CHAPTER I

INTRODUCTION

1.1 FLORISTIC STUDY

The first ever floristic work on plants of Manipur was carried out by

Clarke (1889) in ‘Plants of Kohima and Munneypore ’ and recorded 248

species under 57 families of Dicotyledonous plants which also includes

many Acanthaceae species. Manipur as habitat was cited in Hooker (1885)

‘Flora of British India ’ on the basis of collection by Clarke in (1885) from

Manipur. Watt (1889 -1890) also made collection of economic plants from

Manipur. Kingdom Ward (1948, 1952), described the characteristics of

some species including Strobilanthes asymmetrica from Manipur. Deb

(1961) studied the flora of Manipur from 1951 to 1955 and published

‘Dicotyledonous plants of Manipur territory ’. Singh et al. (2002) published

the flora of Manipur which does not include the family Acanthaceae. Some

floristic works have also been done on the flora of some districts by

Manipur University, (Singh 1987; Singh 1990) in Tengnoupal and

Tamenglong districts of Manipur. Recently, some floristic exploration had

also been done by (Devi 2004; Devi 2006) for the state and (Devi 2008;

Devi 2010) for Thoubal district. These localized studies have resulted in

the publication of many papers, describing taxonomic studies of many

species, ethno-botanical studies and many new reports of taxa for the state

flora. With the Jhuming cultivation in practiced, anthropogenic activities

and over exploitation of natural resources, thereby contributes to the rapid

degradation of forest causing irreversible loss of species diversity in

Manipur. Hence, revision work of this rich diversity is urgent necessity for

taking up concrete steps for protection and conservation.

The state Manipur lies in the northernmost part of the Indian sub-

continent which falls under the Indo-Burma (IBR) which is one of the

biodiversity hot spots of the globe (Myers et al. 2000). IBR is an accurate

sedimentary belt with North - South trend of folded mountains chains,

formed by Cenozoic rock with Triassic metamorphic basement. The sub-

duction between the Indian and Burmese plate bisects the state Manipur

approximately into two equal halves (Sahu et al. 2006). Eastern Himalaya,

which includes Manipur, has a high plant endemism (Behera et al. 2002).

Moreover, Indo-Myanmar and South East Asian region has been a major

source of origin of many Acanthaceae species found around the globe

(Meyer & Lavergne 2004). There has been inflow and outflow of flora

between the two plates naturally which have naturalized in the state flora.

Manipur is situated on the easternmost part of Northeast India, possesses

all the major vegetation types. It is one of the floristically richest states in

India with unique composition of plants including a number of endemic

species due to its physical geographic structure coupled with varied nature

of rainfall, temperature and altitude. Though the state enjoys luxuriant

vegetation including many endemic, endangered and vulnerable species, a

complete account of plants in the state is still lacking. The area of the state

is about 22,372 Km2 of which about 1,813 Km2 form the central valley. The

major portion of the sate consists of ranges of hills with a north and south

general trend. In the centre lies the valley which is about 48.3 km in length

and 32.2 km in breadth. The valley districts comprises of Imphal East,

Imphal West, Thoubal and Bishnupur (Vedaja 1998; Singh 2010). [Map.

1A].

The family Acanthaceae is a large flowering plant family that

comprises about 220 genera and ca. 4000 species distributed in pan -

tropical and sub-tropical, with a few species in temperate regions. Clarke

(1885) reported 49 genera and 501 species of Acanthaceae from the then

British India. Since then a large number of new species have been

discovered and several ornamental and economically important plants

have been introduced to India. In Karthikeyan et al. (2009) ‘Flowering

Plants of India’ reported 47 genera, 475 species and 118 varieties of

Acanthaceae in India. The family is considered as a compl icated one by

most taxonomists (Remadevi & Binojkumar 2009). Classification of

Acanthaceae is largely based on minute characters. The interrelationships

among them are not sufficiently understood possibly due to the diverse

taxonomical characters posses by many species. The identifying characters

are often technical and many species appear to be so much similar that it

becomes difficult to group into separate genera and tribes. The taxonomic

status of this family is unclear and many taxa are not yet finally delimited.

The family is of varied economic importance as medicinal, taboos and

socio-religious aspects. Sinha (1996) reported 16 genera with 20

medicinally important Acanthaceae species from the state . Many others

ethno-botanical aspects were also published (Singh & Singh 1995a,b;

Singh & Singh 1996; Singh et al. 1988). Species of a few genera mostly

tropical are cultivated as ornamentals. In spite of their importance in

biodiversity, medicinal and economic values, very little attention was paid

to them by taxonomist.

No comprehensive study for this family has been taken up in

Manipur. Deb (1961) reported 17 genera with 44 species of Acanthaceae

from Manipur. Singh (1990) reported 12 genera with 24 species of

Acanthaceae from Tamenglong district. Singh (1987) also reported 10

genera with 17 Acanthaceae species from Tengnoupal district. In spite of

this with some others ethno-botanical published papers no significant

work on Acanthaceae has been done so far. In ‘Flora of Manipur’ published

by Botanical Survey of India, the family Acanthaceae was not included or

neglected. With the objective in view, this revision of the family

Acanthaceae is a must after (Deb 1961) a gap of 50 years for flora of the

valley district of Manipur. In ensuring 50 years enormous changes have

taken place in taxonomy and nomenclature of flowering plants. The

present revision study was done by intensive plant exploration in various

areas of the valley districts whose floristic wealth was unknown earlier

and this taxonomic works results in a better understanding of

nomenclatural updates and taxonomic clarification of Acanthaceae species,

new additions to the existing flora, rediscovery of rare species, new re cord

of pliestesial or mass flowering species. Moreover, this research work will

give a complete account of the members under this family, distribution,

morphological accounts, relationship among taxa and ethno-botanical of

the species found in valley districts.

