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ISSN: 2287-688X
Original Research Article
*Corresponding Author:
Dr. M. Sabu,
Professor & Head,
Department of Botany,
University of Calicut,
Kerala, 673 635, India.
Page| 260
Notes on the Rediscovery and Taxonomic Status of
M. flaviflora
N.W.Simmonds and M. thomsonii (King ex Schumann) A.M.Cowan & Cowan
(Musaceae) From India
Alfred Joe, Sreejith PE and M Sabu*
Department of Botany, University of Calicut, Calicut University P.O., Kerala- 673 635, India
Received for publication: May 13, 2013; Accepted: June 21, 2013.
Introduction
The genus Musa L. (Linnaeus, 1753)
belongs to the family Musaceae with major
centre of distribution in South and South-East
Asian countries, Africa and Pacific Islands.
The Family includes two other genera, Ensete
Bruce ex Horaninow and Musella (Franchet)
Wu and around 150 taxa in the world. In
India about 20 taxa have been reported under
two genera, Ensete and Musa (Karthikeyan et
al.,1989, Uma et al., 2005). India is well
known for its vast genetic diversity of
bananas comprising seeded wild species to
seedless cultivars. This in support of the fact
that the origin of Musa is South and South-
East Asia including Indo-Myanmar region
(Singh et al., 2001). In India wild Musa
species are largely distributed in North-
Eastern States, Western Ghats, Eastern Ghats
and Andaman and Nicobar Islands, because,
these areas consists of tropical rain forests,
wet evergreen forests, moist deciduous forest
of low rainfall zones, which are favorable for
its growth. Although India is considered as
one of the centres of origin of family
Musaceae, the taxonomy of many taxa is still
in a state of flux. The endemism of many taxa
in inaccessible forests, bulky nature of the
pseudostem, etc. made the study difficult. No
comprehensive work on Musaceae in India
was made after Baker (1894) in Hooker’s
‘Flora of British India’. Karthikeyan et al.,
(1989) enumerated 21 species under two
genera in ‘Florae Indicae Enumeratio
Monocotyledonae’. Recently some new taxa
were described from India viz. M. velutina
subsp. markkuana M.Sabu et al., (2013a), M.
velutina var. variegata A.Joe et al., (2013),
M. sabuana Prasad et al., (2013) etc. Besides
these there are some new additions to the
wild banana flora of India. Sabu et al.,
(2013b) and Alfred et al., (2013) recorded
the occurrence of M. chunii Häkkinen and M.
laterita Cheesman respectively from India.
During intensive explorations in North-
East India, authors could collect curious
specimens of wild banana species and found
that taxonomy statuses of many species are
still in confusion and several species need
typification and clarification about its identity.
This paper focuses mainly on the taxonomic
history, correct identity and about the
rediscovery of M. flaviflora N.W.Simmonds
and M. thomsonii (King ex Schumann)
A.M.Cowan & Cowan.
Baker (1893) mentioned four wild
‘forms’ in Sikkim under the heading M.
sapientum subsp. 3. M. seminifera Lour,
distinguished by Dr. King as pruinosa, dubia,
hookeri and thomsoni and added “Dr. King
thinks the two later forms as likely to be
distinct specifically from sapientum. His
hookeri is probably M. sikkimensis Kurz”. He
gave a small description to thomsoni and the
diagnosis was as follows: “4. thomsoni
(Kergel of Lepchas). Stem green, 12–15 ft.
Abstract:
The taxonomy of Musa flaviflora and M. thomsonii has been in a state of flux
since their inception. Due to their close resemblance and lack of proper types and
protologues they were treated as conspecific. Both species have been studied in detail
based on live collection and protologues for their taxonomic status and
circumscription. Here both the species are correctly identified and provided detailed
description after the type collection. Photographs, distribution, types and ecological
notes of both species are also provided.
Keywords: Musa, M. flaviflora, M. thomsonii, Musaceae, North-East India,
rediscovery, taxonomic status.
