Chapter

11

Classification of Compositae
Vicki A. Funk, Alfonso Susanna, Tod F. Stuessy and Harold Robinson

INTRODUCTION

The Compositae (Asteraceae) family is nested high in the Angiosperm phylogeny in Asterideae/Asterales. The family contains the largest number of described, accepted, species of any plant family, ca. 24,000, with estimates of the total number reaching 30,000. There are 1600–1700 genera distributed around the globe except for Antarctica. Assuming that there are 250,000–350,000 species of flowering plants, then one out of every eight to twelve species is in Compositae (about 10%). That the family is monophyletic has never been in question. Every early worker in plant classification recognized Compositae as a group at some level (i.e., Tournefort 1700; Berkhey 1760; Vaillant 1719–1723) and in every type of analysis the family is monophyletic (i.e., Small 1919; Bremer 1987; Jansen and Palmer 1987; Hansen 1991; Michaels et al. 1993; Lundberg and Bremer 2003). The family is characterized by florets arranged on a receptacle in centripetally developing heads and surrounded by bracts, by anthers fused in a ring with the pollen pushed or brushed out by the style, and by the presence of achenes (cypselas) usually with a pappus (Fig. 11.1). Although the family is well-defi ned, there is a great deal of variation among the members: the habit varies from annual and perennial herbs to shrubs, vines, or trees, although few are true epiphytes; species grow in just about every type of habitat from forests to high elevation grasslands, however, they are less common in tropical wet forests and more common in open areas. Most groups in the family contain some useful and some noxious species as well as common and rare taxa. However,

the general perception of this family as “weedy” is not correct. Certainly there are members that benefit from disturbance, such as a few species of dandelions and thistles, and a few global pests (e.g., Chromolaena odorata (L.) R.M. King & H. Rob.), but most species have a restricted distribution and just about every ‘at risk’ habitat in the world contains members of this family that are an important part of the flora. From the beginning, those who studied this family thought that presence of both ray and disk florets (Fig. 11.1A) represented the basic head pattern. In his classic illustration, Cassini (1816; Chapters 1, 6 and 41) placed Heliantheae at the center, Vernonieae and Eupatorieae at one end, and Mutisieae and Cichorieae (Lactuceae) at the other. The treatment by Bentham (1873a) had 13 tribes (the most frequently used suprageneric rank), which remained more or less the same until the 1980s; although some of the concepts changed, especially in Vernonieae, Liabeae, Senecioneae and Helenieae. Bentham’s work was developed independently from that of Cassini (e.g., 1826), but the 13 tribes of Bentham do correspond to many of the 19–20 tribes of Cassini. Hoff mann’s treatment of Compositae (1890–1894) essentially repeated the classification of Bentham (Turner 1977; Bremer 1994). Bentham (1873b), and more recently Cronquist (1955, 1977) and Turner (1977), all thought that Heliantheae were the most primitive tribe of the family, and accordingly assumed that the ancestor was a perennial herb with opposite leaves and a yellow-flowered, radiate capitulum. In 1977 Cronquist changed his mind and allowed for a woody ancestor, as suggested by Carlquist (1966, 1976). Carlquist (1966) proposed changes to the system of Bentham, but

and both erred in their placement of Eupatorieae because of its superficial resemblance to Vernonieae. sometimes the style branches are recurved and come in contact with the style shaft. b). C Some of the achene (cypsela) and pappus types found in Compositae.2). At the same time. which is monophyletic and the sister group to the rest of the family. long believed to be closely related. for the most part agreed with the molecular fi ndings. Carlquist (1976). achene drawings from Robinson 1981.1. showing that part of Mutisieae was the basal branch of the family and that the tribe Heliantheae s. England (Heywood et al. it was based on the molecular work by Jansen and Palmer (1987. It illustrates the current thinking about the relationships among the major tribes and subfamilies in Compositae. which contains less than 1% of the species in Compositae. Intercalated between the two monophyletic subfamilies are groups that used to be included in the subfamily . disc florets in the center. a tree of trees (Chapter 44). which contains ca. Stuessy and Robinson Fig. and Jeffrey (1978) all divided Compositae into two groups approximating Asteroideae and non-Asteroideae of recent treatments. The basal group. 11. where he mentions that the pollen structure of Mutisieae is more like closely related families and very different from Heliantheae. were actually in separate parts of the phylogeny. (1991a. A summary tree shows the position of the major branches (Fig. They literally turned the Compositae phylogeny upside down. their work showed that Vernonieae and Eupatorieae. Jansen et al. Bremer’s analysis (1987) based mostly on morphological data. a meeting on “The Biology and Chemistry of the Compositae” was held in Reading.l. Understanding the phylogeny of Compositae has come a long way since the papers of Jansen and Palmer (1987) and Bremer (1987). Poljakov (1967) provided a fair approximation of the modern two groups of tribes with Cichorieae-Arctotideae more basal and Anthemideae-Heliantheae derived. Robinson (1973). Also monophyletic is the highly nested subfamily Asteroideae.] they were not often used and acceptance of the basic 13 tribes continued. Characters of Compositae. In 1975. Using published trees for the family as a whole. only Carlquist and Jeffrey applied the subfamily names Asteroideae and Cichorioideae in their revised sense. Bremer’s analysis placed Eupatorieae close to Astereae and not Heliantheae. 65% of the species in the family. we have constructed a metatree for the family. was nested far up in the tree. The biggest change in Composite systematics took place in the late 1980s and early 1990s. and an involucre with bracts (phyllaries) surrounding the outermost florets. The proceedings from that meeting are interesting in that the book used the Bentham system. [Drawings by Alice Tangerini (US). 1988). 1977). which are fused at the margins. However. is the distinctive subfamily Barnadesioideae. figure from Funk et al. B The pollen is released via the style pushing out through the anthers. A The head with ray florets arranged around the perimeter. but several of the chapters made it clear that the data did not fit this classification. and published and unpublished trees for individual clades.172 Funk. Furthermore. and Jansen and Kim (1996). 2005. Abandonment of the stand-alone Lactuceae in the subfamily Cichorioideae was inevitable. Of these authors. Wagenitz (1976). A good example is the pollen paper by Skvarla (1977). The advent of new techniques and new markers has greatly increased the amount of sequence data available. Susanna. 11.

also biennial or perennial herbs. all ligulate (ligulate capitulum) or all bilabiate. fi laments usually free. rarely the florets variously otherwise arranged. 2008) that recognized new and previously described subfamilies and tribes so that now there are 12 and 43. style solitary. Unit of inflorescence a capitulum (head). 35% of the species in the family) and that vary in their morphological and molecular characters. shrubs.l. the upper part of the fi lament usually with thick-walled cells. ligulate with an apically 5-dentate abaxial ligule. with rare exceptions surrounded by an involucre of one to several series of protective bracts (phyllaries). chaff ) subtending some or all of the florets. However. capitula sometimes solitary at the apices of more or less leafless stems (scapes) but usually few to many in often corymbiform cymose inflorescences (capitulescences. Our current classification is built on the foundation laid by others from Cassini to Kadereit and Jeff rey. tissues usually with schizogenous secretory canals (resin-ducts) and/or with articulated lacticifers. usually in several series. respectively. it broke up Heliantheae into twelve tribes that reflected the current state of knowledge. Each of these works advanced our knowledge of the family and with this volume we hope to do the same. or variously zygomorphic. usually echinate (spiny). synflorescence) of various types. other areas of the family were still in flux (e. occasionally all the florets pistillate or staminate and the plants dioecious or monoecious. equal in number to and alternating with the corolla lobes. and the inner regular. The treatments in this book are based on our attempt to combine the morphological and molecular data into a meaningful classification. forming a split cylindrical or balusterform anther-collar (or fi lament-collar). usually with an apical appendage. capitulum homogamous) and all regular (capitulum discoid). two or three are not supported by currently available morphological data and it remains to be seen whether or not these few taxa will be accepted by the Compositae community (see Chapter 12). rarely whorled. more or less regular (actinomorphic) and equally or unequally (3–)5(–6)-lobed or -toothed with the lobes or teeth valvate. pollen mostly tricolporate. sessile or subsessile. pales. capitulum heterogamous) with the inner (disc florets) regular (or rarely bilabiate) and perfect (bisexual. Stamens with the fi laments inserted on the corolla-tube. sometimes echinolophate or lophate (with a pattern of raised ridges) or spinulate (microechinate. Panero and Funk 2002. For instance. perfect or functionally staminate (capitulum disciform). tapetum integumentary. dehiscent by longitudinal slits. or the outer fi liform pistillate. calcarate or tailed at the base. of (3–)5(–6) united petals. rounded.g. both of which were based on a mixture of morphological and molecular data. dithecal. often pistillate (female) or sometimes sterile (neuter). 2007. The Kadereit and Jeff rey treatment included the latest information available at the time. apically divided (except sometimes . in one or more series (capitulum radiate). COMPOSITAE GISEKE (1792) [ASTERACEAE MARTYNOV (1820)] Family description (adapted from Jeff rey 2007) Often annual herbs. anisomorphic. sometimes succulent. exstipulate. spinulose).) and while some groups such as Pertyeae and Dicomeae were recognized on an informal level. introrse. fi mbrilliferous with non-vascularized fi mbrils or scale-like processes surrounding the bases of the florets.. rarely the corolla absent from the pistillate florets. Bremer (1994) and Kadereit and Jeff rey (2007). often caveate. monocarpic or polycarpic. or fi liform with the lobes reduced or absent or with a minute ray. 1–1000 or more per capitulum. usually terrestrial (rarely epiphytic or aquatic). usually with one or more of various types of glandular and eglandular hairs. hermaphrodite) or functionally staminate (male) and the outer (ray florets) radiate. Florets (flowers) small. or paleate with persistent or caducous vascularized scales (paleae. others did not have sufficient data available to make a decision on their placement. the florets either all alike (homomorphic. The Bremer book retained the three subfamilies but acknowledged that Cichorioideae were most likely paraphyletic. rarely connate.Chapter 11: Classification of Compositae 173 Cichorioideae (ca. Since that time there have been only two treatments of the whole family. corolla gamopetalous. pseudobilabiate with an unlobed internal (adaxial) lip and a 4-lobed external (abaxial) lip. 2002. calyx a pappus associated with the fruit (see below). commonly the glandular biseriate and the eglandular uniseriate. bilabiate with a 2-lobed internal (adaxial) lip and a 3-lobed external (abaxial) lip. or radiate with an abaxial 0–3(–4)-dentate ray. isomorphic. Leaves alternate or opposite. usually simple but often lobed or divided. Receptacle either naked or hairy and smooth. anthers united into a tube surrounding the style. nectary a thickened scale or cup surrounding the style base. sometimes aggregated into often involucrate capituliform syncephalia of the second or even third order. leptocaul or sometimes pachycaul. Mutisieae s. the different types variously arranged within the capitulum. areolate with polygonal areoles or alveolate with depressions in which the florets are inserted. subshrubs. elongating through the anther-tube and extruding the pollen at its summit. vines or trees. Recently a new higher classification system was proposed for the family (Baldwin et al. or of more than one type (heteromorphic. Chapter 1 covers the work of many of the scientists who provided early classifications of the family up until 1977 when the Heywood et al. very rarely free. sagittate. volumes were published.

