PHYTOTAXONOMY

Vol. 1,2001, pp. 5-17

Study of floristics and plant taxonomy

D.B. Deb
Central National Harbarium, Indian Botanic Garden, Howrah-711 103.

Some aspects of taxonomic work of the author, particularly on genera Nymphaea, Aldrovanda,
Solarium, Clarkella and Polyura are reviewed. Distinguishing characters between the genera
Keenania, Mycetis and Myrioneuron are summarised.
Keywords: Nymphaea; Aldrovanda; Solanum; Clarkella; Keenania; Mycetis; Myrioneuron;
Polyurs; Tribe Clarkellaeae
My revered teacher, Padma Bhushan, Prof. typed for publication. Then, he pointed out the genus
A.K.. Sharma, formerly President, Indian National Dracaena in the thesis and told me that the correct
Science Academy and General President, Indian name is Pleomele, and asked me to find out if a
Science Congress, Dr. S.K. Jain, FNA, Director, new combination would arise. In 1957,1 wrote to
Institute o f Ethnobiology and President, and Fr. Santapau seeking his views if the Flora of
members of Association for Plant Taxonomy, my Tripura State could be a thesis for D.Sc. He wrote
colleagues, officers and staff of Botanical Survey back “Every word you write should be worth a Naye
of India, ladies and gentlemen: Paisa”. I tried to follow his advice. As a token of
I feel highly honoured with the award of the my regard for him, I jointly with Rasamoy Dutta
first Dr. H. Santapau M edal instituted by the dedicated a new genus “Pauia” in his honour.
Association for Plant Taxonomy. Fr. Santapau On this occasion, I am duty bound to name
initiated Indian workers to Botanical Latin and Krishna Chandra Malick with whom I initiated
International code of Botanical Nomenclature, and taxonomic revision of the genus Rubia in 1965 and
encouraged them to undertake intensive field my Ph.D. students in Plant Taxonomy* Syamali
studies. I am now reminded of my contact with him. Dasgupta, Ratna Dutta, Durga Charan Mondal,
It was a spring morning in the last week of Mohan Gangopadhyay and Ramesh Chandra Rout
M arch 1955, I went to Dr. I. Banerji, Head, who in course o f their studies helped me in
Department of Botany, Ballygunge Science College, developing my taxonomic concepts.
with a copy of my Ph.D. thesis to be submitted. Prof. Introduction
G.P. Mazumder was with him. Dr. Banerji told me In the D.M. College, Imphal, established in
that the newly appointed Chief Botanist, BSI, Fr. 1945 by the then M aharaja o f M anipur, Arts
Santapau would be visiting the Department at that subjects, including Mathematics, were taught up to
time. He desired to show him my thesis and asked the Honours course. In 1951, Government o f
me to sit in the anteroom of the Library where tea Manipur, then a part C State, started Intermediate
party was to be held. Sunanda Banerjee (now Dr. Science classes in four subjects: Chemistry, Physics,
Mrs. Pal) also sat there with her research material. Botany and Biology. The appointees were asked to
Fr. Santapau asked me to give him the thesis on his join on 15 June, during the Summer vacation of the
way back and to meet him in the Indian Museum College for organizing the laboratories. I was
after two days. When I met him there, he gave me a appointed in Botany. A bearer for the Department,
typed paper and showed me how a paper was to be Shri Ibodi Singh, was appointed earlier. In the
First H. Santapau Medal lecture, delivered at Calcutta on 24 February 1999.
6 PHYTOTAXONOMY, VOL. 1,2001
course of organizing the Laboratory, I felt the
n ecessity o f know ing the local plants for
dem onstration in teaching. This collection o f
specimens became my essential duty for proper
teaching. This gradually resulted in exploration of
Manipur State and working out the Floristic study
of Manipur. This was the first Ph.D. thesis on
Floristics of a state from any Indian university.
In the course o f exploration o f M anipur,
Tripura, Mizoram, Arunachal Pradesh (Tirap Dist.)
Khasi & Jaintia Hills, Garo Hills, North Cachar
H ills, N orth East Assam , N agaland, B hutan
(Eastern), Sea coasts and South Indian Hill Ranges,
I co llected over 12000 F ield N um bers, 4-6
specimens each. Efforts to collect more than a single
specimen failed in the case of Sapria himalayana
and Rauvolfia chinensis; in the case of the latter, a
single flowering plant could be collected, there
being no other plant growing nearby.
These explorations yielded three new genera,
13 new species and four infra-specific taxa. Of these,
I have already mentioned the mono-specific genus,
a Solanaceous plant. M atured fruits were not
available. I could not visit the locality for the second
time, nor could I get it collected by anybody in BSI
or the Forest Department of Anmachal Pradesh.
Ripe fruits still now remain unknown. Some of these
studies are summarized here.
On Nymphaea
The first scientific note entitled ‘New Record
of epiphyllous bud on Nymphaea stellata Willd’.
was published in Set & Cult., Dec. 1952. While
com m unicating the note, I used a note o f
interrogation after the name, as I had doubt about
the determination. But the authorities of the journal
deleted the question mark. I was not convinced. I
forwarded the specimen to RBG, Kew, where it was
determined as Nymphaea micrantha Guill. & Perr.
On consulting literature, I noted the synonymy as
follows:
Nymphaea micrantha Gull. & Perr., FI. Seneg.
Tent. 1: 16. 1830, A. Coerulea Guill., & Perr. l.c
non Savigny, 1802 N. rufescens Guill. & Perr. l.c.
