chromosome studies in lamiaceae -...

CHROMOSOME STUDIES IN LAMIACEAE

The dicotyledonous family Lamiaceae is a very large natural group of aromatic

and medicinal plants (Morton, 1962). Lindley (1830) was the first to propose

the name, *Lamiaceae' and later on in 1935 the International Code of Botanical

Nomenclature approved that it can be used as an alternative to Labiatae. The

name .Labiatael, coined by de Jussieu (1789) originates from the Latin noun

labium (lip), referring to the bilipped corolla of their zygomorphic flowers,

a salient feature of the family.

Lamiaceae enfolds 220 genera (Hedge, 1992) and almost 5000 species (Mabberley,

1987), with a very wide range of morphological diversity. The historical

background of Lamiaceae is of much importance with regard to it's phylogeny.

The ancestors of the family might have been originated in the tertiary

tropical forests (Chengyih and Hsiwen, 1982). The evolution and specialization

of the family began in the ancient Mediterranean region. The phylogeny of the

family is detailed just as the centres of origin. Zeven and Zhukovsky (1975)

proposes nine different gene centres, which formed cradles of Lamiaceae

cultivation as, the Chinese-Japanese centre, the Indo-Chinese-Indonesian

centre, the Australian centre, the Hindustani centre, the Central Asian

centre, the Near Eastern centre, the Mediterranean centre, the European

Siberian centre and the African centre.

Majority of these plants are annual and perennial herbs, cosmopolitan, growing

under a great variety of soil and climate and constitute a scrub type of

vegetation (Simpson and Conner - Ogorzaly, 1986). Though the members have a

global distribution (Epling, 1940b; Harley and Reynolds, 1992), the prime

centre of concentration lies in the Mediterranean basin and Asia Minor, both

in regards the number of genera and species (Rivera Nunez and Obon de Castro,

1992). The other main centres of diversity include Europe, Middle East,

Tropical and Central Africa, South Africa, Asia, Australasia, North America,

Arctic and Antartic regions (Morley, 1970).

Members of Ocimeae flourishes in tropics and subtropics, represented in both

hemispheres (Rendle, 1925) by 32 genera and 970 species (Morton, 1962).

Mentheae includes the largest group of plants among Lamiaceae, which are

cosmopolitan in distribution and posses 42 genera and about 540 species (EI-

Gazzar and Watson, 1967). The various taxa of Salveae are distributed in the

temperate and warmer regions of both hemispheres with 10 genera and more than

738 species (Bailey, 1960). The aromatic herbs of Nepeteae are concentrated in

subtropical and temperate Asia (Gill, 1979) and are represented by 9 genera

and 393 species (Willis, 1973). Lanieae consists of a large group of plants,

cosmopolitan in distribution with about 37 genera and about 850 species

(Craner, 1981). Members of Prasieae have a limited distribution, chiefly in

tropical Asia and the Ha~iai Islands (Rendle, 1925), represented by 5 genera.

(Bentham 1876). Prostanthereae includes a small group of plants confined to

Australia and Tasmania (Lawrence, 1951), represented by a few genera and 200

species (Conn, 1992). The various plants of Ajugeae are cosmopolitan in

distribution (Lawrence, 1951) with about 8 genera (Cantino, et al. 1992). In

India, the family is represented by about 69 genera and 425 species of which

261 are endemic. The temperate Himalayas, the Deccan, North Western India and

South India are the chief centres of concentration. South India is one of the

major centres of distribution since it support over 139 species (Mukherjee,

1940).

The Lamiaceae members are characterized by their peculiar morphology and

aromaticity that they have been well recognized for centuries and is one of

the oldest families known (Swingle, 1946).

The family Lamiaceae is of great importance from a commercial standpoint

(Heywood, 1978). Besides numerous aroma chemicals, perfume products,

medicinals, pot herbs and honey pasturage, the family yield a number of

choice ornamentals (Stanford, 1937). Many members of Lamiaceae find use in

cosmetic industry (Fleisher and Fleisher, 1991 b; Knaut and Richtler, 1990),

for the preparation of detergents, fabric conditioning mixtures (Behan and

Clements, 1990; Ito and Kikuchi, 1991; Nishida, 1991), deodorants and bath

preparations [Kawasaki and Tsunakawa, 1990; Nishida, 1990; Ueyama, et al.,

1990; Morgan and Blagdon, 1991) long lasting fragrant preparations and

perfumes (Tsukasa, 1990; Yoshihara, et al., 1990; Ikimoto and Kuwaori, 1991;

Okada, et al., 1991), herbal preparations (Muller, 1990, 199l),non-irritating

cleansers (Morganti, 1990), hairgels (Lochhead, 1990) and in baking industry

(Bassiouny, et al., 1990; Economou, et al., 1991). Some species find use in

the preparation of dentifrices, mouthwashes, toothpowders, chewing gum and

lozenges, which act as anti-plaque and anti-tartar agents of teeth (Anonymous,

1991; Reuss, 1991; Mori and Yokosuka, 1991). Antifungal and antibacterial

activities have been reported on many species. (Gangrade, et al., 1989a,

1989b) This property is due to their volatility and ephemeral nature (French,

1985) and is of much importance since herbal fungicides are non-toxic (Fawcett

and Spencer, 1970) and easily bio-degradable (Mahadevan, 1982).

The position affinity and supra-familial relationship of the family Lamiaceae

in different systems of classification has been a subject of considerable

debate from time immemorial. Linnaeus (1737) placed the family under 2

classes. Didynamia and Tetra dynamia. However de Jussieu (1789). Gray (1858)

and Eichler (1875-1878) grouped them together under Sympetalae. Bentham and

Hooker (1862-1883) classified Lamiaceae in the order Lamiales, followed by

Bessey (1915). Hutchinson (19261, Cronquist (19681, Takhtajan (1969) and

Dahlgren (1975). On the basis of morphological parameters, Engler and Prantle

(1888-1897) placed Lamiaceae in the order Tubiflorae. Later on Hallier (1905).

Wettstein (19241, Rendle (1925) and Pulle (1938) followed the same pursuit.

June11 (1934) and Erdtman (1945) established their systems of classification

on the basis of evidence from gynoecial and pollen characters. In a more

recent system of classification proposed by Goebel (1984) Lamiaceae is placed

in the order Labiatiflorae. Recent research suggests that the family Lamiaceae

is polyphyletic (Abu-Asab and Cantino, 1989, 1992; Cantino, 1992 a, b).

Bentham (1832-1836, 1848, 1876), and Briquet (1895-1897) have made major

contributions towards the infra-familial classification of the family. Later

on modified systems have been proposed by Hillson (1959), Melchior (1964).

Wunderlich (1967), El-Gazzar and Watson (1970a), Dahlgren (1980). Taktajan

(1980), Thorne (1983), Cantino and Sanders (1986) as well as Goldberg (1986).

Bentham (1876) Sub divided Lamiaceae into eight tribes, which are Ocimoideae

(*Ocimeae), Satureineae (*Mentheae), Monardeae (*Salvieae), Nepeteae, Lamieae

(*Stachydeae) Prasieae, Prostanthereae and Ajugoideae (*Ajugeae).

The familial relationship of Lamiales have been evaluated by Cantino (19821,

Cantino et al. (1992) Cronquist (1981) and Thorne (1976) and supra-ordinal

relationships by Lu (1990). The family Lamiaceae is readily distinguished from

all the other families except Verbenaceae by the distinctive gynoecium

(Lawrence, 1951). It is considered to be closely resembling Verbenaceae in

having similar vegetative as well as floral characteristics. The presence of

* Correct names proposed by International Code of Botanical Nomenclature

(Stafleu, et al., 1978; Greuter, 1981).

these intergrading characters make it difficult, if not impossible to separate

all members of one family from the other by any single charac.ter or a

combination of characters (Rendle, 1925). Thus the family Verbenaceae is

phylogenetically related to Lamiaceae (Bessey, 1897). However, Hutchinson

(1924) considered that the two are unrelated, but the two families seem rather

similar because both of them represent climaxes of evolution in their

respective groups. The complicated floral structure among the gamopetalous

families, corroborating with various adaptations for entomophily show that

Lamiaceae can be considered undoubtedly as the culmination of the sympetalous

herbaceous line of Herbaceae (Hutchinson, 1926).

In addition to this Lamiaceae resembles Scrophulariaceae and Acanthaceae by

showing more or less parallel floral structure (Hutchinson, 1924), as well as

the occasional quadrangular nature of stem and opposite phyllotaxy (Rendle,

1925). However according to Evans (1936) Lamiaceae differs from both by the

presence of verticillaster inflorescence, gyno-basic style and the 4-lobed

nature of the ovary as well as fruit.

Lamiaceae also bears relationship with the Boraginaceae, but is set apart by

the technical character of the ovule. When the micropyle is inferior and raphe

inward in Lamiaceae, it is just the reverse in Boraginaceae (Lawrence, 1951).

More over Lamiaceae possess fewer number of stamens than the corolla lobes

(Evans, 1936). Thus Lamiaceae is entitled to a rather high rank among the

Metachlamydeae by reason of their irregularity, tetracyclic condition, reduced

number of stamens as well as carpels and highly specialized type of fruit

(Haupt, 1946).

The perennial growth habit, simple leaves, heterogenous medullary rays,

bisexual hypogynous flowers, anatropous ovules and entomophilous pollination

are some of those primitive features shown by the family (Coulter and

Chamberlain, 1903, Eames, 1961; Metcalfe and Chalk, 1950; Takhtajan, 1969).

The herbaceous nature, exstipulate, opposite or whorled leaves, small

zygomorphic flowers, bilabiate calyx and corolla showing cohesion, reduced

number of stamens, exhibiting epipetaly, nearly monothecous condition of

anthers, bicarpellary syncarpous pistil showing axile placentation, reduction

of the number of ovules to one in each loculus, simple fruits, non-endospermic

seeds and presence of small sized vessels and fibres with simple pits are some

of the advanced characters exhibited by the family (Bessey, 1893; Cronquist,

1968; Erdtman, 1952; Hallier, 1905; Metcalfe and Chalk, 1950; Nageli, 1884;

Takhtajan, 1959; Bailey and Tupper, 1918).

Some of the common genera found in India include Ocimum L.; Acrocephalus

Benth.; Orthosiphon Benth.; Plectranthus L'Herit.; Coleus Lour.; Anisochilus

WalL, Hyptis Jacq.; Pogostenon Desf.; Eusteralis Rafin.; Mentha L.; Salvia L.;

Anisomeles R.Br.; Leucas R.Br.; Leonotis R.Br. and Teucrium L.

Ocimum L., the common *Basil' derives its name from Greek, where 'ozo' means

smell (Hereman, 1868). It possess 160 species with the maximum members

originating from the tropical forests of Africa (Sobti and Pushpangadan,

1982). Other main centres of diversity include South America and Asia

(Krishnamoorthy, (1989). The medicinal and pharmacological properties of this

genus are very well described, in the treatises *Pharmacographia Indica'

(Dymock, et al; 1890) and *Indian Materia Medica' (Nadkarani, 1976 a, 1976 b).

It is well known in perfumery, flavour and cosmetic industries (Kirtikar and

Basu, 1935; Guenther, 1952; Dastur 1962).

The plant of little beauty, Acrocephalus Benth, is named after .akros1 -

summit and *kephalel - head; on account of the flowers being on top of the

branches (Hereman, 1868). This tropical genus, with about 100 species

(Santapau and Henry, 1973) is distributed in the warmer parts of Asia and

Africa (Hooker, 1885). Some species are used in folk medicines (Chopra, et

al., 1956).

The generic name Or-thosiphon Benth., originates from 'orthos' - straight and

-siphon1 - tube; refers to the straight corolla tube (Bailey, 1960). It is

represented by more than 100 species (Santapau and Henry, 1973), mainly annual

or perennial herbs and are distributed in the tropics from Africa to Australia

(Bailey, 1960). Some species are employed in Ayurvedic and Unani medicines

(Uphof, 1959).

Plectranthus L' Herit., an Old World genus (Rivera Nunez & Obon de Castro,

1992), is named from *plektronl-Cock's spur and <anthos1-flower; alluding to

the swollen corolla base and hence the common name, .Cock's spur' (Hereman,

1868). A genus with about 300 species (Richardson, 1992) of fleshy herbs and

under shrubs (Bower, 1956) are distributed chiefly in the tropical and

subtropical regions of Africa, and Asia, spreading eastwards to Australia,

Malaysia, Japan and the Pacific region (Bailey, 1960).

The name Coleus Lour., araises from *coleos'-sheath; referring to the

monadelphous stamens (Plowden, 1968). This Old World genus, commonly known as

the -Flame-nettle' (Fyson, 1986) possess 150 species (Morley, 1970), that are

mainly distributed in the Eastern Hemisphere, being especially abundant in

Africa, India, the Malayan Archipelagoes, extending to Australia and the

Pacific islands (~aile~, 1960). Coleus Lour. yields a variety of pot and

garden ornamentals, and plants with edible tubers (Swingle, 1946). Some

members are found to be of very high medicinal value (CSIR, 1986).

Anisochilus Wall., derived its name from *anises' unequal and *cheilos' - lip;

with regard to the zygomorphic corolla of the flowers (Hereman, 1868). It is a

cosmopolitan genus and holds about 20 species Cramer, 19811, distributed in

India and Tropical Africa (Fyson, 1986). Medicinal and pharmacological

properties are enumerated in -Pharmacographia Indica' (Dymock et al., 1890)

Hyptis Jacq. is named after .hyptios' - resupinate; since the limb of the

corolla is turned on its back (Hereman 1868). It is a large genus with as many

as 400 species and occur in different parts of the world (Richardson, 1992),

the chief centre of concentration being America (Hooker, 1885). The medicinal

properties are revealed by Chopra, et al. (1956).

The name Pogostemon Desf., is originated from -pogon'-beard and .stemon' -

stamen; owing to the tuft of hairs present in the stamens (Bailey, 1960). The

genus comprises a group of aromatic herbs and shrubs (Bower, 1956) with about

40 species (Cramer, 1981) and is mainly distributed in India and Eastern Asia

(Fyson, 1986). Many species are described in *Pharmacographia Indica' for

their medicinal qualities (Dymock, et al., 1890).

Eusteralis Rafin. is a very close ally of Pogostemon Desf. It is a woddy

aromatic herb very common on the rocky slopes, growing in the crevices. 21

species have been reported so far in this genus (Cramer, 1981).

The name Mentha L., bestowed upon by the Greek philosopher Theopharastus

(Jeans, 1973) after *Menthael, Latin name of a nymph; due to the aquatic

habitat it prefers (coats, 1968). It includes several perennial herbaceous

plants (Ranade, 1982), thriving very well in marshy places and propagate by

means of runners or stolons (Stanford, 1937). It is commonly called as mint

(Gray, 1858). Index Kewensis (1893 - 1947) lists about 900 binomials widely

distributed in the temperate and subtropical regions of the World. However,

Harley and Brighton (1977) estimated that there are probably some 25 species.

The medicinal and aromatic properties of many species are described in

'Materia Indica' (Ainslie, 1826) and 'Pharmacographia Indica' (Dymock, 1890).

The name -a L., is derived from the Latin word 'salvo' which means-I save;

referring to the medicinal qualities (Plowden, 1968). The 'common sage' (Gray,

1858) is the most primitive genus of the family although L. is

considered by many authors (Fujita, 1970). This genus is the largest in the

family Lamiaceae (Morley, 1970), with about 1000 species (Estilai, et al,

1990) and widely distributed in the temperate and warmer regions. 'Materia

Indica' (Ainslie 1826') gives a detailed account of its medicinal uses.

The genus' Anisomeles R.Br. is named from 'anisos' unequal and 'melos' -

member; in collaboration with the corolla character (Hereman, 1868). This

small genus with 7 species (Cramer, 1981) is distributed in tropical and

subtropical Asia and Australia (Hooker, 1885). Medicinal properties are

described by Kirtikar and Basu (19351, Uphof (1959), Dastur, (1962) and CSIR

(1986).

Leucas R.Br. is named after 'leukos' - white; in accordance with the downy

white flowers (Hereman, 1868). The genus commonly known as 'Dead-nettle'

(Fyson, 1986) includes wooly, villous pubescent herbs or shrubs with about 60

species (Cramer, 1981) and is concentrated in Asia and Africa (Fyson, 1986).

Medicinal and pharmacological notes are given by Chopra, et al. (1956).

The name Leonotis R.Br. is derived from 'leon' - lion and 'ous' - ear'

referring to the resemblance of the flower to lion's ear. ( Plowden, 1968:. It

consists of annual or perennial herbs or shrubs with about 20 species found

mainly in tropical and South Africa (~ailey, 1960). Medicinal aspects are

mentioned in 'Pharmacographia Indica' (Dymock, et al., 1890).

Teucrium L., is named after -Teucer' of Greek mythology, who discovered it's

medicinal uses (Plowden, 1968). The genus is commonly known as *Germander1

(Gray, 1858). Many of its species are natives of the Mediterranean (Morley,

1970) and chiefly distributed in Temperate and Southern Europe (Fyson, 1986).

The genus consists of more than 100 species (Gleason, 1963).

During the past few decades, the cytological data of many families have been

widely used in comparative and evolutionary studies of various taxa. In

plants, the cytological data of taxonomic importance is mainly pertaining to

the chromosome number and structure during mitosis. Chromosome number will

serve as useful taxonomic markers to distinguish between different taxa.

Additionally it can also solve the problems of dispute between different taxa.

Therefore, these factors are used as classificatory criteria in the same

manner as the morphological characters since the chromosomes have a direct

relation to the genetic system of which they are an integral part (Den Hartog,

et al. 1979).

Sufficient information is available on the Indian members of Lamiaceae by

virtue of the major cytological surveys conducted by Gill (1970, 1971a, 1971b,

1971~) Mehra and Gill (1972), Bhattacharya (1976, 1978a, 1978b) Vij and

Kashyap (1976b) Singh and Sharma (1981a, b, 1982) Vembu and Sampathkumar

(1981), Bir and Saggoo (1984, 1985), Saggoo and Bir (1983a, 1985, 19861, Kundu

and Sharma (1988a). However, almost all these cytological studies have

concentrated mainly on the determination of chromosome number and provided

scarcely any data on chromosome morphology and structure in detail.

Karyomorphological studies are of paramount importance as they often provide

authentic information pertaining to chromosome structure, number and in

general their gross morphology (Darlington and La Cour, 1970). Detailed

studies pertaining to chromosome length, disparity index of chromosomes,

variation coefficient among chromosome complement and total forma percentage

are necessary in order to throw light on the phylogenetic relationships among

the taxa of flowering plants, since chromosome number tend to exhibit a

greater consistency than any other character. (Yoshikane and Naohiro, 1991).

Moreover,Jones (1978) pointed out that in order to determine the polarity of

karyotype evolution, it is useful to integrate the study of karyotype

morphology with other aspects of the organism, like chemical constitution.

Therefore in the present investigation karyomorphological analysis is

conducted on fifty four members of South Indian Lamiaceae with the aid of "---- " ,-..

1 .' $8 &. d the improved techniques (Sharma and Sharma, 1980) in order to .kt& . yp

number, structure and behaviour of their

correlation between the chromosome

quantitative changes of their essential oil

Fifty four taxa of Lamiaceae has been collected from different parts of South

India, situated between the north latitudes 8 and 20 and the east longitudes

74 and 86. (vide. Table 1; Fig.1).

Table 1 --

List of aerbers investigated

S1. No.

1.

2.

3.

4.

5 .

6.

7 .

8.

9.

10.

11.

12.

13.

14.

15.

Name of the taxa

Ocimum adscendens Willd.

0- americanum L.(syn 0. canum Sims)

0. basilicum L.var. glabratum Benth.

- 0. basilicum L.var pilosum Benth.

- 0. basilicum L.var. purpurascens Benth.

O, basilicum L.var thyrsiflorum Benth.

0. - L. -

O, gratissimum L.var. suavis H0ok.f.

- 0. tenuiflorum L.f.(syn. - 0. sanctum L.) cv. green

- 0. tenuiflorum L.f.cv. purple

O, tenuiflorum L.f. cv.purple-green

- 0. tenuiflorum L.f.var. hirsuta H0ok.f.

Acrocephalus capitatus Benth.

Orthosiphon glabratus Benth.

& glabratus var. parviflorus Benth.

Locality of collection

Cherthalai

Calicut

Cherthalai

Mysore

Wynad

Kodaikanal

Cochin

Bangalore

Kottayam

Munnar

Cochin

Ooty

Cochin

Kodaikanal

Palani

Altitude in metres (approx) .

Sea Level

45

Sea Level

450

1200

2100

Sea Level

920

80

1050

Sea Level

2290

Sea Level

2100

800

- a. No.

- 16.

17.

18.

19.

20.

21.

22.

23.

24.

25.

26.

27.

28.

29.

30.

31.

32.

33.

34.

35.

36.

37. -

Name of the taxa

0. srandiflorus Bold(syn O, stamineus Benth. ) - cv. lilac

0. grandiflorus Bold cv. white - 0. pallidus Royle - 0. thymiflorus Roth. - Plectranthus nummularius Briq.

P. wightii Benth. - Coleus aromaticus Benth. (syn. amboinicus

Lour. )

C. aromaticus Benth. var. variegata - C. blumei Benth. var. verschaffeltii - C. forskohlii Briq. - C. laciniatus Benth.

C. parviflorus Benth. - C. rehneltianus Berger - C. zeylanicus(Benth.) Cramer - Anisochilus carnosus (L.f.1 Wall.

A. eriocephalus Benth. - Hyptis capitata Jacq.

H. suaveolens (L.) Poit. - Pogostemon benghalensis (Burm.f.)Kuntze

P. heyneanus Benth. - P. purpurascens Dalz. - P. vestitus Benth. -

Palode

loty

Kodaikanal

Ponmudi

Yunnar

Cochin

Palani

Bangalore

Cochin

Trivandrul

Palghat

Idukki

Kottayam

Wynad

Cochin

Munnar

Ponmudi

Palghat

Cochin

ltitude n metres approx) .

180

2290

2100

900

1050

Sea Level

800

920

Sea Level

50

150

1000

80

1200

Sea Level

1050

900

150

Sea Leve:

1 . Ocimum adscendens Willd. (Fig.2a) Annual herb; stem fleshy, erect or

prostrate, quadrangular; leaves somewhat fleshy, broadly serrate, ovate-

obovate, membraneous; inflorescence a lax terminal raceme; flowers small,

S1. No.

38.

39 .

40 .

41 .

42.

43 .

44.

45.

46.

47.

48.

49 .

50.

51.

52.

53 .

54 .

blue; tropical.

Altitude in metres (approx) .

45

2290

1800

Sea Level

450

1050

1000

Sea Level

900

Sea Level

1050

45

1220

800

900

Name of the taxa

Eusteralis quadrifolia (Benth.) Panigrahi

Mentha arvensis L.

- M. rotundifolia (L.) Huds.

M. spicata (L.) Huds.

Salvia coccinea Juss.

& leucantha Cav.

- S. plebeia R.Br.

- S. splendens Ker-Gawl.

Anisomeles indica (L.) Kuntze

- A. malabarica R.Br.

Leucas aspera Spreng.

Z, cephalotes Spreng.

- L. linifolia Spreng.

- L. stricta Benth.

- L. vestita Benth.

Leonotis nepetifolia (L.) Ait.f.

Teucrium plectranthoides Gamble

Locality of collection

Calicut

Ooty

Coonoor

Cochin

Mysore

Munnar

Idukki

Cochin

Ponmudi

Cochin

Munnar

Calicut

Devikolam

Palani

Thekkadi

2. 0. americanum L. (Fig. 3a) 'Hoary basil'; origin - Tropical Africa;

perennial herb; stem 4 - angled, grooved; leaves glabrous, simple,

elliptic, slightly crenate; inflorescence a terminal raceme; flowers

white; tropical.

3. 0. basilicum L.var. glabratum Benth. (Fig. 4a) 'French basil'; origin -

France; perennial herb; stem four angled, woody; leaves ovate, glabrous,

margins serrate, acuminate at apex; inflorescence a long raceme; flowers

yellow; tropical and subtropical.

4. 0. basilicum L. var pilosum Benth. (Fig. 5a) Perennial herb; stem

quadrangular, highly pubescent; leaves ovate, acute and pilose with

glandular hairs, margins sparingly serrate; inflorescence a slender

raceme, axis pilose; flowers white; tropical and subtropical.

5. 0. basilicum L. var. purpurascens Benth. (Fig. 6a) 'Sweet basil'; origin

- Tropical Asia, Africa and Pacific Islands; perennial herb; stems

square, tender parts purple; leaves opposite, petioles long, blades

broad, ovate-acute, coarsely crenate, inflorescence a spike, terminal or

axillary; flowers pink or purple; tropical and subtropical.

6. 0. basilicum L. var. thyrsiflorum Benth. (Fig. 7a) Perennial herb; stem

stout, tetragonous; branchlets glabrascent; leaves elliptic - lanceolate, thick, chartaceous, margin serrate, apex narrowly acute; inflorescence

typical thyrse - like racemes; flowers pink, tropical and subtropical.

7. 0. gratissimum L. (Fig.8a) 'Shrubby basil'; origin-Tropical Africa,

America and India; perennial undershrub; stem 4 - angled, 4 - grooved,

young branches pubescent; leaves large, obovate, acute at apex, cuneate

Fig. 2a

Fig. 5a I!---,, Fig. 6a

- Fig. 4a

I I Fig. 7a

Fig. 10a

at base, crenate-serrate, long-petioled, glabrous except the nerves;

inflorescence a terminal raceme; flowers pale greenish yellow; tropical.

8. 0. gratissinn L. var. suavis H0ok.f. (Fig.9a) Perennial undershrub;

stern-&angular; leaves elliptic-lanceolate, acute, coarsely crenate,

pubescent and gland-dotted on both sides, base cuneate, petioles slender;

inflorescence a raceme; flowers greenish-yellow, pedicels softly hairy;

tropical and subtropical.

9. 0. tenuiflonm L.f. cv. green (Fig. 10a) 'Holy basil'; origin-Tropics of

the Old World; perennial herb; stem woody at base, branches clothed with

soft pubescent hairs, greenish, leaf blade elliptic-oblong, apex obtuse,

margins subterrete, hairy on both surfaces; inflorescence a thyrsus,

slender; flowers in close whorls, purplish-pink; tropical.

10. 0. tenuiflorm L.f. cv. purple (Fig. lla) 'Sacred basil'; perennial herb;

stem woody and purplish; leaves ovate-oblong; crenate-serrate, purple,

hairy; inflorescence a lax raceme of verticils; flowers purple in colour;

tropical.

11. 0. tenuiflonm L.f. cv. purple-green (Fig. 12a) Perennial herb; stem

solid, 4-angled, branched, hairy, woody below and herbaceous above;

leaves simple, opposite and decussate, ovate, serrate, acute, purplish-

green; inflorescence a raceme of thyrses; flowers small, pink; tropical.

12. 0. tenuiflorm L.f. var. hirsuta H0ok.f. (Fig. 13a) Perennial herb; stem

slender, hairy, quadrangular, branched; leaves simple, opposite,

petioled, narrowly ovate, toothed, green; inflorescence a thyrsus, axis

pilose; flowers yellow, small; tropical and subtropical.

13. Acrocephalus capitatus Benth. (Fig. 14a) Annual herb; stem erect,

branching from the base, bluntly 4-angled, glabrous; leaves small,

lanceolate, subacute, coarsely serrate, tapering at the base;

inflorescence a terminal capitate head, subtended by a pair of foliar

bracts; flowers small, pale lilac; tropical.

14. Orthosiphon glabratus Benth. (Fig. 15a) Erect perennial undershrub; stem

glabrous, tetragonous, stout; leaves broadly ovate-cordate, membraneous,

chartaceous, margin deeply crenate, scarcely tomentose on the veins;

inflorescence a lax raceme; flowers medium sized, pinkish white; tropical

and subtropical.

15. 0. glabratus var. parviflorus Benth. (Fig. 16a) Perennial undershrub;

stem glabrous, quadrangular, 4-grooved; leaves broadly ovate, coarsely

serrate, nearly glabrous; inflorescence a short raceme; flowers small,

narrow , pale purplish; tropical.

16. 0. grandiflorus Bold cv. lilac (Fig. 17a) 'Kidney Tea plant' or 'Java

Tea'; origin-Tropical Asia and Malayan Archipelago; erect perennial herb;

stem 4-angled, 4-grooved, stout, purple to green coloured; leaves ovate-

acute, margin coarsely toothed, blade smooth and glabrascent;

inflorescence a large flowered raceme, terminal, whorls crowded; flowers

tubular, pale lilac-blue with long protruding stamens; tropical.

17. 0. grandiflorus Bold cv. uMte(Fig. 18a) Perennial herb; stem

tetragonous, sinused; leaves glabrous, olivegreen; broadly ovate,

crenate-serrate; inflorescence a raceme; flowers large, showy, white with

conspicuous very long filiform stamens; tropical.

Fig. 14a

Fig. 17a

Fig. 12a

Fig-. 18a

Fig . 13a

Fig. 16a

Fig . 19a

18. 0. paJJ.idus Royle (Fig. 19a) A low perennial herb; stem glabrous;

branches spreading; leaves small, ovate, toothed, petiolate, pale green;

inflorescence a small terminal raceme; flowers white; tropical and

subtropical.

