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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
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 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: Arctiunt lappa - Deciduous pappus scars on the top of cypsela, x 50. Fig. 8l: Carlina vulgaris - Plumose bristles ol' pappus. x 25. Fig. 82 Centaurea cyanus - Subulate pappus scales in 3 rows, x 50. Fig.83: C. nnculosa - Subulate scales in -i rows. x 50. Fig. 84. Catananche caerulea - Scarious scales, x 25. Fig. 85 Sonchus brachyotus - Apical part of inner rorv ol pappus bristle s, x 3200. Fig. 86: Vernonia cistrfulia - Double pappus, x 50. Fig. 87'. Lepidaploa gracilis - Double pappus. x 5 0 Fig. 88. L'ernonia scorpioides - Double pappus, x 100. mounting, labelling and gold coating. SEM images Table I gives the Conspectus of pappus features were photograbhed by uping 35 mm (125 ASA) of 52 species in some basal tribes of the Asteraceae. film. Magnification of the photographs can be of studied species and the source of material ,List'is measured from thlbar diagrams of the photographs. as follows: Terminology pappus for description of follows ' Ttibe Pertyeae: Ainsliaea latifolia (D. Don) (1977) cornmon usage such as in Heywood et al. Sch.Bip., S. Mukherjee 17 (KAL); A. reflexa Merr. and Bremer Q99\. This study covers selected taxa var. nimborum Hand.-Mazz., Yuh Fong Chen 3300 from seven tribes of the subfamily Cichorioideae (rAr). sensu Bremer (1994). The tribe Liabeae is not Dicomeae: Maclediunt sessiliflorunt included, neither the recently distinguished Tribe (Harv.) S. ssp. sessili/Iorum, A. R. Torre l3 Gundelieae, Gymnarrheneae, Tarchonantheae, Ortiz (Lrsc). Hecastocleideae and Gochnatieae, nor is the subfamily Barnadesioideae (tribe Barnadesieae) Tribe Mutisieae: Gerbera jamesonii Bolus considered. ex Adlam, Nr. 397 (Z). DIVERSITY OF PAPPUS STRUCTURE IN SOME TRIBES OF THE ASTERACEAE 37 Tribe Cardueae: Arctium lappa L., Nr. 343 Torre 118 (LISC); V scorpioides (Lam.) Pers., SN (Z); Carduus defloratus L., Nr. 359 (Z): Carlina 251 (RB); V senegalensr's Less., Schlieben 2457 acanthifolia All. ssp. cynara (Pourret ex Duby) (Lrsc). Rouy, Nr. 360 (Z); C. vulgaris L. ssp. vulgaris, Tribe Arctoteae: Arctotheca calendula (L) Nr. 361 (Z); Centaurea cyanus L., 36a Nr. Q); Levyns, N. N. Donner 8541 (AD); Arctotis venusta C. macrocephala Puschk. ex Willd., Nr. 366 (Z); T. N6rl., Nr, 3a5 (Z); Berkheyo zeyheri Harv. & C. maculosa Lam. ssp. maculosa, 369 (Z); Nr. Sond. ssp., zeyheri, A. R. Torre 6907 (LISC). Cirsium japonicum DC., Nr. ,378 (Z): 'C. valgare (Savi) Ten., s. tr., s. coll. (BRI); Echirlops Discusiions sphaerocephalus L., Nr. 386 (Z); Leuzea As mentioned in the Intrgduction, most recent rhapontica (L.) Holub, Nr. 413 (Z); Ptilostemon workers have adopted the view that the pappus is diacanthus (Labill.) Greuter, Nr. 377 (Z). a modified calyx, e.g., Cronquist (1977), Thorne (1983) Tribe Lactuceaez Actites megalocarpa (Hook. and Takhtajan (1997). All species studied here