S = Senecioneae. A = Arctotideae. CARDU. = Hyalideae. Stylidiaceae (245) Alseuosmiaceae (10) Phellinaceae (11) Argophyllaceae (20) Menyanthaceae (60) Goodeniaceae (440) Calyceraceae (60) Barnadesieae (91) Onoserideae (52) Nassauvieae (313) Mutisieae (254) Stifftia (8) Hyaloseris clade (7) Gongylolepis clade (29) Hyalis clade (3) Leucomeris clade (3) Wunderlichia (5) Stenopadus clade (30) Gochnatieae (70) Hecastocleideae (1) Dicomeae (75-100) Cardueae (2500) Tarchonantheae (13) Oldenburgieae (4) Pertyeae (70) Gymnarrheneae (1) Warionia C1-3 (313) Cichorieae C4 (900+) Cichorieae C5 (200+) Eremothamneae (3) A-Arctotidinae (76+) Heterolepis (3) A-Gorteriinae (131+) Platycarpheae (3) Liabeae (174) Distephanus (40) Moquinieae (2) Vernonieae (1500+) .. = Wunderlichieae.174 MUT. MUT. The numbers by the terminal taxa reflect the number of species in that clade. A summary tree based on the metatree (Figs. Mexico and North America) and that the red color in Vernonieae represents Tropical America.3–44. WUNDER. Susanna.7). Outgroups Funk. 11. Subfamilies that have more than one tribe are indicated in capital letters (see classification in this chapter). Hya. = Carduoideae.2. The branches and internodes were colored according to the distribution of the taxon or the optimization of those distributions. Wun. = Wunderlichioideae. = Mutisioideae. Wun. Note that some areas have been combined (e. Stuessy and Robinson WUNDER. The tribes or clades have been represented by one to four branches. CICHORIOIDEAE Cichorieae Stifftieae Fig. CARDU.g. Hya. 44.

ovary inferior. usually an achene (cypsela). very rarely a drupe. the apices of the style arms acute to rounded. fruit. Mexico Central America. southern South America General South America Africa Southern Africa Madagascar. Europe Eastern & central Asia South America Brazil Guiana Shield North & central Andes Southern Andes. scales (squamae. stigmas dry. . basal ovule. distinguished by the form of its cells and the texture of its surface. New Zealand Widespread or ambiguous in functionally staminate or apomictic florets) into two (rarely three) short to long branches (style arms) with stigmatic areas on their inner (adaxial) surfaces. indistinct to prominent. with one erect. southern Europe General Africa North America North America. forming a thin layer around the embryo. unitegmic. endosperm scant. indehiscent. papillose. New Guinea. embryo straight. unilocular. crowned by a pappus formed of (1–)2 to many awns. truncate or with various appendages. abscission scar surrounded by a carpopodium. homomorphic or heteromorphic.Chapter 11: Classification of Compositae 175 ASTEROIDEAE Senecioneae (3500) Corymbieae (9) Doronicum (40) Abrotanella (20) S-Tussilagininae Grade S-Othonninae S-Senecioninae Calenduleae (120) Gnaphalieae (1240) Astereae (3080) Anthemideae (1800) Inuleae (687) Athroismeae (55) Feddeeae (1) Helenieae (120) Coreopsideae (550) Neurolaeneae (153) Tageteae (267) Chaenactideae (29) Bahieae (83) Polymnieae (3) Heliantheae (1461) Millerieae (380) Perityleae (84) Eupatorieae (2200) Madieae (203) Asia-Eurasia Eurasia. tenuinucellate. sometimes apparently absent. Mediterranean. squamulae). of one to many rows of cells. of two (rarely three) united carpels. setae or hairs in one or more series. ovule anatropous. New Caledonia. Caribbean Australia and the Pacific Australia. tropical Africa Northern Africa. or by a more or less coroniform or auriculiform structure. 1-seeded. or the pappus caducous or completely absent.

Don) Bremer & Jansen (1992) 1. Barnadesioideae (D. Cass. shrubs. often villous. emarginated to bilobed. anther apical appendage rounded to obtuse or acute. ligulate. (1829) 24.A. and Wunderlichieae were not validly published in 2002. orange. doubly split.* Wunderlichioideae Panero & V. Recent investigations show that Corymbieae. Within the subfamily Asteroideae there is the well recognized clade. (1819) 21. leaves alternate except for most Liabeae. Carduoideae p. pollen with or without depressions. Cichorieae Lam. 3-lobed “true ray corollas” are found only in Liabeae and some Arctotideae. (1994) Heterolepis incertae sedis XI. Gnaphalieae (Cass. Pertyeae. Funk (2002) 9. Tarchonantheae Kostel. lophate or echinate (Liabeae. Pertyeae Panero & V. disc corollas in many forms but lobes usually longer than wide (deeply lobed). yellow. occasionally paleous or glabrous. chartaceous.A. (1819) 25. Liabeae (Cass.* Stifftieae D. (1819) 30. Helenieae Lindl. Funk (2002) VII. thecae with or without spurs (calcarate or ecalcarate) and with or without tails (caudate or ecaudate). 1-numerous. Funk (2002) VI. short descriptions of tribes with diagnostic features and geographic distribution are provided below. some Cardueae). Characters follow a uniform order.) Lindl. Oldenburgieae S. calcarate) and with tails (caudate). Gochnatioideae (Benth. Mutisioideae (Cass. (1901) 42. opposite. Neurolaeneae Rydb. The tribes are broken into two large groups. Eupatorieae Cass. & Brettell (1973) 19. Susanna. Coreopsideae Lindl.Ort.A.A.A. (1819) IV. & DC.A.A. corollas tubular.) Lecoq.A. Rob. Barnadesieae D. (1929) 41. Gymnarrhenoideae Panero & V. heads discoid or radiate or ligulate. Hecastocleidoideae Panero & V. Pertyoideae Panero & V. Funk (2002) 23.G. Anthemideae Cass. Astereae Cass. Herrera. Tageteae Cass. Corymbieae Panero & V. split. (1829) 33. Wunderlichieae Panero & V. fasciculate or whorled. Millerieae Lindl. f.A. Group 1: ‘Non-Asteroideae’ grade Tribes not contained in the subfamily Asteroideae usually have: stigmatic surface solid on inner surface of style branch. Gochnatieae (Benth.A. the Non-Asteroideae (a grade) and the subfamily Asteroideae (a clade).A.A. Gymnarrheneae Panero & V. Funk (2002) X.) Panero & V. Rob. & Hook.A. Madieae Jeps. Don (1830) III. (1819) 29. (1819) 4. Ortiz (2009) 12. white. pink. Gymnarrheneae. (1819) 36.G. Nassauvieae Cass. The primary providers of the information for most of the tribes are listed. Baldwin (2002) 38. Funk (2007) 6. (1819) 40.176 Current classification Funk. rarely bilabiate (3 + 2). Funk (2002) 15.) I. + denotes small tribes with only one or two genera. Hecastocleideae. Don (1830) II. Athroismeae Panero (2002) “Heliantheae alliance” 31. receptacle usually pilose. (1819) 26. Don) Panero (2007) 2. Baldwin (2002) 37.* Stifftioideae (D. & Hook. ex Sweet (1826) 10.) Lindl. Baldwin (2002) 43. Funk (2002) 11. Moquinieae H. Funk (2007) 5. Cichorioideae ( Juss.A.) Panero (2002) 39. (1806) 17. Base chromosome number x = 9 or 10. Heliantheae Cass.* Hyalideae Panero (2007) V. ex Dumort. (1819) 28. Mutisieae Cass. Funk (2002) XII. A republication of these suprageneric names with a Latin diagnosis will be submitted shortly. perennial or annual herbs without latex. & Juillet (1831) 27. Platycarpheae V. An online key to the tribes can be found on www. f. Dicomeae Panero & V. Funk (2007) 7. (1829) 3. subshrubs. Rob. Anderb.compositae. florets usually hermaphroditic. those without attribution are by V. the Heliantheae Alliance. (1819) 18. Chaenactideae B.p. Funk (2002) 14. Vernonieae Cass. Polymnieae (H. & Franc. bracts in several series.A. Cardueae Cass. see Chapter 12. some Lactuceae. Senecioneae Cass. leaves alternate. Funk & H. pollen bearing part of anther extending well below the insertion of the fi lament (spurred. Corymbioideae Panero & V. P. purple to violet. Hecastocleideae Panero & V. 1. Arctotideae Cass. Funk. Inuleae Cass.org. (1819) VIII.G. . pollen psilate. subbilabiate (4 + 1).A. Bahieae B. Eremothamneae H. Feddeeae Pruski. (1829) 34. Onoserideae (Bentham) Panero & V. (1927) 35.) Rydb. Funk (2002) 8. Asteroideae (Cass. (1833) 13. Stuessy and Robinson (* = under discussion. (2009) 22. Funk (2002) Catamixis incertae sedis IX. Perityleae B. Rob. — (information provided by Estrella Urtubey): Trees. Barnadesieae.) Panero & V. (2008) 32. (1819) As an aid to understanding the system of classification. (1828) 16.) Chevall. Calenduleae Cass. (1927) 20. involucre from cylindrical to widely campanulate.