19. 0. thymiflorus Roth. (Fig. 20a) Perennial herb; stem 4-angular,

glabrascent; leaves broadly ovate to cordate acute, chartaceous, margin

serrate; inflorescence a terminal raceme; flowers pale purple; tropical.

20. Plectranthus nltmmularius Briq. (Fig.2la) 'Swedish Ivy'; origin-Australia

and Pacific Islands; perennial herb; stem weak, creeping; leaves

leathery, metallic green, waxy, rounded, deeply crenate; inflorescence a

lax spike; flowers white; tropical and subtropical.

21. P. wightii Benth. (Fig.22a) Tall robust; stem scaberulous; leaves broadly

ovate-cordate, crenate-toothed, rigid; inflorescence a large pyramidal

panicle; flowers blue; tropical and subtropical.

22. Coleus armticus Benth. (Fig. 23a) 'Wild Oregano' or 'Indian Borage';

origin-Malaya and Indonesia; fleshy perennial herb; stem weak, obturely

4-angled, pubescent; ,leaves succulent, ovate-elliptic, margins finely

serrate, light green, puberulous, petioles slender; tropical and

subtropical.

23. C. arovlticus Benth. var.variegata (Fig. 24a) 'Variegated Borage',

perennial herb; stem fleshy, puberulous; leaves showy, variegated, thick,

margin crenate, obovate, finely tomentose; petioles greenish white;

tropical and subtropical.

24. C. bl-i Benth. var verschaffeltii (Fig. 25a) 'Painted nettle'; origin-

Java; perennial herb; stem square, half hardy, purple; leaves ovate,

margin toothed, showy, blade shows crimson coloured patches, purple in

centre, border narrow green; petiole purplish-white; inflorescence a

branched spike; flowers pale blue; tropical.

25. C. forskohlii Briq. (Fig. 26a) Perennial herb; stem obtusely 4-angled,

woody at base, pubescent; leaves large, thick, ovate-oblong, hispidly

tomentose on both sides, margins crenate; tropical and subtropical.

26. C. laciniatus Benth. (Fig. 27a) Perennial herb; stem fleshy; leaves soft,

torn or slashed into narrow lobes, attractively coloured; inflorescence a

branched lax spike; flowers light purple; tropical.

27. C. parviflorus Benth. (Fig. 28a) 'Country potato', perennial herb; stem

tetragonous, fleshy, produces tubers at the base; leaves long petioled,

membraneous, chartaceous, margin crenate, rarely pubescent, grassy green;

tropical.

28. C. rehneltianus Berger (Fig. 29a) 'Red Trailing Queen'; perennial herb;

robust, plant with half-hardy, quadrangular, purple stem; leaves broad-

ovate, margins slightly lobed, blade deep purple coloured with carmine in

the centre and a broad border of dullgreen; tropical.

29. C. zeylanlcus (Benth.) Cramer (Fig. 30a) An aromatic, perennial herb;

stem obscurely tetragonous, pubescent, light green in tender parts,

slightly brownish when old; leaves large, thin, soft, velvetty, rounded,

puberulous, yellowish green; tropical and subtropical.

Fig. 23a Fig. 24a

Fig . 22a

F i g . 26a Fig. 27a

30. Anisochilu carnosus (L.f.) Wall. (Fig. 31a) A gregarious annual

subshrub; stem stout, bluntly 4-angled, glabrous, often tinged with red;

leaves rather fleshy, ovate-cordate, pubescent, margin serrate,

deciduous; inflorescence a strobiloid head like spike; flowers

palepurple; tropical.

31. A. eriocephalu Benth. (Fig. 32a) Annual herb; stem and branches

puberulous; leaves ovate-orbicular, crenate, dark green; inflorescence a

dense cylindrical spike; flowers lavender blue; tropical.

32. Hyptis capitata Jacq. (Fig. 33a) Annual subshrub; stem half-hardy,

distinctly tetrangular, sparingly pubescent; leaves large, ovate-oblong,

margin crenate-serrate, coarse, dark green; inflorescence a capitate

head, axillary; flowers minute white; tropical and subtropical.

33. H. suaveolens (L . ) Poit. (Fig. 34a) Origin-Tropical America and West

Indies; aromatic sweet smelling annual herb; stem tetragonal, roughly

hairy; leaves ovate-cordate, denticulate, strigose; inflorescence a

verticillaster of stalked cymes; flowers small, blue; tropical.

34. Pogostemon benghalensis (Burm.f.1 Kuntze (Fig. 35a) Perennial herb; stem

and branches subquadrangular, covered by grey tomentose; leaves ovate-

oblong, doubly serrate, puberulous; inflorescence an erect terminal

panicle formed of many short spikes; flowers pink-purple; tropical and

subtropical.

35. P. heyneanus Benth.(Fig. 36a) Perennial undershrub; stem and branches

glabrous, nodes swollen, obliquely 4-angled, brownish green; leaves

large, ovate, sparingly puberulous, greyish green; tropical and

subtropical.

4 Fig. 29a

Ffg. 33a

Fig. 31a

Fig. 37a

36. P. purpurascens Dalz. (Fig. 37a) Perennial herb; stem half-hardy,

branchlets hispid; leaves in unequal pairs, ovate-acuminate, inciso-

serrate, pubescent; inflorescence a compact panicle, formed of flattened

spikes; flowers purplish-white; tropical and subtropical.

37. P. vestitus Benth (Fig. 38a) Perennial herb; stem densely wooly; leaves

ovate-obtuse, serrate, pubescent; inflorescence a unilateral panicle,

bracts foliaceous; flowers purple, small; tropical.

38. Eusteralis quadrifolia (Benth.) Panigrahi (Fig. 39a) An annual aromatic

woody herb; leaves in whorls of four, narrowly oblong-obtuse, clothed by

dense greyish tomentose; inflorescence a long dense terminal spike;

flowers rosy pink, minute; tropical.

39. Hentha amensis L. (Fig. 40a) 'Japaneese mint' or 'Field mint'; origin

Japaneese Archipelago; perennial aromatic herb with running leafy

stolons; stem square, branches ascending; leaves broadly oblong,

puberulent, sharply serrate; temperate and subtropical.

40. M. rotundifolia (L.) Huds. (Fig. 41a) 'Round-leaved mint'; origin-

Temperate Asia; perennial herb with running root stock; stem slender

without much branches, glabrous; leaves nearly round, crenate, obtuse,

dark green; temperate and subtropical.

41. H. spicata (L.) Huds. (Fig. 42a) 'Spear mint'; origin-Temperate Europe

and Asia; perennial herb; rootstocks spreading; stem erect; leaves

oblong-lanceolate, serrate, much wrinkled, chartaceous, finely pubescent

on the veins; temperate and subtropical.

42. Salvia coccinea Juss. (Fig. 43a) 'Red sage' or 'Texas sage'; origin-

Central and Tropical Aierica; perennial undershrub; stem sparsely hairy;

leaves subcordiform, puberulous, crenate-serrate, gray-green;

inflorescence a lax raceme; flowers deep red; tropical and subtropical.

43. S. leucantha Cav. (Fig. 44a) 'Mexican bush sage'; origin-Central Mexico;

perennial wooly undershrub; stem and branches tomentose; leaves, narrow,

oblong-lanceolate, crenate, blade white lanate beneath; inflorescence a

long slender spike, floral whorls remote at the base; flowers purple,

covered by downy felt; subtropical.

44. S. plebeia R.Br. (Fig. 45a) A roughly pubescent annual subshrub; stem

stout, fastigiately branched, obtusely 4-angled and grooved; leaves

oblong-lanceolate, crenate-serrate, rugose; inflorescence a panicle of

compact racemes; flowers small, blue; tropical.

45. S. splendens K&- awl. (Fig. 46a) 'Scarlet sage'; origin - Brazil;

perennial undershrub; stem half hardy, branches glabrous; leaves ovate-

acuminate, crenate-serrate; inflorescence a terminal spike like raceme;

flowers showy, scarlet-red; tropical.

46. Adscaeles indica (L.)Kuntze (Fig. 47a) Perennial under shrub; stem

acutely tetragonous, covered with greyish tomentose, branches softly

hairy; leaves obovate, chartaceous, crenate, pubescent; inflorescence,

terminal or axillary dense spikes formed of sessile cymes; flower

purplish - white; tropical.

47. A lalabarica R.Br. (Fig. 48a) Perennial undershrub; stem obtusely

tetragonous, pubescent; leaves oblong-lanceolate, sub-coriaceous;

crenate-serrate, blade pale green above and white below, softly wooly;

inflorescence a dense spike; flowers pale purple, pubescent; tropical.

F i g . 44a

Fig* 3%.

Fig. 42a

Fig. 45a F i g . 46a

48. Leuas aspera Spreng. (Fig. 49a) Coarse annual herb; stem square, grooved

on all sides, rugose; leaves linear-oblong, toothed, coarse hairy;

inflorescence a verticillaster; flowers white, sessile; tropical.

49. L. cephalotes Spreng. (Fig. 50a) Annual herb; stem sparsely pubescent,

tetragonous; leaves linear-oblong, toothed, roughly hairy; inflorescence

a verticillaster, slightly pendant; flowers white; tropical.

50. L. linifolia Spreng. (Fig. 51a) Annual herb; stem 4-angled, pubescent,

branched below; leaves linear-lanceolate, narrow, almost entire, covered

by appressed hairs; inflorescence a verticillaster; flowers white,

shortly stalked, finely pubescent; tropical.

51. L. stricta Benth. Annual herb; stem upright, stout; leaves narrow,

coarse, elliptic-lanceolate; densely hairy; inflorescence a

verticillaster; flowers white; tropical.

52. L. vestita Benth. (Fig. 53a) Annual herb; stem quadrangular, tawny

villous with brownish felt; leaves lanceolate, crenate-serrate,

pubescent; inflorescence a verticillaster; flowers white, bearded;

subtropical.

53. Leo~tis nepetifolia ( L . ) Ait.f. (Fig. 54a) Origin-Tropical America;

perennial glabrous undershrub; stem 4-angled, stout, erect; leaves ovate-

elliptic, soft, pubescent, crenate-serrate; inflorescence a

verticillaster formed of gorgeous many flowered whorls; flowers showy

orange- red with hooded downy corolla and spinescent calyx; tro.pica1.

54. Teucrium plectranthoides Gamble (Fig. 55a) Perennial undershrub; stem

branches robust, quadrangular, glabrous; leaves long, greyish green,

finely serrate; inflorescence a spreading panicle of open racemes;

flowers yellowish white; tropical.

Fig. 50a

Fig. 48a Fig. 49a

47a-Habit of Anisomeles indica (L.) Kuntze, 48a-A.malabarica - R . B r . , 49a-Lcucas aspera Spreng., 50a-L. cephalotes Spreng., 51a-L.linifo1ia Spreng., 52a-L.stricta Benth. 53a-%.vestita Benth., 54a-Leonotis nepetifolia (L.) Ait.f., 55a-Teucrium plectranthoizes Gamble.

METHODS

A. Cytological studies

1. Mitotic squash experiments

Somatic chromosome studies have been conducted on fifty four members

of South Indian Lamiaceae with the help of improved cytotechniques

(Sharma and Sharma, 1980). Squash experiments were carried out on

root tip meristem at mitotic metaphase stage. Young healthy root

tips are collected from the vegetative cuttings planted in the

experimental botanical garden, at the periods showing peak mitotic

frequency, i.e., 9.00 A.M. to 10.30 A.M. In plant cutting with a

lesser rooting ability, seeds are germinated in petridishes lined

with moist filter paper under laboratory conditions. The root tips

are then washed thoroughly in distilled water and pretreated with

different pretreating chemicals. Saturated aqueous paradichloro

benzene solution with a trace of aesculine is found to be most

suitable for the members of Lamiaceae. The root tips are initially

chilled at 0 - 5 O ~ for 4-6 minutes and then kept at 12-14O~ for 1-3

hours for obtaining the best results.

The pretreated root tips are then washed thoroughly with distilled

water and fixed in 1:3 acetic acid - ethyl alcohol mixture (Carnoy,

1886) overnight, followed by 3-7 minutes treatment in 45% acetic

acid and stained with 2% aceto-orcein - 1 N HC1 mixture (9:l) for 2

112 to 3 112 hours. While staining, the vial is initially warmed

over a flame for effective results. The stained root tips are then

squashed in 45% acetic acid and the preparation was then temporarily

sealed.

To remove the excess oil deposits, rather common in Lamiaceae, it

has been found preferable to add a little saponin along with the

pretreatment mixture and shake thoroughly.

2. Meiotic smear experiments

pollen mother cell (PMC) analysis were carried out on those members

which bloomed frequently in the experimental garden. For this flower

buds are fixed in modified Carnoy's fluid (1 acetic acid : 3

chloroform : 6 ethyl alcohol) for 1-2 days and stored in 70% ethyl

alcohol. Young anthers are smeared with 2% aceto-carmine stain to

obtain PMC's showing divisional stages.

For better results, the fixing mixture is replaced by fresh Carnoy's

fluid one or two times, which helpes to remove the excess cell

inclusions.

B. Detailed Karyaorphometrical analysis

1. Drawings - and Wotoricrographs

Karyotype drawings are made with the help of a Leitz-BIOMED tracing

&vice with a magnification of X 2400 (approximately). The

photomicrographs are taken using 125 ASA 35mm ORWO film in a Leitz

photographic attachment and suitably enlarged. The idiograms are

prepared after comparing atleast five well spread metaphase plates.

2. Karyomorphological studies

The total length as well as the short arm length of all the

chromosomes of the fifty four cytotypes are measured accurately

using micrometers. In all the karyotypes, ratio of the short arm to

the total length of the chromosome in percentage, F% (Forma

percentage or Centromeric index) is determined after Krikorian, et

al. (1983). On the basis of F% the nature of primary constriction in

the chromosomes are classified as follows:

The disparity index (DI) of chromosomes in a karyotype is

calculated after Mohanty, et al. (1991), by the formula:

F%

50

49.9 - 37.6 37.5 - 25.1

25

24.9 - 18.76

18.75 - 12.6

12.5

12.4 - 6.26

6.25 - 1 1

DI = Longest chromosome - Shortest chromosome x 100 Longest chromosome + Shortest chromosome

Nature of Primary Constriction

Median

Nearly median

Nearly submedian

Submedian

Nearly submedian

Nearly subterminal

Subterminal

Nearly subterminal

Extremely subterminal

Terminal

-

The variation coefficient among chromosome complements (VC) is

determined after Verma (1980) as follows:

VC = Standard deviation X 100 Mean length of chromosomes

The total forma percentage or the mean centromeric index value (TFX)

is calculated in each taxa after Huziwara (1962), by the formula :

TF% = Total sum of short arm lenth X 100 - - - - - - - - Total sum of chromosome lenth

For the calculation of polyploidy, the base numbers given in

Darlington and Wylie (1955), Morton (1956a, 1962), as well as Love

and Love (1961b), are usually followed. In those cases where the

basic numbers are not mentioned in these references, the latest

literature is consulted.

In the present investigation, fifty four members of South Indian Lamiaceae including

twelve cytotypes in the genus Ocimum L., one in Acrocephalus Benth., six in

Orthosiphon Benth., two in Plectranthus L' Herit., eight in Coleus Lour., two in

Anisochilus Wall., two in Hyptis Jacq., four in Pogostenon Desf., one in Eusteralis

Rafin., three in Mentha L., four in Salvia L., two in Anisomeles R.Br., five in

Leucas R.Br., one in Leonotis R.Br. and one in Teucrium L. were analysed. The

normal somatic chromosome number ranges from 2n = 16+1-OB to 96. However,

numerical variations have been recorded in many species. The ploidy level exhibited

by different taxa range from diploidy to octaploidy. The chromosome pair with

secondary constriction is found to range between one and three. The karyotypes are

characterized by the presence of comparatively small chromosomes, ranging from 4.2 p

to 1.0 p in length. The chromosomes in each karyotype decrease in size progressively

and bear nearly median to nearly submedian primary constriction.

The general description of the common chromosome types is given below, followed by a

karyotype description for the members investigated.

Type A : Comparatively long chromosome with two constrictions, one median and the

other nearly sibmedian in position (4.2 to 1.4 p).

Type B : Comparatively long chromosome with nearly median to nearly submedian

primary constriction and a satellite at the distal end of shorter arm,

joined by a SAT thread (2.8 to 2.2 p) .

Type C : Relatively long chromosomes (4.2 - 2.0~) with nealy median to nearly

submedian primary constriction.

Type D : Medium to short chromosomes (<2.0p) with nearly median to nearly

submedian primary constriction.

Diagrammatic representation of the different chromosome types observed in the South Indian taxa of Lamiaceae is shown below.

L-__r . 'I'y A 'I 'y IN: II

Diagrammatic representation of different chromosome

types found in the taxa investigated in Laoiaceae

Chromosome count on pollen mother cell

Normal somatic chromosome number

Somatic variation numbers

Chromosome pairs with secondary constriction :

Range of chromosome length

Average chromosome length

Total chromatin length

Chromatin length of basic complement

Disparity index (DI)

Variation coefficient (VC)

TF value (%)

n = 11 (Fig. 2j)

2n = 22 (Fig. 2b, 2-2)

2n = 24 (Fig. 2e)

1

3 . 2 ~ to 1 . 8 ~

2.31~

50.

25.4 p

28.00

14.93

40.08

Table 2 --

Detailed karyotype analysis (Pig. 2d)

Chromo- some type

A

C

C

C

C

D

No. of pairs

1

4

1

2

2

1

Total length in AI

3.2

2.4

2.4

2.2

2.0

1.8 --

Nature of primary constriction

-

Nearly median

Nearly submedian

Nearly median

Remarks

Chromosome pair with secondary constriction

-

-

- -

-

Short arm length in AI

0.6 0.8

1.0

0.8

0.8

0.8

0.8

' FX

18.75 25.00

41.67

33.33

36.36

40.00

44.44 --

Chromosome count on pollen mother cell : n = 36 (Fig 3 j )

Normal somatic chromosome number : 2n = 72 (Figs. 3b, 3c)

Somatic variation numbers : 2n=18, 40 and 48 (Figs. 3e,3f ,3g)

Chromosome pair with secondary constriction : 2

Range of chromosome length : 4.2 p to 1.4 p

Average chromosome length : 2 . 1 5 ~

Total chromatin length : 154.8 p

Chromatin length of basic complement : 25.8 p

Disparity index (DI) : 50

Variation coefficient (VC) : 28.28

TF value (%) : 43.04

Table 3 --


Chromo- some type

A

A

C

C

C

C

C

C

C

D

D

D

Short arm length in p

1 .0 1.2

0.8 1.0

1.4

1.2

1 . 0

1 .0

1.0

0 .8

0.8

0 . 8

0.6

0.6

No. of pairs

1

1

1

1

2

2

4

3

5

7

7

2

F%

23.81 28.57

23.53 29 .41

43.75

40.00

35.71

38.46

41.67

36.36

40.00

44.44

37.50

42.86

Total length in u

4.2

3.4

3.2

3.0

2.8

2.6

2.4

2.2

2.0

1.8

1 .6

1.4


-

-

Nearly median

Nearly submedian

Nearly median

Nearly submedian

Nearly median

Nearly submedian

Nearly median

Remarks


-

- -

-

-

-

-

-

-

-

A\ p?

O c h u m basilicum L.var. glabratum Benth. (2n=4x=48=A4C12D32)

Chromosome count on pollen mother cell : n = 24 (Fig. 4j)

Normal somatic chromosome number : 2n = 48 (Figs. 4b,4c)



Average chromosome length : 1.73 p



Disparity index (DI) : 29.41

Variation coefficient (VC) : 16.57 s ,

TF value (%I : 40.57.'-. . ,

Table 4 --

Detailed karyotype analysis (Fig.4d)

Remarks


-

-

-

-

-

-

-

Chromo- some type

A

A

C

C

D

D

D

D

D

F%

18.18 27.27

18.18 36.36

45.45

40.00

33.33

44.44

37.50

42.86

33.33


-

-

Nearly median

Nearly submedian

Nearly median

Nearly submedian

Nearly median

Nearly submedian

No. of pairs

1

1

1

5

2

2

6

5

1

Total length in p

2.2

2.2

2.2

2.0

1.8

1.8

1.6

1.4

1.2


0.4 0.6

0.4 0.8

1.0

0.8

0.6

0.8

0.6

0.6

0.4

Fig. 2c

Fig. 3g

Fig. 3c

Fig. 3f

# Fig. 4c

Fig. 2c, 2d - Karyotype and idiogram of -- Ocimum adscendens Willd. (2n=22)x2400 approx., 3c, 3d, 3f, 3g - - 0. americanum L.: 3c; 3d Karyotype and idiogran ( 2 ~ 7 2 ) x 2400 amrox.. 3f. 3~ somatic variants with 211-40 and 2n=48 , - respectively; 4c, ii - bryotype and idiograrn of - 0 basilicum L.var.&abratun Benth. (2n=48) x 2400 approx.

-

Fig. 2b .Fig. 2e Fig. 2j

- Fig. 3b Fig. 3 j

- -

Fig. 4b Fig. 4j Pig. 5b

Scale: r 8 wJ

Figs. 2b, 2e, 2 j - Oci mum adscendens Willd. : 2b mitotic metaphase (2n=22), - 212 - somatic variant (2n=24), 2j - meiotic anaphase I (n=ll); 3b, 3e, 33 - 2. americanum L.: 3b - mitotic metaphase ( 2 ~ 7 2 1 , 3e - somatic variant (2nX18), 3j - meiotic anaphase I (abnormal seggregation); 4b, 4j - 0. basilicum L. var. - labracum Benth.: 4b - mitotic metaphase (2n=48), 4j - meiotic metaphase I

fn=24); 5b - - 0 . bnsilicum L. var. pilosum Benth. - mitotic metaphase (2n=48).

Oc-um basilium L.var. pilosum Benth. (2n=4~=48=A4CZOD24)



Chromosome pair with secondary constriction :







TF value (%)

n = 24 (Fig. 5j)

2n = 48 (figs. 5b, 5c)

2

3.2 p to 1.4 p

1.96 1.1

94 1.1

23.5 p

39.13

19.76

41.14

Table 5 --

Detailed karyotype analysis (Pig. M)

Remarks


-

-

-

-

-

-

F%

25.00 31.25

23.08 30.77

41.67

36.36

40.00

44.44

37.50

42.86

Chromo- some type

A

A

C

C

C

D

D

D


-

-

Nearly median

Nearly submedian

Nearly median

Nearly submedian

Nearly median

Total length in p

3.2

2.6

2.4

2.2

2.0

1.8

1.6

1.4

No. of pairs

1

I

1

4

5

5

6

1


0.8 1.0

0.6 0.8

1.0

0.8

- 0.8

0.8

0.6

0.6

O c i n n basillcum L. var . purpurascens Benth. (211=4~=48=A4C28D16)



Chromosome pairs with secondary constriction : 2

Range of chromosome length : 2.8 p to 1.6 1.1



Chromatin length of basic complement : 2 5 . 1 ~



TF value (%) : 42.79%

Table 6 -- Detailed karyotype analysis (Pig. 6d)

Chromo- some type

A

A

C

C

C

C

C

D

D

No. of pairs

1

1

1

3

4

1

5

6

2

Total length in 1.1

2.8

2.6

2.6

2.4

2.2

2.2

2.0

1.8

1.6

Short arm length in 0

0.6 0.8

0.6 0.8

1.0

1.0

1.0

0.8

0.8

0.8

0.6

F%

21.43 28.57

23.08 30.77

38.46

41.67

45.45

36.36

40.00

44.44

37.50


-

-

Nearly median

Nearly submedian

Nearly median

Nearly submedian

Remarks


-

- -

- -

- -

Ocimum basilicum L.var. thyrsiflorum Benth. (2n=4~=48=A4C22D22)

Chromosome count on pollen mother cell : n = 24 (Fig. 7 j )

Normal somatic chromosome number : 2n = 48 (Figs. 7 b , 7c)


Range of chromosome length : 3.4 fl to 1.4 f l

Average chromosome length : 2.02 /.I





TF value (%) : 40.39

Table 7 -- Detailed Laryotype analysis (Pig. 7d)

Chromo- some type

A

A

C

C

C

C

D

D

D

D

No. of pairs

1

1

1

2

2

6

2

3

5

1

Total length in p

3.4

2.8

2.6

2.4

2.2

2.0

1.8

1.8

1.6

1.4


0.8 1.0

0.6 0.8

1.0

0.8

1.0

0 .8

0.6

0.8

0.6

0.6 --

F%

23.53 29.41

21.43 28.57

38.46

33.33

45.45

40.00

33.33

44.44

37.50

42.86


-

-

Nearly median

Nearly submedian

Nearly median

Nearly submedian

Nearly median

Nearly submedian

Nearly median

Remarks


-

-

- - - -

- -

Fig. 5c Fig. 7c

Fig. ,6c

Fig. 5c, 5d - Karyotype and idiogram of Ocimum basilicum L.var.pilosum Benth. i2n=48) x 2400 approx. 6c,bd - Karyotype and idiogram of 0 . basilicum L. var.purpurascen Benth. ( 2 ~ 4 8 ) x 2400 approx.; 7c,7d - ~aryoty~e-and idiogram of 0 . basilicum L var. thyrsiflorum Benth ( 2 ~ 4 8 ) x 2400 approx. - .- -



Somatic variation numbers : 2n=34 and 42 (Figs. 8 e , 8 f )

Chromosome pairs with secondry constriction : 2




Chromatin length of basic complement : 2 0 . 1 ~



TF value (%) : 40.66

Table 8 --


Remarks


Chromosome pair with a satellite

-

- -

-

-

-

Chromo- some type

A

B

C

C

C

D

D

D


-

-

Nearly median

Nearly submedian

Nearly median

Nearly submedian

Nearly median

Nearly submedian

No. of pairs

1

1

2

2

5

2

3

4

Total length in p -

2.8

2.8

2.4

2 .2

2.0

1 . 8

1 .8

1.6


0 . 6 0 .8

0 . 8 0 .8

1 .0

0 . 8

0 .8

0 . 6

0 .8

0 . 6

F%

21.43 28.57

28.57 28.57

41.67

36.36

40.00

33.33

44.44

37.50

- -- - ---

Fig. 5j Fig. 6b Fig. 6 j

-- - -

Fig. 7b Fig. 7j Fig. 8b

Fig. 8e Fig. 8j Fig. 8m

Scale: I.----r BP' - --!

Fig. 5j - Ocimum basilicum L. var. pilosum Benth.- meiotic metaphase I ( ~ 2 4 ) ; 6b, 6 j - - 0 . basilicum L. var. purpurascens Benth.: 6b - mitotic metaphase (2n=48), 6j-meiotic metaphase I (n=24); 7b, 7j - - 0. basilicum L. var. thyrsiflorum Benth.: 7b - mitotic metaphase (2n=48), 7j - meiotic anaphase I (n=24); 8b, 8e, 8j, 8m - - 0. gratissimum L.: 8b - mitotic metaphase (2nz40), 8e - somatic variant (2n = 34), 8j - meiotic metaphase I (abnormal with 14 bivalents and 12 univalents), 8m - normal tetrad.

Ocimum gratissimum L.var. suavis H0ok.f. (2n=4~=48=A2B2C20D24)

Chromosome count on pollen mother cell : n = 24 (Fig 9j)



: 2n = 48 (Fig. 9b, 9c)

: 2 n ~ 3 6 , 38and42 (Figs. 9e,9f ,9g)








TF value (%) : 42.64


Chromo- some type

A

B

C

C

C

D

D

D


-

-

Nearly median

Nearly submedian

Nearly median

Remarks


Chromosome pair with a satellite

- -

-

-

-

-

No. of pairs

1

1

1

2

7

6

4

2


0.6 0.8

0.6 0.6

1.0

1.0

0.8

0.8

0.6

0.6

Total length in p

2.6

2.2

2.4

2.2

2.0

1.8

1.6

1.4

F%

23.08 30.77

27.27 27.27

41.67

45.45

40.00

44.44

37.50

42.86

Fig. 8c

Fig. 9e

Fig. 8f

Fig. 9f

Fig. 9c

Fig. 9g

Fig. 8c,8d,8f - Ocimum gratissimum L.: 8c, 8d - Karyotype and idiogram ( 2 ~ 4 0 ) x 2400 approx., 8f - somatic variant (2n=42); 9c, 9d, 9e, 9f. 9g - 0. gratissirnum L.var.suavis H0ok.f. : 9c,9d - Karyotype and idiogram (2n=48) x 2400 approx., 9e,9f,9g - somatic variants with 2n=36, 2n=38 and 2n=42 respectively.

Ocimm tenuiflonm L. f . cv. green (2n=4x=32=A4C4D24)










TF value (%)

n = 16 (Figs. 10j)

2n = 32 (Figs. lob, 10c)

2

2.0 @ to 1.2 p

1.65 ,u

52.8 p

13.2 p

25.00

16.87

43.18

Table 10 --

Detailed Karyotype analysis (Fig. 10d)

Remarks

Chromosome pair with secondary constriction.