Alfred Joe et al.: Annals of Plant Sciences, 2013, 02 (08), 260-267
Page | 261
long, leaves glaucous only when young,
conspicuously cuspidated at the apex, bracts
ovate, outside with vertical streaks of yellow
and purplish-brown, yellow inside; fruits 2/5
in. long, ¾ in. diam. faintly ribbed; seeds
few, black soft, 1/5 in. diam. Surrounded by
copious sweet pulp. Does not rise above 1500
feet”. King collected the specimens from
Sikkim known as ‘Kergel’, a vernacular name
used by the tribe ‘Lepchas’ in Sikkim. The
distinguishable characters from any other
Musa are about the bract colouration and
cuspidate nature of leaf apex. May be inspired
from Baker’s last sentence about the last two
forms, Cowan and Cowan (1929) treated the
first two forms as varieties of M. sapientum
and the last two as two separate species viz.
M. thomsoni and M. hookeri and the
description of M. thomsoni as follows “In
sheltered spots in the Lower Hill Forest
“L.H.F.” up to 1,500 ft. Stem 12–15 ft. high,
7–9 ins. in diameter at ground level. Stem
green, except at the base, where it has brown
spots. Seeds few, soft, black; pulp soft,
sweet”. But before Cowan and Cowan (1929),
Schumann (1900) gave it to a varietal status
under M. sapientum subsp. seminifera. After
Cowan and Cowan this species was unknown
to most botanists. Cheesman (1948b) came
across King’s four forms and about materials
of those. He adds “ There is materials of
these forms in the Kew Herbarium, which
may help to settle the nomenclature when the
plants have been further studied on living
specimens, but the first requirement is to
determine by re-collection how many of the
large-seeded species there are in Sikkim”.
During the Taxonomic revision of Musaceae in
India, authors collected this species from
Meghalaya, quite far from the type locality.
After King, there has not been any collection
of M. thomsonii and hence the present study
forms the rediscovery of the taxon after lapse
of more than a century.
Cheesman (1948a) described a plant
flowered at Imperial College of tropical
Agriculture, Trinidad and Tobago, which was
introduced from Assam, by P.H. Carpenter
Esq., of the Indian Tea Association at Tocklai
Experimental Station. He called it as ‘Mariani
form’ of M. acuminata. The description was
based on a specimen (I.R. 209) which was
collected from Mariani Range of Assam,
North-East India and the description was
“Male bud in advanced blooming acute, the
bracts convolute at the tip, the whole bud
usually aborting before the fruits are mature
and falling off; bracts rather bright red,
moderately glaucous outside, yellow and
shining within. Male flowers pale orange-
yellow, the teeth of the compound tepal deep
orange and about 5mm long…..”
He also added “ if this form has to be
separated from M. acuminata it will probably
prove conspecific with a plant from Tagwin,
Myitkyina, Upper Burma (I.R. 183), which at
present pending cytological studies. The
Tagwin plant closely resembles the Mariani
form….”
Subsequently, during the ‘Banana
Expedition trip’ to India at 1954–55,
Simmonds (1956) elevated ‘the Mariani form’
to species level by keeping Cheesman’s I.R.
209 in Herb. Kew. as the type specimen. He
also remarked that “This species resembles
M. acuminata and on purely morphological
grounds would be best treated as the
northernmost subspecies of it”. The recorded
distribution in India include Assam, Manipur,
Meghalaya and Nagaland. He gave several
vernacular names for the species. After
Simmonds the identity of this species is
unclear to most botanists and they just
enumerate the name in their works and some
are mere compilation of Simmonds’ work
(Karthikeyan et al., 1989, Hore et al., 1992,
Noltie 1994, Uma et al., 2005, Ude et al.,
2002; Liu et al., 2010; Singh 2010; Li et al.,
2010). Some workers even treated M.
flaviflora as a synonym of M. thomsonii. This
schism starts after Simmonds (1956).
Actual problem starts when Simmonds
(1956) treated doubtfully M. sapientum
subsp. seminifera form thomsonii “thomsoni”
King Mss ex Baker and M. thomsonii
“thomsoni” (King Mss ex Baker) Cowan and
Cowan as synonyms, when he described M.
flaviflora. He criticized the validity of M.
thomsonii, which he thought it was
inadequately described. Simmonds adds:
“Baker states that the bracts have “vertical
streaks of yellow and purplish brown” outside
which does not agree with the bright red of M.
flaviflora. His publication of the name,
incidentally, by Arts, 42 and 43 of the code, is
probably invalid”. But Simmonds never
include the notes regarding the inclusion of M.
thomsonii as a synonym of his species, yet he
was sure about the two taxa are different.
