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1 H M m m v i m v i liiitiiiiiir mkh umdaiii mmi

2 BIODIVERSITY Interrelation between Flora, Fauna and Human Proceedings of the U.G.C. sponsored National Conference Organized by Departments of Anthropology, Botany, and Zoology Mrinalini Datta Mahavidyapith Birati, Kolkata-51 aw In collaboration with Department of Anthropology West Bengal State University, Barasat on September 29th and 30th, 2013

3 Biodiversity: Interrelationship between Flora, Fauna and Human 2014, Mrinalini Datta Mahavidyapith Published by Mrinalini Datta Mahavidyapith, Vidyapith Road, Birati, Kolkata West Bengal, India. Phone No: Website: The presentations and opinions expressed in the articles are exclusively the matters of the authors concerned. The editors and publisher are by no way responsible. All rights reserved. No part of this book may be reproduced, except for brief quotations, without the prior written permission from the publisher. ISBN: Printed and bounded at Vibgyor

4 Proceedings on Biodiversity: Interrelationship between Flora, Fauna and Human Pp (ISBN: ) COMPARATIVE MORPHO-ANATOM1CAL STUDIES OF CYPSELAS OF SIX SPECIES OF THE TRIBE ANTHEM1DEAE (COMPOSITAE) Bidyut Kumar Jana and Sobhan Kr. Mukherjee Department of Botany, University of Kaiyani, Kaiyani , West Bengal, India Abstract This paper deals with the detail morpho-anatomical structures of cypselas of six species (Anthemis maritima, Artemisia vulgaris, Tanacetum parthenium, Tanacetum vulgare, Tripleurospermum inodorum and Tripleurospermum maritimum ) of the tribe Anthemideae. Except the cypselas of Tanacetum vulgare and Tripleurospermum inodorum, the remaining 4 studied cypselas are heteromorphic. In the cypsela of Anthemis maritima, bristelate pappus is present, whereas in remaining 5 studied cypselas pappus are absent. Stylopodium is very prominent, enlarged and dome shaped in the cypsela of Anthemis maritima, whereas in remaining studied cypselas stylopodia are inconspicuously developed. In the cypsela of Anthemis maritima, Tanacetum vulgare, Artemisia vulgaris and Tripleurospermum inodorum true carpopodia are present i.e. histologically carpopodial cells are clearly distinct from the remaining part of cypsela. Remaining 2 studied cypselas, carpopodial cells are not clearly distinct from the remaining part of cypsela i.e. pseudocarpopodia. Anatomically all the studied cypselas are elliptical to trigonos to tetragonus in cross sectional configuration. In the cypsela of Anthemis maritima and Tripleurospermum inodorum, mesocarpic region is heterogeneously developed, whereas in remaining 4 studied cypselas, mesocarpic region is homogenously developed. Vellicular cavity is present in the mesocarpic region of the cypsela of Anthemis maritima and Tripleurospermum maritimum. Cypselas of Artemisia vulgaris, vellicular cavity is present at the lower region of testa, near the lateral lobe region of cypsela. Secretary ducts are present in the mesocarpic region of the cypselas of Tanacetum parthenium. Testal layer is uniseriately developed in case of all the studied cypselas. In the cypsela of Tripleurospermum inodorum and Anthemis maritima endosperm layers are uniseriately develop but in 4 studied cypselas, endosperm layer is biseriately developed. The morpho-anatomical features of the above studied cypselas have been investigated to establish their potential usefulness in taxonomy. Keywords : Anthemideae; Cypselar morpho-anatomy; Compositae Proceedings 30