- -

- -

-

I I I


-

Nearly median

Nearly submedian

Nearly median

Nearly submedian


0.6 0.6

0.8

0.8

0.6

0.6

0.4

Total length in ,u

2.0

2.0

1.8

1.6

1.4

1.2

Chromo- some type

A

C

D

D

D

D

F%

30.00 30.00

40.00

44.44

37.50

42.86

33.33

No. of pairs

2

2

4

2

4

2

Ocinn tenuiflonm L.f. cv. purple (2n=4x=32=A4D28)










TF value (%)

n = 16 (Fig. llj)

2n = 32 (Figs. llb, llc)

2

1.8 ,u to 1.0 ,u

1.49 p

47.6 p

11.9 u

28.57

18.36

35.93

Table 11 --

Detailed Karyotype analysis (Fig. lld)

Remarks


- -

-

- -


-

-

Nearly submedian

D 1 2 1 1.0 I 0.4 1 40.00 I Nearly median

F%

22.22 33.33

25.00 25.00

33.33

37.50

28.57

33.33


0.4 0.6

0.4 0.4

0.6

0.6

0.4

0.4

Total length in p

1.8

1.6

1.8

1.6

1.4

1.2

Chromo- some type

A

A

D

D

D

D

No. of pairs

1

1

4

2

4

2

-- -- - - - - .- -

Fig. 8n Fig. 80 Fig. 8 p

- -. -- Fig. 9b Fig. 9 j Fig. lob

Fig. 1 O j - -- Fig. l l b

Scale: - 73 I C ~

Fig. l l j

Figs. an , 80, 8p - Ocimum gratissimum L. showing polyspory: 8n - pentad, 80 - hexad, 8p - octad; 9b, 9j - - 0 . - gratissimum L.var. suavis Hook. f.: 9b - mitotic metaphase (2n=48), 9 j - meiotic anaphase I (abnormal seggregation); lob, 10 j - 0 . tenuiflorum L . f . cv. p e e n : lob - mitotic metaphase (2n=32), 10j. - meioti; meta~hase I (n=16) ; I l b , 11 j - 0 . tenuiflorum ~ . f . cv. purple: l l b - mitotic metaphase I (2n=32), 11 j - diakiyesis (n=16).

39

Ocimm tenuiflonm L.f. cv. purple-green (2n=4x=36=A4D32)










T F value (%)

n = 18 (Fig. 12j )

2n = 36 (Figs. 12b, 12-2)

2

1 . 8 p to 1 . 0 ,u

1 . 3 8 p

Table 12 --

Detailed Karyotype analysis (Pig. 12d)

Chromo- some type

A

D

D

D

D

D

I l l I I I

No. of pairs

2

2

2

2

2

2

Total length in p

1 . 8

1 .8

1 .6

1 . 4

1 . 2

1 . 0


0 . 4 0 . 6

0 . 6

0 . 6

0 . 6

0 . 4

0 . 4

F%

22.22 33 .33

33.33

37.50

42.85

33.33

40.00


-

Nearly submedian

Nearly median

Nearly submedian

Nearly median

Remarks

Chromosome with secondary constriction.

- - - -

-










TF value ( X I

n = 18 (Fig. 1 3 j )

2n = 36 (Fig. 13b, 13c)

2

Table 13 --


Remarks


-

-

-

-

- -

Chromo- some type

A

A

C

D

D

D

D

D -

F%

20.00 30.00

22.22 33.33

40.00

44.44

37.50

42.86

33.33

40.00


-

-

Nearly median

Nearly submedian

Nearly median

Nearly submedian

Nearly median

No. of pairs

1

1

2

2

3

6

2

1

Total length in p

2.0

1.8

2.0

1 .8

1.6

1.4

1.2

1.0


0.4 0.6

0.4 0.6

0.8

0.8

0.6

0.6

0 .4

0.4

Fig. 1oc Fig. llc

Fig. 12c Fig. 13c

Fig. 10d J8 88, ,#I 11, 11 8 l 1 1 8 ) 8 ) 4 1 I1 11 Il I 1 18,

Fig. lld jfi dd, J1 4; 18 11 18 11 I1 I t I 8 81 81 88 18,

Fig. 10c, 10d - Karyotype and idiogram of -- Ocinun tenuiflorum L.f. cv ereen (2n=32) x 2400 approx., llc, lld - Karyotype and idiogram of 2. tenuiflorm L.f. cv.purple f2n=32) x 2400 approx., 12c, 12d - karyotype and idiogram of 0 tenuiflorum L.f cv purple-green (211-36) x 2400 approx.; 13c, 13d - - Karyotype and idiogran of 2. tenuiflorum L.f. var. hirsuta (2n=36) x 2400 approx

Acrocephalus capitatus Benth. (2n=Zx=l8=AZD16)


Chromosome pair w*.th secondary constriction :







TF value (%)

2n = 18 (Figs. 14b, 14c)

1

1 . 6 ~ to 1.0~

1.36~

24.b

12.w

23.08

13.47

40.62

Table 14 -- Detailed karyotype analysis (Fig. 14d)

Remarks


-

- - -

Chromo- some type

A

D

D

D

D

F%

25.00 25.00

37.50

42.86

33.33

40.00


-

Nearly median

Nearly submedian

Nearly median

No. of pairs

1

1

4

2

1

Total length in p

1.6

1.6

1.4

1.2

1 .O


0.4 0.4

0.6

0.6

0.4

0.4

Orthosiphon glabratus Benth. (2n=Zx=26=AZC24)










TF value (%)

n = 13 (Fig. 15j)

2n = 26 (Figs. 15b, 15c)

1

3.6 p to 2.2 p

2.74 p

71.2 p

35.6 p

24.14

13.24

39.93

Table 15 --


Remarks


-

- -

Chromo- some type

A

C

C

C

F%

22.22 33.33

40.00

38.46

36.36


-

Nearly median

Nearly submedian

No. of pairs

1

4

6

2

Total length in pl

3.6

3.0

2.6

2.2

Short arm length in &I

0.8 1.2

1.2

1.0

0.8

Orthosiphon glabratus var. parviflorus Benth. (2n=2~=26=A2ClZD12)










TF value (%)

n = 13 (Fig. 16j)

2n = 26 (Figs. 16b, 16c)

1

2.8 ,u to 1.4 p

2.06 ,u

53.6 ,u

26.8 ,u

33.33

20.64

41.22

Table '16 --


Remarks


- - - -

Chromo- some type

A

C

C

D

D

F%

21.43 28.57

38.46

36.36

44.44

42.86


-

Nearly median

Nearly submedian

Nearly median

No. of pairs

1

2

4

4

2

Total length in ,u

2.8

2.6

2.2

1.8

1.4


0.6 0.8

1.0

0.8

0.8

0.6

-- --- Fig. 12b Pig. 12; Fig. 13b

--- - - .- - -. - Fig. 13j Fig. 14b Fig. 15b

-- - - - --

Fig. 15j Fig. 16b Fig. 16j

Scale: 811h

Figs. 12b, 12j - Ocimum tenuiflorum L.f. cv.purple-green: 12b - mitotic metaphase (2n=36), 12j-meiotic anaphase I (n=18); 13b, 13j - - 0. tenuiflorum L . f . var. hirsuta H0ok.f.: 13b - mitotic netapllase (2n=36), 13j - meiotic metaphase 1: (n=18); 14b - Acrocephalus capitatus Benth. - mitotic metaphase (2n=,18); lSb , 1 5 j - Orthosiphon glabratus Benth.: 15b - mitotic metaphase (2n=26), 15j - meiotic anaphase 1 (~13); 16b, 1 6 j - 0. glabratus var. parviflorus Benth. : 16b - mitotic metaphase (2~261, 16 j-. meiotic metaph- 'ase I (n=13)

Orthosiphon grandiflorus Bold cv. lilac (Zn=4~=28=A2ClZD14)










TF value (X)

n = 14 (Fig. 17j)

2n = 28 (Figs. 17b, 17c)

1

2.8 ,u to 1.4 p

2.04 p

57.2 p

14.3 p

33.33

21.66

38.6

Table 17 -- Detailed Karyotype analysis (Pig. 17d)

Remarks


- - -

- -

Chromo- some type

A

C

C

C

D

D

D I I

FX

21.43 35.71

38.46

36.36

40.00

33.33

37.50

No.of pairs

1

3

2

1

3

3

0.6 42.86 I Nearly median I -


-

Nearly median

Nearly submedian

Nearly median

Nearly submedian

Total length in p

2.8

2.6

2.2

2.0

1.8

1.6

Shortarm length in p

0.6 1.0

1.0

0.8

0.8

0.6

0.6

Orthosiphon grandiflorus Bold cv. white (2n=4~=28=AZC18D8)










TF value (%)

n = 14 (Fig. 181)

2n = 28 (Figs. lab, 18~)

1

3.2 p to 1.6 ,u

2.2 cl

61.6 p

15.4 p

33.33

22.00

42.1

Table I8 -- Detailed Karyotype analysis (Fig. 18d)

Chromo- some type

A

C

C

C

C

C

D

I I I I I I

No. of pairs

1

1

2

2

2

2

4

Total length in p

3.2

2.8

2.6

2.4

2.2

2.0

1.6


0.6 1.0

1.2

1.2

1.0

1.0

0.8

0.6

FX

18.75 31.25

42.86

46.15

41.67

45.45

40.00

37.50


-

Nearly median

Nearly submedian

Remarks


-

-

- - - -

Fig. 14c Fig. 15c Fig. 16c

Fig. 18c

Fig. 14d Jt a8 88 88 81 w #I #I,

Fig. 16d 81 41 11 11 11 1) I! .!I 41 I J II 8 8 18, - Fig. 17d 41 81 11 I J I J JJ 81,

A C D

Fig. 18, @ 11 11 l t 11 - 11 jd dd If, 41 81 - 81 11, K c. D

Figs. 14c, 14d - Karyotype and idiogram of Acrocephalus capitatus Benth. (2n=18) x 2400 approx.; 15c, 15d - Karyotype and idiogram of Orthosiphon glabratus Benth. ( 2 ~ 2 6 ) x 2400 approx., 16c, 16d - Karyotype and idiogram of 2. glabratus var. parviflorus Benth. (2n=26) x 2400 approx., 17c, 17d - karyotype and idiograrn of 0.grandiflorus Bold cv.lilac (2n=28) x 2400 approx.; 18c, 18d - Karyotype and - - idiogram of - 0. srandiflorus Bold cv. white (2n=28) x 2400 approx.

Orthosiphon pallidus Royle (2n=4x=28=A4C8D16)










TF value (Z)

n = 14 (Fig. 19j)

2n = 28 (Figs. 19b, 19c)

2

2.6 p to 1.2 p

1.73 P.I

48.4 P.I

12.1 P.I

36.84

25.33

39.68

Table 19 -


Chromo- some type

A

A

C

D

D

No. of pairs

1

1

4

4

4

I I I I I

Total length in P.I

2.6

2.4

2.0

1.6

1.2

Short arm length in pl

0.6 0.8

0.6 0.8

0.8

0.6

0.4

F%

23.08 30.77

25.00 33.33

40.00

37.50

33.33


-

-

Nearly median

Nearly submedian

Remarks


-

- -

Orthosiphon thymiflorus Roth. (2n=4x=24=A4C20)










TF value (%)

n = 12 (Fig. 20j)

2n = 24 (Figs. 20b, 20c)

2

3.6~ to 2.0~

2.53~

60.8~

15.2~

28.57

15.59

45.07

Table 20 --

Detailed taryotype analysis (Fig. 20d)

Remarks


-

-

-

-

-

-

Chromo- some type

A

A

C

C

C

C

C

C

F%

22.22 27.78

21.43 28.57

42.86

38.46

33.33

41.67

36.36

40.00


-

-

Nearly median

Nearly submedian

Nearly median

Nearly submedian

Nearly median

No. of pairs

1

1

1

2

3

1

2

1

Total length in p

3.6

2.3

2.8

2.6

2.4

2.4

2.2

2.0


0.8 1 .'O

0.6 0.8

1.2

1.0

0.8

1.0

0.8

0.8

Plectranthus numrmularius Briq. (2n=4x=28=A4D24)










TF value (%)

n = 14 (Fig. 21j)

2x1 = 28 (Figs. 21b, 21~)

2

1.8 p to 1.0 ,u

1.34 p

37.6 ,u

9.4 p

28.57

19.89

40.57

Table 3


Remarks


-

-

-

-

Chromo- some type

A

D

D

D

D

Total length in p

1.8

1.6

1.4

1.2

1.0

No. of pairs

2

2

3

4

3


0.4 0.6

0.6

0.6

0.4

0.4

F%

22.22 33.33

37.50

42.86

33.33

40.00


-

Nearly submedian

Nearly median

Nearly submedian

Nearly median

--

F i g . 17b F i g . 1 7 j - - -

Fig. 18b

-.

Fig. 18j Fig. 19b Fig. 19j

- I F i g . 20b

- -- - - Fig. 2 0 j Fig. 21b

Scale : - g*

Figs. 17b, 1 7 j - Orthosiphon grandiflorus Bold. cv. lilac: 17b - mitotic metaphase (2n=28), 17j - meiotic metaphase 1 (11~14); 18b, 18j - - 0. grandiflorus Bold. cv. white: 18b - mitotic rnetaphase (2n=28), 18j - -- diakinesis (n=14); 19b, 1 9 j - 0, pallidus Royle: 19b - mitotic metaphase (2n=28), 1 9 j - meiotic metaphaso-I (n=14); 20b, 2 0 j - 0. thymiflorus Roth.: 20b-, mitotic metaphase (2n=24), 20j - meiotic anaphase I (n=12); 21b - Plectranthus nummularius B r i q . - mitotic metaphase (2n=28).

Plectranthus wightii Benth.(2n=4~=24=A4C4DI6)










TF value (%)

n = 12 (Fig. 22j)

2n = 24 (Figs. 22b, 22c)

2

2.2 p to 1.4 p

1.78 p

42.8 p

10.7 p

22.22

12.53

42.56


Remarks


-

-

-

-

I

Chromo- some type

A

A

C

D

D

D


0.4 0.8

0.4 0.6

0.8

0.8

0.6

0.6

I

No. of pairs

1

1

2

3

4

1

I

F%

18.18 36.36

20.00 30.00

40.00

44.44

37.50

42.86

I

Total length in p

2.2

2.0

2.0

1.8

1.6

1.4

I


-

-

Nearly median

Nearly submedian

Nearly median

I

Coleus arematicus Benth. (2n=4x=32=A4C22D6)










TF value (%I

: 2n = 32 (Figs. 23b, 23c)

: 2n = 16, 24, 30, 34 and 48 (Figs. 23e, 23f ,23g, 23h,23i)

Table 23 -- Detailed karyotype analysis (Fig. 233)

Remarks


- -

- -

-


-

-

Nearly median

Nearly submedian

F%

20.00 33.33

23.08 30.77

42.86

41.67

45.45

44.44

37.50

Chromo- some type

A

A

C

C

C

D

D

Total length in p

3.0

2.6

2.8

2.4

2.2

1.8

1.6

No. of pairs

1

1

2

3

6

1

2


0.6 1.0

0.6 0.8

1.2

1.0

1.0

0.8

0.6

F i g s . 19c

Fig . 21c

Fig . 20c

F ig . 22c

F ig . 22d 88 8Cf l 8 8 18 8 l 11 o 81 8 1 18, A C D

F i g s . 19c, 19d Karyotype and id iogram of Orthosiphon p a l l i d u s Royle (211~28) x 2400 approx. ; 20c, 20d - Karyotype and idiograrn of 0 . t h y o i f l o r u s Roth . ( 2 ~ 2 4 ) x 2400 approx . ; 21c, 21d - Karyotype and idiogram,f P l e c t r a n t h u s - nummularius Br iq . (2n=28) x 2400 approx . , 22c 22d - Karyotype and idiograrn of - P. w 9 h t i i Benth. (2n=24; x 2400 approx .

Coleus aromaticus Benth. var. variegata (2n=4x=32=A4C16D12)










TF value ( % I

2n = 32 (Figs 24b, 24~)

2u = 16, 24, 30 and 44 (Figs.Z4e,24f,24g,24h)

Table 24 -- Detailed karyotype analysis (Pig.24d)

Remarks


-

-

-

-

Chromo- some type

A

A

C

C

C

D

D / 3 1 . 6 0.6 I 3 7 . 5 0 1 ~ .. ~- -- __I_. __ I -

No.of pairs

-

1

1

1

3

4

3

Total length in p

2.8

2.4

2.4

2.2

2.0

1.8


0.6 1.0

0.6 0.8

0.8

0.8

0.8

0.6

F%

21.43 35.71

25.00 33.33

33.33

36.36

40.00

33.33


-

-

Nearly submedian

Nearly median

Nearly submedian

Fig. 23c

Fig. 23h

Fig. 23e

Fig. 23i

Fig. 23f

Fig. 24c

Figs. 23c, 23d. 23e, 23f. 23h, 23i - Coleus aromaticus Benth.: 23c, 23d - Karyotype and idiogram (2n=32) x 2400 approx.;23e, 23f, 23h, 23i - somatic variants with 2n=16, 2n=24, 2 ~ 3 4 and 2n=48 respectively; 24c, 24d - karyotype and idiogram of - C. aromaticus Benth. var. variegata (2n=32) x 2400 approx.

Coleus bl-i Benth. var. verschaffeltii (2n=4x=48=A4C8D36)










TF value (%)

2n = 48 (Figs. 25b, 2%)

2n=46 (Fig. 25e)

2

2.6 p to 1.2 p

1.76 p

84.4 I.I

21.1 p

36.84

17.65

41.44

Table 25 -- Detailed karyotype analysis (Fig. 2 M )

Remarks


- -

-

-

-

-

-


-

-

Nearly median

Nearly submedian

Nearlymedian

Nearly submedian

Chromo- some type

A

A

C

D

D

D

D

D

D


0.6 0.8

0.6 0.8

0.8

0.8

0.6

0.6

0.6

0.4

0'. 4

F%

23.08 30.77

25.00 33.33

40.00

44.44

33.33

37.50

42.86

28.57

33.33

No. of pairs

1

1

4

7

1

5

3

1

1

Total length in p

2.6

2.4

2.0

1.8

1.8

1.6

1.4

1.4

1.2

Fig. 21j Fig. 22b F i g . 22j

- - - -- Fig. 23b Fig. 23g Fig. 2 4 7 --

P

Fig. 24f --

F i g . 24h Fig. 25b

Fig. 21j - Plectranthus nummularius Briq. - meiotic metaphase I (abnormal with 13 bivalents and 2 univalents); 22b, 22j - P. wightii Benth.: 22b - mitotic metaphase (2n=24), 22 j - meiotic anaphase (n=i2); 23b, 23g - Coleus aromaticus Benth. : 23b -mitotic metaphase (2n=32), 23g - somatic variant (2n=30) ; 24b , 24f, 24h - C. aromaticus Benth. var. variegata : 24b - mitotic metaphase (2n=32),, 24r - somatic variant (2n~24); 24h - somatic variant (2n=44); 25b - - C. blumei Benth. var. verschaffeltii - mitotic metaphase (2nZ48).

Coleus forskoblii Briq. (2n=4x=28=A4C22D2)










TF value (%)

2n = 28 (Figs. 26b, 26c)

2n = 26 and 30 (Figs. 26e, 26f)

2

3.4 p to 1.6 u

2.36 ,u

66 P

16.5 ,u

36

17.89

42.11

Table 26 --


Chromo- some type

A

A

C

C

C

C

No.of pairs

1

1

1

5

2

3

D I 1

/ 1.6 / 0.6 / 37.50 / Nearly submedian -

Remarks


-

-

-

-

Total length in p

3.4

2.8

2.8

2.4

2.2

2.0

Shortarm length in u

0.6 1.0

0.6 0.8

1.0

1.0

1.0

0.8

F%

17.65 29.41

21.43 28.57

35.71

41.67

45.45

40.00


-

-

Nearly submedian

Nearly median

Fig. 24c

Fig. 25e

Fig. 24g

- 3 \I Fig. 26c

Fig. 25c

Fig. 26e

~ --

Figs. 24e,24g - Somatic variants of Coleus aromaticus Benth. var. variegata with 2n=16 and 2n=30 respectively; 25c,25d,25e - - C -- blumei Benth. var. verschaffeltii : 25c, 25d - Karyotype and idiogram ( 2 ~ 4 8 ) x 2400 approx., 25e - somatic variant (2~46); 26c, 26d, 26e - C. forskohlii Briq.: 26c, 26d - Karyotype and idiogram ( 2 ~ 2 8 ) x 2400 approx., 26e1 somatic variant (2n=26)

Coleus lacidatus Benth. (2n=4x=48=A4D44)






Total chromatin length .




TF value (%)

2n = 48 (Figs. 27b, 27~)

2n = 42 (Pig. 27e)

2

2.0 p to 1.2 p

1.49 p


Chromo- some type

A

A

D

D

D

D

No. of pairs

1

1

2

5

12

3 I

Total length in p

2.0

1.8

1.8

1.6

1.4

1.2 I


-

-

Nearly median

Nearly submedian

Nearly median

Nearly submedian I

Remarks


-

-

-

- I


0.4 0.6

0.4 0.6

0.8

0.6

0.6

0.4 I

F%

20.00 30.00

22.22 33.33

44.44

37.50

42.86

33.33 I

5 5

Coleus parviflorus Benth. (Zn=6x=72=AbD66)










TF value (%)

2n = 72 (Figs. 28b, 28c)

2n = 24,32,36,48 and 68 (Figs. 28e,28f ,28g,28h,281)

Table 28 --

Detailed karyotype analysis (Fig. 28d)

Chromo- some type

A

A

D

D

D

D

No.of pairs

2

1

1

6

22

4

Total length in p

2.0

1.8

1.8

1.6

1.4

1.2


0.6 0.6

0.4 0.6

0.6

0.6

0.6

0.4

F%

30.00 30.00

22.22 33.33

33.33

37.50

42.86

33.33


-

-

Nearly submedian

Nearly median

Nearly submedian

Remarks


-

-

- -

Fig. 26f Fig. 27c Fig. 27e

Fig. 28c Fig. 28e Fig. 28g

Fig. 26f - Somatic variant of Coleus forskohlii Briq. with 2n=30; 27c,27d,27e - C.laciniatus Benth.: 27c, 27d - Karyotype and idiogram (2n=48) x 2400 approx., 27e - - somatic variant with 2n=42; 28c, 28d, 28e, 28g - Coleus parviflorus Benth.: 28c 2nd - Karyotype and idiogran (2n=72) x 2400 approx. 2 8 e 288 - somatic variants with 2n=24 and 2n=36 respectively

Coleus rehneltianus Berger (Zn=4~=48=A4CZD42)










TF value (%)

2n = 48 (Figs. 29b, 29c)

2n=50 (Fig. 29e)

2

2.2 p to 1.2 p

1.6 p

76.8 p

19.2 p

29.41

15.73

40.33

Table 29 --


Chromo- some type

A

A

C

D

D

D

D

D

No. of pairs

1

1

1

3

2

6

8

2

Total length in p

2.2

2.0

2.0

1.8

1.8

1.6

1.4

1.2


0 .'4 0.8

0.4 0.6

- 0.8

0.6

0.8

0.6

0.6

0.4

F%

18.18 36.36

20.00 30.00

40.00

33.33

44.44

37.50

42.86

33.33


-

-

Nearly median

Nearly submedian

Nerarly median

Nearly submedian

Nearly median

Nearly submedian

Remarks


-

-

-

-

-

-

Coleus zeylanicus (Benth.) Cramer (2n=4x=28=A4C8D16)










TF value (%)

2n = 28 (Figs. 30b, 30c)

2n = 14, 30, 32 and 56 (Figs. 30e, 30f, 30g, 30h)

Table 30 --


Chromo- some type

A

A

C

C

D

D

D

No.of pairs

1

1

2

2

4

2

2

I

Total length in p

2.8

2.2

2.2

2.0

1.8

1.6

1.4

I


-

-

Nearly submedian

Nearly median

Nearly submedian

Nearly median

I

Remarks

Chromosome pair [with secondary constriction

-

-

-

-


0.6 0.8

0.4 0.8

0.8

0.8

0.8

0.6

0.6

I

F%

21.43 28.57

18.18 36.36

36.36

40.00

44.44

37.50

42.86

I

Fig. 28i @ Fig. 29c Fig. 29e

Fig. 30c Fig. 30e Fig. 30f

Fig 281 Sonatic variant of - Coleus E--. arviflorus Benth. (2~68); 29c,29d,29e C rehneltianus Berzer 29c,29d - Karyotype and idiogram (2n=48) x 2400 approx., 29e - somatic variant (2n-50); 30c,30d:30e,30f - C. zeylanicus .-

(Benth., Cramer 30c, 30d - Karyotype and idiogram (2n-28, x 2400 appro~. , 30e 30f somatic variants with 2n=30 and 2n=32 respectively.

dnisochilus carnosus ( L . f . ) Wall. (2n=Zx=34=A4C14D16)









TF value (%)

: 2n = 34 (Figs. 31b, 31c)

: 2

: 3.4,u to 1.2p

: 2.1@

: 73.61~

: 36.41

: 47.83

: 30.00

: 42.24


Chromo- some type

A

A

C

C

C

D

D

D

D

F%

17.65 29.41

18.75 31.25

42.86

41.67

40.00

44.44

37.50

42.86

33.33

No. of pairs

1

1

3

3

1

3

2

2

1


-

-

Nearly median

Nearly submedian

Nearly median

Nearly submedian

Total length in p

3.4

3.2

2.8

2.4

2.0

1.8

1.6

1.4

1.2

Remarks


-

-

- -

-

- -

Short arm length in j~

0.6 1.0

0.6 1.0

1.2

1.0

0.8

0.8

0.6

0.6

0.4

A n i s ~ u s eriocepbalus Benth. (2n=2x=32=A4C12D16)







Disparity index (Dl)


TF value (%)

2n = 32 (Figs. 32b, 32c)

2

3 . 2 p to 1.2 p

1 .91 p

61.2 u

30.6 p

45.45

28.91

41.45

Table 32 --


Chromo- some type

A

A

C

C

C

D

D

D

No. of pairs

1

1

2

1

3

3

4

1

Total length in p

3.2

2.8

2.4

2 .2

2 .0

1.6

1.4

1.2

Short arm length in &I

0.8 , 1 . 0

0 . 6 0 .8

1 . 0

0 .8

0 .8

0 . 6

0.6

0 .4

F%

25.00 31.25

21.43 28.57

41.67

36.36

40.00

37.50

42.86

33.33


-

-

Nearly median

Nearly submedian

Nearly median

Nearly submedian

Nearly median

Nearly submedian

Remarks


- -

-

- -

--- Fig. 26b

. * ', 'v'

Fig. 27b

- ~ - -~

Fig. 28f Fig. 28h Fig. 29b

-- -. --

~ .-

Fig. 30b

T( f 9 / it' ,R/ 3 4

Fig. 31b -

Fig. 32b

Scale: - SP

Fig. 26b - Coleus forskohlii Briq. - mitotic metaphasc (211'281, 27b-C. lachiatus Benth. - mitotic metaphase (2n=48); 28b, 28f ; X h - C. parviflor& Benth.: 28b - mitotic metaphase (2nZ72), 28f - somatic variant Dn=32), 28h - somatic variant (2n=48); 29b - C. rehnelcianus Serger - mitotic metaphase (2n=48); 30b - C. zeylanicus (~eni-h.) Cramer - sitotic metaphase (2n=28); 31b - Anisochilus carnosus (L . f . ) Wall. - mitotic mctaphase (2n=34); 32b - - A. - eriocephalus Benth. - mitotic metaphase (?n=32).

Eyptis capitata Jacq. (2n=4x=32=A4D28)



Range of chromosome length : 1.81-1 to 1.0~

Average chromosome length : 1.35~

Total chromatin length : 43.q~

Chromatin length of basic complement : 10.8~



TF value (%) : 39.68

Table 33 --

Detailed taryotype analysis (Pig. 33d)

Chromo- some type

A

D

D

D

D

D

No. of pairs

2

1

1

3

7

2

Total length in p

1.8

1.8

1.6

1.4

1.2

1.0


0.4 0.6

0.8

0.6

0.6

0.4

0.4

F%

22.22 33.33

44.44

37.50

42.86

33.33

40.00


-

Nearly median

Nearly submedian

Nearly median

Nearly submedian

Nearly median

Remarks


-

-

- -

-

Fig. 30g

Fig. 32c

& 4 - \ #

Fig. 30h

Fig. 31c

Fig. 33c

33d &jp,Jt I t I# rr #I rt r ~ ~ t #I JI 18 J# B, ,b --

Figs 30g, 30h - - Coleus . zeylanicus (Benth.) Cramer: 30g - somatic variant (2nz32), 30h - somatic variant (2n=56); 31c, 31d - karyotype and idiogram of Anisochilus carnosus ( L . f . ) Wall. ( 2 ~ 3 4 ) x 2400 approx.;32c, 32d - Karyotype and idiogram of A. eriocephalus Benth. (2n=32) x 2400 approx.; 33c, 33d - Hyptis capitata Jacq. - (2n=32) x 2400 approx.