N. stellata var. bulbifera Planch, in F I. Serres.
8:286. 1852. N. vivipara Lehmann, Nov. PI. Hort.
Bot. Hamb. 11. 1852. N. guineensis Schumach &
Thonn. Beskr. Ghip P 1. 218. 1829
It is evident form the above that N. stellata var.
bulifera Planch, is also one of the synonyms.
On Aldrovanda
Aldrovanda vesiculosa L. Sp. PI. 1: 281. 1753
& Gen. PL ed. 5, no. 350. 1754. Wt. & Ann. Prodr.
34: 1834; C.B. Clarke in Hook. f. FI. Brit. Ind. 2:
424. 1878; Prain. Beng. Pl. 1:342 1903 & in Rec.
Bot. Surv. India 2: 305. 1903 & 3: 210. 1905;
Sengupta in Sci. & Cult. 3: 97. 1937; Ven Steenis
FI. Males Ser. 1, 4(4): 381. 1953; Deb in Curr. Sci.
26: 229 1957 and in Bull. Bot. Surv. Ind. 3: 327.
1961. Basak in Bull. Bot. Surv. Ind. 17 (1-4) : 97.
1975; Kunda etal. in J. Econ. Tax. Bot. 20 (3): 719-
724. 1996. A.verticillata Roxb. FI. Ind. (ed. Carey)
2:113,1832 & Icon 1129 Lenticulapalustris Indica
Plukenet, Photographia 1691-1692 & Alamagestum
Botanicum 1696.
In July 1953, it was raining heavily in Manipur.
The rivers were overflowing the banks. Most parts
of Imphal valley were submerged. The vast college
surrounding was flooded. On 2nd August, the
Sunday, along w ith Shri Ibodi Singh, the
departmental bearer, I waded through the flooded
foot tracks in Thangamaiband to look for plants
growing in and floating with flood water. We
proceeded towards the newly renovated college
Reserve Tank. To my surprise, I saw colonies of
Aldrovanda vesiculosa submerged and floating here
and there along the sides of the pond. Earlier, I saw
this plant preserved in a jar in the Science College,
Ballygunge Circular Road, Calcutta. Just on seeing
the plant, I could identify it. The colonies were of
two types: one green with greenish white flowers
and the other pink, with pinkish stem, leaves and
flowers. On observing the plants o f different
populations, I felt that there m ust be genic
differences in the seeds producing the plant
populations, even though it may not attain a
taxonomic difference. I collected a large number of
plants, grew them in the laboratory in that water and
preserved in jars in preservatives. I sent specimens
to Prof. P. Maheshwari, Delhi University, to Indian
 STUDY OF FLORISTICS AND PLANT TAXONOMY
Botanic Garden and to RBG, Kew for views on the
colour differentiation. Director, RBG Kew, as
expected, wrote back that the differences in
p ig m en tatio n were not taxonom ically
distinguishable. The plant did not grow in the
laboratory. Nor did it reappear in subsequent years.
This plant was initially sent from Calcutta in
1690 by Du-Bioss of East India Company to his
friend James Petiver, a famous aporthecary in
London. He gave it to Leonard Plukenet, Royal
P ro fesso r o f B otany, who w orked it out as
Lenticularia palustris Indica in Phytographia 1691-
92 and in Alamagestum Botanicum, 1696, the
specimen being deposited later by Sir Hans Slonne
in the British Museum. In 1947, Gaetano Carle
Amadi collected such a plant form Bologna, Italy,
and it was named after the Italian Naturalist, Ulisse
Aldrevandi, by Mont. This was identical with the
Indian material and Carolus Linnaeus described
these gatherings as Aldrovanda vesiculosa Linnaeus
(1753-1754).
William Roxburgh described and illustrated it
in 1812 as Aldrovanda verticillata Roxb. FI Ind.
(ed. Carey) 2: 112. 1832 & Icon no. 1129 (CAL).
Thomson was the first botanist to collect the plant
from salt pans in South of Calcutta (now Salt Lake)
and Malta river at Port Canning on 10 November
1855. He did not see the plant flowering. S.Kurz
collected the plant from Malta river (Canning) in
November 1869, while M.S. Ramaswami collected
it again from Saltlake margin on 12 July 1916. All
these gatherings are from brackish water. J.C.
Sengupta (1937) collected on August 1936 between
Tarpassa and Boloi, Vikrampur, Dacca (now
Bangladesh) growing in sweet water. I was informed
by my friend, Md. Allarakha, in 1946, that this plant
was found in Rajshahi (now Bangladesh). I wonder
if it appeared there in the same year in which
Sengupta reported it form Dacca. The Professor of
Botany, Rajshahi College could not throw any light
on the matter in reply to my letter. One Dr. Biswas
working oii a thesis under Professor R.S. Mishra of
Banaras Hindu University reported the occurrence
of this plant in Krishna Nagar, West Bengal. Prof.
Mishra, in reply to my letter, informed me that Dr.
7
Biswas was then in USA on a post-doctoral
fellowship at that time. He must have collected the
plant when he mentioned. I had a feeling that it
was copied from Bengal Plants by D. Prain by
changing the name of the locality. That area has
been explored very thoroughly, but no such record
is there by anybody else.
Roxburgh himself did not see the plant in field.
His observation that it was swimming on ponds of
water over Bengal during the hot and cold seasons
appears to be erroneous. David Prain cited the plant
in three publications (1902, 1903, a, b) based on
collections by earlier workers cited above. How it
received the vernacular name “Malacca Changi” in
Bengali could not be traced out. From South East
Asia it was reported from Timor only. Basak (1975)
reported on the distribution of the plant in the world.
Kundu et al. (1996) attem pted a review and
speculated on the germ plasm collection and
conservation at Sunderban Biosphere Reserve:
All the collections of the plant made from West
