Indian J. Applied & Pure Bio. Vol. 29(2), 291-301 (2014).

Inter tribal variations of fruit structure among four tribes of the family Compositae

Bidyut Kumar Jana* and Sobhan Kr. Mukherjee

Taxonomy and Biosystematics Laboratory, Department of Botany, University of Kalyani, Kalyani, Nadia, West Bengal (India) Email: *[email protected], [email protected]

Abstract

Variations of fruit structure of six species (Aster albanicus, Carpesium cernuum, Helianthus debilis, Leptorhynchos elongates, Podotheca angustifolia and Zinnia haageana), under four tribes- Astereae, Heliantheae, and Gnaphaleae of the family Compositae have been studied to observe the detailed morpho-anatomical features, among them. In all the studied fruits, there are great variations of morpho-anatomical features. Morphologically, special attention has been taken to the shape, size, colour, pappus type, stylopodia, carpopodia of the fruits. Among the studied cypselas, in case of Carpesium cernuum and Helianthus debilis, pappus bristles are absent. In the cypsela of Zinnia haageana, insteade of pappus bristle, awn like structure is present. In the cypsela of Carpesium cernuum, surface is glabrous than the remaining five studied cypselas, where the surface is pubescent. Morphologically, phytomelanin layer is exist in the cypselar wall of Helianthus debilis and Zinnia haageana. As, the two species are belongs to the tribe Heliantheae and phytomelanin layer is an important characteristic features of the tribe Heliantheae. Anatomically, mesocarpic region is well studied than another region of the pericarp due to the greait cellular variations in mesocarpic region. In the cypsela of Aster albanicus, Leptorhynchos elongates and Zinnia haageana, vellicular cavity is exists in the mesocarpic region. In the cypsela of Carpesium cernuum, secretary cavity is present in the mesocarpic region. Among the studied cypselas, testal layer is also interesting. In the cypsela of Leptorhynchos elongates and Podotheca angustifolia, testal layer is made up of U-shaped parenchyma cells. The orientation of cotyledons in relation to the cypselar wall is taxonomically significant.The number of resin ducts in each cotyledon is greatly variable.

Key words: Variations of fruit structure; four tribes; Compositae (292)

orientation with the surface, made up of body The family Compositae is one of the and basal cells. Surface containing 6 ribs, largest and highly evolved family among the alternating with furrows. Furrows wider than dicotyledons, consisting of 43 tribes, 1600- the ribs. The tip portion of body cell with 5 1700 genera and 24000 species . In India, there bifercation, arranged in different plane. At the are about 177 genera and 1,052 species upper portion of cypsela, stylopodium present; belonging to this family17 and which is very inconspicuous, fully immersed into the nectary. interesting to the taxonomist, due to the great Pappus homomorphic, represented by 15-20, diversity of habit, habitat, morphology and yellowish, unequal, serrulate-setose type of histology of vegetative and reproductive pappus bristles. At the basal region of cypsela, structures2. The members of the family carpopodium present; narrow than the base, Compositae are easily recognized by some symmetric. Carpopodial cells with thick-walled, characteristic features, which are pseudoanthial large, rectangular-square, arranged in 3 rows. heads with a specialized type of pollen presentation mechanism and pappus structure14, Table-1. Showing the number of studied 4 species and their sources nature of fruit and their particular array of Name of studied taxa Sources chemical weapons6. In spite of the presence 1. Aster albanicus Botanic Garden of of these aforesaid features of this family, in Degen. the University of the present study, special attention has been Copenhagen, Danmark. taken to the morpho-anatomical features of 211S2000-0862*A fruits of seven species of this family.There 2. Carpesium Botanischer Garten is a requirement of detailed information about cernuum L. der Universitat Zurich, the cypselar features for the all the tribes of Zollikerstrasse 107, . In this connection, the present CH-8008, Zurich, study has been taken. Switzerland. XXOZ- 20051457 Dry, mature, identified, disease free 3. Helianthus debilis North Central fruits were procured from the different foreign Nutt. Regional herbaria of the world, which are given in the Introduction Station, Ames. table below. 4. Leptorhynchos Herbarium CBG, elongates DC. CBG- 8906276 Aster albanicus : 5. Podotheca Botanic Garden of Morphology (Fig. 1 A-E) angustifolia Adelaide, South (Labill.) Less. Australia. Cypsela homomorphic, 8 mm x 1 mm 6. Zinnia haageana Botanic Garden of including pappus, 4 mm x 1 mm excluding Regel the University of pappus, yellow brown, slightly curved, upper Copenhagen, Danmark. part truncate whereas lower part tapered, 497 E2493-0005*AG ellipsoidal in cross sectional configuration. Cypselas were processed following the Surface pubescent. Surface hair ascending in methodology of Jana and Mukherjee7. (293)