have f.) Lander, A. A. Munir 5512 (AD); Catananche a well defined pappus, though it is absent caerulea L., Nr. 363 (Z); Cicerbitq cyanea (D. in many other genera in different tribes (Achyrothalamus, Adenocaulon, Arnoseris, Don) Beauv., S. Mukherjee 18 (KAL); C. macrorhiza (Royle) Beauv., S. Mukherjee l9 Cephalopappus, Cullumia, Panphalea, Tarchonanthus, (IGL); Crepis pyrenaica (L.) Greuter, Nr. 383 etc.). (Z); C. vesicaria L., N. N. Donner 8607 (AD); Pappus is often persistent, but a deciduous or Hieracium villosum Jacq., Nr 404 (Z); Hypochaeris caducous pappus exists in many studied taxa of the glabra L., A. A. Munir 8601 (AD); H. radicata tribe Cardueae and a few other genera (Sonchus, L., s. n., s. coll. (BRI); Lactuca serriola L., s.n., Bothriocline, etc,). Pappus may be homomorphic s. coll. (BRI); Leontodon autumnalis L., Nr. 409 or heteromorphic, but the latter type is predominant (Z); Sonchus braclryotus DC., S. Mukherjee 20 in Cardueae and Lactuceae, (KAL); S, schweinfurthii Oliv:&.Hiern, M. Mavi On the basis of this observation, pappus can 8 (SRGH); Taraxacum fficinale G. Weber, A. A. be classified into four major types which are shown Munir 5500 (AD); Tragopogon porifolizs L.r, N. in Table 2. N. Donner 8606 (AD) . Among the pappus types, scabrous barbellate Tribe Vernoniea e: Baccharoides anthelmintica is most prevalent in the studied taxa. Length of (L.) Moench, S. Mukherjee I (KAL); B. pappus bristles varies from taxon to taxon and within hymenolep,s (A. Rich.) Isawumi, El-Ghazaly & B. taxa, and also within a single floret. So, in general, Nord., G. Pope 1930 (SRGH); Bothriocline laxaN. this feature is less important, but may sometimes E. Br. ssp. laxa, M. Mavi ll (SRGH); Centrapalis be employed for separation of taxa (e.g., Arctotis, kirkii (Oliv. & Hiern) H. Rob., F. A. Mendonca Berkheya, etc.). The longest pappus is recorded in 2033 (LISC); Elephantopus scaber L., SN 257 (RB); Cirsium. Colour of pappus and structure of apical Lepidaploa gracilis (H.B.K.) H. Rob., SN 250 (RB); cells of the pappus elements may or may not be Linzia melleri (Oliv. & Hiern) H. Rob., R. Santos taxonomically significant. 2051 (LISC); Polydora bainesii (Oliv. & Hiern) H. Pappus features are specially significant in the Rob., G. Pope 1929 (SRGH); P. poskeana (Vatke following cases: & Hildebrand| H. Rob., A. R. Torre & Paiva 11332 (LISC); Rolandra fruticosa (L.) Kuntze, SN 255 i. In case of plumose pappus, the length of the (RB); Vanillosmopsis capitata (Spreng.) Sch.Bip., lateral projection of the pappus bristles is SN 248 (RB); Vernonanthura gondensata (Baker) diacritical for the identification of species H. Rob., SN 249 (RB); V. dffisr) (Less,.) H; Rob.; (Ainsliaea). SN 254 (RB); .Vernonia cistifolia O. Hoffrn," G ii. In .Centoure4, pappus is generally double, Pope 1931 (SRGH); V glabra (Wjlld.) Vatke, M. sometimes represented by three rows of scales Mavi 12 (SRGH); V petersii Oliv. & Hiern, A. R. ' or bristles. The innermost scales are shortest and PHYTOTAXONOMY, VOL. 8, 2OO8 the