APIIDAE M. J. Donoghue et P. D. Cantino

Donoghue et Cantino in Taxon 56: E31. Aug 2007


[Campanulales+[Escalloniaceae+[Bruniales+Dipsapiidae]]]


CAMPANULALES Juss. ex Bercht. et J. Presl

Berchtold et Presl, Přir. Rostlin: 253. Jan-Apr 1820 [‘Campanulaceae’]

Habit Usually bisexual (sometimes monoecious, gynomonoecious, andromonoecious, polygamomonoecious, dioecious, gynodioecious, or androdioecious), usually perennial, biennial or annual herbs (sometimes evergreen, rarely deciduous, suffrutices, shrubs, trees or lianas). A large number of species are xerophytes and some species are succulent; others are aquatic or helophytes. C4 and/or CAM physiology sometimes present.

Vegetative anatomy Phellogen ab initio usually superficial (sometimes cortical). Vascular tissue atactostele-like, with scattered stem bundles. Cortical and/or medullary vascular bundles sometimes present. Endodermis sometimes with thick-walled cells. Cambium storied or non-storied. Secondary lateral growth usually normal or absent (sometimes anomalous via concentric cambia or cylindrical cambium). Vessel elements usually with simple (sometimes scalariform or reticulate) perforation plates; lateral pits usually alternate (sometimes opposite, rarely scalariform), usually bordered (sometimes simple) pits. Vestured pits present. Imperforate tracheary xylem elements libriform fibres, tracheids or fibre tracheids with simple or bordered pits, septate or non-septate (often also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular or heterocellular, or absent. Axial parenchyma apotracheal diffuse or absent, or paratracheal scanty, aliform, lozenge-aliform, winged-aliform, confluent, vasicentric, or banded (sometimes absent). Wood elements sometimes storied. Tyloses sometimes frequent. Intraxylary phloem rarely? present. Sieve tube plastids S type. Nodes usually ≥3:≥3, trilacunar, pentalacunar or multilacunar with three or more leaf traces (rarely 1:1, unilacunar with one trace). Phloem sometimes with articulated laticifers containing coloured, white or colourless latex rich in triterpenes and/or tissues with schizogenous secretory resinous canals often lined with epithelial cells (scattered latex cells sometimes present as well as resinous canals). Sclerenchymatous idioblasts often present. Parenchyma sometimes with prismatic calciumoxalate crystals (acicular crystals, druses, styloids, crystal sand, etc.).

Trichomes Hairs unicellular or multicellular, uniseriate or biseriate, simple or branched, flagellar hairs, T-shaped, stellate, candelabra-shaped, dendritic, arachnoid, peltate, lepidote, or vesicular; glandular hairs with unicellular or multicellular stalk and unicellular or multicellular head present or absent; laticiferous hairs sometimes present.

Leaves Usually alternate (spiral or rarely distichous; sometimes opposite, rarely verticillate), usually simple (sometimes pinnately or palmately compound), entire or pinnately lobed (sometimes repeatedly pinnately lobed), sometimes coriaceous, with conduplicate, supervolute, revolute or involute ptyxis. Stipules absent; leaf sheath usually absent. Petiole vascular bundle transection usually arcuate (sometimes annular). Venation pinnate, palmatopinnate, parallelopinnate, parallelodromous, craspedodromous, semicraspedodromous, camptodromous or palmate (rarely flabellate or acrodromous). Stomata usually anomocytic (sometimes anisocytic or helicocytic, rarely paracytic). Cuticular wax crystalloids when known partly as large glabrous more or less inrolled scales, partly as reticulate to annular threads or small scales (sometimes rodlets or platelets). Secretory cavities (with resin and/or latex) present or absent. Mesophyll sometimes with idioblasts containing sclereids. Leaf margin serrate, serrate-dentate, crenate, lobate or entire, sometimes glandular serrate or with hydathodes. Extrafloral nectaries rarely present.

Inflorescence Terminal or axillary, fasciculate, paniculate, corymbose, raceme-, spike-, head- or umbel-like, or raceme, dense capitula, or capitulate pseudanthia with involucre of bracts (rarely helicoid cyme; flowers sometimes solitary axillary, rarely solitary terminal).

Flowers Actinomorphic or zygomorphic (sometimes resupinate). Hypanthium sometimes present. Usually epigyny (rarely half epigyny or hypogyny). Sepals (two to) five (to eight), with valvate, imbricate or open aestivation, usually persistent, often connate, or modified into usually accrescent scales or hairs, pappus, or absent. Petals (three to) five (to ten), usually with valvate (rarely imbricate or open) aestivation, usually caducous, usually more or less connate into tubular, urceolate, campanulate, infundibuliform, spathulate, or (uni- or) bilabiate (rarely free) corolla, sometimes fringed (rarely absent), with apex inflexed. Nectariferous or non-nectariferous disc intrastaminal, annular, scale-like, tubular, or absent (sometimes with nectariferous glands alternating with stamens).

Androecium Stamens (two to) five (to ten), haplostemonous, antesepalous, alternipetalous. Filaments usually free from each other (sometimes connate), usually adnate to petals (epipetalous). Anthers free (sometimes connivent) or connate into tube around style, basifixed or dorsifixed, usually non-versatile, usually tetrasporangiate (rarely disporangiate), usually introrse, longicidal (dehiscing by longitudinal slits); introrse anthers in Asteraceae and in some Goodeniaceae and Campanulaceae entirely or partially connate, syngenesy. Tapetum ab initio cellular, with multinucleate cells, later usually amoeboid-periplasmodial by dissolution of cell walls, or secretory with binucleate or multinucleate cells. Female flowers with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3(–6)-colporate (sometimes tri- to polycolpate, rarely tri- to polyporate), usually shed as monads (rarely tetrads), usually tricellular (sometimes bicellular) at dispersal. Exine tectate or semitectate, with columellate infratectum, punctate, perforate, microreticulate, or lophate, often caveate (fenestrate; sometimes reticulate, rugulate or striate, rarely imperforate), echinate, verrucate, spinulate, scabrate, psilate, or granulate.

Gynoecium Pistil composed of (one or) two (to five) connate carpels. Ovary usually inferior (rarely semi-inferior or superior), unilocular or bilocular (to quinquelocular), sometimes incompletely septate. Style single, usually bilobate or simple (rarely trilobate), usually hairy. Stigma(s) capitate, truncate, clavate, discoid, or stigma bilobate (rarely trilobate or quinquelobate), often with stigmatic surface on adaxial side of lobes (sometimes with appendages on apex of stylar branches), usually papillate, Dry or Wet type. Male flowers with pistillodium. Secondary pollen display with several different types of stylar adaptations present in Asteraceae, Calyceraceae, Goodeniaceae and Campanulaceae, such as pollen pump mechanisms, plunger or brush pollination, pollen collecting hairs, cupular structure outside gradually and strongly expanding style, and stigma late pollen receptive.

Ovules Placentation basal or axile (sometimes apical, rarely parietal or free central). Ovule one per ovary, or two to numerous per carpel, usually anatropous (rarely campylotropous or hemianatropous), ascending (sometimes pendulous or horizontal), epitropous (sometimes apotropous), unitegmic, tenuinucellar. Integument rarely vascularized. Hypostase absent. Endothelium present. Archespore usually unicellular (sometimes multicellular, rarely bicellular). Megagametophyte usually monosporous, Polygonum type (rarely disporous or tetrasporous). Synergids sometimes with a filiform apparatus. Antipodal cells usually ephemeral (sometimes persistent, occasionally proliferating and haustorial). Endosperm development usually cellular (rarely nuclear). Endosperm haustoria chalazal and/or micropylar, or absent. Embryogenesis usually asterad (sometimes chenopodiad or solanad). Agamospermy sometimes abundant.

Fruit A loculicidal (sometimes septicidal, poricidal or irregularly dehiscing) capsule, berry, drupe or achene (cypsela) with seed wall adnate to pericarp and with persistent and accrescent calyx (rarely a schizocarp with nutlike mericarps or a pyxidium).

Seeds Aril absent. Exotestal cells often thickened, usually palisade, sometimes flattened, cuboidal, fibriform or indistinct. Endotesta often developed into endothelium, integumentary tapetum. Perisperm not developed. Endosperm copious to sparse, oily and proteinaceous (starch usually absent; sometimes with hemicellulose or absent). Embryo large or small, usually well developed, straight, without chlorophyll. Embryo suspensor often filamentous. Cotyledons usually two (rarely absent). Germination phanerocotylar.

Cytology n = 6–18 (rarely 2, or up to 120; 9 and 17 most frequent) – Polyploidy, aneuploidy and agamospermy frequently occurring. Protein bodies present in nucleus.

DNA Plastid genome with large number of inversions. Mitochondrial gene rpl2 lost.

Phytochemistry Flavonols (kaempferol, quercetin), O-methylflavonols, afzelechin, flavonoid sulfates, biflavonoids, cyanidin, coumarins, p-coumaric acid, dammaranes, Group VI secoiridoids (e.g. secologanin), Group VII secoiridoids (e.g. sweroside), Group X secoiridoids (e.g. loganin), cantleyoside, oleanolic acid derivatives, diterpenoids, lupeol and its derivative lupenylacetate, pentacyclic triterpene alcohols, terpenoid ethereal oils, balsams, triterpene acetates, furanoeremophilane sesquiterpenes, sesquiterpene lactones, ursolic acid, caffeic acid esters (verbascosides), chlorogenic acid, ellagic acid (rare), tannins, proanthocyanidins (prodelphinidins, rare), pyrrolizidine alkaloids as macrocyclic aliphatic monocarboxylic diesters, iridoid alkaloids, caurane alkaloids and other alkaloids (rarely benzylisoquinoline alkaloids, homoerythrine alkaloids: homoerythrine, homoerythroidine, homoazaerythrine, holidine, etc.), cyanogenic glycosides (linamarin, lotaustralin, proacacipetalin, prunasin, sambunigrin, triglochinin, zierin), phenylalanine- and tyrosine-derived cyanogenic compounds (rare), triterpene saponins, fatty acid derived polyacetylenes (substitute for iridoids?), aliphatic tetrahydropyrane derivatives, amides, asarone, germacrane-like compounds, myo-inisitol, chiro-inositols (pinitol, quebrachitol), lignans (pinoresinol), arbutin, sinapic acid, chelidonic acid, polyacetate-derived arthroquinones, stigmasterol, polysterols, steroids, and stearic acid. Carbohydrates usually stored as oligo- or polyfructosans (i.a. inulin in subterranean parts of perennials) with isokestose linkages (starch usually absent).

Systematics Campanulales are sister to the clade [Escalloniaceae+[Bruniales+Dipsapiidae]] (Tank & Donoghue 2010).

One possible topology of Campanulales is as follows: [[Rousseaceae+Campanulaceae]+ [Pentaphragmataceae+[[Stylidiaceae+[Phellinaceae+[Argophyllaceae+Alseuosmiaceae]]]+ [Menyanthaceae+[Goodeniaceae+[Calyceraceae+Asteraceae]]]]]].

Rousseaceae and Campanulaceae share the characters large flowers and free stamens (Stevens 2001 and onwards). Campanulales except Rousseaceae and Campanulaceae have the potential synapomorphy corolla lobes provided with marginal wings.

The clade [Phellinaceae+[Alseuosmiaceae+Argophyllaceae]] is characterized by the synapomorphies: woodiness; lamina serrate and gland-toothed; and x = 8. Moreover, Phelline and Argophyllaceae share, according to Stevens (2001 onwards): subepidermal phellogen; spiny pollen grains, with rugulate exine; short style; and apotropous ovules.

The clade [Menyanthaceae+[Goodeniaceae+[Calyceraceae+Asteraceae]]] have the following potential synapomorphies: vessel elements with simple perforation plates; inflorescence with one terminal flower, single flowers and cymes below; petals connate, with early tube formation and with strong fused marginal (commissural) veins joining median vein near apex; filaments epipetalous; tapetum with bi- or multinucleate cells; pollen grains psilate; pistil composed of two carpels; integument more than ten cell layers thick; antiraphal vascular bundle proceeding to micropyle; absence of endosperm haustoria; embryo long; x = 9; and presence of inulin and caffeic acid.

The clade [Goodeniaceae+[Calyceraceae+Asteraceae]] share the unique features: pollen grains with bifurcating columellae; stigma papillate, Dry type; secondary pollen presentation (protandry, anthers connivent at dehiscence, style elongating following pollen deposition); calyx persistent in fruit; and x = 8. Finally, Calyceraceae and Asteraceae possess the synapomorphies (Stevens 2001 onwards): involucrate and capitate inflorescence (forming pseudanthium); small sessile flowers; tubular corolla, with connate commissural veins (median veins sometimes absent); presence of filament collar; presence of pollenkitt; pollen grains with intercolpal depressions; pistil composed of a single carpel; unilocular ovary; fruit a cypsela, with persistent and modified calyx, involved in fruit dispersal.

Phylogeny of Campanulales based on DNA sequence data (Tank & Donoghue 2010). Some analyses (e.g. Olmstead & al. 2000; Bremer & al. 2002) place Campanulaceae as sister to Stylidiaceae (except Donatia, which appears in a different clade), and Pentaphragmataceae also appear in this lineage. Several other analyses (e.g. Lundberg & Bremer 2003) identify the clade [Rousseaceae+Pentaphragmataceae+Campanulaceae] as sister-group to the remaining Campanulales with moderate support.

ALSEUOSMIACEAE Airy Shaw

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Airy Shaw in Kew Bull. 18: 249. 8 Dec 1965

Alseuosmiales Doweld, Tent. Syst. Plant. Vasc.: liv. 23 Dec 2001; Platyspermatiaceae Doweld, Tent. Syst. Plant. Vasc.: liv. 23 Dec 2001 [’Platyspermataceae’]; Alseuosmiineae Reveal in Kew Bull. 66: 47. Mar 2011

Genera/species 5/10

Distribution New Guinea, southeastern Australia, Queensland, New Caledonia, New Zealand.

Fossils Unknown.

Habit Usually bisexual (in Wittsteinia sometimes functionally unisexual), evergreen shrubs (sometimes epiphytic).

Vegetative anatomy Phellogen ab initio superificial. Endodermis present, uniseriate. Young stem with separate vascular bundles. Vessel elements usually with scalariform (sometimes also simple) perforation plates; lateral pits usually opposite or alternate (sometimes scalariform, with numerous cross-bars), bordered pits. Imperforate tracheary xylem elements tracheids with simple or bordered pits, septate or non-septate (as mature living and starchy). Wood rays usually absent (in Crispiloba uniseriate to multiseriate, heterocellular; in Wittsteinia sometimes multiseriate). Axial parenchyma usually absent (sometimes apotracheal or paratracheal scanty). Starch-storing fibres usually present. Pericyclic fibres weakly developed. Sieve tube plastids S type? Nodes 3:3, trilacunar with three leaf traces. Calciumoxalate crystals not found.

Trichomes Hairs unicellular, bicellular or multicellular, uniseriate (in Platyspermation with reddish persistent base), usually confined to leaf axils (in Platyspermation also on other parts of plant).

Leaves Alternate (spiral), simple, entire, sometimes coriaceous, with conduplicate ptyxis. Stipules and leaf sheath absent. Tufts of uniseriate multicellular hairs with dark red pigment present in leaf axils. Petiole vascular bundle transection arcuate; petiole usually with uniseriate endodermis. Venation pinnate. Stomata anomocytic. Cuticular wax crystalloids? Sclereids usually present. Leaf margin serrate or entire.

Inflorescence Usually axillary (sometimes terminal), fasciculate, raceme- or umbel-like, cymose, or flowers solitary axillary.

Flowers Actinomorphic. Hypanthium sometimes present. Usually epigyny (sometimes half epigyny). Sepals (four or) five (to seven), with valvate or open aestivation, free. Petals (four or) five (to seven), with valvate aestivation, connate into urceolate, campanulate or infundibuliform corolla; corolla lobe margins erose, with fimbriate or notched wings (absent in Platyspermation). Nectariferous disc intrastaminal or absent. Tanniniferous cells present in flowers.

Androecium Stamens (four or) five (to seven), haplostemonous, antesepalous, alternipetalous. Filaments free from each other, usually adnate to corolla tube (epipetalous, sometimes only at base). Anthers basifixed to almost dorsifixed, non-versatile, tetrasporangiate, usually introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate cells. Staminodia present in functional female flowers in Wittsteinia. Secondary pollen display absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolpate or tricolporate, usually shed as monads (sometimes tetrads), tricellular at dispersal. Exine tectate, with columellate intratectum, microreticulate to punctate?, tuberculate.

Gynoecium Pistil composed of two (or three) carpels. Ovary usually inferior (sometimes semi-inferior), usually bilocular (sometimes trilocular). Style single, simple, narrow or stout. Stigma capitate, clavate or discoid, often bilobate (sometimes trilobate), little expanded, type? Pistillodium present in functional male flowers in Wittsteinia.

Ovules Placentation axile. Ovules two to numerous (sometimes one) per carpel, anatropous, unitegmic, tenuinucellar. Integument ? cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development cellular. Endosperm haustoria? Embryogenesis?

Fruit Usually a berry (in Platyspermation a capsule), usually with persistent calyx.

Seeds Seeds small (in Platyspermation flattened). Aril absent. Exotestal cells little thickened, lignified (Alseuosmia). Mesotesta persistent. Endotesta? Perisperm not developed. Endosperm copious. Embryo small, straight, chlorophyll? Cotyledons two. Germination?

Cytology n = 9 (Alseuosmia)

DNA Mitochondrial gene rpl2 absent?

Phytochemistry Insufficiently known. Flavonols (quercetin, kaempferol), p-coumaric acid, lupeol and its derivative lupenylacetate, triterpene acetates, condensed tannins, ellagitannins, proanthocyanidins, caffeic acid, triterpene saponins, stigmasterol, and stearic acid present (Alseuosmia). Inulin? Iridoids and alkaloids not found.

Use Ornamental plants.

Systematics Platyspermation (1; P. crassifolium; New Caledonia); Crispiloba (1; C. disperma; northeastern Queensland), Alseuosmia (4; New Zealand), Wittsteinia (3; W. papuana: Papua New Guinea; W. vacciniacea: eastern Victoria; incl. Periomphale?), Periomphale (1; P. balansae; New Caledonia; in Wittsteinia?).

Alseuosmiaceae are sister-group to [Argophyllaceae].

Platyspermation is sister to the remaining Alseuosmiaceae. The corolla tube is very short in Platyspermation, and the lobes are reflexed and provided with papillae along the margins. Tufts of uniseriate hairs are particularly abundant in the leaf axils, as in other Alseuosmiaceae, but in Platyspermation uniseriate hairs cover practically the entire plant. The reddish hair bases somewhat resemble glands.

Cladogram of Alseuosmiaceae based on morphology and DNA sequence data (Kårehed & al. 1999).

ARGOPHYLLACEAE (Engl.) Takht.

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Takhtajan, Sist. Magnoliof. [Systema Magnoliophytorum]: 208. 24 Jun 1987

Corokiaceae Kapil ex Takht., Divers. Classif. Fl. Pl.: 374. 24 Apr 1997

Genera/species 2/17–21

Distribution Eastern Australia, New Caledonia, New Zealand, Lord Howe, Rapa Island.

Fossils Unknown.

Habit Bisexual, evergreen trees or shrubs. Often densely tomentose.

Vegetative anatomy Phellogen ab initio subepidermal? (Corokia). Vessel elements with scalariform perforation plates (often with numerous cross-bars); lateral pits usually alternate (sometimes opposite), bordered pits. Imperforate tracheary xylem elements tracheids (absent in Argophyllum and some species of Corokia) with simple and/or bordered pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma usually absent (rarely apotracheal diffuse, or paratracheal scanty). Sieve tube plastids S type? Nodes usually 3:3, trilacunar with three leaf traces (in Corokia sometimes 1:1, unilacunar with one trace; in Argophyllum sometimes 5:5, pentalacunar with five traces). Crystals absent.

Trichomes Hairs T-shaped, usually with multicellular uniseriate stalk (provided with slits) and very long terminal bifid cell with narrowing apices.

Leaves Alternate (spiral), simple, entire, with supervolute-curved ptyxis (Corokia macrocarpa). Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate. Venation pinnate. Stomata anomocytic, with guard cells raised above epidermis. Cuticular wax crystalloids? Leaf margin coarsely glandular serrate to entire (in Corokia entire).

Inflorescence Terminal or axillary, panicle or corymb (Argophyllum) or paniculate, raceme-shaped or few-flowered fasciculate (Corokia), or flowers solitary axillary (Corokia).

Flowers Actinomorphic. Half epigyny (Argophyllum) or epigyny (Corokia). Sepals (four or) five (to eight), with valvate aestivation, persistent, connate at base. Petals (four or) five (to eight), with valvate aestivation, sometimes with wings at margins, free or connate at base; each corolla lobe usually with adaxial fringed ligule immediately above corolla tube mouth (absent in Corokia macrocarpa). Nectariferous disc present in Corokia. Tanniniferous cells present in flowers.

Androecium Stamens (four or) five (to eight), haplostemonous, antesepalous, alternipetalous. Filaments free from each other and from tepals. Anthers dorsifixed, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Micosporogenesis simultaneous? Pollen grains tricolporate, shed as monads, bicellular (Argophyllum) or tricellular (Corokia) at dispersal. Endoapertures complex and H-shaped. Exine tectate, with columellate? infratectum, finely perforate, coarsely scabrate (Argophyllum) or spinulate (Corokia).

Gynoecium Pistil composed of (one or) two or three (to five) connate carpels. Ovary semi-inferior or inferior, bilocular or trilocular (to quinquelocular) (Argophyllum), or unilocular or bilocular (to quinquelocular) (Corokia). Style single, simple, short. Stigma capitate, punctate, or bilobate or trilobate (to quinquelobate), papillate?, Wet type. Pistillodium absent.

Ovules Placentation axile (Argophyllum) or apical-axile (Corokia). Ovule one (Corokia) or several (Argophyllum) per carpel, anatropous, pendulous (Corokia), apotropous, unitegmic, tenuinucellar. Integument approx. six cell layers thick. Megasporangium with massive base. Megagametophyte monosporous, Polygonum type. Endosperm development cellular (Corokia). Endosperm haustoria chalazal and micropylar (Corokia). Embryogenesis chenopodiad (Corokia).

Fruit A loculicidal capsule (Argophyllum) or a unilocular or bilocular drupe with persistent calyx and style (Corokia).

Seeds Aril? Exotestal cells with inner walls heavily thickened and lignified (Argophyllum), or all walls somewhat thickened (Corokia). Endotesta? Perisperm not developed. Endosperm fleshy, with hemicellulose. Embryo small (Argophyllum) or medium-sized (Corokia), straight, chlorophyll? Cotyledons two. Germination?

Cytologi n = 9 (Corokia)

DNA Mitochondrial gene rpl2 absent?

Phytochemistry Flavonols (kaempferol, quercetin), cyanidin, Group VI secoiridoids (secologanin), triterpenes, ellagic and gallic acid, tannins, and caffeic acid, present. Inulin? Nickel accumulated in two species of Argophyllum.

Use Ornamental plants.

Systematics Argophyllum (11–15; eastern Queensland, eastern New South Wales, New Caledonia), Corokia (6; eastern Queensland, eastern New South Wales, New Zealand, Lord Howe, Rapa Island).

Argophyllaceae are sister to Alseuosmiaceae.

ASTERACEAE Bercht. et J. Presl

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Berchtold et Presl, Přir. Rostlin: 254. Jan-Apr 1820 [‘Astereae’], nom. cons.

Cichoriaceae Juss., Gen. Plant.: 168. 4 Aug 1789 [‘Cichoraceae’], nom. cons.; Compositae Giseke, Prael. Ord. Nat. Plant.: 538. Apr 1792, nom. cons. et nom. alt.; Cynarineae Raf., Anal. Nat.: 191. Apr-Jul 1815 [‘Cynarea’]; Cnicaceae Vest, Anleit. Stud. Bot.: 273, 297. 1818 [’Cnicoideae’]; Tanacetaceae Vest, Anleit. Stud. Bot.: 273, 298. 1818 [’Tanacetoideae’]; Xanthiaceae Vest, Anleit. Stud. Bot.: 273, 298. 1818 [‘Xanthoideae’]; Adenostylidaceae Bercht. et J. Presl, Přir. Rostlin: 254. Jan-Apr 1820 [‘Adenostyleae’], nom. illeg.; Ambrosiaceae Bercht. et J. Presl, Přir. Rostlin: 254. Jan-Apr 1820, nom. cons.; Anthemidaceae Bercht. et J. Presl, Přir. Rostlin: 254. Jan-Apr 1820 [‘Anthemideae’]; Arctotidaceae Bercht. et J. Presl, Přir. Rostlin: 255. Jan-Apr 1820 [‘Arctotideae’]; Artemisiaceae Martinov, Tekhno-Bot. Slovar: 48. 3 Aug 1820 [’Artemisiae’]; Athanasiaceae Martinov, Tekhno-Bot. Slovar: 56. 3 Aug 1820 [’Athanasiae’]; Calendulaceae Bercht. et J. Presl, Přir. Rostlin: 254. Jan-Apr 1820; Carduaceae Bercht. et J. Presl, Přir. Rostlin: 255. Jan-Apr 1820; Centaureaceae Bercht. et J. Presl, Přir. Rostlin: 255. Jan-Apr 1820 [‘Centaureae’]; Echinopaceae Bercht. et J. Presl, Přir. Rostlin: 255. Jan-Apr 1820 [‘Echinopseae’]; Eupatoriaceae Bercht. et J. Presl, Přir. Rostlin: 253. Jan-Apr 1820 [‘Eupatoriae’]; Helianthaceae Bercht. et J. Presl, Přir. Rostlin: 254. Jan-Apr 1820; Inulaceae Bercht. et J. Presl, Přir. Rostlin: 254. Jan-Apr 1820; Lampsanaceae Martinov, Tekhno-Bot. Slovar: 356. 3 Aug 1820 [‘Lampsana’], nom. illeg.; Picridaceae Martinov, Tekhno-Bot. Slovar: 482. 3 Aug 1820 [‘Picrides’]; Santolinaceae Martinov, Tekhno-Bot. Slovar: 560. 3 Aug 1820 [‘Santolinae’]; Senecionaceae Bercht. et J. Presl, Přir. Rostlin: 254. Jan-Apr 1820 [‘Senecioneae’]; Serratulaceae Martinov, Tekhno-Bot. Slovar: 577. 3 Aug 1820 [’Serratulae’]; Tussilag[in]aceae Bercht. et J. Presl, Přir. Rostlin: 254. Jan-Apr 1820 [‘Tussilagineae’]; Heleniaceae Raf. in Cincinnati Lit. Gaz. 2: 28. 24 Jul 1824 [‘Helenidia’]; Acarnaceae Link, Handbuch 1: 684. 20 Apr 1829, nom. illeg.; Ambrosiales Link, Handbuch 1: 816. 4-11 Jul 1829 [‘Ambrosiaceae’]; Anthemidales Link, Handbuch 1: 752. 4-11 Jul 1829 [’Anthemideae’]; Asterales Link, Handbuch 1: 731. 4-11 Jul 1829 [’Asteroideae’]; Calendulales Link, Handbuch 1: 776. 4-11 Jul 1829 [’Calendulaceae’]; Cichoriales Link, Handbuch 1: 779. 4-11 Jul 1829 [’Cichoriaceae’]; Coreopsidaceae Link, Landbuch 1: 786, 20 apr 1829 [‘Coreopsideae’], nom. illeg.; Echinopales Link, Handbuch 1: 814. 4-11 Jul 1829 [‘Echinopeae’]; Helichrysaceae Link, Handbuch 1: 712. 20 Apr 1829 [‘Elichryseae’], nom. illeg.; Partheniaceae Link, Handbuch 1: 816. 20 Apr 1829, nom. illeg.; Perdiciaceae Link, Handbuch 1: 728. 20 Apr 1829 [‘Perdicieae’], nom. illeg.; Eupatoriineae Link, Handbuch 1: 729. 4-11 Jul 1829; Gnaphaliaceae Link ex F. Rudolphi, Syst. Orb. Veg.: 46. 5-12 Jul 1830 [‘Gnaphalieae’]; Cynaraceae Spenn., Handb. Angew. Bot. 1: 296. 1-19 Jul 1834 [‘Cynareae’]; Mutisiaceae Burnett, Outl. Bot.: 934, 935, 1094, 1111. Feb 1835; Asterineae Burnett, Outlines Bot.: 901, 1111. Feb 1835 [‘Asterosae’]; Nassauviaceae Burmeist., Handb. Naturgesch. 1: 290. 12-17 Dec 1836; Vernoniaceae Burmeist., Handb. Naturgesch. 1: 296. 12-17 Dec 1836; Aposeridaceae Raf., New Fl. N. Amer. 4: 106. med 1838 [’Aposerides’]; Asteropsida Brongn., Enum. Plant. Mus. Paris: xvii, 32. 12 Aug 1843 [’Asteroideae’]; Xeranthemaceae Döll, Rhein. Fl.: 498. 24-27 Mai 1843 [‘Xeranthemeae’]; Matricariaceae Voigt, Hort. Suburb. Calcutt.: 400. Aug-Dec 1845; Cichoriineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 856, 944. 1846 [‘Cichoraceae’]; Mutisiineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 855, 941. 1846; Nassauviineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 856, 943. 1846; Senecionineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 855, 877. 1846 [‘Senecionideae’]; Vernoniineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 855, 856. 1846 [‘Vernonieae’]; Lactucaceae Drude in J. A. Schenk, Handb. Bot. 3(2): 369. 1886; Carduales Small, Fl. S.E. U.S.: 1148. 22 Jul 1903; Grindeliaceae Reichb. ex A. Heller in Muhlenbergia 2: 330. 30 Dec 1907; Madiaceae (Greene) A. Heller in Muhlenbergia 2: 332. 30 Dec 1907

Genera/species 1.566–1.576/22.480–23.100

Distribution Cosmopolitan except Antarctica.

Fossils Pollen grains are known from the Oligocene onwards of North and South America, and Europe, although pollen probably originating from Asteraceae have been found in Paleocene layers of western South America. The earliest unambiguous pollen fossils of Asteraceae are Oligocene in origin. Macrofossils of Asteraceae have also been reported from the mid-Eocene onwards, i.a. the Patagonian fossil Raiguenrayun (Barreda & al. 2010, 2012; Panero & al. 2014).

