Exomorphology and Internal Structure of Cypselas of Seven Species of the Tribe Cardueae (Compositae)

J. Indian bot. Soc. ISSN - 0019 - 4468 Vol. 93 (1 & 2) 2014 : 50- 5059 EXOMORPHOLOGY AND INTERNAL STRUCTURE OF CYPSELAS OF SEVEN SPECIES OF THE TRIBE CARDUEAE (COMPOSITAE)

BIDYUT KUMAR JANA AND SOBHAN KR. MUKHERJEE Taxonomy and Biosystematics Laboratory, Department of Botany, University of Kalyani, Kalyani-741235, Nadia, West Bengal, India.

Cypselar morpho-anatomy is very helpful for the separation of taxa, when flowering condition is unavailable i.e. fruiting condition is available. For this purpose, cypselas of seven species belonging to five genera (Arctium lappa, Cirsium arvense , Cirsium candelabrum, Cirsium heleniodes, Mantisalca salmantica, Saussurea fastuosa and Xeranthemum annuum) of the tribe Cardueae have been investigated. The important morphological features are the presence of surface structure, pappus bristles, stylopodia, carpopodia with cellular orientation etc. Except in the cypselas of Mantisalca salmantica and Arctium lappa, the remaining 5 studied cypselas are homomorphic. In the cypsela of Xeranthemum annuum, surface is pubescent , whereas in remaining 6 studied cypselas, surface is rough and glabrous. In the cypsela of Arctium lappa, Cirsium arvense, Cirsium heleniodes and Cirsium candelabrum, stylopodia are prominent and enlarge than the remaining studied cypselas where they are inconspicuous. Carpopodia are the basal, meristematic zone of cypselas. In the cypsela of Cirsium arvense, Arctium lappa, Mantisalca salmantica and Cirsium candelabrum, pseudocarpopodia are present, whereas in remaining studied cypselas carpopodial cells are arranged from 1 layer (Cirsium heleniodes, Xeranthemum annuum ) to 2 layers (Saussurea fastuosa). Pappus structures help in the dispersal of cypselas. In the cypsela of Saussurea fastuosa, Cirsium heleniodes and Cirsium candelabrum, plumose type of pappus bristles are present , whereas in the cypsela of Man- tisalca salmantica barbellate pappus bristles are present. In the cypsela of Xeranthemum annum, scaly pappus is present. Among the studied cypselas, in Arctium lappa and Cirsium arvense, pappus are absent. Anatomically, the mesocarpic region is very interesting than other layer of pericarp. In the cypsela of Saussurea fastuosa , Cirsium heleniodes and Xeranthemum annuum, mesocarpic regions are heterogeneously developed than the other studied cypselas where mesocarpic regions are homogenously developed. In the cypsela of Arctium lappa, crystal is present in the mesocarpic region but in six studied cypselas crystals are absent. Presence of secretary duct is also an important taxonomic character and which is present in the mesocarpic region of the cypsela of Mantisalca salmantica and Cirsium candelabrum. In remaining five studied cypselas, secretary ducts are absent. In the cypsela of Cirsium candelabrum, internal to the testal region vellicular cavity is present , whereas in the cypsela of Xeranthemum annuum, vellicular cavity is present within the mesocarpic region. In the cypsela of Saussurea fastuosa and Xeran- themum annuum, testal layers are made up of round-elongated, uni-seriately arranged, parenchyma cells , whereas in rest studied cypselas, testal layers are made up of vertically arranged, thick-walled, palisade parenchyma cells.