middle ones longest. Dittrich (1977) reported primitive (Norlindh, 1977). Such type of that the pappus bristles are longer towards the arrangement is noticed in the tribe Cardueae. Bremer centre in Centaureinae, sometimes with reduction (1994) considered this group as fairly primitive of the innermost row. among the tribes of the subfamily. He noted the iii. Regarding the pappus structure of Vernonieae, plumose pappus found in many genera of this tribe Jones (1977) pointed out that external feature as essential for generic delimitation. Regarding this of pappus under LM and SEM studies yielded tribe, Dittrich (1968, 1970) emphasized that pappus valuable information for certain genera, features may not be reliable and should be employed particularly Vernonia. This statement is verified with caution, because different species of some by the preserit study. Out of 7 studied species genera have both scabrid barbellate and plumose of Vernonia and Vernonanthuro,5 species have bristles. a double pappus of scales and bristles, but Bremer (1,994) considered the Arctoteae the pappus scales are absent in Vernonia ' highest evolved [ribe, as may be inferred from the senegalensls and Vernonanthura condensata. present study. According to Heywood et al. (1977), iv. In Arctoteae, in addition to general features of Vernonieae is one of the most primitive tribes in the pappus, the shape and size of its elements Asteraceae, as was suggested by Augier & du Merac are taxonomically diacritical (Roessler, 1959). (1951), although this view was not accepted by This opinion is correct for the threi: studied taxa. Cronquist (1955). Pappus structure clearly indicates v. Apical part of pappus bristles can be used for the position of the Vernonieae as fairly advanced discrimination of different species of Sonchus. as has been noticed by Bremer et al. (1992). Tips of inner row of bristles either possess Norlindh (1977) has shown that Arctoteae is recurved lateral cells, or recurved and upwardly closely related to Cynareae or present Cardueae and directed cells, while the tips of outer row of the pappus structure distinguishes these two tribes. bristles have upwardly directed lateral cells or In Arctoteae, the pappus is characteristically scaly uniseriate parenchyma cells without lateral and never plumose, whereas pappus is commonly projections. plumose in C5mareae. The positions of these two vi. Pappus plays an important role for protection tribes are quite distant in Bremer's system. of ray and disc cypselas in Hypochaeris glabra. Mutisieae .is closely allied to Cardueae (Karis, Outer row of bristles is more or lesq identical 1992; Karis et al., 1992; Jansen et al., 1990, 199 I ). in ray and disc cypselas. Irurer row, of bristles Both these tribes oommonly have plumose bristles. in ray cypsela have tuft of .long tomentose hairs I,actuceae towards the base which are absent in inner row is probably the highest evolved tribe of bristles in disc cypsela. (Heywood et al., 1977). vii.Vascular traces are not usually