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Bulbophyllum

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Bulbophyllum
Bulbophyllum echinolabium
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Monocots
Order: Asparagales
Family: Orchidaceae
Subfamily: Epidendroideae
Subtribe: Dendrobiinae
Genus: Bulbophyllum
Thouars, 1822[1]
Type species
Bulbophyllum nutans
Species

See List of Bulbophyllum species

Diversity
2167 species
Synonyms[2]
List of synonyms
  • Acrochaene Lindl.
  • Adelopetalum Fitzg.
  • Anisopetalon Hook.
  • Blepharochilum M.A.Clem. & D.L.Jones
  • Bolbophyllaria Rchb.f.
  • Bolbophyllopsis Rchb.f.
  • Bolbophyllum Spreng.
  • Bulbophyllaria S.Moore
  • Canacorchis Guillaumin
  • Carparomorchis M.A.Clem. & D.L.Jones
  • Chaseella Summerh.
  • Cirrhopetalum Lindl.
  • × Cirrhophyllum Anon.
  • Cochlia Blume
  • Codonosiphon Schltr.
  • Dactylorhynchus Schltr.
  • Didactyle Lindl.
  • Diphyes Blume
  • Drymoda Lindl.
  • Ephippium Blume
  • Epicranthes Blume
  • Epicrianthes Blume
  • Ferruminaria Garay, Hamer & Siegerist
  • Fruticicola (Schltr.) M.A.Clem. & D.L.Jones
  • Genyorchis Schltr.
  • Hamularia Aver. & Averyanova
  • Hapalochilus (Schltr.) Senghas
  • Henosis Hook.f.
  • Hippoglossum Breda
  • Hordeanthos Szlach.
  • Hyalosema Rolfe
  • Ichthyostomum D.L.Jones, M.A.Clem. & Molloy
  • Ione Lindl.
  • Jejosephia A.N.Rao & Mani
  • Katherinea A.D.Hawkes
  • Kaurorchis D.L.Jones & M.A.Clem.
  • Lepanthanthe (Schltr.) Szlach.
  • Lyraea Lindl.
  • Macrolepis A.Rich.
  • Malachadenia Lindl.
  • Mastigion Garay, Hamer & Siegerist
  • Megaclinium Lindl.
  • Monomeria Lindl.
  • Monosepalum Schltr.
  • Odontostylis Blume
  • Odontostylis Breda
  • Oncophyllum D.L.Jones & M.A.Clem.
  • Osyricera Blume
  • Oxysepala Wight
  • Pachyrhachis A.Rich.
  • Papulipetalum (Schltr.) M.A.Clem. & D.L.Jones
  • Pedilochilus Schltr.
  • Pelma Finet
  • Peltopus (Schltr.) Szlach. & Marg.
  • Phyllorkis Thouars
  • Rhytionanthos Garay, Hamer & Siegerist
  • Saccoglossum Schltr.
  • Sarcobodium Beer
  • Sarcopodium Lindl. & Paxton
  • Serpenticaulis M.A.Clem. & D.L.Jones
  • Sestochilos Breda
  • Spilorchis D.L.Jones & M.A.Clem.
  • Sunipia Lindl.
  • Synarmosepalum Garay, Hamer & Siegerist
  • Tapeinoglossum Schltr.
  • Taurostalix Rchb.f.
  • Trachyrhachis (Schltr.) Szlach.
  • Trias Lindl.
  • Tribrachia Lindl.
  • Tripudianthes (Seidenf.) Szlach. & Kras
  • Vesicisepalum (J.J.Sm.) Garay, Hamer & Siegerist
  • Xiphizusa Rchb.f.
  • Zygoglossum Reinw.

Bulbophyllum is a genus of mostly epiphytic and lithophytic orchids in the family Orchidaceae. It is the largest genus in the orchid family and one of the largest genera of flowering plants with more than 2,000 species, exceeded in number only by Astragalus. These orchids are found in diverse habitats throughout most of the warmer parts of the world including Africa, southern Asia, Latin America, the West Indies, and various islands in the Indian and Pacific Oceans.[3] Orchids in this genus have thread-like or fibrous roots that creep over the surface of trees or rocks or hang from branches. The stem is divided into a rhizome and a pseudobulb, a feature that distinguished this genus from Dendrobium. There is usually only a single leaf at the top of the pseudobulb and from one to many flowers are arranged along an unbranched flowering stem that arises from the base of the pseudobulb. Several attempts have been made to separate Bulbophyllum into smaller genera, but most have not been accepted by the World Checklist of Selected Plant Families.

