Orchids of Bhutan
The Genus Vanda
STIG DALSTRÖM, DHAN BAHADUR GURUNG, CHOKI GYELTSHEN AND NIMA GYELTSHEN
1
The genus Vanda Jones ex R.Br., in
Bhutan is currently represented by a handful
of what must be considered as “disputed”
species where the status of all taxa have
been questioned in both past and present
by various authors for various reasons. The
generic name Vanda was proposed in 1795
by William Jones in asiatic Researches
but it was not linked to a specific epithet.
Robert Brown (1820) formally established
the genus in the Botanical Register when
he described Vanda roxburghii, the plant
from east Bengal that Jones originally
had in mind (Pearce and Cribb 2002).
The actual name “Vanda” is from sanskrit
and refers to certain parasitic mistletoes
(Loranthaceae) and some orchids, including
Vanda roxburghii, of similar habit (Pearce
and Cribb 2002). up to 2012, some 50
species were added to the genus, which
is distributed from sri Lanka and India in
the west to China, the Malay Archipelago,
the Philippines, Australia, new guinea
and the solomon Island in the southeast.
Leslie garay (1986) removed Vanda alpina
and Vanda griffithii to establish the genus
Trudelia together with a new species;
Trudelia chlorosantha garay. he had
received photos of what he considered to
represent three different species from the
swiss photographer niklaus Trudel, who
34 ORCHIDS JANUARY 2015
WWW.AOS.ORG
apparently collected all three specimens in
southwest Bhutan. garay did not include
Vanda cristata Lindl. or Vanda pumila
hook. f., due to a lack of information,
but senghas (1988a) added those species
to Trudelia. Later, Christenson (1992)
moved them all back into Vanda again.
Recent molecular analysis demonstrates
that Trudelia is embedded in the main
bulk of Vanda species and cannot be
accepted as a separate genus according
to modern nomenclatural rules (gardiner
2012). The same molecular analysis also
demonstrated that many other species in
previously distinct genera also could be
transferred to Vanda, which was executed
by gardiner (2012). Although the transfer
of all species of genus ascocentrum schltr.
to Vanda may seem acceptable based on
similarities in vegetative plant morphology,
such as conduplicate (folded) leaves, the
species Vanda (ascocentrum) himalaicum
(Deb, sengupta & Malick) L.M.gardiner,
in contrast, differs significantly in general
morphology from all the others in the
genus by having pendent rattail leaves.
We therefore suspect that this orchid
may not have been DnA sequenced at all
due to its rarity, and transferred based on
“nomenclatural features” only, which is
risky and can be highly misleading. We
suggest that, in future botanical treatments
and transfers, a note is added for each
species as a standard procedure, describing
whether it has been DnA sequenced or
not.
Va n d a a m p u l l a c e a ( R o x b . )
L.M.gardiner is one of the recent additions
to the genus, which was transferred by
gardiner from ascocentrum based on
molecular analysis. It grows epiphytically
and usually very exposed to bright light along
the seasonally (June through september)
very hot and humid southern border
with India. This species has a “mottled”
nomenclatural history like so many other
orchids. It was originally described as
aerides ampullacea by Roxburgh (1832).
It was transferred to Saccolabium Blume by
Lindley (1838). Kuntze (1891) moved it to
Gastrochilus D.Don, and schlechter (1913)
moved it to ascocentrum. We can only hope
that the current nomenclatural position in
Vanda will remain for a while so we all
have a chance to learn its new “home.” The
pollinator of V. ampullacea is unknown to
us but the flower morphology with a deep
and nectar-filled spur (gardiner, 2012)
and the rosy color suggest bee or butterfly
pollination. This species flowers from
March to May.
