Orchids of Bhutan III: The in Bhutan by stig Dalström and dhan bahadur gurung

In The Orchids of Bhutan (Pearce and Cribb 2002), we can read the following: “The genus Epipogium was established (as Epipogum) in 1747 by S. [sic. J.]G. Gmelin in his Flora Siberica [sic. Sibirica]. Since the original description, there have been no less than four different published spellings for the genus, causing a confusion that was reviewed by Sprague and Green (1937). Most authors have since used Epipogium, the earliest use of which was in 1792 by Moriz Balthasar Borkhausen in his Tentamen Dispositionis Plantarum Germaniae.” Pearce and Cribb (2002) also mention that the genus includes 1 2

“about” five species S tig Dalström S tig Dalström with Epipogium aphyllum Sw. other scientifically described “names,” [1] Epipogium aphyllum, myco- reported from a including varieties and forms, have few locations in received the same treatment. parasitic habit in leaf Stig Dalström Bhutan; Epipogium of Epipogium were for a long mold, Högberget, Sweden. roseum (D.Don) time considered to be “saprophytic” [2] Epipogium aphyllum, with Lindl. from Darjeeling and Sikkim, ; (living on dead material), and existing and Epipogium sessanum S.N.Hegde & in a harmonious symbiotic relationship nonresupinate flowers, Hög- A.N. Rao from the West Kameng district with a mycorrhizal fungus. This view berget, Sweden of Arunachal Pradesh, India. An additional has changed in recent times, however, Epipogium taxon was discovered by and a more common view today is that Clarke in the Sind Valley of Kashmir and the orchid really feeds on the fungus span, and therefore have no known ability described as Epipogium tuberosum Duthie and gives nothing back. It appears that to produce their own food. The plants (Duthie 1906), but is listed as a synonym the orchid allows the fungus to enter its seem to be completely dependent on their of E. roseum by Pearce and Cribb, albeit root hairs and to develop tiny ”balls” of a fungus partners and really parasitize them, with a question mark. The “fifth” species thread-like structure known as pelotons in presumably without killing them. There that Pearce and Cribb refer to is most the cells. The pelotons are then digested appears to be no hard evidence that the certainly Epipogium japonicum Makino, by the orchid as an energy source through fungus receives any reward for its role which is not reported from Bhutan, or a process known as mycotrophy (Zettler (Zettler 2005). from the Himalayan region. The World 2005). It is believed that all orchids utilize Due to the development of this Checklist of Selected Plant Families (also a fungus partner during some stage of mycoparasitic habit, the Epipogium plants known as the Kew Monocot Checklist), their lives, particularly when the seed lack leaves and the pale inflorescences on the other hand, lists only three valid germinates and before the seedling has the have an ethereal and “mystical” quality species altogether in Epipogium: E. ability to produce and its own about them wherever they appear, often aphyllum, E. japonicum and E. roseum, energy or food through photosynthesis. in deep shade under large trees. They are placing E. sessanum and E. tuberosum as But some orchid species have taken this also unpredictable in terms of when they synonyms of E. roseum. These considered relationship much further and in a “single- flower. Only when the conditions are synonyms are by no means the only “taxa” minded” way. Species of Epipogium lack favorable will the inflorescences develop. treated as such by Kew. No less than 17 chlorophyll entirely throughout the life It is believed that plants of E. aphyllum