Bengal were from salt pans, now* known as Salt
Lake, and the river in Canning. There is no record
of the plant growing for two or more successive
years in nature. Caspery’s report on cultivation in
cultural media is not clear. Wherefrom did the plant
come to Imphal?. The seeds must have been carried
by the w ater current flow ing from the sub-
Himalayan rivers and rivulets connecting Barak that
flows through the state. Sporadic occurrence of this
plant cannot be properly explained with the
available information.
Ecology: Sengupta reported on its occurrence
in association w ith Utricularia flexuosa,
Myriophyllum sp., Mariscus cyperoides, Hygrorhiza
sp., Riccia sp., etc. Since the small plant is slightly
submerged and floating on water at margins of water
pools, it is only a coincidence that due to slow
movement of water due to wind or flow of water or
any other reasons it comes close to some aquatic
plants only. It may not have any obligatory
significance at all.
It appears to me that the seed remains dormant
for an exceedingly long period and geminates only
in a microclimate that does not prevail in any locality
8 PHYTOTAXONOMY, VOL. 1,2001
every year. Probably, the attainment of a particular
microclimate or environmental condition is the
limiting factor for appearance of the plant.
On th e O rig in s of Egg P la n t (Solatium
melongena)
Linneous (1755) described S. melongena on the
basis o f plants cultivated in Asia, Africa and
America. C.B. Clarke treated S. incanum and S.
insanum as synonym ous with S. melongena.
Bhaduri (1951) observed that S. incanum and S.
insanum are the nearest ancestors of S. melongena.
I treated S. incanum and S. insanum as varieties of
S. melongena (Deb 1978) and presented the
distribution of these varieties in a map of India. I
was requested by Mr. M. Ikebe of Tokyo, the leader
of a Japanese exploration party, to accompany them
along my distribution map in South India. We
conducted field studies in Karnataka, Kerala and
Tamil Nadu and studied over 100 populations and
grew some plants in IBG and at Salt Lake. Plants in
varying habitat, growth and phenological conditions
were studied in the field. The plant varied in habit
from small prostrate or procumbent herbs to bushy
shrubs (PI. 1) The growth is stunted in rocky or
sandy soil. The same plant bears fruits of the same
colour, form and size or of varying colour, forms
and sizes on different branches or on two sides of
the main stem. Some plants bear white fruits of the
same form and size as the hen’s egg (PI. 1, Ph.7)
or others bearing white fruits on one side of some
branches, and violet or white with greenish patches
on the other side, the form and size remaining same
as the hen’s egg or in another population the colour
may be violet or yellowish. Likewise, the form may
be roundish or oblong or elongated. These variations
throw light on v ariab ility , taxonom y and
relationship of taxa to such an extent as to obliterate
the known distinctions observed on assessment of
herbarium specimens or on cultivating plants in
field.
Feral forms of S. melongena are encountered
which are characterised by more prickly stem, leaf
and calyx, reduction in pulp of the fruit and poor
edibility.
Sometimes, it is procum bent or diffused,
scarcely branching or bushy, variable in density of
prickles, pubescens size, form, thickness, location
and colour of leaves, branching of the peduncle,
number of bisexual flowers or axillary or extra-
axillary peduncle, flowers bisexual or unisexual,
and/or moncecious, usually pentamerous, some­
times, tetra or hexamerous. Corolla is more or less
violet or white with violet veins. Fruits are white,
golden, yellow or tinged with green, round, ovoid
or obovoid or egg-shaped, White pendulous fruits
of the size, shape and form of a hen’s egg on the
plant at one reminds one as to how appropriate the
name egg plant is. The variations are evident not
only in the same population but also on the same
plant, eliminating thereby the distinction between
taxa even at the varietal level. Such a conception of
variability cannot be had on the study of herbarium
specimens only. This leads me to believe that in
critical cases, toxonomic study cannot be done
properly without thorough field knowledge.
From this field study, I am fully convinced that
Solanum incanum is the wild form of Solanum.
melongena, or, in other words, Solanum melongena
in cultivation arose from S. incanum and that S.
insanum is synonymous with S. incanum.
T rib a l P o sitio n of th e G en u s Clarkella
(Rubiaceae)
J.D .. H ooker (1880) p ostulated the
monospecific genus Clarkella in honour of C.B.
Clarks. The genus is based on Ophiorrhiza nana
Edgeworth (1860), which stands out as very
different from the character states of the genus
Ophiorrhiza L. While describing the genus, J.D.
Hooker observed “A very singular little plant
requiring examination in a living state as to the mode
o f grow th” . He treated the genus in tribe
‘Oldenlandeae’ (Hedyotideae). Bremekamp (1966
: 28) treated this genus along with Argostemma
Wall, and Neuroclayx Hook. f. in Argostemmatideae
differing from the Hedyotideae in the connivent
anthers opening with a short slit or a pore and in the
insertion of the stamens at the base of the corolla
tube. In Clarkella, anthers are not connivent.
Robbrecht (1988, 1993) kept the genus among the
genera incertae sedis.
 STUDY OF FLORISTICS AND PLANT TAXONOMY 9