5 Introduction The family Asteraceae is one of the most advanced, highly evolved family among the dicotyledons, containing 43 tribes, genera and species by Funketal.1. The Anthemideae is one of the oldest tribe of the Asteraceae, where capsular features were used for better evaluation and understanding of tax. According to Bremer et al.2, Cypsela provides much information in this tribe, Bremer & Humphries3 have made a cladistic analysis of the genera of this tribe. The use of capsular features in the classification of Asteraceae has been employed since the work of Schultz - Bipontinus4. Kynclova5 has provided a detailed investigation of cypselar structure in some genera of the tribe Anthemideae. The most stricking features of cypselas of this tribe are a. Apical part of cypsela bears scales or corona like pappus. b. Pappus bristles are usually absent which are present in other major tribes of this family. In addition to this, apical part of cypsela possesses persistent stylar base in association with glandular nectary, known as stylopodium. Differences in size, shape, colour of cypsela, orientation of cypsela in relation to receptacle, nature of apical margin of the corona, number and distribution of vascular traces and ribs and distribution of carpopodial cells differ from taxa to taxa. The present study is aimed to supplement the previous works ( Bhar and Mukherjee 6, Kallersjo7 etc.), for better understanding of taxa, which may help in reshaping the existing system of classifications. The present study deals with the detailed structure of cypsela in 6 species belonging to 4 genera of the tribe Anthemideae. Materials and Methods The present work is based on the Herbarium specimens obtained from 3 herbaria of the world which were given in the table-1. Table-1: Showing the name of studied taxa of the tribe Anthemideae and their sources SI. No. NAME OF TAXA SOURCES 1. Anthemis maritima L. Botanischer Garten der Universitat Zurich. XXOBRISS Artemisia vulgaris L. Botanic Garden Of The University, Jyeangontie 2, SF Helsinki, Finland. 3. Tanacetum parthenium Sch. Bip. Botanischer Garten der Universitat Zurich. XXOZ Tanacetum vulgare L. Humboldt- Universitat zu Berlin. Institutfur Biologie. Spezielle Botanik u. Arboretum. Berlin, Germany. 5. inodorum (L.) Sch. Bip. 6. maritimum (L.) W. D. J. Koch Botanic Garden of The University, Jyrangontie 2, SF Helsinki, Finland Humboldt - Universitat zu Berlin. Institut fur Biologie. Spezielle Botanik u. Arboretum. Berlin, Germany. Proceedings 31

6 For morphological observation, mature cypselas were softened by dipping in 4% NaOH solution and after that they were stained in % aqueous safranin - light green combination to observe the different morphological parts under simple dissecting microscope and stereo dissecting binocular microscope. For anatomical study, cross sections were done from the middle portion of cypselas, with the help of sharpe blade to observe the different anatomical regions with the help of compound light microscope. Results and Discussion Comparative morphological and anatomical features of six studied species were given in. the table - 2 and table - 3 respectively. Table - 2 : Showing the morphological features of studied cypselas Anthemis maritima (Fig 1 A-C) Artemisia vulgaris (Fig 1 D-F) Tanacetum parthenium (Fig 1 G-H) Tanacetum vulgare (Fig 11-J) inodorum (Fig 1K) maritimum (Fig 1L-M) Cypsela heteromorphic. Ray cypsela 2mm x 0.5 mm; disk cypsela 1.5mm x 0.5mm. Cypsela Cypsela heterom6rphic. Ray cypsela 1.5 mm x 0.5 mm; disk cypsela 1 mm X 0.5 mm. heteromorphic. Ray cypsela 1 mm x 0.05 mm; disk cypsela 1.05 mm x 0.05 mm. Cypsela homomorphic, 2 mm x 0.5 mm. Cypsela homomorphic, 2 mm x 0.05 mm. Cypsela heteromorphig. ray cypsela 2 mm x 1 mm; disk cypsela 2.05 mm x 0.5 mm. Ray cypsela yellow brown, oblong, slightly curved; disk cypsela yellow brown, straight, obovate. Ray cypsela pale brown, wide oblong, slightly curved; disk cypsela pale yellow, ovate, slightly curved. Ray cypsela pale yellow, straight, obovate; disk cypsela pale yellow, linear-ovate, straight. Pale yellow, linear, straight. Upper part truncate, lower part rounded. Black, fistulose, straight, upper part truncate, lower part tapered Ray cypsela pale yellow, ovate, upper part truncate, lower part tapered Surface glabrous, containing 9 ribs. Surface rough and glabrous, containing 4-5 ribs. Surface rough and glabrous, containing 10 ribs. Surface rough and glabrous, containing 5 ribs. Surface glabr-ous, containing 4 ribs Surface rough and glabrous, containing 3 ribs. Pappus bristelate type. Pappus absent. Pappus absent. Pappus absent. Pappus absent. Pappus absent. Stylopodium prominent Stylopodium inconspicuous. Stylopodium inconspicuous. Stylopodium inconspicuous. Stylopodium inconspicuous. Stylopodium inconspicuous. Proceedings 32