www.AOS.org December 2013 Orchids 735 3 yeltshen G yeltshen N gawang need a mild and humid spring in order also tropical . We are going to [3] Epipogium roseum habititat to flower, which takes place later, during take a closer look at these species, but near Saleng, studied by Stig the summer. If the conditions are less than also question some of the listed Kew perfect, many of the flower buds, which synonyms. Dalström. were initiated the previous fall, will die The second taxon on our list is E. [4] Epipogium roseum, myco- prematurely and the plant will remain roseum, which is mentioned by Sithar hidden. After flowering, parts of the plant Dorji in his field guide to the orchids of parasitic plant habit in leaf die but the coralloid root system can Bhutan (Dorji 2008), but no preserved mold, Saleng, Bhutan. produce long “stolons” where new plants specimens were apparently made. The eventually develop and which may appear author died prematurely the same year and quite a distance from the original plant. no field journals have been discovered that catch, or at least photograph, the busy Sometimes several years pass by before may reveal where he found this species. little visitor, but without success. Before a the beautiful and ghost-like flowers appear Since then, a couple of new populations second attempt could be made, a generous again, and at a different site (Mossberg of E. roseum have been documented by portion of the plant cluster had ended up in and Nilsson 1977). the TDOCP team. In May of 2010 a large the herbarium press due to some seriously Before the Thunder Dragon Orchid population was discovered near the little dutiful efforts by the “herbariumeer” in Conservation Project (TDOCP) was village of Saleng, in the district of Mongar. charge at the time, and no more visiting initiated in 2008 as a collaborative effort Inflorescences were scattered over a rather insects would dare to approach the site. between the team of the first author and wide area, growing in deep shade among In 2006, An Illustrated Guide to the Thomas Höijer of Jakobsberg, Sweden, leaf mold in a mixed semideciduous forest. Orchids of Bhutan was authored by Dhan and the National Biodiversity Centre in A second population was discovered Bahadur Gurung (2006). Much of the Serbithang, Bhutan, only E. aphyllum was in May of 2013, in the Sakten Wildlife in this little handbook follows reported from Bhutan by Pearce and Cribb Sanctuary of Trashigang. A large bundle the 2002 treatment by Pearce and Cribb, (2002). Since then two more taxa have of plants was growing on and around a with a few exceptions. One such exception been added through the TDOCP, which rotten tree stump, where also a fungus had concerns a plant that appears at the end correspond to Kew’s number of accepted produced a rich amount of fruiting bodies of the field guide, under “undetermined species. Rather amazingly they represent (). The identity of the fungus, species” on page 165. This plant was all accepted species of a genus that is which is suspected to be the fungal partner tentatively identified as Epipogium aff. distributed from the extreme western parts for the orchid, is uncertain at this time. roseum by Gurung, and was discovered of , throughout temperate all The identity of a bee-like insect that in flower on October 10, 1999, along the way to and Korea, including the appeared to be pollinating the flowers the Chuselumpa mountain ridge near , and possibly, but doubtfully, is also unknown. Efforts were made to Hetsho Thangka. This ridge is located

736 Orchids December 2013 www.AOS.org flower of E. sessanum is described as lacking a spur altogether. This eliminates this taxon from the equation and we obviously need to expand our search in order to find a positive identification for the Chuselumpa plant. Epipogium tuberosum Duthie was described in the Annals of the Royal Botanic Garden, Calcutta (Duthie 1906), and is based on a plant collected by C.B. Clarke (No. 31230) at 6,000 feet (ca. 2,000 m) in the Sind Valley of Kashmir. The description reads as follows: “Whole plant 1 to 3 dm. high, glabrous. Root tuberous, not coralloid. Scape usually swollen above the base and tapering upwards, bearing a few membranous broad-based bracts. Raceme laxly 3–5 flowered, floral bract ovate-lanceolate, acuminate, entire, longer than the slenderly stalked subglobose , 5-nerved. and 8 mm. long, broader than those of E. aphyllum, all 3-nerved. Lip without side lobes, 9 mm. long, its upper surface with three carunculate ridges, margins not erose. Spur as in E. aphyllum, but more slender and with a less rounded apex. Stigma occupying a deep cavity on the anterior face of the column. Pollinia globose, with caudicles equal to them in length, glands minute globular.” Then follows: “Very similar to the preceding [E. aphyllum, authors’ note] in general appearance, but specifically distinct by reason of its tuberous non- coralloid root, its entire lip and by the very different pollinia and column.” 4 The description of E. tuberosum fits tig Dalström S tig Dalström the Chuselumpa plant very well as can be a few kilometers southeast of the city of with and slightly curved toward the ovary. seen in Gurung’s photo, and we have little Wangdue in central Bhutan, toward the The Chuselumpa plant is therefore clearly doubt that it is an accurate identification Tsirang highway and east of the river not the same as Epipogium roseum, which of a distinct species. The only question Punatsang Chhu. The plant was growing is illustrated with a line drawing in The that remains to be answered is whether among decaying wood chips in a stand of Orchids of Bhutan (Pearce & Cribb 2002). any other listed name also correlates with Cupressus corneyana, at an elevation of When we compare the Chuselumpa plant this species and may have been described about 7,800 feet (2,600 m). This particular with E. aphyllum on the other hand, earlier, thus having nomenclatural area is dominated by Cupressus forest and we can see that both have flowers with priority? with oaks as codominants. forward-curved spurs, but in the case of In 1904, two years prior to the The photo in the field guide shows a E. aphyllum the flower is nonresupinate description of E. tuberosum, Tomitaro pale inflorescence with a few resupinate (lip uppermost), and the lip has distinct Makino described Epipogium japonicum (with the lip lowermost) flowers and some side lobes, whereas the Chuselumpa plant based on a plant collected by Viscount additional buds near the apex. Although has a cordate-ovate lip without distinct Nobumitsu Aoki in a shady forest at few morphological details can be observed side lobes. Another significant difference the eastern foot of Mount Nyohō, in the in the photo, one particular feature stands can be observed underground. Epipogium province of Shimotsuke, Japan (Makino out that helps us along the way toward a aphyllum has an irregular and coral- 1904). Although no original plant material positive identification. At least it clearly shaped whereas the Chuselumpa or illustrations have been seen by us, the tells us what it is not. The large and plant has a thick and ovoid tuber, similar overall description and the mentioning of curved spur near the base of the lip is bent to E. roseum. the “patent-nutant” (nodding) flowers with forward and toward the lamina of the lip. Our conclusion is that the Chuselumpa a descending (as opposed to ascending) Epipogium roseum has a large spur that plant represents something different from spur distinguish this taxon from both E. points in the other direction and is parallel both E. aphyllum and E. roseum. The aphyllum and E. roseum, and corresponds