visible in the On the basis of pappus features, it has affrnities pappus element structure, except in Arctotis and with Cardueae which has been established from the Catananche. work of Kim et al. (1992) based on rbcL sequence This work isnnot sufficient to correlate or data. The position of tribe Lactuceae should be an discuss the evolutionary relationships among the intermediate location in an evolutionary scale. studied tribes. But still it is possible to show some Structure and arrangement of pappus elements relationships among the tribes of the subfamily in a particular species are more or less stable and Cichorioideae. Small (1919), Robinson (1981) and definite and the basic pattern is not much determined Bremer (1987) considered a pappus of many parts by external factors of environment. Morphological as primitive, whereas Cronquist (1955) stated that variations ofpappus are nearly constant at generic the chafl pappus of five elements is most primitive. and, to some extent, up to tribal level too. Since Pappus is often arranged in I or 2 rows, occasionally pappus characteristics are ultimately controlled by in 3 or more rows. Arrangement of pappus in 2- genes, they are diacritical for taxonomic distinction 3 rows has sometimes been considered as most at or below the tribal level along with other 'srnuclun-t DTvERSITv oi ,orru, rN soME riueps oF THE ASTERACEAE 39 associated vegetative and floral features. Dittrich, M. 1958. Morphologische Untersuchungen an den Frtlchten der Subtribus Cardueae-Centaureineae Acknowledgements (Compositae). Willdenowia 5: 67-107. 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S. & Hind, Plrytomorpholog. | : 391404. D. J.N. (eds.). Compositae: Biology & Utilization, Proceed. Ramiah, N. & Sayeeduddin, M. 1958. Homology of the pappus Intem. Compositae Conf. Kew 1994, Vol. 2 : 8l-91. Royal in the light of trichome distribution. Curr Sci.27: 402404. Bot. Gard. Kew. Robinson, H. 1977. Au analysis of the characters and Takhtajan, A. 1997. Dirersity and Classification of Flowering relationships of the tribes Eupatorieae and Vernonieae Plants. New York: Columbia University Press. (Asteraceae). Systematic Botany 2: 199-208. Taliew, W. 1895. Ueber das hygroscopische Gewebe Robinson, H. 1981. A revision of the tribal and subtribal limits des Compositen-Pappus. Dissert. Abstr. Bol. Cent. Bd. of the Heliantheae (Asteraceae). Smithsonian Contrib. Bot. 63: 320. 5l: l-102. Thorne, R.F. 1983. Proposed new realignments in the R. D. 1973. Studies in the Senecioneae Robinson, H. & Brettell, Angiosperms. Nordic. .1. Bot. 3: 85-117. (Asteraceae) IV. The genera Mesadenia, Syneilesis, . Miracacalia, Koyamacaila, and Sinacalia. Phytologia. ,ftael, V.O. & Singh, B. P. 1975. Vascularization, morphological 27: 265---276. nature and trends of evolution i4 the pappus of the Compositae. Abstr. ln: All India Synposium on Form, Roessler, H. 1959. Revision der Arctotideae-Gorteriineab Structure andFinction in Plants. Sardar Patel Univ., India. (Compositae) . Mi tt. B ot. Staat s samml. Miinchen. 