Description

[edit]

Plants in the genus Bulbophyllum are epiphytic or lithophytic sympodial herbs with thread-like or fibrous roots that creep over the surface on which they grow. The stem consists of a rhizome and a pseudobulb, the latter with one or two usually fleshy or leathery leaves. The flowers are arranged on an unbranched raceme that emerges from the pseudobulb, usually from its base. The dorsal sepal is free from the lateral sepals which themselves may be free or fused to each other. The petals are also free from each other and smaller than the lateral sepals. The labellum is often fleshy, curved and hinged to the base of the column.[1][4][5][6]

Distribution and habitat

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The center of diversity of this genus is in the montane forests of Papua New Guinea (more than 600 species) which seems to be the evolutionary homeland,[7] though the genus is pantropical and widespread, occurring in Australia, Southeast Asia (with over 200 species in Borneo), India, Madagascar (with 135 species, some endemic), Africa and in tropical central and South America.[6]

The erect to pendent inflorescence arises laterally from the base of the pseudobulb. The flower form has a basic structural blueprint that serves to identify this genus. But this form can be very diverse : compound or single, with few to many flowers, with the resupinate flowers arranged spirally or in two vertical ranks. The sepals and the petals can also be very varied : straight or turned down, without footstalk or with a long claw at the base. They are often hairy or callous. There are two to four hard and waxy pollinia with stipes present or absent. The fruits are beakless capsules.[6]

Taxonomy and naming

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The genus Bulbophyllum was first formally described in 1822 by Louis-Marie Aubert du Petit-Thouars in his book Histoire particulière des plantes orchidées recueillies sur les trois Iles Australes d'Afrique, de France, de Bourbon et de Madagascar in which he described eighteen species of Bulbophyllum.[1][8]

There are now more than 2,800 records (accepted names and synonyms) for this genus. This large number and the great variety of its forms make the genus a considerable challenge for taxonomists: 120 sections and subgenera have been listed.[6]

The genus name (Bulbophyllum) is derived from the Greek bolbos meaning 'bulb' and phyllon, 'a leaf', referring to the pseudobulbs on top of which the leaf grows.[9][6]

In 2014, Alec Pridgeon and others proposed merging the genus Drymoda with Bulbophyllum in the Genera Orchidacearum and the change is accepted by Plants of the World Online and the World Checklist of Selected Plant Families.[10][11][12] The former species of Drymoda included D. digitata (now B. digitatum),[13] D. gymnopus (now B. gymnopus),[14] D. latisepala (now B. capillipes),[15] D. picta (the type species of Drymoda, now B. drymoda),[16] and D. siamensis (now B. ayuthayense).[17]

Evolution and biogeography

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Molecular phylogenetic studies place the origin of the orchid genus Bulbophyllum into the early Miocene. Biogeographic analyses and ancestral area reconstructions identified the Asia-Pacific region as the ancestral area of Bulbophyllum and suggest an early-to-late Miocene scenario of 'out-of-Asia-Pacific' origin and progressive (east-to-west) dispersal-mediated diversification, resulting in three additional radiations in Madagascar, Africa and the Neotropics, respectively.[18]

Sections

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In 2019 Bulbophyllum were separated into four clades Neotropics, Africa, Madagascar, and the Asia-Pacific.[19][20]

Africa Clade

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Image Section Type species
Bifarium Bulbophyllum bifarium
Carnosisepala Bulbophyllum carnosisepalum
Chaseela Bulbophyllum pseudohydra
Comata Bulbophyllum comatum
Denticulata Bulbophyllum denticulatum
Genyorchis Bulbophyllum apetalum
Genyorchis Bulbophyllum deshmukhii
Gilgiana Bulbophyllum gilgianum
Megaclinium Bulbophyllum falcatum
Ptiloglossum Bulbophyllum barbigerum
Oreonastes Bulbophyllum oreonastes

Madagascar Clade

[edit]
Image Section Type species
Alcistachys Bulbophyllum occlusum
Bifalcula Bulbophyllum implexum
Bulbophyllum Bulbophyllum nutans
Elasmotopus Bulbophyllum oxycalyx
Ikongoense Bulbophyllum ikongoense
Inversiflora Bulbophyllum cardiobulbum
Kinethrix Bulbophyllum mirificum
Lichenophylax Bulbophyllum lichenophylax
Lupulina Bulbophyllum occultum
Lyperocephalum Bulbophyllum lyperocephalum
Moratii Bulbophyllum moratii
Pachychlamys Bulbophyllum pachypus
Pantoblepharon Bulbophyllum pantoblepharon
Ploiarium Bulbophyllum coriophorum
Polyradices Bulbophyllum petrae

