cap, side view; L. Pollinarium, side view; M. Anther cap, dorsal view; N. Pollinia, O. Pollinarium dorsal view. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone after P. O’Byrne, Orchideen J., 16, 129 (2009).

Phytochemistry

Pollination

No reports have been found on phytochemistry of Eclecticus. (NV, RG)

No information on pollination of Eclecticus is available. ( JW)

Phylogenetics

Uses

Eclecticus is member of grade K, in which it falls in an isolated position, not sister to any specific genus but rather to clade L. (AK)

No uses have been reported for Eclecticus; a few plants are in cultivation. ( JW)

Ecology

Cultivation

Nothing is known of the habitat of this species. ( JW)

Group 1 (see subtribal treatment). (MM) 187

VA N D E A E

Fig. 652.2.  Distribution map of Eclecticus.

Taxonomic notes Eclecticus looks superficially like a species of Kingidium, now included in Phalaenopsis. Closer examination reveals characters that indicate an affinity with disparate genera in Aeridinae. The thickened rachis is found in a number of genera including Chroniochilus and Gastrochilus, but those have different pollinia. The inflorescence and pollinia of Eclecticus resemble those found in Thrixspermum sect. Dendrocolla, but that section has ephemeral flowers with a different labellum structure. The flower shape might show an affinity with Lesliea (now included in Phalaenopsis) or Smitinandia, but both of those have a different arrangement of calli on the labellum. The elongated rostellum projection resting across the stigmatic cavity resembles Rhynchogyna, which is allied to Cleisostoma and has a large callus on the back wall of the spur. ( JW)

653.  G AS TRO C H ILUS Gastrochilus D.Don, Prodr. Fl. Nepal., 32 (1825). Type species: Gastrochilus calceolaris (Buch.-Ham. ex Sm.) D.Don (basionym: Aerides calceolaris Buch.-Ham. ex Sm.) Haraella Kudô, J. Soc. Trop. Agric., 2, 26 (1930), syn. nov. Type species: Haraella odorata Kudô Luisiopsis C.S.Kumar & P.C.S.Kumar, Rheedea, 15, 46 (2005). Type species: Luisiopsis inconspicua (Hook.f.) C.S.Kumar 188

& P.C.S.Kumar (basionym: Saccolabium inconspicuum Hook.f.)

Derivation of name From the Greek gaster, belly, and cheilos, lip, an allusion to the swollen labellum. ( JW)

Description (Plates 76, 77; Fig. 653.1, 653.2) Epiphytic herbs. Stem ascending or pendent, enclosed in basal sheaths of leaves. Leaves usually distichously arranged, narrowly elliptic or strap-shaped, margin sometimes undulate, apex unequally bilobed, acute or rarely with three setae, slightly coriaceous, flat, articulate to an amplexicaul-sheathing base, sometimes twisted at base and lying in one plane. Inflorescence lateral, racemose, often dense and appearing subumbellate, few to many-flowered. Flowers resupinate or with the labellum pointing toward the axis. Sepals and petals free, similar, spreading. Labellum firmly attached to base of column, divided into a semi-globose to saccate or conical hypochile with side lobes reduced to edges of the sac, and a flabellate, often broadly triangular, often hairy or papillose epichile, margin entire to fimbriate. Column foot absent; anther operculate, semi-globose, apex narrower than base, pollinia two, waxy, subglobular, porate or rarely cleft, or rarely four, solid, attached by a common, longer-than-wide stipe to a bilobed viscidium; rostellum bilobed. ( JW)

GASTROCHILUS

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Fig. 653.1.  Gastrochilus patinatus (Ridl.) Schltr. A. Habit; B. Flower, front view; C. Dorsal sepal; D. Petal; E. Lateral sepal; F. Flower with sepals and petals removed, side view; G. Labellum and column, longitudinal section; H. Anther cap, dorsal view; I. Anther cap, side view; J. Anther cap, ventral view showing pollinarium; K. Pollinarium; L. Pollinia. M. Ovary, transverse section. Drawn by Chan Chew Lun from a plant cultivated at Tenom Orchid Centre, Sabah. Reproduced with permission from Chan et al. (1994), Orchids of Borneo. Volume 1. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

Distribution (Fig. 653.3)

Phytochemistry

Gastrochilus includes about 48 species collectively distributed from China, India, and Sri Lanka through eastern Asia and Japan south to Indonesia and the Philippines. There are 29 species in China. ( JW)

One of two species of Gastrochilus examined by Lüning (1974) contained alkaloids at levels ≥ 0.1%, although the species names were not recorded. An earlier survey by the same author (Lüning 1967) noted that alkaloids were not detected in G. inconspicuus (Hook.f.) Kuntze (as Luisia inconspicua (Hook.f.) King & Pantl.). Additional records are those published under the former generic name Saccolabium (Lüning 1964, 1967), comprising positive results for G. dasypogon (Sm.) Kuntze (as S. dasypogon (Sm.) Lindl.) and G. distichus (Lindl.) Kuntze (as S. distichum Lindl.), and a negative result for G. acutifolius (Lindl.) Kuntze (as S. acutifolium Lindl.). According to Kaiser (2011), the floral fragrance of G. japonicus (Makino) Schltr. has a spicy, clove-like character due to its content of eugenol (27.0%), which is the major component, and (E )-isoeugenol (3.5%). Other contributions to the scent

Palynology Schill and Pfeiffer (1977) described pollen tetrads of G. fusco­ punctatus (Hayata) Hayata (= G. pseudodistichus (King & Pantl.) Schltr.) as even (neither concave nor convex) and partly sunken with a laevigate surface. (AP)

Cytogenetics Brandham (1999) and Felix and Guerra (2010) reported chromosome numbers of 2n = 30, 34, 38, 40; the latter also reported 2n = 76. ( JW)

189

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O A Fig. 653.2.  Gastrochilus retrocallus (Hayata) Hayata [formerly Haraella retrocalla (Hayata) Kudô]. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum, from above; H. Labellum, longitudinal section; I. Column and labellum, side view; J. Column, front view; K. Column apex, anther cap removed, front view; L. Column, side view; M. Anther cap, dorsal view; N. Anther cap, ventral view; O. Pollinarium; P. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from a photograph by L.M. Mason and Kew Spirit Collection no. 48523.

are made by a combination of decanal (7.5%), (Z)-dec-4-enal (0.4%), and (Z)-hex-3-en-1-ol (5.2%). ( NV, RG)

Phylogenetics Gastrochilus falls in clade H, in which it is well supported as sister to Pomatocalpa (Topik et al. 2005). The former monotypic Haraella is now considered a member of Gastrochilus (Carlsward et al. 2006b). The general view of Seidenfaden (1988) that the genus is not well understood is still true today, and a systematic study will help clarify relationships within the genus and delimitation of species. (AK) 190

Ecology Species are epiphytic in forests (lithophytic in G. gongshanensis Z.H.Tsi), ranging from 0 to 3000 m but less frequent in the lowlands. ( JW)

Pollination There is no information available on the pollination of Gastrochilus. Gastrochilus retrocallus (Hayata) Hayata (often cultivated as Haraella odorata) from Taiwan has an insect-like labellum, possibly indicative of pollination through sexual deception. Most other species have flowers resembling those

GROSOURDYA

Fig. 653.3.  Distribution map of Gastrochilus.

of certain Epipactis species and may be pollinated by wasps. (AS)

Uses No uses have been reported for Gastrochilus; several species are cultivated. ( JW)

Kumar, C. S. and Kumar P. C. S. (2005). An orchid digest of Manipur, northeastern India. Rheedea, 15, 1–74. Seidenfaden, G., (1988). Orchid genera in Thailand XIV. Fifty-nine vandoid genera. Opera Botanica, 95, 1–398. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

Cultivation Group 1 (see subtribal treatment). (MM)

Taxonomic notes The generic position of G. inconspicuus is unclear. This species combines the habit of Luisia with the floral morphology of Gastrochilus. Given that vegetative characters in orchids appear less prone to convergent evolution than floral characters, it could be supposed that it is in fact a species of Luisia with Gastrochiluslike flowers (L. inconspicua). Kumar and Kumar (2005) placed it in the monospecific genus Luisiopsis. (AS)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogeny of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86.

6 5 4 .   GROS OU R DYA Grosourdya Rchb.f., Bot. Zeitung (Berlin), 22, 297 (1864). Type species: Grosourdya elegans Rchb.f. Ascochilopsis Carr, Gard. Bull. Straits Settlem., 5, 21 (1929), syn. nov. Type species: Ascochilopsis myosurus (Ridl.) Carr (basionym: Saccolabium myosurus Ridl.) Ascochilus Ridl., J. Linn. Soc., Bot., 32, 374 (1896), syn. nov. Type species: Ascochilus siamensis Ridl. Theana Aver., Taiwania, 57, 136 (2012). Type species: Theana vietnamica Aver.

Derivation of name The generic name commemorates René de Grosourdy (floruit 1836-1864), a plant hunter specializing in medicinal plants of the Antilles and tropical South America. ( JW) 191

VA N D E A E

Description (Plate 78; Fig. 654.1, 654.2) Epiphytic herbs. Stem enclosed in basal leaf-sheaths. Leaves 1–6, basal, distichous, flat, narrowly oblong to lanceolate, twisted at base so as to lie in one plane, apex often unequally bilobed, articulate and short-sheathing at base. Inflorescences lateral, usually shorter than leaves, often many borne simultaneously on a plant, usually erect; peduncle longer than rachis, both usually pricklyhairy, with 1–6 flowers open at a time; floral bracts persistent. Flowers resupinate, 0.5–1.5 cm across, ephemeral, opening successively, widely opening. Sepals and petals free, similar, spreading. Petals narrower than sepals. Labellum motile on column foot or

rigidly attached to base of column, usually trilobed, spurred, side lobes narrow, erect, midlobe often replicate on spur, sometimes with two basal lobules on both sides, with a median tooth, giving a ‘four-lobed’ appearance; spur prominent, usually spreading at open end, rarely pendent. Column bent forward at an obtuse angle at base of stigma, as long as or longer than column foot, or foot absent; pollinia two, waxy, subglobular to ovoid, entire, or rarely four and the pollinia then completely split into two unequal bodies, each of the pollinia often attached to stipe by an auriculate caudicle, stipe cuneate or narrowly triangular-cuneate to spatulate, viscidium triangular or ovate; rostellum beaked. ( JW)

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Fig. 654.1.  Grosourdya appendiculata (Blume) Rchb.f. A. Habit; B. Flower, side view; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum, from above; G. Labellum, oblique view; H. Labellum, longitudinal section; I. Column and labellum, side view; J. Column, side view; K. Column apex, anther cap removed; L. Column, side view; M. Anther cap, dorsal view; N. Anther cap, ventral view; O. Pollinarium; P. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Kerr 0794 (K) and Kew Spirit Collection no. 46990.

192

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Fig. 654.2.  Grosourdya emarginata (Blume) Rchb.f. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum, from above; H. Labellum, longitudinal section; I. Column and labellum, side view; J. Column, anther cap removed, front view; K. Column, anther cap removed, side view; L. Pollinarium; M. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Flora of Siam 4059 (K) and Kew Spirit Collection no. 10453.

Distribution (Fig. 654.3)

Phylogenetics

The 12 species of Grosourdya are collectively distributed from the Andaman Islands, Burma, Thailand, and Indochina to Malaysia, Indonesia, and the Philippines. ( JW)

Grosourdya has been expanded and now includes the former Ascochilopsis, Ascochilus, and some species of Biermannia (clade C). Grosourdya s.l. is sister to Brachypeza s.l. (AK, MC)

Cytogenetics No chromosome numbers have been recorded for Grosourdya. ( JW)

Phytochemistry No reports have been found on the phytochemistry of Grosourdya. (NV, RG)

Ecology All species of Grosourdya are epiphytes from 0 to 1600 m. Grosourdya appendiculata (Blume) Rchb.f. is recorded as epiphytic on tree trunks in broad-leaved evergreen forest in China, on cultivated fruit trees near sea level in Thailand, and as a common twig epiphyte in Sumatra (Comber 2001) and also in Java (Comber 193

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Fig. 654.3.  Distribution map of Grosourdya.

1990). Grosourdya bicornuta J.J.Wood & A.L.Lamb is recorded from forests at c. 1300 m in Sabah. Grosourdya incurvicalcar (J.J.Sm.) Garay is epiphytic on branches of small trees such as Diospyros L. (Ebenaceae) at c. 30 m in Peninsular Malaysia. Grosourdya quinquelobata (Schltr.) Garay is reported from remnant riverine forests at 800–900 m in Sulawesi. Grosourdya tripercus (Ames) Garay is known from forests at sea level in the Philippines. ( JW)

Pollination There is no information available on the pollination of Grosourdya. ( JW)

Uses No uses have been reported for Grosourdya; it is uncommon in cultivation. ( JW)

Cultivation Group 1 (see subtribal treatment). Species are best grown under warm to intermediate conditions with high humidity and moderate light levels on slabs of cork with some moss. Plants should be misted frequently but allowed to dry out between waterings. (AS, MM)

Taxonomic notes The recently described monotypic genus Theana Aver. from Vietnam (Averyanov 2012a) agrees in most respects, especially 194

in the abruptly swollen rachis of the inflorescence, with species formerly included in the genus Ascochilopsis but differs in having four strongly unequal pollinia, a ligulate callus at the mouth of the spur, and a glabrous inflorescence. Recognizing the morphological diversity in many other genera in the Aeridinae, we consider it likely that Theana belongs here. (AS)

Taxonomic literature Averyanov, L. (2012a). New orchid taxa and records in the flora of Vietnam. Taiwania, 57, 127–52. Comber, J. B. (1990). Orchids of Java. Bentham-Moxon Trust, Royal Botanic Gardens, Kew. Comber, J. B. (2001). Orchids of Sumatra. Royal Botanic Gardens, Kew. Kocyan, A. and Schuiteman, A. (In press). New combinations in Aeridinae. Phytotaxa.

6 5 5 .  GU NNA R EL L A Gunnarella Senghas, Orchidee (Hamburg), 39, 71 (1988). Type species: Gunnarella carinata (J.J.Sm.) Senghas (basionym: Chamaeanthus carinatus J.J.Sm.)

Derivation of name The name commemorates the Danish diplomat and orchid specialist, Gunnar Seidenfaden (1908–2001), specializing in the orchid flora of Southeast Asia; -ella, Latin diminutive suffix. (AS)

H OLCOGLOSSUM

Description (Fig. 655.1)

6 5 6 .   H OL C OGL OS S U M

Epiphytic herbs. Roots terete, not or sparsely branching, glabrous, smooth. Stem patent, rooting in basal part. Leaves conduplicate, articulate, distichous, narrowly oblong to linear, straight to subfalcate, twisted at base so as to lie in one plane with stem, often tinged red-purple, apex asymmetric, bilobed with obtuse lobes, margins smooth. Inflorescence lateral, almost wiry, solitary, erect, patent or pendent, racemose, many-flowered; peduncle much shorter than rachis. Flowers resupinate, c. 0.5 cm wide, ephemeral, opening simultaneously, quaquaversal, pale yellow or whitish, sometimes tinged purple. Sepals entire, free or shortly connate at base, linear to ovate, obtuse. Petals similar to sepals. Labellum hinged with apex of column foot, not spurred, not fleshy, often folded lengthwise, sometimes with a basal callus, entire or trilobed, the midlobe, if present, laterally flattened. Column semiterete, column foot distinct, as long as column or longer; anther cucullate, pollinia four, in two, subglobose, unequal pairs, stipe about as long as pollinia, linear to spatulate, viscidium rounded. (AS)

Holcoglossum Schltr., Repert. Spec. Nov. Regni Veg. Beih., 4, 285 (1919). Type species: Holcoglossum quasipinifolium (Hayata) Schltr. (basionym: Saccolabium quasipinifolium Hayata) Ascolabium S.S.Ying, Coll. Illustr. Indig. Orch. Taiwan, 1, 51 (1977). Type species: Ascolabium pumilum (Hayata) S.S.Ying (basionym: Saccolabium pumilum Hayata) Penkimia Phukan & Odyuo, Orchid Rev., 114, 330 (2006). Type species: Penkimia nagalandensis Phukan & Odyuo Chenorchis Z.J. Liu, K.W. Liu & L.J.Chen, Acta Ecol. Sin., 28, 2435 (2008). Type species: Chenorchis singchii Z.J.Liu, K.W.Liu & L.J.Chen Paraholcoglossum Z.J.Liu, S.C.Chen & L.J.Chen, PLoS ONE, 6(10), 6 (2011). Type species: Paraholcoglossum amesianum (Rchb.f.) Z.J.Liu, S.C.Chen & L.J.Chen (basionym: Vanda amesiana Rchb.f.) Tsiorchis Z.J.Liu, S.C.Chen & L.J.Chen, PLoS ONE, 6(10), 7 (2011). Type species: Tsiorchis kimballiana (Rchb.f.) Z.J.Liu, S.C.Chen & L.J.Chen (basionym: Vanda kimballiana Rchb.f.)

Distribution (Fig. 655.2) Gunnarella is a genus of 10 species from New Guinea, the Solomon Islands, Vanuatu, and New Caledonia. (AS)

Cytogenetics Felix and Guerra (2010) reported a chromosome number of 2n =38 for G. robertsii (Schltr.) Senghas. (AP)

Phytochemistry No reports have been found on phytochemistry of Gunnarella. (NV, RG)

Phylogenetics Gunnarella robertsii together with Rhinerrhizopsis and Dryadorchis forms the New Guinea clade. Although not a large genus, Gunnarella is clearly polyphyletic; another species, G. begaudii (N.Hallé) Senghas (from New Caledonia), is sister to the rest of clade E. (AK, MC)

Ecology All species are epiphytes in forests at 0 to 900 m. (AS)

Pollination

Derivation of name From the Greek holkos, strap, and glossa, tongue, in reference to the strap-like spur on the labellum. ( JW)

Description (Plate 79; Fig. 656.1–656.3) Epiphytic herbs. Stems erect or pendent, completely enclosed by distichously arranged, persistent leaf-sheaths, rooting mostly from base or lower part. Leaves distichous, duplicate, terete, semi-terete, broadly semi-terete, or triquetrous, adaxial surface channelled, articulate, dilated and sheathing at base, deciduous. Inflorescence axillary, racemose, few- to many-flowered; floral bracts shorter than pedicellate ovary. Flowers resupinate, usually widely opening, often white or yellowish. Sepals free, subsimilar, dorsally usually carinate, lateral sepals often slightly larger, oblique. Petals similar to dorsal sepal. Labellum sessile, firmly attached to column base, saccate or spurred, trilobed, lateral lobes erect beside entrance to spur, midlobe larger, often narrowed and with appendages at base; spur cylindrical, attenuate toward tip, usually arcuate, interior unornamented. Column prominently winged, foot absent, clinandrium deeply cleft in front; pollinia two, solid, waxy, globular, provided with a narrow, elongate pore, attached by a common linear to subelliptic, tapering stipe to a broad viscidium; rostellum acute, bifid. ( JW)

Nothing is known about pollination in Gunnarella. (AS)

Distribution (Fig. 656.4)

Uses

Holcoglossum comprises 16 species collectively distributed mainly in China, extending to Taiwan, Vietnam, Laos, Thailand, Burma, and northeastern India. ( JW)

No uses have been reported for Gunnarella; it is rare in cultivation. (AS)

Cultivation Group 1 (see subtribal treatment). (MM)

Cytogenetics Holcoglossum amesianum (Rchb.f.) Christenson, H. flavescens (Schltr.) Z.H.Tsi, H. kimballianum (Rchb.f.) Garay, H. lingulatum 195

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Fig. 655.1.  Gunnarella robertsii (Schltr.) Senghas. A. Habit, ×0.67; B. Flower, ×6; C. Anther cap, dorsal and ventral views, ×18; D. Pollinia, ×18; E. Labellum, side view, ×14; F. Column, ×18; G. Labellum, ×14; H. Dorsal sepal, ×12; I. Petal, ×12; J. Lateral sepal, ×12. All magnifications as originally published. Drawn by Sue Wickison from Cribb and Wheatley 5. Reproduced with permission from B. A. Lewis and P. J. Cribb (1991), Orchids of the Solomon Islands and Bougainville. Royal Botanic Gardens, Kew.

196

H OLCOGLOSSUM

Fig. 655.2.  Distribution map of Gunnarella.

(Aver.) Aver., H. nujiangense X.H.Jin, S.Chen & D.Z.Li, H. rupestre (Hand.-Mazz.) Garay, H. sinicum Christenson, H. subulifolium (Rchb.f.) Christenson, H. wangii Christenson, and H. weixiense X.H.Jin & S.C.Chen all have a chromosome number of 2n = 38. Jin et al. (2007) reported that H. tsii T.Yukawa is a tetraploid with 2n = 76 and suggested that H. rupestre, with similar but smaller flowers, may be one of its ancestors. They remarked that ‘the low frequency of polyploidy in Holcoglossum in the Hengduan Mountains region supports the conclusion that chromosome stasis during rapid speciation is common there’. ( JW, AS)

Phytochemistry No reports have been found on phytochemistry of Holcoglossum. (NV, RG)

Phylogenetics DNA-based phylogenetic analyses by Fan et al. (2009) and Liu et al. (2011) have provided little support for the infrageneric classifications of Holcoglossum proposed by Christenson (1987) and Jin (2005). However, Penkimia and Ascolabium were not included in these analyses, which therefore do not present a complete picture of the relationships of the genus. Moreover, the study of Liu et al. (2011) deviates from other molecular analyses of Aeridinae in that it shows Papilionanthe nested within Holcoglossum,

which would render Holcoglossum paraphyletic. In addition, the name Papilionanthe would have priority at genus level. In Liu et al. (2011), the position of Papilionanthe within Holcoglossum s.l. could be due to the fact that the authors used pseudogene copies of matK. Xiang et al. (2012) recognized the problems with that study and transferred the segregate genera Paraholcoglossum, Penkimia, and Tsiorchis back into Holcoglossum. The same authors also moved Ascolabium pumilum and Ascocentrum himalaicum into Holcoglossum. Holcoglossum is member of clade F. At a larger scale, Holcoglossum s.l. is sister to Vanda s.l. (Kocyan et al. 2008; Gardiner et al. 2013), and then this pair is sister to Luisia and Papilionanthe in clade F, which also includes Seidenfadenia and Rhynchostylis. (AK, MC, AS)

Ecology Species of Holcoglossum are epiphytes in forests between 1000 and 3200 m, except for H. calcicola Schuit. & P.Bonnet, which is found in Laos as a lithophyte on bare vertical limestone cliffs in open deciduous forest at 450 m. ( JW, AS)

Pollination Based on studies performed in China, more is known about pollination in Holcoglossum than in any other genus of Aeridinae. Jin et al. (2005) reported that Holcoglossum rupestre is pollinated by a 197

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Fig. 656.1.  Holcoglossum himalaicum (Deb, Sengupta & Malick) Aver. [formerly Ascocentrum himalaicum (Deb, Sengupta & Malick) Christenson]. A. Habit; B. Leaf, transverse section; C. Flower, lateral sepals and petal removed, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum, from above; H. Column and labellum, side view; I. Column, anther cap removed, front view; J. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Forrest 21630 (K).

beetle, Hybovalgus bioculatus Kolbe, which was found to be the only effective pollinator even though another beetle species was a much more frequent visitor. Ants and bees were also observed as visitors, frequently so in the case of ants, but not as pollinators. Unusually, pollinia were deposited on two different parts of the pollinator’s body: the front legs and the posterior end. It is suggested that the ‘strong buttered tea smell’ of the orchid discourages bees from entering the spur in search of nectar. Liu et al. (2006) observed that H. amesianum is self-pollinating; in this species the anther cap falls off, and the stipe then bends through 180°, depositing the pollinia on the stigmatic surface. 198

Holcoglossum nujiangense X.H.Jin & S.C.Chen is pollinated by an apid bee, Micraphis andreniformis Smith (Jin et al. 2007). The spur of this species does not contain ‘obvious nectar’. According to Jin et al. (2007), two other species, H. weixiense and H. flavescens, are also pollinated by bees, whereas H. sinicum is autogamous. Holcoglossum nagalandense (Phukan & Odyuo) X.H.Jin forms a symbiosis with ants of the genus Temnothorax (Liu et al. 2008, as Chenorchis singchii Z.J.Liu, K.W.Liu & L.J.Chen). This species is at first sight different from other species of Holcoglossum because of its much smaller, inconspicuous flowers. In this connection Jin et al. (2005) reported that ants commonly visited H. rupestre

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F O Fig. 656.2.  Holcoglossum kimballianum (Rchb.f.) Garay. A. Habit; B. Flower; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum; G. Column and labellum. side view; H. Column, front view; I. Column, side view; J. Column apex, anther cap removed, front view; K. Column apex, anther cap removed, back view; L. Anther cap, dorsal view; M. Anther cap, ventral view; N. Pollinarium; O. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Henry 13545 (K) and Kew Spirit Collection no. 71860.

without ever being pollinators. This would indicate that H. nagalandense could have evolved from large-flowered, bee- or beetlepollinated species of Holcoglossum through specialization towards ant-pollination, and demonstrates how plastic floral morphology can be even in closely related species of orchids, an observation that has serious implications for the many, morphologically narrowly circumscribed genera of Aerideae. (AS, JW)

Uses No uses have been reported for Holcoglossum; several species are cultivated. ( JW)

Cultivation Group 4 (see subtribal treatment). Plants should be grown as for cool-growing species of Aerides, except for H. calcicola, which needs more warmth. Species with pendent leaves are best grown on pieces of cork. (AS, MM)

Taxonomic notes Holcoglossum may be recognized by the relatively short stems with crowded, terete to semi-terete leaves. Seidenfadenia and Paraphalaenopsis are similar vegetatively, but Seidenfadenia differs 199

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H A D Fig. 656.3.  Holcoglossum nagalandense (Phukan & Odyuo) X.H.Jin [formerly Penkimia nagalandensis Phukan & Odyuo]. A. Habit; B. Inflorescence; C. Flower; D. Bract; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum; I. Labellum and spur, opened; J. Column, front view; K. Column, side view; L. Anther cap; M. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Redrawn by Judi Stone after S. Phukan and N. Odyuo, Orchid Review, 114, 330 (2006).

in the crest-shaped rostellum that extends upward to the dorsal side of the column, and Paraphalaenopsis differs in the presence of a well-developed column foot and the spurless labellum. (AS)

Taxonomic literature Christenson, E. A. (1987). An infrageneric classification of Holcoglossum (Orchidaceae: Sarcanthinae) with a key to the genera of the Aerides–Vanda alliance. Notes from the Royal Botanic Garden, Edinburgh, 44, 249–56. Fan, J., Qin, H-N., Li, D-Z., and Jin, X-H. (2009). Molecular phylogeny and biogeography of Holcoglossum (Orchidaceae: Aeridinae) based on nuclear ITS, and chloroplast trnL-F and matK. Taxon, 58, 849–61.

200

Gardiner, L. M., Kocyan, A., Motes, M., Roberts, D. L., and Emerson, B. C. (2013). Phylogenetic patterns in the genus Vanda and related genera (Orchidaceae). Botanical Journal of the Linnean Society, 173, 549–72. Jin, X-H. (2005). Generic delimitation and a new infrageneric system in the genus Holcoglossum (Orchidaceae: Aeridinae). Botanical Journal of the Linnean Society, 149, 465–8. Kocyan, A., de Vogel, E. F., Conti, E., and Gravendeel, B. (2008). Molecular phylogeny of Aerides (Orchidaceae) based on one nuclear and two plastid markers: a step forward in understanding the evolution of the Aeridinae. Molecular Phylogenetics and Evolution, 48, 422–43. Liu, Z-J., Chen, L-J., Chen, S-C., Cai, J., Tsai, W-C., Hsiao, Y-Y., Rao, W-H., Ma, X-Y., and Zhang, G-Q. (2011). Paraholcoglossum

H YMENORCHIS

Fig. 656.4.  Distribution map of Holcoglossum.

and Tsiorchis, two new orchid genera established by molecular and morphological analyses of the Holcoglossum alliance. PLoS ONE, 6(10), e24864. doi:10.1371/journal.pone.0024864. Xiang, X., Li, D., Jin, X., Hu, H., Zhou, H., Jin, W., and Lai, Y. (2012). Monophyly or paraphyly – the taxonomy of Holcoglossum (Aeridinae: Orchidaceae). PLoS ONE, 7(12), e52050. doi:10.1371/journal.pone.0052050.

657.   HY ME N O RC H IS Hymenorchis Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 994 (1913). Type species: Hymenorchis javanica (Teijsm. & Binn.) Schltr. (basionym: Oeceoclades javanica Teijsm. & Binn.)

Derivation of name

leaf-sheaths smooth or papillose-rugose, margins entire or fimbriate. Inflorescence lateral, solitary or sometimes in sparse fascicles, patent or pendent, racemose; rachis almost umbellate, rarely elongating over time and producing flowers over a long period. Flowers usually opening simultaneously, quaquaversal, thin-textured, usually translucent white or greenish with a green labellum. Sepals free, usually relatively broad and obtuse, sometimes narrower and acute, margins denticulate to serrulate. Petals similar to sepals. Labellum not motile, spurred, concave, entire, margins entire or denticulate to serrulate; spur up to as long as sepals. Column semi-terete, often dilated apically, lacking a column foot; anther cucullate, pollinia two, subglobose, with a small pore, stipe often tapering toward apex, longer than pollinia, viscidium elliptic to rectangular; rostellum bidentate. Ovary glabrous or rarely pubescent. Capsule terete, sometimes angular. (AS)

From the Greek hymen, a membrane, and orchis, orchid, so named for the thin-textured flowers, especially when dried. (AS)

Distribution (Fig. 657.2)

Description (Plate 80; Fig. 657.1)

Hymenorchis is a genus of 12 species collectively from Vietnam, Java, Philippines, New Guinea, and New Caledonia. It may be expected to occur also in Sulawesi, the Moluccas, the Solomon Islands, and Vanuatu. (AS)

Epiphytic herbs. Roots terete, not or sparsely branching, glabrous, smooth. Stem patent, rooting in the basal part. Leaves conduplicate, articulate, distichous, linear-oblong to elliptic-ovate, often twisted at base so as to lie in one plane with stem, usually fleshy, surface rugulose; margins smooth or more frequently denticulate to serrulate; apex obtuse to emarginate, minutely bidentate or mucronate;

Cytogenetics There are no published reports of chromosome numbers for the genus. (AP) 201

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Fig. 657.1.  Hymenorchis nannodes Schltr. and Hymenorchis saccata Schltr. A. Habit; B. Flower; C. Dorsal sepal; D. Lateral sepal; E. Lateral sepal, showing keel on back; F. Petal; G. Labellum; H. Column and labellum, side view; I. Column and ovary, front view; J. Column apex, anther cap removed; K. Anther cap, dorsal view; L. Anther cap, ventral view; M. Ovary, transverse section; N. Pollinarium. A–M. H. nannodes; N. H. saccata. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone. A-M from Kew Spirit Collection no. 31007; N from Kew Spirit Collection no. 41755.

Phytochemistry

Ecology

No reports have been found on phytochemistry of Hymenorchis. (NV, RG)

Species are epiphytes in forests, usually on thin, lichen-covered branches and twigs of large or isolated trees, from 300 to 2600 m, usually above 1700 m. (AS)

Phylogenetics Hymenorchis is a member of clade E, and the two sequenced species are monophyletic and sister to a subclade that includes Amesiella, Ceratocentron, Cryptopylos, Macropodanthus, Porrorhachis, Trachoma, and Tuberolabium. This overall topology was also supported by Topik et al. (2005). After further study, including type species, some alterations to the circumscription of these genera can be expected. (AK, MC) 202

Pollination Nothing is known about pollination in Hymenorchis. (AS)

Uses No uses have been reported for Hymenorchis; it is uncommon in cultivation. (AS)

JEJEWOODIA

Fig. 657.2.  Distribution map of Hymenorchis.

Cultivation

Description (Fig. 658.1)

Group 1 (see subtribal treatment), but requiring somewhat drier and brighter conditions. The roots should be able to dry off rapidly. Plants should be misted frequently. Most species are coolgrowing. (MM, AS)

Epiphytic herbs. Stem up to c. 20 cm long, simple or branching, rooting at base, leafy. Leaves distichous, ensiform, flattened. Inflorescence lateral, axillary, borne from upper portion of stems, one-flowered, sometimes four or five flowers open simultaneously in succession per stem; floral bracts less than 1/8 the length of pedicel, brownish. Flowers up to 1 cm in diameter, usually opening widely, sepals and petals white, less often yellow or greenish yellow, labellum white, sometimes with a yellow central patch at base of midlobe. Sepals and petals free, spreading. Sepals narrowly elliptic, acute. Petals broadly elliptic and obtuse or narrowly oblong and subacute. Labellum trilobed, pandurate, spurred or slightly saccate, constricted near middle, hypochile canaliculate, thickened at the margins, margins erect, epichile transversely elliptic, emarginate with a broadly triangular tooth in sinus, or rounded, thick; spur (when present) narrow, shallowly conical, entrance glabrous. Column foot absent; anther apically elongate, pollinia four, ovoid, unequal, viscidium transversely elliptic, stipe linear; rostellum finger-like. ( JW)

Taxonomic notes Although a distinctive taxon, Hymenorchis belongs to a problematic group of genera closely allied to Pteroceras; generic delimitation in this group needs further study. (AS)

Taxonomic literature Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

658.   J E JE WO O D IA Jejewoodia Szlach., Fragm. Flor. Geobot. Suppl., 3, 135 (1995). Type species: Jejewoodia jiewhoei (J.J.Wood & Shim) Szlach. (basionym: Ceratochilus jiewhoei J.J. Wood & Shim)

Distribution (Fig. 658.2)

Derivation of name

Cytogenetics

The genus is named after Jeffrey J. Wood (1952–), former Curator of the Orchid Herbarium at the Royal Botanic Gardens, Kew. (JW)

No chromosome numbers have been recorded for Jejewoodia. ( JW)

The six species of Jejewoodia are endemic to Borneo. ( JW)

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Fig. 658.1.  Jejewoodia jiewhoei (J.J.Wood & Shim) Szlach. A. Plant; B. Flower, front view; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum, front view; G. Column and labellum, side view; H. Column and labellum, longitudinal section; I. Anther cap with pollinia, front view; J. Pollinarium. Drawn as Ceratochilus jiewhoei J.J.Wood & Shim by Mark Fothergill from Lamb AL 611/86, Beaman 10673, and Lamb AL 58/83. Reproduced with permission from Chan et al. (1994), Orchids of Borneo. Volume 1. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

Phytochemistry

Ecology

No reports have been found on phytochemistry of Jejewoodia. (NV, RG)

Jejewoodia crockerensis J.J.Wood & A.L.Lamb occurs in forests on sandstone at 1200–1700 m. Jejewoodia jiewhoei is a twig epiphyte in forests at elevations of 900–1800 m. Jejewoodia jongirii J.J.Wood & A.L.Lamb and J. rimauensis J.J.Wood & A.L.Lamb are twig epiphytes in forests at 1300–1600 m. Jejewoodia linusii J.J.Wood & A.L.Lamb is epiphytic on small branches in forests at 1200–1700 m. Jejewoodia longicalcarata (Ames & C.Schweinf.) J.J.Wood & A.L.Lamb is reported from forests, sometimes on ultramafic substrate, at 1100–1500 m. (JW)

Phylogenetics Jejewoodia is a member of clade I. The genus is sister to Microsaccus, from which it differs in the number of flowers per inflorescence open at one time, one versus two, but flower number is variable in some cases, so this appears not to be a good basis for generic distinction. Adenoncos, which differs among other traits in an absence of a spur, is weakly supported as sister to this pair. Topik et al. (2005) found a different set of relationships for these genera. (AK, MC) 204

Pollination There is no information available on the pollination of Jejewoodia. ( JW)

LUISIA

Fig. 658.2.  Distribution map of Jejewoodia.

Uses

Derivation of name

No uses have been reported for Jejewoodia; it is rare in cultivation. ( JW)

The name commemorates the Spanish explorer Don Luis de Torres (-1493) who was an interpreter for Christopher Columbus on his first voyage to the Caribbean. ( JW)

Cultivation Group 1 (see subtribal treatment). (AS)

Taxonomic literature Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

659.   LUIS IA Luisia Gaudich. in Freycinet, Voy. Uranie, 426 (1829). Type species: Luisia teretifolia Gaudich. Mesoclastes Lindl., Gen. Sp. Orch. Pl., 44 (1830). Type species: Mesoclastes brachystachys Lindl., lectotype selected here. Birchea A.Rich., Ann. Sci. Nat., Bot., sér. 2, 15, 66, t. 10 (1841). Type species: Birchea teretifolia A.Rich. Trichorhiza Lindl. ex Steud., Nom., ed. 2, 2, 702 (1841). Type species: Trichorhiza teretifolia (Gaudich.) Lindl. ex Steud. (basionym: Luisia teretifolia Gaudich.) Lockia Aver., Orchids, 81, 363 (2012). Type species: L. sonii Aver.

Description (Plates 81, 82; Fig. 659.1) Epiphytic or lithophytic herbs. Roots flattened, borne from lower nodes. Stem elongate, erect or climbing, often branched at base giving a tufted habit, some with a single shoot, terete, usually stiff, enclosed in leaf-bases. Leaves many, well-spaced, linear, terete, distichous, quaquaversal or secund, articulated to a tubular sheathing base. Inflorescence lateral, erect or ascending, racemose, subsessile, fewer than 10-flowered, with few to several flowers opening simultaneously, sometimes reduced to a solitary flower; peduncle and rachis attenuate, rachis congested. Flowers resupinate, usually greenish with a maroon labellum, continuing to increase in size for some days after they open, long lasting. Sepals and petals free, similar or petals longer and narrower, spreading, lateral sepals often cymbiform and dorsally carinate or narrowly winged toward apex. Labellum firmly adnate to base of column, spur absent, pendent, flat, often distinctly divided by a transverse, curved furrow into a narrow basal hypochile and a broader apical epichile of different thickness and texture, hypochile often concave, base sometimes with lateral lobes embracing column, epichile often extending forward, adaxial 205

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Fig. 659.1.  Luisia curtisii Seidenf. A. Plant; B. Flower, front view; C. Pollinarium, front view; D. Pollinarium, side view; E. Anther; F. Labellum; G. Column and labellum, longitudinal section; H. Ovary, transverse section. Drawn by Shim Phyau Soon from living plant collected from Lohan River, Mt. Kinabalu. Reproduced with permission from Chan et al. (1994), Orchids of Borneo. Volume 1. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

surface often longitudinally wrinkled, grooved or tuberculaterugose. Column straight, semi-cylindrical, column foot absent, clinandrium shallowly excavate; anther cap terminal, operculate, pollinia two, waxy, globular, solid, with a short slit, caudicles absent, attached by a short and broad stipe to a single short, broad viscidium; rostellum apex subtruncate, entire or slightly emarginate; stigma transverse. Ovary triangular in cross-section. ( JW)

Distribution (Fig. 659.2) The approximately 40 species of Luisia are collectively distributed in Australia, Sri Lanka, India, Bhutan, China, Thailand, Indochina, Korea, Japan, Malaysia, Indonesia, the Philippines, New Guinea, and east to Fiji and Samoa in the Pacific islands. ( JW) 206

Palynology Schill and Pfeiffer (1977) studied pollen of L. teretifolia Gaudich. (= L. tristis (G.Forst) Hook.f.) and an unknown entity named ‘L. birdean’, which might have been L. birchea Blume (= L. tenuifolia Blume). Pollen tetrads were convex with a laevigate surface. (AP)

Cytogenetics Brandham (1999) listed chromosome numbers for the genus as 2n = 38, 40. Felix and Guerra (2010) also listed n = 18. ( JW)

Phytochemistry In early work, Boorsma (1902) noted that L. brachystachys (Lindl.) Blume contained traces of alkaloids but no saponins. Alkaloids

LUISIA

Fig. 659.2.  Distribution map of Luisia.

have been detected in L. trichorrhiza (Hook.) Blume, L. tristis (as L. teretifolia), and L. volucris Lindl. (Lüning 1967; Lawler and Slaytor 1969). These are present at relatively low levels (< 0.1%); a later survey indicated that of eight species tested, none contained alkaloids at ≥ 0.1% (Lüning 1974). (NV, RG)

Phylogenetics Luisia, Papilionanthe, and Paraphalaenopsis form a strongly supported subclade in clade F, which is sister to that of Holcoglossum s.l. and Vanda s.l. Though florally distinct, the first three genera have terete leaves. Seidenfaden (1971) cited flower size as an important character within Luisia, but Kocyan (personal communication) found no support for this with molecular data. (AK, MC)

and Clinteria ducalis (Scarabaeidae) were the only flower visitors observed to carry pollinaria of this species. The hypothesis of specialized cantharophily was further supported by the presence of 2-methylbutyric acid and caproic acid in the floral scent. (AK)

Uses Subedi et al. (2011) reported the use in Nepal of a paste made from the leaves of L. trichorrhiza (Hook.) Blume applied externally against muscular pain and of the sap of L. tristis to treat chronic wounds. Similar and other medicinal uses have been reported for Luisia species from India, Sri Lanka, China, and Vietnam (Lawler 1984). Various species of Luisia are sometimes cultivated. (AS)

Cultivation Ecology

Group 4 (see subtribal treatment). (MM)

Species are epiphytic in forests, sometimes lithophytic, at 0 to 2000 m. ( JW)

Taxonomic notes

Pollination Pedersen et al. (2013) studied a population of Luisia curtisii Seidenf. in northern Thailand. They demonstrated a high level of self-compatibility experimentally in contrast to low natural fruit set (1.4–1.9%), indicating that the species depends on external agents for pollination. Their observations showed that L. curtisii is pollinated by beetles, as Lema unicolor (Chrysomelidae)

Luisia is a distinctive and relatively homogeneous genus that is unlikely to be confused with any other genus in Aeridinae. The recently described genus Lockia Aver. (Averyanov 2012b) is possibly a natural hybrid between a Luisia and a Vanda species. (AS)

Taxonomic literature Averyanov, L. (2012b). Lockia sonii and Schoenorchis scolopendria. Orchids, 81, 362–71.

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Description (Plate 83; Fig. 660.1)

Seidenfaden, G. (1971). Notes on the genus Luisia. Dansk Botanisk Arkiv, 27, 1–101.

Usually pendent, epiphytic herbs. Stem to about 10 cm long, obscured by leaf-sheaths, leafy. Leaves narrow, strap-shaped, apex unequally bilobed, coriaceous. Inflorescences often clustered, racemose, as long as the leaves, c. 7–25-flowered; rachis terete or laterally compressed, sometimes slightly keeled or winged, the keels or wings tapering upward and mutually continuous at their base, leaving a concavity where the flower has fallen; floral bracts scale-like, obtuse or acute. Flowers showy, long-lasting, opening simultaneously, sometimes sweetly scented, white or yellow, marked with red, lilac-pink or orange-brown. Dorsal sepal free. Lateral sepals and petals adnate

660.  MAC RO P O DA N TH US Macropodanthus L.O.Williams, Bot. Mus. Leafl. Harvard Univ., 6, 103 (1938). Type species: Macropodanthus philippinensis L.O.Williams

Derivation of name From the Greek makros, long, pous, podos, foot, and anthos, flower, in reference to the long column foot. ( JW) E

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C O Fig. 660.1.  Macropodanthus sabahensis J.J.Wood & A.L.Lamb. A. Habit; B. Flower, side view; C. Flower, back view; D. Floral bract; E. Dorsal sepal; F. Lateral sepal; G. Lateral sepal apex showing keel on back; H. Petal; I. Labellum, oblique view; J. Labellum, from above; K. Pedicel, ovary, labellum, and column, side view; L. Pedicel, ovary, and column, oblique view; M. Anther cap, dorsal view; N. Anther cap, ventral view; O. Pollinia. Drawn by Linda Gurr from Lamb AL 1462/92 (holotype). Reproduced with permission from J. J. Wood et al. (2011), The Orchids of Mount Kinabalu, Volume 2. Natural History Publications (Borneo), Kota Kinabalu, Sabah.

208

MAC ROPODANTHUS

to column foot. Labellum articulate to column foot and bent upward at an angle, motile, distinctly clawed, trilobed, carinate, side lobes linear or tooth-like, midlobe triangular, acute, sometimes reduced to a low fleshy keel, sometimes with a pair of raised linear appendages in front of side lobes; spur distinct, about 1 cm long, saccate, ecallose. Column slightly broadened at truncate apex, with small apical stelidia, column foot long; rostellar projection terminal, about as long as the column, filiform, closely appressed to inner surface of column; anther cap ovate, cucullate, apex often triangular, acute, pollinia two, globose, sulcate, stipe to 5 mm long, viscidium obovate or oblong. ( JW)

Distribution (Fig. 660.2) Macropodanthus includes eight species collectively distributed from the Andaman Islands, Thailand, and Peninsular Malaysia, east to Indonesia and the Philippines. ( JW)

Cytogenetics There are no chromosome counts published for the genus. (AP)

Phytochemistry No reports have been found on the phytochemistry of Macropodanthus. (NV, RG)

Phylogenetics Macropodanthus is well supported as closely related to Amesiella, Cryptopylos, and Dyakia in clade E. Topik et al. (2005) identified a similar relationship. (AK, MC)

Ecology Macropodanthus philippinensis L.O.Williams occurs in hill forests at an elevation of 500 m in the Philippines. In Borneo, M. rimauensis J.J.Wood is reported from lower montane forests at elevations of 1500–1600 m, and M. sabahensis J.J.Wood & A.L.Lamb is recorded from mixed lower montane forests on sandstone ridges. Macropodanthus teysmannii (Miq.) H.Æ.Pedersen occurs at elevations of 1200–1900 m in Sumatra. ( JW)

Pollination No information has been published on pollination of Macropodanthus. ( JW)

Uses No uses have been reported for Macropodanthus; some species are sometimes cultivated. ( JW)

Cultivation Group 1 (see subtribal treatment). (MM)

Fig. 660.2.  Distribution map of Macropodanthus.

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Taxonomic notes

6 6 1 .  M I C ROP ER A

Macropodanthus is similar to Pteroceras but may be distinguished by the long-lasting flowers opening simultaneously on the inflorescence. The clawed labellum is bent upward so as to form a right angle (or less often a more shallow angle) with the elongate column foot. The rostellum is beaked and nearly always as long as the column. ( JW)

Micropera Lindl., Bot. Reg., 13, sub t. 1522 (1832). Type species: Micropera pallida (Roxb.) Lindl. (basionym: Aerides pallida Roxb.) Camarotis Lindl., Gen. Sp. Orch. Pl., 219 (1833). Type species: Camarotis purpurea Lindl.

Derivation of name From the Greek mikros, small, and pera, pouch, in reference to the labellum. ( JW)

Taxonomic literature Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

Description (Plate 84; Fig. 661.1) Climbing, epiphytic herbs. Stem simple or branching, up to 1 m long, internodes to about 2 cm long, bearing many support

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Fig. 661.1.  Micropera callosa (Blume) Garay. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Ovary, labellum, and column, longitudinal section; H. Anther cap, dorsal view; I. Pollinarium, front view; J. Pollinarium, side view; K. Ovary, transverse section. Drawn by Chan Chew Lun and Lucy Liew Fui Ling from Lamb AL 1150/89. Reproduced with permission from J. J. Wood et al. (2011), The Orchids of Mount Kinabalu, Volume 2. Natural History Publications (Borneo), Kota Kinabalu, Sabah.

210

MICROPERA

roots at intervals. Leaves linear, to 17 × 2 cm, usually much less. Inflorescence lateral, leaf-opposed, bending through a right angle so the rachis points either straight up or straight down, racemose, to c. 15 cm long, few- to many-flowered. Flowers usually non-resupinate, fleshy, usually yellow with purple markings, or pink. Sepals oblong or obovate, obtuse, often reflexed, lateral sepals connate for a short distance at base. Petals linearobovate or oblong, obtuse, often reflexed. Labellum porrect, saccate, spurred, apex trilobed, midlobe much smaller than sac or spur; spur saccate, with a back wall callus, longitudinal septum, and usually a bilobed callus on front wall. Column variable, usually extended into a proboscis-like structure; pollinia four, as two unequal masses, stipe linear, viscidium ovate. (JW)

Phytochemistry Alkaloids are present in M. fasciculata (Lindl.) Garay (as Camarotis keffordii (F.M.Bailey) J.J.Sm.) at levels of approximately 0.01% (Lawler and Slaytor 1969). (NV, RG)

Phylogenetics

Micropera comprises about 20 species collectively distributed from India (Sikkim), Bhutan, China (Hainan), and Indochina, south to Malaysia and Indonesia, east to the Philippines, New Guinea, Australia, and the Solomon Islands. (JW)

Micropera has had an unstable taxonomic history. It was established in 1832 by Lindley, who placed it in Camarotis the following year. It was later treated as a section of Sarcochilus and then again considered a member of Camarotis. Only Tang and Wang (1951) and Garay (1972) accepted it as a distinct genus. Micropera belongs to clade L, but exactly how it fits among these taxa is unclear (low levels of support). Given the weak support, it is not surprising that Topik et al. (2005) and Carlsward et al. (2006b), who both sequenced Micropera pallida, found different topologies, neither of which is found here. Micropera pallida and M. philippinensis appear to be related to Uncifera, some species of Stereochilus, and Smitinandia, and this set of taxa is in some manner related to the mess that is Cleisostoma. The final status of Micropera cannot be determined until Cleisostoma has been thoroughly investigated. (AK, MC)

Cytogenetics

Ecology

Brandham (1999) and Felix and Guerra (2010) reported a chromosome number of 2n = 38 for the genus. (AP)

Species are epiphytes in swamp and broad-leaved forests from 0 to 1200 m. Micropera pallida climbs up tree trunks where light

Distribution (Fig. 661.2)

Fig. 661.2.  Distribution map of Micropera.

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levels are high along roadsides, on isolated trees in scrub, and on cultivated village trees in Sumatra and Java. Micropera sterrophylla (Schltr.) Garay is epiphytic on the branches of large roadside trees at 600–1200 m in Sulawesi. ( JW)

1.5–3.0 times the diameter of the pollinia, viscidium narrowly elliptic. ( JW)

Distribution (Fig. 662.2)

Pollination

Microsaccus includes 13 species collectively distributed from Burma to Indonesia and the Philippines. ( JW)

There is no information available on pollination of Micropera. ( JW)

Cytogenetics

Uses

No chromosome numbers have been recorded for Microsaccus. ( JW)

No uses have been reported for Micropera; it is uncommon in cultivation. ( JW)

Phytochemistry

Cultivation Group 2 (see subtribal treatment). (MM)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogeny of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Garay, L. A. (1972). On the origin of the Orchidaceae. II. Journal of the Arnold Arboretum, 53, 202–15. Tang, H. C. and Wang, F. T. (1951) Contribution to the knowledge of eastern Asiatic orchids 2. Acta Phytotaxomica Sinica, 1, 305–26. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

662.  MIC RO SAC C US Microsaccus Blume, Bijdr., 367 (1825). Type species: Microsaccus javensis Blume

No reports have been found on phytochemistry of Microsaccus. (NV, RG)

Phylogenetics Microsaccus is closely related to Jejewoodia in clade I, from which it differs having two flowers per inflorescence open at a time rather than one. Adenoncos is also related, but it lacks a spur, whereas the other two genera have one. Topik et al. (2005) found Microsaccus to be related to Abdominea and Cleisomeria, but this relationship lacked support. (AK, MC)

Ecology Species are epiphytes in forests, open deciduous and evergreen swamp forests from 0 to 2000 m. ( JW)

Pollination There is no information available on pollination of Microsaccus. ( JW)

Uses No uses have been reported for Microsaccus; it is rare in cultivation. ( JW)

Derivation of name

Cultivation

From the Greek mikros, small, and sakkos, sac, in reference to the small, saccate labellum. ( JW)

Group 2 (see subtribal treatment). (MM)

Description (Plate 85; Fig. 662.1) Epiphytic herbs. Stem up to 20 cm long, often curved, simple or branching, leafy, rooting at base. Leaves laterally flattened, distichously arranged, usually imbricate at base. Inflorescences arising from leaf axils, one- or two-flowered, often borne opposite one another; floral bracts linear and ovate. Flowers white, fleshy. Sepals and petals usually not widely spreading. Sepals usually free, laterals sometimes joined at base to and decurrent along spur. Petals free. Labellum firmly adnate to base of column, entire, emarginate or slightly bilobed, ecallose, pitcher-shaped and quadrangular in outline, oblong, spurred; spur globose or narrow and shallowly conical, sometimes transversely compressed, ecallose. Column foot absent; pollinia four, equal, stipe linear-clavate, 212

Taxonomic literature Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

6 6 3 .  M OB I L A B I U M Mobilabium Rupp, North Queensland Naturalist, 13, 2 (1946). Type species: Mobilabium hamatum Rupp

Derivation of name From the Latin mobilis, mobile, and labium, lip, describing the labellum that is motile on a short claw. ( JW)

MOBILABIUM

D 3 mm

A

1 cm

3 cm

1 mm

C

E

B

1 cm 5 mm

2 mm

2 mm

G

F

I

H

K 3 cm

L

M

J

P 2 mm

0.5 mm

1 cm

N

Q

3 mm

O

Fig. 662.1.  Microsaccus Blume drawings. A–E. M. griffithii (C.S.P.Parish & Rchb.f.) Seidenf. A. Plant; B. Apical portion of flowering stem; C, D. Inflorescence; E. Pollinarium. F–I. M. sumatranus J.J.Sm. F. Plant; G. Flower; H. Column and labellum; I. Labellum with spur (all after J. J. Smith). J–M. M. ampullaceus J.J.Sm. J. Flowering stem (after Holttum); K. Flower; L. Column and labellum; M. Column (after J. J. Smith). N–Q. M. javensis Blume. N. Flowering stem (after Holttum); O. Flower; P. Column and labellum; Q. Pollinarium. O–Q after J. J. Smith. Drawn by G. Seidenfaden. Reproduced with permission from G. Seidenfaden (1992), Orchids of Peninsular Malaysia and Singapore: A Revision of R.E. Holttum: Orchids of Malaya. Olsen & Olsen, Fredensborg, Denmark.

Description (Fig. 663.1) Epiphytic herbs. Stem 3–60 cm long, erect or straggling, branching when long. Leaves 3–12 or more, widely spaced, oblong, apex acute, usually uncinate, margins often decurved, stiff, coriaceous, yellow-green. Inflorescence 2–6 cm long, racemose, 5–15-flowered; peduncle usually shorter than rachis. Flowers 5–7 mm wide, remaining open for over two days, not opening widely, cream, pale green or brownish with purple or red markings on labellum and column. Sepals concave. Petals subfalcate, slightly shorter and narrower than sepals. Labellum hinged to and hanging from upturned apex of column foot so that the disc is vertical, side lobes vertical, well developed, suboblong, acute, proximal halves of front margins strongly incurved;

disc thick and fleshy, with a prominent, upwardly directed, transverse septum with a cavity on top, the cavity containing a sticky substance; spur subovoid or cylindrical. Column slightly curved and projected forward over labellum, foot at a right angle to column proper, apex uncinate; anther cap with a slightly upturned beak, pollinia four, in two closely appressed pairs on short caudicles attached to a short stipe, viscidium pyriform; stigma quadrate or circular; rostellum decurved, bifid. ( JW)

Distribution (Fig. 663.2) Mobilabium is monospecific and endemic to northeastern tropical Australia. ( JW) 213

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Fig. 662.2.  Distribution map of Microsaccus.

Cytogenetics

Cultivation

No chromosome numbers have been recorded for M. hamatum. ( JW)

Group 2 (see subtribal treatment). This species is easily grown on a slab in bright light with frequent misting, high humidity, and brisk air movement. (PA, MM)

Phytochemistry A survey of alkaloid distribution in Orchidaceae gave a negative result for M. hamatum. (Lawler and Slaytor 1969). (NV, RG)

6 6 4 .  OM OEA

Phylogenetics

Omoea Blume, Bijdr., 359 (1825). Type species: Omoea micrantha Blume

Mobilabium is a member of the Australia clade where it is strongly supported as sister to Drymoanthus. (AK)

Derivation of name

Ecology

From the Greek omoios, alike, referring to a resemblance with the genus Ceratochilus (= Trichoglottis). (AS)

The species is epiphytic on rain forest trees on the plateaux at 500–1300 m. It is locally common along streams, roads, fences, and relictual trees, sometimes in exposed situations, where it is pendent and attached by a few roots. (PA, JW)

Description (Fig. 664.1)

Pollination There is no information available on pollination of Mobilabium. ( JW)

Uses No uses have been reported for Mobilabium; it is occasionally cultivated. ( JW) 214

Epiphytic herbs. Roots terete, not branching, glabrous, smooth or papillose (O. philippinensis Ames). Stem elongate, patentascending to erect, leafy, rooting at base, sparsely branching, the branches well above base in O. micrantha. Leaves conduplicate, articulate, distichous, linear, almost semi-terete, apex obtuse; leaf-sheaths finely ribbed or transversely wrinkled. Inflorescences lateral, solitary or in pairs, racemose, 1–6-flowered. Flowers resupinate or not, 0.3–0.6 cm wide, opening simultaneously, quaquaversal, sepals and petals green with maroon markings or greenish yellow, labellum yellowish green with brownish side

OMOEA

B

C

P

N O

Q

K L M

J E A G

F D

H I Fig. 663.1.  Mobilabium hamatum Rupp. A. Habit; B. Flower, front view; C. Flower, side view; D. Bract; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum from above; I. Labellum, longitudinal section; J. Column and labellum, side view; K. Column, front view; L. Column apex, anther cap removed; M. Column, side view; N. Anther cap, dorsal view; O. Anther cap, ventral view; P. Pollinarium; Q. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from A.W. Dockrill, Australian Indigenous Orchids 2, 943 (1992) and Kew Spirit Collection no. 28235.

lobes or white with violet markings. Sepals entire, free, obtuse or acute. Petals narrower than sepals. Labellum spurred, trilobed, rigidly attached to column; spur conical to cylindrical, bilobed at apex. Column foot absent; anther cucullate, short-rostrate, pollinia two, angular-ovoid, stipe about as long as pollinia, linear. (AS)

Distribution (Fig. 664.2) Omoea is a genus of only two species—O. micrantha and O. philippinensis—collectively from Sumatra, Java, and the Philippines (Luzon). (AS)

Cytogenetics There are no published chromosome counts for the genus. (AP)

Phytochemistry No reports have been found on phytochemistry of Omoea. (NV, RG)

Phylogenetics Omoea occupies an isolated position in clade L, but until relationships of the various subclades of Cleisostoma are better 215

VA N D E A E

Fig. 663.2.  Distribution map of Mobilabium.

understood, the status of Omoea cannot be robustly assessed. (AK, MC)

Ecology Both species are epiphytes in forests at 800–2000 m. (AS)

Pollination Nothing is known about pollination in Omoea. (AS)

Uses No uses have been reported for Omoea; it is rare in cultivation. (AS)

Cultivation Group 1 (see subtribal treatment). (MM)

665.  O PH IO G LO SSE LLA Ophioglossella Schuit. & Ormerod, Kew Bull., 53, 742 (1998). Type species: Ophioglossella chrysostoma Schuit. & Ormerod

Derivation of name From the Greek ophis, serpent, and glossa, tongue, referring to the forked rostellum that resembles a snake’s tongue, and the Latin diminutive suffix -ella. (AS) 216

Description (Plate 86; Fig. 665.1) Epiphytic herbs. Roots terete, not branching, glabrous, smooth. Stem pendent, rooting at base. Leaves 3–5, conduplicate, articulate, distichous, elliptic to linear-elliptic, subfalcate, twisted at base so as to lie in one plane with stem, margins denticulate near the unequally bidentate apex. Inflorescence lateral, solitary, racemose, many-flowered; peduncle elongate, patent; rachis continuing the line of the peduncle, flexuose, angular. Flowers resupinate, about 1.5 cm wide, longevity unknown, opening a few at a time, quaquaversal, light purple, with orange-yellow and deeper purple markings on labellum. Sepals entire, free, elliptic or obliquely ovate, obtuse. Petals narrower than sepals, obovate-oblong. Labellum hinged with apex of column foot, not spurred, concave, trilobed, with two parallel calli in centre of labellum between lateral lobes, midlobe elliptic, with a raised crest along median, the midlobe connected to lateral lobes by a pair of lamellae, one on each side of central crest. Column curved, dilated apically, column foot distinct, about half as long as column; anther helmet-shaped, long-rostrate, pollinia four, separate, obliquely ovoid, stipe longer than pollinia, narrowly obovate-oblong; rostellum elongate, bifurcate at apex after removal of pollinarium. (AS)

Distribution (Fig. 665.2) Ophioglossella is monospecific from New Guinea. (AS)

OPH IOGLOSSELLA

C

A

B

H D

F

I

E

G

K J

Fig. 664.1.  Omoea micrantha Blume. A. Habit; B. Flower, front view; C. Flower and bract, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum; H. Column and labellum, side view; I. Column and labellum, longitudinal section; J. Column, anther cap removed, front view; K. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone; I and K redrawn after M. Kromohardjo in J.J. Smith, Die Orchideen von Java, Figuren-Atlas, Fig. 481 (1914); others from Comber 1673 (K).

Cytogenetics

Ecology

There are no published chromosome numbers for the genus. (AP)

The species is epiphytic in forests from 800 to 2300 m. (AS)

Phytochemistry

Nothing is known about pollination in Ophioglossella. (AS)

No reports have been found on phytochemistry of Ophioglossella. (NV, RG)

Uses

Phylogenetics There is no phylogenetic information available for this genus. (AK)

Pollination

No uses have been reported for Ophioglossella; it is not known to be in cultivation. (AS)

Cultivation Group 1 (see subtribal treatment). (MM) 217

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Fig. 664.2.  Distribution map of Omoea.

Taxonomic notes

Derivation of name

Ophioglossella has not yet been included in phylogenetic studies. Based on morphology and biogeography, it may be conjectured that Dryadorchis is among its nearest relatives. (AS)

From the Latin papilio, butterfly, and the Greek anthos, flower. ( JW)

Description (Plate 87; Fig. 666.1) Taxonomic literature Schuiteman, A. and de Vogel, E. F. (2006). Flora Malesiana: Orchids of New Guinea, vol. IV; Genera Kuhlhasseltia to Ophioglossella. CD-ROM. ETI, Amsterdam, Nationaal Herbarium Nederland, Leiden.

666.  PAPILIO N A N TH E Papilionanthe Schltr., Orchis, 9, 78 (1915). Type species: Papilionanthe teres (Roxb.) Schltr. (basionym: Dendrobium teres Roxb.) Taxa at the rank of section referable to Papilionanthe: Vanda sect. Teretifoliae Pfitzer, in Engler & Prantl, Natürl. Pflanzenfam. 2, 6, 214 (1889). Type species: Vanda teres (Roxb). Lindl. (basionym: Dendrobium teres Roxb.) Aerides sect. Phalaenidium Pfitzer, in Engler & Prantl, Natürl. Pflanzenfam. 2, 6, 217 (1889). Type species Aerides vandarum Rchb.f. Vanda sect. Teretivanda Kuntze, in Post & Kuntze, Lex. Gen. Phan., 584 (1903), nom. illeg. Type species: Vanda teres (Roxb). Lindl. (basionym: Dendrobium teres Roxb.) 218

Scrambling terrestrial or epiphytic herbs. Stem climbing or pendent, terete, elongate, branching or not. Leaves distichous, fleshy, terete, channelled adaxially, articulate to a sheathing base; sheaths coriaceous, tightly enclosing stem, persistent. Inflorescence lateral, 1- to several-flowered. Flowers resupinate. Sepals and petals free, subsimilar, spreading, petals usually larger. Labellum broadly adnate to column foot, non-articulate, spurred, variously trilobed, side lobes erect, either parallel to or embracing column, midlobe often dilated and bi- or trilobed; spur funnel-shaped, conical to long horn-shaped, without interior callosities. Column subterete, foot prominent; pollinia two, shortly cleft, waxy, attached by a broadly triangular or subquadrate stipe to a large viscidium; rostellum beaked. ( JW)

Distribution (Fig. 666.2) Papilionanthe consists of 12 species collectively distributed in India, China, Southeast Asia, and the Malay Archipelago. ( JW)

Cytogenetics Felix and Guerra (2010) listed chromosome numbers of 2n = 38, 49, 76. ( JW)

PAPILIONANTHE

B

C

L M

N O I

J K

D

F

P

Q

E

A H

G

Fig. 665.1.  Ophioglossella chrysostoma Schuit. & Ormerod. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum, from above; H. Labellum, longitudinal section; I. Column and labellum, side view; J. Column, front view; K. Column apex, anther cap removed; L. Column, side view; M. Anther cap, dorsal view; N. Anther cap with pollinia, ventral view; O. Anther cap, ventral view; P. Pollinarium; Q. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Kew Spirit Collection no. 46222.

Phytochemistry

Phylogenetics

Extracts of stems of P. teres (as Vanda teres) contain (2R)-2-(phydroxybenzyl)malic acid (eucomic acid) together with vandaterosides I–III (Fig. 666.3), which are glucopyranosyloxybenzyl eucomate derivatives (Simmler et al. 2011). The last are also present in leaves and roots of P. teres. In bioassays, eucomic acid and vandateroside II were shown to stimulate cellular respiratory functions in keratinocytes, suggesting that they might be used as components of preparations to combat skin aging (Simmler et al. 2011). Several species contain alkaloids at levels of 0.001–0.01% (Lüning 1964), including P. teres (as Vanda teres), P. uniflora (Lindl.) Garay (as Aerides longicornu Hook.f.), and P. vandarum (Rchb.f.) Garay (as Aerides vandarum Rchb.f.). (NV, RG)

Papilionanthe shares a complicated taxonomic history with Vanda. However, they are not closely related within clade F. It is strongly supported as related to Paraphalaenopsis and Luisia, with which it shares the terete to semi-terete leaves. Carlsward et al. (2006b) found a similar topology. Unification in an enlarged genus might be a possibility, but floral differences are numerous. (AK, MC)

Ecology Species are terrestrial in open swamps or epiphytic in forests at 0–2400 m. Papilionanthe hookeriana (Rchb.f.) Schltr. occurs as terrestrial in full sun in swamps near sea level in Peninsular 219

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Fig. 665.2.  Distribution map of Ophioglossella.

Malaysia and Borneo, usually in deep peaty soils, often growing with shrubs and tall grasses for support. The swamps are subject to seasonal flooding when the roots become submerged; this species has become scarce in both areas due to over-collection and burning of peat-swamps. Papilionanthe teres is epiphytic on tree trunks in open forests or at forest margins at 500–900 m in China; it is recorded as an epiphyte on tall forest trees at 200– 2400 m in Bhutan and occurs on trees in scrub or on rocks at 200 m and as an epiphyte on Butia Becc. (Arecaceae) at 500 m in Thailand. ( JW)

Pollination Van der Pijl and Dodson (1966) suggested pollination by carpenter bees (Xylocopa spp.). They noted that dead bees have been found in the flowers of P. (Vanda) teres, and its fragrance was similar to that of specialized Xylocopa-pollinated flowers. ( JW)

Uses Subedi et al. (2011) reported the use in Nepal of a paste made from P. teres applied externally to treat dislocated bones. In Peninsular Malaysia, a decoction of P. (Vanda) hookeriana has been used as a poultice to treat rheumatism and pains in the bones and joints (Lawler 1984). Papilionanthe is horticulturally significant and has often been used in hybridization. (AS, JW) 220

Cultivation Group 4 (see subtribal treatment). (MM)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogeny of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Christenson, E. A. (1999). Proposal to conserve the name Dendrobium teres (Orchidaceae) against Limodorum subulatum. Taxon, 48, 587–88. Garay, L. A. (1972). On the systematics of the monopodial orchids II. Botanical Museum Leaflets, Harvard University, 23, 369–72.

6 6 7 .  PA P I L L I L A B I U M Papillilabium Dockrill, Austral. Sarcanth., 31, t. 7 (1967). Type species: Papillilabium beckleri (F.Muell. ex Benth.) Dockrill (basionym: Cleisostoma beckleri F.Muell. ex Benth.)

Derivation of name From the Latin papilla, nipple, and labium, lip, referring to the small warts that cover the labellum. ( JW)

PAPILLILABIUM

L

B

J

G

K

I

C E

A

H

D

F

Fig. 666.1.  Papilionanthe teres (Roxb.) Schltr. A. Habit; B. Flower; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum; G. Column and labellum, side view (half of labellum removed); H. Column side view; I. Column apex, anther cap removed; J. Anther cap, dorsal view; K. Anther cap, ventral view; L. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Ross, cult. s.n. (1895) (K) and Pantling 45 (K).

Description (Fig. 667.1) Epiphytic herbs. Stem up to 4 cm long, but usually much shorter. Leaves 2–6, linear, acute. Inflorescence racemose, 1–4 cm long, 3–8-flowered; peduncle and rachis equal. Flowers 5–7 mm across, resupinate, green, sometimes marked purplish brown

or crimson, labellum white, green or greenish yellow, strongly fragrant, lasting longer than one day. Dorsal sepal free. Lateral sepals adnate to base of column foot, falcate-oblong. Petals falcate-linear. Labellum rigidly joined to apex of column foot, trilobed, spurred; disc papillose, the papillae extending into spur, 221

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Fig. 666.2.  Distribution map of Papilionanthe.

Fig. 666.3.  Phenolic constituents of Papilionanthe teres.

222

PAPILLILABIUM

B

A

C

I

M

N

J D F O

K

E

L

H

G

Fig. 667.1.  Papillilabium beckleri (F.Muell.ex Benth.) Dockrill. A. Habit; B. Flower, front view; C. Flower and bract, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum, from above; H. Labellum, longitudinal section; I. Column and labellum, side view; J. Column, front view; K. Column, side view; L. Column apex, anther cap removed, side view; M. Anther cap, dorsal view; N. Anther cap, ventral view; O. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone; habit redrawn after R. D. Fitzgerald, Australian Orchids (1875–1894) and others from Kew Spirit Collection no. 53110.

side lobes crescent-shaped, free portion spreading, subdeltoid, obtuse, midlobe decurved, bifid or emarginate; spur 2.0–2.5 mm long, dilated in proximal half with distal half cylindrical when viewed from the front, obtuse to almost truncate, posterior wall formed by column foot, ecallose, anterior wall papillose near mouth. Column 1.5 mm long, erect, not winged, column foot in line with column, 2.0–2.5 mm long; anther cap reniform, with a rostrum deflexed at a right angle, pollinia four, obovoid, in two closely appressed, subequal pairs, stipe about 1½ times as long as pollinia, slightly dilated near apex, curling and twisting after removal from rostellum, viscidium ovate; stigma deeply set, rostellum decurved. ( JW)

Distribution (Fig. 667.2) Papillilabium is monospecific and confined to Australia (southern Queensland and northern New South Wales). ( JW)

Cytogenetics No chromosome numbers have been recorded for Papillilabium. ( JW)

Phytochemistry No reports have been found on phytochemistry of Papillilabium. (NV, RG) 223

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Fig. 667.2.  Distribution map of Papillabium.

Phylogenetics

Taxonomic literature

Papillilabium is part of the Australia clade, but relationships within this clade are not well supported. (AK, MC)

Riley, J. J. and Banks, D. P. (2002). Orchids of Australia. Princeton University Press.

Ecology

6 6 8 .  PA R A PH A L A ENOPS I S

Papillilabium beckleri is a twig epiphyte up to 1000 m, especially common as populations overhanging watercourses along humid, well-lit rain forest margins and growing on sclerophyll shrubs. (PA, JW)

Paraphalaenopsis A.D.Hawkes, Orquídea (Rio de Janeiro), 25, 212 (1963). Type: Paraphalaenopsis denevei (J.J.Sm.) A.D.Hawkes (basionym: Phalaenopsis denevei J.J.Sm.)

Pollination There is no information available on pollination of Papillilabium. ( JW)

Uses No uses have been reported for Papillilabium; it is rare in cultivation. ( JW)

Cultivation Group 1 (see subtribal treatment). As for Sarcochilus twig epiphytes but with medium shade and reduced watering in winter. (PA) 224

Derivation of name From the Greek para, near or beside, and Phalaenopsis, a genus of tropical Asiatic orchids with similar flowers. (PC)

Description (Plate 88; Fig. 668.1) Pendent epiphytic or rarely lithophytic herbs. Stem unbranched, leafy. Leaves few, pendent, coriaceous, terete, with a longitudinal groove along their length, acute. Inflorescence axillary, racemose, unbranched, subclavate, spreading horizontally, few-flowered; peduncle and rachis terete. Flowers resupinate, with spreading perianth; pedicel terete, elongate, usually longer than floral segments; bracts triangular-ovate, cucullate, shorter than ovary. Sepals free, spreading, oblanceolate to elliptic, the laterals oblique. Petals spreading, oblanceolate. Labellum porrect, firmly attached to column foot, clawed

PARAPH ALAENOPSIS

2 cm 2 mm

E

B

1 cm

A

D F

2 mm

G 10 cm

2 mm

H 1 cm

C Fig. 668.1.  Paraphalaenopsis laycockii (M.R.Hend.) A.D.Hawkes. A. Plant; B. Flower, front view; C. Column and labellum, longitudinal section; D. Labellum, spread; E. Anther; F. Pollinarium, front view; G. Pollinarium, side view. H. Ovary, transverse section. Drawn by Shim Phyau Soon and Chan Chew Lun from cultivated material at Tenom Orchid Centre, Sabah. Reproduced with permission from Chan et al. (1994), Orchids of Borneo. Volume 1. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

at base, trilobed, lacking a spur, with a lobed, raised callus between side lobes, midlobe porrect or slightly deflexed, broadening at tip, side lobes erect on either side of column. Column terete-clavate, with a foot; anther-cap obovoid, pollinia two, deeply cleft, stipe broadly spatulate, viscidium subcircular-cordate; stigma a ventral cavity; rostellum bifid. Ovary six-ridged, rarely angular. (PC)

Distribution (Fig. 668.2) Paraphalaenopsis is a genus of four species endemic to Borneo. (PC)

Cytogenetics Brandham (1999) and Felix and Guerra (2010) reported chromosome counts of 2n = 38 for the genus. (AP)

Phytochemistry In surveys of alkaloid content, both P. denevei (as Phalaenopsis denevei) and P. serpentilingua (J.J.Sm.) A.D.Hawkes (as Phalaenopsis serpentilingua J.J.Sm.) gave positive results (Lüning 1964, 1967). Chemical composition data are available for the floral fragrance of P. laycockii (M.R.Hend.) A.D.Hawkes (Kaiser 2011). The main component is 6-methoxyeugenol (22.0%); other aromatics including elemicine (5-allyl-1,2,3-trimethoxybenzene), 1,2,3,5-tetramethoxybenzene, and 3,4,5-trimethoxybenzaldehyde are present at lower levels (all < 5.0%), although only 6-methoxyeugenol and elemicine are said to contribute to the spicy, aromatic quality of the scent. Aldehydes, including octanal (13.5%), decanal (0.7%), and (Z)-dec-4-enal (1.5%) are responsible for an ‘aldehydic citrus-peel’ aspect of the fragrance, 225

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Fig. 668.2.  Distribution map of Paraphalaenopsis.

together with dodecanoic and undecadienoic acids. Relatively large amounts of the ketone pentadecan-2-one (9.2%) were also found (Kaiser 2011). (NV, RG)

Phylogenetics Paraphalaenopsis is a member of clade F in a well-supported subclade with Luisia and Papilionanthe characterized by terete to semi-terete leaves. Carlsward et al. (2006b) found Paraphalaenopsis also to be related to Luisia and Papilionanthe. (AK, MC)

Ecology Paraphalaenopsis denevei is found in lowland riverine forests below 300 m elevation. Paraphalaenopsis serpentilingua occurs in swamp and lowland forests, where it is occasionally found growing on rocks. Paraphalaenopsis laycockii and P. labukensis Shim, A.L.Lamb & C.L.Chan grow in forests as epiphytes at elevations up to 1000 m. (PC)

Pollination There is no information available on pollination of Paraphalaenopsis. (AS)

Uses No uses have been reported for Paraphalaenopsis; it is frequently cultivated and used in hybridization. (AS) 226

Cultivation Group 4 (see subtribal treatment). Plants may be grown in baskets with a well-draining medium or on slabs of cork with some moss and humus around the roots. These plants require high humidity, regular watering throughout the year, and moderate light levels. The roots should be allowed to dry out between waterings. (AS, MM)

Taxonomic notes The floral morphology of Paraphalaenopsis is similar to that of some Phalaenopsis species, and the first three species to be discovered were described as Phalaenopsis. However, their distinctive habit with hanging, terete leaves differs markedly from those of Phalaenopsis, a fact recognized by horticulturists who called them ‘rat-tailed phalaenopsis’. (PC)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogeny of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Schuiteman, A. (In press). Paraphalaenopsis, an endangered orchid genus from Borneo. Malesian Orchid Journal.

6 6 9 .  P EL ATA NT H ER I A Pelatantheria Ridl., J. Linn. Soc., Bot., 32, 371 (1896). Type species: Pelatantheria ctenoglossum Ridl.

PELATANTHERIA

Derivation of name From the Greek pelates, neighbour, approacher, and the Latin anthera, anther, possibly referring to the stelidia that enfold the anther cap and approach each other in so doing. ( JW)

Description (Plate 89; Fig. 669.1) Epiphytic or lithophytic herbs. Stem elongate, clambering, rooting from nodes, usually slightly compressed-trigonous, manynoded, rigid, enclosed in persistent leaf-sheaths, sometimes branched. Leaves many, usually densely distichous, flat, rarely semi-cylindrical, bilobulate, rarely obtuse, coriaceous, articulated

to a sheathing base. Inflorescence lateral, racemose, few-flowered. Flowers resupinate. Sepals and petals free, similar, petals slightly smaller, both with longitudinal purple lines. Labellum rigidly adnate to column, spurred at base, trilobed, side lobes erect, midlobe often shortly caudate, adaxial surface thickened and cushion-like centrally; spur narrowly conical, with a longitudinal septum or ridge on inner surface, and with a tough appendage on back wall near entrance. Column lacking a column foot, apex with two long and incurved stelidia; pollinia four, arranged in two pairs, slightly unequal, solid, waxy, semi-globular, stipe irregular, much broader than long viscidium crescent-shaped; rostellum abbreviated. ( JW)

B C J

P K E

M G

L F

H

I

A O Q

N

D

Fig. 669.1.  Pelatantheria ctenoglossum Ridl. A. Habit; B. Flower, side view; C. Flower, front view; D. Bract; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum, from above; I. Labellum, longitudinal section; J. Column and labellum, side view; K. Column, front view; L. Column, anther cap removed, front view; M. Column, side view; N. Anther cap, dorsal view; O. Anther cap with pollinia, ventral view; P. Pollinarium; Q. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Kerr 0883 (K) and Kew Spirit Collection 60523.

227

VA N D E A E

Distribution (Fig. 669.2)

Ecology

The eight species of Pelatantheria are collectively distributed in India and Southeast Asia, China, Korea, and Japan south to Sumatra. ( JW)

Species of Pelatantheria are epiphytes or lithophytes in forests at 700–1100 m. ( JW)

Cytogenetics

Pollination

Brandham (1999) and Felix and Guerra (2010) reported a chromosome number of 2n = 38 for the genus. ( JW)

There is no information available on pollination of Pelatantheria, although the labellum of P. insectifera (Rchb.f.) Ridl. would seem to indicate deception. ( JW, FR)

Phytochemistry

Uses

Low levels of alkaloids have been detected in P. ctenoglossum (Lüning 1967). In a later survey, Lüning (1974) noted that two species were tested for alkaloid content, but neither contained these compounds at levels ≥ 0.1%. (NV, RG)

Phylogenetics As far as it has been sampled (three species), Pelatantheria is monophyletic and a member of clade L. It falls in a wellsupported subclade that includes Cleisomeria, two species of Cleisostoma, Rhynchogyna, and Schoenorchis. The taxonomic disposition of Pelatantheria and these other genera cannot be robustly assessed until the larger clade (L) is better understood. Topik et al. (2005) found generally similar relationships for Pelatantheria. (AK, MC)

No uses have been reported for Pelatantheria; it is occasionally cultivated. ( JW)

Cultivation Group 2 (see subtribal treatment). (MM)

Taxonomic notes The taxonomic status of the genera closely allied to Cleisostoma, such as Pelatantheria, Rhynchogyna, Sarcoglyphis, Sarcophyton, Stereochilus, and others, is still unresolved, and Pelatantheria is only tentatively maintained here. Within this alliance, Pelatantheria may be recognized by the elongate stems, relatively small leaves, and short, few-flowered inflorescences. (AS)

Fig. 669.2.  Distribution map of Pelatantheria.

228

PENNILABIUM

Taxonomic literature

Description (Plate 90; Fig. 670.1)

Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

Epiphytic herbs. Stem leafy. Leaves few, clustered, ligulate or linearelliptic, often subfalcate, often twisted at base, unequally bilobed, fleshy, to 11 × 3 cm. Inflorescence lateral, emerging from base of leaf-sheaths, racemose, 3–8 cm long; rachis complanate, with one or two distichously arranged flowers opening in succession; floral bracts distichous. Flowers lasting for a day or two, white, cream, yellow or orange. Sepals and petals subequal, 1–2 cm long. Petals sometimes toothed. Labellum trilobed, rigidly attached to base of column, spurred, internal callosities absent, side lobes either well developed and truncate or reduced to ear-like lobes, when present often fimbriate or toothed; midlobe fleshy and solid or reduced to a fleshy lobe; spur ecallose and aseptate, often with raised edges near entrance. Column compressed ­dorsally, column foot absent; pollinia two, nearly round, entire, stipe broader below pollinia, often spatulate, 3–5 times the diameter of the

670.   PE N N ILA BIUM Pennilabium J.J.Sm., Bull. Jard. Bot. Buitenzorg, sér. 2, 13, 47 (1914). Type species: Pennilabium angraecum (Ridl.) J.J.Sm. (basionym: Saccolabium angraecum Ridl.)

Derivation of name From the Latin penna, feathered, and labium, lip, in reference to the erose or fimbriate margins of the labellum side lobes in some species. ( JW)

L

K 2 mm

J 2 mm 4 mm

G

A

5 mm

4 mm

F

H

I

1 mm

D 6 mm

B 6 mm

C

E

Fig. 670.1.  Pennilabium struthio Carr. A. Habit; B. Flower, front view; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Pedicel with ovary, lower portion of labellum showing spur, and column, side view’ G. Labellum and column, longitudinal section; H. Labellum margin showing hairy appendages; I. Lower portion of labellum and column, anther cap removed, front view; J. Anther cap, dorsal view; K. Pollinarium; L. Ovary, transverse section. Drawn by C. L. Chan from Lamb AL 382/85. Reproduced with permission from J. J. Wood (2003), Orchids of Borneo. Volume 4. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

229

VA N D E A E

pollinia; rostellar beaked; stigma covering almost the entire front side of column. ( JW)

once greater sampling of the species has been accomplished. (AK, MC)

Distribution (Fig. 670.2)

Ecology

With about 15 species, Pennilabium is distributed from northern India (Assam) through Thailand, Vietnam, and Malaysia to Indonesia and the Philippines. ( JW)

Cytogenetics

Pennilabium kidmancoxii J.J.Wood is an epiphyte in forests at 1300– 1500 m in Borneo. Pennilabium struthio is recorded as an epiphyte in forests at 200–500 m in Borneo. ( JW)

Pollination

No chromosome counts for Pennilabium have been reported. ( JW)

The erect, lamellate keels at the entrance to the spur in, for example, P. struthio, are transparent and may serve as guides for the maxillae of visiting moths. ( JW)

Phytochemistry

Uses

No reports have been found on phytochemistry of Pennilabium. (NV, RG)

Phylogenetics In clade E, Pennilabium (two species sampled) is well supported as a member of a subclade that includes one of the two sampled species of Biermannia, Chroniochilus, and Spongiola. Pennilabium struthio Carr is moderately to well supported as sister to the single species in Spongiola, so it would appear that some changes to the taxonomy of this clade will be required

No uses have been reported for Pennilabium, and it is not commonly cultivated. ( JW)

Cultivation Group 2 (see subtribal treatment). (MM)

Taxonomic notes The generic delimitation of the genera closely allied to Pteroceras, including Pennilabium, needs further study, and Pennilabium is only

Fig. 670.2.  Distribution map of Pennilabium.

230

PERISTERANTHUS

tentatively accepted here. It may be recognized by the usually distichous, ephemeral flowers that open in succession, usually with relatively large, spreading lateral lobes and a slender spur. The column lacks a foot, the stipe is markedly dilated near the pollinia, and the rostellum is long and narrow. (AS)

Derivation of name

671.   PE R IS TE R A N TH US

Epiphytic, semi-pendent herbs. Stem up to 25 cm long, usually shorter, rigid, becoming pendent with the weight of the leaves. Leaves 3–10, oblong, unequally emarginate, flat, coriaceous. Inflorescence 5–25 cm long, racemose, pendent, many-flowered, flowers radiating in all directions; rachis c. 1 mm in diameter;

From the Greek peristera, dove, and anthos, flower, referring to the fanciful resemblance of the flowers to a dove. ( JW)

Description (Fig. 671.1)

Peristeranthus T.E.Hunt, Queensland Naturalist, 15, 17 (1954). Type species: Peristeranthus hillii (F.Muell.) T.E.Hunt (basionym: Saccolabium hillii F.Muell.)

N M

B

I

D K

J

F

L

O

E

P

A

G

C

H

Fig. 671.1.  Peristeranthus hillii (F.Muell.) T.E.Hunt. A. Habit; B. Flower and bract, side view; C. Bract; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum, from above; H. Labellum, longitudinal section; I. Column and labellum, side view; J. Column, front view; K. Column apex, anther cap removed; L. Column, side view; M. Anther cap, dorsal view; N. Anther cap with pollinia, ventral view; O. Pollinarium; P. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Weinthal s.n. Sep. 1927 (K) and Kew Spirit Collection no. 53526.

231

VA N D E A E

floral bracts 1 mm long, deltoid. Flowers 4–7 mm in diameter, lasting longer than two days, pale green spotted crimson, fragrant. Sepals and petals subsimilar, narrowly oblong to narrowly spatulate, lateral sepals adnate, at least in part, to base to column foot. Labellum hinged to apex of column foot, bilobed, spurred, side lobes erect, broad at base, tapered to a decurved tip at apex, midlobe vestigial, or possibly modified to form the distal part of disc; disc broad, thick, with an erect, digitiform callus at distal side of spur entrance; spur slightly projecting forward, 1 mm long, broad, internally evenly tapered from a wide orifice to a blunt point at the apex, walls thick, externally much broader laterally than anterior-posterior-aligned, apex often notched. Column 1.0–1.5 mm long, sharply incurved from about the middle, column foot in line with proximal half of column proper; anther cap shallow, with an elongate, narrow, slightly decurved beak, pollinia four, in two closely appressed, subequal pairs, each pollinium attached by an elastic caudicle to a long, narrow stipe; stigma deeply set, subrhombic to almost circular, rostellum decurved, bifid. ( JW)

Distribution (Fig. 671.2) Peristeranthus is monospecific and endemic to eastern tropical and subtropical Australia (northern Queensland to northern New South Wales). ( JW, PA)

Cytogenetics No chromosome numbers have been recorded for Peristeranthus. ( JW)

Phytochemistry Alkaloids have been detected at low levels in P. hillii (Lawler and Slaytor 1969). (NV, RG)

Phylogenetics Peristeranthus is a member of the Australia clade, within which it has weak support as sister to Drymoanthus/Mobilabium. (AK, MC)

Ecology Peristeranthus hillii is epiphytic on trunks and branches of trees in coastal and near-coastal rain forests and also wet tropical rain forests at 0–1000 m. (PA, JW)

Pollination Peristeranthus is visited by a small beetle, Metriorrhynchus rufipennis Fabricius (Wallace 1980; Forster 1988), but a full pollination sequence has not been reported (Adams and Lawson 1993). ( JW, PA)

Fig. 671.2.  Distribution map of Peristeranthus.

232

PH ALAENOPSIS

Uses No uses have been reported for Peristeranthus; it is rare in cultivation. (JW)

Cultivation Group 3 (see subtribal treatment). This species is relatively easy to grow on a slab in a greenhouse with warm humid conditions and brisk air movement. (PA, MM)

672.   PH ALA E N O P S IS Phalaenopsis Blume, Bijdr., 294 (1825). Type species: Phalaenopsis amabilis (L.) Blume (basionym: Epidendrum amabile L.) Polychilos Breda in Kuhl and van Hasselt, Gen. Sp. Orch., 1 (1827). Type species: Polychilos cornu-cervi Breda Doritis Lindl., Gen. Sp. Orch. Pl., 178 (1833). Type species: Doritis pulcherrima Lindl. Synadena Raf., Fl. Tellur., 4, 9 (1836, publ. 1838). Type species: Synadena amabilis (L.) Raf. (basionym: Epidendrum amabile L.) Stauroglottis Schauer, Nov. Act. Acad. Nat. Cur. 19, suppl., 1, 432 (1843). Type species: Stauroglottis equestris Schauer Polystylus Hassk., Natuurk. Tijdschr. Ned. Indië, 10, 3 (1856). Type species: Polystylus cornu-cervi (Breda) Hassk. (basionym: Polychilos cornu-cervi Breda) Stauritis Rchb.f., Hamburger Garten-Blumenzeitung, 18, 34 (1862). Type species: Stauritis violacea (Hort. Bogor ex Witte) Rchb.f. (basionym: Phalaenopsis violacea Hort. Bogor ex Witte) Ornithochilus (Lindl.) Wall. ex Benth., Gen. Pl., 3, 478 (1883). Type species: Ornithochilus difformis (Wall. ex Lindl.) Schltr. (basionym: Aerides difforme Wall. ex Lindl.) Hygrochilus Pfitzer in Engler & Prantl, Natürl. Pflanzenfam. Nachtr., 1, 112 (1897). Type species: Hygrochilus parishii (Veitch & Rchb.f.) Pfitzer (basionym: Vanda parishii Veitch & Rchb.f.) Kingiella Rolfe, Orchid Rev., 25, 197 (1917). Type species: Kingiella taenialis (Lindl.) Rolfe (basionym: Aerides taenialis Lindl.) Grafia A.D.Hawkes, Phytologia, 13, 306 (1966). Type species: Grafia parishii (Rchb.f.) A.D.Hawkes (basionym: Phalaenopsis parishii Rchb.f.) Kingidium P.F.Hunt, Kew Bull., 24, 97 (1970). Type species: Kingidium taeniale (Lindl.) P.F.Hunt (basionym: Aerides taenialis Lindl.) Sedirea Garay & H.Sweet, Orchids S. Ryukyu Is., 149 (1974). Type species: Sedirea japonica (Linden & Rchb.f.) Garay & H.R.Sweet (basionym: Aerides japonica Linden & Rchb.f.) Lesliea Seidenf., Opera Bot., 95, 190 (1988). Type species: Lesliea mirabilis Seidenf. Nothodoritis Z.H.Tsi, Acta Phytotax. Sin., 27, 58 (1989). Type species: Nothodoritis zhejiangensis Z.H.Tsi Grussia M.Wolff, Orchid. Atlas, 165 (2007). Type species: Grussia appendiculata (Carr) M.Wolff (basionym: Phalaenopsis appendiculata Carr)

Taxa at the rank of section referable to Phalaenopsis: Aerides sect. Ornithochilus Lindl., Gen. Sp. Orchid. Pl., 242 (1833). Type species: Aerides difforme Wall. ex Lindl. Phalaenopsis sect. Doritis (Lindl.) J.J.Sm., Repert. Spec. Nov. Regni Veg., 32, 366 (1933), nom. inval. Type species: Phalaenopsis pulcherrima (Lindl.) J.J.Sm.

Infrageneric classification After Christenson (2001), emended by Cribb and Schuiteman (2012) in Renziana, 2, 14–15 and Kocyan and Schuiteman (in press). Phalaenopsis subgenus Parishianae (Sweet) Christenson, Phalaenopsis – A Monograph, 63 (2001). Type species: Phalaenopsis parishii Rchb.f. Phalaenopsis subgenus Parishianae (Sweet) Christenson sect. Aphyllae Sweet, Amer. Orchid Soc. Bull., 37, 872 (1968). Type species: Phalaenopsis stobartiana Rchb.f. Phalaenopsis subgenus Parishianae (Sweet) Christenson sect. Deliciosae Christenson, Selbyana, 9, 167 (1986). Type species: Phalaenopsis deliciosa Rchb.f. Phalaenopsis subgenus Parishianae (Sweet) Christenson sect. Esmeralda Rchb.f., Gard. Chron., n.s., 11, 398 (1879). Type species: Phalaenopsis esmeralda Rchb.f. Phalaenopsis subgenus Parishianae (Sweet) Christenson sect. Parishianae Sweet, Amer. Orchid Soc. Bull. 37, 872 (1968). Type species: Phalaenopsis stobartiana Rchb.f. Phalaenopsis subgenus Phalaenopsis sect. Phalaenopsis. Type species: Phalaenopsis amabilis (L.) Blume (basionym: Epidendrum amabile L.) Phalaenopsis subgenus Phalaenopsis sect. Polychilos (Breda) Rchb.f., Bot. Zeit. 22, 298 (1864). Type species: Phalaenopsis cornu-cervi (Breda) Blume & Rchb.f. (basionym: Polychilos cornu-cervi Breda) Phalaenopsis subgenus Hygrochilus (Pfitzer) Kocyan & Schuit., Phytotaxa (in press). Type species: Phalaenopsis marriottiana (Rchb.f.) Kocyan & Schuit. (basionym: Vanda parishii Veitch & Rchb.f. var. mariottiana Rchb.f.) Phalaenopsis subgenus Ornithochilus (Lindl.) Kocyan & Schuit., Phytotaxa (in press). Type species: Phalaenopsis difformis (Wall. ex Lindl.) Kocyan & Schuit. (basionym: Aerides difforme Wall. ex Lindl.).

Derivation of name From the Greek phalaina, moth, and opsis, likeness, in reference to the appearance of the flower of P. amabilis, hence the common name of ‘moth orchid’. ( JW)

Description (Plates 91-96; Fig. 672.1-672.4) Terrestrial, lithophytic, and epiphytic herbs. Roots terete or flattened. Stem leafy, concealed by overlapping persistent leaf-sheaths, rooting mostly at base. Leaves alternate, distichous, succulent, persistent or sometimes deciduous, oblong to broadly elliptic, sometimes marbled or suffused with purple or silver. Inflorescence axillary racemes or panicles, axillary, erect to pendent; peduncle 233

VA N D E A E

1 cm

C 10 cm

F

A 1 mm

1 cm

E

5 mm

D

B

Fig. 672.1.  Phalaenopsis amabilis (L.) Blume. A. Plant; B. Labellum, spread; C. Column and labellum, longitudinal section; D. Pollinarium; E. Anther; F. Ovary, transverse section. Drawn by Chan Chew Lun from Lamb AL 120/83. Reproduced with permission from Chan et al. (1994), Orchids of Borneo. Volume 1. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

terete; rachis terete or bilaterally compressed, rarely swollen relative to peduncle; floral bracts usually inconspicuous, persistent, succulent or papery. Flowers 1–30, resupinate, produced simultaneously or in succession, often long-lasting, sometimes fragrant, plain or variously spotted, marbled, or barred. Sepals and petals free, spreading, subsimilar to dimorphic, lateral sepals usually oblique and larger than dorsal sepal. Labellum trilobed, clawed, continuous with column foot or rarely hinged at base of a footless column (species formerly included in Hygrochilus), sometimes saccate-spurred or subsaccate, side lobes erect and subparallel, often callose, midlobe oblong-elliptic to obtrullate, rarely transverse, sometimes pubescent to villose, apex sometimes bearing a pair of tendril-like appendages (cirrhi), callus uni-, bi- or triseriate, longitudinal, rarely transverse; sac or spur, if present, without internal ornamentation. Column often subtended by a pair of fleshy, knee-like protrusions, without wings, usually with a 234

column foot, usually dilated lateral to stigma; pollinia two or four, on a common spatulate stipe and viscidium; rostellum sometimes beaked. Ovary pedicel terete, shallowly six-sulcate. ( JW)

Distribution (Fig. 672.5) There are 45–50 species of Phalaenopsis collectively distributed from India (including Andaman and Nicobar Islands) to southern China, Korea, Japan, Thailand, Indochina, Malaysia, and Indonesia to the Philippines, Australia, and New Guinea. The majority of species occur in Indonesia and the Philippines. ( JW)

Anatomy Carlsward et al. (2006a) examined three species of Phalaenopsis. Their results are summarized below.

PH ALAENOPSIS

M

B

C

K

J N Q

I O E G

P

F

L

H A D

Fig. 672.2.  Phalaenopsis japonica (Rchb.f.) Kocyan & Schuit. A. Habit; B. Flower, front view; C. Flower, side view; D. Bract; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum, from above; I. Labellum, longitudinal section; J. Column and labellum, side view; K. Column, front view; L. Column, side view; M. Column apex, anther cap removed; N. Anther cap, dorsal view; O. Anther cap, ventral view; P. Pollinia; Q. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone after W. H. Fitch in Curtis’s Botanical Magazine, t. 5798 (1869) and from Kew Spirit Collection no. 13537.

Leaf

Root

Cuticle smooth to ridged along the contours of the epidermal cells. Ad- and abaxial cuticle 2.5 µm thick. Hairs absent. Stomata abaxial. Outer ledges thin to moderate, inner ledges moderate to thick; cuticular horns inconspicuous. Substomatal chambers small, irregularly shaped. Epidermal cells periclinally oriented to isodiametric. Hypodermis absent. Fibre bundles absent. Mesophyll homogeneous, 15 cells wide. Water-storage cells with birefringent walls absent. Cells above midrib distinctly modified. Vascular bundles collateral, in one row. Thin-walled sclerenchyma associated with xylem and phloem poles. Bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with phloem and xylem sclerenchyma.

Velamen two cells thick. Epivelamen cells isodiametric to tangentially flattened in P. deliciosa and P. stobartiana Rchb.f., radially elongate in P. wilsonii Rolfe. Endovelamen cells angular, isodiametric to radially elongate. Distinct endovelamen thickenings absent. Cover cells present over short cells of exodermis. Hairs present in P. deliciosa. Exodermal cells radially elongate to isodiametric. Long cell walls ∩-thickened in P. stobartiana and P. wilsonii, strongly ∩-thickened in P. deliciosa. Proliferations present in P. deliciosa and P. wilsonii. Cortex 10–21 cells wide. Starch grains present in P. deliciosa. Cells isodiametric to tangentially elongate in P. hainanensis. Water-storage cells with birefringent walls infrequent in P. wilsonii and absent in P. deliciosa and P. stobartiana. 235

VA N D E A E

I

H

J

M

B

N

K

P

O

C

F

E D

L

A G Fig. 672.3.  Phalaenopsis marriottiana (Rchb.f.) Kocyan & Schuit. A. Habit; B. Flower; C. Dorsal sepal; D. Lateral sepal, front view; E. Lateral sepal, back view; F. Petal; G. Labellum, from above; H. Labellum, longitudinal section; I. Column and labellum, side view; J. Column, front view; K. Column apex, anther cap removed; L. Column, side view; M. Anther cap, dorsal view; N. Anther cap, ventral view; O. Pollinarium; P. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Kerr 16 (K) and Kew Spirit Collection no. 53432.

Aeration units present. Endodermal cells heavily ○-thickened in P. deliciosa. Pericyclic cells thin-walled opposite xylem and thickwalled opposite phloem. Vascular cylinder 7–9-arch. Vascular tissue embedded in sclerenchyma. Pith parenchymatous to thick-walled and sclerenchymatous in P. deliciosa and P. stobartiana. (BC)

Palynology Schill and Pfeiffer (1977) reported on the pollen of P. amabilis, P. lueddemanniana Rchb.f., and Doritis pulcherrima (= P. pulcherrima). 236

Tetrads were convex with a laevigate surface and a rounded sexine (calymmate in P. pulcherrima) 1–2 μm thick. (AP)

Cytogenetics Brandham (1999) listed published chromosome numbers for Phalaenopsis as 2n = 38, 55, 57, 69+Bs, c. 74, 76, 92, 100, 114, 152±2. He suggested x = 19 as the basic number, yielding 2x = 38 (most common), 3x = 57, 4x = 76, 6x = 114, 8x = c. 152. For other genera now included in Phalaenopsis, Brandham (1999) reported similar counts of 2n = 38, 40, 57, 76 for Doritis, 2n = 36, 38 for Kingidium, and 2n = 38 for Ornithochilus. (AP)

PH ALAENOPSIS

L

B

A H

G

E

I

C

K D

J F Fig. 672.4.  Phalaenopsis zhejiangensis (Z.H.Tsi) Schuit. [formerly Nothodoritis zhejiangensis Z.H.Tsi]. A. Habit; B. Flower; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum; G. Column and labellum, side view; H. Column, side view; I. Anther cap, ventral view; J. Anther cap, side view; K. Pollinarium; L. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone after drawing in Chen Sing-chi et al., Native Orchids in China in Colour, 298 (1999) and from Tsi 86-006 (K).

Phytochemistry Leaf extracts of P. cornu-cervi and P. pulcherrima (as Doritis pulcherrima) contain flavone C-glycosides (C. Williams 1979). Examples of this class of flavonoids are also found in the flowers of P. schilleriana Rchb.f., principally as saponarin and a 7-O-glucoside of vicenin-2 (Griesbach 1990). The main anthocyanin pigments of the red-purple flowers of P. equestris, P. ×intermedia Lindl., P. ×leucorrhoda Rchb.f., P. sanderiana Rchb.f., and P. schilleriana ‘Pink Butterfly’ are acylated cyanidin glucosides (Tatsuzawa et al. 1997). Two of these (Fig. 672.6, 1 and 2) were identified as cyanidin 3-O-[6-O-(malonyl)glucoside]-7,3∙-di-O-[6-O-(E-sinapoyl)

glucoside] (26.0–64.4% total anthocyanins), and its demalonyl derivative (9.2–33.3% total anthocyanins). Although an earlier report described the presence of a trisinapoylated cyanidin 3,7,3∙-triglucoside in the flowers of P. schilleriana, the complete structure of the pigment was not elucidated (Griesbach 1990). Two other anthocyanin pigments described in the later study of Tatsuzawa et al. (1997) are cyanidin 3,7,3∙-triglucosides triacylated either with ferulic, sinapic and malonic acids, or ferulic (2) and malonic acids; however, the sites of acylation were not determined. Information on other phenolic constituents of Phalaenopsis is limited to reports of phenanthropyran derivatives 237

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Fig. 672.5.  Distribution map of Phalaenopsis.

Fig. 672.6.  Phenolic constituents of Phalaenopsis. Anthocyanins: cyanidin 3-O-[6-O-(malonyl)glucoside]-7,3∙-diO-[6-O-(E-sinapoyl)glucoside] (1), cyanidin 3-O-glucoside-7,3∙-di-O-[6-O(E-sinapoyl)glucoside] (2). Stilbenoids: 2,6,7-trihydroxy-5H-phenanthro[4,5bcd]pyran (3) and its 6-methyl ether (4). Phenolic glucoside: parishin (5).

(Fig. 672.6, 3 and 4) from leaves of P. equestris (Manako et al. 2001) and phenolic glucosides from P. marriottiana (studied as Vanda parishii Veitch & Rchb.f.). The latter were characterized as tris[4(β-d-glucopyranosyloxy)benzyl] citrate (parishin; Fig. 672.6, 5) and 4-(β-d-glucopyranosyloxy)benzyl alcohol (Dahmén and Leander 1976). However, the authors suggested that the benzyl alcohol derivative is probably a hydrolysis product of parishin formed during the isolation of this compound. 238

The alkaloid chemistry of Phalaenopsis is relatively well studied. Several early researchers detected alkaloids in stem and root material of P. lueddemanniana Rchb.f. (De Wildmann 1892; De Droog 1896; Wester 1921), and in all tissues of P. amabilis (Wester 1921). Many species records for alkaloid content are available in later surveys (Lüning 1964, 1967; Lawler and Slaytor 1969). In summary, Lüning (1974) noted that 17 of 35 species of Phalaenopsis tested had an alkaloid content ≥ 0.1%; comparable

PH ALAENOPSIS

data for a small number of other species were presented under the former generic names—Doritis (1/2), Kingiella (1/1), and Ornithochilus (0/2). Alkaloids isolated by Swedish researchers in the late 1960s and early 1970s comprise cornucervine from P. cornu-cervi (Brandänge et al. 1971), phalaenopsine Is from P. equestris (Brandänge et al. 1972), and phalaenopsines La and T from P. mannii Rchb.f. and P. amabilis, respectively (Brandänge and Lüning 1969). These are esters of 1-hydroxymethylpyrrolizidine diastereoisomers with methyl esters of 2-benzylmalic acid (phalaeonopsines) or 2-isobutylmalic acid (cornucervine). Both of the malic acid derivatives possess the (R)-configuration (Brandänge et al. 1973). The suffixes Is, La, and T as applied to phalaeonopsines refer to isoretronecanol, laburnine, and trachelanthamidine, respectively, which are also known as necine bases (Fig. 672.7). In a wider survey, the distribution of pyrrolizidine alkaloids in 18 species of Phalaenopsis, including P. pulcherrima (as Doritis pulcherrima) and P. taenialis (Lindl.) Christenson & Pradhan (as Kingiella taenialis), was reported (Brandänge et al. 1972). Frölich et al. (2006) observed that phalaeonopsines Is and T were present as a mixture of free base and N-oxide forms in Phalaenopsis hybrids and occurred in all tissues. The highest concentrations were found in young and developing tissues, peripheral tissues, and reproductive organs. Studies using the interspecific hybrid P. equestris × (P. aphrodite Rchb.f. × P. mannii ) reveal that homospermidine synthase, the first pathway-specific enzyme in pyrrolizidine alkaloid biosynthesis, is expressed mainly in aerial root tips and young flower buds (Anke et al. 2008). The evolutionary history of pyrrolizidine alkaloid biosynthesis in Phalaenopsis has been compared with that in other monocotyledons by analyzing cDNA sequences coding for homospermidine synthase and the paralogous deoxyhypusine synthase enzyme, respectively. These data indicate that the ability to synthesize pyrrolizidine alkaloids is an old feature in both lineages (Nurhayati et al. 2009). Information about other types of alkaloid in Phalaenopsis is limited, although a small amount of the pyrrolidine alkaloid hygrine was detected in P. marriottiana (studied as Vandopsis parishii (Veitch & Rchb.f.) Schltr.), using GC-MS (Brandänge and Granelli 1973). The floral fragrance of P. bellina (Rchb.f.) Christenson comprises geraniol (43.0%) and linalool (49.0%) together with geranial

(0.5%), neral (0.4%), and other minor components (Kaiser 2011). A fragrance described as ‘aromatic-floral’ by Kaiser (2011) is produced from the highly scented flowers of P. violacea H.Witte. This is due to a combination of (E)-cinnamaldehyde (3.2%), (E)-cinnamyl acetate (12.3%), (E)-cinnamyl alcohol (4.8%), and methyl salicylate (1.0%), with geraniol (7.2%), geranyl acetate (1.3%), and linalool (27.7%); large amounts of elemicine (25.0%) are also present (Kaiser 2011). The major components of the floral fragrance of P. japonica (Rchb.f.) Kocyan & Schuit. (studied as Sedirea japonica (Rchb.f.) Garay & H.R.Sweet) are nerol (48.8%) and methyl benzoate (12.0%). Other contributions to the scent are made by (E)-2,3-dihydrofarnesol, (E,E)-farnesal, (E,E)-farnesol, geranial, geraniol, methyl salicylate, and neral (Kaiser 2011). (NV, RG)

Phylogenetics For Phalaenopsis (the sole member of clade B), we have accepted a broad circumscription, which includes not only those genera previously thought to have a close relationship to Phalaenopsis (i.e., Doritis, Kingidium, and Nothodoritis) but also Hygrochilus, Ornithochilus, and Sedirea. Topik et al. (2005) did not place Sedirea, but they found Hygrochilus and Ornithochilus closely related to Arachnis, Sarcoglyphis, Chiloschista, Cleisocentron, and one Cleisostoma species, a topology that is likely due to the highly incomplete generic sampling in that study. There are significant differences in floral characters (including column foot, pollinia number, and rostellum structure), and if we can accept this degree of morphological heterogeneity in the case of Phalaenopsis, then many other larger genera can also be created in Aeridinae after more extensive species sampling. (AS, AK, MC)

Ecology Phalaenopsis species are mainly epiphytes at 0–2500 m and occur primarily in three distinct habitats: seasonally dry areas, seasonally cool areas, and constantly warm and humid areas. The plants show adaptations to each of these (Christenson 2001). Species confined to monsoonal areas with a pronounced wet-and-dry cycle have adapted to the stress of the dry season in several

Fig. 672.7.  Pyrrolizidine alkaloids of Phalaenopsis.

239

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ways. These include increased succulence in species such as P. cornu-cervi, in which the roots and leaves are thicker than other species of similar size. There are even unsubstantiated reports that P. cornu-cervi may be semi-deciduous in part of its range. Christenson (2001), however, pointed out that this may not be a direct adaptation to seasonal dryness because the sister species P. pantherina Rchb.f. is recorded from high in the forest canopy, where it is exposed to bright, diffuse light unlike most other species in Borneo that occur toward the base of trees under low-light conditions. Increased succulence in P. cornu-cervi may simply be a response to degree of exposure independent of seasonal dryness. The most extreme form of adaptation in Phalaenopsis native to seasonally dry areas is a deciduous habit, which eliminates excessive transpiration. This is seen primarily in several mainly Himalayan groups of Phalaenopsis, including P. sections Aphyllae and Parishianae. Species adapted to seasonally cool to cold conditions also occur in regions that are seasonally dry. It is difficult to say which adaptation came first because many adaptations apply to both extremes equally. The only truly cool- or cold-growing species belong to P. section Aphyllae. Phalaenopsis taenialis has been recorded as high as 2500 m in China and the Himalayan region, an elevation more typical of cold-growing genera. All these cold-growing species of Phalaenopsis are either deciduous or semi-deciduous in habit, accompanied by a strong dormancy, which affords some protection from cold damage. The majority of species in the genus, belonging to P. ­subgenus Phalaenopsis occur in areas that are uniformly warm and humid throughout the year. Most species occur in evergreen forests without a significant dry season. Within these forests is a mosaic of habitats, and Phalaenopsis species occur in most of them (Christenson 2001). Some species, such as P. gigantea J.J.Sm. and P. pantherina, occur relatively high in the canopy where conditions are exposed. These have leathery leaves to prevent desiccation and will tolerate higher light levels than many. Other species, such as P. fuscata Rchb.f. and P. tetraspis Rchb.f., grow under extremely low light levels. Many others occur in riparian habitats, such as along streams and rivers, which provide constantly high humidity regardless of local fluctuations in rainfall. Phalaenopsis violacea H.Witte occurs in swamps where humidity levels are also high. Species such as P. lowii and P. sumatrana Korth. & Rchb.f. are able to grow either as epiphytes or opportunistically as lithophytes. Christenson (2001) suggested that ‘life on large boulders may provide the plants with a steadier moisture supply or may take advantage of the light gap often associated with rock outcroppings in otherwise continuous forest’. The three species of P. section Esmeralda (formerly placed in the genus Doritis) and some Australian populations of P. amabilis (L.) Blume subsp. rosenstromii (F.M.Bailey) Christenson are unusual in being obligate terrestrials or lithophytes. Christenson (2001) commented that ‘both appear to be derived conditions within the genus, and an epiphytic habit is accepted as the ancestral character for the genus as a whole. In the cases of P. buyssoniana Rchb.f., P. pulcherrima, and P. regnieriana Rchb.f., the terrestrial habit appears to be an adaptation, in part, to an extremely pronounced dry season. Direct access to the soil and accumulated leaf litter presumably provide supplemental water during the dry 240

season. Life on the ground for these three species, which are native to more or less deciduous forests, also presumably provides some shade even when the forest canopy is leafless. These species also show perhaps the strongest tendency for the leaves to turn reddish purple under intense solar radiation or water stress, another defence under extreme conditions’. ( JW)

Pollination Phalaenopsis amabilis is visited by carpenter bees of the genus Xylocopa. Christenson commented that ‘pollination events are frequent and successful in P. amabilis, judging by the high percentage of fruit set in the wild (up to 50 per cent of flowers are pollinated). It is reasonable to assume that similar large bees pollinate other species of section Phalaenopsis with comparable lip and callus structure.’ Christenson (2001) also suggested that ‘most species of subgenus Polychilos are pollinated by a group of bees smaller than the massive Xylocopa. These Phalaenopsis species, such as P. bellina and P. sumatrana, have strongly day-fragrant flowers typical of bee-pollinated flowers. The longevity of the flowers are [sic] in part a result of the infrequent pollination events in nature. Nothing is known of the function of the variably expressed spur in section Aphyllae or the mobile lip of subgenus Parishianae and how these structures relate to pollination’. Jin et al. (2012) investigated the pollination system of P. pulcherrima (as Doritis pulcherrima). Phalaenopsis pulcherrima is fooddeceptive and pollinated by the bee Amegilla nigritar. To prevent pollinator-mediated selfing, many orchid species bring pollinaria into the correct position for pollination by bending the stipe a few minutes after removal. However, in P. pulcherrima pollinaria that have been removed show no modifications of shape, and only the unchanged orientation of the pollinaria on the bee’s body allows successful pollination. ( JW, AK)

Uses Phalaenopsis is currently the most important and popular orchid genus in cultivation, with species of P. sections Phalaenopsis and Esmeralda being particularly valuable in breeding programmes for the houseplant and cut-flower trade. No uses have been reported apart from some former medicinal use of P. aphrodite Rchb.f. and P. schilleriana Rchb.f. in the Philippines (Lawler 1984). (AS)

Cultivation Group 1 (see subtribal treatment). Plants are best grown in small pots or on slabs of cork with some moss and humus around the roots. Most species need high humidity, moderate light levels, and regular watering throughout the year, at the same time preventing sogginess of the medium. Species from continental Asia usually require a distinct cooler and drier resting period in the winter; some shed their leaves in the dry season. Phalaenopsis pulcherrima, as a lithophyte in exposed places, needs more light and less humidity than most other species. Artificial hybrids, which are much more commonly cultivated than the species, are usually more tolerant to drought and low humidity than most of the species. (AS, MM)

PH RAGMORCHIS

Taxonomic notes

Description (Fig. 673.1)

In light of the phylogenetic studies mentioned above, the genus Phalaenopsis appears less uniform than it was long believed to be. Characters thought to be critical for generic delimitation in subtribe Aeridinae, such as number of pollinia, presence or absence of a column foot, motility of the labellum, and presence or absence of a spur, are now known to be variable within some genera, including Phalaenopsis. Vegetative characters are on the whole more stable within genera, and this is true of Phalaenopsis, which may to some extent be recognized by the short stems with few, closely-spaced, relatively broad leaves. However, plants with a similar habit occur in Brachypeza and a few other genera. Among the more dependable floral characters in Phalaenopsis are the trilobed, not fleshy labellum with erect lateral lobes and a relatively large midlobe, almost always with a callus, appendage or crest at the base of the midlobe, and often with additional callosities between and on the lateral lobes. (AS)

Epiphytic herbs. Roots from basal part of stem, shortly papillosehairy. Stem elongate, leafy. Leaves distichous, ascending, angular-terete, tapering, acute, with a sheathing base; leaf-sheaths transversely wrinkled. Inflorescence axillary, opposite each leaf, one-flowered. Flower not opening widely. Sepals free, cucullate, obovate-elliptic. Petals narrowly elliptic, acute, porrect. Labellum adnate to base of column, fleshy, trilobed, side lobes erect, acute in front, midlobe porrect, ovate, calli two, near base of midlobe; spur pendent, cylindrical, slightly bidentate at tip, as long as labellum. Column fleshy, terete, lacking a column foot; anther obovoid, pollinia two, globose, each slightly cleft, stipe spatulate, viscidium elliptic. Ovary and pedicel shorter than sepals and petals, terete. (PC)

Taxonomic literature Christenson, E. A. (2001). Phalaenopsis – A Monograph. Timber Press, Portland, Oregon. Cribb, P. and Schuiteman, A. (2012). [Phalaenopsis] Classification. Renziana, 2, 14–15. Kocyan, A. and Schuiteman, A. (In press). New combinations in Aeridinae. Phytotaxa. Padolina, J., Linder, C. R., and Simpson, B. B. (2005). Phylogeny of Phalaenopsis using multiple chloroplast markers. Selbyana, 26, 155–8. Schuiteman, A. (2012). Nothodoritis zhejiangensis transferred to Phalaenopsis. Renziana, 2, 48–50. Shim, P. S. (1982). A new generic classification in the Phalaenopsis complex (Orchidaceae). Malayan Nature Journal, 36, 1–28; reprinted in 1984 in Malayan Orchid Review, 18, 48–61. Sweet, H. R. (1968). Revision of the genus Phalaenopsis, parts 1 and 2. American Orchid Society Bulletin, 37, 867–77, 1089–104. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84. Tsai, C. C., Chiang, Y. C., Huang, S. C., Chen, C. H., and Chou, C. H. (2010). Molecular phylogeny of Phalaenopsis Blume (Orchidaceae) on the basis of plastid and nuclear DNA. Plant Systematics and Evolution, 288, 77–98. Yukawa, T., Kita, K., Handa, T., and Topik, H. (2005). Molecular phylogenetics of Phalaenopsis (Orchidaceae) and allied genera: re-evaluation of generic concepts. Acta Phytotaxonomica et Geobotanica, 56, 141–61.

