Orchids of Bhutan The Genus Phaius 484 ORCHIDS AUGUST 2015 WWW.AOS.ORG 1 2 STIG DALSTRÖM Plants of the genus Phaius loureiro, have been a favorite of the first author for many years. It was one of the very first orchids that he acquired in the early 1970s from a small backwoods nursery in sweden. Perhaps this nursery was the last place where you would expect to find tropical orchids in those days, but the owner was an old-timer with “green thumbs” and a keen interest in “different” and exotic plants. he had therefore accumulated a variety of rather unusual plants over the years. he also had a bench full of healthy looking Coelogyne cristata lindl., which were divisions from a single plant obtained some time in a misty past. one of these divisions went home to a budding windowsill collection together with the Phaius, and they founded an interest in himalayan orchids that has never diminished. It actually took some time to figure out that it really was a Phaius that had been purchased. But with the help of a classic swedish orchid book by axel holzhausen (1929), it was soon concluded that the correct name probably was Phaius tankervilleae (Banks ex l’héritier) Bl. the recommended culture for this unusual looking orchid was to grow it in a warm place, where the heat should come from underneath. hmm? how was this to be accomplished? the problem was solved by placing the large pot on top of a refrigerator that in turn was placed near a window. the light would then reach the large leaves, and the heat at the top of the refrigerator would warm up the medium in the pot. It worked! the plant produced a strong new growth that eventually produced a tall spike with some awesomely beautiful flowers. It was a spellbinding experience for a young orchid novice! there are four species of Phaius known from Bhutan. In May of 2014 a team from the national Biodiversity Centre in serbithang, consisting of Choki Gyeltshen, nima Gyeltshen, Dupchu Wangdi and the first author, set out to find them all and if possible, bring back live plants for the rapidly developing orchidarium in the Royal Botanical Garden. We had great help from the second author of this article with finding STIG DALSTRÖM STIG DALSTRÖM, DHAN BAHADUR GURUNG, CHOKI GYELTSHEN AND NIMA GYELTSHEN STIG DALSTRÖM 3 DHAN BADAHUR GURUNG some of the localities since he had seen them all while preparing his field guide to the orchids in Bhutan (Gurung 2006). the expedition was successful and healthy clumps of all four species were brought back to serbithang. finding the plants was a small challenge compared to figuring out their accurate nomenclature. as many of us orchid enthusiasts are painfully aware of by now, orchid taxonomy can be a real brain teaser. and the nomenclatural maze of Phaius is no exception. Phaius flavus (Blume) lindl. was originally described as Limodorum flavum by Karl ludwig von Blume (1825) based on a plant from Java. John lindley (1831) transferred it to Phaius in his Genera and Species of Orchidaceous Plants, on the subsequent page after describing the very same species as Phaius maculatus lindl. the type specimen for this latter taxon was based on a plant obtained by nathanial Wallich in nepal. this attractive and very distinctive orchid was then again described up to 17 times under different names by various authors (seidenfaden 1986, World Checklist of selected Plant families 2015). let us not pursue this nomenclatural quagmire any further here, but rather look at what this species looks like in Bhutan. Phaius flavus is a very stately plant that can reach more than 3–3.5 feet (1 m) in height. It often grows in rather shady and humid locations with the fleshy root system buried deeply in humus-rich leaf mulch. the plants are easily distinguished by the whitish spots on the large leaves, which appear to be genetically inherited. Virtually identical spot patterns can be seen in all the Bhutanese Phaius species, but in those cases the patterns are most likely caused by fungal (or bacterial) infections, and the spots can be present on plants in nature as well as in cultivation. Phaius flavus grows