Vanda bicolor griff. is an attractive and
NIMA GYELSHEN
UNLESS OTHERWISE CREDITED, PHOTOGRAPHS BY STIG DALSTRÖM
relatively common epiphytic orchid that is
often considered endemic to Bhutan. no
doubt, however, it occurs in some of the
neighboring states of India, particularly
the state of Assam, since it has been
observed very close to the border. The
taxonomic status of V. bicolor has also
been considered as “doubtfully distinct”
from Vanda tessellata (Roxb.) Loddiges,
by Motes (1997). Pearce and Cribb (2002)
on the other hand treat Vanda bicolor as
distinct, based on a different coloration
of the flower, and rounded lateral lobes
of the lip versus triangular acute lobes in
V. tessellata. We agree with Pearce and
Cribb and accept this differentiation, but it
should also be noted that the color of at least
the sepals and petals in both species vary
and should not be considered as a reliable
feature of distinction. Vanda bicolor was
originally described as a Vanda and is
the only Bhutanese species in the genus
that is relatively unaffected by taxonomic
disagreements. Vanda bicolor flowers
from February through May in Bhutan
and, though no active pollination of this
species has been observed by us in the field,
visits by large bees are frequent on plants
in cultivation.
Vanda cristata Wall. ex Lindl. is
probably the most common and widespread
Vanda in Bhutan. It can be found in many
different types of habitats, from rather
hot lowland areas to surprisingly cool
mountain forests. This species, together
with V. griffithii Lindl. can also be found
in shadier locations than the other species
in the genus, with the exception of the
questionable V. himalaica. This ecological
preference, or perhaps “tolerance” is
a better choice of words, may be the
result of adaptation to beetle pollination
(cantarophilly). The quite variable and
rather peculiar looking flowers with large,
fleshy and blackish-purple lip lamina may
appear strange and even unattractive to
us at first. Both the odd shape and the
coloration may function as an attractor to
a certain species of an unidentified beetle,
which was observed and photographed
while successfully pollinating flowers
of V. cristata in cultivation at the Royal
Botanical garden in serbithang, Thimphu.
What appears to be the same species of
beetle also visited flowers of V. griffithii
and effectively removed the pollinarium.
The placement of the pollinarium differed
though, with V. cristata placing the pollinia
package near the scutellum (the “mid
back”) while V. griffithii placed the pollinia
package on the frons (the “head”) of the
beetle. Whether a successful pollination
of V. griffithii also took place in the
2
3
photographed specimen was unfortunately
not recorded. The big mystery behind these
observations, however, is how the beetles
could find the cultivated flowers in the first
place since no natural Vanda habitat exists
anywhere near where the plants are kept.
Beetle pollination of V. cristata correlates
with an observation by Pradhan (1983).
Vanda cristata flowers in April through
June.
Vanda griffithii Lindl. is named for
William griffith who traveled extensively
in the himalayan region in the early 19th
century and made many original scientific
collections of plants in Bhutan. griffith
[1] Vanda ampullacea growing in habitat and
a closeup (inset) picture of the flowers.
[2] Vanda bicolor growing in habitat along
with a closeup (inset) taken in situ along
with a likely pollinator. Dalström
[3] Vanda cristata growing in habitat along
with a closeup (inset) showing the flowers.
WWW.AOS.ORG
JANUARY 2015
ORCHIDS 35
36 ORCHIDS JANUARY 2015
WWW.AOS.ORG
5
Averyanov (1988) and senghas (1988b)
independently transferred Saccolabium
himalaicum to Holcoglossum instead. The
taxonomic controversy over this species
is probably due to the combination of
morphological features. The color and
appearance of the flowers do not really
resemble any others in Holcoglossum,
but has a certain superficial resemblance
to particularly Vanda (ascocentrum)
ampullacea. The plant habit of strict
pendent and terete leaves, however, makes
the position in Vanda or ascocentrum seem
like a “thumb in the eye.” The placement
of this odd species in Holcoglossum on the
other hand seems more likely, based on
the plant habit and also the basic flower
morphology. The rather unique plant
habit, with strictly pendent rattail leaves,
readily separates Vanda himalaica from all
other sarcanthinae species that are known
from Bhutan, where it flowers in October
NIMA GYELSHEN
4
NIMA GYELSHEN
later became superintendent of the Calcutta
(today called Kolkata) Botanic garden
(Pearce and Cribb 2002). The Vanda that
is named for this distinguished botanist is
the subject of some taxonomic controversy,
however. Traditionally, three very similar,
but considered distinct species, have been
described (Vanda alpina Lindl., Vanda
chlorosantha (garay) e.A.Christenson
and V. griffithii) that occur sympatrically
(growing together) in Bhutan, or at least in
the same southwest region (garay 1986).