Key Words : Cypselar exomorphology and internal structures, Cardueae, Compositae

The broadly defined tribe Cardueae consists of of cypselas but also the internal characters are four subtribes ( Bentham, 1873), which are Echinop- also helpful for better taxonomic treatment. The sidinae, Carlinae, Carduinae and Centaureinae. objective of the present paper is to show the Bremer (1996) has recognized the tribe Cardueae cypselar features of this tribe in detail and to as subfamily Carduoideae. The exomorphology show the relationship on the basis of present ob- and internal structure of cypselas of this tribe has servation. a great importance for the better classification of taxa. Lavialle (1912) has studied the cypselar fea- MATERIALS AND METHODS tures of about 15 genera and 83 species of this Mature, dry, identified cypselas were collected tribe. He pointed out the importance of crystals from 4 Herbaria of the world and 1 specimen and secretary ducts which are present in the from Sikkim, by first author, which are given in mesocarpic region of cypselas of this tribe. Pres- the Table -1. ence of calcium oxalate crystals has been reported by Hanausek (1911) and Singh and Pandey For morphological observation, cypselas were sof- (1984) etc. in this tribe. Importance of cypselar tened by NaOH (2 % ) solution and after that exomorphology and internal structure of cypselas they were stained by using safranin-light green in this tribe has been reported by authors like combination (4% ) and finally observed under sim- Mukherjee (2000) Fourment and Rouzet (1957), ple dissecting microscope ( Meopta, Praha, 65301, Carlquist (1958), Chauhan (1972), Jana and Muk- made in Czechoslovakia) and stereo biocular dis- herjee (2012), Zarembo and Boyko (2008). Accord- secting microscope (Olympus, made in Tokyo, ing to Roth (1977) not only the exomorphology Japan). For anatomical observation, hand sections Received on August 24, 2013 Accepted on February 6, 2014

BIDYUT KUMAR JANA AND SOBHAN KR. MUKHERJEE 51

Table-1: List of taxa studied and their sources. Sl. No. Name of Taxa Source 1 Arctium lappa L. Humboldt- Universitat zu Berlin. Institut fur Biologie, Spezielle Botanik u. Arboretum, Berlin, Germany. Collection Number- 704.

2 Cirsium arvense (L.) Scop. Conservatoire Et Jardin Botaniques Geneve- Geneva, Switzerland. Col- lection Number- 107. 3 Cirsium candelabrum Griseb. Botanic Garden and Museum of the University of Copenhagen ( Natural History Museum of Denmark). Collection Number- 268 S2000-0876 A W Kit Tan 24698. 