The Families of Flowering Plants

L. Watson and M. J. Dallwitz

Compositae Giseke

Alternatively Asteraceae Dum. (nom. altern.)

Including Acarnaceae Link, Ambrosiaceae Dum. & Link (reduced, anemophilous Heliantheae), Anthemideae (Anthemidaceae) Link, Arctotidaceae Bessey, Calendulaceae Link, Carduaceae Dum., Cassiniaceae Sch. Bip., Cichoraceae (Cichoriaceae) Juss., Coreopsideae (Coreopsidaceae) Link, Cynaraceae Juss., Echinopaceae Dum., Echinopiaceae Link corr. Bullock, Echinopsidaceae Link, Eupatoriaceae Link, Heleniaceae Bessey, Helianthaceae Bessey, Helichrysaceae Link corr. Bullock, Inulaceae Bessey, Lactucaceae Bessey, Mutisiaceae Burnett, Nucamentaceae Hoffmgg., Partheniaceae Link, Perdicieae (Perdiciaceae) Link, Ritroneae (Ritronaceae) Hoffmgg. & Link, Senecionidaceae (Senecionaceae) Bessey, Spurionucaceae Dulac, Vernoniaceae Bessey

Habit and leaf form. Herbs (mostly), or trees (rarely), or ‘arborescent’, or shrubs (rarely), or lianas (rarely); laticiferous, or non-laticiferous and without coloured juice; bearing essential oils, or without essential oils; resinous, or not resinous. ‘Normal’ plants, or switch-plants. Leaves well developed, or much reduced (sometimes). Plants non-succulent, or succulent (a few). Annual, biennial, and perennial; with a basal aggregation of leaves, or with neither basal nor terminal aggregations of leaves. Self supporting (usually), or climbing (rarely); the climbers stem twiners, or scrambling; Mikania twining anticlockwise. Hydrophytic (very rarely, e.g. species of Bidens, Cotula), or helophytic, or mesophytic, or xerophytic; the hydrophytes rooted. Leaves of hydrophytes submerged and emergent. Leaves alternate (usually), or opposite (less often), or whorled (rarely); when alternate, spiral; ‘herbaceous’, or leathery, or fleshy, or membranous, or modified into spines; petiolate to sessile; sheathing, or non-sheathing. Leaf sheaths with free margins. Leaves gland-dotted, or not gland-dotted; aromatic, or foetid, or without marked odour; simple, or compound; peltate (occasionally), or not peltate. Lamina dissected, or entire; pinnatifid, or palmatifid, or runcinate, or spinose. Leaves exstipulate (nearly always), or stipulate (rarely). Lamina margins entire, or crenate, or serrate, or dentate; flat, or revolute, or involute. Leaves without a persistent basal meristem; becoming compound from primordial lobes. Domatia recorded (only from a Vernonia species); represented by pits.

General anatomy. Plants with laticifers (articulated, mainly in the Cichorieae), or without laticifers. The laticifers in leaves, or in stems, or in roots, or in flowers, or in the fruits (in all or different combinations of these).

Leaf anatomy. Hydathodes present (occasionally), or absent.

Lamina with secretory cavities, or without secretory cavities. Secretory cavities containing resin, or containing latex. Minor leaf veins with phloem transfer cells (58 genera), or without phloem transfer cells (Barnadesia and Eupatorium only).

Stem anatomy. Secretory cavities present, or absent; with resin, or with latex. Nodes unilacunar, or tri-lacunar, or multilacunar. Cortical bundles present, or absent. Medullary bundles present, or absent. Internal phloem present, or absent. Secondary thickening absent (?), or developing from a conventional cambial ring, or anomalous; when anomalous, via concentric cambia, or from a single cambial ring. ‘Included’ phloem present (e.g. Chrysanthemoides, Stoebe), or absent. Xylem with libriform fibres; with vessels. Vessel end-walls simple (commonly), or scalariform and simple, or simple and reticulately perforated. Vessels without vestured pits. Wood storied, or partially storied, or not storied; parenchyma typically rather sparse and paratracheal. Sieve-tube plastids S-type. Pith with diaphragms, or without diaphragms.

Reproductive type, pollination. Plants gynomonoecious (commonly, with hermaphrodite disk florets and female ray florets), or hermaphrodite, or monoecious, or dioecious, or androdioecious (rarely), or gynodioecious (rarely), or polygamomonoecious. Entomophilous (mostly), or anemophilous (in the Anthemideae, which include significant hayfever plants). Pollination mechanism conspicuously specialized (via stylar modification, with active presentation, and the Cynareae also with irritable stamens), or unspecialized (anemophilous forms).

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’; in heads. The terminal inflorescence unit racemose (but the primary capitula sometimes in cymose secondary heads). Inflorescences nearly always indeterminate heads, but sometimes primary ‘heads’ reduced to single florets are grouped into secondary heads — e.g. Echinops; with involucral bracts; usually more or less pseudanthial. Flowers bracteate (the bracts forming an involucre in one to several series); minute to small; regular, or regular and somewhat irregular, or regular and very irregular (often combining central actinomorphic and marginal ‘ray’ florets); (4–)5 merous; cyclic; tricyclic, or tetracyclic.