1.2 OBJECTIVES

1. To develop full taxonomic criteria, nomenclatural changes,

contribution of additional data to the existing information and most

updated classification of the family Acanthaceae.

2. To have a better understanding of taxonomic status, field status,

distribution and conservation status of the Acanthaceae species.

3. Cladistic and phenetic analysis of relationships among the genera and

species of Acanthaceae.

4. To explore significant ethno-botanical aspects of Acanthaceae.

1.3 SYSTEMATICS

Systematics is a science that includes and encompasses traditional

taxonomy, the description, identification, nomenclature, and classification

of organisms and that has its primary goal the reconstruction of phylogeny,

or evolutionary history of life. The definition of systematics is not novel,

but neither is it universal (Simpson 2006). The use of systematics depends

on how specialist in the field use the term, an example being the journal

Systematic Botany, which contains articles both in traditional taxonomy

and phylogenic reconstruction. Plant systematic is studied by acquiring,

analyzing, and synthesizing information about plants and plant parts . It

includes discussion about all recent developments in various botanists

work.

Although systematic was recognized as a formal major field of study

during the latter half of twentieth century, the term had been in use for a

considerable period. Derived from the Latin word Systema (organized

whole), forming the title of the famous work of Linnaeus Systema naturae

(1735), the term Systematics first appeared in his Genera plantarum

(1737), though Huxley is often credited to have made the first use of the

term in his article in nature on the systematic of birds. Simpson (1961)

defined systematic as a scientific study of the kinds and diversity of

organisms and of any and all relationship between them. According to

(Simpson 1961; Heywood 1967 and Mayr 1969) have treated taxonomy as

a broad field of biological science which includes four main subjects of

discussion such as i) comparative study of organisms i.e. systematic ii)

taxonomic systems iii) nomenclature and iv) documentation.

Systematic research is the basis for acquiring, cataloguing, and

retrieving information about life’s diversity. Essential to this research is

documentation, through collection and storage of references specimens e.g.

in accredited herbarium. Computerized data entry of this co llection

information is now vital to cataloguing and retrieving the vast amount of

information dealing with biodiversity.

Systematics is also an integrative and unifying science. The study of

systematic provides the scientific basis for defining or delimiting species

and infra-specific taxa (sub species or varieties) and for establishing that ,

these are distinct from others, closely related and similar taxa. Such

studies are especially important today in conservation biology. In addition,

understanding the history of evolution and geography may aid in

conservation and management decisions, where priorities must be set as to

which regions or areas should be preserve.

Finally, perhaps the primary motivation for many in the field of

systematic has been the joy of exploring the intricate complexity and

incredible diversity of life. Systematic also can be challenging intellectual

activity, generally requiring acute and patient skills of observation.

Reconstruction of phylogeny relationship can be especially ch allenging and

rewarding. Today we faced a moral issue: the tragic and irrevocable loss of

species, particularly accelerated by rampant and destruction of habit, such

as deforestation. We can all try to help, both on personal and professional

level, systematic, which has been called simply ‘The study of biodiversity’

is the major tool for documenting that biodiversity and can be a major tool

for helping to save it.

1.4 ACANTHACEAE

The family Acanthaceae is a large flowering plant family that

comprises about 220 genera and ca. 4000 species distributed in pan-

tropical and sub-tropical, with a few species in temperate regions. From

related families, the plant of Acanthaceae are distinguished by a number of

characters, notably, the usually presence of cystoliths in vegetation organs,

the presence and development of floral bracts and bracteoles. Bilabiate

corolla associated bilocular ovary, the generally bivalvate elastically

dehiscing capsules, and usually by the curved retinacula supporting the

seeds. The anthers and stamens provide many characters diagnostic of

groups of genera.

1.5 REVIEW OF LITERATURE

1.5.1 Earlier Works

In Linnaeus (1753) in his ‘Species plantarum’ described 17 species and

placed Acanthaceae under class Didynamia. It was Jussieu (1789) in his ‘Genera

plantarum’ gave the name ‘Acanthi’ for Acanthaceae after separating it from its

related families. The appellation ‘Acanthaceae’ was used for the first time by Robert

Brown (1810). Nees von Essenbeck (1832, 1847) after an in-depth study based on

the presence or absence of rectinacula, recognized two major division of

Acanthaceae such as Anechmatacanthaceae and Echmatacanthaceae. Anderson

(1867) published an enumeration of Indian species of Acanthaceae. After a detailed

study of this family Bentham & Hooker (1876) classified it into 5 tribes, viz.

Thunbergieae, Nelsonioideae, Mendoncioideae, Thunbergioideae and Acanthoideae.