4 Cirsium heleniodes (L.) Hill Botanischer Garten der Universitat Zurich. Collection Number- CHOZ -20031567 5 Mantisalca salmantica (L.) Botanic Garden and Museum of the University of Copenhagen ( Natural Brig. Et casill. History Museum of Denmark). Collection Number- 389 E 2867-B001 A G 6 Saussurea fastuosa (Decne.) Sikkim. Collection Number- 52. Sch. Bip. 7 Xeranthemum annuum L. Botanischer Garten der Universitat Zurich. Collection Number- XXOZ 20040897 were done by sharp razor blade from the middle spectively. Pericarp thick, differentiated into part of cypselas and finally the good sections epicarp, mesocarp and endocarp. Epicarp uni-seriate, were taken for anatomical observation under com- thin - walled, triangular – quadrangular, compactely pound light microscope ( Metzer). arranged, parenchyma cells provided with cuticle. Internal to the epicarp, mesocarp present; made up of OBSERVATIONS AND DISCUSSION thin - walled, rounded, compactely arranged, parenchyma cells containing crystals. Internal to the meso- Arctium lappa Morphology ( Fig 1 A-E ) carp, endocarp present; thin walled, parenchymatous, Cypsela heteromorphic. Ray cypsela 5 mm x 2 mm, uniseriately arranged. Testa attached with cypselar creamy brown, blotched, narrow oblanceolate, wall, approximately 0.04 mm thick, made up of slightly curved, upper part truncate whereas lower thick-walled, vertically placed, palisade parenchyma part narrow. Disk cypsela 6 mm x 2 mm, offwhite, cells. Endosperm persists in mature cypsela, uniseri- blotched, straight, upper part truncate, whereas lower ate, thick-walled, parenchymatous. Mature embryo part tapered. Ellipsoidal in cross sectional configura- occupies a major part of cypsela; cotyledons 2 in tion. Surface rough and glabrous containing 14-18 number, arranged oblique to the axis of cypsela, con- ribs, alternating with furrows. Furrows wider than the taining 6 resin ducts (3 ducts in each cotyledon ). ribs. Pappus absent. At the upper part of cypsela, sty- Cirsium arvense lopodium present; enlarge, cylindric. At the basal re- Morphology ( Fig 1 F-H ) gion of cypsela, carpopodium present, narrower than Cypsela homomorphic, 3 mm x 0.5 mm, black brown, the base, quadrangular. Carpopodial cells are not obovate, slightly curved, upper part truncate whereas clearly distinct from the remaining part of cypselas. lower part tapered, ellipsoidal in cross sectional con- Anatomy ( Fig 2 A ) figuration. Surface glabrous, ribs absent. Pappus absent. At the upper portion of cypsela, stylopodium Cypsela elliptic in cross section. Ribs present; 14-18 present, enlarge, dome shape, free. At the basal re- in number, conspicuous. Cypselar wall 0.04 mm gion of cypsela, carpopodium present, narrow than and 0.06 mm wide at ribs and furrow region re- the base, irregular ring like. Carpopodial cells not