673.   PH R AG MO RC H IS Phragmorchis L.O.Williams, Bot. Mus. Leafl., Harvard Univ., 6, 52 (1938). Type: Phragmorchis teretifolia L.O.Williams

Derivation of name From phragmites, reed, and orchis, orchid, in allusion to the vaguely reed-like plant habit of this orchid. (AS)

Distribution (Fig. 673.2) Phragmorchis is monospecific and endemic to the Philippines (Luzon). (PC)

Cytogenetics Chromosome numbers for P. teretifolia have not been published. (AP)

Phytochemistry No reports have been found on phytochemistry of Phragmorchis. (NV, RG)

Phylogenetics There is no phylogenetic information available for this genus. (AK)

Ecology Nothing is known of the habitat of the species. (AS)

Pollination Pollination of P. teretifolia has not been studied. (AP)

Uses No uses have been reported for Phragmorchis; it is not known to be in cultivation. (AS)

Taxonomic notes According to Williams (1938), Phragmorchis is allied to Schoenorchis. It has not been included in any morphological or DNA-based analysis of Aeridinae. In its papillose roots, fleshy leaves, wrinkled leaf-sheaths, and overall floral morphology, Phragmorchis teretifolia (known only from the type) resembles Omoea philippinensis Ames. If fresh material of Phragmorchis becomes available, a critical comparison may show that they are not generically distinct. A record by Christenson (1992) of Phragmorchis teretifolia from the 241

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C

D

G

F

A E

B

Fig. 673.1.  Phragmorchis teretifolia L.O.Williams. A. Habit; B. Stem, close-up; C. Flower; D. Column and labellum, side view; E. Column, anther cap removed; F. Anther cap; G. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Loher 14744 (K) and redrawn after G. W. Dillon, Botanical Museum Leaflets, Harvard University, 6, 55 (1938).

island of Sumbawa in Indonesia is based on a misidentification; the specimen in question represents a species of Thrixspermum, possibly the little known T. angustifolium (Blume) Rchb.f. (AS, PC)

Williams, L. O. (1938). A new genus of the Sarcanthinae. Botanical Museum Leaflets, Harvard University, 6, 52–7.

Taxonomic literature

Plectorrhiza Dockrill, Austral. Sarcanth., 27, t. 14 (1967). Type species: Plectorrhiza tridentata (Lindl.) Dockrill (basionym: Cleisostoma tridentata Lindl.)

Christenson, E. A. (1992). Notes on Asiatic orchids. Lindleyana, 7, 88–94.

242

6 7 4 .  P L EC T OR R H I ZA

PLECTORRHIZA

Fig. 673.2.  Distribution map of Phragmorchis.

Derivation of name

Distribution (Fig. 674.2)

From the Greek plektos, twisted, and rhiza, root, referring to the tangled roots. ( JW)

The three species of Plectorrhiza are distributed in temperate and tropical eastern Australia (northern Queensland to eastern Victoria), with P. erecta (Fitzg.) Dockrill endemic to Lord Howe Island. ( JW, PA)

Description (Plate 97; Fig. 674.1) Epiphytic or occasionally lithophytic, rarely terrestrial herbs. Roots many, often tangled. Stem pendent or erect. Leaves oblong, narrowly ovate, elliptic or narrowly ovate, obtuse, acute or acuminate, sometimes unequally emarginate, persistent. Inflorescence racemose, usually few-flowered; peduncle and rachis wiry; floral bracts minute. Flowers resupinate, lasting longer than two days, strongly fragrant. Sepals and petals free, similar in length, spreading. Labellum rigidly joined to apex of column foot, either in line with foot or at a right angle to it, saccate, spurred, trilobed; disc not readily differentiated from spur and midlobe, side lobes not in contact with column or column foot, variously shaped, midlobe not readily differentiated from front wall of spur, appearing to be an extension of it, variously shaped; spur either subcylindical and curved slightly forward or hemispherical, a digitiform or claviform callus covered with trichomes arising from its distal side close to orifice and directed toward its apex. Column without wings, column foot less than half the length of column proper, not readily differentiated from it; anther cap shallow, pollinia four, in two unequal, closely appressed pairs, stipe longer than wide, viscidium set at an acute angle to stipe; rostellum beaked, porrect or slightly upcurved, apex bifid. ( JW)

Cytogenetics No chromosome numbers have been recorded for Plectorrhiza. ( JW)

Phytochemistry A survey of alkaloid distribution in Orchidaceae gave a negative result for P. tridentata (Lawler and Slaytor 1969). (NV, RG)

Phylogenetics There is no phylogenetic information available for this genus. (AK)

Ecology Two of the species are epiphytes at 0–1300 m. Plectorrhiza brevilabris (F.Muell.) Dockrill is widespread in Queensland and northern New South Wales in rain forest on trees and small shrubs (Jones 2006). Plectorrhiza tridentata (Lindl.) Dockrill is common along almost the entire eastern Australian coast in 243

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J C

I A

L

B D

F

H E

K

G Fig. 674.1.  Plectorrhiza brevilabris (F.Muell.) Dockrill. A. Habit; B. Inflorescence; C. Flower; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum; H. Column and labellum, side view; I. Column, anther cap removed, front view; J. Column, anther cap removed, side view; K. Pollinarium (no information about size); L. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Dockrill s.n.23/12/1962 and Kew Spirit Collection no. 30984; pollinia redrawn after W. H. Nicholls, Orchids of Australia, pl. 471 (1969).

humid rainforest, on tree branches and outer twigs, especially in deep gullies and along creeks and in swamps. Large plants are often suspended by a large mass of tangled roots. There are occasional natural hybrids with Sarcochilus species (Peter Adams, unpublished). The third species, Plectorrhiza erecta (Fitzg.) Dockrill, grows on rocks or occasionally as a terrestrial or epiphyte in dense shade at 0–800 m on Lord Howe Island. (PA, JW)

Pollination There is no information available on pollination of Plectorrhiza. ( JW) 244

Uses Plectorrhiza tridentata is common in cultivation in Australia and used in hybridization to produce small plants with brightly coloured flowers; some intergeneric miniature hybrids have been produced using P. erecta. Other species are grown occasionally. (PA)

Cultivation Group 1 (see subtribal treatment). Plants should be grown as for temperate twig epiphytes in the genus Sarcochilus, with high humidity and brisk air movement. (PA, MM)

POMATOCALPA

Fig. 674.2.  Distribution map of Plectorrhiza.

Taxonomic literature Jones, D. L. (2006). A complete guide to native orchids of Australia including the island territories. Reed New Holland, Sydney. Riley, J. J. and Banks, D. P. (2002). Orchids of Australia. Princeton University Press.

675.   PO MATO C A LPA Pomatocalpa Breda, Gen. Sp. Orch. Asclep., Fasc. 3, t. 15 (1829). Type species: Pomatocalpa spicatum Breda

Derivation of name From the Greek poma (-atos), drinking, and kalpis, pitcher, in reference to the flask-shaped labellum. ( JW)

Description (Plates 98, 99; Fig. 675.1) Rambling (up to 5 m long), epiphytic, occasionally lithophytic herbs. Stem pendent or ascending, sometimes clambering, simple, with a few to many nodes covered by leaf-sheaths. Leaves distichous, linear to oblong, sometimes falcate, usually unequally bilobed or retuse, flat, coriaceous, articulate to a sheathing base. Inflorescences lateral, penetrating the leaf-sheaths, racemose or paniculate, with the flowers clustered in the distal portion of each racemose branch, erect or pendent, many-flowered; peduncle shorter or longer than rachis (or the terminal rachis branch in panicles), glabrous, minutely papillose or distinctly

pubescent; rachis with flowers opening from base of inflorescence; floral bracts persistent, triangular, acute to acuminate, erose. Flowers resupinate or not, usually widely opening, rarely scented. Sepals and petals free, similar, spreading. Labellum firmly attached to column, trilobed, shortly spurred, side lobes erect, deltoid or triangular, the back edge of each lobe usually producing a right angle to the front edge, midlobe fleshy, often recurved, usually suborbicular or ovate-triangular; spur bucket-shaped, front wall thicker than back wall, back wall with a projecting erect, rectangular, bifid to subtruncate, ligulate appendage reaching or exserted beyond the entrance to spur. Column erect or slightly recurved, column foot absent; anther cap shortly beaked, pollinia four, arranged as two unequal pairs, waxy, semi-globular, solid, with a long common stipe and a single, usually concave viscidium; rostellum hammer-shaped, bifid after removal of the pollinarium, shorter than diameter of column. ( JW)

Distribution (Fig. 675.2) The 13 species of Pomatocalpa are collectively distributed from India and Sri Lanka through Malesia and northern Australia to Fiji, north to northeastern Himalayas and China (Hainan and Taiwan). (JW)

Cytogenetics Brandham (1999) and Felix and Guerra (2010) listed a count of 2n = 38 for the genus. ( JW) 245

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E

F 1 mm

I

H

3 mm

A

10 cm

3 mm

G

D

3 mm

C

B 5 mm

Fig. 675.1.  Pomatocalpa kunstleri (Hook.f.) J.J.Sm. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum and column, longitudinal section; H. Anther cap, dorsal view; I. Pollinarium, front and back views. Drawn by Chan Chew Lun and Liew Fui Ling from Lamb AL 1153/89. Reproduced with permission from J. J. Wood (2003), Orchids of Borneo. Volume 4. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

Phytochemistry

Ecology

Low levels of alkaloids (0.01%) are found in P. marsupiale (Kränzl.) J.J.Sm., but P. macphersonii (F.Muell.) T.E.Hunt does not contain them (Lawler and Slaytor 1969, 1970). According to Lüning (1974), one of eight species of Pomatocalpa tested for alkaloids contained them at levels ≥ 0.1%, although species names were not given. (NV, RG)

All species are epiphytes or lithophytes at 0–750 m in swamp and coastal rocky forests, mangrove and beach forests, and dipterocarp forests. ( JW)

Phylogenetics Pomatocalpa is well supported as a member of clade H, within which it is sister to Gastrochilus, a result also found by Topik et al. (2005). A phylogenetic study by Watthana (2007) found that Gastrochilus retrocalla is nested within Pomatocalpa, which was not found here, although bootstrap support for monophyly of Gastrochilus is not strong. (AK) 246

Pollination Watthana (2007) observed that the flowers seem to fit the beepollination syndrome of van der Pijl & Dodson (1966). In Australia, Jones (1981) observed a small bee (Trigona sp.) with the dark pollinarium of P. macphersonii attached to its head. ( JW)

Uses No uses have been reported for Pomatocalpa; it is occasionally cultivated. ( JW)

PORRORHACHIS

Fig. 675.2.  Distribution map of Pomatocalpa.

Cultivation Group 1 (see subtribal treatment). (MM)

Taxonomic literature Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84. Watthana, S. (2007). The genus Pomatocalpa (Orchidaceae): a taxonomic monograph. Harvard Papers in Botany, 11, 207–56.

676.   PO R RO R H AC H IS Porrorhachis Garay, Bot. Mus. Leafl., Harvard Univ., 23, 191 (1972). Type species: Porrorhachis galbina (J.J.Sm.) Garay (basionym: Saccolabium galbinum J.J.Sm.)

rooting at base. Leaves conduplicate, articulate, distichous, linear, apex obtuse, apiculate. Inflorescence lateral, solitary, racemose, patent, many-flowered; rachis continuing the line of the peduncle, straight, elongate, glabrous, thicker than peduncle. Flowers resupinate about 0.4–0.6 cm wide, opening in succession, one or a few at a time, quaquaversal, green or ochre-yellow. Sepals entire, free, obtuse, lateral sepals strongly oblique. Petals shorter than sepals. Labellum rigidly attached, subentire, saccate, fleshy at apex. Column with foot hardly developed; pollinia two, spherical, stipe suborbicular, about as long as the pollinia. Capsule longer than unfertilized ovary, narrowly fusiform, almost ­rostrate. (AS)

Distribution (Fig. 676.2) Porrorhachis is a genus of two species—P. galbina and P. macrosepala (Schltr.) Garay—from Java, Borneo, and Sulawesi. (AS)

Derivation of name

Cytogenetics

From the Greek porrho, forward, and rhachis, rachis, referring to the straight inflorescence. (AS)

There are no published chromosome counts for these species. (AP)

Description (Plate 100; Fig. 676.1)

Phytochemistry

Epiphytic herbs. Roots terete, not branching, glabrous, smooth. Stem elongate, leafy, pendent or patent-ascending to erect,

No reports have been found on phytochemistry of Porrorhachis. (NV, RG) 247

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Fig. 676.1.  Porrorhachis galbina (J.J.Sm.) Garay. A. Habit; B. Flower, front view; C. Flower, oblique view; D. Ovary, transverse section; E. Rachis and flower (dorsal sepal, lateral sepal, petal, and anther cap removed), side view; F. Labellum and column, with lateral sepal and petal attached, longitudinal section; G. Dorsal sepal; H. Lateral sepal; I. Petal; J. Anther cap, dorsal view; K. Pollinarium. Drawn by C. L. Chan and Lucy F. L. Liew from Lamb AL 1133/89. Reproduced with permission from J. J. Wood (1997), Orchids of Borneo. Volume 3. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

Phylogenetics

Uses

Bispecific Porrorhachis is embedded in Trachoma (clade E), which is an indication that the two genera should be combined; however, bootstrap support for this relationship is weak. Before making this move, more sequencing of other species of the latter are needed. Garay (1972) segregated Porrorhachis from Saccolabium based on the differences in the column, rostellum, and habit. (AK, MC)

No uses have been reported for Porrorhachis; it is rare in cultivation. (AS)

Ecology

Taxonomic notes

Both species are epiphytes in mossy forests and scrub from 1100 to 1700 m. (AS)

The generic delimitation of the genera closely allied to Pteroceras, including Porrorhachis, needs further study. Porrorhachis is only tentatively accepted here. It may be distinguished from Trachoma by the lax, elongate rachis of the inflorescence, with the flowers not opening in clusters. (AS)

Pollination Nothing is known about pollination in Porrorhachis. (AS) 248

Cultivation Group 2 (see subtribal treatment). (MM)

PTEROCERAS

Fig. 676.2.  Distribution map of Porrorhachis.

Taxonomic literature

Derivation of name

Garay, L. A. (1972). On the origin of the Orchidaceae. II. Journal of the Arnold Arboretum, 53, 202–15.

From the Greek pteron, wing, and keras, horn, in reference to the two narrow, wing-like appendages at the base of the labellum. ( JW)

677.   PTE RO C E R A S Pteroceras Hassk., Flora, Jena, 25(2), Beibl., 6 (1842). Type species: Pteroceras radicans Hassk. Ornitharium Lindl. & Paxton, Paxt. Fl. Gard., 1, 188 (1851). Type species: Ornitharium striatulum Lindl. & Paxton Proteroceras J.Joseph & Vajr., J. Indian Bot., 53, 189 (1974), nom. illeg. Type species: Proteroceras holttumii J.Joseph & Vajr. Taxa at sectional rank referable to Pteroceras: Dendrocolla Blume sect. Tubera Blume, Bijdr., 288 (1825). Type species: not designated. Aerides Lour. sect. Tubera (Blume) Lindl., Gen. Sp. Orchid. Pl., 238 (1833). Type species: not designated. Sarcochilus R.Br. sect. Tubera Benth. in Benth. & Hook.f., Gen. Plant., 3, 575 (1883), p.p., nom. illeg. Type species: not designated. Sarcochilus R.Br. sect. Pteroceras (Hassk.) Hook.f., Fl. Brit. Ind., 6, 33 (1890), p.p. Sarcochilus R.Br. sect. Eusarcochilus Schltr., Repert. Spec. Nov. Regni Veg., Beih., 1, 964 (1913), p.p., nom. illeg. Type species: not designated.

Description (Plate 101; Fig. 677.1) Epiphytic or rarely lithophytic herbs. Stem producing leaves from apex while gradually dying from its base, terete, glabrous, erect to pendent, usually unbranched. Leaves sessile, articulate, up to c. 17 per flowering shoot, distichously arranged, spreading, conduplicate, coriaceous, rarely semi-terete, linear to narrowly elliptic, unequally bilobed to acute. Inflorescence racemose, rarely oneflowered with the only flower being terminal, up to 30 per shoot; peduncle sub-terete, glabrous to prickly; rachis (if present) (sub) terete or laterally compressed, glabrous to prickly, sometimes dilated at floral nodes into semi-lanceolate wings or keels; floral bracts persistent, shorter than ovary. Flowers to about 1.5 cm across, variously coloured, usually sweetly scented, up to c. 190 per rachis, spirally or distichously arranged, usually resupinate, usually developing successively. Sepals and petals free, spreading, often concave and/or carinate, truncate to acute, sometimes prickly on exterior; lateral sepals adnate to column foot for at least some distance. Labellum sessile or nearly so, continuing the line of the column foot to which it is movably hinged, trilobed, central part of blade developed into a porrect spur, side lobes 249

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Fig. 677.1.  Pteroceras teres (Blume) Holttum. A. Habit; B. Flower, side view; C. Ovary, labellum, and column, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum, front view; H. Column, side view; I. Column apex with anther cap, front view; J. Column apex with anther cap, oblique view; K. Anther cap, ventral view; L. Pollinarium, front and side views. Drawn by Susanna Stuart-Smith from Lamb AL 1163/89. Reproduced with permission from N. R. Pearce and P. J. Cribb (2002), The Orchids of Bhutan. Royal Botanic Garden, Edinburgh and Royal Government of Bhutan.

erect, sometimes partly embracing column, midlobe strongly modified, appearing as a ridge-, crest-, cone-, or saddle-shaped callus, a tiny tooth, or simply a slight thickening of upper wall; up to three small calli (or a high transverse wall) present above spur entrance, between bases of side lobes and midlobe; spur sac-like to elongate, outer surface smooth to pustulate, glabrous to hairy, inner surface never developed into calli but sometimes hairy; surface of all parts of labellum finely rugose-papillose in some species. Column continuing the line of ovary, subterete to clavate, column foot distinct, sometimes finely rugose-papillose; anther terminal, strongly incumbent, pollinia two, sulcate (very 250

rarely four, entire, closely appressed in two pairs), stipe longer than viscidium, viscidium variously shaped; stigmatic cavity deep; rostellum deeply bifid after removal of pollinarium. Ovary glabrous to prickly. Capsule cylindric to winged-triangular, often with a short beak. ( JW)

Distribution (Fig. 677.2) Pteroceras presently comprises 27 species distributed from India and Sri Lanka to Maluku. The centre of distribution is Borneo. ( JW)

PTEROCERAS

Fig. 677.2.  Distribution map of Pteroceras.

Cytogenetics According to Löve and Solbrig (1965), Pancho reported 2n = 38 in P. longicalcarum (Ames & Rolfe) Garay and P. teres (Blume) Holttum. ( JW)

Phytochemistry Alkaloids are present in P. pallidum (Blume) Holttum (as Sarcochilus pallidus (Blume) Rchb.f.) at levels > 0.1% (Lüning 1967). (NV, RG)

Phylogenetics Pedersen (1993) revised Pteroceras and summarized its problematic taxonomic history. He noted the possibility that Pteroceras might not be monophyletic, given the characters that were studied and problems in defining floral characters (long column foot, fleshy lip with a spur, short rostellum, and short-lived flowers) that are exhibited by other genera. In an unpublished molecular study of Pteroceras, Kocyan et al. demonstrated that the genus needs extensive reorganization. As a first step, some species of Pteroceras have been moved to Brachypeza (clade C). However, other Pteroceras species, including the type (P. teres) are members of clade E; before further generic alterations of this clade are undertaken, more species need to be sequenced. (AK, MC)

Ecology Pedersen (1993) noted that most species of Pteroceras seem to be lowland plants with an upper elevational limit of 1000 m.

Pteroceras species are typically found in evergreen rain forests (lowland, hill, and upper montane) but are also recorded from mangrove swamps, scrub, rocky seashores, and isolated trees in open fields. ( JW)

Pollination No information has been published on pollination of Pteroceras, but they all seem to exhibit the syndrome of melittophilous flowers (Pedersen 1993). The flowers are zygomorphic and usually sweetly scented, diurnal, horizontal, white or variously coloured (but never pure red). The labellum provides a prominent landing platform, and its side lobes are often provided with nectar guides. It seems evident that the erect labellum side lobes, together with the elastic junction of the labellum to the column foot, ensure that insects visiting the flower will assume a position and behaviour favourable to pollination (Pedersen 1993). ( JW)

Uses No uses have been reported for Pteroceras; some species are sometimes cultivated. ( JW)

Cultivation Group 1 (see subtribal treatment). Most species can be grown under warm to intermediate conditions with high humidity and moderate light levels in small pots with a well-draining medium, or else on slabs of cork with some moss and humus around the 251

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roots. Plants should be misted frequently but allowed to dry out between waterings. (AS, MM)

Taxonomic notes Pedersen (1993) included Pteroceras within a discrete group of genera having a column foot and two sulcate pollinia. Other genera in this group are Brachypeza, Dimorphorchis, and Macropodanthus. He pointed out that ‘this group of genera may turn out to be highly artificial – but it is, nevertheless, convenient for purposes of determination’. Phylogenetic analyses by Kocyan (see subtribal treatment) have shown that this is an artificial group. Pteroceras as treated by Pedersen appears to be polyphyletic, with the type species more closely related to such genera as Tuberolabium and Macropodanthus, whereas P. pallidum and P. semiteretifolium H.A.Pedersen belong to a different clade with Brachypeza. Florally, the two clades are remarkably similar, but they can be separated morphologically on the basis of a character of the rachis. In Pteroceras s.s. the pedicel is inserted at the base of a sharply delimited, almost winged, concavity in the rachis, whereas in Brachypeza s.l. (including species formerly treated as Pteroceras) there is no such concavity with winged margins. (JW, AS)

distichous, often oblong, unequally bilobed, coriaceous, conduplicate, articulated to a sheathing base. Inflorescence lateral, paniculate, occasionally racemose, many-flowered. Flowers resupinate or not, widely opening, flat, predominantly bright red or orange, sometimes yellow with red markings or bright purple. Sepals and petals free, dorsal sepal and petals similar, spreading, lateral sepals usually broader than dorsal sepal and petals, often clawed, margins undulate, subparallel. Labellum rigidly attached to column base, sac-shaped, usually much smaller than sepals and petals, trilobed, saccate or spurred, without interior callosities, side lobes erect, adaxial surface with a basal callus, midlobe often ligulate and recurved, with lamellate basal calli; spur conical. Column without foot; pollinia two or four, about equal or slightly unequal, waxy, reniform, solid, each pair with an elastic thread at base, attached by a common linear stipe to a transversely elliptic viscidium. ( JW)

Distribution (Fig. 678.2) Renanthera comprises 20 species distributed from eastern India (Assam) through China to the Philippines and south to Malaysia, Indonesia, New Guinea, and the Solomon Islands. ( JW)

Cytogenetics Taxonomic literature Pedersen, H. Æ. (1993). The genus Pteroceras - a taxonomic revision. Opera Botanica, 117, 1–64.

678.  R E N A N TH E R A Renanthera Lour., Fl. Cochinch., 516 (1790). Type species: Renanthera coccinea Lour. Nephranthera Hassk., Tijdschr. Natuurl. Gesch. Physiol., 9, 145 (1842). Type species: Nephranthera matutina (Poir.) Hassk. (basionym: Epidendrum matutinum Poir.) Renantherella Ridl., J. Linn. Soc., Bot., 32, 354 (1896). Type species: Renantherella histrionica (Rchb.f.) Ridl. (basionym: Renanthera histrionica Rchb.f.) Ascoglossum Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 974 (1913), syn. nov. Type species: Ascoglossum calopterum (Rchb.f.) Schltr. (basionym: Saccolabium calopterum Rchb.f.) Porphyrodesme Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 982 (1913), syn. nov. Type species: Porphyrodesme papuana Schltr. (basionym: Saccolabium porphyrodesme Schltr.)

Derivation of name From the Latin ren, kidney, and anthera, anther, in reference to the kidney-shaped pollinia of the type species. ( JW)

Description (Plate 102; Fig. 678.1) Epiphytic or lithophytic (rarely terrestrial) herbs. Stem elongate, often up to several metres in length, often climbing, sometimes branched, with many nodes and long internodes. Leaves many, 252

Brandham (1999) listed published chromosome counts for the genus as 2n = 38, 76, 114, 115 with a basic number of x = 19, yielding 2x = 38 (the most common), 4x = 76, and 6x = 114. He noted that this is one of the few series with no major deviations except at the hexaploid level. (AP)

Phytochemistry Flavone C-glycosides occur in the leaves of R. imschootiana Rolfe (C. Williams 1979). Preliminary results on the anthocyanins of the blue and purple flowers of Renanthera hybrids R. Lena Rowell, R. Meda Arnold, and R. Oricil indicate that the major pigment (‘Vanda A’) is an acylated 3-glycoside of cyanidin comprising arabinosyl and glucosyl residues (Lowry and Keong 1973). Several species have been tested for their alkaloid content (Lüning 1974), with the highest levels (> 0.1%) found in R. storiei Rchb.f. (Lüning 1967). Other species giving positive or weakly positive results are R. coccinea, R. edelfeldtii F.Muell. & Kränzl., and R. monachica Ames (Lüning 1964, 1967; Lawler and Slaytor 1970). Records are also available for two unnamed species under the former generic names Ascoglossum and Porphyrodesme; neither contained alkaloids at levels ≥ 0.1% (Lüning 1974). (NV, RG)

Phylogenetics Renanthera (clade G) is the oldest generic name in subtribe Aeridinae, but few combinations have ever been made within it. Three genera that have always been thought to be closely allied, Ascoglossum, Porphyrodesme, and Renantherella, have been sunk in Renanthera s.l. The other genera included in clade G are Aerides and Arachnis. (AK, MC)

RHINERRHIZA

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Fig. 678.1.  Renanthera bella J.J.Wood. A. Plant; B. Flower, front view; C. Column and labellum, longitudinal section; D. Column and labellum, side view; E. Anther cap; F. Pollinarium, side view; G. Pollinarium, front view; H. Ovary, transverse section. Drawn by Shim Phyau Soon from cultivated material at Tenom Orchid Centre, Sabah. Reproduced with permission from Chan et al. (1994), Orchids of Borneo. Volume 1. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

Ecology

Cultivation

Species of Renanthera are epiphytes or lithophytes in forests at 0–1400 m. Renanthera citrina Aver. also occurs as a terrestrial in grassland and scrub at 400–800 m in Sumatra. ( JW)

Group 3 (see subtribal treatment). (MM)

Pollination No information on pollination of Renanthera is available. ( JW)

Uses Lawler (1984) reported the (former) use of the pickled or salted young leaves of R. moluccana Blume for flavouring on the island of Ambon in the Moluccas. The genus is popular in cultivation and important in hybridization programmes. ( JW, AS)

Taxonomic literature Kocyan, A. and Schuiteman, A. (In press). New combinations in Aeridinae. Phytotaxa. Rice, R. (2003). A preliminary checklist of the genus Renanthera (Orchidaceae) and its distribution. Orchids Australia, 15, 48–52. Rice, R. (2005). Notes on the genus Renanthera (Orchidaceae). Oasis the Journal, Supplement, 4, 11. Rice, R. (2008). Renanthera of Borneo. Malesian Orchid Journal, 1, 71–6.

6 7 9 .   R H I NER R H I ZA Rhinerrhiza Rupp, Victorian Naturalist, 67, 206 (1951). Type species: Rhinerrhiza divitiflora (F.Muell. ex Benth.) Rupp (basionym: Sarcochilus divitiflorus F.Muell. ex Benth.) 253

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Fig. 678.2.  Distribution map of Renanthera.

Derivation of name From the Greek rhine, file, rasp, and rhiza, root, in reference to the scabrid, rasp-like roots of the single known species. ( JW)

Description (Plate 103; Fig. 679.1) Epiphytic herbs. Roots scabrous, rasp-like, broad, flat, grey. Stem usually a single growth, occasionally up to 10 cm long, erect, horizontal or pendent, the portion below the leaves covered with the imbricate scarious remains of the leaf-sheaths. Leaves 3–6, oblong or narrowly oblong, apex acute or oblique, margins sometimes undulate, coriaceous, smooth or harsh and dry to the touch, sometimes strongly suffused with purple. Inflorescence many-flowered, pendent, the flowers opening in one or a few episodes, simultaneously or in groups, fugaceous; peduncle shorter than rachis; floral bracts minute. Flowers resupinate, short-lived (1–3 days), usually opening widely, pale orange with red spots and blotches, labellum white, or ochre-yellow irregularly and heavily blotched and spotted brown. Sepals and petals free, subsimilar and usually of equal length, spatulate or almost filiform. Labellum hinged to apex of column foot, trilobed, spurred, side lobes erect, incurved and overlapping each other at distal end but not enclosing column, midlobe situated at top of base of spur, tooth-like, disc with three calli, a central centrally grooved one and a tooth-like one at base of and joined to each side lobe opposite central callus; spur situated at distal 254

end of labellum, hollow, obtuse, broader than deep. Column with foot set at an acute angle with column proper; anther cap with a beak, pollinia four, in two closely appressed subequal pairs sessile on stipe, viscidium oblong or subobovate; stigma deeply set; rostellum composed of two widely separated, decurved, toothlike arms. ( JW)

Distribution (Fig. 679.2) Rhinerrhiza is monospecific and confined to Australia (northeastern Queensland, northern New South Wales). ( JW)

Cytogenetics No chromosome numbers have been recorded for Rhinerrhiza. ( JW)

Phytochemistry Alkaloids are present in R. divitiflora at levels > 0.1% (Lawler and Slaytor 1969). (NV, RG)

Phylogenetics Rhinerrhiza is a member of the Australia clade. Topik et al. (2005) found a similar grouping that included, among others, Sarcochilus and Bogoria. (AK, MC)

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Fig. 679.1.  Rhinerrhiza divitiflora (F.Muell.ex Benth.) Rupp. A. Habit; B. Inflorescence; C. Flower; D. Bract; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum, from above; I. Labellum, from below; J. Labellum, longitudinal section; K. Column and labellum, side view; L. Column, anther cap removed, side view; M. Anther cap, dorsal view; N. Anther cap with pollinia, ventral view; O. Pollinarium; P. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Weinthal s.n. Aug. 1928 (K) and Kew Spirit Collection no. 24732.

Ecology Rhinerrhiza divitiflora is epiphytic on tree trunks or occasionally lithophytic, common and widespread in rain forests, open forests, and sheltered areas at 0–1200 m. (PA, JW)

Pollination There is no information available on pollination of Rhinerrhiza. ( JW)

often with Sarcochilus to provide longer-lasting flowers. It is occasionally cultivated. (SL, PA)

Cultivation Group 1 (see subtribal treatment). Plants should be grown as for Sarcochilus, with moderate shading and regular watering throughout the year. (PA, MM)

Uses

Taxonomic literature

No uses have been reported for Rhinerrhiza except in horticulture, where it imparts its yellow colour in intergeneric crosses,

Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK

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Fig. 679.2.  Distribution map of Rhinerrhiza.

and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

680.  R H IN E R R H IZ O P S IS Rhinerrhizopsis Ormerod, Oasis, Suppl., 1, 3 (2001). Type species: Rhinerrhizopsis moorei (Rchb.f.) Ormerod (basionym: Thrixspermum moorei Rchb.f.)

Derivation of name From the orchid genus Rhinerrhiza, and the Greek opsis, likeness. (AS)

Description (Plate 104; Fig. 680.1) Epiphytic herbs. Roots terete, not branching, glabrous, smooth. Stem patent, rooting at base. Leaves conduplicate, articulate, distichous, oblong, twisted at base so as to lie in one plane with stem, apex oblique. Inflorescence lateral, solitary, racemose, many-flowered; peduncle pendent or erect; rachis continuing the line of peduncle, straight, not thicker than peduncle. Flowers resupinate, c. 2 cm wide, lasting less than a day, opening all simultaneously, quaquaversal, fragrant, sepals and petals ochre-yellow, usually with red-brown spots, labellum creamy white with reddish and orange or red-brown markings. Sepals entire, free, broadly ovatespatulate, rounded. Petals similar to but narrower than sepals. 256

Labellum hinged with apex of column foot, spurred-saccate, trilobed, lateral lobes rectangular, erect; spur conical, obtuse, continuing the line of column foot. Column with foot about as long as column; pollinia four, round, in two pairs, stipe about as long as pollinia, linear-spatulate. (AS)

Distribution (Fig. 680.2) Rhinerrhizopsis includes three species from New Guinea, Australia, and the Solomon Islands. (AS)

Cytogenetics There are no published chromosome counts for this species. (AP)

Phytochemistry Alkaloids have been detected in R. moorei (as Sarcochilus moorei Rchb.f.), but only at levels of < 0.01% (Lawler and Slaytor 1969). (NV, RG)

Phylogenetics Rhinerrhizopsis is well supported as sister to one species of Gunnarella (another falls in clade E), and this pair is sister to Dryadorchis in the New Guinea clade. (AK, MC)

RH INERRHIZOPSIS

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Fig. 680.1.  Rhinerrhizopsis moorei (Rchb.f.) Ormerod. A. Habit, ×0.67; B. Flower, ×2; C. Column, ×8; D. Anther cap, dorsal and ventral views, ×8; E. Pollinia, ×8; F. Labellum, side view, ×6; G. Labellum, 6; H. Labellum, transverse section ×6; I. Dorsal sepal, ×4; J. Petal, ×4; K. Lateral sepal, ×4. All magnifications as originally published. Drawn as Rhinerrhiza moorei (Rchb.f.) M.A.Clem., B.Wallace & D.L.Jones by Sue Wickison from Wickison 68. Reproduced with permission from B. A. Lewis and P. J. Cribb (1991), Orchids of the Solomon Islands and Bougainville. Royal Botanic Gardens, Kew.

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Fig. 680.2.  Distribution map of Rhinerrhizopsis.

Ecology

Derivation of name

Rhinerrhizopsis moorei is epiphytic in rain forests, beach and mangrove forests, often along streams, at 0–600 m. (AS)

From the Greek rhynchos, snout, and gyne, female, referring to the beak-like rostellum. (AS)

Pollination Jones (1981; as Sarcochilus moorei (Rchb.f.) Schltr.) reported pollination by ‘small bees of the genus Carbonaria’ (Trigona sp.) in Australia. He noted that a bee will probe the yellow mass on the labellum and farther down in the sac. Within a minute the bee backs out, usually bearing a pollinarium, and then spends several minutes cleaning its antennae, mouth parts, head, and front legs. (AS)

Uses No uses have been reported for Rhinerrhizopsis; it is rare in cultivation. (AS)

Cultivation Group 1 (see subtribal treatment). (MM)

681.  R H Y N C H O G Y N A Rhynchogyna Seidenf. & Garay, Bot. Tidsskr., 68, 88 (1973). Type species: Rhynchogyna luisifolia (Ridl.) Seidenf. & Garay (basionym: Saccolabium luisifolium Ridl.) 258

Description (Fig. 681.1) Epiphytic herbs. Roots terete, not or sparsely branching, glabrous, smooth. Stem elongate, pendent to ascending, rooting in basal part. Leaves articulate, distichous, terete. Inflorescence lateral, solitary, paniculate, pendent, many-flowered; rachis continuing the line of peduncle, straight, glabrous. Flowers resupinate on a pendent inflorescence, about 0.6–1.0 cm wide, many opening simultaneously, quaquaversal, sepals and petals yellowish or brownish, labellum partly pinkish purple. Sepals entire, free, obtuse. Petals shorter than sepals. Labellum rigidly attached to column foot, spurred, trilobed; spur saccate to cylindrical, almost closed at mouth by a callus at back wall opposite a swelling below midlobe, interior without aseptum. Column with a well-developed column foot; pollinia four, compressed in two unequal pairs, those of a pair unequal, stipe linear-spatulate, longer than pollinia. (AS)

Distribution (Fig. 681.2) Rhynchogyna is a genus of three species from Thailand and Peninsular Malaysia. (AS)

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I Fig. 681.1.  Rhynchogyna luisifolia (Ridl.) Seidenf.& Garay. A. Habit; B. Portion of inflorescence; C. Flower, side view; D. Dorsal sepal, front view; E. Dorsal sepal, side view; F. Lateral sepal; G. Petal; H. Labellum, from above; I. Labellum, longitudinal section; J. Column and labellum, side view; K. Column, anther cap removed, side view; L. Anther cap, dorsal view; M. Anther cap, ventral view; N. Pollinarium. Single bar = 1 mm; double bar = 1 cm. Drawn by Judi Stone from Flora of Siam 3210 (K).

Cytogenetics There are no published chromosome counts for these species. (AP)

before alterations to the taxonomy of this genus can be contemplated Cleisostoma needs to be much better understood. More study is required to clarify generic boundaries in clade L. (AK, MC)

Phytochemistry

Ecology

No reports have been found on phytochemistry of Rhynchogyna. (NV, RG)

All three species are epiphytes in lowland forests at 200–700 m. (AS)

Phylogenetics This genus is clearly not monophyletic (clade L). Rhynchogyna fallax (Guillamin) Seidenf. and R. luisifolia are not sister species, but

Pollination Nothing is known about pollination in Rhynchogyna. (AS) 259

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Fig. 681.2.  Distribution map of Rhynchogyna.

Uses No uses have been reported for Rhynchogyna; it is rare in cultivation. (AS)

Cultivation

Anota Schltr., Orchideen, 587 (1914). Type species: Anota densiflora (Lindl.) Schltr. (basionym: Vanda densiflora Lindl.)

Derivation of name

Group 4 (see subtribal treatment). (MM)

From the Greek rhynchos, snout, and stylos, column, in reference to the beak-like column of the type species. ( JW)

Taxonomic notes

Description (Plate 105; Fig. 682.1)

The taxonomy of the genera closely allied to Cleisostoma, such as Pelatantheria, Rhynchogyna, Sarcoglyphis, Sarcophyton, Stereochilus, and others is still unresolved, so that Rhynchogyna is only tentatively maintained here. According to Seidenfaden (1988), Rhynchogyna differs from terete-leaved species of Cleisostoma mainly in the absence of a septum inside the spur and the long, beak-like rostellum. (AS)

Epiphytic herbs. Stem robust. Leaves many, distichous, often lorate, channelled, unequally bilobed or toothed, usually with pale green stripes, articulate to a sheathing base. Inflorescence lateral, racemose, erect or pendent, many-flowered. Flowers resupinate, with a ‘soapy’ fragrance. Sepals and petals free, similar, spreading, lateral sepals often wider and oblique. Petals smaller than sepals. Labellum rigidly adnate to column foot, entire or slightly trilobed, midlobe dorsiventrally flattened, spurred at base; spur laterally compressed and backward-pointing, without internal callosities. Column with a column foot; anther cap beaked; pollinia two, waxy, globular, cleft, solid, stipe linear, apex slightly dilated, viscidium ovate; rostellum beaked. ( JW)

Taxonomic literature Seidenfaden, G. (1988). Orchid genera in Thailand XIV. Fifty-nine vandoid genera. Opera Botanica, 95, 192–3.

682.  R H Y N C H O S TY LIS

Distribution (Fig. 682.2)

Rhynchostylis Blume, Bijdr., 285 (1825). Type species: Rhynchostylis retusa (L.) Blume (basionym: Epidendrum retusum L.)

Rhynchostylis includes three species collectively distributed from Sri Lanka and India through mainland Southeast Asia, China, and the Philippines south through Malaysia and Indonesia. ( JW)

260

RH YNCHOSTYLIS

C

A B

P K

N O M

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J L E

H

G

F

D

Fig. 682.1.  Rhynchostylis retusa (L.) Blume. A. Habit; B, C. Flower, three views; D. Ovary, labellum, and column, side view (floral bract detached); E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum, front view; I. Labellum with detail of hairs, longitudinal section; J. Column, front view; K. Column with anther cap, side view; L. Column, anther cap removed, side view; M. Anther cap, ventral view; N. Pollinarium; O. Capsule; P. Capsule, transverse section. Single bar = 1 mm; double bar = 1 cm. Drawn by Susanna Stuart-Smith from Koelz 22584 (A & O–P) and Trudel 584 (B–N), both (K). Reproduced with permission from N. R. Pearce and P. J. Cribb (2002), The Orchids of Bhutan. Royal Botanic Garden, Edinburgh and Royal Government of Bhutan.

261

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Fig. 682.2.  Distribution map of Rhynchostylis.

Palynology Schill and Pfeiffer (1977) examined the pollen of R. coelestis (Rchb.f.) A.H.Kent, R. gigantea (Lindl.) Ridl., and an unidentified species. Tetrads were convex with laevigate sculpturing and a rounded sexine (partly calymmate in the unnamed species) about 1.5–2.0 μm thick. (AP)

Cytogenetics Felix and Guerra (2010) listed chromosome numbers of n = 19–20 and 2n = 38 for the genus. ( JW)

Phytochemistry Alkaloids have been detected in R. coelestis, R. gigantea (Lindl.) Ridl. (as Saccolabium giganteum Lindl.), and R. retusa, including an additional record for the latter under the synonym Anota violacea (Lüning 1964, 1967). The main components of the floral fragrance of R. coelestis are (E)-ocimene (47.0%) and (E,E)-farnesal (34.4%). The latter occurs together with its regiosiomer, (Z,E)farnesal (5.9%), and the related (E,E)- and (Z,E)-farnesols (3.8 and 0.3%, respectively); other minor components that contribute to the scent include anethole, benzyl acetate, geranial, linalool, and α-terpineol (Kaiser 1993, 2011). (NV, RG)

and Vanda. A similar result was obtained by Carlsward et al. (2006b). (AK, MC)

Ecology All three species are epiphytes in forests at 0–1800 m. Rhychostylis coelestis (Rchb.f.) A.H.Kent occurs in deciduous forests and on small trees in open scrub at 200 m in Thailand. Rhychostylis gigantea is epiphytic on tree trunks in open forests at 1000 m in China (Hainan); it also occurs in deciduous forests at 200 m in Thailand, coastal forests on limestone in Sabah, and in the Philippines at 500 m. Rhychostylis retusa grows in open forests or at forest margins at 300–1500 m in China, beside streams in savannas in Sri Lanka, on Acacia Mill. (Fabaceae) at 600 m in Burma, on mango (Mangifera indica L.; Anacardiaceae) in India, in deciduous forests at 300–800 m in Thailand, and in open forests at 0–1000 m in Java, where it is particularly common in teak plantations (Tectona grandis L.f.; Lamiaceae). Rhychostylis gigantea subsp. violacea (Lindl.) Christenson grows on Mangifera indica in the Philippines. ( JW)

Pollination No information on pollination of Rhynchostylis is available. ( JW)

Phylogenetics

Uses

Rhynchostylis is sister to the rest of clade F, which includes Holcoglossum, Luisia, Papilionanthe, Paraphalaenopsis, Seidenfadenia,

Subedi et al. (2011) reported the medicinal use of R. retusa in Nepal: the sap of the roots is applied to cuts and wounds, leaf powder

262

ROBIQUETIA

is used to treat rheumatism, and the dried flowers are used as an insect repellent and to induce vomiting. Similar and additional medicinal uses have been reported for this species from India, where the root is known as rasna, and Sri Lanka (Lawler 1984). All species of Rhynchostylis are popular in cultivation. (AS)

Cultivation Group 2 (see subtribal treatment). (MM)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogeny of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86.

683.   RO BIQUE TIA Robiquetia Gaudich., Freyc. Voy. Uranie Phys. Bot., 426 (1829). Type species: Robiquetia ascendens Gaudich. Malleola J.J.Sm. & Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 979 (1913), syn. nov. Type species: not designated (note: the lectotype proposed by Garay (1972), Malleola sphingoides J.J. Sm., is not a valid type as this species is not mentioned in the protologue) Abdominea J.J.Sm., Bull. Jard. Bot. Buitenzorg, sér. 2, 14, 52 (1914), syn. nov. Type species: Abdominea micrantha J.J.Sm. Megalotus Garay, Bot. Mus. Leafl., Harvard Univ., 23, 184 (1972), syn. nov. Type species: Megalotus bifidus (Lindl.) Garay (basionym: Saccolabium bifidum Lindl.) India A.N.Rao, J. Econ. Taxon. Bot., 22, 701 (1998, publ. 1999), syn. nov. Type species: India arunachalensis A.N.Rao Samarorchis Ormerod, Taiwania, 53, 106 (2008), syn. nov. Type species: Samarorchis sulitiana Ormerod

Derivation of name The name commemorates Pierre Jean Robiquet (1780–1840), a French chemist who made numerous important and useful chemical discoveries, including caffeine and morphine. ( JW)

Description (Plates 106–108; Fig. 683.1, 683.2) Epiphytic herbs. Stem usually elongate, simple or sometimes branching, spreading or pendent, leafy. Leaves few to many, distichous, oblong to elliptic, obtusely or acutely unequally bilobed, coriaceous, deciduous, articulated to a sheathing base, sheaths (and occasionally blades) sometimes speckled purple. Inflorescences lateral at nodes, perforating leaf-sheaths, racemose or (less often) paniculate, many-flowered, usually pendent. Flowers resupinate, quaquaversal. Sepals and petals free, similar, dorsal sepal arched over column, lateral sepals spreading. Petals usually slightly smaller than sepals. Labellum rigidly adnate to column, trilobed, spurred, side lobes joined to column at posterior margin, often thickened anteriorly within, ascending, midlobe porrect, incurved, or recurved, linear or attenuate, basal portion concave, gradually becoming more fleshy and convex

distally, subacute to acuminate, sometimes furcate; spur apical, frequently retrorse, often curved, cylindric, inflated, frequently slightly to moderately constricted near middle, obtuse, with or without interior callosities or scales on front or back wall. Column without a foot; anther terminal, operculate, prominently rostrate, pollinia two or rarely four, when four in two unequal pairs, when two entire or partially cleft, solid, waxy, subglobose, commonly reclining on rostellum into clinandrium, stipe linear, spatulate, often uncinate, longer than pollinia, viscidium variable in size; stigma transverse, deeply recessed; rostellum bifid after removal of pollinarium. ( JW)

Distribution (Fig. 683.3) Robiquetia comprises 70 species collectively distributed from Sri Lanka, India, and the Himalayan region through Malaysia and Indonesia to the Philippines, northern Australia, New Guinea, and the Pacific islands, east to Fiji and Tonga, with a few species extending to Indochina and China. The centre of distribution lies in the Malay Archipelago. ( JW)

Cytogenetics According to Brandham (1999) and Felix and Guerra (2010), chromosome counts published for the genus are all 2n = 38. ( JW)

Phytochemistry Alkaloids have been detected in R. gracilistipes (Schltr.) J.J.Sm. (studied as R. tierneyana (Rupp) Dockrill), R. penangiana (J.J.Sm.) Kocyan & Schuit. (as Malleola penangiana (Hook.f.) J.J.Sm. & Schltr.), R. spathulata (Blume) Rchb.f. (as Cleisostoma spicatum Lindl.), and R. succisa (Lindl.) Seidenf. & Garay (as Saccolabium buccosum Rchb.f.) but not in R. wassellii Dockrill (Lüning 1964, 1967; Lawler and Slaytor 1969). (NV, RG)

Phylogenetics Robiquetia is a member of clade L, and as treated here includes Abdominea, India, Malleola, Megalotus, and Samarorchis. This condensation of genera was also supported by the results of Carlsward et al. (2006b), who found Malleola and Robiquetia (as sister taxa) related to Cleisostoma and allied genera. A similar result was obtained by Topik et al. (2005), who included only Abdominea and Malleola. All these taxa have a densely flowered inflorescence. (AK, MC)

Ecology All species are epiphytes in wet evergreen and other forests at 0–2000 m. ( JW)

Pollination Hardly any information on the pollination of Robiquetia is available. Carr (1928) reported that R. sylvestris (Ridl.) Kocyan & Schuit. (as Saccolabium undulatum Ridl.) is pollinated by small moths. ( JW, AS) 263

VA N D E A E

B C

I

J

H K D N F

L

E G

A

M

Fig. 683.1.  Robiquetia bifida (Lindl.) Kocyan & Schuit. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. lateral sepal; F. Petal; G. Labellum, from above; H. Labellum, longitudinal section; I. Column and labellum, side view; J. Column, anther cap removed, front view; K. Column, anther cap removed, side view; L. Anther cap, dorsal view; M. Anther cap, ventral view; N. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Wenzel 134 (K).

Uses No uses have been reported for Robiquetia; it is occasionally cultivated. ( JW)

Cultivation

Kocyan, A. and Schuiteman, A. (In press). New combinations in Aeridinae. Phytotaxa. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

Group 1 (see subtribal treatment). (MM)

6 8 4 .  S AC C OL A B I OPS I S

Taxonomic literature

Saccolabiopsis J.J.Sm., Bull. Jard. Bot. Buitenzorg, sér. 2, 26, 93 (1918). Type species: Saccolabiopsis bakhuisenii J.J.Sm.

Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogeny of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86.

Derivation of name

264

From Saccolabium, an orchid genus, and the Greek opsis, likeness. ( JW)

SAC COLABIOPSIS

C D

E

A

F

J

K I

G

H B Fig. 683.2.  Robiquetia transversisaccata (Ames & C.Schweinf.) J.J.Wood. A. Habit; B. Flower, side view; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum, spur removed; G. Pedicel with ovary, labellum, and column, side view; H. Pedicel with ovary, labellum, and column, longitudinal section; I, Anther cap, side view; J. Stipe and viscidium, front view; K. Pollinia. Single bar = 1 mm, double bar = 1 cm. Drawn by Eleanor Catherine from Clemens 50340 and Lamb AL 196/84. Reproduced with permission from J. J. Wood (1997), Orchids of Borneo. Volume 3. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

Description (Fig. 684.1) Epiphytic herbs. Leaves few, oblong-lanceolate, obtusely bilobed, thin-textured. Inflorescence lateral, few- to many-flowered, racemose, rarely subpaniculate, producing flowers at intervals facing apex of raceme, lasting at least two days, generally green; peduncle

sometimes thickened and angled near apex. Flowers thin-textured. Sepals and petals oblong or obovate, ringent. Petals slightly shorter than sepals. Labellum firmly adnate to base of column, trilobed, saccate or spurred, with a wide, broad opening, lacking interior ornamentation, although a transverse ridge sometimes separates 265

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Fig. 683.3.  Distribution map of Robiquetia.

spur from midlobe, side lobes generally erect, sometimes rudimentary, midlobe variable, from concave to convex; spur in line with column, cylindrical or cornute. Column cylindrical, with narrow apical wings, column foot absent; anther cap cucullate, narrowly rostrate, pollinia four, unequal, on an elongate stipe; rostellum porrect from apex of column, elongate, bifid at apex. ( JW)

species of the genus has been sequenced. It falls in clade L as the strongly supported sister to Cleisostoma lohii P.O’Byrne & J.J.Verm. Because Cleisostoma is large and polyphyletic, more research is needed before alteration of generic limits in this clade. (AK, MC)

Ecology Distribution (Fig. 684.2)

Species are epiphytes at 0–1500 m in forests and dry scrub. ( JW)

Saccolabiopsis includes 14 species collectively distributed from India, Bhutan, and China south to Thailand, through the Malay Archipelago, and east to New Guinea, Australia, and Fiji. ( JW)

Pollination

Cytogenetics

Uses

No chromosome counts for Saccolabiopsis have been published. ( JW)

No uses have been reported for Saccolabiopsis; it is rare in cultivation. ( JW)

Phytochemistry

Cultivation

A survey of alkaloid distribution in Orchidaceae gave a negative result for S. armitii (F.Muell.) Dockrill (Lawler and Slaytor 1969). (NV, RG)

Phylogenetics Saccolabiopsis has already been enlarged by the inclusion of some species originally described under Malleola, but so far only one 266

No information on pollination of Saccolabiopsis is available. ( JW)

Group 2 (see subtribal treatment). (MM)

6 8 5 .  S AC C OL A B I U M Saccolabium Blume, Bijdr., 292 (1825). Type species: Saccolabium pusillum Blume Saccochilus Blume, Fl. Jav., 1, viii (1828), nom. invalid. Type species: not designated.

SACCOLABIUM

B A

D

F

I

H J

E

C

G Fig. 684.1.  Saccolabiopsis pusilla (Lindl.) Seidenf. & Garay. A. Habit; B. Flower, oblique view; C. Ovary, labellum, and column, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum with spur, oblique view; H. Column, anther cap removed, oblique view; I. Anther cap, ventral and side views; J. Stipe and viscidium (left), pollinarium (right). Single bar = 1 mm; double bar = 1 cm. Drawn by Susanna Stuart-Smith from Pantling 112 (K). Reproduced with permission from N. R. Pearce and P. J. Cribb (2002), The Orchids of Bhutan. Royal Botanic Garden, Edinburgh and Royal Government of Bhutan.

Taxa at the rank of section referable to Saccolabium: Saccolabium sect. Eusaccolabium Cordem., Fl. Ile Réun., 195 (1895), nom. illeg. Saccolabium sect. Microsaccolabium J.J. Sm., Bull. Jard. Bot. Buitenzorg, sér. 2, 13, 46 (1914). Type species: Saccolabium pusillum Blume

Derivation of name From the Latin saccus, bag, sac, and labium, lip, in reference to the bag-like shape of the labellum. ( JW)

Description (Fig. 685.1) Epiphytic herbs. Stems 10–43 cm long, simple or branching, leafy, horizontal to semi-pendent, rooting at base and turning up distally. Leaves flat, oblong or lanceolate, distichous, unequally bilobed or acutely bidentate. Inflorescence lateral, one-flowered. Flowers resupinate or non-resupinate, not opening widely, 4–6 mm wide, fugaceous. Sepals and petals free, or lateral sepals sometimes adnate to column foot, often incurved. Labellum obscurely trilobed, distinctly spurred, midlobe sometimes 267

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Fig. 684.2.  Distribution map of Saccolabiopsis.

trilobed; disc sometimes with a transverse lamella behind side lobes and pressed against column foot; spur ecallose, sometimes hairy inside. Column with a short foot; pollinia two, entire. ( JW)

(members of clade E here). Saccolabium was not included in the study here. (AK, MC)

Distribution (Fig. 685.2)

Species occur as epiphytes in forests at 900–2000 m. ( JW)

Ecology

The five species of Saccolabium are collectively distributed in Sumatra and Java. ( JW)

Pollination

Cytogenetics

There is no information available on pollination of Saccolabium. ( JW)

Brandham (1999) reported published chromosome numbers of 2n = 30, 36–44, 38, 76 with a basic number of x = 19, yielding 2x = 38 (most common) and 4x = 76. (AP)

Phytochemistry Surveys of alkaloid content published by Lüning (1964, 1967) for Saccolabium are now referred to other genera (Gastrochilus, Rhynchostylis, Robiquetia, and Vanda). No other reports on the phytochemistry of this genus are available. (NV, RG)

Phylogenetics Topik et al. (2005) sequenced one species of Saccolabium, which falls together with Pennilabium, Hymenorchis, Tuberolabium, Amesiella, Ceratocentron, Dyakia, Cryptopylos, and Macropodanthus 268

Uses No uses have been reported for Saccolabium; it is rare in cultivation. ( JW)

Cultivation Group 2 (see subtribal treatment). (MM)

Taxonomic notes Saccolabium s.l. has been a major source of taxonomic confusion. Originally proposed by Blume in 1825, subsequent authors have proposed 27 sections, 6 series, and added over 300 names to the genus. As Christenson (1986) pointed out, few of these

SANTOTOMASIA

B

L M N I C

J E

D K

G F

A

H

O Fig. 685.1.  Saccolabium sigmoideum J.J.Sm. A. Habit; B. Flower; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum, from above; G. Labellum, longitudinal section; H. Column and base of labellum, side view; I. Column, front view; J. Column apex, anther cap removed; K. Column, side view; L. Anther cap, dorsal view; M. Anther cap, ventral view; N. Pollinarium; O. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Kessler & de Vogel 8712 (K).

subsequent additions are related to the type. Saccolabium, as currently understood and here accepted, comprises four named and one undescribed species from Sumatra and Java. A full account of the taxonomic history of the genus together with a full list of excluded species was provided by Christenson (1986). ( JW)

Taxonomic literature Christenson, E. A. (1986). An historical review of Saccolabium (Orchidaceae) and excluded species. Kew Bulletin, 41, 833–53. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK

and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

6 8 6 .   S A NT OT OM A S I A Santotomasia Ormerod, Taiwania, 53, 162 (2008). Type species: Santotomasia wardiana Ormerod

Derivation of name The genus was named after Mt. Santo Tomas on the island of Luzon in the Philippines where the type species was discovered. (AS) 269

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Fig. 685.2.  Distribution map of Saccolabium.

Description (Fig. 686.1)

Phytochemistry

Epiphytic herbs. Vegetative parts incompletely known. Stem said to be stout, elongate. Leaves conduplicate, articulate, distichous, ligulate to oblong, coriaceous, apex unequally obtusely bilobed. Inflorescence lateral, paniculate, many-flowered; rachis continuing the line of peduncle, straight. Flowers about 1 cm wide, longevity unknown, opening several at a time, quaquaversal, creamy white, labellum with two dark reddish spots inside. Sepals entire, free, obtuse to subacute. Petals slightly shorter than sepals, obovate, obtuse. Labellum rigidly attached to base of column, spurred-saccate, entire; spur conical, with two thickenings, one below the other, on the front wall and one thickening on each of the side walls. Column without a foot; anther helmet-shaped, long-rostrate, pollinia two, obovoid, incompletely cleft, stipe longer than pollinia, strap-shaped; rostellum bidentate. (AS)

No reports have been found on the phytochemistry of Santotomasia. (NV, RG)

Distribution (Fig. 686.2) Santotomasia is monospecific and from the Philippines (Luzon). (AS)

Cytogenetics Chromosome numbers for the species have not been published. (AP) 270

Phylogenetics There is no phylogenetic information available for this genus. (AK)

Ecology The habitat of S. wardiana is unknown. (AS)

Pollination Nothing is known about pollination in Santotomasia. (AS)

Uses No uses have been reported for Santotomasia; it is not known to be in cultivation. (AS)

Taxonomic notes Floral morphology of this genus is similar to that of Tuberolabium, but the paniculate inflorescences, robust plant habit, and the morphology of the pollinia indicate an affinity with Cleisostoma. (AS)

SARCANTHOPSIS

B I

H J

C F

E

A D

G

Fig. 686.1.  Santotomasia wardiana Ormerod. A. Inflorescence branch; B. Flower; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Column and labellum, side view; G. Labellum, longitudinal section; H. Column, side view; I. Pollinarium; J. Viscidium with stipe removed. Single bar = 1 mm, double bar = 1 cm. Redrawn by Judi Stone after P. Ormerod, Taiwaniana, 53, 162, Fig. 4 (2008).

687.   SARC A N TH O P SIS

Description (Fig. 687.1)

Sarcanthopsis Garay, Bot. Mus. Leafl., Harvard Univ., 23, 198 (1972). Type: Sarcanthopsis nagarensis (Rchb.f.) Garay (basionym: Sarcanthus nagarensis Rchb.f.)

Epiphytic or lithophytic herbs. Roots terete, not or sparsely branching, glabrous, smooth. Stem to more than 3 m tall, erect or overhanging with apical part erect, leafy, rooting along its length. Leaves conduplicate, articulate, distichous, oblong to linearoblong, not twisted at base, coriaceous, apex rounded to emarginate or unequally bilobed with rounded lobes, margins entire. Inflorescences lateral, arising near apex of stem, solitary, erect, paniculate with up to two levels of branching, many-flowered,

Derivation of name From the orchid genus Sarcanthus (now known as Cleisostoma), and the Greek opsis, likeness. (AS)

271

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Fig. 686.2.  Distribution map of Santotomasia.

bracts subtending branches relatively large and broad; floral bracts similar but smaller, patent, yellowish. Flowers resupinate, 2–3 cm in diameter, quaquaversal, usually yellowish with brown spots. Sepals free, oblong to obovate, margins entire. Petals obovate. Labellum rigidly attached to column, not spurred, trilobed, divided into a concave hypochile with the lateral lobes and a flat epichile at right angles or at a sharp angle to the hypochile (the labellum knee-like bent in lateral view), hypochile with a central, cushion-like, laterally flattened callus, epichile with two calli, one basally and one just below midlobe, which forms apical part of epichile. Column lacking a foot; anther cucullate, pollinia four, cohering closely in two bodies, those in each body unequal, stipe linear, about as long as pollinia, viscidium broadly oblong; rostellum bidentate after removal of pollinarium. Capsule oblong. (AS)

Distribution (Fig. 687.2)

Phytochemistry Alkaloids are present in S. warocqueana (Rolfe) Garay (as Vandopsis warocqueana (Rolfe) Schltr., although given as ‘V. wracquianum’) at levels < 0.01% (Lawler and Slaytor 1970). (NV, RG)

Phylogenetics Sarcanthopsis is sister to two included species of Diploprora plus Diplocentrum, and this clade is one of those in grade K. (AK, MC)

Ecology All species are epiphytes or lithophytes in coastal vegetation and lowland rain forests at 0–1000 m, often in exposed positions, for example on Cocos nucifera L. (Arecaceae), Calophyllum inophyllum L. (Callophyllaceae) along the seashore and on coral rock. They sometimes grow close to the sea and are subjected to saltwater spray. (AS)

Sarcanthopsis is a genus of five morphologically similar species collectively from New Guinea, Palau, Solomon Islands, Vanuatu, New Caledonia, Fiji, and the Horne Islands. (AS)

Pollination

Cytogenetics

Uses

Chromosome numbers for Sarcanthopsis have not been reported. ( JW)

No uses have been reported for Sarcanthopsis; it is rare in cultivation. (AS)

272

Nothing is known about pollination in Sarcanthopsis. (AS)

SARCANTHOPSIS

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E

H

F

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I

J B A K

Fig. 687.1.  Sarcanthopsis nagarensis (Rchb.f.) Garay. A. Habit, ×0.67; B. Labellum, ×3; C. Flower with petal and lateral sepal removed, ×3; D. Dorsal sepal, ×3; E. Labellum, longitudinal section, ×4; F. Lateral sepal, ×3; G. Petal, ×3; H. Flower, ×3; I. Anther cap, dorsal and ventral views, ×6; J. Pollinia, ×10; K. Pollinia, ×10. All magnifications as originally published. Drawn by Sue Wickison from Wickison 140. Reproduced with permission from B. A. Lewis and P. J. Cribb (1991), Orchids of the Solomon Islands and Bougainville. Royal Botanic Gardens, Kew.

Cultivation Group 2 (see subtribal treatment). Sarcanthopsis plants are among the largest in Aeridinae, rivalled only by Vandopsis and some Vanda species. Apart from considerable space, they require ample warmth, light, humidity, and air movement. As the roots

cannot be confined in a container, the plants are best grown in a basket or tied to a support and sprayed regularly. (AS, MM)

Taxonomic literature Wood, J. J. and Ormerod, P. (2003). Sarcanthopsis hansemannii. Curtis’s Botanical Magazine, 20, 152–60.

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Fig. 687.2.  Distribution map of Sarcanthopsis.

688.  SARC O C H ILUS Sarcochilus R.Br., Prodr. Fl. Nov, Holl., 332 (1810). Type species: Sarcochilus falcatus R.Br. Gunnia Lindl., Edwards’s Bot. Reg., sub t. 1699 (1834). Type species: Gunnia australis Lindl. Parasarcochilus Dockrill, Austr. Sarcanthinae, 22 (1967). Type species: Parasarcochilus spathulatus (R.S.Rogers) Dockrill (basionym: Sarcochilus spathulatus R.S.Rogers) Monanthochilus (Schltr.) R.Rice, Oasis, Suppl., 3, 2 (2004). Type species: Monanthochilus chrysanthus (Schltr.) R.Rice (basionym: Sarcochilus chrysanthus Schltr.)

Derivation of name From the Greek sarx (sarkos), meaning flesh, and cheilos, lip, in allusion to the fleshy labellum of the type species. (PC)

Description (Plate 109; Fig. 688.1, 688.2) Epiphytic or lithophytic herbs. Roots smooth. Stem leafy, covered with leaf bases. Leaves few, distichous, coriaceous, linear, ligulate, obovate or oblong-elliptic, often falcate, unequally bilobed at apex, articulated at base to sheath. Inflorescences axillary, erect to pendent, usually from basal part of stem, simple, few- to several-flowered, usually as long as or longer than leaves, rarely much shorter; bracts triangular, sheathing. Flowers usually flat or campanulate; pedicel and rachis as long as or longer than sepals. 274

Sepals and petals similar, free, spreading, white to olive, sometime spotted with red. Labellum smaller than sepals, trilobed, spurred at base, side lobes erect around column, hairy or glabrous; spur tapering to rounded tip, callus lobed, on back wall within spur. Column terete; pollinia four, ovoid, in two unequally sized pairs, stipe narrowly to broadly spatulate, viscidium elliptic to oblong. (PC)

Distribution (Fig. 688.3) Sarcochilus is a genus of 25 species collectively distributed mostly in eastern Australia from Cape York to Tasmania, with four species in New Caledonia. Sarcochilus sect. Monanthochilus Schltr. (Monanthochilus (Schltr.) R.Rice) with four species is endemic to New Guinea. The delimitation of some Australian species needs further study. (PC, AS, PA)

Palynology Schill and Pfeiffer (1977) described pollen tetrads of S. hartmannii F.Muell. as convex with laevigate to weakly rugulate sculpturing and a rounded sexine. (AP)

Cytogenetics Brandham (1999) listed published chromosome numbers for the genus as 2n = 36, 38, 76 with a basic number of x = 19, yielding 2x = 28 (the most common), and 4x = 76. (AP)

SARCOCHILUS

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Fig. 688.1.  Sarcochilus falcatus R.Br. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal, E. Lateral sepal; F. Petal; G. Labellum; H. Labellum, from above; I. Labellum, from below; J. Column and labellum, side view; K. Column and labellum, from below; L. Column, front view; M. Column, side view; N. Column apex, anther cap removed; O. Anther cap with pollinia, ventral view; P. Anther cap, dorsal view; Q. Pollinarium; R. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Weinthal s.n. Nov. 1927 (K) and Kew Spirit Collection no. 52488.

Phytochemistry In an early survey, Boorsma (1902) detected traces of alkaloids, but no saponins, in one species of Sarcochilus. Numerous records on alkaloid content of the genus (including two entries for the former Parasarcochilus) were published subsequently (Lüning 1967; Lawler and Slaytor 1969). According to a later review, 2 of 21 species of Sarcochilus (and neither of two species of Parasarcochilus) contained alkaloids at levels ≥ 0.1% (Lüning 1974). (NV, RG)

Phylogenetics Topik et al. (2005) found that Sarcochilus is not ­monophyletic, and Carlsward et al. (2006b) had it sister to Taeniophyllum,

which is probably an artefact of incomplete generic sampling. Here, Sarcochilus (one species included) is a member of the Australia clade, perhaps as sister to the rest of that clade. (AK, MC)

Ecology Most species grow in and around rain forests and wet sclerophyll forests as shaded epiphytes, often in large populations, or less commonly as lithophytes (Sarcochilus hartmannii F.Muell., S. fitzgeraldii F.Muell., and S. ceciliae F.Muell.). Sarcochilus hillii (F.Muell.) F.Muell., S. dilatatus F.Muell., and S. minutiflos F.M.Bailey are twig epiphytes found in moist or dry rain forest scrub on 275

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Fig. 688.2.  Sarcochilus spathulatus R.S.Rogers [formerly Parasarcochilus spathulatus (R.S.Rogers) Dockrill]. A. Habit; B. Flower, front view; C. Flower and bract, side view; D. Flower, longitudinal section; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum; I. Column and labellum, side view; J. Column, front view; K. Column, side view; L. Anther cap, dorsal view; M. Anther cap with pollinia; N. Pollinarium; O. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from W. H. Nicholls, Orchids of Australia, pl. 463 (1969) and Kew Spirit Collection no. 28260.

small trees and low shrubs, often by watercourses. Sarcochilus spathulatus R.S.Rogers is localized on rain forest trees, often as a twig epiphyte, and along creeks, and S. weinthallii F.M.Bailey is found around moister rain forests and in dry vine thickets. Sarcochilus australis (Lindl.) Rchb.f. is a twig epiphyte in temperate Australian mainland forests, often along watercourses; it is the most southerly species, growing in cool temperate forests on Nothofagus Blume (Nothofagaceae) in Tasmania. Natural hybrids occur occasionally where species grow close together, and there are at least two intergeneric hybrids with Plectorrhiza (Upton 1992; Adams, unpublished). (PA, PC, AS) 276

Pollination There are no reports of pollination in Sarcochilus (but see under Rhinerrhizopsis). (AS)

Uses Sarcochilus is commonly cultivated, especially in Australia. Hybrids are used increasingly in the international pot-plant trade and for exhibition. The long racemes of white-flowering S. hartmannii and pink-red flowering S. fitzgeraldii and their hybrids are used in floristry, especially in wedding bouquets. (PA)

SARCOCHILUS

Fig. 688.3.  Distribution map of Sarcochilus.

Cultivation

Taxonomic notes

Larger species are relatively easy to grow in temperate climates in a cool greenhouse or shade house with brisk air movement, moderate to high humidity, shady conditions, good drainage, and weak fertilizer applied frequently except in the coldest months. In winter, overwatering should be avoided. Most species do not tolerate drying out; misting assists in summer. Watering is best in the morning in winter and in the evening in summer (Upton 1992). There are three main cultural groups:

Most species of Sarcochilus have calli within the spur. On this basis, Dockrill (1967) removed Sarcochilus spathulatus R.S.Rogers and S. weinthalii F.M.Bailey from the genus to establish his new genus Parasarcochilus because of their lack of calli in the spur. At the same time he added a third species, S. hirticalcar (Dockrill) M.A.Clem. & B.J.Wallace, to the genus. Subsequent authors have not upheld Parasarcochilus (e.g. Clements 1989; Upton 1992). Rice elevated S. section Monanthochilus Schltr., which is endemic to New Guinea, to generic level. This section is characterized by the abbreviated, one-flowered inflorescences; its status needs further analysis. (PC, AS)

1) lithophytes: Sarcochilus hartmannii, S. fitzgeraldii, and S. ceciliae are grown under cool conditions, mainly in pots with weathered bark/stone mix under 60% shade, winter minimum above 3° C. Sarcochilus ceciliae succeeds in pure stone and tolerates warmer temperatures. 2) larger epiphytes: Sarcochilus falcatus and S. olivaceus Lindl. require slab culture, 70% shade, minimum temperature 2° C, attached firmly to a slab and lying flat on a bench until established. Watering is required more frequently than with pot culture. These species grow well attached to shady trees in suitable climates. 3) small (twig) epiphytes: these are mainly cool-growing and more difficult to cultivate, requiring a slab or mount, with constantly moist and humid conditions. Tropical S. minutiflos F.M.Bailey and S. hirticalcar (Dockrill) M.A.Clem. & B.J.Wallace require intermediate conditions, minimum 12° C. (PA, SL)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogeny of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Clements, M. A. (1989). Catalogue of Australian Orchidaceae. Australian Orchid Research, 1, 1–155. Dockrill, A. W. (1967). Australasian Sarcanthinae. Australasian Native Orchid Society. Sydney, Australia. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

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Upton, W. T. (1992). Sarcochilus orchids of Australia. Double ’U’ Orchids, West Gosford, New South Wales, Australia.

689.  SARC O G LY P H IS Sarcoglyphis Garay, Bot. Mus. Leafl., Harvard Univ., 23, 200 (1972). Type species: Sarcoglyphis mirabilis (Rchb.f.) Garay (basionym: Sarcanthus mirabilis Rchb.f.)

Derivation of name

and future research will hopefully clarify the status of Sarcoglyphis. Earlier workers with much sparser sampling of genera found different topologies. Carlsward et al. (2006b) produced a topology similar to that provided here, whereas Topik et al. (2005) placed Sarcoglyphis with Arachnis; the latter is distant from clade L. (AK, MC)

Ecology Species are epiphytes in primary, riverine, lowland, and hill forests at 100–1000 m. ( JW)

From the Greek sarx, flesh, and glyphon, carving, in reference to the prominent, fleshy rostellum that rests on the clinandrium like a carved ornament, upon which the pollinia recline so as to be in a dorsal position. ( JW)

Pollination

Description (Plate 110; Fig. 689.1)

No uses have been reported for Sarcoglyphis; it is occasionally cultivated. ( JW)

Epiphytic herbs. Stem few-leaved. Leaves distichously arranged, ligulate, flat, unequally bilobed, with sheathing base, articulate, duplicate. Inflorescence arising from lower portion of stem, axillary, pendent, branched or unbranched, many-flowered. Flowers resupinate. Sepals and petals free, similar. Petals usually smaller and narrower than sepals. Labellum rigidly adnate at base to column, spurred, trilobed, lateral lobes suberect; spur subconical, inside septate and with a callus on back wall. Column without foot; clinandrium low; anther cap semi-globose, apex beaked, pollinia four, appearing as two almost equal pollen masses, waxy, compressed-globular, separate, each attached to a common long stipe through a caudicle at a right angle, viscidium ovate; stigma suborbicular; rostellum raised, laterally compressed, helmet-like in side view, with a longitudinal furrow along its edge into which stipe and dorsally positioned pollinia recline. ( JW)

Distribution (Fig. 689.2) Sarcoglyphis includes 12 species distributed from India, China, and Burma south and east to the Indonesian Archipelago. ( JW)

Cytogenetics No chromosome counts for Sarcoglyphis have been recorded. ( JW)

Phytochemistry No reports have been found on phytochemistry of Sarcoglyphis. (NV, RG)

Phylogenetics Sarcoglyphis (clade L) is characterized by a bent rostellum. Three of the species included fall together with strong support, but another species (S. masiusii Miadin, A.L.Lamb & Emoi) is well supported as sister to Cleisostoma suaveolens, and so belongs in a different genus. However, clade L is a difficult clade that includes many poorly delimited genera (e.g. Stereochilus, Cleisostoma, Cleisocentron), 278

No information on pollination of Sarcoglyphis is available. ( JW)

Uses

Cultivation Group 2 (see subtribal treatment). (MM)

Taxonomic notes The taxonomy of the genera closely allied to Cleisostoma, such as Pelatantheria, Rhynchogyna, Sarcoglyphis, Sarcophyton, Stereochilus, and others, is still unresolved, and Sarcoglyphis is only tentatively maintained here. Within this alliance, Sarcoglyphis may be recognized by the raised, crest-like rostellum, which is similar to that found in Seidenfadenia. (AS)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogeny of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

6 9 0 .  S A RC OP H YT ON Sarcophyton Garay, Bot. Mus. Leafl., Harvard Univ., 23, 201 (1972). Type species: Sarcophyton crassifolium (Lindl. & Paxton) Garay (basionym: Cleisostoma crassifolium Lindl. & Paxton)

Derivation of name From the Greek sarx, flesh, and phyton, plant, in reference to the highly fleshy nature of all plant parts. ( JW)

Description (Plate 111; Fig. 690.1) Epiphytic or lithophytic herbs. Stems erect, elongate, with many leaves. Leaves distichous with sheathing bases, coriaceous, conduplicate, unequally bilobed, sessile, articulate to sheath. Inflorescence

SARCOPHYTON

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Fig. 689.1.  Sarcoglyphis potamophila (Schltr.) Garay & W.Kittr. A. Habit; B. Leaf; C. Flower, side view; D. Rachis, transverse section; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum and column, anther cap removed, front view; I. Labellum, longitudinal section; J. Column with pollinarium, anther cap removed, side view; K. Anther cap, front view; L. Pollinarium, two views. Single bar = 1 mm, double bar = 1 cm. Drawn by Susanna Stuart-Smith after O’Byrne in Malayan Orchid Review, 32, 52, 69 (1998). Reproduced with permission from J. J. Wood (2003), Orchids of Borneo. Volume 4. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

axillary, racemose or paniculate, lateral, many-flowered. Flowers resupinate. Sepals similar, free, spreading. Petals smaller and narrower than sepals, spreading. Labellum adnate at base to column, trilobed, side lobes erect, midlobe recurved, adaxial surface usually conspicuously wrinkled, base with two calli at spur entrance; spur cylindrical. Column without foot; anther cap rostrate; pollinia four, waxy, globose, almost equal in size, separate from each other, attached by a

caudicle to a linear-spatulate stipe at an angle, viscidium subglobose; stigma suborbicular; rostellum arcuate acute or bilobed. ( JW)

Distribution (Fig. 690.2) Sarcophyton consists of three species collectively and discontinuously distributed in India (?), Burma, China, Taiwan, and the Philippines (Luzon). ( JW) 279

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Fig. 689.2.  Distribution map of Sarcoglyphis.

Cytogenetics

Cultivation

No chromosome counts for Sarcophyton have been recorded. ( JW)

Group 2 (see subtribal treatment). (MM)

Phytochemistry

Taxonomic notes

No reports have been found on phytochemistry of Sarcophyton. (NV, RG)

Phylogenetics Sarcophyton is a member of clade L, which includes the problematic genus Cleisostoma. Until a solution is determined for Cleisostoma, which is grossly polyphyletic in the results shown here, the generic disposition of Sarcophyton cannot be resolved. (AK, MC)

Ecology All species of Sarcophyton are epiphytes or lithophytes from the lowlands up to 800 m elevation. ( JW)

Pollination No information on the pollination of Sarcophyton is available. ( JW)

Uses No uses have been reported for Sarcophyton; it is rare in cultivation. ( JW) 280

The taxonomy of the genera closely allied to Cleisostoma, such as Pelatantheria, Rhynchogyna, Sarcoglyphis, Sarcophyton, Stereochilus, and others is still unresolved, and Sarcophyton is only tentatively maintained here. (AS)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogeny of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

6 9 1 .  S C H I S T OT YLU S Schistotylus Dockrill, Austral. Sarcanth., 29, t. 43 (1967). Type species: Schistotylus purpuratus (Rupp) Dockrill (basionym: Cleisostoma purpuratum Rupp)

SC HISTOTYLUS

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Fig. 690.1.  Sarcophyton crassifolium (Lindl. & Paxton) Garay. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum; H. Labellum, longitudinal section; I. Column and labellum, side view; J. Column, front view; K. Column, side view; L. Anther cap, dorsal view; M. Anther cap, ventral view; N. Pollinarium, O. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone habit after Paxton’s Flower Garden, 3, pl. 99 (1852) and from Kew cult. s.n. June 1988 (K).

Derivation of name From the Greek schistos, split, separated, and tylos, callus, referring to the callus aligned perpendicular to the spur entrance. ( JW)

Description (Fig. 691.1) Epiphytic herbs. Stem up to 3 cm long. Leaves 3–6, linear, often falcate. Inflorescence racemose, 2–4 cm long, 4–8-flowered; peduncle about same length as rachis; rachis flexuous; floral bracts minute. Flowers 5–6 mm long, resupinate, green heavily blotched purple, pleasantly scented, lasting for about two days. Sepals and petals free.

Dorsal sepal ovate, concave, lateral sepals falcate-subovate, asymmetric, shortly clawed, obtuse. Petals falcate-subovate, asymmetric, acute. Labellum rigidly joined to base of column, saccate, spurred, trilobed, side lobes erect, incurved in front, sublunate, distally narrowly triangular or tooth-like, midlobe concave-deltoid, the walls thick, the thick floor produced backwards part way across and connate to sides of anterior section of spur entrance as a thick, downcurved, deeply bifid callus, with divergent arms; spur cylindrical, compressed anterior-posteriorly, without interior calli or septa. Column slender at base, dilated above, column foot absent, wings absent; pollinia four, in two closely appressed, 281

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Fig. 690.2.  Distribution map of Sarcophyton.

subequal pairs, stipe lorate, viscidium set at an acute angle to stipe; stigma deeply set; rostellum downcurved, apex bifid. ( JW)

and ridges, and on Leptospermum J.R.Forst. & G.Forst. (Myrtaceae) in sclerophyll forest at 600–1000 m. (PA, JW)

Distribution (Fig. 691.2)

Pollination

Schistotylus is monospecific and endemic to Australia (southern Queensland and northern New South Wales). ( JW)

Cytogenetics No chromosome numbers have been recorded for Schistotylus. ( JW)

Phytochemistry Low levels of alkaloids (< 0.01%) have been detected in S. purpuratus (Lawler and Slaytor 1969). (NV, RG)

Phylogenetics Schistotylus is a member of the Australia clade. (AK, MC)

There is no information available on pollination of Schistotylus. ( JW)

Uses No uses have been reported for Schistotylus; it is rare in cultivation. ( JW)

Cultivation Group 1 (see subtribal treatment). Plants may be grown as for Sarcochilus twig epiphytes in constantly cool and humid conditions with minimal drying out. It is difficult to grow over many years. (PA)

Ecology

6 9 2 .  S C H OENORC H I S

Schistotylus purpuratus is a twig epiphyte, localized on shrubs in humid areas of wet sclerophyll and other drier forests, on slopes

Schoenorchis Reinw., in Hornschuh., Syll. Pl. Nov., 2, 4 (1825). Type species: Schoenorchis juncifolia Blume

282

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Fig. 691.1.  Schistotylus purpuratus (Rupp) Dockrill. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum, from above; H. Labellum, longitudinal section; I. Column and labellum, side view; J. Column, front view; K. Column, side view; L. Column, back view; M. Anther cap; N. Pollinarium, front view; O. Pollinarium, side view; P. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone; habit, anther, and pollinarium redrawn after A. W. Dockrill, Australian Indigenous Orchids (1969); others from Kew Spirit Collection no. 28493.

Bidoupia Aver. & Christenson ex Senghas, Orchideen, ed. 3, 2758 (2002), nom. invalid. Type species: Bidoupia bicamerata Aver. & Christenson ex Senghas

Derivation of name From the Latin schoenus, reed or rush, in reference to the rushlike leaves of the type, and the Greek orchis, orchid. ( JW)

Description (Plate 112; Fig. 692.1) Epiphytic herbs. Stems up to 30 cm long, pendent or ascending, simple or sometimes branching, with several to many nodes.

Leaves many, distichously arranged, glabrous, dorsiventrally flattened, semi-terete or terete, articulate to a sheathing base, deciduous, duplicate. Inflorescence lateral, racemose or paniculate, many-flowered. Flowers resupinate, usually not opening widely, usually white or red-purple. Sepals similar, free, often dorsally carinate. Petals free, smaller than sepals. Labellum rigidly adnate to base of column, about as long as petals or longer, trilobed, spurred at base, side lobes erect, midlobe slightly larger, often spatulate; spur usually parallel to ovary, lacking interior calli, sometimes with a longitudinal dividing wall inside near apex. Column without foot; pollinia four, arranged as two pairs, waxy, about equal, flat-globular or semi-globular, solid, attached by a 283

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Fig. 691.2.  Distribution map of Schistotylus.

common stipe to a narrowly elliptic to ovate viscidium; stigma at base of column; rostellum pointed, geniculate. ( JW)

S. micrantha (as S. densiflora Schltr.) gave positive results (Lüning 1967; Lawler and Slaytor 1969). (NV, RG)

Distribution (Fig. 692.2)

Phylogenetics

Schoenorchis comprises 23 species collectively distributed from the Himalayan region, southern India, and Sri Lanka to southern China and the Philippines, Thailand, south to Indonesia and east to New Guinea, Australia, and the Pacific islands including Samoa. ( JW)

Schoenorchis is a member of clade L, in a subclade with Cleisomeria, polyphyletic Cleisostoma, Pelatantheria, and one of the included species of Rhynchogyna. Morphological differences within the genus include two forms of leaf characters/habit, suggesting the possibility of two genera. However, Carlsward et al. (2006b) sequenced two Schoenorchis species, one of each of the respective groups, which were sister taxa. (AK, MC)

Palynology Schill and Pfeiffer (1977) studied the pollen of S. juncifolia Reinw. ex Blume and S. micrantha Reinw. ex Blume (as Ascocentrum micranthum (Reinw. ex Blume) Holttum) and reported that the tetrads were convex with laevigate sculpturing (calymmate in S. micrantha like Ascocentrum species). The sexine was rounded and 0.8–2.0 μm thick. (AP)

Cytogenetics Brandham (1999) listed a chromosome number of 2n = 36 for S. gemmata (Lindl.) J.J.Sm., and Felix and Guerra (2010) cited 2n = 38 for the same species. ( JW)

Ecology Species of Schoenorchis are epiphytes in gallery and wet evergreen forests, sometimes on ultramafic substrate, at 0–2500 m. Schoenorchis sarcophylla Schltr. grows on the outermost branches of Tristania exiliflora F.Muell. (Myrtaceae) overhanging rapidly flowing creeks in Queensland, Australia. ( JW)

Pollination There is no information available on pollination of Schoenorchis. ( JW)

Phytochemistry

Uses

Two species have been investigated for alkaloid content; both S. gemmata (as Cleisostoma gemmatum (Lindl.) King & Pantl.) and

No uses have been reported for Schoenorchis; it is sometimes cultivated. ( JW)

284

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Fig. 692.1.  Schoenorchis buddleiflora (Schltr. & J.J.Sm.) J.J.Sm. A. habit; B. Leaf, basal portion and transverse section; C. Portion of inflorescence; D. Flower, side view; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Pedicel with ovary, labellum, and column, side view; I. Portion of labellum, column, and portion of ovary, oblique view; J. Column, front view; K. Column, side view; L. Anther cap, side view; M. Anther cap and pollinarium, side view; N. Pollinarium, side view. Single bar = 1 mm, double bar = 1 cm. Drawn by Susanna Stuart-Smith from Argent C. 14844. Reproduced with permission from J. J. Wood (2003), Orchids of Borneo. Volume 4. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

Cultivation Group 2 (see subtribal treatment). (MM)

Taxonomic notes The authorship of Schoenorchis is somewhat in doubt. Carl Ludwig von Blume (1796–1862) first introduced the name into print in

his Catalogus published in 1823, where it appears as a nomen nudum. He commented that it had been described by Caspar Reinwardt (1773–1854) but omitted details of where and when. Two independent descriptions were subsequently published in 1825, viz. by Reinwardt (in Hornschuh’s Nova plantarum) and by Blume (in his Bijdragen). Seidenfaden (1988), supported by Stafleu (personal 285

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Fig. 692.2.  Distribution map of Schoenorchis.

communication), considered Reinwardt’s publication to predate Blume’s. Garay (1972), however, explained that Blume’s figures of S. juncifolia Reinw. ex Blume, which is the type of the genus and published in the Platen of Bijdragen, appeared in June 1825, casting further uncertainty over priority. Seidenfaden maintained that, in the light of Blume’s original statement in his Catalogus, Reinwardt should have the benefit of the doubt. ( JW)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogeny of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Garay, L. A. (1972). On the systematics of the monopodial orchids I. Botanical Museum Leaflets, Harvard University, 23, 149–212. Seidenfaden, G. (1988). Orchid genera in Thailand XIV. Fifty-nine vandoid genera. Opera Botanica, 95, 168–81.

693.  SE ID E N FA D E N IA Seidenfadenia Garay, Bot. Mus. Leafl., Harvard Univ., 23, 203 (1972). Type species: Seidenfadenia mitrata (Rchb.f.) Garay (basionym: Aerides mitrata Rchb.f.)

Derivation of name The name commemorates the Danish diplomat and orchid taxonomist Gunnar Seidenfaden (1908–2001), who was Danish Ambassador to Thailand from 1955 to 1959. (PC) 286

Description (Plate 113; Fig. 693.1) Epiphytic or lithophytic herbs. Roots stout, basal. Stem covered in sheathing leaf bases. Leaves pendent, semi-terete, sulcate, articulate to sheathing base. Inflorescence axillary, elongate, pendent to suberect, racemose, multiflowered, borne toward basal part of stem below leaves. Flowers resupinate, whitish with purple labellum. Sepals and petals free, subequal or petals smaller, spreading or reflexed. Labellum attached to column base, trilobed, side lobes horn-like, midlobe prominent, flat, callus keeled; spur laterally compressed, semi-lunate from the side, obtuse, callose within. Column fleshy, dilated toward base, clinandrium dorsal; pollinia two, sulcate, stipe linear-spatulate, viscidium prominent; rostellum erect, crest-shaped, bidentate. Capsule fusiform-clavate. (PC)

Distribution (Fig. 693.2) Seidenfadenia is monospecific and from Burma, Thailand, and Laos. (PC)

Palynology Schill and Pfeiffer (1977) reported that pollen tetrads of S. mitrata (studied as Aerides mitrata) were convex with laevigate sculpturing and a rounded sexine. (AP)

Cytogenetics Felix and Guerra (2010) listed a chromosome count of 2n = 38 for S. mitrata. (AP)

SEIDENFADENIA

O

P

C

B

I

M

H N

L

K

D

F

A

J

E

G

Fig. 693.1.  Seidenfadenia mitrata (Rchb.f.) Garay. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum, from above; H. Labellum, longitudinal section; I. Column and labellum, side view; J. Column, front view; K. Column, side view; L. Column apex, side view; M. Anther cap, dorsal view; N. Anther cap, ventral view; O. Pollinia; P. Viscidium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Kerr 0921 (K) and Kew Spirit Collection no. 26591.

287

VA N D E A E

Fig. 693.2.  Distribution map of Seidenfadenia.

Phytochemistry

Taxonomic notes

Alkaloids are present in S. mitrata (as Aerides mitrata) at levels of 0.001–0.01% (Lüning 1964). (NV, RG)

Although originally referred to Aerides, Garay (1972) suggested that this orchid bears only a passing resemblance to that genus because of its saccate spur that projects obliquely forward beneath the labellum. Its column and pollinia are reminiscent of those of Rhynchostylis, but its habit, clinandrium, stigma, and shape and position of the rostellum are distinctive. The crestshaped rostellum that projects across the clinandrium and protrudes dorsally on the column is a unique feature. (PC)

Phylogenetics In clade F, Seidenfadenia is sister to the rest, with the exception of Rhynchostylis. In Topik et al. (2005), Seidenfadenia is sister to the same groups but also including Aerides, which is likely due to the sparser generic sampling in that study. (AK, MC)

Ecology On trees or rocks in tropical deciduous forest; 300–1050 m. (PC)

Pollination There is no information available on pollination of Seidenfadenia. (AS)

Uses No uses have been reported for Seidenfadenia; it is occasionally cultivated. (AS)

Cultivation Group 2 (see subtribal treatment). (AS) 288

Taxonomic literature Garay, L. A. (1972). On the origin of Orchidaceae. II. Journal of the Arnold Arboretum, 53, 202–15. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

6 9 4 .  S EI DENFA DENI EL L A Seidenfadeniella C.S.Kumar, Cat. Indian Orchids, 43 (1994). Type species: Seidenfadeniella rosea (Wight) C.S.Kumar (basionym: Sarcanthus roseus Wight)

Derivation of name The name commemorates Danish diplomat and orchid taxonomist Gunnar Seidenfaden (1908–2001), who was Danish

SEIDENFADENIELLA

Ambassador to Thailand from 1955 to 1959, and the Latin diminutive -ella. (PC)

Description (Fig. 694.1) Epiphytic herbs. Roots basal. Stem elongate, covered in sheathing leaf bases. Leaves terete, aciculate, articulate to sheathing base. Inflorescence axillary, pendent or spreading, racemose, multiflowered; bracts triangular, recurved. Flowers resupinate. Sepals free, subequal, spreading, narrowly elliptic, rounded at apex. Petals broadly obovate-subcircular, rounded at apex. Labellum attached to column base, trilobed, side lobes erect, rounded at

tip, midlobe prominent, porrect-deflexed, ovate, callus keeled; spur inflated, with a thickened back wall and horizontal bridge. Column with two cleft pollinia, stipe linear, transparent; rostellum with two, two-pronged arms, the lower pointing downward and almost covering stigma, the upper pointing upward. (PC)

Distribution (Fig. 694.2) Seidenfadeniella is a genus of only two species—S. rosea and S. filiformis (Rchb.f.) Christenson & Ormerod—from southern India and Sri Lanka. (PC)

B

C

H L

D

F

E

I

K A

J

G

Fig. 694.1.  Seidenfadeniella filiformis (Rchb.f.) Christenson & Ormerod. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum; H. Column and labellum, side view; I. Column, from above; J. Column, side view; K. Anther cap, dorsal view; L. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Bear Shola 296 (K).

289

VA N D E A E

Fig. 694.2.  Distribution map of Seidenfadeniella.

Cytogenetics

Cultivation

A count of n = 19 has been recorded for the species (Vatsala 1964; Abraham and Vatsala 1981). (AP)

Group 2 (see subtribal treatment). (AS)

Phytochemistry

Taxonomic notes

No reports have been found on the phytochemistry of Seidenfadeniella. (NV, RG)

DNA sequences are not as yet available for Seidenfadeniella. The genus is distinguished by its characteristic, branched rostellum. (PC)

Phylogenetics

6 9 5 .  S I NGC H I A

There is no phylogenetic information available for this genus. (AK)

Ecology Both species are epiphytic in stunted evergreen tropical and subtropical moist broadleaf forests at 1000–2600 m. (PC)

Pollination There is no information available on pollination of Seidenfadeniella. (AS)

Uses No uses have been reported for Seidenfadeniella; it is virtually unknown in cultivation. (AS) 290

Singchia Z.J.Liu & L.J.Chen, J. Syst. Evol., 47, 600 (2009). Type species: Singchia malipoensis Z.J.Liu & L.J.Chen

Derivation of name The name commemorates Chinese orchid taxonomist, Professor Chen Singchi, emeritus professor at the Chinese Academy of Sciences. ( JW)

Description (Fig. 695.1) Epiphytic herbs. Stem 6–8 cm long, suberect. Leaves four or five, narrowly oblong, obtusely unequally bilobed, with a mucro in the sinus, flat, thickly coriaceous. Inflorescence lateral, up to 4 cm long, 9- to 12-flowered; floral bracts minute, ovate. Flowers white, sepals and petals tinged pale purple dorsally, spur white tinged pale

SINGCHIA

H G B K J

I

C

D A

F

E

Fig. 695.1.  Singchia malipoensis Z.J.Liu & L.J.Chen. A. Habit; B. Flower, front view; C. Flower with sepal and petal removed, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Column and labellum, side view; H. Column and labellum with lateral lobe artificially recurved; I. Column and labellum, longitudinal section; J. Pollinarium, front view; K. Pollinarium, back view. Single bar = 1 mm, double bar = 1 cm. Redrawn by Judi Stone after Z. J. Liu and L. J. Chen, Journal of Systematics and Evolution, 47, 602, Fig. 2 (2009).

purple. Dorsal sepal and petals free. Lateral sepals adnate to base of column foot. Labellum spurred, trilobed, the basal margins rigidly attached to lower part of column foot, side lobes almost as long as midlobe, subtriangular to semi-orbicular, obtuse, midlobe broadly ovate, with a saddle-shaped basal callus; spur subconical-cylindrical, thin-walled, ecallose. Column with a pendent foot; pollinarium cruciform, pollinia two, unequally and deeply split, caudicle distinct, stipe narrowly linear, viscidium broadly oblong-ovate. ( JW)

Cytogenetics

Distribution (Fig. 695.2)

Phylogenetics

Singchia is a monospecific and endemic to southwestern China. ( JW)

There is no phylogenetic information available for this genus. However, the authors of the original description proposed that

No chromosome numbers have been recorded for Singchia. ( JW)

Phytochemistry No reports have been found on phytochemistry of Singchia. (NV, RG)

291

VA N D E A E

Fig. 695.2.  Distribution map of Singchia.

it is closely related to Pteroceras, which is clearly polyphyletic here, so this is probably difficult to evaluate. However, it may be well be a member of the Vanda clade based on its morphology. (AK, MC)

Ecology Singchia malipoensis is epiphytic in broad-leaved forests at 1000 m. ( JW)

Pollination There is no information available on pollination of Singchia. ( JW)

Uses No uses have been reported for Singchia; it is not known to be in cultivation. ( JW)

Cultivation Group 1 (see subtribal treatment). (MM)

Taxonomic notes Singchia has little in common with Pteroceras, with which Liu and Chen (2009) compared it; it appears more closely related to Vanda. As floral 292

morphology is only an imperfect guide to relationships in Aeridinae, molecular data are needed to determine the status of Singchia. (AS)

Taxonomic literature Liu, Z-J. and Chen, L-J. (2009). Singchia and Gunnaria, two new genera of Orchidaceae. Journal of Systematics and Evolution, 47, 599–604.

6 9 6 .  S M I T H S ONI A Smithsonia C.J.Saldanha, J. Bombay Nat. Hist. Soc., 71, 73 (1974). Type species: Smithsonia straminea C.J.Saldanha Micropera Dalzell, in Hooker, J. Bot. Kew Gard. Misc., 3, 282 (1851), nom. illeg. (non Micropera Lindl. 1832). Type species: not designated. Loxoma Garay, Bot. Mus. Leafl., Harvard Univ., 23, 183 (1972), nom. illeg. (non Loxsoma R.Br. ex Cunn.). Type species: Loxoma maculatum (Dalzell) Garay (basionym: Micropera maculata Dalzell) Loxomorchis Rauschert, Taxon, 31, 561 (1982). Type species: Loxomorchis maculata (Dalzell) Rauschert (basionym: Micropera maculata Dalzell)

Derivation of name The generic name commemorates James Smithson (1754–1829), founder of the Smithsonian Institution (Washington, D.C., USA). (CSK)

SMITHSONIA

Description (Plate 114; Fig. 696.1) Epiphytic herbs. Roots many, arising from base of stem, up to 100 or more cm long, 0.4–0.5 cm in diameter, sometimes flattened and closely adhering to trunk or branch. Stem with distichous leaves. Leaves 2–6, 2.5–10.0 × 0.5–2.4 cm, oblong to linear-oblong or elliptic-lanceolate, sometimes slightly falcate, articulate, sheathing at base, green or rarely mottled with purple or dark green, unequally or obtusely bilobed at apex with (S. straminea) or without a sharp median mucro in between. Inflorescence spreading or pendent, elongate or (sub)umbellate raceme with up to 20 flowers, 5–15 cm long, axillary; bracts 2 mm long, broadly oblong-ovate or ovatetriangular, acute or obtuse. Flowers c. 0.6–1.2 cm wide, resupinate,

greenish to yellow, either with a solitary reddish brown spot (S. maculata (Dalzell) C.J.Saldanha), one or two violet spots (S. straminea) or without spots on sepals and petals, labellum white, often marked with pink or violet. Dorsal sepal obovate, obovate-oblong, or oblanceolate, often narrowed at base and gland-dotted, 3–5-veined, obtuse or acute at apex. Lateral sepals obovate or broadly obovate to obovate-oblong, narrowed at base (S. maculata) or not, 3–5-veined, gland-dotted (S. viridiflora (Dalzell) C.J.Saldanha) or not, obtuse at apex. Petals obovate, oblong or obovate-oblong, 3–5-veined, obtuse at apex. Labellum white, trilobed, shortly or deeply saccate at base, and immovably attached to column base, side lobes triangular, erect, sometimes connate below, narrow and recurved,

M

L

N

O

D A F H E K

G

C I B J

Fig. 696.1.  Smithsonia maculata (Dalzell) C.J.Saldanha. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum, from above; H. Labellum, longitudinal section; I. Column and labellum, side view; J. Column and labellum from above, anther cap removed; K. Column, side view; L. Anther cap and pollinarium; M. Anther cap, ventral view; N. Pollinarium; O. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Dalzell s.n. (K).

293

VA N D E A E

midlobe entire or weakly trilobed; spur subconical, straight, 3–4 mm long, naked within or with a tuft of white hairs (S. maculata). Column 1–2 mm long, semi-terete, without a foot; anther conical, pollinia two, deeply and unequally cleft, stipe narrow, viscidium oblong, bifid; rostellum bilobed. Ovary with pedicel 3–7 mm long. Capsule 3–5 cm long, oblong or fusiform. (CSK)

Distribution (Fig. 696.2)

Ecology The species of this genus occur as epiphytes at 50–950 m. (CSK)

Pollination No information is available on the pollination of Smithsonia. (CSK)

Smithsonia is a genus of four species endemic to India (Western Ghats from Maharashtra, Karnataka, Goa, Tamil Nadu, and Kerala). (CSK)

Uses

Palynology

Cultivation

Nothing is known about the pollen of Smithsonia. (CSK)

Cytogenetics Felix and Guerra (2010) listed a chromosome count of 2n = 38 for both S. maculata and S. viridiflora. (AP)

Phytochemistry No reports have been found on the phytochemistry of Smithsonia. (CSK, NV, RG)

Phylogenetics There is no phylogenetic information available for this genus. (AK)

No uses have been recorded for Smithsonia. (CSK)

Group 1 (see subtribal treatment). Species of this genus are easy to cultivate. When tied to slabs of burnt wood or coconut husk and hung, plants are easily grown in tropical climates. Gurukula Botanical Garden at Wayanad and Jawaharlal Nehru Tropical Botanic Garden & Research Institute at Trivandrum, India, grow all the species. (CSK, MM)

Taxonomic notes The first two species of the genus were described by Dalzell in 1851 as Micropera maculata Dalzell and M. viridiflora Dalzell but without a generic description. Also, Lindley had already used that generic name in 1832, making Micropera Dalzell a later homonym. Since then they have been transferred to Sarcochilus by Pfitzer,

Fig. 696.2.  Distribution map of Smithsonia.

294

SPONGIOLA

Saccolabium by J. D. Hooker, and Gastrochilus by Kuntze. Garay (1972) found Micropera maculata distinct from Gastrochilus in having a reclining column, short column foot, and distinctly divided pollinia. Unfortunately, Garay’s name Loxoma for the new genus was a later homonym of that of the fern genus Loxsoma R.Br. ex Cunn. At the same time, Garay transferred Micropera viridiflora to Aerides, believing it to be close to A. japonica Rchb.f. This confusion was cleared up by Saldanha (1976), who created a new genus Smithsonia typified by a new species, S. straminea C.J.Saldanha, and transferred Dalzell’s two Micropera species to it. A fourth species, S. saldanhae C.S.Kumar & Theuerkauf (Kumar and Theuerkauf, in press), was recently described from the Western Ghats, where the genus is endemic. (CSK)

Taxonomic literature Garay, L. A. (1972). On the systematics of the monopodial orchids I. Botanical Museum Leaflets, Harvard University, 23, 149–212. Kumar, C. S. and Theuerkauf, W. D. (In press). A new species in the genus Smithsonia C.J.Saldanha (Orchidaceae). Rheedea. Pande, S., Sant, N., Vishwasrao, V., and Datar, M. (2010). Wild orchids of northern Western Ghats. Ela Foundation, Pune 411 109, India. Saldanha, C. J. (1976). Orchidaceae. Flora of Hassan District, Karnataka, India (ed. C. J. Saldanha and C. J. Nicolson), pp. 850–52. Amerind Publishing Co., New Delhi.

697.   SMITIN A N D IA

Cytogenetics Feliz and Guerra (2010) listed a chromosome number of 2n = 38 for S. micrantha. (AP)

Phytochemistry No reports have been found on the phytochemistry of Smitinandia. (NV, RG)

Phylogenetics Smitinandia is a member of clade L and is closely allied with Micropera, Uncifera, and one species of Stereochilus (which receives high posterior probability but low parsimony bootstrap support). However, Topik et al. (2005) placed Smitinandia together with some species of Cleisostoma, Pelatantheria, and Schoenorchis, but that result could be due to different sampling. (AK, MC)

Ecology Smitinandia helferi (Hook.f.) Garay is epiphytic in mixed deciduous forests in Thailand. Smitinandia micrantha grows in riverine forests on limestone and ultramafic substrate and secondary forests in subtropical valleys. It is also epiphytic on Syzygium Gaertn. (Myrtaceae) and Mangifera L. (Anacardiaceae) in India. Both species occur at 0–1000 m. ( JW)

Smitinandia Holttum, Gard. Bull. Singapore, 25, 105 (1969). Type species: Smitinandia micrantha (Lindl.) Holttum (basionym: Saccolabium micranthum Lindl.)

Pollination

Derivation of name

Uses

Nothing is known about pollination in Smitinandia. ( JW)

The name commemorates Thai forest botanist Tem Smitinand (1920–1995), associated with the Forest Herbarium (BKF) and co-author with Gunnar Seidenfaden of their account of the orchids of Thailand. ( JW)

Pant (2011) reported the use of a powder made from the roots of S. micrantha as a tonic in Nepal and noted that the stems are considered to have antiseptic properties. Species of Smitinandia are sometimes cultivated. (AS)

Description (Plate 115; Fig. 697.1)

Cultivation

Erect to pendent epiphytic herbs. Stem simple or branching, compressed, stout, covered by leaf-sheaths. Leaves distichous, sessile, articulate, narrow, ligulate, apex truncate, unequally bilobed, or acute, distant. Inflorescence lateral, racemose, many-flowered. Flowers with sepals and petals white flushed rose-pink or greenish yellow, labellum purple or white. Sepals and petals spreading, petals narrower than sepals. Labellum spurred, lacking ornaments, trilobed, side lobes triangular, midlobe oblong or rhomboid, obtuse, margin sometimes erose; spur with a mouth closed by a high transverse wall at base of midlobe. Column cylindric, broadened at base, foot not distinct; pollinia four, in unequal pairs, the smaller half in each pair detaching as a free flat disc, stipe broadened upwards, about twice the length of diameter of pollinia; viscidium ovate. ( JW)

Group 2 (see subtribal treatment). (MM)

Distribution (Fig. 697.2) Smitinandia consists of three species distributed in Bangladesh, India, Nepal, Bhutan, Burma, Thailand, Laos, Cambodia, Vietnam, Peninsular Malaysia, Borneo, and Sulawesi. ( JW)

Taxonomic literature Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

6 9 8 .   S P ONGI OL A Spongiola J.J.Wood & A.L.Lamb in Chan et al., Orchids of Borneo, 1, 283 (1994). Type species: Spongiola lohokii J.J.Wood & A.L.Lamb

Derivation of name From the Latin spongia, sponge, in reference to the form of the labellum midlobe. ( JW) 295

VA N D E A E

H C A

2 mm

I 5 mm 5 cm

G

2 mm

D

E F

B 5 mm

2 mm

Fig. 697.1.  Smitinandia micrantha (Lindl.) Holttum. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Pedicel with ovary, labellum, and column, longitudinal section; H. Anther cap, dorsal view; I. Pollinarium. Drawn by C. L. Chan and Liew Fui Ling from Lamb AL 1148/89. Reproduced with permission from J. J. Wood (2003), Orchids of Borneo. Volume 4. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

Description (Fig. 698.1) Pendent epiphytic herbs. Stem leafy, obscured by persistent leaf bases, producing roots at the nodes. Leaves oblong or oblongelliptic, apex obtuse and asymmetric, coriaceous. Inflorescence lateral, racemose, occasionally branched, pendent, much longer than leaves, up to about 25-flowered, usually only five or six open at a time, progressing downward in succession, sometimes opening in middle of raceme; floral bracts minute, ovate. Flowers resupinate, short-lived, each lasting one day only, sepals and petals semi-translucent pale yellow to yellowish green, margins and tips semi-translucent whitish, labellum white with pale to dark purple spots, side lobes flushed purple on inner surface, spur pale yellow. Sepals and petals free, oblong-elliptic, obtuse, spreading. Labellum rigidly attached to column base, trilobed, spurred, ecallose, side lobes auriculate, midlobe longer than side lobes and resembling a sponge-like pouch, hollow near mouth of spur, solid below, shallowly concave on underside, papillose-verrucose; spur 2 296

mm long, conical, obtuse, aseptate. Column without foot; anther cap terminal, operculate, apex acuminate, sometime obscurely tridentate, pollinia two, ovoid, entire, stipe spatulate, broadened below pollinia, viscidium cucullate; rostellum prominent. Ovary with pedicel terete, straight. ( JW)

Distribution (Fig. 698.2) Spongiola is monospecific and endemic to Borneo (Sabah only). ( JW)

Cytogenetics No chromosome counts for Spongiola have been recorded. ( JW)

Phytochemistry No reports have been found on phytochemistry of Spongiola. (NV, RG)

STEREOCHILUS

Fig. 697.2.  Distribution map of Smitinandia.

Phylogenetics

Group 1 (see subtribal treatment). (MM)

Porrorhachis, Robiquetia, and Tuberolabium among others. It is distinguished from those by the spongy, pouched midlobe of the labellum. Labellum structure is relatively simple compared with many genera in Aeridinae, having neither calli nor a septate spur. Its habit is similar to that of Brachypeza, particularly B. archytas (Ridl.) Garay from Christmas Island near Java, and certain species of Pteroceras, Robiquetia, and Sarcochilus. Spongiola is distinguished from Robiquetia by the large sponge-like midlobe of the labellum and the long, slender rostellar projection. In Robiquetia the midlobe is small, conical or linear, never spongelike, and the rostellum is short. The labellum of Spongiola is not articulate on a long column foot as in Grosourdya, Macropodanthus, Pteroceras, and Sarcochilus. Grosourdya may also be distinguished by the long, bent column and the labellum midlobe bent downward and appressed to the prominent spur. In Macropodanthus, the long-lasting flowers open simultaneously and are distinguished by the labellum being bent upward so as to form a right angle with the long column foot. The labellum midlobe of Pteroceras is much smaller than the side lobes, and the two pollinia are sulcate into subequal halves. In Sarcochilus, the flowers are longer lasting and have a callose labellum and four unequal pollinia. Generic delimitation of the genera closely allied to Pteroceras, including Spongiola, needs further study. Spongiola is only tentatively accepted here. (JW, AS)

Taxonomic notes

6 9 9 .   S T ER EOC H I LU S

Spongiola belongs to the group of genera in Aeridinae having a column without a foot and two pollinia, including Pennilabium,

Stereochilus Lindl., J. Proc. Linn. Soc., Bot., 3, 38 (1859). Type species: Stereochilus hirtus Lindl.

Spongiola is a member of a well-supported subclade of clade E, which includes one species of Biermannia plus Pennilabium and Chroniochilus. (AK, MC)

Ecology This species is epiphytic on the branches of small trees in damp, shaded and humid situations in riverine forest on limestone and sandstone ridges at elevations at 300–500 m. ( JW)

Pollination Nothing is known about pollination in Spongiola. ( JW)

Uses No uses have been reported for Spongiola, and it is not commonly cultivated. ( JW)

Cultivation

297

VA N D E A E

J A 1 mm 5 cm

E D 4 mm

F

C

2 mm

H I

2 mm

B

5 mm

G

2 mm

Fig. 698.1.  Spongiola lohokii J.J.Wood & A.L.Lamb. A. Habit; B. Flower, front view; C. Lateral sepal; D. Dorsal sepal; E. Petal; F. Labellum; G. Column and labellum, longitudinal section; H. Anther cap, dorsal view; I. Pollinarium. J. Ovary, transverse section. Drawn by Chan Chew Lun from Lamb AL 426/85 (holotype, K). Reproduced with permission from Chan et al. (1994), Orchids of Borneo. Volume 1. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

Derivation of name From the Greek stereos, solid, and cheilos, lip. ( JW)

Description (Plate 116; Fig. 699.1) Epiphytic herbs. Stem suberect. Leaves several to many, distichous, coriaceous, articulate to a sheathing base, deciduous, duplicate. Inflorescences lateral, racemose, many-flowered, often pendent; peduncle and rachis sometimes glandular-pubescent; floral bracts shorter than ovary and pedicel. Flowers resupinate, often widely opening. Sepals and petals spreading. Lateral sepals adnate to base of labellum. Petals smaller than sepals. Labellum rigidly fused to column base, saccately spurred at base, indistinctly trilobed, side lobes erect, midlobe larger; spur with longitudinal septum inside and often one 298

or two calli on back wall. Column erect, foot absent; pollinia four, arranged in two nearly equal pairs, ellipsoid-oblong to ovoid, waxy, with short but distinct caudicles, attached by a clavate stipe to an ovate viscidium; rostellum subulate-lanceolate, porrect. ( JW)

Distribution (Fig. 699.2) Stereochilus comprises seven species collectively distributed in northeastern India, Bhutan, Burma, Thailand, Vietnam, and China. ( JW)

Cytogenetics Felix and Guerra (2010) listed a chromosome count of 2n = 38 for S. erinaceus (Rchb.f.) Garay. (AP)

TAENIOPHYLLUM

Fig. 698.2.  Distribution map of Spongiola.

Phytochemistry

Cultivation

A survey of alkaloid distribution in Orchidaceae gave a negative result for S. hirtus (Lüning 1964). (NV, RG)

Group 2 (see subtribal treatment). (MM)

Phylogenetics

The taxonomy of the genera closely allied to Cleisostoma, such as Pelatantheria, Rhynchogyna, Sarcoglyphis, Sarcophyton, Stereochilus, and others is still unresolved. Within this alliance, Stereochilus may be recognized by the slender column and elongate rostellum. (AS)

Stereochilus is a member of the highly problematic clade L, within which Cleisostoma is grossly polyphyletic. Stereochilus is at least biphyletic, and the eight species included thus far are closely related to some species of Cleisostoma, Micropera, Rhynchogyna, Sarcoglyphis, and Uncifera (this was also found by Carlsward et al. 2006b). Clade L needs a great deal more work to sort out the status of these genera. (AK, MC)

Ecology Species are epiphytes in forests at 300–1800 m. ( JW)

Pollination There is no information available on pollination of Stereochilus. ( JW)

Uses No uses have been reported for Stereochilus; it is occasionally cultivated. ( JW)

Taxonomic notes

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogeny of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86.

7 0 0 .   TA ENI OP H YL LU M Taeniophyllum Blume, Bijdr., 355 (1825). Type species: Taeniophyllum obtusum Blume Alwisia Thwaites ex Lindl., J. Proc. Linn. Soc., Bot., 3, 42 (1858). Type species: Alwisia minuta Thwaites ex Lindl. Cryptorchis Makino, Bot. Mag. (Tokyo), 7, 118 (1893), nom. invalid. Type species: Cryptorchis aphylla Makino Geissanthera Schltr. in K.M. Schumann and C.A.G. Lauterbach, Nachtr. Fl. Deutsch. Schutzgeb. Südsee, 231 (1905). Type species: Geissanthera papuana Schltr. 299

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Fig. 699.1.  Stereochilus brevirachis Christenson. A. Habit; B. Flower, front view; C. Flower, side view; D. Rachis, close-up; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum, from above; I. Labellum, longitudinal section; J. Column and labellum, side view; K. Column, side view; L. Column, anther cap removed, front view; M. Anther cap, dorsal view; N. Anther cap, ventral view; O. Pollinarium; P. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from photograph in Orchid Digest, 62, 123 (1998) and Kew Spirit Collection no. 60360.