And he conclude “M. thomsoni is so ill
described and typified that it may well be
rejected; if it can ever be certainly identified
Alfred Joe et al.: Annals of Plant Sciences, 2013, 02 (08), 260-267
Page | 262
then less confusion will result from the
treatment adopted here than by taking up
thomsoni for the Assam species if they are
not in fact the same’.
After Simmonds many botanists used
M. thomsoni as synonym of M. flaviflora
(Simmonds 1960; Simmonds & Weatherup
1990; Shepherd 1999). But some thought
about the rule of priority and without looking
the difference in two species, treated M.
flaviflora as the synonym of M. thomsoni
(Häkkinen & Väre, 2008). Some treated both
as different species (Karthikeyan et al., 1989;
Noltie 1994). This adds a great confusion
among these two species.
As part of taxonomic revision of
Musaceae in India, authors visited
Mariani, Assam, the type location of the
species (M. acuminata ‘Mariani form’). But the
whole area is entirely changed. Now Mariani
is a small town near Jorhat, and surrounding
areas were acquired for tea plantations and
couldn’t collect the species from its type
locality. But a plant collected from Zunheboto
District, Nagaland, in non-flowering condition
was grown in Calicut University Botanical
Garden (CUBG) and flowered in CUBG and it
was identified as M. flaviflora. Since
Simmonds (1956), no live collection of this
species has been made and hence the present
collection forms the rediscovery of the taxon
after a lapse of more than half a century. The
authors also came across the two species and
identified the two are different taxa and
therefore here submit the two species as
different.
The descriptions provided below from
notes, live plants grown in CUBG as well as
photos taken from living plants in the wild by
completing the entire INIBAP Musa Descriptor
List (IPGRI INIBAP/CIRAD 1996). The
descriptive terminology here follows that used
in the traditional banana taxonomic works by
Simmonds (1962, 1966) and Argent (1976).
Musa flaviflora N.W.Simmonds, Kew
Bull., 11(3): 471. 1956; In. Champion, Les
bananiers et leur culture, 33. 1967;
Karthikeyan et al.,, Flora Indica Enum.
Monocot., 104. 1989; Hore et al.,, J. Econ.
Tax. Bot., 16(2): 451. 1992; Noltie, Flora of
Bhutan, 3(1): 182.1994; Häkkinen & Väre,
Adansonia, 30(1): 77. 2008. Fig. 1.
Type: INDIA. Assam, Mariani hills, Cheesman E.E., I.C.T.A., I.R. 209 (spirit collection) K!),
(Lectotype designated by Häkkinen & Väre, 2008)
Alfred Joe et al.: Annals of Plant Sciences, 2013, 02 (08), 260-267
Page | 263
Musa acuminata ‘Mariani form’ Cheesman,
Kew Bulletin, 3(1): 28. 1948; Simmonds,
Journ. Genetics, 51, 32-40. 1952.
Plants slender, suckers 3–4, close to
the main shoot, 10–15 cm away, vertically
arranged. Mature pseudostems 1.8–2.1 m
high, 20–30 cm circumference at the base,
green with some black blotches, covered with
old brown leaf sheaths except apex region,
apex waxy, sap watery. Leaf habit
intermediate, laminae 150–155 × 36–51 cm,
oblong-lanceolate, apex oblique, truncate,
adaxially dark green with black blotches or
dots, dull, abaxially medium green,
appearance shiny, leaf bases symmetric, both
rounded, midrib adaxially light green and
abaxially yellow-green. Petioles 43–50 cm
long, glaucous, petiole margins wide with
erect, with blackish-brown scarious margin,
and black-brown blotches at the base, petiole
base winged and clasping the pseudostem.