Chromosome pair with secondary constriction , :







TF value (%)

2n = 28 (Figs. 34b, 34c)

2

1.6 p to 1 . 0 p

1 . 3 p

36 .4 p

9.1 p

23.08

12.67

39.04

Table 31r -- Detailed karyotype analysis (Fig. 34d)

Chromo- some type

A

A

D

D

D

D

I I I I I

No. of pairs

1

1

1

3

7

1

Total length in p

1.6

1.4

1.6

1.4

1.2

1.0

Short arm 'length in p

0.4 0 .4

0 .4 0.4

0.6

0.6

0 .4

0 .4


-

-

Nearly submedian

Nearly median

Nearly submedian

Nearly median

F%

25.00 25.00

28.57 28.57

37.50

42.86

33.33

40.00

Remarks


-

- -

-

Pogostemon benghalensis (Burm.f.) Kuntze (2n=4x=64=A4D60)










TF value ( X )

2n = 64 (Figs. 35b, 35c)

32 and 66 (Figs. 35e, 35f)

2

1.8 p to 1.0 p

1.24 cl

79.6 pl

19.9 p

28.57

14.94

37.77

Table 35 --

Detailed karyotype analysis (Pig. 3%)

Chromo- some type

A

A

D

D

D

D

I l l I I

No. of pairs

1

1

2

5

17

6

Total length in p

1.8

1.4

1.6

1.4

1.2

1.0

Short arm length in w

0.4 0.6

0.4 0.4

0.6

0.6

0.4

0.4


-

-

Nearly submedian

Nearly median

Nearly submedian

Nearly median

F%

22.22 33.33

28.57 28.57

37.50

42.86

33.33

40.00

Remarks


-

-

-

-

Pogostemon heyneanus Benth. (2n=4x=64=A4D60)










TF value (%)

2n = 64 (Figs 36b, 36c)

2n=68 (Fig. 36e)

2


Chromo- some type

A

A

D

D

D

D

I l l I I I

No.of pairs

1

1

7

7

12

4

Total length in p

1.8

1.6

1.6

1.4

1.2

1.0


0.4 0.6

0.4 0.4

0.6

0.6

0.4

0.4

F%

22.22 33.33

25.00 25.00

37.50

42.86

33.33

40.00


-

-

-

Nearly submedian

Nearly median

Nearly submedian

Nearly median

Remarks


- -

- -

Fig. 34c

B -

Fig. 35f

Fig. 35c

Fig. 35e

Fig. 36e

Fig. 34d88 48 81 11 11 11 I1 I 1 81 I t I 1 18 II I! -\

Fig.34c, 34d - Karyotype and idiogram of Hyptis suaveolens ( L . ) Poit. (2n=28) x 2400 approx.; 35c, 35d, 35e, 35f - Pogostemon benghalensis (Burm.f.) Kuntze : 35c, 35d - Karyotype and idiogram (2n=64) x 2400 approx., 35e - somatic variant (2n=32), 35f - somatic variant (2n=66j; 36c, 36d, 36e - P. heyneanus Benth.: 36c, 36d - Karyotype and idiogram i2n=64) x 2400 apcrox., 36e - somatic varlant (211~68)

Pogosteron purpurascens Dalz . (2n=Zx=32=A4CZD26)


Somatic variation numbers : 2n = 30 (Fig. 37e)








TF value (%)

Table 37 --

Detailed karyotype analysis (~ig. 37d)

Chromo- some type

A

A

C

D

D

Total length in p

2.2

2.0

2.0

1.6

1.4

No. of pairs

1

1

1

5

7

1

I I I


0.4 0.6

0.4 0.6

0.8

0.6

0.6

33.33

O.* I I I

F%

18.18 27.27

20.00 30.00

40.00

37.50

42.86

Nearly submedian -


-

-

Nearly median

Nearly submedian

Nearly median

Remarks


-

-

-

Pogostenon vestitus Benth . (2"=2~=32=A4D28)










TF value (%)

2n = 32 (Figs. 38b, 38c)

34 (Fig. 38e)

2

2.0 p to 1.0 p

1.39 p

44.4 p

22.2 p

33.33

20.01

40.53

Table 38 --


Remarks


-

- -

D I 2 ( 1.0 / 0.4, /4O.OO/Nearlymedlan 1 -


0.4 0.6

0.4 0.6

0.8

0.6

0.6

0.4

Total length in p

2.0

1.8

1.8

1.6

1.4

1.2

Chromo- some type

A

A

D

D

D

D

No. of pairs

1

1

1

1

5

5

F%

20.00 30.00

2 2 . 2 2 33.33

44.44

37.50

42.86

33.33


-

-

Nearly median

Nearly submedian

Nearlymedian

Nearly submedian

6 6

Zusteralis quadrifolia (Benth3 Panigrahi (2n=2x=30=A4D26)









TF value ( % I

2n = 30 (Figs. 39b, 3%)

2

1.6 ,u to 1.0 p

1.16 p

34.8 p

17.4 p

23.8

16.89

38.67

Table 39 - -


Chromo- some type

A

D

D

No.of pairs

2

6

7

Total length in p

1.6

1.2

1.0


0.4 0.4

0.4

0.4

F%

25.00 25.00

33.33

40.00


-

Nearly submedian

Nearly median

Remarks


-

-

Fig. 38c

Fig. 37c Fig. 37e

Fig. 38e Fig. 39c

Fig. 39d $8 ,tt 11 81 18 81 18 11 # t 11 11 11 11 11, /

- - A D

Figs. 37c 37d, 37e Pogostemon Krpurnscens Dalz.: 37c, 37d - karyotype and -- - -- idiogram (2n=32) x 2400 approx., 37e - somatic variant (2~30); 38c, 38d. 3& - P. vestitus Benth.: 38c, 38d - Karyotype and idiogram (2n=32) x 2400 - approx., 38e - somatic variant (2n=34); 39c, 39d - Karyotype and idiogram of Eusteralis quadrifolia (Benth.) Panigrahi !2n=30) x 2400 approx.










TF value (%)

2n = 96 (Figs. 40b, 40c)

2n=72 (Fig. 40e)

2

1.8 p to 1.0 p

1.17 p

112.4 p

14.05 p

28.57

17.79

37.98


Total lengtt in P

Chromo- some type

No. of pairs

Short arm length in ~ . r



F% Remarks

Nearly submedian

Nearly median

Nearly submedian

-

- -

1 Nearly median - -

- -- -- -

Fig. 33b

-

Fig. 36b

-

Fig. 34b

-- --

Fig. 37b

Fig. 35b

Fig. 38b

Fig. 39b Fig. 40b Fig. 40e

Scale : - G P

Fig. 33b - Hyptis capitata Jacq. - mitotic metaphase (2n=32); 34b - H. suaveolens ( L . ) Poit. - mitotic metaphase (2n = 28); 35b - ~ogostemon benghalensis (Burm.f.1 Kuntze - mitotic metaphase (211~64); 36b - P. heyneanus Benth. - mitotic metaphase (2n=64); 37b - - P. purpurascens ~alz: - mitotic netaphase (2n=32); 38b - P. vestitus Benth. - mitotic metaphase (2~32); 39b - Eusteralis quadrifolia T~enth. ) Panigrahi - mitotic metaphase (2ns30) ; 40b, 40e - Mentha arvensis L.: 40b - mitotic metaphase (2n=96); 40e - somatic variant (2n=72);

Mentha rotuudifolia (L.) Huds.(211=2~=24=A4D20)










TF value (%)

2n = 24 (Figs. 41b, 41c)

2n=36 and 48 (Figs. 41e. 41f)

2


Chromo- some type

A

A

D

D

No. of pairs

1

1

2

8

Total length in p

1.6

1.4

1.2

1.0


0.4 0.4

0.4 0.4

0.4

0.4

F%

25.00 25.00

28.57 28.57

33 .33

40.00


-

-

Nearly submedian

Nearly median

Remarks


- -

llentha spicata(L.) Huds. (2n=4x=48=A4D44)


Somatic variation numbers : 2n=24, 64 and 72 (Figs. 42e, 42f, 42g)








TF value (%) : 40.53

Table 42 --


Chromo- some type

A

A

D

D

No. of pairs

1

1

3

19

Total length in p

1.8

1.4

1.2

1.0


0.4 0.6

0.4 0.4

0.4

0.4

F%

22.22 33.33

28.57 28.57

33.33

40.00


-

-

Nearly submedian

Nearly median

Remarks


-

-

Fig. 40c Fig. 41c

Fig. 41f Fig. 42c

Fig. 41e

Fig. 42e

Fig. 40c,40d - Karyotype and idiogram of Mentha arvensis L.(2n=96) x 2400 approx.; 41c 41d,41e 41f - - M. rotundifolia (L.) Huds.: 41c,41d - Karyotype and idiogram (2n=24> x 2400 approx , 41e, 41f - somatic variants with 2n=36 and 2n=48 respectively.; 42c,42d,42e - M. scats (L. ) Huds.: 42c, 42d - Karyotype and - -. idiogran ( 2 ~ 4 8 ) x 2400 approx., 42e somatic variant with 2n=24

S a l a coccinea Juss. (2n=Zx=22=A2C8D12)





Range of Chromosome length






TF value (%)

n = 11 (Fig. 433)

2n = 22 (Figs. 43b, 43c)

2n = 20 and 24 (Figs. 43e, 43f)

1

2.6 p to 1.6 p

1.95 p

42.8 p

21.4 p

23.81

18.08

37.92

Table 43 --



-

Nearly submedian

Nearly median

Nearly submedian

Chromo- some type

A

C

C

D

D

Remarks


-

- -

-

Total length in fl

2.6

2.4

2.0

1.8

1.6

No. of pairs

1

2

2

2

4


0.6 0.8

0.8

0.8

0.6

0.6

F%

23.07 30.77

33.33

40.00

33.33

37.50

Salvia leucantha Cav. (2n=2x=22=A2C18D2)







Total chromatin length .




TF value (%)

n = 11 (Fig. 44j)

2n = 22 (Figs. 44b, 44~)

2n = 23 and 24 (Figs. 44e, 44f)

1

2.8 p to 1.6 p

2.47 p

54.4 !J

27.2 ,u

27.27

14.35

40.95

Table 3 - Detailed Karyotype analysis (Pig. 44d)

-- - . . ,

Fig. 41b Fig. 42b Fig. 42g

- - -- --

Fig. 43b Fig. 43e Fig. 43j

Fig. 44b Fig. 44e Fig. 44 j

Scale: U BP

Fig. 41b - Mentha rotundifolia ( L . ) Huds, - mitotic metaphase, (2n=24); 42b, 42g - M. spicata (L.) Huds.: 42b - mitotic metaphase (2n=48), 42g - somatic variant (2n=72); 43b, 43e, 43j - Salvia coccinea Juss.: 43b - mitotic metaphase (2n=22), 43e - somatic variant (2n=20), 43 j - diakinesis (n=ll); 44b, 44e, 44j - S. leucantha Cav.: 44b - mitotic metaphase (2n=22), 44e - somatic varia.nt (%=23), 441 - meiotic anaphase I (n=ll)

Salvia plebeia R.Br. (2n=2~=16+0-IB=A2C4DlO)

Chromosome count on pollen mother cell : n = 8+0-1B (Fig. 453)

Normal somatic chromosome number : 2n = 16+0-1B (Figs. 45b, 45c)









TF value (%)

2n=14 & 18 (Figs. 45e, 45f)

1

2.6 ,u to 1.4 p

1.83 p

29.2 p


Chrono- some type

A

C

D

D

D

No. of pairs

1

2

1

3

1

Total length in p

2.6

2.0

1.8

1.6

1.4


0.6 0.8

0.8

0.6

0.6

0.6

F%

23.08 30.77

40.00

33.33

37.50

42.86


-

Nearly median

Nearly submedian

Nearly median

Remarks


-

-

-

-

Fig. 42f

Fig. 44c

Fig. 43c Fit.43f

Fig. 44f

Fig. 43d & II I I 11. .II ir JI 18 rt rr A C D

Fig. 44d

Fig. 45d

Fig. 45c

Fig. 42f - -- Mentha spicata ( L . ) Huds.-somatic variant (2n=64); 43c, 43d, 43f - Salvia coccinea Juss.: 43c, 43d - karyotype and idiogram i2n-22) x 2400 appxox., 43f - somatic variant (2n=24); 44c 44d, 44f - 5.leucantha Cav.: 44c, 44d - karyotype and idiogram (2n=22) x 2400 approx.; 44f - somatic variant (2n=24); 45c, 45d - karyotype and idiogram of plebeia R.Br. [2n=16+0-1 B :" B Chromosome)] x 2400 approx.

Salvia splendens ker - Gawl. (2n=4x=44=A4C2D38)











TF value (%)

n = 22 (Fig. 46j)

2n = 44 (Figs. 46b, 46c)

2n=22 and 48 (Figs. 46e, 46f)

2

2.2 p to 1.2 p

1.63 p

71.6 p

17.9 p

29.41

17.07

41.19

Table 46 --

Detailed karyotype analysis (Pig- 46d)

Chromo- some type

A

A

C

D

D

D

D 1 3 1 1.2 / 0.4 I33.33/Nearlysubmedian I -

No. of pairs

1

1

1

5

7

4

Total length in p

2.2

2.0

2.2

1.8

1.6

1.4


0.6 0.6

0.4 0.6

1.0

0.8

0.6

0.6

Remarks


-

-

-

-

F%

27.27 27.27

20.00 30.00

45.45

44.44

37.50

42.86


-

-

Nearly median

Nearly submedian

Nearly median

Anisomeles indica (LJ Kuntze (2n=Zx=34=A4D30)









TF value (%) : 39.19

Table 47 -- Detailed Karyotype analysis (Pig. 47d)

Chromo- some type

No. of pairs

1

1

2

7

6

Total length in p

1.6

1.4

1.4

1.2

1.0


0 .'4 0.4

0.4 0.4

0.6

0.4

0.4

.

Remarks


-

-

-

F%

25.00 25.00

28.57 28.57

42.86

33.33

40.00


-

-

Nearly median

Nearly submedian

Nearly median

Fig. 45e

Fig. 46e

Fig. 45f

Fig. 46f

Fig. 46c

Fig. 47c

Fig. 45e, 45f - Salvia plebeia R. Br.: 45e - somatic variant (2n=14), 45f - somatic variant (2n=18); 46c, 46d, 46e, 46f - S. splendens Ker - Gawl.: 46c, 46d - karyotype and idiogram (2n=44) x 2400 approx., 46e - somatic variant (2n~7.3; 46E - somatic variant (2~1~48); 4732, 47d - Anisomeles indica (L.) Kuntze.-karyotype and idiogram (in-34) x 2400 approx.









TF value ( X )

2n = 32 (Figs. 48b, 48c)

Table 48 --


Remarks


-

-

-

-

I I I I I I

F%

20.00 30.00

22.22 33 .33

44.44

42.86

33 .33

40.00


0 .4 0.6

0.4 0 .6

0.8

0.6

0 .4

0 .4

Chromo- some type

A

A

D

D

D

D


-

-

Nearly median

Nearly submedian

Nearly median

No. of pairs

1

1

2

4

4

4

Total length in p

2.0

1.8

1.8

1.4

1.2

1.0

Fig. 45b

- - - -

Fig . 463

- - - -

Fig. 45 j Fig. 46b

. - -

Fig. 47b F i g . 48b

- I--- - -- -- - -

Fig. 49b Fig. 50b Fig. 51b

F i g . 45b , 4 5 j - Salvia p l e b e i a R .Br . : 45b-mitotic metaphase [2n=16+0-1~ (* B chromosome) 1, 45j - meiotic anaphase I [n=8+0-16 (* B chromosome) 1; 46b, 4 6 j - S . s p l e n d e n s Ker. Gawl.: 46b - mitotic metaphase ( 2 ~ 4 4 1 , 46j - meiotic - metaphase (abnormal airi in^ of bivalents): 47b - Anisomeles indica (L.) Kuntze - - <

- . - mitotic metaphase (211~34); 48b. - A. malabarica R . B r . - mitotic metaphase (2n=32) ; 49b - Leucas aspera spreng. - mitotic metaphase ( 2 ~ 2 2 ) ; 50b - 4. -

cephalotes Spreng. - mitotic metaphase (2n = 2 2 ) ; 51b - - L. linifolia Spreng. - mitotic rnetaphase (2n=22)

Leucas aspera Spreng. (2n=Zx=22=A4D18)









TF value (%)

2n = 22 (Figs. 49b, 49c)

2

2.4 p to 1.0 p

1.44 p

31.6 p

15.8 p

41.18

26.36

37.48

Table 49 --

Detailed Karyotype analysis (Fig. 4%)

Chromo- some type

A

A

D

D

D

D

I I I I I I

No. of pairs

1

1

1

4

2

2

Total length in p

2.4

1.8

1.6

1.4

1.2

1.0

Short arm lengfh in p

0.6 0.8

0.4 0.6

0.6

0.4

0.4

0.4

F%

25.00 33.33

22.22 33.33

37.50

28.57

33.33

40.00


-

-

Nearly submedian

Nearly median

Remarks


-

-

-

-

Le- cephalotes Spreng. (2n=Zx=22=A4D18)









TF value (%)

2n = 22 (Figs. 50b, 50c)

2

2.4 p to 1.0 p

1.47 p

32.4 p

16.2 p

41.18

26.76

38.53

Table 50


I I I I I I


-

-

Nearly submedian

Nearly median

Chromo- some type

A

A

D

D

D

D

Remarks


-

- -

-


0.6 0.8

0.6 0.6

0.6

0.4

0.4

0.4

No. of pairs

1

1

2

2

4

1

F%

25.00 33.33

30.00 30.00

37.50

28.57

33.33

40.00

Total length in p

2.4

2.0

1.6

1.4

1.2

1 .O

Fig. 48d

Fig. 49d

Fig. 50d

Fig. 51d

Fig. 50c

Fig. 49c

Fig. 51c

Fig. 48c, 48d - Karyotype and idioeram of Anisomeles malabarica R.Br. ( 2 ~ 3 2 ) x 2400 approx.; 49c, 49d - Karyotype and idiogram of Leucas aspera Spreng. (211-22) x 2400 approx.; 50c, 50d - Karyotype and i d i o x o f L.cephalotes Spreng. (2n=22) x 2400 approx.; 51c, 51d - Karyotype and idiogram of L.linifolia Spreng. (Zn=22) x 2400 approx. .-

78

Leucas linifolia Spreng. (2n=2x=22=A4D18)









TF value (%)

2n = 22 (Figs. 51b, 51c)

2

1.6 ,u to 1.0 ,u

1.27 ,u

28.0 p

14.0 ,u

23.08

16.82

37.95

Table 51 --


Chromo- some type

A

D

D

D

No. of pairs

2

1

6

2

Total length in ,u

1.6

1.6

1.2

1.0

Short arm length in ,u

0.4 0.4

0.6

0.4

0.4

F%

25.00 25.00

37.50

33.33

40.00


-

Nearly submedian

Nearly median

Remarks


-

-

-

Leucas stricta Benth. (2n=Zx=22=A4D18)



Range of chromosome length : 2.0 p to 1 . 0 ~






TF value (%) : 37.09

Table 52 --


Chromo- some type

A

A

D

D

D

D

No.of pairs

1

1

1

2

4

2

I I I I

Total length in p

2.0

1.6

1.6

1.4

1.2

1.0


0.4 0.6

0.4 0.4

0.6

0.4

0.4

0.4

F%

20.00 30.00

25.00 25.00

37.50

28.57

33.33

40.00


-

-

Nearly submedian

Nearly median

Remarks


-

-

-

-

Leucas vestita Benth. (2n=2~=22=A2ClODlO)









TF value (XI

2n = 22 (Figs. 53b, 53c)

1

2.4 p to 1.4 p

1.95 p

42.8 p

21.4 p

26.32

14.56

43.64

Table 53 -- Detailed karyotype analysis (Fig. 5M)

Chromo- some type

A

C

C

D

D

D

NO. of pairs

1

3

2

3

1

1

Total length in p

2.4

2.2

2.0

1.8

1.6

1.4


0.4 0.8

1.0

0.8

0.8

0.6

0.6

F%

16.67 33.33

45.45

40.00

44.44

37.50

42.86


-

Nearly median

Nearly submedian

Nearly median

Remarks


-

-

-

-

-

Leonotis nepetifolia (L.) Ait.f. (2n=4x=28=A2C24D2)









TF value (%)

: 2n = 28 (Figs. 54b, 54c)

: 1

Table 54 --


Chromo- some type

A

C

C

C

C

C

Short arm length in IJ

0.8 1.2

1.2

1.2

1.0

1.0

0.8

No. of pairs

1

1

1

4

4

2

D I 1 1.6 I 0.6 I 37.50 Nearly submedian

Total length in IJ

3.6

3.2

2.8

2.6

2.4

2.0 ‘

F%

22.22 33.33

37.5

42.86

38.46

41.67

40.00

-


-

Nearly submedian

Nearly median

Remarks


-

- -

-

-

~- - Fig . 52b

I Fig. 53b

-- - -

F i g . 54b F ig . 55b

Scale: - % P

Fig. 52b - Leucas stricta Benth. - mitotic metaphase (2n - 22); 53b - E. v e s t i t a Benth. - mitotic metaphase (2n-22); 54b - Leonotis nepetifolia (L.) Ait.f. - mitotic metaphase (2n=28); 55b - Teucrium plectranthoidee Gamble - mitotic metaphase (2n = 32)

Teucrium plectranthoides Gamble (2n=4x=32=A4C2D26)









TF value (%)

2n = 32 (Figs. 55b, 55c)

2

2.2 p to 1.2 p

1.55 p

49.6 ,u

12.4 p

29.41

16.13

42.27

Table 55 --

Detailed Karyotype Analysis (Pig. 5 M )

Chromo- some type

A

A

C

D

D

No. of pairs

1

1

1

4

8

Total length in p

2.2

1.8

2.0

1.6

1.4

D , 1 , 1.2 , 0.4 33.33 Nearly submedian

F%

27.27

22.22 33.33

40.00

37.50

42.86


0.6 0.6

0 .4 0 .6

0 . 8

0.6

0.6

-


-

-

Nearly median

Nearly submedian

Nearly median

Remarks


-

-

-

Fig. 52c

Fig. 54c

Fig. 53c

Fig. 55c

Fig. 52d L-A

Fig. 53d , 4 48 4, 4 , I J I4 14 I,,

Fig. 54d

___JV

A C D

~ i g . 55d $1 $0, fi 11 81 t8 # f #8 #I 88 11 88 11 81 I t A C D

Fig. 52c, 52d - Karyotype and idiogram of Leucas stricta Benth. (211~22) x

2400 approx.; 53c, 53d - Karyotype and idiogram of L.vestita Benth. (2n=22) x 2400 npprox. ; 54c, 54d - karyotype and idiogram of -~eonotis nepetifolia ( L . ) Ait.f. (2n=28) x 2400 approx.; 55c, 55d - Teucrium plectranthoides Gamble (2n=32) x 2400 approx.

Table 56 -- sl l rary of the karymrphetrical analysis on the fifty four e b e r s of South Indian Lariaceae investigated

S1. Name of the taxa 2n n PL KF CPSC RCL ACL TCL CLBC DI VC TF% No. in p in f i in p in p

-- -

Ocimum adscendens Willd.

0. americanum L. - 0. basilicum L.var. - glabratum Benth.

0. basilicum L.var. pilosum Benth.

0. basilicum L.var. - purpurascens Benth.

0. basilicum L.var. - thyrsiflorum Benth.

0. gratissimum L. -

0. gratissimum L.var. - suavis Hook. f . 0. tenuiflorum L.f. - cv. green

0. tenuiflorum L.f. - cv. purple

0. tenuiflorum L.f. - cv. purple-green

S1. Name of the taxa 2n n PL KF CPSC RCL ACL TCL CLBC DI VC TF% No. in p in p in p in pl

12. 0. tenuiflorum L.f. var. hirsuta H0ok.f.

13. Acrocephalus capitatus Benth.

14. Orthosiphon glabratus Benth.

15. - 0. glabratus var. parviflorus Benth.

16. - 0. grandiflorus Bold cv. lilac -

17. - 0. grandiflorus Bold cv. white

18. 0- pallidus Royle

19. - 0. thymiflorus Roth.

20. Plectranthus nummularius Briq.

21. - P. wightii Benth.

22. Coleus aromaticus Benth.

23. - C. aromaticus Benth. var. variegata

S1. Name of the taxa 2n n PL KF CPSC RCL ACL TCL CLBC D I V C T F % No. in &I in p in p in p

2 4 . C. blumei Benth. -- var. verschaffeltii

25. - C. forskohlii Briq.

26. C. laciniatus Benth. - 27. - C. parviflorus Benth.

28. - C. rehneltianus Berger

2 9 . - C. zeylanicus(Benth.) Cramer

30. Anisochilus carnosus (L.f.1 Wall.

31. & eriocephalus Benth.

32. Hyptis capitata Jacq.

33. - H . suaveolens (L.) Poit.

3 4 . Pogostemon benghalensis (Burm.f.) Kuntze

35. - P. heyneanus Benth.

36. & purpurascens Dalz.

37. P, vestitus Benth.

S1. Name of the taxa 2n n PL KF CPSC RCL ACL TCL CLBC D I V C T F % No. in p in p in p in p

38. Eusteralis quadrifolia - (~enth.) Panigrahi 30 - 2x A4D26 2 1.6-1.0 1.16 34.8 17.4 23.08 16.89 38.67

39. Mentha arvensis L. 96 - 8x A4D92 2 1.8-1.0 1.17 112.4 14.05 28.57 17.79 37.98

40. M. rotundifolia (L. ) Huds. 24 - 2x A4D20 - 2 1.6-1.0 1.12 26.8 13.4 23.08 17 .03 41.15

41. M. spicata (L.) Huds. 48 - 4x A4D44 2 1.8-1.0 1.08 51.6 12.9 28.57 16.75 40.53

42. Salvia coccinea Juss. 22 11 2x A2C8D12 1 2.6-1.6 1 .95 42.8 21.4 23.81 18.08 37.92

43. S. leucantha Cav. - 22 11 2x A2C18D2 1 2.8-1.6 2.47 54.4 27.2 27.27 14.35 40.95

44. - S. plebeia R.Br . 16+ 8+ 2x A2C4D10 1 2.6-1.4 1.83 - 29.2 14 .6 30.00 19.27 40.32 0-1B 0-1B 00 u-

45. - S. splendens ker- awl. 44 22 4x A4C2D38 2 2.2-1.2 1 .63 71.6 17 .9 29.41 17.07 41.19

Anisomeles indica (L.) Kuntze

47. - A. malabarica R.Br. 32 - 2x A4D28 2 2.0-1.0 1.36 43.6 21.8 33.33 23.38 41.19

48. Leucas aspera Spreng. 22 - 2x A4D18 2 2.4-1.0 1.44 31.6 15.8 41.18 26.36 37.48 - 49. - L. cephalotes Spreng. 22 - 2x A4D18 2 2.4-1.0 1.47 32.4 16.2 41.18 26.76 38.53

50. 4, linifolia Spreng. 22 - 2x A4D18 2 1.6-1.0 1.27 28.0 14.0 23.08 16.82 37 .95

51. & stricta Benth. 22 - 2x A4D18 2 2.0-1.0 1.35 29.6 14.8 33.33 21.04 37.09

S1. Name of the taxa 2n n PL KF CPSC RCL ACL TCL CLBC D. I. V.C . T.F.% No. in p in in p in p

52. - L. vestita Benth. 22 - 2x A2ClODlO 1 2;4-1.4 1.95 42.8 21.4 26.32 14.56 43.64

53. Leonotis nepetifolia (L.) Ait.f. 28 . - 4x A2C24D2 1 3.6-1.6 2.51 70.4 17.6 38.46 18.98 40.99

54. Teucrium plectranthoides Gamble 32 - 4x A4C2D.26 2 2.2-1.2 1.55 49.6 12.4 29.41 16.13 42.27

Abbreviations used: PL - Ploidy level; K I : Karyotype formula; CPSC - Chromosome pair with secondary constriction; RCL - Range of chromosome length; ACL - Average chromosome length; TCL - Total chromatin length; CLBC - Chromatin length of basic complement; DI - Disparity index; VC - Variation coefficient; TF% - Total forma % (mean centromeric index)

Table 57

Previous and present chromosome counts on different members of the genera investigated in Jimiaceae. The arrangement upto the generic level is essentially based on the corrected version of Bentham's (1876) classification (Sanders and Cantino 1984) 1

Taxon

Tribe - Ocimeae

Sub tribe - Plectranthinae Genus - Ocimum L. 0 . adscendens Willd. -

0. americanum L. -

0 . basilicum L. - ----

Author/~

Sanjappa

Cherian & Kuriachan

Krishnappa & Basavaraj

Saggoo & Bir

Bir & Saggoo

Pushpangadan, et al.

Vij & Kashyap

Singh

Pushpangadan & Sobti

Singh & Sharma

Sobti & Pushpangadan

Singh

Vaarama

Morton

Count Dip- loid number (2n)

-

22

22

-

-

72

' 6 4

84

72

72, 84

72

72, 84

48

48

Year/s

1979

1981

1982

1982, 1983

1985

1975

1975

1980

1982

1982, 1983

1982

1985

1947a

1962

Previous

Hap- loid number (n)

32

-

-

11

11

-

-

-

-

-

-

-

-

-

Present Count

Dip- loid number (2n)

-----

-

-

-

-

-

22

-

-

-

-

-

-

-

72

-

-

Hap- loid number (n)

-

-

-

-

-

11

-

-

-

-

-

-

-

36

-

-

var . " glabratum Benth.

s. var. difforme

s . var. purpurascens var. minima

var. minimum

var. pilosum Benth.

var. purpurascens Benth.

var. thyrsiflora

Sz - Borsos

Mehra & GI11

San jappa



Saggoo & Bir

Singh & Sharma

Gill

Ma, et al.