The specific distinction between V. alpina,
V. chlorosantha and V. griffithii according to
garay is mostly based on the lip shape, and
particularly the apex with its fleshy ventral
hump. Vanda chlorosantha, however,
appears to be distinguished mainly by a lack
of anthocyanin (purple pigment) since no
particular morphological distinction can be
observed. Through field observations by the
authors of this paper, and in particular the
second author (gurung) who has extensive
experience with orchids in Bhutan, the
natural variations in size, shape and
coloration of the flowers of this group are
such that consistent specific distinctions
hardly can be made. We therefore favor
treating V. alpina, V. chlorosantha and V.
griffithii as representatives of the same
variable species, in partial agreement with
Joseph hooker, in the orchid part of his
Flora of British India in 1890 (garay 1986).
since V. griffithii was described first, this
name has nomenclatural priority and the
two others represent synonyms. Although
hooker’s opinion was based on a lack of
additional collections of V. griffithii, and
therefore considered it a synonym of V.
alpina, the conclusion is the same. Plants of
V. griffithii flower from March to July but
probably at other times of the year as well.
A plant that lacked anthocyanin, similar
to the type of V. chlorosantha, flowered in
september.
Vanda himalaica (Deb, gupta &
Malick) L.M.gardiner is another peculiar
species that really does not seem to fit
well in the current taxonomic position.
The first scientific description of this
species as Saccolabium himalaicum was
made by the team of Deb, sengupta and
Malick (1968). It is based on a collection
from “upper Burma” (Myanmar), sima,
by shalik Mokin (no. 13). What appears
to be the same species from Yunnan in
China was then described as Holcoglossum
junceum by Tsi (1982). This time, it is
based on a collection from an unknown
location by M. K. Li (no. 1798) in 1939.
Christenson (1987) transferred Saccolabium
himalaicum to ascocentrum in a revision of
Holcoglossum, but the following year, both
6
[4] Vanda griffithii showing typical habitat
along with a closeup (inset) showing
flowers from two plants growing side by
side.habitat.
[5] Vanda griffithii, without anthocyanin and
similar to “Vanda chlorosantha.”
[6] Vanda griffithii being visited by a beetle
carrying pollinia from another V. griffithii
[7] Vanda (Holcoglossum) himalaica displaying habitat typical for the species and
a closeup (inset) of the flowers of the
species. Note the elongated nectary of
this species.
[8] Vanda testacea flowering in habitat; typical for the species and a closeup (inset)
of the flowers.
through november.
Plants named Vanda testacea (Lindl.)
Rchb.f. in various literatures are small and
rather insignificant in habit, with yellow
flowers and bluish-purple specks on the
lip in various amounts. And yet, this
taxon is the source of some impressive
taxonomic confusion. The main problem
is that there are two rather similar but yet
morphologically different taxa that hide
under the name Vanda testacea. They
differ slightly in color, size and shape of
the flowers, and probably in geographical
distribution as well. The basic difference
can be seen in the structure of the lip,
particularly the spur, which is rather short,
broad and straight in the larger form from
Bhutan and sikkim, but being longer,
narrower and curved forward in plants from
sri Lanka, main parts of India, Myanmar
and Thailand. The larger-flowered form
also has a well developed bilobed callus
hump on the lip near the entrance of the
spur, which appears to be less developed
in the smaller flower. The sepals and petals
also appear paler yellow in the Bhutanese
(himalayan) form, but this observation can
also be due to a lack of experience with the
smaller form by the authors. since there
are several specific names published under
both aerides Loureiro, and later Vanda that
currently are lumped together as Vanda
testacea, we need to analyze the types of
each of these names in order to straighten
out the convoluted taxonomic history.
This entangled story is well illustrated by
Reichenbach’s comments in his transfer of
aerides testaceum Lindl. to Vanda testacea
(Reichenbach 1877):
“One might be doubtful as to
the oldest name, since a. testaceum
and wightianum were published
with descriptions at the same
time. One might say that aerides
wightianum, being a Wallichian
name, had the priority. My opinion
is that Wallich’s Catalogue gives no
authority for priority, for the names
there given have not won sanction
by an accompanying description.