EXOMORPHOLOGY AND INTERNAL……. 52

Figure 1. A-V. Cypselar morphology of studied species. Figs. A-E. Arctium lappa: A- Ray cypsela, B- Upper part of cypsela, C- Lower part of cypsela, D- Carpopodial cells, E- Disk cypsela; F-H- Cirsium arvanse: F-Cypsela, G- Upper part of cypsela, H- Lower part of cypsela; I-K: Circium candelabrum : I- Cypsela, J- Upper part of cypsela, K- Upper part of pappus bristles; L-N: Cirsium heleniodes: L- Cypsela, M-Upper part of cypsela, N- Carpopodial cells; O-Q- Montisalea salmantica: O- Ray cypsela, P- Disk cypsela, Q- Upper part of pappus bristles; R-Cypsela of Saussurea fastuosa; S-V- Xeranthemum annuum : S-Cypsela, T- Surface hair, U- Upper part of cypsela, V- Carpopodial cells.

BIDYUT KUMAR JANA AND SOBHAN KR. MUKHERJEE 53 clearly distinguish from the remaining part of cypsela cells, containing secretary duct. At the basal region of i.e. pseudocarpopodium. secretary duct, vascular trace present. Testa attached with cypselar wall, approximately 0.029 mm thick, Anatomy ( Fig 2 B ) uniseriate, made up of thick walled, vertically placed, palisade parenchyma cells. Internal to the testal re- Cypsela elliptic in cross section. Ribs absent. Pericarp gion, vellicular cavity present. Endosperm persists in thin, on an average 0.02 mm, differentiated in to mature cypsela; biseriate. Outer layer made up of epicarp and mesocarp. Epicarp uniseriate, made up of thick walled, rectangular, parenchyma cells with vas- thin walled, compactely arranged, rectangular, paren- cular trace whereas inner layer made up of thick chyma cells, provided with cuticle. Internal to the walled, barrel shaped, compactely arranged, paren- epicarp, mesocarp present; homogenous, made up of chyma cells. Mature embryo occupies a major part of thick-walled, rounded, compactely arranged, paren- the cypsela; cotyledons two in number, arranged chyma cells. Testa attached with cypselar wall, ap- oblique to the axis of cypsela, containing 10 resin proximately 0.02 mm thick, made up of thick-walled, ducts ( 5 ducts in each cotyledon). elongated, vertically oriented, palisade parenchyma cells, uniseriately arranged. Endosperm persists in Cirsium heleniodes mature cypsela, biseriate, outer layer made up of Morphology ( Fig 1 L-N ) crusted layer of parenchyma cells whereas inner layer Cypsela homomorphic, 24 mm x 1 mm with pappus, made up of thick walled, barrel shape parenchyma 6 mm x 1 mm without pappus, light brown, oblanceo- cells. Mature embryo occupies a major part of the late, curved, margin entire, upper part truncate where cypsela; cotyledons two in number, arranged at right as lower part compressed, ellipsoidal in cross sec- angle to the axis of cypsela, containing 6 resin ducts tional configuration. Surface rough and glabrous con- ( 3 ducts in each cotyledon ). taining 6 ribs, alternating with furrow. Furrows wider than ribs. At the upper part of cypsela, stylopodium Cirsium candelabrum present, enlarge, cylindric. Pappus homomorphic, Morphology ( Fig 1 I-K ) represented by 18-22, unequally arranged, white, plu- Cypsela homomorphic, 4 mm x 1 mm, brownish, mose type of pappus bristles. At the basal region of oblong, straight, margin entire, upper part truncate cypsela, carpopodium present, narrow than the base, whereas lower part slightly tapered. Surface glabrous. biconvex, symmetric. Carpopodial cells with thick- Ribs absent. Stylopodium prominent, enlarge, dome walled, medium, not pitted, arranged in single row. shape. Pappus homomorphic, represented by 20-29, unequally arranged, plumose type of pappus bristles, Anatomy ( Fig 2 D ) brownish, arranged in single whorl. At the basal region of cypsela, carpopodium present, symmetric, Cypsela elliptic in cross section. Ribs present; 6 in biconvex. Carpopodial cells not clearly distinct from number, inconspicuous. Cypselar wall 0.19 mm and the remaining part of cypsela i.e. pseudocar- 0.14 mm wide at ribs and furrow region respectively. popodium. Pericarp thick, differentiated into two zones- epicarp and mesocarp. Epicarp uniseriate, made up of thin Anatomy ( Fig 2C ) walled, rectangular, compactely arranged, parenchyma cells, provided with cuticle. Internal to the Cypsela elliptic in cross sectional configuration. Ribs epicarp, mesocarp present, heterogenous, made up of absent. Pericarp thick, on an average 0.02 mm wide, thin walled, circular, compactely arranged paren- differentiated into two zones- epicarp and mesocarp. chyma cells and thickwalled, circular, compactely Epicarp uniseriate, made up of thin walled, cubical, arranged, sclerenchyma cells containing vascular parenchyma cells, provided with cuticle. Internal to trace. Testa attached with cypselar wall, approxi- the epicarp, mesocarp present; made up of thick mately 0.02 mm thick, made up of vertically placed, walled, rounded, compactely arranged, parenchyma thick-walled, palisade parenchyma cells, uniseriately