7 Anthemis Artemisia Tanacetum Tanacetum maritima vulgaris parthenium vulgare inodorum maritimum (Fig 1A-C) (Fig 1 D-F) (Fig 1 G-H) (Fig 11-J) (Fig 1 K) (Fig 1 L-M) Carpopodium Carpopodium Carpopodium Carpopodium Carpopodium Pseudocarpopodium symmetric. symmetric, symmetric, symmetric, symmetric, bi- present. Carpopodial ring like. carpopodial 4lobed. convex. Carpcells arranged Carpopodial cells not clearly Carpopodial opodial cells in single row. cells arranged distinct from the cells arranged arranged in 3-4 in single row remaining part in single row. rows. of cypsela i.e. pseudocarpopodium. L M ~ 0.C5 mm. C, F, J 1mm. A, B, D, E, G, H, I, K, L, M Fig.1 Morphology of studied cypselas Proceedings 33

8 A-C-Anthemis maritima: A-Ray cypsela, B- Disk cypsela, C- Carpopodial cells; D-F- Artemisia vulgaris: D-Disk cypsela, E- Ray cypsela, F- Carpopodial cells; G-H Tanacetum parthenium: G- Ray cypsela, H- Disk cypsela; l-j- Tanacetum vulgare: l-cypsela, J- Carpopodial cells; K- Cypsela of inodorum; L-M- maritimum: L-Ray cypsela, M- Disk cypsela. Table-3: Showing the anatomical features of studied cypselas Anthemis Artemisia Tanacetum. Tanacetum maritima vulgaris parthenium vulgare inodorum maritimum (Fig. 2A) (Fig. 2B,C) (Fig. 2D) (Fig. 2E) (Fig. 2F) (Fig. 2G) Cypsela elliptic Cypsela elliptic Cypsela elliptic Cypsela elliptic in Cypsela Cypsela in cross in cross section. in cross section. cross section. tetragonous in tetragonous in section. cross section cross section Cypselar wall Cypselar wall Cypselar wall Cypselar wall Cypselar wall Cypselar wall mm x mm x mm x mm x mm x 0.03 mm x 0,02 mm wide at mm wide at mm wide at ribs mm wide at ribs mm wide at ribs mm wide at ribs ribs and furrow ribs and furrow and furrow and furrow and furrow region and furrow region region region respec- region respec- region respec- respectively respectively. respectively. tively. tively tively Epicarp Epicarp Epicarp Epicarp Epicarp Epicarp uniseriate, uniseriate, uniseriate, uniseriate, made uniseriate, made uniseriate, made up of thin made up of thin made up of thin up of thin walled, up of thick- made up of thin walled, rectangular, walled, walled, rectangular, walled, rectangu- walled, rectangu- compactely rectangular, rectangular, compactely lar - oval, lar, arranged parencompactely compactely arranged compactely compactely chyma cells, arranged, arranged parenchyma arranged, arranged provided with parenchyma parenchyma cells, provided parenchy-ma parenchyma cuticle cells. cells,provided with cuticle cells, provided cells, provided with cuticle. with cuticle. with cuticle Mesocarp Mesocarp Mesocarp Mesocarp Mesocarp Mesocarp heterogenous, homogenous, homogenos, homogenous, heterogenous, homogenous, containing both made up of only made up of sclerenchyma- containing parenchymatous, parenchyma sclerenchyma parenchyma tous. parenchyma and containing vellicular and scleren- cells. cells containing sclerenchyma cavity. chyma cells. secretary duct. cells. Mesocarpic region containing vellicular cavity Proceedings 34