in warmer to intermediate conditions in Bhutan, in tropical to subtropical, seasonally extremely wet areas. But with an area of distribution that covers much of southeastern asia, a certain tolerance to differing growing conditions can be expected, as well as a morphological variability. this should be true of all four species included here. Phaius mishmensis (lindl. & Paxton) Rchb.f. was originally described as Limatodes mishmensis by John lindley and Joseph Paxton, based on a plant collected by William Griffith on the Mishmee hills in assam, India (lindley and Paxton 1852). It was then transferred to Phaius by Reichenbach with a simple note: “obs. Limatodes mishmensis 4 lindl. = Phajus mishmensis Rchb. fil.” (Reichenbach 1857). this species has “only” been redescribed five to seven times (seidenfaden 1986; WCsPf 2015). like Phaius flavus, Phaius mishmensis is a rather distinctive orchid that is easily recognized by the tall stems that can reach well over 3 feet (1 m) in height and where the rose-purple flowers emerge from the side of the stem on erect few-flowered racemes. It is an attractive species in a modest way. It grows under similar conditions as Phaius flavus in Bhutan, and they are sympatric in some areas, [1] Phaius flavus in situ, Gomphu, eastern Bhutan. The inset is a close-up of the naturally spotted leaves of this species. This moderately sized species inhabits the humid, shady forest floor over a very wide geographic region. In this habitat, the plants are nearly constantly moist. [2] Phaius flavus in close-up, Gomphu. Flowers of this species are about 3 inches (7.5 cm) across. Photograph taken in situ. [3] Phaius mishmensis taken in situ with Choki Gyeltshen, Rimchu, Bhutan. WWW.AOS.ORG AUGUST 2015 ORCHIDS 485 486 ORCHIDS AUGUST 2015 WWW.AOS.ORG 6 STIG DALSTRÖM 5 STIG DALSTRÖM growing side by side. It can be difficult to distinguish smaller plants of the two without flowers because they both can have spotted leaves, but for different reasons as mentioned above. Phaius tankervilleae represents the third species documented from Bhutan, and constitutes a real taxonomic challenge. It was originally described in Sertum Anglicum by the french botanist Charles l’heritier de Brutelle (1789) as Limodorum “tancarvilleae” based on a Chinese plant brought to england by John fothergill around 1778 (Brutelle 1789, seidenfaden 1986). the description by Brutelle was based on an engraving in the Joseph Banks Museum (sims 1817), and for this reason the authorship is usually attributed to Banks (seidenfaden 1986). Banks’ illustration was not included in the original description, but a plate prepared by the english naturalist and illustrator James sowerby under the name “Limodorum tankervilliae” is included in the third fascicle of William aiton’s Hortus Kewensis, which was published later the same year. the brief description that accompanies the illustration refers to the plant introduced by John fothergill in “about 1778” (aiton 1789). sowerby’s great illustration depicts well the plant that today is commonly known as Phaius tankervilleae. this species was subsequently redescribed at least 22– 25 times (seidenfaden 1986, WCsPf 2015) under different names by various authors. Limodorum “tankervilliae” was transferred to Phaius by Blume (1856), where it remains today. the correct spelling, as tankervilleae, honors the Right honourable Countess of tankerville, and was clarified by Gunnar seidenfaden (1986). Phaius tankervilleae has had a wide nomenclatural journey with a plethora of names of various spellings through the years, which we will not delve into too deeply here, except for two cases. let us look at these before returning to Phaius tankervilleae. Phaius grandifolius loureiro was described based on a plant that, according to the Portuguese author Joao de loureiro, was cultivated in China and Cochinchina (Vietnam) at the time, and the description was published in his Flora Cochinchinensis (loureiro 1790). nine years later olof swartz, the swedish botanist and pupil of Carolus linnaeus, listed “Phayus” grandifolius loureir. Cochin. 