www.AOS.org December 2013 Orchids 737 738 aasa and aeg itit f rnca Pradesh Arunachal of West the district in Kameng Tipi from reported also is 1992). Wood and in and 276), plate 2010, Sinicae Popularis Republicae of Blume, as this species is illustrated in looks plant very the similar to however, illustration, type the from Judging treatment. specific separate a justifying significance taxonomic have can well must very feature we this that then conclude spur, a without flowers have plants both if And clones. separate two represent they that here assumed is it roseum to (similar rhizome coralloid divided a individual than rather tubers separate have Since plants morphology. with assumingly similar found, was plant one than more that suggests it observed, were (plural) “plants” that mention Since authors the base. the near side each on angle minor a for except sepals, the resembling and callus visible a without “primitive,” The spur. general shape a of the lip also appears lacks rather that lip dissected a with a flower the shows description accompanies original that illustration The Stereosandra than rather characters genus anther and column the to belonged they that found was it critically, further studied were specimens] [sic. specimen Rao wecanreadthefollowing: and Hegde by description original the In interesting taxon that needs further scrutiny. at thistime. from each other, but that is unknown to us taxa separated geographically but similar very these distinguish that exist features over priority has name this first, described was Because plant. Chuselumpa the like much very looks illustration this and 278), plate 2010, Sinicae Popularis Republicae Flora of Committee (Editorial published recently the in as identified is well with morphological features.” of lip the unlike lip the of spur lacked specimen the that resembling found was it observation close Upon roseum plants, Epipogium few saprophytic a Kameng Pradesh, Arunachal West district, Sessa, Sanctuary, Orchid at inspection field a “During ecember D Orchids hn olw, Hwvr we the when “However, follows, Then ioim sessanum Epipogium E. tuberosum n dfeet from different and Eioil omte o Flora of Committee (Editorial E. tuberosum Epipogium ” (HegdeandRao1982). Stereosandra h Ocis f Peninsular of Orchids The . japonicum E. Singapore . It is possible that some eesnr javanica Stereosandra Stereosandra javanica ee noticed. were . A drawing of what 2013 but simulating but fnly i an is finally, , lora of Flora n t gross its in (Seidenfaden Epipogium aphyllum E. . japonicum E. can be seen be can www.AOS.org China Flora in E. ),

tig Dalström et o i fr xmnto blns to belongs examination for him to sent sessanum E. of publication original the in Rasmussen from how clue to a perhaps phenomenon interesting the lacks and typical petals the than broader ( specimen the of labellum “the publication: original the in quoted then is Rasmussen 1982). to belonged flower) (one him to sent specimen determined a that however, Rasmussen, Finn anther characteristics,isunconvincing. and column unspecified to referring Rao, and Hegde of by mentioned differentiation congeneric members least at as or conspecific, as treated be strongly should taxa these that that suggest similarities morphologic sessanum is of illustration comparison type the between critical made a Cribb and When (Pearce 2002). Rao and Hegde by hr i n epaain rvdd by provided explanation no is There Epipogium Epipogium Stereosandra n of and ge s.n Hegde f h te lwr ht was that flower the why of roseum E. likeancestors.” pr hs s highly a is This spur. javanica S. could have evolved have could Stereosandra ) s ol) slightly (only) is .) Hge n Rao and (Hegde i reveals it , The . E.