Anatomy (Fig. 3 A-B) : Anatomy: (Fig. 3 C-D) :

Cypsela elliptic in cross-sectional Cypsela round to ellipsoidal in cross configuration. Ribs present; 6 in number, sectional configuration. Ribs present, 15 in conspicuous. Cypselar wall 0.04 mm and 0.01 number, conspicuous. Cypselar wall 0.03 mm mm thick in ribs and furrow region respectively. and 0.02 mm wide at ribs and furrow region Pericarp thick and devided in to epicarp and respectively. Pericarp thick, differentiated into mesocarp.Epicarp uni-seriate, made up of epicarp and mesocarp. Epicarp uni-seriate, thick-walled, rectangular-quadrangular, paren- made up of thin walled, rectangular, horizontally chyma cells, provided with cuticle. Internal to placed, parenchyma cells, provided with cuticle. the epicarp, mesocarp present. Outer region Internal to the epicarpic region, mesocarp of mesocarp made up of dark colour, pigment present. Outer region of mesocarp made up containing tissue region. Inner region of of thick walled, pentangular-hexagonal, mesocarp made up of penta-hexagonal, compactely arranged, sclerenchyma cells, sclerenchyma cells, containing vascular trace. containing secretary cavity just below the furrow Internal to the mesocarpic region, vellicular region. In the ribs region, this cavity absent. cavity present. Testa attached with cypselar wall, Inner region of mesocarp made up of, dark approximately 0.008 mm thick, made up of coloured, pigment containing, sclerenchyma rectangular, thick-walled, horyzontally placed, tissue region. Tesata attached with cypselar parenchyma cells, uni-seriately arranged. wall, approximately 0.01 mm thick, differen- Endosperm layer not clearly observed. Mature tiated into outer and inner testal region. Outer embryo occupies a major part of cypselas. testal layer made up of elongated, thick-walled, Cotyledons 8 in number, arranged at right angle horizontally placed, parenchyma cells whereas to the axis of cypselas, containing 8 resin ducts inner testal layer made up of crusted layer of (4 ducts in each cotyledon). parenchyma cells. Endosperm persists in mature cypselas, made up of, thick walled, Carpesium cernuum : horizontally placed, parenchyma cells, Morphology: (Fig. 1 F-J) : uniseriately arranged. Mature embryo occupies a major part of cypselas. Cotyledons 2 in number, Cypsela homomorphic, 2 mm x 0.5 arranged oblique to the axis of cypselas, mm, light brown, slightly curved, upper part containing 6 resin ducts (3 ducts in each blunted whereas lower part slightly tapered, cotyledon). round to ellipsoidal in cross sectional confi- guration. Surface glabrous, containing 15 ribs, Helianthus debilis : alternating with furrows. Furrows wider than Morphology (Fig. 1 L-O) : the ribs. Pappus absent. Stylopodium inconspicuously developed. At the basal region Cypsela heteromorphic. Ray cypsela of cypsela, stylopodium present; narrow than 8 mm x 3 mm while disk cypsela 8 mm x 2.5 the base, symmetric, triangular. Carpopodial mm. Ray cypsela white brown, straight, upper cells with thick walled, rectangular, horizontally part truncate whereas lower part blunted. Disk placed, uniseriately arranged. cypsela black brown, straight, upper part truncate (294) whereas lower part slightly tapered. Surface layer not clearly observed. slightly pubescent, containing 12 ribs, alternating with furrows. Furrows wider than the ribs. Leptorhynchos elongates : Surface hair made up of body and basal cells. Morphology (Fig. 1 P-T) : The tip portion of body cells with bifurcation Cypsela homomorphic, 5 mm x 0.5 and arranged in different