Microtatorchis Schltr. in K.M. Schumann and C.A.G. Lauterbach, Nachtr. Fl. Deutsch. Schutzgeb. Südsee, 224 (1905). Type species: Microtatorchis perpusilla Schltr. Ankylocheilos Summerh., Bot. Mus. Leafl., Harvard Univ., 11, 168 (1943). Type species: Ankylocheilos coxii Summerh.

Derivation of name From the Greek taenio-, originally a hair band, but applied to the tapeworm, and phyllon, leaf, in reference to the flattened, photosynthetic roots. ( JW) 300

Description (Plates 117, 118; Fig. 700.1, 700.2) Epiphytic or lithophytic herbs. Roots flattened and ribbon-like or rarely subterete, appressed to substrate or pendent and spreading, green and photosynthetic in rainy season, whitish or greyish in dry season. Stem inconspicuous. Leaves absent or rudimentary and reduced to tiny brown scales, or basal, distichous, few, articulate to sheath, blade usually oblong-obovate, membranous, conduplicate in bud, deciduous. Inflorescence lateral, racemose, glabrous or muricate, few- to many-flowered; rachis slowly elongating, terete or flattened, bearing flowers in

TAENIOPHYLLUM

Fig. 699.2.  Distribution map of Stereochilus.

succession, one or two at a time; floral bracts alternate, persistent, sometimes with aristate, stipule-like appendages on both sides decurrent to form wings on peduncle and rachis. Flowers distichously arranged, ephemeral or lasting only a few days, usually resupinate, pale green to yellowish white. Sepals and petals free and widely spreading or connate at base or below middle into a tube, this fissured on the side with labellum, the free portions erect or weakly spreading, slightly dissimilar. Labellum entire or trilobed, rigidly attached to base of column, spurred or saccate, disc naked or with a single callus; spur globose to cylindrical, often retrorse, with the opening often partially or entirely covered by a hyaline septum, frequently with a globular or dactyliform appendage within, base often concave, lateral margins inflexed, apex subacute to acute and sometimes provided within with a reflexed tooth or bristle-like appendage, or truncate and unappendaged. Column usually with two lobes on adverse sides of apex projecting into concave base of labellum, foot absent, clinandrium shallowly excavate; anther cap proclined, often beaked; pollinia two or four, free, waxy, subglobular, often obovoid or ellipsoid, borne on a common stipe frequently dilated distally, viscidium oblong or elliptic; stigma transverse, deeply recessed; rostellar projection apex bifid. ( JW)

Distribution (Fig. 700.3) There are 236 species of Taeniophyllum distributed from tropical Africa (only one species) through tropical Asia—Sri Lanka,

India, China, and Japan southeast into Malesia, and from northern Australia, New Zealand, Micronesia, and the Pacific islands east to the Austral Islands. ( JW)

Anatomy Carlsward et al. (2006a) examined six species of Taeniophyllum, including T. iboetii (J.J.Sm.) Kocyan & Schuit. (studied as Microtatorchis iboetii J.J.Sm.) Their results are summarized below.

Root Velamen absent to two cells thick. Epivelamen cells isodiametric to tangentially flattened. Endovelamen cells angular, isodiametric to radially elongate. Distinct endovelamen thickenings absent or ridged in some species. Cover cells absent. Hairs present in T. biocellatum J.J.Sm., T. fasciola (G.Forst.) Seem. and T. iboetii. Exodermal cells radially elongate to isodiametric. Long cell walls ∩-thickened in T. fasciola and T. smithii Kores & L.Jonss., ○-thickened in T. biocellatum and M. iboetii. Proliferations present in T. biocellatum and T. iboetii. Cortex 4–24 cells wide. Cells isodiametric to radially and tangentially elongate. Starch grains and hyphae present in T. biocellatum. Water-storage cell walls birefringent and smooth to pitted in T. biocellatum, T. fasciola, T. smithii, and T. iboetii, with birefringent bands in T. smithii. Aeration units present or absent depending on species. Endodermal cells ○-thickened. Pericyclic cells thin-walled opposite xylem and thick-walled opposite phloem. Vascular cylinder 3–8-arch. 301

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Fig. 700.1.  Taeniophyllum coxii (Summerh.) Summerh. A. Habit, ×4.5; B. Inflorescence, ×9; C. Flower, side view, ×12; D. Flower (enlarged), ×21; E. Sepals and petals, ×21; F. Column (anther cap removed) and labellum, side view, ×21; G. Column (anther cap removed) and labellum from above, ×21; H. Anther cap and pollinia, ×21. Drawn by Judi Stone from Kew Spirit Collection no. 51549.

Xylem arms without distinct metaxylem elements in T. fasciola. Vascular tissue embedded in thick-walled sclerenchyma. Pith sclerenchymatous. (BC)

explaining the 2n = 24, 36 counts, or x = 19, yielding 2x = 38, 4x = 56. Felix and Guerra (2010) noted that x = 19 is the probable basic number. (AP)

Cytogenetics

Phytochemistry

Brandham (1990) listed chromosome numbers of 2n = 24, 36, 38, 40, 56 and suggested two basic numbers, either x = 6 or 12,

According to Lüning (1974), three out of ten species tested for alkaloids contained them at levels ≥ 0.1%. However, named

302

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Fig. 700.2.  Taeniophyllum fasciola (G.Forst.) Seem. A. Habit; B. Inflorescence; C. Flower; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum, from above; H. Labellum, longitudinal section; I. Column and labellum, side view; J. Column, front view; K. Column, side view; L. Column, anther cap removed, front view; M. Column, anther cap removed, back view; N. Anther cap, dorsal view; O. Anther cap, ventral view; P. Pollinarium; Q. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Kew Spirit Collection no. 53141.

records are few. Alkaloids were not detected in T. lobatum Dockrill (as T. flavum Dockrill). Results for T. muelleri Lindl. ex Benth. are conflicting; a specimen examined did not contain alkaloids, whereas another cited under its synonym, T. wilkianum T.E.Hunt, gave a positive result (Lawler and Slaytor 1969). (NV, RG)

Phylogenetics The species of Taeniophyllum are generally leafless. However, the species of Microtatorchis have leaves, and the two included Microtatorchis species occur at different positions within

Taeniophyllum (clade D), which means that presence/absence of leaves is evolutionarily labile. Here, we include Microtatorchis in Taeniophyllum. Taeniophyllum was previously subdivided into two subgenera and nine sections. However, due to the lack of living study material, Taeniophyllum s.l. is far from being well understood. (AK, MC)

Ecology Species are epiphytic, lithophytic, or rarely terrestrial in subtropical and tropical wet evergreen forests and subtropical forests, dry 303

VA N D E A E

Fig. 700.3.  Distribution map of Taeniophyllum.

forests, mangrove and swamp forests, rubber plantations, and hill forests on ultramafic substrate at 0–3600 m. ( JW)

Pollination Little is known about pollination of Taeniophyllum. Some species such as T. hasseltii Rchb.f. are autogamous (van der Cingel 2001). (AP)

Uses Parham (1943) recorded that the Fijian islanders considered T. parhamiae L.O.Williams (= T. fasciola) of medicinal value. The photosynthetic roots are crushed and rubbed on the affected area in treating lumbago and other pains of the lower back, and a drink made of the plant is used internally for abdominal pains. Taeniophyllum is rare in cultivation. ( JW)

Cultivation Group 2 (see subtribal treatment), but some of the New Guinea species at higher elevations require more constantly humid and cool conditions. (AS, MM) 304

Taxonomic literature Carlsward, B. S., Whitten, M., Williams, N. H, and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Carr, C. E. (1932) The genus Taeniophyllum in the Malay Peninsula. The Gardens’ Bulletin Singapore, 7, 61–86. Kocyan, A. and Schuiteman, A. (In press). New combinations in Aeridinae. Phytotaxa. Seidenfaden, G. (1988) Orchid genera in Thailand XIV: fifty-nine vandoid genera. Opera Botanica, 95, 1–398. Senghas, K. (1990). Taeniophyllum. In Rudolf Schlechter Die Orchideen, 3rd ed., (ed. F. G. Brieger, R. Maatsch, and K. Senghas), pp. 1412–18. Paul Parey, Berlin.

7 0 1 .  TA P ROBA NEA Taprobanea Christenson, Lindleyana, 7, 90 (1992). Type species: Taprobanea spathulata (L.) Christenson (basionym: Epidendrum spathulatum L.)

Derivation of name From Taprobanea, the ancient Greek name for Sri Lanka. (LG)

TH RIXSPERMUM

Description (Fig. 701.1) Climbing, epiphytic herb. Roots usually emerging from nodes near base or lower portion of stem. Stem climbing, with elongated internodes. Leaves many, distichous, rigid, slightly decurved or held horizontally to stem, elliptic, shortly sheathing at base, deciduous, apex unequally but smoothly bilobed. Inflorescence axillary, unbranched, racemose, few-flowered, much longer than leaves. Flowers showy, resupinate, widely opening, uniform bright sulphuryellow throughout. Sepals and petals free, similar, obovate, slightly reflexed. Labellum immobile, trilobed, with a short spur, side lobes erect and connate to midlobe, midlobe spreading. Column lacking a foot; pollinia two, porate, stipe spatulate, viscidium bilobed. (LG)

Distribution (Fig. 701.2) Taprobanea is monospecific and distributed in southern India and Sri Lanka. (LG)

Cytogenetics The diploid chromosome number for Taprobanea spathulata is 2n = 114 (Storey et al. 1963; Felix and Guerra 2010). Most taxa in Vandeae have a diploid chromosome number of 2n = 38, so Taprobanea spathulata is thought to be hexaploid. Abraham and Vatsala (1981) also recorded n = 38 for south Indian populations. (LG)

Phytochemistry No reports have been found on phytochemistry of Taprobanea. (NV, RG)

Phylogenetics Taprobanea spathulata was transferred into Vanda by Sprengel (1826) but elevated to generic rank by Christenson (1992). DNAbased phylogenetic analyses by Gardiner et al. (2013) have shown that the species falls well outside the Vanda clade. (LG)

Ecology Taprobanea spathulata is a common, vine-like plant growing in direct sunlight at low elevations, scrambling through low, drybrush vegetation (Jayaweera 1981). (AS, LG)

Pollination No pollinators have been recorded for the genus. (LG)

Uses Lawler (1984) reported that the crushed stems and leaves of T. spathulata were once used in India and Sri Lanka to treat skin diseases and scabies, and powdered leaves and flowers were used against various diseases. The genus is also involved in breeding vandaceous hybrids. (AS, LG)

Cultivation Group 3 (see subtribal treatment). Taprobanea spathulata is not commonly cultivated. The species grows as a scrambling vine

and needs full sun to bloom. It has been used in the breeding of vandaceous hybrids for producing pure, sulphur-coloured flowers. Taprobanea is cold-sensitive, which limits its horticultural value in subtropical areas. (MM, LG)

Taxonomic notes Formerly placed in Vanda, the habit of Taprobanea spathulata is distinctly different, growing as a tall, vine-like plant with long internodes. In addition, leaves of T. spathulata are unequally bilobed at their tips rather than premorse, and the perianth parts are not clawed as in most Vanda species. (LG)

Taxonomic literature Christenson, E. A. (1992). Notes on Asiatic orchids. Lindleyana, 7, 88–94. Gardiner, L. M., Kocyan, A., Motes, M., Roberts, D. L., and Emerson, B. C. (2013). Phylogenetic patterns in the genus Vanda and related genera (Orchidaceae). Botanical Journal of the Linnean Society, 173, 549–72. Motes, M. R. (1997). Vandas: their botany, history, and culture. Timber Press, Portland, Oregon. Sprengel, K. P. J. (1826). Caroli Linnei: Systema Vegetabilum. Ed. 16. Vol. 3. Sumptibus Librariae Dieterichianae, Göttingen.

7 0 2 .   T H R I X S P ER M U M Thrixspermum Lour., Fl. Cochinch., 519 (1790). Type species: Thrixspermum centipeda Lour. Dendrocolla Blume, Bijdr., 286 (1825). Type species: Dendrocolla hystrix Blume Saccochilus Blume, Fl. Javae, Praef.: viii (1828). Type species: not designated Orsidice Rchb.f., Bonplandia (Hannover), 2, 93 (1854). Type species: not designated Cylindrochilus Thwaites, Enum. Pl. Zeyl., 307 (1861). Type species: Cylindrochilus pulchellus Thwaits Ridleya (Hook.f.) Pfitzer, in Engl. & Prantl, Naturl. Pflanzenfam., Nachtr., 2, 16 (1900) . Type species: Ridleya notabilis (Hook.f.) Pfitzer (basionym: Sarcochilus notabilis Hook.f.) Cordiglottis J.J.Sm., Bull. Jard. Bot. Buitenzorg, sér. 3, 5, 95 (1922). Type species: Cordiglottis westenenkii J.J.Sm. Cheirorchis Carr, Gard. Bull. Straits Settlem., 7, 40 (1932). Type species: Cheirorchis breviscapa Carr Thylacis Gagnep., Bull. Mus. Hist. Nat. (Paris), sér. 2, 4, 599 (1932). Type species: not designated

Derivation of name From the Greek thrix, hair, and sperma, seed. ( JW)

Description (Plate 119; Fig. 702.1) Epiphytic, lithophytic, or rarely terrestrial herbs. Stem ascending, climbing or pendent, simple, either with several closely spaced leaves or many well-spaced leaves. Leaves distichously arranged, flat, conduplicate, coriaceous, terete or laterally compressed, articulate to a tubular, sheathing base. Inflorescence lateral, axillary, 305

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F Fig. 701.1.  Taprobanea spathulata (L.) Christenson. A. Habit; B. Flower; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum; G. Column and labellum, side view; H. Column, anther cap removed, front view; I. Column apex with anther cap, side view; J. Anther cap, ventral view; K. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Madras Herbarium 72932 (K), Matthew 24426 (K), and Herbarium Hookerianum 2798 (K).

borne on or between nodes of stem, perforating the leaf-sheaths, racemose, erect or ascending, few- to rarely many-flowered, a few opening at a time; rachis either fractiflex with the flowers biseriate, or straight and congested with flowers omnilateral; floral bracts laterally compressed or not, either distichous and persistent on a flattened rachis or facing all ways on terete rachis. Flowers 306

resupinate, mostly developing sequentially with 1–3 undergoing anthesis at a time, ascending, usually ephemeral, often fully open for only half a day, from a few mm to several cm across. Sepals and petals free, subsimilar, spreading. Lateral sepals decurrent along column foot. Labellum rigidly attached to column foot by a broad base, trilobed, concave, saccate but not truly spurred,

TH RIXSPERMUM

Fig. 701.2.  Distribution map of Taprobanea.

side lobes erect, midlobe frequently thick and fleshy distally, disc usually with a partly hairy or papillose front-wall apical callus or with one or more thickenings at various locations. Column sometimes winged, not dilated distally, foot well developed, usually broad, frequently concave, not sharply differentiated from base of labellum, clinandrium shallowly excavate; anther cap terminal, operculate, hardly or not rostrate, pollinia four, in two unequal pairs, waxy, semi-globular or linear-oblong, attached by a common stipe, viscidium solitary, often peltate; rostellum shortly bidentate; stigma directly below rostellum, deeply recessed. ( JW)

Distribution (Fig. 702.2) Thrixspermum includes about 168 species collectively distributed from Sri Lanka and the Himalayan region and Taiwan, southeast through Malesia to northern Australia and the Pacific islands, and east to Fiji and Samoa. The centre of distribution appears to be Sumatra. ( JW)

Cytogenetics Felix and Guerra (2010) listed chromosome numbers of several species of Thrixspermum as 2n = 38. Thrixspermum japonicum (Miq.) Rchb.f. has a chromosome number of 2n = 36, 38. ( JW)

Phytochemistry Alkaloids have been detected in T. amplexicaule (Blume) Rchb.f., T. centipeda (as T. arachnites (Blume) Rchb.f.), and T. calceolus (Lindl.)

Rchb.f. at levels of 0.01–0.1% (Lüning 1967); a later survey indicated that of six species tested, none contained alkaloids at levels ≥ 0.1% (Lüning 1974). Chemical composition data for the floral fragrance of T. arachnites indicate that two major components predominate, the furanoid form of cis-linalool oxide (46.0%) and (E)-nerolidol (37.0%). Kaiser (1993) also emphasized the contribution to the scent made by several minor components, including (E,Z)-2,4-decadienal, γ-decalactone, and uniquely among orchids, ethyl (Z)-4-decenoate and ethyl (E,Z)-2,4-decadienoate. (NV, RG)

Phylogenetics The exact position and relationships of Thrixspermum are different in the Bayesian and parsimony results. In the former, it is well supported as sister to Dimorphorchis, but this pair of genera comprises one of the four major clades in the large polytomy at the base of Aeridinae; in the parsimony analysis Dimorphorchis is sister to Chiloschista, and this pair is then sister to Thrixspermum in clade A. Clade A is weakly supported in the parsimony analysis and does not exist in the Bayesian analysis. With different sampling of genera, Topik et al. (2005) also found that Thrixspermum was related to Dimorphorchis but also to Microsaccus, Abdominea, Sarcochilus, and Cleisomeria; Carlsward et al. (2006b) produced yet a different result and had Thrixspermum related to Vanda. Three sections were previously recognized in Thrixspermum, two of which (T. sects. Thrixspermum and Dendrocalla) appear to be 307

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Fig. 702.1.  Thrixspermum tortum J.J.Sm. A. Habit; B. Leaf, adaxial surface; C. Flower with lateral sepal and petal removed, front view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum, longitudinal section; H. Ovary and column, anther cap removed, side view; I. Column, front view; J. Anther cap, dorsal view; K. Anther cap, ventral view; L. Pollinarium; M. Pollinia. Single bar = 1 mm; double bar = 1 cm. Drawn by Linda Gurr from Wood 818 (K) and Lamb s.n. (K). Reproduced with permission from J. J. Wood (2003), Orchids of Borneo. Volume 4. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

monophyletic, but T. section Katocalla may be embedded in one of these (Breitkopf and Kocyan, personal communication). Cordiglottis was always thought to be closely allied to Thrixspermum but was kept apart because of its terete leaves and labellum characters, which are variable in both genera. However, Thrixspermum species with terete leaves have been described, and the results presented here support inclusion of Cordiglottis in Thrixspermum. (AK, MC)

Ecology Species of Thrixspermum are epiphytes or lithophytes, rarely terrestrials, occurring in rain forests, peat swamp forests, roadside vegetation, and broad-leaved deciduous, riverine, and wet evergreen forests at 0–3400 m. ( JW) 308

Pollination Flowers are ephemeral, usually lasting for less than one day or two days at the most. The short duration of anthesis is compensated by gregarious flowering, similar to that found in Dendrobium crumenatum Sw. Carr (1928) reported that T. hystrix (Blume) Rchb.f. (as Dendrocolla alba Ridl.) is pollinated by small bees. The callus inside the labellum causes the insect to rise relative to the floor of the labellum and press its thorax against the viscidium; pollinia are attached as a result. More recently, Ong (2011) observed that this species (as T. trichoglottis (Hook.f.) Kuntze) is pollinated by Trigona and Halictus bees. (JW, AS)

Uses No uses have been reported for Thrixspermum. Several species are cultivated by amateur growers. ( JW)

TRACHOMA

Fig. 702.2.  Distribution map of Thrixspermum.

Cultivation Group 2 (see subtribal treatment). (MM)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogeny of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Kocyan, A. and Schuiteman, A. (In press). New combinations in Aeridinae. Phytotaxa. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

703.   TR AC H O MA Trachoma Garay, Bot. Mus. Leafl., Harvard Univ., 23, 207 (1972). Type species: Trachoma rhopalorrhachis (Rchb.f.) Garay (basionym: Dendrocolla rhopalorrhachis Rchb.f.)

Derivation of name From the Greek trachoma, roughness, in reference to the ‘rough’ inflorescence with its remnants of densely spaced floral bracts. (JW)

few-flowered, flowers borne in small clusters throughout the year. Flowers resupinate, ephemeral. Sepals and petals free, spreading. Sepals slightly longer than petals, similar in shape. Labellum rigidly attached to base of column, laterally compressed, calceoliform, slightly spurred or saccate, interior ecallose, tuberculose or pulvinate-thickened in front. Column lacking a foot; pollinia two, globose, entire or obscurely sulcate, stipe linear, viscidium distinct; stigma suborbicular; rostellum erect, bifid after removal of pollinarium. ( JW)

Distribution (Fig. 703.2) Trachoma is a genus of 14 species collectively distributed in India, Peninsular Malaysia, Thailand, and Indochina, eastward through Indonesia to the Philippines, New Guinea, Australia, Solomon Islands, Vanuatu, New Caledonia, Fiji, Niue, Cook Islands, and Austral Islands (including Rapa). ( JW)

Cytogenetics Brandham (1999) listed published a chromosome count of 2n = 38 for the genus. ( JW)

Description (Plate 120; Fig. 703.1)

Phytochemistry

Epiphytic herbs. Stem leafy. Leaves coriaceous, linear-falcate, distichous. Inflorescence axillary, racemose, rachis distinctly fleshy,

No reports have been found on phytochemistry of Trachoma. (NV, RG) 309

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Fig. 703.1.  Trachoma rhopalorrhachis (Rchb.f.) Garay. A. Habit; B. Inflorescence; C. Flower; D. Bract; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum, from above; I. Labellum, longitudinal section; J. Column and labellum, side view; K. Column, front view; L. Column, side view; M. Column apex, anther cap removed, front view; N. Column apex, anther cap removed, side view; O. Anther cap, dorsal view; P. Anther cap, ventral view; Q. Pollinarium; R. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Comber 1234 (K) and Kew Spirit Collection no. 56546.

Phylogenetics

Pollination

Two species of Trachoma were sequenced for this project, and one of the two species of Porrorhachis, P. galbina (J.J.Sm.) Garay, is weakly supported as sister to T. stellatum M.A.Clem. & al.; the two genera are strongly supported as a subclade of clade E; other members of clade E include Amesiella, Ceratrocentrum, Cryptopylos, Dyakia, Macropodanthus, and Tuberolabium. (AK, MC)

Ong (2012) reported how a single individual of T. rhopalorrhachis was pollinated by a bee of the genus Halictus. Pollinia were deposited on the head of the bee as it forced its head into the mouth of the spur. However, although the flowers were pollinated, no fruit-set was observed afterwards, which would indicate that cross-pollination is required. (AS)

Ecology Species are epiphytes in forests at 0–1800 m. ( JW) 310

Uses No uses have been reported for Trachoma, but several species are cultivated. ( JW)

TRICHOGLOTTIS

Fig. 703.2.  Distribution map of Trachoma.

Cultivation Group 2 (see subtribal treatment). (MM)

Taxonomic notes The generic delimitation of the genera closely allied to Pteroceras, including Trachoma, needs further study. Trachoma is only tentatively accepted here. The flowers are similar to those of Tuberolabium, but in that genus the flowers open simultaneously along an elongate rachis, whereas in Trachoma they open in successive clusters on an abbreviated rachis. (AS)

Taxonomic literature Garay, L. A. (1972). On the origin of the Orchidaceae. II. Journal of the Arnold Arboretum, 53, 202–15. WCSP. (2013). World Checklist of Selected Plant Families. Facilitated by the Royal Botanic Gardens, Kew. Published on the Internet; http://apps.kew.org/wcsp/ Retrieved 18 March 2013. Wood, J. J. (1990). Notes on Trachoma, Tuberolabium and Parapteroceras (Orchidaceae). Nordic Journal of Botany, 10, 481–6.

Synptera Llanos, Fragm. Pl. Filip., 98 (1851). Type species: Synptera subviolacea Llanos Stauropsis Rchb.f., Hamburger Garten-Blumenzeitung, 16, 117 (1860). Type species: Stauropsis philippinensis (Lindl.) Rchb.f. (basionym: Trichoglottis philippinensis Lindl.) Staurochilus Ridl., J. Linn. Soc., Bot., 32, 351 (1896). Type species: Staurochilus fasciatus (Rchb.f.) Ridl. (basionym: Trichoglottis fasciata Rchb.f.) Sarothrochilus Schltr., Repert. Spec. Nov. Regni Veg., 3, 50 (1906). Type species: Sarothrochilus dawsonianus (Rchb.f.) Schltr. (basionym: Cleisostoma dawsonianum Rchb.f.) Ventricularia Garay, Bot. Mus. Leafl., Harvard Univ., 23, 210 (1972). Type species: Ventricularia tenuicaulis (Hook.f.) Garay (basionym: Saccolabium tenuicaule Hook.f.)

Derivation of name From the Greek thrix, trichos, hair, and glotta, tongue, in reference to the pubescent appendage on the back wall of the labellum spur or sac. ( JW)

704.   TR IC H O G LO TTIS

Description (Plates 121–123; Fig. 704.1, 704.2)

Trichoglottis Blume, Bijdr., 359 (1825). Type species: Trichoglottis retusa Blume Ceratochilus Blume, Bijdr., 358 (1825). Type species: Ceratochilus biglandulosus Blume

Climbing epiphytic herbs. Stem pendent or climbing, internodes elongate. Leaves many, distichously arranged, coriaceous, linear, lorate to elliptic, apex usually unequally bilobed, articulated to a sheathing base. Inflorescence axillary, either fasciculate with one to 311

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Fig. 704.1.  Trichoglottis biglandulosa (Blume) Kocyan & Schuit. A. Habit; B. Flower and bract; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum, from above; G. Labellum, longitudinal section; H. Column and labellum, side view; I. Column, front view; J. Column apex, anther cap removed; K. Column, side view; L. Anther cap, dorsal view; M. Anther cap with pollinia, ventral view; N. Pollinarium; O. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Kew Spirit Collection nos. 46635 and 72736.

several flowers in a cluster or a many-flowered raceme or panicle. Flowers opening widely, lasting about a week, resupinate, usually yellowish with light brown or purple markings. Sepals and petals free, petals slightly smaller than sepals. Labellum firmly fused to column, spurred or saccate at base, trilobed, side lobes erect, midlobe sometimes trilobed, often hairy or papillate; sac or spur often thickened, with a hairy ligulate appendage on back wall just below base of column, completely covering the entrance to the nectariferous cavity. Column cylindric, sometimes broadened at base so as to appear to have a foot-like extension, often with 312

hairy stelidia, foot absent; pollinia four, arranged in two pairs, unequal, solid, waxy, attached by a common linear-oblong stipe to an ovate or elliptic viscidium. ( JW)

Distribution (Fig. 704.3) There are 85 species of Trichoglottis collectively distributed from India, Sri Lanka, and the Nicobar Islands east to New Guinea, Australia, and the Solomon Islands, north to China, the Ryukyu Islands, and Thailand. The centre of diversity lies in Indonesia and the Philippines. ( JW)

TRICHOGLOTTIS

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Fig. 704.2.  Trichoglottis smithii Carr. A. Plant; B. Flower; C. Column and labellum; D. Dorsal sepal; E. Petal; F. Lateral sepal; G. Pollinarium; H. Anther, dorsal view; I. Anther, ventral view. Drawn by Chan Chew Lun and Jaap J. Vermeulen from cultivated material at Tenom Orchid Centre, Sabah. Reproduced with permission from Chan et al. (1994), Orchids of Borneo. Volume 1. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

Anatomy

Root

Carlsward et al. (2006a) examined T. atropurpurea Rchb.f. Their results are summarized below.

Velamen three cells thick. Epivelamen cells radially elongate. Endovelamen cells angular, isodiametric to radially elongate; cells in the outer layer thicker-walled than those of the inner layers. Distinct endovelamen thickenings absent. Cover cells present over short cells of exodermis. Exodermal cells radially elongate to isodiametric. Long cell walls primarily ○-thickened. Proliferations absent. Cortex 20 cells wide. Water-storage cells with birefringent finely banded walls, often slightly thickened. Aeration units present. Endodermal cells ○-thickened. Pericyclic cells thin-walled opposite xylem and thick-walled opposite phloem. Vascular cylinder 21-arch. Vascular tissue embedded in thick-walled sclerenchyma. Pith sclerenchymatous. (BC)

Leaf Cuticle smooth to ridged along the contours of the epidermal cells. Hairs rare, multicellular, glandular. Stomata ad- and abaxial. Epidermal cells periclinally oriented to isodiametric. Hypodermis absent. Fibre bundles absent. Mesophyll homogeneous. Waterstorage cells with birefringent, finely banded walls. Cells above midrib distinctly modified. Vascular bundles collateral, in one row. Sclerenchyma associated with both xylem and phloem poles. Bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with phloem and xylem sclerenchyma.

313

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Fig. 704.3.  Distribution map of Trichoglottis.

Cytogenetics Brandham (1999) and Felix and Guerra (2010) listed published chromosome numbers of 2n = 38, 40. ( JW)

Phytochemistry Alkaloids are present at low levels (< 0.01%) in T. fasciata and T. philippinensis (Lüning 1967). A later survey indicated that two of seven species tested contained alkaloids at levels ≥ 0.1% (Lüning 1974). The major components of the floral fragrance of T. philippinensis are (E,E)-α-farnesene (40.5%) and linalool (19.2%). Minor components that contribute to the scent include benzaldehyde (5.0%), p-methylanisole (7.2%), 3-phenylpropanal, and 3-phenylpropanol, although the last two compounds are present only in trace amounts (Kaiser 1993). (NV, RG)

Phylogenetics Trichoglottis is the principal member of clade J, and it has been substantially expanded here by the inclusion of Ceratochilus, Staurochilus, and Ventricularia (Kocyan and Schuiteman, in press). In particular, the subject of Staurochilus versus Trichoglottis has been one of almost endless debate (Seidenfaden 1988); long, often branched, many-flowered inflorescences in the former versus truncated with 1-few flowers allowed easy separation of the two genera. Topik et al. (2005) found Trichoglottis and Staurochilus as sister; Carlsward 314

et al. (2006b) identified a sister relationship between Trichoglottis and Ceratochilus, but Staurochilus was placed together with Acampe, which is here sister to Cottonia. Cottonia/Acampe are then sister to Trichoglottis. (AK, MC)

Ecology All species are epiphytes in swamp and other forests at 0–1800 m. ( JW)

Pollination There is no information available on pollination of Trichoglottis. ( JW)

Uses No uses have been reported for Trichoglottis. A few species are occasionally cultivated. ( JW)

Cultivation Group 3 (see subtribal treatment). (MM)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogeny of Vandeae (Orchidaceae) and

UNCIFERA

the evolution of leaflessness. American Journal of Botany, 93, 770–86. Kocyan, A. and Schuiteman, A. (In press). New combinations in Aeridinae. Phytotaxa. Seidenfaden, G. (1988). Orchid genera in Thailand XIV: fifty-nine vandoid genera. Opera Botanica, 95, 1–398. Senghas, K. (1989). Trichoglottis. In Rudolf Schlechter Die Orchideen, 3rd ed., (ed. F. G. Brieger, R. Maatsch, and K. Senghas), pp. 1310–14. Paul Parey, Berlin. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

705.   TUBE RO LA BIUM Tuberolabium Yamam., Bot. Mag. Tokyo, 38, 209 (1924). Type species: Tuberolabium kotoense Yamam. Parapteroceras Aver., Bot. Zhurn. (Moscow & Leningrad), 75, 723 (1990). Type species: Parapteroceras elobe (Seidenf.) Aver. (basionym: Pteroceras elobe Seidenf.)

Phytochemistry No reports have been found on phytochemistry of Tuberolabium. (NV, RG)

Phylogenetics Tuberolabium is well supported as sister to Ceratocentron in clade E, and then this pair is related to Amesiella, Cryptopylos, Dyakia, Hymenorchis, Macropodanthus, and Trachoma in clade E. (AK, MC)

Ecology Species of Tuberolabium are epiphytes in forests at 500–1500 m. ( JW)

Pollination There is no information available on pollination of Tuberolabium. ( JW)

Derivation of name

Uses

From the Latin tuber, swelling, tuber, and labium, lip, in reference to the swollen labellum that resembles a tuber. ( JW)

No uses have been reported for Tuberolabium. Several species are cultivated. ( JW)

Description (Plate 124; Fig. 705.1)

Cultivation

Epiphytic herbs. Stem ascending or sometimes pendent, enclosed in leaf-sheaths. Leaves few to several, distichous, conduplicate, apex acute, acuminate or unequally bilobed, coriaceous, articulate to a sheathing base. Inflorescence axillary, racemose, arching to pendent, flowers borne simultaneously along whole inflorescence in one season, facing all directions; rachis alate and sulcate. Flowers resupinate, not ephemeral, sometimes strongly scented. Dorsal sepal and petals free. Lateral sepals usually larger, base adnate to column foot. Petals often smaller than sepals, sometimes erose-serrulate. Labellum rigidly attached to column foot, conical to saccate, laterally compressed, trilobed, ecallose inside, sometimes hirsute, the anterior wall much thickened toward apex, side lobes entire or bilobulate, ascending, midlobe reduced, spreading forward; spur conical or saccate, often spreading horizontally, almost continuing the line of column foot, laterally compressed, narrowed toward apex. Column with an obliquely decurrent foot; pollinia two, waxy, subglobose to ovoid, entire, solid, stipe rudimentary, viscidium distinct; rostellum bifid, with a median tooth, the two teeth sometimes strongly retrorse and fitting into two cavities on viscidium. (JW)

Group 1 (see subtribal treatment). (MM)

Distribution (Fig. 705.2) Tuberolabium comprises 11 species distributed in Taiwan, Thailand, Sumatra, Java, Lombok, Sulawesi, Maluku, and the Philippines ( JW)

Cytogenetics Tuberolabium kotoense has a chromosome number of 2n = 30 (reported as Saccolabium kotoense (Yamam.) Yamam.; Felix and Guerra 2010). ( JW)

Taxonomic notes The generic delimitation of the genera closely allied to Pteroceras, including Tuberolabium, needs further study. Tuberolabium is only tentatively accepted here. Its vegetative habit and inflorescence closely resemble those of Dyakia, but flowers of Tuberolabium are far more similar to those of Trachoma, possibly reflecting convergent pollination strategies. (AS)

Taxonomic literature Choltco, T. C. (2006). The genus Tuberolabium with a new species from the Philippines. Orchids, 75, 922–5. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84. Wood, J. J. (1990). Notes on Trachoma, Tuberolabium and Parapteroceras (Orchidaceae). Nordic Journal of Botany, 10, 481–6.

7 0 6 .   U NC I FER A Uncifera Lindl., J. Proc. Linn. Soc., Bot., 3, 39 (1859). Type species: Uncifera acuminata Lindl.

Derivation of name From the Latin uncus, hook, and -fer, bearing, in reference to the retrorse spur on the labellum. ( JW) 315

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Q A I Fig. 705.1.  Tuberolabium odoratissimum (J.J.Sm.) Garay. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum, from above; H. Labellum, longitudinal section; I. Column and labellum, side view; J. Column, front view; K. Column, side view; L. Column, anther cap removed, front view; M. Column, anther cap removed, side view; N. Anther cap, dorsal view; O. Anther cap with pollinia, ventral view; P. Pollinia; Q. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Kew cult. 2005–1001 (K) and Kew Spirit Collection no. 77601.

Description (Plate 125; Fig. 706.1) Epiphytic herbs. Roots not piercing leaf-sheaths. Stems usually pendent, leafy. Leaves overlapping, distichous, conduplicate, oblong to lanceolate, apex unequally bilobed or premorse, articulate to an amplexicaul, sheathing base, spreading in one plane, deciduous. Inflorescence lateral, racemose, few- to many-flowered, usually pendent. Flowers resupinate. Sepals and petals free, similar, concave, lateral sepals slightly oblique. Petals slightly smaller than sepals. Labellum rigidly attached to column base, funnel-shaped, spurred, trilobed; side lobes suberect, midlobe fleshy, adaxially concave; spur hooked or curved, often abruptly swollen near 316

apex, without interior callosities. Column terete, bent down over spur entrance, foot absent; anther cap conical, apex elongate and narrowed, pollinia two, waxy, globular, deeply cleft, separated by a hyaline flange, attached by a common long, linear stipe to a viscidium, stipe wings borne on a ridge formed by basal part of rostellum; rostellum beaked. ( JW)

Distribution (Fig. 706.2) The six species of Uncifera are collectively distributed from northeastern India, Nepal, Bhutan, Burma, and China to Thailand and Vietnam. ( JW)

VANDA

Fig. 705.2.  Distribution map of Tuberolabium.

Cytogenetics

Uses

Brandham (1999) and Felix and Guerra (2010) recorded chromosome numbers of 2n = 38 for the genus. (AP)

No uses have been reported for Uncifera. It is rare in cultivation. ( JW)

Phytochemistry

Cultivation

No reports have been found on phytochemistry of Uncifera. (NV, RG)

Group 2 (see subtribal treatment). (MM)

Phylogenetics Uncifera belongs to a subclade of clade L, in which it is related to Micropera, at least one of the species of Stereochilus, and Smitinandia. A solution to the generic delimitation problems in clade L requires resolution of relationships in grossly polyphyletic Cleisostoma. (AK, MC)

Ecology All species of Uncifera are epiphytes in forests at 1000–2000 m. ( JW)

Pollination There is no information available on pollination of the genus. ( JW)

7 0 7 .   VA NDA Vanda R.Br., Bot. Reg., 6, t. 506 (1820). Type species: Vanda tessellata (Roxb.) Hook. ex G.Don (basionym: Epidendrum tessellatum Roxb.) Ascocentrum Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 975 (1913). Type species: Ascocentrum miniatum (Lindl.) Schltr. (basionym: Saccolabium miniatum Lindl.) Euanthe Schltr., Orchideen, 567 (1914). Type species: Euanthe sanderiana (Rchb.f.) Schltr. (basionym: Esmeralda sanderiana Rchb.f.) Finetia Schltr., Beih. Bot. Centralbl., 36, 140 (1918), nom. illeg. Type species: Finetia falcata (Thunb.) Schltr. (Basionym: Orchis falcata Thunb.) Neofinetia Hu, Rhodora, 27, 107 (1925). Type species: Neofinetia falcata (Thunb.) Hu (basionym: Orchis falcata Thunb.) 317

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Fig. 706.1.  Uncifera acuminata Lindl. A. Habit; B. Flower with detached floral bract, side view; C. Ovary, labellum, and column, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Detail of labellum from above; H. Column with anther cap, side view; I. Column, anther cap removed, side view; J. Anther cap, side view; K. Pollinarium, two views; L. Capsule. Single bar = 1 mm; double bar = 1 cm. Drawn by Susanna Stuart-Smith from Hooker 194 (K). Reproduced with permission from N. R. Pearce and P. J. Cribb (2002), The Orchids of Bhutan. Royal Botanic Garden, Edinburgh and Royal Government of Bhutan.

Nipponorchis Masam., Mem. Fac. Sci. Taihoku Imp. Univ., 11, 592 (1934), nom. illeg. Type species: Nipponorchis falcata (Thunb.) Masam. (basionym: Orchis falcata Thunb.) Eparmatostigma Garay, Bot. Mus. Leafl., Harvard Univ., 23, 178 (1972). Type species: Eparmatostigma dives (Rchb.f.) Garay (basionym: Saccolabium dives Rchb.f.) Trudelia Garay, Orchid Digest, 50, 73 (1986). Type species: Trudelia alpina (Lindl.) Garay (basionym: Luisia alpina Lindl.) 318

Christensonia Haager, Orchid Digest, 57, 40 (1993). Type species: Christensonia vietnamica Haager Ascocentropsis Senghas & Schildh., J. Orchideenfr., 7, 289 (2000). Type species: Ascocentropsis pusilla (Aver.) Senghas & Schildh. (basionym: Ascocentrum pusillum Aver.) Gunnaria Z.J.Liu & L.J.Chen, J. Syst. Evol., 47, 602 (2009). Type species: Gunnaria pusilla (Aver.) Z.J.Liu & L.J.Chen (basionym: Ascocentrum pusillum Aver.)

VANDA

Fig. 706.2.  Distribution map of Uncifera.

Derivation of name From the Sanskrit word vanda, meaning epiphyte. (LG)

Description (Plates 126–130; Fig. 707.1–707.4) Epiphytic or occasionally lithophytic herbs. Roots usually emerging from nodes near base or lower portion of stem. Stem usually stiffly erect, completely enclosed by distichously arranged persistent leaf-sheaths. Leaves distichous, rigid, usually decurved or held erect, linear or oblong, adaxial surface channelled shallowly to deeply, jointed and sheathing at base, deciduous, apex premorse. Inflorescence axillary, racemose, few- to many-flowered, floral bracts triangular. Flowers resupinate, usually widely opening, exhibiting a wide range of colours and patterning. Sepals and petals free, similar, elliptic-obovate, twisted or undulate, often tessellated, margins often reflexed, often narrowed at base. Labellum usually rigidly attached but occasionally articulate, trilobed, side lobes orbicular to pointed, midlobe simple, deltoid or bilobed to fimbriate, usually shortly spurred but occasionally no spur or with an elongate, nectiferous spur; spur often with thickenings at entrance. Column usually lacking a foot; pollinia two, waxy, spherical, grooved; rostellum shelf-like. (LG)

Distribution (Fig. 707.5) Vanda comprises 74 species collectively distributed from India, Nepal, Bhutan, Burma, Thailand, Indochina, southern China,

Taiwan, Japan, Korea, the Philippines, and Indonesia, to New Guinea, northern Australia, and the Solomon Islands. Many species appear to be narrow (island) endemics, with the highest species diversity in the Southeast Asian archipelagos and the Himalayan–Indochinese region. (LG)

Anatomy The following treatment includes both Vanda and the former Neofinetia (Carlsward et al. 2006a). Differences in character-states between the two genera are noted.

Leaf Cuticle smooth to ridged along contours of epidermal cells. Hairs absent. Stomata abaxial. Substomatal chamber small, usually wedged between hypodermal fibres. Epidermal cells conical. Hypodermis ad- and abaxial, composed of one or two rows of thick-walled fibrous idioblasts interspersed with thin-walled chlorenchyma. Fibre bundles absent. Mesophyll homogeneous. Birefringent water-storage cells smooth to pitted. Thick-walled fibrous idioblasts scattered throughout abaxial mesophyll. Cells above midrib slightly modified. Vascular bundles collateral, in one row. Sclerenchyma associated with both xylem and phloem poles. Bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with phloem and xylem sclerenchyma. 319

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Fig. 707.1.  Vanda jennae P.O’Byrne & J.J.Verm. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum, from above; H. Labellum, longitudinal section; I. Labellum base with callus on back wall; J. Column and labellum, side view; K. Column, front view; L. Anther cap; M. Pollinarium (left), front view, and pollinia (right), side view. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Schlechter 20458 (K) (habit) and others redrawn after J. J. Smith, Icones Bogorienses, 3, t. 243 (1907).

Root Velamen three or four cells thick. Epivelamen cells radially elongate. Endovelamen cells angular, isodiametric to radially elongate; cells 320

of outer layer thicker-walled than those of the inner layer. Distinct endovelamen walls absent. Cover cells present over short cells of exodermis. Exodermal cells radially elongate to isodiametric. Long

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Fig. 707.2.  Vanda christensoniana (Haager) L.M.Gardiner [formerly Ascocentrum christensonianum Haager]. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum; H. Column and labellum, side view; I. Column, front view; J. Column, side view; K. Column apex, anther cap removed; L. Anther cap with pollinia; M. Anther cap, dorsal view; N. Anther cap, ventral view; O. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from a living plant cultivated at Kew (accession 2003-360) and Kew Spirit Collection no. 70829.

cell walls primarily ○-thickened. Proliferations absent (present in Neofinetia). Cortical cells often irregularly shaped. Thick-walled fibrous idioblasts abundant throughout. Water-storage cell walls birefringent and smooth to pitted. Single modified cortical layer of thin- to thick-walled cells surrounding endodermis. Aeration units present (absent in Neofinetia). Endodermal cells ○-thickened.

Pericyclic cells thin-walled opposite xylem and thick-walled opposite phloem. Vascular cylinder 31-arch (9–12-arch in Neofinetia). Xylem rays without distinct metaxylem elements. Vascular tissue embedded in sclerenchyma, cell walls of embedding sclerenchyma thickest around phloem clusters. Pith mainly sclerenchymatous with scattered thin-walled parenchyma in the centre. (BC) 321

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Fig. 707.3.  Vanda dives (Rchb.f.) L.M.Gardiner [formerly Eparmatostigma dives (Rchb.f.) Garay]. A. Habit; B. Flower and bract; C. Flower; D. Labellum base, close-up; E. Column; F. Column apex with raised rostellum; G, H. Pollinarium, two views. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from photographs in Blumea, 53, Fig. 1 (2008) and redrawn after G. Seidenfaden in Opera Botanica, 114, Fig. 300 (1992).

Palynology

Cytogenetics

Schill and Pfeiffer (1977) included four species of Vanda in their pollen survey of Orchidaceae: V. coerulescens Griff., two accessions of V. lilacina Teijsm. & Binn. (one as V. laotica Guillaumin), V. suavis Lindl. (= V. tricolor Lindl. var. suavis (Lindl.) Rchb.f.), and V. stangeana Rchb.f. They also reported on pollen of Ascocentrum ampullaceum (Roxb.) Schltr., A. miniatum (Lindl.) Schltr., and A. micranthum (Reinw. ex Blume) Holttum (= Schoenorchis micrantha Reinw. ex Blume, q.v.). Tetrads were convex with laevigate sculpturing and a calymmate sexine (0.8–2.0 μm thick in Ascocentrum). (AP)

The most common diploid chromosome number for species in Vanda is 2n = 38 (Woodward 1952; Storey 1952; Jones 1967; Narayan et al. 1989; Jones et al. 1998; Kumar and Sharma 2002; Chen Xinqi and Bell 2009; Chen Xinqi and Wood 2009). Counts of 2n = 18, 20, 28, 36, 38–42, 57, 72, 76, 95 have also been reported (Jones 1967; Brandham 1999; Kumar and Sharma 2002; Chen Xinqi and Wood 2009). Brandham (1999) suggested a basic number of x = 19, giving 2x = 38, 3x = 57, 4x = 76, 5x = 95, 6x = 114.

322

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Fig. 707.4.  Vanda nana L.M.Gardiner [formerly Ascocentropsis pusilla (Aver.) Senghas & Schildh.]. A. Habit; B. Flower, front view; C. Flower, side view; D. Column and labellum, side view; E. Labellum base, longitudinal section. F. Pollinarium; G. Capsule. Single bar = 1 mm, double bar = 1 cm. Redrawn by Judi Stone after K. Senghas and H. Schildhauer, J. Orchideenfreund, 7, 290 (2000).

Tanaka and Kamemoto (1961) and Kamemoto (1963) studied the chromosomal relationships between many of the taxa within and closely related to the Vanda alliance, including taxa at the time placed in Vanda, Ascocentrum, Euanthe, Aerides, Neofinetia, Phalaenopsis, Renanthera, Trichoglottis, and Arachnis. This work found little support for the genus Euanthe as distinct from Vanda in light of the perfect chromosomal pairing of Euanthe sanderiana with Vanda species and hybrids; the resulting hybrids exhibit high fertility. The chromosomes of species formerly belonging to the genera Ascocentrum and Neofinetia were found to be similar to those of Vanda species in size and morphology, pairing well during meiosis. (LG, JW)

Phytochemistry The anthocyanin pigments of the red-purple flowers of V. tricolor Lindl. var. suavis (Lindl.) Rchb.f. and V. ustii Golamco, Claustro & de Mesa are acylated 3,7,3∙-triglucosides of cyanidin. These are also present as minor components in the violet-blue flowers of V. coerulea Griff. ex Lindl. and V. coerulescens, together with the corresponding delphinidin derivatives, which are the major pigments (Tatsuzawa et al. 2004). The acylating groups comprise ferulic, sinapic, and malonic acids. Cyanidin 3-O-[6-O-malonyl) glucoside]-7,3∙-di-O-[6-O-(E-sinapoyl)glucoside], which is the major pigment of both V. tricolor var. suavis and V. ustii, is also 323

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Fig. 707.5.  Distribution map of Vanda.

Fig. 707.6.  Phenolic constituents of Vanda. Anthocyanins: delphinidin 3-O-[6-O-(malonyl) glucoside]-7,3∙-di-O-[6-O-(E-sinapoyl) glucoside] (1), delphinidin 3-O-glucoside7,3∙-di-O-[6-O-(E-sinapoyl)glucoside] (2). Stilbenoids: 9,10-dihydrophenanthropyrans (3–6), 9,10-dihydrophenanthrenes (7 and 8).

found in Phalaenopsis (Fig. 672.6). However, a crucial distinction between the two genera is the lack of delphinidin derivatives in the latter, according to those species surveyed to date (Tatsuzawa et al. 1997, 2004). Among the delphinidin glycosides of V. coerulea and V. coerulescens are the 3-O-[6-O-(malonyl)glucoside]-7,3∙-­ di-O-[6-O-(E-sinapoyl)glucoside] and its demalonyl analogue 324

(Fig 707.6, 1 and 2). The distribution of acylated 3,7,3∙-triglucosides of cyanidin and delphinidin in numerous cultivars and hybrids of Vanda has been documented (Tatsuzawa et al. 2004) and complements earlier work in which the anthocyanin structures were not defined (Arditti and Fisch 1977). Little is known about other aspects of the flavonoid chemistry of Vanda,

VANDA

although flavone C-glycosides were detected in the leaves of V. merrillii Ames & Quisumb. (C. Williams 1979). Whole plant extracts of V. testacea (Lindl.) Rchb.f. (as V. parviflora Lindl.) and V. tessellata contain the 9,10-dihydrophenanthropyran derivatives, parviflorin and tessalatin (Fig. 707.6), respectively (Anuradha and Rao 1998a,b). Stilbenoids have also been reported from stem extracts of V. coerulea (Fig. 707.6), comprising 9,10-dihydrophenanthropyrans (flavidin and imbricatin), 9,10-dihydrophenanthrenes (coelonin and 6-methoxycoelonin), and the bibenzyl, gigantol (Simmler et al. 2010). Bate-Smith (1968) found ferulic and sinapic acids in acid-hydrolyzed leaf extracts of V. tricolor var. suavis. In early work on the microchemical localization of alkaloids in Orchidaceae first published in 1896 but later reprinted (De Droog 1906), the author failed to detect alkaloids either in V. tricolor Lindl. or V. tricolor var. suavis, although a footnote indicated that Molle was able to demonstrate their presence in young tissues of V. tricolor. According to Lüning (1964), alkaloid content of this species is between 0.001 and 0.01%. Levels of alkaloids in many other species of Vanda have been determined (Lüning 1964, 1967; Lawler and Slaytor 1969, 1970), including a few records under the former generic name, Ascocentrum (as Saccolabium); conversely, some of the original entries for Vanda are now referred to other genera (Dimorphorchis, Papilionanthe, Phalaenopsis, and Vandopsis). Information on the types of alkaloid present is limited. Extracts of the whole plant of V. cristata Wall. ex Lindl. afforded laburnine acetate (Lindström and Lüning 1969), which also occurs in V. helvola Blume (together with laburnine) and V. hindsii Lindl. (Brandänge and Granelli 1973). An acetate either of laburnine or its enantiomer has been detected by GC-MS in extracts of V. luzonica Loher ex Rolfe (Brandänge and Granelli 1973). The pyrrolizidine alkaloids of Vanda are similar to those of Vandopsis (Fig. 708.3). Although esters of laburnine (a pyrrolizidine alkaloid) and its diastereoisomers are also found in Phalaenopsis (Fig. 672.7), the esterifying acids are relatively complex in structure. Several constituents have been isolated from saponified extracts of the whole plant of V. tessellata (as V. roxburghii R.Br.), including sterols (sitosterol and stigmasterol) and long-chain alkanes and alcohols (Basu et al. 1971). Chemical composition data for floral fragrances of V. coerulescens, V. denisoniana Benson & Rchb.f., V. hindsii, and V. tessellata are available from the work of Kaiser (1993, 2011). The major components of the scent of V. coerulescens are methyl anthranilate (23.5%) and methyl (E,Z,Z)-deca-2,4,7-trienoate (33.0%). Minor components of interest include 2-aminobenzaldehyde, and the (E,Z) and (E,E)-isomers of methyl deca-2,4-dienoate (Kaiser 2011). Methyl anthranilate (1.5%) and 2-aminobenzaldehyde (< 1%) are also present in the scent of V. denisoniana, although linalool (66.0%) and (E)-ocimene (14.0%) predominate (Kaiser 1993). In contrast, the scent of V. hindsii, which has been described as ‘aromatic-floral’ (Kaiser 2011), comprises mainly methyl salicylate (48.0%), (E)-ocimene (12.5%), and (E)-cinnamaldehyde (10.8%). The ‘aromatic-floral’ character also applies to the fragrance emitted by V. tessellata, which consists of large amounts of methyl benzoate (61.5%) and linalool (23.0%) together with numerous minor components, notably benzyl acetate, (E)-cinnamaldehyde, p-cresol, indole, α-ionone,

methyl (E)-cinnamate, and 3-phenylpropanal (Kaiser 1993, 2011). In a related study, Ono and Miyazawa (1999) investigated headspace constituents of the flowers of Vanda falcata (Thunb.) Beer (as Neofinetia falcata) using GC-MS. The major components were linalool (40.5%), methyl benzoate (13.4%), cis-3-hexenyl tiglate (12.5%), α-farnesene (9.0%), methyl 2-pentanoate (7.7%), and cis-3-hexen-1-ol (4.2%). Hydrodistillation of fresh flowers yielded a yellowish green oil (0.01%) that contained at least 40 volatile components (Miyazawa and Ono 2000). These comprised mainly terpenes (66.2%) and long chain alkanes (22.9%), including geranyl geraniol (53.0%), heptacosane (5.3%), linalool (4.5%), pentacosane (4.0%), squalene (3.9%), ethyl arachidate (3.8%), and α-farnesene (3.1%). (NV, RG)

Phylogenetics DNA-based phylogenetic analyses by Gardiner et al. (2013) have provided strong support for the monophyly of Vanda sensu lato, incorporating the former genera Ascocentrum, Ascocentropsis, Christensonia, Eparmatostigma, and Neofinetia; these are members of clade F. Overall phylogenetic resolution within the genus is poor, although a number of potential subgenera/sections have relatively strong bootstrap support. These potential subgroupings are not congruent with the previous morphological groups suggested by Lindley, Christenson, Senghas, Seidenfaden, and others, and a new subgeneric classification of Vanda, based on this phylogenetic work, is being developed by Motes, Roberts, and Gardiner. The phylogenetic results also show that species in the genus Vanda are closely related and exhibit low levels of divergence, particularly in that part of the tree comprising the Southeast Asian archipelago species; this result is compatible with rapid divergence of species having taken place. Taprobanea spathulata (L.) Christenson (syn. Vanda spathulata) has been shown to fall well outside the Vanda clade, but the study has also confirmed the previous incorporation of the small genera Trudelia and monospecific Euanthe into Vanda. These results are congruent with the higher level study of Topik et al. (2005) and studies with a narrower sampling of Vanda species, concentrating on Aerides (Kocyan et al. 2008), Holcoglossum Schltr. (Fan et al. 2009), leafless vandoids (Carlsward et al. 2006b), and Phalaenopsis (Padolina et al. 2005). Topik et al. (2005) stated that ‘morphological diversification and possible parallelism of vegetative and reproductive characters’ has probably been a significant contributor to the difficulty of resolving relationships within Aeridinae, and this is reflected in the necessity to use a broader concept for the genus Vanda based on molecular phylogenetic data. (LG)

Ecology Vanda species are usually epiphytes or lithophytes but may also grow as terrestrials and can form large plants with extensive aerial root systems on trees. Plants usually flower when the growing tip of the plant overtops or falls clear of any supporting branch or rock. The thick roots are the primary water storage organs, allowing plants to withstand extensive periods of drought in a semi-dormant condition. In some species the roots 325

VA N D E A E

can extend for six or seven metres or more and find sustaining pockets of leaf mould or detritus. Some species will withstand nearly full winter sun in deciduous forests, but all require some shade in summer. Most species occur at elevations of 800–1600 m. Some species such as V. coerulea, V. cristata, V. javierae D.Tiu ex Fessel & Lückel, and V. tricolor var. suavis come from elevations of 2000 metres or more, where subzero temperatures are not uncommon. Others such as V. sanderiana and the wide ranging V. lamellata come from low elevations and in some cases are found in mangrove communities. Species formerly placed in the genus Ascocentrum, such as V. ampullacea (Roxb.) L.M.Gardiner, V. miniata (Lindl.) L.M.Gardiner, and V. garayi (Christenson) L.M.Gardiner, may also be found at lower elevations, from near sea-level. (LG, MM)

members of Aeridinae (Motes 1997). Ideally grown in baskets with their roots allowed to hang freely, high light levels and brisk air movement around the plants are needed for healthy growth and flowering. Predominantly warm-growing, ideal temperatures for growing most Vanda species and hybrids are 15–32° C (60–90° F). Temperatures below 10° C (50° F) will result in root and leaf growth being checked, and plants will become semi-dormant, except for species from higher elevations that are adapted to occasional subzero temperatures. Vanda roots should be allowed to dry until the velamen is silvery white before being fully soaked until dark green and no more water is being absorbed as it is applied. (MM, LG)

Pollination

As Christenson (1987) stated, a ‘poorly known, imprecisely defined group held together by symplesiomorphies … Vanda requires a complete taxonomic revision’. This taxonomic revision is currently in progress by Gardiner, Roberts, and Motes, incorporating the previously accepted genera Ascocentrum, Ascocentropsis, Christensonia, Eparmatostigma, and Neofinetia as morphological subgenera/sections. Premorse leaf tips and a trilobed labellum are reliable morphological characters for Vanda (Seidenfaden 1988), but the diversity in floral morphology seen across the spectrum of taxa is extremely high, with a wide range of sizes, shapes, colours, and labellum structures (such as the presence or absence of a spur, column foot, midlobe ornamentation, etc.). It is likely that this floral diversity is a result of different pollination syndromes developing within the genus, with the relatively low levels of divergence revealed in the recent phylogenetic work representing the rapid diversification of new, closely related species able to adapt and occupy a wider range of ecological niches. (LG, MM)

Few pollination studies of Vanda species have been carried out. Many Vanda species with large, showy, colourful flowers, often with an open, spreading perianth, are visited by bees and butterflies in cultivation. Holttum (1953) concluded that ‘typical’ Vanda species are pollinated by bees. Beetle pollination has been reported for V. cristata (Pradhan 1983), and beetles may also pollinate V. denisoniana Benson & Rchb.f. (Christenson 1992). The species formerly placed in Neofinetia (V. falcata, V. richardsiana (Christenson) L.M.Gardiner, V. xichangensis (Z.J.Liu & S.C.Chen) L.M.Gardiner) are thought to be pollinated by moths based on floral morphology (white flowers, long, thin, nectariferous spurs). Similarly, the species formerly placed in Ascocentrum (V. ampullacea, V. aurantiaca (Schltr.) L.M.Gardiner, V. aurea (J.J.Sm.) L.M.Gardiner, V. christensoniana (Haager) L.M.Gardiner, V. curvifolia (Lindl.) L.M.Gardiner, V. garayi, V. himalaica (Deb, Sengupta & Malick) L.M.Gardiner, V. insularum (Christenson) L.M.Gardiner, V. miniata, V. nana L.M.Gardiner, V. rubra (Lindl.) L.M.Gardiner, V. semiteretifolia (Seidenf.) L.M.Gardiner) are thought to be birdpollinated, given their small, brightly coloured flowers with short, nectariferous spurs (Dressler 1981, 1993). (LG)

Uses Subedi et al. (2011) reported the use of V. cristata in Nepal to prepare a root paste to treat boils and dislocated bones, a leaf powder as an expectorant, and a leaf paste for cuts and wounds. Roots of V. tessellata were used to treat scorpion stings, bronchitis, and rheumatism, and a leaf paste of this species was used to suppress fever. Similar and additional medicinal uses have been reported for these species from Sri Lanka and India, especially V. tessellata, which, like Rhynchostylis retusa (L.) Blume, is locally known as rasna. Vanda coerulea was also used medicinally in India (Lawler 1984). Many species of Vanda are cultivated, and the genus is the one of the five most horticulturally important in the Orchidaceae. It is widely used in breeding programmes for the cut-flower industry. (AS, LG)

Cultivation Group 2 (see subtribal treatment). Vanda cultivation is similar to that of Cattleya, Phalaenopsis, and Dendrobium, as well as other 326

Taxonomic notes

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogeny of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Chen Xinqi and Bell, A. 2009. Vanda. In Flora of China, 25 (ed. Wu Z., P. H. Raven, and Hong D.), pp. 471–4. Science Press, Beijing & Missouri Botanical Garden Press, St. Louis. Chen Xinqi, and Wood, J. (2009). Flora of China, 25 (ed. Wu Z., P. H. Raven, and Hong D.). Science Press, Beijing & Missouri Botanical Garden Press, St. Louis. Christenson, E. A. (1987). Taxonomy of Aerides and related genera. In Proceedings of the 12th World Orchid Conference (ed. K. Saito and R. Tanaka), pp. 35–40. 12th World Orchid Conference Organizing Committee, Tokyo. Christenson, E. A. (1992). Notes on Asiatic orchids. Lindleyana, 7, 88–94. Fan, J., Qin, H. N., Li, D. Z., and Jin, X. H. (2009). Molecular phylogeny and biogeography of Holcoglossum (Orchidaceae: Aeridinae) based on nuclear ITS, and chloroplast trnL-F and matK. Taxon, 59, 849–61. Gardiner, L. M. (2012). New combinations in the genus Vanda (Orchidaceae). Phytotaxa, 61, 47–54. Gardiner, L. M., Kocyan, A., Motes, M., Roberts, D. L., and Emerson, B. C. (2013). Phylogenetic patterns in the genus

VANDOPSIS

Vanda and related genera (Orchidaceae). Botanical Journal of the Linnean Society, 173, 549–72. Kocyan, A., de Vogel, E. F., Conti, E., and Gravendeel, B. (2008). Molecular phylogeny of Aerides (Orchidaceae) based on one nuclear and two plastid markers: a step forward in understanding the evolution of the Aeridinae. Molecular Phylogenetics and Evolution, 48, 422–43. Kumar, L. Y., and Sharma, J. (2002). Orchids of India - III. Biodiversity and status of Vanda Jones ex R.Br. Daya Publishing House, Delhi. Motes, M. R. (1997). Vandas: their botany, history, and culture. Timber Press, Portland, Oregon. Padolina, J., Linder, C. R., and Simpson, B. B. (2005). A phylogeny of Phalaenopsis using multiple chloroplast markers. Selbyana, 26, 23–7. Pradhan, G. M. (1983). Vanda cristata. American Orchid Society Bulletin, 52, 464–8. Schuiteman, A. and Bonnet, P. (2008). Eparmatostigma dives (Orchidaceae), a new generic record for Laos. Blumea, 53, 341–4. Seidenfaden, G. (1988). Orchid genera in Thailand XIV. Fifty-nine vandoid genera. Opera Botanica 95, 1–398. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

708.   VAN D O P SIS Vandopsis Pfitzer in Engler & Prantl, Natürl. Pflanzenfam., 2(6), 211 (1889). Type species: Vandopsis lissochiloides (Gaudich.) Pfitzer (basionym: Fieldia lissochiloides Gaudich.) Fieldia Gaudich., Voy. Uranie, Bot., 424 (1829), non Cunningh. (1825). Type species: Fieldia lissochiloides Gaudich.

Derivation of name From the genus Vanda, and the Greek opsis, likeness, implying a resemblance to Vanda. ( JW)

Description (Plate 131; Fig. 708.1) Epiphytic, lithophytic, or terrestrial herbs. Stem rigid, decumbent or pendent, sometimes branched. Leaves many, distichous, coriaceous, conduplicate, articulated to a sheathing base, glabrous, deciduous. Inflorescences axillary, suberect or pendent, racemose, rarely shortly branched, several- to many-flowered. Flowers flat, resupinate, usually white or pale yellow, often with reddish spots, labellum sometimes with purple-red striations at base. Sepals and petals free, similar, spreading. Labellum smaller than petals, firmly attached to column, geniculately bent, channelled, concave or gibbous at base, trilobed, spur absent, with a transverse basal lamella, midlobe laterally flattened, adaxial surface usually longitudinally ridged or keeled. Column foot absent; pollinia two, deeply grooved with the halves unequal, waxy, sessile, stipe broadly oblong, viscidium transverse, prominent; rostellum inconspicuous. ( JW)

Distribution (Fig. 708.2) Vandopsis consists of four species collectively distributed in Burma, China, Thailand, Laos, Vietnam, the Philippines, and the Malay Archipelago to New Guinea. ( JW)

Cytogenetics Brandham (1999) and Felix and Guerra (2010) listed a chromosome number of 2n = 38 for the genus. ( JW)

Phytochemistry Alkaloids are present at levels of 0.01–0.1% in V. lissochiloides (as V. batemannii Lindl.) and V. gigantea (Lindl.) Pfitzer (Lüning 1964). These species contain the alkaloids laburnine and lindelofidine (Fig. 708.3), which are diastereoisomers of 1-hydroxymethylpyrrolizidine, together with their acetates (Brandänge and Granelli 1973). Little is known about other aspects of the phytochemistry of Vandopsis, other than the observation by C. Williams (1979) that flavone C-glycosides occur in the leaves of V. lissochiloides (as Vanda lissochiloides (Gaudich.) Lindl.). (NV, RG)

Phylogenetics Vandopsis is a member of grade K. Earlier studies found Vandopsis sister to Trichoglottis (Topik et al. 2005) or Adenoncos/Aerides (Carlsward et al. 2006b). Vandopsis undulata (Lindl.) J.J.Sm. does not come out with the two species in grade K but rather in an isolated position between clades H and I. It probably represents a new genus (Kocyan and Schuiteman, in preparation). (AK, MC)

Ecology Vandopsis gigantea (Lindl.) Pfitzer is epiphytic on tree trunks in open forest or at forest margins at 800–1700 m in China. Vandopsis lissochiloides is lithophytic or terrestrial on limestone and schist outcrops among patches of coastal vegetation in Sulawesi. It is also recorded as a lithophyte on limestone rocks beside the sea in New Guinea and growing on boulders in semi-deciduous secondary forest at 300 m in Laos. (JW, AS)

Pollination There is no information available on pollination of Vandopsis. ( JW)

Uses No uses have been reported for Vandopsis. Several species are cultivated. ( JW)

Cultivation Group 2 (see subtribal treatment). (MM)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogeny of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Topik, H., Yukawa, T., and Ito, M. (2005). Molecular phylogenetics of subtribe Aeridinae (Orchidaceae): insights from plastid matK and nuclear ribosomal ITS sequences. Journal of Plant Research, 118, 271–84.

327

VA N D E A E

J

C

B

D F

E

K

G I

M H P

L

N

Q

O

A Fig. 708.1.  Vandopsis gigantea (Lindl.) Pfitzer. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum, from above; H. Labellum, longitudinal section; I. Column and labellum, side view; J. Column, front view; K. Column, side view; L. Column, anther cap removed, side view; M. Column apex, detail; N. Anther cap, dorsal view; O. Anther cap with pollinia, ventral view; P. Pollinarium, pollinia removed; Q. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Kerr 0300 (K) and Kew Spirit Collection no. 74684.

709.  XE N IKO P H Y TO N

Description (Fig. 709.1)

Xenikophyton Garay, Bot. Mus. Leafl., Harvard Univ., 23, 374 (1974). Type species: Xenikophyton smeeanum (Rchb.f.) Garay (basionym: Saccolabium smeeanum Rchb.f.)

Erect, epiphytic herbs. Stem simple or with an occasional short branch, leafy. Leaves distichous, coriaceous, obtusely unequally bilobed, articulate to a sheathing base; sheaths rugose, falcate. Inflorescence erect, racemose or paniculate, many-flowered; floral bracts minute, acuminate. Flowers about 4 mm long, resupinate, mostly greenish white streaked with brown, labellum greenish white or white, sometimes with violet around entrance to sac. Sepals and petals similar, free. Dorsal sepal connivent with petals to form a galea. Labellum sessile, base scrotiform, apex

Derivation of name From the Greek xenikos, strange, and phyton, plant, in reference to the strange admixture of characters from Cleisomeria and Sarcophyton. ( JW) 328

XENIKOPHYTON

Fig. 708.2.  Distribution map of Vandopsis.

Fig. 708.3.  Pyrrolizidine alkaloids of Vandopsis.

recurved, provided with a swelling, entrance almost closed, back wall callus absent. Column fleshy, obtuse, stelidia obscure, foot absent, clinandrium distinctly excavate, dorsally reclinate; anther cap mitre-shaped, pollinia four, free, globose, stipe linear, replicate, viscidium elliptic; stigma with a vertical margin, narrowly elliptic; rostellum vertical, pointing upward, prominently bifurcate. ( JW)

Distribution (Fig. 709.2) The two species of Xenikophyton—X. smeeanum and X. seidenfadenianum M.Kumar, Sequiera & J.J.Wood—are limited to southern India. ( JW)

Cytogenetics No chromosome counts for Xenikophyton have been recorded. ( JW)

Phytochemistry No reports have been found on phytochemistry of Xenikophyton. (NV, RG)

Phylogenetics There is no phylogenetic information available for this genus. (AK) 329

VA N D E A E

M K

L

C

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I

O J

D

F

E

A H B

G

Fig. 709.1.  Xenikophyton smeeanum (Rchb.f.) Garay. A. Habit; B. Portion of inflorescence; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum; H. Labellum, longitudinal section; I. Column and labellum, side view; J. Column (anther cap removed) and labellum, side view; K. Column, front view; L. Column, side view; M. Anther cap (front view); N. Anther cap, ventral view; O. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Meebold 10650 (K) and Kew Spirit Collection no. 13797.

Ecology

Cultivation

Both species are epiphytes at 900–1350 m in India. ( JW)

Group 2 (see subtribal treatment). (MM)

Pollination

Taxonomic notes

No information on pollination of Xenikophyton is available. ( JW)

Uses No uses have been reported for Xenikophyton; it is rare in cultivation. ( JW) 330

This genus has so far not been included in DNA-based phylogenetic analyses, and therefore its systematic position within the subtribe is still undetermined. The floral morphology is similar to that of Schoenorchis, which may be its nearest relative. They agree in having minute flowers with the labellum distinctly projecting beyond the sepals and petals, the apex of the labellum

AGROSTOPHYLLUM

Fig. 709.2.  Distribution map of Xenikophyton.

having a swelling underneath, and the long, bifurcate rostellum being abruptly curved upwards. Both may have freely branching inflorescences. Plants of Xenikophyton are more robust with thick, papillose-hairy roots, and the four equal-sized pollinia are not held in two pairs. In Schoenorchis, the four pollinia are distinctly unequal in size and clearly arranged in two pairs. (AS)

SUB T RIBE AGRO S TO P H Y L L IN AE Agrostophyllinae Szlach., Fragm. Flor. Geobot. Supplementum, 3, 66 (1995). Type: Agrostophyllum Blume

Description Epiphytic, lithophytic, or terrestrial herbs with a creeping rhizome. Roots glabrous. Stems caespitose or well spaced, reed-like, not pseudobulbous, leafy, or pseudobulbous and completely obscured by persistent leaf-sheaths (e.g. Earina valida Rchb.f.). Leaves many, duplicate, distichous, articulate at apex of tubular or imbricate sheaths; sheaths often deeply split, sometimes with blackish margins, sometimes with two acicular to filiform appendages. Inflorescence terminal, rarely lateral, one- to manyflowered, fascicled into a globose capitulum, racemose or paniculate. Flowers resupinate or non-resupinate, not showy, white, greenish white or yellow, sometimes self-pollinating. Sepals free. Labellum immobile, entire or trilobed, often saccate at base,

lateral margins of hypochile (when present) sometimes adnate to column foot, rim connecting lobes of the hypochile often with a callus, or disc ecallose. Column with or without stelidia, column foot rudimentary or absent; anther erect or incumbent, ellipsoid to ovate; pollinia four or eight, clavate; stigma concave; rostellum triangular, erect to suberect, remnant after removal of pollinia notched to obtuse; viscidium solitary. ( JW)

Distribution Agrostophyllinae comprise two genera distributed from the Seychelles and tropical Asia to New Caledonia, Solomon Islands, and Vanuatu, eastward to Fiji and Samoa, south to New Zealand, with a centre of distribution in New Guinea. ( JW)

Artificial key to the genera of Agrostophyllinae ( JW)

●

●

Pollinia four; column foot absent . . . . . . . . . . . 711. Earina Pollinia eight; column foot rudimentary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 710. Agrostophyllum

7 1 0 .   AGROS T OPH YL LUM Agrostophyllum Blume, Bijdr. Fl. Ned. Indië, 368 (1825). Type species: Agrostophyllum javanicum Blume 331

VA N D E A E

Diploconchium Schauer, Nov. Actorum Acad. Caes. Leop.-Carol. Nat. Cur., 19 (Suppl. 1), 428 (1843). Type species: Diploconchium inocephalum Schauer Chitonochilus Schltr. in K.M. Schumann & C.A.G. Lauterbach, Fl. Schutzgeb. Südsee, Nachtr., 134 (1905). Type species: Chitonochilus papuanus Schltr. Appendiculopsis (Schltr.) Szlach., Fragm. Florist. Geobot., Suppl., 3, 119 (1995). Type species: Appendiculopsis stipulata (Griff.) Szlach. (basionym: Appendicula stipulata Griff.)

with a membranous, narrowly ovate bract. Fruit ellipsoid with elaters. ( JW)

Derivation of name

Infrageneric treatment

From the Greek agrostis, grass, and phyllon, leaf, in reference to the grass-like leaves of many species. ( JW)

A. sect. Agrostophyllum. Rhizome creeping and rooting along its entire length; inflorescence a usually dense capituliform cluster of one- to few-flowered racemes, sometimes reduced to a single flower. Type species: A. javanicum Blume A. sect. Appendiculopsis Schltr. (as ‘Apendiculopsis’). Leaves numerous, less than 3.5 cm long, and usually abruptly contracted between blade and sheath (leaves longer and not abruptly contracted in A. laterale); inflorescences terminal or lateral (A. laterale). Type species: A. stipulatum (Griff.) Schltr. A. sect. Dolichodesme Schltr. Inflorescence terminal, a single, elongated raceme or panicle (or at least seemingly so). Type species: A. paniculatum J.J.Sm. A. sect. Oliganthe Schltr. Rhizome forming a dichotomously branched, pendent sympodium, rooting at its base; inflorescence a dense terminal cluster of few-flowered racemes. Type species: A. superpositum Schltr. ( JW)

Description (Plate 132; Fig. 710.1) Epiphytic or occasionally lithophytic, sympodial herbs with a creeping rhizome. Roots along entire rhizome, glabrous. Stems caespitose or well-spaced, without pseudobulbs, erect or pendent, bilaterally flattened, of many internodes, simple or branching, sometimes fleshy, many-leaved. Leaves articulate, distichous, papyraceous to thinly coriaceous, usually twisted at base to lie in one plane with stem axis, mostly at right angles to stem (mostly in A. sect. Appendiculopsis), sometimes marked by impressions of adjacent leaves, base truncate (or not), sometimes contracted between sheath and blade, margins entire, apex usually conspicuously obtusely or acutely bilobed, sometimes mucronate; sheaths enveloping the stem, tubular, glabrous, sometimes with two acicular to filiform appendages, entire or deeply longitudinally split opposite blade and then with blackish or brown margins along the slit. Inflorescences normally terminal, occasionally individual inflorescences arising laterally from stems (e.g. A. laterale J.J.Sm.), one- to many-flowered, usually fascicled into a globose capitulum on an elongate axis, the base of the fascicle with several persistent, distichous, short-lived bracts that eventually disintegrate into fibres, or paniculate; peduncle enveloped by two or three membranous, subacute, tubular bracts. Flowers resupinate or not, fleshy, especially labellum, white or yellow, exterior of sepals with scattered brownish scale-hairs, labellum epichile often papillose, otherwise glabrous. Sepals and petals similar, free, petals narrower. Dorsal sepal often concave. Lateral sepals forming an obtuse mentum, connate in basal part on adaxial side of mentum, often keeled outside along midvein. Petals linear to lanceolate. Labellum not motile, entire or trilobed, hypochile concave, with lateral margins free from column foot, or saccate, with lateral margins adnate to column foot, which then appears to be absent, containing nectar, sulcate below, conspicuously bilobed in front, the rim connecting lobes of hypochile often with a callus; epichile broadly ovate to transversely elliptic, convex, often minutely papillose, often with a longitudinal median concavity. Column semi-terete, stelidia present or absent, column foot rudimentary; anther cap minute, pollinia eight, clavate, flattened, attached without caudicles to a solitary, brown viscidium; stigma concave; rostellum either short, bidentate or triangular, or oblong and entire and then overlapping clinandrium. Ovary and pedicel with scattered brownish scale-hairs, six-ribbed, terete, 332

Distribution (Fig. 710.2) The genus includes 50–60 species distributed in the Old World tropics from the Seychelles and tropical Asia east to Samoa, with a centre of distribution in New Guinea, but absent from Australia. ( JW)

Anatomy Reports of anatomical features of Agrostophyllum are scattered in the literature. Khasim and Mohana Rao (1990) briefly described the anatomy and habitat tolerance of orchids, including Agrostophyllum. Pridgeon (1987) and Porembski and Barthlott (1988) studied roots of A. planicaule (Wall. ex Lindl.) Rchb.f. (as A. khasianum Griff.), and Porembski and Barthlott (1988) also those of A. superpositum (as A. congestum Schltr.). Møller and Rasmussen (1984) reported conical silica bodies in A. callosum (Blume) J.J.Sm.) and A. stipulatum (Griff.) Schltr.

Leaf transverse section

Mesophyll homogeneous with water-storage cells and fibre bundles. Vascular bundles with fibrous cap at phloem pole. Conical silica bodies present throughout the shoot system.

Root transverse section

Velamen 2–8 layers thick. Exodermal cells thickened on outer walls. Some cortical cells reticulately thickened. Endodermal cells thickened on radial and inner walls. (WS)

Palynology Nothing is known about pollen of Agrostophyllum. (AP)

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Fig. 710.1.  Agrostophyllum glumaceum Hook.f. A. Habit; B. Inflorescence; C. Floral bract; D. Flower, side view; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum and column, anther cap removed, front view; I. Labellum and column, anther cap removed, side view; J. Labellum, showing hypochile and lower portion of epichile; K. Column with anther cap, front view; L. Anther cap, dorsal view; M. Anther cap, ventral view; N. Pollinia; O. Capsule. Single bar = 1 mm, double bar = 1 cm. Drawn by Linda Gurr from Clemens 27481 (A, B), Lamb SNP 1992 (C-L), and Leche s.n. (M-P). Reproduced with permission from J. J. Wood (2003), Orchids of Borneo. Volume 4. The Sabah Society, Kota Kinabalu, in association with the Royal Botanic Gardens, Kew.

Cytogenetics Brandham (1999) and Felix and Guerra (2010) listed chromosome counts of 2n = 38, 40, and 46 for species of Agrostophyllum, with 2n = 40 the most common. (AP)

Phytochemistry Alkaloids have been detected at low levels (< 0.1%) in A. majus Hook.f., A. myrianthum King & Pantl., and A. stipulatum subsp. bicuspidatum (J.J.Sm.) Schuit. (cited as A. bicuspidatum J.J.Sm. and A. callosum), but neither A. brevipes King & Pantl. nor A. planicaule (as A. khasianum Griff.) contained them (Lüning 1964, 1967). A later survey indicates that of 23 species tested, none had an alkaloid content ≥ 0.1% (Lüning 1974).

Three species—A. brevipes, A. callosum Rchb.f., and A. planicaule (cited as ‘A. khasiyanum’, probably meant to be A. khasianum)—are rich in stilbenoids, some of which are found only in this genus (Fig. 710.3). Constituents isolated from extracts of the whole plant of A. planicaule, which was the first of the three species to be investigated, comprise the phenanthropyrans agrostophyllin, agrostophyllidin, and agrostophylloxidin, the phenanthropyranones agrostophyllone and agrostophylloxin, the diastereoisomeric phenanthropyran derivatives agrostophyllanthrol and isoagrostophyllanthrol, the phenanthrene dimers agrostonin and agrostonidin, and a number of common stilbenoids (Majumder and Sabzabadi 1988; Majumder et al. 1996b, 1998, 2008c). The last comprises the bibenzyl moscatilin, the phenanthropyran imbricatin, and the phenanthropyranones flaccidinin and isoflaccidinin 333

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Seychelles

Fig. 710.2.  Distribution map of Agrostophyllum.

(Majumder et al. 1996b). A comparable study on extracts of the whole plant of A. callosum afforded callosin, a 9,10-dihydrophenanthrene derivative also present in Coelogyne flaccida Lindl., callosinin, a 9,10-dihydrophenanthropyran derivative, and the known stilbenoids flaccidin, flaccidinin, imbricatin, iso-oxoflaccidin, 6-methoxycoelonin, orchinol, and oxoflaccidin (Majumder et al. 1995). Other novel constituents isolated from A. callosum include callosumin, callosuminin, and callosumidin (Majumder et al. 1996a), and the diastereoisomeric 9,10-dihydrophenanthropyran derivatives agrostophyllol and isoagrostophyllol (Majumder et al. 1999c). This species also contains agrostophyllin, agrostonin, and agrostonidin, in common with A. planicaule (Majumder et al. 1995, 1998). The stilbenoid constituents of the whole plant of A. brevipes are agrostophyllin, callosinin, flaccidin, flaccidinin, flavanthrinin, imbricatin, 6-methoxycoelonin, and nudol (Majumder et al. 2003). Information on other aspects of Agrostophyllum phytochemistry is limited, although the lanostane triterpenoids agrostophyllinol and agrostophyllinone occur in A. brevipes and agrostophyllinone in A. callosum (Majumder et al. 2003). (NV, RG)

Phylogenetics Schuiteman (1997) considered A. sect. Appendiculopsis as probably monophyletic. He cited the following characters as 334

synapomorphies of the section: production of lateral inflorescences on an undamaged leafy stem, simple column, and creeping rhizome, which probably evolved independently in some New Guinea species of A. sect. Agrostophyllum. ( JW)

Ecology Species of Agrostophyllum occur as epiphytes or terrestrials in lowland, hill, and lower montane forests, mossy forests, and treefern savannas from sea level to 2900 m. ( JW)

Pollination Autogamy has been reported for several species of Agrostophyllum by Schlechter (1914, transl. 1982) and Smith (1928). ( JW)

Uses Christensen (2002) reported that A. bicuspidatum J.J.Sm. is used to protect against curses in the Kelabit Highlands of Sarawak, Borneo. Whenever a person carries a part of this plant, he or she is protected against curses. Species are not in general cultivation. ( JW)

EARINA

Fig. 710.3.  Stilbenoid constituents of Agrostophyllum.

Taxonomic literature Brieger, F. G. (1975). Agrostophyllum. In Rudolf Schlechter Die Orchideen, 3rd ed. (ed. F.G. Brieger, R. Maatsch, and K. Senghas), p. 378, Paul Parey, Berlin. Schuiteman, A. (1997). Revision of Agrostophyllum section Appendiculopsis (Orchidaceae), with notes on the systematics of Agrostophyllum. Orchid Monographs, 8, 1–20.

or spreading. Labellum sessile, not motile, erect, entire or weakly trilobed, usually channelled, the base often saccate, distal portion reflexed; disc ecallose, glabrous. Column semi-terete, column foot absent, clinandrium deeply excavate, the margins entire or weakly bilobed; anther cap terminal, decumbent, operculate, two-celled, pollinia four, waxy, joined to a common viscidium; stigma directly below rostellum, transverse, often deeply recessed. ( JW)

711.   EAR IN A

Distribution (Fig. 711.2)

Earina Lindl., Edwards’s Bot. Reg., 20, sub t. 1699 (1834). Type species: Earina mucronata Lindl.

Derivation of name From the Greek earinos, of springtime, vernal, referring to the season of flowering of the type species, E. mucronata, in New Zealand. ( JW)

Description (Plate 133; Fig. 711.1) Epiphytic or terrestrial herbs with an abbreviated rhizome. Stems well spaced or caespitose, erect, simple, leafy, conspicuously elongated or condensed and fleshy, pseudobulbous and completely obscured by persistent leaf-bases. Leaves distichous, articulate, conduplicate in bud, blade ligulate, coriaceous; base sheathing, conspicuously enlarged and imbricate or relatively widely spaced and tubular. Inflorescence terminal, distichous, racemose or more commonly paniculate, many-flowered; floral bracts imbricate. Flowers resupinate or not, usually white or greenish white, sometimes heavily scented. Sepals free, erect or spreading, lateral sepals oblique. Petals similar to dorsal sepal or slightly narrower, free, erect

Earina consists of six or seven species distributed in New Zealand, New Caledonia, Solomon Islands, and Vanuatu, eastward to Fiji and Samoa. ( JW)

Anatomy Curtis (1917) gave a thorough account of the anatomy of two New Zealand species of Earina—E. mucronata and E. autumnalis (G.Forst.) Hook.f. (as E. suaveolens Lindl.). Porembski and Barthlott (1988) studied roots of E. autumnalis. Møller and Rasmussen (1984) reported conical silica bodies in E. mucronata.

Leaf TRANSVERSE SECTION

Cuticle thick. Stomata abaxial. Mesophyll heterogeneous with fibre bundles. Vascular bundles surrounded by a sclerenchyma sheath. Conical silica bodies present.

Stem transverse section

Cuticle thick. Cortex surrounded internally by complete or incomplete sclerenchyma ring continuous with vascular bundles. 335

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Fig. 711.1.  Earina deplanchei Rchb.f. A. Base of plant, ×0.66; B. Inflorescence, ×0.66; C. Floral bud, 4.2 × 2.5 mm; D. Flower, side view; E. Flower, face view; F. Lateral sepal, 6.0 × 7.0 mm; G. Petal, 6.0 × 1.2 mm; H. Labellum, face view, 6.0 × 3.0 mm; I. Column, 4 mm; J. Anther, 0.7 mm; K. Pollinia, 0.5 mm; L. Ovary, transverse section, diam. 1.5 mm. Magnifications as originally published. Drawn by N. Hallé from MacKee 24502, except Cribs 1286 (A), and Buchholz 1263 (H, L). Reproduced with permission from N. Hallé (1977), Flore de la Nouvelle Calédonie et dépendances, no. 8, Orchidacées, plate 130, p. 313. Muséum d’Histoire Naturelle, Paris.

Vascular bundles and fibre bundles also scattered in starch-containing ground tissue.

Root

Cytogenetics Felix and Guerra (2010) listed chromosome counts of 2n = 40 for the genus. (AP)

transverse section

Velamen 3–5 layers thick. Radial and outer walls of exodermal cells thickened. Inner cortical cells reticulately thickened, whereas central cells are spirally thickened. Hyphae and raphides present throughout cortex. Roots 11–13-arch. (WS)

Palynology There are no reports on pollen of Earina species. (AP) 336

Phytochemistry The phenanthraquinone bulbophyllanthrone is a constituent of E. autumnalis and showed antibacterial and cytotoxic activity in bioassays (Hinkley and Lorimer 1999). This compound was first reported in Bulbophyllum odoratissimum (Smith) Lindl. ex Wall. (Majumder and Sen 1991). (NV, RG)

EARINA

Fig. 711.2.  Distribution map of Earina.

Ecology

Pollination

Earina autumnalis from New Zealand occurs as a lithophyte on rocks, occasionally on the ground when trees fall or branches break, or when the orchid itself falls, and as an epiphyte low down on trees, sometimes growing on exposed uppermost branches (Johns and Molloy 1983; St George 1999; de Lange 2007). Earina deplanchei Rchb.f. is recorded as a terrestrial on red soils, or rarely an epiphyte in forest, at elevations of 400–1200 m in New Caledonia (Hallé 1977). Earina floripecten Kränzl. has been recorded as a terrestrial or lithophyte on rocks, maquis on serpentine substrate, ‘coteaux broussailleux’, at elevations of 450–1100 m in New Caledonia (Hallé 1977). Earina mucronata Lindl. from New Zealand is an epiphyte on trees in lowland forest, covering well-lit trunks and branches, or occasionally on fallen logs and brick or concrete walls in urban areas (Johns and Molloy 1983; St George 1999; de Lange 2007). Earina valida Rchb.f. is recorded as an epiphyte in dense or open forest or on trees at the edges of grassland at elevations of 700–1100 m in Fiji (Kores 1991). Hallé (1977) reported it from humid forest or rarely maquis at 500–1450 m in New Caledonia. It is epiphytic in rain forest at 300–1070 m in Vanuatu (Lewis and Cribb 1989) and epiphytic in forests at 500–900 m in Samoa (Cribb and Whistler 1996). ( JW)

Information on pollinators of species of Earina is limited, but what little there is indicates at least visitation by crane flies (Tipulidae) to flowers of E. mucronata and E. autumnalis (Scanlen 1996). (AP)

Uses There are no known uses for any of the species, and they are not in common cultivation. (AP)

Taxonomic literature Cribb, P. and Whistler, W. A. (1996). Orchids of Samoa. Royal Botanic Gardens, Kew. De Lange, P. J. (2007). Wild orchids of the lower North Island: field guide. Department of Conservation, Wellington Conservancy, Wellington, New Zealand. Hallé, N. (1977). Orchidacées. In Flore de la Nouvelle-Calédonie et Dependances 8 (ed. A. Aubréville, A. and J-F. Leroy), Muséum National D’Histoire Naturelle, Paris. Johns, J. and Molloy, B. (1983). Native orchids of New Zealand. A. H. & A. W. Reed, Wellington, New Zealand.

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Kores, P. J. (1991). Family 32. Orchidaceae. In Flora vitiensis nova: a new flora of Fiji (spermatophytes only) (ed. A. C. Smith). Volume 5, pp. 322–575. National Tropical Botanical Garden, Lawai, Kauai, Hawaii. Lewis, B. and Cribb, P. (1989). Orchids of Vanuatu. Royal Botanic Gardens, Kew. St George, I. (1999). The nature guide to New Zealand native orchids. Godwit Press, Auckland, New Zealand.

S UB TR IBE A NGR A E C I NAE Angraecinae Summerh., Kew Bull., 20, 188 (1966); Schltr., Notizbl. Bot. Gart. Berlin, 9, 590 (1920), sine descr. Type: Angraecum Thouars Aerangidinae Summerh., Kew Bull., 20, 188 (1966); Schltr., Notizbl. Bot. Gart. Berlin, 9, 591 (1920), sine descr. Type: Aerangis Rchb.f. Podanginae Brieger in Schlechter, Orchideen ed. 3, 128 (1971), nom. provis. sine descr. Type: Podangis Rchb.f., syn. nov. Bolusiellinae Szlach., Fragm. Flor. Geobot., Suppl., 3, 91 (1995). Type: Bolusiella Schltr., syn. nov. Listrostachyinae Szlach., Fragm. Flor. Geobot., Suppl., 3, 92 (1995). Type: Listrostachys Rchb.f., syn. nov. Calyptrochilinae Szlach., Fragm. Flor. Geobot., Suppl., 3, 92 (1995). Type: Calyptrochilum Kränzl., syn. nov. Rhaesterinae Szlach., Fragm. Flor. Geobot., Suppl., 3, 91 (1995). Type: Rhaesteria Summerh., syn. nov.

Description Epiphytic, lithophytic, or rarely terrestrial, monopodial, sometimes psygmoid herbs. Roots elongate with a prominent, silvery velamen. Stem usually covered by distichously arranged leaf-bases or rarely with cataphylls. Leaves present, rarely absent, fleshy to coriaceous, conduplicate, distichous, often articulated to a conduplicate or sheathing leaf-base. Inflorescence lateral, axillary, one- to many-flowered, simple or rarely branching, peduncle usually cylindrical, sometimes covered by sterile bracts, rachis cylindrical; bracts usually shorter than pedicel and ovary, sometimes sheathing at base or amplexicaul. Flowers resupinate or non-­resupinate, usually unicolorous, white, green, yellow-brown, orange or red, sometimes marked with green, yellow, brown, or red on labellum. Sepals and petals free, spreading or reflexed, rarely porrect. Labellum entire or trilobed, ecallose or with a callus of 1–3 basal, fleshy ridges, spur usually present, side lobes erect and enclosing column or spreading, midlobe usually larger, recurved or porrect; spur cylindrical, clavate or S-shaped or geniculate, sometimes saccate or rarely absent. Column usually fleshy, less commonly elongate and cylindrical; anther operculate, pollinia two, porate or cleft, attached by one or two stipes (tegulae) to one or two viscidia; rostellum cleft or protruding-deflexed and bifid or trifid with central lobe shorter to longer than side lobes. Ovary cylindrical, six-angled or triangular and sometimes winged. (PC)

Distribution The subtribe comprises 49 genera with some 760 species and is centred on Madagascar, Africa, and the neighbouring islands. 338

Two genera with 76 species are confined to the American tropics. (PC)

Anatomy Carlsward et al. (2006a) examined the leaf and root anatomy of 47 genera in Vandeae. They focused primarily on the Angraecinae, using Aeridinae and Polystachyinae as outgroups. Phylogenetically informative characters included tilosomes and the loss of mucilage as well as development of spherical silica bodies in stegmata, which defined the clade of Aeridinae + Angraecinae. Inner tangential wall thickenings of the endovelamen, which appear superficially as tilosomes using light microscopy, were also found to be phylogenetically informative among angraecoids. Aeration units, as defined by Benzing et al. (1983), were present in the root cortex for most Angraecinae and Aeridinae but were absent in Polystachyinae. Detailed comments, if available, are organized by genus below. (BC)

Phylogenetics The two African and Malagasy subtribes of Vandeae, Angraecinae and Aerangidinae, were originally circumscribed by Summerhayes (1966) based on rostellum shape and chromosome number. These differences, however, seem to be phylogenetically misleading. Chromosome counts from several sources (Charard 1963; Jones 1967; Arends et al. 1980; Jonsson 1981; Arends and Van der Laan, 1983, 1986) support a general trend of x = 25 for Aerangidinae and x = 19 for Angraecinae, as indicated by Summerhayes (1966), but within each subtribe there is ample variation. In the most recent molecular phylogenetic analyses of Vandeae (Carlsward et al. 2006b; Micheneau et al. 2008), Aerangidinae and Angraecinae were interdigitated, but together they formed a well-supported clade (Fig. B.4). Therefore, Carlsward et al. (2006b) proposed recognizing a broadly circumscribed subtribe Angraecinae that included Aerangidinae. Based on the work of Carlsward et al. (2006b), the two leafless species of Solenangis (S. aphylla (Thouars) Summerh. and S. cornuta (Rchb.f.) Summerh.) were more closely related to species of Microcoelia than to the leafy species of Solenangis and were therefore transferred to Microcoelia. Chauliodon and Encheiridion are two other African leafless genera that Summerhayes (1943) recognized as segregates of Microcoelia, but their vegetative and floral similarity to Microcoelia is a strong indication that they should be included in Microcoelia (Jonsson 1981). Rhipidoglossum has traditionally been segregated from Diaphananthe by two key features: a column foot and a distinct conical projection or ‘tooth-like’ callus at the opening of the spur (Summerhayes 1960; Garay 1972; Senghas 1986a). However, Summerhayes (1960) and Cribb (1989) observed a complete continuum in these floral character states within Diaphananthe and Rhipidoglossum. Sequence data support the inclusion of Rhipidoglossum with Cribbia and Diaphananthe millarii (Bolus) H.P.Linder, whereas most species of Diaphananthe are more closely related to Chamaeangis (Carlsward et al. 2006b; Fig. B.4). It is clear from these preliminary molecular analyses that Rhipidoglossum and Diaphananthe

ANGRAECINAE

are not monophyletic. However, given the relatively small percentage of Diaphananthe and Rhipidoglossum species sampled in their study (Carlsward et al. 2006b), no nomenclatural changes were made. Species of Aerangis form a monophyletic group with the inclusion of Microterangis hariotiana (Kränzl.) Senghas (the

type species) and M. hildebrandtii (Rchb.f.) Senghas (Carlsward et al. 2006b; Fig. B.4). Before Senghas erected Microterangis (1985), most of its members were included in Chamaeangis section Microterangis (Schlechter 1918). Morphologically, flowers of Microterangis look more similar to Chamaeangis than to Aerangis, but molecular data clearly indicate this to be pollinator

Fig. B.4.  Phylogenetics of Angraecinae. One of 2688 trees resulting from a heuristic search of the combined molecular data (ITS, trnL-F, and matK) from Carlsward et al. (2006b). Filled circles represent clades not present in the strict consensus. Fitch lengths are shown above branches; bootstrap percentages (1000 replicates) are shown below branches.

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convergence. The type species (M. hariotiana) and M. hildebrandtii should have been transferred to Aerangis, but doing so would have left the five species not sampled in their molecular study in a state of nomenclatural limbo, and Carlsward et al. (2006b) felt that the genus needed further study before making any name changes. Angraecum was found to be polyphyletic (Carlsward et al. 2006b; Micheneau et al. 2008), with over 200 species throughout Madagascar and Africa. Bonniera, Oeoniella, and Sobennikoffia are embedded within a clade composed primarily of Angraecum, whereas Jumellea and Aeranthes each seem to be monophyletic (Carlsward et al. 2006b; Micheneau et al. 2008). Generic circumscriptions for Cyrtorchis, Listrostachys, Podangis, Rangaeris, Tridactyle, and Ypsilopus (which form a clade using plastid and nuclear sequence data) may need to be reevaluated with denser taxon sampling than Carlsward et al. (2006b) provided. Species of Cyrtorchis were monophyletic, and Listrostachys and Podangis may also be distinct genera (only one of the two species of Listrostachys was sampled by Carlsward et al. (2006b), and Podangis is monospecific). Some species of Rangaeris were more closely related to Podangis and Tridactyle than they were each other, but there was no clear pattern of relationships seen in the analyses of Carlsward et al. (2006b). The two species of Ypsilopus sampled in that study were more closely related to Tridactyle than they were to one another, so Ypsilopus should probably be transferred to Tridactyle. However, as only six of the 43 species were sampled by Carlsward et al. (2006b), more taxon sampling within Tridactyle is needed to draw any taxonomic conclusions. Mystacidium is monophyletic but embedded in a larger clade that includes several species of Angraecopsis and one species of Sphyrarhynchus (Carlsward et al. 2006b; Fig. B.4). As with much of Angraecinae s.l., more extensive taxon sampling of Mystacidium and Angraecopsis is needed for generic recircumscription. Neotropical genera Campylocentrum and Dendrophylax, as redefined by Carlsward et al. (2003), form well-supported sister clades. Other angraecoid genera from the Old World that form monophyletic groups include Eurychone, Ancistrorhynchus, and Bolusiella (Carlsward et al. 2006b; Micheneau et al. 2008). Cyrtorchis, Oeonia, Neobathiea, Lemurella, Beclardia, and Lemurorchis are all small genera with only one species sampled from each (Carlsward et al. 2006b), and they are likely monophyletic as well. (BC)

Cultivation Most Angraecinae are epiphytes from mainland Africa, Madagascar, and surrounding islands. They occupy a great range of elevations and climates; many have adapted to localized microenvironments. An almost full range of climates and habitats can be covered by the different species in one genus, all requiring a different cultural regime. Even within one species one can find distinct forms from moist evergreen forest, exposed rock, to virtual desert; Aerangis ellisii (B.S.Williams) Schltr., Angraecum sesquipedale Thouars, and A. eburneum Bory are just a few examples. Other species have evolved on

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different islands and require different environments in cultivation. Angraecum leonis (Rchb.f.) André, for example, from the Comoro Islands needs warmth and humidity, whereas those from Madagascar will thrive in a drier atmosphere. In some species groups these forms have developed into distinct species; Angraecum pseudofilicornu H.Perrier/A. scottianum Rchb.f. and the species in the Angraecum germinyanum Hook.f. group (A. germinyanum, A. conchoglossum Schtr., and A. arachnites Schltr.) are just a few examples. Some species from harsh, desert-like areas such as those in Sobennikoffia need to be grown almost as succulents with little water and constant exposure to high light levels. In some genera almost all species prefer shade. For example, virtually all Aerangis and Aeranthes species grow best in wellshaded places. Angraecum and Jumellea have many species that can be found in full sun but others in deep shade. Many species have a distinct flowering season in the wild. This is often related to shorter or longer days or to the onset of the rainy season or a combination of both. Flower initiation is also influenced by a seasonal drop in temperature. Inflorescences of many species can take several months to develop. Some can be reluctant to flower in cultivation, and often a boost in light levels or a drop in temperature can initiate flowering. Angraecum sororium Schltr. and some lithophytic species of Jumellea are just a few troublesome examples. Maintaining a fresh and moving atmosphere around the plants is essential. In the wild, air movement is constant. A gentle breeze is normal and strong winds not uncommon. Ventilation through air-vents in a glasshouse is usually not sufficient to provide this level of air movement for plants; an additional source of movement from fans or cooling apparatus is often necessary to keep leaf temperature down. Additional humidity will also be necessary during the warmer summer months and during the winter when artificial heat decreases humidity levels. Some humidifier and misting devices increase air movement, which can be beneficial to the plants. Stagnant, humid air should be avoided at all times, especially at night; bacterial or fungal infections can set in rapidly. Most Angraecinae and particularly those started off as seedlings can be grown in an open compost mix in clay or plastic pots. Others thrive in the more open conditions of a woodenslatted basket, filled with a mixture of moss, fibre or other materials. The rampant aerial roots of many angraecoids need plenty of space and must be allowed to drain quickly and dry out between waterings. Many plants grow well in cultivation in the natural way—clinging to a piece of cork bark, fibre or suitable wood. The plants are tied tightly to the host with nylon fishing line or some other durable material, incorporating a small pad of moss or other water-retentive material. This method leaves free rein for the aerial roots. Mounted plants can be sprayed or dunked when they become dehydrated. This type of cultivation is essential for twig epiphytes such as Lemurella, Microcoelia, Solenangis, Taeniorrhiza, and many of the smaller angraecoids. Slices of cork or twigs make ideal mounts.

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In general, water should drain quickly, and dry periods between successive watering will approximate the growing conditions of many species and promote healthy growth. Plants in this group often grow slowly, so they will need only small quantities of dilute fertilizer during the growing season. After flowering the rate of growth commonly slows down and stops altogether in the dry season, when there is sometimes a partial leaf-fall. At this stage light can be increased a little and water decreased. The usual orchid pests and diseases will attack Angraecinae. Leaf burn and fungal rot can be problematic, especially as the leaves remain on plants for many years. More specific cultural recommendations can be found below under respective genera. ( JH)

Taxonomic notes Schlechter (1918) attempted a ‘natural’ classification of AfroMadagascan angraecoid orchids, recognizing 32 genera and providing a key to them. He originally proposed Angraecinae and Aerangidinae, two African and Malagasy subtribes of Vandeae, but did not validate them (Schlechter 1926). Summerhayes (1966) circumscribed them based on rostellum shape and chromosome number. Szlachetko (1995) further subdivided the Afro-Madagascan angraecoid orchids into six subtribes: Aeranginae, Angraecinae, Bolusiellinae, Listrostachyinae, Calyptrochilinae, and Rhaesteriinae. His interpretation of relationships was based on column morphology, in particular rostellum and pollinarium structures. Thus his Bolusiellinae (including Bolusiella, Chamaeangis, Cyrtorchis, Diaphanathe, Eggelingia, Eurychone, Nephrangis, Podangis, Rangaeris, Taeniorrhiza, and Tridactyle) have a long, wide, flat, firm, tridentate rostellum, bent down or sometimes U-curved with the middle tooth reduced and finger-like; a large, flat, membranous, entire or split tegula; a large, lamellate viscidium; and two cleft or porate pollinia. His Listrostachyinae (Bonniera, Lemurorchis, Listrostachys, Oeoniella, and Sphyrarhynchus) have a short, fleshy, massive, erect rostellum; two lamellate to thread-like tegulae; and a single large, lamellate viscidium. His Calyptrochilianae (Calyptrochilum only) have a short, triangular, fleshy, massive, acute rostellum, bent towards the tip; a single linear to oblong, elongate tegula; a single elliptic, lamellate viscidium; and two porate pollinia. Finally, his Rhaesteriinae (Rhaesteria only) have a finger-like, long, fleshy, acute rostellum; two thread-like fleshy tegulae; two elliptic, lamellate, small viscidia and two porate viscidia. Calytrochilum and Rhaesteria were not included in Carlsward’s study, but there is no reason to think that they would not fall in the same clade. Schlechter (1918) prepared a complete account of the AfroMadagascan angraecoid orchids at the generic level. Senghas (1986b) updated it, providing good black-and-white line illustrations and photographs of one species of each genus. Szlachetko (1995, 2003a) revised the subtribal delimitations and provided detailed line drawings of the column structure of a species from every genus. (PC)

Taxonomic literature Arends, J. C., van der Burg, W. J., and Van der Laan, F. M. (1980). Notes on African orchids. Miscellaneous papers, Landbouwhogeschool, Wageningen, 19, 23–36. Arends, J. C. and Van der Laan, F. M. (1983). Cytotaxonomy of the monopodial orchids of the African and Malagasy regions. Genetica, 62, 81–94. Arends, J. C. and Van der Laan, F. M. (1986). Cytotaxonomy of the Vandeae. Lindleyana, 1, 33–41. Carlsward, B. S., Whitten, W. M., and Williams, N. H. (2003). Molecular phylogenetics of Neotropical leafless Angraecinae (Orchidaceae): reevaluation of generic concepts. International Journal of Plant Sciences, 164, 43–51. Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Chase, M. W., Cameron, K. M., Barrett, R. L., and Freudenstein, J. V. (2003). DNA data and Orchidaceae systematics: a new phylogenetic classification. In Orchid conservation (ed. K. W. Dixon, S. P. Kell, R. L. Barrett, and P. J. Cribb), pp. 69–89. Natural History Publications, Kota Kinabalu, Sabah, Malaysia. Charard, R. (1963). Contributions a l’étude cyto-taxonomique des Orchidées. Revue de Cytologie et de Biologie Végétales, 26, 1–58. Cribb, P. (1989). Flora of tropical East Africa, Orchidaceae (Part 3). A.A. Balkema, Rotterdam, The Netherlands. Garay, L. A. (1972). On the systematics of the monopodial orchids I. Botanical Museum Leaflets (Harvard University), 23, 149–212. Jonsson, L. (1981). A monograph of the genus Microcoelia (Orchidaceae). Borgströms Tryckeri AB, Motala, Sweden. Micheneau, C., Carlsward, B. S., Fay, M. F., Bytebier, B., Pailler, T., and Chase, M. W. (2008). Phylogenetics and biogeography of Mascarene angraecoid orchids. Molecular Phylogenetics and Evolution, 46, 908–22. Schlechter, R. (1918). Versuch einer natürlichen Neuordnung der afrikanischen agraekoiden Orchidaceen. Beihefte zum Botanischen Centralblatt. Abt. II., 36, 62–181. Schlechter, R. (1926). Das System der Orchidaceen. Notizblatt des Botanischen Gartens und Museums zu Berlin-Dahlem, 9, 590–1. Senghas, K. (1985). Cribbia und Microterangis, zwei neue Orchideen Gattungen. Orchidee, 36, 19–22. Senghas, K. (1986a). Rhipidoglossum. Orchidee, 18, 1110–12. Senghas, K. H. (1986b). 15. Tribus: Vandeae. In Rudolf Schlechter Die Orchideen (ed. F. G., Brieger, R. Maatsch, and K. H. Senghas), 3rd ed., pp. 973–1130. Paul Parey, Berlin. Summerhayes, V. S. (1943). African orchids: XIII. Botanical Museum Leaflets (Harvard University), 11, 137–70. Summerhayes, V. S. (1960). African orchids: XXVII. Kew Bulletin, 14, 126–57. Summerhayes, V. S. (1966). African orchids: XXX. Kew Bulletin, 20, 165–99. Szlachetko, D. L. (1995). Systema orchidalium. Fragmenta Floristica et Geobotanica Supplementum, 3, 1–152. Szlachetko, D. L. (2003a). Gymnostemia orchidalium 3. Acta Botanica Fennica, 176, 1–311.