Perianth with distinct calyx and corolla, or petaline (calyx sometimes absent — e.g. Ambrosia and relatives); (1–)3–35; 1 whorled, or 2 whorled. Calyx when present, (1–)2–30 (of scales, awns or bristles constituting the ‘pappus’); represented by bristles (commonly), or not represented by bristles; 1 whorled; persistent, or not persistent; often accrescent (becoming the pappus); open in bud. Corolla 1–3 (ligulate florets), or (4–)5 (disk florets); 1 whorled; gamopetalous; valvate; unequal but not bilabiate (ligulate), or regular, or unequal but not bilabiate and regular, or bilabiate (in Mutisieae).

Androecium 3–5. Androecial members adnate; coherent; 1 whorled. Stamens 3–5; oppositisepalous (inserted on the corolla tube, alternating with the lobes); filantherous (with short filaments). Anthers cohering (nearly always, forming a tube around the style — with a few exceptions among anemophilous Anthemideae); basifixed; dehiscing via longitudinal slits; tetrasporangiate (usually), or bisporangiate (occasionally); usually appendaged. The anther appendages apical, or apical and basal. Endothecium developing fibrous thickenings. Anther epidermis persistent. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral, or decussate. Anther wall initially with one middle layer; of the ‘dicot’ type. Tapetum amoeboid, or glandular (rarely). Pollen grains aperturate, or nonaperturate (perhaps, occasionally?); mostly 3 aperturate; colporate (commonly), or colpate, or porate, or zoniaperturate; lophate (notably in most Lactucoideae), or not lophate (usually spinulose); 3-celled.

Gynoecium 2 carpelled. The pistil 1 celled. Gynoecium syncarpous; synovarious to synstylovarious; inferior. Ovary 1 locular. Gynoecium median. Epigynous disk usually present (around the base of the style). Styles 2; partially joined; attenuate from the ovary. Stigmas 2; dry type; papillate; Group II type. Placentation basal. Ovules in the single cavity 1; ascending; non-arillate; anatropous; unitegmic; tenuinucellate. Endothelium differentiated. Embryo-sac development Polygonum-type, or Chrysanthemum-type (or unspecified). Polar nuclei fusing prior to fertilization, or not fusing (when apomictic). Antipodal cells formed; 2, or 3 (sometimes becoming multinucleate); proliferating (rarely, with up to 60 cells), or not proliferating. Embryogeny asterad.

Fruit non-fleshy; indehiscent; a cypsella (almost invariably), or a drupe (occasionally). The drupes with one stone. Dispersal unit the remains of the flower (i.e. the cypsella). Dispersal commonly by wind, via the hairy pappus. Seeds non-endospermic (or endosperm ‘very thin’). Cotyledons 2 (expanded); occasionally rolled (but usually planoconvex). Embryo achlorophyllous (17/21); straight (oily).

Seedling. Germination phanerocotylar.

Physiology, biochemistry. Cyanogenic (rarely), or not cyanogenic. Cynogenic constituents phenylalanine-derived, or of Hegnauer’s ‘Group C’. Polyacetylenes recorded. Alkaloids present, or absent. Iridoids not detected. Arthroquinones detected (3 genera); polyacetate derived. Verbascosides detected (Echinacea). Proanthocyanidins absent (with the exception of Cosmos bipinnatus, with a trace of cyanidin). Flavonols present (mostly), or absent; kaempferol, or quercetin, or kaempferol and quercetin. Ellagic acid practically always absent (absent from 40 species and 28 genera, with a positive record only for Tagetes patula). Arbutin present, or absent. Ursolic acid present. Saponins/sapogenins present, or absent. Aluminium accumulation not found (but many accumulate selenium). Sugars transported as sucrose, or as sugar alcohols + oligosaccharides + sucrose. Inulin recorded (very widespread). C₃, or C₄, or CAM, or C₃-C₄ intermediate. C₃ physiology recorded directly in Acanthospermum, Achillea, Ambrosia, Arctium, Artemisia, Aster, Baccharis, Bahia, Bebbia, Bidens Blainvillea, Centaurea, Ceruana, Chrysactinea, Chrysanthemum, Cirsium, Clappia, Coreocarpus, Coreopsis, Cosmos, Dicoma, Dyssodia, Echinops, Encelia, Erodiophyllum, Eupatorium, Flaveria, Franseria, Galinsoga, Goldmanella, Grangea, Grangeopsis, Grindelia, Guizotia, Gymnolaena, Haplopappus, Helianthus, Heterospermum, Hidalgoa, Hofmeisteria, Hymenoclea, Isocoma, Iva, Lactuca, Lagenophora, Leucactinia, Liatris , Matricaria, Myriactis, Nicolletia, Oligochaeta, Parthenium, Peucephyllum, Plagiocheilus, Pluchia, Pulicaria, Rhynchospermum, Schizotrichia, Solenogyne, Solidago, Sonchus, Stephanomeria, Strotheria, Tagetes, Taraxacum, Thelesperma, Tragopogon, Trichoptilium, Trichospira, Urbinella, Varilla, Vernonia, Xanthium, Zinna. C₄ physiology recorded directly in Chrysanthellum, Eryngiophyllum, Flaveria, Glossocardia, Glossogyne, Isostigma, Pectis. CAM recorded directly in Aeonium, Aster, Kleinia, Notonia, Othonna, Senecio. C₃-C₄ intermediacy in Flaveria (10 species) and Parthenium hysterophorus. Anatomy C₄ type (Chrysanthellum, Flaveria, Parthenium, Pectis), or non-C₄ type (numerous genera).

Geography, cytology. Frigid zone, temperate, sub-tropical, and tropical. Cosmopolitan. X = 2–19(+). Supposed basic chromosome number of family 9.