EXOMORPHOLOGY AND INTERNAL……. 54 arranged. Endosperm persists in mature cypselas, Saussurea fastuosa uniseriate, made up of thick walled, horizontally Morphology ( Fig 1 R) placed, parenchyma cells. Mature embryo occupies a major part of cypsela, cotyledons 2 in number, ar- Cypsela homomorphic, 15 mm x 1.5 mm including ranged oblique to the axis of cypsela, containing 6 pappus, 6 mm x 1.5 mm excluding pappus, black resin ducts ( 3 ducts in each cotyledon). brown, linear, straight, upper part truncate whereas lower part tapered. Irregular in cross sectional con- Mantisalca salmantica figuration. Surface rough and glabrous, containing 10 Morphology ( Fig 1 O-Q ) ribs, alternating with furrow. Furrows wider than the ribs. At the upper portion of cypsela, stylopodium Cypsela heteromorphic. Ray cypsela 5 mm x 1 mm present; inconspicuously arranged, fully immersed in including pappus, 3 mm x 1 mm excluding pappus, the nectar. Pappus represented by 10-17, unequally yellow brown, oblanceolate, straight, upper part trun- arranged, plumose type of pappus bristles, light yel- cate, whereas lower part slightly tapered with entire low in colour. At the basal region of cypsela, car- margin. Disk cypsela 3 mm x 0.5 mm, light yellow, popodium present, narrow than the base, irregular oblanceolate, straight, middle part slightly swollen, ring like. Carpopodial cells with thick walled, biseri- tapered at both ends. Ellipsoidal in cross sectional ately arranged. configuration. Surface glabrous.At the upper part of cypsela, stylopodium present, inconspicuously de- Anatomy ( Fig 2 F ) velop, fully immersed into the nectar. Pappus homo- Cypsela irregular in cross section. Ribs present, 10 in morphic, represented by 19-24, unequally arranged, number, conspicuous. Cypselar wall 0.05 mm and barbellate setose type of pappus bristles, white yel- 0.01 mm wide at rib and furrow region respectively. low, arranged in 3 rows. Carpopodium basal in posi- Pericarp thick, differentiated in to two zones-epicarp tion, symmetric, biconvex. Carpopodial cells not and mesocarp. Epicarp uniseriate, made up of thin clearly observed from the remaining part of cypsela walled, rectangular, compactely arranged, paren- i.e. pseudocarpopodia. chyma cells provided with cuticle. Internal to epicarp, Anatomy ( Fig 2 E ) mesocarp present; heterogenous, made up of thin walled, circular, compactely arranged, parenchyma Cypsela elliptic in cross section. Ribs absent. Pericarp cells and thick walled, circular, compactely arranged, thick, approximately 0.02 mm, differentiated into sclerenchyma cells. Testa attached with cypselar wall, three zones- epicarp, mesocarp and endocarp. Epicarp approximately 0.014 mm thick, made up of thick uniseriate, made up of thin walled, rectangular, com- walled, round to square, compactely arranged, paren- pactely arranged, parenchyma cells, provided with chyma cells, uniseriately arranged. Endosperm per- cuticle. Internal to the epicarp, mesocarp present, sists in mature cypsela, biseriate. Outer cells wide- made up of thick walled, rounded, compactely ar- barrel shaped and inner cells narrow-barrel shaped. ranged, parenchyma cells, with secretary duct. Inter- Cells of both the layers thich walled, parenchyma- nal to the mesocarpic region, endocarp present, paren- tous, compactely arranged. Mature embryo occupies a chymatous, uniseriately arranged. Testa attached with major part of the cypsela. Cotyledons 2 in number. cypselar wall, approximately 0.06 mm thick, made Cotyledonary resin ducts are not clearly distinguish. up of, vertically arranged, thickwalled, palisade parenchyma cells. Endosperm persists in mature cyp- Xeranthemum annuum sela, uniseriate, made up of thick walled, barrel Morphology ( Fig 1 S-V ) shaped, parenchyma cells. Mature embryo occupies a Cypsela homomorphic, 6 mm x 0.5 mm including major part of cypsela; cotyledons 2 in number, ar- pappus, 4 mm x 0.5 mm excluding pappus, black ranged oblique to the axis of cypsela, containing 16 brown, oblanceolate, straight, upper part truncate resin ducts ( 8 ducts in each cotyledon).

BIDYUT KUMAR JANA AND SOBHAN KR. MUKHERJEE 55

Figure 2. A-G. Cypselar anatomy of studied species. A. Arctium lappa, B . Cirsium arvanse , C. Circium candelabrum , D. Cirsium heleniodes E. Montisalea salmantica , F. Saussurea fastuosa , G. Xeranthemum annuum

List of abbreviations: Ep.- Epicarp, Me.- Mesocarp, T.- Testa, End- Endocarp, E- Endosperm, Pa- Parenchyma, Scl- Sclerenchyma, V.T.- Vascu- lar trace, V.C.- Vellicular cavity, S.D.- Secretary duct, Scb- Sclerenchyma bundle.