Plate Is Wild brinjal (Solanum melongena L. syn. S. incanum L. and S. insanum L.)
(1) A branch with pentam erous violet flower and stout recurved prickles (Bangalore to Ootacamund).
(2) A branch bearing a violet flower with 4 stamens (Ootacamund to Cochin). (3) An erect
branch with light violet flowers and less prickly leaves. (4) A branch with globose yellow fruit.
(5) A branch with oblong fruit. (6) an erect branch with white flowers and less prickly leaves.
(7) A branch bearing fruit of the form and colour of a hen’s egg. (8) A branch bearing white
flowers, and ablong fruit, white tinged with green. (9) A branch bearing white flowers and
fruits white or white tinged with green lines (patches). (10) A branch with white and light violet
flowers, and oblong fruits mostly white in colour with greenish tinge.
 10 PHYTOTAXONOMY, VOL. 1,2001
T uberous epiphytes o f the R ubiaceae
representing the tribe Psychotrieae have long been
a subject o f interest because o f their complex
chambered tubers, which are often inhabited by
insects. The presence o f a tuber in Clarkella
distinguishes the genus form others in Hedyotideae,
Argostemmatidae and Ophiorrhizeae. The tuber of
Clarkella has not been studied so far. No insect
inhabits this tuber.
Generico-Specific Description
Clarkella nana (Edgew.) Hooker f. PI. Brit. Ind.
3 :46.1880; Schum. in Engl. & Prantl, Pfanzanfam.
4 (4): 31. 1891 William in J. Bot. 44: 377, 1908,
Ophiorrhiza nana Edgew. in Trans. Linn. Soc. 20:
60. 1860 (Type : North India, Halluyoan, on rocks,
1500-1800 m, 1842, Edgeworth s.n. kl) Clarkella
siamensis Craib in Kew Bull. 1931: 216. 1931.
(annotated and treated by L.A. Lawner and D.K.
Ferguson (E) in 1972 in sched., as a synonym, and
confirmed by Mrs. Diane M. Bridson (K) in Lit.).
Herbs 1.0-12 cm long, very delicate, simple
rarely branched, sometimes curved upwards, hispid,
with a tuber, 1.5-2.8 cm * 1.3-1.8 cm ovoid,
cylindric or elliptic, narrow ed at both ends,
sometimes slightly irrfegular in form with a dense
cushion o f fibrous roots (PI. II) some of which
adhere to the surrounding particles of stones; stem
quadrangular, winged at angles, extending to the
base of the petiole connecting the stipules. Leaves
opposite, 1-3 pairs, petiolate, the lowest the largest
Plate II: Clarkella nana (Edgew.) Hook.f. L.S. of
a tuber showing two parts
with the longest petiole; one leaf of the lowest pair
is often suppressed, the other very large, sometimes
3 leaves at a node, the third being derived from
modification of stipule. Radical leaves with larger
petioles 2-3 cm long, solitary, orbicular-oblong,
ovate or cordate, obtuse or acute, attenuated and
sheathing at base on the top o f the tuber,
Membranous; lateral nerves 3-6 pairs, spreading,
arching; petiole 1-6 cm long, slender, a single pair
of small leaves close to the cyme; stipules ovate-
lanceolate, ca. 3 mm long; bracteoles linear-
lanceolate, 2-3 mm long. Inflorescence terminal,
peduncled, bracteolate cymes. Flowers 1-6, erect,
1.25 long; pedicels upto 3 mm long, erect. Calyx
persistent. Hypanthium rigid, obconic; limb-tube
very short, dilated above, distantly reticulated, 5-7
toothed, corolla white, infundibuliform, ca. 12 mm
long, tube long; slender; lobes 5, lanceolate, valvate.
Stamens 5, attached at the base o f the corolla tube;
filaments slender, 0.2 mm long, anthers ca. 1 mm
long, linear-oblong, dersifixed; pollen 3-4 colporate.
Ovary 2-loculed, obconic {ca. 40-50% of the ovules
form seeds); ovules many on ascending placenta,
attached to the septum below its middle, style 1mm
long, arms 2, slender, ca 1.3 mm, pubescent.
C apsule obconic, subcoriaceous, 5-7 ribbed
crowned with dilated calyx limb, dehiscing seeds
through minute pores. Seeds many, minute tectum
perforatum. Embryo with the pair of cotyledons
remaining coherent above, forming a heart-shaped
structure. The hypocotyl forms the tuber. Ffe.;June
- October; Frts; : July-March.
Distribution: A monospecific genus distributed
in Western Himalayas and Thailand. Saxicolous on
limestone rocks.
Morphological study: (/) It is evident from some
herbarium specimens that the hypocotyl swells and
forms a protuberance that gradually grows in size
and forms a small tuber. It is brown in colour with
greenish patches here and there and is fleshy in
texture. The surface is often areolate. Alveoli appear
here and there on the surface, through which