Artificial key to the genera of Angraecinae (PC)  1. Column with a rostellum deeply indented with auriculate side lobes and sometimes with a small central tooth, ●

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always shorter than side lobe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2   Column with a rostellum in which central lobe is longer than side lobes, the latter often being obscure . . . . . . . 24  2. Plants lacking leaves but sometimes with papery brownish scale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Plants with green leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . 4  3. Inflorescence with many flowers densely and distichously arranged on rachis; flowers only a few mm in diameter; pollinaria one or two; spur rarely up to twice as long as labellum lamina . . . . . . . . 721. Campylocentrum (in part)   Inflorescence 1- to few-flowered, if many-flowered then flowers developing and opening in succession; flowers more than 1 cm in diameter, often much larger; pollinaria two; spur more than twice as long as labellum lamina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 727. Dendrophylax  4. Flowers bright orange with darker sepals and petals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 745. Ossiculum   Flowers not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5  5. Pollinaria two per flower . . . . . . . . . . . . . . . . . . . . . . . . . 14   Pollinarium only one per flower . . . . . . . . . . . . . . . . . . . . 6  6. Column with a foot; spur entrance narrow and at a right angle, spur geniculate . . . . . . . . . . . . 720. Calyptrochilum   Column lacking a foot; spur of different form or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7  7. Labellum with a claw, hence spur entrance distant from column base; spur entrance narrow, sticking out horizontally from labellum; inflorescence secund (flowers turned to the same side) in two ranks and densely many-flowered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 737. Listrostachys   Labellum lacking a claw, hence spur mouth directly at column base or spur absent; inflorescence otherwise . . . . . 8  8. Labellum lacking a spur . . . . . . 717. Angraecum (in part)   Labellum with a spur . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9  9. Labellum simple or tripartite, if tripartite, then inflorescence not with many larger flowers . . . . . . . . . . . . . . . . 10   Labellum tripartite or more; inflorescence many-flowered; flowers more than 1 cm in diameter . . . . . . . . . . . . . . . . 12 10. Petal base plicate; inflorescence 1-flowered; labellum entire, sagittate to pandurate; pollinarium lacking stipe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 734. Jumellea Petal base not plicate; the other features never combined in this form . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11. Flowers resupinate; labellum often with a median keel; spur entrance usually narrow; flowers mostly larger than 1 cm in diam., occasionally smaller, but then mostly greenish . . . . . . . . . . . . . . . . . . . . . . . . 717. Angraecum (in part) Flowers non-resupinate; labellum lacking a keel; spur entrance relatively broad, lamina of labellum gradually merging into spur; flowers smaller than 1 cm in diameter, usually white . . . . . . . . . . 721. Campylocentrum (in part) ●

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12. Rostellum not bidentate . . . . . . . . . . . . . 736. Lemurorchis Rostellum shortly bidentate . . . . . . . . . . . . . . . . . . . . . . . 13 13. Stipe one, broad, agglutinate; column auriculae rounded; spur pointing backward, markedly longer than labellum lamina . . . . . . . . . . . . . . . . . . . . . . . . . . 751. Sobennikoffia Stipes two, separate; column auriculae like pliers; spur nearly adnate to ovary for a few mm . . . . . 744. Oeoniella ●

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14. Column with a boat-shaped foot, at its end a mostly forward-pointing spur (but A. ecalcarata nearly lacks a spur, and A. tricalcarata has three spurs); sepals and petals of distinctly different form . . . . . . . . . . . . . . . . . . . . . . . . . . 15 Column lacking a foot (if present, obscure); sepals and petals of similar form and size . . . . . . . . . . . . . . . . . . . . 16 15. Inflorescence with flowers appearing simultaneously; rachis not wiry . . . . . . . . . . . . . . . . . . . . . . 732. Erasanthe Inflorescence 1- to many-flowered but flowers borne successively; rachis wiry . . . . . . . . . . . . . . . . . . 713. Aeranthes 16. Column with a foot and without auriculae; rostellum tridentate . . . . . . . . . . . . . . . . . . . . . . . 724. Cribbia (in part) Column lacking a foot but with auriculae; rostellum without or with one middle tooth or two middle teeth . . . . 17 17. Inflorescence with many white flowers; shoots with distichously arranged leaves; column auriculae missing or only weakly developed . . . . . . 721. Campylocentrum (in part) Inflorescence laxly flowered, occasionally only 1-flowered; column auriculae well developed . . . . . . . . . . . . . . 18 18. Labellum entire, often with median keel; lamina abruptly leading to spur . . . . . . . . . . . . . 717. Angraecum (in part) Labellum tripartite or more; lamina gradually leading into spur . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 19. Labellum tripartite, partition far in frontal part, basal part tubular-involute, embracing column . . . . . . 714. Ambrella Labellum tripartite or more, partition more toward base and not involute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 20. Flowers non-resupinate; viscidium geniculate, stipe attached at bend . . . . . . . . . . . . . . . . . . . . . 735. Lemurella Flowers resupinate (or only partly so if inflorescence pendent or drooping); viscidium not geniculate . . . . . . . . . 21 21. Petals clawed, the lamina horizontal, broader than long; emarginate apically; labellum 4–8-partite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 725. Cryptopus Petals not clawed; labellum tripartite (except in Neobathiea filicornu and N. keraudrenae) . . . . . . . . . . . . . . . . . . . . . . . . 22 22. Viscidium glabrous; inflorescence 1-flowered; rostellum evenly tridentate; auriculae like the papillose middle tooth in length and width; labellum midlobe apically emarginate . . . . . . . . . . . . . . . . 743. Oeonia (O. madagascariensis) Viscidium papillose; inflorescence many-flowered; rostellum with a distinct elongate middle tooth, but shorter than column auriculae; labellum mostly of a different form . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 ●

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23. Labellum evenly trilobed (entire in N. filicornu and N. keraudrenae); labellum not embracing column; perianth parts reflexed . . . . . . . . . . . . . . . . . . . . . . . . . . . 741. Neobathiea Labellum tripartite, its midlobe horizontally broad and apically emarginate or still more divided; labellum embracing column; perianth parts not reflexed . . . . . . . . . . . . . . . . . . . . . . . . . . 743. Oeonia (except O. madagascariensis) 24. Plants lacking foliage leaves, sometimes with brownish scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 Plants with green leaves . . . . . . . . . . . . . . . . . . . . . . . . . . 28 25. Stem elongate . . . . . . . . . . . . . . 739. Microcoelia (in part) Stem abbreviated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 26. Inflorescence 1-flowered; flowers 2.5 cm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 755. Taeniorrhiza Inflorescence few- to many-flowered; flowers markedly smaller . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 27. Labellum with a high, fleshy, erect, tooth-like callus at spur entrance . . . . . . . . . . . . . . . . . . . . . . . . . . . 723. Chauliodon Labellum always lacking a callosity at spur entrance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 739. Microcoelia (in part) 28. Pollinaria two per flower; viscidia two, separate . . . . . . 29 Pollinarium one per flower; viscidium one, shared by one or two stipes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 29. Spur tip biglobular . . . . . . . . 753. Angraecopsis (in part) Spur thread-like or clavate . . . . . . . . . . . . . . . . . . . . . . . . 30 30. Sepals and petals connate for a third to half their length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 749. Rhaesteria Sepals and petals usually not connate (but see Angraecopsis where the petals are often adnate to lateral sepals) . . . . 31 31. Rostellum 6-partite, with three teeth pointing upward and three downward . . . . . . . . . . . . . . 756. Triceratorhynchus Rostellum of different shape . . . . . . . . . . . . . . . . . . . . . . 32 32. Rostellum bipartite, but teeth strictly parallel pointing upward . . . . . . . . . . . . . . . . . . . . . . . . . . . . 730. Distylodon Rostellum of different shape . . . . . . . . . . . . . . . . . . . . . . 33 33. Rostellum tridentate, lateral teeth markedly papillose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 740. Mystacidium Rostellum of different shape . . . . . . . . . . . . . . . . . . . . . . 34 34. Flowers not fully opening, appearing nearly globose; labellum entire, roundish, without callosity, with a clavate or subspherical spur and a narrow spur entrance; rostellum tridentate with a long middle tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 738. Margelliantha Flowers of different form . . . . . . . . . . . . . . . . . . . . . . . . 35 35. Column with a foot; rostellum only weakly dentate . . . 36 Column lacking a foot; rostellum always prominent . . . 37 36. Labellum mostly circular to subquadrate and about as long as broad, sometimes with a basal tooth; inflorescence mostly pendent, and then flowers non-resupinate . . . . . . . . . . . . . . . . . . . . . . . . . . . . 750. Rhipidoglossum (in part) ●

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Labellum narrowly ovate, at least twice as long as broad, without a basal callus; inflorescence erect, flowers only partially resupinate . . . . . . . . . . . . . 724. Cribbia (in part) Labellum lacking side lobes (but partially incised at its front); rostellum with a mid-tooth; petals and lateral sepals not adnate or coalescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 750. Rhipidoglossum (in part) Labellum markedly tripartite; rostellum remnant bi- or tripartite; petals coalescent with lateral sepals along their basal upper side . . . . . . . . . . . . . . . . . . . 716. Angraecopsis Labellum with a claw . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 Labellum without claw . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 Flowers only a few mm in diameter, resupinate; labellum clawed with a transverse lamina, deeply incised at its tip, lacking a callus, with a spur; spur less than 1 cm long, with two side auricles beside mouth; stem with loosely arranged leaves, rounded at tip; rostellum remnant bilobate; pollinaria with a bifurcate stipe (or two separate stipes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 742. Nephrangis Flowers 8 cm or more in diameter, non-resupinate; labellum erect, clawed at base, obovate with an elongate tip; spur 17–25 cm long, gently spiralling; calli 2, small, within the spur entrance another callus; stem rather short, with densely arranged flat, very long and broad leaves; rostellum long, tooth-like, pointing forward and downward; pollinaria with 2 separate stipites . . 746. Plectrelminthus Pollinaria with 2 separate stipes . . . . . . . . . . . . . . . . . . . 41 Pollinaria with a common stipe . . . . . . . . . . . . . . . . . . . . 49 Labellum clearly or weakly tripartite . . . . . . . . . . . . . . . . 42 Labellum entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 Rostellum with 2 lateral and a shorter middle tooth; flowers white, spreading like a star; sepals and petals are rather similar; column slender; viscidium large, almost oblong; petals never fused with the lateral sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 748. Rangaeris Rostellum entire, subclavate or hamate; flowers not stellate, often subcampanulate; viscidium elliptic or ovate; petals fused with the lateral sepals . . . . . . . . . . . . . . . . . 43 Stem psygmoid, covered unifacial leaves; flowers scarcely open, non-resupinate, white; spur shorter than the labellum lamina; sepals and petals similar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 719. Bolusiella (in part) Stems not arranged psygmoid, leaves bifacial, conduplicate; flowers usually fully open, white, whitish green to yellowish green, pale salmon; spur short to very long; sepals and petals usually of different size and shape; petals often fused with the lateral sepals along their basal side . . . . . . . . . . . . . . . . . . . . . . . . 753. Sphyrarhynchus (in part) Inflorescence usually densely capitate; flowers only half open; lip bag- or funnel- shaped, usually surrounding the column; rostellum with double teeth, teeth first bend downwards, then upward at sharp angle, frequently with a middle tooth . . . . . . . . . . . . . . . . . . 715. Ancistrorhynchus

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47. Habit psygmoid; inflorescence stalked; rostellum deeply bifid in front . . . . . . . . . . . . . . . . . . . . . . . . . 747. Podangis Habit not psygmoid; inflorescence not umbellate; rostellum usually not deeply bifid . . . . . . . . . . . . . . . . . . . . . . . 48 ●

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48. Sepals and petals reflexed; labellum gradually narrowing to mouth of spur; flowers stellate, white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 726. Cyrtorchis Sepals and petals not reflexed; labellum suddenly narrowing to spur mouth; flowers not stellate, usually with greenish, yellowish, bronze, salmon and similar hues . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 728. Diaphananthe ●

●

49. Labellum concave, cordiform; with an auriculate base, clasping the column; inflorescence 1-flowered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 722. Cardiochilos Labellum of a different shape; inflorescence usually manyflowered but, if 1-flowered, then with long spur . . . . . . 50 ●

●

50. Habit psygmoid; leaves imbricate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 719. Bolusiella (in part) Habit not psygmoid; leaves conduplicate, bifacial . . . . . 51 ●

●

51. Rostellum bidentate, both teeth bending first down, then at narrow angle again upwards, in between often shorter middle tooth . . . . . . . . . . 715. Ancistrorhynchus (in part) Rostellum of a different shape . . . . . . . . . . . . . . . . . . . . 52 ●

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52. Labellum tripartite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53 Labellum entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57 ●

Stipe forked, Y-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . 56 56. Labellum concave, kidney-shaped, incised apically, lateral lobules on either side of spur mouth erect; spur not exceeding length of labellum lamina; leaves not grass-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 742. Nephrangis Labellum obscurely divided near its middle, the side lobes not sharply distinct; labellum lamina flat; spur several times the length of labellum; leaves grass-like, division between sheath and leaf blade oblique, a pointed sheath remaining after leaf has fallen . . 758. Ypsilopus (in part) 57. Flowers 4 cm in diameter; spur gradually narrowing, with a clavate extension near tip . . . . . . . . . . . . 733. Eurychone Flowers mostly much smaller, if of similar size, then with a delicate spur, tapering to acuminate or filiform tip . . 58 58. Stipe forked, Y-shaped . . . . . . . . 758. Ypsilopus (in part) Stipe undivided . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59 59. Flowers only half open, non-resupinate, about 2 cm in diameter; labellum strongly concave and marginally undulate; spur entrance only gradually tapering . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 718. Beclardia Flowers of different appearance, mostly smaller; spur entrance abruptly narrow . . . . . . . . . . . . . . . . . . . . . . . . .60 ●

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●

●

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60. Rostellum consisting of one tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 712. Aerangis (in part) Rostellum bidentate, bilobed or tridentate . . . . . . . . . . . 61 ●

●

61. Spur 2.5–20 cm long; viscidium calceolate; labellum concave, fleshy, auriculate at base; rostellum tridentate with middle tooth longer than lateral teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 754. Summerhayesia Spur up to 2 cm long; viscidium flat; labellum flat, lacking auricules; rostellum bidentate or bilobate . . . . . . . . . . . 62 ●

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62. Flowers fully extended; stipe oblong, completely covering viscidium that is only visible from the back; rostellum bidentate; stem up to 40 cm long . . . . . . . 731. Eggelingia Flowers only half open; stipe lanceolate, sitting on top of a much larger viscidium; rostellum bilobate; stem up to 25 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 752. Solenangis (except S. clavata) ●

●

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53. Labellum with three blunt or rounded lobes of equal size; pollinia sitting almost at middle of stipe; roots borne almost to tip of stem . . . . . . . . . . . . . . . 729. Dinklageella Labellum of different shape, if tripartite, then lobes pointed; pollinia attached to tip of stipe; roots only at stem base, if higher then only occasional . . . . . . . . . . . 54 ●

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54. Labellum tripartite with pointed parts; labellum divided almost always in middle or even nearer tip, almost always clearly auriculate at base, also auriculate when labellum is undivided . . . . . . . . . . . . . . . . . . . . . . . . . . . 757. Tridactyle Labellum entire, obscurely tripartite or, if tripartite, then minute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55 ●

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55. Stipe single, linear . . . . . . . . . 752. Solenangis (S. clavata) ●

344

7 1 2 .  A ER A NGI S Aerangis Rchb.f., Flora, 47, 190 (1865). Type species: Aerangis flabellifolia Rchb.f. Radinocion Ridl., Bol. Soc. Brot., 5, 200 (1887). Type species: Radinocion flexuosum Ridl. Rhaphidorhynchus Finet, Bull. Soc. Bot. France, 54(9), 32 (1907), pro parte. Lectotype: Rhapidorhynchus bilobus (Lindl.) Finet (basionym: Angraecum bilobum Lindl.) Barombia Schltr., Orchideen, 600 (1914). Type species: Barombia gracillima (Kränzl.) Schltr. Microterangis (Schltr.) Senghas, Orchidee (Hamburg), 36, 22 (1985). Type species: M. hariotiana (Kränzl.) Senghas (basionym: Mystacidium hariotianum Kränzl.)

AERANGIS

Derivation of name

Leaf

From the Greek, aer, air, and angos, a vessel, in allusion to the epiphytic nature of the plant. (PC)

TRANSVERSE SECTION

Description (Plates 134–136; Fig. 712.1, 712.2) Epiphytic herbs with tough stems bearing numerous, elongate, aerial roots in lower part. Stem usually unbranched, covered with the remains of overlapping leaf-bases, bearing few toseveral leaves apically. Leaves in two rows, usually thick and fleshy, sometimes leathery during dry season, sheathing at base, much longer than broad and usually wider in upper half, unequally bilobed at apex. Inflorescence lateral, racemose (rarely branched), few- to many-flowered; bracts often amplexicaul. Flowers resupinate, white or variously tinted with green or brown, pale green, brown or orange. Sepals and petals free, spreading or reflexed. Labellum entire, often similar to sepals and petals, spurred at base, ecallose or with two low ridges at mouth of spur; spur cylindrical, elongate, decurved or rarely rising above ovary. Column often narrowed toward base and enlarged at level of stigma; androclinium straight or sloping, anther-cap sometimes beaked, pollinia two, sessile on a single stipe attached to a variously shaped viscidium; stigma an oval or rhombic sticky depression; rostellum entire, elongate, deflexed or porrect. Ovary straight or curved. Capsule cylindric or ellipsoid, often elongated. (PC)

Distribution (Fig. 712.3) Aerangis is a genus of 51 species of which at least 26 occur in Africa and the remainder in Madagascar and the Comoro Islands. One species is recorded from both East Africa and Sri Lanka (possibly introduced). (PC)

Infrageneric classification Senghas (1986b) and Szlach. and Olszewski (2001) attempted to subdivide the genus. The latter’s treatment recognized three subgenera as follows: A. subgenus Aerangis A. subgenus Barombia Szlach. & Olszewski, Fl. Cameroun, 36, 832 (2001). Type species: B. gracillima (Kränzl.) Schltr. A. subgenus Uniflorae Senghas, Orchideen ed. 3, 1, 16–18, 1051 (1986). Type species: A. fastuosa (Rchb.f.) Schltr. This treatment does not include the species previously included in Microterangis. See Cribb and Carlsward (2012) for new combinations associated with the synonymization of Microterangis. Stewart (1979, 1986) and Stewart et al. (2006) revised the African and Madagascan species of Aerangis sensu stricto. (PC)

Anatomy Carlsward et al. (2006a) examined 11 species of Aerangis and also the former Microterangis hildebrandtii (Rchb.f.) Senghas, now Aerangis hildebrandtii (Rchb.f.) P.J.Cribb & Carlsward. Their results are summarized below.

Trichomes infrequent; when present glandular, multicellular, frequently sunken; supported by a raised buttress of several epidermal cells in A. coriacea Summerh. and A. ugandensis Summerh. Stomata generally abaxial but ad- and abaxial in A. coriacea, A. somalensis (Schltr.) Schltr., and A. verdickii (De Wild.) Schltr. Hypodermis absent. Fibre bundles absent. Mesophyll homogeneous or heterogeneous with columnar anticlinal adaxial cells and isodiametric abaxial cells in A. verdickii. Waterstorage cells smooth to pitted, banded, or thickened along the edges. Vascular bundle sheath distinct, often containing large and darkly stained chloroplasts. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Velamen 2–6 cells thick. Epivelamen cells isodiametric to radially elongate; walls ∪-thickened with fine anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate. Inner endovelamen cells thin-walled with undulate radial walls and angular tangential walls. Distinct endovelamen wall thickenings absent. Cover cells (Leitgeb 1864; Solereder and Meyer 1930) present over short cells of the exodermis. Exodermal long cells ∩- to ○-thickened. Water-storage cells smooth to banded. Aeration units present in A. biloba (Lindl.) Schltr., A. kirkii (Rchb.f.) Schltr., A. ugandensis, and A. hildebrandtii. Endodermal cell walls primarily ○-thickened to infrequently ∪-thickened. Vascular cylinder 6–24-arch. Vascular tissue embedded in sclerenchyma. Pith sclerenchymatous or parenchymatous. (BC)

Palynology Schill and Pfeiffer (1977) examined pollen of three species of Aerangis and characterized the tetrads as concave-convex with laevigate sculpturing and a calymmate sexine. (AP)

Cytogenetics Arends and Van der Laan (1983) recorded chromosome numbers of 2n = 42–54 for eight species of Aerangis. Brandham (1999) suggested that the basic chromosome number is x = 25 to account for the many records of 2n = 50. He speculated that the high basic number might have an allopolyploid origin, supported by the occurrence of dysploids around the 2n = 50 number. Known polyploids in this genus reach 8x = 200. (AP)

Phytochemistry Chemical composition data are available for the floral fragrances of A. appendiculata (De Wild.) Schltr., A. biloba, A. brachycarpa (A.Rich.) Durand & Schinz, A. confusa J.Stewart, A. cryptodon (Rchb.f.) Schltr., A. distincta J.Stewart & la Croix, A. fastuosa (Rchb.f.) Schltr., A. kirkii, A. kotschyana (Rchb.f.) Schltr., 345

VA N D E A E

A

B

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K

C

D E J

F

G H

Fig. 712.1.  Aerangis distincta J.Stewart & la Croix. A. Habit; B. Flower, side view; C. Dorsal sepal; D. Petal; E. Lateral sepal; F. Labellum; G. Column, front view; H. Column, side view; I. Column, anther cap removed; J. Anther; K. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from living plant, RBG Kew cult.503–87 (A), slide 871 by I. La Croix, and Kew Spirit Collection no. 51484. Reproduced with permission from G. V. Pope (ed.) (1998), Flora Zambesiaca: Mozambique, Malawi, Zambia, Zimbabwe, Botswana. Volume 11, Part 2. Royal Botanic Gardens, Kew.

A. modesta (Hook.f.) Schltr., and A. somalensis, which are emitted during the night (Kaiser 1993, 2011). Of particular interest is the presence of the so-called ‘aerangis lactone’ in the scents of A. confusa and A. kirkii, where it is found at levels of 2–5% and 20–30%, respectively (Fig. 712.4). This unique δ-lactone is a 346

cis-4-methyl-5-decanolide (Kaiser 1993), which is present only as the (4S,5S)-stereoisomer (Bartschat et al. 1995). Enantioselective syntheses of the (4S,5S)-, (4R,5R)-, and (4S,5R)-stereoisomers of 4-methyl-5-decanolide have been achieved (Brenna et al. 2001; Wu et al. 2002; Sabitha et al. 2007). Although literature sources

AERANGIS

C

H I

K J D F

L M

B

N E

G

A

Fig. 712.2.  Aerangis hariotiana (Kränzl.) P.J.Cribb & Carlsward. A. Habit; B. Portion of inflorescence, enlarged; C. Flower; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum; H. Column and labellum, side view; I. Column, front view; J. Column apex, anther cap removed; K. Column, side view; L. Anther cap, dorsal view; M. Anther cap, ventral view; N. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Labat/Yahaha 3137 (K).

commonly refer to these as (–)-cis-, (+)-cis- and (+)-trans-aerangis lactones, respectively, only the first mentioned occurs in Aerangis (Bartschat et al. 1995). Small amounts of methyl 3-methyloctanoate (Fig. 712.4), which may be a metabolite of a precursor of (–)-cis-aerangis lactone, are also found in the floral scents of A. confusa (3.2%) and A. kirkii (1.9%) but not the other species examined (Kaiser 1993, 2011). The structurally related γ-lactone, cis-3-methyl-4-decanolide, occurs only as a minor component of A. kirkii. Synthetic routes to the (3S,4S)- and (3R,4R)stereoisomers of cis-3-methyl-4-decanolide are available (Wu et al. 2002). (NV, RG)

Phylogenetics Senghas (1985, 1986b) separated the Malagasy and Comorian species previously ascribed to Chamaeangis to his new genus Microterangis, which he distinguished by its pollinarium structure with the pollinia attached to a single tegula (stipe) rather than each pollinium being attached to its own tegula. Otherwise, he recognized Microterangis in the same key couplet as Aerangis, distinguishing the former by its smaller yellow, green, or orange flowers rather than predominantly white flowers. He acknowledged the similarity of Microterangis to Aerangis in the name he chose. Szlachetko (2003a) illustrated the floral structure of both 347

VA N D E A E

Fig. 712.3.  Distribution map of Aerangis.

Fig. 712.4.  Scent constituents of Aerangis confusa and Aerangis kirkii.

Aerangis and Microterangis, showing their similarity in column, stigma, rostellum, and pollinarium shape. Flower colour is also not as clear-cut as Senghas (1985, 1986b) suggested because some of the smaller-flowered Aerangis species have a pale yellow or green perianth, and even the larger-flowered ones have a white perianth variously flushed with pink, salmon-pink, brown, or green. Carlsward et al. (2006b) included 16 species of Aerangis and two of Microterangis, including the type species, M. hariotiana Senghas, in their DNA-based analyses of vandaceous

348

orchids. The Microterangis species were nested within Aerangis, sister to the Malagasy species, A. punctata Stewart (1986). It seems likely that Microterangis are just small-flowered species of Aerangis that evolved in the Comoros or northern Madagascar from a common ancestor of larger-flowered Malagasy Aerangis. (PC, BC)

Ecology Most species are found in humid evergreen and semi-deciduous forests and woodland, in shade or sunlight from sea level to 2000 m. Aerangis ellisii (B.S.Williams) Schltr. can grow in both forest in the shade and on rocky outcrops in full sun. Aerangis decaryana H.Perr. grows in dry spiny forest of Tamarindus L. (Fabaceae), Alluaudia Drake (Didiereaceae), Euphorbia L. (Euphorbiaceae), and other succulents in southern Madagascar. Aerangis pulchella (Schltr.) Schltr. inhabits dry deciduous forest in western Madagascar. (PC)

AERANTHES

Pollination Nilsson et al. (1987) found pollinia of Aerangis fuscata (Rchb.f.) Schltr. attached to the hawkmoth species, Panogena lingens. As the moth hovers against the opening of the spur, the viscidium attaches to its frons and/or palpi. This same hawkmoth species as well as another, Agrius convolvuli, visited Aerangis ellisii as determined by the scales/hairs of the moths left on the stigma (Nilsson et al. 1988). The relationship among proboscis length, nectar position in the spur, and spur morphology limits pollination to these two hawkmoth species. Martins and Johnson (2007) studied pollination biology of Rangaeris amaniensis (Kränzl.) Summerh. and four species of Aerangis (A. brachycarpa, A. confusa, A. thomsonii (Rolfe) Schltr., and A. kotschyana) in Kenya. All but A. confusa have spur lengths > 10 cm and were pollinated by hawkmoth species Agrius convolvuli and Coelonia fulvinotata. Aerangis confusa with a spur length of c. 4 cm was pollinated by the short-tongued hawkmoths Hippotion celerio and Daphnis nerii. Nectar often filled the spur in some of the species, and sugar concentration varied from 1% at the mouth of the spur to 20% at the tip. Such gradients were most pronounced in species with long, straight spurs so that the shorttongued pollinators cannot gain access to the most concentrated nectar. Individual visits lasted no more than five seconds. The hawkmoths followed the fragrance trail and approached flowers upwind, circled the tree or bushes a few times, and then hovered while unrolling their proboscides to probe individual flowers for no more than three seconds. When a moth did insert its proboscis fully into a flower, its head would end up flush with the column. In some cases, the viscidium would be attached to either the proboscis or regions of the head where scales were missing. As the hawkmoth probed another flower, the attached pollinia would be deposited on the sticky mucilage of the concave stigma. Fruit set was 5.0–17.5%. Bagged, unmanipulated flowers did not produce capsules. (AP)

Uses A paste made from A. thomsonii was said to ensure hatching of hen’s eggs in East Africa and was also used to treat abscesses and hernias. Leaves of A. biloba were used as an emetic in Gabon, whereas its roots were sometimes fashioned into guitar strings there (Lawler 1984). Many species are common in hobbyist collections (see cultivation notes in subtribal treatment). (AP)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Cribb, P. J. and Carlsward, B. (2012). New combinations in Aerangis, Diaphananthe and Podangis (Orchidaceae, subtribe Angraecinae). Phytotaxa, 71, 42–7. Micheneau, C., Carlsward, B. S., Fay, M. F., Bytebier, B., Pailler, T., and Chase, M. W. (2008). Phylogenetics and biogeography of Mascarene angraecoid orchids. Molecular Phylogenetics and Evolution, 46, 908–22.

Senghas, K. H. (1985). Cribbia und Microterangis, zwei neue Orchideengattungen. Orchidee, 36, 19–22. Senghas, K. H. (1986b). Tribus: Vandeae. In Rudolf Schlechter Die Orchideen, 3rd ed. (ed. F. G. Brieger, R. Maatsch, and K. H. Senghas), pp. 973–1130. Blackwell, Berlin. Szlachetko, D. and Olszewski, T. (2001). In Flore du Cameroun 36. Orchidacees, vol. 3 (ed. G. Achoundong and P. Morat), pp. 790–808. Ministere de la Recherche Scientifique et Technique, Yaounde, Cameroun. Stewart, J. (1979). A revision of the African species of Aerangis (Orchidaceae). Kew Bulletin, 34, 239–319. Stewart, J. (1986). Stars of the islands. A new look at the genus of Aerangis in Madagascar. American Orchid Society Bulletin, 55, 792–802, 902–9, 1008–15, 1117–1125. Stewart, J., Hermans, J., and Campbell, D. (2006). Angraecoid orchids. Timber Press, Portland, Oregon. Szlachetko, D. L. (2003a). Gymnostemia orchidalium 3. Acta Botanica Fennica, 176, 1–311.

7 1 3 .   A ER A NT H ES Aeranthes Lindl., Bot. Reg., 10, t. 817 (1824). Type species: Aeranthes grandiflora Lindl. (as A. grandiflorus Lindl.)

Derivation of name From the Greek, aer, air, and anthos, flower, from the epiphytic nature of the plant. (PC)

Description (Plates 137, 138; Fig. 713.1) Epiphytic herbs with leafy stems. Roots wiry, branching. Leaves linear, distichous, coriaceous. Inflorescence racemose or rarely branching, wiry, usually bearing flowers in succession, rarely simultaneously. Flowers translucent, white, pale yellow, or green. Sepals long-attenuate. Petals smaller than sepals, inserted on column foot. Labellum attached to column foot, entire, often square to oblong, broader than sepals and petals, long-attenuate in front, hypochile inflated and spurred, epichile trullate to ovate. Column with an inflated foot; pollinia two, each attached by a short stipe (tegula) to its own linear viscidium; rostellum trifid with a short central tooth, deeply emarginate. (PC)

Distribution (Fig. 713.2) Aeranthes is a genus of 43 species in Madagascar, the Mascarenes, and the Comoros with two species in Zimbabwe; 37 species occur in Madagascar. (PC)

Anatomy Carlsward et al. (2006a) examined five species of Aeranthes. Their results are summarized here.

Leaf TRANSVERSE SECTION

Trichomes multicellular, glandular. Stomata abaxial only. Hypodermis adaxial, composed of one or two rows of large, thin-walled, birefringent, usually pleated water-storage cells. Fibre bundles distributed in a single row along the abaxial

349

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C M

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Fig. 713.1.  A–J. Aeranthes africana J.Stewart. A. Habit; B. Flower, front view; C. Floral bud, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Column and labellum, side view; H. Column, anther cap removed; I. Anther cap; J. Pollinia. Drawn by Judi Stone from J.S. Ball 1283 (K). K–M. Aeranthes parkesii G.Will. K. Habit; L. Flower; M. Labellum. Drawn by Judi Stone from slide by A. Cambitzis. Single bar = 1 mm, double bar = 1 cm. Reproduced with permission from G. V. Pope (ed.) (1998), Flora Zambesiaca: Mozambique, Malawi, Zambia, Zimbabwe, Botswana. Volume 11, Part 2. Royal Botanic Gardens, Kew.

mesophyll, composed of several thick-walled lignified cells surrounding 1–3 thin-walled cells. Mesophyll mainly homogeneous; heterogeneous with columnar adaxial cells and smaller isodiametric abaxial cells in A. ramosa Rolfe; isobilateral in A. peyrotii Bosser, composed of isodiametric cells near ad- and abaxial 350

surfaces and anticlinally oriented cells in the central mesophyll. Water-storage cells smooth to banded. Vascular bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma and fibre bundles.

AMBRELLA

(Kaiser 1993). The relative proportions of these diketones as constituents of the scent of a specimen analysed in 2003 were 10.2%, 3.9%, and 0.2%, respectively, although cis-linalool oxide (furanoid form, 38.9%) and benzaldehyde (14.6%) were still the most prominent components (Kaiser 2011). (NV, RG)

Ecology Most species are found in deep shade in coastal, riverine, and wet forests from sea level to 2000 m but usually above 500 m. Aeranthes crassifolia Schltr. and A. parvula Schltr. are found in semideciduous woodland on Tamarindus L. (Fabaceae). Aeranthes sambiranoensis Schltr., A. schlechteri Bosser, and A. tropophila Bosser can be found in seasonally dry, deciduous and semi-deciduous woodland. (PC)

Pollination Although Aeranthes clearly fits the sphingophilous syndrome, pollination studies have not yet been published. (AP) Fig. 713.2.  Distribution map of Aeranthes.

Root TRANSVERSE SECTION

Velamen one or two cells thick. Epivelamen cells radially elongate to isodiametric; walls ∪-thickened with fine anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate. Distinct endovelamen thickenings ridged when present. Cover cells present over short cells of exodermis. Exodermal long cell walls ∩-thickened. Water-storage cell walls smooth to pitted. Single modified cortical layer of thin- to thickwalled cells surrounding the endodermis in A. ramosa. Aeration units present in A. arachnites (Thouars) Lindl., A. caudata Rolfe, and A. grandiflora. Endodermal cell walls ○-thickened. Vascular cylinder 6–16-arch. Vascular tissue embedded in sclerenchyma. Pith sclerenchymatous. (BC)

Palynology According to Schill and Pfeiffer (1977) who studied three species of Aeranthes, pollen tetrads are convex with laevigate sculpturing and a calymmate sexine. (AP)

Cytogenetics Arends and Van der Laan (1983) published a chromosome count of 2n = 38 for A. caudata Rolfe. (AP)

Phytochemistry The floral fragrance of the night-scented A. grandiflora comprises mainly cis-linalool oxide (furanoid form, 29.8%), benzaldehyde (24.6%), and 4-oxo-3-hexanol (10.5%). From an olfactory point of view, important contributions to the scent are made by the diketones pentane-2,3-dione (1.8%), hexane-3,4-dione (< 1%), and 1-phenylpropane-1,2-dione (< 1%), as well as by benzaldehyde

Uses Some species of Aeranthes are occasionally cultivated (see cultivation notes in the subtribal treatment).

Taxonomic notes Carlsward et al. (2006b) and Micheneau et al. (2008) included 12 taxa of Aeranthes in their DNA analyses of the angraecoid orchids. Aeranthes comprised a clade sister to Jumellea. Stewart et al. (2006) illustrated several species with colour photographs. (PC)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Micheneau, C., Carlsward, B. S., Fay, M. F., Bytebier, B., Pailler, T., and Chase, M. W. (2008). Phylogenetics and biogeography of Mascarene angraecoid orchids. Molecular Phylogenetics and Evolution, 46, 908–22. Stewart, J., Hermans, J., and Campbell, D. (2006). Angraecoid orchids. Timber Press, Portland, Oregon.

7 1 4 .   A M B R EL L A Ambrella H.Perrier, Bull. Soc. Bot. France, 81, 655 (1934). Type species: Ambrella longituba H.Perrier

Derivation of name The generic name is derived from Montagne d’Ambre in northeastern Madagascar, where the type was collected. (PC)

Description (Plate 139; Fig. 714.1) Epiphytic herbs, up to 10 cm tall. Leaves five or six, elliptic or obovate-oblong, unequally bilobed at tip. Inflorescence equalling 351

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A F Fig. 714.1.  Ambrella longituba H.Perrier. A. Habit; B. Flower, side view; C. Flower, from above; D. Open flower, from above; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum; I. Anther cap. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from a photograph (D), and others redrawn after H. Perrier de la Bâthie, Bulletin de la Société Botanique de France, 81, 655 (1934).

leaves, one-flowered; bracts amplexicaul. Flower tubular, green. Sepals and petals free, not spreading widely, linear or narrowly oblanceolate, acute. Labellum trilobed in apical part, as long as the sepals or slightly longer, completely enveloping column and rolled into a narrow tube in lower two-thirds, ecallose, spurred at base, spur recurved in middle, cylindrical-tapering, 3.5 cm long. Column with pollinia two, each attached by its own stipe (tegula) to a separate viscidium; rostellum trilobed with an elongate midlobe. (PC)

Distribution (Fig. 714.2) Ambrella is a monospecific genus endemic to Madagascar. (PC) 352

Anatomy Vegetative anatomy of this species has not been studied. (BC)

Palynology Nothing is known about the pollen of Ambrella. (AP)

Cytogenetics Chromosome counts of A. longituba have not been published. (AP)

ANC ISTRORHYNCHUS

Cephalangraecum Schltr., Beih. Bot. Centralbl., 36, 137 (1918). Type species: not designated. Phormangis Schltr., Beih. Bot. Centralbl., 36, 103 (1918). Type species: Phormangis schumannii (Kränzl.) Schltr. (basionym: Angraecum schumannii Kränzl.)

Derivation of name From the Greek ankistron, fishhook, and rhynchos, snout, in reference to the form of the column apex. (PC)

Description (Plate 140; Fig. 715.1)

Fig. 714.2.  Distribution map of Ambrella.

Phytochemistry No reports have been found on the phytochemistry of Ambrella. (NV, RG)

Ecology The species is epiphytic on Calliandra alternans Benth. (Fabaceae) at about 800 m. (PC)

Pollination Pollination vectors of this species are as yet unknown. (AP)

Uses There are no known uses of A. longituba, and it is not in common cultivation. (AP)

Epiphytic herbs. Roots clustered toward base of stem. Stem covered by persistent sheathing leaf bases. Leaves imbricate, suberect, spreading or recurved, ligulate or tapering to apex, unequally bilobed at apex, lobes sometimes toothed, coriaceous or fleshy. Inflorescence axillary from lower leaves, almost sessile, usually forming globose or ellipsoidal heads; bracts chartaceous to membranous, as long as flowers. Flowers mostly white, often marked with green on labellum. Sepals and petals free, subsimilar. Labellum oblong to orbicular, entire to trilobed, spurred at base, lacking a callus; spur straight to sigmoid or geniculate, sometimes swollen at apex and/or opening. Column with anther cap hemispherical, produced in front into a truncate appendage, pollinia two, globose, stipes either two and subspatulate or one and bifid, viscidium long and narrow; rostellum projecting down then abruptly geniculate in middle and upcurved in apical half, bilobed after removal of pollinarium, each lobe falcate and acute. (PC)

Distribution (Fig. 715.2) Ancistrorhynchus is a genus of 17 species widespread in tropical Africa. Nine species have been recorded from East Africa. (PC)

Anatomy Carlsward et al. (2006a) examined three species of Ancistrorhynchus. Their results are summarized below.

Taxonomic notes

Leaf

Ambrella resembles Aerangis vegetatively, but its inflorescence is one-flowered and floral morphology distinctive. It is apparently an isolated and morphologically unique genus that Senghas (1986b) linked with Calyptrochilum. (PC)

TRANSVERSE SECTION

Taxonomic literature Senghas, K. H. (1986b). Ambrella H.Perr. In Rudolf Schlechter Die Orchideen, 3rd ed. (ed. F. G. Brieger, R. Maatsch, and K. Senghas), pp. 1015–16. Paul Parey, Berlin.

715.   A N C IS TRO R H Y N C H U S Ancistrorhynchus Finet, Mem. Soc. Bot. France, 9, 44–47 (1907). Type species: Ancistrorhynchus recurvus Finet

Trichomes glandular, multicellular. Stomata abaxial in A. clandestinus (Lindl.) Schltr. and A. metteniae (Kränzl.) Summerh.; ad- and abaxial in A. refractus (Kränzl.) Summerh. Hypodermis ad- and abaxial, composed of a single row of fibrous idioblasts scattered among thin-walled chlorenchyma cells. Fibre bundles absent. Mesophyll heterogeneous with columnar anticlinal adaxial cells and isodiametric abaxial cells in A. metteniae; isobilateral in A. clandestinus and A. refractus with columnar anticlinal cells situated on either side of a central row of isodiametric cells. Water-storage cells smooth to pitted or spirally thickened. Vascular bundle sheath distinct or indistinct in A. clandestinus and A. refractus. Stegmata contain spherical, roughsurfaced silica bodies found in association with vascular bundle sclerenchyma. 353

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Fig. 715.1.  Ancistrorhynchus tenuicaulis Summerh. A. Habit; B. Inflorescence; C. Bract; D. Flower, front view; E. Flower, side view; F. Dorsal sepal; G. Lateral sepal; H. Petal. I. Labellum; J. Column and labellum; K. Column, front view; L. Column, side view; M. Anther cap; N. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Kew Spirit Collection no. 50070. Reproduced with permission from G. V. Pope (ed.) (1998), Flora Zambesiaca: Mozambique, Malawi, Zambia, Zimbabwe, Botswana. Volume 11, Part 2. Royal Botanic Gardens, Kew.

Root TRANSVERSE SECTION

Velamen 2–7 cells thick. Epivelamen cells isodiametric to radially elongate; walls ∪-thickened with fine anastomosing radial bands. Endovelamen cells angular and isodiametric to radially elongate. 354

Distinct endovelamen wall thickenings absent. Cover cells present over short cells of exodermis. Exodermal long cells ∩- to ○-thickened. Water-storage cells smooth to pitted or banded. Aeration units present in A. metteniae and A. refractus. Endodermal cells ○-thickened to infrequently ∪-thickened in A. metteniae.

ANGRAECOPSIS

Uses There are no ethnobotanical uses for these species, but a few, especially A. metteniae and A. clandestinus, are occasionally available in the trade. (AP)

Taxonomic notes Ancistrorhynchus is sister to a clade that includes Bolusiella and Microcoelia (Carlsward et al. 2006b). Stewart et al. (2006) illustrated several species. (PC)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Stewart, J., Hermans, J., and Campbell, D. (2006). Angraecoid orchids. Timber Press, Portland, Oregon.

Fig. 715.2.  Distribution map of Ancistrorhynchus.

Vascular cylinder 12–18-arch. Vascular tissue embedded in sclerenchyma, cell walls of embedding tissue in A. metteniae thickest around phloem clusters. Pith sclerenchymatous. (BC)

Palynology There is no published information of pollen of any species of this genus. (AP)

Cytogenetics Arends and Van der Laan (1983) reported chromosome counts of 2n = 48, c. 50, 72, 96. Brandham (1999) suggested a basic number of x = 24 based on these records, but he noted that the occurrence of sterile triploids would be obviated if the basic number were x = 12, in which case all records would be of fertile polyploids. (AP)

7 1 6 .   A NGR A EC OPS I S Angraecopsis Kränzl., Bot. Jahrb. Syst., 28, 171 (1900). Type species: A. tenerrima Kränzl. Coenadenium (Summerh.) Szlach., Ann. Bot. Fenn., 40, 70 (2003). Type species: A. breviloba Summerh., syn nov. Holmesia P.J.Cribb, Kew Bull., 32, 175 (1977). Type species: H. parva P.J.Cribb Microholmesia P.J.Cribb, Mabberley, The Plant Book, 371 (1987). Type species: M. parva (P.J.Cribb) P.J.Cribb

Derivation of name From the Indonesian anggrek, orchid, and the Greek opsis, appearance, in allusion to the resemblance of the type species to an Angraecum. (PC)

Description (Plate 141; Fig. 716.1)

Species of Ancistrorhynchus are epiphytic or rarely lithophytic in humid, evergreen and semi-deciduous forests at 100–2100 m elevation. (PC)

Epiphytes or rarely lithophytic herbs. Roots flexuous, terete or flattened, simple or branching. Leaves distichous, ligulate to oblanceolate, often falcate, unequally bilobed at apex, sometimes twisted at base and articulated to imbricate sheathing bases. Inflorescence axillary, one to several, racemose, few- to many-­ flowered, peduncle wiry, often longer than rachis. Flowers white, pale green or yellow-green. Sepals similar or dissimilar; lateral sepals often oblanceolate and produced at one side at base. Petals often adnate to base of lateral sepals and obliquely triangular. Labellum trilobed or rarely entire, ecallose, spurred at base, spur sometimes inflated at apex. Column fleshy; pollinia two, globose, stipites two, linear or rarely oblanceolate, viscidia two (rarely one), elongate; rostellum trilobed. (PC)

Pollination

Distribution (Fig. 716.2)

There are no reports on pollination of any species of this genus. (AP)

Angraecopsis is a genus of 22 species occurring mostly in tropical Africa with one in Madagascar. (PC)

Phytochemistry Information on chemical constituents of this genus is minimal, comprising only a record that two species were tested for alkaloids but did not contain them at levels ≥ 0.1% (Lüning 1974). (NV, RG)

Ecology

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G H E Fig. 716.1.  A–J. Angraecopsis parviflora (Thouars) Schltr. A. Habit; B. Flower; C. Dorsal sepal; D. Lateral sepal and petal; E. Column and labellum, side view; F. Labellum, front view; G. Column with pollinaria, front view; H. Column with anther removed; I. Anther cap; J. Pollinaria. Drawn by Maureen E. Church from la Croix 49. K–U. Angraecopsis amaniensis Summerh. K. Habit; L. Flower; M. Dorsal sepal; N. Lateral sepal and petal; O. Column and labellum, side view; P. Labellum, front view; Q. Column, side view; R. Column and pollinaria, front view; S. Column, pollinaria removed; T. Anther cap; U. Pollinarium with back and side views of stipe and viscidium. Drawn by Maureen E. Church from la Croix 168. V. Angraecopsis breviloba Summerh. Flower. Drawn by Maureen E. Church from unidentified source material. Single bar = 1 mm, double bar = 1 cm. Reproduced with permission from P.J. Cribb (1989), Flora of Tropical East Africa, Orchidaceae (Part 3). A.A. Balkema, Rotterdam/Brookfield.

356

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Cytogenetics Arends and Van der Laan (1983) published chromosome counts of 2n = 50 for both A. gracillima (Rolfe) Summerh. and A. pusilla Summerh. Brandham (1999) also reported 2n = 48. (AP)

Phytochemistry Flowers of A. amaniensis Summerh. emit a strong scent after sunset, the composition of which was analyzed by Kaiser (1993). The fragrance is mainly due to a combination of the major components, benzyl acetate (54.5%), (Z)-3-hexenyl acetate (15.2%), methyl benzoate (8.4%), and methyl 2-methoxybenzoate (8.1%), together with geraniol (2.1%). Although no chemical data are available, the scent composition of A. breviloba Summerh. is said to be similar (Kaiser 1993). (NV, RG)

Ecology Species are epiphytic or lithophytic in evergreen and semi-deciduous forests at elevations from sea level to 1800 m. Smaller species can occur on smaller axes. (PC) Fig. 716.2.  Distribution map of Angraecopsis.

Anatomy Carlsward et al. (2006a) examined three species of Angraecopsis. Their results are summarized here.

Leaf TRANSVERSE SECTION

Trichomes multicellular, glandular. Stomata abaxial in A. parviflora (Thouars) Schltr.; ad- and abaxial in A. breviloba Summerh. Hypodermis absent. Fibre bundles absent. Mesophyll homogeneous. Water-storage cells smooth to pitted. Mucilage globules present throughout mesophyll of A. parviflora. Vascular bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Velamen two or three cells thick. Epivelamen cells isodiametric; walls ∪-thickened with fine anastomosing radial bands. Endovelamen cells angular and isodiametric to radially elongate. Distinct endovelamen wall thickenings absent. Cover cells present over short cells of exodermis in A. amaniensis Summerh. and A. parviflora. Exodermal long cells primarily ○-thickened. Mucilage abundant throughout in A. parviflora. Water-storage cell walls smooth to pitted or thickened along the edges. Aeration units present in A. amaniensis, A. breviloba, and A. parviflora. Endodermal cell walls ○-thickened or thin-walled (A. parviflora). Vascular cylinder 6–10-arch. Vascular tissue embedded in sclerenchyma, cell walls of embedding sclerenchyma thickest around phloem clusters in A. breviloba. Pith sclerenchymatous. (BC)

Palynology Pollen of this genus has not been studied. (AP)

Pollination There is no pollination information for the genus. (AP)

Uses There are no uses known for species of Angraecopsis, but a few such as A. amaniensis and A. gracillima are occasionally cultivated. (AP)

Taxonomic notes Summerhayes (1951) revised Angraecopsis and recognized three sections: Angraecopsis, Cardiochilos Summerh., and Coenadenium Summerh. Carlsward et al. (2006b) included species from two sections in their DNA analyses. Angraecopsis parviflora of A. sect. Angraecopsis, clearly closely related to the type species of the genus (A. tenerrima Kränzl.), is sister to a clade that includes Mystacidium, Sphyrarhynchus, and Angraecopsis sect. Coenadenium. Szlachetko (2003b) raised Coenadenium to generic rank. Angraecopsis, as currently circumscribed, is undoubtedly paraphyletic (Carlsward et al. 2006b). However, Angraecopsis tenerrima Kränzl., the type species, has not so far been sequenced. Further DNA sequencing will be necessary before any taxonomic revision of Angraecopsis is made, and the genus clearly needs revision. Angraecopsis comprises only those species of A. sect. Angraecopsis, recognized by their oblanceolate lateral sepals incumbent on the column foot, triangular or dolabriform petals, and pollinia attached by separate tegulae to distinct viscidia, as well as species of A. sections Cardiochilos and Coenadenium. Rice (2005) included Margelliantha P.J.Cribb, Cribbia Senghas, Coenadenium (Summerh.) Szlach., and some species of Diaphananthe (D. sect. Rhipidoglossum) in a more broadly circumscribed Angraecopsis based upon his observations of the drawings of floral dissections by Summerhayes and those by Szlachetko of column and pollinarium structure (Szlachetko 2003b). However, 357

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this genus is then extremely diverse in habit and floral morphology. He later included Rhipidoglossum and Angraecopsis sensu Rice (2005) within an even more broadly defined Mystacidium Lindl. (Rice 2006b). (PC)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Rice, R. (2005). A preliminary checklist and survey of the subtribe Aerangidinae (Orchidaceae). Oasis Publications, Dora Creek, New South Wales, Australia. Rice, R. (2006b). Notes on the subtribe Aerangidinae (Orchidaceae). Oasis, 3, 12–15. Summerhayes, V. S. S. (1951). New orchids from Africa. Botanical Museum Leaflets, Harvard University, 14, 215–61. Szlachetko, D. (2003b). Coenadenium (Summerh.) Szlach. Annales Botanici Fennici, 40, 70.

717.  AN G R A E C UM Angraecum Bory, Voy. Iles Afrique, 1, 359, t. 19 (1804). Type species: Angraecum eburneum Bory Angorkis Thouars, Nouv. Bull. Sci. Soc. Philom. Paris, 1, 318 (1809). Superfluous substitute for Angraecum Bory Angorchis Thouars, Cahier des six plances (1804–19), orth. var. of Angorkis Aerobion Kaempf. ex Spreng., Syst. Veg., 3, 679 (1826). Type not designated. Macroplectrum Pfitzer in Engl. & Prantl, Natur. Pflanzenf., 2, 6, 214 (1889). Type species: Macroplectrum sesquipedale (Thouars) Pfitzer (basionym: Angraecum sesquipedale Thouars) Bonniera Cordem., Rev. Gén. Bot., 11, 416 (1899). Type species: not designated Lepervenchea Cordem., Rev. Gén. Bot., 11, 416 (1899). Type species: Lepervenchea tenuifolia (Frapp. ex Cordem.) Cordem. (basionym: Angraecum tenuifolium Frapp. ex Cordem.) Pectinaria (Benth.) Cordem., Rev. Gen. de Bot., 11, 412 (1899), nom. illeg. Type species: Pectinaria pectinatum (Thouars) Cordemoy (basionym: Angraecum pectinatum Thouars) Ctenorchis K.Schum., Just’s Bot. Jahresber., 27, 467 (1901). Type species: Ctenorchis pectinata (Thouars) K.Schum. (basionym: Angraecum pectinatum Thouars) Monixus Finet, Bull. Soc. Bot. Fr., 54(9), 15 (1907). Lectotype: Angraecum striatum Thouars (vide Garay, Kew Bull., 28, 496. 1974) Dolabrifolia (Pfitzer) Szlach. & Romowicz, Richardiana, 7, 54 (2007). Type species: Dolabrifolia disticha (Lindl.) Szlach. & Romowicz (basionym: Angraecum distichum Lindl.)

Derivation of name The generic name is derived from the Indonesian word anggrek, orchid. (PC)

Description (Plates 142–145; Fig. 717.1) Epiphytic or less commonly lithophytic herbs, sometimes psygmoid. Stem erect to pendent, leafy. Leaves distichously arranged 358

or in a fan, dorsiventrally flattened, V-shaped in cross-section, unequally bilobed at tip. Inflorescences one to many in a fascicle, axillary, solitary- or many-flowered, usually unbranched. Sepals and petals free, subsimilar or petals smaller than sepals, usually tapering. Labellum entire, with or without a basal linear callus, with a globose, clavate or filiform spur. Column lacking a foot; pollinia two, each attached to its own linear or elliptic viscidium; rostellum deeply cleft, sometimes with a median tooth. (PC)

Distribution (Fig. 717.2) Angraecum comprises 221 species, occurring mainly in Africa, Madagascar, and the adjacent islands of the Indian Ocean. Angraecum zeylanicum Lindl. occurs in the Seychelles and Sri Lanka. (PC)

Infrageneric classification (PC) Schlechter (1918) established six sections: A. sect. Pectinaria, A. sect. Lepervenchea, A. sect. Conchoglossum, A. sect. Pseudojumellea, A. sect. (Eu-)Angraecum, and A. sect. Hadrangis. Garay (1973) developed this and produced a key to the infrageneric classification of Angraecum in which he recognized 19 sections. His key is modified below:  1. Inflorescence 1-flowered, rarely 2-flowered . . . . . . . . . . . 2   Inflorescence many-flowered . . . . . . . . . . . . . . . . . . . . . 10  2. Plants with a few, basal leaves . . . . . . . . . A. sect. Acaulia   Plants several to many leaves along stem . . . . . . . . . . . . . 3  3. Ovary lacking a stalk; inflorescence stalk scarcely developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4   Ovary with a distinct stalk; inflorescence stalk distinctly developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5  4. Leaves laterally compressed, densely imbricate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. sect. Dolabrifolia   Leaves with a distinct blade, or linear, never bilaterally compressed, laxly distichous . . . . . . . . A. sect. Pectinaria  5. Inflorescence stalk covered entirely with 3 or 4, laterally compressed sheathing bracts; ovary triangular in crosssection . . . . . . . . . . . . . . . . . . . A. sect. Perrierangraecum   Inflorescence stalk covered at base only with 1 or 2 adpressed, but not compressed sheathing bracts; ovary terete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6  6. Inflorescence stalk shorter than internodes . . . . . . . . . . . 7   Inflorescence stalk longer than internodes . . . . . . . . . . . 8  7. Spur of labellum as long as the ovary or commonly shorter, rarely surpassing it . . . . A. sect. Conchoglossum   Spur of labellum many times longer than the ovary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. sect. Filangis  8. Spur cylindrical or club-shaped . . . A. sect. Angraecoides   Spur filiform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9  9. Flowers resembling those of Jumellea; labellum rarely apiculate; sepals and petals usually as long as labellum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. sect. Pseudojumellea   Flowers not as above; labellum ovate to transversely oblong, apiculate; sepals and petals often much longer than labellum . . . . . . . . . . . . . A. sect. Arachnangraecum ●

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Fig. 717.1.  A–I. Angraecum geniculatum G.Will. A. Habit; B. Flower; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum; G. Column, anther cap removed, front view; H. Column, anther cap removed, side view; I. Anther cap. Drawn by Judi Stone from Holmes 088 (K). J–P. Angraecum chimanimaniense G.Will. J. Habit; K. Flower, side view; L. Dorsal sepal; M. Lateral sepal; N. Petal. O. Labellum; P. Column, anther cap removed, side view. Drawn by Judi Stone from Ball 412 (K) and painting by J.S. Ball, Southern African Epiphytic Orchids (1978), based on J.S. Ball 412. Single bar = 1 mm, double bar = 1 cm. Reproduced with permission from G. V. Pope (ed.) (1998), Flora Zambesiaca: Mozambique, Malawi, Zambia, Zimbabwe, Botswana. Volume 11, Part 2. Royal Botanic Gardens, Kew.

10. Flowers thin-textured, diaphanous . . . . . . . . . . . . . . . . . . 9 Flowers fleshy in texture . . . . . . . . . . . . . . . . . . . . . . . . . 15 11. Stem distinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 ●

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12. Inflorescences borne in lower leaf axils . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. sect. Gomphocentrum ●

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Fig. 717.2.  Distribution map of Angraecum.

Inflorescences in axils of upper leaves . . . . . . . . . . . . . . 13 13. Inflorescences as long as or longer than the leaves; rachis flexuous . . . . . . . . . . . . . . . . . . . . . . A. sect. Lepervenchea Inflorescences shorter than the leaves; rachis hardly flexuous . . . . . . . . . . . . . . . . . . . . . . . . . . . A. sect. Lemurangis 14. Raceme densely flowered, normally one-sided . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. sect. Nana Raceme laxly flowered, all round the rachis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. sect. Boryangraecum 15. Stem not obvious, usually absent . . . . . . . . . . . . . . . . . . 16 Stem prominent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 16. Spur slenderly cylindrical with a narrow mouth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. sect. Chlorangraecum Spur broadening at mouth to resemble a cornet . . . . . . 17 17. Spur conical or saccate . . . . . . . . . . . . A. sect. Hadrangis Spur filiform . . . . . . . . . . . . A. sect. Humblotiangraecum 18. Leaves iridiform or dorsiventrally flattened; labellum usually broadest in apical half; spur erect at base then arching and pendent in upper part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. sect. Afrangraecum Leaves dorsiventrally flattened; labellum usually broadest in basal half; spur pendent only . . . . . A. sect. Angraeum ●

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A. sect. Acaulia Garay, Kew Bull., 28, 498 (1973). Type species: Angraecum rhynchoglossum Schltr. A. sect. Afrangraecum Summerh., Kew Bull., 13, 276 (1958). Type species: Angraecum reygaertii De Wild. A. sect. Angraecoides (Cordem.) Garay, Kew Bull., 28, 500 (1973). Type species: Angraecum pingue Frapp. Mystacidium sect. Angraecoides Cordem., Rev. Gén. Bot., 11, 413 (1899) 360

Epidorkis sect. Angraecoides (Cordem.) Kuntze in Post & Kuntze, Lex. Gen. Phan., 199 (1903) Angraecum sect. Baroniangraecum Schltr., Repert. Sp. Nov. Regni Veg., Beih. 33, 309 (1925). Type species: Angraecum rostratum Ridl. (selected by Garay 1973) A. sect. Angraecum Angraecum sect. Euangraecum Benth. in Benth. & Hook. f., Gen. Pl., 3, 585 (1883). Type species: Angraecum eburneum Bory Angraecum sect. Aerobion (Kaempf. ex Spreng.) Pfitzer in Engl. & Prantl., Natürl. Pflanzenfam., 2 (6), 214 (1889). Type species: Angraecum superbum (Thouars) Spreng. Angorchis sect. Euangorchis Kuntze in Post & Kuntze, Lex. Gen. Phan., 30 (1903) Angorchis sect. Aerobion (Kaempf. ex Spreng.) Kuntze in Post & Kuntze, Lex. Gen. Phan., 31 (1903) Angraecum sect. Macroura Benth., J. Linn. Soc., Bot., 18, 337 (1881). Type species: Angraecum sesquipedale Thouars (selected by Garay 1973) Angorchis sect. Aerobion (Kaempf. ex Spreng.) Kuntze in Post & Kuntze, Lex. Gen. Phan., 31 (1903). Type species: Angraecum sesquipedale Thouars Angraecum sect. Subulatae Kränzl. in Rchb.f., Xenia Orch., 3, 74 (1890). Type species: Angraecum scottianum Rchb.f. (selected by Garay 1973) A. sect. Arachnangraecum Schltr., Repert. Sp. Nov. Regni Veg., Beih., 33, 309 (1925). Type species: Angraecum ramosum Thouars (selected by Garay 1973) A. sect. Boryangraecum Schltr., Beih. Bot. Centralbl., 33, 308 (1925). Type species: Angraecum pumilio Schltr. (selected by Garay 1973) A. sect. Chlorangraecum Schltr., Repert. Sp. Nov. Regni Veg., Beih., 33, 310 (1925). Type species: Angraecum chloranthum Schltr. (selected by Garay 1973) A. sect. Conchoglossum Schltr., Beih. Bot. Centralbl., 36, 2, 157 (1918). Type species: Angraecum viride Kränzl. (selected by Garay 1973) A. sect. Dolabrifolia (Pfitzer ex Rchb.f.) Garay, Kew Bull., 28, 498 (1973). Type species: Angraecum distichum Lindl. Aeranthus sect. Dolabrifolia Rchb.f., Walp., Ann. Bot. Syst., 6, 901 (1864), nom. nud. Mystacidium sect. Dolabrifolia Pfitzer ex Rchb.f. in Engl. & Prantl., Natürl. Pflanzenfam., 2 (6), 216 (1889) Epidorkis sect. Dolabraria Kuntze in Post & Kuntze, Lex. Gen. Phan., 200 (1903), nom. illeg. A. sect. Filangis Garay, Kew Bull., 28, 500 (1973). Type species: Angraecum filicornu Thouars A. sect. Gomphocentrum (Benth.) Garay, Kew Bull., 28, 501 (1973). Type species: Angraecum caulescens Thouars Mystacidium sect. Gomphocentrum Benth., J. Linn. Soc., Bot., 18, 337 (1881) Epidorkis sect. Gomphocentrum (Benth.) Kuntze in Post & Kuntze, Lex. Gen. Phan., 200 (1903) Angraecum sect. Hildebrandtiangraecum Schltr., Repert. Sp. Nov. Regni Veg., Beih., 33, 308 (1925). Type species: Angraecum calceolus Thouars (selected by Garay 1973) A. sect. Hadrangis Schltr., Beih. Bot. Centralbl., 36, 158 (1925). Type species: Angraecum striatum Thouars

ANGRAECUM

Angraecum sect. Thouarsiangraecum Schltr., Repert. Sp. Nov. Regni Veg., Beih., 33, 310 (1925), nom. illeg. Type species: Angraecum striatum Thouars A. sect. Humblotiangraecum Schltr., Repert. Sp. Nov. Regni Veg., Beih., 33, 310 (1925). Type species: Angraecum leonis Rchb.f. A. sect. Lemurangis Garay, Kew Bull., 28, 501 (1973). Type species: Angraecum madagascariense (Finet) Schltr. A. sect. Lepervenchea (Cordem.) Schltr., Beih. Bot. Centralbl., 36, 157 (1918). Type species: Angraecum tenuifolium Frapp. ex Cordem. A. sect. Nana (Cordem.) Garay, Kew Bull., 28, 501 (1973). Type species: Angraecum nanum Frapp. (selected by Garay 1973) Mystacidium sect. Nana Cordem., Rev. Gén. Bot., 11, 415 (1899) Angraecum sect. Micrangraecum Schltr., Repert. Sp. Nov. Regni Veg., Beih., 33, 308 (1925). Type species: Angraecum pusillum Lindl. A. sect. Pectinaria (Benth.) Schltr., Beih. Bot. Centralbl., 36, 157 (1918). Type species: Angraecum pectinatum Thouars Mystacidium sect. Pectinaria Benth. in Benth. & Hook. f., Gen. Pl., 3, 585 (1883) Epidorkis sect. Pectinaria (Benth.) Kuntze in Post & Kuntze, Lex. Gen. Phan., 200 (1903), nom. illeg. A. sect. Perrierangraecum Schltr., Repert. Sp. Nov. Regni Veg., Beih., 33, 309 (1925). Type species: Angraecum triquetrum Thouars A. sect. Pseudojumellea Schltr., Beih. Bot. Centralbl., 36, 157 (1918). Type species: Angraecum mauritianum (Poir.) Frapp. (selected by Garay 1973)

Anatomy Carlsward et al. (2006a) examined 19 species of Angraecum. Their results are summarized below.

Leaf TRANSVERSE SECTION

Angraecum distichum Lindl., A. subulatum Lindl., and A. teres Summerh. possess semi-terete leaves with an adaxial groove. In A. teres the adaxial groove is reduced to a sulcus of six or seven cells, so the leaf superficially appears terete. Trichomes multicellular, glandular; absent in A. birrimense Rolfe and A. eichlerianum Kränzl. Stomata abaxial, slightly sunken in A. moandense De Wild. Hypodermis ad- and abaxial, forming a single row of cells and composed of fibrous idioblasts in groups of one to several cells interrupted by chlorenchyma in A. birrimense, A. cultriforme Summerh., A. eichlerianum, A. erectum Summerh., A. moandense, and A. multinominatum Rendle; ∩-thickened fibres forming an uninterrupted row in A. sesquipedale Thouars. Hypodermis only adaxial and composed of one to several rows of smooth to pleated water-storage cells in A. appendiculatum Frapp. ex Cordem. (syn. Bonniera appendiculata (Frapp. ex Cordem.) Cordem.), A. conchiferum Lindl., A. eburneum ssp. superbum (Thouars) H.Perrier, A. gabonense Summerh., A. germinyanum Hook.f., and A. sacciferum Lindl.; ad- and abaxial in A. calceolus Thouars, A. eburneum ssp. superbum, A. eburneum ssp. xerophilum H.Perrier, A. pungens Schltr., and A. subulatum. Fibre bundles

absent; present in a single abaxial peripheral row and scattered among vascular bundles in A. teres; alternating with vascular bundles in a single abaxial row in A. subulatum; solitary bundles present only at leaf margins in A. distichum and A. sesquipedale. Bundles composed only of thick-walled sclerenchyma in A. distichum, A. sesquipedale, and A. teres; composed of abaxially thick-walled cells and adaxially thinner-walled cells in A. subulatum. Mesophyll most commonly homogeneous; heterogeneous with columnar adaxial cells and isodiametric abaxial cells in A. appendiculata (syn. Bonniera appendiculata), A. conchiferum, A. distichum, A. dives Rolfe, A. eburneum ssp. superbum, A. eburneum ssp. xerophilum, A. gabonense, A. germinyanum, A. pungens, A. sesquipedale, A. subulatum, and A. teres; heterogeneous with empty adaxial cells and smaller, assimilatory abaxial cells in A. calceolus and A. erectum. Water-storage cells with smooth to pitted walls in most species; banded in A. birrimense, A. cultriforme, A. dives, A. eichlerianum, and A. erectum; thickened along the edges in A. erectum. Scattered fibrous idioblasts throughout mesophyll in A. gabonense and A. subulatum. Vascular bundles generally in one row; two rows in A. teres (one central ring of large bundles surrounded by a ring of smaller bundles interspersed with fibre bundles). Vascular bundle sheath distinct. Stegmata of vascular bundles associated with vascular bundle sclerenchyma and fibre bundles (encircling fibre bundles in A. distichum, A. sesquipedale, and A. teres; found only on the abaxial surface of fibre bundles in A. subulatum).

Root TRANSVERSE SECTION

Velamen 2–4 cells thick. Epivelamen cells isodiametric to radially elongate; walls ∪-thickened with fine anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate; outer layers usually thicker-walled than those of inner layers. Distinct endovelamen thickenings generally absent; when present appear ridged in A. distichum or smooth in A. eburneum, A. eburneum ssp. giryamae (Rendle) Senghas & P.J.Cribb, A. eburneum ssp. superbum, A. eburneum ssp. xerophilum, and A. sesquipedale. Cover cells usually present over short cells of exodermis. Exodermal long cells ∩- to ○-thickened. Water-storage cell walls smooth to pitted or banded. Single modified cortical layer of thin- to thick-walled cells surrounding the endodermis in A. distichum and A. subulatum. Aeration units present in A. calceolus, A. conchiferum, A. cultriforme, A. eburneum, A. eburneum ssp. superbum, A. eburneum ssp. xerophilum, A. eichlerianum, A. erectum, A. germinyanum, A. pungens, A. sacciferum, and A. teres. Endodermal cells primarily ○-thickened to infrequently ∪-thickened in A. eburneum, A. eburneum ssp. superbum, and A. multinominatum. Vascular cylinder 7- to 28-arch. Vascular tissue embedded in sclerenchyma. Pith sclerenchymatous with scattered thin-walled parenchyma cells in A. germinyanum. (BC)

Palynology Schill and Pfeiffer (1977) studied the pollen of 11 species of Angraecum and described the tetrads as concave-convex with laevigate sculpturing and a calymmate sexine. (AP) 361

VA N D E A E

Cytogenetics Brandham (1999) summarized chromosome numbers reported in the literature for Angraecum as 2n = 38, 40, 42, 42–48, 46, 50, 63, c. 76, 92. He suggested a basic number of x = 19, which is common in Angraecinae but argued that there must be dysploid variation to explain the numbers that do not fit that series. Many other angraecoid genera are based on x = 24, and if this were the case ancestrally for Angraecum, then there would be dysploid reductions as well as increases. (AP, MC)

containing a higher content than those from the Comoro Islands (Kaiser 2011). Linalool is also the major constituent of the scents of A. magdalenae (64.5%) and A. rutenbergianum (62.0%). In addition, these species contain relatively large amounts of germacrene D (21.1%) and (E)-nerolidol (28.5%), respectively. Two compounds of special interest that have a limited distribution among natural scents are 3-(4-methoxyphenyl)-propan-1-ol and its acetate, which were detected as minor components of the fragrance of A. rutenbergianum (Kaiser 2011). (NV, RG)

Phytochemistry

Phylogenetics

Alkaloid levels ≥ 0.1% have been recorded in A. falcatum (Thunb.) Lindl. (Lüning 1964, 1967). According to a later assessment, 3 of 12 species of Angraecum tested had an alkaloid content ≥ 0.1%, although no species names were given (Lüning 1974). Flavone C-glycosides occur in leaves of A. eburneum and possibly in A. eichlerianum (C. Williams 1979). Chemical composition data are available for the floral fragrances of A. aporoides Summerh., A. eburneum (2 subspecies), A. eichlerianum, A. leonis (Rchb.f.) André, A. magdalenae Schltr. & H.Perrier, A. rutenbergianum Kränzl., A. sesquipedale, and A. sesquipedale var. angustifolium Bosser & Morat (cited as A. bosseri Senghas), which are night-scented moth-pollinated species (Kaiser 1993, 2011). Measurement of the time-dependence of fragrance composition in A. sesquipedale indicates that changes occur as the flower matures; for example, indole is most prominent as an olfactory component during the first two nights after anthesis. According to Kaiser (1993), the fragrance has two attributes, a ‘basic scent’ to which benzyl alcohol (14.8%), cinnamic alcohol (< 1%), phenylethyl alcohol (2.5%), derivatives of these alcohols, and indole (< 1%) contribute, and ‘scent modifiers’, which are oximes, notably 3-methylbutyraldoxime (34.0%) and phenylacetaldoxime (2.0%). The latter are thought to be metabolites of the amino acids leucine and phenylalanine, respectively. Oximes are also present in the floral fragrance of A. sesquipedale var. angustifolium, albeit at much lower levels (< 1%). The particular character of this scent derives from a mixture of eugenol, isoeugenol, chavicol, and isochavicol, although linalool (75.0%) and phenylethyl alcohol (6.8%) are the major components. Floral fragrances of A. eburneum subspecies can be distinguished by their oxime content. In the case of A. eburneum subsp. eburneum, 3-methylbutyraldoxime (9.9%) predominates, whereas for A. eburneum subsp. superbum, large amounts of 2-methylbutyraldoxime (26.8%) are present. Important contributions to the scent of A. eburneum subsp. eburneum are also made by benzaldehyde (12.0%), benzyl acetate (34.6%), indole (< 1.0%), methyl benzoate (6.4%), phenylacetaldehyde (3.0%), and phenylethyl alcohol (7.1%). In contrast, the fragrance of A. eburneum subsp. superbum contains only one other major component, (Z)-3-hexenol (53.0%). The character of the scent of A. aporoides is attributable to anisaldehyde (1.5%) and benzaldehyde (31.0%), whereas that of A. eichlerianum derives from a combination of indole (1.9%), linalool (13.0%), methyl benzoate (26.7%), and methyl salicylate (4.8%) (Kaiser 1993). The major scent constituents of A. leonis are linalool (49.2%), chavicol (16.1%), and (E)-nerolidol (15.0%), although the proportion of chavicol varies according to geographical origin, with plants from Madagascar

An analysis of over 50 species of Angraecum in 16 sections by Micheneau et al. (2008) demonstrated that Angraecum in its previously accepted sense is polyphyletic. A large clade, including the type species A. eburneum and more than 40 other species, also includes the genus Bonniera, of which two species were included in their analysis. Bonniera species have a similar habit and inflorescence to Angraecum ramosum and its allies (A. sect. Arachnagraecum sensu Garay in part), but the flowers have a petaloid labellum and lack a spur. It seems likely that Bonniera flowers are semi-peloric with a petaloid labellum. In this main clade, a few sections are identifiable as subclades in their analysis, notably A. sects. Filangis (1 species examined), Hadrangis (3 species), Humblotiangraecum (3 species), Lemurangis (1 species), Lepervenchea (3 species), and Pseudojumellea (1 species). The remaining sections are polyphyletic, although A. section Gomphocentrum is monophyletic if A. chloranthum (A. sect. Chlorangraecum) is included, and A. sect. Perrierangraecum is monophyletic if A. cornigerum (A. sect. Filangis) is included. Most sections appear to be unnatural. Nine African Angraecum species included in the analysis fall outside the main Angraecum clade. Five species, namely A. aff. eichlerianum, A. birrimense, A. infundibulare Lindl., A. aff. chevalieri (=A. moandense), and A. cultriforme (the last two from A. sect. Angraecoides), belong to a clade that includes both tropical American genera, Campylocentrum and Dendrophylax. The first three form a grade relative to the American genera. Four species—A. aporoides and A. distichum (both in A. sect. Dolabrifolia), and A. subulatum and A. aff. doratophyllum Summerh. (both in A. sect. Pectinaria)—are sister to a clade that includes Aerangis, Angraecopsis parviflora (Thouars) Schltr., Mystacidium flanaganii (Bolus) Bolus, and Solenanagis aphylla (Thouars) Summerh. This clade in turn is sister to that including the tropical American genera. Unfortunately, A. pectinatum Thouars and A. hermannii (Cordem.) Schltr., both in A. sect. Pectinaria, and A. obversifolium Frapp. ex Cordem., A. pingue Frapp. ex Cordem., and A. sedifolium Schltr. in A. sect. Angraecoides, all nest in the main Angraecum clade (Micheneau et al. 2008). It seems likely, therefore, that A. sect. Dolabrifolia and parts of A. sects. Angraecoides, Arachnangraecum, and Pectinaria might be separated from Angraecum sensu stricto when further information becomes available. (PC)

362

Ecology Angraecum species occupy a wide range of habitats but are most frequent in evergreen humid forests and scrub from

BECLARDIA

the lowlands into the mountains. Angraecum eburneum and A. sesquipedale Thouars are among the species that are frequent in coastal forests. A number of species, including A. coutrixii Bosser, A. longicalcar (Bosser) Senghas, A. magdalenae Schltr. & H.Perr., A. protensum Schltr., A. sororium Schltr., and A. madagascariense Schltr. grow lithophytically on rocky outcrops in the highlands of Madagascar. Some species, including A. aloifolium Hermans & P.J.Cribb, A. decaryanum H.Perr., A. dollii Senghas, A. eburneum var. xerophilum H.Perr., A. microcharis Schltr., A. myrianthum Schltr., A. popowii Braem, and A. sesquipedale var. angustifolium Bosser & P.Morat, are found in dry, deciduous forests in western and southern Madagascar. Some species can be found on particular trees or scrubs: Angraecum leonis (Rchb.f.) André can occasionally be found on species of Adansonia L. (Malvaceae); A. palmicolum Bosser grows on Dypsis decipiens (Becc.) Beentje & J.Dransf. (Arecaceae); and A. rubellum Bosser grows on species of Pandanus Parkinson (Pandanaceae). (PC)

Pollination Darwin’s (1862) deductive account of the probable pollination of A. sesquipedale is now legendary. Flowers sent to him by James Bateman had nectaries ‘eleven and half inches long, with only the lower inch and a half filled with nectar’. He argued against his experience with smaller English sphingid moths that there must be moths in Madagascar with proboscises between 10 and 11 inches (25–28 cm) long, long enough to reach the nectar at the bottom of the spur and be rewarded for its efforts. That such large moths existed, at least in Brazil, was proven by Fritz Müller, who published a letter with a drawing by his brother Hermann in Nature in 1873 on page 223 (Darwin 1862). Not until 1903, however, was such a moth found in Madagascar and named Xanthopan morgani praedicta. Xanthopan morgani had been known from East Africa for a much longer period. Pollination of A. sesquipedale has not been observed or documented unequivocally in the field. Phil DeVries of the University of New Orleans filmed a visitation by the moth, which inserted its unfurled proboscis into the spur but apparently did not remove or deposit pollinia. Wasserthal (1997) photographed the removal and deposition of pollinia by Xanthopan morgani but under controlled conditions in a large flight tent with introduced plants and captured moths. He did find, however, pollinia of A. sororium attached to the tongues of the hawkmoths Coelonia solani and Panogena lingens; he photographed pollinia transfer by C. solani with subsequent fruit set under the same controlled conditions. Panogena lingens also removed pollinia from A. compactum Schltr. in the flight tent. Nilsson et al. (1985), however, did document field-pollination of A. arachnites Schltr. by Panogena lingens in central Madagascar. Pollinaria were attached to the ventral side of the proboscis. Out of 66 individuals captured, 65% carried orchid pollinia, and, among those, 65% carried pollinia of A. arachnites. In those 28 moths, 50% carried pollen loads of other orchid species as well. The vast majority (97) of captured individuals were males. There were 105 viscidia in toto on the captured moths, but only 36

pollinia remained, indicating that two-thirds of the pollinia had already been deposited. Of the 23 visited flowers, about 78% were pollinated, and pollinia were removed from 87% of them. Fruit set was at least 41%, perhaps higher later. Nilsson et al. (1985, 1987) also recorded visitation to flowers of A. compactum by male individuals of Panogena lingens. Viscidia were attached dorsally on the proboscis. Once again, the moths carried mixed pollen loads. In both A. arachnites and A. compactum, the moths followed a zig-zag path from downwind, indicating long-distance attraction by floral fragrances followed by a straighter and more rapid approach using visual orientation. (AP)

Uses Various species of Angraecum have been used in Africa to treat inflammation, haemoptysis, and sores (Lawler 1984). More importantly, many species such as A. sesquipedale, A. eichlerianum, A. leonis, A. germinyanum, A. eburneum, and A. magdalenae have been hybridized and propagated artificially for hobbyist collections (see cultivation notes in the subtribal treatment). (AP)

Taxonomic literature Garay, L. A. (1973). Systematics of the genus Angraecum (Orchidaceae). Kew Bulletin, 28, 495–516. Micheneau, C., Carlsward, B. S., Fay, M. F., Bytebier, B., Pailler, T., and Chase, M. W. (2008). Phylogenetics and biogeography of Mascarene angraecoid orchids. Molecular Phylogenetics and Evolution, 46, 908–22. Schlechter, R. (1918). Versuch einer natürlichen Neuordnung der afrikanischen angraekoiden Orchidaceen. Beihefte zum Botanischen Centralblatt, 36, 62-181.

7 1 8 .   B EC L A R DI A Beclardia A.Rich., Mem. Soc. Nat. Hist. Paris, 4, 69 (1828). Type species: Beclardia macrostachya (Thouars) A.Rich. (basionym: Epidendrum macrostachys Thouars) “Brachystepis”, alternative epithet used by Thouars in Orch. Iles. Austral. Afr. (1822), occasionally mistaken for a generic name.

Derivation of name Beclardia was named for Augustine Beclard, Professor of Anatomy at the Faculty of Medicine in Paris and plant collector of the early 19th century. (PC)

Description (Plate 146; Fig. 718.1) Erect or spreading, epiphytic herbs. Stem covered by sheathing leaf bases, flattened in cross-section. Leaves distichous, conduplicate, coriaceous, ligulate, unequally bilobed at the tip. Inflorescence lateral, axillary, elongate; bracts ovate-triangular, much shorter than ovary. Flowers not opening widely, white with a green mark on labellum disc. Sepals oblong, recurved. Petals spreading, obovatespatulate. Labellum trilobed but appearing with the apical lobe 363

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I

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Fig. 718.1.  Beclardia macrostachya (Thouars) A.Rich. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum; H. Labellum, longitudinal section; I. Column and labellum, side view; J. Column, front view; K. Column, side view; L. Column, anther cap removed, side view; M. Anther cap, dorsal view; N. Anther cap, ventral view; O. Pollinarium; P. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Malcomver et al. 1402 (1992) and Kew Spirit Collection no. 30374.

itself bilobed, ecallose but hairy on disc, spurred at the base, side lobes erect around column, midlobe emarginate at tip; spur funnel-shaped at base, cylindrical-rounded at tip. Column with the anther rostrate, pollinia two, deeply cleft, attached by a spatulate stipe (tegula) to a semi-elliptic viscidium; stigma deeply concave, transversely oblong; rostellum bifid at the acute tip after removal of pollinarium. (PC)

Distribution (Fig. 718.2) Beclardia is a genus of only two species, B. grandiflora Bosser and B. macrostachya, endemic to Madagascar and the Mascarenes. (PC) 364

Anatomy Carlsward et al. (2006a) examined Beclardia macrostachya. Their results are summarized below.

Leaf TRANSVERSE SECTION

Trichomes multicellular, glandular. Stomata primarily abaxial, rarely adaxial. Hypodermis absent. Fibre bundles absent. Mesophyll homogeneous; adaxial cells generally larger with fewer chloroplasts than cells closest to the abaxial surface. Waterstorage cells infrequent, smooth-walled, usually found within adaxial mesophyll. Vascular bundle sheath distinct. Stegmata

BOLUSIELLA

clade that includes Lemurella, Neobathiea, Cryptopus, and Oeonia (Micheneau et al. 2008). (PC)

Ecology Beclardia grandiflora and B. macrostachya are epiphytic in humid and semi-deciduous montane and plateau forests at 900–2000 m. (PC)

Pollination Pollination of these two species has not been investigated. (AP)

Uses There are no known uses of either species, and they are rare in cultivation. (AP)

Taxonomic literature Fig. 718.2.  Distribution map of Beclardia.

contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Velamen two or three cells thick. Epivelamen cells mainly isodiametric; ∪-thickened with fine anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate. Distinct endovelamen wall thickenings absent. Cover cells present over short cells of the exodermis. Exodermal long cells ∪-thickened. Water-storage cells large, radially elongate, and thickened along the edges. Aeration units present. Endodermal cells strongly ○-thickened. Vascular cylinder hexarch. Vascular tissue embedded in sclerenchyma, cell walls of embedding sclerenchyma thickest around phloem clusters. Pith sclerenchymatous. (BC)

Palynology Pollen of Beclardia has not been studied. (AP)

Cytogenetics Chromosome counts of neither species have been reported. (AP)

Phytochemistry No reports have been found on the phytochemistry of Beclardia. (NV, RG)

Phylogenetics Allied to Oeonia, also a Madagascan endemic genus, Beclardia differs in its habit and column structure. Beclardia is sister to a

Micheneau, C., Carlsward, B. S., Fay, M. F., Bytebier, B., Pailler, T., and Chase, M. W. (2008). Phylogenetics and biogeography of Mascarene angraecoid orchids. Molecular Phylogenetics and Evolution, 46, 908–22.

7 1 9 .   B OLU S I EL L A Bolusiella Schltr., Beih. Bot. Centr., 36, 106 (1918). Type species: Bolusiella maudiae (Bolus) Schltr. (basionym: Angraecum maudiae Bolus), designated by R. Rice, Preliminary checklist survey subtribe Aerangidinae (Orchidaceae), 23 (2005)

Derivation of name Bolusiella commemorates Harry Bolus (1834–1911), a wellknown botanist who specialized in the flora of South Africa. (PC)

Description (Plate 147; Fig. 719.1) Erect, epiphytic or occasionally lithophytic, psygmoid herbs. Roots tufted. Stem abbreviated, simple, leafy above. Leaves equitant, imbricate, unifacial, arranged in a plane forming a fan, narrowly elliptic or ensiform to linear-ligulate, obtuse to acute, rigid and articulated to a sheathing base, fleshy, sometimes sulcate-canaliculate and falcate-recurved. Inflorescence exceeding leaves, manyflowered, arising from axils of old leaf-sheaths; peduncle shorter than or equalling rachis, rarely longer; bracts imbricate and partially concealing flowers, membranous, grey to blackish brown or olive. Flowers resupinate, white. Sepals and petals free, oblong or narrowly elliptic, obtuse to acuminate, subequal, spreading. Labellum entire or obscurely trilobed, spurred, oblong or ovateelliptic, obtuse to acuminate, sometimes recurved; spur cylindrical, ellipsoid or conical, obtuse, usually shorter than labellum. Column oblong, constricted above, without a foot; anther quadrate, cucullate, pollinia two, oblong or ellipsoid, each attached by a linear stipe to an ovate viscidium; rostellum subulate, hamate. (PC) 365

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Fig. 719.1.  A–L. Bolusiella maudiae (Bolus) Schltr. A. Habit; B. Inflorescence; C. Flower, side view; D. Bract; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum; I. Column and labellum, side view; J. Column, front view; K. Anther cap; L. Pollinarium. Drawn by Judi Stone from Kew Spirit Collection no. 30289. M–Y. Bolusiella iridifolia (Rolfe) Schltr. subsp. picea P.J.Cribb. M. Habit; N. Leaf, transverse section; O. Inflorescence; P. Flower, side view; Q. Bract; R. Dorsal sepal; S. Lateral sepal; T. Petal; U. Labellum; V. Column and labellum, side view; W. Column, front view; X. Anther cap; Y. Pollinarium. Drawn by Judi Stone from Kew Spirit Collection no. 33800. Single bar = 1 mm, double bar = 1 cm. Reproduced with permission from G. V. Pope (ed.) (1998), Flora Zambesiaca: Mozambique, Malawi, Zambia, Zimbabwe, Botswana. Volume 11, Part 2. Royal Botanic Gardens, Kew.

Distribution (Fig. 719.2)

Anatomy

Bolusiella comprises five species distributed in tropical and southern Africa. (PC)

Carlsward et al. (2006a) examined three species of Bolusiella. Their results are summarized below.

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Palynology Schill and Pfeiffer (1977) examined pollen of B. batesii and found it similar to that of Angraecum with no further details. (AP)

Cytogenetics Brandham (1999) reported a chromosome count of 2n = 50 for the genus. (AP)

Phytochemistry Little is known about the phytochemistry of this genus other than a brief record that one species was tested for alkaloids, but did not contain them at levels ≥ 0.1% (Lüning 1974). (NV, RG)

Phylogenetics Carlsward et al. (2006b) included three species of Bolusiella in their analysis of angraecoid orchids based on ITS sequences. Bolusiella is sister to Microcoelia. (PC) Fig. 719.2.  Distribution map of Bolusiella.

Leaf TRANSVERSE SECTION

Bolusiella batesii (Rolfe) Schltr. and B. maudiae possess terete leaves with no discernible adaxial epidermis. Leaves of B. iridifolia (Rolfe) Schltr. are deeply sulcate with the adaxial epidermis restricted to the sulcus. Trichomes absent. Stomata ad- and abaxial in dorsiventral leaves of B. iridifolia. Substomatal chamber forming a distinct gap between hypodermal fibres in B. batesii (Rolfe) Schltr. Hypodermis composed of a single row of fibrous idioblasts in B. batesii and B. iridifolia; absent in B. maudiae. Water-storage idioblasts thick-walled and tightly packed in B. batesii, thin-walled and interspersed among chlorenchyma cells in B. iridifolia. Fibre bundles absent. Mesophyll homogeneous; central suture of flattened cells in B. batesii and B. maudiae. One to three thick-walled fibrous idioblasts present near the adaxial pole in B. maudiae. Vascular bundles arranged in a ∪-shaped arc. Vascular bundle sheath indistinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Velamen 2–4 cells thick. Epivelamen cells radially elongate to isodiametric; walls ∪-thickened with fine anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate; strongly thick-walled in B. batesii. Distinct endovelamen wall thickenings absent. Cover cells present over short cells of exodermis in B. iridifolia. Exodermal long cells primarily ○-thickened to infrequently ∩-thickened. Water-storage cell walls smooth to pitted. Aeration units present in B. iridifolia. Endodermal cells heavily ○-thickened in B. batesii and B. iridifolia. Vascular cylinder 7–12-arch. Vascular tissue embedded in sclerenchyma, thick-walled in B. batesii and B. iridifolia. Pith sclerenchymatous, thick-walled in B. batesii. (BC)

Ecology Species of the genus occur from sea level to 2500 m in humid lowland and evergreen forests, riverine forests, plantations (e.g. coffee), or on moss- and lichen-covered trees on rocky outcrops. (PC)

Pollination There no reports on pollination of any of the species. (AP)

Uses There are no ethnobotanical uses known for the species, and they are all rare in cultivation. (AP)

Taxonomic notes Verlynde et al. (2013) have recently revised the genus.

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Verlynde, S., Dubuisson, J.-Y., Stévart, T., Simo-Droissart, M., Geerinck, D., Sonké, B., Cawoy, V., Descourvières, P., and Droissart, V. (2013). Taxonomic revision of the genus Bolusiella (Orchidaceae: Angraecinae) with a new species from Cameroun, Burundi and Rwanda. Phytotaxa, 114, 1–22.

7 2 0 .   C A LYP T ROC H I LU M Calyptrochilum Kränzl., Bot. Jahrb. Syst., 22, 30 (1895). Type species: Calyptrochilum preussii Kränzl. (=Calyptrochilum emarginatum (Afzel ex Sw.) Schltr.; basionym: Limodorum emarginatum Afzel ex Sw.) 367

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Fig. 720.1.  Calyptrochilum christyanum (Rchb.f.) Summerh. A. Habit; B. Bract; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum, from above; H. Column and labellum, side view; I. Column, side view; J. Column, front view; K. Anther cap; L. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Morris 76A (K) and Kew Spirit Collection no. 28451. Reproduced with permission from G. V. Pope (ed.) (1998), Flora Zambesiaca: Mozambique, Malawi, Zambia, Zimbabwe, Botswana. Volume 11, Part 2. Royal Botanic Gardens, Kew.

Derivation of name

Description (Plate 148; Fig. 720.1)

From the Greek calyptra, veil, and chilos, labellum, in reference to the labellum that encloses the column and anther. (PC)

Epiphytic or lithophytic herbs. Stem cylindrical, spreading to pendent, covered by sheathing leaf bases. Leaves coriaceous, alternate,

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cells angular, isodiametric to radially elongate; outer layers possess thicker-walled cells than those of inner layers. Distinct endovelamen thickenings absent. Cover cells present over short cells of exodermis. Exodermal long cells ○-thickened or ∪-thickened. Water-storage cells banded, often slightly lignified. Single modified cortical layer of thin- to thick-walled cells surrounding the endodermis in C. christyanum and C. emarginatum where lignification is most pronounced. Aeration units present in C. christyanum and C. emarginatum. Endodermal cells ○-­thickened. Vascular cylinder 13–35-arch. Vascular tissue embedded in sclerenchyma; sclerenchyma forming caps of thick-walled cells over phloem clusters in C. emarginatum. Pith sclerenchymatous. (BC)

Palynology Pollen of these two species has not been studied. (AP)

Cytogenetics Fig. 720.2.  Distribution map of Calyptrochilum.

distichous, twisted to lie in one plane, unequally bilobed at tip, articulated to sheathing base. Inflorescence axillary, few-flowered, shorter than leaves; bracts distichous, cucullate. Flowers white, sometimes with a green or yellow mark on labellum. Sepals and petals subsimilar, free, spreading. Labellum porrect, trilobed, side lobes erect around column, spurred at base; spur geniculate in middle, clavate at tip. Column with two pollinia, operculate, attached to a single linear stipe and that to an ovate viscidium. (PC)

Distribution (Fig. 720.2) Calyptrochilum is a genus of only two species, C. christyanum (Rchb.f.) Summerh. and C. emarginatum, distributed throughout tropical Africa. (PC)

Anatomy Carlsward et al. (2006a) examined both species of Calyptrochilum. Their results are summarized below.

Leaf TRANSVERSE SECTION

Trichomes multicellular, glandular; absent in C. emarginatum. Stomata abaxial only. Hypodermis absent. Fibre bundles absent. Mesophyll homogeneous in C. emarginatum; heterogeneous with columnar adaxial cells and isodiametric abaxial cells in C. christyanum. Water-storage cells slightly lignified and banded. Vascular bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Velamen 3–7 cells thick. Epivelamen cells radially elongate; walls ∪-thickened with fine, anastomosing, radial bands. Endovelamen

Brandham (1999) reported a chromosome count of 2n = 38 for Calyptrochilum. Arends and Van der Laan (1983) noted that its basic number of x = 19 was different from the basic number of x = 23, 24, 25 of other Aerangidinae, where it was formerly placed, and more properly belonged in Angraecinae with the basic number of x = 19. Now that Aerangidinae and Angraecinae have been merged on molecular evidence, their argument is well taken but moot. (AP)

Phytochemistry No reports have been found on the phytochemistry of Calyptrochilum. (NV, RG)

Phylogenetics In the study of Carlsward et al. (2006b), Calyptrochilum was sister to all other angraecoid orchids. (PC)

Ecology Both species are found in humid evergreen forest and humid woodland from sea level to 1200 m. Calyptrochilum christyanum also grows on trees in savannas and on rocks. (PC)

Pollination Nothing is known about pollination in Calyptrochilum. (AP)

Uses There are no uses known for the genus, and it is rare in cultivation. (AP)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86.

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721.  C AMP Y LO C E N TRUM Campylocentrum Benth., J. Linn. Soc., Bot., 18, 337 (1881). Type species: Campylocentrum schiedei (Rchb.f.) Benth. ex Hemsley (basionym: Angraecum schiedei Rchb.f.; originally published as Todaroa micrantha A.Rich & Galeotti) Todaroa A.Rich. & Galeotti, Ann. Sci. Nat. Bot., ser. 3, 3, 28 (1845). Type species: Todaroa micrantha A.Rich. & Galeotti

Derivation of name From the Greek kampylox, crooked, and kentron, spur. (BC)

Description (Plate 149; Fig. 721.1) Epiphytic herbs. Roots photosynthetic in all species, forming the main plant body in most leafless species. Stem erect to pendent, sprawling in leafy species, erect and less than 1 cm in most leafless species. Leaves articulate, sheathing, conduplicate, emarginate and well developed in most species; subterete and caducous in C. poeppigii (Rchb.f.) Rolfe; reduced to nonphotosynthetic scales in most leafless species. Inflorescence axillary, unbranched, distichous, usually congested with many open flowers, often not extending leaves. Flowers usually resupinate, white, occasionally tinged with orange. Sepals and petals connivent, free, lanceolate, equal in size. Labellum entire or lobed with a saccate nectar spur that is straight to incurved. Column without a foot; pollinia two, with one or two stipes; rostellum deeply divided. Ovary narrowly cylindrical and ridged. (BC)

Distribution (Fig. 721.2) There are currently 65 Neotropical species of Campylocentrum, ranging from Florida and the Caribbean throughout most of Central and South America as far south as Argentina. More than half of the species diversity is in Brazil. (BC)

Anatomy Carlsward et al. (2006a) studied four leafless species of Campylocentrum and one leafy species. Their observations are summarized below.

Leaf Transverse section

Hairs absent. Stomata abaxial. Hypodermis ad- and abaxial, composed of one row of fibrous idioblasts scattered among mesophyll chlorenchyma. Fibre bundles absent. Mesophyll heterogeneous with several rows of ad- and abaxial columnar cells surrounding central isodiametric cells. Water-storage cells with birefringent walls absent. Bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with phloem and xylem sclerenchyma.

Root

wall thickenings; endovelamen cells angular, isodiametric to radially elongate. Cover cells present over exodermal passage cells. Distinct endovelamen thickenings absent. Exodermal cells ∩-thickened to rarely ∩- or ○-thickened. Water-storage cells with birefringent walls that vary from smooth to pitted or banded. Aeration units present. Endodermal cells ○-thickened. Vascular cylinder 6–9-arch. Vascular tissue embedded in sclerenchyma. Pith sclerenchymatous. (BC)

Palynology Schill and Pfeiffer (1977) examined an unknown species of Campylocentrum and found smooth, convex tetrads of pollen grains with smooth walls. (BC)

Cytogenetics Most species investigated have a chromosome count of 2n = 38 (Daviña et al. 2009; Felix and Guerra 2010), although Brandham (1999) listed 2n = 44 from other reports. (BC)

Phytochemistry Surveys of the alkaloid content of Orchidaceae indicate that low levels (< 0.01%) are found in C. brenesii Schltr., C. micranthum (Lindl.) Rolfe, and C. ornithorrhynchum (Lindl.) Rolfe (Lüning 1964, 1967). (NV, RG)

Phylogenetics Achille Richard and Henri Guillaume Galeotti first described Campylocentrum in 1845 as Todaroa, but that name had already been used for another genus in Apiaceae. In 1881, Bentham renamed the species of Todaroa. Todaroa micrantha is now known as C. schiedei and not C. micranthum (Lindl.) Maury, which was originally described as Angraecum micranthum Lindl. in 1835 by Lindley (Ackerman 2004). As with many other orchid genera, the original taxonomy of Campylocentrum was historically based on floral morphology, especially for leafless species with reduced stems. The taxonomic history of the genus was traced by Bogarin and Pupulin (2009). Most species were originally placed in either Aeranthus or Angraecum by early workers and later separated from the Palaeotropical taxa. Two Neotropical genera, Dendrophylax and Campylocentrum, were later determined to be sister taxa (Carlsward et al. 2003). Based on phylogenetic analyses of molecular data (ITS, matK, and trnL-F regions), Carlsward et al. (2003) transferred the small-flowered Harrisella porrecta, Campylocentrum macrocarpum Dod, C. helorrhizum (Dod) Nir, C. ariza-juliae Ames, and C. filiforme (Sw.) Cogn. ex Kuntze into Dendrophylax. In general, species of Campylocentrum have smaller flowers that are more numerous than those of the closely related Dendrophylax and occur in congested distichous inflorescences. (BC)

Transverse section

Ecology

Velamen of 1–3 layers; epivelamen cells isodiametric to tangentially flattened, ∪-thickened with fine anastomosing radial

Species of Campylocentrum occur from sea level to about 2940 m. Many are found in shaded sites with high humidity but may also

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Fig. 721.1.  Campylocentrum huebneri Mansf. A. Habit; B. Flowers, side view; C. Flower, dissected; D. Labellum and base of spur; E. Column with pollinaria; F. Anther, pollinaria removed; G. Pollinaria. Drawn by G. C. K. Dunsterville. Originally published in Venezuelan Orchids Illustrated, 3, 50 (1965), Andre Deutsch, London. Reproduced with permission of the Oakes Ames Orchid Herbarium, Harvard University Herbaria.

be found in xeric habitats on twigs of shrubs or small trees or on limestone rocks. The most widespread species is C. micranthum, which occurs from Mexico to Bolivia across a wide range of elevations (Todzia 1980). (BC)

Pollination Pollination by various bees (halictid bees in C. aromaticum and stingless bees in C. burchellii) has been reported by Singer and Cocucci (1999). Campylocentrum micranthum has been reported to be autogamous (Damon and Salas-Roblero 2007). (BC)

Uses Plants of Campylocentrum are infrequently cultivated. (BC)

Cultivation The most commonly cultivated species is the leafy C. micranthum, which does well in a mossy substrate and high humidity. Leafless species are challenging to grow but do well under the same conditions as the related Dendrophylax species (in empty, plastic, hanging baskets with no growing medium, high humidity, and frequent watering). (BC)

Taxonomic literature Ackerman, J. D. (2004). Notes on the Caribbean orchid flora. V. New species, combinations and records. Lankesteriana, 4, 47–56. Bogarin, D. and Pupulin, F. (2009). The genus Campylocentrum (Orchidaceae: Angraecinae) in Costa Rica: some critical

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Description (Fig. 722.1) Epiphytic herbs. Roots branching, clustered in lower part of stem. Stem short. Leaves distichous, arcuate-spreading, linear, unequally roundly or obtusely bilobed at apex, conduplicate or twisted at base to lie in one plane, articulated at base to persistent, imbricate leaf bases. Inflorescence shorter than leaves, one- or rarely two-flowered; peduncle terete, elongate, bearing two sterile sheathing bracts along length. Flowers green to ochre-orange, non-resupinate, scabrid on outer surface. Sepals and petals subsimilar; lateral sepals fused at their oblique bases to enclose base of spur. Labellum obscurely trilobed, concave, ecallose, spurred at base. Column with two pollinia, subglobose, stipe solitary, linear, viscidium reniform; rostellum bifid, dilated and scutelliform at apex. (PC)

Distribution (Fig. 722.2) Cardiochilos is a monospecific genus known only from Malawi and adjacent Tanzania. (PC)

Anatomy Nothing is known of the anatomy of this species. (AP)

Palynology There are no reports on the pollen of this species. (AP)

Cytogenetics Chromosome numbers for the genus have not been reported. (AP) Fig. 721.2.  Distribution map of Campylocentrum.

questions and a few answers. In Proceedings of the Second Scientific Conference on Andean Orchids (ed. A. M. Pridgeon and J. P. Suárez), pp. 32–45. Universidad Técnica Particular de Loja, Loja, Ecuador. Carlsward, B. S., Whitten, W. M., and Williams, N. H. (2003). Molecular phylogenetics of Neotropical leafless Angraecinae (Orchidaceae): reevaluation of generic concepts. International Journal of Plant Sciences, 164, 43–51. Todzia, C. A. (1980). A revision of the Costa Rican species of Campylocentrum (Orchidaceae). Brenesia, 18, 117–136.

722.  C AR D IO C H ILO S Cardiochilos P.J.Cribb, in Kew Bull., 32, 183 (1977). Type species: Cardiochilos williamsonii P.J.Cribb.

Derivation of name From the Greek cardia, heart, and chilos, labellum, from its distinctive labellum shape. (PC) 372

Phytochemistry No reports have been found on the phytochemistry of Cardiochilos. (NV, RG)

Ecology Cardiochilos williamsonii occurs as an epiphyte in mist forests on Protea L. (Proteaceae) at about 2400 m. (PC)

Pollination Pollination of this species has not been investigated. (AP)

Uses There are no known uses for C. williamsonii. (AP)

Taxonomic notes Cardiochilos is similar in habit to Tridactyle species such as T. furcistipes Summerh. and T. tricuspis (Bolus) Schltr., but it differs in having a one- or rarely two-flowered inflorescence, a spatulate rostellum that is dilated at the tip, and a broad, heart-shaped, concave labellum with a straight spur emerging from its base. (PC)

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Fig. 722.1.  Cardiochilos williamsonii P.J.Cribb. A. Habit; B. Flower, front view; C. Flower, side view; D. Leaf, transverse section; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum; I. Column and labellum, side view; J. Column, front view; K. Column, side view; L. Column, back view; M. Anther cap; N. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Kew Spirit Collection no. 33735. Reproduced with permission from G. V. Pope (ed.) (1998), Flora Zambesiaca: Mozambique, Malawi, Zambia, Zimbabwe, Botswana. Volume 11, Part 2. Royal Botanic Gardens, Kew.

723.  CH AULIO D O N

Derivation of name

Chauliodon Summerh., Bot. Mus. Leafl., Harvard Univ., 11, 163 (1943). Type species: Chauliodon buntingii Summerh. (=Chauliodon deflexicalcaratum (De Wild.) L.Jonss.; basionym: Angraecum deflexicalcaratum De Wild.)

From the Greek chauliodous, with prominent teeth, in reference to the labellum callus with its prominent callus teeth. (PC)

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Phylogenetics Although not included in the DNA analysis of Carlsward et al. (2006b), they commented on its vegetative morphological similarity to Microcoelia and suggested that it might belong there. (PC)

Ecology Chauliodon deflexicalcaratum occurs up to 100 m on branches of trees growing in light shade on granitic outcrops or on the margins of dense, moist, mature forest of Autranella A.Chev. & Aubrév. (Sapotaceae) and Microberlinia A.Chev. (Fabaceae) on heavy clay, and also on Citrus L. (Rutaceae) in dense shade. (PC)

Pollination There are no reports concerning pollination in this species. (AP)

Uses Fig. 722.2.  Distribution map of Cardiochilos.

Description (Fig. 723.1) Leafless herbs. Roots elliptic or circular in cross-section. Stem up to 2.5 cm long, 0.3 mm in diameter. Inflorescence up to 25 cm long, unbranched, few-flowered. Flowers resupinate, brownish rose. Sepals reflexed, oblong. Petals tapering to tip. Labellum transversely elliptic, strongly cochleiform, with a tooth-like callus in front of spur mouth; spur elongate, geniculate in middle, the apical part deflexed. Column upcurved and broadest across apex, hammer-like; anther incumbent; pollinia two, porate, attached by a simple oblanceolate or spatulate tegula, viscidium horseshoeshaped; rostellum triangular, acute or subacute. (PC)

Distribution (Fig. 723.2) Chauliodon is a monospecific genus in West Africa ranging from Liberia to Nigeria. (PC)

Anatomy The vegetative anatomy of this species has not been studied. (AP)

Palynology Pollen of this species has not been investigated. (AP)

Cytogenetics

There are no ethnobotanical uses known for the species, and it is not in common cultivation. (AP)

Taxonomic notes In habit, this genus resembles Microcoelia, but its flowers are different, characterized by the concave labellum with a large erect callus at the apex, the geniculate spur with a broad mouth, the column bent back toward the tip, and the pollinarium in which the elongate, elliptic pollinia lie on the apical, expanded part of the spatulate tegula. (PC)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86.

7 2 4 .  C R I B B I A Cribbia Senghas, Orchidee (Hamburg), 36, 19 (1985). Type species: Cribbia brachyceras (Summerh.) Senghas (basionym: Rangaeris brachyceras (Summerh.) Summerh.) Azadehdelia Braem, Schlechteriana, 1, 34 (1988), nom. superfl. Type species: Azadehdelia brachyceras (Summerh.) Braem (basionym: Aerangis brachyceras Summerh.)

Derivation of name

Chromosome counts of C. deflexicalcaratum have not been reported. (AP)

The generic name commemorates Phillip Cribb, former Curator of the Orchid Herbarium at the Royal Botanic Gardens, Kew. (PC)

Phytochemistry

Description (Plate 150; Fig. 724.1)

No reports have been found on phytochemistry of Chauliodon. (NV, RG)

Erect, epiphytic herbs. Roots in basal part of stem. Stem leafy. Leaves conduplicate, subcoriaceous, lorate, unequally bilobed

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at tip, bases sheathing-imbricate. Inflorescence suberect to subpendent, as long as or longer than leaves, many-flowered; bracts triangular. Flowers resupinate, translucent, buff-coloured, yellowish or pale green. Sepals free, spreading, lineartapering. Petals spreading, linear. Labellum entire, ovate to

obovate, ecallose, spurred at base; spur cylindrical, incurved or rising above ovary. Column with an obovoid anther, pollinia two, porate, each attached by a linear stipe (tegula) to an elliptic viscidium; rostellum trilobed, each lobe a short tooth. (PC)

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embedded in sclerenchyma, cells thick-walled in C. confusa. Pith sclerenchymatous. (BC)

Palynology Pollen of this genus has not been studied. (AP)

Cytogenetics Arends and Van der Laan (1983) reported a chromosome number of 2n = 50 (as Rangaeris brachyceras). (PC)

Phytochemistry No reports have been found on the phytochemistry of Cribbia. (NV, RG)

Phylogenetics

Fig. 723.2.  Distribution map of Chauliodon.

Distribution (Fig. 724.2) Cribbia includes four species in tropical Africa. (PC)

Anatomy Carlsward et al. (2006a) examined two species of Cribbia. Their results are summarized below.

Leaf TRANSVERSE SECTION

Trichomes multicellular, glandular, basal cell sunken into a buttress of raised epidermal cells. Stomata abaxial. Hypodermis most prominent on the adaxial surface with one to two complete rows of smooth to pleated water-storage cells, less prominent abaxially with scattered water-storage cells interspersed within mesophyll chlorenchyma. Fibre bundles absent. Mesophyll homogeneous. Water-storage cell walls smooth to pitted. Vascular bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Velamen two or three cells thick. Epivelamen cells isodiametric and thin-walled to slightly ∪-thickened. Endovelamen cells angular, isodiametric to radially elongate, and thin-walled. Distinct endovelamen thickenings absent. Cover cells present over exodermal short cells in C. brachyceras. Exodermal long cells primarily ○-thickened. Water-storage cell walls smooth to pitted. Aeration units present in C. brachyceras and C. confusa P.J.Cribb. Endodermal cells strongly ○-thickened to occasionally ∩-thickened. Vascular cylinder 7–11-arch. Vascular tissue 376

Carlsward et al. (2006b) included C. brachyceras and C. confusa in their ITS rDNA analysis. They were nested in a clade with Rhipidoglossum, sister to R. kamerunense (Schltr.) Garay. Their pollinarium and rostellum structure are similar to but less prominent than those of Rhipidoglossum. Further DNA sampling of Cribbia and Rhipidoglossum species are needed to increase bootstrap support and resolve their relative positions. (PC)

Ecology Species of Cribbia are epiphytic on trunks and larger branches of trees in forests at elevations of 600–2100 m. (PC)

Pollination There is no published information on pollination of these species. (AP)

Uses Cribbia species are not in common cultivation. (AP)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86.

7 2 5 .  C RYP T OPU S Cryptopus Lindl., Bot. Reg., 10, sub t. 817 (1824). Type: Cryptopus elatus (Thouars) Lindl. (basionym: Angraecum elatum Thouars)

Derivation of name From the Greek kryptos, hidden, and pus, foot, because John Lindley believed that the pollinia were hidden in a bag-like structure. (PC)

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Fig. 724.1.  A–K. Cribbia thomensis la Croix & P.J.Cribb. A. Habit; B. Flower, side view; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum; G. Column and labellum, side view; H. Column, side view; I. Column apex, anther cap removed; J. Anther cap and pollinarium; K. Pollinarium. Drawn by Judi Stone from Brune ST38 (K). L–V. Cribbia pendula la Croix & P.J.Cribb. L. Habit; M. Dorsal sepal; N. Lateral sepal; O. Petal; P. Labellum; Q. Column and labellum, side view; R. Column, front view; S. Column, side view; T. Column apex, anther cap removed; U. Anther cap and pollinarium; V. Pollinarium. Drawn by Judi Stone from a drawing by I. la Croix, a photograph, and spirit material no. ST38A. Single bar = 1 mm, double bar = 1 cm. Reproduced with permission from Kew Bulletin, 52, 746 (1997).

Description (Plate 151; Fig. 725.1) Scrambling or scandent, epiphytic herbs. Roots distributed along stems. Stem branching, leafy. Leaves conduplicate, distichous, coriaceous, not twisted at base, articulated to a sheathing base.

Inflorescence axillary, simple or branched, many-flowered; peduncle and rachis terete. Flowers flat, white, sometimes marked with red on labellum, sepals and petals rarely olive-yellow or brown. Sepals and petals free, the latter often lobed. Labellum three- or 377

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paniculatus. Endodermal cells primarily ○-thickened to infrequently ∪-thickened. Vascular cylinder 7–13-arch. Vascular tissue embedded in sclerenchyma. Pith sclerenchymatous. (BC)

Palynology There are no known studies on the pollen of the genus. (AP)

Cytogenetics Arends and Van der Laan (1983) recorded a chromosome count of 2n = 76 for C. elatus. Brandham (1999) also reported 2n = 95 from the literature, with a basic number of x = 19 for the genus, so that 4x = 76 and 5x = 95. (AP)

Phytochemistry No reports have been found on the phytochemistry of Cryptopus. (NV, RG) Fig. 724.2.  Distribution map of Cribbia.

Phylogenetics

four-lobed. Column with two pollinia, each attached by a short stipe (tegula) to its own viscidium; rostellum trilobed, dependent. (PC)

Carlsward et al. (2006b) included Cryptopus paniculatus and C. elatus plus Oeonia rosea in their DNA analysis of Afro-Madagascan angraecoid orchids. Oeonia rosea was nested between C. elatus and C. paniculatus H.Perrier, making Cryptopus paraphyletic. More sampling of taxa is needed before taxonomic decisions can be made with authority. (PC)

Distribution (Fig. 725.2) Cryptopus comprises four species, endemic to Madagascar and the Mascarenes. (PC)

Anatomy Carlsward et al. (2006a) examined two species of Cryptopus. Their results are summarized below.

Leaf TRANSVERSE SECTION

Trichomes multicellular, glandular; absent in C. elatus. Stomata abaxial. Small globules of a brown ergastic substance (probably tannins) present in scattered ad- and abaxial epidermal cells in C. paniculatus. Hypodermis absent. Fibre bundles absent. Mesophyll homogeneous. Water-storage cells banded. Vascular bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Velamen two or three cells thick. Epivelamen cells isodiametric to tangentially flattened; walls ∪-thickened with fine anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate. Distinct endovelamen thickenings absent. Cover cells present over short cells of exodermis. Exodermal long cells ∩- to ○-thickened. Water-storage cell walls smooth to pitted or banded. Aeration units present in C. elatus and C. 378

Ecology Species of Cryptopus occur in humid and evergreen forests from sea level to 1200 m. Cryptopus paniculatus is found on rocks by the coast in littoral forest. (PC)

Pollination Nothing is known about the pollination of these species. (AP)

Uses Root fibres were once used to catch prawns in rivers of the Mascarenes. Cryptopus elatus and C. paniculatus are occasionally cultivated. (AP)

Taxonomic notes Cryptopus is morphologically similar to Oeonia, differing by the column not enclosed by the side lobes of the labellum and the lobed petals that resemble the labellum. (PC)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86.

CYRTORCHIS

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Fig. 725.1.  Cryptopus elatus (Thouars) Lindl. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum; H. Column and labellum, side view; I. Column, front view; J. Column, side view; K. Column, anther cap removed, from above; L. Anther cap, dorsal view; M. Anther cap, ventral view; N. Pollinia; O. Viscidia. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from P. Thouars, Herbarium Hookerianum 1867 and Kew Spirit Collection no. 56059.

726.  CY RTO RC H IS

Description (Plates 152, 153; Fig. 726.1, 726.2)

Cyrtorchis Schltr., Orchideen, 596 (1914). Type species: Cyrtorchis arcuata (Lindl.) Schltr. (bsasionym: Angraecum arcuatum Lindl.) Homocolleticon (Summerh.) Szlach. & Olszewski, Fl. Camer., 36, 727 (2001), syn. nov. Type species: Homocollecticon monteiroae (Rchb.f.) Szlach. & Olszewski (basionym: Cyrtorchis monteiroae (Rchb.f.) Schltr.)

Epiphytic or rarely lithophytic herbs. Roots emerging all along stem, branching. Stem erect or pendent, covered by distichous leaf bases. Leaves distichous, coriaceous or fleshy, flat or conduplicate, much longer than broad, unequally bilobed at apex, articulated to a sheathing base, deciduous and leaving obvious sharp edges of leaf-bases. Inflorescence usually shorter than leaves, axillary in upper part of stem; peduncle often covered by sterile, sheathing bracts. Flowers stellate, sweetly scented, white, sometimes with a greenish, pinkish, or brownish spur. Sepals and petals subsimilar, free, linear-lanceolate, acuminate, usually recurved. Labellum similar, entire or obscurely trilobed, ecallose, with a long, tapering spur at base; spur sigmoid or incurved. Column

Derivation of name From the Greek kyrtos, curved, and orchis, orchid, in reference to the long, apparently curved rostellum. (PC)

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Root TRANSVERSE SECTION

Velamen 2–4 cells thick. Epivelamen cells isodiametric to radially elongate; walls ∪-thickened with fine anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate; walls of outer layers thicker than those of inner layers. Distinct endovelamen thickenings ridged in C. praetermissa and C. ringens. Cover cells present over short cells of the exodermis. Exodermal long cells primarily ○-thickened. Water-storage cell walls smooth to pitted or banded. Mucilage present in cortical cells of C. arcuata, C. arcuata ssp. whytei, and C. chailluana. Aeration units present in C. arcuata, C. chailluana, and C. praetermissa. Endodermal cells primarily ○-thickened to rarely ∪-­thickened. Vascular cylinder 17–34-arch. Vascular tissue embedded in sclerenchyma, cell walls of embedding sclerenchyma thickest around phloem clusters in C. arcuata ssp. whytei and C. praetermissa. Pith sclerenchymatous. (BC)

Palynology Fig. 725.2.  Distribution map of Cryptopus.

with an elongate anther cap at apex, pollinia two, stipes two, oblanceolate, viscidium either hyaline or comprising an indurate, saddle-shaped upper part and a hyaline lower part; rostellum trilobed, pendent, outer lobes much longer than midlobe, often papillate toward apex. Capsule 3–6-winged. (PC)

Distribution (Fig. 726.3) Cyrtorchis is a genus of 18 species confined to tropical and southern Africa. (PC)

Anatomy Carlsward et al. (2006a) examined five species of Cyrtorchis. Their results are summarized below.

Leaf TRANSVERSE SECTION

Trichomes multicellular, glandular; absent in C. chailluana (Hook.f.) Schltr. Stomata abaxial. Hypodermis absent or adaxial in C. praetermissa Summerh. and C. ringens (Rchb.f.) Summerh. (composed of fibrous idioblasts arranged in a single row and interspersed among mesophyll chlorenchyma cells). Fibre bundles absent. Mesophyll homogeneous in C. arcuata and C. arcuata ssp. whytei (Rolfe) Summerh.; heterogeneous with columnar adaxial cells and isodiametric abaxial cells in C. aschersonii (Kränzl.) Schltr., C. praetermissa, and C. ringens. Water-storage cells smooth to pitted. Fibrous idioblasts scattered throughout mesophyll in C. aschersonii, C. praetermissa, and C. ringens. Vascular bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma. 380

Schill and Pfeiffer (1977) described the pollen tetrads of C. ringens as convex-concave with laevigate-foveolate sculpturing and a calymmate sexine. (AP)

Cytogenetics Arends and Van der Laan (1983) recorded chromosome counts of six taxa of Cyrtorchis as 2n = 46, 92, 138. In addition, Brandham (1999) reported counts of 2n = 44, 45, 50, c. 150. He suggested a basic number of x = 23, yielding 2x = 46 (the majority of chromosome numbers), 4x = 92, and 6x = 138. (AP)

Phytochemistry One of four species of Cyrtorchis analyzed by Lüning (1974) is recorded as having an alkaloid content ≥ 0.1%, but no species names were given. (NV, RG)

Ecology Species of Cyrtorchis are epiphytic in the open, in shade, or in partial shade in semi-deciduous and evergreen forests and riverine forests, or epiphytic or lithophytic in Brachystegia Benth. (Fabaceae) and savanna woodlands, ranging from sea level to 2000 m. (PC)

Pollination Little is known about pollination in Cyrtorchis except that C. aphylla may be autogamous based on observations by Smith (1907) that one stigma lobe is either in contact with pollinia or situated in such a way as to receive falling pollen (Catling 1990). (AP)

Uses Lawler (1984) reported that an extract of one unspecified species has been used as an antimalarial agent. Several species, especially C. arcuata and C. praetermissa, are often cultivated. (AP)

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M L Fig. 726.1.  Cyrtorchis crassifolia Schltr. A. Habit; B. Leaf, transverse section; C. Flower, side view; D. Flower, front view; E. Bract; F. Dorsal sepal; G. Lateral sepal; H. Petal; I. Labellum, spur removed; J. Column and labellum, side view; K. Column, front view; L. Column, side view; M. Anther cap with pollinia; N. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Kew Spirit Collection no. 25764. Reproduced with permission from G. V. Pope (ed.) (1998), Flora Zambesiaca: Mozambique, Malawi, Zambia, Zimbabwe, Botswana. Volume 11, Part 2. Royal Botanic Gardens, Kew.

Taxonomic notes Summerhayes (1948) revised the genus and recognized two sections: Cyrtorchis (as Heterocollecticon) and Homocollecticon. The former has a viscidium of two parts, the part nearest the anther

indurate, the upper part hyaline. In section Homocollecticon, the viscidium is entirely hyaline. Szlachetko and Olszewski (2001) raised the latter to generic rank as Homocollecticon, but the distinction seems to be based upon a single character, and Cyrtorchis is otherwise morphologically homogeneous. (PC) 381

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Fig. 726.2.  Cyrtorchis glaucifolia Summerh. A. Habit; B. Flower; C. Flower, sepals and petals removed; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Anther cap; H. Column, anther cap removed; I. Column, anther cap removed, front view; J, K. Pollinarium, two views. Single bar = 1 mm, double bar = 1 cm. Drawn by Mary Grierson from Schelpe & Leach 6980 (K). Reproduced with permission from G. V. Pope (ed.) (1998), Flora Zambesiaca: Mozambique, Malawi, Zambia, Zimbabwe, Botswana. Volume 11, Part 2. Royal Botanic Gardens, Kew.

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86.

382

Summerhayes, V. S. S. (1948). African orchids XVIII. Kew Bulletin, 1947, 277–302. Szlachetko, D. and Olszewski, T. (2001). In Flore du Cameroun 36. Orchidacees, vol. 3 (ed. G. Achoundong and P. Morat), pp. 790–808. Ministere de la Recherche Scientifique et Technique, Yaounde, Cameroun.

DENDROPHYLAX

Distribution (Fig. 727.2) There are currently 14 recognized species of Dendrophylax, with a centre of distribution in the Greater Antilles, particularly Hispaniola. The northernmost range is in Florida (D. lindenii (Lindl.) Benth. ex Rolfe and D. porrectus) and the westernmost range in El Salvador and Mexico (D. porrectus). (BC)

Anatomy Carlsward et al. (2006a) described the root anatomy of eight species of Dendrophylax. Their observations are summarized below.

Root TRANSVERSE SECTION

Fig. 726.3.  Distribution map of Cyrtorchis.

727.  DE N D RO P H Y LA X Dendrophylax Rchb.f. in Ann. Bot. Syst. (Walpers), 6, 903 (1864). Type species: Dendrophylax varius (J.F.Gmel.) Urb. (basionym: Orchis varia J.F.Gmel.) Polyrrhiza Pfitzer, Die Nat. Pflanzenf., 2, 6, 208 (1888). Type species: Polyrrhiza funalis (Sw.) Pfitzer Harrisella Fawc. & Rendle, J. Bot., 47, 265–6 (1909). Type species: Harrisella porrecta (Rchb.f.) Fawc. & Rendle (basionym: Aeranthes porrecta Rchb.f.) Polyradicion Garay, J. Arn. Arb., 50, 466 (1969). Type species: Polyradicion lindenii (Lindl.) Garay (basionym: Angraecum lindenii Lindl.)