646 as a synonym of Limodorum “tankervilliae” (swartz 1799). What he based this conclusion on is unknown to us. Perhaps swartz had a chance to examine loureiro’s type specimen, wherever it is deposited (we have not seen it). In any case, lindley accepted swartz’s opinion of treating Limodorum tankervilleae and Phaius grandifolius as conspecific, but he reversed the nomenclatural priority and listed the former as a synonym of the latter (lindley 1831). the conclusion to synonymize these two names has not been shared by all authors over the years. Blume accepted them as separate species and transferred Limodorum tankervilleae to Phaius as a separate species (Blume 1856). the current opinion, however, is that Phaius grandifolius should be listed as a synonym of Phaius tankervilleae (seidenfaden 1986). We will follow this opinion here, but with a question mark, which refers to the following species. Phaius wallichii lindl. was based on a plant collected by M.R. (Mr.?) smith in sylhet, Bangladesh. It was described by lindley in nathanial Wallich’s Plantae Asiaticae Rariores (lindley in Wallich 1831), which features a good colored illustration of the plant. the flowers have a rather long and distinctly curved spur. this is a key feature to the identification of this species in Bhutan, as apart from Phaius tankervilleae, which has a very short and rather straight spur. sometimes, however, flowers from other areas labeled Phaius tankervilleae can be seen with a curved spur as well (sarawak, see Beaman et al. 2001; sumatra, see Comber 2001). But at least in the latter case, it seems that the depicted flowers really represent Phaius wallichii instead. lindley compares Phaius wallichii with Phaius grandifolius/tankervilleae and distinguishes it by “…its very acuminate bractea; sepals, and petals, and especially in the labellum not only tapering to a very acute point, as in Brassavola nodosa, but being destitute of the rich yellow and red marking of the Chinese species” (lindley in Wallich 1831). In other words, lindley separates Phaius wallichii from Phaius grandifolius/tankervilleae (which he had synonymized) mostly based on the shape and coloration of the sepals, petals and lip. By looking at the illustration in the original publication we know what Phaius wallichii looks like. We also know what Phaius tankervilleae looks like, and that they are rather different, particularly in the shape of the flower spur, but also in the coloration and shape of the lip. But we do not really know what the type of Phaius grandifolius looks like! the case gets a twist in Jean and lucien linden’s Lindenia, volume 4, plate 188 (linden and linden 1888), where a beautiful 7 illustration of what appears to be Phaius wallichii is featured with the name Phaius grandifolius. this raises the question whether the original Phaius grandifolius may actually be the true and oldest name for Phaius wallichii instead of being a synonym of Phaius tankervilleae? Without a close examination of loureiro’s holotype of Phaius grandifolius from “Cochinchina” it is not possible to say. What adds weight to this speculation is the fact that both Phaius tankervilleae and Phaius wallichii have been documented from China (the latter as Phaius magniflorus Z.h.tsi & s.C.Chen, in Chen et al. 1999) and from Vietnam (Cribb et al. 2004). and because both species occur throughout much of [4] Phaius mishmensis in close-up, Rimchu. The brightly colored 1 ¾ –2-inch (4.4–5-cm) flowers of this species often do not open fully. The inflorescences of this species are few-flowered and typically less than 2 feet (60 cm) tall. [5] Phaius tankervilleae taken in situ, Thinleygang, Bhutan. Thinleygang village in western Bhutan lies at about 6,000 feet (1,850 m). [6] Phaius tankervilleae in close-up, Thinleygang. [7] The type illustration of Phaius wallichii published in Wallich’s Plantae Asiaticae Rariores. WWW.AOS.ORG AUGUST 2015 ORCHIDS 487 488 ORCHIDS AUGUST 2015 WWW.AOS.ORG 8 9 DHAN BADAHUR GURUNG southeastern asia (Cribb et al, 2004) and are closely sympatric in Bhutan, this line of thinking is worth considering. loureiro, in the original