Taxonomy. Subclass Dicotyledonae; Tenuinucelli. Dahlgren’s Superorder Asteriflorae; Asterales. Cronquist’s Subclass Asteridae; Asterales. APG (1998) Eudicot; core Eudicot; Asterid; Euasterid II; Asterales. Species about 13000. Genera about 1500; Aaronsohnia, Abrotanella, Acamptopappus, Acanthocephalus, Acanthocladium, Acanthodesmos, Acantholepis, Acanthospermum, Acanthostyles, Achillea, Achnophora, Achnopogon, Achyrachaena, Achyrocline, Achyropappus, Achyrothalamus, Acilepidopsis, Acilepis, Acmella, Acomis, Acourtia, Acrisione, Acritopappus, Acroptilon, Actinobole, Actinoseris, Actites, Adelostigma, Adenanthellum, Adenocaulon, Adenocritonia, Adenoglossa, Adenoon, Adenopappus, Adenophyllum, Adenostemma, Adenothamnus, Aedesia, Aegopordon, Aequatorium, Aetheolaena, Aetheorhiza, Ageratella, Ageratina, Ageratinastrum, Ageratum, Agoseris, Agrianthus, Ainsliaea, Ajania, Ajaniopsis, Alatoseta, Albertinia, Alcantara, Alciope, Aldama, Alepidocline, Alfredia, Aliella, Allagopappus, Allardia, Alloispermum, Allopterigeron, Almutaster, Alomia, Alomiella, Alvordia, Amauria, Amberboa, Amblyocarpum, Amblyolepis, Amblyopappus, Amboroa, Ambrosia, Ameghinoa, Amellus, Ammobium, Amolinia, Ampelaster, Amphiachyris, Amphiglossa, Amphipappus, Amphoricarpos, Anacantha, Anacyclus, Anaphalioides, Anaphalis, Anaxeton, Ancathia, Ancistrocarphus, Ancistrophora, Andryala, Angelphytum, Angianthus, Anisochaeta, Anisocoma, Anisopappus, Anisothrix, Antennaria, Anthemis, Antillia, Antiphiona, Antithrixia, Anura, Anvillea, Apalochlamys, Aphanactis, Aphanostephus, Aphyllocladus, Apodocephala, Apopyros, Aposeris, Apostates, Arbelaezaster, Archibaccharis, Arctanthemum, Arctium, Arctogeron, Arctotheca, Arctotis, Argyranthemum, Argyroglottis, Argyrovernonia, Argyroxiphium, Aristeguietia, Arnaldoa, Arnica, Arnicastrum, Arnoglossum, Arnoseris, Arrhenechthites, Arrojadocharis, Arrowsmithia, Artemisia, Artemisiopsis, Asaemia, Asanthus, Ascidiogyne, Aspilia, Asplundianthus, Aster, Asteridea, Asteriscus, Asteromoea, Asteropsis, Asterothamnus, Astranthium, Athanasia, Athrixia, Athroisma, Atractylis, Atractylodes, Atrichantha, Atrichoseris, Austrobrickellia, Austrocritonia, Austroeupatorium, Austrosynotis, Axiniphyllum, Ayapana, Ayapanopsis, Aylacophora, Aynia, Aztecaster, Baccharidopsis, Baccharis, Baccharoides, Badilloa, Baeriopsis, Bafutia, Bahia, Bahianthus, Baileya, Balduina, Balsamorhiza, Baltimora, Barkleyanthus, Barnadesia, Barroetea, Barrosoa, Bartlettia, Bartlettina, Basedowia, Bebbia, Bedfordia, Bejaranoa, Bellida, Bellis, Bellium, Belloa, Berardia, Berkheya, Berlandiera, Berroa, Berylsimpsonia, Bidens, Bigelowia, Bishopalea, Bishopanthus, Bishopiella, Bishovia, Blainvillea, Blakeanthus, Blakiella, Blanchetia, Blennosperma, Blennospora, Blepharipappus, Blepharispermum, Blepharizonia, Blumea, Blumeopsis, Boeberastrum, Boeberoides, Bolanosa, Bolocephalus, Boltonia, Bombycilaena, Borkonstia, Borrichia, Bothriocline, Brachanthemum, Brachionostylum, Brachyactis, Brachyclados, Brachyglottis, Brachylaena, Brachyscome, Brachythrix, Bracteantha, Brickellia, Brickelliastrum, Bryomorphe, Buphthalmum, Burkartia, Cabreriella, Cacalia, Cacaliopsis, Cacosmia, Cadiscus, Caesulia, Calea, Calendula, Callicephalus, Callilepis, Callistephus, Calocephalus, Calomeria, Calostephane, Calotesta, Calotis, Calycadenia, Calycoseris, Calyptocarpus, Camchaya, Campovassouria, Camptacra, Campuloclinium, Canadanthus, Cancrinia, Cancriniella, Cardopatium, Carduncellus, Carduus, Carlina, Carminatia, Carpesium, Carphephorus, Carphochaete, Carramboa, Carterothamnus, Carthamus, Cassinia, Castanedia, Castrilanthemum, Catamixis, Catananche, Catatia, Cavalcantia, Cavea, Celmisia, Centaurea, Centaurodendron, Centauropsis, Centaurothamnus, Centipeda, Centratherum, Cephalipterum, Cephalopappus, Cephalorrhynchus, Cephalosorus, Ceratogyne, Ceruana, Chacoa, Chaenactis, Chaetadelpha, Chaetanthera, Chaetopappa, Chaetoseris, Chamaechaenactis, Chamaegeron, Chamaemelum, Chamaepus, Chaptalia, Chardinia, Cheirolophus, Chersodoma, Chevreulia, Chiliadenus, Chiliocephalum, Chiliophyllum, Chiliotrichiopsis, Chiliotrichum, Chimantaea, Chionolaena, Chionopappus, Chlamydophora, Chloracantha, Chondrilla, Chondropyxis, Chresta, Chromolaena, Chromolepis, Chronopappus, Chrysactinia, Chrysactinium, Chrysanthellum, Chrysanthemoides, Chrysanthemum, Chrysanthoglossum, Chrysocephalum, Chrysocoma, Chrysogonum, Chrysolaena, Chrysoma, Chrysophthalmum, Chrysopsis, Chrysothamnus, Chthonocephalus, Chucoa, Chuquiraga, Cicerbita, Ciceronia, Cichorium, Cineraria, Cirsium, Cissampelopsis, Cladanthus, Cladochaeta, Clappia, Clibadium, Cnicothamnus, Cnicus, Coespeletia, Coleocoma, Coleostephus, Colobanthera, Columbiadoria, Comaclinium, Comborhiza, Commidendrum, Complaya, Condylidium, Condylopodium, Conocliniopsis, Conoclinium, Conyza, Coreocarpus, Coreopsis, Corethamnium, Correllia, Corymbium, Cosmos, Cotula, Coulterella, Cousinia, Cousiniopsis, Craspedia, Crassocephalum, Cratystylis, Cremanthodium, Crepidiastrum, Crepis, Critonia, Critoniadelphus, Critoniella, Critoniopsis, Crocidium, Cronquistia, Cronquistianthus, Croptilon, Crossostephium, Crossothamnus, Crupina, Cuatrecasanthus, Cuatrecasasiella, Cuchumatanea, Cullumia, Cuspidia, Cyanthillium, Cyathocline, Cyathomone, Cyclolepis, Cylindrocline, Cymbolaena, Cymbonotus, Cymbopappus, Cynara, Cyrtocymura, Dacryotrichia, Dahlia, Damnamenia, Damnxanthodium, Dasycondylus, Dasyphyllum, Daveaua, Decachaeta, Decastylocarpus, Decazesia, Delairea, Delamerea, Delilia, Dendranthema, Dendrocacalia, Dendrophorbium, Dendrosenecio, Dendroseris, Denekia, Desmanthodium, Dewildemania, Diacranthera, Dianthoseris, Diaphractanthus, Diaspananthus, Dicercoclados, Dichaetophora, Dichrocephala, Dichromochlamys, Dicoma, Dicoria, Dicranocarpus, Didelta, Dielitzia, Digitacalia, Dimeresia, Dimerostemma, Dimorphocoma, Dimorphotheca, Dinoseris, Diodontium, Diplazoptilon, Diplostephium, Dipterocome, Dipterocypsela, Disparago, Dissothrix, Distephanus, Disynaphia, Dithyrostegia, Dittrichia, Doellingeria, Dolichoglottis, Dolichorrhiza, Dolichothrix, Dolomiaea, Doniophyton, Dorobaea, Doronicum, Dracopis, Dresslerothamnus, Dubautia, Dubyaea, Dugesia, Duhaldea, Duidaea, Duseniella, Dymondia, Dyscritogyne, Dyscritothamnus, Dysodiopsis, Dyssodia, Eastwoodia, Eatonella, Echinacea, Echinocoryne, Echinops, Eclipta, Edmondia, Egletes, Eirmocephala, Eitenia, Ekmania, Elachanthus, Elaphandra, Elephantopus, Eleutheranthera, Ellenbergia, Elytropappus, Embergeria, Emilia, Emiliella, Encelia, Enceliopsis, Endocellion, Endopappus, Engelmannia, Engleria, Enydra, Epaltes, Epilasia, Episcothamnus, Epitriche, Erato, Erechtites, Eremanthus, Eremosis, Eremothamnus, Eriachaenium, Ericameria, Ericentrodea, Erigeron, Eriocephalus, Eriochlamys, Eriophyllum, Eriothrix, Erlangea, Erodiophyllum, Erymophyllum, Eryngiophyllum, Erythradenia, Erythrocephalum, Espejoa, Espeletia, Espeletiopsis, Ethulia, Eucephalus, Euchiton, Eumorphia, Eupatoriastrum, Eupatorina, Eupatoriopsis, Eupatorium, Euphrosyne, Eurybiopsis, Eurydochus, Euryops, Eutetras, Euthamia, Evacidium, Ewartia, Ewartiothamnus, Exomiocarpon, Facelis, Farfugium, Faujasia, Faxonia, Feddea, Feldstonia, Felicia, Femeniasia, Fenixia, Ferreyranthus, Ferreyrella, Filago, Filifolium, Fitchia, Fitzwillia, Flaveria, Fleischmannia, Fleischmanniopsis, Florestina, Floscaldasia, Flosmutisia, Flourensia, Flyriella, Formania, Foveolina, Freya, Fulcaldea, Gaillardia, Galactites, Galeana, Galeomma, Galinsoga, Gamochaeta, Gamochaetopsis, Garberia, Garcibarrigoa, Garcilassa, Gardnerina, Garhadiolus, Garuleum, Gazania, Geigeria, Geissolepis, Geraea, Gerbera, Geropogon, Gibbaria, Gilberta, Gilruthia, Gladiopappus, Glaziovianthus, Glossarion, Glossocardia, Glossopappus, Glyptopleura, Gnaphaliothamnus, Gnaphalium, Gnephosis, Gochnatia, Goldmanella