Nelsonioideae was removed from Acanthaceae and transferred it to a new sub

family Rhinanthoideae under the family Scrophulariaceae by Bremekamp (1944,

1955), based on various palynological characters. Cramer (1989) studied the

Hygrophila complex and pointed out that species belonging to Hygrophila from

India and Ceylon, show high degree of polymorphism in the characters of

inflorescence and other floral characters. He also studied the nomenclatural

changes of Acanthaceae with special reference to India and Ceylon (Cramer 1992).

There were certain important contributions to the family Acanthaceae.

Several new genera had been reported viz. Apassalus (Kobuski 1928); Trichaulax

from Africa (Vollensen 1992); Pranceacanthus from Brazil (Wasshausen 1884 a).

Several new species of Acanthaceae were also reported under different

genera Pseudocalyx (Breteler 1994); Strobilanthes (Bremakamp 1953, Carine et al.

2000; Wood et al. 2003) Rungia (Champluvier 2000); Parareullia (Chen et al.

2009); Anisacanthus (Daniel 1982); Justicia (Durkee & McDade 1996; Kelbessa

1995, 1999, 2009); Wasshausen 2000; Wasshausen 2002); Asystasia ( Enserma

1989; Endress et al. 2000); Lepidagathis (Kameyama 2008); Barleria (Malombe &

Darbyshire 2010); Ruellia ( Ramamoorthy 1988); Trichaulax (Vollensen 1992);

Habracanthus (Wasshausen 1884 b) and Odontonema (Wasshausen 1993).

Several significant Taxonomic and Systematics studies of Acanthaceae were

done for different genera were published viz., Hypoestes (Balkwill & Norris 1985);

Dicliptera (Balkwill et al 1988; Darbyshire & Vollensen 2007); Barleria (Balkwill &

Kevin 1997; Balkwill & Balkwill 1998; Darbyshire 2008); Thunbergiodeae (Borg et

al. 2008 ); Justicia (Brummitt 1990; Graham 1988; Hedren 1988; Mc Dade et al.

2000 a,b; 2008; Wood 2009; Wood 2010); Strobilanthes (Bremekamp, 1994; Carine

& Scotland 2002; Moylan et al. 2002, 2004a; Wood & Scotland 2000, 2003a, 2003b,

2006; Bennett & Scotland 2003; Fei 2007); Metarungia (Darbyshire et al. 2008);

Hygrophila (Cramer, 1989; Cuda & Sutton, 2000; Ningombam & Singh 2010;

Carlowrightia (Daniel 1983); Tetramerium ( Daniel 1986); Ruellia (Daniel 1999;

Ezcurra 1993; Long 1964, 1973); Asystasia ( Furness & Vollensen 1991; Kiew &

Vollensen 1997); Nelsonia (Hossain, 1984; Hossain & Emumwen 1981);

Lepidagathis (Kameyama 2008); Odontonema (Wasshausen 1993); Peristrophe (Hao

2007).

Cytogenetic studies of Acanthaceae on epidermal structure and ontogeny

(Grant 1955; Inamdar 1970; Sharma et al. 2010). Moylan et al. (2004 b) studies the

floral anatomy of Strobilanthes. Schonenberger & Endress (1998) studied the

structure and development of flowers in Mendoncia, Pseudocalyx and Thunbergia

and their systematics application. Wilson (1988) studied the mechanism of seed

expulsion in Acanthaceae. Study on chromosomes of Acanthaceae was done (Daniel

et al. 1990; Piovano & Bernardello 1991; Daniel 1993, 2000, 2006).

Various Floristic works on Acanthaceae were carried for the flora of Siamese

(Imlay 1939); Panama (Woodson et al. 1978); Africa (Heine 1962; Troupin 1982);

Colombian Acanthaceae (Wood 1988); Almas, Bahai & Brazil (Wasshausen 1993);

Ceylon (Wood 1995); Panama (Daniel & McDade 1995); South America

(Wasshausen 1995); Srilanka (Cramer 1996); South Eastern United States

(Wasshausen 1998); Central America (Durkee 1999); Mexico (Daniel 2003); Guinea

and Gabon (Champluvier et al. 2003); South East Asia (Wood & Scotland 2000,

2003a, 2003b); Somalia (Hedren 2006) and Pakistan (Perveen & Qaiser 2010).

Corolla aestivation pattern are highly conserved across genera of

Acanthaceae (Anderson 1863, 1867; Balkwill & Norris 1988; Bentham & Hooker

1876; Bremekamp 1965; Burkill & Clarke 1899 – 1900; Clarke 1885; Lindau 1893,

1895; Nees 1832, 1847. Lindau (1895) there are at least five distinct corolla

aestivation types: ascending-cochlear, left-controted, quincuncial, open and

descending-cochlear. Schonenberger (1998) further showed that Thunbergia is

polymorphic for corolla aestivation.

Due to the enormous range of pollen variation within Acanthacae (Radlkofer

1883; Lindau 1893, 1895; Raj 1961, 1973; Bremekamp 1944, 1965; Furness 1985,

1989, Balkwill & Norris 1988; Scotland 1990, 1991, 1992a, 1992b, 1993; Carine &

Scotland 1999) emphasized pollen characters particularly at the tribal and sub-

tribal levels. The extreme diversity of pollen types within Acanthaceae means that

pollen variation has the potential to provide useful evidence for the discovery of

taxa. However, it remains unclear how best to treat pollen variation in any analysis

encompassing the whole family; decision concerning characters and characters

states are problematic Scoltland & Vollensen 2000; Perveen & Qaiser 2010.