Vembu

Bir & Saggoo

Singh


Singh

Vaarama


Vaarama

Cherian & Kuriachan

Singh



1982

1992

1962

1970

1971

1975

1976b

1979

1981

1981b

1982a

1982

1982

1982

1982

1983

1983

var. thyrsiflorum Benth.

0. campechianum Miller -

0. canum Sins --

-

48

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

52

54

24

22- 32, 34

24, 2 6

- -

24, 26

22

24, 26

2 4 , 26

-

24, 56

-

24

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

6, 11- 14, 17 42

-

-

32+ 0- 4B 32

40

-

40+ 0- 1 B

-

-

-

13

13, 40+ 0- 1 B

-


Harley & Heywood

Morton

Sharma

Pal


Vij & Kashyap

Sanjappa

Saggoo & Bir

Singh & Sharma

Bir & Saggoo




Saggoo & Bir

Singh & Sharma

0. carnosum Link et Otto -

0. gratissimum L. -

embu

ir & Saggoo

ingh

aggoo & Bir

,arley & Heywood

'ushpangadan, et al.

iobti & pushpangadan

jingh & Sharma

aosla & Sobti

Zhosla

rischler

Morton

Mitra & Datta

Pal

Bhattacharya

Singh

Singh & Sharma


Vembu

Khosla, et al.

Khosla & Sobti

var. suavis Hook. f.

0 . irvinei Mort. - 0 . kilimandscharicum Guerke -

0. nicranthum - 0 . selloi Benth. -

0 . suave Willd. --

Singh

Singh & Sharma

Khosla

Morton

Choudhury, et al.

Kumar, et al.

Bose & Choudhury

Mehra & Gill

' Pushpangadan, et al.


Saggoo & Bir

Singh & Sharma

I Gill

Bir & Saggoo

Singh

I Sobti & Pushpangadan

I Harley & Heywood

de Wet

I Morton

, Sobti & Pushpangadan

I Renard, et al. / Khosla & Sobti

1 Khosla

0. v i r i d e Willd. -

Genus - Acrocepbalus Benth.

A . c a p i t a t u s Benth. -

Genus - Orthosiphon Benth.

0. d i f f u s e s Benth. -

0. glabratum Benth. -

0. g l ab ra tus Benth. -

var. pa rv i f lo rus Benth.

0. g rand i f lo rus Bold -

cv. lilac

;ingh

'ushpangadan, e t a l .

jobti & Pushpangadan

jingh & Sharma

(hosla, e t a l .

(hosla & Sobti

(hosla

:herian & Kuriachan


Cherian & Kuriachan


Saggoo & B i r

B i r & Saggoo

Basavaraj & Krishnappa


cv. white

0 . incisus Chev. -

0 . incurvus Benth. -

0 . pallidus Royle -

0 . rubicundus (Don) Benth. -

0 . scapiger Benth. - 0. stamineus Benth. -

0 . suffrutescens (Thoun. ) - - Mort.

0 . thymiflorus Roth. -

0 . tomentosus Benth. -

var. tomentosa Hook. -

0 . tubiformis Good --

28

26

28

24

28

28

22

22

-

-

-

28

26

26

48

28

-

-

-

-

-

28

-

-

-

-

-

-

-

-

14

14

14

-

-

-

-

-

14

13

13

13

13

-

28

-

-

-

-

-

-

-

-

-

-

-

28

-

-

-

-

- -

24

-

-

-

-

-

14

-

-

-

-

-

-

-

-

-

-

-

14

-

-

-

-

- -

12

-

-

-

-

-

Morton

Kundu and Sharma

Morton

Chopde

Mehra & Gill

Bhatt

Vembu

Vembu & Sampathkumar

Saggoo & Bir

Gill

Bir & Saggoo

Morton


Kundu and Sharma

Cherian & Kuriachan

Morton

Rao & Mwasumbi

Saggoo & Bir

Bir & Saggoo

Saggoo & Bir

Bir & Saggoo

de Wet

1962

198&

1962

1965

1968b, 1972

1974, 1976

1980

1980

1983b

1984

1985

1962

1982

1988a

1984

1962

1981

1982

1985

1983

1985

1958b

0. viscosus Benth. -

Genus - Plectranthus L'Herit.

P. amicorum Blake -

P. argentatus Blake -

P. assurgens (Bak.) Mort. -

P. barbatus -

P. calycinus Benth. -

P. ciliatus Mey. et Benth. - -

P. coesta Buch.- Ham. -

var. macraei Hook.

P. dolichopodus Briq. -

P. dregei Codd -

P. ecklonii Benth. -

P. elegantulus Briq. -

P. esculentus Brown -

P. fruiticosus L'Herit. -

P. gerardianus Benth. -

var. graciliflorus Hook.

P. glandulosus Hook. f. -

P. grallatus Briq. -

P. grandidentatus Guerke -

P. halli Mort. -- P. harrissi Mort. - P. hirtus Benth. --

P. incanus Link - P. insignis Hook. f. - P. japonicus Koidz. -

P. kamerunensis Guerke -

P. laxiflorus Benth. - -

P. macraei Benth. -

P. maddenii Benth. -

P. mollis (Ait.) Spreng. --

P. nilghiricus Benth. -

P. nummularius Briq. -

P. oertendahlii Fries -

P. pachyphyllus Guerke et - Cooke

P. parviflorus Willd. -

P. peglerae Cooke -

P. petiolaris Mey. et Benth. -

P. punctatus L'Herit. -

P. purpuratus Harv. -

de Wet

Morton

Saggoo & Bir

Bir & Saggoo

Morton

Saggoo & Bir

Bir & Saggoo

Cherian & Kuriachan

Saggoo & Bir

Bir & Saggoo

de Wet

Morton

Morton

de Wet

Morton

Hendersson

de Wet

Morton

de Wet

Morton

P. rehmannii Guerke -

P. repens Wall. -

P. rugosus Wall. -

P. saccatus Benth. -

P. scrophularioides Wall. -

P. spectabilis Blake -

P. spicatus May.et Benth. -

P. stocksii Hook. f. -

P. striatus Benth. -

P. strigosus Benth. -

28

28

-

24

24

-

-

28

28

-

-

28

28

-

24

-

24

-

28

( 2 8

-

1 2

-

-

12

12

- -

12

17

-

-

12

-

12

-

12

-

-

P. succulentus Dyer et Bruce -

P. suluensis Cooke - --

-

-

-

28

28

28

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

- -

-

-

-

-

-

-

-

-

-

-

-

- -

-

-

deWet

Morton

Saggoo & Bir

Mehra & Gill

Vij & Kashyap

Bir & Saggoo

Gill

deWet

Morton

Saggoo & Bir

Hendersson

deWet

Morton

Saggoo & Bir

Mehra & gill

Bir & Saggoo


Gill

deWet

Morton

-

-

-

deWet

Morton

deWet

p. tenuicaulis(Hook. f.)Mort. -

P. ternifolius Don -

P. thunbergii Benth. -

P. tomentosus Benth. -

P. villosus Cooke -

P. wightii Benth. -

P. woodii Guerke -

Genus - Coleus Lour. C. amboinicus Lour. -

C. aromaticus Benth. -

3hra & Gill

L11

e Wet

or t on

e Wet

orton

e Wet

.orton

:a japathy

kishnappa & Basavaraj

laggoo & Bir

lir & Saggoo

le Wet

(orton

samachandran

Mehra & Gill

Krishnappa & Basavara:

Gill

Scheel

Reddy

Morton

Saggoo & Bir

var. variegata

C. barbatus Benth. -

C. blumei Benth. -

var. verschaffeltii

C. carnosus Hassk. -

C. comosus Hochst. -

C. forskohlii Briq. -

C. frederici Taylor - C. hybridus Hort. -

C. laciniatus Benth. -

C. lanuginosus Hochst. - C. malabaricus Benth. -

C. parviflorus Benth. -

C. pentheri Guerke -

de Wet

Morton

Scheel

Reddy

Morton

Borgnann

Morton

Cherian & Kuriachan

de Wet Morton

Cherian & Kuriachan

Bir & Saggoo


Saggoo & Bir

Cherian & Kuriachan

Saggoo & Bir

Cherian & Kuriachan

H. alata (L.f.) Shinners - - s.sp. rugosula (Briq.)

Harley

H. albida Kunth. -

H. argyrophylla Harley -

H. atrorubens Poit. -

H. brevipes Poit. - H. capitata Jacq. -

H. colombiana Epling -

H. conferta Pohl et Benth. -

var. angustifolia Benth.

H. cuneata Pohl et Benth. - --

H. cuniloides Epling - H. emoryi Torr. -

H. eriocephala Benth. -

H. fasciculata Benth. -

H. floribunda Briq. - H. fruticosa Salzm. et Benth. -

H. homalophylla Pohl et - Benth. I

H. inodora Sehrank -

H. irwinii Harley -

H. lanceolata Poir. -

Harley & Heywood

Morton

Harley & Heywood

Chuang, et al.

Reinhard

Harley & Heywood

Harley & Heywood

Baker & Parfitt

Harley & Heywood

Coleman

Harley & Heywood

Morton

H. lantanifolia Poir. -

H. lappulacea Mart. et Benth. - I H. leptostachys -

s.sp. caatingae Harley 1 H. lewocephala Mart. e t -

Benth.

H. lorentziana Hoffm. -

H. macrocephala Mart. et Gal. -

H. macrostachys Benth. - H. martiussi Benth. -

H. microphylla Pohl et Benth. - H. mutabilis (Rich.) Briq. -

H. nudicaulis Benth. -

H. oblongifolia Benth. -

H. pachycephala Epling - H. paludoza St. Hil.et Benth. - H. parkeri Benth. - H. pectinata (L. ) Poit. -

H. pinheiroi Harley -

H.~latanifolia Mart.et Benth. -

H. propinqua Epling - I H. racemulosa Mart. et Benth. - I H. rhomboidea Mart. et Gal. - I H. selloi Benth. - - I H. sidifolia (L'Herit. )Briq. -

H. silvinae Harley -

H..sinuata Pohl et Benth. - I H. spicigera Lam. -

H. stachydifolia Epling - H. suaveolens (L.) Poit. -

Diers

Harley & Heywood 1 1992

Harley & Heywood

Morton

Harley & Heywood

Miege

Morton

Harvey

Pal

Gill & Abubaker

Vij & Kashyap

Bir & Saggoo

Coleman


Saggoo & Bir

Vembu

1986

1992

1992

1986

1992

1992

1963

1967

1984

1985

1981

1984

1981

1982

1983

1984

-

-

-

-

-

-

-

-

-

- - -

-

-

-

- -

- -

H. subtilis Epling -

H. tomentosa Poit. -

H. umbrosa Saz: et Benth. - H. urticoides H.B.K. -

H. villicaulis Epling - H. villosa Pohl et Benth. -

Tribe - Uentheae Sub tribe - Pogostemoninae Genus - Pogosteu~n Desf.

(= Dysophylla Blume)

P. auricularia Blume -

P.benghalensis (Burm.f.) - kuntze

P. cablin Benth. -

P. sardneri Hook. -

P. heyneanus Benth. -

Vembu & Ayyangar

Harley & Heywood

Harley & Heywood

Sundberg & Dillon

Harley & Heywood

Chuang, et al.

Hsu

Cherian & Kuriachan

Bir & Saggoo

Lavania

Cherian & Kuriachan



Cherian & Kuriachan

----

-

14

16

16

8

20

-

-

-

- 6 -

-

-

-

16

16

32

16

16

-

24

28

32

-

16

40

63

90, 96

34

34

34

32

32

-

64

-

-

64

-

-

28

-

- - -

-

-

-

- -

-

64

-

-

- -

- -

64

P. rnollis -

P. paludosus Benth. -

P. paniculatus Benth. -

P. parviflorus Benth. -

P. plectranthoides Desf. -

"

P. pubescens Benth. -

P. purpurascens Dalz. -

P. speciosus Benth. -

P. verticillata Benth. -

P. vestitus Benth. -

P. wightii Benth. -

-

-

- -

32

32

64

64

-

- -

64

-

-

-

-

-

32

34

34

34

72

-

----

16

17

17

16

- -

- -

32

17

32

-

32

16

16

16

16

-

-

-

-

-

16

-

-

-

-

- - - -

-

- -

-

-

-

-

-

- - -

32

-

-

-

32

-

-

-

-

-

-

-

- -

-

- -

- -

-

-

-

- - -

-

-

-

-

Cherian & Kuriachan

Saggoo & Bir

Bir & Saggoo

Cherian & Kuriachan



Mehra & Gill

Gill

Gill

Mehra & Gill

Gill

Pal

Vij & Kashyap

Saggoo & Bir

Bir & Saggoo

Saggoo & Bir


Cherian & Kuriachan



Borgmann

Cherian & Kuriachan

1981

1982

1985

1981

1982

1983

1968 b

1971 a

1984

1968 b

1971 a, 1984

1971

1975, 1976 b

1982, 1983

1985

1981, 1982a

1981

1982

1984

1982

1983

1964

1981

Genus - Eusteralis Rafin. E. quadrifolia (Benth. ) -

Panigrahi

Sub tribe - Origaninae Genus - Mentha L.

M. alopecuroides Hull. -

M. aquatica L. -

-

-

64

36

-

36

96

96

96

96

96

96

96

96

96

96

96

60, 96

96

96

---- 17

17

-

-

18

-

- - -

-

-

- -

-

-

-

-

- -

-

-

-

-

30

-

- - - - -

-

-

- - -

-

-

-

-

-

-

-

-

-

-

- - - - -

- - - -

-

-

-

-

-

-

-



Cherian & Kuriachan

Morton

Gill

Schurhoff

Ruttle

Tischler

June11

Rohweder

Graham

Love & Love

Morton

Murray

Gadella & Kliphuis

Olsson

Ouweneel

Ikeda & Ono

Harley & Brighton

Van Loon & de Jong

1982

1983

1984

1956 a

1981 b

1929 b

1931 a

1934

1937

1937

1954

1956 b

1956 a

1958

1963,

1968 b

1967

1968

1969

1977

1978

M. arvensis L. -

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

- -

-

96 +lo5 -

96

96

96

72

72

72, 96

12, 60- 62, 7 2

72

64, 90, 92

72

90

72, 84, 96

72, 96

72

24, 72, 90

36, 92

96

72

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

- -

-

----

-

-

-

-

-

-

- -

- -

-

-

-

-

- -

- -

-

Tucker, et al.

Roy,etal.

Fernandes & Leitao

Singh

Schurhoff

Lietz

Ruttle

Junell

Rohweder

Nagao

Morton

Sharma & Bhattacharya

Ikeda & Seiroku

Ikeda & Udo

Olsson

Ouweneel

Taylor & Mulligan

Belyaeva & Kovineva

Harley & Brighton

1980

1983

1984

1984, 1985

1929 b

1930

1931 a

1937

1937

1941 a

1956 a

1959 a

1966

1967

1967

1968

1968

1972

1977

1980

1981 b

1984

1985

1985

1986

1990

1977

1982 a

1971 b, 1975 1986

1965

1985

1965

1972

1972

1929 b

1929

1980

Tucker, et al.

Gill

Fernandes & Leitao

Kundu & Sharma

Parfenov & Dimitrieva

Pogan, et al.

Tucker & Fair brothers

Harley & Brighton

Love & Love

Sobti

Tyagi

Sobti

Kundu & Sharma

Sobti

Belyaeva & Kovineva

Ono

Schurhoff

Wolf

Tucker, et al.

-

- -

-

-

-

-

-

-

- -

-

- -

-

-

-

-

-

-

-

-

-

-

-

-

96

-

-

-

-

-

-

-

- -

-

-

-

---- -

48

-

-

-

- -

-

-

- -

-

-

-

- - -

-

-

f. piperascens Holmes

M. arvensis L. -

s.sp. borealis (Michx.) Taylor et MacBryde

s.sp. haplocalyx var. piperascens

Holm.

M. arvensis - var . javanica (Blume)

H0ok.f.

var. piperascens Holm.

M. canadensis L. -

72

-

24, 72, 90

96

72, 74

36

72

96

72

96

96

72

60

96

96

96

54

54

96

M. cervina L. -

M. citrata - M. cordifolia -

M. crispa L. -

M. cunninghamii Benth. -

M. dumetorum -

M. gattefossei Maine -

M. gentilis L. -

36

36

36

96

3 6 , 48

48

84

4 8 , 84

72

96

40

20

48

+ 32 -

54 , 60 8 4 , 96 ,

108, 120

54

7 2

72

54

54 , 60 8 4 , 96 ,

108 , 120

- -

-

-

-

- -

-

- -

-

-

-

-

-

- - -

-

-

---- -

-

-

-

-

- -

-

-

-

-

-

-

-

-

- - -

-

-

-

-

-

-

-

- -

-

-

-

-

-

- -

-

-

-

-

-

-

Harley & Brighton

Tucker, et al.

Fernandes & Leitao

Sobti

Sobti

Murray

Harley & Brighton

Tucker, et al.

Hair & Beuzenberg

Sobti

Morton

Sobti

Makarov & Reznikova

Harley & Brighton

Morton

Sobti

Baquar

Tucker, et al.

Gill

Fernandes & Leitao

1977

1980

1984

1965

1962 b, 1965

1960 b

1977

1980

1960

1965

1956 a

1965

1972

1977

1956 a

1965

1967

1980

1981 b

1984

M. gracilis Sole. -

M. japonica Makino - M. lamarckii -

M. lavandulidora Sacco -

M. longifolia (L.) Huds. -

-

-

- -

-

- - -

- -

-

- -

-

-

-

-

-

-

-

-

-

54

60 , 7 2 , 8 4 , 96

49

36

96

18

18

24

24

18

48

48

24

36 , 48

24

18, 27

24 36 , 48

36

24

24

24

24

---- -

-

- -

-

- -

- -

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

- -

-

-

-

-

- -

-

-

-

-

-

-

-

Gill

Tucker & Fairbrothers

NagAo

Delay

Shimizu, et al.

Schurhoff

Lietz

Ruttle

June11

Heimans

Nagao

Polya

Tsuda

Morton

Murray

Sobti

Zhukova

Ouweneel

Majovsky, et al.

Sobti

Markowa & Iwanowa

1981 b

1990

1941 a

1947

1967

1929 b

1930

1931 a

1937

1938

1941 a

1950

1954

1956 a

1958, 1960 b

1962 b

1965

1967 b

1968

1970 a

1971 a,b

1972

s.sp. hymalaiensis Briq.

s.sp. lavandulacea Schimp ex Briq.

s.sp. longifolia

s.sp. typhoides Briq.

var. cyprica (Braun) Briq.

M. longifolia (L.) Huds. -

var. grandis Briq.

var. typica Fiori

M. microphylla Koch. -

M. muelleriana Schultz -

M. nemorosa Willd. -

M. niliaca Jacq.et Briq. -

M. parniroalaica Boriss -

M. piperita L. -

Wolf

Ruttle

Nagao

Morton

Murray


Sobti

Sobti

Baquar

Ouweneel

Belyaeva, et al.

Belyaeva & Kovineva

Bugaenko & Reznikova

Gill

Roy, et al.

Kundu & Sharma

M. sacchalinensis (Briq.) - Kudo

M. sapida -

M. satureioides R. Br. - -

M. scotica Graham - var. lacerata

M. smithian? Graham -

M. spicata (L. ) Huds. -

Belyaeva & Kovineva

Sobti

Harley & Brighton

Morton

Sobti

Ruttle

Nagao

Suzuka & Koriba

Morton

Murray

Sobti

Baquar

Harley & Brighton

Bugaenko & Reznikova

Pucker, et al.

:ill

coy, et al.

?ernandes & Leitao

Cundu & Sharma

Tucker & Fairbrothers

Harley & Brighton

Tucker, et al.

Harley & Brighton

Tucker, et al.

Harley & Brighton

Fernandes & Leitao

Silvestre

Harley & Brighton

Harley & Brighton

Schurhof f

Ruttle

Tischler

Rohweder

Delay

Bir & Saggoo

Gill

Saggoo & Bir

Heimans

Schurhof f

Lietz

var. acutifolia (Sm. ) Fraser

M. villosonervata Opiz. -

M. viridis L. -

var. typica

f. brevipetiolata (Rehb. ) Fiori

42 , 84 ,

120, 132

7 2 , 8 4 , 96,

120

96

7 2 , 7 8 , 84

7 8 , 8 4 , 90,

120, 132

84

48

36

36

32, 48

-

-

48

-

-

- -

-

-

- - -

-

24

24

-

----

---- -

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

- -

-

-

-

-

-

Morton

Baquar & Reese

Sobti

Suoninen

Olsson

VanLoon & de Jong

Morton

Schurhoff

Delay


Saggoo & Bir

Bir & Saggoo

Sacco, et al.

1956 a

1965

1965

1966

1967

1978

1956 a

1929 b

1947

1959 a

1983

1984

1968

Tribe : Salvieae I Genus : Salvia L. I S. aegyptiaca L. -

S. aethiopis L. -

S. albimaculata Hedge et - Huber - Morath

S. alborosea Epling et Jativa

S. amarissima Ort. -

S. amasiaca Freyn et Bornm. - S. amplexicaulis Lam. -

S. argentea L .

S. armenekensis Rech. -

S. armeniaca -

Delestaing

Bhattacharya, et al.

Humphires, et al.

Markova & Ivanova

Afzal - Rafii Sopova, et al.

Fernandes & Leitao

Singh

Afzal-Rafii

Harley & Heywood

Mercardo, et al.

Aryavand

Markova & Penka

Strid & Franzen

Markova & Ivanova

Markova & Thu

Afzal-Rafii

Strid & Franzen

Markova & Ivanova

Fernandes & Leitao

Afzal-Rafii

Patudin, et al.

S. atropatana Bge. - S.austriaca - S. azurea Lam. -

S. beckeri -

S. brachysiphon Stapf. - S. breviflora Moc. et Sesse -

S. cadmica Boiss - S. caespitosa Montbr. et -

Heldr .

S. campanulata Wall. - var. hirtella Stibal

var. nepalensis Stibal

S. candelatrum -

S. candidissima Vahl -

S. carduacea -

S. castanea Dielz.

S. ceratophylla L. -

S. chloroleuca Rech.f. et - Allen

S. chrysophylla Stapf. -

S. cilicica Boiss. -

I Afzal-Rafii

1 Markova & Ivanova

Scheel

1 Patudin, et al.

Afzal - Rafii

Harley & Heywood

Afzal - Rafii

Saggoo & Bir

Rosua & Gabriel

Van Loon & Snelders

Afzal - Rafii

Stewart

Epling, et al.

Gill

Afzal - Rafii

Fernandes & Leitao

Haque

Sugiura

Hruby

Delestaing

Erbrich

Harvey

Bhattacharya

Haque 1 1981

Krishnappa & Basavaraj 1982 I Saggoo & Bir

Gill

Bir & Saggoo

Harley & Heywood

Haque & Ghoshal I lg80a

Scheel

Gill

Bir & Saggoo

Gill

S. columbariae -

S. columbariae -

S. crassifolia Sibth. et Sm. -

S. cryptantha Montbr.et Auch - S. deserta - S. desoleana Atzei et Picci - S. discolor Kunth - S. dorisiana Fernald -

S. dumetorum Andrz. -

S. elegans Vahl. - var. sonorensis Fernald I

S. farinacea Benth. -

cv. " Lavender Blue'

cv. 'Royal Blue' I

cv. 'White'

Haque & Ghoshal

Stewart

Epling, et al.

Afzal - Rafii

Diana - Corrias

Harley & Heywood

Sugiura

Delestaing

Harley & Heywood

Sugiura

Gill

Bir, et al.

Gill

Haque

Haque & Ghoshal

S. fluviatilis Fern. -

S. forskohlei L. -

S. frigida Boiss.

S. fruticosa Mill. -

S . gerneriflora Lindl - Paxton

S. glutinosa L. -

S. grahamii Benth. -

Mercardo, et al.

Afzal-Rafii

Markova & Penka

Markova & Ivanova

Afzal-Rafii

Harley & Heywood

Scheel

Hruby

Polya

Linnert

Majovsky, et al.

Markova & Ivanova

Skalinska, et al.

Afzal-Rafii

Haque & Ghoshal

Strid & Franzen

Love & Love

Markova & Ivanova

Singh

Bir & Saggoo

Gill

Raman & Kesavan

Haque & Ghoshal

Haque

Haque

Saggoo & Bir

Bir & Saggoo

Afzal - Rafii

Markova & Ivanova

Palomino, et al.

Harley & Heywood

Ward

Scheel

Gill

Vir-Jee & Kachroo

Palomino, et al.

Afzal-Rafii

Haque & Ghoshal

Haque

Mercardo, et al.

Estilai, et al.

Linnert

Afzal-Rafii

Haque & Ghoshal

S. hydrangea DC ex Benth. - S. hypargeia Fisch. et Mey. - S. hypoleuca Benth. -

S. interrupta - s.sp. pani

S. involucrata -

S. itatiaiensis Dusem - S. judaica Boiss.

S. jurisicii Kosanin - .-

S. lanata Roxb. -

S. lanigera Poir. -

S. lavandulifolia Vahl -

S. lavandulifolia Vahl -- s.sp. amethystea

s.sp. blancoana

s.sp. maurorum

s.sp. mesatlantica

s. sp. Eyrenaeorum

s.sp. vellerea

s.sp. vellerea var. lagascana

Sopova, et al.

Afzal-Rafii

Aryavand

Afzal-Rafii

Rosua & Gabriel

Gill

Harley & Heywood

Afzal-Rafii

Sopova, et al.

Mehra & Gill

Gill

Sarkar, et al.

Kliphuis

Ubera

Rosua & Gabriel

S. lavanduloides Kunth

S. leucantha Cav.

S. limbata Mey. -

S. longispicata Mart. et -

S. lupulina Fern. -

S. macilenta Boiss. -

S.macrosiphon Boiss. -

S. merjamie Forsk. -

S. mexicana L. -

S. microphylla Kunth -

S. microstegia Boiss. et -

Palomino, et al.

Carlson & Stuart

Gill

Vij & Kashyap

Bhattacharya

Haque & Ghoshal

Bir & Saggoo

Haque

Gill

Afzal - Rafii

Mercardo, et al.

Palomino, et al.

Afzal-Rafii

Delestaing

Aryavand

Hedberg & Hedberg

Palomino, et al.

Harley & Heywood

Afzal-Rafii

S. molesta Mitchell -

S. montbretti -

S. moorcroftiana Wall. -

S. multicaulis Vahl

S. munzii Epling -

S. napifolia Jacq. -

S. nemorosa L. -

s. sp. nemorosa

I Kuriachan

/ Bhatt, et al. 1 Bir & Saggoo

Gill I Aryavand

Afzal-Rafii

Baker & Parfitt

Harley & Heywood

Afzal-Rafii

Hruby

Benoist

Tischler

Linnert

Chauhan & Abel

Majovsky, et al.

Aryavand

Afzal-Rafii

Afzal-Rafii

Markova & Ivanova

Markova & Penka

Van Loon & de Jong

S. nemorosa L. var. pseudo-sylvestris (Stapf.) Bornm.

S. nilotica Juss. et Jacq. - S. nubicola Benth. -

S. nutans - S. occidentalis Sw. -

S. ochrantha Epling - S. officinalis L. -

S. ombrophila Dusen - S. ~alestina Benth. -

S. persepolitana Boiss. -

14

32

12- 24

16 , 18 , 20 , 22, 24

22

18

32

14

14

14

14

14

14

14

14

14

14

-

240

20

-

-

-

-

-

-

-

-

-

-

- -

- -

-

- 7 -

- -

-

7

- -

10

-

-

-

-

-

-

-

- -

-

-

-

-

-

-

-

-

- - -

-

-

-

-

-

- -

- -

-

-

-

-

-

-

-

-

-

-

-

- -

-

Afzal - Rafii

Hedberg & Hedberg

Pal

Bhattacharya

Markova & Ivanova

Harley & Heywood

Scheel

Hruby

Suzuka & Koriba

Linnert

Butterfass

Love & Love

Gill

Af zal-Rafii

Haque & Ghoshal

Van Loon

Fernandes & Leitao

Gill

Harley & Heywood

Af zal-Rafii

Aryavand

S. phlomoides Asso. -

S. pinnata L. -

S. pisidica Boiss. et Heldr. -

S. plebeia R.Br. -

S. polystachya Ort. -

S. pomifera L. - s. sp. calycina (Sm. )

Hayek

s.sp. pomifera

S. potentillaefolia Boiss.et - Heldr .

S. pratensis L. -

S. pseudococcinea Jacq. -

20

22

14

16

16

16+ 1 B

16

-

22

14

14

14

16

16

18

18

18

18

18, 32

-

-

-

-

-

-

-

-

8+ 1 B

-

- -

-

- -

-

-

-

- 8 -

-

-

11

-

-

-

-

-

-

-

-

16+ 0-1B

-

- -

-

-

-

- -

-

-

-

-

-

-

-

-

-

-

-

-

8+ 0-1B

-

-

- -

-

-

-

-

-

-

-

-

-

Afzal - Rafii

Chauhan & Abel

Markova & Cerneva

Af zal-Rafii

Mehra & Gill

Gill

Vij & Kashyap

Gill

Harley & Heywood

Von Bother

Afzal - Rafii

Van Loon

Frey, et al.