, Gongrostylus, Gongylolepis, Goniocaulon, Gonospermum, Gorceixia, Gorteria, Gossweilera, Goyazianthus, Grangea, Grangeopsis, Graphistylis, Gratwickia, Grauanthus, Grazielia, Greenmaniella, Grindelia, Grisebachianthus, Grosvenoria, Guardiola, Guayania, Guevaria, Guizotia, Gundelia, Gundlachia, Gutierrezia, Gymnanthemum, Gymnarrhena, Gymnocondylus, Gymnocoronis, Gymnodiscus, Gymnolaena, Gymnopentzia, Gymnosperma, Gymnostephium, Gynoxys, Gynura, Gypothamnium, Gyptidium, Gyptis, Gyrodoma, Haastia, Haeckeria, Haegiela, Handelia, Haplocarpha, Haploesthes, Haplopappus, Haplostephium, Harleya, Harnackia, Hartwrightia, Hasteola, Hatschbachiella, Hazardia, Hebeclinium, Hecastocleis, Hedypnois, Helenium, Helianthella, Helianthus, Helichrysopsis, Helichrysum, Heliocauta, Heliomeris, Heliopsis, Helipterum, Helminthotheca, Helogyne, Hemisteptia, Hemizonia, Henricksonia, Heptanthus, Herderia, Herodotia, Herrickia, Hesperevax, Hesperodoria, Hesperomannia, Heteracia, Heteranthemis, Heterocoma, Heterocondylus, Heterocypsela, Heteroderis, Heterolepis, Heteromera, Heteromma, Heteropappus, Heteroplexis, Heterorhachis, Heterosperma, Heterothalamus, Heterotheca, Hidalgoa, Hieracium, Hilliardia, Hinterhubera, Hippia, Hippolytia, Hirpicium, Hispidella, Hochstetteria, Hoehnephytum, Hoffmanniella, Hofmeisteria, Holocarpha, Holocheilus, Hololeion, Hololepis, Holozonia, Homognaphalium, Homogyne, Hoplophyllum, Huarpea, Hubertia, Hughesia, Hulsea, Humeocline, Hyalis, Hyalochaete, Hyalochlamys, Hyaloseris, Hyalosperma, Hybridella, Hydroidea, Hydropectis, Hymenocephalus, Hymenoclea, Hymenolepis, Hymenonema, Hymenopappus, Hymenostemma, Hymenothrix, Hymenoxys, Hyoseris, Hypacanthium, Hypericophyllum, Hypochaeris, Hysterionica, Hystrichophora, Ichthyothere, Idiothamnus, Ifloga, Ighermia, Iltisia, Imeria, Inezia, Inula, Inulanthera, Inulopsis, Iocenes, Iodocephalus, Iogeton, Ionactis, Iostephane, Iotasperma, Iphiona, Iphionopsis, Iranecio, Irwinia, Ischnea, Ismelia, Isocarpha, Isocoma, Isoetopsis, Isostigma, Iva, Ixeridium, Ixeris, Ixiochlamys, Ixiolaena, Ixodia, Jacmaia, Jaegeria, Jalcophila, Jaliscoa, Jamesianthus, Jaramilloa, Jasonia, Jaumea, Jefea, Jeffreya, Jessea, Joseanthus, Jungia, Jurinea, Jurinella, Kalimeris, Karelinia, Karvandarina, Kaschgaria, Kaunia, Keysseria, Kinghamia, Kingianthus, Kippistia, Kirkianella, Kleinia, Koanophyllon, Koehneola, Koelpinia, Krigia, Kyrsteniopsis, Lachanodes, Lachnophyllum, Lachnorhiza, Lachnospermum, Lactacella, Lactuca, Lactucella, Lactucosonchus, Laennecia, Laestadia, Lagascea, Lagedium, Lagenithrix, Lagenophora, Laggera, Lagophylla, Lamprachaenium, Lamprocephalus, Lamyropappus, Lamyropsis, Langebergia, Lantanopsis, Lapsana, Lapsanastrum, Lasianthaea, Lasiocephalus, Lasiolaena, Lasiopogon, Lasiospermum, Lasthenia, Launaea, Lawrencella, Layia, Lecocarpus, Leibnitzia, Leiboldia, Lembertia, Lemoorea, Leontodon, Leontopodium, Lepidaploa, Lepidesmia, Lepidolopha, Lepidolopsis, Lepidonia, Lepidophorum, Lepidophyllum, Lepidospartum, Lepidostephium, Leptinella, Leptocarpha, Leptoclinium, Leptorhynchos, Leptostelma, Lescaillea, Lessingia, Lessingianthus, Leucactinia, Leucanthemella, Leucanthemopsis, Leucanthemum, Leucheria, Leucoblepharis, Leucocyclus, Leucogenes, Leucomeris, Leucophyta, Leucoptera, Leunisia, Leuzea, Leysera, Liabellum, Liabum, Liatris, Libanothamnus, Lidbeckia, Lifago, Ligularia, Limbarda, Lindheimera, Lipochaeta, Litogyne, Litothamnus, Litrisa, Llerasia, Logfia, Lomatozona, Lonas, Lopholaena, Lophopappus, Lordhowea, Lorentzianthus, Loricaria, Lourteigia, Loxothysanus, Lucilia, Luciliocline, Lugoa, Luina, Lulia, Lundellianthus, Lycapsus, Lychnophora, Lycoseris, Lygodesmia, Machaeranthera, Macowania, Macrachaenium, Macraea, Macroclinidium, Macropodina, Macvaughiella, Madagaster, Madia, Mairia, Malacothrix, Mallotopus, Malmeanthus, Malperia, Mantisalca, Marasmodes, Marshallia, Marshalljohnstonia, Marticorenia, Matricaria, Mattfeldanthus, Mattfeldia, Matudina, Mauranthemum, Mausolea, Mecomischus, Megalodonta, Melampodium, Melanodendron, Melanthera, Merrittia, Metalasia, Metastevia, Mexerion, Mexianthus, Micractis, Microcephala, Microglossa, Microgynella, Microliabum, Micropsis, Micropus, Microseris, Microspermum, Mikania, Mikaniopsis, Milleria, Millotia, Minuria, Miricacalia, Miyamayomena, Mniodes, Monactis, Monarrhenus, Monogereion, Monolopia, Monoptilon, Montanoa, Monticalia, Moonia, Moquinia, Morithamnus, Moscharia, Msuata, Mulgedium, Munnozia, Munzothamnus, Muschleria, Mutisia, Mycelis, Myopordon, Myriactis, Myriocephalus, Myripnois, Myxopappus, Nabalus, Nananthea, Nannoglottis, Nanothamnus, Nardophyllum, Narvalina, Nassauvia, Nauplius, Neblinaea, Neja, Nelsonianthus, Nemosenecio, Neocabreria, Neocuatrecasia, Neohintonia, Neojeffreya, Neomirandea, Neomolina, Neopallasia, Neotysonia, Nesomia, Nestlera, Neurolaena, Neurolakis, Nicolasia, Nicolletia, Nidorella, Nikitinia, Nipponanthemum, Nivellea, Nolletia, Nothobaccharis, Nothocalais, Noticastrum, Notobasis, Notoseris, Nouelia, Novenia, Oaxacania, Oblivia, Ochrocephala, Oclemena, Odixia, Odontocline, Oedera, Oiospermum, Oldenburgia, Olearia, Olgaea, Oligactis, Oliganthes, Oligocarpus, Oligochaeta, Oligoneuron, Oligothrix, Olivaea, Omalotheca, Omphalopappus, Oncosiphon, Ondetia, Onopordum, Onoseris, Oonopsis, Oparanthus, Ophryosporus, Opisthopappus, Oreochrysum, Oreoleysera, Oreostemma, Oritrophium, Orochaenactis, Osbertia, Osmadenia, Osmiopsis, Osmitopsis, Osteospermum, Otanthus, Oteiza, Othonna, Otopappus, Otospermum, Outreya, Oxycarpha, Oxylaena, Oxylobus, Oxypappus, Oxyphyllum, Oyedaea, Ozothamnus, Pachylaena, Pachystegia, Pachythamnus, Pacifigeron, Packera, Pacourina, Palafoxia, Paleaepappus, Pamphalea, Pappobolus, Pappochroma, Paracalia, Parachionolaena, Paragynoxys, Paralychnophora, Paranephelius, Parantennaria, Parapiqueria, Paraprenanthes, Parasenecio, Parastrephia, Parthenice, Parthenium, Pasaccardoa, Pechuel-Loeschea, Pectis, Pegolettia, Pelucha, Pentacalia, Pentachaeta, Pentanema, Pentatrichia, Pentzia, Perdicium, Perezia, Pericallis, Pericome, Peripleura, Perityle, Perralderia, Pertya, Perymeniopsis, Perymenium, Petalacte, Petasites, Peteravenia, Petradoria, Petrobium, Peucephyllum, Phacellothrix, Phaenocoma, Phaeostigma, Phagnalon, Phalacrachena, Phalacraea, Phalacrocarpum, Phalacroseris, Phaneroglossa, Phanerostylis, Phania, Philactis, Philoglossa, Philyrophyllum, Phoebanthus, Phyllocephalum, Phymaspermum, Picnomon, Picradeniopsis, Picris, Picrosia, Picrothamnus, Pilosella, Pilostemon, Pinaropappus, Pingraea, Pinillosia, Piora, Pippenalia, Piptocarpha, Piptocoma, Piptolepis, Piptothrix, Piqueria, Piqueriella, Piqueriopsis, Pithecoseris, Pithocarpa, Pittocaulon, Pityopsis, Pladaroxylon, Plagiobasis, Plagiocheilus, Plagiolophus, Plagius, Planaltoa, Planea, Plateilema, Platycarpha, Platypodanthera, Platyschkuhria, Plazia, Plecostachys, Plectocephalus, Pleiotaxis, Pleocarphus, Pleurocarpaea, Pleurocoronis, Pleurophyllum, Pluchea, Podachaenium, Podanthus, Podocoma, Podolepis, Podotheca, Poecilolepis, Pogonolepis, Pojarkovia, Pollalesta, Polyachyrus, Polyanthina, Polyarrhena, Polycalymma, Polychrysum, Polymnia, Polytaxis, Porophyllum, Porphyrostemma, Praxeliopsis, Praxelis, Prenanthella, Prenanthes, Printzia, Prionopsis, Prolobus, Prolongoa, Proteopsis, Proustia, Psacaliopsis, Psacalium, Psathyrotes, Psathyrotopsis, Psednotrichia, Pseudelephantopus, Pseudobahia, Pseudoblepharisper, Pseudobrickellia, Pseudocadiscus, Pseudoclappia, Pseudoconyza, Pseudognaphalium, Pseudogynoxys, Pseudohandelia, Pseudojacobaea, Pseudokyrsteniopsi, Pseudoligandra, Pseudonoseris, Pseudostifftia, Psiadia, Psiadiella, Psilactis, Psilocarphus, Psilostrophe, Psychrogeton, Psychrophyton, Pterachaenia, Pterocaulon, Pterocypsela, Pteronia, Pterothrix, Pterygopappus, Ptilostemon, Pulicaria, Pycnocephalum, Pyrrhopappus, Pyrrocoma, Pytinicarpa, Quelchia, Quinetia, Quinqueremulus, Radlkoferotoma, Rafinesquia, Raillardella, Raillardiopsis, Rainiera, Raoulia, Raouliopsis, Rastrophyllum, Ratibida, Raulinoreitzia, Rayjacksonia, Reichardia, Relhania, Remya, Rennera, Rensonia, Revealia, Rhagadiolus, Rhamphogyne, Rhanteriopsis, Rhanterium, Rhetinolepis, Rhodanthe, Rhodanthemum, Rhodanthemum, Rhynchopsidium, Rhynchospermum, Rhysolepis, Richteria, Riencourtia, Rigiopappus, Robinsonecio, Robinsonia, Rochonia, Rojasianthe, Rolandra, Roldana, Rosenia, Rothmaleria, Rudbeckia, Rugelia, Ruilopezia, Rumfordia, Russowia, Rutidosis, Sabazia, Sachsia, Salmea, Santolina, Santosia, Sanvitalia, Sarcanthemum, Sartorina, Sartwellia, Saussurea, Scalesia, Scariola, Scherya, Schischkinia, Schistocarpha, Schistostephium, Schizogyne, Schizoptera, Schizotrichia, Schkuhria, Schlechtendalia, Schmalhausenia, Schoenia, Sciadocephala, Sclerocarpus, Sclerolepis, Sclerorhachis, Sclerostephane, Scolymus, Scorzonera, Scrobicaria, Selloa, Senecio, Sericocarpus, Seriphidium, Serratula, Shafera, Sheareria, Shinnersia, Shinnersoseris, Siapaea, Siebera, Sigesbeckia, Siloxerus, Silphium, Silybum, Simsia, Sinacalia, Sinclairia, Sinoleontopodium, Sinosenecio, Sipolisia, Smallanthus, Soaresia, Solanecio, Solenogyne, Solidago, Soliva, Sommerfeltia, Sonchus, Sondottia, Soroseris, Spaniopappus, Sparganophorus, Sphaeranthus, Sphaereupatorium, Sphaeromeria, Sphagneticola, Spilanthes, Spiracantha, Spiroseris, Squamopappus, Stachycephalum, Staehelina, Standleyanthus, Staurochlamys, Stebbinsoseris, Steiractinia, Steirodiscus, Stemmacantha, Stenachaenium, Stenocephalum, Stenocline, Stenopadus, Stenophalium, Stenops, Stenoseris, Stenotus, Stephanochilus, Stephanodoria, Stephanomeria, Steptorhamphus, Stevia, Steviopsis, Steyermarkina, Stifftia, Stilpnogyne, Stilpnolepis, Stilpnopappus, Stoebe, Stokesia, Stomatanthes, Stomatochaeta, Stramentopappus, Streptoglossa, Strotheria, Stuartina, Stuckertiella, Stuessya, Stylocline, Stylotrichium, Sventenia, Symphyllocarpus, Symphyopappus, Symphyotrichum, Syncalathium, Syncarpha, Syncephalum, Syncretocarpus, Synedrella, Synedrellopsis, Syneilesis, Synotis, Syntrichopappus, Synurus, Syreitschikovia, Taeckholmia, Tagetes, Takeikadzuchia, Takhtajaniantha, Talamancalia, Tamananthus, Tamania, Tamaulipa, Tanacetopsis, Tanacetum, Taplinia, Taraxacum, Tarchonanthus, Tehuana, Teixeiranthus, Telanthophora, Telekia, Telmatophila, Tenrhynea, Tephroseris, Tessaria, Tetrachyron, Tetradymia, Tetragonotheca, Tetramolopium, Tetraneuris, Tetranthus, Tetraperone, Thaminophyllum, Thamnoseris, Thelesperma, Thespidium, Thespis, Thevenotia, Thiseltonia, Thurovia, Thymophylla, Thymopsis, Tiarocarpus, Tietkensia, Tithonia, Tolbonia, Tolpis, Tomentaurum, Tonestus, Tourneuxia, Townsendia, Tracyina, Tragopogon, Traversia, Trichanthemis, Trichanthodium, Trichocline, Trichocoronis, Trichocoryne, Trichogonia, Trichogoniopsis, Trichogyne, Tricholepis, Trichoptilium, Trichospira, Tridactylina, Tridax, Trigonospermum, Trilisa, Trimorpha, Trioncinia, Tripleurospermum, Triplocephalum, Tripteris, Triptilion, Triptilodiscus, Trixis, Troglophyton, Tuberostylis, Tugarinovia, Turaniphytum, Tussilago, Tuxtla, Tyleropappus, Tyrimnus, Uechtritzia, Ugamia, Uleophytum, Unxia, Urbananthus, Urbinella, Urmenetea, Urolepis, Uropappus, Urospermum, Ursinia, Vanclevea, Varilla, Varthemia, Vellereophyton, Venegasia, Verbesina, Vernonia, Vernoniopsis, Viereckia, Vieria, Vigethia, Viguiera, Villanova, Vilobia, Vittadinia, Vittetia, Volutaria, Waitzia, Wamalchitamia, Warionia, Wedelia, Welwitschiella, Wendelboa, Werneria, Westoniella, Whitneya, Wilkesia, Willemetia, Wollastonia, Wulffia, Wunderlichia, Wyethia, Xanthisma, Xanthium, Xanthocephalum, Xanthopappus, Xeranthemum, Xerolekia, Xylanthemum, Xylorhiza, Xylothamia, Yermo, Youngia, Zaluzania, Zandera, Zexmenia, Zinnia, Zoegea, Zyrphelis, Zyzyxia.