Derivation of name From the Greek dendro, tree, and phylax, guard, referring to the clasping epiphytic roots. (BC)

Description (Plates 154, 155; Fig. 727.1) Leafless, epiphytic herbs. Roots photosynthetic, comprising most of plant. Stem erect, short (usually less than 1 cm). Leaves reduced to nonphotosynthetic scales. Inflorescence usually solitary, axillary, unbranched, 1–10 flowered (usually with one flower open at a time), much longer than stems. Flowers usually resupinate in largeflowered species and nonresupinate in small-flowered species; colour may be white, green, or green with white labellum. Sepals and petals free, lanceolate, reflexed, equal in size or sepals wider than petals. Labellum entire or 2–4-lobed; often deeply spurred; spur usually several times larger than entire flower; rarely bilobed (as in D. porrectus (Rchb.f.) Carlsward & Whitten); ridged callus at base of labellum. Column without a foot; pollinia two, with one or two stipes; rostellum deeply divided. Ovary narrowly cylindric or rarely globose (D. porrectus) and ridged. (BC)

Velamen 1–4 layered; epivelamen cells radially elongate to isodiametric, ∪-thickened with fine anastomosing radial wall thickenings; endovelamen cells angular with undulate to straight-sided radial walls and isodiametric to radially elongate, outer layers with thicker cell walls than inner layers. Cover cells present over exodermal passage cells. Distinct endovelamen thickenings absent. Exodermal long cells ∩ n-thickened to rarely ○-thickened. Water-storage cells with birefringent bands in walls or thickened along the edges in D. porrectus. Aeration units present. Endodermal cells ○-thickened. Vascular cylinder 4–13-arch. Vascular tissue embedded in sclerenchyma. Pith sclerenchymatous. (BC)

Palynology Dendrophylax varius has two globular pollinia with smooth, slightly convex tetrads. The wall sculpturing is smooth to slightly plicate, with a sexine that is 1.3 µm thick (Schill and Pfeiffer 1977). (BC)

Cytogenetics Dendrophylax funalis (Sw.) Benth. ex Rolfe is reported to have 2n = 42 (Shindo and Kamemoto 1963). Felix and Guerra (2010) also reported 2n = 44 for D. monteverdi (Rchb.f.) Ackerman & Nir (as Campylocentrum). (BC)

Phytochemistry No reports have been found on the phytochemistry of Dendrophylax. (NV, RG)

Phylogenetics Because of the reduced nature of these leafless plants, much of the original taxonomy was historically based on floral morphology. Most species were originally placed in Aeranthus Lindl. by early workers and were later separated from the Palaeotropical taxa. In his treatment of Antillean Orchidaceae, Nir (2000) transferred Polyradicion and Campylocentrum constanzense Garay into Dendrophylax and transferred Harrisella into Campylocentrum Benth. This left two Neotropical genera, Dendrophylax and Campylocentrum, 383

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Fig. 727.1.  Dendrophylax porrectus (Rchb.f.) Carlsward & Whitten. A. Habit; B. Inflorescence; C. Flower, front view; D. Flower, side view; E. Capsule; F. Dorsal sepal; G. Lateral sepal; H. Petal; I. Labellum; J. Column and labellum, side view; K. Column, side view; L. Column, front view; M. Anther cap, dorsal view; N. Anther cap, ventral view; O. Pollinarium; P. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Cribb s.n. (K) and Kew Spirit Collection no. 50625.

which were later determined to be sister taxa (Carlsward et al. 2003). Based on phylogenetic analyses of molecular data (ITS, matK, and trnL-F regions), Carlsward et al. (2003) confirmed Nir’s (2000) inclusion of Polyradicion and Campylocentrum constanzense in Dendrophylax. They also transferred the small-flowered Harrisella, Campylocentrum macrocarpum Dod, and Campylocentrum filiforme (Sw.) Cogn. ex Kuntze in Dendrophylax. In general, species of Dendrophylax have fewer flowers and a more lax inflorescence than the closely related Campylocentrum. (BC)

Ecology Species of Dendrophylax occur from sea level to about 1600 m (D. macrocarpus (Dod) Carlsward & Whitten). Many are found in 384

shaded sites with high humidity on tree trunks (D. lindenii) but may also be found in xeric habitats on twigs of shrubs or small trees (D. porrectus) or on limestone rocks (D. funalis). (BC)

Pollination Pollination by sphingid moths has been hypothesized by Luer (1972) for D. lindenii. Ongoing research by J. J. Sadler, L. W. Zettler, and L. W. Richardson in southern Florida is attempting to verify Luer’s hypothesis. According to C. V. Covell, Jr., Florida Museum of Natural History (personal communication to LWZ), the giant sphinx moth (Cocytius antaeus) is the most likely candidate as the primary pollinator of D. lindenii, based on proboscis length, distribution, flight times that overlap flowering, and

DIAPHANANTHE

Description (Plate 156; Fig. 728.1, 728.2)

Fig. 727.2.  Distribution map of Dendrophylax.

associated larval food sources in the region (Annona glabra L., Annonaceae). (BC)

Uses Plants of Dendrophylax are infrequently cultivated for their large, showy flowers. (BC)

Cultivation Most species of Dendrophylax are difficult to cultivate in a greenhouse, and many seem to have a relatively short lifespan. Also because of their leafless habit, these plants are not often found in cultivation. The most commonly cultivated species (and the easiest to grow) is D. funalis from Jamaica. Most species are best grown in plastic hanging baskets without a medium but with high humidity and frequent watering. Some species such as D. fawcettii can be asexually propagated via plantlets produced on old inflorescence axes. Plants tend to do poorly with algaecide or fungicide application. (BC)

Taxonomic literature Carlsward, B. S., Whitten, W. M., and Williams, N. H. (2003). Molecular phylogenetics of Neotropical leafless Angraecinae (Orchidaceae): reevaluation of generic concepts. International Journal of Plant Sciences, 164, 43–51. Nir, M. A. (2000). Orchidaceae Antillanae. DAG Media Publishing, Inc., New York.

728.  DIAPH A N A N TH E Diaphananthe Schltr., Orchideen, 593 (1914). Type species: D. pellucida (Lindl.) Schltr. (basionym: Angraecum pellucidum Lindl.) Chamaeangis Schltr., Beih. Bot. Centralbl., 36, 107 (1918). Type species: C. gracilis (Thouars) Schltr. (basionym: Angraecum gracile Thouars)

Derivation of name From the Greek, diaphanes, translucent, and anthos, flower, in allusion to the translucent nature of its flowers. (PC)

Epiphytic or rarely lithophytic herbs. Roots emerging all along stem through leaf-bases, often prominent and rarely branched. Stem usually unbranched and covered by sheathing leaf-bases. Leaves distichous, coriaceous or fleshy, rarely thin-textured, unequally bilobed at apex, articulated at base to a sheathing leafbase, often twisted at base to lie in one plane. Inflorescences one to many, emerging through the sheathing leaf-bases in upper part of stem, usually several-flowered; bracts usually amplexicaul, rarely prominent. Flowers white, pale green or yellow, rarely pinkish, usually translucent, rarely showy. Sepals free, subsimilar. Petals free. Labellum entire or obscurely lobed, spurred at base, usually with a tooth-like or transverse callus in mouth of spur. Column porrect; pollinia two, stipites two, viscidia one or two; rostellum pendent or reflexed, linear or tapering and bifid to pendent, clavate and obscurely trilobed. (PC)

Distribution (Fig. 728.3) Diaphananthe is a genus of about 33 species confined mostly to tropical Africa. (PC)

Anatomy Carlsward et al. (2006a) examined eight species of Diaphananthe (including four formerly treated as Chamaeangis). Their results are summarized below.

Leaf TRANSVERSE SECTION

Trichomes multicellular and glandular; absent in D. fragrantissima (Rchb.f.) Schltr. and species once ascribed to Chamaeangis. Stomata abaxial or both ad- and abaxial. Hypodermis absent. Fibre bundles absent. Mesophyll homogeneous or rarely isobilateral. Water-storage cells smooth to pitted or banded. Vascular bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Velamen 2–4 cells thick. Epivelamen cells isodiametric to radially elongate; walls ∪-thickened with fine anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate; outermost layers of cells thicker-walled than inner layers. Distinct endovelamen thickenings usually ridged. Cover cells present over short cells of exodermis except in D. fragrantissima. Exodermal long cells ∩-thickened or ○-thickened. Water-storage cell walls smooth to pitted or banded. Single modified cortical layer of thin- to thick-walled cells surrounding the endodermis in D. bidens and D. lorifolia. Aeration units often present. Endodermal cells primarily ○-thickened; rarely ∪-thickened in D. bidens. Vascular cylinder 9–28-arch. Vascular tissue embedded in sclerenchyma, cell walls of embedding sclerenchyma occasionally thickest around phloem clusters. Pith usually sclerenchymatous, but parenchymatous in D. bidens. (BC) 385

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Fig. 728.1.  Diaphananthe odoratissima (Rchb.f.) P.J.Cribb & Carlsward. A. Habit; B. Flower, side view; C. Flower, spur removed, front view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum; H. Column, front view; I. Column, side view; J. Anther cap; K. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from La Croix 1104 (K) and Kew Spirit Collection no. 45129. Reproduced with permission from G. V. Pope (ed.) (1998), Flora Zambesiaca: Mozambique, Malawi, Zambia, Zimbabwe, Botswana. Volume 11, Part 2. Royal Botanic Gardens, Kew.

Palynology

Cytogenetics

Schill and Pfeiffer (1977) examined the pollen of Chamaeangis vesicata (Lindl.) Schltr. (=D. vesicata (Lindl.) P.J.Cribb & Carlsward) and described it as similar to that of Angraecum. (AP)

Arends and van der Laan (1983) published a chromosome count of 2n = 96 for C. vesicata (=D. vesicata). Jones (1967) and Arends and Van der Laan (1983) reported chromosome counts of

386

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Fig. 728.2.  Diaphananthe pellucida (Lindl.) Schltr. A. Habit, ×2/3; B. Dorsal sepal, ×3; C. Lateral sepal, ×3; D. Petal, ×3; E. Labellum, ×3; F. Spur and portion of labellum, side view, ×4; G. Column, side view, ×8; H. Column, front view, ×8; I. Anther cap, ×8; J. Pollinarium, ×8. Magnifications as originally published. Drawn by Maureen E. Church from Linder 1375 (K) and Smith 604 (K). Reproduced with permission from P. J. Cribb (1989), Flora of Tropical East Africa, Orchidaceae (Part 3). A.A. Balkema, Rotterdam/Brookfield.

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(2003a) illustrated the similarity of the column and pollinarium morphologies of Chamaeangis odoratissima and Diaphananthe fragrantissima, showing that both have a large stigma, tapering bifid rostellum, and two porate pollinia, each attached to its own tegula and a single viscidium. Carlsward et al. (2006b) included three species of Diaphananthe sensu stricto and four species of Chamaeangis in their phylogenetic analyses of Afro-Malagasy vandaceous orchids. Diaphananthe was embedded in a paraphyletic Chamaeangis with strong bootstrap support, so there is no doubt that the latter should be synonymized under the earlier name, Diaphananthe. (PC)

Ecology Most species are epiphytes of riverine and humid evergreen forests from sea level to 2400 m. (PC)

Pollination

Fig. 728.3.  Distribution map of Diaphananthe.

Johansson (1974) reported moths of the genus Euchromia visiting D. pellucida. Otherwise, nothing is known about pollination in this genus. (AP)

Uses 2n = 50, 100 for Diaphananthe. Brandham (1999) suggested a high basic number of x = 25 for the latter. (AP)

Phytochemistry Flavone C-glycosides are present in the leaves of C. odoratissima (Rchb.f.) Schltr. (=D. odoratissima (Rchb.f.) P.J.Cribb & Carlsward) according to C. Williams (1979). Lüning (1974) recorded that of eight species of Diaphananthe tested for alkaloids, one contained them at levels ≥ 0.1% (species names not given). The floral fragrance of D. pellucida (Lindl.) Schltr. is dominated by large amounts of phenylacetonitrile (62.0%) together with 3-methylbutyronitrile (3.5%) and 2-methylbutyronitrile (< 1%). These are degradation products of the oximes, phenylacetaldoxime (2.2%), 3-methylbutyraldoxime (< 1%), and 2-methylbutyraldoxime (0.1%), respectively, which are also found in other angraecoid orchids (Kaiser 1993). (NV, RG)

Phylogenetics Schlechter (1918) established the genus Chamaeangis for a number of African angraecoid orchids that were distinguished by having a many-flowered inflorescence of small flowers, often borne in whorls along the rachis. However, some Diaphananthe species, such as D. fragrantissima (Rchb.f.) Schltr., also bear their flowers in whorls along the rachis. Furthermore, the similarity in the floral structure of the African species of Chamaeangis with those of Diaphananthe has been long recognized by orchid taxonomists such as Senghas (1986b), who grouped the two genera in the same key couplet. He distinguished Chamaeangis from Diaphananthe by its short stem, lack of callus at the base of the labellum in the mouth of the spur, and oval labellum. None of these characters alone is diagnostic for the genus. Szlachetko 388

Lawler (1984) reported that the long aerial roots of Diaphananthe bidens (as Mystacidium productum Kränzl.) are used in Fernando Po to tie bundles of sticks for yam cultivation and to weave baskets there as well as in Equatorial Guinea. (PC)

Taxonomic notes Diaphananthe, as recognized here, is confined to those species in which the tegulae are attached to a single viscidium; thus, Rhipidoglossum is considered distinct. Although the latter was established by Schlechter (1918), it has not generally been accepted as distinct by most authors (Summerhayes 1968; Cribb 1989; La Croix and Cribb 1998). Szlachetko and Olszewski (2001) resurrected the genus as distinct from Diaphananthe and distinguished it by its pollinia that are each attached by a linear stipe to a discrete viscidium and by the rostellum that has a prominent, fleshy, pendent midlobe. (PC)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Cribb, P. J. (1989). Flora of tropical East Africa. Orchidaceae, 3. Balkema, Rotterdam. Cribb, P. J. and Carlsward, B. (2012). New combinations in Aerangis, Diaphananthe and Podangis (Orchidaceae, subtribe Angraecinae). Phytotaxa, 71, 42–7. La Croix, I. and Cribb, P. J. (1998). Flora Zambesiaca. Volume 11, Part 2. Orchidaceae. Royal Botanic Gardens, Kew. Schlechter, F. R. R. (1918). Versuch einer natürlichen Neuordnung der afrikanischen angraekoiden Orchidaceen. Beihefte Botanisches Centralblatt, 36, 62–181.

DINKLAGEELLA

Senghas, K. H. (1986b). Tribus: Vandeae. In Rudolf Schlechter Die Orchideen, 3rd ed., (ed. F. G. Brieger, R. Maatsch, and K. Senghas), pp. 973–1130. Blackwell, Berlin. Summerhayes, V. S. S. (1968). 42. Diaphananthe Schltr. In Flora of west tropical Africa, Orchidaceae (ed. F. N. Hepper), pp. 260–3. Crown Agents for Overseas Governments and Administrations, London. Szlachetko, D. L. (2003a). Gymnostemia orchidalium 3. Acta Botanica Fennica, 176, 1–311. Szlachetko, D. and Olszewski, T. (2001). Flore du Cameroun. 36. Orchidacees, volume 3. Ministere de la Recherche Scientifique et Technique, Yaounde, Cameroun.

729.  DIN KLAG E E LLA Dinklageella Mansf., Repert. Sp. Nov. Regni Veg., 36, 63 (1934). Type species: Dinklageella liberica Mansf. Lacroixia Szlach. & Olszewski, Ann. Bot. Fenn., 40, 69 (2003). Type species: Lacroixia minor (Summerh.) Szlach. & Olszewski (basionym: Dinklageella minor Summerh.), syn. nov.

Derivation of name The generic name commemorates M. Dinklaage, a late 19th century German merchant and orchid enthusiast in Cameroon. (PC)

Description (Fig. 729.1) Epiphytic herbs. Roots emerging individually through sheathing leaf bases. Stem cylindrical, leafy. Leaves distichous, coriaceous, oblongelliptic, unequally roundly bilobed at tip, with sheathing cylindrical leaf bases. Inflorescence lateral, axillary, emerging through sheathing leaf bases, longer than leaves, few-flowered. Flowers resupinate, creamy white. Sepals free, spreading, elliptic to oblong, rounded at tip. Petals free, oblong-elliptic, obtuse. Labellum trilobed, recurved with reflexed lateral lobes, calcarate, lateral lobes obovate or narrowly ovate, midlobe obovate and truncate or narrowly ovate; spur clavate or sigmoid, cylindrical and dilated toward tip. Column with two pollinia, cleft, attached near centre of a single, elliptical stipe (tegula) to a rectangular viscidium; rostellum bilobed. Ovary with pedicel longer than floral bracts. (PC)

Distribution (Fig. 729.2) Dinklageella is a genus of four species ranging from Liberia across to Cameroon. (PC)

Anatomy Nothing is known about the vegetative anatomy of the genus. (BC)

Palynology Pollen of these species has not been studied. (AP)

Cytogenetics Chromosome counts for Dinklageella have not been reported. (AP)

Phytochemistry No reports have been found on the phytochemistry of Dinklageella. (NV, RG)

Ecology Species occur from sea level to 1000 m in exposed situations on margins of lowland rainforest and on granite boulders in wet regions. They are also known from coffee plantations. (PC)

Pollination Nothing is presently known about the pollination of these species. (AP)

Uses Dinklageella species are not commonly cultivated. (AP)

Taxonomic notes Mansfeld established Dinklageella based on the scandent west African species D. liberica. It was distinguished by its 1) trilobed labellum with broad lobes, 2) pollinia attached near the centre of the ovate, upper part of a single elliptic tegula, bent and dilated in the middle, and attached to a convex elliptic viscidium, and 3) short, bifid, porrect rostellum. Summerhayes (1960) described a second species, D. minor Summerh., with smaller flowers with a short, S-shaped spur. Szlachetko (2003b) established a new genus Lacroixia for D. minor, distinguishing it by its S-shaped spur with a clavate tip, unbent tegula, and semi-elliptic viscidium. There are no molecular data to support the split. The differences outlined by Szlachetko and Olszewski seem to be matters of degree rather than distinctions sufficient to recognize the two species in different genera. Rice (2006b) included Dinklageella (in the sense used here) as a section of Solenangis, basing his decision on their similar habit and pollinia attached to a single tegula and viscidium. However, the labellum lobing and pollinarium structures are different in the two genera. The tegula in Solenangis is linear not elliptic, and the pollinia are porate rather than cleft as in Dinklageella. (PC)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Micheneau, C., Carlsward, B. S., Fay, M. F., Bytebier, B., Pailler, T., and Chase, M. W. (2008). Phylogenetics and biogeography of Mascarene angraecoid orchids. Molecular Phylogenetics and Evolution, 46, 908–22. Rice, R. (2006a). An overview and three new species of the Solenangis aphylla alliance. Oasis, 3, 2–11. Rice, R. (2006b). Notes on the subtribe Aerangidinae (Orchidaceae). Oasis, 3, 12–15. Summerhayes, V. S. (1960). African orchids: XXVII. Kew Bulletin, 14, 126-157.

389

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Fig. 729.1.  Dinklageella liberica Mansf. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum; H. Column and labellum, side view; I. Column, front view; J. Column, side view; K. Column, anther cap removed, front view; L. Anther cap, dorsal view; M. Anther cap, ventral view; N. Pollinarium, front view; O. Pollinarium, from below; P. Pollinarium, back view; Q. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Perez-Vera 726 (K) and Kew Spirit Collection no. 13604.

Szlachetko, D. (2003b). Genera et species orchidalium. 7. Vandeae. Annales Botanici Fennici, 40, 67–70.

730.  D IS TY LO D O N Distylodon Summerh., Kew Bull., 20, 197 (1966). Type species: Distylodon comptum Summerh.

Derivation of name From the Greek di, two, and stylos, pillar, in allusion to the unusual rostellum shape. (PC) 390

Description (Fig. 730.1) Epiphytic herbs. Roots produced along length of stem. Leaves distichous, conduplicate, elliptic, twisted at base to lie in one plane. Inflorescence axillary, one-flowered. Flowers pale green, not opening widely. Sepals and petals free, lanceolate-tapering. Labellum free, concave, enclosing column base, trilobed, lateral lobes shorter than midlobe, acute, spurred at base; spur cylindrical, straight. Column with two porate pollinia, each attached by an oblanceolate tegula to a linear, elongate viscidium; stigma oblong; rostellum at first reflexed then porrect, bilobed, each lobe linear. (PC)

EGGELINGIA

and distinctive rostellum and androecium and from the latter in its column and rostellum structure. There are no molecular data as yet to indicate its affinities. (PC)

7 3 1 .  EGGEL I NGI A Eggelingia Summerh., Bot. Mus. Leafl., Harvard Univ., 14, 235 (1951). Type species: Eggelingia clavata Summerh.

Derivation of name The generic name commemorates William Eggeling, who collected plants for the Royal Botanic Gardens, Kew, in Uganda. (PC)

Description (Plate 157; Fig. 731.1)

Distylodon is a monospecific genus known only from western Uganda and Rwanda. (PC)

Erect to pendent epiphytic herbs. Roots mostly near base of stem. Stem elongate. Leaves distichous, coriaceous, ligulate to linear-oblong, unequally bilobed at tip, sheathing at base with a tapering ligule opposite leaf at apex of sheath. Inflorescence axillary, sessile, one- to few-flowered. Flowers white. Sepals and petals subsimilar, free, elliptic-ovate, subacute. Labellum entire, ecallose, calcarate; spur clavate, about as long as ovary or shorter. Column with obovoid anther, acute at tip, pollinia two, porate, attached to a common spatulate tegula and it to an oblong or elliptic viscidium; rostellum triangular. (PC)

Anatomy

Distribution (Fig. 731.2)

There are no available reports on the vegetative anatomy of this species. (BC)

This is a genus of three species, ranging from Cote d’Ivoire in West Africa across to Uganda and south to Zambia. (PC)

Fig. 729.2.  Distribution map of Dinklageella.

Distribution (Fig. 730.2)

Palynology Pollen of this species has not been investigated. (AP)

Cytogenetics Chromosome counts for this species have not been reported. (AP)

Phytochemistry No reports have been found on the phytochemistry of Distylodon. (NV, RG)

Ecology Distylodon comptum is an epiphyte in rain forests from 700–1270 m in elevation. (PC)

Pollination

Anatomy Carlsward et al. (2006a) examined E. ligulifolia Summerh. Their results are summarized below.

Leaf TRANSVERSE SECTION

Trichomes multicellular, glandular. Stomata abaxial. Hypodermis ad- and abaxial, composed of fibrous idioblasts in one discontinuous row scattered amongst mesophyll chlorenchyma cells. Fibre bundles absent. Mesophyll heterogeneous, adaxial cells columnar, abaxial cells isodiametric. Water-storage cells smooth to pitted. Solitary fibrous idioblasts scattered throughout mesophyll. Vascular bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root Roots have not been studied. (BC)

The pollination biology of D. comptum is unknown. (AP)

Taxonomic notes Superficially, Distylodon resembles a small-flowered Angraecum or Angraecopsis, but it differs from the former in its trilobed labellum

Palynology Schill and Pfeiffer (1977) described the pollen tetrads of E. liguifolia as concave-convex with laevigate sculpturing and a calymmate sexine. (AP) 391

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G Fig. 730.1.  Distylodon comptum Summerh. A. Habit, ×2; B. Flower, side view, ×4; C. Flower, front view, ×4; D. Dorsal sepal, ×6; E. Lateral sepal, ×6; F. Petal, ×6; G. Flower with sepals and petals removed, ×4; H. Labellum, one side flattened, ×6; I. Column, oblique view; J. Column, anther cap raised and pollinaria removed; K. Column, anther cap removed, ×8; L. Anther cap, ×8; M. Pollinarium, ×12. Magnifications as originally published. Drawn by Maureen E. Church from Eggeling 5381 (K). Reproduced with permission from P. J. Cribb (1989), Flora of Tropical East Africa, Orchidaceae (Part 3). A.A. Balkema, Rotterdam/Brookfield.

392

ERASANTHE

7 3 2 .  ER A S A NT H E Erasanthe P.J.Cribb, Hermans & D.Roberts, Adansonia, 29, 28 (2007). Type species: Erasanthe henrici (Schltr.) P.J.Cribb, Hermans & D.Roberts (basionym: Aeranthes henrici Schltr.)

Derivation of name The generic name is an anagram of Aeranthes, the genus in which the type species was formerly placed. (PC)

Description (Plate 158; Fig. 732.1) Epiphytic herb. Stem leafy. Leaves distichous, leathery, elliptic-obovate, obliquely bilobed at apex. Inflorescence pendent, as long as leaves or longer, few-flowered. Flowers white with green on labellum. Sepals and petals spreading but not widely, free, lanceolate, acuminate, petals inserted on column foot. Labellum entire, lacerate, with a spur longer than labellum. Column with a prominent saccate foot; pollinia two, porate, each attached by a short stipe to its own elongate viscidium; rostellum deeply emarginate. (PC) Fig. 730.2.  Distribution map of Distylodon.

Cytogenetics Brandham (1999) reported a chromosome count of 2n = 46 for the genus. (AP)

Phytochemistry Little has been published on the phytochemistry of Eggelingia, other than the fact that alkaloids are present in one unnamed species at levels ≥ 0.1% (Lüning 1974). (NV, RG)

Distribution (Fig. 732.2) Erasanthe is a monospecific genus endemic to Madagascar, the solitary species being represented by two subspecies, one in the north and the other in the southwest. (PC)

Anatomy Nothing is known about the vegetative anatomy of this species. (BC)

Palynology Pollen data for these two species are not available. (AP)

Ecology Species of Eggelingia occur from sea level to 1950 m in humid evergreen forests, often forming large clumps on the larger branches of large trees, rarely on old lava flows. (PC)

Cytogenetics Chromosome counts for Erasanthe have not been published. (AP)

Pollination

Phytochemistry

There are no reports concerning pollination of the genus. (AP)

No reports have been found on the phytochemistry of Erasanthe. (NV, RG)

Uses Species of Eggelingia have no known ethnobotanical uses and are not in common cultivation. (AP)

Taxonomic notes Members of Eggelingia are distinguished from Tridactyle, which they superficially resemble, by the distinct ligule at the apex of the leaf sheath opposite the leaf blade, the entire labellum that lacks thickened basal auricles, and a pollinarium in which the elliptical to oblong viscidium is larger and linear, nearly as long as the tegula. There are as yet no molecular data to indicate the phylogenetic affinities of the genus. (PC)

Phylogenetics Recent DNA studies, based on DNA sequences of four plastid markers by Micheneau (2005), have indicated that Erasanthe henrici (as Aeranthes) is sister to Beclardia, Cryptopus, plus Oeonia and is distant from the other Aeranthes species, which form a strongly supported sister to Jumellea, another predominantly Madagascan genus. (PC)

Ecology Erasanthe henrici grows in deep shade in lowland and montane evergreen forests and along streams in sandstone massifs at 250–1000 m. (PC) 393

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Fig. 731.1.  Eggelingia clavata Summerh. A. Habit; B. Flower, side view; C. Flower, front view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum; H. Column and labellum, side view; I. Column, front view; J. Column, anther cap removed, front view; K. Column, anther cap removed, side view; L. Anther cap; M. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Morris 155 (K) and Kew Spirit Collection no. 45128. Reproduced with permission from G. V. Pope (ed.) (1998), Flora Zambesiaca: Mozambique, Malawi, Zambia, Zimbabwe, Botswana. Volume 11, Part 2. Royal Botanic Gardens, Kew.

394

EURYCHONE

Micheneau, C., Carlsward, B. S., Fay, M. F., Bytebier, B., Pailler, T., and Chase, M. W. (2008). Phylogenetics and biogeography of Mascarene angraecoid orchids. Molecular Phylogenetics and Evolution, 46, 908–22.

7 3 3 .  EU RYC H ONE Eurychone Schltr., Beih. Bot. Centralbl., 36, 134 (1918). Type species: Eurychone rothschildiana (O’Brien) Schltr. (basionym: Angraecum rothschildianum O’Brien)

Derivation of name From the Greek eurys, broad, and choane, funnel, in reference to the broad opening to the spur of the labellum. (PC)

Description (Plate 159; Fig. 733.1)

There is no published information on pollination of these two species, although they conform to the sphigophilous syndrome of characters. (AP)

Epiphytic herbs. Roots basal, elongate. Stem with a few leaves, distichously arranged, not coriaceous, elliptic-obovate, unequally bilobed at tip. Inflorescence axillary, pendent or spreading, shorter than leaves. Flowers resupinate, large, showy, white with a green or browinish rose mark on labellum. Sepals and petals free, subsimilar, spreading. Labellum infundibuliform or trumpet-shaped, simple or obscurely trilobed, ecallose, spurred at base, spur dilated above and then tapering to a narrow isthmus. Column dilated at base; pollinia two, porate, attached by a fusiform stipe (tegula) to a broadly ovate or triangular viscidium; stigma cavity circular; rostellum spatulate, notched at tip. (PC)

Uses

Distribution (Fig. 733.2)

Erasanthe henrici is often cultivated as Aeranthes henrici in specialist orchid collections. (AP)

Eurychone is a genus of two species in West and Central Africa, from Guinea and Sierra Leone eastward to Uganda and south to Angola. (PC)

Fig. 731.2.  Distribution map of Eggelingia.

Pollination

Taxonomic notes Erasanthe henrici is morphologically distinct from Aeranthes in having stouter roots, oblanceolate leaves with undulate margins, and stouter inflorescence axes, bearing simultaneously several large, white flowers marked with green on the labellum rather than fibrous roots, wiry inflorescences, and smaller translucent flowers. Its large flowers are also distinct for its lacerate labellum and long, slender spur, neither of which is present in other Aeranthes species. In its roots, inflorescence, and spur morphology it is morphologically more similar to Aerangis than to Aeranthes. Aeranthes polyanthemus Ridl. also belongs here by reason of its many-flowered inflorescences in which the flowers open simultaneously and in having stout rather than wiry roots. (PC)

Anatomy Carlsward et al. (2006a) examined E. rothschildiana. Their results are summarized below.

Leaf TRANSVERSE SECTION

Trichomes multicellular, glandular. Stomata abaxial. Hypodermis absent. Fibre bundles absent. Mesophyll homogeneous. Waterstorage cells smooth to pitted. Vascular bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Micheneau, C. (2005). Pollination biology and phylogenetics of the orchids of Réunion. Ph.D. thesis, Université de la Réunion.

Velamen two or three cells thick. Epivelamen cells isodiametric; walls ∪-thickened with fine anastomosing radial bands Endovelamen cells angular, isodiametric to radially elongate. Distinct endovelamen thickenings absent. Cover cells infrequent over short cells of the exodermis. Exodermal long cells primarily ○-thickened. Water-storage cells smooth to pitted. Aeration units present. Endodermal cells ○-thickened. Vascular cylinder 395

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Fig. 732.1.  Erasanthe henrici (Schltr.) P.J.Cribb, Hermans & D.L.Roberts. A. Habit; B. Flower, lateral sepals and petals removed, side view; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Flower, lateral sepals and petals removed, side view; G. Labellum, dorsal view; H. Labellum, ventral view; I. Column, anther cap removed, side view; J. Column, pollinarium removed, side view; K. Anther cap; L. Pollinia; M. Ovary, transverse section. Scale bars: A–H, 30 mm; I–M, 5 mm. Drawn by Juliet Beentje from Perrier de la Bâthie 8128, Hermans 374 (Kew Spirit Collection no. 63582). Reproduced with permission from Adansonia, sér. 3, 29, 29 (2007), Publications Scientifiques du Muséum, Paris.

11-arch. Vascular tissue embedded in sclerenchyma. Pith sclerenchymatous. (BC)

Palynology Pollen of Eurychone has not been examined. (AP)

Cytogenetics Brandham (1999) reported a chromosome count of 2n= 50 for the genus. (AP) 396

Phytochemistry The cinnamon-like floral scent of E. rothschildiana, which is emitted after sunset, contains benzaldehyde (25.0%), (E)-cinnamaldehyde (25.9%), 3-phenylpropan-1-ol (15.0%), and (E)-cinnamyl alcohol (8.0%) as the major components (Kaiser 2011). (NV, RG)

Phylogenetics In the molecular study by Carlsward et al. (2006b), Eurychone was strongly supported as monophyletic and related to Aerangis and the Diaphananthe–Tridactyle–Cyrtorchis group. (AP)

JUMELLEA

Derivation of name The generic name commemorates Professor Henri Jumelle (1866–1935), Professor of Botany and Director of the Botanic Garden in Marseille, France. (PC)

Description (Plate 160; Fig. 734.1)

Fig. 732.2.  Distribution map of Erasanthe.

Epiphytic or lithophytic herbs. Stems leafy, sometimes forming large clumps. Leaves distichous, borne in a fan or in apical part of stem, articulate to a sheathing base, rarely terete, bilobed at apex, articulated at base to distichous, sheathing leaf-bases. Inflorescence one-flowered, axillary; peduncle covered by sheathing bracts. Flowers white, stellate. Sepals and petals subsimilar, free, spreading to reflexed. Labellum entire, broader than other perianth parts, usually dilated above base, spurred at base, sometimes with a keel-like longitudinal callus at base; spur usually as long as or longer than labellum, rarely shorter, often geniculately bent near base. Column with two pollinia, globose or subglobose, stipes two, elliptic, each attached to its own viscidium, rarely attached to a common viscidium. Ovary with pedicel erect or erect-spreading, elongate, cylindrical. (PC)

Distribution (Fig. 734.2) Ecology Eurychone rothschildiana grows epiphytically on trunks of large trees in wet forests and also in forests on granite outcrops and coffee plantations from sea level to 1200 m. Eurychone galeandrae (Rchb.f.) Schltr. grows epiphytically on savanna trees. (PC)

Pollination The pollination biology of this genus is unknown but likely sphingophilous given the coloration, presence of a spur, and floral fragrance. (AP)

Uses There are no known uses for either of these species. (AP)

Taxonomic notes Eurychone is a distinctive genus characterized by its large flowers with a broadly conical, ecallose, entire labellum with a conical spur, two cleft pollinia attached to an obovate tegula and a single viscidium, and a peg-like, pendent rostellum. (PC)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86.

734.  J UME LLE A Jumellea Schltr., Orchideen, 609 (1914). Type species: Jumellea fragrans (Thouars) Schltr. (basionym: Angraecum fragrans Thouars)

Jumellea is a genus of 57 species in Madagascar, the Comores, and Mascarenes, with two species in tropical Africa (Rakotoarivelo et al. 2012). (PC)

Anatomy Carlsward et al. (2006a) examined seven species of Jumellea. Their results are summarized below.

Leaf TRANSVERSE SECTION

Trichomes multicellular, glandular; absent in J. confusa (Schltr.) Schltr. Stomata abaxial. Hypodermis ad- and abaxial, composed of fibrous idioblasts in a single incomplete row scattered among chlorenchyma cells. Fibre bundles absent. Mesophyll homogeneous in J. arachnantha (Rchb.f.) Schltr. and J. phalaenophora (Rchb.f.) Schltr.; heterogeneous with columnar adaxial cells and isodiametric abaxial cells in J. arborescens H.Perrier, J. confusa, and J. flavescens H.Perrier; isobilateral with columnar cells surrounding a central region of isodiametric cells in J. filicornoides (De Wild) Schltr. (=J. walleri (Rolfe) la Croix). Water-storage cells with smooth to pitted walls. Fibrous idioblasts scattered throughout mesophyll in J. arachnantha, J. arborescens, J. flavescens, J. phalaenophora, and J. walleri. Vascular bundle sheaths cells distinct, walls occasionally slightly lignified in J. arachnantha, J. flavescens, and J. phalaenophora. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Velamen two or three cells thick. Epivelamen cells isodiametric to radially elongate; walls ∪-thickened with fine, anastomosing, radial bands. Endovelamen cells angular, isodiametric to radially 397

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C Fig. 733.1.  Eurychone galeandrae (Rchb.f.) Schltr. A. Habit; B. Flower; C. Bract; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum; H. Column and labellum, side view; I. Column, front view; J. Column, side view; K. Column apex, anther cap removed, front view; L. Anther cap; M. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Redrawn by Judi Stone after Curtis’s Botanical Magazine, 18, 159 (2001).

elongate; cells of the outer layers often thicker-walled than those of the inner layers. Distinct endovelamen thickenings ridged. Cover cells present over short cells of exodermis. Exodermal long cells ∩-thickened or ○-thickened. Water-storage cell walls smooth to pitted or banded. Single modified cortical layer of thinto thick-walled cells surrounding the endodermis in J. arachnantha, J. arborescens, and J. phalaenophora. Aeration units present in all species except J. flavescens. Endodermal cells usually ○-thickened to infrequently ∪-thickened. Vascular cylinder 13–38arch. Vascular tissue embedded in sclerenchyma; cell walls of embedding tissue thickest over phloem groups in J. arachnantha, 398

J. confusa, J. flavescens, and J. phalaenophora. Pith sclerenchymatous. (BC)

Palynology Schill and Pfeiffer (1977) surveyed the pollen of 13 species of Jumellea and described the tetrads as convex with laevigate sculpturing and a calymmate sexine about 0.5 μm thick. (AP)

Cytogenetics Brandham (1999) cited chromosome numbers of 2n = 38, 38–40 for Jumellea. (AP)

LEMURELLA

al. 2008), Jumellea was strongly supported as sister to Aeranthes. More recently, Rakotoarivelo et al. (2012) performed phylogenetic analyses of Jumellea based on combined plastid (matK, trnL-F, rps16, ycf1) and nuclear (nrITS) data from 60 specimens representing 47 species, four subspecies, and two varieties of Jumellea. In addition to raising the number of species to 57, they found no support for the informal groups of Jumellea, as all the characters tested are evolutionarily labile. (AP)

Ecology Species of Jumellea are epiphytic in all types of wet and semideciduous woodland and forests from sea level to 2000 m. Jumellea papagensis H.Perrier is found in coastal forests, but most species are found above 500 m. A number of species, including A. brevifolia H.Perrier, A. densifoliata Senghas, A. gregariiflora H.Perrier, J. ibityana Schltr., J. lignosa (Schltr.) Schltr., J. pandurata Schltr., J. rigida Schltr., and J. spathulata (Ridl.) Schltr. are found on granite and gneiss outcrops, especially on the Madagascan plateau. (PC) Fig. 733.2.  Distribution map of Eurychone.

Pollination

Phytochemistry

Nilsson et al. (1987) showed that J. teretifolia Schltr. was pollinated by the hawkmoth Panogena lingens with viscidia attached dorsally to the proboscis. (AP)

The leaves of J. fragrans contain considerable amounts of coumarin and can be used to prepare an infusion known as ‘Faham tea’. According to 19th-century sources, this was consumed by the inhabitants of Mauritius and Réunion in preference to China tea; they also add it locally distilled rum, which is known as rhum arrangé. The dried leaves of J. fragrans (cited as Angraecum fragrans Thouars in older literature) were also imported from these islands to France for use as a beverage and a flavouring for custards and ices (Gobley 1850a,b; Jackson 1866). Interest in Faham tea as a commodity prompted several early investigations on the chemical constituents of the leaves, resulting in the isolation of coumarin and some of its derivatives (Gobley 1850a,b; Zwenger 1872; Trillich 1899). Hegnauer (1963) has commented on problems relating to the identity of the coumarin derivatives obtained by Zwenger (1872), and their relationship to similar substances from Melilotus officinalis (Fabaceae). Analysis of the volatile components of the dried leaves of J. fragrans using GC-MS (gas chromatography coupled to mass spectrometry) revealed a content of free coumarin of 85% (Sing et al. 1992). Almost 100 minor components were also identified. Little is known about other aspects of the phytochemistry of Jumellea. Of two species tested for the presence of alkaloids, neither contained them at levels ≥ 0.1% (Lüning 1974); however, they were present at lower levels (≈ 0.01%) in J. rigida (Lüning 1967). Some information on historic and contemporary medicinal usage of J. fragrans on Mauritius and Réunion is available (Véra et al. 1990). (NV, RG)

Uses Apart from the use of Jumellea fragrans to prepare a beverage like tea (see Phytochemistry above) and to flavour rum, several Jumellea species are occasionally cultivated in specialist collections. (AP, FR)

Taxonomic notes Schlechter (1914) established Jumellea as a segregate genus of Angraecum distinguished by the position of sepals and petals, the labellum that is narrow at the base and often clawed but which never enfolds the column, and the filiform spur with a narrower mouth. (PC)

Taxonomic literature Micheneau, C., Carlsward, B. S., Fay, M. F., Bytebier, B., Pailler, T., and Chase, M. W. (2008). Phylogenetics and biogeography of Mascarene angraecoid orchids (Vandeae, Orchidaceae. Molecular Phylogenetics and Evolution, 46, 908–22. Rakotoarivelo, F. P., Razafimandimbison, S. G., Mallet, B., Faliniaina, L., and Pailler, T. (2012). Molecular systematics and evolutionary trends and relationships in the genus Jumellea (Orchidaceae): implications for its species limits. Taxon, 61, 534–44. Schlechter, R. (1914). Die Orchideen, 2nd ed. Paul Parey, Berlin.

Phylogenetics

7 3 5 .  L EM U R EL L A

In a comprehensive phylogenetic study of Mascarene angraecoid orchids involving four plastid gene regions (Micheneau et

Lemurella Schltr., Repert. Spec. Nov. Regni Veg. Beih., 33, 366 (1925). Type species: Lemurella ambongensis (Schltr.) Schltr.; 399

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Fig. 734.1.  Jumellea walleri (Rolfe) la Croix. A. Habit; B. Dorsal sepal; C. Lateral sepal; D. Petal; E. Labellum; F. Column and labellum, side view; G. Column, side view; H. Column, front view; I. Column, back view; J. Column, anther cap removed, side view; K. Anther cap; L. Anther cap, pollinaria removed; M. Pollinaria. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Luke & Robertson 1826 (K) and Kew Spirit Collection no. 54560. Reproduced with permission from G. V. Pope (ed.) (1998), Flora Zambesiaca: Mozambique, Malawi, Zambia, Zimbabwe, Botswana. Volume 11, Part 2. Royal Botanic Gardens, Kew.

400

LEMURELLA

Leaf TRANSVERSE SECTION

Trichomes rare; multicellular, glandular. Stomata abaxial; superficial to slightly raised. Hypodermis absent. Fibre bundles absent. Mesophyll heterogeneous with columnar adaxial cells and isodiametric abaxial cells. Water-storage cells smooth to pitted. Red ergastic substance (possibly tannins) scattered in epidermal cells and present in mesophyll cells adjacent to epidermal surfaces. Vascular bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Fig. 734.2.  Distribution map of Jumellea.

basionym: Angraecum ambongense Schltr. (=Lemurella culicifera (Rchb.f.) H.Perrier)

Derivation of name The generic name is derived from the Latin lemur, ghost (given to a group of endemic primates in Madagascar), and -ella, a diminutive. (PC)

Description (Plate 161; Fig. 735.1) Epiphytic herbs. Stem covered by compressed leaf bases. Leaves oblong to falcate-oblong, acute to obtuse at unequally bilobed tip. Inflorescence axillary, one- to several-flowered, simple; peduncle much longer than rachis; bracts ovate. Flowers resupinate, green to pale yellow-green. Sepals and petals free, spreading to reflexed at tip, acute. Labellum trilobed in basal half, ecallose, spurred at base, lateral lobes erect, enclosing column, disc sometimes shortly pubescent; spur cylindrical-filiform from a broad infundibuliform mouth, straight or rising above ovary. Column with a subquadrate anther, pollinia two, porate, inserted directly on separate narrowly elliptic or linear viscidia; rostellum trilobed, lateral lobes broad, auriculate, the midlobe more slender, linear, as long as lateral lobes. (PC)

Velamen two cells thick. Epivelamen cells tangentially elongate and scalloped; walls ∪-thickened with fine, anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate. Distinct endovelamen thickenings absent. Cover cells present over short cells of exodermis. Exodermal long cells ∩-thickened. Water-storage cell walls smooth to pitted. Aeration units present. Endodermal cells strongly ○-thickened. Vascular cylinder 8-arch. Vascular tissue embedded in thick-walled sclerenchyma. Pith sclerenchymatous. (BC)

Palynology Schill and Pfeiffer (1977) described the pollen of L. ambongoensis (= L. culicifera) as laevigate to weakly foveolate with a calymmate sexine. (AP)

Cytogenetics There are no reports on chromosome numbers for Lemurella. (AP)

Phytochemistry No reports have been found on the phytochemistry of Lemurella. (NV, RG)

Phylogenetics Carlsward et al. (2006b) included L. pallidiflora in their ITS rDNA study and showed that it is sister to the clade including Cryptopus, Oeonia, and Neobathiea, with Beclardia sister to all of them, although there is no bootstrap support for those sister relationships. (AP)

Ecology

Lemurella is a genus of four species endemic to Madagascar. (PC)

Most species are found in mossy forests between 800 and 1200 m elevation. Lemurella culicifera is, in contrast, found growing as a trunk epiphyte in seasonally dry deciduous forest and woodland from sea level to 700 m. (PC)

Anatomy

Pollination

Carlsward et al. (2006a) examined L. pallidiflora Bosser. Their results are summarized below.

There is no published information on pollination in this genus. (AP)

Distribution (Fig. 735.2)

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K A G H Fig. 735.1.  Lemurella culicifera (Rchb.f.) H.Perrier. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum; H. Column and labellum, side view; I. Column, front view; J. Column, side view; K. Column from above, anther cap removed; L. Anther cap, dorsal view; M. Anther cap, ventral view; N. Pollinaria; O. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Roberts et al. TPP075 and Kew Spirit Collection no. 60967.

Uses

Taxonomic literature

There are no known ethnobotanical uses for the four species, and they are not commonly cultivated. (AP)

Bosser, J. (1970). Contribution a l’étude des orchidées de Madagascar. XIV. Adansonia, ser. 2, 10, 367–73. Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86.

Taxonomic notes Schlechter established Lemurella as a segregate of Angraecum based on his Angraecum ambongense. Bosser (1970) revised the genus and reduced the type species to the synonymy of L. culicifera. Lemurella species resemble some of the green-flowered Angraecum species but are distinguished by their trilobed labellum and trilobed rostellum. (PC) 402

7 3 6 .  L EM U RORC H I S Lemurorchis Kränzl., Bot. Jahrb. Syst., 17, 58 (1893). Type species: Lemurorchis madagascariensis Kränzl.

LEMURORCHIS

Root TRANSVERSE SECTION

Velamen frequently absent or two cells thick. Epivelamen cells isodiametric; walls ∪-thickened with fine anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate. Distinct endovelamen thickenings ridged. Cover cells present over short cells of exodermis. Exodermal long cells ○-thickened. Water-storage cell walls smooth to pitted. Aeration units absent. Endodermal cells ∪- to ○-thickened. Vascular cylinder 18-arch. Vascular tissue embedded in sclerenchyma. Pith sclerenchymatous. (BC)

Palynology Pollen of this species has not been studied. (AP)

Cytogenetics Chromosome counts of L. madagascariensis have not been published. (AP) Fig. 735.2.  Distribution map of Lemurella.

Derivation of name The generic name is derived from the Latin lemur, ghost (given to a group of endemic primates in Madagascar), and orchis (orchid). (PC)

Description (Plate 162; Fig. 736.1) Epiphytic herbs, often forming clumps. Stem indistinct or absent. Leaves distichous, arranged in a fan, linear-ligulate. Inflorescence axillary, unbranched, cylindric, many-flowered. Flowers yellowish. Sepals free. Petals similar to dorsal sepal. Labellum enclosing column, obscurely trilobed in front; spur cylindrical, longer than labellum. Column lacking a foot; pollinia two, each with a rigid stipe and indistinct viscidium; rostellum bifid. (PC)

Distribution (Fig. 736.2) Lemurorchis is a monospecific genus endemic to Madagascar. (PC)

Anatomy Carlsward et al. (2006a) examined L. madagascariensis. Their results are summarized below.

Leaf TRANSVERSE SECTION

Trichomes multicellular, glandular. Stomata abaxial. Hypodermis absent. Fibre bundles absent. Mesophyll heterogeneous with greater numbers of large adaxial water-storage cells and fewer small abaxial assimilatory cells. Water-storage cells with banded thickenings. Vascular bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies primarily associated with vascular bundle sclerenchyma.

Phytochemistry No reports have been found on the phytochemistry of Lemurorchis. (NV, RG)

Phylogenetics Nothing is known of the phylogenetic relationships of this species. (AP)

Ecology The lone species occurs in shady, mossy forests at 2200 m elevation. (PC)

Pollination The pollination biology of this species has not been investigated. (AP)

Uses There are no known uses for L. madagascariensis. (AP)

Taxonomic notes Lemurorchis is a distinctive angraecoid orchid with a short stem; elongate, linear, distichous leaves canaliculate at their imbricate bases; and an inflorescence with an erect peduncle but pendent rachis of many densely clustered flowers in a cylindrical raceme. The flowers resemble those of Microterangis but have a trilobed labellum with large, erect lateral lobes and a bifid rostellum. (PC)

Taxonomic literature Perrier de la Bâthie, H. (1941). Flore de Madagascar. Orchidées, II (ed. H. Humbert). Tananarive Imprimerie Officielle, Madagascar.

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Fig. 736.1.  Lemurorchis madagascariensis Kränzl. A. Habit; B. Flower; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum; G. Column, front view; H. Column, side view; I. Column, longitudinal section; J. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Redrawn by Judi Stone after Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie, 17, t. 3 (1893).

737.  LIS TRO STAC H Y S

Description (Plate 163; Fig. 737.1)

Listrostachys Rchb.f., Bot. Zeitung (Berlin), 10, 930 (1852). Type species: Listrostachys jenischiana Rchb.f. (=Listrostachys pertusa (Lindl.) Rchb.f.)

Epiphytic herbs. Roots toward base. Stem leafy, covered by distichous leaf-bases. Leaves coriaceous, linear, unequally bilobed at tip. Inflorescence suberect, elongate, many-flowered; rachis with flowers arranged distichously in two rows with labellum and spur pointing outward; bracts sheathing rachis. Flowers resupinate, sparsely hairy, white sometimes with minute red spots toward base and with a red spur. Dorsal sepal erect to reflexed, spatulateobovate, apiculate; lateral sepals recurved, ovate, keeled on back. Petals obliquely triangular. Labellum oblong, ecallose, spurred at

Derivation of name From the Greek listron, shovel, and stachyos, spike, in reference to the many-flowered inflorescences in which the dorsal sepal is shovel-shaped. (PC) 404

MARGELLIANTHA

thickenings absent. Cover cells absent. Exodermal long cells primarily ○-thickened. Water-storage cell walls smooth to pitted. Aerations units present. Endodermal cells strongly ○-thickened. Vascular cylinder 7–17-arch. Vascular tissue embedded in sclerenchyma. Pith sclerenchymatous. (BC)

Palynology Schill and Pfeiffer (1977) described the pollen of L. pertusa as having laevigate sculpturing with a calymmate sexine. (AP)

Cytogenetics Arends et al. (1980) reported a chromosome count of 2n = 46 for L. pertusa. (AP)

Phytochemistry No reports have been found on the phytochemistry of Listrostachys. (NV, RG)

Phylogenetics Fig. 736.2.  Distribution map of Lemurorchis.

base; spur clavate. Column upcurved, distally dilated; anther semicircular, pollinia two, slightly cleft, each attached by a broadly spatulate stipe to a common, reniform viscidium; stigma subcircular. (PC)

Distribution (Fig. 737.2) Listrostachys is a monospecific genus in tropical Africa from Sierra Leone to Gabon and Zaire. (PC)

Anatomy Carlsward et al. (2006a) examined L. pertusa. Their results are summarized below.

Leaf TRANSVERSE SECTION

Trichomes multicellular, glandular. Stomata abaxial. Hypodermis ad- and abaxial, composed of fibrous idioblasts occurring singly or in groups of two or three, forming a single row with interspersed chlorenchyma. Fibre bundles absent. Mesophyll heterogeneous with columnar adaxial cells and isodiametric abaxial cells. Water-storage cells with birefringent walls rare. Solitary fibrous idioblasts concentrated in abaxial mesophyll, generally thinner-walled than hypodermal idioblasts. Vascular bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Velamen three or four cells thick. Epivelamen cells primarily isodiametric to infrequently radially elongate; walls ∪-thickened with fine anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate. Distinct endovelamen

Carlsward et al. (2006b) included L. pertusa in their ITS rDNA analysis. It was sister to Tridactyle, but there was no bootstrap support greater than 50% for the relationship. (AP)

Ecology The species occurs in shady places on large branches of large evergreen trees such as Lophira alata Banks ex C.F.Gaertn. (Ochnaceae) and Coula edulis Baill. (Olacaceae) in humid forests from sea level to 200 m. (PC)

Pollination The pollination biology of L. pertusa is unknown. (AP)

Uses There are no known ethnobotanical uses of the species, and it is not in common cultivation. (AP)

Taxonomic notes Listrostachys is now considered to be monospecific but has at various times been expanded to include species in diverse African angraecoid genera, including Ancistrorhynchus, Angraecopsis, Bolusiella, Chamaeangis, Cyrtorchis, Diaphananthe, Plectrelminthus, Podangis, and Tridactyle. (PC)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86.

7 3 8 .  M A RGEL L I A NT H A Margelliantha P.J.Cribb, Kew Bull., 34, 329 (1979). Type species: Margelliantha leedalii P.J.Cribb. 405

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L A Fig. 737.1.  Listrostachys pertusa (Lindl.) Rchb.f. A. Habit; B. Portion of inflorescence; C. Flower, front view; D. Flower, side view; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum; I. Column and labellum, side view; J. Column, front view; K. Column, side view; L. Column, anther cap removed, back view; M. Anther cap with pollinarium, dorsal view; N. Anther cap with pollinarium, ventral view; O. Anther cap, pollinarium removed, ventral view; P. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from van der Burgt & Loan 176 (K) and Kew Spirit Collection no. 47930.

Derivation of name From the Latin margelis, pearl, and the Greek anthos, flower, in reference to the pearl-like flowers of the type species. (PC)

Description (Plate 164; Fig. 738.1) Epiphytic herbs. Stem covered in persistent leaf bases. Leaves distichous, coriaceous, falcate, linear or narrowly oblanceolate, unequally bilobed at apex, twisted at base to lie in one plane, articulated at base to a persistent sheathing leaf base. Inflorescence 406

spreading, racemose, few-flowered, axillary, emerging through leaf bases; peduncle and rachis terete, wiry. Flowers campanulate, white to yellow or pale green. Sepals and petals free; dorsal sepal concave, lateral sepals falcate, concave. Petals elliptic or subcircular. Labellum concave, entire, lacking a callus, spurred at base; spur clavate to globose. Column sometimes with an obscure foot; pollinia two, globose, stipites two, linear, viscidia two, circular or elliptic; stigma concave, with a raised posterior margin; rostellum trilobed, midlobe longer than lateral lobes. (PC)

MICROCOELIA

Uses No ethnobotanical uses for the species have been recorded, and they are not in common cultivation. (AP)

Taxonomic notes Margelliantha was established as a segregate genus of Diaphananthe, distinguished by its pearly white flowers with a characteristic green anther cap, concave and labellate labellum, and clavate spur with a narrow mouth (Cribb 1979). The South African Margelliantha caffra (Mystacidium caffra Bolus) and the similar Mystacidium millarii Bolus were transferred to Diaphananthe by Linder (1989). Carlsward et al. (2006b) included it as Diaphananthe millarii (Bolus) H.P.Linder in their ITS rDNA study, in which the species fell in a clade with Angraecopsis, Mystacidium, Cribbia, and Rhipidoglossum. Rice (2005, 2006b) included Margelliantha, along with Angraecopsis and Rhipidoglossum, in Mystacidium based on suggested similarities of their pollinaria and rostellum. (PC)

Taxonomic literature Fig. 737.2.  Distribution map of Listrostachys.

Distribution (Fig. 738.2) Margelliantha is a genus of seven species in tropical and southern Africa. Three species have been recorded from tropical East Africa. (PC)

Anatomy Carlsward et al. (2006a) studied the vegetative anatomy of M. millarii (Bolus) P.J.Cribb (see below) as Diaphananthe millarii (q.v.) (AP)

Palynology There is no published information on the pollen of these species. (AP)

Cytogenetics Chromosome counts for Margelliantha have not been reported. (AP)

Phytochemistry No reports have been found on the phytochemistry of Margelliantha. (NV, RG)

Ecology Species are epiphytic on branches and twigs mostly in forests from 200 to 2000 m, but M. millarii is found in coastal forests. (PC)

Pollination Pollination biology of the genus is unknown. (AP)

Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Cribb, P. J. (1979). New or little known orchids from East Africa. Kew Bulletin, 34, 321-40. Micheneau, C., Carlsward, B. S., Fay, M. F., Bytebier, B., Pailler, T., and Chase, M. W. (2008). Phylogenetics and biogeography of Mascarene angraecoid orchids. Molecular Phylogenetics and Evolution, 46, 908–22. Linder, H. P. (1989). Notes on southern African angraecoid orchids. Kew Bulletin, 44, 318. Rice, R. (2005). A preliminary checklist and survey of the subtribe Aerangidinae (Orchidaceae). Oasis Publications, Dora Creek, New South Wales, Australia. Rice, R. (2006b). Notes on the subtribe Aerangidinae (Orchidaceae). Oasis, 3, 12–15.

7 3 9 .  M I C ROC OEL I A Microcoelia Lindl., Gen. Spec. Orch. Pl., 50 (1830). Type species: Microcoelia exilis Lindl. Gussonea A.Rich., Mém. Soc. Hist. Nat. Paris, 4, 67, t. 11 (1828). Type species: Gussonea aphylla (Thouars) A.Rich. (basionym: Angraecum aphyllum Thouars). Obs: not Gussonea J.Presl & C.Presl (July 1828), Cyperaceae. Dicranotaenia Finet, Bull. Soc. Bot. France, 54, Mém. 9, 47 (1907). Type species: Dicranotaenia dahomeensis Finet Encheiridion Summerh., Bot. Mus. Leafl., Harvard Univ., 11, 161 (1943). Type species: Encheiridion macrorrhynchium (Schltr.) Summerh. (basionym: Angraecum macrorrhynchium Schltr.)

Derivation of name From the Greek, mikros, small, and koilos, hollow, in allusion to the globular spur of the type species. (PC)

Description (Plates 165, 166; Figs. 739.1, 752.1) Often leafless and epiphytic or rarely lithophytic herbs. Roots often dense, terete or less commonly dorsiventrally flattened, 407

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Fig. 738.1.  Margelliantha leedalii P.J.Cribb. A. Habit, ×1; B. Bract, ×6; C. Flower, ×6; D. Dorsal sepal, ×8; E. Lateral sepal ×8; F. Petal, ×8; G. Labellum, ×8; H. Column, side view, ×24; I. Column, front view, ×24; J. Anther cap, ×20; K. Pollinarium, ×20. Magnifications as originally published. Drawn by Maureen E. Church from Archer 548 (K). Reproduced with permission from P. J. Cribb (1989), Flora of Tropical East Africa, Orchidaceae (Part 3). A.A. Balkema, Rotterdam/Brookfield.

408

MICROCOELIA

except M. aphylla (Thouars) Summerh. Exodermal long cells ∩-thickened. Microcoelia aphylla, M. caespitosa (Rolfe) Summerh., M. exilis, and M. megalorrhiza (Rchb.f.) Summerh. possess exodermal radial wall swellings; M. macrantha (H.Perrier) Summerh., M. obovata Summerh., and M. physophora (Rchb.f.) Summerh. possess exodermal radial wall grooves. Water-storage cell walls smooth to pitted, banded, or thickened along the edges. Single layer of thin- to thick-walled cells surrounding the endodermis in M. exilis, M. obovata, M. perrieri (Finet) Summerh., and M. smithii (Rolfe) Summerh. Aeration units present in all species examined. Endodermal cells primarily ○-thickened; thin-walled only in M. bulbocalcarata L.Jonss., M. corallina Summerh., and M. stolzii (Schltr.) Summerh. Vascular cylinder 3–15-arch. Vascular tissue embedded in sclerenchyma, cell walls of embedding sclerenchyma thickest around phloem clusters in M. globulosa (Hochst.) L.Jonss. and M. stolzii. Pith usually sclerenchymatous, except parenchymatous in M. corallina. (BC)

Palynology Fig. 738.2.  Distribution map of Margelliantha.

smooth or rarely verrucose, unbranched or with a few branches, usually elongate. Stem ‘woody’, terete, with acute to rostrate scale-leaves apically. Inflorescence axillary, racemose, concentrated in apical part of stem; rachis terete or angular, smooth or with processes; bracts sheathing or not. Flowers sessile or pedicellate, usually white variously tinged with green, brown or pink. Sepals and petals free, subsimilar. Labellum entire or obscurely trilobed, free, usually with fleshy calli at base on either side of spur mouth; spur globose, cylindrical or variously swollen. Column with a hemispherical anther cap, often elongated at apex, pollinia two, subglobose to pyriform, stipe (tegula) linear to oblanceolate, entire or bird-like at the column apex, viscidium linear or oblong; rostellum bifid after removal of pollinia, no longer than column. (PC)

Distribution (Fig. 739.2) Microcoelia comprises about 30 species in tropical and southern Africa, Madagascar (11 species), the Comores, and Mascarene islands. (PC)

Anatomy Carlsward et al. (2006a) examined 14 species of Microcoelia. Their results are summarized below.

Root TRANSVERSE SECTION

Velamen two or three cells thick. Epivelamen cells isodiametric to radially elongate; walls ∪-thickened with fine anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate. Distinct endovelamen thickenings absent. Cover cells present over short cells of the exodermis in all species

Schill and Pfeiffer (1977) studied the pollen of M. exilis, M. guyoniana (Rchb.f) Summerh. (= M. globulosa), an unidentified species, and Solenangis aphylla (Thouars) Summerh. (now M. aphylla (Thouars) Summerh.). Sculpturing is laevigate. The sexine is calymmate, wavy, and about 0.3 μm thick. In his elegant monograph of the genus, Jonsson (1981) examined pollinium ontogeny with electron microscopy; he identified specialized viscin-secreting cells that degenerate at anthesis, leaving a cavity through which viscin threads connect to the pollinium to the tegula. (AP)

Cytogenetics Jonsson (1981) and Arends and Van der Laan (1983) reported chromosome counts of 2n = 48 for Microcoelia; Brandham (1999) also listed 2n = 47. (AP)

Phytochemistry No reports have been found on the phytochemistry of Microcoelia. (NV, RG)

Phylogenetics Microcoelia is monophyletic with strong support if the former Solenangis aphylla (Thouars) Summerh. and S. cornuta (Rchb.f.) Summerh.—both of them leafless species—are included as M. aphylla (Thouars) Summerh. and M. cornuta (Ridl.) Carlsward (Carlsward et al. 2006b). Carlsward et al. (2006b) showed that it is related to Bolusiella and Ancistrorhynchus, although support for that relationship is low. (AP)

Ecology Species of Microcoelia are epiphytes, often twig epiphytes, less commonly on rocks, in sun or shade in bushland, thickets, savanna woodlands, and lowland, riverine, and semi-deciduous and evergreen forests from sea level to 2300 m. (PC) 409

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Fig. 739.1.  Microcoelia koehleri (Schltr.) Summerh. A. Habit, ×1; B. Flower, one lateral sepal and petal removed, side view, ×4; C. Dorsal sepal, ×6; D. Lateral sepal, ×6; E. Petal, ×6; F. Labellum, spread, ×6; G. Column, side view, ×12; H. Column, front view, ×12; I. Column, anther removed, side view, ×12; J. Column, anther removed, front view,. ×12; K. Anther, ventral view, ×12; L. Pollinarium, front view, ×12; M. Pollinarium, pollinia removed, side view, ×12. Magnifications as originally published. Drawn by Stella Ross-Craig. Reprinted from J. D. Hooker, Icones Plantarum.

Pollination Pollen vectors for Microcoelia are unknown, although Jonsson (1981) noted strong diurnal fragrances and nectariferous spurs in M. globulosa, M. stolzii, M. elliotii (Finet) Summerh., and M. gilpinae (Rchb.f. & S.Moore) Summerh. that are indicative of 410

butterfly-pollinated flowers. However, Jonsson (1981) also noted that other species such as M. koehleri (Schltr.) Summerh., have a nocturnal fragrance more suitable for moth pollination. (AP)

MYSTACIDIUM

Szlachetko, D. and Olszewski, S. (2001). Microcoelia Lindley. In Flore du Cameroun 36. Orchidacees, vol. 3 (ed. G. Achoundong and P. Morat), pp. 790–808. Ministere de la Recherche Scientifique et Technique, Yaounde, Cameroun.

7 4 0 .  M YS TAC I DI U M Mystacidium Lindl. in Hook. Comp. Bot. Mag., 2, 206 (1837). Type species: Mystacidium filicorne Lindl., nom. illeg. (=Mystacidium capense (L.f.) Schltr.; basionym: Epidendrum capense (L.f.) Schltr.)

Derivation of name From the Greek mystax, moustache, perhaps in reference to the papillose rostellum (Schultes and Pease 1963). (AP)

Description (Plate 167; Fig. 740.1)

There are no known uses for any of the species except as occasionally cultivated curiosities. (AP)

Epiphytic herbs. Stem covered by imbricate, sheathing leaf-bases. Leaves distichous, coriaceous, articulated to a sheathing leaf-base. Inflorescence axillary, spreading to pendent, two- to many-flowered; peduncle wiry. Flowers usually stellate, white, pale green, or pale yellow. Sepals and petals free, subequal, spreading or subconnivent. Labellum entire or trilobed, spurred at base, ecallose; spur tapering from a broad mouth. Column with two pollinia, globose, sulcate, stipes two, linear, viscidia two, rarely adherent, oblong or squamiform; rostellum pendent, trilobed, outer lobes often much longer than midlobe and papillose. (PC)

Taxonomic notes

Distribution (Fig. 740.2)

Fig. 739.2.  Distribution map of Microcoelia.

Uses

Summerhayes (1943) revised Microcoelia, recognizing 25 species in three sections: M. sect. (Eu)Microcoelia, M. sect. Brachyglossa, and M. sect. Dicranotaenia. He also established the genus Encheiridion, based upon Microcoelia macrorrhynchia (Schltr.) Summerh. as a distinct monospecific genus. The genus was later revised by Jonsson (1981) who placed Encheiridion back in Microcoelia. The genus is currently divided in two subgenera (Szlachetko and Olszewski 2001) as follows: M. subgenus Microcoelia M. subgenus Encheiridion (Summerh.) Szlach. & Olszewski, Fl. Cameroun, 36, 792 (2001). Type species: Microcoelia macrorrhynchia (Schltr.) Summerh. (basionym Angraecum macrorrhynchium Schltr.) The latter is distinguished by its flowers in which the labellum has basal, erect auricles or lateral lobes, the free part of the rostellum is longer than the gynostemium and tegula, and viscidium is also long and narrow. (PC)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Jonsson, L. (1981). A monograph of the genus Microcoelia (Orchidaceae). Symbolae Botanicae Upsaliensis, 23, 1–151. Summerhayes, V. S. (1943). African orchids XIII. The African leafless orchids. Botanical Museum Leaflets (Harvard University), 10, 137–72.

Mystacidium is a genus of 15 species mainly in southern, Central and East Africa as far north as Tanzania. (PC)

Anatomy Carlsward et al. (2006a) examined three species of Mystacidium. Their results are summarized below.

Leaf TRANSVERSE SECTION

Trichomes multicellular, glandular. Stomata abaxial; ad- and abaxial in M. flanaganii (Bolus) Bolus. Hypodermis absent. Fibre bundles absent. Mesophyll homogeneous; heterogeneous with columnar adaxial cells and isodiametric abaxial cells in M. braybonae Summerh. Water-storage cell walls smooth to pitted. Vascular bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Velamen 2–4 cells thick. Epivelamen cells radially elongate; walls ∪-thickened with fine anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate. Distinct endovelamen thickenings absent. Cover cells present over short cells of the exodermis. Exodermal long cells ○-thickened to infrequently 411

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R U Fig. 740.1.  A–J. Mystacidium gracile Harv. A. Habit; B. Flower; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum; G. Column and labellum, side view; H. Column, side view; I. Anther cap; J. Pollinarium. Drawn by Judi Stone from Greatrex GHS 29831 (K) and Kew Spirit Collection no. 51064; K–U. Mystacidium venosum Harv. ex Rolfe. K. Habit; L. Flower; M. Dorsal sepal; N. Lateral sepal; O. Petal; P. Labellum; Q. Column and labellum, side view; R. Column, side view; S. Anther cap; T. Pollinarium; U. Column apex showing rostellum. Drawn by Judi Stone from de Koning et al. 8789 (K) and Kew Spirit Collection no. 35108. Single bar = 1 mm, double bar = 1 cm. Reproduced with permission from G. V. Pope (ed.) (1998), Flora Zambesiaca: Mozambique, Malawi, Zambia, Zimbabwe, Botswana. Volume 11, Part 2. Royal Botanic Gardens, Kew.

412

NEOBATHIEA

Pollination The pollination biology of these species is unknown, although the syndrome of floral features would indicate butterflies and/ or moths as vectors. (AP)

Uses There are no recorded ethnobotanical uses of the species, but many are cultivated in specialist collections, usually mounted on slabs. (AP)

Taxonomic notes Mystacidium is characterized by its distinctive trilobed, papillose rostellum, in which the outer lobes are longer than the midlobe or of almost equal length. (PC)

Taxonomic literature

Fig. 740.2.  Distribution map of Mystacidium.

∩-thickened in M. braybonae. Water-storage cell walls smooth to pitted and often with edge thickenings. Aeration units present. Endodermal cells ○-thickened. Vascular cylinder 8–10-arch. Vascular tissue embedded in sclerenchyma. Pith sclerenchymatous. (BC)

Palynology Schill and Pfeiffer (1977) studied the pollen of M. capense and compared it to that of Angraecum with no further details. (AP)

Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86.

7 4 1 .  NEOBAT H I EA Neobathiea Schltr., Repert. Spec. Nov. Regni Veg. Beih., 33, 369 (1925). Type species: Neobathiea perrieri (Schltr.) Schltr. (basionym: Aeranthes perrieri Schltr.) Bathiea Schltr., Beih. Bot. Centralbl., 36, 104 (1918), non Drake (1902).

Derivation of name

Cytogenetics

From the Greek neos, new, and Bathiea, a genus of orchids named in honour of Henri Perrier de la Bâthie, the eminent French botanist. (PC)

Arends and Van der Laan (1983) and Brandham (1999) listed a chromosome count of 2n = 48 for the genus. (AP)

Description (Plate 168; Fig. 741.1)

Phytochemistry No reports have been found on the phytochemistry of Mystacidium. (NV, RG)

Phylogenetics Carlsward et al. (2006b) included four species of Mystacidium in their ITS rDNA study. It has strong bootstrap support as sister to Angraecopsis. (AP)

Ecology Species are epiphytes in evergreen forests and woodlands. The East African species are found in forests and woods from 1300 up to 2500 m elevation. Mystacidium tanganyikense Summerh. has been found in flower growing on the cones of Cupressus L. (Cupressaceae) and also on the native Juniperus procera (Hochst.) ex Endl. (Cupressaceae). (PC)

Epiphytes or rarely lithophytic herbs. Stem with few leaves in a fan or rarely elongate and leafy in apical half. Leaves obovate to oblanceolate, often twisted at base to lie in one plane. Inflorescence one- to few-flowered. Flowers white or pale green with a white labellum. Sepals and petals subsimilar, spreading or reflexed. Labellum entire to trilobed, spurred at base; spur filiform with a wide mouth. Column biauriculate at apex, each auricle attached to a linear viscidium; pollinia two, viscidia two; rostellum trilobed, deeply emarginate with a median linear ligule. (PC)

Distribution (Fig. 741.2) Neobathiea is a genus of five species endemic to Madagascar and the Comoros. (PC)

Anatomy Carlsward et al. (2006a) examined N. grandidierana. Their results are summarized below. 413

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Fig. 741.1.  Neobathiea perrieri (Schltr.) Schltr. A. Habit; B. Flower, front view; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum and portion of spur; G. Column and labellum, side view; H. Column, front view; I. Column, side view; J. Column from above, anther cap removed; K. Anther cap, dorsal view; L. Anther cap, ventral view; M. Pollinarium; N. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Humbert 32.684 (Plantes de Madagascar) and Kew Spirit Collection no. 60809.

Leaf TRANSVERSE SECTION

Trichomes absent. Stomata abaxial. Hypodermis absent. Fibre bundles absent. Mesophyll homogeneous. Water-storage cells smooth to pitted. Vascular bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies associated with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Velamen two cells thick. Epivelamen cells isodiametric; walls ∪-thickened with fine anastomosing radial bands. Endovelamen 414

cells angular, isodiametric to radially elongate. Distinct endovelamen thickenings absent. Cover cells present over short cells of exodermis. Exodermal long cells ∩-thickened. Water-storage cell walls smooth to pitted. Aeration units present. Endodermal cells ○-thickened. Vascular cylinder 5–8-arch. Vascular tissue embedded in sclerenchyma. Pith sclerenchymatous. (BC)

Palynology Pollen of Neobathiea has not been studied. (AP)

Cytogenetics There are no reported chromosome counts for the genus. (AP)

NEPHRANGIS

of that of Oeonia. Like Aeranthes it has a distinctive, flat labellum, and it is markedly separated or segmented at the free spurmouth. Unfortunately, the generic name Bathiea had previously been used for a genus in a different family. (PC)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Schlechter, R. (1918). Versuch einer natürlichen Neuordnung der Afrikanischen angraekoiden Orchidaceen. Beihefte zum Botanischen Centralblatt, 36, 62–181.

7 4 2 .  NEPH R A NGI S Nephrangis (Schltr.) Summerh., Kew Bull., 2, 301 (1948). Type species: Nephrangis filiformis (Kränzl.) Summerh. (basionym: Listrostachys filiformis Kränzl.) Fig. 741.2.  Distribution map of Neobathiea.

Phytochemistry No reports have been found on the phytochemistry of Neobathiea. (NV, RG)

Phylogenetics Carlsward et al. (2006b) included N. grandidieriana (Rchb.f.) Garay in their ITS rDNA study, in which it was sister to Cryptopus/ Oeonia. (AP)

Ecology Neobathiea grandidieriana and N. keraudrenae Toill.-Gen. & Bosser are trunk or branch epiphytes in forests from 1000–1700 m elevation. Neobathiea hirtula H.Perrier, N. perrieri (Schltr.) Schltr., and N. spatulata H.Perrier are found in semi-deciduous and deciduous western forests in Madagascar up to 350 m elevation; the last is also found occasionally on limestone rocks. (PC)

Pollination Nilsson et al. (1987) found pollinia of N. grandidieriana on the hawkmoth Panogena lingens. (AP)

Derivation of name From the Greek nephros, kidney, and angos, vessel, in reference to the kidney-shaped labellum of the type species. (PC)

Description (Plate 169; Fig. 742.1) Pendent, epiphytic herbs. Roots mostly toward stem base. Stem simple or branched, leafy, covered by sheathing leaf bases. Leaves distichous, coriaceous, terete, linear, aculiform, acute. Inflorescence lateral, shorter than leaves, racemose, few-flowered. Flowers semitranslucent, sepals and petals ochre-brown to brown-green, labellum white. Sepals and petals spreading, free, subsimilar. Labellum deflexed, shortly clawed then abruptly dilated into a tranversely reniform lamina, ecallose; spur cylindrical. Column lacking a foot; anther broadly obovoid, pollinia two, cleft, each attached by a linear or Y-shaped tegula to a single oblong to cordate viscidium; stigma obovate or triangular; rostellum bifid, curving forward at tip. Ovary with pedicel shorter than spur, pustular. (PC)

Distribution (Fig. 742.2) Nephrangis is a genus of two species in tropical Africa from Liberia eastward to Uganda and Tanzania and south to Zambia. (PC)

Anatomy

Uses

Nothing is known of the vegetative anatomy of Nephrangis. (AP)

There are no known ethnobotanical uses for these species, and they are not commonly cultivated. (AP)

Palynology Pollen of these two species has not been studied. (AP)

Taxonomic notes Schlechter (1918) established the genus Bathiea to accommodate Aeranthes perrieri Schltr.; he considered it to be aberrant in Aeranthes because of its column structure, which reminded him

Cytogenetics Chromosome counts have not been published for the genus. (AP) 415

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Fig. 742.1.  Nephrangis filiformis (Kränzl.) Summerh. A. Habit; B. Flower; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Column and labellum; G. Column, side view; H. Anther cap; I. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Cherry Ann Lavrih from Pfennig 1287 (K). Reproduced with permission from P. J. Cribb (1989), Flora of Tropical East Africa, Orchidaceae (Part 3). A.A. Balkema, Rotterdam/Brookfield.

Phytochemistry

Ecology

No reports have been found on the phytochemistry of Nephrangis. (NV, RG)

Nephrangis filiformis (Kränzl.) Summerh. is a canopy epiphyte on Piptadeniastrum africanum (Hook.f.) Brenan (Fabaceae) in moist semi-deciduous forests and on Cryptosepalum pseudotaxus Baker f. (Fabaceae) in woodlands on sand in riverine forest and on Uapaca Baill. (Euphorbiaceae) trees in savanna woodlands. Nephrangis bertauxinana Szlach. & Olszewski is found in woodlands on the

Phylogenetics Nothing is known of the phylogenetic relationships of this genus. (AP) 416

OEONIA

7 4 3 .  OEONI A Oeonia Lindl. in Edwards’s Bot. Reg., sub t. 817 (1824). Type species: Oeonia aubertii Lindl. Epidorkis Thouars, Nouv. Bull. Sci. Soc. Philom. Paris, 19, 318 (1809), nom. invalid. Type species: not designated Perrieriella Schltr., Repert. Spec. Nov. Regni Veg. Beih., 33, 365 (1925). Type species: Perrieriella madagascariensis Schltr.

Derivation of name From the Greek oionos, bird of prey, the column of the flower reminding the author of a bird of prey. (PC)

Description (Plate 170; Fig. 743.1)

Fig. 742.2.  Distribution map of Nephrangis.

edges of savannas. Both range from 500 to 2200 m in elevation. (PC)

Pollination

Scrambling or epiphytic herbs. Stem flexuose, climbing or branching. Leaves distichous, oblong-elliptic. Inflorescence axillary, one- to few-flowered. Flowers predominantly white or with yellowish or yellow-green, sometimes with red markings on the labellum. Sepals and petals obovate. Labellum three- or five-lobed, lateral lobes erect or enclosing column, midlobe deeply emarginate, spurred but lacking a callus. Column enclosed by infolded lateral lobes of labellum; pollinia two, waxy, cleft, tegulae elliptic, obscurely papillose, viscidia two, almost as large as tegulae; rostellum deeply emarginate with a peg-like central lobe. (PC)

Distribution (Fig. 743.2)

The pollination biology of Nephrangis has not been studied. (AP)

Oeonia is a genus of five species endemic to Madagascar and the Mascarenes. (PC)

Uses

Anatomy

There are no known uses for these species. (AP)

Carlsward et al. (2006a) examined O. rosea. Their results are summarized below.

Taxonomic notes

Leaf

Kränzlin originally described the type species as Listrostachys filiformis. Schlechter (1918) transferred it to Tridactyle in its own subgenus Nephrangis, distinguished by its distinctive, transversely reniform labellum and broad mouth to the spur. Summerhayes (1948) raised it to generic rank as Nephrangis, allied to Tridactyle, distinguished by the above features and also its reflexed, clavate rostellum, cleft (rather than porate) pollinia, the two tapering tegulae, and cordate viscidium (Szlachetko et al. 2004). Like some Tridactyle species, it has terete leaves and few-flowered racemes. (PC)

Taxonomic literature Schlechter, R. (1918). Versuch einer natürlichen Neuordnung der Afrikanischen angraekoiden Orchidaceen. Beihefte zum Botanischen Centralblatt, 36, 62–181. Summerhayes, V. S. (1948). African orchids XVIII. Kew Bulletin, 1947, 301. Szlachetko, D., Sawicka, M., and Kras-Lapinska, M. (2004). 37. Nephrangis (Schltr.) Summerh. In Flore du Gabon 37, Orchidaceae II (ed. P. Morat), pp. 369–72. Muséum National d’Histoire Naturelle, Paris.

TRANSVERSE SECTION

Trichomes rare; multicellular, glandular. Stomata abaxial. Hypodermis absent. Fibre bundles absent. Mesophyll homogeneous. Water-storage cells smooth to pitted. Vascular bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Velamen two cells thick. Epivelamen cells radially elongate; walls ∪-thickened with fine anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate. Distinct endovelamen thickenings absent. Cover cells present over short cells of exodermis. Exodermal long cells primarily ○-thickened. Water-storage cell walls smooth to pitted. Aeration units present. Endodermal cells ○-thickened. Vascular cylinder 7-arch. Vascular tissue embedded in sclerenchyma. Pith sclerenchymatous. (BC)

Palynology There are no published data on pollen of Oeonia. (AP) 417

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Fig. 743.1.  Oeonia volucris (Thouars) Spreng. A. Habit; B. Flower, front view; C. Flower, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum; H. Labellum, spread; I. Column and labellum, side view; J. Column, front view; K. Column, side view; L. Column, anther cap removed, from above; M. Anther cap, dorsal view; N. Anther cap, ventral view; O. Pollinarium; P. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Humblot 667 (K) and Kew Spirit Collection no. 60989.

Cytogenetics Chromosome counts for these species have not been reported. (AP)

Phytochemistry No reports have been found on the phytochemistry of Oeonia. (NV, RG)

angraecoid orchids. More DNA sampling of taxa is needed to show the true relationship between Oeonia and Cryptopus. (AP)

Ecology Species are found in coastal and humid, mossy, evergreen forests from sea level to 2000 m. (PC)

Phylogenetics

Pollination

Carlsward et al. (2006b) and Micheneau et al. (2008) included Oeonia rosea in their DNA analysis of Afro-Madagascan

There are no published data on the pollination biology of Oeonia. (AP)

418

OEONIELLA

at tip. Inflorescence axillary, suberect-spreading, racemose, manyflowered; peduncle terete; bracts amplexicaul. Flowers distichously arranged on rachis, white. Sepals and petals spreading, incurved at tip, linear-tapering, acuminate. Labellum free, tubular-conical and enclosing column, trilobed at tip, midlobe long-apiculate, ecallose, spurred; spur cylindrical, obtuse, with a narrow mouth. Column with a convex anther, truncate, erose at tip, pollinia two, stipes (tegulae) two, linear, elongate, viscidium cordate-quadrate; stigma transversely oblong; rostellum decurved, trilobed, side lobes longest and incurved-falcate, midlobe minute. Ovary subsessile. (PC)

Distribution (Fig. 744.2) Oeoniella is a genus of only two species—O. polystachys and O. aphrodite (Balf.f. & S. Moore) Schltr.—confined to Madagascar and the Mascarene Islands. (PC)

Anatomy Carlsward et al. (2006a) examined O. polystachys. Their results are summarized below. Fig. 743.2.  Distribution map of Oeonia.

Uses No ethnobotanical uses for these species are available in the literature, and they are rare in cultivation. (AP)

Taxonomic notes Oeonia is closely allied to Cryptopus being distinguished by its entire petals and 3- or 5-lobed labellum in which the basal lobes enclose the column or are erect on either side of it. (PC)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Micheneau, C., Carlsward, B. S., Fay, M. F., Bytebier, B., Pailler, T., and Chase, M. W. (2008). Phylogenetics and biogeography of Mascarene angraecoid orchids. Molecular Phylogenetics and Evolution, 46, 908–22.

744.  OE O N IE LLA Oeoniella Schltr., Beih. Bot. Centralbl., 36, 101 (1918). Type species: Oeoniella polystachys (Thouars) Schltr. (basionym: Epidendrum polystachys Thouars). Vide Senghas, Orchidee, 14, 215 (1963).

Derivation of name

Leaf TRANSVERSE SECTION

Trichomes multicellular, glandular. Stomata abaxial. Hypodermis absent. Fibre bundles absent. Mesophyll homogeneous. Waterstorage cells banded. Vascular bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Velamen two or three cells thick. Epivelamen cells radially elongate; walls ∪-thickened with fine, anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate. Distinct endovelamen thickenings absent. Cover cells present over short cells of exodermis. Exodermal long cells ∩-thickened to ○-thickened. Water-storage cells with pitted to banded walls, often slightly thickened. Aeration units present. Endodermal cells ○-thickened. Vascular cylinder 18–25-arch. Vascular tissue embedded in parenchyma and/or sclerenchyma. Pith parenchymatous or thin-walled and sclerenchymatous. (BC)

Palynology Schill and Pfeiffer (1977) described the pollen tetrads of Oeoniella polystachys as convex-concave with laevigate-foveolate sculpturing and a calymmate sexine. (AP)

Cytogenetics

The generic name is derived from Oeonia, a Madagascan orchid genus, and -ella, a diminutive, meaning small Oeonia. (PC)

Brandham (1999) reported a chromosome count of 2n = 38 for the genus. (AP)

Description (Plate 171; Fig. 744.1)

Phytochemistry

Epiphytic herbs. Roots emerging along stem. Stem leafy. Leaves coriaceous, conduplicate, spreading, oblong-linear, unequally bilobed

The main constituents of the floral scent of O. polystachys (Thouars) Schltr. are methyl benzoate (34.0%), methyl salicylate 419

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Fig. 744.1.  Oeoniella aphrodite (Balf.f. & S.Moore) Schltr. A. Habit; B. Bract; C. Flower, front; D. Flower, back; E. Ovary, transverse section; F. Dorsal sepal; G. Lateral sepal; H. Petal; I. Flower, dorsal sepal removed, side view; J. Labellum, spread; K. Column, side view; L. Column, dorsal view; M. Anther cap, dorsal view; N. Anther cap with pollinarium, ventral view; O. Pollinarium; P. Pollinia. Single bar = 1 mm, double bar = 1 cm. Drawn by Linda Gurr from Bosser 21.572 (K).

(24.0%), (E)-isoeugenol (20.0%), benzyl benzoate (10.3%), and benzyl salicylate (2.7%), although contributions to the overall fragrance are also made by several minor components, including p-cresol, (Z)-hex-3-en-1-yl benzoate, jasmone, methyl cis-(Z)jasmonate, and vanillin (Kaiser 2011). (NV, RG)

Phylogenetics There are no published DNA sequence data for Oeoniella. (AP) 420

Ecology Both species occur in coastal and evergreen forests from sea level to 100 m. (PC)

Taxonomic notes Schlechter (1918) established Oeoniella as a split from Angraecum distinguished by its distinctive column structure, the pollinia each attached to an elongate stipes (tegula), and the funnel-shaped

OSSICULUM

Anatomy Gasson and Cribb (1986) and Carlsward et al. (2006a) examined the leaf anatomy of O. aurantiacum. Below are their observations.

Leaf TRANSVERSE SECTION

Trichomes absent. Sunken areas in epidermis abundant. Stomata abaxial. Hypodermis absent. Fibre bundles present singly along each margin, composed of many thick-walled fibres surrounding 2–5 small thin-walled cells. Mesophyll homogeneous. Waterstorage cells banded, often slightly thickened. Vascular bundles arranged in a V-shaped pattern as a result of leaf TS shape. Bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found associated with phloem sclerenchyma only; encircling fibre bundles.

Root Roots were not studied. (BC)

Palynology Fig. 744.2.  Distribution map of Oeoniella.

Pollen of this species has not been studied. (AP)

labellum with a trilobed tip and apiculate midlobe that enfolds the column. (PC)

Cytogenetics

Taxonomic literature

Brandham (1999) listed a chromosome count of 2n = 34 for Ossiculum. (AP)

Schlechter, R. (1918). Versuch einer natürlichen Neuordnung der Afrikanischen Angraekoiden Orchidaceen. Beihefte zum Botanischen Centralblatt, 36, 62–181.

745.  OS SIC ULUM Ossiculum P.J.Cribb & Laan, Kew Bull., 41, 823 (1985). Type species: Ossiculum aurantiacum P.J.Cribb & Laan.

Derivation of name From ossiculum, a small bone, named after the Dutch botanist Henk Beentje who collected the type species and whose name means ‘small bone’ in Dutch. (PC)

Description (Fig. 745.1) Epiphytic herbs. Roots mostly toward base of stem. Stem leafy. Leaves distichously arranged, canaliculate, acute, sheathing bases imbricate. Inflorescence axillary, lateral, many-flowered, shorter than leaves. Flowers non-resupinate, bright orange with a paler labellum. Sepals subsimilar, ovate-elliptic, acuminate. Petals oblanceolate, acute. Labellum entire, oblong, apiculate, ecallose, with a basal spur; spur clavate. Column with an obovoid anther; pollinia two, porate, attached by an oblanceolate tegula to a horseshoeshaped viscidium; stigma transversely elliptic; rostellum shallowly semi-lunate with a horseshoe-shaped, apical notch. Ovary subsessile. (PC)

Phytochemistry No reports have been found on the phytochemistry of Ossiculum. (NV, RG)

Phylogenetics Ossiculum has the lowest chromosome number of any angraecoid orchid. In its flower colour and labellum shape it is reminiscent of some tropical Asiatic vandaceous orchids. In its pollinarium and rostellar structures it appears to be allied to Calyptrochilum but differs from that genus in its size and habit, leaf orientation and structure, inflorescence and flower size, morphology, and colour. Szlachetko (1995, 2003a) placed it in his subtribe Calyptrochilinae with Calyptrochilum, Chauliodon, Dinklageella, Lacroixia, and Neofinetia. The last does not belong to Angraecinae (Carlsward et al. 2006b). DNA sequences of this species have not been available for phylogenetic analysis. (PC)

Ecology Plants of this species are epiphytic on thick branches of trees in primary moist evergreen forests below 200 m. (PC)

Pollination The pollination biology of this species is unknown. (AP)

Distribution (Fig. 745.2)

Uses

Ossiculum is a monospecific genus in Cameroon. (PC)

There are no known uses for O. aurantiacum. (AP) 421

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A Fig. 745.1.  Ossiculum aurantiacum P.J.Cribb & Laan. A. Habit, ×2/3; B. Inflorescence, ×2; C. Flower, side view, ×4; D, F. Lateral sepals, ×4; E. Dorsal sepal, ×4; G, H. Petals, ×4; I. Labellum with spur, ×4; J. Column and labellum (portion), dorsal view, ×16; K. Anther cap, dorsal view, ×16; L. Column, side view, ×16; M. Column, anther cap removed, ×16; N. Pollinarium, front view, ×16; O. Pollinarium, side view, ×16; P. Column, anther removed, ×16; Q. Capsule, side view, ×2; R. Capsule, transverse section, ×2. Magnifications as originally published. Drawn by Wil Wessel-Brand from Beentje 1460A and van der Laan 718 spirit, WAG). Reprinted with permission from Kew Bulletin, 41, 825 (1986).

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86.

422

Szlachetko, D. L. (1995). Systema orchidalium. Fragmenta Floristica et Geobotanica Supplementum, 3, 1–152. Szlachetko, D. L. (2003a). Gymnostemia orchidalium 3. Acta Botanica Fennica, 176, 1–311.

PLEC TRELMINTHUS

Anatomy Carlsward et al. (2006a) examined P. caudatus. Their results are summarized below.

Leaf TRANSVERSE SECTION

Trichomes multicellular, glandular. Stomata abaxial; rarely adand abaxial. Hypodermis ad- and abaxial; composed of fibrous idioblasts forming a single row and occurring singly or in pairs scattered among mesophyll chlorenchyma cells. Adaxial fibrous idioblasts larger than abaxial idioblasts. Fibre bundles absent. Mesophyll heterogeneous with columnar adaxial cells and isodiametric abaxial cells. Water-storage cell walls not birefringent. Thick-walled fibrous idioblasts concentrated near hypodermis. Vascular bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Plectrelminthus Raf., Fl. Tellur., 4, 42 (1936). Type species: Plectrelminthus caudatus (Lindl.) Summerh. (basionym: Angraecum caudatum Lindl.) Leptocentrum Schltr., Orchideen, 600 (1914). Type species: Leptocentrum caudatum (Lindl.) Schltr. (=Plectrelminthus caudatus (Lindl.) Summerh.)

Velamen four or five cells thick. Epivelamen cells radially elongate; walls ∪-thickened with fine, anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate; outer layers composed of smaller, thicker-walled cells than inner layers. Distinct endovelamen thickenings absent. Cover cells present over short cells of the exodermis. Exodermal long cells primarily ∩-thickened or ○-thickened. Water-storage cell walls banded. Aeration units present. Endodermal cells usually ○-thickened to rarely ∪-thickened. Vascular cylinder 17–40arch. Vascular tissue embedded in thin-walled parenchyma or in sclerenchyma. Pith sclerenchymatous. (BC)

Derivation of name

Palynology

From the Greek plektron, spur, and minthion, worm, in reference to the worm-like spur (Schultes and Pease 1963). (AP)

Pollen of Plectrelminthus has not been studied. (AP)

Fig. 745.2.  Distribution map of Ossiculum.

746.  P LE C TR E LMIN TH U S

Description (Plate 172; Fig. 746.1) Erect or spreading epiphytic herbs. Stem leafy. Leaves distichous, coriaceous, spreading-arcuate, linear, unequally bilobed at tip, imbricate and sheathing at base. Inflorescence longer than leaves, spreading, several-flowered. Flowers non-resupinate, pedicel twisted proximally. Sepals and petals subsimilar, spreading, lineartapering. Labellum erect, narrowly clawed at base with thickened margins or auricles; lamina oblong-pandurate, trilobed at apex, long-apiculate; spur cylindrical, spiral apically, 15–25 cm long, pendent. Column fleshy toward base, dilated at apex; anther obovoid, pollinia two, porate, attached by a oblanceolate stipe (tegula) to an elliptic viscidium with a notched tip; stigma kidney-shaped; rostellum tapering to a shallowly notched tip. (PC)

Distribution (Fig. 746.2) Plectrelminthus is a monospecific genus of West and Central Africa, from Guinea and Sierra Leone to Zaire and Central African Republic. (PC)

Cytogenetics Arends and Van der Laan (1983) reported a chromosome count of 2n = 46 for P. caudatus. (AP)

Phytochemistry The floral fragrance of P. caudatus is composed mainly of tiglate esters, among which hexyl tiglate (58%) predominates (Kaiser 1993). More minor components include tiglyl tiglate (6.9%), butyl tiglate (5.0%), hexyl butyrate (3.4%), and methyl caproate (3.0%). The scent, which is emitted at night, has been described as ‘probably unique in the whole realm of flowering plants’ (Kaiser 1993). (NV, RG)

Ecology The species is epiphytic in light shade on trunks and large branches of forest trees in damp lowland evergreen forests, open forests, and solitary trees in grassland, also on granite rocks in grassland in areas of high rainfall, occurring from sea level to 1000 m. (PC) 423

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Fig. 746.1.  Plectrelminthus caudatus (Lindl.) Summerh. A. Habit; B. Flower; C. Attachment of lateral sepals (dorsal sepal and petals removed); D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum; H. Column and labellum, side view; I. Column, front view; J. Column, side view; K. Column apex, anther cap removed; L. Anther cap, dorsal view; M. Anther cap, ventral view; N. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Cable 32 (K) and Kew Spirit Collection no. 70291.

Pollination There is no information on the pollination biology of this species, but given its nocturnal scent it is probably sphingophilous. (AP)

petals, an erect, obovate labellum with a long apicule, and a long, pendent, spirally twisted spur 17–25 cm long. There have been no phylogenetic analyses using DNA sequences of this species. (PC)

Uses

7 4 7 .  P ODA NGI S

There are no known ethnobotanical uses for the species, but it is occasionally cultivated in specialist collections. (AP)

Podangis Schltr., Beih. Bot. Centralbl., 36, 82 (1918). Type: Podangis dactyloceras (Rchb.f.) Schltr. (basionym: Listrostachys dactyloceras Rchb.f.). Neowolffia O.Gruss, Orchid. Atlas, 239 (2007). Type species: Neowolffia rhipsalisocia (Rchb.f.) O.Gruss (basionym: Angraecum rhipsalisocium Rchb.f.)

Taxonomic notes Plectrelminthus caudatus is distinctive with its spreading racemes of non-resupinate flowers with stellately arranged green sepals and 424

PODANGIS

Stomata abaxial. Hypodermis abaxial; fibrous idioblasts forming an almost continuous layer infrequently interrupted by chlorenchyma cells. Fibre bundles absent. Mesophyll heterogeneous with several rows of columnar cells surrounding a central suture of isodiametric to periclinally flattened cells. Water-storage cell walls smooth to pitted, concentrated in central suture of mesophyll. Vascular bundles forming a circle around central suture. Bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Fig. 746.2.  Distribution map of Plectrelminthus.

Derivation of name From the Greek pous, foot, and angos, vessel, from the spur of the labellum that looks like a foot. (PC)

Velamen two or three cells thick. Epivelamen cells radially elongate; walls ∪-thickened with fine anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate; outer layer composed of thicker-walled cells than inner layer. Distinct endovelamen thickenings absent. Cover cells present over short cells of exodermis. Exodermal long cells primarily ○-thickened. Water-storage cells smooth to pitted. Aeration units present. Endodermal cells strongly ○-thickened. Vascular cylinder 20–30-arch. Vascular tissue embedded in sclerenchyma. Pith sclerenchymatous. (BC)

Palynology Pollen of these species has not been examined. (AP)

Description (Plates 173, 174; Fig. 747.1)

Cytogenetics

Epiphytic psygmoid herbs. Roots basal. Stem leafy. Leaves fleshy, bilaterally flattened, arranged in a fan, linear-tapering, acute, articulated to basal, imbricate leaf sheaths. Inflorescence shorter than leaves, unbranched, many-flowered in heads; bracts triangular. Flowers resupinate, campanulate, pure white with a green anther, pedicel and ovary white. Sepals and petals free, subsimilar, elliptic. Labellum concave, fan-shaped, entire or obscurely trilobed, ecallose, spurred at base; spur pendent, clavate or tapering from a broader mouth. Column clavate; pollinia two, porate, each attached by a linear stipe (tegula) to an oblong viscidium; stigma circularoblong; rostellum porrect but upcurved and bifid at tip. (PC)

Arends et al. (1980) reported a chromosome count of 2n = 46 for Podangis. (AP)

Distribution (Fig. 747.2) Podangis is a genus of two species, P. dactyloceras and P. rhipsalisocia (Rchb.f.) P.J.Cribb and Carlsward, from West and Central Africa, ranging from Senegal, Guinea, and Sierra Leone to Uganda and Tanzania, and south to Angola. (PC)

Anatomy Carlsward et al. (2006a) examined P. dactyloceras. Their results are summarized below.

Leaf TRANSVERSE SECTION

Podangis dactyloceras possesses terete leaves with no discernible adaxial epidermis. Trichomes rare; multicellular, glandular.

Phytochemistry Although Lüning (1974) noted that one unnamed species of Podangis had an alkaloid content ≥ 0.1%, nothing further has been published on the phytochemistry of this genus. (NV, RG)

Phylogenetics Carlsward et al. (2006b) included Podangis dactyloceras in their DNAbased analysis of angraecoid orchids. It was sister to Rangaeris rhipsalisocia (Rchb.f.) Summerh. with strong bootstrap support, and both were part of a larger clade that included Tridactyle, Listrostachys, and Ypsilopus. Podangis dactyloceras and Rangaeris rhipsalisocia both have distichously arranged, imbricate leaves borne on a short stem, a similar column and pollinarium structure, and a common chromosome number of 2n = 46 (Arends et al. 1980). In contrast, the latter has less in common with the rest of Rangaeris, which is polyphyletic. For these reasons R. rhipsalisocia was transferred to Podangis as P. rhipsalisocia (Cribb and Carlsward 2012). (PC)

Ecology These two species are epiphytes in shady places in semi-­ deciduous rain forests and gallery forests, often near water, from sea level to 1900 m. (PC) 425

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Fig. 747.1.  Podangis dactyloceras (Rchb.f.) Schltr. A. Habit; B; Flower; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum; G. Column with anther; H. Column, anther removed; I. Anther cap; J. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Cherry Ann Lavrih from Lock s.n. (unpublished illustration) and Meyer K8 (K). Reproduced with permission from P. J. Cribb (1989), Flora of Tropical East Africa, Orchidaceae (Part 3). A.A. Balkema, Rotterdam/Brookfield.

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RANGAERIS

Description (Fig. 748.1) Epiphytic or rarely lithophytic herbs. Roots emerging along stem opposite leaves, unbranched or rarely branching. Stem entirely covered with persistent leaf-bases. Leaves coriaceous, distichous, conduplicate, linear or oblong, unequally bilobed at apex. Inflorescence pendent or spreading to suberect, bracts longer than pedicels. Flowers usually white or yellowish. Sepals and petals free, subsimilar, spreading to recurved. Labellum entire or obscurely trilobed, ecallose; spur pendent, filiform, elongate from a narrow mouth. Column glabrous or puberulent, lacking a foot; anther cucullate, produced in front and truncate at the apex, pollinia two, pyriform, stipes two, linear or oblanceolate, viscidium oblong or cordate; rostellum bifid. (PC)

Distribution (Fig. 748.2) Rangaeris is a genus of six species confined to tropical and ­southern Africa. (PC)

Fig. 747.2.  Distribution map of Podangis.

Pollination There is no published information on the pollination biology of these species. (AP)

Uses There are no uses recorded for Podangis other than occasionally being cultivated. (AP)

Taxonomic literature Arends, J. C., van der Burg, W. J., and Van der Laan, F. M. (1980). Notes on African orchids. Miscellaneous Papers, Landbouwhogeschool Wageningen. Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Cribb, P. J. and Carlsward, B. (2012). New combinations in Aerangis, Diaphananthe and Podangis (Orchidaceae, subtribe Angraecinae). Phytotaxa, 71, 42–7.

Anatomy Carlsward et al. (2006a) examined four species of Rangaeris. Their results are summarized below.

Leaf TRANSVERSE SECTION

Trichomes infrequent, multicellular, glandular. Stomata abaxial; ad- and abaxial in R. muscicola; slightly sunken in R. muscicola. Hypodermis adaxial only in R. longicaudata (Rolfe) Summerh.; ad- and abaxial in R. amaniensis (Kränzl.) Summerh., R. muscicola, and R. schliebenii (Mansf.) P.J.Cribb; composed of fibrous, thinto thick-walled idioblasts, distributed in a single row below the epidermis interrupted by chlorenchyma. Fibre bundles absent. Mesophyll homogeneous in R. amaniensis and R. muscicola; heterogeneous with columnar adaxial cells and isodiametric abaxial cells in R. longicaudata and R. schliebenii. Lignified, moderately thickened fibrous idioblasts scattered throughout mesophyll in R. schliebenii. Water-storage cells with birefringent walls found only above the midrib in R. muscicola; water-storage cell walls not birefringent in all other species examined. Vascular bundle sheath indistinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

748.  RAN G A E R IS Rangaeris Summerh., Kew Bull., 4, 435 (1949). Type species: Rangaeris muscicola (Rchb.f.) Summerh. (basionym: Aeranthes muscicola Rchb.f.) Barombiella Szlach., Ann. Bot. Fenn., 40, 69 (2003). Type species: Barombiella schliebenii (Mansf.) Szlach. (basionym: Leptocentrum schliebenii Mansf.), syn nov.

Derivation of name Rangaeris is an anagram of Aerangis, a closely related genus. (PC)

Velamen 2–4 cells thick. Epivelamen cells radially elongate; walls ∪-thickened with fine, anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate; outer layer composed of thicker-walled cells than inner layer. Distinct endovelamen thickenings absent. Cover cells present over short cells of the exodermis. Exodermal long cells primarily ○-thickened. Mucilage globules abundant in the cortex of R. longicaudata. Water-storage cell walls banded. Aeration units present in R. amaniensis and R. muscicola. Endodermal cells ○-thickened. Vascular cylinder 15–29-arch. Vascular tissue embedded in sclerenchyma, cell walls of embedding sclerenchyma thickest around 427

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Fig. 748.1.  Rangaeris muscicola (Rchb.f.) Summerh. A. Habit; B. Leaf apex, transverse section; C. Flower, spur removed, front view; D. Flower, side view; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum; I. Column and labellum, side view; J. Column, side view; K. Column, front view; L. Column, anther cap removed; M. Anther cap; N. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Schelpe 2887 (K) and Kew Spirit Collection no. 42275. Reproduced with permission from G. V. Pope (ed.) (1998), Flora Zambesiaca: Mozambique, Malawi, Zambia, Zimbabwe, Botswana. Volume 11, Part 2. Royal Botanic Gardens, Kew.

phloem clusters in R. amaniensis and above phloem clusters in R. schliebenii. Pith sclerenchymatous. (BC)

Palynology Pollen of Rangaeris has not been investigated. (AP) 428

Cytogenetics Brandham (1999) listed chromosome numbers of 2n = 46, 50, 92, c. 100, c. 108 for Rangaeris and suggested two possible basic numbers of x = 23 and x = 25. The difference might be explained by the fact that Rangaeris as presently understood is not

RHAESTERIA

Pollination See reference by Martins and Johnson (2007) under Aerangis. (AP)

Uses There are no ethnobotanical uses reported for Rangaeris, although a few species are occasionally cultivated. (AP)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Cribb, P. J. and Carlsward, B. (2012). New combinations in Aerangis, Diaphananthe and Podangis (Orchidaceae, subtribe Angraecinae). Phytotaxa, 71, 42–7. Rice, R. (2006b). Notes on the subtribe Aerangidinae (Orchidaceae). Oasis, 3, 12–15.

Fig. 748.2.  Distribution map of Rangaeris.

monophyletic according to the results of Carlsward et al. (2006b). (AP)

Phytochemistry The scent of R. amaniensis is due to a combination of farnesol (27.9%), linalool (2.4%), methyl cinnamate (20.1%), methyl salicylate (5.8%), and phenylethyl alcohol (8.5%) (Kaiser 1993). One unnamed species of Rangaeris was reported to have an alkaloid content ≥ 0.1% by Lüning (1974). (NV, RG)

Phylogenetics Rangaeris was established by Summerhayes to include some anomalous angraecoid orchids with their pollinia attached by filiform tegulae to a common, oblong viscidium. In their DNA-based phylogenetic analysis of angraecoid orchids, Carlsward et al. (2006b) showed that Rangaeris is not monophyletic. Rangaeris rhipsalisocia (Rchb.f.) Summerh. is sister to Podangis dactyloceras (Rchb.f.) Schltr., and they share a psygmoid habit. The former species was transferred to Podangis (Cribb and Carlsward 2012). Two accessions of the type species of Rangaeris, R. muscicola, were sister to each other, and they in turn were sister to a grade including R. amaniensis and species of Podangis (including R. rhipsalisocia), Cyrtorchis, Tridactyle, Listrostachys, and Ypsilopus. More DNA sequencing is needed to establish the boundaries of Rangaeris and related genera. Rice (2006b) transferred R. trilobata Summerh. to Solenangis on the basis of its habit, but its leaf texture and morphology and floral structure are distinct. (PC)

Ecology Species of Rangaeris are epiphytes and occasional lithophytes in humid, wet, evergreen and semi-deciduous forests and woodlands from sea level to 2000 m. (PC)

7 4 9 .  R H A ES T ER I A Rhaesteria Summerh., Kew Bull., 20, 191 (1966). Type species: Rhaesteria eggelingii Summerh.

Derivation of name From the Greek rhaister, hammer, in allusion to the hammershaped rostellum of the type species. (PC)

Description (Plate 175; Fig. 749.1) Epiphytic herbs. Stem leafy. Roots emerging all along stem. Leaves distichous, conduplicate. Inflorescence axillary, appearing fasciculate but truly successive, few-flowered; bracts sheathing. Flowers resupinate, yellowish. Sepals and petals not spreading, forming a tube in basal third. Labellum free, concave, enclosing base of column, spurred; spur cylindrical, with a broad mouth. Column with a convex, hemispherical anther, pollinia two, stipes (tegulae) two, viscidia distinct; stigma transversely oblong; rostellum decurved, hammer-shaped. Ovary sessile. (PC)

Distribution (Fig. 749.2) Rhaesteria is a monospecific genus in Uganda and Rwanda. (PC)

Anatomy Nothing is known of the vegetative anatomy of this species. (AP)

Palynology Pollen of Rhaesteria has not been studied. (AP)

Cytogenetics There are no published chromosome counts for this species. (AP) 429

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Fig. 749.1.  Rhaesteria eggelingii Summerh. A. Habit; B. Inflorescence; C. Flower, side view; D. Flower, front view; E. Fused sepals and petals; F. Flower with sepals and petals removed; G. Column with anther cap; H. Column, anther cap removed, side view; I. Column, anther cap removed, front view; J. Anther cap, ventral view; K. Anther cap, dorsal view. Single bar = 1 mm, double bar = 1 cm. Drawn by Mary Grierson from Eggeling 5849 (K). Reproduced with permission from P. J. Cribb (1989), Flora of Tropical East Africa, Orchidaceae (Part 3). A.A. Balkema, Rotterdam/Brookfield.

Phytochemistry

Pollination

No reports have been found on the phytochemistry of Rhaesteria. (NV, RG)

The pollination biology of this species is unknown. (AP)

Ecology

Uses

Rhaesteria eggelingii is an epiphyte on trees in savannas and grasslands on the margins of forests and woodlands at elevations of 1300–2300 m. (PC)

There are no known uses for R. eggelingii. (AP)

430

RH IPIDOGLOSSUM

oblong, ligulate, linear or elliptic, unequally bilobed at tip, with a sheathing base. Inflorescence axillary, spreading to pendent. Flowers often translucent, white, pale yellow, pale green, orange or brown. Sepals and petals free, subsimilar or not, spreading, rarely reflexed. Labellum deflexed or porrect, entire or trilobed or pandurate, spurred at base, ecallose or with a simple, fleshy, basal callus in mouth of spur; spur clavate to cylindrical. Column with two pollinia, porate, each attached to a linear tegula and those to a circular or elliptic viscidium; rostellum dependent, fleshy, peglike. Ovary and pedicel longer than bracts. (PC)

Distribution (Fig. 750.2) Rhipidoglossum is a genus of about 40 species found in tropical and southern Africa. (PC)

Anatomy Carlsward et al. (2006a) examined eight species of Rhipidoglossum. Their results are summarized below. Fig. 749.2.  Distribution map of Rhaesteria.

Leaf TRANSVERSE SECTION

Taxonomic literature

Trichomes absent in R. curvatum (Rolfe) Garay and R. obanense (Rendle) Summerh.; when present, glandular, multicellular with the basal cell sunken in a buttress of raised epidermal cells. Stomata mostly abaxial; ad- and abaxial in R. obanense. Hypodermis absent. Fibre bundles absent. Mesophyll homogeneous or heterogeneous with columnar anticlinal adaxial cells and isodiametric abaxial cells in R. bilobatum (Summerh.) Szlach. & Olszewski, R. pulchellum (Summerh.) Garay, and R. xanthopollinium. Waterstorage cells banded or not birefringent. Vascular bundle sheath indistinct in R. curvatum, R. kamerunense (Schltr.) Garay, and R. obanense. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Rice, R. (2006b). Notes on the subtribe Aerangidinae (Orchidaceae). Oasis, 3, 12–15.

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Taxonomic notes In habit and flower shape, Rhaesteria superficially resembles a small-flowered Angraecum, but it is distinct in that sepals and petals are united in their basal third to form a short tube. This is found elsewhere in Africa only in the leafless, epiphytic genus Taeniophyllum. Rice (2006b) suggested that Rhaesteria should be transferred to Mystacidium, but the floral morphology would indicate that the two genera should be kept separate. There are no DNA data to establish relationships of the species. (PC)

750.  RH IPID O G LO SSUM Rhipidoglossum Schltr., Beih. Bot. Centralbl., 36, 80 (1918). Type species: Rhipidoglossum xanthopollinium (Rchb.f.) Schltr. (basionym: Aeranthus xanthopollinia Rchb.f.) (vide Summerhayes in Blumea, Suppl. 1, 80. 1937) Crossangis Schltr., Beih. Bot. Centralbl., 36, 141 (1918). Type species: Crossangis polydactyla (Kränzl.) Schltr. (basionym: Listrostachys polydactyla Kränzl.) Sarcorhynchus Schltr., Beih. Bot. Centralbl., 36, 104 (1918). Type species: not designated

Derivation of name From the Greek rhipis, fan, and glossa, tongue, in reference to the fan-like labellum of several species. (PC)

Description (Plate 176; Fig. 750.1) Epiphytic or lithophytic herbs. Roots sometimes more prominent leaves. Stem leafy. Leaves distichously arranged, coriaceous,

TRANSVERSE SECTION

Velamen 2–4 cells thick. Epivelamen cells isodiametric in R. curvatum and R. kamerunense, radially elongate in all other specimens examined; walls ∪-thickened with fine, anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate; outermost layers of cells thicker-walled than inner layers. Distinct endovelamen thickenings ridged in R. obanense, absent in all other species examined. Cover cells present except in R. obanense and R. pulchellum. Exodermal long cells ○-thickened or ∩-thickened in R. kamerunense, R. obanense, and R. xanthopollinium. Water-storage cell walls smooth to pitted. Single modified cortical layer of thin- to thick-walled cells surrounding the endodermis in R. curvatum and R. pulchellum. Aeration units present in R. bilobatum, R. kamerunense, R. obanense, R. subsimplex (Summerh.) Garay, R. xanthopollinium. Endodermal cells ○-thickened. Vascular cylinder 7–20-arch. Vascular tissue embedded in sclerenchyma, cell walls of embedding sclerenchyma thickest around phloem clusters in R. pulchellum, R. rutilum (Rchb.f.) Schltr., R. subsimplex, and R. xanthopollinium. Pith usually sclerenchymatous but parenchymatous in R. kamerunense. (BC) 431

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F Fig. 750.1.  Rhipidoglossum stolzii (Schltr.) Garay. A. Habit, ×1; B. Flower, ×2; C. Lateral sepal, ×2; D. Petal, ×2; E. Dorsal sepal, ×2; F. Column and labellum, side view, ×3; G. Column with anther cap removed, side view, ×8; H. Anther cap, dorsal view, ×8; I. Anther cap, ventral view, ×8; J. Pollinaria, ×8. Magnifications as originally published. Drawn by Judi Stone from Chapman 8332 (K) and Kew Spirit Collection no. 49259. Reproduced with permission from G. V. Pope (ed.) (1998), Flora Zambesiaca: Mozambique, Malawi, Zambia, Zimbabwe, Botswana. Volume 11, Part 2. Royal Botanic Gardens, Kew.

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Pollination Nothing is known about the pollination biology of these species. (AP)

Uses There are no known ethnobotanical uses of Rhipidoglossum, and plants are rare in cultivation. (AP)

Taxonomic notes Schlechter (1918) established the genus Rhipidoglossum as distinct from Diaphananthe, distinguishing it by its pollinia each being attached by a tegula to its own viscidium, whereas in the latter the pollinia and stipites are attached to a common viscidium. Nevertheless, many authors have considered the genera to be congeneric (Summerhayes 1960; Cribb 1984). The phylogenetic studies of Carlsward et al. (2006b) have upheld the recognition of Rhipidoglossum but brought into question recognition of Cribbia. (PC)

Fig. 750.2.  Distribution map of Rhipidoglossum.