Habit Usually bisexual (sometimes monoecious, gynomonoecious, polygamomonoecious, dioecious, androdioecious, or gynodioecious), usually perennial, biennial or annual herbs (sometimes evergreen, rarely deciduous, shrubs, trees or lianas). Some species are succulent. Numerous representatives are spiny or prickly. Many Asteraceae have stem or root nodules or lignotuber. A large number of species are xeromorphous. Some species have C4 and/or CAM physiology. Often with a strong scent or odour.

Vegetative anatomy Phellogen ab initio usually superficial (sometimes deeply seated). Cortical and/or medullary vascular bundles sometimes present. Medulla in some species of Senecioneae stratified by diaphragms. Vascular bundles usually separated (in woody species cylinder). Cambium often storied. Wood elements sometimes storied. Secondary lateral growth usually normal (also in numerous herbaceous representatives; sometimes anomalous from concentric cambia or a cylindrical cambium). Vessel elements usually with simple (sometimes scalariform or reticulate) perforation plates; lateral pits usually alternate (sometimes opposite), usually bordered (sometimes simple) pits. Vestured pits present. Imperforate tracheary xylem elements libriform fibres with small simple pits, septate or non-septate (often also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse, or paratracheal scanty, aliform, lozenge-aliform, winged-aliform, confluent, vasicentric, or banded. Tyloses sometimes frequent. Intraxylary phloem rarely? present. Sieve tube plastids S type. Endodermis sometimes with thick-walled cells. Nodes >3:>3, trilacunar, pentalacunar or multilacunar with three or more? leaf traces (rarely unilacunar with one? trace). Phloem (especially in Lactuceae) with articulated laticifers containing latex rich in triterpenes, and/or tissues with schizogenous secretory resinous ducts often lined with epithelial cells (scattered latex cells sometimes present as well as resinous ducts). Parenchyma in some species with prismatic calciumoxalate crystals; acicular crystals, styloids, crystal sand and other types of crystals sometimes present.

Trichomes Hairs unicellular or multicellular, uniseriate or biseriate, simple or branched, flagellar hairs, T-shaped, stellate, candelabra-shaped, dendritic, arachnoid, peltate, lepidote, or vesicular; glandular hairs with unicellular or multicellular head; sometimes laticiferous hairs.

Leaves Usually alternate (spiral; sometimes opposite, rarely verticillate), usually simple (sometimes pinnately or palmately? compound), entire or pinnately lobed (sometimes repeatedly pinnately lobed), often with conduplicate or revolute ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection usually arcuate. Venation pinnate, palmatopinnate, parallelopinnate, parallelodromous, or palmate. Stomata usually anomocytic (sometimes anisocytic or helicocytic). Cuticular wax crystalloids partly as large glabrous partially inrolled scales, partly as reticulate to annular threads or small scales, or as rosettes of platelets (Fabales type). Secretory cavities and ducts (with resin and latex) present or absent. Leaf margin serrate, often serrate-dentate, crenate, lobate or entire, sometimes with hydathodes. Extrafloral nectaries rarely present.

Inflorescence Terminal or axillary, primarily racemose capitulum (pseudanthium) with abaxial involucre consisting of one or several rows of bracts. Capitulum with one or more sessile flowers on common flat to conical or cylindrical receptacle (sometimes, i.a. in Heliantheae, with one bract [floral bract] present adjacent to each flower or with prickles or bristles, e.g. in Carduoideae). Capitula usually organized in compound, often corymbose inflorescence or sometimes in cymose secondary compound capitula (sometimes with secondary involucre; capitulum sometimes consisting of one flower, many capitula aggregated into “super-capitulum”). Capitula homogamous, with all flowers usually bisexual, or heterogamous, with external flowers, ray florets, female, sterile or bisexual, and central flowers, disc florets, bisexual or functionally male. Capitulum without terminal flower, with usually centripetal (peripheral flowers in, e.g., Gorteria centrifugal) development of flowers. Floral prophylls (bracteoles) absent. Extrafloral nectaries sometimes present on bracts.

Flowers Actinomorphic or zygomorphic, usually small. Epigyny. Sepals (and petal bases) modified into persistent and usually accrescent pappus (hairs, scales or bristles), or absent. Petals (three to) five (or six), with valvate aestivation, tubular, spatulate (apically tridentate or quinquedentate) or bilabiate, with upper lip unilobate and lower lip quadrilobate, or upper lip bilobate and lower lip trilobate, connate (rarely absent), usually without midvein (present in, e.g., Barnadesioideae); corolla tube development early or late. Nectariferous disc intrastaminal, annular, scale-like or tubular, surrounding stylar base.

Androecium Stamens (three to) five (or six), haplostemonous, antesepalous, alternipetalous. Filaments usually free from each other (rarely connate into tube), adnate to corolla tube (epipetalous), in upper part usually with ring of thick-walled cells. Anthers usually caudate, with conspicuous apical and basal (calcarate) appendages, usually connate into tube around style, basifixed or dorsifixed, non-versatile, usually tetrasporangiate (rarely disporangiate), introrse, longicidal (dehiscing by longitudinal slits). Tapetum ab initio cellular, with multinucleate cells, later usually amoeboid-periplasmodial by dissolution of cell walls (rarely secretory). Female flowers usually with staminodia. Secondary pollen display present (many species with “pollen pump” or “pollen brush”).

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate (sometimes triporate, rarely pantoporate; in Cichorieae usually strongly lophate, fenestrate; pollen grains often caveate in lineages basal to Corymbium and Asteroideae, but also in Senecioneae, Arctotideae and some Cichorieae), shed as monads, tricellular at dispersal. Exine tectate or semitectate, with columellate infratectum (with furcate columellae), spinulate, echinate, microechinate, scabrate, granulate, psilate, smooth or lophate, often caveate (fenestrate).

Gynoecium Pistil composed of two (or three) connate carpels. Ovary inferior, unilocular. Style single, usually bilobate (rarely trilobate), usually with hairs (in, e.g., Barnadesioideae glabrous) on which pollen grains are deposited (secondary pollen display), occasionally hollow. Stigmatic surfaces present on adaxial side of stylar branches (inner surface of stylar branches stigmatic), papillate, Dry type; apex of stylar branches rounded, obtuse or with different appendages; base of stylar branches often inflated. Male flowers usually with pistillodium.

Ovules Placentation basal. Ovule one per ovary, anatropous, ascending, epitropous, unitegmic, tenuinucellar. Integument six to twenty cell layers thick, rarely vascularized. Archespore usually unicellular (sometimes multicellular, rarely bicellular). Megagametophyte usually monosporous, Polygonum type (sometimes disporous, Allium type, rarely tetrasporous, Chrysanthemum type?, or unspecified). Synergids often elongated (in, e.g., Arctotideae and Calenduleae synergid haustorium present, i.e. synergids sometimes haustorial). Antipodal cells often persistent, proliferating (up to c. 60 cells) and/or multinucleate, and haustorial. Endosperm development usually cellular (sometimes nuclear or variations). Endosperm haustoria usually absent. Embryogenesis asterad. Polyembryony and other types of agamospermy frequent in many lineages.

Fruit Usually an achene, cypsela, with seed wall adnate to pericarp, and often with persistent and accrescent calyx and corolla base modified into pappus, often with hairs, bristles, prickles, hooks or other types of outgrowths (rarely a one-seeded drupe with fleshy pericarp). Phytomelan layer present on surface of cypsela in some groups of Asteroideae (Heliantheae).

Seeds Aril absent. Elaiosome sometimes present. Testa usually vascularized. Exotestal cells thickened, palisade, flattened or indistinct. Endotesta developed into endothelium, integumentary tapetum. Perisperm not developed. Endosperm thin, oily and proteinaceous, or absent. Embryo large, straight, oily, without chlorophyll. Cotyledons usually two (possibly absent in, e.g., Syneilesis). Germination phanerocotylar.

Cytology n = usually 9, 10, 12, 17, or 18 (lowest n = 2, highest n = 120) – Polyploidy, aneuploidy and agamospermy frequently occurring. Protein bodies present in nucleus.

DNA Mitochondrial gene rpl2 absent. “6 bp x 4” inversion present in plastid gene rbcL. Plastid genome in all Asteraceae except Barnadesioideae with inversion of 22 kb. Plastid genome in Lactuca sativa with inversion of 4 kb and, possibly, additional shorter inversion.

Phytochemistry Flavonols (kaempferol, quercetin, quercetagetine based yellow flavonols, etc.), isoflavonoids, chalcones, cyanidin, coumarins, diterpenoids, terpenoid ethereal oils and balsams, sesquiterpene lactones (especially in latex; responsible for bitter taste of numerous Asteraceae), ursolic acid, caffeic acid, verbascosides (rare), pentacyclic triterpene alcohols, ellagic acid (rare), caurane and other alkaloids, triterpene saponins, phenylalanine-derived cyanogenic compounds (rare), lignans, fatty acids (in seeds), fatty acid derived polyacetylenes (heterocyclic, aromatic or with vinylic terminal groups; above all in resinous ducts), arbutin, polyacetate derived arthroquinones, chlorogenic and isochlorogenic acids, and steroids present. Pyrrolizidine alkaloids as macrocyclic aliphatic monocarboxylic diesters (seneciphylline, retrorsine, senecionione, etc.), as well as furanoeremophilane sesquiterpenes abundant in Senecioneae. Carbohydrates stored as oligo- or polyfructosans (e.g. inulin) with isokestose bonds (fructans with unbranched chain); starch absent. Selenium accumulated in some species. Iridoids, tannins, and acetylene compounds not found.

Use Ornamental plants, spices (Artemisia dracunculus), vegetables (Cynara, Helianthus tuberosus, Scorzonera hispanica, Tragopogon porrifolius, Cichorium intybus var. endive, Lactuca sativa, Acmella oleracea, Smallanthus sonchifolius), seed oils (Helianthus annuus, Carthamus tinctorius, Guizotia abyssinica etc.), medicinal plants, insecticides (Tanacetum cinerariifolium, Pyrethrum), cosmetics, perfumes, dyeing substances (Carthamus tinctorius etc.), rubber (Taraxacum bicorne, Parthenium argentatum), timber.

Systematics Asteraceae are sister to Calyceraceae.

A probable topology of Asteraceae is the following: [Barnadesioideae+[Famatinanthoideae+[Mutisioideae+Stifftioideae+[Wunderlichioideae+[Gochnatioideae+[Hecastocleidodoideae+[Carduoideae+[Pertyoideae+[Gymnarrhenoideae+[Cichorioideae+[Corymbioideae+Asteroideae]]]]]]]]]]].

Barnadesioideae (D. Don) K. Bremer et R. K. Jansen in Ann. Missouri Bot. Gard. 79: 415. 6 Mai 1992

9/c 92. Schlechtendalia (1), Doniophyton (2; Andes of Chile and Argentina, Patagonia), Duseniella (1), Fulcaldea (2), Barnadesia (19), Huarpea (1; H. andina; Andes of La Rioja in Argentina ), Dasyphyllum (c 40), Arnaldoa (3), Chuquiraga (23). – Southern South America, Brazil, with their highest diversity in the Andes. Trees, shrubs or herbs. Leaves alternate (spiral), opposite or verticillate. Axillary spines frequent. Capitula discoid, radiate or ligulate. Involucral bracts chartaceous. Corolla tubular, split, ligulate or labiate (usually with upper lip quadrilobate and lower lip simple; rarely with upper lip trilobate and lower lip bilobate). Petal with midvein. Corolla and cypsela bristles with long tricellular hairs (epidermal cell indistinct, basal cell short and thick-walled, third cell elongate and thin-walled). Corolla in Arnaldoa (4+1)-bilabiate. Anther thecae calcarate or ecalcarate and caudate or ecaudate. Pollen grains prolate, spinulate, microechinate, scabrate, granulate, or smooth (rarely lophate and not spinulate), with intercolpar depressions (caveate, with cavity between columellae and foot layer). Style glabrous or papillate below branching point, bilobate, with stylar branches short, widened at apex. Stigma lobate. Cypsela densely villose with typical trichomes. Pappus uniserate, plumose, barbellate, or setaceous (rarely glabrous). x = 9. Flavonoids sparsely present. Flavones absent.

[Famatinanthoideae+[Mutisioideae+Stifftioideae+[Wunderlichioideae+[Gochnatioideae+[Hecastocleidoideae+[Carduoideae+[Pertyoideae+[Gymnarrhenoideae+[Cichorioideae+[Corymbioideae+Asteroideae]]]]]]]]]]

Corolla usually zygomorphic. Style with rigid sweeping pollen-collecting hairs and often long branches. Cypsela with twin hairs (with unicellular to uniseriate base, apical cell simple or equally bifid). Pappus developing late, usually consisting of capillary bristles (sometimes scales, awns etc.). n = 2 to more than 100. Inversion of 22,8 kb in plastid DNA with internal inversion of 3,3 kb.

Famatinanthoideae S. E. Freire, Ariza et Panero in Mol. Phylogen. Evol. 80: 49. Nov 2014

1/1. Famatinanthus (1; F. decussatus; the Andes in Argentina). – Panero & al. (2014) found Famatinanthus to be sister to all Asteraceae except Barnadesioideae.

[Mutisioideae+Stifftioideae+[Wunderlichioideae+[Gochnatioideae+[Hecastocleidoideae+[Carduoideae+[Pertyoideae+[Gymnarrhenoideae+[Cichorioideae+[Corymbioideae+Asteroideae]]]]]]]]]

Mutisioideae (Cass.) Lindl. in Loud., Encycl. Plant.: 1074. 1829

c 44/610–625. South America, Africa, Asia. Usually perennial herbs (sometimes shrubs). Leaves alternate. Pollen grains prolate to spheroidal, with psilate exine. Style bilobate, branches rounded at apex. n = (6–)9 or more; x = 9.

Onoserideae Solbrig in Taxon 12: 231. Jul 1963

6/53. Aphyllocladus (5; the Andes in southern Bolivia, northern Chile and northwestern Argentina), Gypothamnium (1; G. pinifolium; northern Chile), Lycoseris (11; tropical America), Onoseris (32; Mexico and southwards to the Andes), Plazia (3; the Andes from Peru to Chile and Argentina), Urmenetea (1; U. atacamensis; the Andes in northern Chile and northwestern Argentina). – Tropical America, the Andes. Shrubs or herbs. Capitula radiate, with imbricate involucral bracts. Ray corollas bilabiate, disc corollas quinquelobate tubular. Anther thecae calcarate and caudate. Pappus consisting of bi- to quadriseriate bristles.

[Mutisieae+Nassauvieae]

Mutisieae Cass. in J. Phys. Chim. Hist. Nat. Arts 88: 199. Mar 1819

15/c 260. Adenocaulon (>5; southern Asia, North America, Mexico, Guatemala, Argentina), Amblysperma (2; southwestern Western Australia), Brachyclados (3; temperate South America), Chaetanthera (43; southern Peru, Chile, the Andes in Argentina), Chaptalia (c 60; tropical and subtropical America), Chucoa (2; the Andes in Peru), Eriachaenium (1; E. magellanicum; southernmost South America), Gerbera (c 35; northeastern, tropical and southern Africa, Madagascar, southern Asia to China), Leibnitzia (6; southern and eastern Asia, North America), Lulia (1; L. nervosa; Brazil), Mutisia (62; the Andes in Colombia to southern Chile and Argentina, southeastern Brazil, Uruguay, Paraguay, northeastern Argentina), Pachylaena (2; the Andes in Chile and Argentina), Perdicium (2; Western Cape), Trichocline (22; South America, southwestern Western Australia), Uechtritzia (3; Armenia, Central Asia, Himalayas, China). – Africa, Madagascar, Armenia, southern and Central Asia to China, southwestern Australia, America, with their highest diversity in South America. Usually herbs (sometimes shrubs). Capitula usually radiate (sometimes di sciform or discoid), with imbricate involucral bracts. Ray corollas tridentate or bilabiate (rarely tubular) strap-shaped or absent, disc corollas bilabiate or tubular, shallowly lobate. Anther thecae usually ecalcarate and caudate. Style shallowly bilobate, with long papillate branches (papillae rounded). Cypsela often glandular Pappus consisting of uni- or multiseriate scabrid to plumose usually capillary bristles, or absent.

Nassauvieae Cass. in J. Phys. Chim. Hist. Nat. Arts 88: 198. Mar 1819

c 24/300–310. Acourtia (c 40; southern United States, Mexico, Central America, the West Indies), Ameghinoa (1; A. patagonica; Patagonia in Argentina), Berylsimpsonia (2; the Greater Antilles), Burkartia (1; B. lanigera; Patagonia), Calopappus (1; C. acerosus; the Andes in central Chile; in Nassauvia?), Cephalopappus (1; C. sonchifolius; northeastern Brazil), Criscia (1; C. stricta; Brazil), Dolichlasium (1; D. lagascae; the central Andes in Argentina), Holocheilus (7; southern Brazil, Uruguay, Paraguay, northern and central Argentina), Jungia (26; Mexico, Central America, the Andes), Leucheria (46; the Andes from southern Peru to Chile and Patagonia, Falkland Islands), Leunisia (1; L. laeta; the Andes in central Chile), Lophopappus (6; the Andes from southern Peru to northern Chile and northwestern Argentina), Macrachaenium (1; M. gracile; the Andes, Patagonia, Tierra del Fuego), Marticorenia (1; M. foliosa; central Chile), Moscharia (2; central Chile), Nassauvia (c 50; the southern Andes in Bolivia, Chile and Argentina, Patagonia, Falkland Islands; incl. Calopappus?), Oxyphyllum (1; O. ulicinum; northern Chile), Pamphalea (9; southern South America to Uruguay)?, Perezia (32; the Andes from Colombia to southern Patagonia, southern Brazil, Uruguay, Paraguay, northeastern Argentina), Pleocarphus (1; P. revolutus; northern Chile), Polyachyrus (7; Peru to the Andean slopes in central Chile), Proustia (3; Bolivia, Chile, Argentina), Trixis (c 65; southwestern United States, Mexico, Central America, western South America to Chile). – Tropical America, subtropical and temperate South America, with their largest diversity in the Andes. Shrubs, lianas or herbs. Capitula discoid or radiate, with imbricate involucral bracts. Ray corollas bilabiate, disc corollas usually bilabiate. Anther thecae calcarate and caudate. Stylar branches with apical crown of sweeping hairs. Pappus consisting of uni- to multiseriate paleaceous to plumose capillary bristles.

Stifftioideae (D. Don) Panero in Panero et Funk in Phytologia 89: 356. Dec 2007

9/44. Stifftia (8; northeastern Brazil, French Guiana); Achnopogon (2; Venezuela, Guyana), Duidaea (4; Venezuela, Guyana), Glossarion (2; Venezuela, Guyana), Gongylolepis (15; tropical South America, with their largest diversity in Venezuela and Guayana Highlands), Neblinaea (1; N. premontorium; Venezuela, Guyana), Quelchia (4; Venezuela, Guyana), Salcedoa (1; S. mirabiliarum; Hispaniola); Hyaloseris (7; Bolivia, Argentina). – Hispaniola, South America, with their largest diversity in the Venezuela and Guayana Highlands. Trees or shrubs (sometimes lianas). Leaves alternate or opposite. Capitula discoid or ligulate, with multiseriate imbricate involucral bracts. Corolla actinomorphic and quinquelobate (Stifftia) or zygomorphic and usually (3+2)-bilabiate with coiled lobes (Gongylolepis clade) or actinomorphic to zygomorphic and (4+1)-subbilabiate to quinquedentate ligulate (Hyaloseris clade). Anther thecae calcarate and caudate. Pollen grains usually prolate, echinate or psilate. Style bilobate, with rounded to shortly acute apex. Pappus showy, consisting of several series of numerous scabrid capillary bristles or setae. x = 9. – A probable topology is [Stifftia+[Gongylolepis clade+Hyaloseris clade]]. Stifftioideae appear to be sister-group to the remaining Asteraceae (Panero & al. 2014).

[Wunderlichioideae+[Gochnatioideae+[Hecastocleidoideae+[Carduoideae+[Pertyoideae+[Gymnarrhenoideae+[Cichorioideae+[Corymbioideae+Asteroideae]]]]]]]]

Wunderlichioideae Panero et V. A. Funk in Phytologia 89: 357. Dec 2007

8/42. Venezuelan and Guiana Highlands, eastern South America, southwestern China. Style and stylar branches glabrous. x = 9. Deletion in plastid gene rpoB.

Wunderlichieae Panero et V. A. Funk in Phytologia 91: 570. 1 Dec 2009

4/36. Wunderlichia (5; Brazil); Chimantaea (10; Venezuela, Guyana), Stenopadus (15; Guayana Highlands), Stomatochaeta (6; Venezuela, Guyana, northern Brazil). – Northeastern tropical South America. Trees or shrubs. Capitula discoid, with imbricate involucral bracts. Corolla actinomorphic, deeply quinquelobate. Anther thecae calcarate and caudate. Pappus consisting of multiseriate bristles or setae.

Hyalideae Panero in J. L. Panero et V. A. Funk in Phytologia 89: 358. Dec 2007

4/6. Ianthopappus (1; I. corymbosus; South America), Hyalis (2; southern Bolivia, Paraguay, Argentina); Leucomeris (2; Himalayas, Yunnan, Burma, Thailand, Vietnam), Nouelia (1; N. insignis; southwestern China). – Himalayas to southwestern China and Vietnam, South America. Shrubs or small trees. Capitula radiate or discoid, with imbricate involucral bracts. Ray corollas bilabiate (with coiled lobes) or absent, disc corollas tubular, deeply quinquelobate. Anther thecae calcarate and caudate. Pappus consisting of bi- or triseriate scabrid to smooth capillary bristles.

[Gochnatioideae+[Hecastocleidoideae+[Carduoideae+[Pertyoideae+[Gymnarrhenoideae+[Cichorioideae+[Corymbioideae+Asteroideae]]]]]]]

Gochnatioideae Panero et V. A. Funk in Proc. Biol. Soc. Washington 115: 913. 30 Dec 2002

6–7/85–90. Cyclolepis (1; C. genistoides; temperate South America); Gochnatia (7–8; the central Andes), Pentaphorus (2; the southern Andes), Anastraphia (38; Mexico, Cuba, Hispaniola, the Bahamas), Cnicothamnus (2; Bolivia, northwestern Argentina), Moquiniastrum (21; tropical South America, with their highest diversity in Brazil), Richterago (16; southeastern Brazil, Uruguay, northeastern Argentina). – Mountains in Southeast Asia, tropical and subtropical America. Trees, shrubs or perennial herbs. Capitula radiate or discoid, with tri- to decemseriate imbricate involucral bracts. Ray corollas bilabiate or absent, disc corollas deeply quinquelobate tubular. Anther thecae ecalcarate and caudate. Pollen prolate to spheroidal, anthemoid type. Stylar branches usually short and glabrous, with rounded apices. Cypsela villose. Pappus consisting of uni- to triseriate scabrid or plumose capillary (or sometimes flat) bristles. n = 22, 23, 27; x = 9. – Cyclolepis genistoides is often recovered as sister to the remaining Gochnatioideae or, sometimes, as sister to Wunderlichieae (Funk & al. 2014).

[Hecastocleioideae+[Carduoideae+[Pertyoideae+[Gymnarrhenoideae+[Cichorioideae+[Corymbioideae+Asteroideae]]]]]]

Pollen grains spheroidal. Columellae sausage-like. Deletion in plastid gene rpoB.

Hecastocleidoideae Panero et V. A. Funk in Proc. Biol. Soc. Washington 115: 914. 30 Dec 2002

1/1. Hecastocleis (1; H. shockleyi; southern Nevada and adjacent California). – Shrub. Capitula discoid, single-flowered, with imbricate involucral bracts. Corolla actinomorphic, deeply quinquelobate tubular. Anther thecae calcarate and caudate. Pollen grains spheroidal, tricolpate. Stylar branches short, with rounded apex. Pappus consisting of six scales (sometimes fused). x = 8.

[Carduoideae+[Pertyoideae+[Gymnarrhenoideae+[Cichorioideae+[Corymbioideae+Asteroideae]]]]]

Carpels of disc florets superposed (one above the other). Style with pollen collecting hairs (sweeping hairs). x = 10. Deletion and insertion in plastid gene rpoB.

Carduoideae Carduoideae Cass. ex Sweet, Hort. Brit.: 213. Jul-Aug 1826

85–90/2.800–2.900. Cosmopolitan, although mainly in the Northern Hemisphere (especially Eurasia and North Africa). Leaves alternate (spiral), often spinose-serrate. Flowers usually actinomorphic. Pollen grains spheroidal, sometimes psilate, with internal tectum, sometimes with internal foramina. Columellae medium thickness. Style with hair collar below branches. Cypsela usually with twin hairs. Exotestal cells palisade. x = 9–11. – Probable topology is [Cardueae+Dicomeae+[Oldenburgieae+Tarchonantheae]].

Cardueae Cass. in J. Phys. Chim. Hist. Nat. Arts 88: 155. Feb 1819 [‘Carduineae’]

75–80/2.700–2.800. Cardopatium (1–2; the Mediterranean), Cousiniopsis (1; C. atractyloides; Central Asia), Thevenotia (2; southwestern Asia), Tugarinovia (1; T. mongolica; Mongolia), Atractylodes (11; East Asia), Atractylis (14; southern Europe, the Canary Islands, the Mediterranean, temperate Asia), Carlina (28; Europe, Macaronesia, the Mediterranean, western Asia), Echinops (c 120; Europe, the Mediterranean and eastwards to Central Asia, tropical African mountains), Dipterocome (1; D. pusilla; eastern Mediterranean and eastwards to Afghanistan), Acantholepis (1; A. orientalis; western, central and southern Asia), Amphoricarpos (3–4; southeastern Europe to Caucasus), Chardinia (1; C. orientalis; western Asia), Xeranthemum (5–6; the Mediterranean, southwestern Asia), Siebera (2; southwestern Asia to Afghanistan), Berardia (1; B. subacaulis; western Alps), Staehelina (5; the Mediterranean), Hirtellina (3; wart-temperate regions in southeastern Europe and Asia), Shangwua (3; Tibet, western China), Onopordum (c 40; Europe, the Mediterranean, western Asia), Xanthopappus (1; X. subacaulis; Mongolia, northwestern China), Ancathia (1; A. igniaria; Central Asia, Mongolia, China), Synurus (1–2; S. deltoides; East Asia), Alfredia (5; Central to East Asia), Lamyropappus (1; L. shacaptaricus; Central Asia), Syreitschikovia (2; Central Asia), Olgaea (16; Central Asia to northern China), Cynara (8; Canary Islands, the Mediterranean), Ptilostemon (14; the Mediterranean), Galactites (3; the Canary Islands, the Mediterranean), Lamyropsis (8; the Mediterranean, southwestern Asia), Notobasis (1; N. syriaca; the Mediterranean and eastwards to Central Asia), Picnomon (1; P. acarna; the Mediterranean, southwestern Asia), Silybum (2–3; the Mediterranean), ‘Carduus’ (c 90; Europe, Macaronesia, the Mediterranean, East African mountains, temperate Asia; paraphyletic), Cirsium (c 200; temperate regions on the Northern Hemisphere, the Mediterranean), Tyrimnus (1; T. leucographus; the Mediterranean), Arcyna (1; A. tournefortii; the Iberian Peninsula), Takeikadzuchia (1; T. lomonosowii; Mongolia, northern China), Arctium (11; temperate regions in the Old World), Cousinia (600–700; eastern Mediterranean and eastwards to Central Asia and western Himalayas), Schmalhausenia (1; S. nidulans; Tienshan), Hypacanthium (3; Central Asia), Saussurea (c 400; Europe, temperate Asia, with their largest diversity in Himalayas, Tibet and western China), Himalaiella (11; Himalayas), Jurinea (c 200; Central and South Europe, the Mediterranean, northwestern Africa, southwestern and Central Asia, with their largest diversity in Central Asia), Goniocaulon (1; G. glabrum; tropical Africa, India), Schischkinia (1; S. albispina; southwestern to Central Asia), Volutaria (16; Macaronesia, the Mediterranean, southwestern Asia, tropical East Africa), Cheirolophus (c 25; southwestern Europe, Madeira, the Canary Islands, North Africa), Myopordon (6; southwestern to Central Asia), Rhaponticum (26; Tenerife, the Mediterranean, temperate and subtropical Asia, southeastern Australia), Callicephalus (1; C. nitens; southwestern to Central Asia), Mantisalca (4; the Mediterranean, North Africa), Centaurothamnus (1; C. maximus; the Arabian Peninsula), Ochrocephala (1), Serratula (2–4), Klasea (c 65), Stizolophus (2), Centaurodendron (3; Juan Fernandez Islands), Plectocephalus (7; Ethiopia, North America, Mexico, southern South America), Zoegea (3; Egypt, southwestern to Central Asia), Psephellus (85–90; eastern Mediterranean, southwestern to Central Asia, southern Siberia, one species, P. sibiricus, in China), Rhaponticoides (17; Europe, the Mediterranean, temperate Asia), ’Centaurea’ (c 650; Europe, the Mediterranean, tropical African mountains, southwestern and northern Asia; paraphyletic), Carthamus (25–30; the Mediterranean and eastwards to Central Asia; incl. Carduncellus, Femeniasia and Phonus?), Carduncellus (27; the Mediterranean; in Carthamus?), Femeniasia (1; Minorca; in Carthamus?), Phonus (2; the Iberian Peninsula; in Carthamus?). – Unplaced Cardueae Dolomiaea (14; Himalayas, Tibet), Diplazoptilon (2; southwestern China), Polytaxis (2; Central Asia), Crupina (3; the Mediterranean, southwestern Asia and eastwards to China), Amberboa (6; the Mediterranean and eastwards to Central Asia), Tricholepis (18; Central Asia to Burma), Plagiobasis (1; P. centaureoides; Central Asia), Russowia (1; R. sogdiana; Turkestan), Karvandarina (1; K. aphylla; Iran, Pakistan), Oligochaeta (4; southwestern to Central Asia, India), Cavea (1; C. tanguensis; eastern Himalayas), Pilostemon (2; Central Asia to China). – Temperate, arid subtropical and alpine regions in the Old World, with their highest diversity in the Mediterranean and eastwards to Central Asia. Usually herbs (sometimes shrubs, rarely trees). Roots with laticifers. Capitula discoid, with multiseriate involucral bracts (often spiny). Corolla usually actinomorphic, deeply quinquelobate, usually tubular, straight or s-shaped. Anther thecae calcarate and caudate. Pollen grains sometimes with tectum echinate. Style with short hairs above branching point, below with papillose-pilose thickened articulation (“pollen brush”). Cypsela walls with phytomelan. Pappus consisting of scales or bristles. x = 10.