adventitious roots grow and sometimes form a more
or less dense mass.
Phellogen aries de novo in the parenchyma.
 STUDY OF FLORISTiCS AND PLANT TAXONOMY
Initially, it is hook-shaped. It grows in size rapidly
and form s a basal opening. F urther discrete
phellogens arise at several points, each forming a
new cavity and opening outside in the form of holes.
Later cavities may or may not connect with the
existing cavities. Smaller holes to the outside are
also formed. Small cobweb-like chambers or cells
are also present adjacent to the cavities. Two or three
smaller cavities adjacent to the largest one are also
present. Besides, a large number of holes are visible
from the surface all over the tuber. Transverse
section of the tuber shows cobweb-like structure.
In longitudinal division of the tuber, growing points
are seen wherefropi tissues are formed in the tuber,
sometimes lookup like Scales (Plate III). Raphides
are very, common in the tuber.
(ii) Roots: While the hypocotyl forms the tuber,
the radicle continues to grow below it and becomes
ramified. Soon, several adventitious roots larger ind
longer are also formed below the tuber and persist
with it. They are wiry, brown in colour and are
m ostly branching, som etim es thicker or not,
provided with root cap and hairy at the apex, base
and at different heights. The hairs are fine, long or
short, numerous at the apex, looking like a club-
shaped or foliaceous structure under the magnifying
glass, but on higher magnification, 400 times or
more under the light microscope, they are clearly
distinguishable as hairs of various lengths, and very
often remain adherent with the stone particles in
the substratum , sorving as a hold fast, and
simultanously absorbing raw food material for the
growing plant.
As the tuber grows in size, the part of the
seedling just above the tuber grows in length and
thickness. Adventitious roots are also formed on it,
as on the tuber.
(iii) Seedling: The part of the axis above the
tuber produces initially a membranous sheath
bearing the first pair of interpetiolar stipules looking
at an early stage as if the sheath bears four teeth.
Thi^ does not grow in size or form. But the stem
grows through the sheath and forms the vegetative
^arid floral parts o f the plant in due course.
11
(iv) Pericarp: Pericarp of the fruit shows three
distinct zones] the epicarp is composed of a single
layer of compact cells provided externally with
unicellular or 2-3 cellular trichomes. The mesocarp
consists o f several layers o f large cells,
isodiametrial, elongated or irregular cells. ^Towards
the mid-height of the fruit, it is provided with very
large airspaces here and there and bears vascular
bundles. The endocarp is also single layered in
thickness consisting of parenchymatous cells.
(v) Seeds reticulate, paillate; reticula 30-40 pm,
Cell surface psilate, In dry condition, the papillae
shrink, showing a pattern having concentric rings.
Exotesta is one cell in thickness, of rectangular
thick-walled compact cells; the endotesta is of thin
walled parachymatous cells one or more cells in
thickness.
The endosperm, as observed at a matured stage,
is cellular, occupying about two thirds of the seed,
while the embryo occupies the remaining part.
(vi) Microcharacters: Pollen grain 10 x 20 pm
oblate, sp heroidal, m esocolpium 12 pm,
apocarpium 15 pm, anti-circular 4-6 monocolporate,
actocolpium 12 * 1pm, endocolpium2 * 5pmaxine
1-2 pm thick, tectate, collumeliate; asine surfaces
reticulate, heterobrochate, lumina less than 1pm,
muripsilate.
Discussion
In recent years, Huxley (1978, 1993), Huxley
& Jebb (1991 a, b, c) and Jebb (1991) conducted
detailed studies on the tuberous Psychotrieae and
postulated the subtribe Hydnophytinae (Huxley &
Jebb 1991) com prising 5 genera, nam ely,
Hydnophytum Jack, M yrmocodia Jack,
Myrmocophytum Becc., Squamellaria Becc. and
Anthorrhiza Huxley & jebb. These genera are
woody epiphytes on the coastal trees, with large
tuber usually inhabited by ants, hi Hydnophytinae,
the tuber varies from 8x15 to 50 cm across. The
growth is apical, producing cylindrical tubers. The
tuber surface is brown in colour and is often
areolate; otherwise it is smooth. Spines are present
on the tuber and the stem. They vary from simple
to repeatedly branched or regularly stellate or club-
 12 PHYTOTAXONOMY, VOL. 1,2001