Palynology Pollen of Rhipidoglossum has not been investigated. (AP)

Cytogenetics Felix and Guerra (2010) listed chromosome numbers of 2n = 50, 100 for six species (originally treated as Diaphananthe). (AP)

Phytochemistry The floral fragrance of R. pulchellum (cited as Diaphananthe pulchella Summerh.) contains large amounts of methyl benzoate (33.2%) and methyl salicylate (38.5%). Other contributions to the scent are made by benzyl acetate (1.5%) and nerolidol (5.5%), which are among the minor components (Kaiser 1993). One record on alkaloid content for Rhipidoglossum refers to the former genus Sarcorhynchus, but the species in question was not named (Lüning 1974). The compounds were present at levels < 0.1%. (NV, RG)

Phylogenetics As presently understood from ITS rDNA studies (Carlsward et al. 2006b), Rhipidoglossum is paraphyletic. Cribbia brachyceras (Summerh.) Senghas and C. confusa P.J.Cribb were sister to R. kamerunense, and those sister to R. rutilum, R. xanthopollinium, and R. subsimplex. The pollinarium and rostellum structure of Cribbia are similar to those of Rhipidoglossum but differ in details of the rostellum and labellum. Further DNA sequencing is needed to establish the phylogenetic relationship between these two genera. (PC)

Ecology Species occur in evergreen rain forests, seasonally dry forests, and savanna woodlands, on rocks, in shade and in full sun, from sea level to 2000 m. (PC)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Cribb, P. J. (1984). Diaphananthe. In Flora of tropical East Africa. Orchidaceae. Part 3 (ed. R. Polhill), pp. 521–40. Balkema, Rotterdam. Schlechter, R. (1918). Versuch einer natürlichen Neuordnung der Afrikanischen angraekoiden Orchidaceen. Beihefte zum Botanischen Centralblatt, 36, 62–181. Stewart J., Hermans, J., and Campbell, D. (2006). Angraecoid orchids. Timber Press, Portland, Oregon. Summerhayes, V. S. S. 1960. African orchids XXVII. Kew Bulletin, 14, 126–57.

7 5 1 .  S OB ENNI KOFF I A Sobennikoffia Schltr., Repert. Spec. Nov. Regni Veg. Beih., 33, 361 (1925). Type species: Sobennikoffia robusta (Schltr.) Schltr. (basionym: Oeonia robusta Schltr.) Vide F. Butzin, Taxon, 32, 632. (1983)

Derivation of name The generic name commemorates Alexandra Schlechter (nee Sobennikoff), the wife of Rudolf Schlechter and daughter of a Russian tea merchant. (PC)

Description (Plate 177; Fig. 751.1) Lithophytic or epiphytic herbs. Stems leafy, erect, often forming clumps. Leaves distichously arranged, conduplicate, linear or loriform, unequally bilobed at base, with distichous, imbricate, sheathing bases. Inflorescence axillary, erect or suberect, as long as leaves or longer, several-flowered; bracts ovate or triangular and longer than pedicels. Flowers resupinate, white, turning yellow with age. Sepals and petals spreading-recurved, subsimilar, linear-tapering. Labellum 433

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Fig. 751.1.  Sobennikoffia robusta (Schltr.) Schltr. A. Habit; B. Flower and bract; C. Leaf apex; D. Bract; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum; I. Column, labellum, and spur, side view; J. Column, front view; K. Column, side view; L. Column, back view; M. Column apex, anther cap removed, back view; N. Anther cap with pollinarium; O. Pollinarium; P. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Kew Living Collection.

porrect, concave at base, trilobed apically, tapering, acute, midlobe longer than lateral lobes; callus a longitudinal linear ridge; spur cylindrical from a broader mouth, rising above ovary. Column with anther broader than long, pollinia two, cleft, each attached directly to its own linear viscidium; rostellum cleft. (PC) 434

Distribution (Fig. 751.2) Sobennikoffia is an endemic Madagascan genus of four species. (PC)

SOLENANGIS

Cytogenetics There are no reports of chromosome counts for any of the species. (AP)

Phytochemistry The fragrance of S. humbertiana flowers, which resembles that of honeysuckle (Lonicera japonica Thunb.; Caprifoliaceae), is due largely to a combination of linalool (35.0%) and indole (0.8%), together with benzyl acetate, jasmine lactone, (E,E)-farnesene, and germacrene D (Kaiser 2011). Other components of interest include (Z)-hex-3-en-yl tiglate, which occurs in honeysuckle itself, and (Z)-3,4-epoxyhex-1-yl tiglate, the epoxide of the latter. Another ester of S. humbertiana, (Z)-3,4-epoxy-hex-1-yl acetate, also occurs in the scents of three species of Aerangis (Kaiser 2011). (NV, RG)

Ecology Fig. 751.2.  Distribution map of Sobennikoffia.

The species are epiphytes or lithophytes in humid evergreen forests. They are also terrestrial, semi-epiphytic or epiphytic in dry forests and scrub on sandy soils, often at the foot of trees at elevations of 400–2000 m. (PC)

Anatomy Carlsward et al. (2006a) examined two species of Sobennikoffia. Their results are summarized below.

Leaf TRANSVERSE SECTION

Trichomes multicellular, glandular. Stomata abaxial. Hypodermis absent. Fibre bundles absent. Mesophyll heterogeneous with columnar adaxial cells and isodiametric abaxial cells in S. robusta; isobilateral in S. humbertiana H.Perrier, with columnar cells surrounding a central region of isodiametric cells. Water-storage cells banded. Vascular bundle sheath distinct, cell walls occasionally thickened. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Velamen 2–4 cells thick. Epivelamen cells isodiametric to radially elongate; walls ∪-thickened with fine anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate; cells of the outermost layers often thicker-walled than those of the inner layers. Distinct endovelamen thickenings smooth. Cover cells present over short cells of exodermis. Exodermal long cells primarily ○-thickened. Water-storage cells banded. Aeration units present. Endodermal cells ○-thickened. Vascular cylinder 8–18-arch. Vascular tissue embedded in thin-walled parenchyma or thinly lignified cells in S. humbertiana; embedded in thick-walled sclerenchyma in S. robusta. Pith sclerenchymatous, thin-walled in S. humbertiana. (BC)

Palynology No studies of the pollen of Sobennikoffia have been published. (AP)

Pollination There are no published reports on pollination of any of the species of Sobennikoffia. (AP)

Uses No ethnobotanical uses of these species have been published, and plants are rare in cultivation. (AP)

Taxonomic notes Plants of Sobennikoffia are morphologically reminiscent of some of the larger-flowered species of Angraecum, but the flowers differ in having a prominently trilobed labellum and a rostellum with a recurved midlobe. No DNA sequences of Sobennikoffia have been included in phylogenetic analyses. (PC)

Taxonomic literature Perrier de la Bâthie, H. (1938). Sobennikoffia Schltr. Notulae Systematicae. Paris, 7, 134. Perrier de la Bâthie, H. (1951). Sobennikoffia Schltr. Notulae Systematicae. Paris, 14, 164. Schlechter, R. (1925). Sobennikoffia nov. gen. Repert. Sp. Nov. Regni Veg. Beih., 33, 361-3.

7 5 2 .  S OL ENA NGI S Solenangis Schltr., Beih. Bot. Centralbl., 36, 2: 133 (1918). Type species: Solenangis scandens (Schltr.) Schltr. (basionym: Angraecum scandens Schltr.), lectotype selected here. Celatorchis (R.Rice) R.Rice, Oasis, 3, 15 (2007). Type species: Angraecum conicum Schltr. 435

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Neocribbia Szlach., Ann. Bot. Fennici, 40, 69 (2003). Type species: Neocribbia wakefieldii (Rolfe) Szlach. (basionym: Angraecum wakefieldii Rolfe)

embedded in sclerenchyma, cell walls of embedding sclerenchyma thickest around phloem clusters in S. clavata. Pith sclerenchymatous. (BC)

Derivation of name

Palynology

From the Greek solen, channel or gutter, and angos, vessel, from the flower shape of the species. (PC)

Schill and Pfeiffer (1977) described pollen of S. aphylla, now referable to Microcoelia (q.v.) (AP)

Description (Plate 178; Fig. 752.1)

Cytogenetics

Scandent, epiphytic herbs. Stem flexuous, leafy, with roots along its length. Leaves distichously arranged, well spaced along stem, oblong, unequally bilobed at tip. Inflorescences lateral, axillary, fewflowered, longer than leaves. Flowers creamy white, white, or greenish white; peduncle shorter than rachis. Sepals free, spreading. Petals similar to but smaller than sepals. Labellum entire to trilobed in apical part; spur cylindrical or clavate. Column with an obovoid anther, with a triangular apex; pollinia two, ellipsoidal, porate, attached by a spatulate or tapering tegula to a round to ovate viscidium; stigma transversely oblong-reniform; rostellum deflexed but curving forward toward tip, deeply bifid. (PC)

Distribution (Fig. 752.2) Solenangis is a genus of seven species in tropical Africa. (PC)

Anatomy Carlsward et al. (2006a) examined two species of Solenangis. Their results are summarized below.

Leaf TRANSVERSE SECTION

Trichomes multicellular, glandular, with the basal cell sunken in a raised buttress of epidermal cells in S. clavata (Rolfe) Schltr.; absent in S. wakefieldii (Rolfe) P.J.Cribb & J.Stewart. Stomata abaxial; infrequently sunken in S. clavata. Hypodermis ad- and abaxial, composed of thick-walled fibrous idioblasts occurring singly or in groups of two or three cells forming a single layer interspersed among thin-walled chlorenchyma. Fibre bundles absent. Mesophyll homogeneous. Water-storage cells with birefringent walls infrequent or absent. Vascular bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Velamen two to three cells thick. Epivelamen cells isodiametric in S. wakefieldii; radially elongate in S. clavata; walls ∪-thickened with fine anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate. Distinct endovelamen wall thickenings absent. Cover cells present over short cells of exodermis. Exodermal long cells ∪-thickened in S. wakefieldii, ∩-thickened with radial wall swellings in S. clavata. Water-storage cell walls smooth to pitted. Aeration units present. Endodermal cells ○-thickened. Vascular cylinder 11–20-arch. Vascular tissue 436

Arends et al. (1980) reported a chromosome number of 2n = 50 for both S. clavata and S. scandens. (AP)

Phytochemistry No reports have been found on the phytochemistry of Solenangis. (NV, RG)

Phylogenetics Solenangis was established by Schlechter (1918) as a split from Angraecum and comprised two species, S. clavata and S. scandens, characterized by a habit reminiscent of Oeonia but distinguished by their pollinaria and rostellum structure. Solenangis scandens has been selected here as the type. Solenangis aphylla (Thouars) Summerh., S. conica (Schltr.) L.Jonss., S. cornuta (Ridl.) Summerh., and S. wakefieldii were subsequently added to the genus. However, Carlsward et al. (2006b) included S. clavata, S. wakefieldii, S. cornuta, and S. aphylla in their phylogenetic analyses of Afro-Madagascan angraecoid orchids and showed that the first two formed a clade sister to a clade that comprised Ancistrorhynchus, Bolusiella, and Microcoelia. In contrast, S. aphylla was embedded within Microcoelia with S. cornuta sister to Microcoelia. The last two are now included in a more broadly circumscribed Microcoelia. More recently, Rice (2006a) described three new species of Solenangis and made one new combination allied to S. aphylla, all of which now belong in Microcoelia. In a following paper, Rice (2006b) provided a new infrageneric classification of Solenangis in which he included the genus Dinklageella as a new subgenus. These changes were not based on DNA sequence data. He also moved Rangaeris trilobata Summerh. to Solenangis on the basis of its scandent habit, vegetative morphology, and perianth shape (Rice 2006b). However, its leaves are thin-textured and acute rather than coriaceous and bluntly bilobed, and its pollinarium structure and trilobed rostellum are distinctive. (PC)

Ecology Species of Solenangis are epiphytic or lithophytic in montane forests, sometimes on exotic conifers, at elevations of 1300–2250 m. (PC)

Pollination There is no published information on the pollination biology of Solenangis. (AP)

SOLENANGIS

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T R Fig. 752.1.  A–J. Microcoelia aphylla (Thouars) Summerh. [formerly Solenangis aphylla (Thouars) Summerh.]. A. Habit; B. Flower; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum; G. Column and labellum, side view; H. Column, front view; I. Anther cap; J. Pollinarium. Drawn by Judi Stone from Kew Spirit Collection no. 13810; K–U Solenangis conica (Schltr.) L.Jonss. K. Habit; L. Leaf; M. Flower, side view; N. Dorsal sepal; O. Lateral sepal; P. Petal; Q. Labellum; R. Column and labellum, side view; S. Column, front view; T. Anther cap; U. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Kew Spirit Collection no. 44367. Reproduced with permission from G. V. Pope (ed.) (1998), Flora Zambesiaca: Mozambique, Malawi, Zambia, Zimbabwe, Botswana. Volume 11, Part 2. Royal Botanic Gardens, Kew.

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sheathing base. Inflorescence shorter than or as long as leaves, few- to several-flowered, bracts shorter than pedicel. Flowers pale yellow, pale green, or white with a green mark on labellum. Sepals and petals free, subsimilar, spreading. Labellum entire or trilobed at base, deflexed, ligulate, ecallose, spurred at base; spur dependent, cylindrical or clavate. Column with a hemispherical anther, shortly produced in front, pollinia two, spherical or ovoid, porate, each attached to a linear stipe (tegula) to a single viscidium; stigma broadly reniform; rostellum shortly trilobed, oblong-­subquadrate, deflexed, truncate. (PC)

Distribution (Fig. 753.2) Sphyrarhynchus has a single species known only from Tanzania. (PC)

Anatomy Carlsward et al. (2006a) examined material of this species. Their results are summarized below. Fig. 752.2.  Distribution map of Solenangis.

Uses Apart from being rare in cultivation, there are no known uses of these species. (AP)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Rice, R. (2006a). An overview and three new species of the Solenangis aphylla alliance. Oasis, 3, 2–11. Rice, R. (2006b). Notes on the subtribe Aerangidinae (Orchidaceae). Oasis, 3, 12–15. Schlechter, R. (1918). Versuch einer naturlichen Neuordnung der Afrikanischen angraekoiden Orchidaceen. Beihefte Botanisches Centralblatt, 36, 62–181. Stewart J., Hermans, J., and Campbell, D. (2006). Angraecoid orchids. Timber Press, Portland, Oregon.

753.  SPH Y R A R H Y N C H US Sphyrarhynchus Mansf., Notizbl. Bot. Gart. Berlin, 12, 705 (1935). Type species: Sphyrarhynchus schliebenii Mansf.

Leaf TRANSVERSE SECTION

Trichomes glandular and multicellular with basal cell sunken into raised buttress of epidermal cells. Stomata ad- and abaxial. Hypodermis absent. Fibre bundles absent. Mesophyll homogeneous. Water-storage cells with birefringent walls absent. Large raphide idioblasts situated just below the epidermal surfaces, forming raised areas. Vascular bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Velamen two or three cells thick. Epivelamen cells isodiametric; walls ∪-thickened with fine, anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate. Distinct endovelamen wall thickenings absent. Cover cells present over short cells of exodermis. Exodermal long cell walls primarily ○-thickened to slightly ∩-thickened. Water-storage cell walls thickened along the edges. Aeration units present. Endodermal cells usually ○-thickened to infrequently ∪-­thickened. Vascular cylinder 7–9-arch. Vascular tissue embedded in sclerenchyma. Pith sclerenchymatous. (BC)

Palynology Pollen of Sphyrarhynchus has not been examined. (AP)

Derivation of name

Cytogenetics

From the Greek sphyra, hammer, and rhynchos, nose, in allusion to the shape of the rostellum. (PC)

Brandham (1999) reported a chromosome count of 2n = 50 for the genus. (AP)

Description (Plate 179; Fig. 753.1)

Phytochemistry

Epiphytic or lithophytic herbs. Stems clustered, leafy. Leaves fleshy, conduplicate, linear-elliptic, unequally bilobed at apex, with a

No reports have been found on the phytochemistry of Sphyrarhynchus. (NV, RG)

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Fig. 753.1.  Sphyrarhynchus schliebenii Mansf. A, B. Habit; C. Flower; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum, front view; H. Labellum and spur, side view; I. Column with anther cap, side view; J. Column, anther cap removed, side view; K. Column with anther cap, front view; L. Column, anther cap removed, front view; M. Column and rostellum, front view; N. Anther cap; O. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Maureen E. Church from Kew Living Collection and Danahy s.n. (K). Reproduced with permission from P. J. Cribb (1989), Flora of Tropical East Africa, Orchidaceae (Part 3). A.A. Balkema, Rotterdam/Brookfield.

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Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Summerhayes, V. S. S. (1951). New orchids from Africa. Botanical Museum Leaflets (Harvard University), 14, 215–61. Szlachetko, D. (2003a). Gynostemia orchidalium III. Sphyrarhynchus. Acta Botanica Fennica, 176, 256.

7 5 4 .  S U M M ER H AYES I A Summerhayesia P.J.Cribb, Kew Bull., 3, 284 (1977). Type species: Summerhayesia laurentii (De Wild.) P.J.Cribb (basionym: Angraecum laurentii De Wild.)

Derivation of name The name commemorates Victor Summerhayes (1897–1974), botanist and Curator of the Orchid Herbarium, Royal Botanic Gardens, Kew (1930–1968). (PC) Fig. 753.2.  Distribution map of Sphyrarhynchus.

Phylogenetics Carlsward et al. (2006b) included S. schliebenii in their ITS rDNA study of Afro-Madagascan angraecoid orchids. It was sister to Angraecopsis breviloba Summerh. with 73% support, and that pairing sister to A. amaniensis Summerh. with 99% support. Summerhayes (1951) placed these species, and a third, in his section Coenadenium of Angraecopsis, distinguished by a pollinarium in which the tegulae are attached to a common viscidium rather than each to its own and by their subsimilar sepals and non-triangular petals. Szlachetko (2003a) raised the section to generic rank, as Coenadenium, based mainly on its column structure. Carlsward et al. (2006b) showed that a clade that included this small genus and Sphyrarhynchus is sister to Mystacidium. Based on this information and because of their morphological similarity, Coenadenium is likely to be congeneric with Sphyrarhynchus. Unfortunately, Angraecopsis tenerrima Kränzl., the type of the genus, was not included in the molecular study, and until it is, it is premature to redefine Angraecopsis and Spyrarhynchus. (PC)

Ecology Sphyrarhynchus schliebenii is epiphytic in evergreen forests at elevations of 900–2000 m. (PC)

Description (Plate 180; Fig. 754.1) Epiphytic or lithophytic herbs. Roots basal. Stem leafy. Leaves conduplicate, V-shaped in cross section, distichous, linear, unequally bilobed and erose at tip, articulated at base to sheathing, imbricate, acute bracts. Inflorescence spreading, as long as the leaves or longer, few- to many-flowered; rachis fractiflex. Flowers white, non-resupinate. Sepals and petals subsimilar, spreading to reflexed. Labellum uppermost in flower, entire, concave at base, ovate to subtriangular, ecallose, spurred at base; spur pendent, cylindrical, longer than pedicel and ovary. Column with a beak-like apex; anther obovoid, tapering to erose tip, pollinia two, ovoid, cleft, attached by a spatulate tegula to a slipper-shaped, ovate viscidium; rostellum elongate, deflexed then forward-pointed. Ovary with pedicel longer than floral bracts. (PC)

Distribution (Fig. 754.2) Summerhayesia is a genus of two species in tropical Africa from Liberia and Côte d’Ivoire to Tanzania and south to Zimbabwe. (PC)

Anatomy Carlsward et al. (2006a) examined S. zambesiaca P.J.Cribb. Their results are summarized below.

Leaf TRANSVERSE SECTION

Pollination The pollination biology of Sphyrarhynchus has not been investigated. (PC)

Uses The species has no known uses and is rare in cultivation. (AP) 440

Trichomes absent. Stomata abaxial. Hypodermis ad- and abaxial; composed of fibrous idioblasts in one row interrupted by thinwalled chlorenchyma. Abaxial idioblasts thicker-walled than adaxial idioblasts. Fibre bundles absent. Mesophyll heterogeneous with columnar adaxial cells and isodiametric abaxial cells. Waterstorage cells with birefringent walls absent. Fibrous idioblasts scattered throughout mesophyll, similar in appearance to hypodermal

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Fig. 754.1.  Summerhayesia zambesiaca P.J. Cribb. A. Habit; B. Leaf, transverse section; C. Flower, back view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum, spur removed; H. Column and labellum; I. Column, dorsal view; J. Column, ventral view; K. Anther cap, dorsal view; L. Anther cap, ventral view; M. Pollinarium; N. Pollinia. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Williamson & Drummond 1981 (K) and Kew Spirit Collection no. 40741. Reproduced with permission from G. V. Pope (ed.) (1998), Flora Zambesiaca: Mozambique, Malawi, Zambia, Zimbabwe, Botswana. Volume 11, Part 2. Royal Botanic Gardens, Kew.

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Ecology Summerhayesia laurentii is an epiphyte in lowland and secondary forest up to 800 m. Summerhayesia zambesiaca P.J.Cribb grows epiphytically in dry Brachystegia spiciformis Benth. (Fabaceae) woodlands at elevations of 1300–1500 m. (PC)

Pollination The pollination biology of Summerhayesia has not been studied. (AP)

Uses There are no known uses for these species. (AP)

Taxonomic notes Summerhayesia is a split from Aerangis, readily distinguished by its non-resupinate flowers and distinctive flower in which the viscidium is slipper-shaped. (PC) Fig. 754.2.  Distribution map of Summerhayesia.

idioblasts but generally thinner-walled. Vascular bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Velamen four cells thick. Epivelamen cells isodiametric and thin-walled. Endovelamen cells isodiametric to radially elongate, thin-walled with fine, anastomosing radial wall thickenings. Distinct endovelamen wall thickenings absent. Cover cells present over short cells of the exodermis. Exodermal long cells primarily ○-thickened. Water-storage cells banded. Aeration units absent. Endodermal cells usually ○-thickened to infrequently ∪-­thickened. Vascular cylinder 11-arch. Vascular tissue embedded in sclerenchyma. Pith sclerenchymatous. (BC)

Palynology There are no pollen data for these species. (AP)

Cytogenetics Brandham (1999) listed a chromosome count of 2n = 44 for the genus. (AP)

Phytochemistry No reports have been found on the phytochemistry of Summerhayesia. (NV, RG)

Phylogenetics The phylogenetic relationships of these two species are unknown. Presumably they are allied to Aerangis, but DNA sequences for them have not been published. (AP) 442

Taxonomic literature Cribb, P. J. (1977). New orchids from South Central Africa. Kew Bulletin, 32, 137–87.

7 5 5 .  TA ENI OR R H I ZA Taeniorrhiza Summerh., Bot. Mus. Leafl., Harvard Univ., 11, 166 (1943). Type species: Taeniorrhiza gabonensis Summerh.

Derivation of name From the Greek tainia, worm, and rhiza, root, from the tapeworm-like roots of the type species. (PC)

Description (Fig. 755.1) Leafless, epiphytic herbs. Roots elongate, bilaterally compressed or broadly winged. Stem bearing caudate-acuminate cataphylls. Inflorescence axillary from cataphylls, one-flowered; peduncle bearing two bracts at apex. Flowers brown. Sepals and petals spreading. Sepals ovate-lanceolate, lateral sepals larger than dorsal sepal and oblique. Petals lanceolate-ligulate. Labellum narrow at base then dilated, orbicular-ovate, retuse at tip, spurred at base, the veins branching; spur recurved to almost straight, attenuate above, adnate to narrowly winged column foot. Column truncate with a foot; anther oblong, prolonged in front, pollinia two, obliquely elliptic, tegula spatulate with a linear, incurved claw and strongly dilated above with sides incurved around pollinia, viscidium convex, ovate; rostellum convex, triangular, deeply emarginate with an oblong midlobe in sinus. Ovary and pedicel 2.25 cm long. (PC)

Distribution (Fig. 755.2) Taeniorrhiza is a monospecific genus endemic to Gabon in tropical West Africa. (PC)

TAENIORRHIZA

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K Fig. 755.1.  Taeniorrhiza gabonensis Summerh. A. Habit; B. Flower; C. Base of inflorescence; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Column and labellum, side view; H. Column apex, anther cap removed, side view; I. Column apex, anther cap removed, front view; J. Rostellum remnant; K. Rostellum remnant, cut open; L. Pollinarium, front and back views. Single bar = 1 mm, double bar = 1 cm. Redrawn by Judi Stone after drawing of the type specimen and from drawings in D. Szlachetko and S. Olszewski, Flore du Cameroun. 36. Orchidées, 3, pl. 341, 342 (2001).

Anatomy

Phytochemistry

There is no published information on the vegetative anatomy of this species. (AP)

No reports have been found on the phytochemistry of Taeniorrhiza. (NV, RG)

Palynology Pollen of T. gabonensis has not been studied. (AP)

Cytogenetics Chromosomes of the species have not been counted. (AP)

Ecology The species is an epiphyte in lowland evergreen forest. (PC)

Pollination The pollination biology of Taeniorrhiza has not been investigated. (AP) 443

VA N D E A E

Description (Plate 181; Fig. 756.1) Epiphytic herbs. Roots basal. Stem leafy. Leaves distichously arranged, linear, acute at bilobed apex, with a sheathing, distichous base. Inflorescence axillary, longer than leaves, severalflowered; bracts triangular. Flowers resupinate, green. Sepals and petals linear-tapering, not spreading widely. Labellum concave at base, obscurely trilobed, ecallose, spurred at base; spur upcurved above ovary, cylindrical, much longer than ovary. Column with an oblong anther, with a fringed margin; pollinia two, ellipsoidal, each attached to a stipe (tegula) and that to its own oblongelliptic viscidium, twice as long as tegula; rostellum porrect, three-pronged, central prong shorter than outer ones. Ovary and pedicel geniculate-twisted. (PC)

Distribution (Fig. 756.2) Triceratorhynchus is a monospecific genus from Rwanda, Burundi, Uganda, and Tanzania. (PC)

Anatomy Fig. 755.2.  Distribution map of Taeniorrhiza.

Uses There are no known uses of the species. (AP)

Taxonomic notes Summerhayes (1943) established Taeniorrhiza for a strange, leafless, epiphytic angraecoid orchid from Gabon in West Africa. He noted that its flattened roots resembled those of some Asiatic species of Phalaenopsis. Among African leafless orchids, it is distinctive in having single-flowered inflorescences of brown flowers with a distinctively shaped labellum and column structure. The column structure resembles that of Chauliodon, but the cupshaped apical part of the tegula is different, and the viscidium is massive and fleshy. Its phylogenetic relationships are unknown pending DNA sequencing. Szlachetko (2003a) provided a detailed drawing of its column and pollinarium structure. (PC)

Taxonomic literature Summerhayes, V. S. S. (1943). African orchids XIII. Botanical Museum Leaflets (Harvard University), 11, 137–72. Szlachetko, D. (2003a). Gynostemia orchidalium III. Acta Botanica Fennica 176.

Vegetative anatomy of the species has not been investigated. (AP)

Palynology Pollen of T. viridiflorus has yet to be studied. (AP)

Cytogenetics There are no published chromosome counts for the species. (AP)

Phytochemistry No reports have been found on the phytochemistry of Triceratorhynchus. (NV, RG)

Ecology The species is an epiphyte on Olea welwitschii Gilg & G.Schellenb. (Oleaceae) and Podocarpus L’Hér. ex Pers. (Podocarpaceae) in dense, primary evergreen forest at elevations of 1500–1700 m. (PC)

Pollination Nothing is known of the pollination biology of the species. (AP)

756.  TR IC E R ATO R H Y N CH U S

Uses

Triceratorhynchus Summerh., Bot. Mus. Leafl., Harvard Univ., 14, 232 (1951). Type species: Triceratorhynchus viridiflorus Summerh.

There are no published uses for the species, and it is not in common cultivation. (AP)

Derivation of name From the Greek, treis (tri), three, keras, horn, and rhynchos, nose, in allusion to the triangular shape of the column of the flower. (PC) 444

Taxonomic notes Superficially, this species resembles one of the small, greenflowered species of Angraecum. However, it differs markedly in its column structure, in particular its bizarre rostellum and

TRIDACTYLE

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Fig. 756.1.  Triceratorhynchyus viridiflorus Summerh. A. Habit; B. Flower; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum; G. Column with anther cap, side view; H. Column, anther removed, side view; I. Column, front view, showing rostellum; J. Anther cap; K. Pollinia. Single bar = 1 mm, double bar = 1 cm. Drawn by Maureen E. Church from Eggeling 2058 (K) and Eggeling 5492 (K). Reproduced with permission from P. J. Cribb (1989), Flora of Tropical East Africa, Orchidaceae (Part 3). A.A. Balkema, Rotterdam/Brookfield.

Taxonomic literature Rice, R. (2006b). Notes on the subtribe Aerangidinae (Orchidaceae). Oasis, 3, 12–15.

7 5 7 .  T R I DAC T YL E Tridactyle Schltr., Orchideen, 602 (1914). Type: Tridactyle bicaudata (Lindl.) Schltr. (basionym: Angraecum bicaudatum Lindl.) (vide Summerhayes, Kew Bull., 1948, 282. 1948)

Derivation of name From the Greek treis (tri), three, and daktylos, finger, in reference to the trilobed labellum. (PC)

Description (Plates 182, 183; Fig. 757.1)

Fig. 756.2.  Distribution map of Triceratorhynchus.

pollinarium. Rice (2006b) considered it to belong to his enlarged Mystacidium, but his reasoning, based on its malleate rostellum and papillate tegulae, ignores its other distinguishing features. (PC)

Epiphytic or rarely lithophytic herbs. Stem unbranched or littlebranched, erect to pendent, covered by sheathing leaf-bases. Leaves all along stem, distichous, usually twisted at base to lie in one plane, linear to elliptic or narrowly lanceolate, unequally bilobed at apex, articulated at base to a sheathing leaf-base. Inflorescence emerging through leaf-sheaths opposite leaves, twoto many-flowered; rachis often fractiflex; bracts amplexicaul. Flowers white, yellow, orange, or green, usually scented. Sepals and petals subsimilar, free, spreading. Labellum entire to trilobed, auriculate at base, spurred at base. Column lacking a foot; anther 445

VA N D E A E

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G Fig. 757.1.  A–I, Tridactyle virginea P.J.Cribb & la Croix. A. Habit; B. Flower; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum and spur; G. Column, front view; H. Anther cap; I. Pollinarium. Drawn by Judi Stone from la Croix 862 (K). J–R. Tridactyle flabellata P.J.Cribb. J. Habit; K. Flower; L. Dorsal sepal; M. Lateral sepal; N. Petal; O. Labellum and spur; P. Column, front view; Q. Anther cap; R. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Thomas 3763A (holotype, K).

cap thin-textured, smooth or papillate, pollinia two; tegula one, entire or Y-shaped, viscidium circular or elliptic; rostellum elongated with a tapering apex. Ovary with pedicel often lepidote. (PC) 446

Distribution (Fig. 757.2) Tridactyle comprises about 45 species in tropical and southern Africa. (PC)

TRIDACTYLE

of T. bicaudata, T. crassifolia, and T. furcistipes. Endovelamen cells angular, isodiametric to radially elongate; outer layers composed of thicker-walled cells than inner layers. Distinct endovelamen wall thickenings absent. Cover cells present except in T. tanneri. Exodermal long cells ○-thickened to infrequently ∩-thickened. Water-storage cells with banded wall thickenings or thickenings along cell edges. Aeration units present in T. bicaudata, T. furcistipes, T. scottellii, T. tanneri, and T. tridactylites. Endodermal cells primarily ○-thickened to rarely ∪-thickened. Vascular cylinder 11–33-arch. Vascular tissue usually embedded in sclerenchyma; embedding tissue parenchymatous and thin-walled in T. furcistipes. In T. tridactylites, thin-walled parenchyma forms wings of tissue around phloem clusters. Pith sclerenchymatous. (BC)

Palynology Pollen of Tridactyle has not been examined. (AP)

Cytogenetics Fig. 757.2.  Distribution map of Tridactyle.

Anatomy Carlsward et al. (2006a) examined eight species of Tridactyle. Their results are summarized below.

Leaf TRANSVERSE SECTION

Trichomes multicellular, glandular. Stomata abaxial; ad- and abaxial in T. filifolia (Schltr.) Schltr. and T. tridentata (Harv.) Schltr.; slightly sunken in T. crassifolia Summerh. and T. tridentata. Hypodermis ad- and abaxial; composed of thick-walled fibrous idioblasts occurring singly or in groups of two or three forming in a single row interrupted by mesophyll chlorenchyma. Abaxial idioblasts usually thicker-walled than adaxial idioblasts. Fibre bundles absent. Mesophyll homogeneous in T. bicaudata and T. tridactylites (Rolfe) Schltr.; heterogeneous with columnar adaxial cells and isodiametric abaxial mesophyll cells in T. bicaudata, T. crassifolia, T. filifolia, T. furcistipes Summerh., T. scottellii (Rendle) Schltr., T. tanneri P.J.Cribb, and T. tridentata. Water-storage cell walls smooth to pitted or infrequently banded. Fibrous idioblasts scattered throughout mesophyll in all species examined except T. crassifolia. Vascular bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with phloem and xylem sclerenchyma in T. bicaudata, T. crassifolia, T. filifolia, and T. tridentata; only associated with phloem sclerenchyma in T. bicaudata, T. furcistipes, T. scottellii, T. tanneri, and T. tridactylites.

Root TRANSVERSE SECTION

Velamen 2–5 cells thick. Epivelamen cells radially elongate or rarely isodiametric; walls ∪-thickened with fine anastomosing radial bands; cells thin-walled in T. scottellii and T. tridentata. Tufts of several anticlinally oriented cells scattered along the epivelamen

Arends and Van der Laan (1983) reported chromosome counts of 2n = 46, and Jones (1967) recorded 2n = c. 50, c. 100. Brandham (1999) also reported 2n = 44 and suggested a basic number of x = 23, yielding 2x = 46, the most common number. (AP)

Phytochemistry No reports have been found on the phytochemistry of Tridactyle. (NV, RG)

Phylogenetics Carlsward et al. (2006b) included six species of Tridactyle in their ITS rDNA analysis of angraecoid orchids. These clustered in a clade that also included Listrostachys pertusa, Ypsilopus longifolius, Y. viridiflorus, and Rangaeris amaniensis. However, Tridactyle appears not to be monophyletic: T. bicaudata, T. filifolia, T. crassifolia, and T. scottellii formed a clade sister to Listrostachys pertusa, but T. furcistipes was sister to Ypsilopus longifolius. Both of the last two have a Y-shaped tegula. Tridactyle tanneri was sister to Y. viridiflorus, the two forming a clade sister to one that included Listrostachys, the other Tridactyle species, Y. longifolius, Podangis, and Rangaeris amaniensis. Carlsward et al. (2006b) examined relatively few species, but it seems likely, on the basis of the similar pollinarium and rostellum structure of Ypsilopus to Tridactyle furcistipes and its close allies, that the former should be included within Tridactyle. Rangaeris amaniensis may also eventually be lumped with Tridactyle. Its large white flowers have an obscurely trilobed labellum as in Tridactyle, but its pollinaria and rostellum structure differ. Further examination is required to ascertain its relationship to other Rangaeris species and to Tridactyle. (PC)

Ecology Species of Tridactyle are epiphytic in all types of forests and woodlands, occasionally lithophytic on granite boulders, from sea level to 3000 m. (PC) 447

VA N D E A E

Pollination Johansson (1974) reported chrysomelid beetles of the genus Gabonia as visitors to T. tridentata. There are no substantiated reports of pollination. (AP)

Uses Other than as occasionally cultivated plants, there are no known uses of these species. (AP)

Taxonomic notes Schlechter established the genus and subsequently (Schlechter 1918) listed the species, establishing two subgenera, T. subg. Tridactyle and T. subg. Nephrangis, the latter for Tridactyle filiformis (Kränzl.) Schltr., later raised to generic rank by Summerhayes (1948). The genus is characterized by its small flowers with an entire to trilobed labellum, usually lobed near the tip, auricles on each side of the mouth of the spur, and a pollinarium with an entire (rarely Y-shaped) tegula attached to a small viscidium. (PC)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Kränzlin, F. (1895). Listrostachys filiformis Kränzl. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie, 22, 28. Schlechter, R. (1918). Versuch einer natürlichen Neuordnung der Afrikanischen angraekoiden Orchidaceen. Beihefte zum Botanischen Centralblatt, 36, 62–181. Summerhayes, V. S. (1948). African orchids XVIII. Kew Bulletin, 1947, 301. Szlachetko, D., Sawicka, M., and Kras-Lapinska, M. (2004). 37. Nephrangis (Schltr.) Summerh. In Flore du Gabon 37. Orchidaceae II (ed. P. Morat), pp. 369–72. Muséum National d’Histoire Naturelle, Paris.

758.  Y PSILO P US Ypsilopus Summerh., Kew Bull., 4, 439 (1949). Type species: Ypsilopus longifolius (Kränzl.) Summerh. (basionym: Mystacidium longifolium Kränzl.)

Derivation of name From the Greek letter upsilon (ypsilon) and pous, foot, from the Y-shaped tegula that joins the pollinia to the viscidium. (PC)

Description (Plate 184; Fig. 758.1) Erect to pendent epiphytic herbs. Roots from basal part of stem, not branching. Stem less than 6 cm long. Leaves arranged in a fan, linear, conduplicate or twisted at base to lie in one plane, articulated to persistent leaf bases, leaving sharp, acuminate points when leaves fall. Inflorescence usually shorter than leaves, racemose, few- to many-flowered; peduncle and rachis terete; bracts 448

amplexicaul. Flowers white or pale green, with spreading perianth parts, scabrid on outer surface. Sepals and petals free, subsimilar. Labellum entire or obscurely trilobed, ecallose, spurred at base. Column terete; pollinia two, subglobose, tegula solitary, Y-shaped, viscidium reniform or oblong; rostellum geniculate and recurved at base, bilobed, usually dilated and scutelliform at apex. Ovary scabrid. (PC)

Distribution (Fig. 758.2) Ypsilopus is a genus of five species endemic to tropical East and Central Africa. (PC)

Anatomy Carlsward et al. (2006a) examined two species of Ypsilopus. Their results are summarized below.

Leaf TRANSVERSE SECTION

Leaves of Y. viridiflorus P.J.Cribb & J.Stewart are terete along most of their length, except for the leaf sheath that encircles the stem. Adaxial epidermal characteristics were derived from sections of that leaf sheath. Trichomes multicellular, glandular in Y. viridiflorus; absent in Y. longifolius. Stomata abaxial. Hypodermis ad- and abaxial in Y. longifolius, abaxial only in Y. viridiflorus; composed of fibrous idioblasts occurring singly or in groups of two to several cells, distributed among mesophyll chlorenchyma cells in a single row. Fibre bundles absent. Mesophyll homogeneous in Y. viridiflorus; homogeneous or heterogeneous with columnar adaxial cells and isodiametric abaxial cells in different individuals of Y. longifolius. Water-storage cells with birefringent walls rare. Vascular bundles arranged in a single row with a small vascular bundle situated above the larger midrib bundle in the leaf sheath of Y. viridiflorus. During development, the bundles presumably twist to form a single row of variously oriented vascular strands. In all sections examined, the midrib has rotated 90o from its configuration within the leaf sheath (so the xylem and phloem units of different vascular bundles face the leaf margins). Bundle sheath distinct. Stegmata contain spherical, rough-surfaced silica bodies found in association with vascular bundle sclerenchyma.

Root TRANSVERSE SECTION

Velamen 2–4 cells thick. Epivelamen cells thin-walled and isodiametric to radially elongate in Y. longifolius; isodiametric only in Y. viridiflorus, walls ∪-thickened with fine anastomosing radial bands. Endovelamen cells angular, isodiametric to radially elongate. Cell walls thin to slightly thickened; outer layers often thicker-walled than inner layers. Distinct endovelamen thickenings absent. Cover cells present over exodermal short cells. Exodermal long cells ○-thickened or ∪-thickened. Waterstorage cell walls smooth to pitted or banded. Aeration units present. Endodermal cells ○-thickened. Vascular cylinder 8–15arch. Vascular tissue embedded in sclerenchyma, cell walls of embedding sclerenchyma thickest around phloem groups in Y.

YPSILOPUS

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Fig. 758.1.  Ypsilopus erectus (P.J.Cribb) P.J.Cribb & J.Stewart. A. Habit; B. Flower, side view; C. Flower, front view; D. Leaf apex; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Labellum; I. Column and labellum, side view; J. Column, front view; K. Column, side view; L. Column, anther cap removed, front view; M. Anther cap; N. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Morris s.n. 10/4/1965 (K) and Kew Spirit Collection no. 40739. Reproduced with permission from G. V. Pope (ed.) (1998), Flora Zambesiaca: Mozambique, Malawi, Zambia, Zimbabwe, Botswana. Volume 11, Part 2. Royal Botanic Gardens, Kew.

449

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Uses There are no known uses of the genus, and it is rarely cultivated. (AP)

Taxonomic notes The genus is related to Tridactyle, being distinguished by its short stem, usually white flowers, and Y-shaped tegula. However, all these characters are found to some extent in Tridactyle species. Its relationship to that genus needs further investigation. Tridactyle gentilii De Wild. has white flowers with a long spur. The species of Tridactyle allied to T. furcistipes have short stems and a Y-shaped tegula, both considered characteristic of Ypsilopus. (PC)

Taxonomic literature Carlsward, B. S., Whitten, W. M., Williams, N. H., and Bytebier, B. (2006b). Molecular phylogenetics of Vandeae (Orchidaceae) and the evolution of leaflessness. American Journal of Botany, 93, 770–86. Fig. 758.2.  Distribution map of Ypsilopus.

longifolius. Pith mainly sclerenchymatous, central-most region of Y. longifolius parenchymatous. (BC)

Palynology There are no data on pollen of this genus. (AP)

Cytogenetics Chromosomes of Ypsilopus have not been counted. (AP)

Phytochemistry No reports have been found on the phytochemistry of Ypsilopus. (NV, RG)

Phylogenetics Carlsward et al. (2006b) included Y. longifolius in their ITS rDNA study of angraecoid orchids. It was sister to Tridactyle furcistipes Summerh. with strong support. (AP)

Ecology Ypsilopus erectus (P.J.Cribb) P.J.Cribb & J.Stewart and Y. longifolius are epiphytic in evergreen forests, Juniperus L. and Widdringtonia Endl. (Cupressaceae) forests, Brachystegia Benth. (Fabaceae) and Uapaca Baill. (Euphorbiaceae) woodlands, and on Albizia Durazz. (Fabaceae) and Erythrophloem Benth. (Fabaceae). Ypsilopus viridiflorus is also epiphytic in montane forests. All are found at 1450– 2500 m elevation. (PC)

Pollination The pollination biology of Ypsilopus is unknown. (AP) 450

SU BTRIBE POLYSTAC HYINA E Polystachyinae Schltr., Die Orchideen, 292 (1914). Type: Polystachya Hook.

Description Epiphytic, lithophytic or occasionally terrestrial herbs. Stems often pseudobulbous, rarely caulescent or rhizomatous, caespitose or rarely well spaced on rhizome, one- or more-noded; rhizome simple or branched, creeping. Leaves one to many, suberect, spreading or arcuate, fleshy, coriaceous or thin-textured, persistent or deciduous, linear, ligulate, oblong, elliptic, obovate or oblanceolate, acute, obtuse or acuminate; leaf-base sheathing, often with an articulation. Inflorescence terminal or rarely lateral, racemose, simple or branching, one- to many-flowered; peduncle glabrous or pubescent, often bearing one or more sterile bracts; bracts triangular, ovate, lanceolate or setose, glabrous or pubescent. Flowers usually non-resupinate, often not opening widely, often fragrant, pubescent or glabrous on the outer surface. Dorsal sepal ovate, lanceolate or oblong-elliptic, usually curving over column; lateral sepals usually spreading, oblique at base, generally forming a distinct conical mentum with the column foot, rarely not attached to a column foot. Petals linear, oblong, oblanceolate or obovate, smaller than the sepals. Labellum fleshy, usually trilobed, rarely entire, often strongly recurved, lacking a spur, side lobes erect, midlobe often ovate and decurved, disc often callose, callus present or absent; if present, a raised, fleshy mound or a cushion of farinaceous hairs. Column fleshy, lacking wings, usually bearing a prominent column foot; anther terminal, pollinia two, ovoid, deeply cleft, or four and paired, one large and one small in each pair (in Hederorkis), attached at an acute angle to a square, oblong, linear or triangular tegula and that to a viscidium; rostellum bifid after removal of the pollinarium, not prominent, rarely beak-like.

HEDERORKIS

Ovary circular to triangular in cross-section, pubescent to glabrous. Capsule ellipsoidal. (PC)

Distribution Polystachyinae comprise two genera of some 230 species. The greatest diversity of species is found in sub-Saharan tropical Africa from the Gambia and Sierra Leone across to Ethiopia and Eritrea and south to South Africa. The subtribe is also well represented in Madagascar with some 40 species recorded. A few species are found in the tropical Americas and tropical Asia. (PC)

Phylogenetics Most molecular phylogenetic studies (e.g. van den Berg et al. 2005; Górniak et al. 2010) have indicated that Polystachya is related to Vandeae, and we include it and related genera there, in spite of the differences in habit and floral morphology. Russell et al. (2010) showed that Neobenthamia gracilis Rolfe is embedded in Polystachya and should be treated as Polystachya neobenthamia Schltr. Recent DNA sequencing has demonstrated that Hederorkis is sister to Polystachya (including P. neobenthamia; Russell and Chase, unpublished). (MC)

Taxonomic notes Dressler (1981), Chase et al. (2003), and Szlachetko (2003a) included four genera in Polystachyinae—Hederorkis, Imerinaea, Neobenthamia, and Polystachya. It is worth noting that Szlachetko’s drawing of the gynostemium of Polystachya coelogynochila Kränzl., which he notes as peculiar in the genus, illustrates a Dendrochilum flower. Perrier de la Bâthie (1939) included Imerinaea in Liparidinae, but recent DNA analysis placed it sister to Ansellia in Cymbidiinae (Cribb and Pridgeon 2009). (PC)

Taxonomic literature Chase, M. W., Cameron, K. M., Barrett, R. L., and Freudenstein, J. V. (2003). DNA data and Orchidaceae systematics: a new phylogenetic classification. In Orchid conservation (ed. K. W. Dixon, S. P. Kell, R. L. Barrett, and P. J. Cribb), pp. 69–89. Natural History Publications, Kota Kinabalu, Sabah, Malaysia. Cribb, P. J. and Pridgeon, A. M. (2009). Imerinaea Schltr. In Genera orchidacearum, volume 5. Epidendroideae (Part two) (ed. A. M. Pridgeon, P. J. Cribb, M. W. Chase, and F. N. Rasmussen), pp. 73–5. Oxford University Press. Dressler, R. L. (1981). The orchids: natural history and classification. Harvard University Press. Górniak, M., Paun, O., and Chase, M. W. (2010). Phylogenetic relationships within Orchidaceae based on a low-copy nuclear coding gene, Xdh: congruence with plastid DNA results. Molecular Phylogenetics and Evolution, 56, 784–95. Perrier de la Bâthie, H. (1939). Flore de Madagascar. 49. Orchidacées, 1, 254–6. Russell, A., Samuel, R., Safran, M., Rupp, B., Barfuss, M., and Chase, M. W. (2010). Phylogenetics and cytology of a pantropical orchid genus Polystachya (Polystachyinae; Vandeae; Orchidaceae): evidence from plastid DNA sequence data. Taxon, 59, 389–404.

Schlechter, R. (1924). Imerinaea Schltr. Repertorium Specierum Regni Novarum Vegetabilium Beihefte, 33, 153. Szlachetko, D. (2003a). 1.4. Subtribe Polystachyinae Schltr. Genera orchidalium III, Acta Botanica Fennici, 7–15. van den Berg, C., Goldman, D. H., Freudenstein, J. V., Pridgeon, A. M., Cameron, K. M., and Chase, M. W. (2005). An overview of phylogenetic relationships within Epidendroideae inferred from multiple DNA regions and recircumscription of Epidendreae and Arethuseae (Orchidaceae). American Journal of Botany, 92, 613–24.

Key to the genera of subtribe Polystachyinae (PC)

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●

Plant climbing or scandent; stem rhizomatous, covered by sheaths that deteriorate to fibres; inflorescence lateral and arising from a node on rhizome or near tip of shoot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 759. Hederorkis Plant caespitose; stems caulescent or pseudobulbous; inflorescence terminal, arising from axil of terminal leaf . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 760. Polystachya

7 5 9 .  H EDEROR K I S Hederorkis Thouars, Nouv. Bull. Soc. Philom. n. 19, 319 (1809). Type species: Hederorkis scandens Thouars “Scandederis”, alternative epithet used by Thouars in Orch. Iles. Austral. Afr. (1822), occasionally mistaken for a generic synonym.

Derivation of name From the Greek, hedera, ivy, and orchis, an orchid, in reference to the climbing, ivy-like habit of the type species. (PC)

Description (Fig. 759.1) Epiphytic or lithophytic herbs. Stem scandent, sympodial rhizomatous, covered with imbricate sheaths that disintegrate into fibres. Pseudobulbs absent. Leaves two, divergent, inserted at apex of a short shoot of rhizome, limb flat, fleshy, articulated to a sheath. Inflorescence simple, few-flowered, longer than leaves, lateral and arising from a node on rhizome or near tip of shoot. Flowers non-resupinate with labellum uppermost. Sepals and petals free, lateral sepals oblique, petals slightly smaller than sepals. Labellum free to base, trilobed, ecallose, lacking a spur, sparsely hairy on upper surface, side lobes upcurved at the tips, midlobe larger than side lobes, flat. Column free, elongate, semi-terete and incurved, lacking a foot or foot obscure; anther incumbent, caducous, pollinia four, waxy, in two coherent pairs of one large and one small one, attached by a tegula to a viscidium. Ovary triangular in cross-­section. Capsule ellipsoidal. (PC)

Distribution (Fig. 759.2) The two species of Hederorkis are endemic to the Seychelles and Mauritius. (PC)

Anatomy Vegetative anatomy of these two species has not been studied. (AP) 451

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B

H

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D

F

A

Fig. 759.1.  Hederorkis scandens Thouars. A. Habit; B. Flower; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum; G. Column and labellum; H. Column, anther cap removed, side view. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Vaughn 977 (K).

Palynology

Ecology

Nothing is known about the pollen of Hederorkis. (AP)

Both species are epiphytic or lithophytic in savannas and humid evergreen forests at low to high elevations. (PC)

Cytogenetics Chromosome counts for Hederorkis have not been published. (AP)

Pollination There are no published reports on the pollination of Hederorkis. (AP)

Phytochemistry

Uses

No reports have been found on the phytochemistry of Hederorkis. (NV, RG)

There are no known uses for these species, and they are not in common cultivation. (AP)

452

POLYSTACHYA

s

helle Seyc

Mauritius

Fig. 759.2.  Distribution map of Hederorkis.

Taxonomic notes Thouars (1809, 1822) established Hederorkis for a strange Mauritian endemic with a climbing rhizomatous stem. Rolfe (note in Kew Herbarium) considered it to be a Bulbophyllum, some species of which lack pseudobulbs and have scandent rhizomes, and Hunt (1968) agreed. However, Bosser (1976) pointed out that its column and pollinarium structure indicated a close relationship to Polystachya rather than Bulbophyllum. Subsequent authors (e.g. Dressler 1981) have followed Bosser. The habit closely resembles that of some Asiatic species of Epigeneium. Rasmussen (1979) elucidated the nomenclature of the type species, originally described by Thouars (1809). (PC)

Taxonomic literature Bosser, J. (1976). Le genre Hederorkis Thou. (Orchidaceae) aux Mascareignes et aux Seychelles. Adansonia ser. 2, 16, 225–8. Dressler, R. L. (1981). The orchids: natural history and classification. Harvard University Press. Hunt, P. F. (1968). African orchids: XXXII. Kew Bulletin, 22, 489–92. Rasmussen, F. (1979). Nomenclatural notes on Thouars’ works on orchids. Botaniska Notiser, 132, 385–92. Thouars, A. du Petit. (1809). Extrait de trois mémoires. Nouveau Bulletin des Sciences, Publié par la Société Philomatique de Paris, 1, 314–19. Thouars, A. du Petit. (1822). Histoire particulière des plantes orchidées recueillies sur les trois îles australes d’Afrique, de France, de Bourbon et de Madagascar. Paris.

760.  P O LY STAC H YA Polystachya Hook., Exotic Flora, 2, t.103 (1824), nom. con. Type species: Polystachya luteola Hook. (=Polystachya concreta

(Jacq.) Garay and H.R.Sweet; basionym: Epidendrum concretum Jacq.) Dendrorkis Thouars, Nouv. Bull. Sci. Soc. Philom. Paris, 19, 314 (1809), a superfluous substitute of Dendrobium Sw. Epiphora Lindl., in Hook., Comp. Bot. Mag., 2, 201 (1836). Type species: Epiphora pubescens Lindl. Rhopalorrhachis Klotzsch ex Rchb.f., Cat. Orch.-Samml. Schiller, ed. 1, 62 (1857), nom. invalid. Type species: not designated Neobenthamia Rolfe, Gard. Chron., ser. 3, 10, 272, fig. 33 (1891). Type species: Neobenthamia gracilis Rolfe Nienokuea A.Chev., Compt. Rend. Hebd. Séances Acad. Sci., 220, 634 (1945). Type species: Nienokuea microbambusa (Kränzl.) A.Chev. (basionym: Polystachya microbambusa Kränzl.), lectotype selected here. Disperanthoceros Mytnik & Szlach., Richardiana, 7, 65 (2007). Type species: Disperanthoceros anthoceros (la Croix & P.J.Cribb) Mytnik & Szlach. (basionym: Polystachya anthoceros la Croix & P.J.Cribb), syn. nov. Geerinckia Mytnik & Szlach., Richardiana, 7, 62 (2007). Type species: Geerinckia couloniana (Geerinck & Arbonn.) Mytnik & Szlach. (basionym: Polystachya couloniana Geerinck & Arbonn.), syn. nov. Szlachetkoella Mytnik, Richardiana, 7, 57 (2007). Type species: Szlachetkoella mystacioides (De Wild.) Mytnik (basionym: Polystachya mystacioides De Wild.), syn. nov. Epiphorella Mytnik & Szlach., Richardiana, 8, 13 (2008). Type species: Epiphorella elastica (Lindl.) Mytnik & Szlach. (basionym: Polystachya elastica Lindl.), syn. nov. Unguiculabia Mytnik & Szlach., Richardiana, 8, 19 (2008). Type species: Unguiculabia alpina (Lindl.) Mytnik & Szlach. (basionym: Polystachya alpina Lindl.), syn. nov. Isochilostachya Mytnik & Szlach., Acta Soc. Bot. Poloniae, 80, 80 (2011). Type species: Isochilostachya isochiloides (Summerh.) Mytnik & Szlach. (basionym: Polystachya isochiloides Summerh.), syn. nov. Neoburttia Mytnik, Szlach. & Baranow, Polish Bot. J., 56, 46 (2011). Type species: Neoburttia longiscapa (Summerh.) Mytnik, Szlach. & Baranow (basionym: Polystachya longiscapa Summerh.), syn. nov.

Derivation of name From the Greek polys, many, and stachys, an ear of grain or spike, in allusion to the clustered stems of the type species. (PC)

Description (Plates 185–190; Fig. 760.1–760.4) Epiphytic, lithophytic, or occasionally terrestrial herbs. Stem often pseudobulbous, rarely caulescent, caespitose or rarely well spaced on rhizome, one- or more-noded; rhizome simple or branched, creeping. Leaves suberect, spreading or arcuate, coriaceous to thin-textured, persistent or deciduous, linear, ligulate, oblong, elliptic, obovate or oblanceolate, acute, obtuse or acuminate; leaf base sheathing, often with an articulation. Inflorescence terminal, racemose, simple or branching, one- to many-flowered; peduncle glabrous or pubescent, often bearing one or more sterile bracts. Flowers usually non-resupinate, pubescent or glabrous on outer 453

VA N D E A E

I

J

C

K

G H

F E

D

A

B

Fig. 760.1.  Polystachya cooperi Summerh. A. Habit; B. Flower, front view; C. Flower, lateral sepal and labellum removed, side view; D. Dorsal sepal; E. Petal; F. Labellum, side view; G. Labellum, from within; H. Labellum, longitudinal section; I. Anther with pollinarium, dorsal view; J. Anther, ventral view; K. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Mary Grierson from Cooper 2a (K). Reproduced with permission from F. N. Hepper (ed.) (1968), Flora of West Tropical Africa, Volume 3, Part 1–2. Crown Agents for Overseas Governments and Administrations, London.

surface. Dorsal sepal ovate, lanceolate or oblong-elliptic, usually curving over column; lateral sepals usually spreading, oblique at base, often forming a distinct conical mentum with column foot. Petals linear, oblong, oblanceolate, or obovate. Labellum fleshy, 454

usually trilobed, rarely entire, often strongly recurved, side lobes erect around column, midlobe often ovate and decurved, disc often callose, callus a raised, fleshy mound or a cushion of farinaceous hairs. Column fleshy, lacking wings, usually bearing

POLYSTACHYA

G

H

E

I

F

D

A

B

C

Fig. 760.2.  Polystachya meyeri P.J.Cribb & Podz. A. Habit; B. Portion of inflorescence; C. Flower; D. Flower, opened to show column; E. Dorsal sepal; F. Lateral sepal and column; G. Petal; H. Labellum; I. Anther. Single bar = 1 mm, double bar = 1 cm. Drawn by Maureen E. Church from Harrington 541 (K). Reproduced with permission from P. J. Cribb (1989), Flora of Tropical East Africa, Orchidaceae (Part 3). A.A. Balkema, Rotterdam/Brookfield.

a prominent column foot; anther terminal, pollinia two, ovoid, deeply cleft, attached at an acute angle to a square, oblong, linear, or triangular tegula and that to a viscidium; rostellum bifid after removal of the pollinarium, not prominent, rarely beak-like. Capsule ellipsoidal. (PC)

Distribution (Fig. 760.5) Polystachya is a pantropical genus of some 230 species. The greatest diversity of species is found in sub-Saharan tropical Africa from the Gambia and Sierra Leone across to Ethiopia and Eritrea 455

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G

B

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Fig. 760.3.  Polystachya neobenthamia Schltr. A. Habit; B. Dorsal sepal; C. Lateral sepal; D. Petal; E. Labellum; F. Column, ventral view; G. Anther cap, cut open; H. Pollinarium. Single bar = 1 mm, double bar = 1 cm. Drawn by Loura Mason and Diane Bridson from Thulin & Mhoro 3007 (K), Drummond & Hemsley 1975 (K), and living material as Neobenthamia gracilis. Reproduced with permission from P. J. Cribb (1989), Flora of Tropical East Africa, Orchidaceae (Part 3). A.A. Balkema, Rotterdam/Brookfield.

and south to South Africa. The genus is also well represented in Madagascar with some 40 species recorded. A few species are found in the tropical Americas and tropical Asia across to the Philippines, Taiwan, New Guinea, and northeastern Australia, all belonging to section Polystachya. (PC) 456

Infrageneric classification of Polystachya (PC) Mytnik-Ejsmont (2007, 2008, 2012) and Mytnik-Ejsmont and Szlachetko (2007 a,b; 2008 a,b; 2011) proposed several

POLYSTACHYA

E

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F

Fig. 760.4.  Polystachya praecipitis Summerh. A. Habit; B. Flower, side view; C. Column and labellum, side view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum; H. Column apex, ventral view; I. Anther cap, dorsal view; J. Pollinarium; K. Pollinia. Single bar = 1 mm, double bar = 1 cm. Drawn by Stella Ross-Craig from Burtt 4874 (K). Reproduced with permission from P. J. Cribb (1989), Flora of Tropical East Africa, Orchidaceae (Part 3). A.A. Balkema, Rotterdam/Brookfield.

segregate genera based on DNA-based phylogenetic studies by Russell et al. (2010) and Mytnik-Ejsmont (2011). However, their data do not support the splitting of Polystachya, which forms a robust monophyletic clade if Neobenthamia is included.

Mytnik-Ejsmont (2012) also established several new sections. Undoubtedly, further detailed work is needed before a satisfactory infrageneric classification of the genus can be prepared. 457

VA N D E A E

Comoros Mascarene Islands

Fig. 760.5.  Distribution map of Polystachya.

Polystachya sect. Affines Kränzl., Repert. Sp. Nov. Regni Veg. Beih., 39, 93 (1926). Type species: P. affinis Lindl. Polystachya sect. Aporoideae Kränzl., Repert. Sp. Nov. Regni Veg., 39, 10 (1926), emend Cribb, Kew Bull., 32, 746 (1978). Type species: P. mystacioides Kränzl. Polystachya sect. Bicalcaratae Mytnik, Monogr. Subtribe Polystachyinae: 126 (2011). Type species: P. bicalcarata Kränzl. Polystachya sect. Calluniflorae Kränzl., Repert. Sp. Nov. Regni Veg. Beih., 39, 97 (1926). Type species: P. calluniflora Kränzl. Polystachya sect. Polychaetae P.J.Cribb, Kew Bull., 32, 755 (1978). Type species: P. polychaete Kränzl. Polystachya sect. Caulescentes Kränzl., Repert. Sp. Nov. Regni Veg. Beih., 39, 11 (1926). Type species: P. laxiflora Lindl. Polystachya sect. Cultriformes Kränzl., Repert. Sp. Nov. Regni Veg. Beih., 39, 106 (1926). Type species: P. cultriformis (Thouars) Lindl. Polystachya sect. Dendrobianthe Schltr., Bot. Jahrb. Syst., 53, 568 (1915). Type species: P. dendrobiiflora Rchb.f. Polystachya sect. Dimorphocaules P.J.Cribb, Kew Bull., 32, 747 (1978). Type species: P. microbambusa Kränzl. Polystachya sect. Elasticae Kränzl., Repert. Sp. Nov. Regni Veg. Beih., 39, 97 (1926). Type species: P. elastica Lindl. Polystachya sect. Eurychilae Summerh., Mem. New York Bot. Gard., 9, 80 (1954). Type species: P. eurychila Summerh. Polystachya sect. Galeatae Mytnik, Monogr. Subtribe Polystachyinae: 131 (2011). Type species: P. galeata Rchb.f. 458

Polystachya sect. Humiles Summerh., Bot. Mus. Leafl., Harv. Univ., 10, 284 (1942). Type species: P. ottoniana Rchb.f. Polystachya sect. Isochiloides Summerh., Kew Bull., 2, 128 (1948). Polystachya sect. Polystachya Kränzl., Repert. Sp. Nov. Regni Veg. Beih., 39, 48 (1926). Type species: P. luteola (Sw.) Hook. Polystachya sect. Steudnerae Mytnik, Monogr. Subtribe Polystachyinae: 328 (2011), syn. nov. Type species: P. steudneri Rchb.f. Polystachya sect. Oreochares Mytnik, Monogr. Subtribe Polystachyinae: 214 (2011). syn. nov. Type species: P. oreocharis Kränzl. Polystachya sect. Pseudomonophyllae Mytnik, Monogr. Subtribe Polystachyinae: 328 (2011). Type species: P. rosea Ridl. Polystachya sect. Superpositae Kränzl., Repert. Sp. Nov. Regni Veg. Beih., 39, 100 (1926). Type species: P. superposita Rchb.f. Polystachya sect. Kermesinae P.J.Cribb, Kew Bull., 32, 748 (1978), syn. nov. Type species: P. kermesina Kränzl.

Artificial key to the sections of Polystachya (PC) 1.

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Stems with a single terminal leaf subtending the inflorescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Stems with 2 or more leaves per stem or pseudobulb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4



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Leaf grass-like, acute; rostellum elongate, beak-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. sect. Bicalcaratae

POLYSTACHYA



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Leaf oblong, ovate or ligulate; rostellum not beak-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Rachis pubescent; pseudobulbs 2-noded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. sect. Galeatae Rachis glabrous; pseudobulbs several-noded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. section Cultriformes Plants 80–130 cm tall; labellum entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. section Dendrobianthe Plants usually 70 cm or less tall, often much smaller; labellum usually trilobed, sometimes obscurely so . . . . . . . . . 5 Labellum strongly reflexed near base with basal laminate callus and 1 or 2 parallel cushions of hairs above; leaves absent at anthesis . . . . . . . . . . . . . . . . . . P. sect. Elasticae Labellum not strongly reflexed at base with basal or central callus basal and a raised, fleshy mound or farinose mound; leaves usually present at anthesis . . . . . . . . . . . . 6 Leaves absent at anthesis . . . . . . . . . . . P. sect. Eurychilae Leaves usually present at anthesis . . . . . . . . . . . . . . . . . . . 7 New stems produced successively from middle or upper part of preceding stem or pseudobulb . . . . . . . . . . . . . . . 8 New stems produced from base of previous growth or near base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Stems bamboo-like, erect, swollen at base; sepals narrowly lanceolate, 11–16 mm long . . . . P. sect. Dimorphocaules Stems not bamboo-like, usually pendent to spreading; sepals lanceolate to ovate-lanceolate, less than 8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Flowers laxly arranged on inflorescence, usually 5 mm or more long . . . . . . . . . . . . . . . . . . . . . P. sect. Superpositae Flowers densely arranged on inflorescence, usually 4 mm or less long . . . . . . . . . . . P. sect. Calluniflorae (in part) Pseudobulbs discoid, appressed to substrate; inflorescence pendent to arcuate . . . . . . . . . . . . . . P. sect. Affines Pseudobulbs, if present, not as above; inflorescence erect or suberect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .11 Flowers less than 4 mm long, usually arranged in dense cylindrical inflorescences; sepals setose; bracts setosesubulate . . . . . . . . . . . . . . P. sect. Calluniflorae (in part) Flowers often more than 5 mm long, laxly arranged in the inflorescence; sepals acute but not setose; bracts acute but not setose-subulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Stems not pseudobulbous, usually elongate and cylindrical, covered with leaf bases . . . . . . . . . . . . . . . . . . . . . . . 13 Stems with a basal pseudobulb . . . . . . . . . . . . . . . . . . . . 16 Pseudobulbs leafless at anthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. sect. Polystachya (in part) Pseudobulbs leafy at anthesis . . . . . . . . . . . . . . . . . . . . . 14 Pseudobulbs prominent, bearing apical leaves; peduncle not covered in sheathing bracts . . . . . . . P. sect. Humiles Pseudobulbs not prominent and with a stem-like apex formed from cylindrical leaf bases; peduncle often covered in sheathing bracts . . . . . . . . . . . . . . . . . . . . . . . . . . 15 Stems erect . . . . . . . . . . . . . . . . . . . . P. sect. Caulescentes Stems creeping . . . . . . . . . . . . . . . . . . P. sect. Aporoideae

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Anatomy

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Möbius (1887) described the leaf anatomy of P. pubescens (Lindl.) Rchb.f., and Tominski that of P. concreta (as P. luteola). N.Williams (1979) observed subsidiary cells arrangement in three species. Weltz (1897) examined the stem of P. pubescens. Sanford and Adanlawo (1973), Pridgeon et al. (1983), Pridgeon (1987), and Porembski and Bartlott (1988) reported on the velamen and exodermis of the root in several species. Møller and Rasmussen (1984) did not find stegmata in P. concreta in their survey. Carlsward (2004, 2006a) examined the leaves and roots of four taxa including P. neobenthamia Schltr. (as Neobenthamia gracilis). Their findings are summarized here.

8.

13.

16. Inflorescence often branched; leaves not grass-like; stems erect or ascending; labellum callus with pollen-like hairs . . . . . . . . . . . . . . . . . . . . . . . . P. sect. Polystachya (in part) Inflorescence unbranched; leaves grass-like; stems erect to pendent; labellum callus with clavate hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. sect. Isochiloideae

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TRANSVERSE SECTION

Trichomes rare, glandular, multicellular with sunken basal cell in P. modesta Rchb.f.; absent in P. concreta, P. longiscapa Summerh., and P. neobenthamia Schltr. Stomata abaxial. Hypodermis adaxial, composed of 1–3 layers of large, thinwalled, mucilage-containing water-storage idioblasts. Fibre bundles absent. Mesophyll homogeneous. Water-storage cells with birefringent walls absent. Large, mucilage-containing idioblasts scattered throughout. Vascular bundle sheath distinct. Stegmata absent.

Root TRANSVERSE SECTION

Velamen three or four cells thick. Epivelamen cells isodiametric to tangentially flattened, thin-walled in P. concreta, P. modesta, and P. neobenthamia to ∪-thickened in P. longiscapa. Radial wall thickenings fine and anastomosing. Endovelamen cells isodiametric and rounded; walls thin to evenly thickened. Cells of the outer layer thicker-walled than those of the inner layers in P. neobenthamia. Cover cells absent. Exodermal long cells ∩-thickened in P. concreta and P. modesta, ○-thickened in P. longiscapa and P. neobenthamia. Short cells thinwalled to ∩-thickened. Tilosomes webbed. Water-storage cell walls smooth to banded or thickened along the edges. Mucilage-containing idioblasts present throughout. Aeration units absent. Endodermal cells thin-walled to ○-­thickened. Vascular cylinder 11- to 14-arch. Vascular tissue embedded in parenchyma or sclerenchyma. Pith completely parenchymatous to sclerenchymatous. (BC)

Palynology Schill and Pfeiffer (1977) examined the pollen of seven species of Polystachya, including that of Neobenthamia. All species have convex tetrads with laevigate sculpturing. (AP) 459

VA N D E A E

Cytogenetics

Phylogenetics

Russell et al. (2010) argued that the basic chromosome number for the genus is x = 20, and virtually all diploid counts are 2n = 40. However, 2n = 38, 39, 41 have also been reported, leading Brandham (1999) to suggest that 2n = 40 is more likely tetraploid with x = 10. Polyploidy of 2n = 80, 120 is scattered throughout the genus but especially in P. sect. Polystachya and some Malagasy species. Both diploids and tetraploids have been recorded for Polystachya concreta (Jones 1966; Fedorov 1969; Podzorski and Cribb 1979; Reich 2006; Russell et al. 2010). (AP)

Russell et al. (2010) examined 83 species in 12 of the 15 recognized sections of Polystachya in a plastid DNA-based analysis, which showed that the genus is monophyletic only if Polystachya neobenthamia (syn. Neobenthamia gracilis Rolfe) is included, whereas most of the currently recognized sections (see above) are not monophyletic. Neobenthamia was characterized by its habit, capitate inflorescence, and flowers that lack a column-foot and have an entire lip (Cribb 1984); however, these features also occur in species of Polystachya section Dendrobianthe, in which it truly belongs. Of the 12 sections sampled, only sections Dendrobianthe (including P. neobenthamia) and Isochiloides appear to be monophyletic. (PC)

Phytochemistry Flavonoids present in acid-hydrolyzed leaf extracts of 25 species of Polystachya (including one record as Neobenthamia gracilis) were analyzed by C. Williams (1979). Kaempferol was detected in N. gracilis, P. bicarinata Rendle, P. cultriformis (Thouars) Lindl. ex Spreng., and P. pubescens (Lindl.) Rchb.f., and quercetin in N. gracilis, P. bella Summerh., P. bicarinata, P. caespitifica Kränzl. subsp. latilabris (Summerh.) P.J.Cribb & Podz. (as P. latilabris Summerh.), P. laxiflora Lindl., P. mildbraedii var. mildbraedii (as P. megalogenys Summerh.), P. pubescens, P. tenuissima Kränzl. (as P. ashantensis Kränzl. and P. inconspicua Rendle), and P. zambesiaca Rolfe (as P. hislopii Rolfe). Flavone C-glycosides occur in P. anceps Ridl., P. caespitifica Kränzl., P. concreta (Jacq.) Garay & H.R.Sweet (as P. tessellata Lindl.), P. cultriformis, P. dolichophylla Schltr., P. fallax Kränzl., P. golungensis Rchb.f., P. mildbraedii Kränzl. var. angustifolia (Summerh.) Geerinck (as P. angustifolia Summerh.), P. paniculata (Sw.) Rolfe, and P. supfiana Schltr. A survey of alkaloid content in Orchidaceae revealed that of 15 species of Polystachya tested, none contained these compounds at levels ≥ 0.1% (Lüning 1974). The only named record is for P. estrellensis Rchb.f., which has an alkaloid content < 0.01% (Lüning 1964). Chemical composition data are available for the scent constituents of P. campyloglossa Rolfe, P. cultriformis, P. fallax, and P. mazumbaiensis P.J.Cribb & Podz. (Kaiser 1993). The main components of the P. campyloglossa fragrance are isoprenyl acetate (42.0%), isoprenol (16.0%), prenyl acetate (12.0%), and α-pinene (10.0%). Dihydro-β-ionone and β-ionone are also considered to be important contributors to the scent, although present only as minor components. According to Kaiser (1993), the fragrance of the yellow-flowered form of P. cultriformis is reminiscent of lilyof-the-valley due to a combination of dihydrofarnesol, farnesol, and the major component, (E)-nerolidol (39.5%), and also of lime blossom, to which the minor components 2-aminobenzaldehyde, γ-decalactone, β-ionone, and methyl jasmonate contribute. Two scent components of importance in the case of P. fallax are indole (5.5%) and methyl anthranilate (1.5%). Also present are the esters, methyl 2-methylbutyrate, methyl 2-hydroxy-3-methylvalerate (2 diastereoisomers), methyl 3-methyl-2-oxovalerate, and methyl 2-hydroxy-4-methylvalerate. The major component of the P. mazumbaiensis fragrance is (E)-geranylacetone (45.0%). Minor components of interest include anisyl alcohol, anisyl butyrate, and anisyl caproate, as well as (E)-cinnamic aldehyde, dihydro-β-ionone, and 2-N-methylaminobenzaldehyde (Kaiser 1993). (NV, RG) 460

Ecology Polystachya species are mostly epiphytic or less commonly lithophytic and can be found in most forested and woodland areas within their range. They are absent from drier regions, particularly deserts and semi-deserts. Some species are specific in their host plants, for example, Polystachya dendrobiiflora Rchb.f. and P. johnstonii Rolfe are specific to the stems and branches of Xerophyta Juss. (Velloziaceae). Polystachya songaniensis G.Williamson grows in grass around the margins and cracks of rocky outcrops in southern Malawi. Polystachya neobenthamia grows on Xerophyta species but also on dry, sloping, exposed rock faces or on mossy ledges on precipices. (PC)

Pollination Deceit pollination is the rule in nectarless Polystachya and specifically generalized food deception. Labellar food-hairs (“pseudopollen”) of different types characterize the genus (Davies et al. 2002). The most common type is uniseriate, two- to four-celled, and terminated by a clavate or subclavate cell. Some resemble bristles with tapering terminal cells, and still others are uniseriate and moniliform. Although the trichomes of most species contain protein or rarely starch, Jersáková et al. (2006) maintained that there is no direct evidence that pollinators use pseudopollen for nutrition. Vogel (1978) reported that the pseudopollen of P. pobeguinii (Finet) Rolfe is empty and inedible. Goss (1977) described the pollination of P. flavescens (Blume) J.J.Sm. (=P. concreta) by female halictid bees of the species Dialictus creberrimus. They landed on the hinged lip and lateral sepals and collected pseudopollen for a few seconds, storing it in their scopae, and then backed out of the flower. The pollinarium was deposited on the thorax of the bee. On later visits to another flower, the pollinarium was deposited on the stigma. Goss observed 10% seed-set, attributing none of it to autogamy. Pettersson and Nilsson (1993) documented pollination of P. rosea Ridl. on an inselberg in Madagascar. The species is either terrestrial or epilithic, nectarless, and provides no pseudopollen. Both sexes of halictid bees Lasioglossum emirnense and L. nitididorsatum were pollen vectors in this case of non-model deception. Male bees used the plants as landmarks in the search for a mate. The authors speculated that the deceptive system evolved through density-dependent selection caused by revisiting pollinators that avoid individuals with familiar floral stimuli.

POLYSTACHYA

Van der Cingel (2001) reported facultative autogamy in P. concreta and ‘P. fuliformis’ (presumably P. fusiformis (Thouars) Lindl.) and cleistogamy in P. golungensis Rchb.f. (AP)

Uses In West Africa, pseudobulbs of many species of Polystachya were reportedly used in the preparation of aphrodisiacs (Lawler 1984). Occasionally the showier species such as P. bella Summerh., P. galeata (Sw.) Rchb.f., and P. pubescens are available in the horticultural trade. (AP)

Cultivation Plants can be mounted or grown in pots with a medium of fir bark, charcoal, and lava rock or similar components. Provide bright but filtered light, warm to intermediate temperatures, and ample water and fertilizer during the growing season. Reduce watering in the winter. (AP)

Taxonomic literature Cribb, P. J. (1978). Studies in the genus Polystachya (Orchidaceae) in Africa. Kew Bulletin, 32, 746. Cribb, P. J. (1984). Neobenthamia Rolfe. In Flora of tropical East Africa. Orchidaceae. Part two (ed. R. Polhill), pp. 400–2. Royal Botanic Gardens, Kew. Kränzlin, F. (1926). Monographie der Gattung Polystachya Hook. Repertorium Specierum Regni Novarum Vegetabilium Beihefte, 39, 1–116. Mytnik-Ejsmont, J. (2007). Materials to the revision of Polystachyinae Schltr. (Orchidaceae). 1. Szlachetkoella Mytnik gen. nov. Richardiana, 7, 55–60.

Mytnik-Ejsmont, J. (2008). Materials to the revision of Polystachyinae Schltr. (Orchidaceae). 1. Dendrobianthe (Schltr.) Mytnik & Szlach. gen. nov. Richardiana, 8, 23–31. Mytnik-Ejsmont, J. (2011). A monograph of the subtribe Polystachyinae Schltr. (Orchidaceae). Fundacja Rozwoju Uniwersytetu Gdanskiego, Gdansk. Mytnik-Ejsmont, J. and Szlachetko, D. (2007a). Materials to the revision of Polystachyinae Schltr. (Orchidaceae). 2. Geerinckia Mytnik & Szlach. gen. nov. Richardiana, 7, 61–3. Mytnik-Ejsmont, J. and Szlachetko, D. (2007b). Materials to the revision of Polystachyinae Schltr. (Orchidaceae). 3. Dispanthoceros Mytnik & Szlach. gen. nov. Richardiana, 7, 64–6. Mytnik-Ejsmont, J. and Szlachetko, D. (2008a). Materials to the revision of Polystachyinae Schltr. (Orchidaceae). 4. Epiphorella Mytnik & Szlach. gen. nov. Richardiana, 8, 12–17. Mytnik-Ejsmont, J. and Szlachetko, D. (2008b). Materials to the revision of Polystachyinae Schltr. (Orchidaceae). 5. Unguiculabia Mytnik & Szlach. gen. nov. Richardiana, 8, 18–22. Mytnik-Ejsmont, J. and Szlachetko, D. (2011). Isochilostachya Mytnik & Szlach. gen. nov. a new genus of Polystachyinae Schltr. (Orchidaceae). Acta Societatis Botanicorum Poloniae, 80, 79–85. Russell, A., Samuel, R., Rupp, B., Barfuss, M. H. J., Safran, M., Besendorfer, V., and Chase, M. W. (2010). Phylogenetics and cytology of a pantropical orchid genus Polystachya (Polystachyinae, Vandeae, Orchiaceae): evidence from plastid DNA sequence data. Taxon, 59, 389–404. Summerhayes, V. S. (1942). African orchids: XII. Botanical Museum Leaflets (Harvard University), 10, 284. Summerhayes, V. S. (1948). African orchids: XVII. Kew Bulletin, 2, 128. Summerhayes, V. S. (1954). Plants collected in Nyasaland. Orchidaceae. Memoirs of the New York Botanical Garden, 9, 80–2.

461

A DDENDUM Additional Genera O RCH I D OI D EA E CR A N ICHI DE A E CR A N ICHI DI NAE 761.   GALE O G LO SSUM Galeoglossum A.Rich. & Galeotti, Ann. Sci. Nat. ser. 3, 3, 31 (1845). Type species: Galeoglossum prescottioides A.Rich. & Galeotti Pseudocranichis Garay, Bot. Mus. Leafl., 28, 347 (1982). Type species: Pseudocranichis thysanochila (B.L.Rob. & Greenm.) Garay (basionym: Cranichis thysanochila B.L.Rob. & Greenm.)

Derivation of name From the Greek galea, helmet, and glossa, tongue, likely in reference to the shape of the labellum in the type species. (GS)

Description (Plate 191; Fig. 761.1) Terrestrial, acaulescent herbs. Roots fasciculate, fleshy, cylindrical. Leaves forming a basal rosette, sessile or nearly so, present or absent/wilted at flowering time (depending on the species), rarely subtended by imbricating, brown, papery cataphylls (G. cactorum Salazar & Chávez-Rendón), base shortly sheathing, blade spreading, narrowly oblanceolate, elliptic or obovate, acute to obtuse, green, sometimes with a bluish hue (G. thysanochilum (B.L.Rob. & Greenm.) Salazar). Inflorescence glabrous, partially covered by tubular, loosely adpressed, acute bracts; raceme with several flowers opening successively; floral bracts glabrous, green, ovate to lanceolate, acuminate. Flowers nonresupinate, ascending, white with yellowish green calli and a green central line or the sepals and petals greenish white and the labellum deep yellow with a whitish base (G. tubulosum (A.Rich. & Galeotti) Salazar & Soto Arenas), odourless (G. cactorum), agreeably fragrant during the day (G. thysanochilum) or with unpleasant nocturnal odour (G. tubulosum). Sepals glabrous, free except in G. tubulosum, in which they are connate at their bases and adnate with those of ­petals. Dorsal sepal spreading or recurved, lanceolate, obtuse to

rounded. Lateral sepals erect, spreading or recurved, obliquely lanceolate, acute to obtuse. Petals concave, obliquely oblanceolate, erect, spreading or recurved, acute to obtuse. Labellum free, erect, with involute sides, when spread out rhombic in outline, in natural position with its proximal portion embracing column, canaliculate along midvein, with two elliptical areas of spherical, translucent papillae on outer surface, pubescent on inner surface; base clawed, with a rounded, retrorse lobe at each side, these bearing a marginal/apical nectar gland; apex provided with a distinct lobe, which is denticulate in G. tubulosum but prominent, irregularly dentate-fimbriate and ruffled in remaining species. Column white, obtrapezoid to clavate, footless or provided with an inconspicuous column foot at base, sides smooth or provided with a downcurved, longitudinal flap (G. cactorum); anther dorsal, erect, triangular-ovoid, filament free (G. thysanochilum) or fused with column, pollinarium formed by two hairpin-shaped, narrow, laterally compressed yellow pollinia attached at apex to a common, blackish or greyish viscidium; stigma saddle-shaped, consisting of a central dry (sterile) area and two lateral, ear-like receptive areas, these concave, shiny and sticky, rostellum truncate, upon removal of pollinarium bearing a membranaceous apicule. Ovary ascending, clavate, glabrous, green, tricostate, with inconspicuous pedicel. Capsule obovate. (GS)

Distribution (Fig. 761.2) This is a genus consisting of three species, two of which are endemic to the Mexican state of Oaxaca and the third widespread on most mountain ranges in Mexico and adjacent Guatemala. (GS)

Anatomy Some anatomical features of the roots of G. thysanochilum and G. tubulosum were studied by Figueroa et al. (2008). In both species the uniseriate velamen consists of isodiametric cells. Tilosomes are absent in G. thysanochilum, and the cell walls of the velamen lack thickenings but have pores of various sizes. In contrast, Galeoglossum tubulosum has lamellate tilosomes and distinct, parallel wall thickenings. An additional difference is that in G. thysanochilum the cortex has ‘supraendodermal spaces’, i.e. intercellular spaces roughly the size of one of the surrounding cortical cells immediately outside the endodermis, which are absent in G. tubulosum. Such spaces are more common in species of 463

addendum

B

1 mm

2 mm

L

I

H 5 mm

5 mm

M G

F

E A

10 cm

D J

C 4 mm

K

Fig. 761.1.  Galeoglossum thysanochilum (B.L.Rob. & Greenm.) Salazar. A. Habit; B. Flower from side; C. Labellum and column in natural position with the other floral parts excised; D. Dorsal sepal; E. Petal; F. Lateral sepal; G. Labellum; H. Column from side; I. Column from below; J. Column from above with the anther removed; K. Longitudinal section of column (apex to the left); L. Anther from below (apex to the left); M. Developing capsule. Drawn by Marco Antonio López Rosas from Ramírez 424 (AMO). Reproduced with permission from Soto-Arenas, M. A. y Solano-Gómez, A. R. (2007). Información actualizada sobre las especies de orquídeas del PROY-NOM-059-ECOL-2000 (compiler M. A. Soto-Arenas). Instituto Chinoin A.C., Herbario de la Asociación Mexicana de Orquideología A.C.

various subtribes of Cranichideae in seasonally dry habitats and thus may reflect ecological differences between the species of Galeoglossum. (GS)

Palynology

Cytogenetics

Phytochemistry

No information is available on the cytogenetics of Galeoglossum. (GS)

There are no phytochemical data for the genus. (GS)

464

Nothing is known about the pollen of this genus. (GS)

GALEOGLOSSUM

Fig. 761.2.  Distribution map of Galeoglossum.

Phylogenetics A close relationship between the species formerly treated as Prescottia tubulosa (Lindl.) L.O.Williams and Pseudocranichis thysanochila was first suggested by Salazar et al. (2003) based on similarities in labellum and column morphology, but the lack of material suitable for DNA study precluded the inclusion of P. thysanochila in their molecular phylogenetic assessment of Cranichideae at that time. Subsequently, several molecular phylogenetic analyses have recovered those two species as strongly supported sisters of each other, which made Prescottia paraphyletic (Figueroa et al. 2008; Álvarez-Molina and Cameron 2009; Salazar et al. 2009). Monophyly could be achieved either by sinking P. thysanochila in Prescottia, which would render Prescottia structurally and ecologically heterogeneous, or by removing Prescottia tubulosa from that genus. Salazar (2009) and Salazar et al. (2009) argued for the latter approach, which results in the formal recognition of two monophyletic, easily diagnosable sister genera. Salazar (2009) resurrected Galeoglossum, the oldest generic concept available for the clade encompassing the former Prescottia tubulosa and Pseudocranichis thysanochila. Thus redelimited, Galeoglossum is distinguished by the possession of three putative morphological synapomorphies: apically lobed labellum, stigma consisting

of a central sterile surface with two lateral, receptive areas, and ­pollinarium consisting of two hairpin-like pollinia. A third species fitting this concept of Galeoglossum, G. cactorum Salazar & Chávez-Rendón, was recently discovered. Phylogenetic analysis of plastid matK/trnK and nuclear ITS DNA sequences showed that it is most closely related to G. thysanochilum, in agreement with similarities in floral morphology (Salazar et al. 2011). (GS)

Ecology All the species of Galeoglossum are drought-deciduous terrestrials that live in seasonally dry mountain habitats, shedding their leaves during the peak of the dry season (December to April). Galeoglossum tubulosum grows in a variety of soils, both of igneous and sedimentary origin, in oak and pine-oak forests, occasionally found growing epiphytically in humus accumulation on oak trees. Both G. thysanochilum and G. cactorum are restricted to chalk-rich soils in areas with extensive exposure of limestone. Galeoglossum thysanochilum also occurs in oak and pine-oak forests and has been found in at least one location growing side by side with G. tubulosum. On the other hand, G. cactorum occurs in tall, xerophilous scrub dominated by columnar cacti of the genus Nebuxbaumia Backeb. (Salazar et al. 2011). Flowering of G. thysanochilum and 465

addendum

G. cactorum occurs in October and November, while the leaves are still present, but in G. tubulosum anthesis occurs after the wilting of the leaves from December to April (Salazar 2009). (GS)

Pollination Natural pollination has not been observed for any species of Galeoglossum. In all three species nectar droplets are produced at the base of the labellum, and in G. thysanochilum and G. tubulosum the flowers produce distinctive odours. However, the pleasant floral odour of G. thysanochilum is produced during the daylight hours, whereas that of G. tubulosum is nocturnal and unpleasant, reminiscent of naphthalene or some insecticides (Salazar 2009). Therefore, it is likely that these two species are pollinated by different vectors at different times of the day. This, coupled with differences in flowering time, may contribute to maintenance of species identity when they are sympatric. (GS)

Uses No uses have been recorded for species of Galeoglossum. (GS)

Salazar, G. A., Chávez-Rendón, C., Jiménez, R., and de Ávila. (2011). A new species of Galeoglossum (Orchidaceae, Cranichidinae) from Oaxaca, Mexico. Systematic Botany, 36, 261–7.

ORC H ID EAE SU BTRIBE ORC H ID INA E 7 6 2 .  H EM I P I L I OP S I S Hemipiliopsis Y.B.Luo & S.C.Chen, Novon, 13, 450–3 (2003). Type species: Hemipiliopsis purpureopunctata (K.Y.Lang) Y.B.Luo & S.C.Chen (basionym: Habenaria purpureopunctata K.Y.Lang)

Derivation of name From the orchid genus Hemipilia Lindl. and the Greek suffix -opsis, having the appearance of, like. (LY-B)

Cultivation Galeoglossum is practically unknown in cultivation. Plants of G. tubulosum are relatively undemanding, requiring a well-draining soil mix rich in leaf litter and grown in a cool, brightly lit greenhouse suitable for growing other orchids native to tropical mountain habitats (such as Pleione or many Andean Oncidiinae). A slight reduction in the frequency of watering is advisable after leaves have been shed, but the substrate should not be allowed to dry completely. However, both G. thysanochlum and G. cactorum are difficult to maintain for more than one or two seasons in cultivation, likely because of dependence on unknown physicalchemical properties of their chalky substrate in the wild. (GS)

Taxonomic literature Álvarez-Molina, A. and Cameron, K. M. (2009). Molecular phylogenetics of Prescottiinae s.l. and their close allies (Orchidaceae, Cranichideae) inferred from plastid and nuclear ribosomal DNA. American Journal of Botany, 96, 1020–40. Figueroa, C., Salazar, G. S., Zavaleta, H. A., and Engleman, E. M. (2008). Root character evolution and systematics in Cranichidinae, Prescottiinae and Spiranthinae (Orchidaceae, Cranichideae). Annals of Botany, 101, 509–20. Salazar, G. A. (2009). DNA, morphology, and systematics of Galeoglossum (Orchidaceae, Cranichidinae). In Proceedings of the Second Scientific Conference on Andean Orchids (ed. A. M. Pridgeon and J. P. Suárez), pp. 161–72. Universidad Técnica Particular de Loja, Loja, Ecuador. Salazar, G. A., Cabrera, L. I., Madriñán, S., and Chase, M. W. (2009). Phylogenetic relationships of Cranichidinae and Prescottiinae (Orchidaceae, Cranichideae) inferred from plastid and nuclear DNA sequences. Annals of Botany, 104, 403–16. Salazar, G. A., Chase, M. W., Soto, M. A., and Ingrouille, M. (2003). Phylogenetics of Cranichideae with emphasis on Spiranthinae (Orchidaceae, Orchidoideae): evidence from plastid and nuclear DNA sequences. American Journal of Botany, 90, 777–95.

466

Description (Plates 193, 194; Fig. 762.1) Terrestrial herbs. Plant covered with purple spots throughout. Tuber ellipsoid or subellipsoid, fleshy. Stem usually with 2–5 sterile bracts above leaf. Leaf solitary, fleshy, slanting upward, elliptic to ovate-oblong, acuminate or acute, green with purple spots above, light purple beneath. Inflorescence erect, severalflowered, spotted with purple on rachis and peduncle; bracts ovate-lanceolate, with evident purple spots dorsally; pedicel and ovary with purple spots. Flowers spotted with purple except for labellum. Dorsal sepal erect, oblong, concave, forming a hood together with petals. Lateral sepals obliquely ovate-elliptic, usually reflexed. Petals ovate-orbicular. Labellum subobovate or fan-shaped, trilobed at apex, lobes nearly oblong, lateral lobes broader than central one; spur abruptly contracted in basal part and then strongly expanded to form a globose apex. Column erect; anther with two parallel thecae, pollinia sectile, fragile, each proximally attenuated into a caudicle attached to viscidium; rostellum trilobed, with two separate lateral lobes bearing naked viscidia at apex, lateral lobes extending forward, their apices completely separate, central lobe approximately half as long as anther; stigma in a cavity positioned below rostellum, with two elongated pads. Capsule ellipsoid with purple spots. (LY-B)

Distribution (Fig. 762.2) Hemipiliopsis is a monospecific genus distributed from southeastern Xizang to Assam. (LY-B)

Anatomy Nothing has been published on the vegetative anatomy of this species. (LY-B)

H EMIPILIOPSIS

3 mm

A

B

0.5 mm

C D 0.5 mm

1 mm

3 mm

F 3 mm

E 1 cm

G

Fig. 762.1.  Hemipiliopsis purpureopunctata (K.Y.Lang) Y.B.Luo & S.C.Chen. A. Flower, front view; B. Column; C. Pollinarium; D. Column, longitudinal section; E. Flower, side view; F. Dorsal sepal, petal, lateral sepal, and labellum; G. Habit. Drawn by Chen Li-jun from Liu 5590 (NOCC, National Orchids Conservation Center).

467

addendum

Fig. 762.2.  Distribution map of Hemipiliopsis.

Palynology

Taxonomic notes

Pollen of H. purpureopunctata has not been studied. (LY-B)

Based on vegetative and floral morphology, Lang (1980) suggested that Habenaria (=Hemipiliopsis) purpureopunctata appears to represent an intermediate between Hemipilia and Habenaria, which fall into Dressler’s Orchidinae and Habenariinae, respectively (Luo and Chen 2000). However, floral morphological evidence, such as lacking obvious anther canals and strongly convex or stalked stigmas, does not support a close relationship between Hemipiliopsis and Habenaria (Luo and Chen 2003; Luo et al. 2005). In Hemipiliopsis the trilobed rostellum does not have well-developed lobes, the lateral lobes extend straight forward but not along the walls of the cavity, and the viscidia are positioned directly above the spur mouth (Luo et al. 2005). These floral morphological features, especially those of the column, indicate that Hemipiliopsis is more closely related to Brachycorythis than to Hemipilia (Luo et al. 2005). (LY-B)

Cytogenetics Cytological studies of H. purpureopunctata from Nyingchi, southeastern Xizang, China, revealed a chromosome number of 2n = 42. Interphase nuclei are of the simple chromocentre type. At mitotic prophase, most chromosomes had early condensing segments at the proximal region of one or both arms (Luo 2004). (LY-B)

Phytochemistry There are no reports on the phytochemistry of the species. (LY-B)

Ecology This species grows from 2100 to 3400 m in rocky, humus-rich places in conifer or broad-leaved evergreen forest, alpine oak forest, grassy places, and under shrubs in sandy soil along rivers (Luo 2003). (LY-B)

Uses There are no recorded uses of H. purpureopunctata, and it is not in cultivation. (LY-B) 468

Taxonomic literature Lang, K. Y. (1980). A study on the geographical distribution and floristic features of the Xizang (Tibet) orchid flora. Acta Phytotaxonomica Sinica, 18, 391–407. Luo, Y-B. (2003). The true identity of Habenaria purpureopunctata K. Y. Lang, a distinctive orchid from southeast Xizang and northeast India. Orchid Review, 111, 47–9.

NEOLINDLEYA

Luo, Y-B. and Chen, S-C. (2000). The floral morphology and ontogeny of some Chinese representatives orchid subtribe Orchidinae. Botanical Journal of the Linnean Society, 134, 529–48. Luo, Y-B. and Chen, S. C. (2003). Hemipiliopsis, a new genus of Orchidaceae. Novon, 13, 450–3. Luo, Y-B., Zhu, G-H., and Kurzweil, H. (2005). The gynostemium of Hemipiliopsis purpureopunctata and Senghasiella glaucifolia, two taxonomically disputed species of Habenariinae (Orchidaceae). Botanical Journal of the Linnean Society, 147, 191–7.

763.   NE O LIN D LE YA Neolindleya Kränzl., Orch. Gen. Sp., 1, 651 (1899). Type species: Neolindleya decipiens Kränzl. (= N. camtschatica (Cham.) Nevski)

Derivation of name From the Greek neos, new, and John Lindley (1799–1865), the ‘father of orchid taxonomy’. (PC)

Description (Plates 195, 196; Fig. 763.1) Vigorous, terrestrial herbs. Tubers two (one new), elongate, tapering, hairy. Leaves several, cauline, orbicular to broadly elliptic, green, margins crenulate, sheathing at base. Inflorescence many-flowered; bracts lanceolate, acute, longer than flowers, especially at base of inflorescence. Flowers resupinate, reddish purple. Sepals and petals free. Dorsal sepal oblong-ovate, forming a hood over column with petals; lateral sepals similar but spreading. Petals porrect, hiding column, lanceolate, narrower than sepals. Labellum dependent, oblong to obovate, trilobed at apex, ecallose, spurred at base; spur cylindrical, shorter than labellum and incurved. Column short and broad; anther subquadrate, pollinia two, massulate, cleft, enclosed in a bursicle, with caudicles but lacking viscidia; rostellum a porrect beak; stigma trilobed, deeply concave. Ovary twisted. (PC)

Distribution (Fig. 763.2) Neolindleya is a monospecific genus restricted to far northeastern Russia, Korea, and northern Japan. (PC)

Anatomy Vegetative anatomy of N. camtschatica is unknown. (AP)

Palynology Pollen data on this species have not been published. (AP)

Cytogenetics The chromosome number of this species was recorded as 2n = 36, 38 by Sokolovskaya (1960). (AP)

Phylogenetics Using phylogenetic analyses of ITS nuclear DNA sequences, Bateman et al. (2003) showed that Neolindleya was a member of the Pseudorchis–Galearis clade, which, in turn, was sister to the digitate-tubered Dactylorhiza–Gymnadenia clade. Neolindleya occupied a long branch and was placed sister to Platanthera, albeit without significant bootstrap support. However, in a more recent analysis (Bateman et al., unpublished), it was equally parsimonious as sister to either Platanthera or Galearis. Adding morphological characters to the analysis, notably the tuber, perianth, and gynostemium characters, Efimov et al. (2009) suggested that Neolindleya merits recognition at generic rank on the basis of its unique combination of characters and its chromosome number 2n = 36 or 38 rather than 2n = 42 in other members of the non-tuber-bearing clade to which Galearis belongs, and that it shows significantly stronger similarities to Galearis than to Platanthera. Although the interpretation of the tuber by Efimov et al. (2009) is questionable as material examined at Kew shows a distinctly fleshy, tapering tuber (much more like that found in some Platanthera species), in other characters Neolindleya is unusual. (PC)

Ecology Neolindleya camtschatica is a terrestrial species in a wide range of habitats from partly shaded meadows, scrub, forest margins and open woodland, and also in disturbed habitats such as roadsides where it can reach over a metre tall (Efimov et al. 2009). (PC)

Pollination Efimov et al. (2009) suggested that Neolindleya is autogamous on the basis of its non-functional bursicle and the uniform seed set all along the inflorescence. (PC)

Uses There are no known economic uses of Neolindleya, and it is not in common cultivation. (AP)

Taxonomic notes In the past, Neolindleya has been largely overlooked (Vermeulen 1947, 1972; Dressler 1993; Szlachetko and Rutkowski 2000; Pridgeon et al. 2001) and considered congeneric with Platanthera (Kränzlin 1913; Vyshin 1996; Yabukov and Tchernjagina 2004) or Gymnadenia (Hultén 1927; Ohwi 1965; Voroshilov 1982; Lee 1998) and rarely accepted as a valid genus. It was omitted from the second volume of this series, the intention having been to sink it in Platanthera, which it resembles in habit and tuber shape (Schlechter 1919). However, the recent work of Efimov et al. (2009) indicates that it is not closely related to Gymnadenia and better treated as a distinct genus allied to Galearis. (PC)

Phytochemistry

Taxonomic literature

No reports have been found on the phytochemistry of Neolindleya. (NV, RG)

Bateman, R. M., Hollingsworth, P. M., Preston, J., Luo, Y-B., Pridgeon, A. M., and Chase M. W. (2003). Molecular

469

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B

G

C E

D

H

F

A

Fig. 763.1.  Neolindleya camtschatica (Cham.) Nevski. A. Habit; B. Flower; C. Dorsal sepal; D. Lateral sepal; E. Petal; F. Labellum; G. Column and labellum, side view; H. Column, front view. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Museum Botan. Acad. Petrop. Rec. 11/73 (K) and Riabouchinsky s.n. (K).

phylogenetics and evolution of Orchidinae and selected Habenariinae (Orchidaceae). Botanical Journal of the Linnean Society, 142, 1–40. Dressler, R. L. (1993). Phylogeny and classification of the orchid family. Cambridge University Press. Efimov, P. G., Lauri, R. K., and Bateman, R. M. (2009). Neolindleya Kraenzl., an enigmatic and largely overlooked autogamous orchid from temperate East Asia. Kew Bulletin, 64, 661–71. Hultén, E. (1927). Flora of Kamtchatka and the adjacent islands. Kongliga Svenska Vetenskapsakademiens Handlingar, Ser. 3, 5, 1–346.

470

Kränzlin, F. (1913). Orchidaceae Sibiriae. Russkii Botanicheskii Zhurnal, 3-4, 29–60. Lee, Y. N. (1998). Flora of Korea, ed. 3. Kyohak Publishing, Seoul. Ohwi, J. (1965). Flora of Japan. Smithsonian Institution, Washington, D.C. Pridgeon, A. M., Cribb, P. J., Rasmussen, F. N., and Chase, M. W. (2001). Genera orchidacearum. Volume 2. Orchidoideae (Part one). Oxford University Press. Schlechter, R. (1919). Mittelungen über europäischen und mediterrane Orchideen. II. Repertorium Specierum Novarum Regni Vegetabilis, 16, 257–92.

PSEUDERIA

(basionym: Dendrobium simile Schltr.), type species chosen here.

Derivation of name From the Latin pseudo, false, and Eria, an orchid genus, because of the resemblance of the flowers to those of some Eria species. (PC)

Description (Plate 197; Fig. 764.1) Spreading, climbing, or scrambling terrestrial, or rarely epiphytic, shrubby herbs with leafy, branching stems, up to 8 m long, with basal roots, gripping tree trunks and aiding support of stems. Leaves erect-spreading, coriaceous-papery in texture, commonly acuminate, glabrous, glossy green, smaller towards branch tips. Inflorescence pseudolateral, racemose, one- to four-flowered; peduncle indurate after anthesis; bracts leafy. Flowers not showy. Sepals free, ligulate, ringent; lateral sepals falcate. Petals similar to sepals but smaller. Labellum entire, subrhombic to lanceolate, puberulent on upper surface, lacking a spur; callus at base with two keels confluent above. Column arcuate, semi-terete, lacking a foot, apex denticulate; anther ovate-cucullate, cordate at base, keeled on back; pollinia four, of two sizes, a small and large one connivent in pairs. Capsule cylindrical, topped by remains of floral parts. (PC) Fig. 763.2.  Distribution map of Neolindleya.

Distribution (Fig. 764.2) Szlachetko, D. L. and Rutkowski, P. (2000). Gymnostemia orchidalium 1. Acta Botanica Fennica, 169, 1–379. Vermeulen, P. (1947). Studies on dactylorchids. Schotanus & Jens, Utrecht, Netherlands. Vermeulen, P. (1972). Übersicht zur Systematik und Taxonomie der Gattung Orchis s.s. Jahresbericht des Naturwissenschaftlichen Vereins in Wuppertal, 25, 22–36. Voroshilov, V. N. (1982). A manual of the plants of Soviet Far East. Scientific Publications, Moscow. Vyshin, I. B. (1996). Orchidaceae. Vascular plants of the Russian Far East, 8, 301–39. Saint-Petersburg. Yakubov, V. V. and Tscernjagina, O. A. (2004). A catalogue of the flora of Kamtchatka (vascular plants). Petropavlovsk-Kamtchatskiy.

Pseuderia is a genus of some 18 species in the Moluccas, New Guinea, Micronesia, the Solomon Islands, and Fiji. (PC)

Anatomy Nothing has been published on the anatomy of Pseuderia species. (AP)

Palynology There are no reports on the pollen for this genus. (AP)

Cytogenetics

E PI D EN D ROID EA E

Chromosome counts for Pseuderia have not been published. (AP)

PO D O CHIL E A E

Phytochemistry

E R IINAE 764.   PSE UD E R IA Pseuderia Schltr., Repert. Spec. Nov. Regni Veg. Beih., 1, 644 (1912). Type species: Pseuderia similis (Schltr.) Schltr.

Little is known about the phytochemistry of this genus. Determination of the elemental concentrations of chromium, cobalt, manganese, and nickel in leaf material of Pseuderia foliosa (Brongn.) Schltr. indicates that this species is not a hyperaccumulator, in particular of nickel, for which the content (109 μg/g dry weight) was typical of plants growing on ultrabasic substrates. The study material was sourced from a specimen collected at Obi Island, North Moluccas (Wither and Brooks 1977). (NV, RG) 471

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L

J K

B

H

C D

F A E

G M

I

N Fig. 764.1.  Pseuderia similis (Schltr.) Schltr. A. Habit; B. Flower, side view; C. Flower, front view; D. Dorsal sepal; E. Lateral sepal; F. Petal; G. Labellum; H. Column and labellum, side view; I. Column, anther cap removed, front view; J. Anther cap, oblique view; K. Anther cap, side view; L. Anther cap, dorsal view; M. Pollinia; N. Ovary, transverse section. Single bar = 1 mm, double bar = 1 cm. Drawn by Judi Stone from Mitchell 74 (K) and Kew Spirit Collection no. 50788.

Phylogenetics

Pollination

Yukawa et al. (1993, 1996) included Pseuderia smithiana C.Schweinf. in their DNA-based analysis of Dendrobiinae and demonstrated that it fell outside Dendrobiinae in a clade with Podochilus and Eria (both in subtribe Eriinae). Unfortunately, it was inadvertently excluded from that subtribal account in volume 5 of Genera Orchidacearum. (PC)

There are no reports for pollination of any species. (AP)

Ecology

Taxonomic notes

Plants are usually terrestrial, with stems supported by roots on the trunks of trees in hill forests, at 150–1000 m elevation. (PC)

Pseuderia has a distinctive shrubby habit, climbing up the base of forest trees. In habit, plants resemble some of the shrubby New Caledonian species of Dendrobium. Schlechter (1905) originally

472

Uses There are no known uses for any species of this genus, and they are rare in cultivation. (AP)

PSYCHOPSIELLA

Micronesia

Fig. 764.2.  Distribution map of Pseuderia.

described two New Guinea species in his newly established section Pseudo-Eria of Dendrobium as D. frutex Schltr. and D. simile Schltr. However, in his major treatment of his own New Guinea orchid collections, he raised the section to generic rank as Pseuderia (Schlechter 1911–14), based upon its distinctive habit and flowers that lack a column-foot and have a distinct column structure. Dendrobium simile Schltr. (= Pseuderia similis (Schltr.) Schltr.) is here chosen as the type for Pseuderia. The genus continued to be included in the subtribe Dendrobiinae until recently (e.g. Dressler 1981, 1993; Szlachetko and Margonska 2002). (PC)

C YM BID IEAE

Taxonomic literature

7 6 5 .   PS YC H OP S I EL L A

Dressler, R. L. (1981). The orchids: natural history and classification. Harvard University Press. Dressler, R. L. (1993). Phylogeny and classification of the orchid family. Cambridge University Press. Schlechter, R. (1905). XIII. Pseudo-Eria Schltr. In Nachträge zur Flora der deutschen Schutzgebiete in der Südsee (ed. K. Schumann and K. Lauterbach), p. 174. Leipzig. Schlechter, R. (1911–14). Die Orchidaceen von Deutsch-Neu-Guinea (transl. R. S. Rogers, H. J. Katz, and J. T. Simmons (1982)), Australian Orchid Foundation, Melbourne. Szlachetko, D. L. and Margonska, H. B. (2002). Gynostemia orchidalium. II. Acta Botanica Fennica, 173, 1–-275. Yukawa, T., Kurita, S., Nishida, M., and Hasere, M. (1993). Phylogenetic implications of chloroplast restriction site variation in subtribe Dendrobiinae (Orchidaceae). Lindleyana, 8, 211–23. Yukawa, T., Ohba, H., Cameron, K. M., and Chase, M. W. (1996). Chloroplast DNA phylogeny of subtribe Dendrobiinae

Psychopsiella Lückel & Braem, Orchidee, 33, 7 (1982). Type species: Psychopsiella limminghei (E.Morren ex Lindl.) Lückel & Braem (basionym: Oncidium limminghei E.Morren ex Lindl.)

(Orchidaceae): insights from a combined analysis based on rbcL sequences and restriction site variation. Journal of Plant Research, 109, 169–76.

ON C ID IIN AE

Derivation of name Named in reference of its similarity to the genus Psychopsis Raf., which is from the Greek for butterfly (psyche) and -like (opsis), in reference to the resemblance to butterflies. (MC)

Description (Fig. 537.2, Pl. 131 as Psychopsis limminghei – vol. 5) Small (1–5 cm tall, excluding the inflorescence), epiphytic, caespitose, sympodial herbs with ancipitous, often ridged or 473

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furrowed pseudobulbs, with the flattened surface of the unifoliate pseudobulbs appressed to the substrate. Roots many, thin. Leaves bifacial, conduplicate, articulate, eventually deciduous, tough but not succulent, heavily spotted with purplish red. Inflorescences one or two, lateral, unbranched, longer than leaves, one-(two-)flowered, bracts shorter than pedicels. Flowers resupinate, spurless, yellow with red to red-brown spots over all parts, except for the labellum of which basal lobes are spotted but the broad apical lobe with spots only along margin. Sepals and petals free, oblanceolate, the dorsal sepal sometimes wider. Labellum panduriform (trilobed with a narrow isthmus) with side lobes equal in width to apical lobe; callus a trilobed, tuberculate pad of tissue situated at base, often covered with oil, the apical lobe nose-like and smooth. Column straight, with tabula infrastigmatica; anther terminal, anther cap yellow, pollinarium with a stipe that is shorter and wider than pollinia, caudicles large and irregularly shaped, viscidium ventral, massive and hippocrepiform, terminal on rostellum, pollinia two, yellow, obovate; stigmatic cavity ventral, circular, located just under rostellum, rostellum with a pair of fleshy arms or wings just below stigma and another pair with many digitate apices on sides of stigmatic cavity. Ovary glabrous. Capsules unknown. Seeds unknown. (MC)

Distribution (Fig. 765.1) There is only one species of Psychopsiella, but there are a number of distinctive forms seen in cultivation that could be indicative

of a species complex. The type was recorded as being from near Caracas, Venezuela, but in his treatment of orchids in Flora of Venezuela Foldats (1969–70) stated that he had seen no specimens. That this species is naturally occurring in Venezuela is unlikely, but it is well known in Brazil from the vicinity of Rio de Janeiro. (MC)

Anatomy There are no reports on the vegetative anatomy of this species. (AP)

Palynology Nothing is known of the pollen of P. limminghei. (AP)

Cytogenetics As Oncidium, 2n=56 was reported for P. limminghei (Charanasri and Kamemoto 1975). (MC)

Phytochemistry No reports have been found on the phytochemistry of Psychopsiella. (NV, RG)

Phylogenetics The single species of Psychopsiella and those of Psychopsis were for many years included in Oncidium as members of O. sect. Glanduligera, but they differ from the type group of Oncidium in several ways, particularly their habit, chromosome numbers (of some, but not P. limminghei), and particularly in pollinarium structure. On the basis of DNA data, Psychopsiella and Psychopsis are closely related to Trichopilia in clade C (Fig. A.8, volume 5), making this clade another in which differing floral adaptations precluded an accurate assessment of relationships. However, their leaf texture is remarkably similar to that of Trichopilia, so this relationship has support from their habit. Psychopsiella limminghei has the general habit of Psychopsis, but it grows with pseudobulbs and leaves appressed to the substrate. Chase (2005) transferred P. limminghei to Psychopsis, which was supported by analyses of ITS nuclear ribosomal DNA in which results it was sister to Psychopsis. However, analyses of combined ITS and plastid DNA place P. limminghei (two accessions) as sister to Trichopilia with weak support (73% bootstrap; Neubig et al. 2012). Given this uncertainty, it is more appropriate to recognize Psychopsiella. (MC)

Ecology Psychopsiella grows epiphytically in shady sites up to 800 m in wet Atlantic forests in coastal Brazil. (MC)

Pollination Fig. 765.1.  Distribution map of Psychopsiella.

474

As for Psychopsis, Psychopsiella limminghei exhibits the floral traits associated with the oil-bee syndrome, although there are no published observations to support this. (MC)

PSYCHOPSIELLA

Uses Plants of Psychopsiella are known in orchid collections, and no other uses have been reported. (MC)

Cultivation Psychopsiella should be grown on mounts with a bit of moss tied around the roots under warm conditions with high humidity and low light. Its creeping habit with pseudobulbs and leaves appressed to the medium make pot-culture difficult. (MC)

Foldats, E. (1969–70). Flora de Venezuela: Orchidaceae, vol. 15 (parts 1–5). Instituto Botánico de Venezuela, Caracas. Lückel, E. and Braem, G. (1982). Psychopsis und Psychopsiella: eine alte und eine neue Gattung der Oncidium-Verwandtschaft. Orchidee, 33, 1–7. Neubig, K. M., Whitten, W. M., Williams, N. H., Blanco, M. A., Endara, L., Burleigh, J. G., Silvera, K., Cushman, J. C., and Chase, M. W. (2012). Generic recircumscriptions of Oncidiinae (Orchidaceae: Cymbidieae) based on maximum likelihood analysis of combined DNA datasets. Botanical Journal of the Linnean Society, 168, 117–46.

Taxonomic literature Chase, M. W. (2005). Psychopsis limminghei (Orchidaceae). Curtis’s Botanical Magazine, 22, 53–5.

475

REVISED GENERA O RC H ID OI D EA E C R A N ICHI DE A E S UB TR IBE S P I R A NT H I N AE 122 A .   N O TH O STE LE The following is an update to the treatment of Nothostele published in volume 3. Note its transfer from Cranichidinae to Spiranthinae. Nothostele Garay, Bot. Mus. Leafl., 28, 339 (1982). Type species: Nothostele acianthiformis (Rchb.f. & Warm.) Garay (basionym: Pelexia acianthiformis Rchb.f. & Warm.)