Dicomeae Panero et V. A. Funk in Proc. Biol. Soc. Wash. 115: 916. 20 Dec 2002

8/c 100. ‘Pasaccardoa’ (4; tropical and southern Africa; non-monophyletic), Dicomopsis (1; D. welwitschii; Angola), Dicoma (c 35; tropical and southern Africa, Madagascar, Socotra, one species on the Arabian Peninsula, one species in Pakistan and India), Cloiselia (2; Madagascar), Macledium (c 20; tropical and southern Africa); Erythrocephalum (13; tropical East Africa), Pleiotaxis (26; tropical and southern Africa), Gladiopappus (1; G. vernonioides; Madagascar, extinct?). – Tropical and southern Africa, Madagascar, Socotra, the Arabian Peninsula, Pakistan, India. Small trees, shrubs or perennial herbs. Capitula discoid or radiate, with imbricate involucral bracts, often with hyaline margins. Ray corollas bilabiate, disc corollas deeply quinquelobate, tubular. Anther thecae calcarate and caudate. Stylar branches with apical acute sweeping hairs. Cypsela with twin hairs. Pappus consisting of scabrid to plumose multiseriate bristles or scales. x = 11. – Erythrocephalum, Pleiotaxis and Gladiopappus differ from the rest and are provisionally included in Dicomeae.

[Oldenburgieae+Tarchonantheae]

Oldenburgieae S. Ortiz, Compositae Newslett. 47: 2. 15 Apr 2009

1/4. Oldenburgia (4; southern parts of Western and Eastern Cape). – Small trees or shrubs. Capitula radiate, with imbricate involucral bracts, with or without hyaline margins. Ray corollas bilabiate, disc corollas usually actinomorphic, deeply quinquelobulate, tubular. Anther thecae calcarate and caudate. Pollen grains with tectum echinate. Stylar branches usually without sweeping hairs. Pappus consisting of scabrid to plumose multiseriate bristles. x = 9.

Tarchonantheae Vis., Fl. Dalmat. 2: 24, 60. 10 Nov 1847

2/13. Brachylaena (11; tropical and southern Africa, Madagascar, the Mascarene Islands), Tarchonanthus (2; Africa, the Arabian Peninsula). – Tropical and southern Africa, Madagascar, the Mascarene Islands, the Arabian Peninsula. Trees or shrubs. Capitula discoid, with imbricate involucral bracts, with or without hyaline margins. Corolla of male florets actinomorphic, deeply quinquelobate, tubular. Corolla of female florets actinomorphic, tri- to quinquelobate, tubular to filiform. Anther thecae calcarate and caudate. Stylar branches without sweeping hairs. Pappus consisting of uni- or biseriate barbellate bristles (sometimes absent). x = 9.

[Pertyoideae+[Gymnarrhenoideae+[Cichorioideae+[Corymbioideae+Asteroideae]]]]

Pollen grains with unbranched columellae.

Pertyoideae Panero et V. A. Funk in Proc. Biol. Soc. Washington 115: 915. 30 Dec 2002

4/c 80. Pertya (c 25; Afghanistan, China, Japan, Thailand), Macroclinidium (3; Japan), Ainsliaea (c 50; Afghanistan, Himalayas, Bangladesh, East Asia to Japan, Southeast Asia, West Malesia to the Philippines), Myripnois (1; M. dioica; northern China). – Afghanistan, Himalayas to Japan and Southeast Asia, West Malesia to the Philippines. Corolla deeply but unequally divided. Pollen grains spheroidal. Stylar branches “short”, hairy to papillate on abaxial side, with apex rounded to acuminate. Pappus consisting of bristles (sometimes plumose). x = 13, 14. – The position of Catamixis (1) is unresolved. Pertyoideae resemble Mutisieae but the flowers are not bilabiate.

[Gymnarrhenoideae+[Cichorioideae+[Corymbioideae+Asteroideae]]]

Pollen grains with tectum echinate.

Gymnarrhenoideae Panero et V. A. Funk in Proc. Biol. Soc. Washington 115: 912. 30 Dec 2002

1/1. Gymnarrhena (1; G. micrantha; North Africa, southwestern Asia). – Amphicarpic annuals. Capitula aggregated into synflorescences, with florets surrounded by small involucral bracts. Synflorescences disciform, congested in centre of leaf rosette. Corolla in subterranean capitula vestigial, in aerial capitula filiform. Anther thecae ecalcarate and caudate. Pollen grains spheroidal; colpus ends acute; tectum echinate, non-lophate. Stylar branches long. Cypsela of three types, with twin hairs. Pappus consisting of scale-like bristles, or vestigial to absent. x = 10. – Subterranean capitula are homogamous, with entirely female florets, and cleistogamous. Aerial capitula are heterogamous, functionally male florets interspersed with female florets.

[Cichorioideae+[Corymbioideae+Asteroideae]]

Pollen grains at least sometimes caveate; tectal spines 2–5 µm. Stylar branches “medium” to “long”. Deletion in plastid gene ndhF.


Cichorioideae (Juss.) Chevall., Fl. Gén. Env. Paris 2: 531. 5 Jan 1828

c 250/3.130–3.230. Cosmopolitan. Ray florets present in some species, disc florets strap-shaped and deeply lobate. Pollen grains spheroidal, sometimes caveate, with cavities separating columellae from foot layer, or lophate, with internal tectum; columellae aggregated below spines. Carpels collateral. Stylar branches acute. Hairs usually acute. x = (7–)9–10(–13). – [Cichorieae+[Eremothamneae+[Arctotideae+ [Platycarpheae+[Liabeae+[Distephanus+Moquinieae+Vernonieae]]]]]]. – Hetero-lepis (3), shrubs distributed mainly in the Cape provinces, has sometimes been assigned to Arctotideae, but its systematic affiliation is not resolved. It has partially connate involucral bracts, and pappus consisting of biseriate bristle-like scales.

Cichorieae Lam. et DC., Syn. Plant. Fl. Gall.: 255. 30 Jun 1806 [‘Cichoraceae’]

c 85/1.600–1.700 (“macrospecies”). Warionia (1; W. saharae; northwestern Sahara), Scolymus (3; the Mediterranean), Cichorium (6; Europe, the Mediterranean, Ethiopia), Catananche (5; the Mediterranean), Gundelia (2; Cyprus, Turkey to Afghanistan), Hymenonema (2; Greece), Rothmaleria (1; R. granatensis; southern Spain), Acanthocephalus (2; Central Asia), Chondrilla (c 25; Europe, temperate Asia), Crepidiastrum (7; East Asia), Crepis (c 200; the Northern Hemisphere, tropical and southern Africa, South America), Dianthoseris (1; D. schimperi; East and Northeast African highlands), Dubyaea (10; Himalayas, Tibet, western China), Heteracia (2; southwestern Asia and eastwards to China), Heteroderis (1; H. pusilla; southwestern to Central Asia, Pakistan), Ixeridium (13; East and Southeast Asia, Malesia to New Guinea), Ixeris (20; East and Southeast Asia), Lagoseriopsis (1; L. popovii; Central Asia), Lapsana (1; L. communis; Europe, western and southwestern Asia), Lapsanastrum (4; East Asia, North America), Phitosia (1; P. crocifolia; Greece), Soroseris (7; Himalayas, Tibet, western China), Spiroseris (1; S. phyllocephala; Pakistan), Syncalathium (2; Tibet, China), Taraxacum (c 60 “macrospecies”, >1.000 microspecies; temperate regions on the Northern Hemisphere, New Zealand, Stewart Island, Chatham Islands, temperate and subantarctic South America), Willemetia (2; Central and South Europe and eastwards to Iran), Youngia (c 30; Asia), Cephalorrhynchus (14; southwestern to Central Asia), Melanoseris (60–80; Himalayas, Tibet, China), Cicerbita (20–35; Europe, temperate Asia), Mulgedium (12; Europe, temperate Asia), Lactuca (50–75; nearly cosmopolitan except tropical America), Nabalus (15; East Asia, North America), Notoseris (12; China South of Yangtze), Paraprenanthes (11; East and Southeast Asia), Parasyncalathium (1; P. souliei; Himalayas to western China), Prenanthes (c 30; temperate regions on the Northern Hemisphere, African mountains), Stenoseris (5; Central Asia to China), Dendroseris (11; Juan Fernandez), Launaea (54; the Canary Islands, the Mediterranean, Africa, Madagascar, East Asia, the West Indies), Reichardia (8; Macaronesia, the Mediterranean), ’Sonchus’ (c 60; Europe, Macaronesia, the Mediterranean to tropical Africa, Asia to Southeast Asia; non-monophyletic), Thamnoseris (1; T. lacerata; San Ambrosio Island in Chile), Agoseris (11; western North America, Mexico, temperate South America), Krigia (7; North America, Mexico), Microseris (14; southeastern Australia, Tasmania, New Zealand, western North America, Mexico, Peru, Chile), Stebbinsoseris (2; California, Arizona, northwestern Mexico; S. decipiens derived from Microseris bigelovii x Uropappus lindleyi; S. heterocarpa derived from Microseris douglasii x Uropappus lindleyi), Uropappus (1; U. lindleyi; California, western Texas, northern Mexico), Phalacroseris (1; P. bolanderi; California), Picrosia (2; tropical and subtropical South America), Pyrrhopappus (1–5; North America, northern Mexico), Chaetadelpha (1; C. wheeleri; southwestern United States), Lygodesmia (5; North America, Mexico), Marshalljohnstonia (1; M. gypsophila; Mexico), Munzothamnus (1; M. blairii; San Clemente Island in California), Pleiacanthus (1; P. spinosus; western United States), Prenanthella (1; P. exigua; western United States, northern Mexico), Rafinesquia (2; southwestern United States, northwestern Mexico), Shinnersoseris (1; S. rostrata; North America, Mexico), Stephanomeria (c 15; western North America, northwestern Mexico), Anisocoma (1; A. acaulis; southwestern United States, northwestern Mexico), Atrichoseris (1; A. platyphylla; southwestern United States), Calycoseris (2; southwestern United States, northwestern Mexico), Glyptopleura (2; western United States), Malacothrix (21; western North America, northern Mexico), Pinaropappus (7–10; southern United States, Mexico, Central America), Hieracium (c 100 “macrospecies”, c 6.000 “microspecies”; temperate regions on both hemispheres, tropical mountains), Hispidella (1; H. hispanica; the Iberian Peninsula), Hololeion (3; East Asia), Pilosella (20–80?; Europe, the Mediterranean, North Africa, temperate Asia), Tolpis (c 15; Macaronesia, the Mediterranean, Ethiopia, Somalia, southern Africa), Aposeris (1; A. foetida; Central Europe), Arnoseris (1; A. minima; Europe), Garhadiolus (4; southwestern Asia to China), Hedypnois (3; Macaronesia, the Mediteranean and eastwards to Iran), Helminthotheca (4; southeastern Europe and eastwards to Iran), Hyoseris (5; the Mediterranean), Hypochaeris (c 60; Europe, Macaronesia, the Mediterranean, North Africa, Asia, South America), Leontodon (c 25; Europe, the Mediterranean, temperate Asia, southwestern Asia to Iran), Picris (c 40; Europe, the Mediterranean, African mountains, Asia, northern Australia), Rhagadiolus (2; the Mediterranean and eastwards to Iran), Scorzoneroides (c 25; Europe, the Mediterranean, temperate Asia), Urospermum (2; the Mediterranean and eastwards to Pakistan), Epilasia (3; western and Central Asia to China), Geropogon (1; G. hybridus; southern Europe, the Mediterranean, Turkey), Koelpinia (5; the Mediterranean, North Africa and eastwards to East Asia), Pterachaenia (1; P. stewartii; Afghanistan, Pakistan), Scorzonera (175–180; Europe, the Mediterranean and eastwards to western and Central Asia), Tourneuxia (1; T. variifolia; Algeria), Tragopogon (c 110; Europe, the Mediterranean, temperate Asia). – Cosmopolitan, with their highest diversity in temperate regions on the Northern Hemisphere. Usually herbs (rarely shrubs or small trees). Laticifers (with white latex) present. Capitula usually ligulate (not in Gundelia and Warionia), with few outer series of imbricate bracts and one long inner series of bracts, often with hyaline margin. Corolla quinquedentate ligulate. Anther thecae calcarate and caudate. Pollen grains echinolophate or echinate. Style with sweeping hairs. Pappus usually consisting of scales or scabrid to barbellate or plumose bristles (rarely absent). x = 9. - Several genera in Cichorieae are agamospermous, i.a. Taraxacum, with about 2.000 microspecies, and Hieracium, with probably far more than 8.000 microspecies. In the species enumerations above, only the “macrospecies” have been numbered. In many groups (especially in Pilosella and parts of Hieracium) the “species” number is constantly in a state of flux, since the numerous microspecies are not fixed apomicts, but mainly amphiapomictic.

[Eremothamneae+[Arctotideae+[Platycarpheae+[Liabeae+[Distephanus+Moquinieae+Vernonieae]]]]]

Eremothamneae H. Rob. et Brettell in Phytologia 26: 164. 16 Jul 1973

2/3. Eremothamnus (1; E. marlothianus; Namibia), Hoplophyllum (2; Northern and Western Cape). – Western South Africa, southern Namibia. Shrubs. Capitula radiate or discoid, with gradate multiseriate involucral bracts, distally papyraceous and usually with apical spine. Ray corollas shortly tridentate strap-shaped or absent, disc corollas quinquelobate tubular. Anther thecae calcarate and caudate. Style branches with long sweeping hairs consisting of two or three cells separated by longitudinal walls. Pappus consisting of bi- or triseriate scabrid-barbellate capillary bristles. x = 9.

[Arctotideae+[Platycarpheae+[Liabeae+[Distephanus+Moquinieae+Vernonieae]]]]

Arctotideae Cass. in J. Phys. Chim. Hist. Nat. Arts 88: 159. Feb 1819

14/c 200. Heterolepis (4; Western Cape); Arctotheca (5; Mozambique, southern Africa), Arctotis (c 50; Angola, southern Africa), Cymbonotus (3; southeastern South Australia to southeastern Queensland, Tasmania), Dymondia (1; D. margaretae; Bredasdorp in Western Cape), Haplocarpha (10; East Africa to Ethiopia), Berkheya (75–80; tropical and southern Africa), Cullumia (15; Northern and Western Cape), Cuspidia (1; C. cernua; Northern, Western and Eastern Cape), Didelta (2; southern Namibia, Northern and Western Cape), Gazania (17; tropical and southern Africa, with their highest diversity in the Cape Provinces), Gorteria (8; southern Namibia, Northern, Western and Eastern Cape), Heterorhachis (1; H. aculeata; Northern and Western Cape), Hirpicium (13; tropical and southern Africa). – Tropical and southern Africa, southeastern Australia, Tasmania, with their highest diversity in the Cape Provinces. Shrubs or herbs. Latex present (Gorteriinae) or absent (Arctotidinae). Capitula radiate, with imbricate involucral bracts, outer with foliaceous apex, inner with scarious apex. Ray corollas tri- or quadrilobate, disc corollas shallowly quinquelobate. Anther thecae calcarate and caudate or ecaudate. Style with thickened apex and small broadly subulate or clavate patent sweeping hairs, longest in ring below branching point. Cypsela sometimes winged. Pappus consisting of uni- or biseriate scales. x = 9. – The placement of Heterolepis (x = 10) is very unclear. It is positioned as sister-group either to the Arctotideae or to the Platycarpheae, or inside the Arctotideae, depending on the datasets used.

[Platycarpheae+[Liabeae+[Distephanus+Moquinieae+Vernonieae]]]

Platycarpheae V. A. Funk et H. Rob., Compositae Newslett. 47: 25. 15 Apr 2009

1/3. Platycarpha (1; P. glomerata; eastern South Africa), Platycarphella (2; Namibia, South Africa, Botswana). – Southern Africa. Perennial herbs. Capitula discoid, with slightly imbricate involucral bracts. Corollas quinquelobate tubular. Anther thecae calcarate and ecaudate (or caudate). Style usually hairy distally. Pappus consisting of scales. x = 9. – According to Funk & Koekemoer (2011) the distributional pattern of the three species may be explained by fragmentation of a widespread ancestor due to the rise of the Great Escarpment combined with changes in the climate.

[Liabeae+[Distephanus+Moquinieae+Vernonieae]]

Liabeae Rydb. in N. Amer. Fl. 33: 46. 15 Sep 1922

c 18/c 175. Cacosmia (3; Ecuador, Peru), Ferreyranthus (8; Ecuador, Peru), Oligactis (7; Costa Rica, Panamá, Colombia, Ecuador, Venezuela), Dillandia (3; Colombia, Peru), Sampera (8; Colombia, Ecuador, Peru), Liabum (37; Mexico, Central America, the West Indies, tropical South America), Sinclairia (27; Mexico, Central America, Colombia), Stephanbeckia (1; S. plumosa; Bolivia), Microliabum (5; Bolivia, Argentina), Pseudonoseris (3; Peru), Paranephelius (7; Peru, Bolivia, Argentina), Chionopappus (1; C. benthamii; Peru), Philoglossa (5; Colombia, Ecuador, Peru, Bolivia), Erato (5; Costa Rica, Panamá, northwestern South America to Bolivia and Venezuela), ‘Munnozia’ (46; Costa Rica, Panamá, northwestern South America to northern Argentina and Venezuela; paraphyletic; incl. Chrysactinium?), Chrysactinium (8; Ecuador, Peru; in Munnozia?), Bishopanthus (1; B. soliceps; Peru)?, Inkaliabum (1; I. diehlii; Peru). – Mexico, Central America, the West Indies, tropical South America, with their largest diversity in northwestern South America. Small trees, shrubs or annual or perennial herbs. Latex usually present. Leaves usually opposite. Capitula radiate, with graduate involucral bracts. Ray corollas trilobate strap-shaped, disc corollas quinquelobate, tubular. Anther thecae calcarate and caudate. Style with sweeping hairs in distal parts. Pappus consisting of long inner capillary bristles and short outer multiseriate squamellae (sometimes scales, plumose bristles or absent). x = 9.

[Distephanus+Moquinieae+Vernonieae]

Distephaninae S. C. Kelley et H. Rob. in V. A. Funk et al., Syst. Evol. Biogeogr. Compositae: 448. 25 Aug 2009

1/c 35. Distephanus (c 35; tropical and southern Africa, Madagascar, Mauritius). – Trees, shrubs or lianas. Leaves often trinervate or triplinervate. Capitula discoid, with usually persistent involucral bracts. Anther thecae caudate. Stylar base thickened. Pappus consisting of biseriate capillary bristles. n = 10. Elemanolides present.

Moquinieae H. Rob. in Taxon 43: 39. 8 Feb 1994

2/2. Moquinia (2; eastern Brazil), Pseudostifftia (1; P. kingii; Bahia in Brazil). – Brazil. Shrubs. Involucral bracts with hyaline margins. Corollas actinomorphic, quinquelobate. Anther thecae calcarate and caudate. Style swollen and scabrous near branching point, branches scabrous. Achenes with twin hairs. Pappus biseriate-capillary. n = ? Guaianolides present.

Vernonieae Cass. in J. Phys. Chim. Hist. Nat. Arts 88: 203. Mar 1819

c 130/>1.125. Acanthodesmos (1; A. distichus; Jamaica), Gorceixia (1; G. decurrens; southeastern Brazil), Bolanosa (1; B. coulteri; Mexico), Eremosis (20; Mexico, Central America), Leiboldia (1; L. serrata; Mexico), Lepidonia (7; Mexico, Central America), Stramentopappus (1; S. pooleae; Mexico), Albertinia (1; A. brasiliensis; Brazil), Aynia (1; A. pseudascaricida; Peru), Chrysolaena (9; tropical South America), Cololobus (3; Brazil), Cyrtocymura (8; tropical America, with their highest diversity in Brazil), Dasyanthina (2; eastern Brazil), Dipterocypsela (1; D. succulenta; Colombia), Echinocoryne (6; Brazil), Eirmocephala (3; Central America, the Andes in Colombia to Bolivia), Gossweilera (2; Angola), Harleya (1; H. oxylepis; Mexico, Central America), Heterocypsela (1; H. andersonii; Brazil), Lepidaploa (c 115; tropical America), Lessingianthus (c 110; South America, with their highest diversity in Brazil), Manyonia (1; M. peculiaris; Tanzania), Mattfeldanthus (2; Brazil), Pseudopiptocarpha (2; tropical America), Quechualia (4; Bolivia, Argentina), Sparganophoros (1; S. sparganophora; tropical America), Stenocephalum (5; tropical America), Stilpnopappus (24; tropical South America), Tephrothamnus (3; northern tropical South America), Trepadonia (2; Peru), Vernonanthura (>65; tropical America), Vernonia (27; tropical to warm-temperate America), Lepidaploa (c 120; tropical America), Xiphochaeta (1; X. aquatica; northeastern South America), Blanchetia (1; B. heterotricha; northeastern Brazil), Critoniopsis (28; South America, with their largest diversity in the northern Andes), Cuatrecasanthus (3; Ecuador, Peru), Dasyandantha (1; D. cuatrecasasiana; Venezuela), Ekmania (1; E. lepidota; Cuba), Huberopappus (1; H. maigualidae; Venezuela), Irwinia (1; I. coronata; Brazil), Joseanthus (5; Colombia, Ecuador), Piptocarpha (c 45; tropical America), Piptocoma (18; tropical America), Chresta (11; central Brazil), Pithecoseris (1; P. pacurinoides; northern Brazil), Soaresia (1; S. velutina; Brazil), Centratherum (5; northern, eastern and southwestern Australia), Oiospermum (1; O. involucratum; northeastern Brazil), Anteremanthus (1; A. hatschbachi; Brazil), Chronopappus (1; C. bifrons; Brazil), Eremanthus (c 20; Brazil, with their highest diversity in semiarid parts of the cerrado), Lychnophora (c 25; Brazil), Lychnophoriopsis (4; Brazil), Minasia (6; Brazil), Piptolepis (9; Brazil), Prestelia (1; P. eriopus; Brazil), Proteopsis (2; southern Brazil), Bishopalea (1; B. erecta; Brazil), Heterocoma (1; H. albida; Brazil), Sipolisia (1; S. lanuginosa; Brazil), Xerxes (1; X. ekmanianum; Brazil), Caatinganthus (2; Brazil), Elephantopus (c 25; tropical and subtropical regions on both hemispheres), Pseudelephantopus (2; tropical America), Rolandra (1; R. fruticosa; tropical America), Spiracantha (1; S. cornifolia; Central America, Colombia, Venezuela), Stokesia (1; S. laevis; southeastern United States), Pacourina (1; P. edulis; tropical America), Acilepis (33; India to China), Ageratinastrum (2; tropical Africa), Ambassa (1; A. hochstetteri; Ethiopia), Bechium (3; Madagascar), Bothriocline (>30; tropical and southern Africa, Madagascar), Cyanthillium (1–2; C. cinereum; tropical Africa, Madagascar, tropical Asia), Decastylocarpus (1; D. perrieri; Madagascar), Dewildemania (3; tropical Africa), Diaphractanthus (1; D. homolepis; Madagascar), Erlangea (6; tropical and southern Africa, Madagascar), Ethulia (19; tropical Africa, Madagascar, tropical Asia), Gutenbergia (c 25; tropical Africa), Herderia (1; H. truncata; tropical West Africa), Hilliardiella (>8; eastern and southern Africa, the Arabian Peninsula), Hoffmannanthus (1; H. abbotianus; tropical Central and East Africa to Angola, Malawi and Zambia), Hystrichophora (1; H. macrophylla; Tanzania, extinct?), Iodocephalus (2; Thailand, Laos), Jeffreycia (5; Somalia, Kenya, Tanzania, Sri Lanka), Kinghamia (5; tropical West and Central Africa), Koyamasia (1; K. calcarea; Thailand), Lettowia (1; L. nyassae; East Africa), Mesanthophora (1; M brunneri; central Paraguay), Msuata (1; M. buettneri; Congo), Muschleria (1; M. angolensis; Angola), Okia (2; Burma, Thailand), Omphalopappus (1; O. newtonii; Angola), Oocephala (2; tropical Africa), Orbivestus (11; tropical and southern Africa), Phyllocephalum (3; India, Java), Pulicarioidea (1; P. annamica; Burma, Thailand, Laos, Vietnam), Rastrophyllum (2; Tanzania, Zambia), Telmatophila (1; T. scolymastrum; Brazil), Vernoniastrum (8; tropical Africa), Brachythrix (6; Central and East Africa), Cabobanthus (>2; tropical Africa), Centauropsis (8; Madagascar), Centrapalus (1; C. africana; southern Africa), Nothovernonia (2; tropical Africa), Parapolydora (1; P. fastigiata; Nambia, South Africa, Zimbabwe), Asia), Aedesia (3; tropical Africa), Ananthura (1; A. pteropoda; Central Africa), Baccharoides (c 25; tropical and southern Africa, India), Camchaya (5; Southeast Asia), Khasianthus (1; K. subsessilis; India, southern China, Burma), Lachnorhiza (2; western Cuba), Linzia (1; L. accommodata; Zimbabwe), Neurolakis (1; N. modesta; Cameroun), Pleurocarpaea (2; tropical Australia), Vernonella (11; tropical and southern Africa), Brenandendron (3; tropical Africa), Decaneuropsis (12; India, southern China, Southeast Asia, West Malesia), Gymnanthemum (>40; tropical and southern Africa, Madagascar, tropical Asia to southern China and Burma), Hesperomannia (3; the Hawaiian Islands), Kurziella (1; K. gymnoclada; Southeast Asia), Lampropappus (3; tropical Africa), Monosis (7; tropical Asia), Myanmaria (1; M. calycina; Burma), Oliganthes (9; Madagascar), Strobocalyx (6; tropical Asia from India to New Guinea), Tarlmounia (1; T. elliptica; Thailand), Trichospira (1; T. verticillata; tropical America), Uniyala (7; southern India, Sri Lanka). – Tropical and subtropical regions on both hemispheres. Small trees, shrubs, lianas or herbs. Latex sometimes present. Leaves usually alternate. Involucral bracts gradate to subequal. Corollas usually actinomorphic, quinquelobate. Anther thecae calcarate or ecalcarate, caudate or ecaudate. Stylar branches with long sweeping hairs. Pappus usually capillary. x = 10.

[Corymbioideae+Asteroideae]

Pollen grains fully caveate, without columellae spanning space above foot layer, and usually with internal foramina; tectal spines pointed.

Corymbioideae Panero et V. A. Funk in Proc. Biol. Soc. Washington 115: 910. 30 Dec 2002

1/9. Corymbium (9; Western and Eastern Cape). – Perennial herbs. C Leaves long, linear, parallelodromous. Capitulas discoid, with one floret enclosed by two innermost involucral bracts. Corolla quinquelobate, with linear to oblong and patent lobes. Pollen grains spheroidal; internal foramina sometimes absent. Ovary densely hairy. Stylar branches “long”, hairy, with tapering apex. Cypsela densely pubescent. Pappus consisting of basally connate scales and/or thin bristles. n = 16; x = 8.

Asteroideae Lindl. in J. C. Loudon, Encycl. Plant.: 1074. 1829 [‘Astereae’]

1.151–1.156/15.600–16.000. Cosmopolitan. Laticifers absent. Leaves usually alternate (sometimes opposite). Female capitula sometimes uniflorous. Trilobate ray florets frequent (female). Ray corollas tridentate or absent, disc corollas usually actinomorphic. Anthers sometimes free from each other (especially in anemophilous species), often without appendage. Anther thecae often ecalcarate and usually ecaudate. Pollen grains spheroidal, caveate, with cavities separating columellae from foot layer; colpus ends acute. Tectum double. Stylar hairs often rounded, present only at stylar apex. Stigmatic areas in two marginal bands. x = (4–)9–10(–19). ‘4 x 6 bp’ inversion in plastid gene rbcL. Sesquiterpene lactones at biogenetic levels 3 and 4, benzopyrans, benzofurans, and 6,8-deoxygenation of flavonoids present. – The probable topology is [Abrotanella+Doronicum+Senecioneae+[[Calenduleae+[Gnaphalieae+[Astereae+Anthemideae]]]+[Inuleae+[Athroismeae+[Feddeeae+[Helenieae+[Coreopsideae+[[Neurolaeneae+[Tageteae+[Bahieae+Chaenactideae]]]+[[Polymnieae+[Heliantheae+[Millerieae+[Madieae+[Eupatorieae+Perityleae]]]]]]]]]]]]]]]].

Abrotanellinae H. Rob., G. D. Carr, R. M. King et A. M. Powell in Ann. Missouri Bot. Gard. 84: 896. 1998

1/18. Abrotanella (18; mountains on New Guinea, eastern Victoria, Tasmania, New Zealand, southernmost South America). – Perennial herbs (often cushion-plants). Capitula discoid, with biseriate involucral bracts. Pollen grains “senecioid”. Pappus absent. x = 9. – The sister-group relationship of Abrotanella is unresolved.

Doroniceae Panero in Phytologia 87: 1. 5 Apr 2005

1/26. Doronicum (26; Europe except northern parts, the Mediterranean, temperate Asia). – Perennial herbs. Capitula radiate, with bi- or triseriate equal involucral bracts. Ray corollas trilobate strap-shaped, disc corollas quinquedentate tubular. Pappus usually consisting of uni- or biseriate barbellate bristles (sometimes absent). x = 30. – The sister-group relationship of Doronicum is unresolved.