plane. At the upper mm including pappus, 3 mm x 0.5 mm portion of cypsela, stylopodium present; excluding pappus, linear, slightly curved, rounded, partially immersed into the nectar. truncate at both ends. Surface pubescent. Within the surface, phytomelanin layer present. Surface hair inclined in orientation with the Pappus absent. At the basal region of cypsela, surface, made up of body and basal cells. Carpopodium present, rounded, narrow than Surface containing 10- 12 ribs, inconspicuous, the base. alternating with furrow. Furrows wider than the ribs. Stylopodium inconspicuously Anatomy: (Fig. 3 E) : developed, fully immersed into the nectary. At the upper portion of cypsela, pappus Cypsela elliptic in cross sectional present; homomorphic, represented by 25-30, configuration. Ribs present; 12 in number, unequal, serrulate- setose type of pappus inconspicuous. Cypselar wall, 0.08 mm and bristles, white, arranged in a single circle. At 0.07 mm wide at ribs and furrow region the basal region of cypsela, Carpopodium respectively. Pericarp thick, differentiated into present, homogenous, narrow than the base, irregular, ring like. Carpopodial cells with thick epicarp and mesocarp. Epicarp uniseriate, walled, elongated-square, arranged in 3 rows. made up of thick walled, horizontally placed, parenchyma cells. Internal to the epicarp, Anatomy (Fig. 4 H-I) : mesocarp present. Outer region of mesocarp made up of dark coloured, pigment containing Cypsela, elliptic in cross sectional tissue region. Internal to this region, phytome- configuration. Ribs present, 10-12 in number, lanin layer present, continuously arranged. inconspicuous. Cypselar wall, 0.01 mm and Internal to the phytomelanin layer, made up 0.008 mm wide at ribs and furrow region of thick-walled, penta-hexagonal, compactely respectively. Pericarp thick, differentiated into arranged, sclerenchyma cells, containing epicarp and mesocarp. Epicarp uniseriate, made pigment containing tissue zone just below the up of, thick-walled, elongated, horizontally placed, parenchyma cells, provided with cuticle. furrow region. Testa attached with cypselar Internal to the epicarpic region, mesocarp wall, approximately 0.01 mm thick, devided present, made up of thin-walled, elongated, into outer and inner testal region. Outer testal parenchyma cells, arranged in 2-3 rows. Testa layer made up of, thick-walled, uniseriately attached with cypselar wall, approximately arranged, parenchyma cells. Inner testal layer 0.01 thick, made up of thick-walled, U- shape, also parenchymatous but comparatively parenchyma cells, uniseriately arranged. thinner than the outer testal layer. Endosperm Endosperm persists in mature cypselas, (295) biseriate. Outer layer made up of thick-walled, up of thick-walled, vertically placed, palisade elongated, compactly arranged, parenchyma like parenchyma cells. Inner testal layer made cells whereas inner layer made up of compa- up of crusted parenchyma cells. Endosperm ratively thin-walled, elongated, parenchyma persists in mature cypselas, made up of thick- cells. Internal to the endosperm layer, just walled, horizontally placed, paren chyma cells, below the ribs, vellicular cavity present. Mature uni-seriately arranged. Mature embryo embryo occupies a major part cypselas. occupies a major part of cypselas. Cotyledons Cotyledons 2 in number, arranged at right 2 in number, arranged oblique to the axis of angle to the axis of cypselas, containing 10 cypselas, containing 6 resin ducts (3 ducts resin ducts (5 ducts in each cotyledon). in each cotyledon).