Let us add that there is already a
Vanda wightii, and that a. testaceum
comes before a. wightianum in Dr.
Lindley’s book. Those who keep
the oldest name given in the genus
may write Vanda parviflora. This
question is a very difficult one.
(166)”
The first time the name “testacea”
(which refers to the yellow color of
the flower) is used was when Lindley
described aerides testaceum based on a
plant collected by Macrae in sri Lanka
(Lindley 1833). In the original description
Lindley writes: “Hab. in Zeylona [Ceylon],
supra arbores, Macrae” (238). The spur of
the flower is described as “calcare conico
incurvo.” In other words, a conical and
curved spur! Lindley’s type specimen is not
seen by us, but another Macrae collection
of this species from “Ceylon” (today called
sri Lanka) in 1829 and with the number
“49” is deposited in the herbarium of the
Museum of natural history in Vienna (W),
on sheet 38816. The spur of the flower is
rather long, slender and curved forward,
and correlates with Lindley’s original
description of aerides testaceum. There
is also a single flower on sheet 38817,
which seems to come from Lindley’s type
specimen despite a lack of information
other than “aerid. testaceum Ldl!” This
flower also displays a slender and curved
WWW.AOS.ORG
JANUARY 2015
ORCHIDS 37
spur similar to the other Macrae specimen
on sheet 38816 (W).
Immediately following the original
description of aerides testaceum is the
description of aerides wightianum, as
Reichenbach pointed out. Lindley writes:
“Habit. In India Orientali, prope Madras,
Wight.” The Indian city of Madras (today
called Chennai) is also very near Ceylon.
The spur of the flower is described as
“calcare brevi conico,” which translated
says “a broad and conical spur!” There
are three drawings attached on a sheet in
the herbarium in Vienna (W, from here on
referred to as the “three-drawing” sheet)
that show flowers of this species. The left
drawing is labeled “L,” and then “7320
aerides wightianum hb W. Wall” (referring
to Wallich’s herbarium and catalogue, no.
7320, which is mentioned in the original
description of aerides wightianum). The
middle drawing is labeled “L” and aerides
wightianum while the right drawing is
labeled “L” and Vda parviflora = a. wight.”
All three drawings display a flower with a
short conical spur, just like in flowers from
Bhutan. The middle and the right drawings
are copies of (Lindley’s?) drawings that
can be seen on herbarium sheets at Kew.
The problem is that the drawings do not
correlate accurately with the flowers
from which they apparently were drawn
regarding the shape of the spur. Perhaps this
was overlooked as a critical feature? After
analyzing a flower of the type specimen of
aerides wightianum at Kew (K001118001),
it is clear that the spur is both long and
narrow, and curved, thus similar to the
spur of aerides testaceum. since both
collections originate from the same general
area (southeastern India and sri Lanka), the
morphological similarities in the spur shape
support the conclusion that they represent
the same species.
Lindley described Vanda parviflora
based on a specimen he received from
the english nurseryman Loddiges (no.
1567), who apparently had obtained it as
a “native of Bombay” (Lindley 1844).
The spur of the flower of V. parviflora is
described as “calcare angusto obtuso,”
which refers to a narrow and obtuse spur.
The type specimen is mounted on a small
sheet at Kew, which in turn is mounted on
a larger sheet K000895723, and consists
of a single inflorescence with a flower and
two buds. A mounted envelope contains
an additional flower and one bud. There
are also two colored drawings, one with a
lateral view of the lip, and one that shows
the lip lamina from above. These drawings
were apparently copied by Reichenbach
and represent the right drawing on the
38 ORCHIDS JANUARY 2015
WWW.AOS.ORG
9
10
11
[9] Vanda testacea flower grown by Tommy Ljunggren.
[10] Side-by-side comparison of Vanda testacea from Bhutan (left) and the typical form of
Vanda testacea (right), grown by Tommy Ljunggren.