Haque

Love & Love

Markova & Ivanova

Gill

1981

1968

1982, 1984

1980

1968b

1971

1975, 19761,

1984

1992

1970a

1980

1971

1980

1980

1981

1982

1982b

1982a

1984

S. purpurea Cav. -

S. guerceto-pinorum Epling - S. recognita Fisch. et Mey. -

S. relexa Hornem. -

S. reuteriana Boiss. - S. rhyacophila Epling -

S. ringens Sibth. et Smith -

var. macedonica Briq.

S. russegeri Fenzl. -

S. rutilans Carr. - S. sagittata R. & P. - S. sahandica Boiss. et Buhse -

S . scabiosifolia Lam. -

S. sclarea L. -

S. sclareoides Brot. -

S. sessei Benth. -

S. silvarum Epling -

22

22

16

22

20

-

-

12

16

12

14, 16

20

18

22

22

14

22

22

22

22

22

22

20

22

66

-

- - -

10

10

10

6

-

- -

-

9

11

-

- - -

- -

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

- - - -

-

-

-

-

-

-

-

-

-

-

- - - -

-

-

-

-

-

-

- -

-

-

- - -

-

-

-

-

- - -

Harley & Heywood

Afzal-Rafii

Haque & Ghoshal

Haque

Ghaffari

Harley & Heywood

Af zal-Raf ii

Markova & Ivanova

Af zal-Raf ii

Raman & Kesavan

Harley & Heywood

Aryavand

Afzal - Rafii Markova & Ivanova

Fujita

Afzal - Rafii

Van Loon & de Jong

Diana - Corrias Fernandes & Leitao

Elnir, et al.

Fernandes & Leitao

Harley & Heywood

S. tomentella Pohl -

S. tomentosa -

S. tonolea Boiss. - S. triloba L.fil -

S. urica Epling --

S. urnapuana Epling -

S. verbascifolia -

S. verbenacea L. -

22

-

22

22

86

-

22

14

14

22

12

16

64

54

42

59

4 2 , 5 4 , 5 8 , 6 0 , 64

54

4 8 , 5 4 , 58- 6 0 , 64

64

42

-

11

-

-

-

7. 8

- - -

-

-

- - - -

-

-

- -

-

-

-

-

-

-

- -

- - - -

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

- -

- -

- -

- - -

-

-

-

-

-

Haque

Mercardo, et al.

Harley & Heywood

Afzal - Rafii

Ferrarella, et al.

Fernandes & Leitao

Harley & Heywood

Palomino, et a1 .

Af zal-Rafii

Yakovleva

Benoist

Reese

Gadella & Kliphuis

Bjorkqvist, et al.

Bhattacharya, et al.

Dahlgren, et al.

Van Loon, et al.

Kramer, et al.

S. verbenacea L. -

s.sp. multifida

S. verticillata L. -

S. virgata Ait. -

S. viridis L. -

S. viridis L. -

S. viscosa Jacq. -

S. widemanni Boiss. - S. xanthocheila Boiss. - Tribe: Lamieae

Subtribe: Laminae

Genus : Anisomeles R. Br.

A. heyneana -

A. indica (L.) Kuntze -

A. malabarica R.Br. -

16

16

32 , 40

14

22

34

34

34

30

40

34

34

34

34

34

-

-

-

34

32

-

-

-

-

-

-

-

- -

20

-

-

-

-

-

17

17

17

-

-

-

- -

- -

-

- -

-

-

-

- - - -

-

-

-

-

34

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

-

Afzal - Rafii

Markova & Ivanova

Afzal - Rafii


Mehra & Gill

Gill

Sharma

Bir & Sidhu

Vij & Kashyap

Vembu

Cherian & Kuriachan



Saggoo & Bir

Gill

Bir & Saggoo

Kundu & Sharma


1980, 1981

1982a

1980

1972

1981

1982

1968b

1970

1970

1974, 1980

1975, 1976b

1979

1981

1981

1982

1983

1984

1985

1988a

1978,

-

- -

-

-

-

22

-

-

-

-

-

- -

-

-

- -

-

- - -

-

-

1968b

1970

1975, 1976b

1982

1984

1991

1982, 1983

1984

1982

1982, 1983

1983

1980

1980

1980

1984

1982

1982, 1983b

1983

1984

1980

1980

1982

1983b

- -

-

-

-

-

-

- - -

-

-

-

- -

-

-

-

-

-

-

-

-

-

-

-

-

14

11

15

14

-

11

-

-

- -

11

-

11

-

11

-

-

-

11

L. 3halotes Spreng. -

L. chinensis -

L. ciliata -

L. clarkei Hook. f. -

L. deflexa Hook. f. -

L. diffusa Benth. -

L. eriostoma Hook. - L. helianthemifolia Desf. -

L. hirta Spreng. --

Mehra & Gill

Gill

Vij & Kashyap


Gill

Singh, et al.

Saggoo & Bir

Cherian & Kuriachan


Saggoo & Bir

Renard, et al.

Ayyangar & Vembu

Krishnan


Cherian & Kuriachan


Saggoo & Bir

Cherian & Kuriachan

Ayyangar & Vembu



Saggoo & Bir

28

28

22

22

-

-

-

-

22

-

28

22

22

22

-

22

-

22

-

22

22

22

-

L. lamifolia Desf. -

L. lanata Benth. -

L. lanceaefolia Desf. -

L. lavandulaefolia Rees - L. lavandulaefolia Rees -

L. linifolia Spreng. -

L. marrubioides Desf. - L. martinicensis R. Br. -

L. mollissima Wall. -

L. mollissima Wall. - var. scaberula Hook.

-

28

30

-

28

-

22

-

22

-

22

-

-

-

28

28

28

-

- -

11

-

-

14

-

14

-

11

-

11, 11+ 0- 1 B

-

11+ 0- 1 B

14

14

-

-

-

14

14

14

-

-

- -

-

-

-

-

-

-

-

-

22

-

-

-

-

-

-

-

-

138

-

-

-

-

-

-

-

-

-

-

-

-

-

- -

-

- -

- -

Gill

Bhatt, et al.

Bir & Saggoo


Gill


Saggoo & Bir


Saggoo & Bir

Chopde

Saggoo & Bir

Cherian & Kuriachan

- Gill


Gill


Gill

Saggoo & Bir

Bir & Saggoo

1982, 19831,

1970

1975

1981, 1984

1982

1984

1982

1982, 1983b

1982

1983b

1965

1982

1981

1978

1982

1970

1982

1984

1981

1981, 1982a

L. montana Spreng. -

L. nutans Spreng. -

L. procumbens Desf. -

L. stricta Benth. -

L. suffruticosa Benth. -

L. urticaefolia R. Br. - L. vestita Benth. -

L. zeylanica R. Br. -

Genus - Leonotis R.Br.

var. nepetifolia

L. africana Briq. - L. nepetifolia (L.) Ait.f. -

Tribe - A-jugeae

Genus - Teucrium L.

28

-

15

22

28

-

22

22

-

24

28

-

14

-

-

-

11

-

-

11

T. aragonense Loscos et Pardo -

T. arduini L. -

-

-

-

-

-

22

-

-

-

-

22

-

-

52

32

-

-

-

-

-

-

-

-

-

- -

- -

-

-


Bir & Saggoo

Saggoo & Bir

Krishnan


Saggoo & Bir

Cherian & Kuriachan


Saggoo & Bir

Miege

Morton

-

-

1982

1980, 1981, 1984

1982, 1983

1980

1982

1982, 1983

1984

1982

1982, 1983a

-

-

Puech

June11

T. argutum R. Br. -

T. aureum Schreb. -

s.sp. angustifolium (Willk.)Puech

s.sp. aureum

s.sp. latifolium (Willk.)Puech

T. belion Schreb. -

T. bicolor (Sm.) Rees - T. botrys L. -

T. brevifolium Schreb. -

T. brachyandrum Puech - T. canadense -

s.sp. viscidurn (Piper) Taylor et Macbryde

T. capitatum L. -

var. polycephalum (Pomel) Briq.

T. carthaginense -

T. chamaedrys L. -

32

32

-

26

52

26

26

90

-

10

32

30

7 4

32

26

26 , 52

26

52

52

26

32

64

-

-

- -

- -

-

-

- -

- -

-

- -

-

-

- -

-

- -

- -

16

-

-

-

- -

14

- -

-

-

- -

-

-

- -

-

- -

Delay

Gill

Puech

Carmen

June11

Van Loon & Kieft

Strid

Puech

Love & Love

Puech

Fernandes & Leitao

Puech

Puech

Delay

Reese

- - - -

- -

-

-

-

-

-

- -

-

-

-

-

- - -

-

-

T. haenseleri Boiss. - T. halacscynanum -

T. hervieri Barnades - T. hircanicum -

T. lamifolium D'urv. -

T. lucidum L. -

T. mairei Senn. -

T. massilense L. -

T. mesanidum (Lit. et Maire) -- Sauv. et Vindt.

T. montanum L. -

ssp. montanum

T. nuchense -

I

Fernandes & Leitao

' Damboldt .

, Messia & Rejon

i Koudysheva & Akhmed- Zade

Markova & Cerneva

Fernandez, et al.

Puech

Van Loon & de Jong 1 1978 Contandriopoulos 1 1957c Fernandez, et al. 1 1977

Puech

Guinochet & Logeois

Van Loon & de Jong

Fernandez, et al.

Love & Love

Markova & Cerneva

Afzal-Rafii, et al.

Singh

Baksay

1978

Koudysheva & Akhmed - Zade 1 1968

T. orientale L. -

T. plectranthoides Gamble - I T. polium L. -

s.sp. aguilasense Puech I s.sp. capitatum (L.)

Briq.

var. majoricum (Rouy) Wk.

T. polium L. - s. sp. clape Puech I s.sp. gabesianum Le

Houerou

Aryavand 1977

Ga japathy 1962b

Lorenzo-Andreu & Garcia - Sanz.

Reese

Koudysheva & Akhmed - Zade 1 1968 Gill / 197lb

Markova & Thu I lg7* Puech

Puech

Vij & Kashyap

Puech

Sopova, et al.

Afzal - Rafii, et al.

Puech

Fernandez, et al. 1977

Markova & Cerneva 1982

Guinochet & Le Franc 1972

Puech 1 1911

s.sp. polium

s.sp. purpurascens (Benth.) Puech

s.sp. vincentianum (Rouy) Wood

T. pseudochaemopitys L. -

T. quadrifarium Buch - Ham. -

T. guadrifarium Buch - Ham. -

T. rouyanum -

T. royleanum Wall. -

T. salviastrum Schreber - T. sauvagei Le Houerou -

T. scordium L. - T. scordioides Schreber -

T. scorodonia L. -

T. stoloniferum Roxb. -

T. subspinosum Pourr. -

T. tomentosum Heyne -

T. vincentinum Rouy -

- Markova & Cerneva

- Ubera

- Malla, et al.

- Saggoo & Bir

- Gill

- ] Puech

-

- Gill

- 1 Bir & Saggoo

- Gill

- Fernandes & Leitao

- Puech

- Morton


-

- Bhattacharya

- 1 Guinochet & Lefranc

- Krishnappa & Basavaraj

- Cherian & Kuriachan


D..

m i 01

F i q . 56 C m p a r a t i v e id iog ram p l a t e o f d i f f e r e n t t a x a i n v e s t i g a t e d i n Lamiaceae. (Contd.) 12d - - 0. tenu i f l o ru rn L. f . cv . purp le-green, 13d - & t e n u i f l o r u m L.f. var . hirsuta H0ok.f.. 14d - Acrocephalus c a p i t a t u s Benth., 15d - Orthos iphon g l a b r a t u s Benth., 16d - g. q l a b r a t u s v a r . p a r v i f l o r u s Benth., 17d - 2. q r a n d i f l o r u s Bo ld cv. lilac, 18d - & q r a n d i f l o r u s Bo ld cv. white, 19d - g. p a l l i d u s Royle, 20d - 0. t h y m i f l o r u s Roth., 21d - P l e c t r a n t h u s numrnularius Br iq . , 22d - P. w i g h t i i Benth.

F i g . 56 Comparat ive i d i o g r a m p l a t e o f d i f f e r e n t t a x a i n v e s t i g a t e d i n Lamiaceae. (Contd.) 23d-Coleus a r a n a t i c u s Benth., 24d - i . a romat i cus Benth. var . v a r i e g a t a , 25d-C. bluernei Benth. var . v e r s c h a f f e l t i i , 26d- f o r s k o h l i i Br iq . , 27d-5. l a c i n i a t u s Benth., 28d-C. - e a r v i f l o r u s Benth., 29d- C. r e h n e l t i a n u s Berger, 3Dd-C. z e y l a n i c u s (Benth.) Cramer, 3 l d - A n i s o c h i l u s carnosus (L . f ) Wal l , 32d- - - A.er iocephalus Benth., 3 3 d - r n c a p i t a t a Jacq. -

F ig . 56 Comparat ive id iog ram p l a t e of d i f f e r e n t t a x a i n v e s t i g a t e d i n Lamiaceae. (Contd.)

34d-H. - suaveolens (L.) Po i t . , 35d-.Pagostemon benghalens is (6urm.f.) Kuntze, 36d-E. heyneanus Benth., 37d - P, purpurascens Dalz., 38d-c. v e s t i t u s Benth., 39d-Eus te ra l i s q u a d r i f o l i a (Benth.) Pan ig rah i , 40d - Mentha a rvens is L., 41d- - M. r o t u n d i f o l i a (L.) Huds., 42d - - M. spicata (L.) Huds.

F i g . 56 Compara t i ve i d i o g r a m p l a t e o f d i f f e r e n t t a x a i n v e s t i g a t e d i n Lamiaceae. (Contd.)

43d - c o c c i n e a Juss., 44d - 2. l e u c a n t h a Cav., 45d - 8. p l e b e i a R.Br. , 46d - 2. sp lendens Ke r - Gawl, 47d - Anisomeles indica (L.) Kuntze, 48d- A. m a l a b a r i c a R. Br., 49d-Leucas aspe ra Spreng., 50d- L. c e p h a l o t e s Spreng., 5 l d - L - l i n i f o l i a Spreng., 52d - L. s t r i c t a Benth., 53d - L. v e s t i t a Benth., - 54d - L e o n o t i s n e p e t i f o l i a ( L . ) A i t . f . , 55d- - Teucr ium p l e c t r a n t h o i d e s Gamble

16 18 22 24 26 28 30 32 34 36 40 44 48 64 72 96

CHROMOSOME NUMBERS

Fig. 57 Range of chromosome numbers recognized in the present study

6 w t 8 9 10 11 12 13 15 16 17

BASE NUMBERS

Fig. 58 Range of base numbers recognized i n the present study

2 X 4 X 6 X 8 X PLOIDY LEVEL

Fig. 59 Range of ploidy l e v e l recognized i n the present study

I - annual herb I - perennial herb

80 I11 - annual undershrub - perennial undershrub

7 0

6 0

Q 50 a rl 0 rl

E 40 rl 0 a

30 0

w M

2 20 2 0 $4

2 10

0 I I1 111 IV

GROWH HABIT

Fig. 60 Range of polyploids recognized under different growth habits in the present sutdy

~ig. 62: Comparison of the major karyotypic parameters in the cytotypes of Acrocephalus Benth., Orthosiphon Benth. and Plectranthus L' Herit. TCL - Total chromatin length, CLBC - Chromatin length of basic complement, TF% - Total forma percentage, DI - Disparity index, VI - variation coefficient. 13 - Acrocephalus capitatus Benth. (2n=18), 14 - Orthosiphon glabratus Benth. (2n=26), 15 - 0. &labratus var. parviflorus Benth. (2n =26), 16 - 0. grandiflorus Bold cv. lilac (2n=28), 17 - 0. grandTflorus Bold cv. white (2n=28), 18 - 0. pallidus ~oTle (2n=28), 19 - 0. thymiflorus Roth. (2n=24), 20 - Plectranthus - - -- nummularius Briq. (2n=28), 21 - P. wightii Benth. (2~24)

TCL CLBC 0 rn D1 vc

Fig. 63 Comparison of major karyotypic parameters in the cytotypes of Coleus Lour. TCL - Total chromatin length, CLBC - Chromatin length of basic complement, TF% - Total forma percentage, DI - Disparity index, VC - variation coefficient. 22 - Coleus aromaticus Benth. (2n=32), 23 - C. aromaticus Benth. var. variegata (2n=32), 24 - C. blumei Benth. var. verschaffeltii (2n=48), 25 - C. forskohlii Briq. (2n=28), 26 - C. - lacintatus Benth. (2n=48),arviflorus ~enth; (2n=72), 28 - C. rehneltianus ~ e r ~ e r - (2n=48), 29 - - C. zeylanicus (~entF~ramer (2~28)

Fig. 64 Comparison of major karyotypic parameters in the cytotypes of Anisochilus Wall., Hyptis Jacq. and Pogostemon Desf. TCL - Total chromatin length, CLBC - Chromatin length of basic complement, TF% - Total forma percentage, DI - Disparity index, VC - Variation coefficient. 30 - Anisochilus carnosus (L.f.) Wall. (2n=34), 31 - A. eriocephalus Benth. (2n=32), 32 - Hyptis capitata Jacq. (2n=32), 33 - H. suaveolens (L.T Poit. (2n=28), 34 - Pogostemon benghalensis (Burm.f. ) Kuntze (2n=64), 35 P. heyneanus Benth. (2n=64), 36 - P. - purpurascens Dalz. (2n=32), 37 - - P. vestitus Benth. (2n=32)

TCL CLBC TF% DI vc

Fig. 65 Comparison of the major karyotypic parameters in the cytotypes of Eusteralis Rafin., Mentha L. and Salvia L. TCL - Total chromatin length, CLBC - Chromatin length of basic complement, TF% - Total forma percentage,DI - Disparity index, VC - Variation coefficient. 38 - Eusteralis uadrifolia (Benth.) Panigrahi,(211=30),?9-Mentha arvensis L. (2n=96), 40 - M. rotundifolia L . 2 41 - g. spicata (L.) ~udn==48-- Salvia coccinea .Tun;. (2n=22), 43 - S. leucantha Cav. (2n=22), 3 4 -- S. plebeia R. Br., (2n=16+0-1 B) - - 45 - - S. splendens Ker-Gawl. (2~44)

TCL CLBC 0 TF% vc -

Fig.66 Comparison of the major karyotypic parameters in the cytotypes of Anisomeles R.Br., Leucas R.Br., Leonot R.Br. and Teucrium L. TCL - Total chromatin length, CLBC - Chromatin length of basic complement, TF% - Total for) percentage, DI - Disparity index, VC - Variation coefficient 46 - Anisomeles Indica (L.) Kuntze (2n = 34), 47 - A. malabarica R. Br. (2n = 32), 48 - Leucas aspera Spren, (2n = 22), 49 - L. cephalotes Spreng. (2n = 22), 50 - LT linifolia Spreng. (2n = 221, 51 - L. stricta Benth.(2n=2, 52 - L. vestita xenth. (2n=22), 53 Leonotis nepetifoKa (L.) Ait.f. (2n=28), 54 - ~eucrium plectranthoides Gamb. (2n=3T)

Xi - Primary base number

X2 - Secondary base number

PAP - Proto auto ploidy AP - Amphiploidy AD - Ascending dysploidy

DD - Descending dysploidy

2

Fig. 67 Phylogenetical scheme showing the probable evolution of chromosome numbers in the taxa investigated. (Bold numbers in the square's represent chromosome numbers identified)

DISCUSSION

a. Chromosome data in the genera investigated

Chromosome counts are made on fifty four members of Lamiaceae prevailing

in South India. Cytologically the family is variable. The chromosome

spectrum varies from 2n = 16 + 0-1B to 96, with majority of the species

concentrated in the number 2n = 32, followed by 2n = 48, 2n = 28 and

2n = 22 (vide Table 56; Fig. 57) . The presence of identical numbers in

unrelated genera is a very noteworthy feature in this family (Fedorov,

1969). The presence of such widely different series of chromosome numbers

in the species of even the same genus and in genera placed under

different tribes and subtribes, indicates that the different chromosome

numbers can be derived one from the other.

Perusal of the literature (vide Table 57) reveals that with the

exception of a few genera, where the available cytological data is very

meagre, all others exhibit a wide range of chromosome numbers.

Tribe - Ocimeae

The chromosome numbers in different grougs of Lamiaceae has proved to be

of important systematic value (Harley and Heywood, 1992). Thirty three

taxa belonging to seven genera have been examined. Wide range of

chromosome numbers occur in this tribe. But in spite of these

differences, there exist a relationship between the different genera and

species as evidenced by the frequent occurrence of inter and intra

specific variations.

Ocirum L. The genus seems to be highly plastic, both numerically and

structurally. The chromosome number ranges from 2n = 22 in O, adscendens

Willd to 2n = 72 in - 0. americanum L. (vide Table 56). This wide range of

chromosome numbers may be due to the difference in number of chromosomal

biotypes belonging to the different groups (Khosla and Sobti, 1985).

Previous chromosome counts suggest that the genus tolerates a high

chromosomal diversity in it's constitution. Some stabilized diploid

numbers found are 2n = 32, 2n = 40, 2n = 48, 2n = 64 and 2n = 72 (vide

Table 57).

Among the different members of Ocimum L., O, adscendens Willd. and - 0.

americanum L. show intraspecific variations (vide Figs. 2e, 3e, 3f, 3g)

Whereas O, gratissimum L. exhibits both inter and intraspecific

variations (vide Figs. 8b, 8e, 8f, 9b, 9e, 9g) variations in chromosome

number may be due to non-disjunction, somatic reduction or even by

partial endomitosis. Non-disjunction, in the somatic tissue, involves

unequal anaphasic separation, which results in unequal distribution of

chromosomes in the daughter nuclei (Sharma, 1976).

In - 0. basilicum L. the somatic chromosome number of 2n = 48, which seem

to be constant in the four varieties studied along with the absence of

any somatic variation numbers suggest the stability of the chromosomal

biotype. The absence of somatic variation numbers might have been

probably due to the cytologically diploid behaviour and normal

disjunction of chromosomes in these polyploids, during meiosis (Riley and

Chapman, 1958).

On the contrary, - 0. tenuiflorum L.f. is characterized by interspecific

variations. The chromosome number is found to be 211x32 in the cultivars

green and purple. Whereas in the cv. purple-green and the variety hirsuta

Hook.f., it is 2n = 36 (vide Table 56). The occurrence of different

cytotypes in the sane species suggest that chromosome number in this

species is not very stable and may alter either by the duplication or

loss of individual chromosomes (Morton, 1962).

In the closely related genetical strains of - 0. gratissimum L. (2n=40) and

0. gratissirnum L. var. suavis H0ok.f. (2n = 481, there exist two - chromosomal biotypes. The existence of different chromosomal biotypes in

the same species is a remarkable feature exhibited by the genus Ocimum

L.(Pushpangadan and Sobti, 1982).

Acrocephalus Benth. The chromosome count made on the lone member

investigated, A, capitatus Benth. is 2n = 18. Cytological reports are

rare in literature, probably due to the minute chromosome size.

Orthosiphon Benth. In this genus the chromosome number 2n = 28 occur in

three taxa and 2n = 26 in two cytotypes investigated. The somatic number

2n = 24 is rare (vide Table 56). The dominance of the chromosome numbers

2n = 28 and 2n = 26 in Orthosiphon Benth. is very well established in

literature (vide Table 57).

Plectranthus L' Herit. This genus is represented by two species, one with

2n = 28 (P. - nummularius Briq.) and the other with 2n = 26 (P, wightii

Benth.) in the present study. A review of literature reveals that the

chromosome number in Plectranthus L' Herit. ranges from 2n=22 to 2n = 84,

with a high frequency for the number 2n=28 (vide Table 57).

Coleus Lour. As regards the chromosome number this genus seems to be

heterogeneous with the frequent occurrence of the somatic numbers 211~28,

2n = 32 and 2n - 48. A similar trend has also been observed in literature (vide Table 57). Somatic variations are found in most of the members of

Coleus Lour. that are also adopted asexual means of multiplication. In

vegetatively reproducing plants variation leading to speciation is caused

by chromosome alterations in the somatic tissue (Sharma, 1956; Konvicka

and Levan, 1972).

Auisochilua Wall. The chromosome counts made on the genus were 2n = 34 in

A. carnosus (L.f.) Wall and 32 in A. eriocephalus Benth., both falling in - - terms with the previous reports (vide Table 57).

Hyptis Jacq. The chromosome numbers of 2n=32 and 2n = 28 have been found

in - H. capitata Jacq. and - H. suaveolens (L.) poit. respectively. These

numbers come under the wide range of chromosome numbers (211-16 to

96)reported for the genera (Harley and Heywood, 1992). More over the most

common numbers in literature happens to be 2n = 32 followed by 28 (vide

Table 57).

Tribe Hentheae Eight taxa belonging to three genera have been examined.

The chromosome numbers of individual taxa ranges from 211-24 to 2n = 96.

This tribe is also characterized by the frequent occurrence of

intraspecific variations. The chromosome numbers and the associated

variations provides a clue in determining the status of a species

(Sharma, 1976).

Pogost- Desf. The counts made on the genus reveal the codominance of

two chromosome numbers, 2n = 64 and 2n = 32. The dominance of these two

numbers in Pogostemon Desf. is very much evident in literature (vide

Table 57). These two numbers seem to be the multiples of a common

ancestral number and hence appears to have a common origin.

EusteraUs Rafin. The chromosome number 211-30 found in - E. quadrifolia

(Benth.) Panigrahi seems to be novel. Authentic cytological reports are

not met with in this genus.

kntha L. In the present study the counts of 2n=96, 2n = 48 and 2n = 24

shows the chromosomal diversity of the genus. All the three cytotypes

were characterized by variation numbers.

Previous work on the chromosomes of Mentha L. have shown that the

species vary considerably as regards the numerical status (vide Table

57). The genus is polymorphic. Sometimes different chromosome races are

found in the same species (Darlington and Wylie, 1955). Previous records

show that the chromosome number in Mentha L. range from 2n = 18 to 2n=144

(Morton, 1956a; Harley and Brighton, 1977).

Tribe: Salvieae

S a l v i a L. It is the lone genus investigated in this tribe. Out of the

four species studied, two possess 211-22 (S. - coccinea Juss. and - S.

leucantha Cav.), one shows 2n = 16 + 0-1B ( S . - plebeia R. Br.) and the

other 2n = 44 (S, splendens Ker - Gawl). Somatic variations are common

and several chromosomal biotypes are found in a species from which

different chromosome numbers are met with. A scrutiny of the world-wide

data indicates that the chromosome number in the genus Salvia L. ranges

from 2n = 12 to 2n = 240 (Harley and Heywood, 1992). In the genus the

most frequent numbers reported were 2n = 22 followed by 2n = 14, 2n = 16,

2n = 18, 2n = 20, 2n = 32 and 2n = 44 (vide Table 57).

Tribe Larieae

Eight taxa belonging to three genera have been examined. The normal

chromosome number ranges from 2n=22 to 2n = 34. Members of this tribe are

characterized by the absence of somatic variants. Chromosome numbers

certainly appears to have played a contributory role in the evolution of

different groups in the family through speciation and morphological

diversification (Harley and Heywood, 1992).

dniso~eles R.Br. In the two members investigated in this genus, A,

indica (L.) Kuntze possess 2n - 34 and - A. malabarica R. Br., 2n = 32.

These numbers seems to have originated from a common ancestor or one from

the other. Previous studies support the dominance of these two numbers in

Anisomeles R. Br. (Vide Table 57).

Leucas R. Br. The chromosome number was found to be constant (2n=22) for

the five investigated species of the genus. This shows the stability of

the genus at the infrageneric level. 2n = 22 seems to be the predominant

number along with 2n - 28 in the literature (vide Table 57). Leonotis R. Br. The chromosome number in L, nepetifolia (L.) Ait.f. was

found to be 2n=28, which falls in agreement with the earlier reports of

2n = 24, 2n = 26 and 2n = 28 made on the genus. (Vide Table 57).

Tribe djugeae

Teucriln L. The chromosome count made on the single taxa investigated in

the tribe, - T. plectranthoides Gamble is 211~32. This seems to be a

confirmation of an earlier report made on the same taxa (vide Table 57).

b. Basic chromosome numbers in the genera studied

Basic chromosome number forms one of the widely used characters in

formulating phylogenetic speculations and hence can be considered as a

dependable and stable marker of the direction of evolution (Jones, 1970,

1974, 1978). In Angiosperms several basic chromosome numbers are involved

in various polyploid series (Grant, 1981, 1982).

A review of the world-wide data on the basic numbers of Lamiaceae reveal

that they range from X - 5 to X = 64 (Darlington and Wylie, 1955;

Fedorov, 1969; Moore, 1973). However, the most common numbers were found

to be ranging between X = 5 and X = 19 (Leshukova, 1970).