tig Dalström [5] [6] Epipogium roseum Epipogium roseum Bhutan. supinate flowers, Joengkhar, stump, Joengkhar, Bhutan. parasitic habitonoldtree , withre , myco- - 6 5 E. roseum despite the fact that the type plant (Hegde s.n.) demonstrates quite different morphological features, unless specimens of the two species were mixed up somehow. There is a drawing of a single flower of E. roseum included in the type illustration of E. sessanum for comparison, and it seems plausible that a flower of E. roseum may have been sent to Rasmussen for examination by mistake, instead of the intended flower of E. sessanum. Rasmussen’s quote would then refer to the actual type specimen and illustration of what became E. sessanum. In any case, based on the described diagnostic features and what can be seen in the type illustration, we believe that E. sessanum belongs in Stereosandra rather than in Epipogium. In summary, we acknowledge Kew’s listing of E. aphyllum, E. japonicum and E. roseum as valid species of the genus but disagree with placing E. sessanum and E. tuberosum as synonyms of E. roseum. 7 Acknowledgments stig Dalström The authors wish to thank the Government of Bhutan and the Ministry of Agriculture and Forests, and the administration and staff of the National Biodiversity Centre, Serbithang, Thimphu. We also thank Rudolf Jenny for generously providing copies of many of the original species descriptions and Wesley Higgins for viewing and commenting on the manuscript. Special thank yous to Sharon and Russell Stephens of Sarasota, the Friends of Orchid Research Fund (grant number 20132219, administrated by the Community Foundation of Sarasota County), and the Sarasota Orchid Society for funding. References Dorji, S. 2008. The Field Guide to the Orchids of Bhutan. Bhutan Orchid Science Society, Thimphu, Bhutan. Duthie. J. F. 1906. The Orchids of the North-Western Himalaya; Epipogium tuberosum. Ann. Roy. Bot. Gard. (Calcutta) 9(2):151. 8 stig Dalström 9 Editorial Committee of Flora Republicae Popularis Sinicae. Dhan B aha d ur G urung 2010. Flora of China; Illustrations. Volume 25. Science gium. Kew Bull. (9):475–476. Press, Beijing, China and Missouri Botanical Garden Zettler, L.W. 2005. Nature’s Fungal Connoisseurs. New [7] Dhan Bahadur Gurung, au- Press, St. Louis. Insight into the Mysterious Orchid-Fungal Association. thor of An Illustrated Guide Gurung, D.B. 2006. An Illustrated Guide to the Orchids of Orchids 74(4):292–297. Bhutan. DSB Publication, Thimphu, Bhutan. To The Orchids Of Bhutan. Hegde, S.N. and A.N. Rao. 1982. Epipogium sessanum Hegde et Rao — A New Species of Orchid from Stig Dalström is a research associate [8] Epipogium roseum, intimate Arunachal Pradesh, India. J. Econ. Taxon. Bot. 3: at Lankester Botanical Garden, University 597–601. of Costa Rica, Cartago, Costa Rica, and myco-parasitic habit on Makino, T. 1904. Epipogium japonicum Makino, Observa- tions on the Flora of Japan. Bot. Mag. (Tokyo) 18:131. National Biodiversity Centre, Serbithang, old tree stump, Joengkhar, Mossberg, B. and S. Nilsson. 1977. Nordens Orkidéer. Thimphu, Bhutan. 2304 Ringling Boulevard, Bhutan. Note the tiny fungal Wahlström and Widstrand, Stockholm, Sweden Unit 119, Sarasota, Florida 34237. (email Pearce, N.R. and P.J. Cribb (2002). The Orchids of Bhutan. [email protected]; website www. fruiting bodies. Royal Botanic Garden, Edinburgh, and Royal Govern- ment of Bhutan. wildorchidman.com). [9] Epipogium japonicum, inflo- Seidenfaden. G. and J.J. Wood. 1992. The Orchids of Peninsular Malaysia and Singapore. A Revision of rescence of the plant from R. E. Holtum: Orchids of Malaya. Olsen and Olsen, Chuselumpa, Bhutan. Fredensborg. Denmark. Sprague, T.A. and M.L. Green. 1937. Epipogum or Epipo-

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