1.5.2 Acanthaceae studies in India

Rheede (1678–1703) in his monumental work ‘Hortus Indicus Malabaricus’

(in 12 volumes) described 286 species out of which 12 belong to the family

Acanthaceae. The latter half of the 19th century witnessed the publication of Flora

of British India by Hooker (1885). In volume 4 of this work, Clarke (1884 - 85)

described 49 genera and 501 species belonging to this family from the India region

comprising the present India, Pakistan, Sri lanka, Bangladesh, Nepal, Bhutan,

Myanmar and Malayan peninsula. During the first three decades of the 20th century

intensive exploration were done all over the country that resulted in many

publications (Gamble 1924; Sharma 1996; Shendage & Yadav 2010). Vollensen

(1999) reported a new genus Cynarospermum and reported a new species

Cynarospermum asperrimum from India. Justicia burkilliana is reported by

Bhattacharya & Vollensen (2002) and Lepidagathis keralensis species by

Madhusoodanan et al. (1992). Pradeep & Sivarajan (1991) described a new species

Justicia ekakusuma from Kasargod district. Ellis (1966) described a new species

Andrographis nallamalaya from Andrapradesh.

There were certain noteworthy contributions to the family Acanthaceae in

India. Santapau (1944 - 1952) noticed the mass flowering of Strobilanthes in

various part of India. He noticed a reduction in the gap between the flowerings of

Strobilanthes kunthianus to six or seven years in comparison to earlier recorded 12

years. He also studied the phenology of several species of Strobilanthes and

formation of aromatic glandular hairs on the fruit. Matthew (1971) studies the

mass flowering of Strobilanthes in India, especially Strobilanthes kunthianus (Nees)

T. Anders. Agarwal (1979) studied the morphology of fruits and seeds in Barleria

prionitis and Barleria cristata from the angle of their rate of dispersal. Panigrahi &

Das (1981) revised the genus Haplanthodes for India. Ellis (1988) studied the

Rostellularia diffusa complex.

1.5.3 Review works on Acanthaceae in Manipur and Neighbouring

States

Fischer (1932) in Plants of Assam reported one species of Strobilanthes.

Prain (1903) Acanthaceae in ‘Bengal Plants’ reported 78 species under 32 genera.

Kanjilal et al. (1934 – 1940) in ‘Flora of Assam’ recorded 21 genera with 88 species.

Deb (1983) in ‘The Flora of Tripura State’ recorded 198 species under 28 genera.

George Watt, C.B. Clarke, A. Meebold and S.K. Mukerjee were the earliest

plant explorers of Manipur where they collected many plant species of different

families including Acanthaceae. Clarke (1889) “Plants of Kohima and Munneypore”

was the first floristic publication of Manipur where many species of Acanthaceae

were reported. The taxonomic characteristics of many plants were also described

by Kingdom Ward (1948, 1952) among them Strobilanthes asymmetrica is included.

He collected and deposited in the Kew herbarium as type specimen in October 1948

from Manipur. Based on the collection of Watt (1889-1890) and Clarke (1889),

Hooker (1872-1897) in ‘Flora of British India’ described the habitat of

Phologacanthus, Strobilanthes, Justicia and Lepidagathis species in Manipur.

Acanthaceae is one of the dominant families of dicots by Deb (1961) who reported

44 species under 17 genera. Singh (1990) reported 12 genera with 24 species of

Acanthaceae from Tamenglong district. Singh (1987) also reported 10 genera with

17 Acanthaceae species from Tengnoupal district.

1. 5. 4 MOLECULAR SYSTEMATICS OF ACANTHACEAE

There have been four molecular systematic studies specifically addressing

the higher level systematics of Acanthaceae (Hedren & Olmstead 1995; Scotland et

al. 1995; McDade et al. 2000a). The genus used for these studies were rbcL, trnL-

trnL and trnL-trnF combined with ITS, respectively. The rbcL data of Hedren et al.

(1995) was re-analyzed by Scotland et al. (1995) but provided limited resolution

within Acanthaceae. The strict consensus trees of Scotland et al. 1995; McDade &

Moody et al. 2000a resolved Acanthaceae sensu Lindau (1895) as a group.

Sochonenberger & Endress (1998) showed that Mendonia initially possesses two

ovary locules and four ovules, in common with many members of Acanthaceae, but

latter in development forms a 1 – 2 seeded drupe. This confirmed the view of

Brummitt (1989) that separating Mendoncia from Acanthaceae on the basis of

possession of a drupaceous fruit was mistaken. Scotland et al (1995) and McDade et

al. (2000b) confirmed that genera with retinaculate fruits are a monophyletic

group. McDade et al. (2000), Phylogenetic relationship among Acanthaceae

evidence from two genomes. McDade et al. (2000) studied the phylogenetic

relationships within Justicieae from molecular sequences, morphology and cytology

evidences. Moylan et al. (2004a) studied the phylogenetic relationship among

Strobilanthes from ITS nrdna, trnl-fcpdna, and morphology evidences. Borg et al.