Podotheca angustifolia : Zinnia haageana : Morphology: (Fig. 1 U-W) : Morphology: (Fig. 2 X-Z) :

Cypsela homomorphic, 9 mm x 1 mm Cypsela heteromorphic. Ray cypsela including pappus, 3 mm x 1 mm excluding 7.5 mm x 2.5 mm including awn, 5 mm x pappus, brownish, obovate, straight, upper part 2.5 mm excluding awn. Disk cypsela 7 mm truncate whereas lower part tapered, rounded x 1.5 mm including awn, 4.5 mm x 1.5 mm in cross sectional configuration. Surface pubescent, excluding awn. Ray cypsela yellow brown surface hair made up of body and basal cells, in colour whereas disk cypsela black brown ascending in orientation with the surface. With in colour. In both the cypselas, upper part in the surface, ribs absent. At the upper portion truncate whereas lower part tapered, straight. of cypsela stylopodium present, inconspicuous, Surface slightly pubescent. In case of ray fully immersed in to the nectary. Pappus cypsela, surface hair situated in marginal represented by, 5-7, unequal, yellowish, region whereas in case of disk cypsela, surface plumose type of pappus bristles, arranged in hair present in the entire body of cypsela. a single whorl. At the basal region of cypsela, Surface hair made up of body and basal cells. Carpopodium present, narrow than the base, The tip portion of body cell with bifurcation. irregular, reing like. Carpopodial cells not Pappus represented by a single awn, yellow clearly distinguish from the remaining part of brown in colour. Stylopodium prominent, cypselas, i.e. presence of pseudocarpopodium. blunted, partially immersed into the nectary. At the basal region of cypsela, Carpopodium Anatomy: (Fig. 4 J-K) : present, narrow than the base, biconvex. Carpopodial c ells not clearly distinct from the Cypsela rounded in cross sectional remaining part of cypselas. configuration. Pericarp 0.005 mm thick, represented by only epicarp. Epicarp, Anatomy: (Fig. 4 F-G): uniseriate, made up of thin-walled, horizontally placed, parenchyma cells. Mesocarp absent. Cypsela elliptic in cross sectional Testa attached with cypselar wall, devided into configuration. Ribs present, 2 in number, outer and inner testal region. Outer layer made conspicuous. Pericarp thick, differentiated into (296) epicarp and mesocarp.Epicarp uniseriarte, region near the lateral lobe region. Internal to made up of thin walled, rectangular, compactely the mesocarpic region, phytomelanin layer arranged, parenchyma cells. Internal to the present, discontinuously arranged. Testa, epicarp, mesocarp present, made up of thick- parenchymatous, biseriate,. Endosperm walled, compactely arranged, sclerenchyma parenchymatous, uniseriate. cells. Vellicular cavity present in mesocarpic

1 mm. A, E,F,G,H,I,L,M,P,Q,R,U,V, 0.1 mm. B,J. 0.05 mm. B,C,D,J,K,N,O,S,T,W. Fig. 1. Morphology of studied cypselas A-F- Aster albanicus: A-Cypsela, B-Surface, showing surface hair, C-Upper part of pappus bristle, D- Carpopodial cells, E-Lower part of cypsela, F-Upper part of cypsela; G-K- Carpesium cernuum: G- Cypsela, H-Basal part of cypsela, I-Upper part of cypsela, J- Surface of cypsela, K- Carpopodial cells; L- O- Helianthus debilis: L-Ray cypsela, M-Disk cypsela, N-Surface, O-Surface hair; P-T- Leptorhynchos elongates: P-Cypsela, Q-Lower part of cypsela, R-Upper part of cypsela, S-Surface showing, surface hair, T-Part of pappus bristles; U-W- Podotheca angustifolia: U-Cypsela, V-Lower part of cypsela, W- Surface hair. (297)

Fig. 2: Morphology of studied cypsela. 1 mm. X,Y 0.05 mm Z. X-Z- Zinnia haageana: X-Ray cypsela, Y-Disk cypsela, Z- Surface hair