9 Anthemis maritima (Fig. 2A) Artemisia vulgaris (Fig. 2B,C) Tanacetum parthenium (Fig. 2D) Tanacetum vulgare (Fig. 2E) inodorum (Fig. 2F) maritimum (Fig. 2G) Testa 0.01 mm thick, uniseriate,. Testa 0.01 mm thick,uniseriate, parenchymatous Testa 0.01 mm thick, uniseriate, Testa 0.01 mm thick, uniseriate, parenchymatous Testa 0.02 mm thick,uniseriate, Testa 0.01mm thick, uniseriate, Endosperm uniseriate, Endosperm biseriate, parenchymatous Endosperm biseriate, Endosperm biseriate, Endosperm uniseriate, Endosperm biseriate, Cotyledons conta-ining 6 resin ducts (3 ducts in each cotyledon). Cotyledons containing 6 resin ducts (3 ducts in each cotyledon). Cotyledons containing 6 resin ducts (3 ducts in each cotyledon). Cotyledons containing 6 resin ducts (3 ducts in each cotyledon). Cotyledons containing 6 resin ducts (3 ducts in each cotyledon). Cotyledons containing 6 resin ducts (3 ducts in each cotyledon). Fig.2 Anatomy of studied cypselas 0.05 mm A, B, C, D, E, F, G Proceedings 35

10 A- Anthemis maritime, B- Artemisia vulgaris (Through ribs), C- Artemisia vulgaris (Through lateral lobe), D- Tanacetum parthenium, E- Tanacetum vulgare, F- inodorum, G- maritimum Detailed morpho-anatomy of cypselas of 6 species of the tribe Anthemideae have been studied. Regarding the cypselas of this tribe, Bremer et al 8 have indicated a valuable statement "The fruits (cypselas) provide much information in the Anthemideae". Except the cypsela of Anthemis maritima and inodorum, the remaining 4 studied cypselas are homomorphic. Colour quality of cypselas are also variable, though it is not a reliable distinguishing character because it changes with the degree of maturity of cypselas. Among the studied cypselas, in case of Anthemis maritima and maritimum, yellow brown; Tanacetum vulgare, Tanacetum parthenium and Artemisia vulgaris, pale yellow and inodorum, brown black in colour. Shape of studied cypselas are variable from oblong to linear to obovate. Though size, shape, colour are not important taxonomic character, for the separation of taxa. Cypselar surface in the Anthemideae is characterized by the presence of myxogenous trichomes by Kallersjo9. Though, among the studied cypselas, these structures are not observed. Structure of stylopodium is clearly visible in young cypsela. Therefore, this structure may not be valuable in mature cypsela. In the cypsela of Anthemis maritima, stylopodium is elongated, dome shape, whereas in remaining 5 studied cypselas, stylopodia are inconspicuously developed. MukherjeelO, has studied the stylopodial characters of some members of Asteraceae. Actually, stylopodia are the persistent style base with associate nectar. At the basal region of cypselas, carpopodium is present. It is a meristematic zone with variable layers of cellular arrangement. In the cypsela of inodorum, carpopodial cells are arranged in 3-4 layers whereas in the cypsela of Tanacetum parthenium and maritimum, pseudocarpopodia are present. In the cypsela of Tanacetum vulgare, Anthemis maritima and Artemisia vulgaris carpopodial cells are arranged in single row. Actually carpopodium is the basal, abscission zone of cypsela by Mukherjee and Nordenstam 11 and it helps in the detachment of cypselas from the thalamus. Anatomically all the studied cypselas are elliptical to trigonous or tetragonus in cross sectional configuration. Mesocarpic, region exhibits cellular variations. In the cypsela of Anthemis maritima and inodorum, mesocarpic regions are heterogenous. In remaining 4 studied cypselas, this regions are homogenously developed. Heterogenously developed mesocarpic region is also found in some species of the tribe Lactuceae by Jana and Mukherjeel 2.Vellicular cavity is present in the mesocarpic region of cypsela of Anthemis maritima and maritimum whereas in remaining 4 studied cypselas, this cavity is absent. Vellicular cavity is also present in some species (Aster alpinus, Felicia heterophylla) of the tribe Astereae by Jana, Bar and Mukherjeel 3. Another important character is the presence of resin duct in the mesocarpic region of cypsela of Tanacetum parthenium, whereas in remaining 5 studied cypselas, this cavity is absent. Secretary duct is also present in the mesocarpic region of the cypsela of Elephantopus carolinianus of the tribe Moquinieae by Jana, Bar and Proceedings 36