Plate III: Clarkella nana (Edgew). Hook.f. A plant excluding the tuber; B. Flower split open showing
floral parts (stamens at the base of the corolla tube); C. Pistil; D. Capsule with persistent calyx;
E. Pair of cotyledons united above; F. Tuber with roots; G Adventitious root with cluster of
hairs at places; also attached to stone particles all around; H. T. S. of pericarp. I. Seed (part) in
surface view x 450
 STUDY OF FLORISTICS AND PLANT TAXONOMY
shaped. On the stem, they are most developed round
the entrance holes and on ridges.
In Clarkella, as already mentioned, the tuber is
very small and delicate. Young areas are greenish.
Adventitious braching roots on the tuber are never
spines. They bear hairs here and there and at the tip
which appear club-shaped under the magnifying
glass, but on 400 times or more magnification
minute hairs of varying length are distinctly visible,
some remaining adherent to the stone particles all
around and clasping them. Length of hairs in a
cluster depends on the form of the rock particles to
which they cling.
Forbes (1880,1885) showed that in Myrnocodia
tuberosa Jack, the tuber and cavities are formed
even in the absence of ants. This was confirmed by
Treub (1888) from the anatomy of warts that their
function is aeration of the tuber tissues. He was
convinced by cultivation experiments that ants play
no role in the growth of this plant. Retting (1904)
proposed that the cavities insulate the inner parts of
the tuber from excessive heating and also catch and
absorb rain water. Spanner (1938) concluded that
the main function of the tuber is water storage.
Clarkella is saxicolous on limestone rocks in
temperate climate. Flowering is over by October.
The fruit dehisces minute seeds before the onset of
winter. In the habitat of Clarkella, there is scarcely
any water in winter. Whatever water is present there,
solidifies in winter. In that situation, the tuber is
formed by the hypocotyl. Cobweb-like structure
with cavities and pores is formed in the tuber. No
ant inhabits the cavities. Thus, the tuber is an
adaptation for ‘presentation o f the seedling in
unfavourable environmental conditions. It stores
water, air and food material for sustenance of the
dormant embryo. By May, the weather condition
changes, the tuber at the apex bears adventitious
roots like those on the surface of the tuber, and soon
the stem bears leaves and stipules, ultim ately
producing flowers and fruits.
H uxley & Jebb (1991) did not find any
d istin ctio n o f the tuberous P sychotrieae in
reproductive characters from other genera of the
P sy chotrieae, but estab lish ed com m on
13
characteristics of the tuberous genera as almost
certainly homologous in characteristics of tubers
and their cavities, suggestive of having a common
origin and also hom ologous in vegetative
morphology, ecology and distribution. For these
reasons, they treated these genera under a subtribe
Hydnophytinae.
Proximity of Clarkella with Other Genera and
Tribes
H erbaceous habit, presence o f raphides,
interpetiolar stipules, terminal inflorescence, corolla
lobes valvate, flowers isostylous, ovary 2-locular
w ith many ovules present in Clarkella are the
characters common to Hedyotideae, Ophiorrhizeae
and Argestemmatideae which are more closely
allied than other tribes in the subfamily.
Capsule laterally flattened is a unique character
w hich distinguishes O phiorrhizeae from
Hedyotideae and Argostemmatideae.
Monocaulous dwarf, monomorphic flowers,
stamens attached near corolla bases, anthers mostly
united into a cone, fruit dry indehiscent or opening
with operculum are the character states which
distinguish Argostemmatideae form Hedyotideae
and Ophiorrhizeae.
Production of a small tuber with large and small
chambers and holes at the base of the plant for
peVennation o f the seedling in unfavorable
environmental condition; stamens inserted near the
corolla base, capsule obconic with 5-7 ribs and
persistent calyx, dehiscing seeds late through pores,
seeds papillose are the character states of Clarkella,
which do not fit in any of the tribes Hadyotideae,
Ophiorrhizeae or Argostemmatideae. Besides these,
Saxicolous habit o f the genus in temperate climate,
pair of cotyledons undifferentiated above forming
a heart-shaped structure, stem quadrangular and
