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Diversity of Pappus Structure in Some Tribes of the Asteraceae

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Diversity of Pappus Structure in Some Tribes of the Asteraceae See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/270275517 Diversity of Pappus structure in some tribes of the Asteraceae Article · January 2008 CITATIONS READS 30 551 2 authors: Sobhan Kumar Mukherjee Bertil Nordenstam University of Kalyani Swedish Museum of Natural History 260 PUBLICATIONS 876 CITATIONS 92 PUBLICATIONS 1,685 CITATIONS SEE PROFILE SEE PROFILE Some of the authors of this publication are also working on these related projects: indian medicinal plants View project SESONAL VARIATION AND THE DIVERSITY OF THE WEEDS OF NADIA DISTRICTS, WEST BENGAL, INDIA View project All content following this page was uploaded by Sobhan Kumar Mukherjee on 04 January 2015. The user has requested enhancement of the downloaded file. PHYTOTAXONOMY Vol. 8, 2008. pp. 32-46 Diversity of pappus structure in some tribes of the Asteraceae Sobhan K. Mukherjee and Bertit Nordenstamr Department of Botany, University of Kalyani, Kalyani-741235, West Bengal, India The structures of pappus in 52 species belonging to 30 genera of the subfamilies Mutisioideae, Carduoideae, Pertyoideae and Cichorioideae have been studied critically with the help of both light microscope and SEM. The number of studied species in each tribe is indicated in parentheses: Mutisieae (l), Dicomeae (l), Pertyeae (2), Cardueae (12), Lactuceae (15), Vernonieae (18), and Arctoteae (3). Pappus structures can be divided into four categories with the pappus elements consisting of l) scales, 2) bristles, 3) crowns, or coronas, and 4) awns. Combinations of and modifications within the groups occur in the family, e.g., bristles may be smooth, barbellate or plumose. In ths taxa studied, pappus bristle is the most prevalent type, whereas true , coronate type is essentially absent. Pappus may be homomorphic or heteromorphic; eithef ananged in one, two or three rowsl .either persistent or caducous. Sometimes the apical part of the pappus is especially significant for distinction of tar<a (e.g., Sonc'hus spp.) or the-basal part of the pappus is different in ray and disc cypsela (e.g., Hypochaeris glabra). Vascular traces are not usually visible within the pappus structure, but are noted in Arclotis and' Catananche. The role of pappus structure in the evolutionary context is briefly discussed. A table based on the present survey presents the exomorphic variation of pappus and its possible evolutionary pathways. Keywords : Pappus structure; Asteraceae; Gichorioideae s. lat. Introduction A fourth subfamily, the Carduoideae, was soon added the paraphyletic The Asteraceae or Compositae are regarded as by subdivision of Cichorioideae (Bremer, 1996). further step on a fairly advanced family of the dicotyledons (e.g., A route higher taxa was Cronquist, 1988; Takhtajan, 1997). The the towards monophyletic classifications by Bentham (1873a, 1873b), taken by Panero & Funk (2002), who proposed a into Hoffmann (1890) and others have considered classification of the farnily eleven subfamilies these are regarded as pappus as important character in subfamilial, tribal and 35 tribes. All of and infratribal classification. monophyletic except one, the Mutisieae, which in their tree has two branches. Obviously, the last step The number of subfamilies and tribes of in the process of defuring monophyletic subfamilies Asteraceae haq increased dramatically.in the last ind tribes has not yet been taken. fifteen years. In the classification of Bremer (1994), There are two views regarding the morpho- the traditional concept of two about equally large logical nature of puppur. tniiially, a non-calycine subfamilies, Asteloideae and Cichorioideae, was nature of pappus was advocated by Small (1919). and supplemented by a third, the smaller This view was partially adopted by Sattler (1973), geographically more restricted Barnadesioideae. The but has not been generally accepted by recent subfamily Cichorioideae consisted of 6 tribes, 25 workers. The almost universally accepted view is subtribes, 391 genera and about 6700 species that the pappus is calycine in nature, i.e., a modified (Bremer 1994). This subfamily was a paraplyletic calyx. This idea was introduced many years ago group (Bremer, 1996), differentiated from the by Lund (1872) and later it was taken up and subfamily Asteroideae by a number of characters confirmed by a number of workers, e.g., Philipson (Robinson & Brettell, 1973; Wagenitz, 1976, (1953); Carlquist (1957); Ramiah & Sayeeduddin Robinson, 1977; Bremer, 1987, 1994;Kadereit, & (1958); Tiagi & Singh (1975); Cronquist (l9ss. Jeftuey 2007). 