全 文 ：Journal of Systematics and Evolution 46 (2): 230–235 (2008) doi: 10.3724/SP.J.1002.2008.07115
(formerly Acta Phytotaxonomica Sinica) http://www.plantsystematics.com
Taxonomic history and identity of Musa dasycarpa, M. velutina and M.
assamica (Musaceae) in Southeast Asia
1,2Markku HÄKKINEN* 3Henry VÄRE**
1(Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Menglun, Mengla, Yunnan 666303, China)
2(Botanic Garden, University of Helsinki, P.O. Box 44 (Jyrängöntie 2), FI-00014, Finland)
3(Finnish Museum of Natural History, Botanical Museum, University of Helsinki, P.O. Box 7 (Unioninkatu 44), FI-00014, Finland)
Abstract Since the initial description, the name Musa dasycarpa Kurz (1867) has been unclear to most botanists.
It has usually been synonymized with M. velutina H. Wendl. & Drude (1875). However, although the original
diagnosis was very short, “fruits hairy”, it is adequate. Thus, according to Vienna Codes, M. dasycarpa Kurz has
priority over M. velutina H. Wendl. & Drude. The aim of this study is to settle the true identity and to update the
description of M. dasycarpa Kurz. For that purpose the names M. dasycarpa and M. velutina are typified. In addi-
tion, critical notes regarding M. assamica Bull. are given; it is neotypified here, and considered as conspecific
with M. sanguinea Hook. f.
Key words Musa, Musa assamica, Musa dasycarpa, Musa velutina, Musaceae, Rhodochlamys, Southeast Asia,
wild banana.
1. Musa dasycarpa Kurz, J. Agric. Hort. Soc. Ind. 14:
301 (1867). Type: Icon of Musa dasycarpa at Kew
(lectotype, designated here, K!).
Sulpiz Kurz was a German botanist who worked
in India and Myanmar. The name of M. dasycarpa
Kurz (1867) was diagnosed by him very briefly:
“fruits hairy”. There is an icon (Fig. 1) at K, perhaps
drawn by Kurz, with the following comments: Musa
dasycarpa Kurz, Assam (type specimen in Calcutta
herbarium), fruits densely hairy. The Calcutta speci-
men has been lost. Cheesman (1949) noted that he did
not find any collection of M. dasycarpa at Kew. How-
ever, there are two herbarium samples at Kew: sheet
2485 (Fig. 2) of M. dasycarpa predating the descrip-
tion, 3.X.[18]50 (incorporated in Herbarium Hookeri-
anum in 1867). The sheet date of Fig. 2 indicates that
the collector was not Kurz himself because he arrived
to Calcutta, India over a decade later. Kurz (1878)
clearly stated that M. dasycarpa was only known by
its fruits when he described it in 1867. The sheet from
1850 at Kew also has leaves. Therefore it seems very
likely, that the icon at K (Fig. 1) represents the only
original element known today. A third sheet at K,
collected by I. H. Burkhill (no. 35722, Fig. 3), Ahor
Exped. Makum [Assam, India] 21.XI.[19]11, shows
both fruits and leaves also.

2. Musa velutina H. Wendl. & Drude, Gartenflora 65,
t. 823. 1875. Type: Icon in Wendland & Drude, Gar-
tenflora 65, t. 823. 1875 (Fig. 4) (lectotype, designated
here).
Musa velutina was described by a German,
Hermann Wendland, and an Englishman, George




























Fig. 1. Lectotype of Musa dasycarpa (K, 1867). Photo: M. Xanthos, K.

———————————
Received: 30 August 2007 Accepted: 4 December 2007
* Author for correspondence. E-mail: .
** E-mail: .
HÄKKINEN & VÄRE: Taxonomic history and identity of Musa species in Southeast Asia

231

































































Figs. 2–5. 2. Specimen of Musa dasycarpa (K, 2485). Photo: M. Xanthos, K. 3. Specimen of Musa dasycarpa (K, 35722 1911). Photo: M. Häkkinen,
H. 4. Lectotype of Musa velutina (t. 823). Photo: T. Deroin, P. 5. Neotype of Musa assamica (K, 170B/4/36). Photo: M. Häkkinen, H.