(2008) studied the phylogenetic and morphological evolution of Thunbergiodeae.

McDade et al. (2008) studied the phylogenetic relationship among lineages of

Acanthaceae. Recent published molecular studies (Hedren et al. 1995; Scotland et

al. 1992b; Mc Dade & Moody 1999) highlighted the aspects of disagreement

between Lindau (1895) and Bremekamp (1965) classification.

1. 6 CLASSIFICATION

1.6.1 Classification of Acanthaceae (Lindau, 1895).

Lindau (1895) is the most recent taxonomic treatment of Acanthaceae that

accounts for all genera. He recognized four sub families, three of which comprises

genera that do not possess retinaculate fruits (Nelsonioideae, Thunbergiodeae,

Mendonciodeae and Thunbergioideae represented two very similar sub families of

climbing plants separated on the basis of the capsular fruits of Thunbergiodeae or

the drupaceous fruits of Mendoncioiadeae. Acanthoideae, comprising genera with

retinaculate fruits, was subdivided into two groups based on corolla aestivation

pattern (Imbricatae and Contortae).

Acanthoideae

Contortae

Imbricate corolla

Contortedcorolla

Retinacula

Imbricatae

Nels

on

ioid

ea

eT

hu

nbe

rgio

idea

eM

en

don

coio

ideae

Further sub-division was based primarily on pollen morphology.

Nelsonioideae

Thunbergioideae

Mendoncioideae

Acanthoideae

Contortae

Trichanthereae

Louteridieae

Hygrophileae

Petalideae

Strobilantheae

Ruellieae

Barlerieae

Imbricatae

Acantheae

Aphelandreae

Andrographideae

Asystasieae

Graptophylleae

Pseuderanthemeae

Diclipterineae

Odontonemineae

Monothecineae

Porphyrocomineae

Isoglossineae

Justicieae

1.6.2 Classification of Acanthaceae (Bremekamp, 1965).

Bremekamp (1965) Acanthaceae excluded genera that lack retinaculate

fruits . He placed Nelsoniodeae within Scrophulariaceae, recognized Thunbergiaceae

and Mendoniaceae as distinct families and divided his restricted Acanthaceae into

two groups on the basis of presence and absence of cystoliths, articulated stems,

monothecate anthers and colpate pollen

Acanthoideae Ruellioideae

4 Monothecate anthers

Colpate pollen

Articulated stems

Cystoliths

Retinacula

Further sub-division was based primarily on pollen morphology.

Acanthoideae

Haselhoffieae

Rhombochlamydeae

Stenandriopsideae

Aphelandreae

Acantheae

Ruellioideae

Trichanthereae

Whitfieldieae

Louteridieae

Ruellieae

Blechinae

Ruelliinae

Barleriinae

Strobilanthidinae

Petalidiinae

Hygrophylinae

Lepidagathideae

Lepidagathinae

Chroesthidinae

Borneacanthinae

Andrographideae

Justiciinae

Odonteminae

Rhytiglossininae

Justiciinae

1.6.3 Classification of Acanthaceae by Scotland & Vollensen (2000).

The Classification of Acanthaceae by Scotland & Vollensen (2000) represents

a synthesis of morphological and molecular data from 221 accepted genera of

Acanthaceae. Taxa were recognized based on combination of morphological

diagnostic feature, the three - item analysis of morphological data and published

molecular sequence analysis .Twenty of those genera remain unplaced within

Acanthoideae, and the status of some of these genera e.g. Acanthura is Lindau is

doubtful. All of these genera contain few species, and for several species described

by Benoist. Within the subtribe Justiciinae there are 103 genera, compared with 118

in the rest of the family which remains to be determined. The sub tribe Ruellinae,

although smaller than Justiciinae in terms of the number of genera, contains both

Strobilanthes and Ruellia with c 450 and c. 250 species respectively. The genus

Thomandersia is excluded as preliminary research indicates that this genus has

been misplaced within Acanthaceae.

Acanthaceae Juss.

Nelsonioideae Pfeiff.

Thunbergiodeae Kostel.

Acanthoideae Link

Acantheae Dumort.

Ruellieae Dumort.

Ruelliinae Nees

Justiciinae Nees

Andrographiinae Nees

Barleriinae Nees

Nelsoniodeae Thunbergia Retz.

Anisosepalim E.Hossain Acanthoideae

Elytraria Michx. Acantheae

Gynocraterium Bremek. Acanthopsis Harv.

Nelsonia R. Br. Acanthus L.

Ophiorrhiziphyllon Kurz Achyrocalyx Benoist

Saintpauliopsis Staner Aphelandra R.Br.

Staurogyne Wall. Blepharis Juss.

Thunbergioideae Crossandra Salisb.

Anomacanthus R.D. Good Crossandrella C.B. Clarke

Mendoncia Vell. ex Vand. Cynarospermum Vollensen

Meyenia Nees Cyphacanthus Leonard

Pseudocalyx Radlk. Encephalosphaera Lindau

Thunbergia Retz. Acanthoideae

Nelsoniodeae Acantheae

Anisosepalim E.Hossain Acanthopsis Harv.

Elytraria Michx. Acanthus L.