3 : 7 l -5 00. Wagenitz, G 1976. Systematics'irnd phylogeny of the Compositae Sattler, R. 1973. Organogenesis of Flowers. A photographic (Asteraceae). Plant Syst. Evol. 125: 2946. Text-Atlas. Toronto and Buffalo: Univ. of Toronto Press. DIVERSITY OF PAPPUS STRUCTURE IN SOME TRIBES OF THE ASTERACEAE 41 o o c.l o o o o (J I s E t ai a; E a* ! e & & a o, c) - 6) o F C6 o > o o; o o > q)' q) o ! '6 'O >' I C) () c) C) CB o o o (6 o cB o a o' o' o * o O q f-. I oo +*E: a.l qJ :e etcFse I o\ I 'I I 6t \o I t, I gs$ g a .ri 6t E a E o\ \o r- est E r; F- qJ o q) o\ frr T& frr frr T! o o o sFE; ?s ? a m c.t €=- eg aiiY etEt oo d d I d U F t g5 es = 5E 3 fit a I €E gbb ()a m dtr o -oo qd !o EE d -o G) 0.) U) adH clE o c'l ico € (d rd Q c.t rc q rd () ri .o ,o 0) q ' () () c) (ua q I td Ox hl t{ ri R' cl Cd .9nct. cg p a G^ a= 2= ! (.) 3 ! {) F- F d 0 C) '=o o o p 6) C) a o & €E vc) h U ,; & GI G' o C) Q o o H aq t> ah o ! -o ol o Pi ! to e e -oC' Q a_o q_o o () - ri ri ri q ri o. o. n -od tq FC) q te r a .oE tr 3 q) oq) oct qSocd € 'o & / il qo & O o F rE 'r ts F 9'- F F rA q) Hq tr c! q) tr E' tut ile/, F o Sts EO: o a \o 'rOO^Y9I c d...i I I N N (\ ti o ! ts_- o o o C) o o s6E AE EEEEeE ?,i? '4{ FJi 2- f o.r ' F - -=- o- g,i,C=b- Q i a-.:iJ/ t A A o. A IJ a o q) FEEAs -cg eElE. cn + + + + + + + sda.9 l-5 - + 4 .{ .o fil rn s\ a- v $.= E.N FQ t lF.- q) N o 3n $. BI G'O R (Joo * o\o G \€ -8' iri o C.l $.o S)5"; \JP-i $; bo9 R'"; (J:: or S"l ta o-'' 8.9 xbn L n;' \: 4 rrr ao -oo ,< \) ,s.\) E {* q v F \ \ :{4 \, (-) U U -io6Z c.l co + \o r\ € 42 PHYTOTAXONOMY, VOL. 8, 2OO8 d o o I o o o o o o o &I & () C) C) () O o o o c) o o c) (.) (.) ; F c) o J4 l4 .v B JZ B € cd o. E o a a U) U) ' C) .-a h: . j q o\nh !^dH '.d : q F5 q q r-+o ao co Ir F.- t- ,: c.i o loJ > gs _ cal &.&, I o I I I :c.l I I I I 3rJ =cz i-: q n o c o>f4 )&- ; l'\ r; rn + \o 6l C.t x.& oz -j o o o o o O o o ca m o ca ca 'c0 ca m frr fr a x d d x d c) () :-a'r5 ts c)o ,;2 c) ed ci=o< >\4J >,E G- 12^ C! (-A<9 Cgi! o-^ a$ k' c) o =cda) O^ o ir ,r F () O -q) I C;) o:.i 4E a-o tr O() (d(q o (!LO 3$ l6 ..o o_E =5tr-c) c€F a oE .Yq q ri ':>\ 3-o ,ii o €o 7v (.) E EJE .oo €V a? .orE Cn !E .= o oo ar: q hl o'- H- Q gtFt+ --E > g -o C) 3 ai: c)L D a- - ) .i' (d 33gI 9R -o: qo F o4;: LqOEqq =tro- !.1 o Q 'io?3 o=r ot oOa I c) 0) s e5E i :Ye 6 >'cts^ s'- u ao F] 9tr'ecd= =tso -.Oiq i= .= >I0) L O =Eo k ;;" a- FI -oc5().- ()--V- ae Q) .E]6 o3 -o 5 o tr g.e 6 H d= E o Ed C) () c!tr x'- A0-^ cs 3eE aOq 3?, o o & I3E3; =3 E 6-o OJ E tr oc) ti ojjE'i -grl o-vq oro AG 8.9.= .Y o) q olj -e- d=E Gl'= ) ,:o rvL!- ^ =qA ^r= o-E3 3AE d5 83.E J o.-o ts o5; c.t cO (\.l (\l c.l cn (,) c) c) o o c) C) oI o c) :E +r +r c) ir H t-.1 o A a n t..l + + + + + + + + + + + q p 4 ()c \ s c.l :f, soc oo $co P. rS o ia Oh s 9\ *e3 + s*; i \h s S- \? a- ss t (J' sl \ Sc.t SSN s ss $r stl q)u iP shb b: Sr$ sstH:: ts" S E.s s(] sO i GN Bo $iao .IrI. ol) ^o u ? fi $* \) q)\ :h! Srr- dHsr. q tt dr! -EBavii o st! L -t! U * \) U U U \) \Q o\ c.l cO * \o r\ oo o\ DIVERSITY OF.PAPPUS STRUCTURE.IN SOME TRIBES OF THE ASTERACEAE 43 & o () o O o o I cI & o o & o o I & & Q I I o & & C) .:0.) c) c) 4 () - o () >z F E>' o c) o a 6cl a' Fbo 3 6 > o Oo o o o; 7 B EOJ .