11 Mukherjeel 4. Presence of vellicular cavity and resin ducts are most important characters on the basis of taxonomic view point. In all the studied cypselas, testal layers are uniseriately develop. Except the cypselas of Anthemis maritima and inodorum, the remaining studied cypselas endosperm layers are biseriately developed. Key to the studied species 1a. Cypsela homomorphic; pale yellowback (2) 2a. Cypsela pale yellow; surface containing 5 ribs; carpopodial cells arranged in single row; testal layer 0.01 mm in thick Tanacetum vulgare 2b. Cypsela black; surface containing 4 ribs; carpopodial cells arranged in 3-4 rows; testal layer 0.02 mm in thickness inodorum 1b. Cypsela heteromorphic; yellow brown-pale brown-pale yellow 3a.Vellicular cavity present; secretary duct absent (4) 4a. Surface containing 9 ribs Anthemis maritime 4b. Surface containing 3 ribs maritimum 3b. Vellicular cavity absent; secretary duct may or may not be present (5) 5a. Surface containing 4-5 ribs; secretary duct absent Artemisia vulgaris 5b. Surface containing 10 ribs; secretary duct present Tanacetum parthenium List of abbriviations used Ep- Epicarp, Me- Mesocarp, Pa- Parenchyma, Scl- Sclerenchyma, V.C.- Vellicular cavity, S. D.- Secretary duct, T- Testa, E- Endosperm, V. T.- Vascular trace, Pal- Palisade parenchyma. Acknowledgements We are thankful to Dr. Peter Enz, Curator, Botanischer Garten der Universitat Zurich, Zurich; Curator of Humboldt- Universitat zu Berlin. Institut fur Biologie. Spezielle Botanik u. Arboretum. Berlin, Germany and Aune Koponen, Scientific Curator, Botanic Garden of The University, Jyrangontie 2, SF Helsinki, Finland for sending mature, identified seeds for this study. References 1. Funk VA, Susanna A, StuessyTF and Bayer R J 2009, In: Systematics, Evolution and Biogeography of Compositae. International Association for Plant Taxonomy, Vienna, Austria, p Bremer K, Eklund H, Medhanie G, Heidmarsson S, Laurent N, Maad J, Niklasson J and Nordin A 1996, PI. Syst. Evol Bremer K and Humphries C 1993, Bull. Nat. Hist. Mus. London (Bot.). 23 (2) Schultz Bipontinus C H 1844, Phytographia Canariensis Kynclova M 1970, Preslia , 51. Proceedings 37

12 6. Bhar I and Mukherjee SK 2004, J. Econ. Taxon. Bot. 28 (3) Kallersjo M. 1985, Nord. J. Bot. 5 (6) Bremer K, Eklund H, Medhanie G, Heidmarsson S, Laurent N, Maad J, Niklasson J and Nordin A 1996, PI. Syst. Evol Kallersjo M. 1985, Nord. J. Bot. 5 (6) Mukherjee SK 2005, In: Plant Taxonomy. Advances and Relevance. A.K. Pandey, W. Jun and J.V.V. Dogra (eds.), CBS Publishers & Distributors. New Delhi, p Mukherjee SK and Nordenstam B 2004, Comp. Newsl Jana BK and Mukherjee SK 2012, Indian Journal of Fundamental and Applied Life Sciences. 2(1) Jana BK, Bar R and Mukherjee SK 2012, The Journal of Biodiversity, Photon Jana BK, Bar R and Mukherjee SK 2012, TLS. 1(2) 1. Proceedings 38

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