Gynocraterium Bremek. Achyrocalyx Benoist

Nelsonia R. Br. Aphelandra R.Br.

Ophiorrhiziphyllon Kurz Blepharis Juss.

Saintpauliopsis Staner Crossandra Salisb.

Staurogyne Wall. Crossandrella C.B. Clarke

Thunbergioideae Cynarospermum Vollensen

Anomacanthus R.D. Good Cyphacanthus Leonard

Mendoncia Vell. ex Vand. Encephalosphaera Lindau

Meyenia Nees Geissomeria Lindl.

Pseudocalyx Radlk. Holographis Nees

Neriacanthus Benth. Brillantaisia P. Beauv.

Orophochilus Lindau Brunoniella Bremek.

Rhombochlamys Lindau Calacanthus T. Anderson ex Benth.

Salpixantha Hook. Clarkeasia J.R.I. Wood

Sclerochiton Harv. Dischistocalyx T. Anderson ex Benth.

Stenandrium Nees Duosperma Dayton

Streptosiphon Mildbr. Dyschoriste Nees

Strobilacanthus Griseb. Echinacanthus Nees

Xantheranthemum Lindau Epiclastopelma Lindau

Ruellieae Eranthemum L.

Neriacanthus Benth. Eremomastax Lindau

Orophochilus Lindau Eusiphon Benoist

Rhombochlamys Lindau Hemigraphis Nees

Salpixantha Hook. Heteradelphia Lindau

Sclerochiton Harv. Hygrophila R.Br.

Stenandrium Nees Ionacanthus Benoist

Streptosiphon Mildbr. Kosmosiphon Lindau

Strobilacanthus Griseb. Leptosiphonium F. Muell.

Xantheranthemum Lindau Louteridium S. Moore

Ruellieae Mimulopsis Schweinf.

Ruelliinae Pararuellia Bremek.

Acanthopale C.B. Clarke Petalidium Nees

Aechmanthera Nees Phaulopsis willd.

Apassalus Kobuski Physacanthus Benth.

Benoiacanthus Heine & A. Raynal Polylychnis Bremek.

Blechum P.Brown Pseudoruellia Benoist

Bravaisia DC. Ruellia L.

Ruelliospis C.B. Clarke Anisacanthus Nees

Sancheria Riuz & Pav. Anisotes Nees

Satanocrater Schweinf. Anthacanthus Nees

Sautiera Decne. Aphanosperma T.F. Daniel

Spirostigma Nees Ascotheca Heine

Stenosiphonium Nees Asystasia Blume

Stenothyrsus C.B. Clarke Ballochia Balf.f.

Strobilanthes Blume Brachystephanus Nees

Strobilanthopsis S.Moore Calycacanthus K. Schum.

Suessenguthinia Merm. Carlowrightiana A. Gray

Trichanthera Kunth Calerina Benoist

Trichosanchezia Mildbr. Centrilla Lindau

Zygoruellia Baill. Cephalacanthus Lindau

Andrographinae Chamaeranthemum Lindau

Andrographis Wall. ex Nees Chileranthemum Oerst.

Cystacanthus T. Anderson Chlamydocardia Lindau

Diotacanthus Benth. Chlamydostachya Mildbr.

Graphandra J.B. Imlay Chorisochoron Vollesen

Gymnostachyum Nees Clinacanthus Nees

Haplanthodes Kuntze Clistax Mart.

Indoneesiella Sreem. Codonacanthus Nees

Phlogacanthus Nees Conocalyx Benoist

Justiciinae Cosmianthemum Bremek.

Afrofittonia Lindau Cyclacanthus S. Moore

Ambongia Benoist Cylindrosolenium Lindau

Angkalanthus Balf. Danguya Beniost

Dasytropsis Urb. Megalochlamys Lindau

Dichazothece Lindau Megaskepasma Lindau

Dicladanthera F. muell. Mellitacanthus S.Moore

Dicliptera Juss. Metarungia Baden

Ecbolium Kurz Mexacanthus T.F. danniel

Filetia Miq. Mirandea Rzed.

Fittonia Coem. Monechma Hochst.

Forcipella Baill. Monothecium Hochst.

Glossochilus Nees Odontonema Nees

Graptophyllum Nees Oplonia Raf.

Gypsacanthus Lott, Jaramillo & Rzed. Oreacanthus Benth.

Harpochilus Nees Pachystachys Nees

Henrya Nees Pelecostemon Leonard

Herpetacanthus Nees Peristrophe Nees

Hoverdenia Nees Phialacanthus Benth.

Hypoestes Sol. Ex R.Br. Populina Baill.

Ichtyostoma Hedren & Vollensen Pranceacanthus Wassh.

Isoglossa Oerst. Pseuderanthemum Radlk.

Isotheca Turrill Pseudodicliptera Benoist

Jadunia Lindau Psilanthele Lindau

Jurusia Lindau Ptyssiglottis T. Anderson

Justicia L. Pulchranthus V. M. Baum, Reveal & Nowicke

Kalbreyeriella Lindau Razisea Oerst.

Linariantha B. L. Burtt & R.M Sm. Rhinacanthus Nees

Mackaya Harv. Ritonia Benoist

Marcania J.B. Imlay Rungia Nees

Ruspolia Lindau Unplaced within Acanthoideae

Ruttya Harv. Acanthostelma Bidgood & Brummitt

Samuelssonia Urb. & Ekman Acanthura Lindau

Sapphoa Urb. Aphelandrella Mildbr.