EXOMORPHOLOGY AND INTERNAL……. 56 whereas lower part tapered, entire margin. Ellipsoi- in Carlina and Echinops of this tribe. Except in dal in cross sectional configuration. Surface pubes- the cypsela of Xeranthemum annuum, the re- cent. Surface hair ascending-inclined in orientation maining studied cypselas are with rough and gla- with the surface, made up of body and basal cells. brous surface, whereas in the cypsela of Xeranthe- The tip portion of body cells with bifurcation and ar- mum annuum, surface is pubescent, containing ranged in different plain.Surface containing 16-18 twin hairs. Twin hair is also reported in some other ribs, alternating with furrows. Furrows wider than species ( Centaurea cyanus) of this tribe ( Mukher- ribs. The distance between two ribs 0.17 mm. At the jee 2000, Mukherjee and Nordenstam 2012). At the upper portion of cypsela, stylopodium present, incon- upper portion of cypselas, stylopodia are present. spicuous, fully immersed into the nectar. Pappus rep- Among the studied cypselas, in Xeranthemum resented by 4 scarious, subulate, apically pinnate annuum, Saussurea fastuosa and Mantisalca sal- scale, equal. At the basal region of cypsela, car- mantica stylopodia are inconspicuously developed popodium present, narrower than the base, asymmet- but in remaining 4 studied cypselas, stylopodia are ric, irregular ring like. Carpopodial cells with thick very prominent and enlarged. Mukherjee (2005) walled, rectangular, not pitted, arranged in single and Wetter (1983) have studied the stylopodial fea- row. tures in some Asteraceae and they have mentioned the different structures of stylopodia in Anatomy ( Fig 2 G ) Asteraceae. Present observation regarding the stylopodial features are clearly indicated with the stud- Cypsela elliptic in cros section. Ribs present; 16-18 in ies of Mukherjee (2005) and Wetter (1983). At the number, inconspicuous. Cypselar wall 0.09 mm and basal region of cypselas, carpopodia are present. 0.03 mm wide at ribs and furrow region respectively. Carpopodia are usually regular or irregular ring Pericarp thick, differentiated into two zones-epicarp like structure and there are usually symmetric or and mesocarp. Epicarp uniseriate, made up of round some time asymmetric in nature ( Cirsium ar- to ovoid, thin walled, parenchyma cells, provided vense, Cirsium canadelabrum, Mantisalca salman- with cuticle. Internal to the epicarp, mesocarp pre- tica). Diameter of carpopodia may be as wide as sent, made up of round to ovoid, thick walled, paren- the base of cypsela or narrower than the base of chyma cells and thick walled, pentangular, scleren- cypsela as in Arctium lappa, Cirsium arvense etc. chyma cells containing vascular bundle near the ribs Carpopodium is a meristematic zone with variable region. Within the mesocarpic region, vellicular cav- layers of cellular orientation. In the cypsela of ity present. Testa attached with cypselar wall, ap- Cirsium arvense, Arctium lappa, Mantisalca sal- proximately 0.008 mm thick, uniseriate, made up of, mantica and Cirsium candelabrum, pseudocarpopo- thick walled, cubical, parenchyma cells. Endosperm dia are present i.e. carpopodial cells are not dis- persists in mature cypsela, biseriate. Outer layer made tinguishable from other epicarpic cells of cyp- up of, thick walled, rectangular, compactely arranged, selas, whereas in remaining studied cypselas, car- parenchyma cells and inner layer made up of thick popodial cells are arranged from 1 layer (Cirsium walled, barrel shaped, parenchyma cells. Mature em- heleniodes, Xeranthemum annuum ) to 2 layers bryo occupies a major part of the cypsela; cotyledons (Saussurea fastuosa). Carpopodial structures of two in number, arranged at right angle to the axis of some genera ( Echinops, Centaurea ) of this tribe cypselar wall, containing 18 resin ducts ( 9 ducts in have been reported by Haque and Godward (1984). each cotyledon). Mukherjee and Nordenstam (2004), have indicated Cypselar Morphology that majority members of this tribe possess strongly asymmetric ring like carpopodia. In the Shape of cypselas are usually narrow oblanceolate cypsela of Saussurea fastuosa, Cirsium heleniodes – linear- narrow oblong-obovate. Cypsela with nar- and Cirsium candelabrum, plumose type of pappus row oblong is also reported by Mukherjee ( 2000) bristles are present, while the cypsela of Man-