inner longer than outer. usually capillary. or rarely tubular. the innermost slightly apically broadened and longer. capillary bristles. alveolate. anther apical appendages acute. pink. pale yellow. receptacle naked or hirsutulous. the Andes. papillose rarely pilose. plane. bisexual. purplish. leaves alternate. Diagnostic features: woody plants with zygomorphic or bilabiate corollas. multiseriate or few-seriate. — Shrubs or small trees without latex. involucre usually campanulate. thecae with spurs (calcarate) with long. margins not hyaline. truncate. styles thick. receptacle naked. Distribution: southern South America. pollen usually prolate. branches dorsally smooth to slightly rugulose at the apex. pappus of one to more series of bristles. bracts multiseriate. strap-shaped with three teeth. florets isomorphic. bifid. apex rounded to shortly acute. multiseriate. 1–150 florets. capillary to somewhat paleaceous. echinate. microechinate. sometimes dimorphic with the marginal achenes conspicuously larger than the central ones. small to large (6 to ca. Diagnostic features: axillary spines frequently present. smooth to sometimes papillose tails (caudate). plumose. never acute. achenes glabrous to weakly puberulent. glabrous. ascending. setaceous or rarely absent. red or purple. with “barnadesioid trichomes”. apex rounded. involucre corymbose to paniculate. remainder of style glabrous. small to large. rarely subshrubs. florets isomorphic or with transitional corollas in the same head. occasionally some segments partially connate. Sancho): Herbs usually perennial. orange. subprolate. and Brazil. actinomorphic. white with corolla lobes rolled. papillose tails (caudate). inner lip 2-cleft (rarely 3-cleft) and coiled or flexuous. bracts imbricate. often scapose and stout. shrubs or vines. heads ligulate. papillae rounded. pollen prolate. or absent. 3. purple. pollen prolate to spheroidal-subprolate. often glandular. without latex. instead they have “barnadesioid trichomes”. receptacle epaleate. absence of short. scabrate-microspinulate. without hyaline margins. achene oblong to sub-obovate. blue. leaves alternate. without latex. Mutisieae. Diagnostic features: two genera of woody plants of south-central South America with leaves clustered in brachyblasts. capitate glands (sesquiterpenes lacking). and involucral bracts often spine-tipped. heads discoid and one-flowered. styles slender. receptacle epaleate. 1–40 florets. exine almost psilate. anther apical appendages acuminate. the lobes completely coiled or only near the apex. marginal corolla when present. whitish. achenes densely villous. sub-bilabiate (4 + 1 corolla lips). yellow. tubular-funnelform. without latex. bracts imbricate. or homogamous and bilabiate. florets perfect. shallowly divided. or recurved. pollen psilate. and styles bifid and dorsally smooth. clustered in brachyblasts. Distribution: five species are endemic to Brazil and one is endemic to French Guiana. red or reddish. pappus of setae. outer lip 3-lobed. may be smooth or pitted or foveolate or clavate. erect. — Trees or shrubs. Diagnostic features: heads discoid. pappus of scabrid. or red. white. microechinate. the lobes equal or unequal in length. glabrous to pubescent. margins not hyaline. theceae bases calcarate and tailed (caudate). fi laments free or rarely fused. recurving when mature. or almost psilate. glabrous to pubescent. shallowly lobed. the tails long and free or connate with adjacent anthers. ligulate corollas with shallowly 5-dentate limb. anther apical appendage not constricted or demarcated from the thecae (rarely demarcated). shallowly bilobed to bifid with two long branches. pappus in a single whorl. shallowly divided. achenes prismatic or cylindrical or cylindrical-turbinate. scabrid to plumose. 100 capillary bristles in 4–5 series. attenuate to rostrate at the apex. — Trees. magenta. bilabiate. sub-bilabiate. heads discoid. . involucres cylindrical to funnelform. — (information provided by G. acute. deeply 5-lobed. rounded at the apex. style slender. Gongylolepis clade. corollas yellow. stalked. Distribution: Bolivia and Argentina. styles bifid. barbellate. style branches rarely recurving. branches dorsally papillose. cream. rarely bilabiate. usually dorsally papillose only above the style bifurcation point. heads radiate (less commonly disciform or discoid). 2-cleft inner lip that is often much coiled and so is shorter than outer lip. branches dorsally smooth. yellowish or reddish with ca. in 4–6 series. absence of twin hairs on achenes. bracts imbricate but sometimes subequal.Chapter 11: Classification of Compositae 177 spinulate. angled in cross-section. thecae rarely with spurs (calcarate) but with tails (caudate). Stifftieae Stifftia. minutely spinulose. or less commonly trailing shrubs or subshrubs. heads discoid or ligulate. corollas zygomorphic or bilabiate (some actinomorphic in Quelchia). involucre cylindrical or hemispherical or campanulate. pappus of scabrid bristles. never reflexed. white. or white with apices lilac. anther acute or acuminate at apex. or radiate. or homogamous and isomorphic (ligulate or bilabiate) or heteromorphic (ligulate and bilabiate). or red. theceae with spurs (calcarate) and with long laciniate. pappus showy. floret corollas white. 200). surface unknown. corollas whitish. disc corollas bilabiate or tubular. and styles glabrous. bisexual. Hyaloseris clade. 2. glabrous to densely pubescent. styles vary from thick to slender. smooth or rarely lophate. florets actinomorphic. Distribution: predominantly on the Guiana Shield with one monotypic genus in the Dominican Republic. apex rounded to shortly acute. leaves alternate to opposite. granulate. shortly bilobed or bifid. leaves alternate. achenes obovate. subimbricate.

and pappus of 2. 75 florets). styles thick. pink. corollas red. involucre cylindrical to campanulate or turbinate. spinulose. corollas creamcolored to yellow to magenta with five lobes of equal size. yellow. Distribution: Guiana Shield and Brazil. Nassauvieae. Distribution: predominately southern South America with outliers in Africa and Asia. anther apical appendage acute or truncate. leaves alternate. usually dorsally papillose only above the style bifurcation point. corollas tubular. glabrous. leaves alternate sometimes rosulate. 5–100 florets. bracts imbricate to sub-imbricate. thecae calcarate and caudate. glabrous. corollas bilabiate with lobes coiled. Hyalideae. anther apical appendage acute. involucre campanulate to hemispheric. bisexual or female. equal or unequal in length. base tailed. heads radiate. marginal corollas usually bilabiate. the only pubescence is on the arms. bisexual. glabrous. achenes elliptical to turbinate. — (information provided by G. or subshrubs without latex. — (information provided by L. Diagnostic features: marginal corollas usually present. usually in 3–10 series. Distribution: mostly southern South America. pappus of capillary. bisexual. Wunderlichieae. and pappus of 2–4-seriate bristles. without latex. achenes elliptic to turbinate. cylindrical. pappus of 2–4-seriate bristles. pollen psilate. shallowly divided (1. acute to acute-apiculate. thecae with spurs (calcarate) and tails (caudate). blue. never acute. subshrubs. Sancho): Small trees. 6. tails free to connate with tails of adjacent stamens. anther apical appendage apiculate. styles glabrous.178 Funk. orange. styles thick. — (information provided by G. leaves alternate. outer ones shorter. spheroidal and suboblate. purple. style branches never recurving. anther apical appendage acute or truncate.or 3-seriate. styles thick to slender. — Trees. ribbed. outer lip short or long. scabrose. shallowly lobed. lobes equal to unequal. Katinas): Annual or perennial herbs. psilate. receptacle naked or paleate. shrubs. pollen sub-prolate to prolate. pink. deeply 5-lobed. 4. calcarate) with long laciniate or papillose tails (caudate). anthers ecalcarate and caudate. corollas white. or purple. with the ectosexine and the endosexine clearly columellate. medium to large. villose. orange. spurs (calcarate). central florets one (Hyalis) to ca. one to more than one series. Distribution: South America and Asia. achenes fusiform. 7. arms with an apical crown of sweeping hairs. central corollas tubular. florets dimorphic or isomorphic. ray corolla bilabiate (rarely absent). or plumose bristles. ray corolla bilabiate with a 3-toothed outer lip and 1–2-toothed inner lip. sixty. deeply lobed. or subshrubs without latex. marginal florets. red. leaves alternate. when present. achenes prismatic and glabrous to pilose.5 mm). Stuessy and Robinson Diagnostic features: corollas often brightly colored. shrubs. white. corollas whitish. achenes elliptical to turbinate. the lobes equal in length and coiled. scabrid to smooth (occasionally plumose at the apex) capillary bristles. disc corollas with five lobes. alveolate. bracts imbricate or less commonly subimbricate. Sancho): Trees. styles smooth or slightly asperulous at apex. leaves alternate sometimes rosulate. Katinas): Herbs. sometimes scabrous (Leucomeris). the lobes short to long. heads discoid. Diagnostic features: woody. ray corolla bilabiate with a 3-toothed outer lobe and a 1–2-toothed inner lobe. dorsally papillose or dorsally rugulose to papillose much below the bifurcation point. anther apical appendage not constricted or demarcated. small to relatively large (5 to ca. heads discoid or radiate. bracts imbricate. heads . receptacle epaleate to strongly paleate. receptacle naked. radiate or discoid. elliptic. pollen prolate or subprolate. styles smooth or slightly asperulous at apex. anther apical appendages acute to acuminate. Distribution: southern South America but also in the Andes. 5. apex rounded. shrubs or vines without latex. apex rounded.or 3-seriate. rounded at the apex. actinomorphic corollas. thecae with spurs (calcarate) and tails (caudate). corollas often brightly colored. Gochnatieae. rarely yellow. leaves clustered near stem apex. disc corollas tubular with five lobes equal or unequal in length. or echinate with short spines. or bicolor white-purple. or absent. Diagnostic features: involucral bracts imbricate. small to medium size. and style shallowly bilobed to bifid. alveolate. receptacle naked. bracts imbricate. shrubs. shrubs. rarely rounded. Diagnostic features: all corollas bilabiate. which have a unique apical crown of sweeping hairs. bilabiate. disc florets perfect. papillae rounded. glabrous or pubescent. styles thick. exine psilate (to nearly psilate). pappus of 2. flat or alveolate. pappus of multiseriate bristles or setae. subshrubs. Susanna. rarely absent. scabrid or occasionally plumose at the apex capillary bristles with a reduced number of outer shorter bristles. paleaceous. disc corollas bilabiate or tubular. apex truncate. involucre cylindric to campanulate. — (information provided by L. disc corolla bilabiate (rarely with five lobes of equal size). sometimes dioecious. dorsally papillose or dorsally rugulose to papillose much below the bifurcation point. inner bracts longer than outer. glabrous or pubescent. purple. involucre cylindric to hemispherical. sometimes with staminodes (Ianthopappus). or perennial herbs without latex. rarely papillose on abaxial surface of style arms. heads homogamous or heterogamous. deeply cut. Onoserideae. green with darker or lighter margins but not hyaline. pollen exine distinctly bilayered. usually with an extension of the stigmatic surface (Ianthopappus). 8. rarely papillose. violet. anther apical appendages apiculate. thecae sagittate (spurs. style apex truncate. pollen psilate. apex rounded or subrounded.

