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Family Asteraceae) E-ISSN: 2395-6658 Emer Life Sci Res (2015) 1(2): 26-34 P-ISSN: 2395-664X Cypselas Diversity as Novel Taxonomic Marker in the Tribe Astereae (Family Asteraceae) Tulika Talukdar Received: 30 August 2015, Accepted: 14 November 2015 Abstract Introduction Despite the second largest tribe of the family Fruits of angiosperm dicot family Asteraceae are Asteraceae and immense importance of morphology usually termed as cypselae or achenes. The term of cypsela for taxonomic delimitation at tribe level, ‘Cypsela’ was first coined by C.de Mirbel (1815) no work has been done in this direction in the tribe and has often been confused with achenes (Bremer Astereae. In order to evaluate taxonomic 1994). Though, cypsela varies from the achene by implication of cypselas features, diversity of extra coat (perianth) over the pericarp due to the morpho-anatomical features has been carried out in lower location of the ovary (Judd etal. 2002). The the tribe Astereae. Detail macro as well as micro- term cypsela was adopted as a complex, dry, morphology and anatomy of cypselas of nine taxa indehiscent, unilocular fruit, with a single seed not including two species of Aster, two species of adnate to the pericarp (connected only by the Conyza and one species each of Erigeron, funicle) and initiating from an inferior ovary Grindelia, Kalimeris, Solidago and Vittadinia of the (Marzinek et al. 2008). Cypsela and pappus are two tribe Astereae (Family Asteraceae) have been morphological features which are aiding in examined using Light Microscope. Experiments taxonomic classifications at tribal levels of showed that surface characteristics like distribution Asteraceae (Talukdar 2008, Frangiote-Pallone and arrangement, hair type, ribs and their number, Antonio de Souza 2014, Talukdar and Mukherjee pappus diversity and presence and absence of wing 2014). Cypsela and pappus morphology together were taxonomically more important features in with growth form, capitula size, florets, involucral comparison to colour, size and shape of cypsela. bracts and leaf shapes were successfully used in Among anatomical features, mesocarpic characters separating daisy tribe Anthemideae into 12 sub- like type of tissue (whether parenchymatous or tribes (Bremer and Humphries 1993). For the sclerenchymatous), their distribution pattern, and taxonomic delimitation at tribe level, morphological nature of testal layer were found to be significant diversity of cypsela has been used to distinguish the for the tribe Astereae. Finally, involving all these tribe Heliantheae and Eupatorieae of family cypselas features an artificial key to the studied Asteraceae from rest of the tribes by investigating species is constructed. The key based on morpho- carbonized cypselas (Bremer 1994). Another tribe anatomical fearures could be used as reference key Inuleae was usually characterized by the presence to identify taxa of the tribe Astereae solely based on of calcium oxalate crystals in the cypselar its cypsela in absence of their flowering stage. epidermis (Merxmuller and Grau 1977, Breitwieser and Ward 2005). Keywords Astereae, cypsela, mesocarp, pappus, The tribe Astereae is the second largest tribe of surface hair, testa the family Asteraceae, with about 222 genera and ca. 3100 species distributed under 18 sub-tribes with high number of medicinal plants (Nesom and Robinson 2006, Talukdar and Talukdar 2013). The T. Talukdar Department of Botany, APC ROY Govt. College, Matigara, Siliguri, Darjeeling, West Bengal, India [email protected] Talukdar Emer Life Sci Res (2015) 1(2): 26-34 tribe is primarily characterized by anther (tailless Z : Botanischer Garten der Universitat Zurich, anther; ecaudate and ecalcarate anther base) and Switzerland style characters (deltate to triangular or lanceolate Z: Zollikerstrasse 107, CH8008 Zurich, style appendages). The tribe is also chemically Switzerland. distinct from other tribes by the absence of For investigating stable and perfect stage of pentaynene and sesquiterpene lactones (Bremer each character only fully matured and intact 1987, Brouillet et al. 2009). In any larger tribe like cypselas were observed. Investigations were carried Astereae of family Asteraceae, tribe phylogeny out broadly in five categories as follows required a thorough study of each and every organ of all the species without exception. Cladistic Macro-morphological studies of cypselas analysis based on broad morphological features was In situations with integrated cypselas, the initial and done in the tribe (Xiaoping and Bremer 1993). important process was to spot the posterior and However, it is really a fact that morphology of anterior (abaxial) surface of the cypselas. Further, cypselas in most of the tribes of Asteraceae has not 10 dry and FAA (Formalaldehyde, acetic acid, been received much attention as it should be. absolute alcohol) preserved mature cypselas were Besides morphological features, poor knowledge casually taken in glass and graphed slides and seen regarding the anatomical characteristics of cypsela under Olympus stereo dissecting microscope (DM) has hindered progress of taxonomic delimitations and binocular microscope (Olympus Model and tribe interrelationship in Asteraceae (Pandey 611062). Proper images were taken using Zeiss 2003). Structure and anatomical characteristics of Stemi DV4 camera equipped microscope. cypsela have been deeply observed to cover only A careful noting has been done for the shape, color few groups like Anthemideae and Cardueae and and direction of cypselas. Additionally, length and established to be taxonomically significant width of the cypselas were visually measured using (Heywood et al. 1977). In this context, present graphed slides, and in few cases they were scored investigation aims to deal with detailed study on by ocular and stage micrometer. In this work, morphological and anatomical features of nine taxa length of the cypselas was demarcated as the length belonging to the tribe Astereae. Traditional features of the body of cypselas from basal meristematic like size and shape of cypselas, structure of zone (carpopodium) up to apical end excluding stylopodium, nature of surface pubescence, pappus. Moreover, the width of the cypselas was carpopodium, pappus, nature and distribution of measured at the widest part of the cypselar body. In ribs and furrows etc. have been given distinctive case of heteromorphic cypselas, all the features preference. Anatomically, forms and comparative were examined for both the ray and disc cypselas distribution of several tissues in pericarp wall, and were studied separately. Summary drawings of nature of embryo, composition of testa and complete cypsela and diverse cypselar portions endosperm, orientation of cotyledon and number of were drawn by the Mirror type camera lucida. resin duct in each cotyledon etc. were observed in different ways . Finally, a sincere attempt has been Micro-morphological studies of cypselas made to construct an artificial key to studied taxa along with phylogenetic key involving all the For micro-morphological experiments, mature observed characters. cypselas were dipped in 1-5% sodium hydroxide (NaOH) solution for 2-7 d according to the Methodology stiffness. Further, they were moved to saturated chloral hydrate solution for some time, repetitively Collection of plant material washed with water and appropriately stained in 0.2- Plant materials (cypselas) used in this study were 0.5% aqueous Safranin solution. received as herbarium specimens from the Stained specimens were retained in 70% subsequent herbaria, which are mentioned in index phenol glycerine solution and dissected cautiously Herbarium (Holmgren et al. 1981). The list of for reviewing parts of cypselas. Proper photographs collected specimens has been shown in table 1. were taken using Olympus C-310 zoom digital camera (3.2 Megapixel) and Zeiss-stereo DK: Hortus botanicus Hauniensis, Denmark. microscope. emergent 27 Talukdar Emer Life Sci Res (2015) 1(2): 26-34 Nature of ribs, types, distribution and nature of pappus bristles, their number, orientation of hairs, nature of surface cells, other arrangement, colour, length, apex organization etc. epidermal structures, carpopodial cells etc. all were were also studied. judgmentally observed. Pappus characters such as Table 1. Source of materials with collection number Serial No. Name of the Taxa Locality Collection Number 1. Aster radula Aiton Z XXOZ- 20041675 2. Aster tripolium L. DK W DK:06 NJ 0218 3. Conyza bonariensis (L.) Cronquist DK 276E2405-0007*AG 4. Conyza canadensis (L.) Cronquist DK GE 2405-0008 5. Erigeron glabratus Hook. Z CHOZ-20060915 6. Grindelia robusta Nutt. Z XXOZ-20051345 7. Kalimeris mongolica (Franch.) Kitam. Z XXOZ-20041851 8. Solidago virgaurea L. DK W DK:06 NJ 0332 9. Vittadinia triloba DC. DK 488S1968-1357*AZ AU: Kimba (Kynclova 1970, Barthlott 1981) and partially Anatomical studies of cypsela improvised by the author herself. For anatomical studies, fine hand sections of cypselas were utilized for examining the internal Results and Discussion structures. Generally sections were made from the On the basis of morphological observations of middle part of mature cypsela. The cypselas were dipped in different chemicals for different duration cypselas, the tribe was characterized by the following features – cypselas are mostly of times depending upon the hardness of wall, such homomorphic, dorsiventrally compressed, ribbed as – (i) Cypselas were softened by dipping in and/or winged; pubescent with twin, sometimes glandular hairs,
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