In the present investigation the basic chromosome numbers vary widely in

South Indian Lamiaceae. Out of the fifteen genera investgated three have

been found to be polybasic (Ocimum L., Orthosiphon Benth. and Coleus

Lour.), five to be dibasic (Plectranthus L' Herit., Anisochilus Wall.,

Hyptis Jacq. Salvia L. and Anisomeles R. Br.) and three genera to be

monobasic (Pogostemon Desf., Mentha L., Leucas R. Br.) In the current

study, owing to the presence of only one species, it would not be

appropriate to categorize the genera Acrocephalus Benth., Eusteralis

Rafin., Leonotis R.Br. and Teucrium L. as monobasic. Thus it appears that

the family is a highly evolved one.

A great variability in the number of chromosomes in the basic set is a

characteristic feature for the mint family (Leshukova, 1970). This

variability could possibly be the result of aneuploidy at generic level

(Fernandes and Leitao, 1984).

Both primary as well as secondary base numbers are involved in the

evolution of the fifty four taxa studied. The primary base numbers (XI)

range from 6 to 9 and secondary base numbers (X2) from 10 to 13 and 15 to

17. The basic number X2 = 12 was found in majority of the members

(24.07%) followed by the numbers X2 = 11, X1 = 7, X1 = 8 and X2 = 16 with

percentages 16.67, 14.81, 12.96 and 11.11 respectively (vide. Fig. 58).

It is likely that protopolyploidy plays an active role in the evolution

of these basic numbers (Fujita, 1970).

In Lamiaceae, the secondary base numbers might have evolved from the

primary numbers, X1 = 6-9. Grant (1981) proposes the original primary

base numbers of angiosperms to range from X = 7 to 9 and they seem to be

ancestral in the phylogenetic sense. From these primary base numbers many

polyploid series develops both by autopolyploidy and amphipolyploidy. The

latter involves both hyper and hypoploidy (Fernandes and Leitao, 1984).

Thus it can be confirmed that polyploidy had influenzed and played a role

in the evolution of base numbers, mostly of the higher order ususally by

"polyploid drop" (Darlington, 1956) and occasionally by 'polyploid lift'

(Jones, 1970).

Tribe - Ocbeae

The common base numbers of this tribe found in the present

investigation are X = 6-13, 16 and 17. Ten out of the thirty three

members investigated are based on the genome number X = 12. Other

prominent numbers are X = 7 and 8.

Ocirum L. This is a polybasic genus (Bir and Saggoo, 19851, with both

primary and secondary base numbers observed in the present investigation.

A great diversity (X = 8-12) is observed in this regard, with half of the

investigated taxa being based on X = 12. These base numbers might have

evolved from the primitive base number X = 6, through dysploidy and

polyploidy at different levels (Sobti and Pushpangadan, 1977). Literature

evidences reveal the dominance of two common base numbers (X = 8 and

X = 12) in Ocimum L. (Sobti and pushpangadan, 1977; Singh and Sharma,

1982; Khosla, 1989).

Ocimum adscendens Willd. possess the secondary basic chromosome

number, X2 - 11. This might have been formed by amphipolyploidy (Grant,

1981) from the primary base numbers X1 = 5 and 6. The chance for the

evolution through ascending or descending dysploidy (Love, et al., 1957)

from X2 = 10 and X2 = 12 cannot be ruled out. - 0. americanum L. (2n=72)

exhibits secondary major protopolyploidy during the course of evolution

of it's base number X2 = 12 from the corresponding primary number (XI

= 6). Monobasic status (X2 = 12) was evident in 0. basilicum L., with all - the four varieties exhibiting doubling of the primary basic set X1 = 6.

The dibasic nature of - 0. gratissimum L. (2n = 40) with X2 = 10 and - 0.

gratissimum L. var. suavis H0ok.f. (2n = 48) with X2 = 12 might have been

due to independant autopolyploidy (Stebbins, 1971) from primary base

numbers X1 = 5 and 6 respectively.

The different cultivars and a variety of 0. tenuiflorum L.f. are set upon -

two primary base numbers. The basic number X1 = 8 was found in 0. -

tenuiflorum L.f. cv. green and cv. purple ( 2 ~ 3 2 each). Whereas in 0. - -

tenuiflorum L.f. cv. purple -green and 0. tenuiflorum L.f. var. hirsuta -

Hook.f., the basic set was found to be X1 = 9. To establish an ancestral

count among these two primary base numbers is a matter of confusion. The

primary base number X1 = 9 might have originated from X1 = 8 either by

progressive evolution or the vice versa by primary reduction (Jones,

1970).

Acrocephalus Benth. The basic chromosome number found in the lone member

investigated, A, capitatus Benth. was primary in nature with X1 = 9

chromosomes.

Orthosiphon Benth. This genus reveal the existence of three base numbers

out of the six members investigated. They are X1=6 in - 0. thymiflorus

Roth. (2n=24), X1=7 in - 0 . grandiflorus Bold cv. lilac, cv. white and 0,

pallidus Royle (each with 2n=28) and X2-13 in O, glabratus Benth. and - 0.

glabratus var. ~arviflorus Benth. (each with 2n=26). Morton (1962)

considers X1 = 7 as the original primary basic number of this genus. Then

the origin of X1 = 6 can only be through a diminution of the base number

(Stebbins, 1950). The secondary basic number X2=13 found in - 0. glabratus

var . parviflorus Benth. might be a case, evolved through polyploidy

followed by aneuploidy or dibasic polyploidy (Gill, 1970) involving the

primary base numbers X1 6 and 7.

Plectranthus L'Herit. In the present study this genus seems to be dibasic

with the primary base number X1 = 6 in P, wightii Benth. (2n=24) and X1 =

7 in - P. nummularius Briq. (2n=28). The common base number of Plectranthus

L'Herit is X = 6 (Darlington and Wylie, 1955). However deWet (1958b) is

of the opinion that X = 7 also exists. In Plectranthus L'Herit., the

primary basic number X1 = 7 can be the model number from which X1 = 6 is

derived by reduction (Morton, 1962).

Coleus Lour. The genus is polybasic (Bir and Saggoo, 1982a) with both

primary as well as secondary base numbers existing in it's members. The

primary basic number XI = 7 was found in - C. forskohlii Briq. as well as

C. zeylanicus (Benth.) Crarner and X1 = 8 in C. aromaticus Benth. and C. - - -

aromaticus Benth. var. variegata ( 2 ~ 3 2 each). The secondary base number

(X2 = 12) was found in - C. blumei Benth. var. verschaffeltii, - C.

rehneltianus Berger (211x48 each) and in - C. parviflorus Benth. (2n=72).

The primary basic numbers XI = 6, X1 = 7 and X1 = 8 have long been known

in Coleus Lour. (Darlington and Wylie, 1955). The ancestry of these

primary base numbers is a matter of considerable debate. Here X = 6 may

be considered as the primitive model base number from which XI = 7 and X1

= 8 might have been arisen, since an increase in the basic chromosome

number is easier to envisage and is likely to cause less unbalance than a

decrease (Darlington, 1963). The secondary basic number X2 = 12 has been

well established in Coleus Lour. (Bir and Saggoo, 1979, 1982a). This

higher basic number is considered to be derived f r o m lower numbers by

autopolyploidy (Stebbins, 1966).

BnisochFZue Wall. The genus exhibit two secondary basic numbers, one

with X2 = 16 in A, eriocephalus Benth (2n = 32) and the other with X2 =

17 in - A. carnosus (L.f.1 Wall. (2n=34). The secondary basic chromosome

number of X2 = 16 might have derived it's origin by autopolyploidy

(Fernandes and Leitao, 1984) from the primary base figure of X1 = 8.

Where as X2 = 17 might have formed by amphipolyploidy (Grant, 1981) from

the primary numbers XI = 8 and X1 = 9 or either through a 'polyploid

drop' (Darlington, 1956) from the primary base number X1 = 9 or through

a 'polyploid lift' (Jones, 1970) from X1 = 8.

Hyptis Jacq. The basic numbers found in the genus are primary in nature

with X1 = 7 and X1 = 8 and hence the genus appears to be dibasic. Morton

(1962) established these two numbers as the primary basic figures of

Hyptis Jacq. These numbers might have originated one from the other

through an increase (Darlington, 1963) or through diminution (Stebbins,

1950) of primary base figures.

Tribe Hentheae

In this tribe, the common base numbers found are 12, 15 and 16, all of

them being secondary in origin. Auto and amphipolyploidy seem to be

involved in the evolution of various members of this tribe.

Pogostemon Desf. The genus is monobasic with the secondary number X2 =

16, found in all the four investigated members. The existence of these

basic numbers were confirmed by Bir and Saggoo (1982a). This base number

might have been formed through autopolyploidy (Mehra and Bawa, 1969)

during evolution.

Eusteralis Rafin. The basic number found in the lone member investigated,

E. quadrifolia (~enth.) Panigrahi (2n=30) is X2 = 15. This secondary -

basic number is formed either through polyploidy followed by aneuploidy

or dibasic polyploidy (Gill, 1970) involving the primary base numbers

X1 = 7 and X1 = 8.

tkntba L. All the three investigated species are multiples of the

secondary basic chromosome number X2 = 12. Evidences from the literature

(Ikeda and Udo, 1963; Harley and Brighton, 1977) reveal the predominance

of this base number in the genus. This secondary base number can be

formed only through autopolyploidy (Stebbins, 1966) from the primary

basic number, X1 = 6.

Tribe - Salvieae

The basic chromosome numbers found in the investigated members of this

tribe include X1 = 7, X1 = 8, X2 - 11, X2 = 16 and X2 = 17. Both

ascending and descending dysploidy, auto polyploidy and amphipolyploidy

were found to be responsible for the evolution of various taxa coming

under this tribe.

S a l v i a L. The genus appears to be dibasic with both primary as well as

secondary base numbers found in the present investigation. When the

primary number X1 = 8 was found exclusively in S. plebeia R. Br., the - secondary number X2 = 11 was prevalent among S, coccinea Juss., S. -

leucantha Cav. and - S. splendens Ker-Gawl.

The genus Salvia L. is polybasic (Palomino, et al. 1986). Scrutiny of the

world-wide data indicates that the basic numbers range from X = 6 to 19

(Saggoo and Bir, 1986). However, the numbers 6, 7, and 8 are considered

as primary base numbers (Gill 1971 a). According to Afzal - Rafii (1976)

the base numbers prevalent in Salvia are X1 = 7 and X1 = 8, the latter

might have originated from the former as a result of progressive

evolution (Afzal-Rafii, 1971). The primary base numbers X1 = 5 and X1 = 6

seems to be involved in the amphipolyploid origin (Grant, 1981) of the

secondary basic chromosome number X2 = 11. It can also be formed by

ascending or descending dysploidy (Love, et al., 1957).

Tribe - Lamieae

Both primary as well as secondary base numbers seem to be prevalent in

this tribe. Dibasic polyploidy and descending or ascending dysploidy are

involved in the evolution of secondary base numbers.

Anisomeles R. Br. This genus exhibits the secondary basic number X2 = 16

in A, malabarica R. Br. (2~32) and X2 = 17 in - - A. indica (L.) Kuntze

(2~34). The former number arises by autopolyploidy (Stebbins, 1950) of

the primary base number X1 = 8 and the latter by dibasic polyploidy

(Gill, 1970) of the primary base numbers X1 = 8 and X1 = 9. A chance for

the origin of X2 = 17 through a 'polyploid lift' (Jones, 1970) of the

primary number X1 = 8 or even by a 'polyploid drop' (Darlington, 1956)

from XI = 9 cannot be ignored.

Leucas R. Br. All the five cytotypes of the genus possess the same

secondary base number X2 = 11. Literature surveys (Vij and Kashyap,

1976b; Bir and Saggoo, 1979, 1982a; Saggoo and Bir, 1986) confirms the

existence of X2 - 11 in Leucas R. Br. Chance for the occurrence of

amphipolyploidy (Grant, 1981) from the primary base numbers X1 = 5

and X1 = 6 as well as ascending or descending dysploidy (Love, et al.,

1957) from the respective secondary base numbers are equal during the

process of evolution.

Leonotis R. Br. The base number found in L. nepetifolia (L.) Ait.f. is

X1 = 7, which is primary in nature and well supported by an earlier study

(Morton, 1962).

159

Tribe Ajugoideae

Teucrium L. - T. plectranthoides Gamble (2n=32), the lone member

cytologically screened in this tribe reveals the presence of the primary

basic chromosome number X1 = 8, which has been confirmed earlier by

Koudysheva and Akhmed - Zade (1968).

c. Polyploidy in the genera investigated

Polyploidy is a very widespread cytogenetic phenomenon found in over 30%

of dicotyledons and 50% of monocotyledons (Love and Love, 1975). It is a

mechanism which involves multiplication of the whole chromosome

complement and there by an increase of gene number and variety, producing

radically different well adopted genomes (Stebbins, 1950; Gottschalk,

1979).

Polyploids are considered to be more hardy and adaptable to extreme

climatic conditions (Hagerup, 1931; Tischler, 1935; Love and Love, 1942b,

1943). However the theory of hardiness was criticized by Bowden (1940),

Clausen, et al., (1940, 1945) and Nielsen (1947).

Polyploidy is one of the best genetical and evolutionary processes, which

has greatly contributed to speciation and evolution of higher plants

(Gottschalk, 1985). This is mainly due to the ability of polyploids to

increase the chances of fertilization by breaking reproductive barriers,

which permits natural selection and establishment of species even under

adverse environmental conditions (Winge, 1917; Stebbins, 1971, 1974).

There can be no doubt that polyploid species are highly successful. A

comparison of the geographic distribution of polyploids and diploids in

plants show a greater adaptability of polyploids (Love and Love, 1943).

However the evolutionary potentialities of a diploid are likely to be

greater, in the long run (White, 1937).

The family Lamiaceae is characterized by a relatively high frequency of

polyploidy (Morton, 1962; Fernandes and Leitao, 1984). Studies conducted

on fifty four taxa reveals the dominance of polyploids (62.96%) over

diploids in South Indian Lamiaceae. Out of these, 57.41% are tetraploids

followed by hexaploids (3.7%) and octaploids (1.85%) (vide Fig. 59).

The present study reveal that herbaceous elements predominate as compared

with the shrubby elements of the family. Growth habit is one of the

factors which influence the frequency of polyploids in

angiosperms(Baquar, 1976). Correlating polyploidy with life form and

growth habit, it is clear that the highest frequency of polyploids occur

in perennial herbs (79.41%), lowest in annual herbs and undershrubs

(2.94% each) and intermediate in perennial undershrubs (14.71%) vide Fig.

60). In Angiosperms, the highest percentage of polyploidy was found in

perennial herbs while annuals and woody plants have lower percentage of

ploidy (Stebbins, 1938; Fagerlind, 1944) The origin of perennial habit,

as well as accessory methods of vegetative reproduction is considered to

be a direct consequence of polyploidy (Muntzing, 1936; Gustafsson, 1947a,

1948).

The various genera which show a predominance of polyploidy include Ocimum

L., Orthosiphon Benth., Coleus Lour., Hyptis Jacq., Pogostemon Desf.,

Mentha L., Salvia L., Leonotis R.Rr. and Teucrium L.

Tribe - Ocimeae

Different degrees of polyploidy has been observed in this tribe. Out of

the thirty three taxa investigated, twenty seven (81.82%) exhibit

polyploidy, with the majority being tetraploids, found in 25 members. The

role of polyploidy in the mechanism of speciation is obvious in the tribe

Ocimeae (Bhattacharya, 1978b).

Ocimum L. The cytological datas obtained from the genus Ocimum L.

indicated that eleven out of twelve members (91.67%) are polyploids.

Ocimum adscendens Willd. (2n = 22), the lone diploid representative,

exhibits a tetrasomic variant number of 24 (Fig. 2e) This might have been

probably caused by multipolarity that leads to unbalanced chromosome

complements (Brachet, 1957). At the molecular level the predisposition of

polyploidy shown by diploid individuals may be due to a change in the

gene expression and enzyme behaviour which causes a disturbed metabolic

regulation and consequently the abnormality of development. (Albuzzio, et

al., 1978).

Among the polyploid members, tetraploidy is found in all except - 0.

americanum L. (2n = 721, where the ploidy level is 6x. This hexaploid

might have been derived from a polyploid series based on the 'basic set of

12 chromosomes through autopolyploidy (Fernandes and Leitao, 1984).

All the varieties of - 0. basilicum are characterized by the absence of

variants. The stability of the chromosome number ( 2 ~ 4 8 ) indicates that

distinctly well differentiated genomes are involved in the origin of

these polyploids (Koul and Gohil, 1991).

162

0 . gratissimum L. (211-40) and 0. gratissimum L. var suavis Hook. f.(2n = - --

48) exhibit aneuploid somatic variants. Aneuploidy is very common in the

family at both generic and specific levels (Morton, 1962). It is caused

by the gain or loss of one or more chromosomes from the haploid set

(White, 1937).

The cultivars of - 0 . tenuiflorum L.f. exhibit interspecific polyploidy.

This is due to the existence of two different sets of chromosome numbers

2n = 32 and 36 derived through two different series, from the primary

base figures X1 = 8 and X1 = 9, by autopolyploidy (Clausen, et al.,

1945).

A perusal of the literature shows that so far all the investigated =urn

species are at polyploid level. This shows that polyploidy has played a

major role in the speciation of this genus (Morton, 1962; Pushpangadan,

et al., 1975).

Orthosiphon Benth. It is a genus, which was found to exhibit tetraploidy

in four out of the six members (66.67%) investigated. Here the cause for

polyploidy in the two cultivars of - 0 . grandiflorus Bold (2n = 28 each) - 0.

pallidus Royle (2n = 28) - 0. thymiflorus Roth. (2n = 24) may be due to the

parallel evolution of two basic figures X1 = 7 and X1 = 6 and formation

of multiples which develop into two series of polyploids during the

course of evolution (Fernandes and Leitao, 1984).

Plectranthus L' Herit. The two members investigated in the genus, namely

P. nummularius Briq. (211~28) and P. wightii Benth. (2n = 24) are - --

tetraploids. The genus shows predominance of polyploidy (deWet, 1958b).

163

Coleus Lour. The various cytotypes of the genus Coleus Lour. exhibits

polyploidy and varying degrees of somatic variations. The delicately

balanced system of gene interaction in tetraploids is disturbed by the

doubling of chromosomes, resulting in the formation of somatic variants

(Kuckuck and Levan, 1951).

C. aromaticus Benth. and C, aromaticus Benth. var. variegata are - - tetraploids with a basic set of X1 = 8 chromosomes. In both these plants

many somatic variants are present, which are multiples of the lowest one.

When such a phenomenon exists with in a single species of plant, it is

possible that they have arisen by autopolyploidy (White, 1937). The other

variant numbers can be hyperploids (Khush, 1973).

Members like - C. forskohlii Briq. and - C. zeylanicus (Benth.) Cramer

(2n=28 each) are autotetraploids based on the base figure X1 = 7. The

somatic variation numbers of - C. forskohlii Briq. might have been formed

through ascending or descending dysploidy (Love, et al. 1957). - C.

zeylanicus (~enth.), Cramer exhibits both aneuploid variant numbers (2n =

30 and 2n = 32) as well as multiples of the basic set of 7 chromosomes

(2n = 14 and 2n = 56). This shows the role of aneuploidy and

autopolyploidy that have played an important role in the evolution of the

taxa (Ramachandran, 1967).

The tetraploid taxa based on the somatic chromosome number 2 ~ 4 8 ,

blumei Benth. var. verschaffeltii, - C.laciniatus Benth. and C,

rehneltianus Berger may probably have an allopolyploid origin (Hakeem and

Rife, 1966). These three taxa also exhibit dysploid somatic variation

numbers. Dysploidy is quite a common feature found in Lamiaceae (Marrero,

1992).

The vegetatively propagated taxa, - C. parviflorus Benth. exhibits a high

degree of polyploidy (6x), probably due to the failure of cell division

in the first instance (White, 1937). It also exhibits varying degrees of

autopolyploid and aneuploid variants, a common feature of this family

(Fernandes and Leitao, 1984).

Byptis Jacq. The obnoxious weed - H. capitata Jacq. (2n=32) and it's ally

H. suaveolens (L.) Poit. (2n = 28) are tetraploids. There are several -

reports on records, where weedness is often associated with polyploidy

(Cain, 1944).

Tribe llentheae

This tribe was found to attain very high degrees of polyploidy in some

members and four out of eight members exhibit polyploidization. Most of

these polyploids are resorted to vegetative means of reproduction, an

adaptive feature exhibited by polyploids (Fagerlind, 1944; Stebbins,

1947).

Pogostamon Desf. Out of the four taxa investigated in this genus, P,

benghalensis (Burm.f.) Kuntze and P- heyneanus Benth. are tetraploids

with 2 ~ 6 4 . The diploids, P, purpurascens Dalz. and - P. vestitus Benth.

possess a somatic number of 2n = 32 each. Eventhough both the polyploid

and diploid taxa of Pogostemon Desf. are multiples of the basic figure X

= 16, the higher levels of ploidy attained by - P. benghalensis (Burm.f.)

Kuntze and & heyneanus Benth. probably point towards their greater

adaptability. The aneuploid and hypoploid variations found in these

polyploid taxa reveal the ongoing genetic and evolutionary processes

which may help to break the reproductive barriers enabling natural

selection and thereby speciation (Stebbins, 1974).

Hentha L. various levels of ploidy have been observed in the three

cytotypes investigated in this genus. Out of these - M. arvensis L. is an

octaploid, exhibitingfiypo+loid somatic variation numbers.

The octaploid status of & arvensis L. was confirmed earlier by Ikeda and

Seiroku (1966) and Tyagi (1986). Aneuploid numbers in & arvensis L. has

been reported earlier by Sharma and Bhattacharya (1959a), Ouweneel

(1968) as well as Tyagi and Naqvi (1987).

M. rotundifolia (L.) Huds. is characterized by it's diploid status and - occurrence of aneusomatic variation numbers. These numbers are multiples

of the lowest complement there by exhibiting proto-autopolyploidy (White,

1937). Polyploidy, aneuploidy and aneusomaty were found to be effective

in the evolution of various species of Mentha L. (Kundu and Sharma,

1985).

The cytotype - M. spicata (L.) Huds. exhibits tetraploidy (2n=48) as well

as aneuploid, hypoloid and hyperploid variations. Aneuploidy has been

reported on & spicata L. Huds. by Ruttle (1931a) and Nagao (194la).

Moreover the vegetative means of reproduction, which is prevalent in the

investigated taxa seems to bear a correlation with the high degrees of

polyploidy. Polyploidization might have led to the establishment of new

gene combinations that have triggered off new developmental changes

leading to a later shift towards the a sexual mode of reproduction

(Stebbins, 1980; Gustafsson, 1947b). Thus polyploidy and aneuploidy was

found to play an important role in the evolution of Mentha L. (Makarov

and Reznikova, 1972; Ouweneel, 1968; Pogan. et al., 1986).

Tribe - Salvieae

In contrast to the other tribes, it shows a lower percentage of

polyploids among the members studied.

Salvia L. It is a genus of much interest from the cytological point of

view. Eventhough polyploidy is prevalent only in one out of the four

species studied, variants occur in all the species, exhibiting natural

aneuploids from nullisomy to hexasomy. S, splendens Ker-Gawl. was the

only member to show tetraploidy. This was confirmed by earlier findings

(Bhattacharya, 1973; Bir and Saggoo, 1982a, 1985). Dysploidy was also

found in the different members of the genus (Afzal - Rafii, 1972, 1976).

The known trends of evolutionary development in the genus are dysploidy

(descending and ascending) with in the range of the genus and with in the

range of a single taxon, polyploidy and aneuploidy (Markova and Ivanova,

1982a; Palomino, et al., 1986)

Leonotis R.Br. The lone taxa investigated, - L. neptetifolia (L.) Ait. f.

shows tetraploidy from the base figure X1 = 7. Thus it seems to be a case

of autopolyploidy (Clausen, et al., 1945).

Tribe Ajugeae

Teucrium L. The single taxa investigated in the tribe, plectranthoides

Gamble, reveal the existence of autopolyploidy from the basic figure X1 =

8. An increase in the number of chromosomes through autopolyploidy

provides increased possibilities for new gene combinations, which are of

considerable importance in evolution (Reese 1961, 1966).

d) Karyowrphometrical analysis

In Angiosperms, the species of several families, both dicotyledons and

monocotyledons are found to exhibit a direct relationship between their

phylogeny and the chromosome constitution. The chromosomes being the

carriers of heredity, both structural and numerical changes in them can

influence the genetic-evolutionary process at work than do any other type

of changes. A detailed information regarding the chromosome architecture

in higher plants can thus serve as a useful tool to understand their

systematic relationships and for tracing the trend and direction of their

evolution (Love and Love, 1975).

Some of the major karyotype characteristics of considerable evolutionary

and taxonomic significance are (1) differences in the absolute chromosome

size, (2) differences in the position of the centromere, (3) differences

in total chromatin length (4) differences in karyotype formula and (5)

number as well as position of satellites.

The general feature noted in the family Lamiaceae is the wide range of

chromosomes with graded symmetrical karyotypes (vide Fig. 56), as has

been found in an earlier study. (Kundu and Sharma, 1988 a). However the

chromosome complements in South Indian Lamiaceae members differ in minute

karytypic details (vide Table 56). With regards to the gross morphology,

chromosomes are nearly submetacentric to nearly metacentric in nature.

The chromosomes range from 4.2 p to 1.0 p in length. The chromosomes

with secondary constriction ranges from two to six in number. The average

chromosome length (ACL) varies from 2.74 ,u to 1.12 p. The total chromatin

length (TCL) shows a very wide variation with 24.4 ,u being the minimum

and 154.8 p being the maximum value. The disparity index (DI) values

range between 22.22 and 50. The coefficient of variation ranges from

12.53 to 30.00 and total centromeric index (TFX) ranges from 35.83 to

45.07.

These variations found in the karyotypic parameters suggest that South

Indian Lamiaceae members are characterized by symmetrical to slightly

asymmetrical karyotypes. An increase in the range of chromosome length as

well as the increase of submetacentrics at the expense of metacentrics is

accompanied by an increase in the coefficient of variation leading to

asymmetry (Stebbins, 1958). Thus karyomorphological studies are of

considerable importance in order to throw light on the phylogenetic

relationship among taxa of flowering plants (Iwatsubo and Naruhashi,

1991).

Tribe - Ocimeae

Ocirum L. This major genus is characterized by a general homogeneity

observed in the minute details of the karyotype (Singh and Sharma, 1982).

The range of chromosome length (RCL) and their general morphology is the

same as has been noted for the family.

A remarkable feature found in all polyploid taxa of Ocimum L. is the

presence of four chromosomes with satellite or secondary constrictions

inspite of the difference in chromosome number. Thus it is clear that

numerical variations do not involve the satellited chromosomes. However

the micromorphology of these chromosomes varies from cytotype to

cytotype, indicating that structural changes have been operating in their

evolution. In spite of the evidence of the so called minor structural

variations, the presence of four nucleolar chromosomes in all the

polyploid forms of Ocimum L. indicate that these nucleolar chromosomes

may contain vital genes for the survival of the genus, which are

essential and not affected by the structural changes noted so far (Singh

and Sharma, 1981a).

The differences observed in the karyotype formula (KF), average

chromosome length (ACL) and chromatin length of basic complement (CLBC)

among different taxa might have been probably due to structural

alterations of chromosomes (vide Table 56; Fig. 61). Speciation depends

more on chromosomal rearrangements and mutation of individual genes than

on changes in the total amount of genetic content (Stebbins, 1959).

The high values of average chromosome length and chromatin length of

basic complement shown by - 0. adscendens Willd. and 0, americanum L. (vide

Table 56) probably show their primitiveness. Whereas, the lower values

for these parameters found in O,tenuiflorum L.f. cv. purple and CV.

purple-green denotes their highly evolved nature. A decrease in chromatin

length is one of the factors responsible for evolution of higher plants.

(Babcock and Cameron, 1934).

The high disparity index (DI) value found in - 0. americanum L. followed by

0. basilicurn L. var. purpurascens Benth. and var. pilosum Benth. - correspond to the heterogeneous assemblage of chromosomes in these taxa

(vide Fig. 56).Where as, the lower values of DI found in other members

point towards the general homogeneity found in various species of Ocimum

L. Normally a low disparity index value corresponds to the homogeneity of

chromosomes in most of the higher as well as lower plants (Mohanty, et

al., 1991).

In addition to the above, the high mean centromeric index (TF%) value

(vide Table 56) undoubtedly confirms the primitive status of - 0.

americanum L. A high TF% value represent a highly symmetric karyotype,

which is a primitive condition (Stebbins, 1959). However the

comparatively higher variation coefficient and lower TF% values exhibited

by - 0. tenuiflorum L.f. cv. purple shows that it represent the climax of

evolution among the different taxa investigated in Ocimum L.

In - 0. basillcum L., the karyotype of the variety purpurascens Benth.

possess a high number of medium-sized chromosomes (Type C) and low number

of small-sized chromosomes (Type Dl. Whereas, it is just the reverse in

the variety glabratum Benth. (vide Fig.56). A reduction of chromatin

length in - 0. basilicum L. var. glabratum Benth. appeared to have taken

place due to chromosomal rearrangement and the role of heterochromatic

segments during the process of evolution. Here the comparatively inert

heterochromatic segments might have been deleted resulting in the

shortening of the chromosome size (Darlington and Mather, 1950).