Senecioneae Cass. in J. Phys. Chim. Hist. Nat. Arts 88: 195. Mar 1819

c 140/c 3.400. Blennosperma (3; California, Chile), Crocidium (1; C. multicaule; northwestern North America), Ischnea (4; New Guinea), Chersodoma (9; the Andes), Homogyne (3; mountains in Central Europe), Endocellion (2; Siberia), Petasites (c 20; temperate, arctic and alpine regions on the Northern Hemisphere), Tussilago (1; T. farfara; Europe, the Mediterranean, North Africa, temperate Asia), Farfugium (3; East Asia), Miricacalia (1; M. makinoana; Japan), Dendrocacalia (1; D. crepidifolia; Bonin Islands), ’Parasenecio’ (c 70; Russia, East Asia, Alaska; polyphyletic), Sinacalia (4; China; in Parasenecio?), Dicercoclados (1; D. triplinervis; China), Syneilesis (7; East Asia), ’Ligularia’ (>125; eastern Europe, temperate Asia, with their highest diversity in China; paraphyletic), Ligulariopsis (1; L. shichuana; China), ’Cremanthodium’ (70–75; Himalayas, Tibet, western and southern China; polyphyletic), Nemosenecio (6; China, Japan), Sinosenecio (36; China, Southeast Asia, northwestern North America), Hainanecio (1; H. hainanensis; Hainan), Tephroseris (c 50; temperate and arctic regions on the Northern Hemisphere), Psacaliopsis (5; Mexico, Guatemala), Psacalium (c 40; southwestern United States, Central America), Pippenalia (1; P. delphiniifolia; Mexico), Digitacalia (5; Mexico), Rugelia (1; R. nudicaulis; southeastern United States), Arnoglossum (8; eastern United States), Robinsonecio (2; Mexico, Guatemala), Roldana (c 55; southern United States, Mexico, Central America), Nelsonianthus (2; Mexico, Guatemala), Telanthophora (14; Central America), Villasenoria (1; V. orcuttii; Mexico), Pittocaulon (5; Mexico, Central America), Barkleyanthus (1; B. salicifolius; southwestern United States, Mexico, Central America), Yermo (1; Y. xanthocephalus; Wyoming), Rainiera (1; R. stricta; northwestern United States), Cacaliopsis (1; C. nardosmia; western United States), Luina (2; northwestern North America), Tetradymia (10; North America), Lepidospartum (3; southwestern United States), Paracalia (2; Peru, Bolivia), Paragynoxys (15; northwestern tropical South America), Gynoxys (c 130; Central America, northern South America to Peru), Nordenstamia (c 20; western Ecuador, western Peru to Argentina), Aequatorium (c 20; Peru, Bolivia, northern Argentina), Acrisione (2; central Chile), Brachyglottis (c 40; southeastern New South Wales, Tasmania, New Zealand, Three Kings Islands, Chatham Islands, Madagascar?, South America?; paraphyletic?), Centropappus (1; C. brunonis; Tasmania; in Brachyglottis?), Capelio (3; Western Cape), Papuacalia (17; mountains on New Guinea), Brachionostylum (1; B. pullei; New Guinea), Pladaroxylon (1; P. leucadendron; St. Helena), Lachanodes (1; L. arborea; St. Helena), Scrobicaria (2; northeastern South America), Shafera (1; S. platyphylla; eastern Cuba), Mattfeldia (1; M. triplinervia; Hispaniola), Ekmaniopappus (2; Hispaniola), Herodotia (1; H. haitiensis; Hispaniola), Leonis (1; L. trineura; Cuba, Hispaniola), Nesampelos (3; Hispaniola), Zemisia (1; Z. discolor; Jamaica), Antillanthus (17; Cuba), Elekmania (9; Hispaniola), Herreranthus (1; H. rivalis; Cuba), Oldfeltia (1; O. polyphlebia; Cuba), Odontocline (6; Jamaica), Lundinia (1; L. plumbea; Cuba, Hispaniola), Jacmaia (5; Costa Rica, Jamaica), Ignurbia (1; I. constanzae; Hispaniola), Jessea (4; Costa Rica, Panamá), Lordhowea (1; L. insularis; Lord Howe Island), Arrhenechthites (6; mountains on Sulawesi, New Guinea and in eastern New South Wales and Victoria), Bethencourtia (3; the Canary Islands), Pericallis (15; Macaronesia), Cineraria (15–20; tropical and southern Africa, Madagascar, southwestern Arabian Peninsula), Bertilia (1; B. hantamensis; southwestern Northern Cape), Bolandia (5; Western and Eastern Cape, Drakensberg in Lesotho), Mesogramma (1; M. apiifolium; Gariep area in South Africa), Stilpnogyne (1; S. bellidioides; Western Cape), Packera (c 65; Siberia, North America), Dorobaea (3; the Andes), Senecio (c 1.300; cosmopolitan except polar regions), Jacobaea (c 35; Europe, temperate Asia), Solanecio (16; tropical and southern Africa, Madagascar, Yemen), Kleinia (c 40; the Canary Islands, Africa, Madagascar, the Arabian Peninsula and eastwards to southern India and Sri Lanka), Gynura (c 40; tropical regions in the Old World), Erechtites (c 12; America), Io (1; I. amboadrombensis; Madagascar), Crassocephalum (24; tropical and subtropical Africa, Madagascar, the Mascarene Islands, Yemen), Hoehnephytum (3; Brazil), Pentacalia (c 200; Central and South America), Monticalia (c 70; Central and South America, with their largest diversity in the Andes), Dendrophorbium (c 75; South America), Graphistylis (8; southern Brazil), Caxamarca (1; C. sanchezii; northern Peru), Pseudogynoxys (12–14; tropical South America), Arbelaezaster (1; A. ellsworthii; Colombia), Dresslerothamnus (4–5; Central America, Colombia), Cabreriella (2; Colombia), Charadranaetes (1; C. durandii; Costa Rica), Misbrookea (1; M. strigosissima; Peru, Bolivia), Xenophyllum (21; the Andes), Werneria (25–30; the Andes), Oresbia (1; O. heterocarpa; Western Cape), Lamprocephalus (1; L. montanus; Western Cape), Phaneroglossa (1; P. bolusii; Northern and Western Cape), Dendrosenecio (11; mountains in Central and East Africa), Euryops (c 100; Africa, the Arabian Peninsula, Socotra), ’Othonna’ (c 125; tropical and southern Africa, Tasmania, with their highest diversity in southern Africa; diphyletic), Caputia (5; eastern South Africa), Crassothonna (13; southwestern South Africa), Gymnodiscus (2; Northern and Western Cape), Hertia (c 10; northern and southern Africa, southwestern Asia), Lopholaena (c 20; tropical and southern Africa), Cadiscus (1; C. aquaticus; Western Cape), Stenops (2; tropical Africa), Oligothrix (1; O. gracilis; Cedarberg Mountains in Western Cape), Steirodiscus (5–6; Northern and Western Cape), Bafutia (1; B. tenuicaulis; Cameroun), Psednotrichia (3; Angola), Emilia (90–100; tropical and southern Africa, tropical Asia), Emiliella (5; Angola, Zambia), Faujasiopsis (3; the Mascarene Islands), Faujasia (4; Réunion), Eriotrix (2; Réunion), Hubertia (c 25; Madagascar, the Mascarene Islands), Humbertacalia (8; Madagascar, the Mascarene Islands), Parafaujasia (2; the Mascarene Islands), Synotis (50–55; Himalayas, Tibet, China), Mikaniopsis (c 15; tropical and southern Africa), Delairea (1; D. odorata; Western and Eastern Cape, KwaZulu-Natal, Lesotho), Cissampelopsis (10; tropical Asia), Austrosynotis (1; A. rectirama; southern tropical Africa), Dauresia (2; Namibia), Adenostyles (4; Central and South Europe), Caucasalia (4; Turkey, Caucasus), Dolichorrhiza (4; Caucasus, Iran), Iranecio (17; southeastern Europe and eastwards to Iran). – Cosmopolitan except polar regions. Trees, shrubs, lianas, herbs (including epiphytes). Capitula radiate, disciform or discoid, with usually uniseriate (sometimes bi- or multiseriate) equal involucral bracts. Ray corollas often tridentate giving impression of being split zygomorphic (modification of [2:3]-bilabiate corolla, with reduced adaxial lobes), disc corollas quadri- or quinquelobate campanulate or tubular. Anther thecae ecalcarate and caudate or ecaudate. Pollen grains caveate, usually with solid columellae. Style simple or bilobate, with sweeping hairs in apical tuft or on stylar branches. Pappus usually consisting of barbellate bristles (sometimes absent). x = 10. Diester pyrrolizidine alkaloids, eremophilane types sesquiterpene lactones. Polyacetylenes absent.

[[Calenduleae+[Gnaphalieae+[Astereae+Anthemideae]]]+[Inuleae+[Athroismeae+[Feddeeae+[Helenieae+[Coreopsideae+[[Neurolaeneae+[Tageteae+[Bahieae+Chaenactideae]]]+[[Polymnieae+[Heliantheae+[Millerieae+[Madieae+[Eupatorieae+Perityleae]]]]]]]]]]]]]]]

[Calenduleae+[Gnaphalieae+[Astereae+Anthemideae]]]

Calenduleae Cass. in J. Phys. Chim. Hist. Nat. Arts 88: 161. Feb 1819

10/c 105. Calendula (12; Central and South Europe, Macaronesia, the Mediterranean and eastwards to Iran), Oligocarpus (2; O. calendulaceus: Northern, Western and Eastern Cape, KwaZulu-Natal; O. sanctae-helenae: St. Helena), Norlindhia (3; southwesternmost Namibia, western Northern and Western Cape), Monoculus (2; southwesternmost Namibia, western Northern and Western Cape), Inuloides (1; I. tomentosa; southwestern and southern Western Cape), Osteospermum (c 65; tropical eastern and southern Africa, northeastern Africa, Somalia, the Arabian Peninsula, Jordan), Gibbaria (2; Western and Eastern Cape), Nephrotheca (1; N. ilicifolia; Western Cape), Dimorphotheca (9; tropical and southern Africa), Garuleum (8; southern Africa). – Europe, Macaronesia, the Mediterranean, Africa, St. Helena, southwestern Asia, with their largest diversity in the Cape Provinces. Small trees, shrubs or herbs. Leaves alternate or opposite. Capitula radiate, with uni- to triseriate involucral bracts, with or without hyaline margins. Disc corollas quinquelobate. Anther thecae ecalcarate and caudate. Exine without columellae. Style simple or bilobate, with sweeping hairs distally. Cypsela terete or flattened, sometimes curved, rostrate, winged or fenestrate. Exocarp sometimes fleshy. Pappus absent. x = 10.

[Gnaphalieae+[Astereae+Anthemideae]]

Gnaphalieae (Cass.) Lecoq et Juill., Dict. Rais. Term. Bot.: 296. 1831

179/2.020–2.150. Acanthocladium (1; A. dockeri; eastern South Australia, western New South Wales), Achyrocline (32; tropical Africa, Madagascar, South America), Acomis (4; northern Northern Australia, Queensland, northeastern New South Wales), Actinobole (4; southern and central Australia), Alatoseta (1; A. tenuis; Northern and Western Cape), Ammobium (3; southeastern Queensland, eastern New South Wales, eastern Victoria, Tasmania), Amphiglossa (12; southern Africa), Anaphalis (c 110; temperate and alpine Asia and North America), Anaphaloides (7; New Guinea, New Zealand), Anaxeton (9; Western Cape), Ancistrocarphus (2; California), Anderbergia (6; Western Cape), Anemocarpa (3; central and southeastern Australia), Angianthus (18; western, central and southern Australia, Tasmania), Anisochaeta (1; A. mikanioides; Eastern Cape, KwaZulu-Natal), Anisothrix (2; Western Cape), Antennaria (c 45; temperate, arctic and alpine regions on both hemispheres), Antithrixia (1; A. flavicoma; Namaqualand in Western Cape), Apalochlamys (1; A. spectabilis; southeastern South Australia, Victoria, Tasmania), Argentipallium (6; southwestern Western Australia, southeastern South Australia, Victoria, Tasmania), Argyroglottis (1; A. turbinata; southwestern Western Australia), Argyrotegium (4; eastern New South Wales, Victoria, Tasmania, New Zealand), Arrowsmithia (1; A. styphelioides; Eastern Cape), Artemisiopsis (1; A. villosa; tropical and southern Africa), Asteridea (9; southwestern Western Australia, southern South Australia, western Victoria), Athrixia (14; tropical and southern Africa, Madagascar), Atrichantha (1; A. gemmifera; Western Cape), Basedowia (1; B. tenerrima; northern South Australia), Bellida (1; B. graminea; southwesternmost Western Australia), Belloa (9; the Andes in Venezuela to central Chile), Berroa (1; B. gnaphalioides; subtropical South America), Blennospora (3; southwestern Western Australia, southeastern South Australia, western Victoria), Bryomorphe (1; B. aretioides; Western Cape), Callilepis (5; South Africa, Swaziland), Calocephalus (11; Australia, Tasmania), Calomeria (1; C. amaranthoides; eastern New South Wales, Victoria), Calotesta (1; C. alba; Western Cape), Cassinia (18–20; eastern and southeastern Australia, Tasmania), Castroviejoa (2; Corsica, Sardinia), Catatia (2; Madagascar), Cephalipterum (1; C. drummondii; southwestern Western Australia, western South Australia), Cephalosorus (1; C. carpesioides; westernmost Western Australia), Chamaepus (1; C. afghanicus; Afghanistan), Chevreulia (6; South America, Falkland Islands, Tristan da Cunha), Chiliocephalum (2; Ethiopia), Chionolaena (18; central Mexico, Costa Rica, Colombia, southern Venezuela, southern Brazil), Chondropyxis (1; C. halophila; southwestern Western Australia, southern South Australia), Chrysocephalum (8; Australia, Tasmania), Chthonocephalus (6; southern Australia), Cladochaeta (2; Caucasus), Comborhiza (2; Western Cape, KwaZulu-Natal), Craspedia (23; southwestern Western Australia, southeastern Australia, Tasmania), Cremnothamnus (1; C. thomsonii; southern Northern Territory, northern South Australia), Cuatrecasasiella (2; the Andes), Cymbolaena (1; C. griffithii; southwestern to Central Asia), Decazesia (1; D. hecatocephala; westernmost Western Australia), Denekia (1; D. capensis; southern and southeastern Africa), Dielitzia (1; D. tysonii; western Western Australia), Disparago (9; Western and Eastern Cape, KwaZulu-Natal), Dithyrostegia (2; southwestern Western Australia), Dolichothrix (1; D. ericoides; Western and Eastern Cape), Edmondia (3; Western Cape), Elytropappus (10; Northern, Western and Eastern Cape), Epitriche (1; E. demissus; southwesternmost Western Australia), Eriochlamys (2; southeastern and central Australia), Erymophyllum (5; southwestern Western Australia, western South Australia), Euchiton (17–20; East Asia, New Guinea, Australia except northwestern parts, Tasmania, New Zealand), Evacidium (1; E. discolor; Sicily, North Africa), Evax (c 15; the Mediterranean and eastwards to Central Asia), Ewartia (4; southeasternmost New South Wales, eastern Victoria, Tasmania), Ewartiothamnus (1; E. sinclairii; New Zealand), Facelis (4; South America), Feldstonia (1; F. nitens; western Western Australia), Filago (12; temperate and subtropical regions in the Old World), Fitzwillia (1; F. axilliflora; southwestern Western Australia), Galeomma (2; Namibia, Northern Cape, Botswana), Gamochaeta (c 50; tropical and subtropical America), Gamochaetopsis (2; subantarctic southern Andes), Gilberta (1; G. tenuifolia; southwestern Western Australia), Gilruthia (1; G. osbornii; western Western Australia), Gnaphaliothamnus (1; G. lavendulaceum; Mexico, Guatemala), Gnaphalium (c 40; cosmopolitan), Gnephosis (8; Australia), Gnomophalium (1; G. pulvinatum; the Mediterranean), Gratwickia (1; G. monochaeta; South Australia), Haeckeria (3; southeastern South Australia), Haegiela (1; H. tatei; southwestern Western Australia, southeastern South Australia, western Victoria), Haptotrichion (2; Carnarvon District in westernmost Western Australia), Helichrysopsis (1; H. septentrionale; coastal Mozambique, KwaZulu-Natal), Helichrysum (c 600; Europe, Macaronesia, the Mediterranean, Africa, Madagascar, Asia, Australia, with their highest diversity in the Cape Provinces), Hesperevax (3; western United States), Humeocline (1; H. madagascariensis; Madagascar), Hyalochlamys (1; H. globifera; southwestern Western Australia), Hyalosperma (9; southwestern Western Australia, southeastern Australia, Tasmania), Hydroidea (1; H. elsiae; Western Cape), Ifloga (6; the Canary Islands, the Mediterranean, northern to southern Africa, southwestern Asia), Ixiolaena (8; southern Australia), Ixodia (2; southeastern South Australia, Victoria), Jalcophila (3; the Andes in Ecuador, Peru and Bolivia), Lachnospermum (4; Western Cape), Langebergia (1; L. canescens; Western Cape), Lasiopogon (8; Spain, Africa and eastwards to India), Lawrencella (2; southwestern and central Australia), Leiocarpa (10; Madagascar, Australia), Lemooria (1; L. burkittii; Australia)?, Leontopodium (59; mountains in Central Europe and eastwards to China and Burma), Lepidostephium (2; Eastern Cape, KwaZulu-Natal), Leptorhynchos (c 10; southern and southeastern Australia, Tasmania), Leucochrysum (5; Australia, Tasmania), Leucogenes (2; New Zealand, Stewart Island), Leucophyta (1; L. brownii; coastal areas in southwestern Western Australia to eastern Victoria and Tasmania), Leysera (3; southern and northern Africa, the Arabian Peninsula to Pakistan), Logfia (12; Europe, the Mediterranean and eastwards to Afghanistan, North America), Loricaria (20; the Andes), Lucilia (8; South America), Luciliocline (5; the Andes in Peru, Bolivia and Argentina), Macowania (12; Ethiopia, southern Africa, Yemen), Metalasia (57; South Africa, Lesotho, with their highest diversity in Western Cape), Mexerion (2; Mexico), Micropsis (5; Brazil, Uruguay, Argentina), Micropus (5; western Mediterranean and eastwards to Iran, North America), Millotia (16; southern Australia, Tasmania), Mniodes (5; Peru), Myriocephalus (22; Australia), Neotysonia (1; N. phyllostegia; western Western Australia), Nestlera (1; N. biennis; Northern and Western Cape), Odixia (2; Tasmania), Oedera (18; Western and Eastern Cape), Oreoleysera (1; O. montana; Western Cape), Ozothamnus (c 50; Australia except northwestern parts, Tasmania, New Caledonia, New Zealand), Parantennaria (1; P. uniceps; eastern Victoria, southeastern New South Wales), Pentatrichia (4; Namibia, Northern and Western Cape, Mpumalanga), Petalacte (1; P. coronata; Western Cape), Phaenocoma (1; P. prolifera; Western Cape), Phagnalon (43; the Canary Islands, the Mediterranean, southwestern Asia), Philyrophyllum (2; Namibia, South Africa, Botswana), Pithocarpa (4; southwestern Western Australia), Plecostachys (2; Western and Eastern Cape, KwaZulu-Natal, Swaziland), Podolepis (20; Australia, Tasmania), Podotheca (6; southwestern Western Australia, southern South Australia, Victoria, southwesternmost New South Wales), Pogonolepis (2; southwestern and southern Australia), Polycalymma (1; P. stuartii; southern and central Australia), Pseudognaphalium (90–100; temperate and subtropical regions on both hemispheres), Psilocarphus (5; tropical regions on both hemispheres, northwards to western United States, southwards to southern South America), Psychrophyton (10; New Zealand), Pterochaeta (1; P. paniculata; southwestern Western Australia), Pterygopappus (1; P. lawrencei; Tasmania), Quinetia (1; Q. urvillei; southwestern Western Australia, southeastern South Australia, western Victoria), Quinqueremulus (1; Q. linearis; western Western Australia), Rachelia (1; R. glaria; South Island in New Zealand), Raoulia (26; New Guinea, New Zealand), Raouliopsis (2; the Andes in Colombia), Relhania (13; Northern, Western and Eastern Cape, KwaZulu-Natal, Free State, Lesotho), Rhodanthe (46; Australia, Tasmania), Rhynchopsidium (2; Northern and Western Cape), Rosenia (4; Namibia, Northern, Western and Eastern Cape, North-West, Free State, Botswana), Rutidosis (6; Australia), Schoenia (5; Australia except northern parts), Siloxerus (4; southwestern Western Australia, southeastern South Australia, Victoria, southeastern New South Wales, Tasmania), Sondottia (2; southwestern Western Australia), Stenocline (3; Madagascar, Mauritius), Stenophalium (4; Brazil), Stoebe (c 25; tropical and southern Africa, Madagascar, the Mascarene Islands), Stuartina (2; southeastern Australia), Stuckertiella (2; Argentina), Stylocline (7; southwestern United States, northwestern Mexico), Syncarpha (c 30; Western and Eastern Cape), Syncephalum (5; Madagascar), Taplinia (1; T. saxatilis; central Western Australia), Tenrhynea (1; T. phylicifolia; Eastern Cape, KwaZulu-Natal, Northern Province, Mpumalanga, Swaziland), Thiseltonia (2; southern central Western Australia), Tietkensia (1; T. corrickiae; western central Australia), Trichanthodium (4; southern Australia), Trichogyne (9; Namibia, Northern, Western and Eastern Cape, Mpumalanga, Free State), Triptilodiscus (1; T. pygmaeus; southern Australia, Tasmania), Troglophyton (6; Namibia, Northern, Western and Eastern Cape, KwaZulu-Natal, Free State), Vellereophyton (7; Northern, Western and Eastern Cape), Waitzia (5; southern Australia), Xerochrysum (5; Australia, Tasmania). – Subcosmopolitan, with their highest diversity in southern Africa and Australia. Shrubs or herbs. Capitula usually discoid or disciform, with imbricate multiseriate involucral bracts, usually with upper part papery and basal part cartilaginous. Outer corollas usually filiform or absent. Anther thecae ecalcarate and caudate. Ectexine “gnaphalioid”, with outer columellate layer and basal irregularly interlaced layer. Pappus usually consisting of plumose or barbellate to scabrid capillary bristles. x = 7.

[Astereae+Anthemideae]

Astereae Cass. in J. Phys. Chim. Hist. Nat. Arts 88: 195. Mar 1819

210–215/3.000–3.050. Commidendrum (4; St. Helena), Cuniculotinus (1; C. gramineus; western United States), Doellingeria (11; temperate Asia, North America), Ericameria (36; western North America, northwestern Mexico), Eucephalus (11; North America), Eurybia (22; Europe, temperate Asia, North America), Heteroplexis (2; China), Ionactis (5; North America), Melanodendron (1; M. integrifolium; St. Helena), Microglossa (18; tropical and southern Africa, Madagascar, the Mascarene Islands, tropical Asia), Nannoglottis (8; western China), Oclemena (3; North America), Oreostemma (3; western United States), Psiadia (c 60; tropical Africa, Madagascar, tropical Asia), Psiadiella (1; P. humilis; Madagascar), Sarcanthemum (1; S. coronopus; Rodrigues), Tonestus (4; western North America), Vernoniopsis (1; V. caudata; Madagascar), Amellus (12; southern Africa), Chamaegeron (4; Central Asia), Chrysocoma (20; southern Africa, with their highest diversity in Western Cape), Engleria (2; Angola, southern Africa), Felicia (c 85; tropical and southern Africa, Ethiopia, the Arabian Peninsula), Gymnostephium (8; Western Cape), Jeffreya (2; tropical Africa), Lachnophyllum (2; western to Central Asia), Nolletia (14; southern Spain, North and southern Africa), Poecilolepis (2; Western and Eastern Cape), Polyarrhena (4; Western Cape), Roodebergia (1; R. kitamurae; Western Cape), Zyrphelis (13; tropical and southern Africa), Achnophora (1; A. tatei; southeastern South Australia), Aylacophora (1; A. deserticola; Nequén in Patagonian Argentina), Aztecaster (2; Mexico), Blakiella (1; B. bartsiifolia; Colombia, Venezuela), Cabreraea (1; C. andina; La Rioja and Catamarca in Argentina), Celmisia (c 70; southeastern New South Wales, Victoria, Tasmania, New Zealand), Chiliophyllum (3; Mendoza in Argentina), Chiliotrichiopsis (3; southern Peru, Bolivia, northern Argentina), Chiliotrichum (2; Chile, Argentina and southern Bolivia to Tierra del Fuego), Damnamenia (1; D. vernicosum; Auckland Islands, Campbell Islands), Diplostephium (c 90; Costa Rica, the Andes from Colombia to northern Chile), Dysaster (1; D. cajamarcensis; Peru), Floscaldasia (2; the Andes in Colombia and Ecuador), Flosmutisia (1; F. paramicola; Colombia), Guynesomia (1; G. scoparia; central Chile), Haroldia (1; H. mendocina; Mendoza in Argentina), Hinterhubera (8; the Andes), Katinasia (1; K. cabrerae; Mendoza and Nequén in Argentina), Laestadia (6; the West Indies, tropical Andes), Lepidophyllum (1; L. cupressiforme; Chile, Argentina to Tierra del Fuego), Llerasia (14; Colombia to southern Bolivia), Madagaster (5; Madagascar), Mairia (3; Western Cape), Nardophyllum (7; southern Andes in Chile and Argentina), Novenia (1; N. acaulis; the Andes), Ocyroe (1; O. armata; San Juan to Jujuy in Argentina, northern Chile, southern Bolivia), Olearia (c 180; New Guinea, Australia, Tasmania, New Zealand), Oritrophium (c 20; Mexico, the Andes), Pachystegia (1; P. insignis; South Island of New Zealand), Pacifigeron (1; P. rapensis; Rapa Island), Paleaepappus (1; P. patagonicus; Patagonia), Parastrephia (3; central Andes), Pleurophyllum (3; Campbell Islands, Macquarie Island and other islands south of New Zealand), Printzia (6; Western and Eastern Cape, KwaZulu-Natal, Mpumalanga, Free State), Pteronia (70–80; Africa, especially southwestern South Africa), Remya (3; the Hawaiian Islands), Rochonia (4; Madagascar), Westoniella (6; Costa Rica, Panamá), Brachyscome (c 80; New Guinea, Australia, Tasmania, New Zealand), Allittia (2; South Australia, Victoria, southern New South Wales, Tasmania), Hullsia (1; H. argillicola; central Northern Territory), Pembertonia (1; P. latisquamea; westernmost Western Australia), Calotis (28–30; Southeast Asia to Australia), Ceratogyne (1; C. obionoides; southern Australia), Bellis (8; Europe, the Mediterranea), Bellium (4; the Mediterranean), Akeassia (1; A. grangeoides; tropical Africa), Apodocephala (9; Madagascar), Ceruana (1; C. pratensis; Egypt, tropical Africa), Colobanthera (1; C. waterlotii; Madagascar), Cyathocline (3; tropical Asia), Dacryotrichia (1; D. robinsonii; Zambia), Dichrocephala (10; tropical and southern Africa, Madagascar, tropical Asia), Egletes (6; tropical America), Erodiophyllum (2; southern Australia), Grangea (10; tropical and subtropical Africa, Madagascar, tropical and subtropical Asia), Grangeopsis (1; G. perrieri; Madagascar), Grauanthus (2; tropical Africa), Gyrodoma (1; G. hispida; Mozambique), Heteromma (3; mountains in South Africa and Lesotho), Mtonia (1; M. glandulifera; Tanzania), Nidorella (c 15; tropical and southern Africa, Madagascar), Pilbara (1; P. trudgenii; Hamersley Range in Western Australia), Plagiocheilus (7; South America), Keysseria (12; Central and East Malesia, the Hawaiian Islands), Lagenocypsela (2; New Guinea), Lagenophora (14; tropical Asia and eastwards to islands in the Pacific, tropical America), Myriactis (22; Caucasus and eastwards to Japan and New Guinea, Central America), Novaguinea (1; N. radalliae; New Guinea), Pappochroma (9; southeastern South Australia, Victoria, southeastern New South Wales, Tasmania), Phacellothrix (1; P. cladochaeta; East Malesia, Queensland), Piora (1; P. ericoides; New Guinea), Pytinicarpa (3; New Caledonia, Fiji), Rhamphogyne (1; R. rhynchocarpa; Rodrigues), Rhynchospermum (1; R. verticillatum; East and Southeast Asia), Sheareria (2; China), Solenogyne (3; southeastern South Australia to southeastern Queensland, Tasmania), Thespis (3; Southeast Asia), Archibaccharis (32; Mexico to central Panamá), Baccharis (c 360; America), Heterothalamus (8; South America), Asteropsis (1; A. macrocephala; southern Brazil, Uruguay), Camptacra (2; New Guinea, northern and eastern Australia), Dichromochlamys (1; D. dentatifolia; western and central Australia), Dimorphocoma (1; D. minutula; South Australia, western New South Wales), Elachanthus (3; southern Australia), Exostigma (2; southern Brazil and Uruguay to Bolivia, Paraguay and Argentina), Inulopsis (4; South America), Iotasperma (2; northern and central Australia), Isoetopsis (1; I. graminifolia; southern Australia, Tasmania), Ixiochlamys (4; western and central Australia, South Australia), Kippistia (1; K. suaedifolia; southern Australia), Laennecia (18; southwestern United States, Mexico, Central America, South America to Bolivia), Microgyne (1; M. trifurcata; Brazil, Uruguay, Argentina), Minuria (9; Australia), Peripleura (9; Australia; in Vittadinia?), Podocoma (8; Brazil, Argentina), Sommerfeltia (1; S. spinulosa; South America), Tetramolopium (38; New Guinea, eastern Queensland, the Cook Islands, the Hawaiian Islands), Vittadinia (20; New Guinea, Australia, Tasmania, New Caledonia, New Zealand), Arctogeron (1; A. gramineus; Central Asia, Mongolia, China), Aster (c 180; Europe, southeastern Africa, temperate Asia, one species in North America), Asterothamnus (7; Central Asia, Mongolia, China), Callistephus (1; C. chinensis; China), Galatella (c 40; Europe, temperate Asia), Kemulariella (6; Caucasus), Krylovia (4; Central Asia, China), Miyamayomena (5; East Asia), Psychrogeton (20; southwestern to Central Asia), Tripolium (1; T. pannonicum; temperate regions on the Northern Hemisphere), Acamptopappus (2; southwestern United States), Amphiachyris (2; central United States, Texas), Amphipappus (1; A. fremontii; southwestern United States), Bigelowia (2; southeastern United States), Chihuahuana (1; C. purpusii; northern Mexico), Chrysoma (1; C. pauciflosculosa; southeastern United States), Chrysothamnus (9; western North America), Columbiadoria (1; C. hallii; northwestern United States), Eastwoodia (1; E. elegans; southwestern United States), Euthamia (5; North America, northwestern Mexico), Gundlachia (6; Texas, Mexico, Central America to Venezuela), Gutierrezia (28; western North America, South America), Gymnosperma (1; G. glutinosum; southern United States, Mexico, Central America), Lorandersonia (7; North America), Medranoa (1; M. parrasana; Coahuila and Zacatecas in Mexico), Neonesomia (2; Texas, northwestern Mexico), Nestotus (2; northwestern North America), Oreochrysum (1; O. parryi; western North America), Petradoria (1; P. pumila; southwestern United States), Sericocarpus (5; North America), Solidago (c 100; Macaronesia, temperate regions on the Northern Hemisphere, South America), Stenotus (4; western North America), Thurovia (1; T. triflora; southeastern Texas), Toiyabea (1; T. alpina; southern Nevada), Xylovirgata (1; X. pseudobaccharis; Mexico), Pentachaeta (7; southwestern United States, northwestern Mexico), Rigiopappus (1; R. leptocladus; western United States), Tracyina (1; T. rostrata; California), Batopilasia (1; B. byei; Mexico), Boltonia (5; central and eastern North America), Chloracantha (1; C. spinosa; southern United States, Mexico, Central America), Benitoa (1; B. occidentalis; California), Corethrogyne (1; C. filaginifolia; western North America, northwestern Mexico), Grindelia (c 30; central and western North America, Mexico, western South America), Haplopappus (c 70; South America, with their largest diversity in Chile; monophyletic?), Hazardia (13; western North America), Herrickia (4; western United States), Isocoma (16; southwestern United States, Mexico), Lessingia (12; southwestern United States, northwestern Mexico), Machaeranthera (30; western North America, Mexico), Arida (9; western United States, northern Mexico), Dieteria (3; western North America, northwestern Mexico), Xanthisma (17; western North America, Mexico), Olivaea (2; Mexico), Oonopsis (4; central United States), Pyrrocoma (14; western North America), Rayjacksonia (3; North America), Stephanodoria (1; S. tomentella; Mexico), Triniteurybia (1; T. aberrans; northwestern United States), Xanthocephalum (6; southwestern United States, Mexico), Xylorhiza (10; western United States, Mexico), Almutaster (1; A. pauciflorus; western North America, Mexico), Ampelaster (1; A. carolinianus; eastern United States), Canadanthus (1; C. modestus; North America), Psilactis (6; southwestern United States, Mexico, South America), Symphyotrichum (c 90; East Asia, North America), Chaetopappa (11; southwestern United States, northern Mexico), Monoptilon (2; southwestern United States, northwestern Mexico), Astranthium (12; southern United States, Mexico), Dichaetophora (1; D. campestris; southern United States, northern Mexico), Geissolepis (1; G. suaedifolia; Mexico), Townsendia (26–27; western North America), Chrysopsis (11; southeastern United States to Mexico and Bahamas), Croptilon (3; southeastern United States), Heterotheca (25–30; southern North America, Mexico), Noticastrum (c 20; tropical South America), Osbertia (3; western North America, Mexico, Central America), Pityopsis (7; eastern United States to Mexico, Central America), Tomentaurum (1; T. niveum; Mexico), Aphanostephus (4; United States, Mexico), Apopyros (2; southern Brazil, Paraguay, Argentina), Conyza (25–40; temperate to tropical regions on both hemispheres; in Erigeron?), Erigeron (c 400; cosmopolitan except Australia, with their highest diversity in North to Central America; incl. Conyza?), Hysterionica (7; southern Brazil, Uruguay, Argentina), Leptostelma (5; South America). – Cosmopolitan, with their largest diversity in North America, Australia and Africa. Shrubs or herbs. Latex usually absent. Capitula radiate, disciform or discoid, with multiseriate usually imbricate involucral bracts. Ray corollas usually strap-shaped, disc corollas quadri- or quinquelobate, filiform to infundibuliform. Anther thecae ecalcarate and usually ecaudate. Stylar branches scabrous or plumose, pronate, with sweeping hairs on acute appendage. Pappus consisting of uni- to quadriseriate barbellate bristles, or with scales or awns, or absent. x = 9.