Journal of Systematics and Evolution Vol. 46 No. 2 2008 232
Drude, from a flowering plant at Herrenhousen Bo-
tanic Garden at Hannover. It was introduced from
Assam by German Gustav Mann. The diagnosis of M.
velutina (Wendland & Drude, 1875) was as follows:
“Spadix erectus purpureus, pedunculo, rhachi, spathis
floriferis inferioribus dense velutinis. Spatha infima
sterilis, sequentes tres floribus 3–4 feminis axillaribus,
superiores 6–9 floribus totidem masculinis ornatae.
Flores femini germine quam perianthium paulo
longiore vel ei aequilongo velutino anguloso; labio
exteriore tricarinato quinquedentato, dentibus revo-
lutis vel convolutis, labio interiore (labello) obtuso
exterius longitudine aequante eoque latiore; stamini-
bus 2 fertilibus, antheris dimidiatis falcatis; stigmate
crasso lirelliformi. Flores masculini germine rudimen-
tario stipitiformi incurvo glabro; labio exteriore lato
bicarinato dentibus 5 revolutis, interiore acuto bre-
viore; staminibus inaequalibus 5 fertilibus antheras
binas gerentibus; stigmate quam in floribus feminis
minore. Folia basi inaequali in petiolum longe decur-
rentia.”

Musa dasycarpa or M. velutina
Cheesman (1949) was the first who pointed out
that the drawing (lectotype of M. dasycarpa at K)
strongly suggests identity with M. velutina, and he
regarded it as probable that further study may estab-
lish M. dasycarpa Kurz as the valid name for the spe-
cies hitherto called M. velutina. Simmonds (1960)
came to the same conclusion. No other hairy fruited
Musa species have been found from Assam (Singh et
al., 2001; Uma et al., 2006), and such are not known
elsewhere in continental Asia either.
Already Kurz (1878) commented on the name as-
signed by Wendland and Drude (1875): “They pub-
lished a supposed new species which they call M.
velutina.” It seems likely that Kurz recognised that M.
velutina was the same as his M. dasycarpa and he
commented: “I cannot embark here upon a sifting of
the literature and synomy, for such would be of too
technical a character, and will be published in my
revision of the Musaceae under preparation” (Kurz
1878: 163). However, Kurz never published his revi-
sion because he died in Penang Malaysia shortly after
writing those words.
The true schism on the correct name began when
Baker (1893) reduced M. dasycarpa to a synonym of
M. velutina in his synopsis of banana. Subsequent
botanists largely followed his view (e.g., Schumann,
1900; Pucci, 1906; de Wildeman, 1912; Burkill, 1925;
Worsdell, 1941; Walters et al., 1984; Simmonds &
Weatherup, 1990; Hore et al., 1992). Obviously Baker
(1893) was not certain of Kurz’s name (Cheesman,
1949). The reason might be that Wendland and Drude
(1875) did not clearly mention the fruits at all in their
Latin diagnosis, which was the key character of
Kurz’s taxon. However, that did not justify reducing
M. dasycarpa as a synonym of M. velutina, as Baker
(1893) did. At the very end of their article, Wendland
and Drude (1875) stated concerning the figures “die
Samenknospen einhüllenden Haare sind fortgelassen”
(hairs covering the ovaries not drawn in the table 823,
see figure 4). The only other species with hairy ova-
ries is M. hirta Becc. from Borneo, a species of sec-
tion Callimusa Cheesman. Musa dasycarpa belongs to
section Rhodoclamys Baker (Baker, 1893: 205). Musa
dasycarpa from India is one of the five known ba-
nanas of the family Musaceae in which the fruit is
schizocarpic on maturity. The others are M. johnsii
Argent, M. lolodensis Cheesman and M. schizocarpa
Cheesman from Papua New Guinea, and Musella
lasiocarpa (Franc.) C. Y. Wu from China (Franchet,
1889; Baker, 1893; Cheesman, 1950; Simmonds,
1956; Li, 1978; Argent, 2001). The older name M.
dasycarpa has priority over M. velutina according to
Vienna Code (McNeill et al., 2006).
Characterization of Musa dasycarpa
The following description of M. dasycarpa is
based on cultivated living plants in the Botanic Gar-
den of University Helsinki (accession 1998-0017), by
completing the entire INIBAP Musa Descriptor List
(IPGRI-INIBAP/CIRAD, 1996). The descriptive
terms here also follow the tradition of banana taxon-
omy as used by N. W. Simmonds (Simmonds, 1962,
1966).
Plant slender, suckering freely, close to parent
plant, 2–3 suckers, position vertical. Mature pseu-
dostem up to 1.5 m high, 7 cm in diameter at base,
often smaller, underlying colour light green, devoid of
wax and without pigmentations, sap watery. Petiole up
to 30 cm, petiole canal margins straight with erect
margins, petiole bases winged and not clasping the
pseudostem. Leaf habit intermediate, with very corru-
gated lamina, lamina up to 1 m long and 35 cm wide,
truncate at the apex, colour of upper surface dark
green, and lower surface green, appearance shiny,
midrib dorsally green and ventrally red, leaf bases
asymmetric, both sides pointed. Inflorescence erect,
peduncle up to 10 cm long and 2 cm in diameter,
heavily clothed with white pubescence and red in
colour, sterile red bract one, persistent at the opening
of the first female flowers. Female bud ovoid, up to 15
cm long and 6 cm wide, bracts pale pink in external
and internal faces, no imbrications, no wax, lifting two
HÄKKINEN & VÄRE: Taxonomic history and identity of Musa species in Southeast Asia