Neotropical Clade

[edit]
Image Section Type species
Bulbophyllaria Bulbophyllum bracteolatum
Didactyle Bulbophyllum exaltatum
Furvescentia Bulbophyllum nagelii
Micranthae Bulbophyllum micranthum
Napelli Bulbophyllum napellii
Xiphizusa Bulbophyllum chloropterum

Asia-Pacific Clade

[edit]
Image Section Type species
Acrochaene Bulbophyllum kingii
Adelopetalum Bulbophyllum bracteatum
Aeschynanthoides Bulbophyllum dryas
Altisceptrum Bulbophyllum elongatum
Antennata Bulbophyllum fuscopurpureum
Balaenoidea Bulbophyllum balaeniceps
Beccariana Bulbophyllum beccarii
Biflorae Bulbophyllum biflorum
Biseta Bulbophyllum bisetum
Blepharistes Bulbophyllum blepharistes
Brachyantha Bulbophyllum umbellatum
Brachystachyae Bulbophyllum repens
Brachypus Bulbophyllum maxillarioides
Cirrhopetaloides Bulbophyllum longissimum
Cirrhopetalum Bulbophyllum longiflorum
Codonosiphon Bulbophyllum codonanthum
Codonosiphon Bulbophyllum raulersoniae
Desmosanthes Bulbophyllum croceum
Drymoda Bulbophyllum drymoda
Epicranthes Bulbophyllum epicranthes
Ephippium Bulbophyllum lepidum
Eublepharon Bulbophyllum eublepharon
Gongorodes Bulbophyllum digitatum
Hemisterantha Bulbophyllum hemisterranthum
Hirtula Bulbophyllum hirtulum
Hoplandra Bulbophyllum restrepia
Hyalosema Bulbophyllum grandiflorum
Hymenobractea Bulbophyllum infundibuliforme
Intervallatae Bulbophyllum attenuatum
Imitatores Bulbophyllum imitator
Ione Bulbophyllum roseopictum
Lemniscata Bulbophyllum lemniscatum
Leopardinae Bulbophyllum leopardinum
Lepanthanthe Bulbophyllum lepanthiflorum
Lepidorhiza Bulbophyllum amplebracteatum
Macrocaulia Bulbophyllum ovalifolium
Macrouris Bulbophyllum macrourum
Minutissima Bulbophyllum minutissimum
Monanthaparva Bulbophyllum striatellum
Monanthes Bulbophyllum tortuosum
Monomeria Bulbophyllum crabro
Monosepalum Bulbophyllum muricatum
Oxysepala Bulbophyllum clandestinum
Papulipetalum Bulbophyllum papulipetalum
Pedilochilus Bulbophyllum papuanum
Pelma Bulbophyllum absconditum
Peltopus Bulbophyllum peltopus
Phreatiopsis Bulbophyllum phreatiopse
Physometra Bulbophyllum physometrum
Planibulbus Bulbophyllum planibulbe
Piestobulbon Bulbophyllum piestobulbon
Plumata Bulbophyllum plumatum
Polymeres Bulbophyllum tenuifolium
Pseudopelma Bulbophyllum pseudopelma
Racemosae Bulbophyllum careyanum
Rhinanthera Bulbophyllum wrayi
Saurocephalum Bulbophyllum saurocephalum
Schistopetalum Bulbophyllum schistopetalum
Serpenticaulis Bulbophyllum wolfei
Sestochilos Bulbophyllum lobbii
Stachysanthes Bulbophyllum gibbosum
Tapeinoglossum Bulbophyllum centrosemiflorum
Trias Bulbophyllum oblongum
Tripudianthes Bulbophyllum tripudians
Uncifera Bulbophyllum ochroleucum

Ecology

[edit]

Pollination

[edit]

Many Bulbophyllum species have the typical odor of rotting carcasses, and the flies they attract assist in their reproduction through pollination.[6] Nevertheless, some species with mild and pleasant floral fragrance attract Dacini fruit flies (particularly Bactrocera and Zeugodacus species) via methyl eugenol, raspberry ketone or zingerone that also act as floral reward during pollination.[21][22][23][24][25]

To facilitate pollinarium removal and pollinia deposition, this group of orchids, particularly those that attract Dacini fruit flies, possesses a highly modified dynamic lip mechanism - either hinged or see-saw or spring lip (kept either in a close- or open-position depending on resupinate or non-resupinate flower, respectively). When an attracted fly has aligned itself to the opened lip, further probing, feeding and movement towards the lip base result in the lip to snap close, thereby, forcibly tipping the fly in to the column cavity to initiate pollinarium removal or pollinia deposition.[26][27][28]

Use in horticulture

[edit]

Bulbophyllum (abbreviated Bulb. in the horticultural trade)[29] includes species that have been the focus of orchid collectors for over a century. The plants require high humidity combined with good air movement and most of them are ever-blooming - flowering continuously throughout the year. They grow best at moderate light levels, but do not thrive in deep shade. They are considered moderate-to-difficult to cultivate, and require a controlled growing environment to achieve some degree of success. They are not typically suitable as houseplants, and most will not thrive in a Wardian case unless they receive adequate air movement.