winged, etc. add to the distinction of the genus to
be accommodated in a unigeneric tribe Clarkellaeae
Deb tribus nov.
Clarkellaeae Deb tribus nov
Different a Hedyotideae, Ophirrhizeae, quo noc
non Argostemmatideae in plant’s habitu saxicolis,
hypocotylibus tuberasciantibus perquo plantulibue
 14 PHYTOTAXONOMY, VOL. 1,2001
orientibus, capsules crassiusculis, dehiscentibus
peris, seminibus papillis, interalia.
Unigeneric tribe based on unispecific genus
Clarkella Hook.f. growing on limestone rocks at
1800-2000 m in the W estern Him alayas and
Thailand. In India, the plant has been collected in
several localities from Kumaon to Mussourie. It is
likely to occur in the interventing region having
similar habitats.
Confusion in Some G enera
G e n era confused: The genera Keenania ,
Mycetia and Myrioneuron have long been confused
and variously interpreted.
Wild tea was discovered during the Assam Tea
Expedition in 1835-36 by Nathaniel Wallich and
W illiam G riffith from M uttack C hhara near
Dibrugarh, Assam. As a consequence, tea gardens
were soon established in the eighteen eighties,
throughout the eastern and north-eastern ranges of
Lower Hills from Sylhet to Sibsagar district. In the
course of the establishment of tea gardens, the face
of that region suffered a setback due to pulling down
of the tropical rain forest. Once a rain forest is
destroyed on a large scale, it is lost for ever. It is
obvious that herbaceous plants discovered prior to
that period were nearly lost, particularly those which
did not have large communities or were too much
restricted.
This was the fate of Keenania modesta Hook
f. (1880). This plant was originally collected by R.C.
Keenan once attached to the Royal Botanic Garden,
Kew, from Duarbund, Cachar, Assam in 1873 and
named by J.D. Hooker in 1880. Only two specimens
are preserved in Kew, so far as the record goes. J.D.
Hooker did not find pollen in the anthers of the
flower and he assumed it to be a female plant. Now,
we know that protandry is very common in the
Rubiaceae; where pollens are discharged before the
maturity of the gynoecium. Bakhuizen f. (1975) and
Van Steenis correlated Keenania with Mycetia and
described the fruit of Keenania form Malesian
plants. Those fruits cannot represent the genus
Keenania. I explored the locality from which
Keenania was collected. It is full of tea gardens.
The original rain forest is lost leading to the drastic
change of environment in the locality. There is no
trace of the genus Keenania.
Due to inappropriate correlation, a well defined
genus Myrioneuron was ignored and it did not find
a place in Index N om inum G enericorum
(Plantarum). Recently, Diane Bridson (K) drew my
attention, in a personal communication, to the
situation, on examining some Indian m aterial
determined by me as Mymoneuron nutans. She
further observed “Neither Dr. Brummit, nor I find
any problem in accepting Myrioneuron R.Br. ex.
Hook. f. 1873 as valid”, Dr. Dan H. Nicolson,
Nomenclatural Editor, Taxon, advised me in a
personal communication to publish a note on this
genus for inclusion in Index Nominum
Genericorum.
A com parison o f Keenania, Mycetia and
Myrioneruron is made in Table 1.
It is evident from the above comparison that
these three genera are distinct. But a further search
into literature brought to light a different situation.
Keenania is merged with Leptomischus Drake
(1895), which includes Polysolenia Hook. f. and
Indopolysolenia Bennet also as synonyms. Further,
H.S. Lo (1993) reduced our Indopolysonelia
burmanica Deb et Rout (1990) to the type species
Leptomischus primuloides Drake. But the workers
on Leptomischus remained silent on the type species
of Keenania, which deserves a new combination, as
presented below:
Leptomischus Drake in Bull Mus. Nat. Hist.
Natur. Paris 1:117 1895 & in Morot, J. de Bot. 9:208
1895; Pitard in Lecomte, FI. Gen Indo-chine 3:72,
f. 7(10-12) & 8(1-2). 1922; H.S.Lo in Bull. Bot. Res.
6(4):49 1986 & in Acta. Phy to tax. Sin. 31 (3): 273,
1993
Polysolenia Hook. f. in Benth & Hook f. Gen.
PI 2:68; 1873 & FI. Brit. Ind. 3:94 1880 non Ehren
ex Kutzing (1849) a later homonym, nom illeg.
Indopolysolenia Benet in Ind. For. 107 (7): 437
1981
Keenania Hook.f. FI. Brit. Ind 3: 101 1880
Myrioneuron auct., non R.Br. ex Kurz; Bakh.f.
For Bull Thai 9:26 1975
 STUDY OF FLORISTICS AND PLANT TAXONOMY 15