Anthemideae Cass. in J. Phys. Chim. Hist. Nat. Arts 88: 192. Mar 1819

c 104/1.550–1.600. Adenanthellum (1; A. osmitoides; Mpumalanga, northern KwaZulu-Natal, Swaziland), Athanasia (39; South Africa, one species in Namibia), Hymenolepis (8; Western and Eastern Cape), Cotula (50–55; Africa, southwestern and southeastern Australia, Tasmania, Mexico to South America), Leptinella (33; New Guinea, southwesternmost Western Australia, eastern New South Wales, Victoria, Tasmania, New Zealand, Macquarie Island, Campbell Island, Auckland Islands, Antipode Islands, Tierra del Fuego, Antarctic islands), Soliva (8; South America), Eriocephalus (32; southern Africa), Hilliardia (1; H. zuurbergensis; Eastern Cape, KwaZulu-Natal), Hippia (5; Western and Eastern Cape), Inezia (2; Northern Province, Mpumalanga, Swaziland), Inulanthera (10; Angola, Zimbabwe, southern Africa, Madagascar), Lasiospermum (4; Egypt, southern Africa), Lidbeckia (3; Western Cape), Osmitopsis (9; Western and Eastern Cape), Adenoglossa (1; A. decurrens; Northern Cape), Cymbopappus (3; Western and Eastern Cape, Mpumalanga), Foveolina (5; Namibia, Northern and Western Cape), Leucoptera (3; Northern and Western Cape), Marasmodes (4; Western Cape), Myxopappus (2; Namibia, Namaqualand in Northern Cape), Oncosiphon (8; Namibia, South Africa, Lesotho), Pentzia (23; Morocco, Algeria, southern Africa), Rennera (4; Namibia, Northern Cape, North-West, Botswana), Eumorphia (6; South Africa, Swaziland), Gymnopentzia (1; G. bifurcata; Saouth Africa, Lesotho), Phymaspermum (19; Zimbabwe, southern Africa), Schistostephium (12; tropical and southern Africa), Thaminophyllum (3; Western Cape), Ursinia (39; Ethiopia, southern Africa), Ajania (34; Central and East Asia), Ajaniopsis (1; A. penicilliformis; Tibet, western China), Arctanthemum (1; A. arcticum; arctic and subarctic regions), Brachanthemum (10; Central Asia to China), Chrysanthemum (37; Europe, temperate and Central Asia), Elachanthemum (2; China), Hulteniella (1; H. integrifolia; arctic regions), Phaeostigma (3; China), Tridactylina (1; T. kirilowii; eastern Siberia), Artemisia (c 400; temperate regions on the Northern Hemisphere, few species in southern Africa and western South America), Crossostephium (1; C. chinense; East Asia), Neopallasia (3; Central Asia to China), Picrothamnus (1; P. desertorum; North America), Artemisiella (1; A. stracheyi; Himalayas to China), Allardia (8; Cental and East Asia), Cancrinia (4; Central Asia to China), Cancriniella (1; C. krascheninnikovii; Central Asia), Richteria (3; southwestern to East Asia), Trichanthemis (9; Central Asia), Ugamia (1; U. angrenica; Central Asia), Handelia (1; H. trichophylla; Central Asia to China), Lepidolopsis (1; L. turkestanica; Iran, Afghanistan, Central Asia), Polychrysum (1; P. tadschikorum; Afghanistan, Central Asia), Pseudohandelia (1; P. umbellifera; Iran, Afghanistan, Central Asia, China), Sclerorhachis (4; Afghanistan), Hippolytia (19; Central Asia to northern China), Kaschgaria (2; Central Asia to western China), Lepidolopha (9; Central Asia), Leucanthemella (2; Central and southeastern Europe), Microcephala (5; Iran, Central Asia), Nipponanthemum (1; N. nipponicum; Japan), Opisthopappus (2; northeastern central China), Stilpnolepis (2; Mongolia, China), Tanacetopsis (21; Iran, Central Asia), Achillea (c 130; Europe, the Mediterranean, temperate Asia, North America), Anacyclus (12; the Mediterranean), Heliocauta (1; H. atlanticum; Morocco), Leucocyclus (1–2; L. formosus; Turkey), Anthemis (c 175; Europe, the Mediterranean and eastwards to Iran, East Africa), Cota (c 40; Europe, the Mediterranean), Gonospermum (5; the Canary Islands; in Tanacetum?), Nananthea (1; N. perpusilla; Corsica, Sardinia), Tanacetum (c 160; temperate regions on the Northern Hemisphere; incl. Gonospermum?), Tripleurospermum (c 40; temperate regions on the Northern Hemisphere), Brocchia (1; B. cinerea; northern Africa from Mauritania and Mali to Egypt, Sudan and Ethiopia), Castrilanthemum (1; C. debeauxii; southeastern Spain), Hymenostemma (1; H. pseudanthemis; Spain, Morocco), Leucanthemopsis (9; southern Europe, Morocco), Prolongoa (1; P. hispanica; Spain), Matricaria (6; Europe, temperate Asia, northwestern North America), Phalacrocarpum (2; Spain, Portugal), Xylanthemum (9; southern and Central Asia), Aaronsohnia (2; North Africa), Argyranthemum (24; Madeira, the Canary Islands), Glebionis (2; Europe, the Mediterranean, North Africa; incl. Ismelia?), Heteranthemis (1; H. viscidehirta; southwestern Europe, North Africa), Ismelia (1; I. versicolor; Morocco; in Glebionis?), Chamaemelum (2; Europe, the Canary Islands, the Mediterranean), Cladanthus (5; the Mediterranean), Mecomischus (2; Morocco, Algeria), Rhetinolepis (1; R. lonadioides; Algeria, Tunisia, Libya), Santolina (c 10; the Mediterranean), Daveaua (1; D. anthemoides; Portugal, Morocco), Endopappus (1; E. macrocarpus; North Africa), Heteromera (1; H. fuscata; North Africa), Lepidophorum (1; L. repandum; Spain, Portugal), Chlamydophora (1; C. tridentata; southern Europe, North Africa), Chrysanthoglossum (2; North Africa), Coleostephus (3; western Europe, North Africa), Glossopappus (1; G. macrotus; southwestern Europe, North Africa), Leucanthemum (c 30; Europe, northern Asia), Plagius (3; the Balearic Islands, Morocco, Tunisia), Rhodanthemum (12; Spain, Morocco, Algeria), Lonas (1; L. annua; western Mediterranean), Nivellea (1; N. nivellei; Morocco), Otospermum (1; O. glabrum; southwestern Europe, northwestern Africa). – Cosmopolitan, with their highest diversity in southern Africa, Central and East Asia and the Mediterranean. Shrubs or herbs. Leaves usually alternate. Involucral bracts bi- to septaseriate, imbricate, with scarious margins and apex. Ray corollas trilobate, disc corollas uni- to quinquelobate (or lobes absent) to tri- to sexalobate. Upper part of filaments with thickened cell walls, forming anther collar (filament collar). Anther thecae usually ecalcarate and ecaudate. Pollen grains without internal foramina. Stylar branches hairy, with parallel stigmatic surfaces. Pappus not capillary. x = 9.

[Inuleae+[Athroismeae+[Feddeeae+[Helenieae+[Coreopsideae+[[Neurolaeneae+[Tageteae+[Bahieae+Chaenactideae]]]+[[Polymnieae+[Heliantheae+[Millerieae+[Madieae+[Eupatorieae+Perityleae]]]]]]]]]]]

Inuleae Cass. in J. Phys. Chim. Hist. Nat. Arts 88: 193. Mar 1819

c 50/665–670. Duhaldea (14; East Africa, Iran to Himalayas), Caesulia (1; C. axillaris; Sri Lanka, northeastern India, Bangladesh, Nepal, Burma), Blumea (c 100; tropical and southern Africa, tropical Asia, northern Australia), Merrittia (1; M. benguetensis; the Philippines), Schizogyne (2; the Canary Islands), Vieria (1; V. laevigata; Tenerife in the Canary Islands), Rhanteriopsis (4; Lebanon, Syria, Iran, Iraq; in Inula?), Rhanterium (3; northwestern Africa and eastwards to Pakistan), Pulicaria (77; Europe, the Mediterranean, North Africa, temperate Asia), Jasonia (1; J. tuberosa; France, Spain, Morocco), Chiliadenus (10; the Mediterranean), Allagopappus (2; the Canary Islands), Iphiona (11; northeastern Africa, southwestern to Central Asia), Dittrichia (2; the Mediterranean), Limbarda (1; L. crithmoides; southern Europe, the Mediterranean), Anvillea (2; North Africa, southwestern Asia), Lifago (1; L. dielsii; Morocco, Algeria), Asteriscus (9; Macaronesia, the Mediterranean), Pallenis (3; the Mediterranean), Buphthalmum (3; Europe, western Asia), Inula (c 90; Europe, the Mediterranean, Africa, temperate Asia; incl. Carpesium?, Rhanteriopsis?), Carpesium (c 25; Europe, Asia, eastern Queensland, northeastern New South Wales; in Inula?), Stenachaenium (5; southern Brazil, Uruguay, Paraguay, Argentina), Antiphiona (2; tropical and southwestern Africa to Namibia), Calostephane (5; tropical and southern subtropical Africa, Madagascar), Pegolettia (9; Africa, the Arabian Peninsula and eastwards to India), Geigeria (20; tropical and southern Africa), Ondetia (1; O. linearis; Namibia), Sachsia (3; Florida, Bahamas, Cuba), Iphionopsis (3; northeastern and eastern Africa, Madagascar), Cratystylis (3; arid and semiarid regions in western and southern Australia), Pterocaulon (18; Southeast Asia, Malesia, Australia, New Caledonia, tropical and subtropical America), Cylindrocline (2; Mauritius), Epaltes (14; tropical regions on both hemispheres), Litogyne (1; L. gariepina; tropical and southern Africa), Pechuel-loeschea (1; P. leubnitziae; Namibia, South Africa, Swaziland, Botswana, Zimbabwe), Sphaeromorphaea (1; S. australis; Australia except western and southwestern parts), Adelostigma (2; tropical Africa), Pluchea (c 80; tropical and subtropical regions on both hemispheres), Delamerea (1; D. procumbens; northern Kenya), Neojeffreya (1; N. decurrens; tropical Africa, Madagascar), Triplocephalum (1; T. holstii; tropical East Africa), Sphaeranthus (41; Egypt, southern Asia from Iran to Java), ’Pseudoblepharispermum’ (2; P. bremeri: Ethiopia; P. mudugense: Somalia; non-monophyletic), Pseudoconyza (1; P. viscosa; tropical Africa, tropical Asia, Central America), Blumeopsis (1; B. falcata; India to West Malesia), Laggera (17; tropical and southern Africa, the Arabian Peninsula, tropical Asia), Nicolasia (7; southwestern tropical Africa), Doellia (2; tropical Africa, Madagascar, the Arabian Peninsula), Porphyrostemma (4; tropical Africa), Allopterigeron (1; A. filifolius; northern Australia), Nanothamnus? (1; N. sericeus; Bombay). – Temperate to tropical regions on both hemispheres, few species in America. Shrubs or herbs. Capitula radiate, disciform or discoid, with imbricate usually multiseriate involucral bracts. Ray corollas trilobate strap-shaped or absent, disc corollas quinquelobate. Anther thecae usually ecalcarate and caudate. Stylar branches with long acute or obtuse sweeping hairs. Cypsela often with glandular hairs and/or twin hairs; each epidermal cell usually with one elongated crystal. Phytomelan (phytomelanin) not found. Pappus consisting of capillary bristles, of bristles and/or scales, of awns, or absent. x = 10.

[Athroismeae+[Feddeeae+[Helenieae+[Coreopsideae+[[Neurolaeneae+[Tageteae+[Bahieae+Chaenactideae]]]+[[Polymnieae+[Heliantheae+[Millerieae+[Madieae+[Eupatorieae+Perityleae]]]]]]]]]]

Athroismeae Panero in J. L. Panero et V. A. Funk, Proc. Biol. Soc. Washington 115: 917. 20 Dec 2002

7/70–75. Anisopappus (32; tropical and southern Africa, Madagascar, one species, A. chinensis, in India, southern China, Burma and northern Thailand), Welwitschiella (1; W. neriifolia; Angola, Zambia)?, Athroisma (12; tropical regions in the Old World), Blepharispermum (15; Africa, the Arabian Peninsula and eastwards to Sri Lanka), Leucoblepharis (1; L. subsessilis; India), Centipeda (10; Madagascar, tropical Asia, Australia, islands in the Pacific, Chile); Symphyllocarpus (1; S. exilis; eastern Siberia, Manchuria). – Tropical and subtropical regions in the Old World, Chile. Shrubs or herbs. Capitula sometimes aggregated in pseudocephalia. Capitula radiate, disciform or discoid, with few rows of imbricate (often reduced) involucral bracts. Ray corollas tridentate strap-shaped (outer ones often tubular to filiform) or absent, disc corollas quinquelobate. Anther thecae usually ecalcarate and caudate or ecaudate. Stylar branches with short obtuse sweeping hairs. Cypsela sometimes with phytomelan (phytomelanin). Pappus consisting of scales or awns, or absent. x = 10.

[Feddeeae+[Helenieae+[Coreopsideae+[[Neurolaeneae+[Tageteae+[Bahieae+Chaenactideae]]]+[[Polymnieae+[Heliantheae+[Millerieae+[Madieae+[Eupatorieae+Perityleae]]]]]]]]]

Feddeeae Pruski, Herrera, Anderb. et Franc.-Ort. in Syst Bot. 33: 199. 26 Feb 2008

1/1. Feddea (1; F. cubensis; eastern Cuba). – Leaves leathery. Capitula discoid, with multiseriate involucral bracts with resiniferous duct. Corollas quinquelobate. Anther thecae ecalcarate and caudate. Style with inconspicuous hairs. Pappus consisting of uniseriate capillary bristles. n = ?

[Helenieae+[Coreopsideae+[[Neurolaeneae+[Tageteae+[Bahieae+Chaenactideae]]]+[[Polymnieae+[Heliantheae+[Millerieae+[Madieae+[Eupatorieae+Perityleae]]]]]]]]

x = 19.

Helenieae (Cass.) Lindl. in J. C. Loudon, Encycl. Plant.: 1074. 1829

13/110–115. Balduina (3; southeastern United States), Gaillardia (16–20; North America, temperate South America), Helenium (30–35; America), Marshallia (9; the United States), Pelucha (1; P. trifida; Mexico), Plateilema (1; P. palmeri; Texas, Mexico), Psathyrotes (3; southwestern United States, Mexico), Trichoptilium (1; T. incisum; southwestern United States, northwestern Mexico), Amblyolepis (1; A. setigera; Texas, Mexico), Baileya (3; southwestern United States, Mexico), Hymenoxys (25; western North America, Mexico and southwards to Argentina), Psilostrophe (7; southwestern United States, Mexico), Tetraneuris (9; North America, Mexico). – America, with their highest diversity in western North America and Mexico. Usually herbs. Capitula radiate or discoid, with bi- or multiseriate involucral bracts. Ray corollas trilobate strap-shaped or absent, disc corollas quinquelobate tubular. Anther thecae ecalcarate and caudate or ecaudate. Stylar branches with terminal hair tuft. Cypsela cell walls usually with few large crystals. Phytomelan not found. Pappus consisting of scales or bristles. x = 19.

[Coreopsideae+[[Neurolaeneae+[Tageteae+[Bahieae+Chaenactideae]]]+[[Polymnieae+[Heliantheae+[Millerieae+[Madieae+[Eupatorieae+Perityleae]]]]]]]

Coreopsideae (Cass.) Lindl. in J. C. Loudon, Encycl. Plant.: 1074. 1829

27/400–405. Chrysanthellum (13; Mexico, Central America, the West Indies, Galápagos Islands), Diodontium (1; D. filifolium; northern Australia), Glossocardia (12; Southeast Asia, Malesia and eastwards to islands in the Pacific), Isostigma (11; subtropical South America), Trioncinia (1; T. retroflexa; eastern Queensland), Bidens (c 200; cosmopolitan), Coreocarpus (9; southwestern United States, Mexico), Coreopsis (c 35; America), Cosmos (26; tropical and subtropical America, with their highest diversity in Mexico), Cyathomone (1; C. sodiroi; Ecuador), Dahlia (c 35; Mexico, Central America, Colombia), Dicranocarpus (1; D. parviflorus; southwestern United States, Mexico), Ericentrodea (6; the Andes), Fitchia (7; Polynesia), Goldmanella (1; G. sarmentosa; Central America), Henricksonia (1; H. mexicana; Mexico), Heterosperma (7; southwestern United States, Mexico and southwards to South America), Hidalgoa (6; Mexico, Central America), Leptosyne (8; southwestern United States, northwestern Mexico), Moonia (1; M. heterophylla; southern India, Sri Lanka), Narvalina (4; tropical America), Oparanthus (6; Rapa Island, Marquesas Islands), Petrobium (1; P. arboreum; St. Helena), Thelesperma (10–15; western North America, Mexico, southern South America), Koehneola (1; K. repens; eastern Cuba), Pinillosia (1; P. berteroi; Cuba, Hispaniola), Tetraperone (1; T. bellioides; Cuba), Staurochlamys (1; S. burchellii; northern Brazil). – Tropical to warm-temperate regions on both hemispheres, with their highest diversity in America. Shrubs or herbs. Leaves alternate or opposite. Capitula radiate or discoid, with usually bi- or multiseriate involucral bracts with resiniferous ducts (striations). Disc corollas quadri- or quinquelobate. Anther thecae ecalcarate and caudate. Style simple or shortly bilobate, papillate. Pappus consisting of two to 15 barbed bristles or short awns. Cypsela walls with phytomelan (phytomelanin; usually with spines or striations). x = 18.

[[Neurolaeneae+[Tageteae+[Bahieae+Chaenactideae]]]+[[Polymnieae+[Heliantheae+[Millerieae+[Madieae+[Eupatorieae+Perityleae]]]]]]

[Neurolaeneae+[Tageteae+[Bahieae+Chaenactideae]]]

Neurolaeneae Rydb. in N. Amer. Fl. 33: 46. 15 Sep 1922

5/c 140. Enydra (10; tropical and subtropical regions on both hemispheres), Heptanthus (7; Cuba), Calea (c 110; tropical and subtropical America), Greenmaniella (1; G. resinosa; Mexico), Neurolaena (13; tropical America), Tyleropappus (1; T. dichotomus; Venezuela). – Tropical and subtropical regions on both hemispheres, with their highest diversity in tropical America. Shrubs or herbs. Stem hollow. Leaves alternate or opposite. Capitula radiate or discoid, with uni- to octaseriate involucral bracts. Ray corollas very reduced to well developed and tubular or strap-shaped, usually trilobate (sometimes absent), disc corollas usually quinquelobate tubular. Anthers usually black, with ecalcarate and ecaudate thecae. Pollen grains with partial cavea. Stylar branches with terminal hair tuft. Cypsela walls with phytomelan (phytomelanin). Pappus absent or consisting of scales, awns or bristles. n = 11.

[Tageteae+[Bahieae+Chaenactideae]]

Tageteae Cass. in J. Phys. Chim. Hist. Nat. Arts 88: 162. Feb 1819 [‘Tagetineae’]

30/265–280. Clappia (1; C. suaedifolia; Texas, Mexico), Coulterella (1; C. capitata; Baja California), Flaveria (22; southern United States, Mexico, Central and South America, Australia), Haploesthes (3; western Mexico), Sartwellia (4; Mexico), Jaumea (2; western United States, northwestern Mexico, southern South America), Adenophyllum (c 10; southwestern United States, Mexico, Central America), Bajacalia (3; Baja California), Boeberoides (1; B. grandiflora; Mexico), Chrysactinia (6; Mexico), Dysodiopsis (1; D. tagetoides; southern central United States), Dyssodia (c 30; the United States, Mexico, Guatemala, with their highest diversity in Mexico), Gymnolaena (3; Mexico), Harnackia (1; H. bisecta; eastern Cuba), Hydropectis (3; Mexico), Lescaillea (1; L. equisetiformis; western Cuba), Leucactinia (1; L. bracteata; Mexico), Nicolletia (3; southwestern United States, northwestern Mexico), Pectis (85–90; tropical and subtropical America including the Galápagos Islands), Porophyllum (25–30; tropical and subtropical America), Schizotrichia (2–5; Peru), Strotheria (1; S. gypsophila; northern and central Mexico), Tagetes (40–50; tropical and subtropical America), Thymophylla (13; southwestern United States, Mexico), Urbinella (1; U. palmeri; Mexico), Varilla (2; Mexico), Arnicastrum (2; Mexico), Jamesianthus (1; J. alabamensis; Alabama), Oxypappus (2; Mexico), Pseudoclappia (2; southwestern United States, northern Mexico). –Tropical to warm-temperate America, with their largest diversity in Mexico. Shrubs or herbs. Leaves alternate or opposite. Capitula usually radiate, with uni- to quinqueseriate involucral bracts. Ray corollas bi- or trilobate strap-shaped, disc corollas quinque- or sexalobate of equal size or one or two lobes larger. Anther thecae ecalcarate and ecaudate. Stylar branches papillate. Cypsela walls with phytomelan (phytomelanin). Pappus consisting of scales and/or bristles. x = 19.

[Bahieae+Chaenactideae]

Bahieae B. G. Baldwin in B. G. Baldwin, B. L. Wessa et J. L. Panero, Syst. Bot. 27: 192. 4 Mar 2002

21/c 80. Achyropappus (1; A. anthemoides; Mexico), Amauriopsis (1; A. dissecta; western United States, northern Mexico), Apostates (1; A. rapae; Rapa Island), Bahia (c 10; southwestern United States, Mexico, Chile), Bartlettia (1; B. scaposa; Mexico), Chaetymenia (1; C. peduncularis; Mexico), Chamaechaenactis (1; C. scaposa; southwestern United States), Espejoa (1; E. mexicana; Mexico, Central America), Florestina (8; southern United States, Mexico, Guatemala), Holoschkuhria (1; H. tetramera; northern Peru), Hymenopappus (11; southern United States, Mexico), Hymenothrix (5; southwestern United States, Mexico), Hypericophyllum (7; tropical and southern Africa), Loxothysanus (3; eastern Mexico), Palafoxia (12; southern United States, Mexico), Peucephyllum (1; P. schottii; southwestern United States, northern Mexico), Picradeniopsis (2; western and central United States), Platyschkuhria (1; P. integrifolia; southwestern United States), Psathyrotopsis (3; southwestern United States, Mexico), Schkuhria (6; tropical and subtropical America), Thymopsis (2; the West Indies). – Tropical and southern Africa, Rapa Island, tropical to warm-temperate America, with their highest diversity in Mexico. Herbs. Leaves alternate or opposite. Capitula radiate or discoid, with uni- to quadriseriate imbricate involucral bracts often with hyaline margins. Ray corollas usually bi- or trilobate strap-shaped or absent, disc corollas quadri- or quinquelobate. Anther thecae ecalcarate and ecaudate. Stylar branches with short apical hairs. Cypsela walls with phytomelan (phytomelanin; often striate). Pappus consisting of scales with usually thickened base or midrib or of fasciculate or hooked bristles. x = 17.

Chaenactideae B. G. Baldwin in B. G. Baldwin, B. L. Wessa et J. L. Panero in Syst. Bot. 27: 192. 4 Mar 2002

3/20. Dimeresia (1; D. howellii; western United States), Chaenactis (18; western North America, Mexico), Orochaenactis (1; O. thysanocarpha; California). – Western North America, Mexico. Herbs (rarely somewhat woody). Leaves usually alternate. Capitula discoid, with uni- or biseriate involucral bracts. Peripheral corollas sometimes zygomorphic. Anther thecae ecalcarate and ecaudate. Style with short hairs. Cypsela walls with phytomelan (phytomelanin). Pappus consisting of setose to obovate scales without thickened base or midribs, sometimes connate at base into caducous unit. n = 9.

[Polymnieae+[Heliantheae+[Millerieae+[Madieae+[Eupatorieae+Perityleae]]]]]

Tectal spines solid.

Polymnieae (H. Rob.) Panero in Proc. Biol. Soc. Washington 115: 919. 20 Dec 2002

1/3. Polymnia (3; eastern North America). – Perennial herbs. Leaves opposite. Capitula radiate, with bi- or triseriate imbricate involucral bracts. Ray corollas trilobate (mid-lobe longer and wider), disc corollas quinquelobate tubular. Anther thecae ecalcarate and ecaudate. Stylar branches with short hairs. Cypsela walls with phytomelan (phytomelanin). Pappus coroniform or absent. n = 15.