The plants' growth habit produces widely spaced pseudobulbs along cord-like rhizome sections, and most of these plants are best accommodated on plaques. Some species in this genus can get very large, but most are small to medium-sized epiphytes from warm, moist, humid tropical forests. They can grow continuously year round with no apparent dormancy period if they are kept warm, are moderate feeders in cultivation, and must be kept moist all the time. They can tolerate dryness for short periods, but they have fine root systems which require moist conditions all the time.[30]

Some of the smaller species do well in pots with small-diameter bark substrate. The plants produce very fine roots generally, and the roots are easily damaged. The plants react poorly to disturbance of their roots. They are easy to maintain once a good environment is established with high humidity and a fresh, buoyant, lightly circulating atmosphere being critical. Most of these species cannot tolerate cold temperatures or freezing.[30]

The flowers produce various odors resembling sap, urine, blood, dung, carrion, and, in some species, fragrant fruity aromas. Most are fly-pollinated, and attract hordes of flies. Bulbophyllum beccarii in bloom has been likened to smelling like a herd of dead elephants and both this species and Bulbophyllum fletcherianum are variously described as making it difficult to walk into a greenhouse in which they are being cultivated if the plants are in bloom because of their overpowering floral odors.[30]

Species

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Main article: List of Bulbophyllum species

Some species are known for their extreme vegetative and floral forms:

Conservation status

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Some Bulbophyllum species are threatened with extinction, and are recognised as such by the World Conservation Union (IUCN):

[edit]