Distribution: tropical and southern Africa. thecae with spurs (calcarate) and tails (caudate). 12. and Pakistan. Erythrocephalum and Gladiopappus) are rather different from this core group of Dicomeae and their inclusion in this tribe is provisional. cream to pink or mauve. deeply lobed. exine almost psilate. bilabiate corollas. receptacle naked. involucre campanulate to urceolate. bisexual or female. each group of 1–5 heads being subtended by a relatively large spiny whitish or greenish bract. leaves alternate. rarely up to 90 . Diagnostic features: woody plants with coriaceous leaves. margins not hyaline. heads that are single-flowered and re-aggregated on a secondary receptacle. deeply lobed. large (up to 1000 florets). India. obscurely 4–5-nerved. including Madagascar and two species range outward to the Arabian Peninsula. Distribution: restricted to high elevation areas (1500– 2000 m) in southern Nevada and adjacent California including Death Valley. 9. bracts imbricate without hyaline margins. heads discoid. styles slender. — (information provided by S. branches short (0. achene terete. without latex. bracts imbricate in 3–10 series. radiate or discoid (occasionally disciform). leaves alternate. glabrescent. marginal florets. and achenes broadly obconic without twin hairs. corollas white. usually with an extension of the stigmatic surface. apex rounded to subrounded. corollas white to orange (seldom pink. apices rounded. anther apical appendages lanceolate not elongate. involucre cylindric to fusiform. anthers apical appendage acuminate. Pasaccardoa. Distribution: South Africa. bilabiate. anther fi laments and styles. pollen spheroidal to prolate or elliptic. receptacle epaleate. The core group of genera (Dicoma. disc corollas tubular. isomorphic or dimorphic. pappus of six unequal. disc corollas with five lobes of equal size. and styles glabrous. shrubs or small trees without latex. anther tails with subacute to subrounded apex. style rather thick with very short branches. villose. and tails (caudate). large heads. sometimes recurving at maturity. thecae bases with spurs (calcarate) and slightly fi mbriate tails (caudate). alveolate. from southern United States to Argentina and the Caribbean (especially Cuba). small (1 floret). smooth to papillose. anther tails long tapering. anther apical appendages apiculate. pappus of scabrid to plumose bristles. heads discoid. calcium oxalate crystals not star-shaped. heads radiate. whereas Gladiopappus is characterized by phyllaries with rounded apex. rarely with apical acute sweeping hairs. Ortiz): Dwarf cushion-forming shrubs to small trees without latex. anther apical appendages apiculate. anther apical appendage acute to acuminate. Ortiz): Perennial herbs. micro-echinate or echinate. deeply divided.Chapter 11: Classification of Compositae 179 solitary (or 2–3) or several. +Oldenburgieae. Dicomeae. — Small to medium shrub without latex. lilac. rounded at the apex. small to large (ca. — (information provided by S. involucre hemispherical to obconic. involucre oblong to campanulate or turbinate. or purple). Ortiz): Dioecious trees or shrubs. unisexual florets among the bisexual disc florets. style branches separate and often curved. pluriseriate. thecae with spurs (calcarate). 5–300 florets). yellow. deeply or shallowly divided. alveolate. bilabiate or sub-bilabiate (rarely sub-zygomorphic). — (information provided by S. pollen sub-oblate to prolate. Pleiotaxis and Erythrocephalum can be distinguished by the presence of phyllaries not pungent with resin ducts. +Tarchonantheae. corollas white to reddish to greenish white. Macledium. corollas white.or 3-seriate.1–0. bracts imbricate often with hyaline margins: receptacle epaleate (rarely paleate). glabrous. with a subacute to rounded apex and apical or subapical acute sweeping hairs. styles slender and glabrous. pollen psilate and oblong. cream or pinkish-brown. styles thick. leaves alternate. the lobes equal in length. disc corollas often actinomorphic (rarely slightly zygomorphic). The remaining three genera (Pleiotaxis.5 mm). 10. central florets bisexual. style branches straight and adjacent. heads discoid or radiate. and short rounded style branches. echinate. pluriseriate. pollen sub-oblate to prolate. Diagnostic features: stiff holly-like leaves. rarely without them (ecalcarate). tricolpate pollen. pappus of scabrid to plumose bristles or scales. scabrid or occasionally plumose (at the apex) capillary bristles or less commonly some of them relatively flat. and Cloiselia) of this tribe is characterized by: pungent phyllaries without resin ducts (or ducts reduced). achenes obovate. presence of star-shaped calcium oxalate crystals in the corollas. Distribution: America. alveolate. when present. lanceolate or multi-toothed scales sometimes fused to form lacerate crowns. pollen slightly echinate. ray corollas bilabiate (inner lips coiled or not) or true rays with three teeth. usually shallowly divided. disc corollas tubular. pappus of uniseriate or 2. and achenes broadly obconic to cylindrical with twin hairs and without a carpopodium. disc corollas with five lobes of equal size. receptacle naked. Diagnostic features: marginal corollas. 11. deeply 5-lobed. variable in size. leaves alternate. homogamous or heterogamous. small (ca. achenes ellipsoid and with carpopodium. usually with staminodes. achenes often obconic but also narrowly oblong to broadly cylindrical with twin hairs. ray corollas bilabiate with the inner lip coiled lobes. when present. deeply divided. mainly coriaceous. +Hecastocleideae. not recurving at maturity. 30. bracts imbricate often without hyaline margins. marginal florets of the capitula being truly bilabiate with coiled adaxial lobes. thecae with tails (caudate). achenes narrowly ellipsoid to linear.

often subshrubs (less often shrubs or annual herbs. acute to somewhat rounded. anthers reduced or absent. rarely pilose. laciniate. clustered at the median part of the stem or on secondary short shoots. rarely with a few bisexual florets. appearing pseudo-bilabiate or pseudoligulate. dorsally short papillose. small pollen (30 µm). corollas irregularly deeply 5-lobed and therefore zygomorphic by having one sinus deeper than the others. branches bilobed to shortly branched. branches bilobed to shortly branched. dorsally pilose to shortly pilose. achenes usually with hardened pericarp. receptacle naked. 5-lobed but zygomorphic with one split deeper than the others. spherical or more prolate. blackish by presence of phytomelans. Distribution: widespread in Eurasia. at maturity style branches not recurving. distinctive leaves that are obovate. laticiferous ducts present in roots. alveolate. anther apical anther appendages narrow-triangular. and phyllary apices. Susanna): Perennial or monocarpic herbs. especially diverse in the Mediterranean region where they constitute an important characteristic of the vegetation. achenes densely long-setuliferous/sericeous. sinuses varying in depth. styles slender. deeply divided in five lobes of equal size. exine microechinate. pappus setae uniseriate. +Gymnarrheneae. one to ca. sometimes pseudo-bilabiate or pseudo-ligulate. long-barbellate. apices rounded. leaves alternate. dorsally pilose to shortly pilose. female florets with 3–5-lobed actinomorphic. glabrous or pilose.180 Funk. involucre campanulate. styles slender. subshrubs or shrubs. florets usually bisexual and fertile (rarely functionally female/male or cleistogamous). scabrate or echinate. flattened or not. anthers often with laciniate basal appendages. and one species also present in the Arabian Peninsula. 30 florets and heads that are arranged in generally dense racemes or panicles. shaft with some short hairs above the point where the style branches and below with a papillose-pilose thickening (functionally a pollen brush). — (information provided by A. anther fi laments usually papillose. heads discoid. — Shrubs without latex. corolla vestigial. and styles thick. . thecae bases sagittate (calcarate) and tailed (caudate) often with long divisions. Diagnostic features: involucral bracts usually in five rows and spiny-tipped. pollen spheroidal or subprolate. — Perennial herbs. rose or purple florets. seldom blue. often recurved. rarely naked. pappus of numerous barbellate bristles in 1–2 rows or lacking. florets enclosed in small involucral bracts. male florets with deeply 5-lobed actinomorphic. often purplish. leaves alternate forming rosettes. small (6 florets). tubular corollas. usually many-flowered (rarely one-flowered). thecae with spurs (calcarate) and tails (caudate). lignified. thecae with spurs (calcarate) and with tails long and smooth to pilose (caudate). involucre turbinate. alveolate or not. bracts in many rows. Distribution: southeast Asia. white. and stamens often strongly thigmotropic. pappus bristles 1–3-seriate (sometimes absent in functionally male or chasmogamous florets). usually actinomorphic.25 mm). ligulate with five lobes. commonly multiseriate (5–15 rows) to rarely uniseriate of few involucral bracts. often spiny (less frequently unarmed). usually tubular. and southern Africa. Madagascar. straight or s-shaped. female. more seldom emarginated. verrucate. pinnately veined and easily disarticulating. anther apical appendages extending into a rigid. long-acute. florets all fertile or the peripherals sterile. very rarely large shrubs or even treelets). Diagnostic features: dioecious trees or shrubs often aromatic with small discoid capitula of ca. — Amphicarpic annuals with no reports of latex. apices slightly rounded. receptacle epaleate. corollas yellowish or whitish. at maturity style branches not recurving. unisexual. achenes oblong to obovate. without sweeping hairs. more or less alveolate. Susanna. Diagnostic features: small discoid heads of white. without latex. involucre campanulate to obconic. pink or yellow. micro-echinate. Distribution: northern India and adjacent Nepal. oblate. glabrous. style characters reduced. achenes mainly cylindrical to ellipsoid. margins scabrid or plumose. often rosulate. scabrous. bracts scarcely imbricate. +Catamixis incertae sedis. serrate with large teeth. without hyaline margins. bracts imbricate. styles slender. branches relatively short (0. Diagnostic features: small ligulate heads. alveolate. receptacle scaly or more often setose. leaves alternate. thecae with spurs (calcarate) and tails (caudate). sometimes swelling slightly on upper part of style with branches recurved. 13. pollen spinulose with widely dispersed spinules ( < 1 µm in height). heads discoid (very rarely peripheral florets with 5-lobed ligules). flattened branches.or) 10-veined. leaves often spiny. cleistogamous. Distribution: tropical Africa (mainly in the eastern part). corollas purple. leaves alternate frequently forming a rosette. heads ligulate. acute to obtuse at the apex. corollas whitish to pale yellow. Cardueae. style with short. (9. often without hyaline margins. corollas white or pink. or echinate. Stuessy and Robinson florets). 15. pink. some species are cosmopolitan weeds. rounded or acute at the apex. usually imbricate with hyaline margins sometimes ciliolate. lanceolate appendage. pollen oblate-spheroidal to sub-prolate. pappus of scales or bristles. involucre cylindrical to obconic. 13(–16) florets. apices acute. pollen psilate. Pertyeae. tubular to fi liform corollas. style cylindrical with a thickened articulation below the branches bearing a short collar of stiff hairs. receptacle epaleaceous usually glabrous. gradate. subterranean heads homogamous. anther apical appendages truncate or rounded to apiculate. 14. anther apical appendage deltate. variously truncate. making up an elaborate mechanism of pollen presentation.