[Heliantheae+[Millerieae+[Madieae+[Eupatorieae+Perityleae]]]]

Heliantheae Cass. in J. Phys. Chim. Hist. Nat. Arts 88: 189. Mar 1819

114/950–1.000. Ambrosia (c 40; Ameria; incl. Xanthium?), Xanthium (3; tropical and subtropical America; in Ambrosia?), Dicoria (2; southwestern United States, northwestern Mexico), Euphrosyne (5–6; Mexico), Hedosyne (1; H. ambrosiifolia; southwestern United States), Iva (10; North America, Mexico, the West Indies), Parthenice (1; P. mollis; Mexico), Parthenium (16; North America to northern South America, the West Indies), Chromolepis (1; C. heterophylla; Mexico), Dugesia (1; D. mexicana; Mexico), Baltimora (2; tropical America), Blainvillea (10; tropical and subtropical America), Calyptocarpus (2; Texas, Mexico, Guatemala), Clibadium (c 30; tropical America), Damnxanthodium (1; D. calvum; Mexico), Delilia (2; tropical America, the Galápagos Islands), Dimerostemma (12; tropical South America), Eclipta (7; Australia, tropical and subtropical America), Elaphandra (14; tropical America), Eleutheranthera (1; E. ruderalis; tropical America), Exomiocarpon (1; E. madagascariense; Madagascar), Fenixia (1; F. pauciflora; the Philippines), Hoffmanniella (1; H. silvatica; Central Africa, Cameroun), Idiopappus (1; I. saloyensis; Ecuador), Iogeton (1; I. nowickeanus; Panamá), Jefea (5; southern United States, Mexico, Guatemala), Kingianthus (2; Ecuador), Lantanopsis (3; Cuba, Hispaniola), Lasianthaea (12; southwestern United States, Mexico), Leptocarpha (1; L. rivularis; Chile), Lipochaeta (20; the Hawaiian Islands; in Melanthera?), Lundellianthus (8; Central America), Melanthera (c 35; tropical regions on both hemispheres; incl. Lipochaeta?), Monactis (9–10; tropical South America), Oblivia (2; tropical America), Otopappus (14; Mexico, Central America, the West Indies), Oyedaea (16; tropical America), Pascalia (3; South America), Pentalepis (6; northwestern Australia), Perymenium (c 40; Mexico, Central America, the Andes in Colombia to Peru), Plagiolophus (1; P. millspaughii; Yucatán in Mexico), Podanthus (2; Chile, Argentina), Rensonia (1; R. salvadorica; El Salvador), Riencourtia (5–6; northern South America), Schizopsera (1; S. peduncularis; Ecuador), Sphagneticola (4; tropical Asia, tropical America), Steiractinia (12; Colombia, Venezuela, Ecuador), Synedrella (1; S. nodiflora; tropical America), Synedrellopsis (1; S. grisebachii; Argentina), Tilesia (3; tropical America), Trigonopterum (1; T. laricifolium; the Galápagos Islands), Tuberculocarpus (1; T. ruber; Venezuela), Tuxtla (1; T. pittieri; Mexico), Wamalchitamia (5; Central America), Wedelia (c 110; tropical and subtropical Africa, Madagascar, tropical America), Zexmenia (2; Central America), Zyzyxia (1; Z. lundellii; Guatemala, Belize), Encelia (13–14; southwestern United States, Mexico, the Galápagos Islands, Peru, Chile), Enceliopsis (3; western North America), Flourensia (c 30; southern United States, Mexico, South America), Geraea (2; southwestern United States, northwestern Mexico), Helianthella (9; western North America), Agnorhiza (5; Sierra Nevada in California), Balsamorhiza (11–12; western North America), Berlandiera (5; southern United States, Mexico), Borrichia (3; southeastern United States, the West Indies), Chrysogonum (1; C. virginianum; southeastern United States), Engelmannia (1; E. peristenia; North America), Lindheimera (1; L. texana; southern United States, Mexico), Scabrethia (1; S. scabra; western North America), Silphium (12; eastern North America), Vigethia (1; V. mexicana; Mexico), Wyethia (8; western North America, northwestern Mexico), Aldama (2; Central America, tropical South America), Alvordia (4; California, Mexico), Bahiopsis (12; southwestern United States, northwestern Mexico), Calanticaria (4; southern United States, Mexico), Helianthus (c 50; North America, northern Mexico), Heliomeris (5; western North America, northern Mexico), Hymenostephium (c 25; Mexico, Central America, South America to Argentina), Iostephane (4; Mexico), Lagascea (8; Mexico, Central America), Pappobolus (38; Colombia, Ecuador, Peru, Bolivia), Phoebanthus (2; southeastern United States), Rhysolepis (3; Mexico), Scalesia (15; the Galápagos Islands), Sclerocarpus (8; tropical and southern Africa, Mexico, Central America), Simsia (20; southern United States, Mexico, tropical America), Stuessya (3; Mexico), Syncretocarpus (2–3; Peru), Tithonia (11; southwestern United States, Mexico, Central America southwards to Costa Rica), Viguiera (c 150; tropical and subtropical America), Montanoa (c 25; Mexico, Central America, northern South America), Rojasianthe (1; R. superba; Central America), Ratibida (7; North America, northern Mexico), Rudbeckia (23; North America), Acmella (c 30; tropical regions on both hemispheres), Oxycarpha (1; O. suaedifolia; Venezuela), Salmea (10; tropical America), Spilanthes (6; tropical regions on both hemispheres), Tetranthus (4; the West Indies), Podachaenium (2; Mexico, Costa Rica), Squamopappus (1; S. skutchii; southeastern Mexico, Guatemala), Tetrachyron (7–8; eastern Mexico, Guatemala), Verbesina (>200; tropical and subtropical America), Hybridella (2; Mexico), Zaluzania (10; southwestern United States, Mexico), Echinacea (4; eastern United States), Heliopsis (18; tropical America, with their highest diversity in Mexico), Philactis (4; Central America), Sanvitalia (5; southwestern United States, Mexico, Central America), Tehuana (1; T. calzadae; Mexico), Trichocoryne (1; T. connata; Mexico), Zinnia (c 17; United States, Mexico, Central and South America to Peru, with their highest diversity in Mexico). – Temperate to tropical America, few species in the Old World. Trees, shrubs, lianas or herbs. Leaves alternate or opposite. Capitula radiate or discoid, with uni- to septaseriate often foliaceous involucral bracts. Receptacular paleae enclosing cypsela and usually persistent. Ray corollas usually trilobate strap-shaped or absent, disc corollas quinquelobate. Anther thecae usually black, usually ecalcarate and ecaudate. Stylar branches often with tuft of hairs or papillae. Cypsela walls with phytomelan (phytomelanin; often with spines, sometimes with striation). Pappus consisting of awns or scales (rarely absent). x = 19.

[Millerieae+[Madieae+[Eupatorieae+Perityleae]]]

Millerieae (Cass.) Lindl. in J. C. Loudon, Encycl. Plant.: 1074. 1829

31/c 350. Desmanthodium (8; Central and South America), Bebbia (2; southwestern United States, northwestern Mexico), Dyscritothamnus (2; Mexico), Tetragonotheca (4; southeastern United States), Tridax (c 25; tropical to warm-temperate regions in America, with their highest diversity in Mexico), Espeletia (c 88; the Andes in Colombia, Venezuela, Ecuador), Tamananthus (1; T. crinitus; Venezuela), Alepidocline (5; tropical America), Alloispermum (7; America), Aphanactis (8; tropical America), Cuchumatanea (1; C. steyermarkii; Guatemala), Faxonia (1; F. pusilla; Baja California), Galinsoga (13; Mexico, Central America, subtropical and temperate South America), Oteiza (3; Mexico, Guatemala), Sabazia (17; mountains in Central America), Schistocarpha (10; tropical America), Selloa (3; Mexico, Central America), Guardiola (10; Mexico), Jaegeria (6; Mexico, Central America, South America to Uruguay, the Galápagos Islands), Acanthospermum (8; Central and tropical South America), Lecocarpus (3; the Galápagos Islands), Melampodium (c 40; tropical and subtropical America, with their highest diversity in Mexico), Axiniphyllum (5; Mexico), Guizotia (6; tropical and subtropical Africa), Ichthyothere (18; tropical South America), Micractis (3; tropical Africa, Madagascar), Milleria (2; Mexico, Central America to Peru), Rumfordia (12; Mexico, Central America), Sigesbeckia (c 12; tropical Africa and Asia), Smallanthus (c 20; tropical and subtropical America), Stachycephalum (2; S. mexicanum: Mexico; S. argentinum: Argentina), Trigonospermum (5; Mexico). – Tropical and subtropical Africa, Madagascar, tropical Asia, tropical to warm-temperate America, with their highest diversity in Mexico, Central America and northern South America. Trees, shrubs or herbs. Leaves usually opposite, often glandular. Capitula usually radiate, with uni- to quinqueseriate involucral bracts. Ray corollas usually trilobate strap-shaped, disc corollas quadri- or quinquelobate. Anther thecae usually black, ecalcarate and ecaudate. Stylar branches with short hairs. Cypsela walls with phytomelan (phytomelanin; with spines and striation). Pappus consisting of scales or bristles, or absent. x = 19.

[Madieae+[Eupatorieae+Perityleae]]

Madieae Jeps., Fl. W. Middle Calif.: 483, 486. 1901

34/100–105. Arnica (c 30; temperate and arctic-alpine regions on the Northern Hemisphere), Amblyopappus (1; A. pusillus; California, northwestern Mexico, Peru, Chile), Baeriopsis (1; B. guadalupensis; Baja California), Constancea (1; C. nevinii; California), Eriophyllum (13; western North America, Mexico), Lasthenia (18; western North America, Mexico, Chile), Monolopia (5; California), Pseudobahia (3; California), Syntrichopappus (2; southwestern United States), Eatonella (1; E. nivea; western United States), Hulsea (7; western United States), Achyrachaena (1; A. mollis; western North America), Adenothamnus (1; A. validus; Baja California), Anisocarpus (2; western North America, northwestern Mexico), Blepharipappus (1; B. scaber; western United States), Blepharizonia (2; California), Calycadenia (10; western United States), Carlquistia (1; C. muirii; California), Centromadia (4; California, northwestern Mexico), Deinandra (21; southwestern United States, northwestern Mexico), Dubautia (31; the Hawaiian Islands), Harmonia (5; California), Hemizonella (1; H. minima; western United States), Hemizonia (1; H. congesta; southwestern United States, northwestern Mexico), Holocarpha (4; California), Holozonia (1; H. filipes; western United States), Jensia (2; California), Kyhosia (1; K. bolanderi; Oregon, California), Lagophylla (4; western North America), Layia (14; western North America, northwestern Mexico), Madia (10; western North America, Mexico, Chile), Osmadenia (1; O. tenella; California, northwestern Mexico), Raillardella (3; western United States), Venegasia (1; V. carpesioides; California, northwestern Mexico). – Temperate and arctic-alpine regions on the Northern Hemisphere, the Hawaiian Islands, Mexico, Peru, Chile, with their highest diversity in western United States and the Hawaiian Islands. Trees, shrubs, lianas or herbs. Leaves alternate, opposite or verticillate, often glandular. Capitula radiate or discoid, with usually uni- or biseriate subequal involucral bracts. Ray corollas usually (deeply) trilobate strap-shaped, disc corollas quinquelobate. Anther thecae ecalcarate and ecaudate. Style simple to deeply bilobate, branches with short hairs. Cypsela walls with phytomelan (phytomelanin; with spines or striation). Pappus consisting of scales or bristles, or absent. x = 19.

[Eupatorieae+Perityleae]

Eupatorieae Cass. in J. Phys. Chim. Hist. Nat. Arts 88: 202. Mar 1819

166/2.250–2.320. Hofmeisteria (7; Mexico), Ageratina (c 250; eastern United States, Central and western South America), Oxylobus (5; Mexico, Central America, Colombia to Venezuela), Jaliscoa (3; Mexico), Spaniopappus (5; Cuba), Standleyanthus (1; S. triptychus; Costa Rica), Kaunia (14; South America), Jaramilloa (2; northern Colombia), Mikania (c 430; tropical regions on both hemispheres), Trichocoronis (3; southwestern United States, northern Mexico), Sclerolepis (1; S. uniflora; easterna United States), Adenostemma (c 25; tropical Africa, Central and tropical South America, the West Indies), Sciadocephala (5; Panamá, northern South America), Gymnocoronis (2–3; Central and South America), Fleischmannia (c 80; North America, western South America), Sartorina (1; S. schultzii; Mexico?), Acritopappus (16; eastern Brazil), Radlkoferotoma (3; southern Brazil, Uruguay), Ageratum (c 40; Central and South America, the West Indies), Phania (2; Cuba), Phalacraea (4; Colombia, Ecuador, Peru), Blakeanthus (1; B. cordatus; Guatemala, Honduras), Scherya (1; S. bahiensis; eastern Brazil), Ascidiogyne (2; Peru), Cavalcantia (2; Brazil), Teixeiranthus (2; Brazil), Gardnerina (1; G. angustata; Brazil), Ellenbergia (1; E. glandulata; Peru), Guevaria (4; Ecuador, Peru), Ferreyrella (2; Peru), Nesomia (1; N. chiapensis; Mexico), Piqueriella (1; P. brasiliensis; Btrazil), Piqueria (5–6; tropical regions on both hemispheres), Stevia (175–240 [agamospermy]; tropical and subtropical America), Carphochaete (7–8; southwestern United States, Mexico), Macvaughiella (2; Central America), Microspermum (9–10; Central America), Eupatorium (41; Europe, temperate Asia, eastern North America), Austroeupatorium (13; tropical South America to Uruguay), Eutrochium (5; North America), Stomatanthes (15; tropical and southern Africa, Brazil, Uruguay), Criscianthus (1; C. zambiensis; southern tropical Africa), Hatschbachiella (2; southern Brazil), Garberia (1; G. heterophylla; southeastern United States), Liatris (37; North America, Mexico, Bahamas), Carphephorus (7; southeastern United States), Hartwrightia (1; H. floridana; Georgia, Florida), Praxelis (13-14; tropical America), Chromolaena (c 165; tropical and subtropical America), Eupatoriopsis (1; E. hoffmanniana; Brazil), Lomatozona (4; Brazil), Praxeliopsis (1; P. mattogrossensis; Brazil), Eitenia (2; Brazil), Osmiopsis (1; O. plumieri; Haiti), Gyptis (7; tropical South America), Gyptidium (2; Brazil, Argentina), Urolepis (1; U. hecatantha; South America), Barrosoa (11; tropical South America), Dasycondylus (8; Brazil), Diacranthera (2; Brazil), Conocliniopsis (1; C. prasiifolia; Colombia, Venezuela, Brazil), Bejaranoa (2; tropical America), Prolobus (1; P. nitidulus; Bahia in Brazil), Trichogonia (c 30; Brazil, Bolivia, Paraguay), Trichogoniopsis (2; Brazil), Platypodanthera (1; P. melissifolia; eastern Brazil), Neocuatrecasia (12; Peru, Bolivia), Vittetia (2; Brazil), Campuloclinium (14; Brazil, Paraguay, Argentina), Macropodina (3; southern Brazil, Argentina), Conoclinium (4; eastern United States, Mexico), Tamaulipa (1; T. azurea; Texas, northern Mexico), Lourteigia (9; Colombia, Venezuela), Agrianthus (6; Brazil), Catolesia (1; C. mentiens; Bahia in Brazil), Arrojadocharis (2; eastern Brazil), Bahianthus (1; B. viscosus; northeastern Brazil), Semiria (1; S. viscosa; Bahia in Brazil), Lasiolaena (6; eastern Brazil), Stylotrichium (4; eastern Brazil), Bishopiella (1; B. elegans; eastern Brazil), Litothamnus (2; Brazil), Morithamnus (2; eastern Brazil), Acanthostyles (2; eastern South America), Raulinoreitzia (3; eastern South America), Disynaphia (16; South America), Campovassouria (1; C. bupleurifolia; Brazil), Grazielia (10; South America), Symphyopappus (12; Brazil), Ayapana (14; tropical America), Ayapanopsis (15; the Andes), Polyanthina (1; P. nemorosa; Costa Rica, the Andes from Colombia to Bolivia), Gongrostylus (2; Central America), Heterocondylus (13; Central and South America), Condylidium (2; Central America, the West Indies, Colombia and Venezuela to Bolivia), Gymnocondylus (1; G. galeopsifolius; Brazil), Alomiella (2; Brazil), Siapaea (1; S. liesneri; Venezuela), Monogereion (1; M. carajensis; northeastern Brazil), Parapiqueria (1; P. cavalcantei; Brazil), Lepidesmia (1; L. squarrosa; Cuba, Colombia, Venezuela), Isocarpha (5; tropical and subtropical America), Brickellia (c 100; western United States, Mexico, Central America, South America to Argentina), Brickelliastrum (2; southwestern United States, northern Mexico), Flyriella (4; Texas, northern Mexico), Ageratella (1; A. microphylla; Mexico), Asanthus (3; southwestern United States, Mexico), Malperia (1; M. tenuis; California, northwestern Mexico), Pleurocoronis (3; western United States, northwestern Mexico), Alomia (5; Mexico), Kyrsteniopsis (4; Mexico), Steviopsis (10; southwestern United States, Mexico), Carminatia (3; southwestern United States, Mexico, Central America to El Salvador), Dissothrix (1; D. imbricata; Brazil), Austrobrickellia (3; northern South America), Pseudobrickellia (2; Brazil), Goyazianthus (1; G. tetrastichus; Brazil), Leptoclinium (1; L. trichotomum; Brazil), Planaltoa (2; Brazil), Crossothamnus (2; Peru), Helogyne (8; the Andes), Condylopodium (4; Colombia), Critonia (43; tropical America), Critoniadelphus (2; Central America), Urbananthus (2; Cuba, Jamaica), Adenocritonia (1; A. adamsii; Jamaica), Antillia (1; A. brachychaeta; Cuba), Ciceronia (1; C. chaptalioides; eastern Cuba), Eupatorina (1; E. sophiifolia; Hispaniola), Fleischmanniopsis (5; Mexico, Central America), Viereckia (1; V. tamauilpasensis; Mexico)?, Koanophyllon (115–120; tropical America), Eupatoriastrum (5; Mexico, Central America), Sphaereupatorium (2; Brazil, Bolivia), Bishovia (2; Bolivia, Argentina), Nothobaccharis (1; N. candolleana; Peru), Santosia (1; S. talmonii; eastern Brazil), Grisebachianthus (8; eastern Cuba), Lorentzianthus (1; L. viscidus; Bolivia, Argentina), Chacoa (1; C. pseudoprasiifolia; Paraguay, Argentina), Idiothamnus (4; South America), Mexianthus (1; M. mexicanus; Mexico), Peteravenia (5; Mexico, Central America), Critoniella (6; northern Andes), Aristeguietia (21; the Andes), Asplundianthus (17; northern Andes), Austrocritonia (4; Brazil), Badilloa (10; northern Andes), Grosvenoria (4; the Andes), Corethamnium (1; C. chocoense; Colombia), Castenedia (1; C. santamartensis; Colombia), Imeria (2; Venezuela), Malmeanthus (3; Brazil, Uruguay, Argentina), Hughesia (1; H. reginae; Peru), Ophryosporus (37; South America), Cronquistianthus (29; northern Andes), Steyermarkina (4; Venezuela, Brazil), Neocabreria (5; tropical South America), Uleophytum (1; U. scandens; Peru), Amboroa (2; Peru, Bolivia), Tuberostylis (2; Panamá, Colombia), Hebeclinium (20; tropical America), Amolinia (1; A. heydeana; Mexico, Guatemala), Bartlettina (37; tropical America), Decachaeta (8; Mexico, Guatemala), Guayania (6; Guayana Highlands), Neomirandea (27; Mexico, Central America, northern South America to Ecuador). –America, few species in the Old World. Trees, shrubs or herbs. Leaves alternate or opposite. Capitula discoid, with various, persistent to caducous, involucral bracts. Corollas usually actinomorphic. Anthers often glandular. Anther appendages hollow or rudimentary. Anther thecae ecalcarate and ecaudate. Stylar branches usually almost glabrous, with clavate appendages. Cypsela walls with phytomelan (phytomelanin; smooth or with spines), usually with twin hairs. Pappus usually uniseriate, consisting of plumose bristles, scales, or absent. x = 17. Mono-ester pyrrolizidine alkaloids (secreted by nectaries), pentaynene acetylenes, monoterpenes, sesquiterpene lactones, ent-kaurine diterpene glycoside, kolavane derivatives, chromenes, and benzofurans present.

Perityleae B. G. Baldwin in B. G. Baldwin, B. L. Wessa et J. L. Panero, Syst. Bot. 27: 192. 4 Mar 2002

8/c 88. Galeana (3; Central America), Unxia (3; Panamá, northern South America), Villanova (10; Mexico and southwards to Chile), Lycapsus (1; L. tenuifolius; Desventuradas Island in Chile), Amauria (3; southwestern United States, Mexico), Eutetras (2; Mexico), Pericome (2; southwestern United States, Mexico), Perityle (c 65; southwestern United States, Mexico). – Western United States, Mexico, Central America and southwards to Chile, Desventuradas Island, with their highest diversity in western United States and Mexico. Shrubs or herbs. Leaves usually opposite, glandular. Capitula radiate or discoid, with uni- or biseriate subequal navicular involucral bracts. Ray corollas often trilobate, disc corollas quadrilobate (or quinquelobate). Anther thecae ecalcarate and ecaudate. Style deeply bilobate, branches with short hairs. Cypsela walls with phytomelan (phytomelanin). Pappus consisting of usually two bristles and crown of (rudimentary) scales, or absent. n = 19.

Phylogeny of the main clades of Asteraceae based on DNA sequence data (Funk & al. 2009).

Phylogeny of Asteroideae, Corymbium and Cichorioideae based on DNA sequence data showing the tribes (Funk & al. 2009).

CALYCERACEAE R. Br. ex Rich.

( Back to Campanulales )

Richard, Mém. Mus. Natl. Hist. Nat. Paris 6: 74. Nov 1820 [‘Calycereae’], nom. cons.

Boopidaceae Cass. in Bull. Sci. Soc. Philom. Paris 1816: 160. 12 Oct 1816 [’Boopideae’]; Boopidales Cass. ex Bercht. et J. Presl, Přir. Rostlin: 255. Jan-Apr 1820 [’Boopideae’]; Calycerales Link, Handbuch 1: 815. 4-11 Jul 1829 [‘Calycereae’]

Genera/species 5/47–65

Distribution Southeastern United States to South America, the Falkland Islands, with their highest diversity in South America.

Fossils Unknown.

Habit Usually bisexual (rarely monoecious? or gynomonoecious?, in Acicarpha often andromonoecious with male flowers in centre), usually perennial (rarely annual) herbs (occasionally somewhat woody at base).

Vegetative anatomy Phellogen ab initio superficial? Pericyclic fibres absent. Vascular bundles separate. Secondary lateral growth normal or absent. Vessel elements usually with simple (sometimes pseudoscalariform) perforation plates; lateral pits alternate, bordered pits. Imperforate tracheary xylem elements libriform fibres with simple pits, non-septate? (also vasicentric tracheids). Secondary wood rays absent. Axial parenchyma apotracheal diffuse, or paratracheal, vasicentric, or absent. Phloem rays in Acicarpha surrounded by sclerenchyma. Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Secretory cavities absent. Primary cortex and medulla with calciumoxalate crystals.

Trichomes Hairs uniseriate or absent.

Leaves Alternate (spiral), simple or pinnately compound, entire or pinnately lobed, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation pinnate. Stomata anomocytic. Cuticular wax crystalloids partly as large glabrous partially inrolled scales, partly as reticulate to annular threads or small scales. Secretory cavities absent. Leaf margin serrate or entire.

Inflorescence Terminal, probably cymose, head-like or capitulum-like pseudanthia (sometimes compound), surrounded by involucre consisting of one or two whorls of bracts; inflorescences principally developing centripetally or centrifugally.

Flowers Actinomorphic or zygomorphic. Epigyny. Sepals (four or) five (or six), small, thick, with open? aestivation, often aerenchymatous, often modified into spines, persistent, usually connate. Petals (four or) five (or six), with valvate or open aestivation, often persistent, connate into campanulate or tubular corolla, with outer layer separating and photosynthesizing. Nectariferous glands (four or) five (or six; sometimes absent), alternating with stamens. Disc, uniting corolla base with style, surrounding corolla tube and expanding into glandular areoles.

Androecium Stamens (four or) five (or six), haplostemonous, antesepalous, alternipetalous. Filaments at least at base connate into tube, adnate to corolla tube (epipetalous), in upper part usually with ring of thick-walled cells? Anthers connivent to connate at least at base, basifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits), sometimes with appendage. Tapetum secretory, with multinucleate cells. Staminodia absent. Secondary pollen display present.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolporate, with intercolpar depressions, shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum (with furcate columellae), imperforate, spinulate to granulate or almost smooth.

Gynoecium Pistil composed of two? (or five?) connate carpels. Ovary inferior, unilocular (pseudomonomerous by reduction of one locule or perhaps incompletely quinquelocular due to reduction of septa). Style single, simple, narrow, with secondary pollen presentation. Stigma punctate to capitate, indistinctly papillate, Dry type; pollen grains deposited on stigmatic apex and “pumped up” through stylar elongation. Pistillodium absent.

Ovules Placentation apical. Ovule one per carpel, anatropous, pendulous, apotropous, unitegmic, tenuinucellar. Integument more than ten? cell layers thick; outer cell layers of integument containing chloroplasts. Megagametophyte monosporous, Polygonum type. Endosperm development cellular. Endosperm haustoria absent. Embryogenesis solanad.

Fruit An achene with persistent, often accrescent and sometimes lignified calyx (and often corolla). Fruit apex with persistent bases of corolla and style. Fruits of marginal flowers in Acicarpha usually connate.

Seeds Aril absent. Testa indistinct (undifferentiated). Perisperm not developed. Endosperm copious to sparse, oily. Embryo straight, well differentiated, chlorophyll? Cotyledons two. Germination phanerocotylar?

Cytology n = 8 (Acicarpha), 12, 13, 15, 17, 18, 20–22 – Polyploidy frequently occurring (probably dominating).

DNA Mitochondrial gene rpl2 absent (lost).

Phytochemistry Insufficiently known. Flavonols (kaempferol, quercetin), 6-methoxyflavonols, Route I iridoids, Group VI secoiridoids (e.g. secologanin) and caffeic acid present. Ellagic acids, tannins, proanthocyanidins and cyanogenic compounds not found. Carbohydrates stored as oligo- or polyfructosans (e.g. inulin) with isokestose bonds (starch absent).

Use Unknown.

Systematics Boopis (13–25; the Andes, southern Brazil, Argentina), Moschopsis (8; Chile, Patagonia in Argentina), Calycera (15–20; Chile, Argentina), Gamocarpha (6–7; Chile, Argentina), Acicarpha (5; southeastern United States to tropical South America).

Calyceraceae are in most analyses recovered as sister-group to Asteraceae.

Tank & Donoghue (2010) found the topology [Boopis+[Moschopsis+Acicarpha]].

CAMPANULACEAE Juss.

( Back to Campanulales )

de Jussieu, Gen. Plant.: 163. 4 Aug 1789, nom. cons.

Lobeliaceae Juss. ex A. J. A. Bonpland, Descr. Pl. Malmaison: 19, ad t. 7. Sep 1813, nom. cons.; Jasionaceae Dumort., Anal. Fam. Plant.: 28, 30. 1829 [’Jasionideae’]; Lobeliales Link, Handbuch 1: 636. 4-11 Jul 1829 [‘Lobeliaceae’]; Campanulopsida Bartl., Ord. Nat. Plant.: 120, 146. Sep 1830; Cyphiaceae A. DC. in A. P. de Candolle, Prodr. 7(2): 497. late Dec 1839; Nemacladaceae Nutt. in Trans. Amer. Philos. Soc., ser. 2, 8: 254. 15 Dec 1842; Dortmannaceae Rupr., Fl. Ingr.: 649. Mai 1856 [’Dortmanniaceae’]; Cyananthaceae J. Agardh, Theoria Syst. Plant.: 384. Apr-Sep 1858 [’Cyanantheae’]; Cyphocarpaceae (Miers) Reveal et Hoogl. in Phytologia 79: 74. 29 Apr 1996

Genera/species 65–75/c 2.200

Distribution Cosmopolitan except extreme polar regions; few representatives in Malesia, Australia and New Zealand.

Fossils Pollen grains of Campanulaceae are known from the Oligocene onwards. Fossil seeds often assigned to Campanulaceae as Campanula palaeopyramidalis have been found in Miocene layers in Poland.

Habit Usually bisexual (rarely dioecious or gynodioecious), usually perennial (sometimes annual or biennial) herbs (sometimes evergreen shrubs, small trees or lianas). Some species are aquatic. Others are xerophytic.

Vegetative anatomy Phellogen ab initio superficial or inner cortical. Medullary vascular bundles often present (cortical bundles rare). Endodermis often prominent. Vascular cylinder present. Vessel elements usually with simple (sometimes scalariform) perforation plates; lateral pits scalariform to alternate, simple or bordered pits? Imperforate tracheary xylem elements libriform fibres with simple or bordered pits, septate or non-septate? Wood rays uniseriate or multiseriate, heterocellular (Lobelioideae), usually four to eight cells wide (Campanuloideae). Axial parenchyma paratracheal scanty vasicentric (Lobelioideae), or indistinct to absent (Campanuloideae). Sieve tube plastids S type. Nodes usually 1:1, unilacunar with one leaf trace (rarely tri- or pentalacunar with three or five? traces). Phloem with articulated anastomosing laticifers containing white, orange, yellow, dark or colourless (watery) latex. Cystoliths sometimes present. Calciumoxalate crystals acicular.

Trichomes Hairs unicellular or multicellular, usually uniseriate, sometimes stellate (rarely prickles or dendritic hairs), often calcified/silicified; glandular hairs absent.

Leaves Usually alternate (spiral; rarely opposite or verticillate), usually simple (rarely pinnately compound), usually entire (rarely pinnately lobed), with various ptyxis (in Lobelia supervolute). Stipules absent; leaf base rarely sheathing. Petiole vascular bundle transection incurved-arcuate. Venation pinnate, usually craspedodromous or camptodromous (rarely parallelodromous). Stomata usually anomocytic. Cuticular wax crystalloids partly as large glabrous more or less inrolled scales, partly as reticulate to annular threads or small scales. Hydathodes frequent. Leaf margin serrate, crenate or entire, often with hydathodes.

Inflorescence Usually terminal (sometimes axillary) panicle, raceme-, spike-, umbel- or headlike cymose (sometimes pseudanthium with involucre of bracts) particularly in Campanuloideae, or racemose especially in Lobelioideae (flowers sometimes solitary axillary, rarely solitary terminal; in Ruthiella epiphyllous).

Flowers Actinomorphic or zygomorphic (in Lobelioideae usually resupinate). Usually epigyny (rarely half epigyny; in Cyananthus hypogyny). Sepals (three to) five (to ten), with imbricate or valvate aestivation, persistent in fruit, usually connate; median sepal abaxial. Petals (four or) five (to ten), usually with valvate (in Cyphocarpus induplicate-valvate) aestivation, connate into usually campanulate, infundibuliform or tubular corolla (in Nemacladoideae bilabiate, with upper lip trilobate and lower lip bilobate; in Cyphocarpus bilabiate, with upper lip entire and lower lip quadrilobate; in Lobelioideae bilabiate or unilabiate, usually with upper lip bilobate and lower lip trilobate [in resupinated flower]; in Cyphia either bilabiate, with upper lip consisting of three and lower lip of two free petals, or five petals connate into tubular corolla); corolla tube formation early. Nectariferous disc intrastaminal, annular (in Adenophora enlarged, gland-like).

Androecium Stamens (three to) five (to ten), haplostemonous, antesepalous, alternipetalous. Filaments free or more or less connate, free from tepals or adnate to base of corolla tube (epipetalous); filament base often widened and enclosing nectariferous disc (filaments in Nemacladus and Parishella with special multicellular appendages). Anthers free, often connivent, or connate into tube surrounding style and stigma, usually basifixed (in Berenice dorsifixed), non-versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate cells. Staminodia absent. Secondary pollen display via combination of, i.a., protandry, anther tube surrounding style and stigma, anthers dehiscing prior to stylar elongation, and style with pollen-collecting hairs.