[11] Botanical illustration of Vanda testacea, as “Vanda parviflora”, plate from Annals of the
Royal Botanic Garden, Calcutta.
three-drawing sheet in the herbarium in
Vienna (W), and show a short and straight
spur, similar to flowers from Bhutan. The
dried flower on which the drawing is based,
however, reveals a curved spur. There is a
different Vanda specimen of what appears
to be the same species mounted on the
same (large) sheet at Kew. This specimen
is from the J.D. hooker herbarium (no.
183) and has the information “Concan,
W. Law.” Concan is situated on the west
coast of India and correlates geographically
with the type locality of V. parviflora from
Bombay (today called Mumbai). There
is a simple ink (?) drawing of the flower,
which also was copied by Reichenbach
and represents the middle flower on the
three-drawing sheet in Vienna (W). Again,
the drawing shows a straight spur while
the actual flowers reveal a curved spur. It
appears that Lindley identified this latter
specimen as aerides wightianum because
the name is written in his handwriting in the
lower right corner. But this specimen is not
the type of aerides wightianum, which was
collected by Wight near Madras on the east
coast of India.
It can be questioned here whether
Vanda parviflora really was collected
near Bombay or simply shipped from
the port there. Commercial growers and
importers sometime had the habit of
hiding the true origins of their plants, to
taxonomists’ great dismay. What supports
Bombay (western India) as the origin of
Vanda parviflora though is a collection by
Wight at Coimbatore in the southwestern
part of India, of the curved spur form of
this “species” (W, sheet 38813). Lindley
(1853) then sunk his Vanda parviflora into
synonymy with his aerides wightianum,
and later broadened his mind and sunk both
aerides testaceum and Vanda parviflora
into synonymy with aerides wightianum
(Lindley 1859). Our conclusion is that
Lindley first recognized a difference
between aerides testaceum and aerides
wightianum, and possibly also V. parviflora,
but later, gradually, decided that they
were all the same. Reichenbach (1877)
agreed with Lindley’s ultimate transfers
but reasoned that the correct name should
be Vanda testacea, which he believed had
nomenclatural priority. Reichenbach based
his opinion on multiple specimens that he
received from the english nurseryman
Bull, among others. If we analyze some of
these specimens and the drawings thereof,
however, we can see that two different spur
shapes are represented. A specimen from
Day on sheet 38822 (W) has a narrow and
curved spur similar to the type of Vanda
testacea, while a specimen from Bull on
sheet 38812 (W), which Reichenbach refers
to in his publication of Vanda testacea
(Reichenbach 1877) has a shorter, broader
and straighter spur, similar to the himalayan
plants. We therefore suspect that there
exist two geographically separated and
morphologically slightly different “forms”
of this species. The type form with a narrow
and curved spur has been described several
times and is geographically widespread
over much of southeast Asia, while the form
with a shorter and straight spur appears to
be limited to the himalayan region.
King and Pantling (1896) use the
name Vanda parviflora for the himalayan
species in their treatment of the orchids
of sikkim, himalaya. This amazing
publication features a fine illustration of
this species, which correlates with what
occurs in Bhutan.
Plants of Vanda in general, and the
species from Bhutan in particular, should
be treated as extreme epiphytes with no
or very airy medium to grow in. They are
best mounted in open wooden baskets or on
sturdy branches since the thick roots enjoy
spreading out and point in all directions.
Vandas from the himalayan region go
through a very wet and hot monsoon
season with copious watering from June
to september, while enduring a rather
long, dry and cooler winter season from
november to April. The humidity can still
be high though, particularly during night
and early morning. October and May can
be described as “intermediate” months
with a slowly decreasing and increasing
amount of watering, respectively. For
more information regarding cultivation see
Motes (1997).
References
Averyanov, L.V. 1988. new Taxa and nomenclatural
Changes in the Orchidaceae Family of Vietnamese Flora.
Bot. Zhurn. (Moscow and Leningrad) 73(3):432.
Brown, R. 1820. Vanda roxburghii. Bot. Reg. 6: t. 506.
Christenson, e.A. 1987. An Infrageneric Classification of
holcoglossum schltr. (Orchidaceae: sarcanthinae) with
a Key to the genera of the aerides-Vanda Alliance. notes
Roy. Bot. Gard. Edinburgh 44(2):249–256.
_. 1992. notes on Asiatic orchids. Lindleyana 7: 88–94.