Derivation of name

adnate obliquely to ovary apex, clinandrium concave, expanded laterally at each side of the column, expansion membranaceous, translucent, laterally embracing part of the pollinia; anther dorsal, motile, persistent, membranous, apex thickened, filament free near the apex only, pollinia two, soft, clavate, cleft, yellow, viscidium single, terminal, positioned at apex of column, spheroid to ellipsoid, ventrally with a concavity containing apex of rostellar remnant; stigma ventral, entire, deltoid, convex; rostellum triangular, fleshy, apiculate to elongate and acuminate. Ovary fusiform, sparsely pubescent, brown to greenish. (JB, GS)

Distribution (Fig. 122A.2) Nothostele is a genus of two species restricted to the cerrado biome of central Brazil, in the central region of the state of Minas Gerais and the core area of the same biome in the state of Goiás and the Distrito Federal. (JB, GS)

Anatomy

From the Greek nothos, false, and stele, a column or pillar, descriptive of the column of the type, which lacks fusion between the filament and style. (PC)

There are no reports on the vegetative anatomy of either species. (JB, GS)

Description (Plate 192; Fig. 122A.1)

No information on pollen is available. (JB, GS)

Terrestrial, acaulescent herbs. Roots few, fasciculate, tuberous, fleshy, fusiform to cylindrical, pilose. Leaves one or two, basal, fleshy, orbicular, sessile, adpressed to ground, green, absent at anthesis. Inflorescence terminal, erect, racemose, glabrous below, sparsely pubescent above; peduncle provided with tubular sheaths, these imbricating on lower part of peduncle, spaced above; floral bracts lanceolate, acuminate, longer to about as long as pedicel and ovary. Flowers few, ascending, non-resupinate, white to cream-whitish. Sepals spreading. Dorsal sepal glabrous, lanceolate, narrowly acute to acuminate; lateral sepals shortly connate at base, basally adnate to ovary apex and base of dorsal sepal, forming a spur, sparsely pubescent at base, oblique, recurved. Petals obliquely spatulate to oblanceolate, adherent to dorsal sepal. Labellum free, not adherent to column, sessile, conduplicate-concave, ovate to obovate, base pubescent, apex glabrous, acute to cuspidate, gently recurved to strongly reflexed. Column erect, slightly curved, the ventral surface pubescent at base, convex, dorsal side concave, column foot completely 476

Palynology Cytogenetics Nothing is known about chromosome numbers of this genus. (JB, GS)

Phytochemistry There are no reports on the phytochemistry of Nothostele. (JB, GS)

Phylogenetics The phylogenetic position of Nothostele within tribe Cranichideae has been confusing. Since its description, almost every taxonomist who considered the genus expressed a different point of view regarding its generic or subtribal position. Nothostele acianthiformis was originally described as Pelexia acianthiformis Rchb.f. (Reichenbach 1881). Cogniaux (1893–1896) transferred

NOTHOSTELE

F 1 mm 5 mm

G C

2 cm

E 1 mm

A 5 mm 5 mm

D 1 cm

B

H

Fig. 122A.1.  Nothostele acianthiformis (Rchb.f. & Warm.) Garay. A. Inflorescence; B. leaf; C. Flower, side view; D. Perianth, dissected; E. Column, side view; apex of anther cap has withered; F. Pollinarium, dorsal view; G. Pollinarium, ventral view. Nothostele brasiliaënsis J.A.N.Bat., Meneguzzo & Bianch. H. Perianth, dissected. Drawn by Maria Helena Burian.

it to Stenorrhynchos Rich. ex Spreng. and Hoehne (1945) to Centrogenium Schltr., now a synonym of Eltroplectris Raf. Garay (1982) recognized its distinctive characters and created the genus Nothostele. Although all these authors included the genus in subtribe Spiranthinae, Dressler (1993) and Szlachetko and Rutkowski (2000) considered Nothostele as a member of Cranichidinae based on pollinarium morphology. Szlachetko and Rutkowski (2000) described its viscidium as having ‘a hook-like upcurved apex transformed into a hamulus’. However, a reanalysis of gymnostemium morphology shows that the presence of a hamulus was misinterpreted in Nothostele and that the placement of the genus in Cranichidinae based on morphology of the pollinarium is unwarranted. Another difference between Nothostele and

Cranichidinae s.l. is the distinctive leaves, which are sessile, flattened, adpressed to the ground. In contrast, most Cranichidinae are characterized by petiolate leaves arranged in a basal, upright rosette. A recent phylogenetic analysis based on nuclear and plastid DNA sequence data revealed that Nothostele is deeply embedded in Spiranthinae as the strongly supported sister of Eltroplectris within the Stenorrhynchos clade (Batista et al. 2011). Nothostele and Eltroplectris share some morphological features, but the differences are significant and include: shorter and fewer roots; orbiculate, sessile, fleshy leaves adpressed to the ground; nonresupinate flowers; labellum not adherent to the column; and short spur in Nothostele. Thus, despite some similarities, there are 477

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Leaves develop during the rainy season, from December to April, wither, and are eventually lost as the dry season approaches. During the rainy season the leaves are probably protected from direct sunlight by the surrounding grassy vegetation. Bushfires are common in savanna-like habitats with a marked seasonally dry climate and significantly enhance flowering of several Spiranthinae such as Veyretia sagittata (Rchb.f. & Warm.) Szlach., V. rupicola (Garay) F.Barros, Cyclopogon goyazensis (Cogn.) Schltr., and Sacoila lanceolata (Aubl.) Garay. However, this is probably not the case for Nothostele because the flowering time of Nothostele and the bushfire season are coincident, and it is likely that burnings at this time may be disadvantageous to flowering specimens. The two species in the genus are known only from a few collections and can be considered rare. Known populations consist of few plants in small, loose groups. However, suitable habitats occur in many areas. Plants are more easily located when in flower, and the cerrado is not frequently visited by botanists or collectors during the dry season when the plants are in bloom. Therefore it is likely that the species have been overlooked and may be more common than currently believed. (JB, GS)

Pollination

Fig. 122A.2.  Revised distribution map of Nothostele.

No information is available on the pollination of Nothostele. A population with about a dozen flowering specimens of N. acianthiformis, observed in Serra da Calçada, Minas Gerais in August 2009, did not produce any fruit, indicating low reproductive success and the absence of autogamy. (JB, GS)

Uses noticeable morphological differences that support Nothostele as a genus distinct from Eltroplectris. Nothostele is similar in overall morphology of the inflorescence and flowers to Eltroplectris cogniauxiana (Schltr.) Pabst and E. longicornu (Cogn.) Pabst. Like Nothostele, these species are seasonal geophytes restricted to the cerrado of central Brazil, occur in similar habitats, and flower at the same time. The general similarity between Nothostele and these species of Eltroplectris could represent either the ancestral condition in the Nothostele/Eltroplectris clade or convergence in response to similar ecological conditions and probably pollinators. (JB, GS)

Ecology Nothostele occurs in the cerrado and rocky field vegetation of central Brazil. Climate of the cerrado is marked by two welldefined seasons: a rainy season with an average precipitation of 800–2000 mm from October to April and a sharp dry season during the winter from May to September, when the relative humidity reaches less than 20%. Average annual temperatures are 18–28 °C. Nothostele species are deciduous terrestrials and grow in dry grass– herb–subshrub field (campo sujo) vegetation, often associated with rocky fields (campos rupestres), over shallow, sandy or deep, loamy, well-drained soils, at elevations of 700 to 1300 m. Flowering occurs at the peak of the dry season from July to September, when the plants are leafless. At this time the plants have only a few small, underground, tuberous roots that sustain the inflorescence. 478

No uses have been recorded for Nothostele. (JB, GS)

Cultivation Nothostele is unknown in cultivation. Like other deciduous terrestrial species from dry places, cultivation should try to approach the natural conditions in which plants grow. The soils of most of the cerrado and rocky fields are dystrophic, with low pH. They can be superficially humid during the rainy season but completely dry in the dry season. Plants should be grown under strong but filtered light in well-ventilated areas, and the soil should be well drained and never allowed to accumulate water. They should be watered regularly during the hot and wet summer (the rainy season in Central Brazil) and more sparsely during the dry and cold winter (the dry season). Cerrado vegetation is intolerant of waterlogging, and this is probably also true for Nothostele. (JB, GS)

Taxonomic notes Nothostele species are similar to Eltroplectris cogniauxiana (Schltr.) Pabst and E. longicornu (Cogn.) Pabst, and dried material of these species can be easily confused with each other. They have overlapping distributions, occur in similar habitats, flower at the same time, are leafless at anthesis, and have short inflorescences with few flowers with a white perianth. However, Nothostele has non-resupinate flowers (vs. resupinate in Eltroplectris) and a shorter spur (ca. 3 mm long vs 9–11 mm long in E. cogniauxiana and E. longicornu). (JB, GS)

C YANAEORCHIS

Taxonomic literature Batista, J. A. N., Meneguzzo, T. E. C., Salazar, G. A., Ramalho, A. J., and Bianchetti, L. B. (2011). Phylogenetic placement, taxonomic revision, and a new species of Nothostele (Orchidaceae), an enigmatic genus endemic to the cerrado of central Brazil. Botanical Journal of the Linnean Society, 165, 348–63. Cogniaux, A. (1893–1896). Orchidaceae. In Flora Brasiliensis (eds. C. F. P. Martius, A. G. Eichler, and I. Urban), vol. 3, part 4, 1–672. F. Fleischer, München. Dressler, R. L. (1993). Phylogeny and classification of the orchid family. Cambridge University Press. Garay, L. A. (1982). A generic revision of the Spiranthinae. Botanical Museum Leaflets, Harvard University, 28, 277–425. Hoehne, F. C. (1945). Orchidaceas, Centrogenium. In Flora Brasilica (ed. F. C. Hoehne), fasc. 8, vol. 12(2), 281–289. Secretaria da Agricultura, Indústria e Comércio de São Paulo, São Paulo. Reichenbach, H. G. (1881). VII-Novitiae Orchidaceae Warmingianae. Otia Botanica Hamburgensia, 2, 48–65. Szlachetko, D. L. and Rutkowski, P. (2000). Gynostemia orchidalium. Acta Botanica Fennica, 169, 1–379.

D IS E AE

Cyanaeorchis Barb.Rodr., Gen. Sp. Orchid., 1, 112 (1877). Type species: Cyanaeorchis arundinae (Rchb.f.) Barb.Rodr.

Derivation of name From the Greek Kyane, a water nymph, and orchis, orchid, in allusion to the marshy habitat of the type. (PC)

Distribution (Fig. 458A.1) Cyanaeorchis is a genus of three species distributed from the Espinhaço range in the state of Bahia in northeastern Brazil, to central (Distrito Federal, Goiás, Mato Grosso, Mato Grosso do Sul), southeastern (Minas Gerais, São Paulo), and southern Brazil (Paraná, Santa Catarina, Rio Grande do Sul), Argentina (Chaco and Buenos Aires), and Paraguay (Amambay, Caaguazú, Canindeyú and Misiones). Cyanaeorchis arundinae (Rchb.f.) Barb. Rodr. is the most widespread species covering the whole distribution outlined above. Cyanaeorchis minor Schltr. is known only from southern (Paraná and Rio Grande do Sul) and southeastern (São Paulo) Brazil, and Paraguay (Caaguazú). A third undescribed species is restricted to the cerrado of central (Distrito Federal and Goiás) and southeastern (Minas Gerais) Brazil. The report of Cyanaeorchis for the Amazon is equivocal. ( JB, LB)

Phylogenetics

SUB T RIBE DI S I NA E

Cyanaeorchis has been associated with Eulophia and the corresponding subfamiliar ranks based on general resemblance

14 A .  D ISA The genus Schizodium treated in volume 2 can no longer be maintained following molecular systematic studies by Bytebier et al. (2007) and so is now considered a synonym of Disa. (AP)

Taxonomic literature Bytebier, B., Bellstedt, D. U., and Linder, H. P. (2007). A molecular phylogeny for the large African orchid genus Disa. Molecular Phylogenetics and Evolution, 43, 75–90.

E PI D EN D ROID EA E CY MB IDIE AE CATASE TI NA E 458 A .  C YAN A E O RC H IS The following is an update to the treatment of Cyanaeorchis published in volume 5. Note its transfer from Eulophiinae to Catasetinae.

Fig. 458A.1.  Distribution map of Cyanaeorchis.

479

addendum

of the flowers and deciduous terrestrial habit of Cyanaeorchis with some species of Eulophia. For instance, Dressler (1993) included Cyanaeorchis in subtribe Eulophiinae, as did Pridgeon et al. (2009). However, the resemblance is superficial and there are no unique morphological characters that link the two genera. Furthermore, Eulophia and Eulophiinae are mainly from the Old World, whereas Cyanaeorchis is restricted to the New World. Recent molecular phylogenetic analyses (nrITS, matK/trnK, rbcL; Batista et al., unpublished) revealed that Cyanaeorchis belongs to subtribe Catasetinae, where it is strongly supported as sister to Grobya (100% bootstrap and Bayesian posterior probability support). Inclusion of Cyanaeorchis in Catasetinae will require a new morphological circumscription of the subtribe. However, similarly to the inclusion of Cyanaeorchis in Eulophiinae, there are no apparent morphological characters that link Cyanaeorchis to Grobya or other genera of Catasetinae. Cyanaeorchis is similar in habit to the terrestrial species of Galeandra, also in subtribe Catasetinae, but which usually have pseudobulbs. The only exception is Galeandra beyrichii Rchb.f., which seems to be holomycotrophic, however. Like Cyanaeorchis, most terrestrial species of Galeandra are seasonal geophytes in grassland habitats, and the general similarity between Cyanaeorchis and these species of Galeandra could just represent convergence in response to similar ecological conditions. ( JB, LB)

Ecology Species of Cyanaeorchis are terrestrial in humid grasslands, permanent swamps, and marshes, usually from 600 to 1700 m but also near sea level at latitudes above 27o S. Flowering is mainly from October to January, from the beginning to the peak of the rainy season. In central Brazil, C. arundinae can also flower during the peak of the dry season, from July to September. Flowering of the new, undescribed species from the cerrado of central Brazil is associated with bushfires, and flowering specimens have been found only in areas previously burned during the dry season. Cyanaeorchis arundinae is usually found as few scattered individuals, whereas the new, undescribed species forms compact groups with large number of specimens. Plants of Cyanaeorchis have distinct growth and dormancy phases. A new vegetative shoot and terminal flowers are produced in the rainy season during spring and summer. During the dry season or winter the aerial parts wither, and the plant survives as a short underground stem with roots, which makes possible a new cycle of growth in the following rainy season. ( JB, LB)

Taxonomic notes Based on differences in the size of plants, flowers, and colour of the labellum, two species, Cyanaeorchis arundinae and C. minor, were described in the genus. However, these characters overlap between the two species, and the exact separation between them is not clear. Hoehne (1942) suggested that C. minor could correspond to underdeveloped specimens of C. arundinae but accepted the two species. Small plants from central Brazil were previously identified as C. minor by Pabst and Dungs (1975), but now they are known to represent a third species, which is being described as C. praetermissa (Batista et al., unpublished). ( JB, LB) 480

Taxonomic literature Dressler, R. L. (1993). Phylogeny and classification of the orchid family. Dioscorides Press, Portland, Oregon. Hoehne, F. C. (1942). 101. Cyanaeorchis Barb. Rodr. Flora Brasilica Orchidaceas, 12(6), 43−5, t. 26. Pabst, G. F. J. and Dungs, F. (1975). Orchidaceae Brasilienses. Vol. 1. Schmersow, Hildesheim, Germany. Pridgeon, A. M., Cribb, P. J., Chase, M. W., and Rasmussen, F. N. (2009). Genera orchidacearum. Volume 5. Epidendroideae, Part Two. Oxford University Press.

ON C ID IIN AE 4 9 8 A .  B R A S S I A The following is an update to the treatment of Brassia in volume 5 with the additional synonymization of Ada, Brachtia, and Mesospinidium. Brassia R.Br. in W. T. Aiton, Hortus Kew, ed. 2, v. 215 (1813). Type species: Brassia maculata R.Br. Brachtia Rchb.f., Linnaea, 22, 853 (1850). Type species: Brachtia glumacea Rchb.f. Mesospinidium Rchb.f., Bot. Zeit., 10, 929 (1852). Type species: Mesospinidium warscewiczii Rchb.f. Ada Lindl., Fol. Orch., 5, 1 (1853). Type species: Ada aurantiaca Lindl. Oncodia Lindl., Fol. Orchid., 4 (1853). Type species: Oncodia glumacea Lindl. Brassiopsis Szlach. & Górniak, Biodivers. Res. Conservation, 1-2, 12 (2006). Type species: Brassiopsis keiliana (Rchb.f. ex Lindl.) Szlach. & Górniak Taxa at the rank of section referable to Brassia: Oncidium sect. Brassia (R.Br.) Rchb.f., Ann. Bot. Syst. (Walpers), 6, 1112 (1865). Type species: Oncidium maculatum (R.Br.) Rchb.f.

Description (Plates 82, 83, 85, 86, 112; Fig. 495.1, 497.1, 498.1, 523.1 – vol. 5) Small to large (8–80 cm tall), epiphytic, caespitose, sympodial herbs with often strongly flattened, ancipitous, oval to elongate (and then narrow) pseudobulbs. Roots many. Leaves conduplicate, bifacial, often with several leaf-bearing bracts (1–4) subtending each pseudobulb with one or two apical leaves, articulate, eventually deciduous, epetiolate, some species with paler leaves and darker veins (e.g. former species of Mesospinidium), sometimes 1–3 additional nodes occurring at apex of pseudobulb, each bearing a leaf. Inflorescences 1–3, lateral, glabrous, sometimes branched (i.e. in former species of Mesospinidium), often longer than leaves, 5–30-flowered, bracts often shorter than pedicels, partially covering bases of flowers (particularly in species of former Ada and Brachtia). Flowers resupinate, spurless, cream, yellow, orange-yellow, or greenish yellow. Sepals and petals often free but partially fused in some taxa (lateral sepals fused in former species of Mesospinidium), similar in shape but petals often shorter than sepals, often both ‘spidery’ and elongatecurving, spotless or with dark brown to reddish brown spots

BRASSIA

concentrated on their basal halves. Labellum elongate, sometimes almost hastate, unlobed, basalmost quarter paralleling the column but not adnate to it, forming a cavity that does not produce nectar, apical portion downward- or outward-projecting; callus not complex, typically composed of two parallel ridges forming side walls of basal cavity or a single, simple, raised callus, in which there is sometimes a patch of hairs; in some species there are additional pairs of lumps (1–3) around the basal cavity and additional hairs inside cavity. Column straight to arching, anther terminal, without a terminal hood, but sometimes with a pair of broad, basal arms that extend over basal portions of the labellum callus, forming sides of basal cavity; pollinarium with a well-developed stipe usually longer than pollinia and typically flat but sometimes tubularized and with a swollen apex, caudicles irregular, pollinia two, yellow, elongate pyriform to globose; stigmatic cavity entire, oval, immediately under rostellum, viscidium oval, entire, usually smaller than stipe but nearly as long as stipe in some cases. Ovary glabrous. Capsules tri-nerved. Seeds Maxillaria-type. (MC)

Distribution (Fig. 498A.1) This expanded concept of Brassia comprises 64 species from southern Florida through the Caribbean, Mexico, all of Central America, and through South America to Peru and Bolivia. (MC)

Anatomy Anatomical data have been reported for Ada, Brachtia, Brassia s.s., and Mesospinidium (q.v., vol. 5, Genera Orchidacearum). (MC)

Palynology Pollen features have been reported for Ada, Brassia s.s., and Mesospinidium (q.v., vol. 5, Genera Orchidacearum). (MC)

Fig. 498A.1.  Revised distribution map of Brassia.

Cytogenetics Chromosome counts were published for species of Brassia s.s and the former genus Ada (q.v., vol. 5, Genera Orchidacearum). (MC)

Phytochemistry Phytochemical data have been reported for Brassia s.s. and the former genera Ada and Mesospinidum (q.v., vol. 5, Genera Orchidacearum). (NV, RG)

Phylogenetics Brassia s.l. is a member of clade I and sister to the clade of Cischweinfia, Oliveriana, and Systeloglossum. Some species of Brassia make wide hybrids with other genera of Oncidiinae, such as Oncidium, a member of clade G. Brassia is a particularly polymorphic genus vegetatively, with some species having a single terminal leaf and others having several well-developed foliaceous bracts subtending each pseudobulb and several apical leaves. Many species have extra nodes at the apex of the pseudobulb

that bear a leaf. Pollinarium morphology also is heterogenous. Molecular studies have shown that the genus is monophyletic but should include Ada, Brachtia, and Mesospinidium (Neubig et al. 2012); the single species Brassia (Ada) allenii is sister to all of these species, and Chase and Whitten (2011) transferred the species of Ada, Brachtia, and Mesospinidium to Brassia. Szlachetko and Górniak (2006) published Brassiopsis for the glumacous species, which they segregated from Ada, the type of which is A. aurantiaca, but this transfer does not provide for monophyly, even of the new genus, in which A. aurantiaca is embedded. The earlier name, Oncodia, should have been used instead of describing a new genus, Brassiopsis; the type species of Oncodia is O. glumacea, which Szlachetko and Górniak (2006) transferred to Brassiopsis. (MC)

Ecology Species of Brassia grow epiphytically at a wide range of elevations, from sea level in seasonally dry forests up to 2500 m or more in cloud forests. (MC) 481

addendum

Pollination Dodson (1990) described pollination of species of Brassia by females of the wasp genera Pepsis and Campsomeris, which hunt spiders and paralyze them by stinging, after which they take them to a nest and lay eggs on them. The wasp larvae then eat the paralyzed spiders. These wasps seem to mistake Brassia flowers for spiders, clasp the base of the column and sting the surface of the labellum, after which they try to drag away their ‘paralyzed prey’, during which process pollinia are attached to their heads. Similar pollination has been attributed to the former species of Ada, which have similar flowers, but B. (Ada) aurantiaca L.O.Williams ex C.Schweinf. with its brightly-coloured flowers is likely to be hummingbird-pollinated (van der Pijl and Dodson 1966). (MC)

Uses Other than frequent cultivation as an ornamental, no uses are reported. (MC)

Pachyphyllum Kunth, Nov. Gen. Sp. Pl., 338, t. 77 (1816). Type species: Pachyphyllum distichum Kunth Centropetalum Lindl., Sert. Orch., sub. t. 21 (1839). Type species: Centropetalum distichum Lindl. (= Fernandezia subbiflora Ruiz & Pav.) Nasonia Lindl., Benth. Pl. Hartw., 150 (1844). Type species: Nasonia punctata Lindl. (= Fernandezia maculata Garay & Dunst.) Orchidotypus Kränzl., Bot. Jahrb. Syst., 37, 383 (1906). Type species: Orchidotypus muscoides Kränzl. Raycadenco Dodson, Icones Orch. Trop., ser. 2, t. 157 (1989). Type species: Raycadenco ecuadorensis Dodson

Derivation of name The name is dedicated to Gregorio García Fernández, an 18th century Spanish botanist and contemporary of Ruiz and Pavón. (MC)

Description (Plates 101, 125, 134; Fig. 513.1, 532.1, Cultivation Species of Brassia are frequently seen in cultivation; culture in pots with a well-draining medium is effective, and the plants often become large and free-flowering. Since the species come from a wide range of habitats, culture should be modified to fit their provenance. Some species, such as B. caudata (L.) Lindl. and B. maculata R.Br., grow at low elevations and are warm-growing, whereas others such as B. signata Rchb.f. come from much higher elevations and require intermediate-cool conditions. In general, Brassia species are of easy culture, and their hybrids with other genera, such as Miltonia, are commonly grown. (MC)

Taxonomic literature Chase, M. W. and Whitten, W. M. (2011). Further taxonomic transfers in Oncidiinae (Orchidaceae). Phytotaxa, 20, 26–32. Dodson, C. H. (1990). Brassia. In Native Colombian orchids, Vol. 1 (ed. R. Escobar), pp. 52–3. Editorial Colina, Medellín. Neubig, K. M., Whitten, W. M., Williams, N. H., Blanco, M. A., Endara, L., Burleigh, J. G., Silvera, K., Cushman, J. C., and Chase, M. W. (2012). Generic recircumscriptions of Oncidiinae (Orchidaceae: Cymbidieae) based on maximum likelihood analysis of combined DNA datasets. Botanical Journal of the Linnean Society, 168, 117–46. Garay, L. A. (1973). El género Mesospinidium. Orquideología, 8, 165–78. Sweet, H. (1974). El género Brachtia Rchb.f. Orquideología, 9, 3–20. Szlachetko, D. L. and Górniak, M. (2006). New taxa in the subtribe Oncidiinae (Orchidaceae). Biodiversity Research and Conservation, 1-2, 11–13. Williams, N. H. (1972). A reconsideration of Ada and the glumaceous brassias (Orchidaceae). Brittonia, 24, 93–110.

513 A .   FE R N A N D E Z IA The following is an update to the treatment of Fernandezia published in volume 5 with the additional synonymization of Orchidotypus, Pachyphyllum, and Raycadenco. Fernandezia Ruiz & Pav., Prod. Flor. Peruv et Chil., 123, t. 27 (1794). Type species: Fernandezia subbiflora Ruiz & Pav. 482

541.1 – vol. 5) Small to moderate (up to 15 cm) epiphytic, caespitose, erect to upwardly arching, monopodial herbs without pseudobulbs. Roots many, thin. Leaves 4–20, bifacial, articulate, lasting for 3–5 years. Inflorescences 1–3, lateral, glabrous, unbranched, usually only about the same length as leaves, one- or two-flowered, bracts shorter than pedicels and often buried in leaf bases. Flower orientation irregular, but mostly resupinate, unspurred, in some cases not opening completely, green, greenish white, cream, bright red, red-orange, or yellow, some of the last also with reddish brown maculations. Sepals and petals free to fused into a tube, obovate to oblanceolate, similar in size and shape. Labellum triangular to lanceolate, lobed to unlobed, in some cases hastate to panduriform; callus a bilobed, sometimes tuberculate pad in the middle of the basal portion, glabrous to pubescent. Column straight, a tabula infrastigmatica present in F. ecuadorense; anther terminal, usually with a large terminal hood or pair of expanded arms on the sides of the stigmatic cavity, with two types of pollinaria, one (in Fernandezia s.s. and the former Raycadenco) with a pair of stipes that are much longer than the pollinia and small caudicles, the other (in the former species of Pachyphyllum) with long caudicles and no stipe, viscidium oval to almost wedge-shaped, entire, ventral, pollinia two, yellow, pyriform; stigmatic cavity ventral, just under rostellum, sometimes occluded by rostellum, oval. Ovary glabrous. Capsules triangular. Seeds Maxillaria-type. (MC)

Distribution (Fig. 513A.1) There are 51 species of Fernandezia, most of which occur in the higher elevations of the Andes from Venezuela, Colombia, Ecuador, and Peru, with a few species in upper montane regions of Costa Rica and Panama (Central America), and one in Mexico (North America). (MC)

Anatomy Anatomical data have been reported for Fernandezia (q.v., vol. 5, Genera orchidacearum). (MC)

NOTYLIOPSIS

pollination syndromes and are almost identical in habit. In the results of Neubig et al. (2012), Pachyphyllum and Fernandezia are interdigitated, and Raycadenco is sister to this clade. The last could be kept separate, but if we exclude the characters associated with pollinator syndromes, then there is nothing to keep them distinct. (MC)

Ecology These species grow at high elevation (usually above 2000 m) in cloud-forest conditions. (MC)

Pollination The general floral morphology and colours of the yellow- to red-flowered species are compatible with Dressler’s (1981) suggestion that Fernandezia is hummingbird-pollinated, but there are no direct observations to support this. The former single species of Raycadenco exhibits the floral features associated with oil-bee pollination. Those of former Pachyphyllum are less clear based on their floral traits; some set every capsule, indicating that selfpollination may be taking place. (MC)

Uses These species are rarely seen in cultivation, and no other uses have been reported. (MC)

Cultivation These species are probably best grown in pots with a moistureretentive medium under cool conditions. (MC)

Taxonomic literature Fig. 513A.1.  Revised distribution map of Fernandezia.

Palynology Nothing is known about pollen of this genus. (MC)

Dressler, R. L. (1981). The orchids: natural history and classification. Harvard University Press. Neubig, K. M., Whitten, W. M., Williams, N. H., Blanco, M. A., Endara, L., Burleigh, J. G., Silvera, K., Cushman, J. C., and Chase, M. W. (2012). Generic recircumscriptions of Oncidiinae (Orchidaceae: Cymbidieae) based on maximum likelihood analysis of combined DNA datasets. Botanical Journal of the Linnean Society, 168, 117–46.

Cytogenetics Chromosome numbers of this genus have not been reported. (MC)

5 2 7 A .  NOT YL I OP S I S

Phytochemical data, though limited, have been published for the former genus Pachyphyllum (q.v., vol. 5, Genera Orchidacearum). (NV, RG)

The following is an update to the treatment of Notyliopsis published in volume 5. Notyliopsis P.Ortiz, Orquideología, 20, 184 (1996). Type species: Notyliopsis beatricis P.Ortiz Santanderella P.Ortiz, Orquideología, 27, 169 (2011). Type species: Santanderella amadorinconiana P.Ortiz

Phylogenetics

Derivation of name

Fernandezia was previously considered a member of Pachyphyllinae (Dressler 1981), but it was found to be embedded in Oncidiinae, in which subtribe they are treated here (clade E). Fernandezia, Pachyphyllum, and Raycadenco differ only in their

From the generic name Notylia and the Greek -opsis, appearance, in reference to similarity of these species to those of Notylia, although the two genera are not related based on DNA studies. (MC)

Phytochemistry

483

addendum

Description (Plate 116; Fig. 527.1 – vol. 5) Small (10–18 cm tall) epiphytic, caespitose, sympodial herbs with oblong pseudobulbs. Roots many, thin. Leaves one, bifacial, conduplicate, articulate. Inflorescences 1–3, lateral, glabrous, unbranched, pendent, as long or longer than the leaves, 5–15-flowered, bracts shorter than pedicels. Flowers resupinate, cream to whitish, with rose-purple suffusion in the base of the labellum. Sepals and petals free, except for lateral sepals fused to more than ¾ their length in N. beatricis, all elliptic-lanceolate but petals narrower. Labellum trilobed with the side lobes rolled upwards and an acute apex, lamina flatter in N. amadorinconiana (P.Ortiz) M.W.Chase; callus absent (N. amadorinconiana) or a complex structure composed of a raised pad on the base of the labellum that extends out in two flanges toward the apex and in front of which are two, tooth-shaped structures. Column arcuate with a much swollen apex, with or without wings or arms; anther dorsal, anther cap glabrous, pollinarium with a flat stipe longer than the pollinia and expanded apically, caudicles small, viscidium ventral, oval, entire, pollinia two, yellow, highly flattened, discoid; stigmatic cavity slit-like, just below anther, rostellum long. Ovary glabrous. Capsules not seen. Seeds not seen. (MC)

Distribution (Fig. 527A.1) There are two species of Notyliopsis, N. beatricis, known only from Chocó, and N. amadorinconiana, known only from Floridablanca, Santander Department, both in Colombia. (MC)

Anatomy Vegetative anatomy of these species has not been studied. (AP)

Palynology Nothing is known about pollen of this genus. (MC)

Cytogenetics Chromosomes of Notyliopsis have not been counted. (MC)

Phytochemistry No reports have been found in phytochemistry of Notyliopsis. (NV, RG)

Phylogenetics In spite of its name, Notyliopsis is not closely related to Notylia. It is not a member of the twig-epiphyte subclade but instead sister to Gomesa. Notyliopsis amadorinconiana, which was recently described (Ortiz 2011) as the monospecific Santanderella, was also specifically referred to as being morphologically similar to Notylia. The two species differ in habit (slightly), fusion of the lateral sepals, and labellum shape, but they share the details of their column and pollinaria. In analyses of DNA data, Quintanilla-Quintero et al. (2011) found that Notyliopsis amadorinconiana (treated as Santanderella) was sister to Notyliopsis beatricis 484

Fig. 527A.1.  Revised distribution map of Notyliopsis.

with a bootstrap percentage of 94, but then in the discussion they did not consider that it might be placed in Notyliopsis. This seemed to be considered as an ‘outside cluster’, and their results were not considered as suggesting an alternative generic placement for N. amadorinconiana. Chase (unpublished) included the ITS sequence of the new species in the matrix used for the analysis presented in volume 5 of Genera Orchidacearum and confirmed that (with far more outgroups included) this species is sister to N. beatricis. Given the similarities of their columns and pollinaria, this is enough evidence to combine Santanderella with Notyliopsis. (MC)

Ecology Notyliopsis beatricis occurs on the western slopes of the western cordillera in Colombia at around 1300 m. Notyliopsis amadorinconiana was reported in the original description to grow at 1800– 1850 m on Eriobotrya japonica (Thunb.) Lindl. (Rosaceae), but this is not a native tree, leaving in doubt the condition under which it naturally occurs. (MC)

GONGORA

Pollination Pollinators for Notylia are male euglossine bees (van der Pijl and Dodson 1966; Dodson 1967) that attempt to collect floral fragrance compounds. Given the floral similarity of Notyliopsis to Notylia, particularly in the details of the pollinarium and slitshaped stigma, it might be anticipated that these two species are also pollinated by male euglossine bees. (MC)

Uses Plants of Notyliopsis are occasionally seen in cultivation, and no other uses are reported. (MC)

Cultivation Plants of Notyliopsis should be grown on mounts with high humidity and moderate light with a constant watering regime throughout the year. Temperature should be intermediate. (MC)

Taxonomic literature Ortiz, P. (1996). Notyliopsis, un nuevo género afín a Notylia. Orquideología, 20, 184. Ortiz, P. (2011). Santanderella, a Colombian new genus in the Oncidiinae (Orchidaceae). Orquideología, 28, 174–8. (printed as 2011) Quintanilla-Quintero, S., Ortiz, P., Bernal, J. E., and Gómez, A. (2011). Phylogenetic relationships among genera of the subtribe Oncidiinae (Epidendroideaee; Orchidaceae) and a new genus: Santanderella. Phytologia, 93, 388–406.

569 A .  G O N G O R A The following is a revision of the distribution map of Gongora published in volume 5.

Fig. 569A.1.  Revised distribution map of Gongora.

Note added in proof Some genera accepted on the World Checklist of Selected Plant Families () have not appeared in Genera Orchidacearum for one reason or another, usually because they were published after their respective volume appeared or not enough is known about them. They are listed here with their probable systematic placement. Archivea Christenson & Jenny, Orchids, 65, 497 (1996). (Stanhopeinae) Bhutanthera Renz, Edinburgh J. Bot., 58, 99 (2001). (Orchidinae) Cooktownia D.L.Jones, Austrobaileya, 5, 74 (1997). (Orchidinae) Danxiaorchis J.W. Zhai, F.W.Xing & Z.J.Liu, PLOS ONE, 8(4), e60371 (2013). (Calypsoeae)

Devogelia Schuit., Blumea, 49, 362 (2004). (Collabiinae) Didymoplexiopsis Seidenf., Contr. Orchid Fl. Thailand, 13, 13 (1997). (Gastrodieae) Ericksonella Hopper & A.P.Br., Austral. Syst. Bot., 17, 208 (2004). (Caladeniinae) Pheladenia D.L.Jones & M.A.Clem., Orchadian, 13, 411 (2001). (Caladeniinae) Schuitemania Ormerod, Lindleyana, 17, 288 (2002). (Goodyerinae) Sirindhornia H.A.Pedersen & Suksathan, Nordic. J. Bot., 22, 393 (2002 publ. 2003). (Orchidinae) Zeuxinella Aver., Updated Checklist Orchids Vietnam, 96 (2003). (Goodyerinae)

485

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polysaccharide isolated from Dendrobium chrysotoxum Lindl. Journal of Biochemistry and Molecular Biology, 40, 670–677. Zheng, W.-P., Tang, Y.-P., Lou, F.-C., and Zhi, F. (2000a). Studies on the constituents of Dendrobium chryseum Rolfe. Zhongguo Yaoke Daxue Xuebao, 31, 5–7. Zheng, W.-P., Tang, Y.-P., Zhi, F., and Lou, F.-C. (2000b). Dihydroayapin, a new coumarin compound from Dendrobium densiflorum. Journal of Asian Natural Products Research, 2, 301–4. Zwenger, C. (1872). Ueber cumarsäure, hydrocumarin und hydrocumarinsäure. Annalen der Chemie und Pharmacie: Supplement, 8, 23−44.

C U MU L ATI VE GLO SSARY (Prepared by Finn N. Rasmussen)

18S rDNA  a slowly evolving nuclear gene useful in phylogenetic analyses, coding for 18S ribosomal RNA. Abaxial  the surface of an organ directed away from the main axis, in general the lower surface of a leaf, outer surface of a sepal or petal, etc. Acetabuliform  shaped like a saucer or shallow cup. Achlorophyllous  without chlorophyll (i.e. non-photosynthetic). Acicular  needle-shaped. Acrocentric  referring to a chromosome with the centromere lying near one end. Acrodromous  with veins curving toward the apex. Acrotonic  referring to the attachment of the rostellum or viscidium to the apex of the anther. Actinomorphic  radially symmetrical; regular. Acuminate  tapering gradually to an extended point. Acute  terminating in a distinct but not extended point, the converging edges separated by an angle of less than 90°. Adaxial  the surface of an organ directed toward the main axis, in general the upper surface of a leaf, inner surface of a sepal or petal, etc. Adnate  united to a part or organ of a different kind; cf. connate. Adventitious  applied to roots that do not rise from the radicle but from the stem, etc.; also applied to embryo-like structures in a seed that arise from outside the true embryo and often abort. Aestivation  the arrangement (folding, interleaving) of flower parts in the bud, e.g. valvate, imbricate. AFLP  amplified fragment length polymorphism, a PCR-based DNA-‘fingerprinting’ technique. Agamospermy  asexual formation of embryos and seeds without fertilization. Agenous  a form of stomatal development in which the guard-cell mother-cell is surrounded by protoderm cells that do not divide (agene cells). Ageotropic  referring to plant parts growing upward in response to gravity rather than down. Alate  having winglike extensions or parts; winged. Albino  a plant or flower that lacks chlorophyll. Alkaloids  organic compounds with basic chemical properties, usually containing at least one nitrogen atom in a heterocyclic ring and often having a pharmacological effect on animals. Allogamy  cross-fertilization or outbreeding. Allopatric  distributed in separate, non-overlapping geographic areas.

Allopolyploid  a hybrid between two (or more) different species, possessing two (or more) unlike sets of chromosomes from each parent; cf. autopolyploid. Allozyme  variants of an enzyme occurring in a population, distinguishable by electrophoresis (in this case, specifically the various alleles of a single gene). Alveolate  deeply pitted, like a honeycomb. Amorphous  formless, lacking definite shape. Amphistomatous  having stomata on both sides of a leaf. Amphistomaty  the presence of stomata on both surfaces of an organ such as a leaf. Amphivasal  referring to a type of vascular bundle in which xylem surrounds the phloem. Amplexicaul  clasping the stem. Amyloplast  a type of plastid that stores starch. Anastomosing  forming a network by connection of parts. Anatropous  applied to an ovule bent parallel to its funiculus or stalk. Ancipitous  with two edges instead of being round; said of certain flattened stems and pseudobulbs. Androecium  the assemblage of stamens in a flower. Aneuploid  having more or fewer than an exact multiple of the haploid set of chromosomes. Angiosperm  a flowering plant with seeds enclosed in an ovary, a flowering plant. Anisocytic  referring to a stoma surrounded by three ­unequally sized subsidiary cells. Annual  a plant that completes its life cycle within one year and then dies; cf. perennial. Annular thickenings  rings of secondary wall thickenings in tracheary elements. Annulus  a ring-shaped thickening from which the inflorescence emerges from the stem apex in some genera of Pleurothallidinae. Anomocytic  referring to a stoma lacking subsidiary cells. Anther  that part of the stamen in which pollen is produced by microsporogenesis. Anther canal  narrow, basal part of anther enclosing the caudicle, especially in Orchideae. Anther cap  the dry outer wall of anther, remaining after dehiscence and release of the pollinia. Anthesis  specifically, the opening of the stamen(s), but usually meaning the opening and functional period of the flower. Anthocyanin  pigments colouring flowers and other plant parts violet, blue or reddish. 501

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Anticlinal  cell wall or plane of a cell division perpendicular to the nearest surface of a plant organ. Anticous  facing toward the axis of the flower. Antrorse  directed forward and upward. Aperture  a depressed area of the pollen grain through which the pollen tube emerges, characterized by a localized thinning of the exine. Apex (pl. apices, adj. apical)  top, peak, terminal point. Apiculate  ending abruptly with a short point. Apolar  describing a pollen grain without distinct polarity. Apomictic  referring to reproduction without fertilization, or asexual reproduction. Apomixis  asexual reproduction without formation of gametes, the general case including agamospermy (cf. above) and vegetative reproduction. Apomorphy  a derived character state. -arch  referring to the number of ridges of protoxylem (first formed xylem) in steles, e.g. a pentarch stele has five ridges of protoxylem. Arcuate  curved, bent. Aristate  with a bristle-like appendage or point. Articulate  consisting of parts united by joints, jointed. Asparagoids  a large presumably monophyletic group of monocotyledonous plants (usually recognized as order Asparagales) including ‘petaloid’ families such as Amaryllidaceae, Iridaceae, and Orchidaceae. atpB  a slowly evolving plastid gene used in phylogenetic analyses, encoding the β-subunit of ATP synthase. Attenuate  tapering to a slender point. Auricle  a small lobe or ear. Auriculate  shaped like an earlobe. Autapomorphy  a derived character state that occurs in only the clade under study and is therefore uninformative in a more comprehensive cladistic context. Autogamy  self-fertilization. Autopolyploid  an organism with two (or more) sets of chromosomes derived from the same species; cf. allopolyploid. Autotrophic  referring to plants, being capable of synthesizing its own nutrition from inorganic substances using light, a green plant. Axile  a type of placentation in which the ovules are positioned in the middle of the ovary in the angles formed by the septa. B-chromosome  relatively inactive heterochromatin in a nucleus, usually smaller than normal chromosomes, numerically inconstant, and generally of limited taxonomic use. Baculate  rod-shaped; used for instance to describe exine structure in pollen grains. Basic number  the minimum haploid chromosome number in a species series. Basifixed  attached at or by the base. Basifixed anthers are attached by their base to their filaments. Basionym  in nomenclature, the first valid combination in which a certain epithet was used. For example, the basionym of Neottia nidus-avis (L.) Rich. is Ophrys nidus-avis L. Basitonic  referring to the attachment of the rostellum or viscidium at the base of the anther. Bayesian inference  a method of phylogenetic inference from large sets of molecular data employing a model of nucleotide 502

substitution and a posterior probability distribution describing the probability of trees considering the data. Biauriculate  having two ear-like lobes. Bifacial (leaf)  with distinct adaxial and abaxial surfaces and differentiated mesophyll. Bifid  forked or cleft, Y-shaped, also ‘bifurcated’. Bilobulate  having two small lobes (lobules). Bilocular  having two compartments or cavities (locules). Biota  the flora and fauna of a geographical region. Bipartite  deeply cleft or divided almost to the base. Bitegmic  having two integuments surrounding the nucellus and developing into outer and inner seed coats. Bivalent  either a chromosome that has duplicated to form two sister chromatids or a pair of duplicated homologous chromosomes held together at meiosis. Blade  the expanded part of a leaf or petal. Bootstrap support  an estimate derived from a resampling procedure (deleting of data with replacement) of the percentage (frequency of recurrence in the replicates) of clade support in a phylogenetic hypothesis; cf. jackknife support. Bract  a scale-like leaf. Brachyparacytic  referring to a stomatal arrangement in which the subsidiary cells are parallel to the long axis of the guard cells but not completely enveloping them. Bundle cap  a cluster of fibres that covers the outermost side (phloem side) of the vascular bundle. Bursicula, bursicle  a purse-like or pouch-like structure enclosing the viscidium in certain species of Orchideae. Caducous  falling off or being shed at an early stage of development. Caespitose  growing in dense clusters or tufts. Calcareous  composed chiefly of calcium carbonate, or growing on limestone or chalky soil. Calceiform, calceolate  slipper-shaped. Calcicole  a plant that thrives in lime-rich substrate such as chalk, limestone or calcareous sand. Callus  a waxy, fleshy, or other protuberance, as on a labellum. Calyculus  a calyx-like structure found outside the regular calyx (the sepals). Calymmate  a condition of pollen tetrads or polyads in which the ektexine/sexine of the component monads is continuous around the entire pollen unit. Calyx  the assemblage of sepals in a flower. Cambiform  describing cells shaped like the cells of a cambium. Cambium  a layer of actively dividing cells that forms secondary tissues. Campanulate  bell-shaped. Canaliculate  having one or more longitudinal grooves or channels. Capitate  forming a head-like dense cluster. Capitulum  a dense inflorescence (‘head’) composed of sessile flowers on a receptacle. Capsule  a dry fruit splitting along one or more sutures, as in most Orchidaceae. Carina  a keel.

CUMULATIVE GLOSSARY

Carpel  the ovule-bearing organ of the flower, collectively the gynoecium. Cartilaginous  tough and elastic, like cartilage. Casparian strip  a suberized/lignified band in primary cell walls, especially applied to the endodermis of plant roots. Cataphyll  a scale-like leaf, often referring to the first such leaves on a shoot. Caudate  tail-like or with an appendage shaped like a tail. Caudicles  extensions of tissue derived from the anther and connecting the pollinia to stipe or, in orchids without a stipe, directly to the viscidium. Cauline  occurring on the stem. Centric fission  the production of two telocentric chromosomes from one metacentric chromosome by splitting at the centromeric region. Centromere  a specialized, often waist-like chromosome region to which spindle fibres attach during cell division. Chalazal  referring to the region of the ovule opposite the micropyle, where the integuments fuse with the ovule stalk. Chartaceous  papery, paper-like. Chasmogamous  relating to a flower that opens to allow for pollination. Chlorenchyma  parenchyma tissue containing chloroplasts. Chloroplast  a cell organelle in all green plants that contains DNA, chlorophyll, and other pigments involved in photosynthesis; a green plastid. Chromocentre  a tightly coiled chromosome or portion thereof in the interphase nucleus of a cell. Chromosome  a rod-shaped body composed of DNA and histones and other proteins, responsible for transmission of genetic information from cell to cell and generation to generation and also for control of cell function and development. Ciliate  describing a structure fringed with fine hairs, cilia. Ciliolate  having minute cilia. Cincinnus  a form of cymose inflorescence branching alternately on opposite sides of the shoot system (the false axis). Clade  a monophyletic group. Cladistics  a method of systematics used to reconstruct phylogenies of organisms and to construct classifications based on the identification of clades and their interrelationships. Cladogram  a branching diagram of taxa showing relationships as defined by synapomorphies (derivative or novel states). Clathrate  with a lattice-like structure or appearance, having pores, holes or cavities. Clavate  club-shaped. Claw  a stem-like and narrowed basal part of the sepals and petals in some plants. Cleistogamy  referring to self-pollination in flowers that do not open. Clinandrium  the portion of the orchid column underneath the anther; the ‘anther bed’. Clypeus  in entomology, the shield-like part of an insect’s exoskeleton anterior to the front edge of the carapace. Cochleate  twisted like a snail shell. Collateral vascular bundle  a vascular strand in which the xylem is positioned at one side of the phloem, usually the abaxial (outer) side.

Collenchyma  plant tissue composed of elongated cells with non-lignified wall thickenings, often at the corners, providing mechanical support. Colliculate  covered with small, rounded or hillock-like elevations. Colpate  furrowed, as in pollen grains. Columella (pl. columellae)  columnar structural elements of the exine of some pollen grains. Column  an organ of the orchid flower representing the fusion of filaments and style. Column foot  an extension at the base of the column in some orchids, to which the labellum is attached. Commissure  the surface or place along which two structures, such as carpels, are joined. Companion cell  a specialized cell in the phloem of flowering plants. Complanate  flattened or compressed. Concolorous  having the same colour. Conduplicate  folded once longitudinally down the middle. Connate  united with a structure of the same kind, as one sepal to another; cf. adnate. Connective  the part of an anther connecting the locules. Connivent  converging and touching but not fused. Convergence  evolution of superficially similar structures in taxa from completely different antecedent states in their common ancestor; see also homoplasy. Convolute  rolled during development, as some orchid leaves. Coralloid  resembling coral in appearance or form. Corbicula  pollen basket, part of the tibia on the hind legs of certain pollen collecting bees. Cordate, cordiform  heart-shaped. Coriaceous  of a leathery texture, as of some orchid leaves. Corm  a condensed stem, generally underground. Corolla  the assemblage of petals of a flower. Coronescent  remaining on top like a crown, e.g. withered sepals or petals on top of fruit. Cortex  the region of ground tissue between the epidermis and vascular tissues in a stem or root. Cortical  relating to or associated with the cortex. Corymb  a flat-topped inflorescence in which the flowers are born on stalks of different lengths, the longest at the base. Costa  a longitudinal ‘rib’ or ‘nerve’. Crenate  having rounded teeth along the margin. Crenulate  having a margin with small, low, rounded teeth. Crispate  irregularly waved and twisted, curled, or ruffled, as the margins of certain leaves. Crista (pl. cristae)  in palynology, a crest-like ornamentation, taller than wide, characterized by a narrowly curved base and a sharp upper edge. Cruciate  arranged in or forming a cross; overlapping or crossing. Cucullate  shaped like a hood, hooded. Cuneate, cuneiform  wedge-shaped. Cuniculus  a tubular nectary embedded in the ovary. Cupuliform  cup-shaped. Cuticle  a hydrophobic layer composed of cutin on the outer wall of epidermal cells. 503

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Cutin  a wax-like fatty compound impervious to water and making up the cuticle. Cyathiform  spoon- or cup-shaped. Cymbiform  boat-shaped. Cyme  a broad, determinate, repeatedly branching inflorescence, with the oldest flowers at the end of the branch. Dactyliform  finger-shaped. Decussate  pairwise opposite and alternating, like opposite leaves in four rows on a stem; also tetrads of four pollen arranged in two pairs lying across from one another. Deflexed  deflected, bent or turned abruptly downward. Dehiscent  spontaneously opening along certain lines to expose reproductive structures, as in orchid anthers, capsules or seeds, or the dropping of leaves. Deltate, deltoid  triangular. Dendritic  having a tree-like branching structure. Dentate, denticulate  toothed. Dextral  to the right, located on the right side. Diacytic  referring to stomata with two elongate subsidiary cells, their long axis perpendicular to the axis of the stoma. Diaphanous  translucent, transparent; delicate. Dicotyledons  a paraphyletic group comprising all angiosperms that are not monocotyledons. Most dicotyledons have two cotyledons. Dimorphic  occurring in two distinct forms, e.g. dimorphic leaves. Diploid  having two sets of chromosomes, designated ‘2n’. Disc  a region of the labellum, distal to the claw (when it exists), delimited laterally by the lateral lobes of the labellum and distally by the midlobe. The disc usually has a callus, more rarely two calli. Distal  located at the far end, away from centre or main body; cf. proximal. Distichous  in two ranks or rows on opposite sides of an axis, as in leaf arrangement. Dolabriform  shaped like an axe-head. Dorsiventral  flattened and having obvious dorsal and ventral sides, as most foliage leaves. Dropper  a shoot sent downward from near the base of the stem, the growing tip forming a tuber. Druse  a spherical compound crystal. Dune slack  a dip between dunes where water accumulates. Dysploid  referring to differing basic chromosome numbers in populations or species (often confused with aneuploid, in which there are single chromosome changes, i.e. monosomy or trisomy). Ecalcarate  without spur. Ecallose  without a callus. Ecotone  a transition zone between two distinct habitats that contains species from each area. Ectomycorrhiza  a symbiotic relationship between a fungus (the mycobiont) that forms a sheath around root tips of a plant (the phytobiont). The fungal hyphae do not enter the roots of the phytobiont. Edaphic  relating to soil, especially the various physical, chemical, and biological properties of the soil that influence living organisms in association with it. 504

Eglandular  without a gland. Ektexine  the outermost stratum of the exine of pollen grains, comprising tectum, columellae, and foot layer. Elaiophore  oil-secreting gland. Elastoviscin  a highly viscous product of the degeneration of a limited number of tapetal cells keeping pollen together in orchid pollinia. Elaters  spiral thickenings or hairs that help to disperse spores or seeds, often by hygroscopic action. Elfin forest  a type of vegetation dominated by low trees, shrubs and mosses found in tropical montane regions. Emarginate  having a shallow notch at the tip. Embryo sac  the female gametophyte of flowering plants containing the egg cell that is later fertilized in the formation of the embryo. Endemic  restricted to a certain geographical area or country. Endexine  the innermost stratum of the exine of pollen grains. Endintine  the inner zone of the intine in the walls of pollen grains. Endocarpic  inside the fruit. Endodermis  the innermost layer of the cortex in roots and some stems of seed plants. Endogenous  originating or produced inside an organism, a tissue or cell. Endosperm  the nutritive tissue formed in the embryo sac, which is often absorbed by other tissues by the time of germination; absent in nearly all orchids. Endothecial thickening  secondary thickening in a wall layer of the anther. Endovelamen  the layers below the outermost layer of a multiseriate velamen. Ennearch  9-arch; see -arch. Ensiform  shaped like a sword, as the leaf of an iris. Entire  smooth and continuous; said of leaves and leaf-like organs in which the margins are not broken by teeth. Entomogamous  pollinated by insects. Epetiolate  lacking a petiole. Ephemeral  short-lived. Epichile  apical lobe of a labellum differentiated into epi-, (meso-), and hypochile. Epidermis  the outermost layer of cells, usually one cell thick, covering the leaves and parts of a plant. Epigaeous  growing on or close to the ground. Epigenous  developing or growing on a surface, especially on the upper surface of a plant. Epilithic  growing on stones or rocks. Epiphyte  a plant growing on another as its substrate, but not parasitic. Epithelial  referring to the epithelium, a membranous tissue covering internal surfaces. Epitype  a specimen or an illustration selected to serve as an interpretative type when other type-material of a validly published plant name is ambiguous or cannot be critically identified. Epivelamen  the outermost layer of a multiseriate velamen.

CUMULATIVE GLOSSARY

Ergastic substances  materials of cellular activity, including waste products of metabolism and stored nutritional deposits such as starch. Erose  irregularly notched, toothed or indented. Eudicots  a large, presumably monophyletic group (clade) of angiosperms with tricolpate (with three apertures) or tricolpatederived pollen, including most dicots. Eumagnolioids  referring informally to a set of ­early-diverging angiosperm plant families used as reference group in analyses of the monocotyledons. Exalate  without wings. Excavate  hollowed out. Excrescence  an outgrowth, as from the epidermis. Exine  outer wall of pollen grains, composed of sporopollenin and often intricately structured and sculptured. Exintine  an outer layer of the intine in the walls of pollen grains. Exodermis  the cell layer immediately subjacent to the velamen of orchid roots and developmentally the outermost layer of the cortex. Extra-  on the outside of. Extraxylary fibres  sclerenchyma fibres appearing outside the xylem in a plant organ. Falcate  curved like a sickle. Farina  a mealy or powdery covering. Fasciculate  arranged in bundles, tufts or clusters. Fen  an open plant community on generally alkaline or neutral wet peat, characterized by tall herbaceous plants such as reeds. Fibrillar  composed of slender fibres or fibrils. Filament  the stalk supporting an anther. Filiform  shaped like a thread or filament. Fimbriate  fringed. Flabellate  fan-shaped. Flexuous  winding or bending alternately from side to side. Foliaceous  leaflike. Foliage leaf   a fully developed and functioning leaf, different from scale leaves, etc. Foot layer  the innermost layer of the ektexine of pollen grains. Fossulate  with minute grooves, applied to the sculpturing of pollen exine. Foveate  having small depressions or fovea; pitted. Foveolate  with minute pores or pits, applied to the sculpturing of pollen exine. Fractiflex  forming a broken or intermittent zig-zag line. Fugaceous  ephemeral, withering or dropping off early. Fungiform  shaped like a fungus, mushroom-like. Funiculus  the stalk of an ovule. Furfuraceous  covered with scaly particles. Fusiform  spindle-shaped, as some pseudobulbs. Galeate  helmet-shaped, hooded. Gametic  pertaining to gametes (egg and sperm cells). Gametophyte  the haploid, gamete-forming phase in alternation of generations in plants. Gammate  with a hook or perpendicular arm like the Greek letter gamma (upper case). Geitonogamous  pollination of a flower from another on the same plant.

Gemmate  a warty or bumpy surface ornamentation of pollen grains. Geniculate  bent like a knee. Genome  the genetic chromosomal complement of an organism or cell; also refers to the circular DNA molecules found in plastids and mitochondria. Geophyte  a herbaceous plant with underground survival buds and a storage organ. Gibbous  bulging or humpy. Glabrous  smooth or hairless. Gland  a secreting organ or structure. Glenion  a structure at the base of the labellum in some species of Pleurothallidinae. Globose  spherical, ball-shaped. Glomerule  a compact cluster of flowers. Gondwanaland  hypothetical former supercontinent that included South America, Africa, Madagascar, peninsular India, Australia, and Antarctica. Grade  level of adaptation; group of similar organisms not constituting a clade. Granulate  referring to pollen with small, rounded elements on the surface. Guard cells  kidney-shaped cells surrounding a stoma. Guild  a set of organisms that use similar resources and are thus potential competitors. Gymnosperm  a plant with naked seeds, not enclosed within an ovary, such as a conifer. Gynizus  the fertile part of an orchid stigma. Gynoecium  the assemblage of carpels in a flower. Gynostemium  see column. Hamate  hooked at the tip; see also hamulate and uncinate. Hamulate  having small, hook-like processes. Hamulus  a type of stipe representing the recurved apex of the rostellum. Hapaxanthic  describing a shoot or whole plant that dies after flowering. Haploid  having one set of chromosomes, designated ‘n’. Hastate  shape like an arrowhead, spear-shaped. Hemi-  prefix meaning half or partially. Hemiperigenous  a form of stomatal development in which the guard-cell mother-cell is surrounded by agene cells and perigene cells. Hermaphroditic  referring to a bisexual flower with functional stamen(s) and pistil. Heteranthous  flowering from special shoots that do not produce pseudobulbs or foliage leaves. Heteroblastic  having pseudobulbs of a single internode. Heterobrochate  having reticulate sculpturing, such as pollen exine. Heterochromatin  regions of densely staining chromatin that are highly condensed in the interphase nucleus and not transcribed. Heteromodular  with alternating vegetative and reproductive shoots. Heteromycotrophic  referring to a plant with inadequate photosynthesis and partially or entirely dependent on complex organic substances provided by fungi (also ‘mycoheterotrophic’). 505

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Heterotrophic  referring to an organism incapable of synthesizing its own food and thus dependent on consuming/absorbing organic substances for nutrition. Hexaploid  having six sets of paired chromosomes, designed ‘6n’. Hilum  the nucleus of a starch grain; the scar on a seed representing the point of attachment. Hippocrepiform  horseshoe-shaped. Hirsute  covered with rough hairs or bristles. Holomycotrophic  dependent on mycotrophy as sole source of nutrition and hence without or with little chlorophyll. This condition has often been erroneously referred to as ‘saprophytic’. Holotype  the single specimen or illustration designated by the author as the type of a scientific plant name in the rank of species or below; cf. epitype, isotype, lectotype, neotype, syntype. Homoblastic  having pseudobulbs of several internodes. Homogamous  with two sexes, describing a flower with functional stamen(s) and pistil. Homology  trait attributable to inheritance from a common ancestor rather than from independent origins. Homoplasy  resemblance not attributable to inheritance from a common ancestor but to convergence, parallelism, or reversal. Humus  brown or blackish type of soil consisting of partially or wholly decayed organic matter. Hyperincumbent  upside down, reversed. Hypochile  basal lobe of a labellum differentiated into epi(meso-) and hypochile. Hypodermis  a layer or layers of cells beneath the epidermis and distinguishable from underlying layers. Hypophysis  the uppermost cell of the suspensor of the embryo from which the root and root cap are derived. Hypostase  the proliferation of the nucellus at the chalazal end of the embryo sac, often partly lignified or suberized. Hypostomatous  having stomata on the underside of a leaf only. Hypostomaty  the condition of having stomata only on the lower surface of the leaf. Hysteranthous  having leaves produced at a different time than flowers. Idioblast  a cell in a tissue that differs from others in form, size or contents. Idiogram  a diagram of a karyotype (chromosome complement in a cell), often arranged in order of decreasing size. Imbricate  overlapping; referring, for example, to sepals in the flower bud or to leaves on the stem. Inaperturate  describing anthers or pollen grains without apertures. Incumbent  lying upon or bending downwards, as the anther of many orchids during development. Incurved  bent or curved inward. Indehiscent  not opening spontaneously. Indumentum  a hairy, pubescent, scaly, or waxy covering of an organ such as a leaf or sepal. Inflexed  bent inward. Infundibular  funnel-shaped. 506

Insertion-deletion event  length variation in a nucleotide sequence in DNA, also referred to as an indel. Intectate  without a tectum (roof-like layer), with reference to pollen exine. Integument  outer cell layer(s) surrounding the nucellus of the ovule, developing into the seed coat. Internode  the region of an axis between two nodes. Interphase  the period in the cell cycle between one mitosis or meiosis and the next, during which DNA is replicated. Intine  the cellulosic inner wall of a pollen grain, below the exine. Introgression  the incorporation of genes from one species into the gene pool of another through repeated backcrossing of an interspecific hybrid with one of its parents. Introrse  turned or faced inward toward the axis; in anthers, opening toward the centre of the flower. Involute  rolled inward or toward the upper side, as some leaves. Isodiametric  having equal or nearly equal diameters or axes. Isomodular  referring to a plant architecture in which all shoots are equivalent. Isotype  a duplicate of the holotype. An isotype is always a specimen. Isozyme, isoenzyme  any one of several different forms of some enzymes. Isthmus  a narrow strip of tissue joining two larger organs or parts of an organ, ligament. ITS  the internal transcribed spacers of 18S–26S nuclear ribosomal DNA, characterized by tandem repeat structure and high copy number. Jackknife support  an estimate derived from a resampling procedure (deletion of data without replacement) of the percentage (frequency of occurrence in the replicates) of clade support in a phylogenetic hypothesis; cf. bootstrap support. Juxtaposed  placed side by side, e.g. pollinia within an anther; cf. superposed. Karyotype  a representation of the chromosome complement in a cell, with the mitotic chromosomes arranged in pairs in order of size and shape. Keiki  a small plant growing from a node on an inflorescence; a means of asexual reproduction. Keranga  a type of forest with low trees on nutrient-deficient soils in Malaysia and Indonesia. Labellum  the median, modified petal of an orchid flower also known as the lip. Lacerated  having deeply cut edges. Laciniae  narrow, often tapering divisions. Laciniate  deeply slashed into narrow divisions or lobes. Lacunar  with holes or gaps, recessed. Laevigate  smooth, with reference to the sculpture of pollen exine. Lamella (pl. lamellae)  a thin plate or layer. Lamina  the expanded area of a leaf, blade. Lanceolate  widening to a maximum below the middle and tapering toward the apex. Lanuginose  covered with soft short hairs, woolly.

CUMULATIVE GLOSSARY

Lappet  the distal, apiculate portion of the clinandrium in Peristeria. Lectotype  a nomenclatural type of a scientific name designated from the original material (e.g. one of more syntypes) if no holotype was indicated at the time of publication, or if the original holotype is missing or found to belong to more than one taxon. Lenticular  lens-shaped. Ligament  a band of tissue connecting two parts, e.g. a mobile labellum to a column foot. Ligneous  woody. Ligulate  tongue- or strap-shaped. Ligule  tongue. Lilioids  monocotyledonous plants including all ‘petaloid’ families in the orders Asparagales, Dioscoreales, Liliales, and Pandanales. Linguiform  shaped like a tongue. Lip  see labellum. Lithophyte  a plant growing on rocks. Locule  a chamber or cavity, as that bearing ovules in an ovary or pollen in an anther. Long-branch attraction  referring to the tendency of terminals with long branches (i.e. branches with many evolutionary changes) in a phylogenetic tree to be placed artificially close to each other. Lorate  strap-shaped. Marcescent  withering but persisting. Marsupium  a membranous bag enclosing the viscidium in a few members of Oncidiinae. Massula (pl. massulae)  a mass or packet of pollen grains in sectile pollinia. matK  a plastid gene used in phylogenetic analyses, coding for a maturase. Maximum likelihood  a method of phylogenetic inference applied in terms of the probability that a proposed model of evolution and hypothesized history could explain observed data, often but not exclusively used with DNA sequence data. Medulla  pith in stems or roots. Megasporogenesis  in angiosperms, the formation of megaspores containing the embryo sac. Mentum  a chin-like projection in some orchid flowers, formed by the fusion of the lateral sepals and base of the labellum/column. Meristele  one of a number of vascular bundles of a stele. Meristemoid  a cell formed by an unequal division of a protodermal cell. Mesic  adapted to a moderately moist habitat. Mesochile  middle lobe of a labellum differentiated into epi-, (meso-), and hypochile. Mesogene  any cell derived by division of a meristemoid (that is not itself a meristemoid) before the meristemoid divides into two guard cells. Mesoperigenous  a form of stomatal development in which the guard-cell mother-cell is surrounded by both mesogene and perigene cells. Mesophyll  the usually photosynthetic parenchyma cells between the upper and lower epidermis of a leaf. Metacentric  referring to a chromosome with the centromere in the middle. Metaphase  a stage in mitosis or meiosis when the chromosomes are aligned along the equator of the cell with the centromeres along the spindle equator.

Metaxylem  that part of the primary xylem that differentiates after protoxylem and before secondary xylem (if formed). Micropylar  referring to the pore or opening in the integuments of the ovule through which the pollen tube enters, or to the homologous opening in the seed coat; also the distal region of the nucellus. Microsporogenesis  in angiosperms, the formation of microspores, which become the male gametophytes (germinated pollen grain and pollen tube). Mimicry  the advantageous resemblance of one species to another, often unrelated, species. Monad  a single cell, rather than a tetrad, resulting from meiosis; a single pollen grain. Monandrous  with only one functional stamen. Moniliform  with contractions and expansions at regular alternating intervals, giving the appearance of a string of beads. Monocaulescent  having only one main stem. Monocotyledons  a large, presumably monophyletic group of flowering plants sharing a number of distinct characters such as a stele with scattered vascular bundles, deltoid protein inclusions in phloem plastids, often trimerous flowers, and only one cotyledon. The monocots include such plant groups as grasses, palms, lilies, and orchids, although orchids have lost their cotyledon. Monomorphic  occurring or known to exist in only one form. Monophyletic  a group of taxa that are exclusive, e.g. the components are more closely related to each other than any of them is to a member(s) of another such group; a clade. Monopodial  referring to a growth habit in which new leaves develop from the same meristem or growing point as all previous leaves; cf. sympodial. Motile  from a fixed point of connection, moving or being capable of moving, as opposed to moving from one place to another, i.e. mobile. Mucilage  a sticky, slimy or gummy substance that contains protein and polysaccharides. Mucronate  ending abruptly in a sharp point (mucron). Multiseriate  arranged in several parallel rows. Multivalent  a structure formed by the association of more than two chromosomes during meiosis, characterizing some polyploids. Muricate, muriculate  rough with minute hard points or excrescences. Murus  (pl. muri)  a ridge that is part of the ornamentation of pollen grains. Mya  million years ago. Mycoheterotrophic  referring to a plant with inadequate photosynthesis that is partially or entirely dependent on complex organic substances provided by fungi. Mycorrhiza  the association of fungi and roots of higher plants, often termed a symbiosis. Mycotrophic  referring to a plant that utilizes organic substances provided by fungi. Myophilous  pollinated by flies. Myrmecophily  symbiosis with ants. Navicular, naviculate  shaped like a boat. Neighbor-joining  a tree-building algorithm using a matrix of corrected* pairwise distances to produce phylogenetic estimates (*based on a specific model of DNA sequence evolution). 507

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Neotype  a specimen or illustration selected as the type for naming a species or subspecies when there were no types ­originally designated or all type material was subsequently lost. Nexine  in Erdtman’s system of pollen wall stratification, the inner layer of the exine. Nitid  luminous, shining. Node  the region on an axis (stem, rhizome, etc.) to which other plant parts (e.g. leaves) are attached. Nombre fondamental  fundamental number; the total number of chromosome arms in the complete complement of a species. Nototrobic  in pollination, placing of pollinaria on the back of the pollinator; cf. sternotrobic. Nucellus  the tissue that surrounds the megaspore (and embryo sac) in seed plants. Nucleoside  a group of compounds consisting of a purine or pyrimidine base linked to the sugar ribose or deoxyribose, e.g. adenosine. Nucleotide  a phosphate ester of a nucleoside, either a purine or pyrimidine base linked to ribose or deoxyribose phosphates; one of the basic repeating units in DNA or RNA (e.g. adenosine triphosphate). Obtrullate  with widest axis above middle and with straight margins; inversely trowel-shaped. Obtuse  with a blunt or rounded tip. Oligotrophic  deficient in nutrients. Ontogenesis  development of an individual organism from embryo to adult. Operculate  having a lid; said of anthers when the anther wall shrivels and is shed after anthesis, leaving the pollinia exposed. Orbicular  circular. Ornithophilous  pollinated by birds. Osmiophilic  stainable with osmium tetroxide for electron microscopy. Osmophore  a gland, usually floral, that produces fragrances attracting pollinators. Outgroup comparison  in phylogenetics, comparison of character states in the group under study (the ingroup) with another group (the outgroup). The outgroup may be the closest monophyletic relative of the ingroup or a paraphyletic group in which the ingroup is phylogenetically nested. Ovary  the part of the carpel that contains the ovules and becomes the fruit. Ovate (ob-)  an outline shaped as a flat projection of an egg, broadest below the middle (obovate: above middle). Ovoid  egg-shaped. Ovule  the unit of the ovary that contains the egg cell and becomes the seed. Paedomorphosis  evolutionary change in which juvenile and adult characteristics co-occur in the same plant. Paleaceous  chaff-like. Palisade mesophyll  leaf mesophyll characterized by elongate cells arranged with their long axes perpendicular to the surface. Pandurate  shaped like a fiddle, with rounded ends and constricted middle portion. Panicle  a branching inflorescence. Papilla  a rounded projection from a portion of an epidermal cell; a bump or protuberance. 508

Papyraceous  having qualities associated with those of paper. Paracladial  functional similarity between branches of different order, e.g. in an inflorescence when additional racemes are produced as branches of the first raceme while its flowers are fading. Parallelism  similarity in species due to evolution of a trait independently from the same antecedent state in their common ancestor. See also homoplasy. Paraphyletic  referring to a group of species or other higher taxa that is non-exclusive, i.e. not including some of the closest relatives or one or more group members. Parenchyma  a plant tissue composed of thin-walled, relatively unspecialized cells with one or more physiological functions. Parietal  a type of placentation in which the ovary has a single chamber (locule), and ovules are in rows on the ovary wall or its invaginations. Parsimony  a method of phylogenetic inference emphasizing the criterion of the simplest and most economical explanation to explain the observed variation in data; its use does not imply that evolution is parsimonious, but rather the principle applies strictly to simplicity of the explanation relative to available data. Patent  spreading, divergent from the axis at almost 90°. Paucifoliate  having relatively few leaves. PCR  polymerase chain reaction, a method for amplifying a DNA nucleotide sequence using DNA polymerase, an enzyme catalyzing the synthesis of nucleic acids from pre-existing templates. Pedicel  the stalk of an individual flower. Pedicellate  supported by a stalk. Peduncle  the stalk of a solitary flower or an entire inflorescence. Pellicle  a thin, flexible outer layer. Pellucid  translucent. Peloric  referring to a genetically or environmentally induced malformation, such as actinomorphy in a flower that is normally zygomorphic. Peloton  in orchid mycorrhiza, an intracellular coil of fungal hyphae eventually digested. Peltate  shield-shaped, having a flat circular structure attached to a stalk near the centre. Penicillate  having or resembling a tuft or brush of fine hairs. Pentaploid  having five sets of chromosomes, designated ‘5n’. Perennial  living more than one year; cf. annual. Perforate  with holes, in descriptions of pollen grains applied to holes in the tectum less than 1 μm in diameter. Perforation plate  a part of the wall of a vessel element (xylem) that is perforated. Perianth  the sepals and petals of a flower. Pericentric inversion  a chromosome inversion in which the breaks are on either side of the centromere and include it. Periclinal  referring to a cell wall or the plane of cell division parallel to the nearest surface. Pericycle  that part of the stele of a root (one or more layers) between the endodermis and phloem, giving rise to lateral roots. Perigene cell  a daughter cell formed by one or more divisions in a protodermal cell adjacent to the guard-cell mother-cell of a stomatal complex. Petal  any of the whorl of flower parts generally just inside the sepals, usually colourful and showy. Petiolate  borne on a leaf stalk.

CUMULATIVE GLOSSARY

Petiole  leaf stalk. Phalaenophilous  pollinated by butterflies. Phenetics  an approach to classification based on overall similarity of phenotypic characters, not necessarily reflecting phylogenetic relationships. Phloem  the food-conducting tissue of vascular plants. Phototaxis  movement in response to light. Phyllopodium  used in Maxillaria for an abscission layer of the apical leaf often projecting above the pseudobulb in a persistent stalk. Phyllotaxis  the arrangement of leaves on a stem. Phylogenetics  an approach to the study of evolution and classification based on reconstruction of relationships or evolutionary history of organisms (i.e. cladistics); these reconstructions can be based on any type of data, including morphology, DNA information, and chemistry. Pilate  like a hair. Pilose  covered with fine, soft hairs. Pilum  in descriptions of pollen, a rod-like element of the sexine. Pinnate  having parts, e.g. leaflets, arranged on each side of a common axis; feather-like. Pistillate  referring to a unisexual flower lacking stamen(s) but with functional pistil. Pit-field  a thin area in the primary wall of a plant cell in which one or more pits may develop if a secondary wall is formed. Pith  ground tissue (usually parenchyma) in the centre of a stem or root. Placenta  a region(s) in the ovary to which ovules are attached. Placentation  the manner in which ovules are attached in an ovary. Plesiomorphy  an ancestral character state. Pleuridia  falcate or rectangular, liquid-secreting glands in Coryanthes. Plicate  folded or pleated. Plumule  the embryonic shoot of a plant. Plurifoliate  having relatively many leaves. Polar nuclei  one or more (generally two) nuclei in the central cell of a mature embryo sac. Pollen  microspores with highly structured walls, produced by microsporogenesis in anthers and giving rise to the male gametophyte. Pollinarium (pl. pollinaria)  the functional unit of pollen transfer in orchid pollination, consisting of two or more pollinia (sometimes with caudicles), often a stalk or stipe, and viscidium. Pollinium (pl. pollinia)  a coherent mass of pollen grains. Polyphyletic  referring to a group composed of one or more taxa for which the individual components are not each more closely related to each other than they are to multiple members of other groups. Polyploid  having more than two complete sets of chromosomes. Polytomy  three or more branches diverging from one point on a cladogram (also polychotomy). Porate  with a pore or aperture, as in pollen grains. Porrect  extended forward. Premorse  abruptly truncated, as though bitten or broken off. Proembryo  the embryo in its early stages of development, before differentiation of main body and suspensor.

Prosenchyma  plant tissue consisting of elongated cells with tapering ends, occurring in supporting and conducting tissue. Prostrate  growing flat and close to the ground. Protandrous  referring to a flower in which the anther(s) matures and release their pollen before the stigma of the same flower is receptive. Proteranthous  having flowers that appear on a shoot before it develops into a pseudobulb; flowering before the shoot has matured. Protocorm  the ephemeral structure resulting from the germinated orchid seed and from which the first true shoot and root differentiate. Protoderm  the meristematic tissue that produces the epidermis of an organ. Protologue  the description, diagnosis, and citations of material associated with the publication of a scientific name of a taxon. Protuberant  projecting from a surrounding or adjacent surface, often as a rounded mass. Proximal  located closer to the centre or main body, opposite; cf. distal. Pseudobulb  the variously thickened portion of an aerial orchid stem. Pseudocopulation  attempted copulation by the male of a potential insect pollinator with a flower mimicking a female of its species. Psilate  smooth, with reference to the surface of a pollen grain. Psygmoid  shaped like a fan with unifacial (dorsiventrally flattened) leaves. Puberulent  covered with minute hairs or fine down; finely pubescent. Pubescent  covered with short hairs or soft down. Pulverulent  dusty, covered with or crumbling to fine powder or dust. Pulvinate  cushion-shaped or having a cushion-like swelling at the base. Pulvinulus  cushion-like outgrowth. Pustulose  covered with blisters or wart-like swellings. Pyriform  shaped like a pear. Quaquaversal  pointing in all directions. Raceme  an indeterminate inflorescence of stalked flowers. Rachis  that portion of an inflorescence above the peduncle, bearing flowers. Radical leaves  leaves clustered at the base of the stem. Radicle  the embryonic root. Ramentae  thin, brownish, chaffy scales on the leaves or young shoots of some plants. Raphides  intracellular, needle-shaped crystals, usually composed of calcium oxalate. rbcL  the plastid gene that codes for the large subunit of the enzyme ribulose 1,5-bisphosphate carboxylase/oxygenase, which fixes carbon dioxide in photosynthesis. rDNA  ribosomal DNA, sequences encoding ribosomal RNA. Recurved  curved backwards. Reflexed  bend backwards. Reniform  shaped like a kidney. Repent  creeping or prostrate. Restriction site mapping  plotting of the positions at which particular restriction endonucleases cleave a genome; usually 509

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referring to the circular plastid genome or the tandemly repeating l8S–26S region of nuclear ribosomal DNA. Resupinate  referring to an orchid flower with the labellum lowermost relative to the other perianth parts; achieved by several means, including twisting of the pedicel and producing a pendent inflorescence. Reticulate  net-like, forming a network, with parts connected as in a network. Retrorse  directed or turned backward or downward. Retuse having a rounded or obtuse apex with a central shallow notch. Reversal  in phylogenetics, a derived (apomorphic) character state with the appearance of an ancestral (plesiomorphic) state. Revolute  rolled backward; with margin rolled toward lower side, as some leaves. RFLP  restriction fragment length polymorphism, a technique for cutting DNA using restriction enzymes that cleave DNA at specific sites, which are variable among species, thus producing a DNA ‘profile’ of each accession. Rhizodermis  the outer cell layer (epidermis) of a root. Rhizoid  a filamentous outgrowth that functions like a root. Rhizome  the indeterminate axis or system of axes of many plants, such as sympodial orchids, that successively give rise to new shoots and flowers, often horizontal or underground. Rhomboid  diamond-shaped or shaped like a parallelogram with unequal adjacent sides. Rhomboidal  in the description of pollen grains, a tetrad in which the proximal faces of two individual members are in direct contact and the remaining two are separated, giving the tetrad a rhomboidal outline. Ringent  opening wide, gaping. Robertsonian change  a change in chromosome structure as the result of centric fission or centric fusion that does not disrupt the nombre fondamental. Root  in phylogenetics, a designated basal node in a cladogram often determined by use of an outgroup. Rostellum  part of the median stigma lobe of orchid flowers. Rostellum remnant  the often cleft or bilobed part of the rostellum that is left after the viscidium has been removed. Rostrate  having a beak. Rosulate  arranged in a rosette. Ruderal  growing in waste places, rubbish dumps, or similar disturbed sites. Rugulose, rugose  with wrinkles, having a rough, wrinkled surface. Rugulate  describing a type of ornamentation on pollen grains consisting of elongated sexine elements more than 1 μm long, arranged in an irregular pattern. Ruminate  irregularly grooved or ridged. Rupicolous  growing on rocks. s. l.  sensu lato, ‘in the broad sense’, said about taxonomic entities interpreted with a wider circumscription. s.s. or s.str.  sensu stricto, ‘in the strict sense’, said about taxonomic entities interpreted with a narrower circumscription. Saccate  sac-shaped or adorned with a sac-like pouch. Sagittate  shaped like an arrowhead. Saprolignophilous  often found on decaying wood. 510

Sapromyophilous  pollinated by flies and other carrion insects attracted by dark colours and putrid smell. Saprophyte  a plant that derives its nourishment, in whole or part, from dead organic matter (often used incorrectly for achlorophyllous plants parasitic on fungi). Scabrate  rough, or with a covering of stiff hairs or points; describing pollen grain ornamentation with elements smaller than 1µm in all directions. Scalariform  arranged in a ladder-like fashion, as the pitting in some xylem tracheary elements. Scandent  climbing. Scape  a stalk carrying the inflorescence; peduncle; stem. Scarious  thin, membranous, and dry. Sclerenchyma  a cell with thickened, usually lignified secondary walls; a tissue of such cells functioning primarily in support or protection. Sclerification  the process of being changed into sclerenchyma. Sclerotic  hard, firm, indurate. Scobiform  resembling sawdust, appearing as fine grains. Scrotiform  sac- or pouch-like. Sectile  referring to pollinia comprising several ‘packets’ connected by elastoviscin. Secund  directed to one side only, as flowers on an inflorescence. Seep  a spring out of which water or oil oozes. Semelparous  referring to organisms that produce all offspring at one time and then die. Semitectate  referring to the exine of a pollen grain with a discontinuous ‘roof ’ supported by columellae, giving a reticulate appearance to the surface. Sepal  one of the outermost whorl of flower parts, often as colourful and showy as the petals in orchids. Septum  a partition. Sequence data  in phylogenetics, nucleotide sequences; DNA. Seriate  arranged in series or rows. Serpentine  a mineral composed of hydrated magnesium silicate, common in ultramafic soils. Serrate  toothed, as the margin of some leaves and leaf-like organs. Sessile  attached directly to a base without a stalk or petiole. Setaceous, setose  bristly. Sexine  in Erdtman’s system of pollen wall stratification, the outer layer of the exine. Sieve tube  a series of phloem cells joined end to end at sieve plates, forming a tube through which nutrients are conducted. Sigmoid  curved like the letter S. Silica bodies  conical or spherical masses of silica contained in stegmata of many plants, including some orchids. Sinistral  to the left, located on the left side. Sinker  see dropper. Sinuate, sinuous  repeatedly curving in alternate directions, wavy. Sinus  an indentation or recess. Siphonostele  a type of stele in which the vascular cylinder surrounds a pith. Sister groups, sister taxa  in phylogenetics, the two clades resulting from splitting of a lineage (i.e. having a common ancestor), their most recent common ancestor. Sister groups need not be named taxa in a formal classification.

CUMULATIVE GLOSSARY

Somatic  referring to cell types, structures, and processes in an individual other than those giving rise to gametes. Spatulate  shaped as a spatula, with a broad rounded apex and a narrow base. Sphingophilous  pollinated by moths of the family Sphingidae (sphinx or hawk moths). Spicate  with flowers borne in a spike. Spiculate  covered with pointed or needle-like processes. Spike  an indeterminate inflorescence with sessile flowers. Spiranthosomes  specialized amyloplasts characterizing orchids in tribe Cranichideae, among others. Spongy mesophyll  mesophyll characterized by loosely arranged cells and conspicuous intercellular spaces. Sporophyte  the diploid or asexual, (usually) spore-forming phase in alternation of generations in plants; the dominant phase in all seed plants. Sporopollenin  a cyclic alcohol composing the outer wall of pollen grains and spores, highly resistant to decay. Spur  a saccate or tubular extension of the labellum (or other floral parts) in many orchids, often containing nectar. Stamen  the pollen-bearing organ of the flower, composed of anther and filament, collectively the androecium. Staminate  referring to a unisexual flower lacking a pistil but with functional stamen(s). Staminodium, staminode  a sterile stamen. Stegmata  cells lining sclerenchyma fibres and containing silica bodies. Stele  a morphological region of the stem or root containing vascular tissues and associated ground tissues. Stelidium (pl. stelidia)  excrescences on each side of the functioning anther, often slender and elongated, usually interpreted as staminodia (sterile anthers). Sternotrobic  in pollination, placing of pollinaria on the front side (sternum) of the pollinator; cf. nototrobic. Stigma  the upper region of the carpel that is receptive to pollen grains. Stipes (pl. stipites)  a pollinium stalk derived from the rostellum; also called stipe. Stipitate  borne on a stalk. Stolon  a creeping stem that can root and form a new plantlet. Stoma (pl. stomata)  an epidermal pore of many plant parts surrounded by guard cells and functioning in gas exchange. Striate  having fine linear markings or grooves. Suberin  a fatty substance found in endodermal cell walls and cork. Subsidiary cells  epidermal cells associated with stomata and distinguishable from guard cells either by development or morphology. Subulate  tapering into a fine point. Sulcate  with a groove or furrow, as some pollen grains. Superposed  placed on top of each other or at opposite ends; cf. juxtaposed. Suspensor  an extension at the base of the embryo. Sympatric  distributed in the same or in overlapping geographic areas; cf. allopatric. Symplesiomorphy  a shared, ancestral character.

Sympodial  referring to a growth habit in which new shoots arise successively from axillary buds of a rhizome. Synanthous  having flowers and leaves appearing at the same time. Synapomorphy  a shared, derived character. Syndrome  a pattern of functionally correlated characters seen as an adaptation to certain environmental or biotic conditions, for example, a pollination syndrome. Synergid  one of two cells at the micropylar end of the embryo sac and associated with the egg cell. Synorganization  arrangement of parts or organs in a fixed pattern. Synsepal  a floral part formed by the partial or complete fusion of two or more sepals. Syntype  any specimen cited by the author as base of a scientific plant name when no single holotype was designated; cf. lectotype. Systematics  the science of classification based on natural relationships and the study of variation and evolution of taxa. Tabula infrastigmatica  a particular form of enlarged column base with a peculiar structure found in some genera of Oncidiinae. Tannins  a group of phenol, polyphenol, and flavonoid compounds, bound with starches. Taxon (pl. taxa)  any taxonomic group, e.g. species, genus or tribe. Taxonomy  the science of identification and nomenclature of organisms. Tectate  referring to the exine of a pollen grain with a generally continuous ‘roof ’ supported by columellae. Tectate-perforate  referring to a pollen grain having a tectum with perforations smaller than 1µm in diameter. Tectum  layer of the outer wall of pollen grains that forms a “roof ”. Tegula  a pollinium stalk consisting of the modified rostellar epidermis and possibly subtending layers of cells. Telocentric  referring to a chromosome with a terminal centromere. Tenui-, tenuate  thin-, thinly, becoming thin. Tenuinucellate  a type of nucellar organization in which the embryo sac is surrounded by only an epidermis and a single layer of cells. Teratological form  a deviation from the normal type of structure; a monstrosity. Terete  pencil-like; round in cross-section. Terminal node, terminal  in phylogenetics, any entity that appears at the tip of a branch in a cladogram. Terrestrial  referring to a plant growing on the ground with its roots penetrating the soil; cf. epiphytic, epilithic). Tessellated  with a chequered pattern. Testa  the seed coat. Testicular  shaped like a testicle or testicles; cf. scrotiform. Tetracytic  referring to stomata surrounded by four subsidiary cells. Tetrad  as applied to pollen, four grains attached and shed as a unit. Tetraploid  having four complete sets of chromosomes, designated ‘4n’. 511

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Tetrarch  referring to a stele with four ridges of protoxylem. Theca (pl. thecae)  a set of two of the four locules of the anther. Thorax  in insects, the region of the body between the head and the abdomen bearing the legs and wings. Tilosome  a lignified excrescence from the cell wall of the innermost velamen layer of roots adjacent to the thin-walled passage cells of the exodermis. Topology  in phylogenetics, the branching hierarchy and placement of nodes in a cladogram. Torsive  spirally twisted. Torus  the receptacle of a flower. Tracheary element  a xylem cell in a vascular plant conducting water. Tracheid  a tracheary element of the xylem without perforation plates. Trichome  a plant hair. Trigonous  having three longitudinal angles and three plane faces, triangular in cross-section. Trilete  referring to pollen grain or spores with a triradial mark on the outer surface. Tripartite  divided into three parts. Triploid  having three complete sets of chromosomes, designated ‘3n’. Triquetrous  three-edged; having three prominent angles. trnL-F  intergenic spacer in plastid DNA between the transfer RNAs coding for trnL and trnF, and often the intron in the trnL gene as well; used in DNA phylogenetics. Trulliform  shaped like a bricklayer’s trowel, angular-ovate, broadest below the middle. Truncated  shortened, terminated abruptly. Tuber  a thickened, usually underground storage root or stem. Tubercle  a small, rounded protuberance; a nodule on some roots. Tunica  an enclosing membrane or layer of tissue, e.g. withered leaf bases wrapped around a pseudobulb. Type  in nomenclature, that element to which the name of a taxon is permanently attached, whether as a correct name or as a synonym; not necessarily a typical or representative element of the taxon it typifies. Type species  in nomenclature, referring to the type of a name for a taxon above the rank of species.

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Type specimen  in nomenclature, a specimen designated as type for a name of a taxon at the rank of species or below. Ultramafic  referring to soils low in available calcium, potassium, and phosphorus but high in iron and nickel, considered too toxic for many plants. Ultrastructure  fine structure at a level requiring electron microscopy for study. Umbel  a flat-topped inflorescence in which the pedicels arise from the same point. Uncinate  bent at the end like a hook. Undulate  having a wavy outline or appearance. Unguiculate  having a claw-shaped base. Unifacial (leaf)  with a similar structure on both sides, referring to leaves with similar adaxial and abaxial surfaces; cf. bifacial. Uniseriate  occurring in a single row or layer. Unitegmic  having only one integument surrounding the nucellus. Urceolate  urn-shaped. Valvate  meeting by the edges without overlapping. Vascular bundle  A strand of conductive and supportive tissue consisting mainly of xylem and phloem. Velamen  a tissue that arises from root dermatogen, consisting of dead cells at maturity and bordered internally by an exodermis. Venation  the pattern of veins of a leaf. Venose  having noticeable veins or vein-like markings. Ventricose  having a swelling on one side. Vernation  the arrangement of leaves in a bud. Verrucate, verrucose  warty, covered with warts or shaped like a wart. Vessel members one of the cellular components of a vessel, a tracheary element with perforation plates. Vestigial  rudimentary or degenerate, nonfunctional. Villose, villous  covered with soft, shaggy hairs. Viscarium  see viscidium. Viscidium (pl. viscidia)  the sticky portion of the rostellum that is often connected to pollinia. Viscous, viscid  easily flowing. Xerophyte  a plant adapted to dry habitats (e.g. a cactus). Xylem  the water-conducting tissue in vascular plants. Zygomorphic  bilaterally symmetrical.

C O RRI GENDA

Volume 4 Page 3: The name Epidendroideae Lindl. was invalidly published. The correct citation is Epidendroideae Lindl. ex Kostel., Allg. Med.Pharm. Fl., 1, 233 (1831). Page 587: Thelasinae should be Thelasiinae. Plate 160: The species illustrated is not Aphyllorchis montana Rchb.f. but an unidentified species of Lecanorchis. Plate 177: The species illustrated is Epiblastus basalis Schltr., not Epiblastus ornithoides Schltr.

Volume 5 The following contributor’s name was inadvertently omitted from the list in the front matter. We sincerely apologize for the oversight. DR HENRY OAKELEY (HO) Royal College of Physicians, 11 St Andrews Place, London NW1 4LE, UK Page 156: Publication of the name Sudamerlycaste Archila was invalid when first published (Revista Guatemal., 5(2), 26. Nov. 2002) but was validly and effectively published soon after (Revista Guatemal., 5(3), 77. 2002) by providing a Latin diagnosis and predated the publication of Ida A.Ryan & Oakeley (Orchid Digest, 67, 9. 2003). All species of Ida are now referable to Sudamerlycaste. Page 411: The year of birth for Alvin Embree is 1925.

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E F F E C TI VE DATE S OF PUBLICATION

Volume 1 Volume 2 Volume 3 Volume 4 Volume 5 Volume 6

19 August 1999 28 January 2001 24 April 2003 8 December 2005 22 October 2009 6 February 2014

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GENERIC SYN O N YM S As treated in Genera Orchidacearum, Volumes 1–6 (excluding illegitimate, invalid, and rejected names) Abaxianthus (see Dendrobium) Abdominea (see Robiquetia) Abrochis (see Orchis) Acacallis (see Aganisia) Acanthoglossum (see Pholidota) Aceras (see Orchis) Achroanthes (see Malaxis) Ackermania (see Benzingia) Aclinia (see Dendrobium) Acoidium (see Trichocentrum) Acoridium (see Dendrochilum) Acostaea (see Specklinia) Acroanthes (see Malaxis) Acrochaene (see Bulbophyllum) Acropera (see Gongora) Ada (see Brassia) Adactylus (see Apostasia) Adamanthus (see Camaridium) Adelopetalum (see Bulbophyllum) Adenostylis (see Zeuxine) Adipe (see Bifrenaria) Aeridium (see Aerides) Aeridostachya (see Callostylis) Aerobion (see Angraecum) Ala (see Habenaria) Alatiglossum (see Gomesa) Alipsa (see Liparis) Alvisia (see Bryobium) Alwisia (see Taeniophyllum) Amalia (see Laelia) Amalias (see Laelia) Amblostoma (see Epidendrum) Amblyanthe (see Dendrobium) Amblyglottis (see Calanthe) Amblystoma (see Epidendrum) Amesia (see Epipactis) Amparoa (see Rhychostele) Amphigena (see Disa) Amphiglottis (see Epidendrum) Amphiglottium (see Epidendrum) Amphorchis (see Cynorkis) Amphorkis (see Cynorkis) Ampliglossum (see Gomesa) Anacheilium (see Prosthechea) Anaphora (see Crepidium) Andinorchis (see Chaubardia)

Androgyne (see Panisea) Anettea (see Gomesa) Angorchis (see Angraecum) Angorkis (see Angraecum) Ania (see Tainia) Anisopetala (see Dendrobium) Anisopetalon (see Bulbophyllum) Anistylis (see Liparis) Ankylocheilos (see Taeniophyllum) Anneliesia (see Miltonia) Anochilus (see Pterygodium) Anota (see Rhynchostylis) Anthereon (see Pabstiella) Anthosiphon (see Cryptocentrum) Anticheirostylis (see Genoplesium) Antichirostylis (see Genoplesium) Antillanorchis (see Tolumnia) Aopla (see Herminium) Aorchis (see Galearis) Apatostelis (see Stelis) Apaturia (see Pachystoma) Apista (see Podochilus) Aporopsis (see Dendrobium) Aporum (see Dendrobium) Appendiculopsis (see Agrostophyllum) Arachnanthe (see Arachnis) Archineottia (see Neottia) Arethusantha (see Cymbidium) Aretinum (see Cypripedium) Argyrorchis (see Macodes) Arisanorchis (see Cheirostylus) Aristotelea (see Spiranthes) Armodorum (see Arachnis) Arnottia (see Cynorkis) Arrhynchium (see Arachnis) Arthrochilum (see Epipactis) Asarca (see Chloraea) Ascocentropsis (see Vanda) Ascocentrum (see Vanda) Ascochilopsis (see Grosourdya) Ascochilus (see Grosourdya) Ascoglossum (see Renanthera) Ascolabium (see Holcoglossum) Ascotainia (see Tainia) Aspegrenia (see Octomeria) Auliza (see Epidendrum)

Aulizeum (see Epidendrum) Aulostylis (see Calanthe) Aurinocidium (see Grandiphyllum) Australorchis (see Dendrobium) Aviceps (see Satyrium) Baptistonia (see Gomesa) Barombia (see Aerangis) Barombiella (see Rangaeris) Basigyne (see Dendrochilum) Bathiea (see Neobathiea) Beadlea (see Cyclopogon) Bicchia (see Pseudorchis) Bieneria (see Chloraea) Binotia (see Gomesa) Biocornella (see Cynorkis) Birchea (see Luisia) Blephariglottis (see Platanthera) Blepharochilum (see Bulbophyllum) Blumeorchis (see Cleisostoma) Bolbodium (see Dendrobium) Bolbophyllaria (see Bulbophyllum) Bolbophyllopsis (see Bulbophyllum) Bolborkis (see Nervilia) Bollea (see Pescatoria) Bonniera (see Angraecum) Bothriochilus (see Coelia) Bouletia (see Dendrobium) Braasiella (see Tolumnia) Brachtia (see Brassia) Brachystepis (see Beclardia) Brasilidium (see Gomesa) Brasilocycnis (see Lueckelia) Brassiopsis (see Brassia) Brenesia (see Acianthera) Brevilongium (see Otoglossum) Briegeria (see Jacquiniella) Bucculina (see Holothrix) Buesiella (see Cyrtochilum) Burlingtonia (see Rodriguezia) Burnsbaloghia (see Deiregyne) Cadetia (see Dendrobium) Calcearia (see Corybas) Calceolus (see Cypripedium) Calliphyllon (see Epipactis) Callista (see Dendrobium) Callithronum (see Cephalanthera)

517

GENERIC SYNONYMS

Calyea (see Caleana) Camarotis (see Micropera) Camilleugenia (see Cynorkis) Canacorchis (see Bulbophyllum) Cannaeorchis (see Dendrobium) Carenidium (see Gomesa) Carparomorchis (see Bulbophyllum) Carria (see Gomesa) Carriella (see Gomesa) Carteretia (see Cleisostoma) Castroa (see Gomesa) Cattleyopsis (see Broughtonia) Celatorchis (see Solenangis) Centranthera (see Acianthera) Centrochilus (see Habenaria) Centrogenium (see Eltroplectris) Centropetalum (see Fernandezia) Centrosis (see Calanthe) Cephalangraecum (see Ancistrorhynchus) Cepobaculum (see Dendrobium) Ceraia (see Dendrobium) Ceratandropsis (see Ceratandra) Ceratobium (see Dendrobium) Ceratochilus (see Stanhopea) Ceratochilus (see Trichoglottis) Ceratopsis (see Epipogium) Cerochilus (see Hetaeria) Cestichis (see Stichorkis) Chaenanthe (see Comparettia) Chaetocephala (see Myoxanthus) Chamaeangis (see Diaphananthe) Chamaeleorchis (see Oncidium) Chamaerepes (see Chamorchis) Chaseëlla (see Bulbophyllum) Cheiropterocephalus (see Malaxis) Cheirorchis (see Thrixspermum) Chelonanthera (see Pholidota) Chelyorchis (see Rossioglossum) Chenorchis (see Holcoglossum) Chitonanthera (see Octarrhena) Chitonochilus (see Agrostophyllum) Chloidia (see Tropidia) Chlorosa (see Cryptostylis) Christensonia (see Vanda) Chromatotriccum (see Dendrobium) Chrysobaphus (see Anoectochilus) Chrysocycnis (see Mormolyca) Chrysoglossella (see Hancockia) Chrysoglossum (see Collabium) Chusua (see Ponerorchis) Cionosaccus (see Goodyera) Cirrhopetalum (see Bulbophyllum) Cladobium (see Lankesterella) Cladobium (see Scaphyglottis) Clinhymenia (see Cryptarrhena) Cnemidia (see Tropidia) Cochlia (see Bulbophyllum) Cochlioda (see Oncidium) Cocleorchis (see Cyclopogon)

518

Codonosiphon (see Bulbophyllum) Coelandria (see Dendrobium) Coeloglossum (see Dactylorhiza) Coenadenium (see Angraecopsis) Cogniauxiocharis (see Pteroglossa) Cohnia (see Trichocentrum) Cohniella (see Trichocentrum) Coilostylis (see Epidendrum) Collabiopsis (see Collabium) Collare-stuartense (see Oncidium) Concocidium (see Gomesa) Condylago (see Stelis) Conostalix (see Dendrobium) Coppensia (see Gomesa) Coralliokyphos (see Platylepis) Cordanthera (see Telipogon) Cordiglottis (see Thrixspermum) Cordula (see Paphiopedilum) Cordylestylis (see Goodyera) Corisanthes (see Cypripedium) Corunastylis (see Genoplesium) Corymborchis (see Corymborkis) Corysanthes (see Corybas) Costaricaea (see Scaphyglottis) Crinonia (see Pholidota) Criosanthes (see Cypripedium) Crocodeilanthe (see Stelis) Crossangis (see Rhipidoglossum) Cryptanthemis (see Rhizanthella) Cryptoglottis (see Podochilus) Cryptophoranthus (see Acianthera) Cryptosaccus (see Leochilus) Cryptosanus (see Leochilus) Ctenorchis (see Angraecum) Cutsis (see Dichromanthus) Cyanorchis (see Phaius) Cyathoglottis (see Sobralia) Cyclosia (see Mormodes) Cycnauken (see Cycnoches) Cydoniorchis (see Bifrenaria) Cylindrochilus (see Thrixspermum) Cylindrolobus (see Callostylis) Cymbiglossum (see Rhychostele) Cynosorchis (see Cynorkis) ×Cyperocymbidium (see Cymbidium) Cyperorchis (see Cymbidium) Cyrtochilos (see Cyrtochilum) Cyrtoglottis (see Mormolyca) Cyrtopera (see Eulophia) Cytherea (see Calypso) Cytheris (see Nephelaphyllum) Dactylorchis (see Dactylorhiza) Dactylorhynchus (see Bulbophyllum) Dactylostylis (see Zygostates) Darwiniera (see Telipogon) Dasyglossum (see Cyrtochilum) Davejonesia (see Dendrobium) Dendrobates (see Dendrobium) Dendrobianthe (see Polystachya)

Dendrocolla (see Thrixspermum) Dendrolirium (see Callostylis) Dendrorkis (see Dendrobium and Polystachya) Deppia (see Lycaste) Deroemera (see Holothrix) Diacrium (see Caularthron) Diadeniopsis (see Comparettia) Diadeniopsis (see Systeloglossum) Diadenium (see Comparettia) Dialissa (see Stelis) Dicerostylis (see Hylophila) Dichaeopsis (see Dichaea) Dichopus (see Dendrobium) Dicranotaenia (see Microcoelia) Dicrophyla (see Ludisia) Dicrypta (see Heterotaxis) Didactyle (see Bulbophyllum) Didiciea (see Tipularia) Didothion (see Epidendrum) Diglyphis (see Diglyphosa) Dignathe (see Cuitlauzina) Dikylikostigma (see Discyphus) Diothonea (see Epidendrum) Dipera (see Disperis) Diphyes (see Bulbophyllum) Diplacorchis (see Brachycorythis) Diplandrorchis (see Neottia) Diplecthrum (see Satyrium) Diplocaulobium (see Dendrobium) Diplochilus (see Diplomeris) Diploconchium (see Agrostophyllum) Diplodium (see Pterostylis) Diplogastra (see Platylepis) Diplolabellum (see Oreorchis) Dipteranthus (see Zygostates) Diskyphogyne (see Brachystele) Disperanthoceros (see Polystachya) Distichorchis (see Dendrobium) Dithyridanthus (see Deiregyne) Dituilis (see Liparis) Ditulima (see Dendrobium) Dockrillia (see Dendrobium) Dodsonia (see Stenia) Dolabrifolia (see Angraecum) Dolichocentrum (see Dendrobium) Doritis (see Phalaenopsis) Dorycheile (see Cephalanthera) Dothilophis (see Barkeria) Dressleriella (see Jacquiniella) Drymoda (see Bulbophyllum) Dryopeia (see Disperis) Dryorkis (see Disperis) Duboisia (see Myoxanthus) Duboisia-Reymondii (see Myoxanthus) Dungsia (see Cattleya) Durabaculum (see Dendrobium) Eburophyton (see Cephalanthera) Echinella (see Echinosepala) Echioglossum (see Cleisostoma)

GENERIC SYNONYMS

Ecuadoria (see Microthelys) Elasmatium (see Goodyera) Eleutheroglossum (see Dendrobium) Empusa (see Liparis) Empusaria (see Liparis) Encheiridion (see Microcoelia) Endeisa (see Dendrobium) Endresiella (see Trevoria) Eparmatostigma (see Vanda) Ephemerantha (see Dendrobium) Ephippium (see Bulbophyllum) Epicranthes (see Bulbophyllum) Epidanthus (see Epidendrum) Epidendropsis (see Epidendrum) Epidorkis/Epidorchis (see Oeonia) Epigeneium (see Dendrobium) Epilyna (see Elleanthus) Epiphanes (see Didymoplexis) Epiphora (see Polystachya) Epiphorella (see Polystachya) Epipogum (see Epipogium) Epithecia (see Prosthechea) Eriopexis (see Dendrobium) Erporchis (see Platylepis) Esmeralda (see Arachnis) Euanthe (see Vanda) Euchile (see Prosthechea) Eucnemis (see Govenia) Eucosia (see Goodyera) Eulophidium (see Oeceoclades) Euphlebium (see Dendrobium) Euproboscis (see Thelasis) Eurycaulis (see Dendrobium) Evardiana (see Odontochilus) Evardianthe (see Odontochilus) Evelyna (see Elleanthus) Evota (see Ceratandra) Exochanthus (see Dendrobium) Ferruminaria (see Bulbophyllum) Fieldia (see Vandopsis) Finetia (see Vanda) Fingardia (see Crepidium) Fissipes (see Cypripedium) Fitzgeraldia (see Lyperanthus) Flickingeria (see Dendrobium) Forficaria (see Disa) Fractiunguis (see Scaphyglottis) Fregea (see Sobralia) Froscula (see Dendrobium) Fruticicola (see Bulbophyllum) Gabertia (see Grammatophyllum) Galeorchis (see Galearis) Galera (see Epipogium) Garaya (see Mesadenella) Garayanthus (see Cleisostoma) Garayella (see Chamelophyton) Gastroglottis (see Dienia) Gastropodium (see Epidendrum) Geerinckia (see Polystachya)

Geissanthera (see Taeniophyllum) Genyorchis (see Bulbophyllum) Geobina (see Goodyera) Georchis (see Goodyera) Gerlachia (see Stanhopea) Gersinia (see Dendrobium) Ghiesbreghtia (see Calanthe) Gigliolia (see Octomeria) Giulianettia (see Glomera) Glossochilopsis (see Crepidium) Glossorhyncha (see Glomera) Glossula (see Peristylus) Goadbyella (see Microtis) Goldschmidtia (see Dendrobium) Gomphostylis (see Pleione) Goniochilus (see Leochilus) Gonogona (see Goodyera) Gorgoglossum (see Sievekingia) Govindooia (see Tropidia) Gracielanthus (see Aulosepalum) Grafia (see Phalaenopsis) Grastidium (see Dendrobium) Greenwoodia (see Kionophyton) Grussia (see Phalaenopsis) Gudrunia (see Tolumnia) Gularia (see Schiedeella) Gunnaria (see Vanda) Gunnia (see Sarcochilus) Gussonea (see Microcoelia) Gyaladenia (see Brachycorythis) Gyalanthos (see Pabstiella) Gymnadeniopsis (see Platanthera) Gymnochilus (see Cheirostylus) Gyrostachys (see Spiranthes) Habenorkis (see Habenaria) Hadrolaelia (see Cattleya) Haemaria (see Ludisia) Hallackia (see Huttonaea) Hamularia (see Bulbophyllum) Hapalochilus (see Bulbophyllum) Haplochilus (see Zeuxine) Haraella (see Gastrochilus) Harrisella (see Campylocentrum) Harrisella (see Dendrophylax) Hecabe (see Phaius) Helcia (see Trichopilia) Hellerorchis (see Gomesa) Helorchis (see Cynorkis) Hemiperis (see Cynorkis) Hemiscleria (see Epidendrum) Hemohabenaria (see Pecteilis) Henosis (see Bulbophyllum) Herpethophytum (see Dendrobium) Herschelia (see Disa) Herschelianthe (see Disa) Heteroncidium (see Oncidium) Heterozeuxine (see Zeuxine) Hexadesmia (see Scaphyglottis) Hexameria (see Podochilus)

Hexisea (see Scaphyglottis) Hippoglossum (see Bulbophyllum) Hipporkis (see Satyrium) Hirtzia (see Pterostemma) Hispaniella (see Tolumnia) Hoffmansegella (see Cattleya) Hologyne (see Coelogyne) Holopogon (see Neottia) Holmesia (see Angraecopsis) Homocollecticon (see Cyrtorchis) Hordeanthos (see Bulbophyllum) Hormidium (see Prosthechea) Humboldtia (see Stelis) Hyalosema (see Bulbophyllum) Hybochilus (see Leochilus) Hydranthus (see Dipodium) Hygrochilus (see Phalaenopsis) Hymeneria (see Pinalia) Hypodema (see Cypripedium) Hypodematium (see Eulophia) Hysteria (see Corymborkis) Ibidium (see Spiranthes) Ichthyostomum (see Bulbophyllum)

Ida (corrected as Sudamerlycaste, this volume) Iebine (see Liparis) India (see Robiquetia) Inobulbum (see Dendrobium) Ione (see Bulbophyllum) Ionorchis (see Limodorum) Irenea (see Cyrtochilum) Iridorchis (see Cymbidium) Iridorchis (see Oberonia) Iridorkis (see Oberonia) Ischnocentrum (see Glomera) Isochilostachya (see Polystachya) Jamaiciella (see Tolumnia) Jansenia (see Plectrophora) Jejosephia (see Bulbophyllum) Jenmania (see Palmorchis) Jennyella (see Houlletia) Jensoa (see Cymbidium) Josephia (see Sirhookera) Jostia (see Masdevallia) Kalimpongia (see Dickasonia) Kalopternix (see Epidendrum) Katharinea (see Dendrobium) Kaurorchis (see Bulbophyllum) Kegelia (see Kegeliella) Kerigomnia (see Octarrhena) Kinetochilus (see Dendrobium) Kingidium (see Phalaenopsis) Kingiella (see Phalaenopsis) Kitigorchis (see Oreorchis) Kleberiella (see Gomesa) Kochiophyton (see Aganisia) Konantzia (see Ionopsis) Kornasia (see Orestias) Kryptostoma (see Habenaria)

519

GENERIC SYNONYMS

Lacroixia (see Dinklageella) Laelia p.p. (see Cattleya) Laeliopsis (see Broughtonia) Lanium (see Epidendrum) Laricorchis (see Ornithidium) Larnandra (see Epidendrum) Latourorchis (see Dendrobium) Leaoa (see Scaphyglottis) Lectandra (see Poaephyllum) Leioanthum (see Dendrobium) Lemboglossum (see Rhychostele) Lemuranthe (see Cynorkis) Leopardanthus (see Dipodium) Lepanthanthe (see Bulbophyllum) Lepervenchea (see Angraecum) Leptocentrum (see Plectrelminthus) Leptoceras (see Leporella) Leptorchis (see Liparis) Leptorkis (see Liparis) Leptothrium (see Isochilus) Lequeetia (see Limodorum) Lesliea (see Phalaenopsis) Leucohyle (see Trichopilia) Leucolena (see Didymoplexiella) Leucorchis (see Pseudorchis) Leucostachys (see Goodyera) Lichenora (see Porpax) Lichterveldia (see Cuitlauzina) Limatodes (see Phaius) Limnas (see Malaxis) Limnorchis (see Platanthera) Lindleyella (see Rudolfiella) Lindsayella (see Sobralia) Lisowskia (see Orestias) Lissochilus (see Eulophia) Listera (see Neottia) Lobogyne (see Appendicula) Lophiarella (see Trichocentrum) Lophiaris (see Trichocentrum) Lophoglotis (see Cattleya) Loroglossum (see Himantoglossum) Lothiania (see Porroglossum) Loxomorchis (see Smithsonia) Lueranthos (see Andinia) Luerella (see Phloeophila) Luisiopsis (see Gastrochilus) Lyraea (see Bulbophyllum) Lysiella (see Platanthera) Maccraithea (see Dendrobium) Macdonaldia (see Thelymitra) Macrochilus (see Miltonia) Macrolepis (see Bulbophyllum) Macroplectrum (see Angraecum) Macrostomium (see Dendrobium) Macrostylis (see Corymborkis) Maelenia (see Cattleya) Malachadenia (see Bulbophyllum) Malleola (see Robiquetia) Mariarisqueta (see Cheirostylus)

520

Marsupiaria (see Heterotaxis) Masdevalliantha (see Pleurothallis) Mastigion (see Bulbophyllum) Megaclinium (see Bulbophyllum) Megalotus (see Robiquetia) Meliclis (see Coryanthes) Menadena (see Maxillaria) Menadenium (see Zygosepalum) Menezesiella (see Gomesa) Mesoclastes (see Luisia) Mesodactylus (see Apostasia) Mesoglossum (see Rhychostele) Mesoptera (see Liparis) Mesospinidium (see Brassia) Metachilum (see Appendicula) Mexicoa (see Oncidium) Microholmesia (see Angraecopsis) Microlaelia (see Cattleya) Microphytanthe (see Dendrobium) Microtatorchis (see Taeniophyllum) Microterangis (see Aerangis) Microtheca (see Cynorkis) Miltonioides (see Oncidium) Mirandopsis (see Pleurothallis) Mischobulbum (see Tainia) Mitostigma (see Amitostigma) Moerenhoutia (see Platylepis) Monachanthus (see Catasetum) Monadenia (see Disa) Monanthochilus (see Sarcochilus) Monanthos (see Dendrobium) Monixus (see Angraecum) Monochilus (see Zeuxine) Monomeria (see Bulbophyllum) Monorchis (see Herminium) Monosepalum (see Bulbophyllum) Monotris (see Holothrix) Montolivaea (see Habenaria) Monustes (see Spiranthes) Muluorchis (see Tropidia) Myanthus (see Catasetum) Myoda (see Ludisia) Myrobrosma (see Vanilla) Mystacorchis (see Stelis) Nageliella (see Domingoa) Nanodes (see Epidendrum) Nasonia (see Fernandezia) Nauenia (see Lacaena) Neippergia (see Acineta) Neo-urbania (see Ornithidium) Neobartlettia (see Palmorchis) Neobennettia (see Lockhartia) Neobenthamia (see Polystachya) Neoburttia (see Polystachya) Neocribbia (see Solenangis) Neodryas (see Cyrtochilum) Neoescobaria (see Trichopilia) Neofinetia (see Vanda) Neokoehleria (see Comparettia)

Neolauchea (see Isabelia) Neolehmannia (see Epidendrum) Neoruschia (see Gomesa) Neottidium (see Neottia) Neowilliamsia (see Epidendrum) Neowolffia (see Podangis) Nephranthera (see Renanthera) Nidus (see Neottia) Niemeyera (see Apostasia) Nienokuea (see Polystachya) Nigritella (see Gymnadenia) Nipponorchis (see Vanda) Nothodoritis (see Phalaenopsis) Nyctosma (see Epidendrum) Oakes-amesia (see Ornithocephalus) Ochyrella (see Pteroglossa) Octadesmia (see Dilomilis) Odontoglossum (see Oncidium) Odontostylis (see Bulbophyllum) Oerstedella (see Epidendrum) Oestlundorchis (see Deiregyne) Olgasis (see Tolumnia) Ommatodium (see Pterygodium) Oncodia (see Brassia) Oncophyllum (see Bulbophyllum) Onychium (see Dendrobium) Ophidion (see Phloeophila) Orchiastrum (see Spiranthes) Orchidofunckia (see Cryptarrhena) Orchidotypus (see Fernandezia) Orchiodes (see Goodyera) Orchites (see Traunsteinera) Ormerodia (see Cleisostoma) Ormostema (see Dendrobium) Ornitharium (see Pteroceras) Ornithochilus (see Phalaenopsis) Ornithophora (see Gomesa) Orsidice (see Thrixspermum) Orthochilus (see Eulophia) Orthoglottis (see Dendrobium) Orthopenthea (see Disa) Orxera (see Aerides) Osmoglossum (see Cuitlauzina) Osyricera (see Bulbophyllum) Otopetalum (see Kraenzliniella) Oxyanthera (see Thelasis) Oxyglossellum (see Dendrobium) Oxysepala (see Bulbophyllum) Pachychilus (see Pachystoma) Pachyne (see Phaius) Pachyphyllum (see Fernandezia) Pachystele (see Scaphyglottis) Paliris (see Liparis) Palumbina (see Cuitlauzina) Panstrepis (see Coryanthes) Pantlingia (see Stigmatodactylus) Papperitzia (see Leochilus) Papulipetalum (see Bulbophyllum) Paragnathis (see Diplomeris)

GENERIC SYNONYMS

Paraholcoglossum (see Holcoglossum) Parapactis (see Epipactis) Parapteroceras (see Tuberolabium) Parasarcochilus (see Sarcochilus) Parhabenaria (see Pecteilis) Parlatorea (see Sanderella) Pattonia (see Grammatophyllum) Pedilea (see Dienia) Pedilochilus (see Bulbophyllum) Pedilonum (see Dendrobium) Pelma (see Bulbophyllum) Peltopus (see Bulbophyllum) Penkimia (see Holcoglossum) Penthea (see Disa) Perrieriella (see Oeonia) Pesomeria (see Phaius) Petalocentrum (see Oncidium) Petalochilus (see Caladenia) Petronia (see Batemannia) Pfitzeria (see Comparettia) Phaeus (see Phaius) Phajus (see Phaius) Philippinaea (see Orchipedum) Phormangis (see Ancistrorhynchus) Phragmopedilum (see Phragmipedium) Phyllomphax (see Brachycorythis) Phyllorkis (see Bulbophyllum) Phymatochilum (see Miltonia) Physanthera (see Rodriguezia) Physinga (see Epidendrum) Physosiphon (see Stelis) Physothallis (see Stelis) Physurus (see Erythrodes) Pierardia (see Dendrobium) Pilumna (see Trichopilia) Pinelia (see Homalopetalum) Pinelianthe (see Homalopetalum) Piperia (see Platanthera) Pittierella (see Cryptocentrum) Placostigma (see Podochilus) Platyclinis (see Dendrochilum) Platyglottis (see Scaphyglottis) Platysma (see Podochilus) Platystylis (see Liparis) Pleuranthium (see Epidendrum) Plexaure (see Phreatia) Podandria (see Habenaria) Podandriella (see Habenaria) Podanthera (see Epipogium) Podochilopsis (see Adenoncos) Polychilos (see Phalaenopsis) Polycycnopsis (see Polycycnis) Polyradicion (see Dendrophylax) Polyrrhiza (see Dendrophylax) Polystylus (see Phalaenopsis) Polytoma (see Aerides) Porphyrodesme (see Renanthera) Portillia (see Masdevallia) Pristiglottis (see Odontochilus)

Pseudepidendrum (see Epidendrum) Pseuderiopsis (see Eriopsis) Pseudocranichis (see Galeoglossum) Pseudoctomeria (see Specklinia) Pseudoeurystyles (see Eurystyles) Pseudoliparis (see Crepidium) Pseudomacodes (see Macodes) Pseudomaxillaria (see Camaridium) Pseudoperistylis (see Habenaria) Pseudostelis (see Stelis) Psilanthemum (see Epidendrum) Psittacoglossum (see Camaridium) Psychechilos (see Zeuxine) Psygmorchis (see Erycina) Pterochilus (see Crepidium) Pteroglossaspis (see Eulophia) Ptilocnema (see Pholidota) Ptychogyne (see Coelogyne) Pycnantha (see Malaxis) Queteletia (see Orchipedum) Raciborskanthos (see Cleisostoma) Radinocion (see Aerangis) Ramonia (see Scaphyglottis) Raycadenco (see Fernandezia) Reichenbachanthus (see Scaphyglottis) Renantherella (see Renanthera) Renata (see Pseudolaelia) Restrepiopsis (see Pleurothallopsis) Reymondia (see Myoxanthus) Rhamphidia (see Hetaeria) Rhaphidorhynchus (see Aerangis) Rhinocerotidium (see Gomesa) Rhinocidium (see Gomesa) Rhizocorallon (see Corallorhiza) Rhynchadenia (see Macradenia) Rhynchopera (see Pleurothallis) Rhynchophreatia (see Thelasis) Rhytionanthos (see Bulbophyllum) Ridleya (see Thrixspermum) Rodrigueziella (see Gomesa) Rodrigueziopsis (see Gomesa) Roezliella (see Oncidium) Rolfea (see Palmorchis) Rolfeella (see Benthamia) Rusbyella (see Cyrtochilum) Rynchanthera (see Corymborkis) Saccidium (see Holothrix) Saccochilus (see Saccolabium) Saccoglossum (see Bulbophyllum) Sacodon (see Cypripedium) Salacistis (see Goodyera) Salpistele (see Stelis) Samarorchis (see Robiquetia) Santanderella (see Notyliopsis) Sarcanthus (see Acampe) Sarcocadetia (see Dendrobium) Sarcoglossum (see Cirrhaea) Sarcorhynchus (see Rhipidoglossum) Sarmenticola (see Macroclinium)

Sarothrochilus (see Trichoglottis) Sarracenella (see Acianthera) Satyridium (see Satyrium) Saurolophorkis (see Crepidium) Sayeria (see Dendrobium) Scelochiloides (see Comparettia) Scelochilopsis (see Comparettia) Scelochilus (see Comparettia) Schidorhynchos (see Sauroglossum) Schismoceras (see Dendrobium) Schizodium (see Disa) Schlechterella (see Rudolfiella) Schluckebieria (see Cattleyella) Schoenomorphus (see Tropidia) Schomburgkia (see Laelia) Schwartzkopffia (see Brachycorythis) Scleropteris (see Cirrhaea) Scoliochilus (see Appendicula) Scopularia (see Holothrix) Sedirea (see Phalaenopsis) Seidenfia (see Crepidium) Selenipedilum (see Selenipedium) Senghasia (see Kefersteinia) Senghasiella (see Habenaria) Sepalosaccus (see Camaridium) Sepalosiphon (see Glomera) Seraphyta (see Epidendrum) Serpenticaulis (see Bulbophyllum) Serrastylis (see Macradenia) Sestochilos (see Bulbophyllum) Siagonanthus (see Ornithidium) Siederella (see Cyrtochilum) Sigmatochilus (see Chelonistele) Sigmatogyne (see Panisea) Sigmatostalix (see Oncidium) Sinorchis (see Aphyllorchis) Sodiroella (see Telipogon) Solenidiopsis (see Oncidium) Sophronia (see Cattleya) Sophronitella (see Isabelia) Sophronitis (see Cattleya) Spathiger (see Epidendrum) Spathium (see Epidendrum) Sphyrastylis (see Ornithocephalus) Spilorchis (see Bulbophyllum) Stacyella (see Erycina) Stanhopeastrum (see Stanhopea) Stauritis (see Phalaenopsis) Staurochilus (see Trichoglottis) Stauroglottis (see Phalaenopsis) Stauropsis (see Trichoglottis) Stelbophyllum (see Dendrobium) Steliopsis (see Stelis) Stellilabium (see Telipogon) Stellorkis (see Nervilia) Stenocoryne (see Bifrenaria) Stenoglossum (see Epidendrum) Stenopolen (see Stenia) Stevenorchis (see Steveniella)

521

GENERIC SYNONYMS

Stichorchis (see Stichorkis) Stictophyllorchis (see Quekettia) Stictophyllum (see Quekettia) Stigmatosema (see Cyclopogon) Stillifolium (see Trichocentrum) Stimegas (see Paphiopedilum) Strateuma (see Orchis) Sturmia (see Liparis) Styloglossum (see Calanthe) Sunipia (see Bulbophyllum) Symphyglossum (see Oncidium) Synadena (see Phalaenopsis) Synanthes (see Eurystyles) Synarmosepalum (see Bulbophyllum) Synassa (see Sauroglossum) Synptera (see Trichoglottis) Szlachetkoella (see Polystachya) Tadeastrum (see Stanhopea) Tamayorkis (see Malaxis) Tangtsinia (see Cephalanthera) Tankervillia (see Phaius) Tapeinoglossum (see Bulbophyllum) Taurostalix (see Bulbophyllum) Telopogon (see Telipogon) Tetrabaculum (see Dendrobium) Tetragamestus (see Scaphyglottis) Tetrapeltis (see Otochilus) Tetrodon (see Dendrobium) Theana (see Grosourdya) Thelychiton (see Dendrobium) Thelypogon (see Telipogon) Theodorea (see Gomesa) Thicuania (see Dendrobium) Thorvaldsenia (see Chysis)

522

Thylacis (see Thrixspermum) Ticoglossum (see Rossioglossum) Tinaea (see Neotinea) Tinea (see Neotinea) Titania (see Oberonia) Todaroa (see Campylocentrum) Tomotris (see Corymborkis) Trachelosiphon (see Eurystyles) Trachyrhachis (see Bulbophyllum) Trachyrhizum (see Dendrobium) Triaristella (see Trisetella) Trias (see Bulbophyllum) Tribrachia (see Bulbophyllum) Triceratostris (see Deiregyne) Trichochilus (see Dipodium) Trichorhiza (see Luisia) Trichosma (see Eria) Tridachne (see Notylia) Trigonochilum (see Cyrtochilum) Triorchis (see Spiranthes) Tripleura (see Zeuxine) Tripudianthes (see Bulbophyllum) Tritelandra (see Epidendrum) Trophianthus (see Aspasia) Tropilis (see Dendrobium) Trudelia (see Vanda) ×Trudelianda (see Vanda) Tryphia (see Holothrix) Tsaiorchis (see Diphylax) Tsiorchis (see Holcoglossum) Tubilabium (see Myrmechis) Tulotis (see Platanthera) Tussaca (see Goodyera) Tussacia (see Spiranthes)

Tylochilus (see Cyrtopodium) Ulantha (see Chloraea) Unguiculabia (see Polystachya) Uropedium (see Phragmipedium) Urostachya (see Pinalia) Vanillophorum (see Vanilla) Vappodes (see Dendrobium) Ventricularia (see Trichoglottis) Vermeulenia (see Anacamptis) Vesicisepalum (see Bulbophyllum) Vexillabium (see Kuhlhasseltia) Vieillardorchis (see Goodyera) Vonroemeria (see Octarrhena) Wailesia (see Dipodium) Wallnoeferia (see Helonoma) Waluewa (see Gomesa) Warczewitzia (see Clowesia) Warscaea (see Cyclopogon) Winika (see Dendrobium) Xaritonia (see Tolumnia) Xeilyathum (see Oncidium) Xiphizusa (see Bulbophyllum) Xiphophyllum (see Cephalanthera) Xiphosium (see Cryptochilus) Yoania (see Danhatchia) Yolanda (see Brachionidium) Zeduba (see Calanthe) Zetagyne (see Panisea) Zhukowskia (see Sarcoglottis) Zoophora (see Orchis) Zosterophyllanthos (see Pleurothallis) Zosterostylis (see Cryptostylis) Zygoglossum (see Bulbophyllum) Zygopetalon (see Zygopetalum)

SERIES CONTRIBUTORS

We offer our profound thanks to the following individuals for their authoritative contributions to Genera Orchidacearum since its inception. ACKERMAN, JAMES D.