BIDYUT KUMAR JANA AND SOBHAN KR. MUKHERJEE 57 tisalca salmantica pappus bristles are barbellate 1. Testal epidermal cells are palisade like in ap- type. Only in the cypsela of Xeranthemum an- pearance. e.g. Arctium lappa , Cirsium arvanse , num, scaly pappus has been observed. Among the Cirsium candelabrum , Cirsium heleniodes , Man- studied cypselas, Arctium lappa and Cirsium ar- tisalca salmantica and Saussurea fastuosa. vense lacks pappus bristles. Pappus is the main or- gan for the dispersal of cypselas. Dittrich ( 1977) 11. Testal epidermal cells are cuboidal paren- has contributed regarding the pappus structures of chyma cells. e.g. Xeranthemum annuum the tribe Cardueae. In Arctium lappa , Cirsium heleniodes and Man- Cypselar Anatomy tisalca salmantica endosperm layer is uniseriately develop , whereas in remaining 4 studied cypselas, Anatomically, all the cypselas are elliptic to ir- endosperm layers are biseriately oriented. Biseri- regular in cross sectiona. Among the studied ately development of endosperm layer have been cypselas, pericarpic region is differentiated into reported by Singh and Pandey (1984) in some spe- 2-3 zones. Within the pericarpic region, the meso- cies ( Carthamus baeticus and Saussurea hetero- carpic region is more interesting than the other mella ) of this tribe. Number of resin ducts of layers. In the cypsela of Saussurea fastuos , Cir- each cotyledon is more or less fixed and is one sium heleniodes and Xeranthemum annuum, meso- of the diacritical features of cypselas in Composi- carpic regions are heterogeneously developed tae. Each cotyledon has 3 (Cirsium heleniodes, Cir- rather than the other studied cypselas, where sium arvense, Arctium lappa), 5 (Cirsium candela- mesocarpic regions are homogenously developed. brum), 8(Mantisalca salmantica), or 9 In the cypsela of Arctium lappa, crystals are pre- (Xeranthemum annuum) resin ducts but in Saus- sent in the mesocarpic region but in remaining 6 surea fastuosa, resin duct is not clearly visible. studied cypselas, crystals are absent. Zarembo and Variable type of resin ducts have also been re- Boyko (2008) have reported the presence of crys- ported by Mukherjee (2000) in the same tribe. tals in the inner layer of pericarp and testa. They had also mentioned that the crystals are formed From this study, it can be concluded that mor- by calcium oxalate and have complex inner or- pho-anatomical features of cypselas are equally ganization. Dormer ( 1961, 1962 ) has studied the valuable as other taxonomic parameters, which crystal structure in the ovaries of certain Compo- are employed for proper identification of taxa sitae ( Centaurea, Cirsium and others), whereas along with other floral and vegetative characters Mukherjee and Nordenstam (2010), have studied in the tribe Cardueae. the crystal structure in Compositae and they have Key to the studied species mentioned the presence of crystals only in Arctium lappa among the studied taxa of this tribe. Taxo- 1a. Pappus absent; 3 resin ducts in each cotyle- nomically, secretary ducts have great importance don………………………………………(2) in the tribe Cardueae (Dittrich, 1977). Among the studied cypselas Mantisalca salmantica and Cir- 2a. Mesocarp with calcium oxalate crys- sium candelabrum secretary ducts are present tals………………………………… Arctium lappa which are absent in five other studied taxa. Secre- 2b. Mesocarp without calcium oxalate crys- tary ducts are also absent in some genera Amber- tals……………………………….Cirsium arvense boa (Pers.) Less., Yunquea Skottsb. in this tribe as has been reported by Chauhan (1972), Carlquist 1b. Pappus present; resin ducts are either not (1958). On the basis of the structure of testal epi- clearly observed or varies from 3-5-8 in each dermal cells, the studied species could be divided cotyledon……………………………………..(3) into 2 categories:

EXOMORPHOLOGY AND INTERNAL……. 58

3a. Carpopodial cells not clearly distinguishable positae Conference, eds. Hind DJN and Beentje HJ from the epidermal cells, i.e. pseudocarpopodia; Royal Botanic Garden, Kew.1-7. mesocarp homogenous; cotyledonary resin ducts varies from 5-8……………………………………(4) Carlquist S 1958 Anatomy and systematic position of Centaurodendron and Yunque ( Compositae). Britto- 4a. Stylopodium prominent; cotyledonary resin nia 10 78-93. ducts 5 in number… Cirsium candelabrum Chauhan YS 1972 Effect of gamma rays and 2,4-D 4b. Stylopodium inconspicuous; cotyledonary resin on the structure and development of seed coat ducts 8 in number…………… Mantisalca salman- and fruit wall of Carthamus tinctorius L. Pl Sci 4 tica 33-36.

3b. Carpopodial cells distinct from epidermal cells Dittrich M 1977 Cynareae- systematic review. In: of epicarp; pappus may be either plumose or The Biology and Chemistry of the Compositae, scaly type; mesocarp heterogenous; cotyledonary eds. Heywood VH, Harborne JB and Turner BL Aca- resin ducts either not clearly observed or varys demic Press, London. 999-1038. fr om 3 -8 in each cotyledon……………………………………………..(5) Dormer KJ 1961 The crystals in the ovaries of certain Compositae. Annals of Botany 25 241-254. 5a. Pappus plumose type……………………..(6) Dormer KJ 1962 The taxonomic significance of 6a. Carpopodial cells arranged in 1 crystal forms in Centaurea. New Phytologist 61 32 layer…………………… Cirsium heleniodes -35.

6b. Carpopodial cells arranged in 2 lay- Fourment PS and Rouzet M 1957 Fruit structure of ers…………… Saussurea fastuosa Galactites tomentosa Monech. Bull Soc Hist Nat Afr Nord 48 307-317. 5b. Pappus scaly type……………………………… Xeranthemum annuum Hanausek TF 1911 Uber das Pericarp und das Pericarpsekret der Gattung Carthamus. Berichte We are thankful to Prof. (Dr.) Hans V. Hansen, der Deutschen Botanischen Gesellschaft 29 13-18. Curator of the Herbarium, Botanic Garden and Museum of the University of Copenhagen, Den- Haque MZ and Godward MBE 1984 New records mark (DK) ; Dr. Peter Enz, Curator, Botanischer Gar- of the carpopodium in Compositae and its taxo- ten der Universitat Zurich, Zurich, Switzerland (Z) ; nomic use. Bot Jour Linnean Soc 89 321-340. Dr. Thomas Janßen, Curator of Späth-Arboretum, Hu mboldt-Universität zu Berlin, Ber- Jana BK and Mukherjee SK 2012 Diversity of cyp- lin,Germany (BHU) and Curator of Conservatoire selar features and their taxonomic significance in Et Jardin Botaniques Geneve, Geneva, Switzerland three species of the tribe Cardueae of Asteraceae. (G) for sending mature seeds for this study. In: Diversity and Conservation of Plants and Tradi- tional Knowledge 249-257. REFERENCES Lavialle P 1912 Researches sur le development Bentham G 1873 Compositae. In: Genera Planta- de 1’ overie en fruit chezles Composees. Ann Sci rum,eds. Bentham G and Hooker JD, Londini Nat Bot Ser 9 (15) 39-151. Reeve and Co., London. 163-533. Mukherjee SK 2000 Comparative morpho- Bremer K 1996 Major clades and grades of the As- anatomical studies of cypselas of some members teraceae. In: Proceedings of the International Com- of the tribe Cardueae (Asteraceae) by LM and

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