Inflorescences first erect at arising stage,
then horizontal to sub-horizontal, peduncle
13–15 cm long, green, glabrous. Sterile
bracts 1–2, deciduous, c. 30 × 10 cm,
adaxially red and abaxially creamy pink, apex
with leafy appendage. Female buds
lanceolate, convolute, apex slightly imbricate.
Female bracts 15–23.3 × 8–12.4 cm,
moderately grooved, adaxially red with yellow
tip, slightly glaucous, abaxially cream with
pink tinge, shiny, apex acute, lifting one bract
at a time, reflex and revolute before falling,
cincinnus 12–20 flowers. Basal 5–8 hands
female. Female flowers 12–20 per bract in a
two rows, 6.5–8.5 cm long. Compound tepal
3.5–4 × 1.1–1.3 cm, yellow, ribbed at dorsal
angles, lobes 5, 0.4–0.6 × 0.2–0.3 cm, dark
yellow, outer two lobes with small horns. Free
tepal 2.5–2.7 × 1.4–2 cm, boat-shaped,
translucent cream tinged with yellow, apex
corrugated, with a short acumen. Staminodes
5, 1.3–2.1 cm long, cream with brown apex.
Ovary 3.8–4.1 cm long, straight, light green–
yellow-green, with ovules in 2 rows per
locule, style straight, exserted, 2.5–3.4 cm
long, cream with some red dots towards the
apex, surface with stiff hairs, stigma globose,
creamy-grey, sticky. Male bud intermediate,
apex imbricate, top-shaped and convolute at
advanced blooming, rachis with a curve. Male
bracts 10–17 × 7–10.5cm, moderately
grooved, adaxially red with yellow tip,
moderately glaucous, abaxially creamy pink,
shiny, apex acute, lifting one bract at a time,
reflex and revolute before falling, the whole
bud degenerate before maturity of fruits. Male
flowers on average 14–32 per bract in two
rows, 5.8–6.2 cm long, falling with the bract,
bract scars prominent. Compound tepal 3.3–
4.5 × 1.2–1.6 cm, yellow, ribbed at dorsal
angles, and with 5-toothed dark yellow lobes,
0.3–0.4 cm long. Free tepal 1.9–2.6 × 1–1.6
cm, translucent cream tinged with yellow,
much darker at apex and base, boat-shaped,
corrugated at apex, apex with small acumen.
Stamens 5, 3.3–4.7 cm long, exserted,
filament cream, 2–2.4 cm long, anther cream
with pink tinge, 1.1–2.3 cm long. Ovary
straight, rudiment, 0.9–1.5 cm long, creamy
white, style slightly curved at base, inserted,
2.9–3.5 cm long, stigma globose, yellow.
Fruit bunch lax, with 5–8 hands and 12–20
fruits per hand, in a two rows, fingers
arranged perpendicular to the rachis, fruits
glabrous, straight, slightly ridged, apex
pointed, without any floral relicts, immature
fruit peel color green, mature fruit peel color
dull yellow with black blotches, pulp cream.
Habitat: Growing as under growth in wet
evergreen forests.
Distribution: Assam, Nagaland, Manipur,
Meghalaya.
Specimens examined: India, Nagaland:
Zunheboto District, Nagutomi, 20 August
2011, N26
0
08.093’ E094
0
30.299’ 1297 m, A.
Joe & P.E. Sreejith 130726 (CALI!).
Conservation status: The species has been
collected from only one locality of Nagaland at
high altitudes and is very rare. Continued
decline in area of occupancy and the threat to
the population by forest fire may be a reason
for the decline of population. Clearing of hills
for Jhum cultivation and shifting cultivation
can also cause damage to the existing
population.
Phenology: In case of Musa, flowering
season differ in different altitude and in
separate climate conditions. But in our
Garden it flowers throughout the year and
produce fruits.
Etymology: The specific epithet was applied
to the species due to its flowers suffused with
yellow, in virtually complete methylation of
the bract anthocyanins (Simmonds, 1956).
Alfred Joe et al.: Annals of Plant Sciences, 2013, 02 (08), 260-267
Page | 264
Notes: This species is different from other
species in the Section in having comparatively
small pseudostem, and orange-yellow
flowers. But it shows more similarity to M.
acuminata in its appearance of pseudostem,
waxy nature towards the upper portion of
pseudostem, male bud colour and shape.