233
bracts at a time, revolute before falling. Basal flowers
female, hermaphrodite, 3–5 per bract in a single row,
ovary 3 cm long, pale pink, velvety, densely pubes-
cent, arrangements of ovules in two rows per loculus,
compound tepal 3 cm long, orange pink in colour and
the lobes orange, free tepal as long as the compound,
boat shaped, yellowish, strongly grooved, stamens 5,
whitish, light green style with orange stigma. Male
bud ovoid, 12 cm long and 5 cm wide, bracts pale
pink in external and internal faces, lifting two bracts at
a time, revolute before falling, commonly aborting
after producing about a dozen of bracts with male
flowers. Male flowers on average 5 per bract in one
row, falling with the bract, compound tepal 3.8 cm
long, orange-yellow, with a pink flush on the back,
with 5-toothed apex, free tepal 3.7 cm long, translu-
cent white, oblong, with a short broad acumen, sta-
mens 5, whitish, anthers yellow, light green style with
orange stigma, anthers and style at the same level,
ovary straight, orange, without pigmentation.
Fruit bunch compact, with 5 hands and 4 fruits
per hand on average, in 1 row, fruit bright pink,
rounded, 7 cm long, 3–4 cm in diameter, pubescent
like the peduncle, broadly truncate at apex, and sub-
sessile at base, without any floral relicts, pericarp 3–4
mm, thick, splitting at maturity and separating in ir-
regular strips from apex to base, exposing the central
mass of white pulp and seeds.
Seeds black, tuberculate, irregularly angulate-
depressed, 4–6 mm across, 2–3 mm high, 80–90 per
fruit. Chromosome numbers are 2n = 22 (Cheesman,
1947).
Additional specimens examined:
Burma (Myanmar). Myitkina, Tawgin, Ehaung, alt. 800
ft. 24.XI. [19]28, C. E. Parkisson 1765 (K!); Trinidad. Cult.
Imp. Coll. Trop. Agric., Introduction 212, H. 1171/1949, R. E.
D. Baker 23.VI.1949. (K!).
China. Yunnan: Xishuangbanna Tropical Botanical Gar-
den, beside dinner room in XTBG, alt. 570 m, 1.X.2005,
Heliqing 5 (HITBC 111415!).
Distribution and habitat Musa dasycarpa is
distributed at lower altitudes across Assam to Arun-
achal Pradesh in northeast India and northern Myan-
mar. It has become a menace as a weed in agricultural
fields due to its fast suckering and spreading seed
habit by birds and bats (Uma et al., 2006). Its fruits do
not change the colour upon ripening but they dehisce
as a sign of maturity. It has also been disseminated all
over the world as the most ornamental banana. It also
hybridizes well with the other Rhodochlamys species
so there are number of unidentified artificial hybrid
clones in horticulture all over the world (Simmonds,
1962; Shepherd, 1999; Häkkinen & Sharrock, 2002;
Häkkinen, 2005, 2007).
Key to some closely related Rhodochlamys species
1a. Basal flowers hermaphrodite, ovary 3 cm long and pale pink, peduncle dark red and very hairy, male bracts pink at both sides,
fruits 7 cm long, hairy, angular, pink at maturity, self peeling at ripening….…………..…..….…………....…..Musa dasycarpa
1b. Basal flowers female, ovary 4 cm long and light green, peduncle light green, male bracts lilac at both sides, fruits 8 cm long,
glabrous, straight, rounded, light green, becoming pale yellow at maturity…………………………………...……………..….2
2a. Up to 15 pseudostems in the same plant, close to parent plant, inflorescence erect, free tepal as long as compound tepal……..…
………………………………………………………………………………………………………………………….…..M. ornata
2b. Up to 2–3 pseudostems in the same plant, emerging 2 meters from the parent plant, free tepal 1/4 long as compound tepal …3
3a. Inflorescence erect, male flowers 6–10 per bract in two rows, orange in colour, bracts brick red in colour…………M. laterita
3b. Inflorescence horizontal, male flowers 2–3 per bract in one row, light yellow in colour, bracts rose in colour…….……M. mannii