description of Phaius grandifolius (sometimes accidently cited as “grandiflorus”; Cribb et al. 2004), mentions that the species was cultivated in China and “Cochinchina” (Vietnam) as early as 1790. this correlates with what we learn from Cribb et al. (2004), that Phaius wallichii, locally known as the “Kadoorie” Phaius since it was first reported (in recent times) from the Kadoorie farm and Botanic Garden in the new territories, hong Kong, which in turn was established in 1956 by the Kadoorie agricultural aid association (Wikipedia 2015). In reality, Phaius wallichii has been known in China for much longer than that. It was collected by Barretto (no. 146) in hong Kong and was subsequently cultivated at the Royal Botanic Garden at Kew in May 1888 (Cribb et al. 2004). During a meeting of the Royal horticultural society at the Drill hall in london on May 3, 1892, frederik sander exhibited a plant of what was described as “Phaius sanderii,” which received a first Class Certificate by the orchid Committee (hort 1892). the type specimen is deposited at Kew, cited as coming from hong Kong and identified as P. wallichii (Cribb et al. 2004). Joseph Dalton hooker (1890) includes Phaius wallichii as a separate species in The Flora of British India, where he adds at the end of the description: “Possibly as form of the Chinese P. grandifolius, lour.” hooker adds weight to this line of thoughts four years later when he, what appears to be somewhat skeptically, comments on how Veitch distinguished Phaius wallichii from Phaius grandifolius (hooker 1894b). Veitch, however, accepts Phaius grandifolius as a separate name and lists Limodorum “tankervilliae” as a synonym, following lindley on this matter (Veitch and sons 1890). Veitch also lists Phaius bicolor lindl. as a synonym of Phaius wallichii, which we can agree with after viewing the beautiful illustration of the former in lindley’s Sertum Orchidaceum (lindley 1838). the length and shape of the spur is very distinct. But we still don’t know what the original Phaius grandifolius really looked like and where the specimen is deposited, if it still exists. apparently seidenfaden looked for it, but could not find it at the British Museum where other loureiro specimens are placed (seidenfaden 1986). Without a close examination of loureiro’s type specimen of Phaius grandifolius, it is doubtful that a definite decision can be STIG DALSTRÖM 10 NIMA GYELTSHEN [8] Phaius wallichii identified as Phaius grandifolius in Lindenia 4, plate 188. [9] A close-up of Phaius wallichii taken in situ at Thinleygang, Bhutan. Plants of Phaius wallichii are much larger than those of Phaius tankervilleae. The pseudobulbs are up to 4 ¾ inches (12 cm) tall, the inflorescences can reach 6 feet (1.8 m) tall and the nearly 6 inch (15 cm) flowers face forward in contrast to the downward-facing flowers of Phaius tankervilleae. [10] Phaius tankervilleae (or possibly a natural hybrid with Phaius wallichii) taken in situ in Saleng, Bhutan. [11] A close-up of the flower of the plant illustrated in [10] taken in cultivation in Serbithang, Bhutan. 11 WWW.AOS.ORG AUGUST 2015 ORCHIDS 489 made of whether it really is the same as Phaius tankervilleae, or possibly Phaius wallichii. Regardless of this conundrum, it seems clear beyond any doubt that in Bhutan Phaius wallichii is a distinct species in its typical form and separate from Phaius tankervilleae. the difference in flower morphology and appearance in general and the spur shape in particular readily separate the two. the plants are sympatric in Bhutan and occur very closely together in the same habitat near the village of thinleygang. We were not able to see both species in flower in May of 2014, only Phaius tankervilleae, but the second author of this article has, on previous visits. In their typical forms, these species are quite distinctive. there is a possibility that they may hybridize naturally, however, while occurring so close to each other and evidence exists that may support this. a Phaius plant of extraordinary vigor was collected near the village of saleng in 2010 and brought back to serbithang. It