Schaueria Nees Camarotea Scott Elliot

Sebastiano-Schaueria Nees Chlamydacanthus Lindau

Spathacanthus Baill. Dolichostachs Benoist

Sphinctacanthus benth. Golaea Chiov.

Stenostephanus Nees Idianacanthus Leonard

Streblacanthus Kuntze Kudoacanthus Hosok.

Tessmanniacanthus Midbr. Lankesteria Lindl.

Tetramerium Nees Lasiocladus Bojer ex Nees

Thysanostigma J.B. Imlay Leandriella Benoist

Trichaulax Vollensen Morsacanthus Rizzini

Trichocalyx Balf. F. Neuracanthus Nees

Xerothamnella C.T. White Perenideboles Ram. Goyena

Yeatsia Small Pericalypta Benoist

Barleriinae Sphacanthus Benoist

Barleria L. Vavara Benoist

Barleriola Oerst. Vindasia Benoist

Borneacanthus Bremek. Whitfieldia Hook.

Boutonia DC. Excluded genera

Chroesthes Benoist Thomandersia Baill.

Crabbea Harv.

Hulemacanthus S. Moore

Lepidagathis Willd.

Lophostachys Pohl

1. 7 Distribution and range of Acanthaceae

The four main centers of diversity of Acanthaceae species are Indo-Malaya,

Africa, Brazil, and Central America. Members of the acanthaceae are distributed

from the tropics to temperate regions. The genus Acanthus consists of about 20–25

species: tropical and subtropical regions of Old World; 6 species in India.

Andrographis, about 20 species: tropical and subtropical Asia; 26 species in India.

Asystasia, about 40 species: tropical and subtropical regions of Old World; 9 species

in India. Barleria, about 80–120 species: tropical regions of Africa and Asia, one

species in tropical America; 29 species in India. Dicliptera, about 100 species:

tropical and temperate regions worldwide; 25 species in India. Eranthemum, about

15 species: tropical and subtropical Asia; 17 species in India. Hygrophila, about 100

species: widely distributed in tropical and subtropical regions; 18 species in India.

Hypoestes, about 150 species: Old World; 5 species in India. Justicia, about 700

species: tropical and temperate regions worldwide; 50 species in India.

Lepidagathis, about 100 species: tropical to subtropical regions worldwide (mostly

paleotropical); 23 species in India. Nelsonia by 5 species or fewer: widely occurring

in tropical Africa, Asia, Australia, and South America; 1 species in India. Phaulopsis,

22 species: tropical Africa, SE, and S Asia; 1 species in India. Pseuderanthemum,

about 50 species: pantropical; 8 species in India. Phlogacanthus, about 15 species:

mainland Asia; 13 in India. Peristrophe, about 40 species: tropical and subtropical

Africa, Asia, and Madagascar; 1 species in India. Ruellia, about 250 species: tropical

to temperate regions worldwide; 10 species in India. Rungia, about 50 species: Old

World tropics and subtropics; 15 species in India. Strobilanthes, about 400 species:

tropical Asia; 130 species in India. Thunbergia, more than 100 species: Old World

tropical regions; 13 species in India.

1.8 Numerical Taxonomy: Cladistic and Phenetic

The application of numerical methods to taxonomy, dating back to the rise of

biometrics in the late nineteenth century, has received a great deal of attention with

the development of the computer and computer technology. Thus, it is the analysis

of various types of taxonomic data by mathematical or computerized methods and

numerical evolution of the similarities or affinities between taxonomic units, which

are then arranged into taxa on the basis of their affinities. Adanson (1763), a

French botanist, who for first time put forward a plan for assigning numerical

values to the similarity between organisms and proposed that equal weightage

should be given to all the characters while classifying plants. He used as many

characters as possible for classification, and such classification came to known as

Adansonian classification. According to Heywood (1967) the numerical taxonomy

may be defined as the numerical evaluation of the similarity between groups of

organisms and the ordering of these groups into higher ranking taxa on the basis of

theses similarities.

Numerical taxonomy is a developing branch of taxonomy, which received a

great impetus with the development and advancement of computers. This field of

study is known as mathematical taxonomy (Jardine & Sibson 1971), taximetrics

(Mayr 1966), taximetrics (Rogers 1963), multivariate morphometrics (Blackith &

Reyment 1971). The modern methods of numerical taxonomy had its beginnings

from the contribution of Sneath (1957). Michener & Sokal (1957), and Sokal &

Michener (1958) which culminated in the publication of principles of numerical

taxonomy (Sokal & Sneath 1963) with an expanded and update version Numericial

taxonomy ( Sneath & Sokal 1973).

Numerical taxonomy was however largely developed and popularized by

Sneath & Sokal (1973). The application of Andansonian principles and use of

modern methods and electronic data processing techniques, and have helped in the

evolution of several new classification of plants during the past few decades.

Numerical taxonomy or taximetrics, nowadays frequently and perhaps more

appropriately referred to as phenetic, refers to the application of various

mathematical procedures to numerically encoded character state data for

organisms under study.