ribbed. inconspicuous anther collar. rosette shrubs and rosette treelets) with latex. broadly subulate. heads radiate or discoid. corollas yellow. sometimes absent. patent and broadly subulate sweeping hairs. shrublets. — (information provided by H. ray florets often brightly colored. vestigial. Robinson): Erect branching shrubs without latex. receptacle mostly naked. and herbs without latex. ray florets sterile. longer in a ring below the bifurcation. anther apical appendage oblong. dwarf desert annual. style arms long with rounded apex. ovoid. and disc corolla lobes with sclerified margins. 4-lobed. functionally staminate florets in small groups. much longer in a ring below the bifurcation. — (information provided by P. 17. endothecium without lateral wall thickenings or some polarized. corollas usually 4-lobed. thecae with spurs (calcarate) and tails (caudate). thecae with spurs (calcarate) and no tails (ecaudate). glabrous. wrinkled. pollen echinate. strap-shaped with three short teeth. somewhat scale-like bristles. leaves alternate frequently forming a rosette. sometimes papillose or hairy. involucre cylindric. — (information provided by N.O. leaves alternate. upper portion of style shaft and outer surfaces of branches covered with elongate sweeping hairs formed by two or . corollas 3–4-lobed. mostly differentiated into a few imbricate outer series of bracts and a longer inner series (rarely uniseriate).or biseriate scales.O.Chapter 11: Classification of Compositae 181 pappus absent. pollen spinulose with larger spines in intercolpi and smaller crowded spines around poles. each enclosed in a prominent. and pappus scale cells very long and with abaxially reinforced walls. +Eremothamneae. fi laments smooth. with 5-toothed ligule. achenes of three types. 3–4 anthers. small. collar usually inconspicuous. sweeping hairs small patent. branches narrow. anther apical appendage absent. inner with scarious tips. mostly ribbed. pappus of uni. sometimes winged. urceolate or obconic. involucre campanulate. receptacle naked. anther apical appendage short. the spines unevenlydistributed. heads radiate. and exclusive presence of lactiferous canals in both the subterranean and aerial parts. slender branches. receptacle deeply alveolate with projections along alveole margins. sterile ray florets. with spines or longitudinally striate hairs. deeply alveolate receptacles. rarely villose. white and green bract. — (information provided by P. cylindric. stiff. villous. oblate-spheroidal. outer with foliaceous tips. often somehow sculptured. corollas small. Diagnostic features: almost exclusive presence of homogamous capitula with 5-dentate. surface unknown. or fusiform to slenderly obconic. 18. heads ligulate (except for Gundelia and Warionia). achenes somewhat flattened. thecae without spurs (ecalcarate) and with tails (caudate). bracts multiseriate. bracts connate. apical anther appendage fi rm. involucre cylindric to campanulate. ray corollas when present. tiny. acuminate scales. leaves alternate. elongated. with one to over 600 florets but mostly with a few dozen. homogamous. interspersed among the small pistillate florets. shrublets or herbs with latex. sublophate or lophate generally oblate-spheroidal. ligulate flowers. Distribution: southern Africa. sweeping hairs on the shaft and branches. Arctotideae Arctotidinae. Kilian): Annual to perennial herbs (more rarely subshrubs. smooth. apiculate or broadly elongate. attenuate or beaked. frequently heteromorphic. amphicarpic. pollen echinolophate or echinate. soft. thecae calcarate and caudate. pollen echinate and non-lophate. leaves usually alternate. mostly glabrous. small (5 florets in Hoplophyllum) or medium (12–20 rays and 25–30 disk florets in Eremothamnus). apically thickened styles. distally papyraceous and usually with apical spine. gradate. usually female. whitish. herbaceous. with long twin hairs. endothecium radial. corolla predominantly of some shade of yellow or blue. pappus of scales or scabrid to barbellate or plumose bristles. those of pistillate florets numerous. Cichorieae. Diagnostic features: ephemeral. ciliate. involucre campanulate. style slender. achenes cylindrical. or of short. rounded. Distribution: mainly southern Africa. thecae with spurs (calcarate) and with (caudate) or without tails (ecaudate). sometimes compressed or flattened. smooth or shallowly honeycombed. and aerial heads clustered together and with functional male and female florets. Distribution: deserts in northern Africa and the Middle East. female florets solitary. usually ribbed. loosely connected on very short pedicels. anther apical appendage elongate. heterogamous. cylindric or urceolate. Diagnostic features: radial anther endothecium. bracts imbricate. heads generally radiate. perfect. styles slender. disc corollas with five linear lobes of equal size. achenes flattened. ciliate. bracts with or without hyaline margin. Karis): Shrubs. corollas 3-lobed. connate involucral bracts. aerial heads congested in the center of the leaf rosette. disc corollas shallowly 5-lobed. at maturity style branches not recurving. pappus usually of uni. disc corollas with sclerified lobe margins. Distribution: nearly global with centers of diversity in the Mediterranean area and north temperate climates. apically truncate. that of female florets of long-lanceolate. styles with clavate or subulate-ensiform sweeping hairs. naked (rarely with scales or bristles). pollen echinate.or biseriate scales. styles with a thickened apical portion. disciform. Karis): Shrubs. rather short. pappus of three types. both chasmogamous aerial heads and cleistogamous subterranean ones and two different types of achenes. Diagnostic features: latex. usually with long. corolla fi liform. shrubs. 16. receptacle areolate or alveolate. Gorteriinae.

styles slender. outer somewhat shorter. Distribution: tropical and southern Africa. shrubs. non-caveate. latex mostly lacking. Diagnostic features: complicated secondary head structures. occasionally whitish. corollas mostly blue to reddish or purple. involucre with bracts subequal to gradate. alveolate. pappus of many stout scabrid-barbellate capillary bristles in 2–3 series. vines or small trees. bracts somewhat imbricate without hyaline margins. heads radiate (3 to ca. style branches spreading tangentially. 22.to many-flowered. strap-shaped with three lobes. swollen and scabrous above near branches. otherwise glabrous. stolons that emanate from the crown at the base of the secondary head. 19. inner persistent or deciduous. pollen spherical. — (information provided by H. homogamous. leaves mostly alternate. at maturity style branches recurving. achenes prismatic. actinomorphic. and pollen echinate with spines in a regular arrangement or in irregularly arranged ridges surrounding the colpi in a unique bowtie pattern. mauve. bracts graduate. latex present in most taxa. or small trees usually with latex. lilac. . in Platycarphella apex slightly rounded and styles with a few distal hairs. anther apical appendage acute not elongate. somewhat gradate in length. often with T-shaped or stellate hairs.or 5-sided. or white). with baculae not aligned with spines. leaves arranged around edge of secondary head. bracts green-brown with short hairs and small hyaline margins. 320 florets). pappus capillary in ca. spines regularly and irregularly arranged. anther apical appendage acute. Diagnostic features: alternate leaves with pinnate venation. Liabeae. achenes oblong or columnar. with numerous twin hairs. receptacle alveolate. secondary head and leaves lying more or less flat on the ground. lobes elongate. — (information provided by M. heads with- out rays. Distribution: tropical America with the vast majority in Peru and Ecuador. Susanna. one. receptacle naked. 10–17-ribbed. pappus usually capillary. rarely winged. idioblasts and raphids obscure. heads discoid. lobes elongate. Robinson): Perennial herbs (few annuals). shrubs. in Platycarpha apex slightly tapered. Robinson): Shrubs without latex. dark. Diagnostic features: leaves usually opposite. branches terete. pollen echinate. and anther endothecial cells with thickenings on upper and lower ends. southeast Asia to Australia and the Pacific. with faint ribs. anthers apical appendages oblong-ovate. styles slender below. with hairs nearly to tip. thecae subequally sagittate (calcarate) and without tails (sometimes with very short tails) (ecaudate or caudate). sometimes coroniform or lacking. long sweeping hairs on outside extending onto upper style shaft. styles lavender. heads one to many. oblong. thecae spurred (calcarate) and short tailed (caudate). pollen echinate. or pink. Diagnostic features: slender style branches bearing relatively long sweeping hairs divided longitudinally into two or three cells. actinomorphic (rarely zygomorphic). pollen sublophate or lophate. heads with one or five florets. rich in sesquiterpene lactones. branches short. involucres cylindrical. leaves alternate. 5-lobed. 5-lobed (rarely with lobes grouped as four outer and one inner). achenes usually prismatic with 5–10 ribs. anthers purple. not caveate. apex attenuate. without hyaline margins. often with short outer series. corollas lavender to purplish. with long sweeping hairs on outer surface extending onto upper part of style shaft. two series. Vernonieae. pappus of persistent white scales. distally papyraceous and usually with apical spine. leaves usually opposite. without chaff. corollas yellow (rarely red. +Moquinieae. rarely with phytomelanin in walls. styles slender. spherical. achenes densely sericeous. homogamous or with anthers aborted in functionally female florets. leaves that are elongate and prostrate. +Platycarpheae. with continuous stigmatic surface inside. slender tapering style branches. disc corollas with five lobes of equal size. sometimes obcompressed. 20. Dillon): Annual or perennial herbs. glabrous. pollen spinulose with larger spines in intercolpi and smaller crowded spines around poles. Distribution: three species in southern Africa. — (information provided by H. rugose or smooth. Stuessy and Robinson three cells separated by longitudinal walls. non-lophate. and pollen echinate. corollas mostly blue to reddish or purple. usually sublophate or lophate. pollen spherical. purple. 21. at maturity style branches arched but not recurved. at maturity style branches not recurved. receptacles rarely paleate. one group with 5-alkyl-coumarins. without hyaline margins. echinate. achenes 3. gradate. pappus of numerous long inner capillary bristles and short outer series of squamellae. slender and tapering. and the Western Hemisphere. without phytomelanin. — Acaulescent perennial herbs without latex. mostly 35–70 μm in diameter in medium. Distribution: western South Africa and southern Namibia. corollas purple. anther apical appendage oblong-ovate. ray corolla present. achenes usually with twin hairs. corollas usually yellow. with or without hairs. and involucral bracts multiseriate. often tailed. sometimes with scales or plumose bristles or absent. sweeping hairs on upper style shaft and backs of branches. thecae with spurs (calcarate) and tails (caudate) fringed or digitate. involucre subglobose to cylindrical.182 Funk. often with perforated tectum partially or totally missing. with five lobes of equal size. branches in multiflowered heads spreading tangentially. usually with raphids. involucre campanulate. longer than wide. thecae usually spurred. scabrous outside.