References

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  1. ^ a b c "Bulbophyllum". World Checklist of Selected Plant Families (WCSP). Royal Botanic Gardens, Kew.
  2. ^ "Bulbophyllum". Plants of the World Online. Retrieved 18 July 2021.
  3. ^ David G. Frodin (2004). "History and concepts of big plant genera". Taxon. 53 (3): 753–776. doi:10.2307/4135449. JSTOR 4135449.
  4. ^ Weston, Peter H. "Genus Bulbophyllum". Royal Botanic Garden Sydney. Retrieved 5 December 2018.
  5. ^ Groeneveld, Zoe P.; Jones, David L. "Bulbophyllum". Retrieved 12 May 2023.
  6. ^ a b c d e f Dressler, Robert L. (1990). The Orchids, Natural History and Classification. Harvard University Press. ISBN 0-674-87526-5.
  7. ^ Emly S. Siegerist (2001). Bulbophyllums and their allies: a grower's guide. Portland (OR): Timber Press. ISBN 978-0-88192-506-7.
  8. ^ Thouars, Louis-Marie (1822). Histoire particulière des plantes orchidées recueillies sur les trois Iles Australes d'Afrique, de France, de Bourbon et de Madagascar. Paris. pp. 93–110. doi:10.5962/bhl.title.492. Retrieved 5 December 2018.
  9. ^ Quattrocchi, Umberto (2000). CRC world dictionary of plant names : common names, scientific names, eponyms, synonyms, and etymology. Boca Raton, Florida: CRC Press. p. 372. ISBN 0849326753.
  10. ^ "Drymoda". Plants of the World Online. Retrieved 18 July 2021.
  11. ^ "Drymoda". World Checklist of Selected Plant Families. Retrieved 18 July 2021.
  12. ^ Chase, Mark W.; Cameron, Kenneth M.; Freudenstein, John V.; Pridgeon, Alec M.; Salazar, Gerardo; van den Berg, Cássio; Schuiteman, André (2015). "An updated classification of Orchidaceae". Botanical Journal of the Linnean Society. 177 (2): 151–174. doi:10.1111/boj.12234.
  13. ^ "Drymoda digitata". Plants of the World Online. Retrieved 18 July 2021.
  14. ^ "Drymoda gymnopus". Plants of the World Online. Retrieved 18 July 2021.
  15. ^ "Drymoda latisepala". Plants of the World Online. Retrieved 18 July 2021.
  16. ^ "Drymoda picta". Plants of the World Online. Retrieved 18 July 2021.
  17. ^ "Drymoda siamensis". Plants of the World Online. Retrieved 18 July 2021.
  18. ^ Gamisch, A., Comes, H.P. Clade-age-dependent diversification under high species turnover shapes species richness disparities among tropical rainforest lineages of Bulbophyllum (Orchidaceae). BMC Evol Biol 19, 93 (2019). https://doi.org/10.1186/s12862-019-1416-1
  19. ^ Gamisch, Alexander; Comes, Hans Peter (2019-04-24). "Clade-age-dependent diversification under high species turnover shapes species richness disparities among tropical rainforest lineages of Bulbophyllum (Orchidaceae)". BMC Evolutionary Biology. 19 (1). Springer Science and Business Media LLC: 93. Bibcode:2019BMCEE..19...93G. doi:10.1186/s12862-019-1416-1. ISSN 1471-2148. PMC 6480529. PMID 31014234.
  20. ^ Smidt, Eric C.; Borba, Eduardo L.; Gravendeel, Barbara; Fischer, Gunter A.; van den Berg, Cássio (2011). "Molecular phylogeny of the Neotropical sections of Bulbophyllum (Orchidaceae) using nuclear and plastid spacers". Taxon. 60 (4). Wiley: 1050–1064. doi:10.1002/tax.604009. ISSN 0040-0262.
  21. ^ Tan, Keng-Hong; Nishida, Ritsuo (2000). "Mutual reproductive benefits between a wild orchid, Bulbophyllum patens, and Bactrocera fruit flies via a floral synomone". Journal of Chemical Ecology. 26 (2): 533–546. doi:10.1023/A:1005477926244. S2CID 24971928.
  22. ^ Tan, Keng-Hong; Nishida, R (March 2005). "Synomone or kairomone?--Bulbophyllum apertum flower releases raspberry ketone to attract Bactrocera fruit flies". Journal of Chemical Ecology. 31 (3): 497–507. Bibcode:2005JCEco..31..497K. doi:10.1007/s10886-005-2023-8. PMID 15898497. S2CID 39173699.
  23. ^ Tan, Keng Hong; Tan, Lin Tze; Nishida, Ritsuo (3 November 2006). "Floral Phenylpropanoid Cocktail and Architecture of Bulbophyllum vinaceum Orchid in Attracting Fruit Flies for Pollination". Journal of Chemical Ecology. 32 (11): 2429–2441. Bibcode:2006JCEco..32.2429T. doi:10.1007/s10886-006-9154-4. PMID 17082990. S2CID 15812115.
  24. ^ Tan, K.H. (2009). "Fruit fly pests as pollinators of wild orchids". Orchid Digest. 73 (3): 180–187.
  25. ^ Nakahira, Masataka; Ono, Hajime; Wee, Suk Ling; Tan, Keng Hong; Nishida, Ritsuo (December 2018). "Floral synomone diversification of Bulbophyllum sibling species (Orchidaceae) in attracting fruit fly pollinators". Biochemical Systematics and Ecology. 81: 86–95. Bibcode:2018BioSE..81...86N. doi:10.1016/j.bse.2018.10.002. hdl:2433/235528. S2CID 91933950.
  26. ^ Tan, K. H. 2009. Fruit fly pests as pollinators of wild orchids. Orchid Digest, 73(3): 180-187.
  27. ^ Tan, K.H. and Nishida R, 2015 Pollination of bactrocerophilous Bulbophyllum Orchids. pp. 273-279. Proceedings of the 20th World Orchid Conference, Singapore, 2011. Singapore Botanic Gardens.
  28. ^ Tan, K.H. and Tan, L.T. (2018) Movements of floral parts and roles of the tooth on column wall of Bulbophyllum praetervisum (Orchidaceae) flower for pollination by Dacini fruit flies (Diptera: Tephritidae). Journal of Pollination Ecology, 24(17): 157-163.
  29. ^ Alphabetical List of Standard Abbreviations for Natural and Hybrid Generic Names, Royal Horticultural Society, 2017.
  30. ^ a b c Illustrated Encyclopedia of Orchids ISBN 0-88192-267-6

Bibliography

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  • J.J. Vermeulen : Orchid Monographs Vol. 7 (1993), A taxonomic revision of Bulbophyllum, sections Adelopetalum, Lepanthanthe, Macrouris, Pelma, Peltopus, and Uncifera (Orchidaceae). iv + 324 pp., 25 text-figs. + 116 full-page line drawings, 6 pp. colour plates. ISBN 90-71236-17-X
  • Siegerist E.S.: - Bulbophyllums and their allies Timber Press, 2001, 77 colour photos, 296 pp ISBN 0-88192-506-3 - devoted solely to Bulbophyllums, it is an introductory guide for amateur and advanced orchid growers.
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