M. thomsonii (King ex Schumann)
A.M.Cowan & Cowan ‘thomsoni’, Trees of
North Bengal, 135. 1929. Karthikeyan et al.,,
Flora Indica Enum. Monocot., 104. 1989;
Noltie, Flora of Bhutan, 3(1): 181.1994. Fig.
2.
Type: India, Sikkim, Ryang, 28 November 1877, 1500 ft, G. King [Acc. No. H.0014/91-53
(K!)], G. King, (Lectotype designated by Häkkinen & Väre, 2008).
Musa sapientum subsp. seminifera
form 4. thomsoni King ex Baker, Ann. Bot. 7:
214. 1893; Bull. Misc. Info., 1894(92): 256.
1894; Cheesman, Kew Bull. 327. 1948;
Häkkinen & Väre, Adansonia, 30(1): 77.
2008; Musa paradisiaca subsp. seminifera
(Lour.) Baker var. thomsonii “thomsoni” King.
K. Schumann. In A. Engler (ed.),
Pflanzenreich., IV, 45: 21. 1900.
Plants slender, suckers 2–3, close to
the main shoot, 10–13 cm away, vertically
arranged. Mature pseudostems 4–5 m high,
30–45 cm circumference at the base, light
green with reddish-brown patches, covered
with old leaf sheaths except apex region,
apex glabrous, shiny, underlying color creamy
green with red pigmentation, sap watery. Leaf
habit intermediate, laminae 165–170 × 42–
48 cm, oblong-lanceolate, apex oblique,
truncate, cuspidate, adaxially dark green,
dull, abaxially medium green, appearance
Alfred Joe et al.: Annals of Plant Sciences, 2013, 02 (08), 260-267
Page | 265
shiny, leaf bases symmetric, both rounded,
midrib adaxially light green and abaxially
yellow-green. Petioles 46–50 cm long, slightly
waxy, petiole margins wide with erect, with
blackish-brown scarious margin, and black-
brown blotches at the base, petiole base
winged and clasping the pseudostem.
Inflorescences horizontal, peduncle 23–36 cm
long, dark green, glabrous. Sterile bracts 1–2,
deciduous, c. 62 × 16 cm, adaxially yellow-
green, slightly glaucous, abaxially creamy,
shiny, apex with leafy appendage. Female
buds lanceolate, convolute. Female bracts
32–34.2 × 13.8–14.3 cm, moderately
grooved, adaxially brown-purple with yellow
streaks, slightly glaucous, apex acute, yellow,
abaxially cream to creamy yellow, shiny, apex
acute, lifting one bract at a time, persistent
for two or three days, giving an appearance
of lifting several bracts at a time, reflex and
revolute before falling, cincinnus 16–18
flowers. Basal 8–12 hands female. Female
flowers 16–18 per bract in a two rows, 9.8–
10.3 cm long, yellow. Compound tepal 4.3–
4.6 × 1.7–1.9 cm, orange-yellow, ribbed at
dorsal angles, lobes 5, orange, outer two
lobes with small horns. Free tepal 2.9–3 ×
1.7–2 cm, boat-shaped, translucent cream
tinged with yellow, apex corrugated at apex,
with a short acumen, yellow. Staminodes 5,
2.2–2.5 cm long, cream with brown apex,
irregularly arched. Ovary 6–6.5 cm long,
straight, creamy green to light green or
yellow green, with ovules in 2 rows per locule,
style straight, exserted, 3.7–4 cm long,
cream, stigma globose, creamy-grey, sticky.
After female bunch there is a transitional
bunch. Male bud lanceolate, intermediate to
top shaped at advanced blooming, convolute,
rachis falling with a curve. Male bracts 16–18
× 8–12 cm, moderately grooved, adaxially
brown-purple with yellow striations, apex
yellow, moderately glaucous, abaxially cream
to creamy yellow, shiny, apex acute, lifting
one bract at a time, reflex and revolute
before falling. Male flowers on average 16–20
per bract in two rows, 5.2–8.3 cm long,
falling with the bract, bract scars prominent.