. disc florets perfect or functionally male. deeply divided. involucre calyculate or ecalyculate. pink. thecae basally obtuse to sagittate or caudate. but most members of Tussilagininae have a continuous stigmatic surface. sweeping hairs on backs of style branches acute. involucre campanulate or cup-shaped. sometimes with hyaline margins. — (information provided by B. pappus of basally connate short scales and/or discrete fi ne bristles. thecae with short spurs (calcarate) and barely branched tails (caudate). lianas. enclosing the floret. apex tapering gradually to a slightly rounded point. non-caveate. corolla tubular or with a campanulate 4. endothecial tissue radial (Othonninae and Senecioninae) or polarized (Tussilagininae). style branches short. in Senecio and many other genera of Senecioninae the stigmatic surface is divided into two parallel bands. white or strawcolored. anther apical appendage reduced. Nordenstam): Scapose perennial herbs without latex. fi lament collar balusterform (Othonninae and Senecioninae) or cylindrical and straight (Tussilagininae). few-calyculate. polyacetylenes lacking. epiphytes or trees. and closely related tribes. Diagnostic features: ray florets that generally have staminoides and a filiform lobe ventrally in the sinus of the tube. various sizes. subulate. involucre broadly campanulate with 2–3 rows of bracts somewhat connate at base. — Shrubs or subshrubs without latex. strap-shaped with three teeth and four veins but also staminoides and a fi liform lobe ventrally in the sinus of the tube. pollen echinate. continuous stigmatic surface inside. corolla yellow or orange. shrubs. achenes terete or flattened. styles slender. non-lophate. pollen echinate with spines regularly or somewhat irregularly spaced. ray corollas female. caveate. the Andes (Peru to Colombia). leaves alternate. densely hirsute ovary. leaves alternate sometimes rosulate (rarely opposite). leaf blades elliptical to obovate. Distribution: the area occupied by the genus coincides with the Cape Floristic Region in South Africa. small achenes covered with dense white twin hairs. bracts 1–3-seriate. glabrous. 25. Diagnostic features: uniseriate involucre (although not universal). apex slightly rounded with short hairs. non-lophate. and Southeast Asia. ray corollas female fertile or sterile or neuter. pappus biseriate. without hyaline margins. receptacle naked or fi mbriate. Calenduleae. sweeping hairs in apical tuft or distributed abaxially on style branches. pollen echinate. receptacle naked. shrubs. receptacle naked. Senecioneae. pollen spherical. mostly in the Cape Floristic Region. somewhat longer hairs in a ring below the bifurcation. and elongated. sometimes absent. ray florets female. heads. styles slender. and partially connate involucral bracts. bristle-like scales of varying length. anthers without spurs (ecalcarate). corollas pink to purplish or white with five oblong to linear lobes of equal size. pollen echinate. or appendage. heads radiate. corollas yellow. or x = 19 (Heliantheae Alliance). sometimes with tuft. disc corollas with five lobes of equal size. 23. apical appendage flat. Nordenstam): Herbs. involucre cylindric. caveate. when a ray/disc arrangement occurs the ray corollas are 3-toothed ‘true rays’. thecae blackish. Group 2: Asteroideae clade Tribes contained in the subfamily Asteroideae usually have: stigmatic surface in two lines on inner surface of style branch. achenes without phytomelanin. winged or angled. leaves alternate or opposite. pappus of bristle-like scales which form a pom-pomlike shape in older heads. also with centers in Central America. red or purple. endothecial cells with thickenings at upper and lower ends. homogamous or gynodioecious florets. densely pubescent. but in ovules. apically scarious and laciniate. small achenes covered with dense white twin hairs. styles swollen distally with short scabrae. stout. singleflowered capitula. with non-aligned baculae. pappus of barbellate bristles. with pinnate veins. — (information provided by B. +Corymbieae. echinate with spines evenly and deeply separated. +Heterolepis incertae sedis. anthers four or five. Distribution: restricted to Brazil. fertile (rarely sterile). style bifurcate or simple. leaves alternate except for members of Heliantheae s. two bracts. anther apical appendages soft and oblong. red or blue. flat. pencil. radiate. Base chromosome number x = 9 or 10. terete or subcompressed. denticulate or hairy. heads discoid. bracts uniseriate and subequal or sometimes in two or more series. heads radiate. involucres of two bracts. of various sizes. style branches not recurving. Nordenstam): herbs. white. branches apically truncate or obtuse to conical. shaft slightly thickened below the style branches. up to 6 cm in diameter. disc florets usually actinomorphic and corolla lobes are as long as wide (shallowly lobed). purple. di-ester type pyrrolizidine alkaloids and the eremophilane types of sesquiterpene lactones.str.Chapter 11: Classification of Compositae 183 Diagnostic features: hairs simple. Distribution: three species endemic to South Africa. which have opposite leaves (some alternate). of 1–10 yellow-brown. raphids not in walls of achenes. sometimes ribbed. shortly sagittate (calcarate) and without tails (ecaudate). rarely a single scale. achene oblong.or 5-lobed limb. leaves alternate mainly rosulate. at maturity style branches recurving. singleflowered. short hairs on the back side of linear style branches and upper part of shaft. exine with columellae solid (rarely with internal foramina). or small trees without latex. anthers spurred. Diagnostic features: tufted parallel-veined leaves from a silky-hairy rhizome. without latex. receptacles shallowly alveolate. Distribution: worldwide with the most marked center of diversity in South Africa. disciform or discoid. 24. — (information provided by B.

stigmatic lines running up to base of lanceolate to triangular tip. nearly always flat or convex. leaves alternate. and Australia. subshrubs. imbricate and almost always with scarious margins and apex. styles fertile or sterile. rarely entire with characteristic aromatic smell. and the Mediterranean region. sometimes functionally male. 27. ray corollas white. involucre cylindric to campanulate (hemispheric). tubular or funnel-shaped. sometimes purplish-black-tipped. “gnaphalioid” pollen. involucral bracts imbricate in several rows. female outer florets generally fi liform or often absent. stems generally with fibers in phloem and without resin canals. terete or flattened. or discoid. pink. mucilage cells and resin sacs/ducts. and achenes small. pollen spinulose. central Asia. — (information provided by J. persistent or caducous. rarely lacking. tubular florets hermaphrodite or functionally male. pollen with a two-layered ectexine comprising an outer columellate layer and an irregularly interlaced basal layer (“gnaphalioid” type). New Zealand. Diagnostic features: involucral bracts with a papery. shrubs. with a 2-layered ectexine comprising an outer columellate layer and an irregularly interlaced basal layer. style hairs rather short and apically rounded. heads radiate. leaves usually alternate. leaves entire. sometimes curved. actinomorphic with 3–6 apical lobes. and South America. usually small to medium-sized (rarely large). entire. cartilaginous basal part (stereome). exine without baculae. heads disciform or discoid (rarely radiate). central disc corolla yellow or rarely whitish or reddish.184 Funk. Gnaphalieae. sometimes with outer scales or awns. disc corollas hermaphrodite. without latex. achenes compressed and 2-nerved or angular to terete and multi-nerved (rarely obcompressed). stigmatic lines running up to base of lanceolate to triangular tip. subshrubs. bracts in 2–7 rows. subshrubs or shrubs without latex. bracts in 3–5 rows and usually imbricate. sometimes nearly equal. Susanna. Brouillet): Annual or perennial herbs. thecae usually ecalcarate and ecaudate (rarely shortly tailed). pappus absent. 5-lobed. exocarp sometimes fleshy and colored. entire or bilobed. rostrate. 26. thecae without spurs (ecalcarate) but with tails (caudate). thecae rarely with basal tail. and unusual fruit structures such as fenestrate cavities and a fleshy exocarp. Distribution: mainly Africa with a distinct center in South Africa. forming a split cylindrical or balusterform anther collar (fi lament collar). heads various. or pink to blue or purple. generally epaleate. disciform. widespread heterocarpy. 28. style branch with acute appendage. Diagnostic features: leaves generally variously dissected. in one to several rows. Distribution: mainly in southern Africa. and Anatolia eastwards to Iran. receptacle generally flat to convex. or shrubs. . pinkish or reddish). anther apical appendages flat. with five or four (rarely three) equal lobes. or lacking. Distribution: worldwide with major centers of diversity in Africa. perfect or functionally male. scabrous or plumose on outer surface. corolla with 0–5 lobes. sometimes conical or peg-like. Distribution: centers of diversity in southern Africa. purple or blue. pollen echinate (rarely rugose or smooth) and spherical. and style branches pronate. tricolporate and echinate. Anthemideae. generally with a papery upper part (lamina) and a thickened. often with glandular hairs. sterile styles in many taxa. anthers tailed. Stuessy and Robinson yellow to orange or white. not calyculate. Diagnostic features: lack of pappus. yellow. size varies. only scales. limbs with three lobes. pappus generally of plumose or barbellate to scabrid capillary bristles (occasionally of bristles and scales. sweeping hairs on entire outer surface of appendage. with sweeping hairs in a subapical collar or rarely extending down the style branches. with acute appendage. scabrous or plumose on outer surface. disc corollas yellow to white. only Calendula extending outside Africa into Macaronesia. and achenes without a carbonized layer. style branches pronate. achenes various. outer disc florets (if present in disciform capitula) yellow. rarely paleate. style slender. leaves alternate (rarely opposite). generally rounded. receptacles flat. North America. hemispherical or conical. with tails and with endothecial tissue almost always polarized. style branches with hairs apically or sometimes apically and dorsally (rarely dorsally only). style slender. achenes generally small and oblong to obovoid. bisexual or functionally male. glabrous or hairy. central florets generally perfect. Ober prieler): Herbs. often brightly colored lamina and a cartilaginous basal part (stereome). without well-developed hyaline margin. involucres often hemispheric. achenes homoor heteromorphic. squamose or fi mbrilliferous. endothecial tissue polarized. or absent). anther apical appendage flat triangular-ovate. pappus usually of 1–4 series of barbellate or rarely plumose bristles. no tails on anthers. usually strap-shaped (rarely bilabiate). Diagnostic features: receptacle usually naked. Australia. upper part of the fi lament with cells with thickened walls. yellow to orange or reddish. bracts in (2–)3–5(–9) rows. anther thecae without spurs. apex truncate with parallel stigmatic surfaces. Astereae. usually without latex. — (information provided by L. anther apex usually ovate or triangular. lanceolate to deltate. Ward): Herbs. pappus never capillary. fi liform to funnelform or with abruptly ampliate limb. receptacle usually naked. paleate or epaleate. South and Central Europe. — (information provided by C. usually imbricate. ray corollas white and/or yellow (rarely blue-violet. usually female. rarely trees or vines. winged or fenestrate. involucral bracts with hyaline margin. pollen spherical.