Fig. 3: Anatomy of studied cypselas. 0.1 mm. A,C. 0.05 mm B,D,E. A-B- Aster albanicus: A-Diagramatic view, B-Cellular view; C-D- Carpesium cernuum: C-Diagramatic view, D-Cellular view; E- Helianthus debilis (Cellular view) (298)

Fig. 4: Anatomy of studied cypselas 0.1 mm.F,H,J. 0.05 mm G,I,K. F-G- Zinnia haageana: F-Diagramatic view, G-Cellular view; H-I- Leptorhynchos elongates: H-Diagramatic view, I-Cellular view; J-K- Podotheca angustifolia: J-Diagramatic view, K-Cellular view.

Six species, belongs to the four tribes Cypselar morphology : of compositae (Astereae-Aster albanicus, Inuleae- Carpesium cernuum, Heliantheae- Among the studied cypselas, Aster Helianthus debilis, Zinnia haageana, albanicus belongs to the tribe Astereae. Cypsela is homomorphic, 8 mm x 1 mm including Gnaphaleae- Leptorhynchos elongates, pappus, 4 mm x 1 mm excluding pappus. Podotheca angustifolia) have been studied Homomorphism is also present in another to observe the morpho-anatomical variation species (Solldago virgaurea, Solidago pattern among them and as well as among the Canadensis) of this tribe8. Surface is pubescent different tribes, as the studied six cypselas are type. Surface hair is distributed in the furrow belongs to the four different tribes. The species region. At the upper portion of cypsela, pappus examined here demonstrated mature ovarian is present, numerous, free, persistent, structure (Cypsela), on the basis of morpho- represented by serrulate-setose type of anatomical study. pappus bristles. Not only bristilate pappus, (299) scaly type of pappus is found to be present in angustifolia. case of Aster alpinus, of this tribe7. Stylopo- dium is inconspicuous, fully immersed into the Among the studied cypselas, Carpesium nectary. At the basal region of cypsela, cernuum, is belonging to the tribe Inuleae. In carpopodium is present, which is narrow than the tribe Inuleae, pappus is usually absent. the base, symmetric. Carpopodial cells are Though, pappus structure has been observed thick-walled, large, rectangular-square, in some other species (Inula ensifolia, I. 9 arranged in 3 rows. Carpopodium may be Britannica, I. helenium) of this tribe . Surface symmetric or asymmetric type. Within the features of cypselas are diagnostic for studied cypsela in Aster albanicus, carpo- characterization of taxa.Surface is glabrous, podium is symmetric type. The cellular in case of Carpesium cernuum. Multicellular, arrangement in carpopodium is also variable. conical glandular structure is found at the Hare, carpopodial cells are arranged in 3 rows. apical and basal region of cypsela. This observation is closely allied with the Among the studied cypselas, Helianthus observation of Mukherjee and Sarkar. At the debilis and Zinnia haageana belong to the basal region of cypsela, carpopodium is tribe Heliantheae. In both the studied cypselas, present. In the studied cypsela, carpopodial surface is slightly pubescent. At the upper cells are uni-seriately arranged and horizontally portion of the cypsela of Zinnia haageana, placed. Carpopodium may be present or awn like structure is present, whereas in the absent in the cypsela. In the studied cypsela, cypsela of Helianthus debilis, awn like structure carpopodial cells are present. Mukherjee & is absent. Morphologically, phytomelanin Sarkar12, have pointed out that the absence deposition is seen in cypselarv wall during of carpopodium in Carpesium cernuum. morphological study. Presence of phytomelanin layer in the cypselar wall, is an important Cypselar anatomy: features of the tribe Heliantheae, though in case of Helianthus annus, phytomelanin layer Anatomically, all the studied cypselas is absent11. Pandey16, has reported that are round to elliptic in cross-sectional confi- phytomelanin layer is secreted by the glandular guration. Epicarp is uni-seriate, made up of activity of hypodermal cells. In case of both horizontal (Carpesium cernuum, Helianthus the cypselas, pappus is absent. Pappus