1977, I98l), etc. rDepartment of Phanerogamic Botany, Swedish Museum of Natural History SE-104 05 Stockholm, Sweden g:\dp\phynotaxonomy-volume-8\fi nal folder\fi le-7-F.pmd DIVERSITY OF PAPPUS STRUCTURE IN SOME TRIBES OF THE ASTERACEAE 33 Small (1919) recognized many types of pappus The pappus is a taxonomically important and sffucture and in his opinion all are derived from useful organ in Asteraceae, as noted by many the fundamental setose scabrid type. From the synantherologists including, of course, Cassini (e.g., scabrid seta, he derived simple and plumose bristles,' 1827). In tribal and generic revisions and floristic as well as paleaceous, aristate and coroniform work, descriptions of pappus are routinely given, 'superficial, pappus $pes, by fusion, rarnification, splitting and although sometimes brief and trnd the reduction processes (Fig. 1). Most denominations same is often the case in descriptions of infrageneric were adopted from early workers such as Berkhey taxa. An overview of the diversity in pappus (1761), Gaertner (1791) and Cassini (1826-1834). structwe within the subfamilyAsteroideae was given (2001). present paper It is generally considered that the pappus plays by Mukherjee & Sarkar The an impoftant role in the dispersal of the one-seeded aims to briefly survey and summarize the pappus groups dry fruits, called achenes or cypselas. This is variation in the other of the fhnrily, i.e., especially obvious in cypselas with numerous subfamily Cichorioideae sensu Bremer ( 1994)^ or' some and according capillar5, or plumose persistent pappus bristles, as of the basal subfamilies tribes in the common dandelion, Taraxacum sect. to Panero & Funk (2002) . Ruderalia. The transportation in the air of pappose Materials and Methods fi'uits is controlled by the relative humidity of the Pappus structures from 52 species under 30 atmosphere as well as air current (Taliew, 1895; genera belonging to 7 tribes (Pertyeae, Dicomeae, Sheldon & Burrows, 1973: Matlack, 1987; Mutisieae, Cardueae, Lactuceae, Vernonieae and Anderson, 1992, 1993). Not seldom, the pappus is Arctoteae) were studied critically with the help of (deciduous, caducous fugacious) and is detached dissecting binocular microscope, light microscope ' before dispersal of cypsela, although sometimes the and a ferv with SEM. The material of 48 species pappus (Shmida, is inconsistently deciduous 198!). was procur€d, from seven herbaria around the world, Epappose cypselas are conrnon in the.family arid viz., State Herbarium of South Australia, Adelaide may even be a constant feature for an entire tribe (AD); Queensland Herbarium, Queensland (BRI); (e.g., Calenduleae). Already Bentham (1873a) Centro de Botanica da Junta de investigacoes remarked that pappus was not always associated Cientificas do Ultramar, l,isboa (LISC); Jardirn with the dispersal of cypsela. Botanico do Rio de Janeiro, Rio de Janeiro (RB); Pappus structures can be involved in other National Herbarium, Harare, Zimbabwe (SRGH); dispersal mechanisms, such as zoochory. In the The Herbarium, National Taiwan University. Taipei genus Bidens L. (Heliantheae), the pappus awns (TAI); and Botanischer Garten der Universitat are retrorsely barbed, effectively promoting dispersal Zurich, Zurich (Z). In addition, fbur species were by furred or feathered animals. collected by the first author from India. Another function of pappus structure is Whole cypselas or detached pappus elements protection against predation on the fruits or ovaries, were directly stained in 0.5Yo aqueous safranin as discussed by Stuessy & Garver (1996), who solution and were mounted in phenol glycerine suggested that this defensive role of the pappus solution. For some species, pappus elements were may have been original and that the dual role for processed in a saturated solution of sodium protection and dispersal evolved later. This hypochloride for clearing the cellular structure of assumption led them to postulate the paleaceous the pappus. After that, the material was repeatedly pappus as primitive, in contrast to the general viey washed in water and stained as mentioned. At least of a setose pappus as the primitive state (e.g., five samples from each species were studied to cover Bremer, 1994). i the range of variations and to collect proper and average measurements the pappus. Pappus structures may have other frrnctions than of wind dispersal and predation defense, functions still Dry cleaned pappus or cypselas were scanned unknown or little investigated, €.9., pollinator using a Philips Electron Microscope at 15 kV iil attraction and dispersal by ants (myrmecochory). RSIC of Bose Institute, Kolkata, India after 34 PHYTOTAXONOMY, VOL. 8, 2OO8 Plate I re of_different species s. J-5 - A. reflexa var. t. 35 TRIBES OF THE ASTERACEAE STRUCTURE IN SOME DIVERSITY OF BAPPUS Plale2 77 36 PHYTOTAXONOMY, VOL. 8. 2OO8 , Plate 3 ii.' Figs. 80-88. SEM photographs of pappus structures of the Cichorioidear s. lat. Fig 80:
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