(.) c) 0) G) o 6) Q ?o O coF \o o<; .i 6\ l1 oo I () E. lr. tu E! * E. IL I& o o c0 co ca s d d d d x d O >,! q qr CO'lJ 4 q () () q) ! crJ.:o+ ;.1€ C)- q q 3Ea .UEg"E;EE 6 .dH5 F sF L o-L o q) rEC't6E CHiq€{ -C) 5oc) 3EE s o-t: o I I E dtrts desog>> E 3C) =bo () () oO< tr rE=ts L h'5 ! € SE s 6Ov --Lv co d d o t:€s =-o- 6 g ; 3a Ja) ! 6s c6 €-c='tr (d ,o (.) c) "iF lE:E;€; () 5-OE = c) = o SsE i EE€ G.i 9: o9 5 g:.835EE bo-li - - qi -o (.) kE= € 3tr= cq !d (€ o 3J'x 9 5 d aoo o € €crvo o ,^.E I 6Y!-(9 ! F dqb0 a)rq= E ,t? q q 3g E 3 H'- o s] o Y= i -.! o- O :f EEs EE =oiD- OE o o >.3s €'..O--^Ea >,c? 6,= i53E G) € 3 +.e E 3 -o o_!2 L =.=Cg g5E La(E c€ != r(6 t s:iE*; (,) :q q O ,(t >,9 >.: >,b i - >\o=-.Y^- o eFb a.o € .oq=u_^ 2-O=.9 E F9f:IE; 2=Aa E c) ; ETi F F: () 3!t V.,o2 qQ) E - 9.?6 o 9d ;?tr?i )q Oqc,? itt:-b o 'a;9to?E (.)() rc)o- aq !u9=0 E d) I L kii -tr9>, oo:e=L- € - c) o dE ! a- v E€ TFY],iL JO :.9.= ts o EF,gE-i3 o- d5€ fJr ds 3 S$EE?Eg !r d5 t 3 d58tE c05E E;9 >5 a.l NN a.t c.t a.t c\ N a\ a.t () q) o c) o o o aI oI Ji o o + + + AA o. A H A L] c q + + + + + + + + + + + v 3 s \3 B U .E 4 "a U Fn d Q6 S- o^ Xoo oO co \ ^sN s E3 s .9* =c4 ln" s€$ U\O iv) > o.i V2 B* "j "oto\ c)' \co3+ 8g \ S-r LI f.oN tr >bo sca q) s$ .S cn f F; s.s PL q) 'i :.E Bq; R; or$ 5() ;\33o .{ 'a\ (j ol) "S o0 () ():r su QEq d; E .9p $ TrI. ol& o str \ B!! s5b oE SS qJ \ u1 U O $ { o R. .s V) e.l cO t \o F- oo o\ N (\ a.l N a.l al (-.t N c{ N co 44 PHYTOTAXONOMY; VOL. 8, 2008 o () o o b o o q) C) & $ o I & o (.) E c) d dg &F ts I o () o o ts> &o = I oo 6E > ()5 H ' C)' (,) C) o C) () €aE '=e€ c) .=> ' >a)tr r9 _€ s>, 'q o C) o o. C) =.o E046 5q> () I >.o }EI. > >F i ct d dF? o0 hI) o.- o d o. & o- J )d H o oE) oi g5 G) J 5> 8d F- C) rO r; o ovi ?o ,q X\O c'l oi ^o\ \o it; o04oi oo t- r lt + -.:t^ o9o 6l \Y ol a.l oo I I I rno I t. q o ;-i \O ?+ ,2= 9.4 r- r; 9v Xg= ;-i \o &i ct; 84 - z- OcZ & a.t d d q) d d d d +Y d trr' g* d Ed 'd5 >'!2 OF '83 '-t c G< !! tro oi () c) do ?.o d3'E Eo .=q J'* 39 Ect o-oQE i\ d)' ,c) 3 h >EE FAE oo9 q PE ts -6 it .- ^cdr.o (\ a\t a.l cn N N (\l d) ! a L] n ! o I o I I I at I (j C) (.) (.) C) o (.) + :E + O.ArH A a d) A< H o- A o. o. a + + + + + + + + + + + + q) s ,o :E U2 X .\a u1 rS kta s (J B q \ o I s) (l^ q, F osh -R .9 S\o lal s q) S- .v ;".= NN -. G "\. .i \o gR v2 bo q) FE is €+ o ql.oo SR s \ o s \ as *s: }\J!R iln I s\o s p L {!* s600 v!9s ti' $ Il x,; HE i frs L i t .99 o tfo '=$ F\ \ii \ ls l& di d.{' o \) a- { e'!I. I m U $' $ q. o kl { a- q N c.) .s w; \o F- co c.) d o\ a.l cr) c7) cn cr) c.t + !f, = I DIVERSITY OF PAPPUS STRUCTURE IN SOME TRIBES OF THE'ASTERACEAE 45 & o o I o o o o o & o & o D I tr d'= &'= J,&. &ot - z-o o c)/ c)t C)_c ()o ' o g 56 .=> (.) -o -o Eq Eq '== >' 5-c (.) q t o >t >i EB q,) G ro t,o E +g +> .