structure debilis, Leptorhynchos elongates, Podotheca is also found to be present in some other angustifolia, Zinnia haageana) or radially species (Bidens cernua, Bidens pilosa, Tithonia (Aster albanicus) placed, parenchyma cells. diversifolia etc) of the tribe Heliantheae13. Internal to the epicarpic region, mesocarp In both the cypselas, stylopodia are prominently present (except in Podotheca angustifolia, developed. where mesocarp is absent). Mesocarpic region may be continuous (Carpesium cernuum, Among the studied cypselas, the tribe Helianthus debilis, Leptorhynchos elongates, Gnaphaleae has 2 species, which are- Podotheca angustifolia, Zinnia haageana) Leptorhynchos elongates and Podotheca or discontinuous (Aster albanicus). In the (300) cypsela of Zinnia haageana and - Helianthus (Gnaphaleae), and Carpesium cernuum debilis (Tribe-Heliantheae), outer region of (Inuleae), testal layer is bi-seriately arranged mesocarp, phytomelanin layer is present. In but in remaining two studied cypselas the cypsela of Helianthus debilis, phytomelanin [Leptorhynchos elongates (Gnaphaleae) and layer is continuously developed, than the Aster albanicus (Astereae)], testal layer is cypsela of Zinnia haageana, where, uni-seriately arranged. Testal layer is greatly phytomelanin layer is discontinuously variable in structure. In the cypsela of Podotheca developed. Presence of phytomelanin layer is angustifolia, out of two testal layers, outer a very important character in case of the tribe layer is made up of U-shaped, parenchyma Heliantheae. Phytomelanin is a unique type of cells and inner layer is made up of crusted resinous substance, which is usually present layer of parenchyma cells. In the cypsela of in the members of the tribe Heliantheae, Leptorhynchos elongates, testal layer is uni- Helenieae etc. 11. In respect to this features, seriately arranged and is made up of U-shaped, Heliantheae is closely related to the tribe parenchyma cells. In the cypsela of Carpesium Eupatorieae, but apert from other some tribes, cernuum, outer testal layer is parenchymatous such as- Astereae, Inuleae, Anthemideae, and inner testal layer is made up of crusted Arctotideae3. In the cypsela of Zinnia layer of cells. In the cypsela of Aster albanicus, haageana, Leptorhynchos elongates and Zinnia haageana, Helianthus debilis testal Aster albanicus, vellicular cavity is present layer is made up of only parenchyma cells. In in the pericarpic region. In remaining three the cypsela of Aster albanicus and Helianthus studied cypselas (Podotheca angustifolia, debilis, endosperm layer is not clearly Helianthus debilis, Carpesium cernuum ), observed.The information about the endosperm 15 vellicular cavity is absent. The function of layer has been reported by Pandey and Sing . this cavity is still unknown. In the cypsela of Within the cotyledons, resin ducts are also Carpesium cernuum, secretary cavity is variable. The presence of fixed number of resin present in the mesocarpic region. This cavity ducts in each cotyledon has been reported by 15 is absent in remaining five studied cypselas. Pandey and Sing . In Carpesium cernuum (Inuleae), crystal From the above discussion, it can be formation is absent in pericarpic region. concluded that cypselar features are not only Though, crystal formation has been reported variable among the same tribe but also among by Mukherjee and Sarkar12, in some other the different tribes of Asteraceae. species (Buphthalmum, Inula etc) of the tribe Inuleae. Crystal formation in the tribe-Inuleae, We are thankful to Prof. (Dr.) Hana 1 is also reported by Anderberg . Testal layer is Vilhelm Hansen, Curator, University Botanic also variable. In the cypsela of Zinnia Garden of Copenhagen (DK), Denmark; Dr. haageana (Heliantheae), Helianthus debilis Peter Enz, Curator, Botanischer Garten der (Heliantheae), Podotheca angustifolia Universitat Zurich (Z), Switzerland; Director, (301)

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