31. Diagnostic features: white hermaphroditic florets. styles slender or thick. markedly lobed ray corollas. pollen echinate and spherical. caveate with one baculate layer. involucre cylindric to hemispheric. heads disciform. — (information provided by B. anther apical appendage acute. pollen spinulose. bracts without hyaline margins but with median resiniferous duct. . and style frequently with obtuse sweeping hairs extending below style bifurcation. anther apical appendage acute.Chapter 11: Classification of Compositae 185 29. pappus of scales or bristles (rarely absent). receptacle naked. sterile. large crystals. achenes without large elongated crystals. leaves alternate. strapshaped with three teeth. achenes more or less clavate. some achenes have phytomelanin. spinulose. branches usually truncate. at maturity style branches slightly reflexed. anther apical appendage ovate to round. Distribution: warm climate areas worldwide. apex rounded to acute. apex blunt. Heliantheae Alliance Information for all tribes of the Heliantheae Alliance was provided by B. Diagnostic features: involucral bracts usually 1–3-seriate. walls lack phytomelanin and usually include few. Diagnostic features for Inulinae: heads usually radiate and yellow-flowered. of awns. tubular. corollas yellow or white to red or purple. baculate with ramified basal layer. with terminal tuft of hairs otherwise glabrous or papillose. 30. Anderberg): Vine without latex. involucre cylindric. disc corollas with five lobes of equal size. often much reduced. awns or missing. Baldwin and A. achenes subcylindrical. of scales only. style branches recurving at maturity. rarely truly paleate) and sometimes alveolate. — (information provided by A. disciform with fi liform. pappus of scales. tubular. walls without large elongated epidermis crystal and without phytomelanin. pistillate or sterile. leathery entire leaves. in few rows. caveate. discoid or radiate. Distribution: Western Hemisphere. violet or white. walls without phytomelanin. strap-shaped with three lobes. leaves alternate. thecae generally without spurs (ecalcarate) but often with distinct tails (caudate). lobes mostly shorter than wide. style appendages usually shorter than stigmatic portion (except in Eupatorieae). receptacle usually naked (sometimes with palea-like receptacular outgrowths. heads radiate or discoid. involucre cup-shaped to cylindrical. styles usually with sweeping hairs above where the style divides. styles slender. Anderberg): Shrubs or herbs without latex. Inuleae. bracts imbricate without hyaline margins. generally in several rows. receptacle naked or paleate. campanulate or cup-shaped. prismatic or quadrangular. glabrous to densely hairy. — Annual to perennial herbs (rarely shrubs) without latex. disc corollas yellow or red to purple. leaves alternate. corollas white or yellow. large but few-flowered (9–12 florets). pollen spherical. tubular florets. elliptic. at maturity style branches recurving. without internal foramina. and base chromosome number x = 19. margins (at least of outer) usually not hyaline or scabrous. — (information provided by A. glabrous. strap-shaped with three lobes. heads discoid. pollen spinulose. pappus of capillary bristles. multiseriate. multiseriate involucre. often with glandular hairs and/or non-myxogenic twin hairs. ray corolla when present. thecae not blackened. Diagnostic features: heads sometimes in pseudocephalia. Distribution: confi ned to Cuba. achenes generally homomorphic. usually epaleate receptacles. hairs inconspicuous. outer florets often fi liform. involucre cylindrical. bracts in two or more series. Distribution: mainly Africa and Australia. but also in East Africa. Saharo-Sindian. pink. apex slightly rounded with short obtuse sweeping hairs above where the style divides. tubular. small to medium with up to 45 florets (except for Anisopappus which can have 100’s of florets). with short acute or obtuse sweeping hairs above or extending below where the style divides. most taxa with phytomelanin layer in the achene. corolla white with five lobes of equal size. and anthers with tails. sometimes reflexed. Helenieae. +Feddeeae. lobes usually not longer than wide. branches elongate. if present. Athroismeae. Diagnostic features for Plucheinae: heads often purple-flowered. radiate. Baldwin unless indicated otherwise. achenes without large epidermal crystal. small to large ( > 100 flowers). mostly medium-sized to large. ray florets. leaves alternate (rarely opposite). 32. and acute sweeping hairs on style branches. ray corolla when present. sometimes small. outer florets often fi liform. heads. Anderberg): Shrubs or herbs without latex. disciform or discoid. disc corollas with five lobes of equal size. anther thecae often blackened. at maturity style branches recurving. without spurs (ecalcarate) and with or without short sterile tails (caudate or ecaudate). anther apical appendage acute. Diagnostic features: combination of alternate leaves. at least as terminal tuft. receptacle naked or paleate. or absent. styles slender. corollas yellow. pappus of uniseriate tawny capillary bristles. thecae normally without spurs (ecalcarate) but generally with distinct tails (caudate). caveate with one baculate layer. and bracts with resiniferous duct. of bristles and scales. bracts imbricate without hyaline margins. styles slender or thick. thecae without spurs (ecalcarate) and with long tails (caudate). with five lobes of equal size. or with irregularly interlaced inner layer. Distribution: mainly Mediterranean. without spurs (ecalcarate) or tails (ecaudate). achenes with elongated crystal in each epidermal cell.

heads usually radiate (rarely discoid). tubular or strap-shaped. Chaenactideae. styles slender. then achenes usually strongly 9–15-ribbed. receptacle usually naked. Distribution: warm temperate to tropical New World (1 sp. most with glandular pockets (dark spots).186 Funk. bracts 1–2-seriate without hyaline margins. anther apical appendage ovate to deltate. discoid heads. mainly California. mostly southwestern North America. thecae usually blackened. corollas of peripheral florets sometimes zygomorphic. leaves alternate or opposite (rarely whorled). spherical. usually 3-lobed. receptacle with brownish-orange striated paleae. apices of style branches truncate or deltate to acuminate. — Annual or perennial herbs or shrubs (rarely trees) without latex. 37. rarely narrowed toward the apex and beaked. paleae with brownish-orange striations. involucre cylindric to more or less broadly campanulate. apices more or less acute with terminal tuft of hairs. flat to conical. paleae tightly enfolding achenes in aquatic or marsh-dwelling taxa. at maturity style branches recurving. corollas white to reddish or yellow with five lobes of equal or unequal size (peripheral lobes sometimes enlarged). recurving at maturity. anthers usually blackened. corollas mostly yellow or white. heads radiate or discoid. pappus of setose to obovate scales without thickened bases or midribs and sometimes fused at base into deciduous unit. — Annual to perennial herbs (rarely shrubs) without latex. 2–15 cm wide. small to medium-sized. pappus of scales and/or bristles (rarely absent). with or without hyaline margins in 1–4 series. Diagnostic features: epaleate receptacles. receptacles usually naked. antrorsely or retrorsely barbed bristles or short awns. smooth. small to medium-sized. inner with brownish-orange striations and scarious margins. epaleate receptacles. pappus absent or of scales. ray florets pistillate. anther apical appendages usually sclerified. apices usually acute with short hairs. Tageteae. achenes blackened. pistillate. disc corollas with (4–)5 lobes of equal size. thecae without spurs (ecalcarate) and short-tailed (caudate). sometimes deciduous as a unit. awns. bracts 1–8-seriate without hyaline or scabrous margins. bracts usually in two or more continuous series. — Annual or perennial herbs or shrubs (rarely trees) without latex. bracts in 1–5 series. heads radiate or discoid. 35. Craw ford and M. — (information provided by D. styles slender. Stuessy and Robinson truncate style branches. outermost green. more or less terete (not flattened). Diagnostic features: stems fistulose and rooting at nodes or not. thecae not blackened. leaves usually alternate. receptacle usually paleate. involucre cylindric to hemispheric. Diagnostic features: combination of alternate. without spurs (ecalcarate) and without tails (ecaudate). pollen radially symmetrical. with phytomelanin layer. heads discoid. corollas yellow to red or white. mostly small to medium-sized. sometimes large. sometimes pink to purplish. involucre cylindric to hemispheric. characteristic smell (monoterpenes). and if glands lacking. ray florets. apex penicellate or truncate with fringes of papillae. Distribution: centered in North and South America 34. pollen echinate and spherical. receptacle naked (rarely partially paleate). achene walls striate. ray florets neuter or pistillate and then either sterile or fertile. without spurs (ecalcarate) and without tails (ecaudate). — Annual or perennial herbs (rarely subshrubs) without latex. Bahieae. usually variously papillose. pollen echinate and spherical. thecae not blackened. Coreopsideae. with phytomelanin layer. glabrous or hairy below style bifurcation. echinate and tricolporate. pollen echinate and spherical. 36. leaves alternate or opposite. size varies. leaves alternate or opposite. leaves. strap-shaped with usually . tubular. apical appendage ovate. remainder of style glabrous or papillose. corollas usually yellow to orange or white. strap-shaped with 2–3 lobes. walls with phytomelanin layer. receptacle usually paleate. usually with five lobes of equal size not longer than wide. Diagnostic features: outer phyllaries differentiated from inner in color and shape. if present. yellow to white. usually ovate. if present. at maturity style branches recurving. and unribbed and unthickened pappus scales. involucre cylindric to hemispheric. non-carbonized achenes (with large crystals in achene-wall cells). achenes usually not flattened. style undivided or shortly cleft to bifurcate. pale or reddish (not blackened) anthers. striate achenes. achene walls striate or ribbed. recurved at maturity. Neurolaeneae. bracts imbricate. Susanna. isopolar. branches conic with short papillae on the outer surface. of Flaveria in Australia). disc corollas yellow with 5(–6) lobes of usually equal size or 1–2 lobes larger than others. smooth or pitted. styles slender. ray florets. Distribution: New World (mainly southwestern North America). Distribution: western and southwestern North America. 33. often lobed. striations dark green or brownishblack. anther apical appendage more or less round to ovate. sometimes coroniform. pappus of 2–15 smooth. corollas highly reduced or well-developed. inner phyllaries with orange-brown striations (resin ducts). heads radiate or discoid. anthers brown to black. disc corollas yellow. involucre cylindric to obconic or hemispheric. rarely absent. or bristles. and scaly pappus. Distribution: mainly Neotropics (few species in Paleotropics). pistillate. and achenes flat obcompressed to quadrangular. achenes flat to quadrangular. without spurs (ecalcarate) and without tails (ecaudate). linear to lanceolate. Tadesse): Herbs or shrubs without latex. leaves opposite or alternate.