Compound tepal 4.6–6.3 × 1.1–1.5 cm,
orange-yellow, ribbed at dorsal angles, and
with 5-toothed orange lobes, 0.5–0.7 cm
long. Free tepal 2.1–2.8 × 1.2–1.5 cm,
translucent cream, boat-shaped, corrugated
at apex, apex with small acumen, yellow.
Stamens 5, 4.5–5 cm long, exserted, filament
cream, 1.6–2.3 cm long, anther cream with
pink tinge, 2.3–3.4 cm long. Ovary straight,
rudiment, 1.2–2.1 cm long, creamy white,
style straight, inserted, 4–5 cm long, stigma
globose, creamy yellow. Fruit bunch lax, with
8–12 hands and 16–18 fruits per hand, in a
two rows, fingers arranged perpendicular to
the rachis, curved to one side, fruits glabrous,
straight or curved, pronouncedly ridged, apex
pointed, without any floral relicts, immature
fruit peel color green.
Habitat: Growing as under growth in moist
ravine black humous at evergreen forest and
also near water streams.
Distribution: Sikkim and Meghalaya, North-
East India.
Specimens examined: India. Sikkim:
Ryang, 1875–76, 1500 ft, G. King, [Acc. No.
H.0014/91-50 (K!)], 1875–76, 1500 ft, G.
King, [Acc. No. H.0014/91-51 (K!)], 1875–76,
1500 ft, G. King, [Acc. No.H.0014/91-52
(K!)], Drawings (4 sheets), 1500 ft, G.King,
75/51 (CAL!), Meghalaya: Umkiang, Jaintia
Hills, Jamsara, 02 May 2011, 154 m, A. Joe &
P.E. Sreejith 116177 (CALI!)
Notes: This species is different from other
species in the Section in having bracts purple
with yellow striations outside and cuspidate
leaf apex.
Table.1: Major differences between M. thomsonii and M. flaviflora
Characters M. thomsonii M. flaviflora
Pseudostem height 4–5 m high 1.8–2.1 m high
Pseudostem color Light green with reddish-brown patches Green with small black blotches
Bract color Adaxially brown-purple with yellow streaks and
abaxially cream to creamy yellow
Adaxially red and abaxially cream
with pink tinge
Female flower length 9.8–10.3 cm long 6.5–8.5 cm long
Compound tepal in
female flowers
4.3–4.6 cm long, orange-yellow 3.5–4 cm long, yellow
Free tepal in female
flowers
2.9–3 cm long 2.5–2.7 cm long
Staminodes 2.2–2.5 cm long 1.3–2.1 cm long
Ovary 6–6.5 cm long 3.8–4.1 cm long
Style surface Smooth Rough with stiff hairs
Male bud Not degenerative after the maturity of fruits Degenerating before maturity of
fruits
Alfred Joe et al.: Annals of Plant Sciences, 2013, 02 (08), 260-267
Page | 266
Conclusion
“Musa taxonomy is still very obscure
today just as it has been throughout its
history despite attempts to clarify it”
(Häkkinen 2008). Here authors concluded
that both the species have their own identity
and are distinct with the taxonomic confusion
belonging to M. flaviflora and M. thomsonii is
solved, but also suggest further studies of
these two taxa, both by cytogenetically
studies and molecular studies because of
being relation with M. acuminata.
Acknowledgements
The authors are grateful to the
Department of Science and Technology (New
Delhi) for the financial assistance for the
research projects on Indian Marantaceae and
Musaceae (Sanction No. SR/SO/PS-115/09,
dtd 19.08.2010). The authors would also like
to thank to Mr. Markku Häkkinen (Finnish
Museum of Natural History, Botanic Garden,
University of Helsinki) for the valuable opinion
and the identity of these species through
correspondence in June 2011 and further
during his visit to Calicut University in 2012.
Also thankful to the officers of the
Forest Department, Assam, Meghalaya, and
Nagaland for granting permission and
providing necessary help for the field studies
in the forest. Thanks are also due to Dr. S. K.
Chathurvedi for his help during the collection
of specimens.
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Source of support: Department of Science and Technology, New Delhi
Conflict of interest: None Declared