Moreover, the high values of average chromosome length, total chromatin

length, chromatin length of basic complement and TF% found in - 0.

basilicum L. var. purpurascens Benth. and the corresponding low values

found in - 0. basilicum L. var. glabratum Benth. further confirms that both

these are at two extremes in the evolutionary pathway (vide Table 56).

The decrease in chromatin length and the abundance of submetacentric

chromosomes with in a single complement are advanced karyomorphological

features (Babcock and Cameron 1934).

Concrete conclusions cannot be drawn from the karyomorphological features

exhibited by the closely related genetic strains of 0. gratissimum L. -

(2n=40) and - 0. gratissimum L. var. suavis H0ok.f. (2n=48). SQ it seems

likely that they represent two different lines as has been found in the

evolution of their basic numbers.

At the infraspecific level, O, tenuiflorum L.f. cv. purple and cv. purple

green are characterized by an increase in number of small sized

chromosomes (Type D) and the low values for the major karyomorphometrical

parameters except variation coefficient (vide Fig. 56, 61; Table 56).

Thus these observations are in advocacy with their highly evolved status.

The phylogenetic changes occurring in the length of the chromosomes oculd

cause shortening in size of one of the arms leading to the shifting of

the centromeric position as well as reduction in the absolute length of

the chromosome (Delauney, 1926).

Acrocephalus Benth.

In this genus, the low disparity index (DI) value reveals the homogeneity

of the chromosome complement. The high T F % found in conjunction with a

low variation coefficient value (vide Fig. 62; Table 5 6 ) reveal that

A. capitatus Benth. is a primitive plant. A high T F X and low variation - coefficient values correspond to the primitive status in the evolution of

flowering plants (Stebbins, 1971).

Orthosiphon Benth.

The comparatively high values for range of chromosome length ,average

chromosome length, total chromatin length and TF% found along with a low

variation coefficient (vide Fig. 62; Table 56), which occur in the

karyotype of - 0. glabratus Benth. and 0. thymiflorus Roth. point towards -

primitiveness. In addition to these, the small sized chromosomes (Type D)

are found to be absent in their karyotype. This along with a low

disparity index value in them show their homogeneity. Long metacentric

nearly equal chromosomes with a high mean centromeric index value were

found in comparatively primitive taxa (Levitsky, 1931). However, in - 0.

pallidus Royle,the lower values for range of chromosome length, average

chromosome length and chromatin length of basic complement found in

association with a high variation co-efficient and comparatively low TF%

value point towards an evolutionary advance.

At the infraspecific level, a numerical increase in small sized (Type D)

chromosomes and comparatively low values of average chromosome length and

total chromatin length (vide Fig. 56, 62; Table 56) found in Lglabratus

var. parviflorus Benth. and - 0. grandiflorus - Bold. cv. - lilac show their

evolved nature than their allys - 0. glabratus Benth. and - 0. grandiflorus

Bold cv. - white. This has been further confirmed by the higher variation

coefficient value found in O, glabratus var. parviflorus Benth. and a low

TF% value found in - 0 . grandiflorus Bold. cv. lilac. Thus these minute

structural alterations might have been responsible for evolution. The

morphological evolution of chromosomes involves a change from uniform

length of chromosomes to differences, change of median centromeres to

submedian centromeres and reduction in chromosome length (Jones, 1978).

In addition to these, differences are also found in the number of SAT-

chromosomes (vide Fig.56; Table 56). The secondary constriction

constitute an important feature of chromosome in karyomorphological

studies at the infraspecific level (~ewis and John, 1963). The presence

of only one pair of SAT - chromosomes in the tetraploid taxa, - 0.

grandiflorus Bold cv. lilac and cv. white may be due to the loss of one

pair of chromosomes during the course of evolution. It also seems that

the amount of nucleolar genes contained with in the satellited

chromosomes is sufficient to meet the need of the ribonucleoprotein

metabolism as has been found in other plants (Singh and Sharma, 1983).

Plectranthus L'Herit. The lower values for range of chromosome length,

average chromosome length, total chromatin length as well as chromatin

length of basic complement and a comparatively high value of variation

coefficient (vide Fig. 62; Table 56) found in P, nummularius Briq., than

it's relative - P. wightii Benth. probably indicate an evolutionary

advance. Moreover, an increase in chromosome number is observed in 11,

nummularius Briq. (vide Fig. 56). Reduction in chromosome size is

apparently a consequence of polyploidy, since it is an adaptation to a

decrease in size of the cell or to an increase in number of chromosomes

(Darlington, 1956).

Coleus Lour. The different species of Coleus Lour. are exceedingly

variable both in chromosome numbers and in morphometric characters of

their karyotypes. This variability of the genus evidently reflects an

important side of it's evolution, apart from it's morphological

differentiation, on which classification is based.

In C, aromaticus Benth. the higher number of nearly median centromeres

and the higher values for all the other karyomorphological parameters

(vide Fig. 56, 63; Table 56) reveal it's primitiveness. In contrary, the

karyotypic data found in - C. aromaticus Benth. var. variegata speaks off

it's evolutionary advance.

On comparing the karyomorphology of cytotypes with 2n = 28 chromosomes,

C. zeylanicus (Benth.) Cramer exhibits a higher number of short sized - (Type D)chromosomes, higher value for variation co-efficient (V.C.) and

lower values for other karyomorphological data (vide Fig. 56, 63; Table

56) Where as, in - C. forskohlii Briq. it is just the reverse. The advanced

nature of - C. zeylanicus (Benth.) Cramer and the primitive status of - C.

forskohlii. Briq. is very much evident. Similarly - C. laciniatus Benth.

pssesses a higher number of short sized (Type D) chromosomes and lower

values for other karyotypic details (vide Fig. 56, 63; Table 56) when

compared with other members based on 2 ~ 4 8 . Hence - C. zeylanicus (Benth.)

Cramer seems to be evolved. The structural rearrangements related to

favourable position effect and gene linkage, which are likely to confer

adaptive advantage, may increase the asymmetry of karyotype (Stebbins,

1958).

C. parviflorus Benth., a hexaploid is characterized by the presence of 6 -

chromosomes with secondary constrictions (vide Fig. 56). It has been very

well established that each basic set of chromosomes has at least one

chromosome with secondary constriction (de Mol, 1926b; Bhaduri, 1944).

Moreover the lesser range of chromosome length and low variation

coefficient value and the comparatively higher TF% value indicate their

probable primitiveness (vide Table 56; Fig. 63).

Thus in the genus Coleus Lour., structural alterations seem to have a

major role during the course of evolutiqn and these changes might have

been probably due to pericentric inversion and unequal translocation in

the chromosomes (Stebbins, 1971).

175

dnisochilus Wall. Karyotypic analysis conducted on the two diploid

species of this genus reveal the homogeneity, as was evidenced by their

almost similar karyotype formula (KF) and comparatively higher value of

disparity index (DI). However, differences are noted for average

chromosome lenth, total chromatin length, chromatin length of basic

complement, variation co-efficient and TF%. (vide Fig. 64, Table 56). The

higher value among these being shown by A, carnosus (L.f.) Wall., thereby

exhibiting it's primitiveness. Karyotype with lesser range of chromosomal

length is found in comparatively primitive genera (Levitsky, 1931).

Hyptis L. Karyomorphometrical surveys on the genus are rare, probably due

to the small chromosome size. Both the tetraploid taxa investigated are

characterized by homogeneous graded karyotypes. (vide Fig. 56). In - H.

capitata Jacq. all the karyometrical values seem to be comparatively

higher than that in - H. suaveolens (L.) Poit. (vide Fig. 64; Table 56).

Such variations are important from the evolutionary viewpoint. Thus

primitiveness is more pronounced in & capitata Jacq. than suaveolens

(L.) Poit. This is probably due to karyotypic rearrangements which is an

initial step in speciation as has been found in other plants (Clausen,

1951).

Pogost-n Desf.

With in the genus Pogostemon Desf., the tetraploid taxa showed more

similarities among them than differences when compared with the diploid

taxa. The polyploid plants, P, benghalensis (Burm.f.) Kuntze and - P.

heyneanus Benth. showed homogeneous symmetrical karyotypes, a primitive

condition. However, the lesser values of average chromosome length,

chromatin length of basic complement, disparity index and TF% (vide Table

56., Fig. 64) advocates their evolved nature. Among the karyotypes of

the diploid forms, - P. vestitus Benth. exhibited a higher number of short

sized (Type D) chromosomes, comparatively low values of average

chromosome length, total chromatin length as well as TF% along with a

high value of variation coefficient. Therefore - P. vestitus Benth. can be

considered as more advanced than - P. purpurascens Dalz. Structural

alterations of chromosomes have a major role towards speciation (Singh

and Sharma, 1982) .

A remarkable feature noted in the four taxa of Pogostemon Desf. is the

presence of 4 SAT-chromosomes (vide Fig. 56), irrespective of their

ploidy level. Thus it has become undoubtedly clear that in some plants

the number of nucleolar chromosomes maynot increase proportionately with

the concomittant increase in ploidy (Singh and Sharma, 1983).

Eusteralis Rafin.

The karyotype of - E. quadrifolia (Benth.) Panigrahi seems to be formed of

very small chromosomes, symmetrical and with a very low range of

chromosome length (vide Fig. 56). Alongwith these primitive features, it

exhibits some advanced evolutionary features like low values for average

chromosome length, chromosome length, chromatin length of basic

complement and TF% (vide Fig. 65; Table 56). So the evolutionary status

of this taxa appears to be enigmatic.

Mentha L.

In the diploid, tetraploid and octaploid species of Mentha L., the

karyotype seems to be symmetrical and homogeneous (vide Fig. 56). The

presence of only two pairs of chromosomes with secondary constrictions in

a11 the species probably show that two pairs of nucleolar chromosomes are

essential for the existence and stability of the different species of

Mentha L. In some plants, SAT - chromosomes are not affected by numerical variations (Kundu and Sharma, 1985). However,cryptic structural changes

have been operating in their evolution. Among the three taxa, & arvensis

L. was found to exhibit a comparatively low TF% and high variation co-

efficient value (vide Fig. 65, Table 56). This correspond with the

evolutionary advancement of this plant. Karyotypic details also points

out the evolved status of - M. spicata (L . ) Huds. However, the diploid taxa

M. rotundifolia (L.) Huds. with a low average chromosome length, and low -

disparity index value and a high TF% seem to be rather primitive in the

evolutionary history of the genus. The centromeric index is a karyotype

characteristic independant from chromosome number. A low mean centromeric

index value suggest a highly advanced karyotype and a high value

represents a primitive karyotype (Vasil'eva, et al., 1985).

Salvia L. The genus exhibit differences as regards chromosome

morphology. In the diploid taxa based on 2n = 22, S. coccinea Juss. shows - -- comparatively lesser values for range of chromosome length, average

chromosome length, total chromatin length, chromosome length of basic

complement and TF.% (vide Fig. 65, Table 56) than S. leucantha Cav. - Eventhough a low range of chromosome length is a primitive character,

all the other karyomorphological data, along with a high variation

coefficient value reveal it's advanced status.

The karyotype of S. plebeia R.Br. is characterized by a higher variation - coefficient and a lower value for chromatin length of basic complement.

These two parameters are sufficient to place it among the advanced

members of Salvia. L. In the present study, B-chromosome are detected

only in the chromosome complement of Salvia plebeia R. Br., (vide Fig. 45

b) ranging from 0-1 in number. This B-chromosome might have arisen by the

fragmentation of a normal chromosome (Kranz, 1971). B-chromosomes

suppress vegetative vigour and delay the onset of flowering (Jones,

1974), rare features observed, when compared with other members of Salvia

L.

The tetraploid member, - S. splendens Ker - Gawl. exhibit the highest TF%

value noted for the genus. Thus it can be c0nsidered.a~ primitive. Where

as, the diploid forms, S, coccinea Juss., - S. leucantha Cav. and &

plebeia R. Br. exhibit two SAT chromosomes each. Where as, the

tetraploid taxa, - S. splendens Ker-Gawl. exhibit four SAT-Chromosomes. In

plants, normally one satellited chromosome will be present in each basic

set of chromosomes (Bhaduri, 1944).

Adso~eles R. Br. In the two diploid members investigated in the genus

Anisomeles R.Br., A. indica (L.) Kuntze shows low values for all - -

karyomorphological characters. (vide Table 56; Fig.66) Eventhough the

lesser range of chromosome length and lesser variation coefficient value

denotes primitiveness, all the other characters including a low TF% (vide

Fig. 66, Table 56) shows the probale advanced status of A. indica (L.) - -

Kuntze than A- malabaica R. Br. The karyomorphological differences found

among these two taxa fully justify that speciation and evolution in this

genus has been principally effected by structural alteration of

chromosomes as observed in some other plants (Cleland, 1949).

Leucas R. Br. The different species of the genus w s R.Br. are

characterized by the homogeneous karyotypes as was evidenced by the

179

similar karyotype formula shown by all, except L, vestita Benth. Despite

the constancy in the chromosome number, the karyotypic differences among

the varieties were quite distinct (vide Table 56, Fig. 66).

In - L. vestita Benth., the high values obtained for all karyotypic details

except variation coefficient value (vide Fig. 66; Table 56) reveal it's

primitiveness, when compared with all the other cytotypes of Leucas R.Br.

Another remarkable feature shown by L, vestita Benth. is that they

contain only one pair of SAT chromosome when compared with two nucleolar

chromosomes found in & aspera Spreng., - L. cephalotes Spreng.,

linifolia Spreng. and - L. stricta Benth. (vide. Fig. 56). The secondary

constrictions are not of much importance when compared with other

karyotypic details, since they might be lost or gained during the course

of evolution (Lewis and John, 1963).

Leonotis R. Br. The higher karyomorphological values exhibited by L.

nepetifolia (L.) Ait.f. denotes it's primitive status. However the vast

difference of chromosome length and a high disparity index value (vide

Fig. 66; Table 56) denote an advanced heterogeneous nature. The

karyotypes that are heterogeneous both cytologically and genetically are

important in the evolution of species (Stebbins, 1958).

Teucrium L. In a more or less contradicting manner, the karyotype of 2

plectranthoides Gamble is characterized by a slight difference in

chromosome length as well as a high TF% value and hence seems to be

primitive. However, the low values for average chromosome length, total

chromatin length and chromatin length of basic complement (vide Fig. 66;

Table 56) point towards their evolved nature. Therefore the evolutionary

status of this taxa seems to be an enigma.

The karyomorphological diversity found in the different taxa of Lamiaceae

in South India can be best explained by assuming that the group is still

undergoing active speciation.

e) Meiotic Analysis

Meiotic studies are relevant, since they focus on details of pairing

behaviour of chromosomes, recombination frequencies and more importantly

their pattern of disjunction during Anaphase I and I1 which are not

deducible from mitotic studies (John and Lewis, 1965). Meiosis is an

integrated process consisting of a series of events which have been shown

to be under genetic control (Golubovskaya, 1979). Normal meiosis leads to

the formation of functional gametes, where as an irregular meiotic

division leads to abnormalities and there by to sterility (Del-Duca,

1976; Moraes - Fernandes, 1982).

In the present investigation, meiotic studies have been conducted on

twenty four members which bloom frequently in the experimental garden.

Out of these nineteen taxa exhibited normal meiosis and five taxa,

abnormal meiosis. Abnormalities in the normal course of meiosis,

especially failure of chromosome pairing has been reported to be

occurring spontaneously in various plant species (Gottschalk and Kaul,

1980).

Interestingly thirteen out of the nineteen taxa with normal meiosis are

polyploids. This might have been probably due to the absence of

multivalent associations (Swaminathan and Sulbha, 1959). The multivalent

formations which were found to be absent in Lamiaceae owe to the small

size of chromosomes. It is clear that well differentiated genomes are

involved in the origin of these polyploids (Koul and Gohil, 1991)

m b e r s with normal peiosis -

Out of the twentyfour taxa investigated, meiosis was perfectly normal in

six diploids and thirteen tetraploids. The diploids which exhibited

normal meiosis are Ocimum adscendens Willd., Orthosiphon glabratus

Benth., & glabratus var. parviflorus Benth., Salvia coccinea Juss., S,

leucantha Cav. and S. plebeia R. Br. -

The polyploids which exhibited normal meiosis included four varieties of

Ocimum basilicum L., four taxa of - 0. tenuiflorum L.f., two cultivars of

Orthosiphon grandiflorus Bold, & pallidus Royle, & thymiflorus Roth.

and Plectranthus wightii Benth. While considerable chromosomal

irregularities are expected in the metaphase I and anaphase I of this

plant, surprisingly it exhibits normal meiosis. This clearly indicates

the cytological stability attained by the polyploid to maintain it's

level. In all these species only bivalents are observed at diplotene and

metaphase I stages followed by normal disjunction of chromosomes to the

two poles.

This shows that irrespective of the degree of polyploidy, chromosomes

always exhibit bivalent formation. As a consequence polyploid meiosis is

characterized by bivalent formation. This can be used as an argument to

consider these plants as allopolyploids. However autopolyploids of long

standing tend to behave like allopolyploids (Giles and Randolph, 1951).

Another point of argument is that autopolyploids can undergo normal

meiosis as a result of the action of some genes which promote homologous

pairing only and not homeologous pairing (Sears and Okamoto, 1958).

mbers with abnormal meiosis

Abnormal meiotic behaviour is found in five out of the twenty four

members examined by smear experiments. The chances of survival of

polyploids are determined by their meiotic behaviour (Evans, 1988). The

tetraploids which exhibit abnormal meiotic status are Ocimum gratissimum

L., O, gratissimum L. var. suavis Hook. f., Plectranthus nummularius

Briq. and Salvia splendens Ker - Gawl. The hexaploid member - 0. americanum

L. also shows meiotic abnormalities. Meiosis in these polyploids are

characterized by the complete lack of multivalent associations even in

members with some what large chromosomes. This suggests that the size of

the chromosomes is perhaps no guide for multivalent formation, but it is

the genetic factors that controls chromosome pairing in autotetraploids

(Singh, 1991). This is evident from the discovery of genes which affect

different stages of meiosis (Rhoades and Dempsy, 1966)

The major meiotic abnormalities encountered in Ocimum americanum L., O,

gratissimum L. and 0- gratissimum L. var. suavis Hook. f. are the

occurrence of unpaired chromosomes at metaphase I and irregular

seggregation during anaphase I. This has been confirmed by earlier

observations made on the genus Ocimum L. (Sobti and Pushpangadan, 1982).

In Ocimum gratissimum L. and O, gratissimum L. var. suavis Hook. f. the

comparatively high number of univalents (vide Fig. 8j, 9j) reveal the

meiotic instability. A univalent is a chromosome that has not undergone

pairing at zygotene or may be the components of a bivalent separated at

diplotene owing to the absence of chiasma formation (Kihara, 1931;

Rosenberg, 1927). Thus the univalents at the end of first meiotic

division mechanically prevent complete separation of the two main groups

of chromosomes, leading to polyploidy (White, 1937).

In addition to this Ocimum gratissimum L. exhibits aberrant spores in the

form of pentads, hexads, octads etc. (vide Figs. 8n, 80, 8p). This may be

due to the unequal seggregation of chromosomes during telophase I1 or due

to the failure at cytokinesis in the last premeiotic division (Nath and

Nielson, 1963). Thus the occurrence of polyspory might be related to

chromosomal irregularities observed during meiosis (Longley, 1925).

The meiotic abnormalities found in Plectranthus nummularius Briq. (vide

Fig. 21j) might have forced the plant to resort to vegetative means of

reproduction, which was found to be the more prevalent method of

propagation. In many flowering plants exhibiting abnormal meiosis, escape

from the bottleneck of sterility is provided by the apomictic mode of

reproduction (Darlington, 1939).

The species Salvia splendens Ker - Gawl. exhibit partial or loose pairing

(vide Fig. 46j) Meiotic abnormalities were reported in many species of

Salvia L. by various authors (Delestaing, 1954; Linnert, 1955b; Haque and

Ghoshal, 1980a).

f) Cytological evolution ift Lamiaceae

From the cytological observations made in the present investigation it

can be concluded that speciation and evolution with in this family has

been possible as a result of increase in variability through changes in

the base numbers, as well as numerical and structural changes in

chromosome numbers.

The vnrious cytological phcnomenn like proto-auto pl.oidy, amphiploidy,

ascending and descending dysploidy might have resulted in the variability

of base numbers in the family (vide Fig. 67).

The wide range of chromosome numbers observed in many genera (vide

Fig.57) in the present investigation marks a significant role that an

aneuploidy and polyploidy have played in the evolution of various taxa of

the family at the generic and species level. It also appears that various

kinds of aberrations have played a vital role in the evolutionary

diversification of the family. The mitotic and meiotic irregularities

might have lead to structural and numerical variations in the chromosomes

of a species. Induviduals with the same chromosome number but with

differences in karyomorphological details reflect the ongoing

evolutionary processes at micro level.

It seems probable that in South Indian Lamiaceae, Robertsonian changes or

mutations might have also played an important role in the evolution of

karyotype. Drastically mutated induviduals are usually unstable and unfit

to survive in nature because they express various degrees of weakness and

chromosomal aberrations leading to genetic sterility. However some

induviduals carrying the changed chromosomal constitutions are well

within their range of tolerance. This was confirmed by the occurrence of

normal meiosis in some polyploids investigated.

More over, most of the taxa belonging to this family have efficient means

of vegetative propagation. This ensures the survival of these genetically

altered types which otherwise would have faced extinction on account of

sexual sterility imposed as a result of these changes. Accumulation of

such small changes can sometimes lead to a taxonomic divergence in a

species during the process of evolution. Thus meiotic accidents may prove

to be more useful than mitotic aberrations from the evolutionary point of

view, since the meiotic abnormalities are hereditary and likely to

multiply and establish in a population.

Polyploidy is said to have an ancient origin if a taxon includes only

polyploids or very few diploids, and relatively of the recent origin if

diploids are more frequent than polyploids or both are in almost equal

proportion (Stebbins, 1950). On this context among the members

investigated, the various genera, viz., Ocimum L., Orthosiphon Benth.,

Plectranthus L' Herit., Coleus Lour., Hyptis Jacq. and Mentha L.,

polyploidy is assumed to have an ancient origin. However polyploidy seems

to be of recent origin in Pogostemon Desf., Salvia L., Anisomeles R.Br.

and Leucas R.Br. An effective conclusion cannot be reached in the genera

like Acrocephalus Benth., Eusteralis Rafin., Leonotis R.Br. and Teucrium

L., since only one taxa has been investigated in each of them.

Thus the family Lamiaceae seems to be in a fairly active state of

evolution because of the quite common occurrence of polyploidy and

aneuploidy. Mutation of genes, structural changes in chromosomes, non-

disjunction, chromosomal rearrangement and several other abnormalities

are those mechanisms leading to the development of polyploidy and thus to

the differentiation of new taxa in Lamiaceae (Fernandes and Leitao,

1984).

There are still enormous gaps in our knowledge as regards the cytological

evolution of Lamiaceae, and much still remains to be done before a major

cytotaxonomic review may be attempted.

Fifty four taxa representing a total of fifteen genera of Lamiaceae found in

South India are examined cytologically. The chromosome spectrum in South

Indian Lamiaceae ranges from 2n=16+0-1 B to 2n596, with majority of the

species concentrated in the number 2n = 32, followed by 2n = 48, 2n = 28 and

2n = 22. In spite of the wide range of chromosome numbers, there exists a

relationship between the different genera and species as evidenced by the

frequent occurrence of numerical variations. This wide range of chromosome

numbers may be due to the difference in number of chromosomal biotypes

belonging to the different groups. In the present investigation, the genera

like, Ocimum L., and Coleus Lour. exhibit inter, intra and infra specific

variations among chromome numbers. Both inter and intraspecific variations

have been found in Anisochilus Wall., Hyptis, Jacq., Pogostemon, Desf., Mentha

L. and Salvia L. Where as, Orthosiphon Benth., exhibit both inter and infra

specific variations. Inter specific variations alone has been a characteristic

feature shown by Plectranthus L'Herit., Anisomeles R.Br. and Leucas R.Br. The

presence of such widely different series of chromosome numbers in the species

of even the same genus and in genera placed under different tribes and

subtribes, indicate that the different chromosome numbers may be derived one

from the other.

The basic chromosome numbers are found to be widely varied in South Indian

Lamiaceae. This great variability in the number of chromosomes at the basic

level could possibly be the result of aneuploidy at generic level. Out of the

fifteen genera investigated, three (Ocimum L., Orthosiphon Benth. and Coleus

Lour.) have been found to be polybasic, five (Plectranthus L' Herit.,

Anisochilus Wall., Hyptis Jacq., Salvia L. and Anisomeles R.Br.) to be dibasic

and three (Pogostemon Desf. Menthn L. and Leucas R.Br.) to be monobasic. As

the following genera, viz. Acrocephalus Benth., Eusteralis Rafin., Leonotis

R.Br. and Teucrium L. are represented by only one of their species in the

present study, it would not be appropriate to classify them as monobasic. Both

primary as well as secondary base numbers are found to be involved in the

evolution of the fiftyfour taxa investigated. The primary base numbers (Xl)

range from 6 to 9 and secondary base numbers (X2) from 10 to 13 and 15 to 17.

The basic chromosome number X2 a 12 is prevalent in majority of the members

(24.07%). This is followed by the base numbers X2 = 11, X1 = 7, X1 = 8 and X2

= 16 with percentages 16.67, 14.81, 12.96 and 11.11 respectively. Thus it is

likely that protopolyploidy plays an active role in the evolution.

The family Lamiaceae is characterized by a relatively high frequency of

polyploidy. Studies conducted on fifty four taxa reveals the dominance of

polyploids (62.96%) over diploids in South Indian Lamiaceae. Out of these,

57.41% were found to be tetraploids followed by hexaploids (3.7%) and

octaploids (1.85%). The present study also reveals that the herbaceous

elements predominate the shrubby elements of the family. Correlating

polyploidy with life form and growth habit, it is clear that the highest

frequency of polyploids occur in perennial herbs (79.41%) lowest in annual

herbs and undershrubs (2.94% each) and intermediate in perennial undershrubs

(14.71%). The various genera which show predominance of polyploidy include

Ocimum L., Orthosiphon Benth., Coleus Lour., Hyptis Jacq., Pogostemon Desf.,

Mentha L., Salvia L., Leonotis R. Br. and Teucrium L. The role of both

polyploidy and aneuploidy in the mechanism of speciation is obvious in South

Indian Lamiaceae. *

The general feature noted in the family Lamiaceae is the wide range of

chromosomes with graded symmetrical karyotypes. However the chromosome

complements in South Indian Lamiaceae members differ in minute karyotypic

details. With regard to gross morphology chromosomes are nearly submetacentric

in nature. The chromosomes were found to range from 4 . 2 p to 1.0 ,u in length.

The chromosomes with secondary constriction range from two to six in number.

The average chromosome length varies from 2 . 7 4 ~ to 1.12 p. The total

chromatin length shows a very wide variation with 2 4 . 4 ~ being the minimum and

154.8 p being the maximum value. The disparity index values were found to

range between 22.22 and 5 0 . The coefficient of variation ranges from 12.53 to

30 .00 and total centromeric index (TF%) from 35 .83 to 45 .07 . Thus the various

micromorphological details of the karyotype like, differences in absolute

chromosome size, differences in the position of centromere, differences in

total chromatin length, differences in karyotype formula and differences in

the number as well as position of satellites vary from cytotype to cytotype.

These variations found in the karyotypic parameters suggest that South Indian

Lamiaceae members are characterized by symmetrical to slightly asymmetrical

karyotypes. The karyomorphological diversity found in the family shows that

the group is still undergoing active speciation.

Meiotic studies have been conducted on twenty four taxa which frequently

blossom in the experimental garden. Out of these, nineteen taxa exhibited

normal meiosis and five taxa, abnormal meiosis. Interestingly thirteen out of

the nineteen members with normal meiosis are polyploids. Multivalent

associations are found to be invariably absent in the taxa examined. The

absence of multivalent formations owe to the small size of chromosomes. Thus

it is clear that well differentiated genomes are involved in the origin of

these polyploids. It has also been found that irrespective of the degree of

polyploidy, chromosomes always exhibit bivalent formation.Polyploid meiosis

characterized by bivalent formation point towards allopolyploidy. However

autopolyploids on long standing behaves like allopolyploids. The major

aberrations encountered in members which exhibit abnormal meiosis are the

occurrence of unpaired chromosomes at metaphase I, irregular segregation

during anaphase I and the disposition of partial or loose pairing. These

abnormalities reveal the meiotic instability leading to polyploidy in many

taxa of the family.

The presence of a wide range of chromosome numbers, numerical variations and

structural changes of chromosomes found in many genera mark the significant

role that both aneuploidy and polyploidy have played in the evolution of

various taxa of the family at the generic and species level. The variability

in base numbers might have been resulted through proto autoploidy,

amphiploidy, ascending and descending dysploidy. Both mitotic and meiotic

aberrations have played a major role in the evolutionary diversification of

the family. Induviduals with same chromosome number but with differences in

karyomorphological details reflect the ongoing evolutionary processes at

microlevel. It has also been found that in South Indian Lamiaceae,

Robertsonian changes or mutations might have also played a major role in the

evolution of karyotype.

chromosome studies in lamiaceae -...

Documents