flowered for the first time in 2014 and revealed flowers suspiciously intermediate between Phaius tankervilleae and Phaius wallichii. In any case, and not to bring any more confusion into this particular subject, we have now discussed all four known Phaius species of Bhutan. But there is one more taxon that needs to be discussed here. Phaius nanus hook. is included in The Orchids of Bhutan by Pearce and Cribb (2002) because it comes from a neighboring area and can be expected to eventually be found in Bhutan. It was originally described by hooker (1892), based on a plant collected by James sykes Gamble (no. 6672) “in the Buxa Doar” of West Bengal, India. the holotype is at Kew and consists of two poor specimens mounted on sheet k000829050. In the original description by hooker, nothing is mentioned about the plant lacking a pseudobulb, even though no visible pseudobulbs can be seen on the type sheet. this may have led Pearce and Cribb to describe Phaius nanus as “lacking pseudobulbs” (Pearce and Cribb 2002). But in the illustration accompanying the original description a rather typical pseudobulb is illustrated in a horizontal position at the base of the flowering growth. Unfortunately, no details of a possible lip spur can be recognized from the way the flowers are depicted, or in the description. two years later, hooker described the horizontally depicted pseudobulb of Phaius nanus as “rootstock stout, with grass-like sheaths, 490 ORCHIDS AUGUST 2015 WWW.AOS.ORG 12 sometimes much longer than the leaf” (hooker 1894a).” hooker continues: “a very singular species, of which I have seen but two specimens, kindly lent from the Calcutta herbarium by Dr. King” (hooker 1894a). Phaius nanus appears again four years later in The Orchids of the SikkimHimalaya, Part 3, published in the Annals of the Royal Botanic Garden of Calcutta by George King and Robert Pantling (1898). this time the illustration clearly shows flowers that are virtually identical to Phaius tankervilleae, having a short and rather straight spur. this illustration is very similar to the original drawing published by hooker six years earlier and most likely made from the same specimen. It seems that one illustrator copied another because even though the motif is similar the styles are very different. the main difference is how the flowers are shown in King and Pantling’s version, with short and straight spurs. It is also possible to interpret the “rootstock” as some kind of tuber, or thickened rhizome, and not a true pseudobulb. But because Phaius species do not have tubers it seems rather clear that the “clump” at the base of the new growth really is a true pseudobulb. It is therefore our conclusion that Phaius nanus probably represents a smaller (younger?) plant of Phaius tankervilleae, perhaps stunted by growing in bright light or simply collected when beginning to flower. acknowledgments the authors cordially thank the Royal Government of Bhutan, the Ministry of Works and human settlements and the Ministry of agriculture and forests for providing necessary permits. a special thank you to Ugyen tshewang, secretary of the national environment Commission, aiton, W. 1789. Limodorum tankervilliae. Hort. Kew. 3:302. Beaman, t., J.J. Wood, R.s. Beaman, and J.h. Beaman. 2001. Orchids of Sarawak. natural history Publications, (Borneo) Kota Kinabalu, in association with Royal Botanic Gardens, Kew, UK. Blume, K.l von. 1825. Limodorum flavum. Bijdr. Fl. Ned. Ind. 8:375. ____. 1856. Phaius tankervilleae (as “tankervillii”). Mus. Bot. Lugd. 2(12):177. Brutelle, C. l’h. 1789. Limodorum tancarvilleae. Sertum Anglicum. p. 28. Petri-francisci Didot, Paris, france. Chen, s., Z. tsi, and Y. luo. 1999. Native Orchids of China in Colour. science Press, Beijing, China and new York. Comber, J.B. 2001. Orchids of Sumatra. the Royal Botanic Gardens, Kew, UK. Cribb. P., M. sun, and G. Barretto. 2004. Phaius tankervilleae and P. wallichii (orchidaceae), a pair of confused species. Kew Bull. 59:547–554. Gurung, D.B. 2006. An Illustrated Guide to the Orchids of Bhutan. DsB Publication, thimphu, Bhutan. holzhausen, a. 1929. Orchidéer: Deras Förekomst, Odlingshistoria och Skötsel. albert Bonniers förlag, sweden. hooker, J.D. 1890. Phaius wallichii. The Flora of British India. Volume 5. l. Reeve & Co., london, UK. p. 816. _. 