JOHNSON, STEVE

ADAMS, PETER B.

JONES, DAVID

ATWOOD, JOHN

KLEIN, BERT

BAILES, CHRISTOPHER

KOEHLER, SAMANTHA

BARKMAN, TODD J.

KOOPOWITZ, HAROLD

BARROS, FÁBIO DE

KORES, PAUL

BATEMAN, RICHARD M.

KUMAR, C. SATHISH

BATISTA, JOÃO A. N.

KURZWEIL, HUBERT

BIANCHETTI, LUCIANO B.

LAWSON, SHERYL D.

BLANCO, MARIO

LIGHT, MARILYN H. S.

BOWER, COLIN C.

LINDER, PETER

BRANDHAM, PETER

LUO, YI-BO

CAMERON, KENNETH M.

MOLLOY, BRIAN

CARLSWARD, BARBARA S.

MOLVRAY, MIA

CARNEVALI FERNÁNDEZ-CONCHA, GERMÁN

MONTEIRO, SILVANA HELENA

CHASE, MARK W.

MOTES, MARTIN

CLEMENTS, MARK A.

NEILAND, M. RUTH M.

CORREA, MAEVIA N.

OAKELEY, HENRY

COX, ANTONY

OJEDA, ISIDRO

COZZOLINO, SALVATORE

ORMEROD, PAUL

CRIBB, PHILLIP J.

PANSARIN, EMERSON R.

CYWES, SID

NG, YAN PENG

DAFNI, AMOTS

PERNER, HOLGER

D’EMERICO, SAVERIO

PRESTON, JILLIAN

DE VOGEL, ED F.

PRIDGEON, ALEC M.

DODSON, CALAWAY H.

PUPULIN, FRANCO

DRESSLER, ROBERT L.

RAMÍREZ, IVÓN M.

FERNANDO, R. H. S. SURANJAN

RASMUSSEN, FINN N.

FISCHER, GÜNTER

RICHARDS, HELEN

FREUDENSTEIN, JOHN V.

ROTHACKER, ERIK PAUL

GARCÍA, JAVIER

ROMERO-GONZÁLEZ, GUSTAVO A.

GARDINER, LAUREN M.

SALAZAR, GERARDO A.

GERLACH, GÜNTER

SÁNCHEZ, MARCELA I.

GOLDMAN, DOUGLAS H.

SCHUITEMAN, ANDRÉ

GRAVENDEEL, BARBARA

SIEDER, ANTON

GRAYER, RENÉE J.

SINGER, RODRIGO

HÁGSATER, ERIC

SMIDT, ERIC DE CAMARGO

HERMANS, JOHAN

SOLANO GOMEZ, RODOLFO

HIGGINS, WESLEY E.

SOSA, VICTORIA

HOLLINGSWORTH, MICHELLE L.

SOTO ARENAS, MIGUEL ANGEL

HOLLINGSWORTH, PETER M.

SQUIRRELL, JANE

HOPPER, STEPHEN D.

STEINER, KIM E.

523

SERIES CONTRIBUTORS

STENZEL, HAGEN

VERMEULEN, JAAP JAN

STERN, WILLIAM LOUIS

VOGELPOEL, LOUIS

STONE, JUDI

VAN VUGT, ROGIER RODERIK

TAPIA, JOSÉ LUIS

WHITTEN, WILLIAM MARK

TOSCANO DE BRITO, A. L. V.

WIDMER, ALEX

VAN DEN BERG, CASSIO

WOOD, JEFFREY

VEITCH, NIGEL C.

524

Index To Scientific Names Page numbers in boldface refer to relevant illustrations of taxa. See Generic Synonyms elsewhere in the end matter for names not listed here. Acampe 111, 118, 119, 123, 125, 128, 129–32, 168, 314 A. multiflora 129 A. ochracea 131 A. pachyglossa 102, 131 A. papillosa 129, 131 A. praemorsa 130, 131, 132 A. rigida 131, 132, Pl. 57 Ada 480–1 Adenoncos 118, 124, 132–3, 134, 204, 212, 327 A. major 132 A. parviflora 133 A. sumatrana Pl. 58 A. virens 132 Adrorhizinae 102, 104–11 Adrorhizon 104–6 A. purpurascens 104–6, 105, Pl. 54 Aerangis 339, 340, 344–9, 353, 362, 395, 396, 442 A. appendiculata 345 A. articulata Pl. 134 A. biloba 345, 349 A. brachycarpa 345, 349 A. confusa 345, 346, 347, 349 A. coriacea 345 A. cryptodon 345 A. decaryana 348 A. distincta 345, 346 A. ellisii 340, 348, 349 A. fastuosa 345 A. fuscata 349 A. flabellifolia 344 A. gracillima 345 A. hariotiana 347, Pl. 135 A. hildebrandtii 345 A. hologlottis 102 A. kirkii 345, 346, 347 A. kotschyana 345, 349 A. modesta 346 A. monantha Pl. 136 A. pulchella 348 A. punctata 348 A. somalensis 345, 346 A. thomsonii 349 A. ugandensis 345 A. verdickii 345 subgenus Aerangis 345 subgenus Barombia 345 subgenus Uniflorae 345 Aeranthes 340, 342, 349–51, 393, 395 A. africana 350 A. arachnites 351 A. caudata 351 A. crassifolia 351

A. filipes Pl. 137 A. grandiflora 349, 351 A. parvula 351 A. peyrotii 350 A. ramosa 350, 351 A. sambiranoensis 351 A. schlechteri 351, Pl. 138 A. tropophila 351 Aerides 112, 118, 119, 120, 128, 129, 133–7, 141, 327 A. crassifolia 135, 136 A. crispa 135 A. falcata 134, 137 A. fieldingii 134, 135 A. krabiensis 136 A. lawrenceae 135, 136 A. multiflora 113, 135, 136, 137 A. odorata 112, 133, 134, 135, 136, 137 A. quinquevulnera Pl. 59 A. ringens 136 A. rosea 134, 135, 136 Aeridinae 101, 102, 111–331 Agrostophyllinae 101, 104, 331–8 Agrostophyllum 101, 102, 331–5 A. brevipes 333, 334 A. callosum 332, 333, 334 A. glumaceum 333 A. javanicum 331 A. laterale 332 A. majus 333, Pl. 132 A. paniculatum 332 A. planicaule 332, 333, 334 A. stipulatum 332 subspecies bicuspidatum 333, 334 A. superpositum 332 section Agrostophyllum 332, 334 section Appendiculopsis 332, 334 section Dolichodesme 332 section Oliganthe 332 Ambrella 342, 351–3 A. longituba 351–3, 352, Pl. 139 Amesiella 117, 119, 126, 137–9, 148, 203, 209, 268, 310, 315 A. philippinensis 137, 138, 138, 139, Pl. 60 Ancistrorhynchus 340, 343, 344, 353–5, 409, 436 A. cephalotes Pl. 140 A. clandestinus 353, 355 A. metteniae 353, 354, 355 A. recurvus 353 A. refractus 353, 354 A. tenuicaulis 354 Angraecinae 101, 102, 338–450 Angraecopsis 340, 343, 355–8, 407

A. amaniensis 356, 357, 440 A. breviloba 356, 357, 440 A. gracillima 357 A. parviflora 356, 357, 362, Pl. 141 A. pusilla 357 A. tenerrima 355, 357, 440 section Angraecopsis 357 section Cardiochilos 357 section Coenadenium 357, 440 Angraecum 340, 342, 358–63, 402, 444 A. aff. doratophyllum 362 A. aloifolium 363 A. aporoides 362 A. appendiculata 361 A. arachnites 340, 363 A. birrimense 361, 362 A. calceolus 361 A. caulescens 360 A. chimanimaniense 359 A. chloranthum 360, 362 A. compactum 363 A. conchiferum 361 A. conchoglossum 340 A. cornigerum 362 A. coutrixii 362 A. cultriforme 361, 362 A. decaryanum 363 A. distichum 360, 361, 362 A. dives 361 A. dollii 363 A. eburneum 340, 358, 362, 363, Pl. 142 subspecies giryamae 361 subspecies superbum 361, 362 subspecies xerophilum 361, 363 A. eichlerianum 361, 362, 363 A. erectum 361 A. filicornu 360 A. gabonense 361 A. geniculatum 359 A. germinyanum 340, 361, 363 A. hermannii 362 A. infundibulare 362 A. leonis 340, 361, 363 A. longicalcar 363 A. madagascariense 361, 363 A. magdalenae 362, 363, Pl. 143 A. mauritianum 361 A. microcharis 363 A. moandense 361, 362 A. multinominatum 361 A. myrianthum 363 A. nanum 361 A. obversifolium 362

525

I nde x T o S cientific N ames

Angraecum (cont.) A. palmicolum 363 A. pectinatum 361, 362 A. pingue 360, 362 A. popowii 363 A. protensum 363 A. pseudofilicornu 340 A. pumilio 360 A. pungens 361 A. ramosum 360, 362 A. reygaertii 360 A. rhynchoglossum 360 A. rubellum 363 A. rutenbergianum 362 A. sacciferum 361 A. scottianum 340 A. sedifolium 362 A. sesquipedale 340, 361, 362, 363, Pl. 144 var. angustifolium 362, 363 A. sororium 340, 363 A. striatum 360 A. subulatum 361, 362 A. tenuifolium 361 A. teres 361 A. triquetrum 361 A. viguieri Pl. 145 A. viride 360 A. zeylanicum 102, 358 section Acaulia 358, 360 section Afangraecum 360 section Angraecoides 358, 360, 362 section Angraecum 360 section Arachnangraecum 358, 360, 362 section Boryangraecum 360 section Chlorangraecum 360, 362 section Conchoglossum 358, 360 section Dolabrifolia 358, 360, 362 section Filangis 358, 360, 362 section Gomphocentrum 359, 360, 362 section Hadrangis 360, 362 section Humblotiangraecum 360, 361, 362 section Lemurangis 360, 361, 362 section Lepervenchea 360, 361, 362 section Nana 360, 361 section Pectinaria 358, 361, 362 section Perrierangraecum 358, 361, 362 section Pseudojumellea 358, 361, 362 Arachnis 118, 119, 125, 129, 136, 139–41 A. beccarii 141 A. calcaratum 141 A. cathcartii 141 A. clarkei 141 A. flos-aeris 139, 140, 141 A. grandisepala 141, Pl. 61 A. hookeriana 141 A. labrosa 140 A. longisepala 140, 141 A. siamense 140 Archivea 485 Beclardia 340, 344, 363–5, 393, 401 B. grandiflora 364, 365 B. macrostachya 363, 364, 364, 365, Pl. 146 Bhutanthera 485 Biermannia 117, 125, 142, 143, 144, 157, 230, 297 B. bimaculata 142, 143 B. calcarata 142 B. ciliata 142

526

B. quinquecallosa 142 Bogoria 116, 126, 142–4, 145, 146, 254 B. raciborskii 142, 145 B. taeniorhiza 102, 142 Bolusiella 340, 341, 343, 344, 355, 365–7, 409, 436 B. batesii 367 B. iridifolia 366, 367 B. maudiae 365, 366, 367, Pl. 147 Bonniera 340, 341 Brachtia 481 Brachypeza 115, 116, 124, 126, 144–6, 147, 148, 193, 241, 251, 252, 297 B. archytas 144, 145, 297 B. indusiata 145, Pl. 62 B. semiteretifolia 124 B. zamboangensis 145, 147 Brassia 480–2 B. allenii 480 B. aurantiaca 481 B. caudata 482 B. maculata 480, 482 B. signata 482 Bromheadia 101, 102, 104, 106–9 B. alticola 108, 109 B. aporoides 108, 109 B. borneensis 107 B. finlaysoniana 106, 108, 109 B. grandiflora Pl. 55 B. pendek 106 section Aporodes 106, 107, 108 section Bromheadia 106, 107, 108 Bulbophyllum 3–51 B. absconditum 33 B. acutiflorum 43 B. ambrosia 43, 50 B. amplebracteatum subsp. orthoglossum 31, Pl. 1 B. andersonii 42, 44 B. antheae 7 B. apertum 46 B. apetalum 14 B. attenuatum 31 B. baileyi 46 B. balaeniceps 26 B. barbasapientis 25 B. barbigerum 15 B. beccarii 26, 47 B. bicolor Pl. 2 B. bifarium 13 B. biflorum 27 B. bisetoides 21 B. bisetum 27 B. blepharistes 27, 47 B. brachystachyum 16 B. bracteatum 26 B. bracteolatum 41 B. calyptratum 8 B. cardiobulbum 17 B. careyanum 35, 41, 43, 50 B. carnosisepalum 14 B. cheiri 46 B. chloropterum 41 B. clandestinum 43 B. clavatum 43 B. cochleatum Pl. 3 B. codonanthum 28 B. coloratum Pl. 4 B. comatum 14 B. coriaceum 41

B. coriophorum 18 B. corolliferum 35 B. croceum 28 B. dasypetalum 42 B. dayanum Pl. 5 B. delitescens Pl. 6 B. denticulatum 14 B. dichotomum 20, 21 B. digitatum 29 B. dryas 26 B. drymoda 28 B. eberhardtii Pl. 7 B. elatum 45 B. ellipticifolium 24 B. elongatum 26 B. emarginatum 29 B. epibulbon 29 B. epicrianthes 29 B. erythrosema Pl. 8 B. eublepharon 29 B. exaltatum 41 B. falcatum 15 B. farinulentum 22 B. fascinator 24, 45 B. fischeri 42, 43 B. fossatum 25 B. fuscopurpureum 26, 44 B. gibbosum 38, 43, 45 B. gilgianum 14 B. glaucum 25 B. gracile 42, 43 B. gracillimum 46 B. grandiflorum 30 B. graveolens 45 B. griffithii 25, 42, 43 B. guttulatum 44, Pl. 9 B. gymnopus 20, 44 B. helenae 36 B. hemiprionotum 24 B. hemisterranthum 29 B. hirtulum 30 B. hirtum 43 B. ikongoense 17 B. imitator 30 B. implexum 16 B. incisilabrum Pl. 10 B. inconspicuum 44 B. infundibuliforme 30 B. involutum 46 B. ipanamense 46 B. khasyanum 42, 43 B. kingii 25 B. kwantungense 44 B. lemniscatum 31 B. leopardinum 31, 42, 43, 44, 50 B. lepanthiflorum 31 B. lichenophylax 17 B. lindleyanum Pl. 11 B. lipense 25 B. lobbii 38, 42, 43, 45, 50 B. longiflorum 14, 28 B. longipes 30 B. longiscapum Pl. 12 B. longissimum 28, 50 B. lyperocephalum 18 B. macraei 32 B. macrourum 32 B. makoyanum 42, 43

Index To S cientific Names

B. maxillarioides 27 B. maximum 43 B. medusae 47 B. membranifolium 43 B. micholitzianum 43 B. micranthum 41 B. minutissimum 32 B. mirificum 17 B. molossus 42, Pl. 13 B. monomeria 33, Pl. 14 B. moratii 18 B. multiflorum 43 B. muricatum 33 B. muscicola 16, 41 B. nagelii 41 B. napellii 41 B. nummularia Pl. 15 B. nummularioides Pl. 16 B. nutans 4, 17 B. occlusum 16 B. occultum 14, 18, 41, 43 B. ochroleucum 40 B. odoratissimum 42, 44, 45, 336 B. oreonastes 15 B. ornatissimum 43, 45 B. osyricera 9 B. ovalifolium 32 B. oxycalyx 17 B. pachypus 18 B. pantoblepharon 18 B. papuanum 33 B. papulipetalum 33 B. patens 46, Pl. 17 B. pedilochilus Pl. 18 B. peltopus 33 B. penduliscapum 42, 47 B. penicillium 42, 43 B. petrae 19 B. phreatiopse 34 B. piestobulbon 34, Pl. 19 B. planibulbe 34 B. plumatum 34, Pl. 20 B. protractum 45 B. psittacoglossum Pl. 21 B. pseudohydra 14 B. pseudopelma 35 B. pygmaeum 41 B. rariflorum 24 B. repens 27 B. reptans 36, 42, 43, 44 B. restrepia 30 B. retusiusculum 43 B. rolfei 42 B. salaccense 20 B. saltatorium 43 B. sannio 10 B. saurocephalum 37, Pl. 22 B. scabratum 42 B. schinzianum var. phaeopogon 11 B. schistopetalum 37 B. senghasii 16 B. striatellum 32 B. striatum 22 B. subapetalum 24 B. subbullatum 25 B. taeter 22 B. tenuifolium 34 B. thouarsii 43

B. thwaitesii 41 B. tindemansianum 12 B. tortuosum 32 B. tothastes 23, 24 B. trias 40 B. triste 42, 43, 45 B. tuberculatum 41, 42 B. turpe 36 B. umbellatum 27, 42, 45 B. uniflorum 47 B. vaginatum 13, 42, 43, 45, 47 B. vietnamense 22 B. vinaceum 46 B. violaceolabellum 21 B. virescens 43 B. viridescens 22 B. viridiflorum 42, 43 B. weddellii 46 section Acrochaene 21, 22, 25 section Adelopetalum 19, 20, 23, 26 section Aeschynanthoides 19, 24, 25, 26 section Alcistachys 16 section Altisceptrum 20, 22, 23, 26 section Antennata 22, 26 section Balaenoidea 19, 21, 25, 26 section Beccariana 21, 22, 23, 25, 26, 47, 50 section Bifalcula 16 section Bifaria 11, 13–14 section Biflorae 21, 27 section Biseta 19, 20, 21, 27 section Blepharistes 20, 27, 47 section Brachyantha 21, 27 section Brachypus 24, 27, 47 section Brachystachyae 20, 22, 23, 27 section Bulbophyllaria 40, 41 section Bulbophyllum 16 section Carnosisepala 13, 14 section Chaseella 7, 14 section Cirrhopetaloides 21, 24, 25, 28, 50 section Cirrhopetalum 9, 14, 15, 21, 28 section Codonosiphon 23, 28 section Comata 9, 14 section Denticulata 11, 14 section Desmosanthes 20, 21, 23, 28 section Didactyle 40, 41, 47, 48 section Drymoda 25, 28 section Elasmatopus 15, 16 section Emarginatae 20, 29 section Epibulbon 24, 25, 29, 47 section Epicrianthes 23, 29, 47 section Eublepharon 19, 20, 21, 22, 29 section Furvescens 40, 41 section Genyorchis 7, 14 section Gilgiana 12, 14 section Gongorodes 22, 29 section Hemisterrantha 29 section Henosis 20, 30 section Hirtula 19, 21, 22, 24, 30 section Hoplandra 25, 30 section Hyalosema 23, 24, 30, 47 section Hymenobractea 30 section Ikongoense 16 section Imitatores 20, 21, 25, 30 section Intervallatae 19, 31 section Inversiflora 16 section Kinethrix 15, 16, 47 section Lemniscata 20, 23 section Leopardinae 19, 20, 21, 23, 24, 25, 31, 46

section Lepanthanthe 19, 20, 31 section Lepidorhiza 19, 20, 31, 50 section Lichenophylax 16 section Lupulina 9, 11, 14, 15, 16, 18 section Lyperocephalum 15, 18 section Macrocaulia 23, 32, 47 section Macrostelydia 21, 32 section Macrouris 19, 20, 21, 25, 32 section Megaclinium 12, 13, 15 section Micranthae 40, 41 section Minutissima 24, 25, 32, 46 section Monanthaparva 19, 23, 24, 25, 32 section Monanthes 25, 32, 47 section Monomeria 22, 33 section Monosepalum 23, 33 section Moratii 16, 18 section Napellii 40, 41 section Oreonastes 12, 13, 15 section Pachychlamys 16, 18 section Pantoblepharon 15, 16, 18 section Papulipetalum 24, 25, 33 section Pediochilus 23, 33 section Pelma 19, 24, 33 section Peltopus 24, 33 section Phreatiopsis 22, 34 section Piestobulbon 24, 34 section Planibulbus 21, 23, 34 section Ploiarium 15, 16, 18 section Plumata 21, 34 section Polymeres 24, 25, 34, 47 section Polyradices 15, 19 section Pseudopelma 22, 23, 25, 35 section Ptiloglossum 11, 12, 13, 15 section Racemosae 22, 35 section Recurvae 21, 25, 35 section Reptantia 23, 36 section Rhinanthera 20, 36 section Rhytionanthos 21, 36 section Saurocephalum 22, 37 section Schistopetalum 25, 37 section Sestochilus 19, 20, 24, 38, 50 section Stachysanthes 20, 22, 23, 25, 38 section Stenochilus 24, 39 section Sunipia 21, 25, 39 section Tapeinoglossum 25, 39 section Trias 25, 40 section Uncifera 19, 20, 22, 23, 40 section Xiphizusa 40, 41 Calymmanthera 125 146–7, 149, 150 C. major 149 C. tenuis 146 Calyptrochilum 341, 342, 353, 367–9, 421 C. christyanum 368, 369, Pl. 148 C. emarginatum 367, 369 Campylocentrum 340, 342, 362, 370–2, 383, 384 C. ariz-juliae 370 C. aromaticum 371 C. brenesii 370, Pl. 149 C. burchellii 371 C. constanzense 383, 384 C. filiforme 370, 384 C. helorrhizum 370 C. huebneri 371 C. macrocarpum 370, 384 C. micranthum 370, 371 C. ornithorrhynchum 370 C. schiedei 370

527

I nde x T o S cientific N ames

Cardiochilos 344, 372, 373, 374 C. williamsonii 372, 373, 374 Ceratocentron 117, 126, 138, 148, 150, 151, 152, 203, 268, 310, 315 C. fesselii 148, 151 Chamaeangis 338, 339, 341, 347, 385, 386, 388 Chamaeanthus 125, 150–2, 153, 154 C. brachystachys 150, 151, 153 C. wenzelii 151 Chauliodon 343, 373–4, 375, 376, 444 C. deflexicalcaratum 373–4, 375, 376 Chiloschista 114, 125, 126, 152–6, 174, 307 C. exuperei Pl. 63 C. godefroyana 152, 154 C. javanica 154, 155 C. lunifera 153, 154 C. parishii 154 C. phyllorhiza 154 C. ramifera 152 C. segawae 155 C. trudelii 152 C. usneoides 152, 154, 155 C. yunnanensis 155 Chroniochilus 117, 125, 142, 156–8, 188, 230, 297 C. minimus 158 C. thrixspermoides 158 C. tjidadapensis 156 C. virescens 157, 158 Cleisocentron 118, 124, 128, 129, 158–60, 167, 168, 278 C. abasii 160 C. gokusingii 160 C. kinabaluense 160 C. merrillianum 159, 160, Pl. 64 C. pallens 160 C. trichromum 158, 160 Cleisomeria 126, 128, 161, 162, 163, 165, 228, 284, 307 C. lanatum 161 C. pilosulum 161, 162, Pl. 65 Cleisostoma 112, 114, 118, 123, 124, 127, 128, 160, 161–6, 188, 211, 228, 259, 260, 266, 270, 278, 280, 284, 299 C. appendiculatum 163, 164 C. arietinum 163 C. brachystachys 118 C. discolor 164, 164 C. duplicilobum Pl. 66 C. filiforme 164 C. gjelleruptii 165 C. javanicum 163 C. paniculatum 112, 163 C. racemiferum 165 C. sagittatum 118, 161, 163, 165 C. simondii 163, Pl. 67 C. striatum 165 C. suaveolens 278 C. subulatum 163, 164 section Cleistostoma 163 section Complicatum 163 section Echioglossum 163 section Mitriformes 163 section Paniculatum 163 section Pilearia 163 section Subulatum 164 Cleisostomopsis 124, 166–8 C. eberhardtii 166, 166, 167, Pl. 68 C. elytrigera 167

528

Cooktownia 485 Cottonia 118, 124, 131, 168, 169, 170, 314 C. peduncularis 168, 169 Cribbia 338, 342, 343, 357, 374–6, 377, 378, 407 C. brachyceras 374, 376, 433 C. confusa 376, 433, Pl. 150 C. pendula 377 C. thomensis 377 Cryptopus 138, 342, 365, 376–8, 379, 380, 393, 401, 418, 419 C. elatus 376, 378, 379 C. paniculatus 378, Pl. 151 Cryptopylos 117, 126, 148, 168–70, 171, 172, 184, 203, 209, 268, 310, 315 C. clausus 168, 169, 170, 171, Pl. 69 Cyanaeorchis 479–80 C. arundinae 479, 480 C. minor 479, 480 C. praetermissa 480 Cyrtorchis 340, 341, 344, 379–83, 396, 429 C. aphylla 380 C. arcuata Pl. 152 subspecies whitei 380 C. aschersonii 380 C. chailluana 380 C. crassifolia 381 C. glaucifolia 382 C. pratermissa 380 C. ringens 380, Pl. 153 section Cyrtorchis 381 section Homocollecticon 381 Danxiaorchis 485 Deceptor 129, 170–2, 173, 174 D. bidoupensis 170, 173 Dendrobieae 3–100, 4 Dendrobium 3–4, 4, 51–100 D. adae 95 D. aemulum 90 D. aff. bilobum Pl. 26 D. aff. monticola Pl. 40 D. aff. spurium 90 D. affine 95, 96 D. agrostophyllum 63 D. albosanguineum 90 D. amethystoglossum 73, 91, 99 D. amoenum 96 D. amplum 72, 73 D. anceps 74 D. angustifolium 73 D. angustipetalum Pl. 23 D. anosmum 75, 89 D. antennatum 72, 89 D. aphyllum 75, 83, 87, 88 D. archipelagense Pl. 24 D. barbatulum 68 D. bifalce 94 D. bigibbum 71, 85, 96, 98, Pl. 25 D. bilobum 63 D. bracteosum 74, 99 D. brevicaule 94, 97 D. brevicolle 54 D. brymerianum 89 D. bulbophylloides 55, 70, Pl. 27 D. cacuminis Pl. 28 D. callitrophyllum 90 D. canaliculatum 73, 95, 96, Pl. 29 D. capillipes 90

D. capituliflorum 74 D. cariniferum 74, 89 D. catenatum 80, 83, 88 D. chrysanthum 78, 83, 87 D. chryseum 80, 87, 88 D. chrysotoxum 78, 83, 87, 88, 89, 90 D. chrysotropis 73, 74 D. cinnabarinum 98 D. citrinum 95 D. comatum 73, 80 D. conostalix 64 D. crassinode 75 D. criniferum 65 D. crocatum 85 D. crumenatum 83, 91, 94, 95 D. crystallinum 83, 87, 88 D. cunninghamii 73, 74, 94, 112 D. cuthbertsonii 94, 95, 97 D. defuscescens 88 D. dekockii 94, 97 D. delacourii 89 D. ×delicatum 89 D. denneanum 80, 83, 87, 88, 89 D. densiflorum 78, 80, 83, 88, 90, Pl. 30 D. denudans 73 D. derryi 91 D. devonianum 96 D. djamuense Pl. 31 D. draconis 78 D. ellipsophyllum Pl. 32 D. eleutheroglossum 67 D. engae 96 D. falconeri 87, 88 D. falcorostum 94 D. farmeri 80, 83, 88 D. fellowsii 98 D. fimbriatum 78, 80, 83, 88, 89, 94 D. forbesii 97 D. formosum 67, 99, Pl. 33 D. furcatum 95 D. fytchianum Pl. 34 D. gibsonii 78 D. gouldii 96 D. gracilicaule 83, 89, Pl. 35 D. gratiosissimum 78, 83, 88 D. hasseltii 85, 91, 99, Pl. 36 D. herpetophytum 69, 90 D. hymenanthum 91 D. infundibulum 95 D. jenkensii 90 D. jerdonianum 91 D. johannis 96 D. jonesii 74, 95 D. kinabaluense 56 D. kingianum 83, 89, 94, 95, 97, 98 D. lacteum 90 D. lampongense 95 D. lawesii 64, 95, 97 D. lichenastrum 69, 89, Pl. 37 D. lindleyi 90, Pl. 38 D. linguiforme 72, 95 D. lobatum 62 D. loddigesii 78, 87 D. longicornu 83, 87, 88 D. luteocilium 95 D. macrostachyum 95 D. malbrownei 73 D. masarangense 94

Index To S cientific Names

D. melanostictum 62, 90 D. microglaphys 62, 90 D. minjemense Pl. 39 D. mirbelianum 95 D. moniliforme 51, 75, 87, 88, 89, 94, 96 D. monophyllum 70, 98 D. moschatum 79, 83, Pl. 41 D. munificum 74 D. mutabile 91 D. nitidissimum 67 D. nobile 75, 78, 83, 88, 96 D. ochreatum 80 D. officinale 88 D. pachyphyllum 74 D. pallens 73 D. pandaneti 95 D. parishii 75 D. patentilobum 57 D. pendulum 75 D. pinifolium Pl. 42 D. platygastrium 91 D. pleianthum 71, Pl. 43 D. plicatile 83, 87, 88 D. polyanthum 75 D. polysema 96, Pl. 44 D. pseudocalceolum 91 D. pseudoglomeratum 95, Pl. 45 D. rarum 95 D. reflexitepalum 91 D. reginanivus 91 D. revolutum 67 D. rigidum 74 D. rumphiae Pl. 46 D. salaccense 68, 73 D. sanderae 99 D. sanguinolentum 91 D. sarcochilus 70 D. schneiderae Pl. 47 D. schoeninum 89 D. ×schumannianum 96 D. secundum 71, 89, 91, 95, 97 D. senile 74, 90, 91 D. sinense 89, 95 D. smillieae 95 D. solomonense 96 D. spatella Pl. 48 D. speciosum 89, 94, 95, 96, 97, 98 D. spectabile 69, 89 D. spectatissimum 58 D. spurium 68, Pl. 49 D. squamiferum Pl. 50 D. striolatum 94 D. ×suffusum 89 D. sulcatum 90 D. ×superbiens 83, 85, 97 D. teretifolium 95, 98 D. tetragonum 66, 97, 98 D. thyrsiflorum 78, 79, 80, 88 D. torresae 74 D. treacherianum 89 D. triflorum 83 D. trigonopus 78, 83, 87, 88, 89, 91 D. umbellatum 63 D. unicum 89, 95 D. vannouhuysii 97 D. venustum 94 D. verruciferum 59, 94, Pl. 51 D. vexillarius 97

D. virgineum 89 D. viridiflorum 63, 94 D. vonroemeri Pl. 52 D. wardianum 75 D. williamsianum 85 D. williamsonii 89 D. winikaphyllum 73, 94 D. womersleyi 95 D. xantholeucum 83, Pl. 53 D. ×yengliense 96 section Amblyanthus 61, 62, 89, 90, 91, 96, 97 section Aporum 58, 59, 60, 62–3, 74, 89, 90, 91, 96, 97, 98 section Biloba 62, 63, 89, 90, 91, 96, 97 section Brevisaccata 61, 62, 63, 89, 91, 96, 97 section Cadetia 56, 63–4, 73, 74, 89, 90, 91, 94, 96, 97, 99 section Calyptrochilus 57, 61, 62, 64, 89, 90, 91, 94, 95, 96, 97, 99 section Conostalix 59, 60, 64–5, 89, 90, 91, 96, 97, 99 section Crinifera 55, 56, 65, 89, 90, 91, 96, 97, 99 section Dendrobium 55, 58, 60, 61, 65–6, 89, 90, 91, 95, 96, 97, 99 section Dendrocoryne 59, 60, 66–7, 89, 90, 91, 94, 95, 96, 97, 98, 99 section Diplocaulobium 56, 67, 74, 89, 90, 91, 96, 97, 99 section Distichophyllae 61, 67, 89, 90, 91, 96, 97, 99 section Eleutheroglossum 61, 67, 89, 90, 91, 96, 98 section Formosae 60, 67–8, 89, 90, 91, 95, 96, 97, 99 section Fugacia 55, 58, 60, 68, 89, 90, 91, 96, 97 section Fytchianthe 61, 68, 89, 91, 96, 97 section Grastidium 53, 60, 62, 68–9, 89, 90, 91, 95, 96, 97, 99 section Herpethophytum 53, 62, 69, 89, 90, 91, 96, 97 section Latouria 55, 58, 59, 60, 69, 89, 91, 96, 97 section Lichenastrum 56, 69–70, 89, 90, 91, 95, 98 section Macrocladium 58, 62, 70, 89, 90, 91, 94, 96, 97, 98, 99 section Microphytanthe 57, 70, 89, 90, 91, 96, 97 section Monophyllaea 57, 58, 70–1, 89, 90, 91, 95, 96, 98 section Pedilonum 59, 61, 71, 89, 90, 91, 95, 96, 97, 99 section Phalaenanthe 61, 71, 89, 90, 91, 95, 96, 97, 98 section Pleianthe 61, 71–2, 89, 91, 96, 97 section Rhizobium 55, 57, 72, 74, 89, 90, 91, 94, 95, 96, 97, 98 section Sarcopodium 56, 57, 72, 89, 91, 96, 97, 98, 99 section Spatulata 61, 72–3, 89, 90, 91, 95, 96, 97, 98 section Stachyobium 61, 73, 89, 90, 91, 94, 96, 97 Dendrophylax 340, 342, 362, 370, 371, 383–5 D. fawcettii 385 D. funalis 383, 384, 385, Pl. 154 D. lindenii 383, 384, Pl. 155 D. macrocarpus 384 D. monteverdi 383 D. porrectus 383, 384, 384 D. varius 383 Devogelia 485 Diaphananthe 338, 339, 341, 344, 357, 385–9, 396, 407, 433

D. bidens 385, 388 D. fragrantissima 385 D. lorifolia 385 D. millarii 338, 407 D. odoratissima 386, 388 D. pellucida 385, 387, 388, Pl. 156 D. vesicata 386 Didymoplexiopsis 485 Dimorphorchis 114, 125, 154, 172–5, 176, 252, 307 D. beccarii 114, 174 D. breviscapa 174 D. graciliscapa 175 D. lowii 172, 174, Pl. 70 D. rossii 114, 174 Dinklageella 344, 389–90, 391 D. liberica 389, 390 D. minor 389 Diplocentrum 118, 126, 175–7, 178, 272 D. congestum 176, 177 D. recurvum 175, Pl. 71 Diploprora 118, 124, 176, 177–80, 272 D. championii 177, 178, 179, Pl. 72 D. truncata 178 Disa 479 Distylodon 343, 390–1, 392, 393 D. comptum 390–1, 392, 393 Dryadorchis 116, 125, 180–2, 195, 256 D. barbellata 180 D. huliorum 181 D. dasystele Pl. 73 Drymoanthus 116, 124, 127, 182–3, 184, 214, 232 D. adversus 112, 182, 183 D. minimus 182 D. minutus 182, Pl. 74 Dyakia 117, 126, 127, 148, 170, 183–6, 209, 268, 310, 315 D. hendersoniana 113, 183, 184, 185, Pl. 75 Earina 101, 102, 331, 335–8 E. autumnalis 335, 336, 337 E. deplanchei 336, 337 E. floripectin 337 E. fouldenensis 73 E. mucronata 335, 337 E. valida 337, Pl. 133 Eclecticus 118, 126, 186–8 E. chungii 118, 186, 187 Eggelingia 341, 344, 391, 392, 393, 394 E. clavata 391, 394 E. ligulifolia 391, Pl. 157 Eltroplectris 477, 478 Epigeneium 72, 89 Erasanthe 342, 393, 395, 396, 397 E. henrici 393, 395, 396, 397, Pl. 158 Eria 472 Ericksonella 485 Eulophia 479–80 Eurychone 340, 341, 344, 395–7, 398, 399 E. galeandrae 397, 398 E. rothschildiana 395, 396, 397, Pl. 159 Fernandezia 482–3 F. subbiflora 482 Galeandra 480 Galearis 469 Galeoglossum 463–6 G. cactorum 463, 465, 466

529

I nde x T o S cientific N ames

Galeoglossum (cont.) G. prescottioides 463 G. thysanochilum 463, 464, 465, 466 G. tubulosum 463, 465, 466, Pl. 191 Gastrochilus 112, 118, 119, 124, 127, 128, 188–91, 246 G. acutifolius 189 G. calceolaris 188 G. dasypogon 189 G. distichus 189 G. formosanus 112 G. gongshanensis 190 G. inconspicuus 124, 189, 191 G. japonicus 189 G. patinatus 189 G. pseudodistichus 189 G. retrocallus 112, 124, 190, 190, Pl. 76 G. sororius Pl. 77 Gongora 485 Grobya 480 Grosourdya 115, 116, 126, 127, 142, 191–4, 297 G. appendiculata 192, 193 G. bicornuta 194 G. elegans 191 G. emarginata 193 G. incurvicalcar 194 G. muscosa Pl. 78 G. quinquelobata 194 G. tripercus 194 Gunnarella 116, 117, 125, 147, 180, 194–5, 196, 197, 256 G. begaudii 195 G. carinata 194 G. robertsii 116, 195, 196 Gymnadenia 469 Harrisella 383 H. porrecta 370 Hederorkis 102, 104, 451–3 H. scandens 451, 452 Hemipiliopsis 466–8 H. purpureopunctata 466–8, 467, Pl. 193, Pl. 194 Holcoglossum 114, 117, 124, 127, 195, 197–201, 207, 262 H. amesianum 195, 198 H. calcicola 197 H. flavescens 195, 198, Pl. 79 H. himalaicum 198 H. kimballianum 195, 199 H. lingulatum 195 H. nagalandense 198, 200 H. nujiagense 197, 198 H. quasipinifolium 113, 195 H. rupestre 197, 198 H. sinicum 197, 198 H. subulifolium 197 H. wangii 197 H. weixiense 197, 198 Hymenorchis 117, 126, 127, 138, 201–3, 268, 315 H. javanica 201 H. nannodes 202 H. saccata 202 H. tanii Pl. 80 Jejewoodia 118, 124, 127, 132, 203–5, 212 J. crockerensis 204 J. jiewhoei 203, 204, 204 J. jongirii 204 J. linusii 204

530

J. longicalcarata 204 J. rimauensis 204 Jumellea 340, 342, 393, 397–9, 400, 401 J. arachnantha 397, 398 J. arborescens 397, 398 J. brevifolia 399 J. confusa 397, 398 J. densifoliata 399 J. flavescens 397, 398 J. fragrans 397, 399 J. gregariiflora 399 J. ibityana 399 J. lignosa 399 J. pandurata 399 J. papagensis 399 J. phalaenophora 397, 398 J. rigida 399 J. rossii Pl. 160 J. spathulata 399 J. teretifolia J. walleri 397, 400 Lemurella 340, 342, 365, 399, 401–2, 403, 404 L. culicifera 401, 402, Pl. 161 L. pallidiflora 401 Lemurorchis 340, 34, 342, 402–4, 405 L. madagascariensis 402–4, 404, 405, Pl. 162 Listrostachys 340, 341, 342, 404–5, 406, 407, 425, 429 L. pertusa 404–5, 406, 407, 447, Pl. 163 Luisia 112, 114, 117, 124, 197, 205–8, 226, 262 L. brachstachys 206 L. curtisii 206, 207 L. filiformis 112 L. morsei Pl. 81 L. tenuifolia 206 L. teretifolia 205, 206 L. trichorrhiza 207 L. tristis 207, Pl. 82 L. volucris 207 Macropodanthus 117, 126, 127, 138, 148, 170, 184, 203, 208–10, 252, 268, 297, 310, 315 M. philippinensis 208, 209 M. rimauensis 209 M. sabahensis 208, Pl. 83 M. teysmannii 209 Margelliantha 343, 357, 405–7, 408, 409 M. caffra 407 M. leedalii 408, Pl. 164 M. millarii 407 Mesospinidium 480, 481 Microcoelia 338, 340, 343, 355, 367, 374, 407, 409–11, 436 M. aphylla 409, 437 M. bulbocalcarata 409 M. caespitosa 409 M. corallina 409 M. cornuta 409 M. elliotii 410 M. exilis 407, 409 M. gilpinae 410 M. globulosa 409, 410 M. koehleri 410, 410, Pl. 165 M. macrantha 409 M. macrorrhynchia 411 M. megalorrhiza 409 M. obovata 409 M. perrieri 409, Pl. 166

M. physophora 409 M. smithii 409 M. stolzii 409, 410 subgenus Encheiridion 411 subgenus Microcoelia 411 Micropera 112, 118, 128, 210–12, 295, 299, 317 M. callosa 210 M. fasciculata 211 M. pallida 210, 211, Pl. 84 M. philippinensis 211 M. sterrophylla 212 Microsaccus 118, 127, 132, 204, 212, 213, 214, 307 M. ampullaceus 213 M. griffithii 213 M. javensis 212, 213 M. sumatranus 213 M. truncatus Pl. 85 Microterangis 339, 347, 348, 403 M. hariotiana 339, 340, 348 M. hildebrandtii 339, 340 Mobilabium 116, 125, 182, 212–4, 215, 216, 232 M. hamatum 212, 215 Mystacidium 340, 343, 357, 358, 407, 411–13 M. braybonae 411, 413 M. capense 411, 413 M. flanaganii 362, 411 M. gracile 412 M. tanganyikense 413 M. venosum 412, Pl. 167 Neobathiea 340, 343, 365, 401, 413–15 N. grandidieriana 413, 415, Pl. 168 N. hirtula 415 N. keraudrenae 415 N. perrieri 413, 414, 415 N. spathulata 415 Neobenthamia gracilis 451 Neolindleya 469–71 N. camtschatica 469–71, 470, Pl. 195, Pl. 196 Nephrangis 341, 343, 344, 415–17 N. bertauxinana 416 N. filiformis 415, 416, Pl. 169 Nothostele 476–8 N. acianthiformis 476, 477, Pl. 192 Notylia 484, 485 Notyliopsis 483–5 N. amadorinconiana 484 N. beatricis 483 Oeonia 340, 342, 343, 365, 393, 401, 417–19, 436 O. aubertii 417 O. madagascariensis 342, 343 O. rosea 378, 417, 418, Pl. 170 O. volucris 418 Oeoniella 340, 341, 342, 419–21 O. aphrodite 419–20, 420 O. polystachys 419, Pl. 171 Omoea 118, 128, 214–16, 217, 218 O. micrantha 214, 215, 217 O. philippinensis 214, 215, 241 Ophioglossella 125, 216–18, 219, 220 O. chrysostoma 216–18, 219, 220, Pl. 86 Ossiculum 342, 421–3 O. aurantiacum 421–3, 422 Pachyphyllum 483 Papilionanthe 117, 119, 124, 197, 207, 218–20, 221, 222, 226, 262

Index To S cientific Names

P. hookeriana 219, 220 P. teres 218, 219, 220, 221, Pl. 87 P. uniflora 219 P. vandarum 219 Papillilabium 116, 126, 220–1, 223–4 P. beckleri 220–1, 223–4, 223 Paraphalaenopsis 117, 122, 124, 207, 224–6, 262 P. denevei 224, 225 P. labukensis 226, Pl. 88 P. laycockii 225, 226 P. serpentilingua 225, 226 Pelatantheria 118, 128, 161, 165, 226–9, 260, 284 P. ctenoglossum 226, 227, 228 P. insectifera 228 P. rivesii Pl. 89 Pennilabium 117, 126, 128, 142, 157, 229–31, 268, 297 P. angraecum 229 P. kidmancoxii 230 P. struthio 229, 230, Pl. 90 Peristeranthus 116, 129, 231–3 P. hillii 231–3, 231 Phalaenopsis 112, 113, 114, 119, 125, 126, 127, 188, 233–41 P. amabilis 112, 233, 234, 236, 238, 239, 240, Pl. 91 subspecies rosenstromii 240 P. amboinensis Pl. 92 P. aphrodite 240 P. appendiculata Pl. 93 P. bellina 240, Pl. 94 P. buyssoniana 240 P. cornu-cervi 233, 237, 239, 240 P. deliciosa 233, 235, 236 P. difformis 233 P. equestris 237, 238, 239 P. esmeralda 233 P. floresensis Pl. 95 P. fuscata 240 P. gigantea 240 P. japonica 235, 239 P. hainanensis 235 P. ×intermedia 237 P. ×leucorhoda 237 P. lowii 240 P. lueddemanniana 236, 238 P. mannii 239 P. marriottiana 233, 236, 238 P. pantherina 240 P. parishii 233 P. pulcherrima 113, 236, 237, 239, 240, Pl. 96 P. regnieriana 240 P. sanderiana 237 P. schilleriana 237, 240 P. stobartiana 233, 235, 236 P. sumatrana 240 P. taenialis 239, 240 P. tetraspis 240 P. violacea 239, 240 P. wilsonii 235 P. zheijangensis 237 subgenus Hygrochilus 233 subgenus Ornithochilus 127, 233 subgenus Parishianae 233 section Aphyllae 233, 240 section Deliciosae 233 section Esmeralda 233, 240 section Parishianae 233, 240

subgenus Phalaenopsis 233, 240 section Phalaenopsis 233 section Polychilos 233 Pheladenia 485 Phragmorchis 124, 241–2, 243 P. teretifolia 241–2, 242, 243 Platanthera 469 Plectorrhiza 128, 242–5 P. brevilabris 243, 244 P. erecta 243, 244 P. tridentata 242, 243, 244, Pl. 97 Plectrelminthus 343, 423–4, 425 P. caudatus 423–4, 424, 425, Pl. 172 Podangis 340, 341, 344, 424–7 P. dactyloceras 424, 425, 426, Pl. 173 P. rhipsalisocia 425, Pl. 174 Podochilus 472 Polystachya 101, 102, 104, 451, 453–61 P. affinis 458 P. anceps 460 P. bella 460, 461 P. bicalcarata 458 P. bicarinata 460 P. caespitifica 460 P. calluniflora 458 P. campyloglossa 460 P. clareae Pl. 185 P. concreta 453, 459, 460, 461, Pl. 186, Pl. 187 P. cooperi 454 P. cultriformis 458, 460 P. dendrobiiflora 458 P. dolichophylla 460 P. elastica 458 P. estrellensis 460 P. eurychila 458 P. fallax 460 P. fusiformis 461 P. galeata 458, 461 P. goetzeana Pl. 188 P. golungensis 460, 461 P. johnstonii 460 P. laxiflora 458, 460 P. longiscapa 459 P. luteola 458 P. mazumbaiensis 460 P. meyeri 455 P. microbambusa 458 P. mildbraedii 460 P. modesta 459 P. mystacioides 458 P. neobenthamia 451, 456, 459, 460 P. ottoniana 458 P. paniculata 460 P. pobeguinii 460 P. porphyrochila Pl. 189 P. pubescens 459, 460, 461 P. praecipitis 457 P. rosea 458, 460 P. songaniensis 460 P. superposita 458 P. supfiana 460 P. tenuissima 460 P. tsaratananae Pl. 190 P. zambesiaca 460 section Affines 458, 459 section Aporoideae 458, 459 section Bicalcaratae 458 section Calluniflorae 458, 459

section Caulescentes 458, 459 section Cultriformes 458, 459 section Dendrobianthe 458, 459, 460 section Dimorphocaules 458, 459 section Elasticae 458, 459 section Eurychilae 458, 459 section Galeatae 458, 459 section Humiles 458, 459 section Isochiloides 458, 459, 460 section Polystachya 458, 459 section Pseudomonophyllae 458 section Superpositae 458, 459 Polystachyinae 101, 104, 450–61 Pomatocalpa 118, 127, 128, 245–7 P. kunstleri 246 P. macphersonii 246 P. maculosum Pl. 98 P. marsupiale 246, Pl. 99 P. spicatum 245 Porrorhachis 117, 128, 138, 203, 247–9, 297 P. galbina 247, 248, Pl. 100, 310 P. macrosepala 247 Prescottia tubulosa 465 Pseuderia 471–3 P. floribunda Pl. 197 P. foliosa 471 P. similis 471, 472 P. smithiana 472 Pseudocranichis thysanochila 465 Psychopsiella 473–5 P. limminghei 473–5 Psychopsis 474 Pteroceras 114, 116, 117, 126, 127, 139, 146, 158, 170, 210, 230, 249–52, 297 P. longicalcarum 251 P. pallidum 116, 251, 252 P. radicans 249 P. semiteretifolium 116, 252 P. teres 117, 250, 251, Pl. 101 Rangaeris 340, 341, 343, 427–9 R. amaniensis 349, 427, 428, 429, 447 R. longicaudata 427 R. muscicola 427, 428, 429 R. rhipsalisocia 425, 429 R. schliebenii 427 Raycadenco 483 Renanthera 114, 118, 119, 129, 136, 141, 252–3, 254 R. bella 253, Pl. 102 R. citrina 253 R. coccinea 252 R. edenfeldtii 252 R. imschootiana 252 R. matutina 113 R. moluccana 253 R. monachica 252 Rhaesteria 341, 343, 429–31 R. eggelingii 429–31, 430, Pl. 175 Rhinerrhiza 116, 126, 253–6 R. divitiflora 253–6, 255, Pl. 103 Rhinerrhizopsis 116, 126, 180, 195, 256–8 R. moorei 256, 257, 258, Pl. 104 Rhipidoglossum 338, 339, 343, 357, 376, 388, 407, 431–3 R. bilobatum 431 R. curvatum 431 R. kamerunense 376, 431, 433 R. obanense 431

531

I nde x T o S cientific N ames

Rhipidoglossum (cont.) R. pulchellum 431, 433 R. rutilum 431, 433 R. stolzii 432, Pl. 176 R. subsimplex 431 R. xanthopollinium 431, 433 Rhynchogyna 118, 124, 161, 165, 188, 228, 258–60, 284, 299 R. fallax 259 R. luisifolia 258, 258, 259 Rhychostylis 112, 117, 129, 197, 260–3, 288 R. coelestis 262, Pl. 105 R. gigantea 262 R. retusa 112, 113, 260, 261, 262 Robiquetia 118, 127, 129, 263–4, 265, 266, 297 R. angustifolia Pl. 106 R. ascendens 263 R. bifida 264 R. gracilistipes 263 R. penangiana 263 R. rosea Pl. 107 R. spathulata 263 R. succisa 263 R. sylvestris 263 R. wassellii 263 R. witteana Pl. 108 Saccolabiopsis 118, 127, 264–6, 267, 268 S. armitii 266 S. bakhuisenii 264 S. pusilla 267 Saccolabium 112, 128, 266–9, 270 S. pusillum 266 S. sigmoideum 269 Santanderella 484 Santotomasia 129, 269–71, 272 S. wardiana 269–71, 271, 272 Sarcanthopsis 118, 124, 176, 178, 271–3 S. nagarensis 271, 273 S. warocqueana 272 Sarcochilus 114, 116, 125, 126, 128, 129, 254, 274–8, 297, 307 S. australis 275 S. ceciliae 275, 277 S. dilatatus 275, Pl. 109 S. falcatus 113, 274, 275, 277 S. fitzgeraldii 275, 276, 277 S. hartmannii 274, 275, 277 S. hillii 275 S. hirticalcar 277 S. minutiflos 275, 277 S. olivaceus 277 S. spathulatus 276, 276, 277 S. weinthallii 276 section Monanthochilus 274, 277 Sarcoglyphis 118, 126, 160, 165, 228, 260, 278, 279, 280, 299 S. masiusii 278 S. mirabilis 278 S. potamophila 279 S. smithianus Pl. 110 Sarcophyton 118, 128, 165, 228, 260, 278–80, 281, 282 S. crassifolium 278, 281 S. pachyphyllum Pl. 111 Schistotylus 116, 127, 280–2, 283, 284 S. purpuratus 280–2, 283, 284 Schizodium 479 Schoenorchis 118, 124, 127, 129, 228, 241, 282–6

532

S. buddleiflora 285 S. gemmata 284, Pl. 112 S. juncifolia 282, 284, 286 S. micrantha 284, 322 S. sarcophylla 284 Schuitemania 485 Seidenfadenia 117, 124, 197, 262, 286–8 S. mitrata 286–8, 287, Pl. 113 Seidenfadeniella 124, 288–90 S. filiformis 289, 289 S. rosea 288 Singchia 128, 129, 290–2 S. malipoensis 290–2, 291 Sirhookera 101, 102, 104, 109–11 S. lanceolata 109, 110, Pl. 56 S. latifolia 109 Sirindhornia 485 Smithsonia 124, 127, 292–5 S. maculata 293, 293, 294 S. saldanhae 295, Pl. 114 S. straminea 292, 293, 295 S. viridiflora 293, 294 Smitinandia 118, 128, 165, 172, 188, 211, 295, 296, 297, 317 S. helferi 295 S. micrantha 113, 295, 296, Pl. 115 Sobennikoffia 340, 342, 433–5 S. humbertiana 435, Pl. 177 S. robusta 433, 434, 435 Solenangis 338, 340, 344, 389, 435–8 S. aphylla 338, 362, 409, 436, 437 S. clavata 344, 436 S. conica 436 S. cornuta 338, 409, 436 S. scandens 435, 436 S. wakefieldii 436, Pl. 178 Sphyrarhynchus 341, 343, 344, 357, 438–40 S. schliebenii 438–40, 439, Pl. 179 Spongiola 117, 126, 128, 142, 157, 230, 295–7, 298, 299 S. lohokii 295–7, 298, 299 Stereochilus 118, 127, 128, 160, 165, 211, 228, 260, 278, 295, 297–9, 300, 301, 317 S. brevirachis 300 S. dalatensis Pl. 116 S. hirtus 297 Summerhayesia 344, 440–2 S. laurentii 440, 442, Pl. 180 S. zambesiaca 440, 441, 442 Taeniophyllum 111, 112, 116, 125, 127, 299–304, 431 T. alwisii Pl. 117 T. biocellatum 301 T. coxii 102, 302 T. fasciola 301, 302, 303, 304 T. hasseltii 304 T. iboetii 301 T. lobatum 303 T. montanum Pl. 118 T. muelleri 303 T. obtusum 299 T. pusillum 112 T. smithii 301 T. wilkianum 303 Taeniorrhiza 340, 341, 343, 442–4 T. gabonensis 442–4, 443 Taprobanea 129, 304–5, 306, 307, 325

T. spathulata 304–5, 306, 307, 325 Thrixspermum 114, 124, 125, 126, 128, 154, 174, 188, 305–9 T. amplexicaule 307 T. angustifolium 242 T. arachnites 307 T. calceolus 307 T. centipeda 305, 307 T. hystrix 308 T. japonicum 307 T. pinocchio Pl. 119 T. saruwatarii 112 T. tortum 308 section Dendrocalla 307 section Katocalla 308 section Thrixspermum 307 Trachoma 117, 126, 128, 138, 185, 203, 248, 309–11 T. rhopalorrhachis 309, 310 T. speciosum Pl. 120 T. stellatum 310 Triceratorhynchus 343, 444–5 T. viridiflorus 444–5, 445, Pl. 181 Trichoglottis 118, 119, 124, 127, 128, 129, 131, 311–15, 327 T. atropurpurea 313 T. biglandulosa 127, 312 T. fasciata 314 T. kinabaluensis Pl. 121 T. philippinensis 314 T. retusa 311 T. smithii 313, Pl. 122 T. tenera Pl. 123 Trichopilia 474 Tridactyle 340, 341, 344, 396, 405, 417, 425, 429, 445–8, 450 T. bicaudata 445, 447, Pl. 182 T. crassifolia 447 T. filifolia 447 T. filiformis 448 T. flabellata 446 T. furcistipes 447, 450 T. gentilii 450 T. scottellii 447 T. tanneri 447, Pl. 183 T. tridactylites 447 T. tridentata 447, 448 T. virginea 446 subgenus Nephrangis 448 subgenus Tridactyle 448 Tuberolabium 114, 117, 126, 128, 138, 148, 185, 203, 252, 268, 270, 297, 310, 315, 316, 317 T. kotoense 315 T. odoratissimum 316, Pl. 124 Uncifera 118, 128, 129, 211, 295, 299, 315–17, 318, 319 U. acuminata 315, 318 U. thailandica Pl. 125 Vanda 112, 114, 117, 119, 120, 125, 126, 127, 128, 129, 184, 197, 207, 262, 292, 305, 307, 317–27 V. ampullacea 326 V. aurantiaca 326 V. aurea 326 V. christensoniana 321, 326 V. coerulea 323, 324, 326

Index To S cientific Names

V. coerulescens 322, 323, 324, 325 V. cristata 112, 325, 326 V. curvifolia 326 V. denisoniana 325, 326 V. dives 322, Pl. 126 V. falcata 325, 326, Pl. 127 V. flabellata 117 V. garay 326, Pl. 128 V. helvola 325 V. himalaica 326 V. hindsii 325 V. insularum 326 V. javierae 326 V. jennae 320 V. lamellata 326 V. lilacina 322

V. luzonica 325 V. merrillii 325 V. miniata 326 V. nana 323, 326 V. richardsiana 326 V. rubra 326 V. sanderiana 326 V. semiteretifolia 326 V. stangeana 322 V. tessellata 112, 113, 317, 325, 326, Pl. 129 V. testacea 325, Pl. 130 V. tricolor 112, 113, 325 var. suavis 322, 323, 325, 326 V. ustii 323 V. vietnamica 117 V. xichangensis 326

Vandeae 101–461 Vandopsis 112, 118, 125, 327–8, 329 V. gigantea 118, 327, 328, Pl. 131 V. lissochiloides 327 V. undulata 118 Xenikophyton 127, 129, 328–31 X. seidenfadenianum 329 X. smeenanum 328, 329, 330 Ypsilopus 340, 344, 425, 429, 447, 448–50 Y. erectus 449, 450 Y. longifolius 447, 448, 450, Pl. 184 Y. viridiflorus 447, 448, 450 Zeuxinella 485

533

SUBJECT INDEX Page numbers in boldface refer to relevant illustrations. Anatomy of Acampe 129–31 of Adrorhizon 106 of Aerangis 345 of Aeranthes 349–51 of Aeridinae 112–3 of Agrostophyllum 332 of Ambrella 352 of Amesiella 137 of Ancistrorhynchus 353–5 of Angraecinae 338 of Angraecopsis 357 of Angraecum 361 of Beclardia 364–5 of Bolusiella 366–7 of Brassia 481 of Bromheadia 108 of Bulbophyllum 41–3 of Calyptrochilum 369 of Campylocentrum 370 of Cardiochilos 372 of Chauliodon 374 of Chiloschista 153–4 of Cribbia 376 of Cryptopus 378 of Cyrtorchis 380 of Dendrobium 73–4 of Dendrophylax 383 of Diaphananthe 385 of Dinklageella 389 of Distylodon 391 of Earina 335–6 of Eggelingia 391 of Erasanthe 393 of Eurychone 395–6 of Fernandezia 482 of Galeoglossum 463–4 of Hederorkis 451 of Hemipiliopsis 466 of Jumellea 397–8 of Lemurella 401 of Lemurorchis 403 of Listrostachys 405 of Margelliantha 407 of Microcoelia 409 of Mystacidium 411, 413 of Neobathiea 413–4 of Neolindleya 469 of Nephrangis 415 of Nothostele 476 of Notyliopsis 484 of Oeonia 417 of Oeoniella 419

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Ossiculum 421 Phalaenopsis 234–6 Plectrelminthus 423 Podangis 425 Polystachya 459 Pseuderia 471 Psychopsiella 474 Rangaeris 427–8 Rhaesteria 429 Rhipidoglossum 431 Sirhookera 109 Sobennikoffia 435 Solenangis 436 Sphyrarhynchus 438 Summerhayesia 440, 442 Taeniophyllum 301–2 Taeniorrhiza 443 Triceratorhynchus 444 Trichoglottis 313 Tridactyle 447 Vanda 319–21 Ypsilopus 448, 450

cultivation of Acampe 132 of Adenoncos 132 of Aerides 137 of Aeridinae 119–23 of Amesiella 138 of Angraecinae 340–1 of Arachnis 141 of Biermannia 142 of Bogoria 144 of Brachypeza 146 of Brassia 482 of Bromheadia 109 of Bulbophyllum 50 of Calymmanthera 147 of Campylocentrum 371 of Ceratocentron 150 of Chamaeanthus 152 of Chiloschista 156 of Chroniochilus 158 of Cleisocentron 160 of Cleisomeria 161 of Cleisostoma 165 of Cleisostomopsis 167 of Cottonia 168 of Cryptopylos 170 of Deceptor 172 of Dendrobium 97–8 of Dendrophylax 385 of Dimorphorchis 175

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Diplocentrum 177 Diploprora 178 Dryadorchis 181 Drymoanthus 183 Dyakia 184–5 Eclecticus 187 Fernandezia 483 Galeoglossum 466 Gastrochilus 191 Grosourdya 194 Gunnarella 195 Holcoglossum 199 Hymenorchis 203 Jejewoodia 205 Luisia 207 Macropodanthus 209 Micropera 212 Microsaccus 212 Mobilabium 214 Nothostele 478 Notyliopsis 485 Omoea 216 Ophioglossella 217 Papilionanthe 220 Papillilabium 224 Paraphalaenopsis 226 Pelatantheria 228 Pennilabium 230 Peristeranthus 233 Phalaenopsis 240 Plectorrhiza 244 Polystachya 461 Pomatocalpa 247 Porrorhachis 248 Psychopsiella 475 Pteroceras 251–2 Renanthera 253 Rhinerrhiza 255 Rhinerrhizopsis 258 Rhynchogyna 260 Rhychostylis 263 Robiquetia 264 Saccolabiopsis 266 Saccolabium 268 Sarcanthopsis 273 Sarcochilus 277 Sarcoglyphis 278 Sarcophyton 280 Schistotylus 282 Schoenorchis 285 Seidenfadenia 288 Seidenfadeniella 290 Singchia 292

535

SUBJECT INDEX

cultivation (cont.) of Smithsonia 294 of Smitinandia 295 of Spongiola 297 of Stereochilus 299 of Taeniophyllum 304 of Taprobanea 305 of Thrixspermum 309 of Trachoma 311 of Trichoglottis 314 of Tuberolabium 315 of Uncifera 317 of Vanda 326 of Vandopsis 327 of Xenikophyton 330 cytogenetics of Acampe 131 of Adenoncos 132 of Adrorhizon 106 of Aerangis 345 of Aeranthes 351 of Aerides 134 of Agrostophyllum 333 of Ambrella 352 of Ancistrorhynchus 355 of Angraecopsis 357 of Angraecum 362 of Arachnis 140 of Beclardia 365 of Biermannia 142 of Bolusiella 367 of Brachypeza 145 of Brassia 481 of Bromheadia 108 of Bulbophyllum 43 of Calymmanthera 147 of Calyptrochilum 369 of Campylocentrum 370 of Cardiochilos 372 of Ceratocentron 148 of Chamaeanthus 151 of Chauliodon 374 of Chiloschista 154 of Chroniochilus 157 of Cleisocentron 159 of Cleisomeria 161 of Cleisostoma 164 of Cleisostomopsis 167 of Cottonia 168 of Cribbia 376 of Cryptopus 378 of Cryptopylos 170 of Cyrtorchis 380 of Deceptor 172 of Dendrobium 74 of Dendrophylax 383 of Diaphananthe 386, 388 of Dimorphorchis 174 of Dinklageella 389 of Diplocentrum 176 of Diploprora 178 of Distylodon 391 of Dryadorchis 180 of Drymoanthus 182 of Dyakia 184 of Earina 336 of Eclecticus 186 of Eggelingia 393

536

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Erasanthe 393 Eurychone 396 Fernandezia 483 Galeoglossum 464 Gastrochilus 189 Grosourdya 193 Gunnarella 195 Hederorkis 452 Hemipiliopsis 468 Holcoglossum 195, 197 Hymenorchis 201 Jejewoodia 203 Jumellea 398 Lemurella 401 Lemurorchis 403 Listrostachys 405 Luisia 206 Macropodanthus 209 Margelliantha 407 Microcoelia 409 Micropera 211 Microsaccus 212 Mobilabium 214 Mystacidium 413 Neobathiea 414 Neolindleya 469 Nephrangis 415 Nothostele 476 Notyliopsis 484 Oeonia 418 Oeoniella 419 Omoea 215 Ophioglossella 217 Ossiculum 421 Papilionanthe 218 Papillilabium 223 Paraphalaenopsis 225 Pelatantheria 228 Pennilabium 230 Peristeranthus 232 Phalaenopsis 236 Phragmorchis 241 Plectorrhiza 243 Plectrelminthus 423 Podangis 426 Polystachya 460 Pomatocalpa 245 Porrorhachis 247 Pseuderia 471 Psychopsiella 474 Pteroceras 251 Rangaeris 428–9 Renanthera 252 Rhaesteria 429 Rhinerrhiza 254 Rhinerrhizopsis 256 Rhipidoglossum 433 Rhynchogyna 259 Rhychostylis 262 Robiquetia 263 Saccolabiopsis 266 Saccolabium 268 Santotomasia 270 Sarcanthopsis 272 Sarcochilus 274 Sarcoglyphis 278 Sarcophyton 280 Schistotylus 282

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Schoenorchis 284 Seidenfadenia 286 Seidenfadeniella 290 Singchia 291 Sirhookera 109 Smithsonia 294 Smitinandia 295 Sobennikoffia 435 Solenangis 436 Sphyrarhynchus 438 Spongiola 296 Stereochilus 298 Summerhayesia 442 Taeniophyllum 302 Taeniorrhiza 443 Taprobanea 305 Thrixspermum 307 Trachoma 309 Triceratorhynchus 444 Trichoglottis 314 Tridactyle 447 Tuberolabium 315 Uncifera 317 Vanda 322–3 Vandopsis 327 Xenikophyton 329 Ypsilopus 450

derivation of name of Acampe 129 of Adenoncos 132 of Adrorhizon 104 of Aerangis 345 of Aeranthes 349 of Aerides 134 of Agrostophyllum 332 of Ambrella 351 of Amesiella 137 of Ancistrorhynchus 353 of Angraecopsis 355 of Angraecum 358 of Arachnis 139 of Beclardia 363 of Biermannia 142 of Bogoria 142 of Bolusiella 365 of Brachypeza 144 of Bromheadia 106 of Bulbophyllum 6 of Calymmanthera 146 of Calyptrochilum 368 of Campylocentrum 370 of Cardiochilos 372 of Ceratocentron 148 of Chamaeanthus 150 of Chauliodon 373 of Chiloschista 152 of Chroniochilus 156 of Cleisocentron 158 of Cleisomeria 161 of Cleisostoma 162 of Cleisostomopsis 166 of Cottonia 168 of Cribbia 374 of Cryptopus 376 of Cryptopylos 168 of Cyanaeorchis 479 of Cyrtorchis 379

SUBJECT INDEX

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Deceptor 171 Dendrobium 53 Dendrophylax 383 Diaphananthe 385 Dimorphorchis 172–3 Dinklageella 389 Diplocentrum 176 Diploprora 177 Distylodon 390 Dryadorchis 180 Drymoanthus 182 Dyakia 183 Earina 335 Eclecticus 186 Eggelingia 391 Erasanthe 393 Eurychone 395 Fernandezia 482 Galeoglossum 463 Gastrochilus 188 Grosourdya 191 Gunnarella 194 Hederorkis 451 Hemipiliopsis 466 Holcoglossum 195 Hymenorchis 201 Jejewoodia 203 Jumellea 397 Lemurella 401 Lemurorchis 403 Listrostachys 404 Luisia 205 Macropodanthus 208 Margelliantha 406 Microcoelia 407 Micropera 210 Microsaccus 212 Mobilabium 212 Mystacidium 411 Neobathiea 413 Neolindleya 469 Nephrangis 415 Nothostele 476 Notyliopsis 483 Oeonia 417 Oeoniella 419 Omoea 214 Ophioglossella 216 Ossiculum 421 Papilionanthe 218 Papillilabium 220 Paraphalaenopsis 224 Pelatantheria 227 Pennilabium 229 Peristeranthus 231 Phalaenopsis 233 Phragmorchis 241 Plectorrhiza 243 Plectrelminthus 423 Podangis 425 Polystachya 453 Pomatocalpa 245 Porrorhachis 247 Pseuderia 471 Psychopsiella 473 Pteroceras 249 Rangaeris 427 Renanthera 252

of Rhaesteria 429 of Rhinerrhiza 254 of Rhinerrhizopsis 256 of Rhipidoglossum 431 of Rhynchogyna 258 of Rhychostylis 260 of Robiquetia 263 of Saccolabiopsis 264 of Saccolabium 267 of Santotomasia 269 of Sarcanthopsis 271 of Sarcochilus 274 of Sarcoglyphis 278 of Sarcophyton 278 of Schistotylus 281 of Schoenorchis 283 of Seidenfadenia 286 of Seidenfadeniella 288–9 of Singchia 290 of Sirhookera 109 of Smithsonia 292 of Smitinandia 295 of Sobennikoffia 433 of Solenangis 436 of Sphyrarhynchus 438 of Spongiola 295 of Stereochilus 298 of Summerhayesia 440 of Taeniophyllum 300 of Taeniorrhiza 442 of Taprobanea 304 of Thrixspermum 305 of Trachoma 309 of Triceratorhynchus 444 of Trichoglottis 311 of Tridactyle 445 of Tuberolabium 315 of Uncifera 315 of Vanda 319 of Vandopsis 327 of Xenikophyton 328 of Ypsilopus 448 description of Acampe 129, 130, Pl. 57 of Adenoncos 132, 133, Pl. 58 of Adrorhizinae 104 of Adrorhizon 104–5, 105, Pl. 54 of Aerangis 345, 346, 347, Pl. 134–6 of Aeranthes 349, 350, Pl. 137, Pl. 138 of Aerides 134, 135, Pl. 59 of Aeridinae 111 of Agrostophyllinae 331 of Agrostophyllum 332, 333, Pl. 132 of Ambrella 351–2, 352, Pl. 139 of Amesiella 137, 138, Pl. 60 of Ancistrorhynchus 353, 354, Pl. 140 of Angraecinae 338 of Angraecopsis 355, 356, Pl. 141 of Angraecum 358, 359, Pl. 142–5 of Arachnis 139–40, 140, Pl. 61 of Beclardia 363–4, 364, Pl. 146 of Biermannia 142, 143 of Bogoria 142, 145 of Bolusiella 365, 366, Pl. 147 of Brachypeza 144, 147, Pl. 62 of Brassia 480–1 of Bromheadia 106, 107, Pl. 55 of Bulbophyllum 6–7, 7, 8, 9, 10, 11, 12, 13, Pl. 1–22

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Calymmanthera 146, 149 Calyptrochilum 368–9, 368, Pl. 148 Campylocentrum 370, 371, Pl. 149 Cardiochilos 372, 373 Ceratocentron 148, 151 Chamaeanthus 150, 153 Chauliodon 374, 375 Chiloschista 152, 155, Pl. 63 Chroniochilus 156–7, 157 Cleisocentron 158–9, 159, Pl. 64 Cleisomeria 161, 162, Pl. 65 Cleisostoma 162–3, 164, Pl. 66, Pl. 67 Cleisostomopsis 166–7, 166, Pl. 68 Cottonia 168, 169 Cribbia 374–5, 377, Pl. 150 Cryptopus 377–8, 379, Pl. 151 Cryptopylos 169, 171, Pl. 69 Cyrtorchis 379–80, 381, Pl. 152, Pl. 153 Deceptor 171, 173 Dendrobieae 3 Dendrobium 53–4, 54, 55, 56, 57, 58, 59, Pl. 23–53 Dendrophylax 383, 384, Pl. 154, Pl. 155 Diaphananthe 385, 386, 387, Pl. 156 Dimorphorchis 173, 175, Pl. 70 Dinklageella 389, 390 Diplocentrum 176, 177, Pl. 71 Diploprora 177–8, 179, Pl. 72 Distylodon 390, 392 Dryadorchis 180, 181, Pl. 73 Drymoanthus 182, 183, Pl. 74 Dyakia 183–4, 185, Pl. 75 Earina 335, 336, Pl. 133 Eclecticus 186, 187 Eggelingia 391, 394, Pl. 157 Erasanthe 393, 396, Pl. 158 Eurychone 395, 398, Pl. 159 Fernandezia 482 Galeoglossum 463, 464, Pl. 191 Gastrochilus 188, 189, 190, Pl. 76, Pl. 77 Grosourdya 192, 192, 193, Pl. 78 Gunnarella 195, 196 Hederorkis 451, 452 Hemipiliopsis 466, 467, Pl. 193, Pl. 194 Holcoglossum 195, 198, 199, 200, Pl. 79 Hymenorchis 201, 202, Pl. 80 Jejewoodia 203, 204 Jumellea 397, 400, Pl. 160 Lemurella 401, 402, Pl. 161 Lemurorchis 403, 404, Pl. 162 Listrostachys 404–5, 406, Pl. 163 Luisia 205–6, 206, Pl. 81, Pl. 82 Macropodanthus 208–9, 208, Pl. 83 Margelliantha 406, 408, Pl. 164 Microcoelia 407, 409, 410, 437, Pl. 165, Pl. 166 Micropera 210–11, 210, Pl. 84 Microsaccus 212, 213, Pl. 85 Mobilabium 213, 215 Mystacidium 411, 412, Pl. 167 Neobathiea 413, 414, Pl. 168 Neolindleya 469, 470, Pl. 195, Pl. 196 Nephrangis 415, 416, Pl. 169 Nothostele 476, 477, Pl. 192 Notyliopsis 484 Oeonia 417, 418, Pl. 170 Oeoniella 419, 420, Pl. 171 Omoea 214–5, 217 Ophioglossella 216, 219, Pl. 86

537

SUBJECT INDEX

description (cont.) of Ossiculum 421, 422 of Papilionanthe 218, 221, Pl. 87 of Papillilabium 221, 223, 223 of Paraphalaenopsis 224–5, 225, Pl. 88 of Pelatantheria 227, 227, Pl. 89 of Pennilabium 229–30, 229, Pl. 90 of Peristeranthus 231–2, 231 of Phalaenopsis 233–4, 234, 235, 236, 237, Pl. 91–6 of Phragmorchis 241, 242 of Plectorrhiza 243, 244, Pl. 97 of Plectrelminthus 423, 424, Pl. 172 of Podangis 425, 426, Pl. 173 of Polystachya 453–5, 454, 455, 456, 457, Pl. 185–90 of Polystachyinae 450–1 of Pomatocalpa 245, 246, Pl. 98, Pl. 99 of Porrorhachis 247, 248, Pl. 100 of Pseuderia 471, 472, Pl. 197 of Psychopsiella 473–4 of Pteroceras 249–50, 250, Pl. 101 of Rangaeris 427, 428, Pl. 174 of Renanthera 252, 253, Pl. 102 of Rhaesteria 429, 430, Pl. 175 of Rhinerrhiza 254, 255, Pl. 103 of Rhinerrhizopsis 256, 257, Pl. 104 of Rhipidoglossum 431, 432, Pl. 176 of Rhynchogyna 258, 259 of Rhychostylis 260, 261, Pl. 105 of Robiquetia 263, 264, 265, Pl. 106–8 of Saccolabiopsis 265–6, 267 of Saccolabium 267–8, 269 of Santotomasia 270, 271 of Sarcanthopsis 271–2, 273 of Sarcochilus 274, 275, 276, Pl. 109 of Sarcoglyphis 278, 279, Pl. 110 of Sarcophyton 278–9, 281, Pl. 111 of Schistotylus 281–2, 283 of Schoenorchis 283–4, 285, Pl. 112 of Seidenfadenia 286, 287, Pl. 113 of Seidenfadeniella 289, 289 of Singchia 290–1, 291 of Sirhookera 109, 110, Pl. 56 of Smithsonia 293–4, 293, Pl. 114 of Smitinandia 295, 296, Pl. 115 of Sobennikoffia 433–4, 435, Pl. 177 of Solenangis 436, Pl. 178 of Sphyrarhynchus 438, 439, Pl. 179 of Spongiola 296, 298 of Stereochilus 298, 300, Pl. 116 of Summerhayesia 440, 441, Pl. 180 of Taeniophyllum 300–1, 302, 303, Pl. 117, Pl. 118 of Taeniorrhiza 442, 443 of Taprobanea 305, 306 of Thrixspermum 305–7, 308, Pl. 119 of Trachoma 309, 310, Pl. 120 of Triceratorhynchus 444, 445, Pl. 181 of Trichoglottis 311–12, 312, 313, Pl. 121–3 of Tridactyle 445–6, 446, Pl. 182, Pl. 183 of Tuberolabium 315, 316, Pl. 124 of Uncifera 316, 318, Pl. 125 of Vanda 319, 320, 321, 322, 323, Pl. 126–30 of Vandeae 101 of Vandopsis 327, 328, Pl. 131 of Xenikophyton 328–9, 330 of Ypsilopus 448, 449, Pl. 184

538

distribution of Acampe 129, 131 of Adenoncos 132, 134 of Adrorhizinae 104 of Adrorhizon 105–6, 106 of Aerangis 345, 348 of Aeranthes 349, 351 of Aerides 134, 136 of Aeridinae 111 of Agrostophyllinae 331 of Agrostophyllum 332, 334 of Ambrella 352, 353 of Amesiella 137, 139 of Ancistrorhynchus 353, 355 of Angraecinae 338 of Angraecopsis 355, 357 of Angraecum 358, 360 of Arachnis 140, 141 of Beclardia 364, 365 of Biermannia 142, 144 of Bogoria 142, 146 of Bolusiella 366, 367 of Brachypeza 144, 148 of Brassia 481, 481 of Bromheadia 107, 108 of Bulbophyllum 7, 42 of Calymmanthera 146, 150 of Calyptrochilum 369, 369 of Campylocentrum 370, 372 of Cardiochilos 372, 374 of Ceratocentron 148, 152 of Chamaeanthus 150, 154 of Chauliodon 374, 376 of Chiloschista 152–3, 156 of Chroniochilus 157, 158 of Cleisocentron 159, 160 of Cleisomeria 161, 163 of Cleisostoma 163, 165 of Cleisostomopsis 167, 167 of Cottonia 168, 170 of Cribbia 376, 378 of Cryptopus 378, 380 of Cryptopylos 169, 172 of Cyanaeorchis 479, 479 of Cyrtorchis 380, 383 of Deceptor 172, 174 of Dendrobieae 3, 4 of Dendrobium 55, 60 of Dendrophylax 383, 385 of Diaphananthe 385, 388 of Dimorphorchis 173–4, 176 of Dinklageella 389, 391 of Diplocentrum 176, 178 of Diploprora 178, 180 of Distylodon 391, 393 of Dryadorchis 180, 182 of Drymoanthus 182, 184 of Dyakia 184, 186 of Earina 335, 337 of Eclecticus 186, 188 of Eggelingia 391, 395 of Erasanthe 393, 397 of Eurychone 395, 399 of Fernandezia 482, 483 of Galeoglossum 463, 465 of Gastrochilus 189, 191 of Gongora 485 of Grosourdya 193, 194

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Gunnarella 195, 197 Hederorkis 451, 453 Hemipiliopsis 466, 468 Holcoglossum 195, 201 Hymenorchis 201, 203 Jejewoodia 203, 205 Jumellea 397, 401 Lemurella 401, 403 Lemurorchis 403, 405 Listrostachys 405, 407 Luisia 206, 207 Macropodanthus 209, 209 Margelliantha 407, 409 Microcoelia 409, 411 Micropera 211, 211 Microsaccus 212, 214 Mobilabium 213, 216 Mystacidium 411, 413 Neobathiea 413, 415 Neolindleya 469, 471 Nephrangis 415, 417 Nothostele 476, 478 Notyliopsis 484, 484 Oeonia 417, 419 Oeoniella 419, 421 Omoea 215, 218 Ophioglossella 216, 220 Ossiculum 421, 423 Papilionanthe 218, 222 Papillilabium 223, 224 Paraphalaenopsis 225, 226 Pelatantheria 228, 228 Pennilabium 230, 230 Peristeranthus 232, 232 Phalaenopsis 234, 238 Phragmorchis 241, 243 Plectorrhiza 243, 245 Plectrelminthus 423, 425 Podangis 425, 427 Polystachya 455–6, 458 Polystachyinae 451 Pomatocalpa 245, 247 Porrorhachis 247, 249 Pseuderia 471, 473 Psychopsiella 474, 474 Pteroceras 250, 251 Rangaeris 427, 429 Renanthera 252, 254 Rhaesteria 429, 431 Rhinerrhiza 254, 256 Rhinerrhizopsis 256, 258 Rhipidoglossum 431, 433 Rhynchogyna 258, 260 Rhychostylis 260, 262 Robiquetia 263, 266 Saccolabiopsis 266, 268 Saccolabium 268, 270 Santotomasia 270, 272 Sarcanthopsis 272, 274 Sarcochilus 274, 277 Sarcoglyphis 278, 280 Sarcophyton 279, 282 Schistotylus 282, 284 Schoenorchis 284, 286 Seidenfadenia 286, 288 Seidenfadeniella 289, 290 Singchia 291, 292 Sirhookera 109, 111

SUBJECT INDEX

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Smithsonia 294, 294 Smitinandia 295, 297 Sobennikoffia 434, 435 Solenangis 436, 438 Sphyrarhynchus 438, 440 Spongiola 296, 299 Stereochilus 298, 301 Summerhayesia 440, 442 Taeniophyllum 301, 304 Taeniorrhiza 442, 444 Taprobanea 305, 307 Thrixspermum 307, 309 Trachoma 309, 311 Triceratorhynchus 444, 445 Trichoglottis 312, 314 Tridactyle 446, 447 Tuberolabium 315, 317 Uncifera 316, 319 Vanda 319, 324 Vandeae 101, 102 Vandopsis 327, 329 Xenikophyton 329, 331 Ypsilopus 448, 450

ecology of Acampe 131–2 of Adenoncos 132 of Adrorhizon 106 of Aerangis 348 of Aeranthes 351 of Aerides 136 of Agrostophyllum 334 of Ambrella 353 of Ancistrorhynchus 355 of Angraecopsis 357 of Angraecum 362–3 of Arachnis 141 of Beclardia 365 of Biermannia 142 of Bogoria 143 of Bolusiella 367 of Brachypeza 145 of Brassia 481 of Bromheadia 109 of Bulbophyllum 47 of Calymmanthera 147 of Calyptrochilum 369 of Campylocentrum 370–1 of Cardiochilos 372 of Ceratocentron 148 of Chamaeanthus 151 of Chauliodon 374 of Chiloschista 154–5 of Chroniochilus 158 of Cleisocentron 160 of Cleisomeria 161 of Cleisostoma 165 of Cleisostomopsis 167 of Cottonia 168 of Cribbia 376 of Cryptopus 378 of Cryptopylos 170 of Cyanaeorchis 480 of Cyrtorchis 380 of Deceptor 172 of Dendrobium 94 of Dendrophylax 384 of Diaphananthe 388

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Dimorphorchis 174 Dinklageella 389 Diplocentrum 176 Diploprora 178 Distylodon 391 Dryadorchis 181 Drymoanthus 182 Dyakia 184 Earina 337 Eclecticus 187 Eggelingia 393 Erasanthe 393 Eurychone 397 Fernandezia 483 Galeoglossum 465–6 Gastrochilus 190 Grosourdya 193–4 Gunnarella 195 Hederorkis 452 Hemipiliopsis 468 Holcoglossum 197 Hymenorchis 202 Jejewoodia 204 Jumellea 399 Lemurella 401 Lemurorchis 403 Listrostachys 405 Luisia 207 Macropodanthus 209 Margelliantha 407 Microcoelia 409 Micropera 211–12 Microsaccus 212 Mobilabium 214 Mystacidium 413 Neobathiea 415 Neolindleya 469 Nephrangis 416–7 Nothostele 478 Notyliopsis 484 Oeonia 418 Oeoniella 420 Omoea 216 Ophioglossella 217 Ossiculum 421 Papilionanthe 219–20 Papillilabium 224 Paraphalaenopsis 226 Pelatantheria 228 Pennilabium 230 Peristeranthus 232 Phalaenopsis 239–40 Phragmorchis 241 Plectorrhiza 243–4 Plectrelminthus 423 Podangis 425 Polystachya 460 Pomatocalpa 246 Porrorhachis 248 Pseuderia 472 Psychopsiella 474 Pteroceras 251 Rangaeris 429 Renanthera 253 Rhaesteria 430 Rhinerrhiza 255 Rhinerrhizopsis 258 Rhipidoglossum 433

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Rhynchogyna 259 Rhychostylis 262 Robiquetia 263 Saccolabiopsis 266 Saccolabium 268 Santotomasia 270 Sarcanthopsis 272 Sarcochilus 275–6 Sarcoglyphis 278 Sarcophyton 280 Schistotylus 282 Schoenorchis 284 Seidenfadenia 288 Seidenfadeniella 290 Singchia 292 Sirhookera 110 Smithsonia 294 Smitinandia 295 Sobennikoffia 435 Solenangis 436 Sphyrarhynchus 440 Spongiola 297 Stereochilus 299 Summerhayesia 442 Taeniophyllum 303–4 Taeniorrhiza 443 Taprobanea 305 Thrixspermum 308 Trachoma 310 Triceratorhynchus 444 Trichoglottis 314 Tridactyle 447 Tuberolabium 315 Uncifera 317 Vanda 325–6 Vandopsis 327 Xenikophyton 330 Ypsilopus 450

infrageneric treatment of Aerangis 345 of Agrostophyllum 332 of Angraecum 358–61 of Bromheadia 107–8 of Bulbophyllum 7–41 of Cleisostoma 163–4 of Dendrobium 55–73 of Polystachya 456–8 of Vandeae 102, 104 key to Adrorhizinae 104 to Aeridinae 124–9 to Agrostophyllinae 331 to Angraecinae 341–4 to Angraecum 358–60 to Bromheadia 108 to Bulbophyllum 7, 9, 11–13, 15–16, 19–25, 40 to Dendrobieae 4 to Dendrobium 55–62 to Polystachya 458–9 to Polystachyinae 451 to Vandeae 102, 104 palynology of Adrorhizon 106 of Aerangis 345 of Aeranthes 351

539

SUBJECT INDEX

palynology (cont.) of Aerides 134 of Aeridinae 113 of Agrostophyllum 332 of Ambrella 352 of Ancistrorhynchus 355 of Angraecopsis 357 of Angraecum 361 of Beclardia 365 of Bolusiella 367 of Brassia 481 of Bromheadia 108 of Bulbophyllum 43 of Calyptrochilum 369 of Campylocentrum 370 of Cardiochilos 372 of Chauliodon 374 of Chiloschista 154 of Cleisostoma 164 of Cribbia 376 of Cryptopus 378 of Cyrtorchis 380 of Dendrobium 56 of Dendrophylax 383 of Diaphananthe 386 of Dinklageella 389 of Distylodon 391 of Earina 336 of Eggelingia 391 of Erasanthe 393 of Eurychone 396 of Fernandezia 483 of Galeoglossum 464 of Gastrochilus 189 of Hederorkis 452 of Hemipiliopsis 468 of Jumellea 398 of Lemurella 401 of Lemurorchis 403 of Listrostachys 405 of Luisia 206 of Margelliantha 56 of Microcoelia 409 of Mystacidium 413 of Neobathiea 414 of Neolindleya 469 of Nephrangis 415 of Nothostele 476 of Notyliopsis 484 of Oeonia 417 of Oeoniella 419 of Ossiculum 421 of Phalaenopsis 236 of Plectrelminthus 423 of Podangis 425 of Polystachya 460 of Pseuderia 471 of Psychopsiella 474 of Rangaeris 428 of Rhaesteria 429 of Rhipidoglossum 433 of Rhychostylis 262 of Sarcochilus 274 of Schoenorchis 284 of Seidenfadenia 286 of Sirhookera 109 of Smithsonia 294 of Sobennikoffia 435

540

of Solenangis 436 of Sphyrarhynchus 438 of Summerhayesia 442 of Taeniorrhiza 443 of Triceratorhynchus 444 of Tridactyle 447 of Vanda 322 of Ypsilopus 450 phylogenetics of Acampe 131 of Adenoncos 132 of Aerangis 347–8 of Aerides 136 of Aeridinae 113–19, 116, 117 of Agrostophyllum 334 of Amesiella 137–8 of Angraecinae 338–40, 339 of Angraecum 362 of Arachnis 140–1 of Beclardia 365 of Biermannia 142 of Bogoria 143 of Bolusiella 367 of Brassia 481 of Bulbophyllum 46–7, 48, 49 of Calymmanthera 147 of Calyptrochilum 369 of Campylocentrum 370 of Ceratocentron 148 of Chamaeanthus 151 of Chauliodon 374 of Chiloschista 154 of Chroniochilus 157 of Cleisocentron 160 of Cleisomeria 161 of Cleisostoma 165 of Cleisostomopsis 167 of Cottonia 168 of Cribbia 376 of Cryptopus 378 of Cryptopylos 170 of Cyanaeorchis 479–80 of Deceptor 172 of Dendrobieae 3 of Dendrobium 89–91, 92, 93 of Dendrophylax 383–4 of Diaphananthe 388 of Dimorphorchis 174 of Diplocentrum 176 of Diploprora 178 of Disa 479 of Dryadorchis 180 of Drymoanthus 182 of Dyakia 184 of Eclecticus 187 of Erasanthe 393 of Eurychone 396 of Fernandezia 483 of Galeoglossum 465 of Gastrochilus 190 of Grosourdya 193 of Gunnarella 195 of Holcoglossum 197 of Hymenorchis 202 of Jejewoodia 204 of Jumellea 399 of Lemurella 401 of Lemurorchis 403

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Listrostachys 405 Luisia 207 Macropodanthus 209 Microcoelia 409 Micropera 211 Microsaccus 212 Mobilabium 214 Mystacidium 413 Neobathiea 415 Neolindleya 469 Nephrangis 416 Nothostele 476–8 Notyliopsis 484 Oeonia 418 Oeoniella 420 Omoea 215–16 Ophioglossella 217 Ossiculum 421 Papilionanthe 219 Papillilabium 224 Paraphalaenopsis 226 Pelatantheria 228 Pennilabium 230 Peristeranthus 232 Phalaenopsis 239 Phragmorchis 241 Plectorrhiza 243 Podangis 425 Polystachya 460 Polystachyinae 451 Pomatocalpa 246 Porrorhachis 248 Pseuderia 472 Psychopsiella 474 Pteroceras 251 Rangaeris 429 Renanthera 252 Rhinerrhiza 254 Rhinerrhizopsis 256 Rhipidoglossum 433 Rhynchogyna 259 Rhychostylis 262 Robiquetia 263 Saccolabiopsis 266 Saccolabium 268 Santotomasia 270 Sarcanthopsis 272 Sarcochilus 275 Sarcoglyphis 278 Sarcophyton 280 Schistotylus 282 Schoenorchis 284 Seidenfadenia 288 Seidenfadeniella 290 Singchia 291–2 Smithsonia 294 Smitinandia 295 Solenangis 436 Sphyrarhynchus 440 Spongiola 297 Stereochilus 299 Summerhayesia 442 Taeniophyllum 303 Taprobanea 305 Thrixspermum 307–8 Trachoma 310 Trichoglottis 314 Tridactyle 447

SUBJECT INDEX

of Tuberolabium 315 of Uncifera 317 of Vanda 325 of Vandeae 101–2, 103 of Vandopsis 327 of Xenikophyton 329 of Ypsilopus 450 phytochemistry of Acampe 131 of Adenoncos 132 of Adrorhizon 106 of Aerangis 345–7, 348 of Aeranthes 351 of Aerides 134–6 of Agrostophyllum 333–4, 335 of Ambrella 353 of Amesiella 137 of Ancistrorhynchus 355 of Angraecopsis 357 of Angraecum 362 of Arachnis 140 of Beclardia 365 of Biermannia 142 of Bogoria 142 of Bolusiella 367 of Brachypeza 145 of Brassia 481 of Bromheadia 108–9 of Bulbophyllum 43–6, 44 of Calymmanthera 147 of Calyptrochilum 369 of Campylocentrum 370 of Cardiochilos 372 of Ceratocentron 148 of Chamaeanthus 151 of Chauliodon 374 of Chiloschista 154 of Chroniochilus 157 of Cleisocentron 159 of Cleisomeria 161 of Cleisostoma 164–5 of Cleisostomopsis 167 of Cottonia 168 of Cribbia 376 of Cryptopus 378 of Cryptopylos 170 of Cyrtorchis 380 of Deceptor 172 of Dendrobium 74–89, 77–8, 80–1, 86–8 of Dendrophylax 383 of Diaphananthe 388 of Dimorphorchis 174 of Dinklageella 389 of Diplocentrum 176 of Diploprora 178 of Distylodon 391 of Dryadorchis 180 of Drymoanthus 182 of Dyakia 184 of Earina 336 of Eclecticus 187 of Eggelingia 393 of Erasanthe 393 of Eurychone 396 of Fernandezia 483 of Galeoglossum 464 of Gastrochilus 189–90 of Grosourdya 193

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Gunnarella 195 Hederorkis 452 Hemipiliopsis 468 Holcoglossum 197 Hymenorchis 202 Jejewoodia 204 Jumellea 399 Lemurella 401 Lemurorchis 403 Listrostachys 405 Luisia 206–7 Macropodanthus 209 Margelliantha 407 Microcoelia 409 Micropera 211 Microsaccus 212 Mobilabium 214 Mystacidium 413 Neobathiea 415 Neolindleya 469 Nephrangis 416 Nothostele 476 Notyliopsis 484 Oeonia 418 Oeoniella 419–20 Omoea 215 Ophioglossella 217 Ossiculum 421 Papilionanthe 219, 222 Papillilabium 223 Paraphalaenopsis 225–6 Pelatantheria 228 Pennilabium 230 Peristeranthus 232 Phalaenopsis 237–9, 238, 239 Phragmorchis 241 Plectorrhiza 243 Plectrelminthus 423 Podangis 425 Polystachya 460 Pomatocalpa 246 Porrorhachis 247 Pseuderia 471 Psychopsiella 474 Pteroceras 251 Rangaeris 429 Renanthera 252 Rhaesteria 430 Rhinerrhiza 254 Rhinerrhizopsis 256 Rhipidoglossum 433 Rhynchogyna 259 Rhychostylis 262 Robiquetia 263 Saccolabiopsis 266 Saccolabium 268 Santotomasia 270 Sarcanthopsis 272 Sarcochilus 275 Sarcoglyphis 278 Sarcophyton 280 Schistotylus 282 Schoenorchis 284 Seidenfadenia 288 Seidenfadeniella 290 Singchia 291 Sirhookera 110 Smithsonia 294

of Smitinandia 295 of Sobennikoffia 435 of Solenangis 436 of Sphyrarhynchus 438 of Spongiola 296 of Stereochilus 299 of Summerhayesia 442 of Taeniophyllum 302–3 of Taeniorrhiza 443 of Taprobanea 305 of Thrixspermum 307 of Trachoma 309 of Triceratorhynchus 444 of Trichoglottis 314 of Tridactyle 447 of Tuberolabium 315 of Uncifera 317 of Vanda 323–5, 324 of Vandopsis 327, 329 of Xenikophyton 329 of Ypsilopus 450 pollination of Acampe 132 of Adenoncos 132 of Adrorhizon 106 of Aerangis 349 of Aeranthes 351 of Aerides 137 of Agrostophyllum 334 of Ambrella 353 of Amesiella 138 of Ancistrorhynchus 355 of Angraecopsis 357 of Angraecum 363 of Arachnis 141 of Beclardia 365 of Biermannia 142 of Bogoria 144 of Bolusiella 367 of Brachypeza 146 of Brassia 482 of Bromheadia 109 of Bulbophyllum 47, 48, 50 of Calymmanthera 147 of Calyptrochilum 369 of Campylocentrum 371 of Cardiochilos 372 of Ceratocentron 150 of Chamaeanthus 151 of Chauliodon 374 of Chiloschista 156 of Chroniochilus 158 of Cleisocentron 160 of Cleisomeria 161 of Cleisostoma 165 of Cleisostomopsis 167 of Cottonia 168 of Cribbia 376 of Cryptopus 378 of Cryptopylos 170 of Cyrtorchis 380 of Deceptor 172 of Dendrobium 94–6 of Dendrophylax 384–5 of Diaphananthe 388 of Dimorphorchis 174 of Dinklageella 389 of Diplocentrum 176

541

SUBJECT INDEX

pollination (cont.) of Diploprora 178 of Distylodon 391 of Dryadorchis 181 of Drymoanthus 183 of Dyakia 184 of Earina 337 of Eclecticus 187 of Eggelingia 393 of Erasanthe 395 of Eurychone 397 of Fernandezia 483 of Galeoglossum 466 of Gastrochilus 190–1 of Grosourdya 194 of Gunnarella 195 of Hederorkis 452 of Holcoglossum 197–9 of Hymenorchis 202 of Jejewoodia 204 of Jumellea 399 of Lemurella 401 of Lemurorchis 403 of Listrostachys 405 of Luisia 207 of Macropodanthus 209 of Margelliantha 407 of Microcoelia 410 of Micropera 212 of Microsaccus 212 of Mobilabium 214 of Mystacidium 413 of Neobathiea 415 of Neolindleya 469 of Nephrangis 417 of Nothostele 478 of Notyliopsis 485 of Oeonia 418 of Omoea 216 of Ophioglossella 217 of Ossiculum 421 of Papilionanthe 220 of Papillilabium 224 of Paraphalaenopsis 226 of Pelatantheria 228 of Pennilabium 230 of Peristeranthus 232 of Phalaenopsis 240 of Phragmorchis 241 of Plectorrhiza 244 of Plectrelminthus 424 of Podangis 427 of Polystachya 460–1 of Pomatocalpa 246 of Porrorhachis 248 of Pseuderia 472 of Psychopsiella 474 of Pteroceras 251 of Rangaeris 429 of Renanthera 253 of Rhaesteria 430 of Rhinerrhiza 255 of Rhinerrhizopsis 258 of Rhipidoglossum 433 of Rhynchogyna 259 of Rhychostylis 262 of Robiquetia 263 of Saccolabiopsis 266

542

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Saccolabium 268 Santotomasia 270 Sarcanthopsis 272 Sarcochilus 276 Sarcoglyphis 278 Sarcophyton 280 Schistotylus 282 Schoenorchis 284 Seidenfadenia 288 Seidenfadeniella 290 Singchia 292 Sirhookera 111 Smithsonia 294 Smitinandia 295 Sobennikoffia 435 Solenangis 436 Sphyrarhynchus 440 Spongiola 297 Stereochilus 299 Summerhayesia 442 Taeniophyllum 304 Taeniorrhiza 443 Taprobanea 305 Thrixspermum 308 Trachoma 310 Triceratorhynchus 444 Trichoglottis 314 Tridactyle 448 Tuberolabium 315 Uncifera 317 Vanda 326 Vandopsis 327 Xenikophyton 330 Ypsilopus 450

taxonomic literature of Acampe 132 of Adenoncos 133 of Adrorhizon 106 of Aerangis 349 of Aeranthes 351 of Aerides 137 of Aeridinae 123 of Agrostophyllum 335 of Ambrella 353 of Amesiella 139 of Ancistrorhynchus 355 of Angraecinae 341 of Angraecopsis 358 of Angraecum 363 of Arachnis 141 of Beclardia 365 of Bogoria 144 of Bolusiella 367 of Brassia 482 of Bromheadia 109 of Bulbophyllum 50–1 of Calyptrochilum 369 of Campylocentrum 371–2 of Ceratocentron 150 of Chamaeanthus 152 of Chauliodon 374 of Chiloschista 156 of Cleisocentron 161 of Cleisomeria 161 of Cleisostoma 166 of Cleisostomopsis 168 of Cribbia 376

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Cryptopus 378 Cryptopylos 170 Cyanaeorchis 480 Cyrtorchis 382 Deceptor 172 Dendrobieae 4 Dendrobium 99–100 Dendrophylax 385 Diaphananthe 388–9 Dimorphorchis 175 Dinklageella 389–90 Diploprora 178, 180 Disa 479 Dryadorchis 181–2 Dyakia 185–6 Earina 337–8 Erasanthe 395 Eurychone 397 Fernandezia 483 Galeoglossum 466 Gastrochilus 191 Grosourdya 194 Hederorkis 453 Hemipiliopsis 468–9 Holcoglossum 200–1 Hymenorchis 203 Jejewoodia 205 Jumellea 399 Lemurella 402 Lemurorchis 403 Listrostachys 405 Luisia 207–8 Macropodanthus 210 Margelliantha 407 Microcoelia 411 Micropera 212 Microsaccus 212 Mystacidium 413 Neobathiea 415 Neolindleya 469–71 Nephrangis 417 Nothostele 479 Notyliopsis 485 Oeonia 419 Oeoniella 421 Ophioglossella 218 Ossiculum 422 Papilionanthe 220 Papillilabium 224 Paraphalaenopsis 226 Pelatantheria 229 Phalaenopsis 241 Phragmorchis 242 Plectorrhiza 245 Podangis 427 Polystachya 461 Polystachyinae 451 Pomatocalpa 247 Porrorhachis 249 Pseuderia 473 Psychopsiella 475 Pteroceras 252 Rangaeris 429 Renanthera 253 Rhaesteria 431 Rhinerrhiza 255–6 Rhipidoglossum 433 Rhynchogyna 260

SUBJECT INDEX

of Rhychostylis 263 of Robiquetia 264 of Saccolabium 269 of Sarcanthopsis 273 of Sarcochilus 277–8 of Sarcoglyphis 278 of Sarcophyton 280 of Schoenorchis 286 of Seidenfadenia 288 of Singchia 292 of Sirhookera 111 of Smithsonia 295 of Smitinandia 295 of Solenangis 438 of Sphyrarhynchus 440 of Stereochilus 299 of Summerhayesia 442 of Taeniophyllum 304 of Taeniorrhiza 444 of Taprobanea 305 of Thrixspermum 309 of Trachoma 311 of Triceratorhynchus 445 of Trichoglottis 314–15 of Tridactyle 448 of Tuberolabium 315 of Vanda 326–7 of Vandeae 102 of Vandopsis 327 of Ypsilopus 450 taxonomic notes of Acampe 132 of Aeranthes 351 of Aerides 137 of Aeridinae 123 of Ambrella 353 of Amesiella 139 of Ancistrorhynchus 355 of Angraecinae 341 of Angraecopsis 357–8 of Biermannia 142 of Bolusiella 367 of Brachypeza 146 of Calymmanthera 147 of Cardiochilos 372 of Ceratocentron 150 of Chauliodon 374 of Chroniochilus 158 of Cleisocentron 160 of Cleisostoma 165 of Cleisostomopsis 167–8 of Cottonia 168 of Cryptopus 378 of Cryptopylos 170 of Cyanaeorchis 480 of Cyrtorchis 381 of Deceptor 172 of Dendrobium 98–9 of Diaphananthe 388 of Dimorphorchis 175 of Dinklageella 389 of Distylodon 391 of Dyakia 185 of Eclecticus 188 of Eggelingia 393 of Erasanthe 395 of Eurychone 397 of Gastrochilus 191

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Grosourdya 194 Hederorkis 453 Hemipiliopsis 468 Holcoglossum 199–200 Hymenorchis 203 Jumellea 399 Lemurella 402 Lemurorchis 403 Listrostachys 405 Luisia 207 Macropodanthus 210 Margelliantha 407 Microcoelia 411 Mystacidium 413 Neobathiea 415 Neolindleya 469 Nephrangis 417 Nothostele 478 Oeonia 419 Oeoniella 420–1 Ophioglossella 218 Paraphalaenopsis 226 Pelatantheria 228 Pennilabium 230–1 Phalaenopsis 241 Phragmorchis 241–2 Plectrelminthus 424 Polystachyinae 451 Porrorhachis 248 Pseuderia 472–3 Pteroceras 252 Rhaesteria 431 Rhipidoglossum 433 Rhynchogyna 260 Saccolabium 268–9 Santotomasia 270 Sarcochilus 277 Sarcoglyphis 278 Sarcophyton 280 Schoenorchis 285–6 Seidenfadenia 288 Seidenfadeniella 290 Singchia 292 Sirhookera 111 Smithsonia 294–5 Sobennikoffia 435 Spongiola 297 Stereochilus 299 Summerhayesia 442 Taeniorrhiza 444 Taprobanea 305 Trachoma 311 Triceratorhynchus 444–5 Tridactyle 448 Tuberolabium 315 Vanda 326 Xenikophyton 330–1 Ypsilopus 450

Acampe 132 Adenoncos 132 Adrorhizon 106 Aerangis 349 Aerides 137 Agrostophyllum 334 Ambrella 353 Amesiella 138

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Ancistrorhynchus 355 Angraecopsis 357 Angraecum 363 Arachnis 141 Beclardia 365 Biermannia 142 Bogoria 144 Bolusiella 367 Brachypeza 146 Brassia 482 Bromheadia 109 Bulbophyllum 50 Calymmanthera 147 Calyptrochilum 369 Campylocentrum 371 Cardiochilos 372 Ceratocentron 150 Chamaeanthus 152 Chauliodon 374 Chiloschista 156 Chroniochilus 158 Cleisocentron 160 Cleisomeria 161 Cleisostoma 165 Cleisostomopsis 167 Cottonia 168 Cribbia 376 Cryptopus 378 Cryptopylos 170 Cyanaeorchis Cyrtorchis 380 Deceptor 172 Dendrobium 96–7 Dendrophylax 385 Diaphananthe 388 Dimorphorchis 174 Dinklageella 389 Diplocentrum 176 Diploprora 178 Dryadorchis 181 Drymoanthus 183 Dyakia 184 Earina 337 Eclecticus 187 Eggelingia 393 Erasanthe 395 Eurychone 397 Fernandezia 483 Galeoglossum 466 Gastrochilus 191 Grosourdya 194 Gunnarella 195 Hederorkis 452 Hemipiliopsis 468 Holcoglossum 199 Hymenorchis 202 Jejewoodia 205 Jumellea 399 Lemurella 402 Lemurorchis 403 Listrostachys 405 Luisia 207 Macropodanthus 209 Margelliantha 407 Microcoelia 411 Micropera 212 Microsaccus 212 Mobilabium 214

543

SUBJECT INDEX

uses (cont.) of Mystacidium 413 of Neobathiea 415 of Neolindleya 469 of Nephrangis 417 of Nothostele 478 of Notyliopsis 485 of Oeonia 419 of Omoea 216 of Ophioglossella 217 of Ossiculum 421 of Papilionanthe 220 of Papillilabium 224 of Paraphalaenopsis 226 of Pelatantheria 228 of Pennilabium 230 of Peristeranthus 233 of Phalaenopsis 240 of Phragmorchis 241 of Plectorrhiza 244 of Plectrelminthus 424 of Podangis 427 of Polystachya 461 of Pomatocalpa 246

544

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Porrorhachis 248 Pseuderia 472 Psychopsiella 475 Pteroceras 251 Rangaeris 429 Renanthera 253 Rhaesteria 430 Rhinerrhiza 255 Rhinerrhizopsis 258 Rhipidoglossum 433 Rhynchogyna 260 Rhychostylis 262–3 Robiquetia 264 Saccolabiopsis 266 Saccolabium 268 Santotomasia 270 Sarcanthopsis 272 Sarcochilus 276 Sarcoglyphis 278 Sarcophyton 280 Schistotylus 282 Schoenorchis 284 Seidenfadenia 288 Seidenfadeniella 290

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Singchia 292 Sirhookera 111 Smithsonia 294 Smitinandia 295 Sobennikoffia 435 Solenangis 438 Sphyrarhynchus 440 Spongiola 297 Stereochilus 299 Summerhayesia 442 Taeniophyllum 304 Taeniorrhiza 444 Taprobanea 305 Thrixspermum 308 Trachoma 310 Triceratorhynchus 444 Trichoglottis 314 Tridactyle 448 Tuberolabium 315 Uncifera 317 Vanda 326 Vandopsis 327 Xenikophyton 330 Ypsilopus 450