Pollen grains Microsporogenesis simultaneous. Pollen grains (3–)5–10-colpate, (2–)3–6-colporate or 3–8-zonoporate (rarely 12–20-pantoporate), usually shed as monads (in Namacodon as tetrads), usually tricellular (sometimes bicellular) at dispersal. Exine tectate or semitectate, with columellate infratectum, imperforate or perforate, spinulate (Cyphioideae), echinate to verrucate (Campanuloideae, Nemacladoideae), reticulate-striate (Lobelioideae) or almost psilate.

Gynoecium Pistil composed of two, three or five connate carpels. Ovary usually inferior (rarely semi-inferior or superior), usually bilocular to quinquelocular (sometimes unilocular; rarely sexalocular to decemlocular by ingrowth of ovary wall). Style single, simple, usually with hairs on which pollen grains are deposited (secondary pollen display); stylar hairs capable of invaginating and hence dislodging pollen grains from anthers. Stigma usually bilobate, trilobate or quinquelobate, papillate or non-papillate, Dry or Wet type (in Cyphocarpus with abaxial pollen-collecting hairs; in Cyphia with liquid-filled porous cavity). Pistillodium absent.

Ovules Placentation usually axile (rarely apical, basal or parietal [when ovary unilocular]; in Cyphocarpus free central). Ovules usually numerous per carpel, anatropous, horizontal, unitegmic, tenuinucellar. Integument approx. six to eight cell layers thick (sometimes vascularized). Hypostase absent. Megagametophyte monosporous, Polygonum type. Synergids sometimes with a filiform apparatus. Endosperm development cellular. Endosperm haustoria chalazal and micropylar. Embryogenesis solanad. Polyembryony occurring.

Fruit Usually a loculicidal capsule (sometimes septicidal or poricidal capsule or irregularly dehiscing; in, i.a., Burmeistera, Canarina, Clermontia a berry; in, e.g., Craterocapsa, Lysipomia, Parishella pyxidium; in Theodoravia schizocarp).

Seeds Aril absent. Testa one to four cell layers thick, sometimes winged. Operculum formed by endosperm haustorium. Exotesta consisting of cuboidal or fibriform cells with lignified walls. Only outer epidermis persistent. Endotestal cell walls (especially inner ones) thickened. Perisperm not developed. Endosperm copious, usually oily (rarely starchy). Embryo small, straight, without chlorophyll. Cotyledons two. Germination phanerocotylar.

Cytology n = 9 (Nemacladoideae, Cyphioideae); n = 6–21, 23–30, 32, 34–36, 40, 42, 45, 48, 51, 52 (17 most frequent) (Campanuloideae); n = 6–14, 19, 21, 35, 70 (Lobelioideae) – Polyploidy and aneuploidy frequently occurring.

DNA Plastid genome extensively rearranged, with numerous inversions and losses. Plastid inverted repeat expanded into small single copy region and sometimes contracted at end of large single copy region. Deletion of 5 bp in plastid gene ndhF. Plastid gene infA lost/defunct (Campanula with pseudogene, Platycodon, Prismatocarpus, Trachelium, Lobelia). Plastid gene accD, ORF244 and ORF2280 lost in numerous species of Campanuloideae and Lobelioideae. Mitochondrial gene rpl2 lost. Mitochondrial intron coxII.i3 lost.

Phytochemistry Flavonols (quercetin, kaempferol), afzelechin, flavonoid sulphates, biflavonoids, p-coumaric acid (absent in Lobelioideae), cyanidin, dammaranes, cantleyoside, oleanolic acid derivatives, ursolic acid, sesquiterpene lactones, caffeic acid (absent in Lobelioideae) or chelidonic acid (present in Lobelioideae), chlorogenic acid, toxic pyrrolizidine and pyridine alkaloids (lobelin etc., in latex of Lobelioideae), cyanogenic glycosides (linamarin, lotaustralin, proacacipetalin, prunasin, sambunigrin, triglochinin, zierin), tyrosine derived cyanogenic compounds, saponins (platycodin, rare, in, e.g., Platycodon), fatty acid derived 14-C-polyacetylenes, aliphatic tetrahydropyrane derivatives, amides, asarone, germacrane-like compounds, myo-inisitol, chiro-inositols (pinitol, quebrachitol), lignans (pinoresinol), arbutin, sinapic acid, ferulic acid, anthraquinones, acetophenones, and polysterols (absent from Lobelioideae) present. Myricetin, iridoids, ellagic acid, tannins, and proanthocyanidins not found. Carbohydrates stored as oligo- or polyfructosanes (e.g. inulin) with isokestose bonds (starch usually absent, although also present in Cyphia).

Use Ornamental plants, medicinal plants (Lobelia inflata, Codonopsis pilosula, etc.), vegetables (Campanula rapunculus, C. rapunculoides, Centropogon, Platycodon grandiflorus).

Systematics Campanulaceae may be sister-group to Rousseaceae.

A topology of Campanulaceae may be the following: [[Nemacladoideae+Campanuloideae]+[Cyphioideae+Lobelioideae+Cyphocarpus]].

The sister-group relationship of Cyphocarpus (Cyphocarpoideae) is unresolved and this small lineage should perhaps be included in Lobelioideae (cf. i.a. Haberle 1998 and Ayers & Haberle 1999).

[Nemacladoideae+Campanuloideae]

Flowers actinomorphic. Pollen grains spheroidal to oblate-spheroidal, verrucate or beset with spiculae. Style with retractile hairs in long upper part (Nemacladoideae?). Cell nucleus with fibrillar protein bodies.

Nemacladoideae (Nutt.) M. H. G. Gust., Phylogen. Stud. Asterales sensu lato: 34. 1996

2/15. Nemacladus (13; southwestern United States, northwestern Mexico), Pseudonemacladus (1; P. oppositifolius; Mexico). – Southwestern United States, Mexico. Usually annual (sometimes perennial) herbs. Flowers usually zygomorphic, often resupinate. Hypogyny or half epigyny. Corolla with three adaxial and two abaxial lobes. Filaments connate into tube, sometimes adnate to corolla lobes (epipetalous); two dorsal filaments sometimes with groups of small reflexed structures (pseudonectaries) at base; abaxial fimbriate scales sometimes present at insertion point of filaments on corolla lobes. Anthers free, flipping back after pollen release. Pollen grains tricolporate or hexacolpate. Pistil composed of two (or three) connate carpels. Stylar base with prominent glands; pollen display by stylar head bending towards or away from median sepal. Fruit in Parishella a pyxidium. n = 9

Campanuloideae Burnett, Outlines Bot.: 942, 1094, 1110. Feb 1835 [‘Campanulidae’]

40–45/c 900. Mainly northern temperate regions of the Old World (very few in Australia and New Zealand). Usually perennial (sometimes annual) herbs. Roots often thick. Vessel elements sometimes with scalariform perforation plates. Polysterol-rich latex present in laticifers in stem and leaves. Inflorescence usually cymose. Flowers actinomorphic at anthesis. Median calyx lobe adaxial. Petals in, e.g., Phyteuma coherent yet open laterally. Staminal bases concealing nectar. Anthers in Ostrowskia with placentoid. Pollen grains also colpate or porate. Tectum echinate. Pistil composed of (two or) three to five (to ten) connate usually antesepalous (rarely antepetalous) carpels (rarely one carpel) or median carpel adaxial. Ovary in some species of Wahlenbergia almost superior. Style (at least upper part) covered by long retractile pollen-collecting and pollen-presenting hairs with bulbous base. Integument in Ostrowskia massive. Fruit with lateral dehiscence, with pores or slits. Epicotyl and hypocotyl usually not prolonged. n = 6–21, 23–30, 32, 34–36, 40, 42, 45, 48, 51, 52; x = 6 or more (usually 17). Mesophyll cell nuclei sometimes with unique type of fibrillar protein inclusions. Large rearrangements in plastid genome. Plastid DNA in Trachelium with seven to ten extensive insertions (alternatively four insertions and three pseudogenes). Intron absent from plastid gene trnI in at least Campanula garganica. Plastid gene accD, ORF244 and ORF2280 absent in numerous species. Caffeic acid, p-coumaric acid, polyacetylenes (14-C-aliphatic tetrahydropyran derivates) present.

Cyanantheae Meisn., Plant. Vasc. Gen.: Tab. Diagn. 273, Comm. 180. 5-11 Apr 1840

7/60–65. Cyananthus (23; Himalayas); Platycodon (1; P. grandiflorus; northeastern Asia), Cyclocodon (3; tropical Asia), Echinocodon (1; E. lobophyllus; China), Ostrowskia (1; O. magnifica; Turkestan), Canarina (3; the Canary Islands, tropical East Africa); Codonopsis (c 30; Central and East Asia, Southeast Asia, Malesia). – Leaves often opposite. Stamens sometimes three. Pollen grains colp(or)ate. Pistil composed of five carpels. n = 7–9 (17). Platycodin (saponin) present in, i.a., Platycodon.

Campanuleae Dumort., Fl. Belg.: 58. 1827 (under construction)

c 34/810–820. Wahlenbergieae Endl., Gen. Plant.: 514. Jun 1838. Siphocodon (2; Western Cape), Rhigiophyllum (1; R. squarrosum; Western Cape), Feeria (1; F. angustifolia; Morocco), ’Wahlenbergia’ (125–260; warm-temperate to tropical regions on both hemispheres, with their highest diversity on the Southern Hemisphere; polyphyletic), Theilera (2; Western and Eastern Cape; in Wahlenbergia?), Nesocodon (1; N. mauritianus; Mauritius), Heterochaenia (3; the Mascarene Islands), Berenice (1; B. arguta; Réunion), Microcodon (4; Northern and Western Cape), Craterocapsa (5; South Africa, Swaziland, Lesotho, Zimbabwe), Namacodon (1; N. schinzianum; central Namibia), Gunillaea (2; tropical Africa, Madagascar), ’Prismatocarpus’ (c 30; tropical and southern Africa, with their highest diversity in the Cape Provinces; paraphyletic; incl. Merciera?, Roella?), Roella (c 20; Western Cape, one species in Eastern Cape and KwaZulu-Natal; in Prismatocarpus?), Merciera (3–6; Western Cape; in Prismatocarpus?), Treichelia (1; T. longibracteata; the Cape Peninsula in Western Cape). – Campanuleae Dumort., Fl. Belg.: 58. 1827. Jasione (13–16; Europe, the Mediterranean, southwestern Asia), Musschia (2; Madeira), ’Campanula’ (300–420; temperate regions on the Northern Hemisphere, tropical mountains; non-monophyletic), Legousia (6–7; Europe, the Mediterranean; in Campanula?), Triodanis (10; the Mediterranean, China, North America; in Campanula?), Petromarula (1; P. pinnata; Crete; in Campanula?), Phyteuma (c 40; Europe, the Mediterranean, temperate Asia; in Campanula?), Physoplexis (1; P. comosa; the Alps; in Phyteuma/Campanula?), Trachelium (2–7; Europe, the Mediterranean; in Campanula?), Homocodon (2; southwestern China; in Campanula?), Favratia (1; F. zoysii; the Alps), Zeugandra (2; Iran), Peracarpa (1; P. carnosa; tropical Asia; in Campanula?), Heterocodon (1; H. rariflorum; western North America; in Campanula?), Githopsis (4; western North America; in Campanula?), Cryptocodon (1; C. monocephalus; Central Asia), Cylindrocarpa (1; C. sewerzowii; Central Asia), Sergia (2; Central Asia). – Subcosmopolitan. Leaves often spiral. Filaments adnate in lower part to petals (epipetalous). Pollen porate, triangular in polar view. Fruits valvicidal or poricidal, also laterally (due to activity of axicorn on drying; Shulkina & al. 2003). Hypocotyl and epicotyl usually not elongated. n = 6 or more. – The Western Cape clade [Siphocodon+Rhigiophyllum] is sister-group to the remaining Wahlenbergieae, according to Cupido & al. (2013), but Feeria was not included in their analyses. At least the majority of the other genera are nested inside ‘Wahlenbergia’. – Most genera in Campanuleae are nested inside Campanula (Borsch & al. 2009; Haberle & al. 2009; Zhou & al. 2012).

[Lobelioideae+Cyphioideae+Cyphocarpoideae]

Flowers zygomorphic. Pollen grains prolate. Stylar apex present at base of newly opened anthers; hairs restricted to stylar apex.

Lobelioideae Burnett, Outlines Bot.: 942, 1094, 1110. Feb 1835 [‘Lobelidae’] (under construction)

25–30/c 1.200. ‘Lobelia’ (>300; nearly cosmopolitan; paraphyletic), Grammatotheca (1–2; G. bergiana; Western and Eastern Cape, KwaZulu-Natal; in Lobelia?), Dialypetalum (5; Madagascar), Wimmerella (10; southern Africa), Dielsantha (1; D. galeopsoides; tropical Africa), Monopsis (13; tropical and southern Africa), Isotoma (14; Australia), Ruthiella (5; New Guinea), Diastatea (7; Mexico to Peru), Solenopsis (7; the Canary Islands, the Mediterranean), Downingia (13; western North America, one species also southwards to Chile), Porterella (1; P. carnosula; western North America), Legenere (2; California, Chile), Howellia (1; H. aquatilis; western North America), Lysipomia (c 30; the high Andes), Burmeistera (>100; tropical South America), ‘Centropogon’ (c 220; tropical America; non-monophyletic),‘Siphocampylus’ (c 220; tropical America; non-monophyletic), Hippobroma (1; H. longiflora; the West Indies; in Lobelia?), Heterotoma (1; H. lobelioides; Mexico, Central America), Sclerotheca (4; the Cook Islands, the Society Islands), Apetahia (4; the Society Islands, Marquesas, Rapa Island), Trematolobelia (4; the Hawaiian Islands), Brighamia (2; the Hawaiian Islands), Delissea (9; the Hawaiian Islands), ‘Cyanea’ (60; the Hawaiian Islands; non-monophyletic), Clermontia (22; the Hawaiian Islands; in Cyanea?). – Subcosmopolitan, although largely tropical and with their highest diversity in Central America and tropical South America. Annual or perennial herbs, small (sometimes pachycaul) trees. Leaves with supervolute ptyxis (Lobelia). Flowers split zygomorphic, with upper and lower lips separate down to petal base; usually with upper lip bilobate and lower lip trilobate, or upper lip absent and lower lip quinquelobate (sometimes with upper lip trilobate and lower lip bilobate). Flowers usually resupinate by torsion of pedicel (Lobelia and closely allied; flower appearing to have ’normal’ orientation due to resupination, with median corolla lobe abaxial). Filaments connate at least apically. Anthers usually connate into tube surrounding style (sometimes free). Pollen grains usually reticulate, striate. Pistil composed of two connate carpels. Extrafloral nectaries occasionally present on ovary. “Pollen pump” mechanism often present. Style pushing up from inside anther tube and exposing pollen grains; style with brush hairs and pollen in pollen box. Stigma Wet type. Synergids hooked. Antipodal cells usually persistent. Fruit rarely a berry or a pyxidium; or a capsule dehiscing laterally. n = 6–14, 19, 21, 35, 70; x = 6 or more (usually 7 and 14). Plastid DNA in Lobelia with three large deletions including parts of clpP. Plastid DNA with two inversions in at least 25 species of Lobelia and in Sclerotheca jayorum, five inversions in Lobelia erinus and L. fervens, three inversions in Lobelia cardinalis, L. holstii and Monopsis lutea. Plastid gene clpP absent (lost) in at least Lobelia holstii and Monopsis lutea. Plastid gene accD, ORF244 and ORF2280 absent in numerous species. Pyridine and pyrrolizidine alkaloids (in latex), and chelidonic acid present. Caffeic acid, p-coumaric acid and polysterols not found. – It seems that a large part of the above genera in Lobelioideae are nested in Lobelia (Antonelli 2008).

Cyphioideae Walp. in Ann. Bot. Syst. 2: 1037. 12-15 Mai 1852 [‘Cyphiaceae’]

1/c 65. Cyphia (c 65; Africa, the Cape Verde Islands, with their largest diversity in southern Africa). – Perennial herbs (often twining) with tuberous root. Flowers zygomorphic, sometimes almost actinomorphic. Corolla with upper lip bilobate and lower lip trilobate, or upper lip trilobate and lower lip bilobate, or upper lip absent and lower lip quinquelobate. Filaments connate at base. Anthers sometimes slightly connivent. Pollen grains psilate. Pistil composed of two carpels. Ovary semi-inferior. Style bending away from median sepal, not elongating after anther dehiscence; stylar hairs with bulbous base (bristle hairs absent); pollen grains deposited in “pollen box”. Stigma with fluid-filled terminal cavity with usually lateral (sometimes terminal) pore. Fruit a septicidal and loculical capsule (valves bifid). Endosperm also with starch. n = 9.

Cyphocarpoideae Miers in London J. Bot. 7: 61. 1848 [‘Cyphocarpaceae’]

1/3. Cyphocarpus (2; Chile). – Perennial or annual herbs. Leaves deeply lobed. Flowers zygomorphic. Epigyny. Corolla with induplicate-valvate aestivation, with upper (adaxial) lip unilobate, cucullate, with apical appendage, and lower lip quadrilobate. Filaments free from each other, adnate in lower part to corolla lobes (epipetalous). Pistil composed of two carpels. Ovary strongly elongated. Fruit laterally dehiscent. n = 9. Cell nuclei with fibrillar inclusions. – Cyphocarpus should possibly be included in Lobelioideae.

Bayesian majority rule consensus tree of Campanuloideae and Cyphia based on DNA sequence data (Haberle & al. 2009). Cyclocodon is sister-group to Canarina according to Borsch & al. (2009).

Bayesian inference tree (simplified) of Lobelioideae based on DNA sequence data (Antonelli 2008).

GOODENIACEAE R. Br.

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Brown, Prodr. Fl. Nov.-Holl.: 573. 27 Mar 1810 [’Goodenoviae’], nom. cons.

Goodeniales R. Br. ex Bercht. et J. Presl, Přir. Rostlin: 252. Jan-Apr 1820 [‘Goodenoviae’]; Brunoniaceae Dumort., Anal. Fam. Plant.: 19, 21. 1829, nom. cons.; Scaevolaceae Lindl., Intr. Nat. Syst. Bot.: 189. 27 Sep 1830 [‘Scaevoleae’]; Brunoniales Lindl., Nix. Plant.: 20. 17 Sep 1833; Scaevolales R. Br. in C. F. P. von Martius, Consp. Regn. Veg.: 32. Sep-Oct 1835 [‘Scaevoleae’]

Genera/species 7–15/400–440

Distribution Tropical and subtropical coastal areas along the Atlantic, the Indian and the Pacific Oceans, Australia, Tasmania, islands in the Pacific and the Indian Oceans, with their largest diversity in Western Australia and Tasmania.

Fossils Unknown.

Habit Bisexual perennial or annual herbs, suffrutices or shrubs (rarely climbing or arborescent). Many representatives are xerophytic.

Vegetative anatomy Phellogen ab initio subepidermal or cortical. Medullary vascular bundles often present. Secondary lateral growth normal or absent. Vessel elements usually with simple (sometimes scalariform) perforation plates, lateral pits usually alternate (sometimes scalariform or intermediate), usually bordered (in Scaevola sometimes simple) pits. Imperforate tracheary xylem elements tracheids and fibre tracheids with bordered pits, non-septate. Wood rays uniseriate or multiseriate, homocellular or heterocellular, or absent. Axial parenchyma apotracheal diffuse, and/or paratracheal scanty vasicentric, or absent. Sieve tube plastids S type. Nodes usually 1:1, unilacunar with one leaf trace (sometimes 3:3, trilacunar with three traces, or 5:5, pentalacunar with five traces). Laticifers absent. Heartwood with gum-like substances. Idioblasts with branched sclereids present especially in foliar mesophyll. Prismatic calciumoxalate crystals or druses present in parenchyma cells in many species.

Trichomes Hairs simple, dendritic or stellate, often verrucose/strigose, sometimes candelabra-shaped, or absent; in Brunonia with short thin-walled basal cell and bicellular or multicellular arm inserted in right angle at basal cell and pressed to surface; glandular hairs with multicellular stalk (sometimes peltate-lepidote).

Leaves Usually alternate (spiral; in Scaevola section Enantophyllum opposite), simple, entire, often coriaceous, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation pinnate. Stomata usually anomocytic (in Brunonia paracytic with four subsidiary cells, two of which parallel to guard cells). Cuticular wax crystalloids? Mesophyll often with sclerenchymatous idioblasts containing branched sclereids. Leaves often with axillary hair tuft. Leaf margin serrate or entire (rarely sinuate).

Inflorescence Terminal or axillary, racemose or thyrsoid, raceme-, spike- or umbel-like (in Brunonia spicate to capitular, terminal; flowers rarely solitary axillary). Inflorescences sometimes head-like pseudanthia with involucre of bracts.

Flowers Zygomorphic (in Brunonia almost actinomorphic). Usually epigyny or half epigyny (in Brunonia and some species of Goodenia hypogyny). Sepals usually small, usually five (rarely three, sometimes indistinct), with open to imbricate aestivation, often persistent, connate. Petals five, with (induplicate-)valvate aestivation, unilabiate with five lower lobes, or bilabiate with two upper and three lower lobes (rarely with spur), connate, with odd lobe ventral, and usually with wings along margins (except in Brunonia and Selliera) each corolla lobe appearing trilobate. Nectary usually absent; one or two intrastaminal nectariferous glands sometimes present. Disc present or absent.

Androecium Stamens five, haplostemonous, antesepalous, alternipetalous. Filaments free from each other, usually adnate at base to corolla tube (epipetalous; in some species of Goodenia free from tepals). Anthers basifixed, non-versatile?, tetrasporangiate, introrse, longicidal (dehiscing in bud by longitudinal slits), connivent or connate and forming tube around style. Tapetum secretory, with binucleate to multinucleate cells. Staminodia absent. Secondary pollen display present.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate (in Leschenaultia 4–8-por[or]ate), usually shed as monads (in Leschenaultia as rhomboidal tetrads), bicellular at dispersal. Exine tectate, with columellate infratectum (with furcate columellae), imperforate to finely perforate, rugulate (Dampiera, Leschenaultia) or microspinulate.

Gynoecium Pistil composed of usually four (in Brunonia two) connate carpels; usually only two carpels fertile. Ovary usually inferior or semi-inferior (in Brunonia and some species of Goodenia superior), bilocular at base, unilocular at centre, and bilocular above (sometimes entirely unilocular; in species of Scaevola quadrilocular; in Brunonia unilocular with septum at base only). Style single, usually simple, narrow, curved, with pollen-catching and often hairy indusium (enclosing pollen initially; indusium glabrous in, i.a., Brunonia) and collar- or cup-shaped outgrowth at apex immediately below stigma (style in some species of Goodenia bifid to quadrifid, each branch with indusium); style “pumping up” pollen grains during its elongation (pollen in Brunonia caught by stylar hairs). Stigma truncate or bilobate, papillate, Dry type. Pistillodium absent.

Ovules Placentation basal-axile (in Brunonia basal) or parietal (when ovules situated in centre of ovary). Ovules usually several or numerous per carpel (in Brunonia, and some species of Dampiera, Goodenia and Scaevola one per fertile carpel), usually anatropous (rarely campylotropous), usually ascending, unitegmic, tenuinucellar. Integument six to 20 cell layers thick (in Levenhookia four). Hypostase present in at least some species (absent in Brunonia). Megagametophyte monosporous, Polygonum type, with multicellular chalazal process. Synergids elongate, hooked. Antipodal cells sometimes persistent. Endosperm development cellular (in Dampiera rarely nuclear?). Endosperm haustoria absent. Embryogenesis solanad.

Fruit Usually a septicidal (and sometimes loculicidal) laterally dehiscing capsule (sometimes a drupe or achene with persistent calyx, rarely a schizocarp with one-seeded nutlike mericarps).

Seeds Aril usually absent (present in Coopernookia?). Testa sometimes winged, seven to 14 cell layers thick (in Brunonia thin, compressed). Exotestal cells usually palisade, with thickened walls (especially inner walls; not in Brunonia), often containing crystals. Both outer and inner epidermis persistent. Endotesta developed into endothelium? Hypodermal layers sometimes lignified. Perisperm not developed. Endosperm usually copious (in Brunonia thin or absent), oily. Embryo well developed, straight, chlorophyll? Cotyledons two (in Brunonia swollen). Germination phanerocotylar.

Cytology n = 16, 18, 24, 27, 32, 36, 45; x = 7–9 – Polyploidy frequently occurring.

DNA Mitochondrial gene rpl2 possibly lost. rpl16 intron lost. Plastid gene infA lost/defunct (Goodenia). Intron lost in plastid gene rpoC1 in Goodenia and Scaevola.

Phytochemistry O-methyl flavonols, Route I iridoids, Group VI secoiridioids (e.g., secologanin; Brunonia lacks iridoids), ursolic acid, caffeic acid, chlorogenic acid, alkaloids, saponins, cyanogenic compounds, and polyacetylenes present. Flavonols, ellagic acid and proanthocyanidins not found. Carbohydrates stored as oligo- or polyfructosans (e.g. inulin in roots and rhizome) with isokestose bonds.

Use Ornamental plants.

Systematics Leschenaultia (26–27; southern New Guinea, Australia, with their largest diversity in southwestern Western Australia), Anthotium (3; southwestern Western Australia), Dampiera (c 65; Australia, with their highest diversity in southwestern Western Australia); Brunonia (1; B. australis; Australia, Tasmania), Scaevola (120–130; Socotra, Australia, Tasmania, Melanesia, New Zealand, Polynesia to the Hawaiian Islands, Cuba, some species also along coasts of tropical Southeast Asia, Malesia, tropical America, Africa, Madagascar, the Mascarene Islands, southern India and Sri Lanka), Coopernookia (6; southwestern Western Australia, southern South Australia, eastern New South Wales), ‘Goodenia’ (180–185; New Guinea, Australia, one species also in Malesia to southern China, G. konigsbergeri on Java; non-monophyletic), Selliera (1–3; S. radicans; southeastern South Australia, Victoria, southeastern New South Wales, Tasmania, New Zealand, Stewart Island, temperate South America), Velleia (21; New Guinea, Australia, Tasmania), Verreauxia (3; southwestern Western Australia), Pentaptilon (1; P. careyi; western Western Australia).

Goodeniaceae are sister-group to [Calyceraceae+Asteraceae], although Soltis & al. (2007) identified Goodeniaceae as sister to Calyceraceae.

Cladogram of Goodeniaceae (50% majority-rule from a partitioned Bayesian inference analysis) based on DNA data (Jabaily & al. 2012).

MENYANTHACEAE (Dumort.) Dumort.

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Dumortier, Anal. Fam. Plant.: 20, 25. 1829 [’Menyanthideae’], nom. cons.

Menyanthales Bercht. et J. Presl, Přir. Rostlin 1(115): 1. 1825 [‘Menyantheae’]

Genera/species 6/60–70

Distribution Cosmopolitan except polar areas, with their largest diversity in Australia.

Fossils Unknown.

Habit Usually bisexual (rarely monoecious, dioecious or gynodioecious), usually perennial (rarely annual) herbs. Hydrophytes (aquatic) or helophytes.

Vegetative anatomy Phellogen? Vascular tissue atactostele-like, with separate and often scattered stem bundles. Cortical vascular bundles present. Cortex and mesophyll with intercellular canals and spaces. Secondary lateral growth normal or absent. Vessel elements usually with simple (rarely scalariform) perforation plates; lateral pits alternate? Imperforate tracheary xylem elements ? Wood rays absent. Axial parenchyma absent. Sieve tube plastids S type. Endodermis sometimes prominent. Nodes 3:3, trilacunar with three leaf traces, or 5:5, pentalacunar with five traces. Parenchyma in some species with idioblasts containing asterosclereids and projecting into intercellular spaces. Crystals?

Trichomes Hairs absent.

Leaves Alternate (spiral or distichous), usually simple (in Menyanthes trifoliolate), entire, in Nymphoides sometimes anisomorphous, with involute ptyxis. Stipules absent; petiole sometimes sheathing and decurrent. Colleters present. Petiole with lateral wings. Petiole vascular bundle transection arcuate to annular. Leaf base in Nephrophyllidium wide. Venation palmate or pinnate. Stomata anomocytic, often present at adaxial side only. Cuticular wax crystalloids? Mesophyll with or without sclerenchymatous idioblasts containing asterosclereids. Leaf margin usually entire (sometimes slightly serrate or crenate), usually glanduliferous, often with hydathodes.

Inflorescence Terminal panicle, raceme, umbel or capitate, or flowers paired or solitary axillary.

Flowers Actinomorphic, often large. Hypogyny (probably secondary) or half epigyny. Sepals usually five (rarely four), persistent, usually connate in lower part. Petals usually five (rarely four), usually with valvate (rarely weakly imbricate) aestivation, connate in lower part, with margins or adaxial surface often fimbriate or hairy and with wings (broad edges). Lateral petal traces fused, possibly indicating primary zygomorphy. Nectariferous disc intrastaminal, usually surrounding ovary base. Heterostyly frequent.

Androecium Stamens usually five (rarely four), haplostemonous, antesepalous, alternipetalous. Filaments free from each other, adnate at base to corolla tube (epipetalous). Anthers dorsifixed or basifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with multinucleate cells (nuclei fused). Female flowers with staminodia; fimbriate scale-like staminodia sometimes alternating with stamens.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually 3(–6)-colp(or)ate (Menyanthes, Nephrophyllidium) or parasyncolpate (Liparophyllum, Nymphoides, Villarsia), shed as monads, usually tricellular (sometimes bicellular) at dispersal. Exine tectate, with columellate infratectum, imperforate?, striate to rugulate (Menyanthes, Nephrophyllidium), or striate, rugulate or smooth (Liparophyllum, Nymphoides, Villarsia).

Gynoecium Pistil composed of two connate carpels. Ovary superior (probably secondarily) or semi-inferior, unilocular, with nectariferous glands at base. Style single, bifid, or absent. Stigmas lobate or spatulate, papillate, Wet type. Male flowers with pistillodium.