1892. Phaius nanus. plate 2088, In: D. oliver, editor. Icones Plantarum. Volume 21. Dulay & Co., london, UK. _. 1894a. Phaius nanus. p. 192. In: The Flora of British India. Volume 6. l. Reeve & Co., london, UK. _. 1894b. Phaius wallichii. p. 191. In: The Flora of British India. Volume 6. l. Reeve & Co., london, UK. hort. 1892. “Phaius sanderii”. Gard. Chron. 3:598. King, G. and R. Pantling. 1898. the orchids of the sikkim-himalaya, Part 3. Ann. Roy. Bot. Gard. Cal. 8. london, england linden. J. and l. linden. 1888. Phaius grandifolius. Lindenia Volume 4. p. 91, pl. 188. l. linden, Brussels, Belgium. lindley, J. 1831. Phaius (as “Phajus”). The Genera and Species of Orchidacious Plants. Ridgways, london, UK. p. 126–128. _. 1838. Phaius bicolor. Sertum Orchidacearum. Ridgways, london, UK. pl. 23. lindley, J. and J. Paxton. 1852. Limatodis mishmensis lindl. & Paxton. Paxton’s Fl. Gard. 3:36. loureiro, J. 1790. Phaius grandifolius. Flora Cochinchinensis 2:529-530. Pearce, n.R. and P.J. Cribb (2002). The Orchids of Bhutan. Royal Botanic Garden, edinburgh, UK, and Royal Government of Bhutan. Reichenbach, h.G. 1857. Phaius mishmensis. Bonplandia (hannover) 5:43 seidenfaden, G. 1986. orchid Genera in thailand 13. STIG DALSTRÖM for generous support in locating proper lodgings throughout the country, and to the administration and staff of the national Biodiversity Centre, serbithang, thimphu, for overall support. We sincerely thank all the forest rangers in general, and Yeshi Dorji, the station chief in tingtibi in particular, who always go out of their way to be helpful and courteous. and finally, we thank Malli and Vera lee Rao and the sarasota orchid society and all its members, as well as Peter Göttfert of orchidéhuset in sweden for generous financial support, and the Bhutan foundation for facilitating the donations. an ultimate thank you goes to Wesley higgins for viewing and commenting on the manuscript. References 13 14 Opera Bot. 89. sims, J. 1817. Bletia tankervilleae. Curtis’s Bot. Mag. 44:1924. swartz, o.P. 1799. Phayus grandifolius. Dianome Epidendri Generis Linn. — Nov. Act. Reg. Soc. Sci. Upsalensis 6:79. Veitch, J. and sons. 1890. Phaius. p. 10–17. In: A Manual of Orchidaceous plants, Part 5. James Veitch & sons, Royal exotic nursery, , Chelsea, UK. Wallich, n. 1831. Phaius wallichii. Plate 158. In: Plantae Asiaticae Rariores 2. treuttel and Würtz, treuttel Jun. and Richter, soho square, london, UK. Wikipedia 2015. https://en.wikipedia.org/wiki/Kadoorie_ farm_and_Botanic_Garden. accessed June 18, 2015. World Checklist of selected Plant families 2015. http:// apps.kew.org/wcsp/synonomy.do?name_id=151000. accessed June 18, 2015. —Stig Dalström (corresponding author) is a research associate at Lankester Botanical Garden, University of Costa Rica, Cartago, Costa Rica and the National Biodiversity Centre, Serbithang, Thimphu, Bhutan. 2304 Ringling Boulevard, unit 119, Sarasota FL 34237, USA (email: stigdalstrom@gmail.com; website: wildorchidman.com). Dhan Bahadur Gurung, PhD is an Assistant Professor and the Dean of the College of Natural Resources, Royal University of Bhutan (email: dbg2006@gmail.com). Choki Gyeltshen is a Biodiversity Officer at the 15 [12] Phaius wallichii identified as Phaius bicolor in Sertum Orchidaceum, plate 23. [13] The village of Thinleygang preparing for the summer rains. [14] Phaius nanus as illustrated in King and Pantling’s The orchids of the Sikkim-Himalaya, part 3, plate 148. [15] The original illustration of Phaius nanus from Hooker’s Icones Plantarum 21, plate 2088. National Biodiversity Centre, Ministry of Agriculture and Forests, Serbithang, Thimphu, Bhutan (email: chokig@ gmail.com). Nima Gyeltshen is the Biodiversity Supervisor at the Royal Botanical Garden, National Biodiversity Centre, Ministry of Agriculture and Forests, Serbithang, Thimphu, Bhutan (email: nimss409@gmail.com). WWW.AOS.ORG AUGUST 2015 ORCHIDS 491