Table 1 : Significant Differences amongst Allied Genera

Keenania Hook.f. Mycetia Reinw. Myrioneuron R. Br. ex Hook.f.
1. Subherbaceous shrubs; bracers Shrubs; branches with conspicuous Small shrubs; branches with
without spongy swollen spongy swollen corky bark. . conspicuous spongy swollen
corky bark. corky bark.

2. Leaves membranous, without Leaves membranous, with marginal Leaves coriaceous or

marginal stalked glands. stalked glands. subcoriaceous, w ithout
marginal stalked glands.

3. Stipules somewhat recurved Stipules rather large membranous or Stipules large, coriaceous
membranous, entire, without subcoriaceous tardily caducous with erect, bifid above* without
marginal stalked glands. marginal stalked glands, sometimes marginal stalked glands.
toothed and bifid above.

4. ‘ Inflorescence in terminal heads. Inflorescence terminal or axillary Inflorescence terminal or

peniculate. rarely axillary.

5. Peduncle short. Peduncle short or long. Peduncle short, stout.

6. Flowers sessile, heterostylous Flowers pedicelled, bisexual, Flowers short pedicelled,

heterostylous sometimes biformous bisexual, not heterostylous
or triformous.

7. Bracts and floaral parts without Bracts and floral parts with stalked Bracts and floral parts
stalked glands glands. without stalked glands.

8. Calyx lobes 5 or 6, longer than Calyx lobes 4-6, longer or shorter Calyx lobes 5, longer than
corolla. than corolla corolla.

9. Corolla lobes 5. Corolla lobes 4-6 Corolla lobes 5-6.

10. Stamens 5. Stamens 4-6. Stamens 5.

11. Ovary 2-loculed; stigmas 2, Ovary 2-6-loculed; stigmas 2, linear. Ovary 2-loculed; stigmas 2,
ovate.
12. Fruit not known. Fruit a berry, fleshy or homy, 2 or Fruit a berry, 2-cocous; cocci
5-6 loculed. horny.
 16 PHYTOTAXONOMY, VOL. 1,2001
Type\ Leptomischus primuloides Drake
Distrib. 4 species in India, Myanmar, China and
Vietnam. Two species in India.
Lo (1993) treated the synonym of the type
species as follows:
1. Leptomischus primuloides Drake in Bull.
Mus. Nat. Natur Paris 1:117 1895 et in Morot, J.
de Bot. 9:208 1895. Pitard in Lecomte, FI. Gen.
Indo-ChineZ'.ll, et 8(1-2). 1992; C.Y. W uet Wang
in Acta Phytotax Sin. 6(3); 294. 1937; H.S, Lo in
Bull. Bot. Res. 6(4); 49, 1986, (specierum clavis
diagnostica)
Indopolysolenia burmanica Deb & Rout in Kew
Bull 45 (2) :339,f. 1. 1990
Myanmar, Myitkyina dist., Laukhaung, 1200m,
13 May, 1938, C. W.D. Kermode 17374A (holo K);
Kachin Hill, S.M. Toppin s.n. (K), Chandwin,
Bhamo, 1911, S.M. Toppin 3244(CAL). ; Hedyotideae, but are absent in Cinchoneae. Seeds
2. Leptomischus wallichii (Hook.f.) H.S. Lo in
Acta Phytotax Sin. 31(3): 275. 1993. Polysolenia
wallichii Hook.f. in Benth & Hook.f Gen PI. 2:68
1873 & FI. Brit Ind. 3:94 1880 (Type: Khasi Hills,
W.Gomez s.n. ex W all Cat. 8424(K), (CAL);
Kanjilal et al FI. Assam 3:49 1939; Balakrishnan,
FI. Jowai 1:237 1981.
Indopolysolenia wallichii (Hook.f.) Bennet in
Ind. For. 107 (7):437 1981; Deb & Rout in Nayar
& Sastry (ed.) Red Data Book 3 : 2 1 4 1 1. 1990
3. Leptomischus modesta (Hook.f.) Deb comb.
nov.
Basionym: Keenania modesta (Hook.f.) F/. Brit.
Ind. 3:101, 1880
Type: Assam, Cachar, Duarband, April 1873,
R.L. Keenan s.n. (K!)
The species is known on the type gathering
only. The type locality was explored by the author
of this paper in October 1995, but it could not be
traced out. The type specimen was collected by
Keenan in 1873 before organisation of tea gardens
in that surrounding which are very common in that
region, that led to the depletion of the environment
co n d itio n th ereon, leaving no scope for its
continuance.
The Genus Polyura
The genus Polyura is very close to Spiradiclis
in habit, leaves, stipules, inflorescence, minute
monomorphic flowers, persistent calyx, corolla
form, insertion of stamens, bilocular ovary with
num erous ovules, m inute embryo and fleshy
endosperm. Darwin (1976) and Ridsdale (1988),
1993) treated the genus Spiradiclis in Ophiorrhizeae
Bremek, ex Verd.
The genera Neohymenopogon and Dunnia were
studied in detail. Matured seeds of Dunnia assamica
were studied for the first time by Deb (1999). 42
character states were compared. 15 of them are
common to Hedyotideae and Cinchoneae. Presence
of rhaphides in the tissues, peltate placenta borne
on a stalk at the base o f the ovary locule and
parenchymatous exotesta are characteristics of
are slightly winged in Neohymenopogen, but club-
shaped excrescences o f exotesta are present in
Dunnia assamica. Thus, the evaluation of character
states justifies their inclusion in the Hedyotideae.
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