Ovules Placentation (often intrusively) parietal. Ovules numerous per carpel, anatropous, horizontal, unitegmic, tenuinucellar. Integument more than ten? cell layers thick. Hypostase present or absent. Megagametophyte monosporous, Polygonum type. Endosperm development cellular. Endosperm haustoria absent. Embryogenesis asterad.

Fruit Usually a septicidal and/or loculicidal or irregularly dehiscing capsule (in Nymphoides sometimes nutlike; in Liparophyllum a berry).

Seeds Aril absent. Seed coat exotestal. Testa sometimes winged, sometims with air-filled and/or hooked hairs. Elaiosome (caruncle rich in lipids) present in some species. Exotestal cells with thickened outer walls, often with various surface processes. Mesotesta and endotesta usually crushed (sometimes with sclerotized cells). Perisperm not developed. Endosperm copious, oily. Embryo well developed, straight, chlorophyll? Cotyledons two. Germination?

Cytology x = 9 (Menyanthes, Nymphoides, Villarsia); x = 17 (Nephrophyllidium); n = 8, 9, 18, 20, 27, 28, 54 – Polyploidy occurring. Protein bodies present in nucleus.

DNA Intron absent from plastid gene rpl2 (entire gene absent?).

Phytochemistry Flavonols (kaempferol, quercetin etc.), Group VI secoiridoids (e.g. secologanin), Group VII secoiridoids (e.g. sweroside), Group X secoiridoids (e.g. loganin), caffeic acid, iridoid alkaloids, and saponins present. Ellagic acid, tannins, proanthocyanidins and cyanogenic compounds not found. Carbohydrates stored as oligo- or polyfructosans (e.g. inulin in roots and rhizome) with isokestose bonds (starch absent).

Use Ornamental plants, medicinal plants.

Systematics Menyanthes (1; M. trifoliata; temperate regions on the Northern Hemisphere), Nephrophyllidium (1; N. crista-galli; northern Japan, northwestern North America); Ornduffia (7; southwestern Western Australia, South Australia, eastern Queensland to Victoria, Tasmania), Villarsia (3; Western Cape, Thailand, Indochina), Liparophyllum (8; southwestern Western Australia, South Australia, eastern Queensland to Victoria, Tasmania, New Zealand), Nymphoides (c 40; nearly cosmopolitan).

Menyanthaceae are sister to the clade [Goodeniaceae+[Calyceraceae+Asteraceae]].

[Menyanthes+Nephrophyllidium] is sister-group to the remainder, whereas Ornduffia is sister to [Villarsia+[Nymphoides+Liparophyllum]].

Cladogram of Menyanthaceae based on DNA sequence data and morphology (Tippery & Les 2009; Tippery & al. 2009).

PENTAPHRAGMATACEAE J. Agardh

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Agardh, Theoria Syst. Plant.: 95. Apr-Sep 1858 [’Pentaphragmeae’], nom. cons.

Pentaphragmatales Doweld, Tent. Syst. Plant. Vasc.: liv. 23 Dec 2001

Genera/species 1/c 30

Distribution Southeast Asia, southern China, Malesia to New Guinea, with their largest diversity in West Malesia.

Fossils Unknown.

Habit Usually bisexual (rarely dioecious), perennial herbs, large and often somewhat succulent. Rooting at stem base.

Vegetative anatomy Phellogen absent. Endodermis conspicuous. Vessel elements with scalariform perforation plates; lateral pits scalariform. Imperforate tracheary xylem elements fibre tracheids with bordered pits, often septate. Wood rays uniseriate, biseriate, homocellular, or absent. Axial parenchyma absent. Wood not storied. Sieve tube plastids S type? Nodes unilacunar? with ? leaf traces. Laticifers and latex probably absent. Crystals absent.

Trichomes Hairs unicellular and uniseriate, or multicellular and uniseriate or branched. Foliar hairs often multicellular and branched, often with uniseriate branches.

Leaves Alternate (distichous), simple, entire, usually with strongly asymmetrical base, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation pinnate. Stomata anomocytic or anisocytic, surrounded by three or four cells slightly different from normal epidermal cells. Cuticular wax crystalloids? Leaf margins usually serrate (sometimes entire).

Inflorescence Axillary, cymose, usually scorpioid (helicoid). Bracts usually conspicuous, membranous.

Flowers Actinomorphic or slightly zygomorphic. Hypanthium usually present, adnate to ovary by five longitudinal septa creating five nectariferous pits. Epigyny. Sepals two large and three small, with imbricate aestivation, petaloid, persistent, often connate. Petals (four or) five, with valvate aestivation, usually fleshy or cartilaginous, persistent, usually connate at base, with marginal wings. Five nectar-producing antepetalous pores, lacunae, inserted between five longitudinal septa fusing hypanthium with ovary. Disc absent.

Androecium Stamens (four or) five, haplostemonous, antesepalous, alternipetalous. Filaments flat, free from each other, in sympetalous species adnate to corolla tube (epipetalous). Anthers basifixed, non-versatile, tetrasporangiate, extrorse, longicidal (dehiscing by longitudinal septa). Tapetum secretory, with binucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3-colporate or (2–)3-colpate, trilobate and pleurotreme in polar view, with apertures between lobes, shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, imperforate?, smooth; endexine lamellate.

Gynoecium Pistil composed of two (or three) connate carpels. Ovary inferior, usually bilocular (rarely trilocular), inserted at hypanthium by (four or) five narrow longitudinal septa, being continuations of staminal filaments and forming (four or) five nectariferous canals or pits. Style single, simple, short and thick. Stigma capitate or clavate, type? Pistillodium absent.

Ovules Placentation axile (placentae bipartite). Ovules numerous per carpel, anatropous, pendulous, unitegmic, tenuinucellar. Integument approx. three cell layers thick. Hypostase absent. Megagametophyte monosporous, Polygonum type, protruding from micropyle (extramicropylar). Endosperm development cellular. Endosperm haustorium micropylar, unicellular and uninucleate (chalazal endosperm haustorium absent). Embryogenesis solanad.

Fruit A many-seeded berry with persistent perianth.

Seeds Seeds minute. Aril absent. Exotestal cells cuboidal, with lignified inner walls. Endotesta developing into an endothelium? Perisperm not developed. Endosperm copious, starchy. Suspensor short. Embryo straight, well developed, two thirds the length of the seed, chlorophyll? Cotyledons two. Germination?

Cytology n = 54–56

DNA Mitochondrial gene rpl2 absent (lost).

Phytochemistry Very insufficiently known. Inulin and alkaloids not found. Iridoids?

Use Vegetables.

Systematics Pentaphragma (25–30; southern Burma, southern China, Indochina, Malesia to New Guinea, with their largest diversity in West Malesia).

Pentaphragma may be sister to the remaining Campanulales above the [Rousseaceae+Campanulaceae] clade, although the support is sometimes low.

PHELLINACEAE (Loes.) Takht.

( Back to Campanulales )

Takhtajan, Sist. Filogen. Cvetk. Rast. [Syst. Phylog. Magnolioph.]: 343, 374. 4 Feb 1967 [’Phellineaceae’]

Phellinales Doweld, Tent. Syst. Plant. Vasc.: liv. 23 Dec 2001; Phellinanae Doweld, Tent. Syst. Plant. Vasc.: liv. 23 Dec 2001

Genera/species 1/12

Distribution New Caledonia.

Fossils Fossil leaf fragments of Phelline has been reported from the Early Miocene of New Zealand (Pole 2010).

Habit Dioecious, evergreen trees or shrubs.

Vegetative anatomy Phellogen? Vessel elements with scalariform perforation plates (end walls almost vertical); lateral pits scalariform or opposite (in one row), bordered pits. Imperforate tracheary xylem elements tracheids or fibre tracheids with bordered pits, non-septate. Wood rays uniseriate or multiseriate (up to 14 cells wide and often more than 2 cm tall), heterocellular. Axial parenchyma apotracheal diffuse, or paratracheal scanty. Sieve tube plastids S type. Nodes 3?:3?, trilacunar? with three? leaf traces. Acicular calciumoxalate crystals, styloids, crystal sand and other crystal types present.

Trichomes Hairs uniseriate, often reddish-brown, tufted.

Leaves Alternate (spiral), simple, entire, often coriaceous, with ? ptyxis. Stipules and leaf sheath absent. Axillary hairs present. Petiole vascular bundle transection annular. Venation pinnate. Stomata anomocytic, with very large guard cells, with inner and outer ledges. Cuticular wax crystalloids as platelets and rodlets. Sclerenchymatic tissue enclosing veins. Leaf margin crenate-sinuate or entire.

Inflorescence Axillary spike- or raceme-like or paniculate cymose.

Flowers Actinomorphic, small. Hypogyny. Sepals four or five (or six), with more or less open aestivation, minute, persistent, connate at base. Petals four or five (or six), with valvate aestivation, fleshy, with small incurved apex, free. Nectary absent. Disc absent.

Androecium Stamens four or five (or six), haplostemonous, antesepalous, alternipetalous. Filaments free from each other and from tepals. Anthers basi-dorsifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum glandular? Female flowers with staminodia.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolpor(oid)ate, shed as monads, ?-cellular at dispersal. Exine tectate, with columellate infratectum, perforate, microechinulate or microrugulate.

Gynoecium Pistil composed of (two or) four or five (or six) connate carpels. Ovary superior, usually quadrilocular or quinquelocular (sometimes bilocular or sexalocular). Style single, simple, very short or absent. Stigma usually quadrilobate or quinquelobate, type? Male flowers with pistillodium.

Ovules Placentation axile to apical. Ovule one per carpel, somewhat campylotropous to hemitropous, pendulous, apotropous, unitegmic, tenuinucellar. Integument ? cell layers thick. Megagametophyte monosporous, Polygonum type? Endosperm development? Endosperm haustoria? Embryogenesis?

Fruit A drupe with one to five separate pyrenes.

Seeds Aril absent. Exotesta present. Endotesta? Perisperm not developed. Endosperm copious. Embryo small, chlorophyll? Cotyledons two. Germination?

Cytology n = 17

DNA Mitochondrial gene rpl2 absent?

Phytochemistry Insufficiently known. Benzylisoquinoline alkaloids (homoerythrine alkaloids: homoerythrine, homoerythroidine, homoazaerythrine, holidine, etc.) present. Route I iridoids? Carbohydrates stored as oligo- or polyfructosans (e.g. inulin) with isokestose bonds (starch absent).

Use Unknown.

Systematics Phelline (12; New Caledonia).

Phelline is sister to [Argophyllaceae+Alseuosmiaceae].

ROUSSEACEAE DC.

( Back to Campanulales )

de Candolle, Prodr. 7(2): 521. late Dec 1839 [‘Roussaeaceae’]

Carpodetaceae Fenzl in Denkschr. Königl.-Baier. Bot. Ges. Regensburg 3: 155. 28 Nov 1841 [’Carpodeteae’]; Abrophyllaceae Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 243. 20 Jul 1943; Rousseales Doweld, Tent. Syst. Plant. Vasc.: liii. 23 Dec 2001

Genera/species 4/15

Distribution Mauritius, New Guinea, the Solomon Islands, eastern Australia, New Zealand, Stewart Island.

Fossils Unknown.

Habit Usually bisexual (in Carpodetus sometimes gynodioecious), evergreen trees or shrubs (Roussea often climbing).

Vegetative anatomy Phellogen? Young stem with separate vascular bundles. Vessel elements with scalariform perforation plates; lateral pits scalariform, opposite or alternate, bordered pits. Imperforate tracheary xylem elements tracheids or fibre tracheids with bordered pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty (Roussea). Sieve tube plastids S type. Nodes 3:3, trilacunar with three leaf traces. Pericycle and leaf mesophyll in Abrophyllum and Cuttsia with latex idioblasts consisting of vertically elongate cells arranged in longitudinal rows and filled with white brittle content, and ring of sclerenchyma. Parenchyma cells in Carpodetus with rhomboidal crystals.

Trichomes Hairs unicellular, usually curved, thick-walled, verrucose; glandular hairs in Abrophyllum and Cuttsia with short sunken stalk and unicellular head; hairs in Roussea unicellular conical or glandular.

Leaves Usually alternate (spiral; in Roussea opposite), simple, entire, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate or annular. Venation usually semicraspedodromous (in Roussea craspedodromous). Stomata anomocytic (Roussea, Carpodetus), in Abrophyllum and Cuttsia with two almost circular cells. Cuticular wax crystalloids? Petiole with radially elongate schizogenous resinous canals (Roussea). Mesophyll in Carpodetus with crystalliferous idioblasts. Petiole and abaxial side of lamina with multicellular peltate glandular hairs (Roussea); domatia as pockets sometimes present in Carpodetus. Leaf margin more or less glandular serrate.

Inflorescence Usually terminal or axillary panicle (flowers in Roussea few or solitary, axillary).

Flowers Actinomorphic, usually small (in Roussea large). Usually hypogyny (in Carpodetus epigyny to half epigyny). Sepals (four or) five (to seven), with valvate aestivation, unequally long, persistent (Roussea) or caducous, free or connate at base. Petals (four or) five (to seven), with valvate aestivation, thick, persistent (Roussea) or caducous, usually free (in Roussea connate in lower part). Nectariferous disc intrastaminal, in Roussea quadrilobate or quinquelobate, or absent.

Androecium Stamens (four or) five (to seven), haplostemonous, antesepalous, alternipetalous. Filaments free from each other and from tepals, in Abrophyllum and Cuttsia inserted at margin of nectariferous disc, in Roussea inserted on lobes of nectariferous disc. Anthers usually basifixed (in Cuttsia dorsifixed?), non-versatile, tetrasporangiate, usually introrse (in Roussea extrorse), longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate cells. Female flowers with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate (in Roussea penta- or hexapantoporate, or penta- or hexapantobrachycolpate), usually shed as monads (in Carpodetus as tetrads), usually bicellular (in Carpodetus tricellular) at dispersal. Endoapertures in Cuttsia H-shaped. Exine tectate, with columellate infratectum, striate to rugulate (Abrophyllum, Cuttsia), or smooth to somewhat uneven (Carpodetus, Roussea).

Gynoecium Pistil composed of (three to) five (to seven) antepetalous carpels. Ovary usually superior (sometimes inferior or semi-inferior), usually quinquelocular (sometimes tri-, quadri-, sexa- or septalocular; Roussea). Style single, usually simple (in Roussea thick, persistent; in Carpodetus narrow; in Cuttsia apically branched) or absent. Stigma capitate (Carpodetus, Roussea), or (trilobate to) quinquelobate, type? Male flowers with pistillodium.

Ovules Placentation axile. Ovules numerous per carpel, anatropous, unitegmic, tenuinucellar. Integument five to eight cell layers thick. Megagametophyte monosporous, Polygonum type. Suspensor filiform (Roussea). Endosperm development ab initio nuclear. Endosperm haustorium micropylar (Roussea). Embryogenesis?

Fruit Usually a berry, in Roussea sharply quinqueangular (to septangular), in Abrophyllum and Cuttsia with persistent stigma, in Carpodetus (trilocular to) quinquelocular and leathery (in Cuttsia a loculicidal capsule).

Seeds Aril absent. Anticlinal and inner periclinal walls of exotestal cells more or less massively thickened (in Carpodetus entirely). Mesotesta and endotesta crushed (Roussea). Perisperm not developed. Endosperm copious, with aleuron and lipids. Embryo small, straight, chlorophyll? Cotyledons two. Germination?

Cytology n = 14, 15 (Carpodetus)

DNA Mitochondrial gene rpl2 absent?

Phytochemistry Insufficiently known. Flavonols (kaempferol, quercetin), triterpenes (lupeol), and prodelphinidins present (Carpodetus). Tannins absent in Roussea.

Use Unknown.

Systematics Rousseaceae are probably sister-group to Campanulaceae.

Rousseoideae Horan., Char. Ess. Fam.: 110. 17 Jun 1847 [‘Roussaeariae’]

1/1. Roussea (1; R. simplex; Mauritius). – Evergreen small tree or liana. Resiniferous canals present. Hairs tufted-stellate; glandular hairs peltate. Leaves opposite or verticillate. Petiole vascular bundle transection annular (with endodermis?). Leaf margins serrate. Bud scales present. Flowers terminal, solitary. Flowers usually pentamerous (sometimes tetramerous). Hypogyny. Calyx large, with valvate aestivation. Petals large, connate at base. Anthers basifixed, extrorse, entirely fused with straight connective. Pollen grains hexaporate or octaporate. Pollen embedded in slime when liberated. Tectum almost imperforate (complete). Pistil composed of five to seven connate carpels. Style apically expanding, persistent, gradually widening into ovary. Stigmatic lobes narrower, erect. Endosperm haustorium micropylar. Fruit a berry, with persistent calyx with reflexed lobes. Seeds carunculate? Exotestal cell walls thickened. Mesotesta and endotesta crushed. Embryo relatively long. n = ? Phytochemistry virtually unknown.

Carpodetoideae (Fenzl) J. Lundb. in Bot. J. Linn. Soc. 137: 276. 3 Dec 2001

3/14. Carpodetus (2–11; New Guinea, the Solomon Islands, New Zealand, Stewart Island), Abrophyllum (1–2; A. ornans; eastern Queensland, eastern New South Wales), Cuttsia (1; C. viburnea; southeastern Queensland, northeastern New South Wales). – New Guinea, eastern Australia, Melanesia, New Zealand. Evergreen trees. Hairs unicellular, thick-walled, strongly curved, verrucose. Petiole vascular bundle transection arcuate or annular; petiole with additional accessory bundles. Inflorescence terminal panicle. Flowers tetramerous to hexamerous. Calyx caducous. Petals finally free. Filaments free from each other and from petals. Pollen grains in Carpodetus shed in tetrads. Pistil composed of three to six connate carpels. Style usually short (occasionally absent). Stigma capitate (in Cuttsia divided). Fruit dry, baccate, or a loculicidal and septicidal capsule. Seeds numerous. Funicle elongate. Anticlinal and inner periclinal exotestal cell walls strongly thickened (in Carpodetus all walls thickened). Endosperm with hemicellulose (Carpodetus). Mesophyll in Abrophyllum and Cuttsia containing white groups of small unlignified cells, similar to raphide bundles. n = 14, 15. – Carpodetus is sister to [Abrophyllum+Cuttsia] (Kårehed & al. 1999).

Cladogram of Rousseaceae based on morphology and DNA sequence data (Kårehed & al. 1999).

STYLIDIACEAE R. Br.

( Back to Campanulales )

Brown, Prodr. Fl. Nov.-Holl.: 565. 27 Mar 1810 [’Stylideae’], nom. cons.

Stylidiales R. Br. ex Bercht. et J. Presl, Přir. Rostlin: 253. Jan-Apr 1820 [‘Stylideae’]; Candolleaceae F. Muell., Syst. Census Austral. Plant.: 85. late 1882-early 1883, nom. illeg.; Donatiaceae (Engl.) B. Chandler in Notes Roy. Bot. Gard. Edinburgh 5: 44. Nov 1911, nom. cons.

Genera/species 4/240–320

Distribution: South and Southeast Asia, mainly extratropical regions of Australia, Tasmania, New Zealand, southernmost South America.

Fossils Fossil pollen grains, Tricolpites stylidioides, resembling those in Forstera, have been reported from Oligocene layers in southeastern Australia.

Habit Usually bisexual (sometimes monoecious or polygamomonoecious), usually perennial herbs (sometimes dwarf shrubs; in Donatia and Phyllachne cushion-shaped). Some species are helophytes and numerous species are xerophytes.

Vegetative anatomy Phellogen ab initio cortical. Young stem with separate vascular bundles. Secondary lateral growth anomalous (via cylindrical cambium, produced outside vascular bundles, with cambial activitiy mainly unifacial producing tissue towards inner side). Vessel elements usually with simple (in Donatia scalariform) perforation plates; lateral pits alternate, bordered pits. Imperforate tracheary xylem elements fibre tracheids with bordered pits, non-septate. Wood rays absent. Axial parenchyma absent (Stylidium). Wood elements partially storied. Interxylary phloem present. Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Crystals?

Trichomes Hairs in Donatia unicellular, uniseriate, in leaf axils; glandular hairs with stalk and unicellular or multicellular head (some hairs mucilaginous), or absent.

Leaves Alternate (spiral), simple, entire, usually linear, often coriaceous, with ? ptyxis. Stipules and leaf sheath absent. Petiole absent. Venation usually parallelodromous, often flabellate or striate (in Donatia acrodromous). Stomata usually anomocytic (in Donatia magellanica paracytic). Cuticular wax crystalloids? Mesophyll with idioblasts as mucilage cells, without sclerenchymatous idioblasts. Leaf margin entire. Leaf apex sometimes with extrafloral nectary.

Inflorescence Terminal, cymose (often raceme- or spike-like) or raceme, spike etc., or flowers solitary terminal (in, i.a., Donatia).

Flowers Zygomorphic and resupinate (in Levenhookia, Stylidium and most species of Forstera), or actinomorphic (in Donatia, Oreostylidium, Phyllachne and some species of Forstera). Epigyny. Sepals (two to) five (to seven), usually with imbricate (in Donatia open?) aestivation, persistent, usually connate. Petals usually five, with imbricate aestivation, unequally sized; odd corolla lobe laid down ab initio ventrally and gradually reduced (in, e.g., Stylidium), or becoming large dome-shaped labellum (in, e.g., Levenhookia), although flowers usually resupinating (flowers semi-resupinate, split-monosymmetric) labellum thus becoming lateral, remaining four corolla lobes uniform or non-uniform (petals in Donatia five to ten, equally sized), usually connate and tubular (in Donatia free); corolla tube formation early; corolla tube mouth often with row of glandular appendages. Nectary epigynous or absent. Disc extrastaminal, usually bifid (sometimes unilateral; in Donatia discoid).

Androecium Stamen usually two (in Donatia two or three), haplostemonous, alternisepalous, antepetalous. Filaments usually connate and tubular (in Donatia free), free from tepals, usually completely adnate to style into gynostemium (stamens in Donatia free from style). Anthers basifixed? (in Donatia dorsifixed), non-versatile, tetrasporangiate, extrorse, longicidal (dehiscing by longitudinal slits); thecae arranged head-to-head. Tapetum secretory, with binucleate cells. Staminodia absent. Secondary pollen display as androphore-like structure, gynostemium, present as stylar column with stigma and anthers at apex.

Pollen grains Microsporogenesis simultaneous. Pollen grains 2–8-colpate (in Donatia 3–4-colp[or]ate), shed as monads, bicellular or tricellular at dispersal (in Donatia?). Exine tectate, with usually columellate infratectum, imperforate to finely perforate, usually spinulate (in Donatia smooth).

Gynoecium Pistil composed of usually two (in Donatia two or three) connate carpels. Ovary inferior, usually bilocular below, unilocular above and sometimes bilocular distally (sometimes pseudomonomerous due to reduced posterior carpel, or unilocular due to absence of septum; in Donatia bilocular or trilocular). Stylodia usually two, usually connate and adnate to filaments (in Donatia two or three, subulate, free). Stigma bilobate (sometimes bristle-like) or stigmas capitate, papillate, Dry type. Pistillodium absent.

Ovules Placentation usually free central in unilocular part of ovary (sometimes free basal; in Donatia axile in bilocular part of ovary). Ovules usually numerous per carpel (one or four when one locule fertile), anatropous, pendulous, horizontal or ascending, unitegmic, tenuinucellar. Integument four to six cell layers thick. Hypostase present in Donatia. Megagametophyte monosporous, Polygonum type. Synergids sometimes with a filiform apparatus. Antipodal cells persistent. Endosperm development cellular. Endosperm haustoria chalazal and micropylar (micropylar haustorium most aggressive). Embryogenesis solanad.

Fruit Usually a septicidal capsule (in, e.g., Donatia a nutlike indehiscent capsule).

Seeds Aril? Testa one to four cell layers thick. Exotesta? Only outer epidermis persistent. Endotesta developing into an endothelium? Perisperm not developed. Endosperm copious, fleshy, oily. Embryo small, well differentiated, chlorophyll? Cotyledons usually two (rarely one). Germination phanerocotylar.

Cytology n = 24 (Donatia); n = 5–16, 18, (24), 26, 28, 30 (mainly in Stylidium) – Protein bodies present in nucleus (Stylidioideae).

DNA Mitochondrial gene rpl2 absent (lost).

Phytochemistry Flavonols (kaempferol, quercetin), Route I iridoids (Route I carbocyclic iridoid resembling monotropein; in Donatia?), caffeic acid, tannins (in Donatia), proanthocyanidins, and saponins (rare) present. Ellagic acid, alkaloids, and cyanogenic compounds not found. Secoiridoids? Carbohydrates stored as oligo- or polyfructosans (e.g. inulin) with isokestose bonds (starch absent).

Use Ornamental plants.

Systematics Stylidiaceae are sister-group to the clade [Phellinaceae+[Argophyllaceae+Alseuosmiaceae], according to Tank & Donoghue (2010). On the other hand, they have sometimes been recovered as sister to the clade [Menyanthaceae+[Goodeniaceae+[Calyceraceae+Asteraceae]]] or as members of a trichotomy also comprising these two clades.

Donatioideae Mildbr. in H. G. A. Engler, Pflanzenr. 35: 18. 26 Mai 1908

1/2. Donatia (2; D. novae-zelandiae: New Zealand, Stewart Island, Tasmania; D. fascicularis: southernmost Chile and Argentina). – Cushion-shaped dwarf shrubs. Often xerophytes. Phellogen cortical? Older stem with very thick cortex and with vascular tissue as a central narrow cylinder. Mucilage cells present. Hairs uniseriate, axillary. Leaves often ericoid. Venation acrodromous. Stomata usually paracytic. Flowers solitary, terminal. Flowers actinomorphic. Sepals three to seven, free. Petals five to ten, free. Stamens two or three. Filaments free from each other and from petals. Anthers extrorse. Pistil composed of two or three connate carpels. Stylidia free, recurved. Stigmas capitate. Hypostaste present. Fruit indehiscent. Suspensor short. n = 24. Tannins present.

Stylidioideae Kitt. in A. Richard, Nouv. Elém. Bot., ed. 3, Germ. transl.: 827. 1840 [‘Stylideae’]

3/c 240. Levenhookia (10; southwestern Western Australia, southeastern South Australia, Victoria, New South Wales), Stylidium (probably at least 300; Bengal, Sri Lanka, Burma, Vietnam, southern coast of China, the Malay Peninsula, the Philippines, Australia, Tasmania, New Zealand, Stewart Island, with almost all species confined to Australia), Forstera (9; Tasmania, New Zealand, Stewart Island, Auckland Islands, Campbell Island, southernmost South America). – Distribution as for Stylidiaceae. Herbs (sometimes climbing or twining; sometimes cushion-shaped). Phellogen superficial or outer-cortical. Phelloderm developing centrifugally. Secondary lateral growth anomalous (secondary tissue developing exclusively towards inside). Cambium storied, developing from beneath endodermis. Vascular bundles closed, scattered or annular. Intraxylary phloem present. Xylem (sometimes also phloem) developing centripetally. Vessel elements with simple perforation plates. Hairs also axillary. Glandular hairs abundant. Leaves pseudoverticillate or in rosette (rosette often present at distance above ground). Flowers usually zygomorphic (in Stylidium occasionally obliquely zygomorphic; rarely actinomorphic). Sepals connate; median sepal abaxial, yet often semi-resupinate. Petals usually 2+2 (occasionally five; in Levenhookia four plus sensitive labellum), connate into tube, often with glandular appendages forming corona, sometimes with spur. Nectaries also as paired glands. Stamens two, entirely adnate to style, thus forming gynostemium (column). Adaxial staminal pair fertile. Anthers inserted near stigma. Thecae with connivent tips. Entire complex usually sensitive (not in Levenhookia?) and moving when touched by pollinator (cf. ‘trigger plants’). Pollen grains 2–8-colpate, bicellular or tricellular at dispersal. Pistil composed of two connate carpels (adaxial carpel strongly reduced). Stigma small, Dry type. Synergids elongate. Fruit usually a septicidal capsule (sometimes indehiscent). Seed coat exo-endotestal. Exotestal cell walls sclerotized. Suspensor much longer than wide. Cotyledon often single. n = 5–16, 18, (24), 26, 28, 30 – Two monophyletic groups were identified by Wagstaff & Wege (2002). The first one consists of Forstera and Phyllachne. None of these are being monophyletic and hence Forstera shoule be included in Phyllachne.

The second lineage comprises Levenhookia as sister to Stylidium. According to Darnowski & al. (2006), there are signs (protease activity in leaves) indicating that at least a few species may be carnivorous or at least insect-trapping.

The gynostemium consists, at the bending point, of an epidermis together with anterior and posterior parenchymatic layers, the cells of which containing large numbers of starch grains, mitochondria and vacuoles. The cell walls are rich in pectin and their cellulose fibrils are almost exclusively circumferential. When the gynostemium is ready to be “fired”, the anterior parenchyma layer has a high turgor – balanced by stretched cells with longitudinal cellulose fibrils in the posterior parenchyma layer – and large KCl concentration. When the sensitive gynostemium is stimulated by the pollinator, the cell walls at the bending point will recurve extremely fast transversely and thus “fire” the gynostemium. This movement takes only about ten to 30 milliseconds – incrediby fast for a plant.

Cladogram of Stylidiaceae based on DNA sequence data (Wagstaff & Wege 2002).


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Robinson H, Brettell RD. 1973c. Tribal revisions in the Asteraceae V. The relationship of Rigiopappus. – Phytologia 26: 69-70.

Robinson H, Brettell RD. 1973d. Tribal revisions in the Asteraceae VII. The relationships of Isoetopsis. – Phytologia 26: 73-75.

Robinson H, Brettell RD. 1973e. Tribal revisions in the Asteraceae VIII. A new tribe, Ursinieae. – Phytologia 26: 76-86.

Robinson H, Brettell RD. 1973f. Tribal revisions in the Asteraceae IX. The relationship of Ischnea. – Phytologia 26: 153-158.

Robinson H, Brettell RD. 1973g. Tribal revisions in the Asteraceae X. The relationship of Plagiocheilus. – Phytologia 26: 159-162.

Robinson H, Brettell RD. 1973h. Tribal revisions in the Asteraceae XI. A new tribe, Eremothamneae. – Phytologia 26: 163-166.

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Robinson H, Brettell RD. 1973l. Studies in the Senecioneae (Asteraceae) IV. The genera Mesadenia, Syneilesis, Miricacalia, Koyamacalia, and Roldana. – Phytologia 27: 265-276.

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