with phytomelanin layer. pappus of awns or scales. pappus of scales with thickened base or midrib (if scales not thickened. sometimes absent. Millerieae. achene walls usually striate. with phytomelanin layer. without latex. tubular with five equal lobes. without spurs (ecalcarate) or tails (ecaudate). limbs 3-lobed. apices often terminated by tuft of hairs. then disk corollas 4-lobed) or of brownish-to-reddish. pollen echinate and spherical. blackened. apices deltate to acuminate with short hairs. achenes compressed. functionally staminate. small to medium-sized. and radially-arrayed (or absent) pappus scales or bristles. usually subequal phyllaries (often with one . sterile. Diagnostic features: combination of usually opposite. often glandular leaves. at maturity style branches not recurving. or terete. middle lobe longer and wider than lateral lobes. achene walls usually striate. Madieae. pollen echinate and spherical. branches tapering with short hairs. thecae usually blackened. styles slender. strap-shaped with three lobes or limb sometimes absent. 41. obcompressed. styles slender. branches truncate to subulate with short hairs. involucre cylindric or globose. with phytomelanin layer. with 4–5 lobes of equal or unequal size. or trees without latex. Distribution: eastern North America. anther apical appendages deltate. anther thecae pale. remainder of style glabrous. anther apical appendage usually lanceolate to ovate. functionally staminate disc florets. anther apical appendage usually ovate. leaves alternate or opposite. thecae without spurs (ecalcarate) and without tails (ecaudate). Diagnostic features: combination of paleate receptacles. corollas yellow with (4–)5 lobes of equal or sometimes unequal size. heads radiate or discoid (rarely disciform). receptacle naked or paleate (paleae often restricted to periphery of receptacle). pappus absent or coroniform. pappus of scales or bristles. pale anthers. — Annual or perennial herbs or shrubs. pale or reddish (not blackened) anthers. at maturity style branches recurving. the style glabrous below the bifurcation. disc corollas yellow or white. styles slender. often foliaceous. often sagittate (rarely with short tails = caudate). achene walls often striate. leaves opposite. sometimes pale (green in Guardiola). usually striate. involucre campanulate to hemispheric. corollas yellow with five lobes of equal size. ray florets pistillate (rarely sterile). achenes dorsiventrally compressed. paleate receptacles. involucre cylindric to hemispheric. disk florets bisexual or functionally staminate. styles slender.Chapter 11: Classification of Compositae 187 2–3 lobes. blackened achenes. striate. shrubs. more or less terete. sometimes large. Heliantheae. opposite. sometimes clasping ray ovaries. apices of branches often acute to penicillate. +Polymnieae. smooth. with phytomelanin layer. often fasciculate or hooked bristles. sometimes pink to purplish. styles slender. trees. achene walls smooth or striate. and pappus usually of awns or scales. thecae usually blackened. bracts in 1–7 series. ray corollas white. remainder of style glabrous. achenes usually more or less terete. achenes compressed (rarely obcompressed). Distribution: mainly Mexico and northern Andes (also Old World tropics. disc corollas yellow with five lobes of usually equal (rarely unequal) size. and South Pacific). ray corolla usually yellow or white. usually blackened anther thecae. Diagnostic features: combination of epaleate heads. 38. or whorled. usually without spurs (ecalcarate). involucre obconic to hemispheric. strap-shaped with usually three lobes. Diagnostic features: combination of often glandular foliage. sometimes coroniform (rarely absent). receptacle paleate. disk florets yellow. style below bifurcation usually glabrous. pollen echinate and spherical. strap-shaped with usually three lobes. flattened. corollas usually yellow or white. with paleae enfolding ovaries. achene walls smooth. without spurs (ecalcarate) or tails (ecaudate). 39. leaves usually opposite. remainder of style glabrous (except in Blepharipappus). anther thecae pale or purplish. size varies. bracts usually subequal in 1–2 series (rarely gradate in 3–4+ series). leaves alternate. Diagnostic features: combination of opposite leaves. usually blackened anthers. — Annual or perennial herbs. ray florets if present pistillate or sterile. or vines without latex. without hyaline margins. style apices terminated by tuft of papillae. size varies. shrubs. receptacle paleate. 40. anther apical appendage rounded or usually more or less ovate to deltate. scarious paleae. or vines. heads radiate (rarely discoid). corollas usually yellowish. — Annual or perennial herbs. without spurs (ecalcarate) and tails (ecaudate). tropical Africa. at maturity style branches erect or recurving. bracts in 1–5 series. with phytomelanin layer. ray florets pistillate. heads radiate. and pappus usually of basally thickened or costate scales or of fascicled or hooked bristles. branches with separate or fused stigmatic lines. Distribution: mainly warm temperate and tropical New World. bracts imbricate without hyaline margins in 2–3 series. especially Africa). and uncompressed smooth achenes. deeply divided or sometimes undivided. pollen echinate and spherical. heads radiate or discoid (rarely disciform). pollen echinate and spherical. mostly small to medium-sized. — Perennial herbs without latex. usually unflattened achenes. pappus absent or of scales or bristles. receptacle paleate (rarely naked) with paleae enfolding achenes and usually persistent. trees. with short hairs. achenes blackened. sometimes white (rarely orange to reddish). at maturity style branches recurving. Distribution: mainly southwestern North America (also temperate and montane South America. anther apical anther appendage usually ovate. disk florets bisexual or functionally staminate.

B.). Phylogenetic studies in Compositae tribe Mutisieae. American Midland Naturalist 53: 478–511. H. usually 4-lobed disc corollas. Bremer. & Turner. styles slender. Funk. & Hooker. 1966. (eds. The Biology and Chemistry of the Compositae.H. Harborne. an ent-kaurine diterpene glycoside. — (information provided by H. apical appendages not recurving. epaleate receptacles. Schilling. K. R. some monoterpenes and sesquiterpene lactones. Bentham. involucre cylindric to hemispheric. 1987. Asteraceae: Cladistics & Classifi cation. Aliso 8: 465–492. The Compositae revisited. without latex. scrambling shrubs or small trees. Chan. usually with twin hairs. Levrault. Genera Plantarum. Wood anatomy of Compositae. between ray and disc florets). Wessa. 163–533 in: Bentham.. Bremer. Cassini. at maturity style branches recurving. blue or lavender. 1991. mostly 18–25 μm in diam. Nuclear rDNA evidence for major lineages of helenioid Heliantheae (Compositae). Leiden. 1977. chromenes. Opuscules Phytologiques. anther appendages hollow or poorly developed. kolavane derivatives.D. V. leaves usually opposite. anthers often with glands. blackened achenes.. benzofurans. pappus usually uniseriate. Bentham. 1955. Perityleae. G. receptacles usually naked. 1994. actinomorphic (rarely peripheral florets with outer one or three lobes enlarged.. often deeply 3-lobed ray corollas. without raphids. Susanna. echinate. 1826. — Annual or perennial herbs or shrubs of rocky sites. Le Francq van. corollas yellow or white. E. J. pale. H. anther apical appendage usually ovate. florets one to many in a head. Baldwin. 1873a. achenes 4–10-ribbed with phytomelanin in walls. Other diagnostic features: mostly opposite leaves. corollas actinomorphic and whitish to reddish or lavender (never yellow). G. of plumose bristles. Literature cited Baldwin. vol. .L. heads radiate or discoid. 43. K. V. Eupatorieae. Tableau exprimant les affinités des tribus naturelles de famille des Synanthérées. with phytomelanin layer. & Panero. A. S. Berkhey. corollas white to reddish. Distribution: mainly deserts of southwestern North America (also Andes and Desventuradas Islands of South America).). Pp. B. 2. subequal. pollen echinate and spherical. without latex. Academic Press.. Tribal interrelationships of the Asteraceae. London. Garcia-Jacas. Cassini.or 5-lobed). A.188 Funk. R. Henri Cassini.. with or without hairs. caveate. Portland. Opera Botanica 109: 1–50.... and pappus (if present) usually of (1–)2 bristles and rudimentary scales.K.V. greatly enlarged. Compositae. Distribution: mainly western North America (especially California).. Cronquist. J. 1873b. Cronquist. reduced or lacking. Stuessy and Robinson phyllary per ray ovary and with phyllary at least partially clasping ray ovary).A.B. achene walls smooth. N. J. (in medium). Distribution: Western Hemisphere with a few pantropical genera. thecae not spurred or tailed. history and geographical distribution of Compositae.L. 2005. often dark purple (not blackened) anther thecae. disc corollas yellow with 4–5 lobes of equal size. Brittonia 29: 137–153. 1–2. H. epaleate or often partially paleate receptacle (with paleae limited to periphery of receptacle. Journal of the Linnean Society. also with pentaynene acetylenes. pollen spherical.. G.L. leaves opposite or alternate. bracts persistent to totally deciduous. Aliso 6: 25–44. Diagnostic features: all members of the tribe (with a few exceptions) have mono-ester type pyrrolizidine alkaloids secreted by nectaries. Paris. Everywhere but Antarctica: using a supertree to understand the diversity and distribution of the Compositae. Watson. Expositio Characteristica Florum qui Dicuntur Compositi. ed. Reeve. B. A summary. involucre campanulate to cylindrical. blackened disc achenes. 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