Numerical taxonomy does not produce new data or a new classification, but

is rather a new method of organizing data that could help in better understanding

are based either on one or few characters or on one set of data (Singh 2004).

Numerical taxonomy seeks to base classification on a greater number of characters

from many sets of data in an effort to produce an entirely phenetic classification of

maximum predictivity.

Two major bases have been recommended for designing a natural system of

classification. These include:

Phenetics – it involves the similarity between individuals under

consideration, based on a set of phenotypic characters. Thus, phenetics

classification is based on overall affinity as judged by using as many characters as

available. It does not embody any relationship through ancestry. The typical tree

diagram or dendograms, with taxa and groups taxa, being connected at different

levels according to their overall similarity, will not reflect evolutionary

relationships accurately and such dendograms are referred to as phenerograms.

Cladistics – it is based on relationship between individuals with regard to

their evolutionary history. The relationship meant common ancestry. The goal is to

produce objective and repeatable diagrams (evolutionary trees) depicting

hypothetical evolutionary histories. The evolutionary trees with anatomizing

branching leading to various taxa are called cladograms.

Scotland et al (2003) had pointed out the importance of plant morphology in

the phylogeny construction of plants. Farres (1972) used the distance matrices in

estimating phylogenetic trees. Pandit & Gupta (2011) pointed out the role of

computer for comparative study on distance measuring approaches for clustering.

Many papers have been published regarding cladistics as viz. Ashlock (1974)

reported the uses of cladistic for classification of plants; Bawn (1976) states the role

weighting of the character states for cladistic analysis; Crisci & Stuessy (1980)

reported on determining primitive character states for phylogenetic reconstruction;

Dallwitz (1980) describe a general system for coding taxonomic descriptions in the

computer for analysis of the character states; Robinson (1986) points out a key to

overcome common errors in of cladistics; Wilson (1988) pointed out importance of

the application of cladistic taxonomy to plants; Chappill (1992) did cladistic

analysis of Chrysobalanaceae using morphological characters; Scotland (2000)

studied the taxic homology and three-taxon statement analysis based on molecular

and morphological data; Leht (2005) studied the cladistic and phenetic relationship

in Vicia Subgenus Cracca (Fabaceae) based on 89 morphological data.

1.9 Ethnobotanical works of Acanthaceae in Manipur and neighbouring

states.

Globally, many significant ethno-botanical studies were reported, Asano et

al. (1996) studied antiviral activities of Justicia procumbens. Myers & Lavergne

(2004) reported the ornamental species of Acanths and its invasive potential.

Thomas & Yoichiro (2010) reported the in-vitro propagation of rare medicinal plant

Justicia gendarussa.

In India also several works on ethno-medicinal works were done viz., Jamir

et al. (1999) on folklore medicinal plants of Nagaland. Khare (2004) had published

‘Encyclopedia of Indian medicinal plants’ which includes many Acanthaceae species.

Sabu et al. (2001) reported hepatoprotective potential of Andrographis paniculata.

Mukherjee et al. (2009) reported the analgesic activity of Phlogacanthus

thyrsiflorus. Dhankhar et al. (2011) reported Justicia adhatoda as a potential source

of natural medicine. Jain et al. (2011) did a case study on the dietary uses and

conservation concern of edible wetlands plants at Indo- Burma hotspot.

1. 9.1 Manipur

Plant forms an integral part of many rites, rituals, folk tales, taboos,

medicinal, customs and traditions. The meitei community has a rich traditional

knowledge for utilization as home prepared remedies, taboos and folktales. Many,

ethno-botanical work has been done viz., Sinha (1987) studies the ethno-botanical

plants of Manipur; Elangbam (2002) reported the various aspects of wild edible

plants of Manipur; Khan (2005) reported ethno-medicinal plants of Thoubal district

which includes several Acanthaceae species. Several noteworthy publications were

also made on ethno-botanical studies in Manipur viz., Singh & Singh (1995a; 1995b)

on folk-medicinal plants. Singh & Singh (1996); Singh et al. (1996) on indigenous

bio-floklores. Khumbongmayum (2005) on ethno-botanical plants in the scared

grooves. Chakraboty (2003) on wild edible plants. Singh et al. (1997) on ethno-

medico-biological plant. Bahadur (1997); Sharma et al. (2005); Singh et al. (2009)

reported Strobilanthes cusia as a dye yielding plant for imparting black and indigo

colour. Singh & Singh (2003a) on uses of folk-medicinal plants for mother and child

care of meitei community. Singh et al. (2003b) recorded Phlogacanthus thyrsiflorus

as important herbal medicine in Manipur. Devi et al (2009) reported Phlogacanthus

thyrsiflorus and Phlogacanthus tubiflorus as edible flowers found in the valley

districts of Manipur. Devi (2011) also reported Hygrophila phlomoides, Justicia

adhatoda and Phlogacanthus tubiflorus for treatment of diabetes by the meitei

community. Pfoze et al. (2011) reported many wild edible plants from Senapati

district of Manipur.

Though, globally the family Acanthaceae is not considered as economically

important family but with respect to this state this is a very ethno-botanical

important. Remarkable attention has been given to various members of

Acanthaceae due to incredible economic and utilitarian importance. Also a great

many more are recognized and treasured for their novelty or ornamental beauty.