Deb, D.B., g. sengupta, and K.C. Malick. 1968. Saccolabium himalaicum. Contribution to the Flora of Bhutan.
Bull. Bot. Soc. Bengal 22:213.
gardiner. L.M. 2012. new Combinations in the genus
Vanda (Orchidaceae). Phytotaxa 61:47–54.
King, g. and R. Pantling. 1896. Vanda parviflora. ann.
Roy. Bot. Gard. Calcutta 8(3): pl. 286.
Kuntze, C.e.O. 1891. Gastrochilus ampullaceus. Revis.
Gen. Pl. 2:661.
Lindley, J. 1833. aerides testaceum and aerides wightianum. Gen. Sp. Orchid. Pl. 533.
_. 1838. Saccolabium ampullaceum (Roxb.) Lindl., Sert.
Orchid. part 4: t. 17.
_. 1844. Vanda parviflora. Bot. Reg. 30:45, Misc. (genus
57).
_. 1853 to 1855. Folia Orchidacea, an Enumeration of the
Known Species of Orchids, Vol. 1. (Vanda). J. Matthews,
London, uK.
_. 1859. aerides wightianum. Journal of the Proceedings
of the Linnean Society of London, contribution to the
Orchidology of India (2):40.
Motes, M.R. 1997. Vandas, Their Botany, History and
Culture, Timber Press, Portland, Oregon.
Pradhan, g.M. 1983. Vanda cristata. amer. Orch. Soc.
Bull. 52(5):464–468.
Reichenbach, h.g. 1877. new garden Plants, Vanda
testacea. Gard. Chron. 2(8):166.
Roxburgh, W. 1832. aerides ampullacea. Fl. Ind. 3:476.
schlechter, R. 1913. ascocentrum ampullaceum. Repert.
Spec. nov. Regni Veg. Beih. 1:975.
seidenfaden, g.
senghas, K. 1988a. Trudelia garay. Schltr. Orchideen
ed. 1:1211.
_. 1988b. Holcoglossum himalaicum. Schltr. Orchideen
ed. 3, 1(19–20):1194.
Tsi, Z.h. 1982. Holcoglossum junceum. acta Phytotax.
Sin. 20:442–443.
Acknowledgments
We sincer ely thank th e Ro y al
government of Bhutan, and the staff of
the Ministry of Agriculture and Forests
of Bhutan for providing the necessary
permits; the administration and staff
of the national Biodiversity Centre,
serbithang, for organizing the logistics
and particularly Dupchu Wangdi who is
the caretaker of the live orchid collection
and does a great job. We also thank the
very helpful herbarium staff at the Royal
Botanic garden, Kew, for excellent digital
service, and the Museum of natural history
in Vienna. We thank Rudolf Jenny for
providing copies of numerous important
and hard-to-find orchid articles, Wesley
higgins for reviewing and commenting
on the manuscript, Tommy Ljunggren for
supplying live flowers of the two forms of
Vanda testacea and the sarasota Orchid
society for financial support.
— Stig dalström is a research
associate at the Lankester Botanical
Garden, University of Costa Rica, Cartago,
Costa Rica, and the national Biodiversity
Centre, Serbithang, Thimphu, Bhutan. 2304
Ringling Boulevard, Unit 119, Sarasota,
Florida 34237. (stigdalstrom@gmail.com;
www.wildorchidman.com). dhan Bahadur
Gurung (Phd), author of An Illustrated
guide To The Orchids Of Bhutan, is an
assistant Professor and dean of the College
of natural Resources at the Royal University
of Bhutan (dbg2006@gmail.com). Choki
Gyeltshen is a Biodiversity Officer at the
national Biodiversity Centre, Ministry
of agriculture and Forests, Serbithang,
Thimphu, Bhutan (chokig@gmail.com)
and nima Gyeltshen is the Biodiversity
Supervisor at the Royal Botanical Garden,
national Biodiversity Centre, Ministry
of agriculture and Forests, Serbithang,
Thimphu, Bhutan (nimss409@gmail.com).
Correspondence should be sent to Stig
dalström.
WWW.AOS.ORG
JANUARY 2015
ORCHIDS 39