Musa assamica Bull in A retail list of new, beautiful
and rare plants offered by William Bull: 6. 1871.
Type: Britain, Kew. Hort. Kew. 170B/4/36. 3XII.
1884. (neotype, designated here, K!). Fig. 5
Bull (1871) published the name Musa assamica
without proper diagnosis: “This is a peculiarly dwarf-
habited and elegant species, and has been imported
from Upper Assam. The slender pseudostems are
about a foot and half high, green, bearing a crowded
tuft of several elliptic lanceolate leaves, which are
stalked, about a foot in length, remarkably unequal-
sided at the base, acute at the apex, and running out
into a slender tendril-like point. The leaves are green,
with a narrow purple border. It will make good plant
for table decoration, on account of its exotic aspect
and moderate size and stature – 1 guinea”. Also
Cheesman (1949) was aware of this description,
which has been almost neglected ever since.
Baker (1892) included the name in his list of im-
perfectly known species allied to M. sanguinea, with a
short “diagnosis”: “Leaves about 1 ft. very unequal-
sided bright green – Assam”. He did not refer to Bull
(1871) at all, and his name is a younger synonym for
M. assamica. Further, M. assamica Baker could be
considered as an invalid name according to Art. 34.1.
(McNeill et al., 2006), as Baker considered the name
uncertain.
According to Cheesman (1949) there is a note
Journal of Systematics and Evolution Vol. 46 No. 2 2008 234
among the Musa material in Kew Herbarium in which
Mann (who was responsible for collecting altogether
four new species of Musa in Assam) states that he
does not know which of [his] Assam species has been
called “assamica”. Cheesman (1949) continued: “We
may conclude fairly safely that the plants offered for
sale under the name M. assamica were young speci-
mens of either M. aurantiaca, M. mannii, M. san-
guinea or M. velutina. It is quite impossible, from a
description which fits almost any Musa seedling, to
connect M. assamica with any of the four species
from Assam”.
Musa mannii Baker was first introduced to Kew
from Herrenhousen in 1885 and M. aurantiaca Baker
in 1894 (Baker, 1894); both much later than Bull’s
sales catalogue. That eliminates the possibility that the
plants for sale under the name M. assamica were ei-
ther M. mannii or M. aurantiaca. It is no longer possi-
ble to determine which of the other two species men-
tioned by Cheesman (1949) in his quotation was M.
assamica. The material associated with that name
probably become extinct in cultivation many years
ago (Baker, 1892, 1893, 1894; Hooker, 1893; Fawcett,
1913; Champion, 1967; Häkkinen & Sharrock, 2002;
Häkkinen & Väre, 2007). Chittenden and Synge
(1956) speculated in The Royal Horticultural Society
Dictionary of Gardening that “M. assamica is proba-
bly identical with M. sanguinea”. That view is
adopted here, and consequently, M. assamica Bull. is
neotypified with M. sanguinea Hook. f. collection at
K. The neotype was collected from a plant cultivated
at Kew Gardens, of which Bull perhaps collected his
seed material.
Baker must be cited as an author alone concern-
ing M. mannii, as he refers to an unpublished manu-
script by Hermann Wendland (Baker, 1892: 263).
Thus the publication by Wendland was not effective
(McNeill et al., 2006; Art. 29.1). The genus Musa was
printed as a part of Fl. Brit. Ind. in 1892 (Stafleu &
Cowan, 1979), not in 1893 as often cited.

Conclusion
Musa taxonomy is still very obscure today just as
it has been throughout its history despite attempts to
clarify it. The Kew superintendent William Watson
wrote in 1894 in Garden and Forest, New York as:
“Mr. J. G. Baker, keeper of the Herbarium at Kew, has
recently prepared a Synopsis of the Genera and Spe-
cies of Musa, which was published in Annals of Bot-
any [1893].” He continued: “The genus Musa, I be-
lieve, gave Mr. Baker considerable trouble. He admits
thirty-two species, and divides them into three sub-
genera: Physocaulis, Eumusa and Rhodochlamys.”
The next classification revision was made by E. E.
Cheesman in his monumental series “Classification of
the Bananas, Critical Notes on Species” published in
Kew Bulletin 1947–1950. He divided wild bananas
into four sections, viz., Australimusa, Callimusa, Eu-
musa and Rhodochlamys. N. W. Simmonds continued
the taxonomical work in 1950–1960 by describing
several new species, also published in Kew Bulletin.
This article is part of the revision work with the old
taxa and the authors do believe that the taxonomical
question regarding M. dasycarpa versus M. velutina
and M. assamica is now solved. However, it might
prove necessary to conserve M. velutina H. Wendl. &
Drude.
Acknowledgements The authors greatly appreciate
the help and support from the staff of the Kew herbar-
ium, London and the Muséum National d’Histoire
Naturelle, Paris, Mrs. Sirkka Sällinen, librarian at
Finnish Museum of Natural History, Helsinki who
made a great effort in searching articles for us, and
Emory Walton from California Rare Fruit Growers,
Thousand Oaks for proof reading this article.
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