(! ! ca o. &/a ! x= & A q) ^o og sfa o- Qo rlo d6 \.n ?o X je; oq r; qF Xgr xva o<; I l1 \Toi ?+ c..i qci ^^l ,,? ro ..!. -o n I p4+ !o +J o tv ol $l'\ Nl ?+ o&&,P4Tod I c.; n4 c.- l6 l6 ld) t- od q'l -o LJH4;/, 84 sp cto <.- o o C0 >ro Ir e+ ();i= (d E. &,& cla frr l\ Er frr t\ l\ r{FEHE L-./ 6)x= o.l E.Y d d d N )& I € r- oo oo E3 q; tJ( () ,b33 ara ko ge.i gg*.1'F A o o- o F }9dL E cn o E6 o bo-o d 6); t)a S= t'* -o c€ 5SEs5 4C()O () 6 a)l i o9- (t i i O! F JO 9 0--='i -c, o 3€ Q-Ea H b -6= "i.2 i! q 9g -oo-b0 tr X cgtc) s 9a F O.cg (g do - o -o .=5 cql !b- E td i= H (oh"o-> () do i.=-o -trc) :YO- 9>i- >\F o: ar:'- Y;g f-l ; !.= _> 9EEEQ -o i oP 3r 6uc)o =;!(d r; iggj g(Jcd F 2E--> C) C.)X o C)- iEoSEeBE g.= !?o5 q) =Ee!XdoStit g.5 F 9LL o-f!= id ? =? q ooh d .9 blE € 6(D236 O@c) EEerl-^€ t@Kl ? 9S XD Q diuA ii l{E 5 qc s -r. c)= o o-Y J o 5-6 o=@= o=iEE (d.= -9o 9oQ r6 I .H F3T3E = 'a6-t 59- '5oH o (r)!CL SETF d-c-a-- o-- 5ll c H odo=n = o'E ; ri E=rE9 O.() Fi k di 5 E2.: E-6 'o € !o= !f- ,Ee tr l!o<€a tX qr *>,di o o i-S a9 s'- ,^6 €ad "G*9 -='-= d !S3EP (h - aoo & 4Or'-rd 6 '-i 'i -= 50 oq) J4()o 3gEE3 O. *i -51 o.E X Fr O-- C) H E *dE15 3 c,-== q a =Ea .3. H e r-2. =c) =7Er o5= dE *EE-b ,o.V c) IVLE; Js di5 od d'H5bE g l(6 o F o.>ei fi ss o.-o-o= A€E EO6 a-o PrctOqO SE dE )tri o: = cd g (^.t a.t c.t N a.l 6l a.t c.l a t-l F o I I I oI aI o o C) o tsG) o c) H >Jr llr +l +t +t o. Or Or A o. o- Or or p. + + + + + + + + + + s c \ s :a Wt q) .O , :s:] O\ \) U1 v) .S:^ -tn 4 S\o o .S*oo .: f N dSr: :\o s :', :; q oo;. S $t'- ll\N \o \ F d- x r.r cs s P-bo *Sr c Oi- s* xh s .F,9 >ri o- 9EO\O .s $,$ r$ 9r q fo o S:* o \ d* E* oE S'".fo \ \. lr< q\) \ er! () -V @;: L .U \ q) rq) s N a c.l + \o t- 6 o\ + $ * s s \t sf 6i 46 PHYTOTAXONOMY, VOL. 8, 2OO8 Table 2. Ilpes of pappus elements and their probable evolution Combination of sciirious scales Combination of plumose bristles and scabrous barbellate to and subplumose bristles scabrous subplumose bristles (Hypochaeris glabra) (Vernonia) I I I I Plumose bristles, definite number, Paleaceous scales to fonn corona 1-roq significantly widened (Arctotheca, Arctotis etc.) towards b ase (L e o nto do n) A I I I I Plumose bristles, one row significantly widened towards basr Scarious scales connate to form (Tragopogon) multifid cup likg laciniate corona (Rolandra) t I l I 1 Plumose bristles, one row, not widened towards base (Carlina, Scarious subulate scales connate Cirsium etc.\ to form corona (Echinops) 'L I I I Plumose bristles, two-three rows (Macledium, Ptilos temon etc.) I I Scarious scales apically I prolonged into scabrid barbellate Subplumose bristles (Leuzea) ' bristles (setae) (Catananche, Elephantopus) I I I 'Scabrous barbellate bristles, one row (B othriocline, Centratherum etc.) Scabrous subulate s0ales free Scabrous barbellate bristles, two or (Centaurea) more rows (Arctium,t Crepis etc.) I I Scabrous to capillary bristles, two rows (Cicerbita, Lactuca etc.) t leaceous) line Bqistle (setose) line Scaly origin View publication stats