Heywood, Harborne and Turner 1977.

Economic uses, etc. Sources of foodstuffs include Lactuca (lettuce), Cynara (globe artichoke), Cichorium (chicory and endive), Tragopogon (salsify). Insecticides from Pyrethrum, safflower dye from Carthamus. At least 200 genera are widely planted as ornamentals; and some are notorious hayfever plants (‘ragweed’ (Ambrosia), etc.).

Illustrations. • Lactuceae. • Lactuceae. • Lactuceae. • Cynareae. • Cynareae (thistles). • Eupatorieae, Heliantheae, Inuleae. • Inuleae. • Senecioneae. • Anthemideae. • Anthemideae. • Anthemideae. • Astereae. • `Cineraria lanata'. • Centaurea glastifolia. • Technical details (Taraxacum). • Technical details (Tragopogon, Cichorium, Nassauvia). • Technical details (Centaurea cyanus). • Technical details (`Marigold' and `Thistle'). • Technical details (various genera). • Technical details (various genera). • Technical details (Vernonia).

Quotations

Wele by reason men it call may
The dayeseye or ellis the eye of day
(Chaucer, ‘Legend of Good Women’, Prologue (c. 1385))

Without the bed her other fair hand was
On the green coverlet, whose perfect white
Showed like an April daisy on the grass
(‘Rape of Lucrece’)

Your pleasures spring like daisies in the grass,
Cut down, and up again as blithe as ever
(Walter Savage Landor, ‘Ianthe’)

The marigold, that goes to bed wi’ the sun,
And with him rises weeping.
(‘The Winter’s Tale’ - Calendula officinalis)

Her eyes, like marigolds, had sheathed their light,
And canopied in darkness sweetly lay,
‘Til they might open to adorn the day
(‘Rape of Lucrece’ - Calendula officinalis)

And winking marybuds begin
To ope their golden eyes
(‘Cymbeline’, ii. 3 - Calendula officinalis (marigold) buds)

Nay, friar, I am a kind of bur, I stick
(‘Measure for Measure’, iv. 3 - Arctium)

Ragwort thou humble flower with tattered leaves
I love to see thee come and litter gold
(John Clare c.1835, ‘The Ragwort’ — Senecio jacobaea)

And iron-weed, content to share
The meanest spot that spring can spare:
E’en roads where danger hourly comes
Are not without its purple blooms;
(John Clare, quoted by Ann Pratt 1857 — of Centaurea scabiosa)

Then did we question of the down-balls, blowing
To know if some slight wish would come to pass
(Quoted by Ann Pratt, ‘Wild Flowers’ (1857), unattributed - Taraxacum)

If the down flyeth off Coltsfoot, Dandelion, and Thistle, when there is no winde, it is a signe of rain
(Coles, quoted by Ann Pratt, ‘Wild Flowers’ (1857))

-The remaining quotations contributed by N.S. Lander:

‘They are nice flowers’, he said, her emotional tones putting a constraint on him.
‘You know that a daisy is a company of florets, a concourse, become individual. Don’t the botanists put it highest in the line of development? I believe they do.’
‘The compositae, yes, I think so’, said Ursula . . .
‘Explain it so, then’, he said. ‘The daisy is a perfect little democracy, so it’s the highest of flowers, hence its charm.’
‘No,’ she cried, ‘no - never. It isn’t democratic.’
‘No’, he admitted. ‘It’s the golden mob of the proletariat surrounded by a showy white fence of the idle rich.’
‘How hateful - your hateful social orders!’ she cried.
‘Quite! It’s a daisy - we’ll leave it alone.’
(David H. Lawrence, ‘Women in Love’ (1920)

The word is ‘Day’s eye’ (O.E. ‘daeges eage’), and the flower is so called because it closes its pinky lashes and goes to sleep when the sun sets, but in the morning expands its petals to the light.
(Ebenezer Cobham Brewer, revised by Ivor H. Evans, ‘Brewer’s Dictionary of Phrase and Fable’, Centenary Edn (1970))

Daisy. An emblem of deceit. Greene (‘Quip for an Upstart Courtier’) speaks of the ‘dissembling daisie’. ‘Light of love wenches’ are warned by it ‘not to trust every fair promise that such amorous bachelors make them’. Ophelia in Shakespeare’s ‘Hamlet’ gives the queen a daisy to signify, ‘that her light and fickle love ought not to expect constancy in her husband’.

Daisies, those pearled Arcturi of the earth,
The constellated flower that never sets.
(Percy Bysshe Shelley, ‘The Question’)

Ennewed your colour
Is like the daisy flower
After the April shower.
(John Skelton 1460?-1529, ‘To Mistress Isabell Pennell’)

When you can tread on nine daisies at once, spring has come.
(Proverb, 19th century)

When you can put your foot on seven daisies summer is come.
(Proverb, quoted in M. Grieve, ‘A Modern Herbal’ (1931) 247)

Of all the floures in the meade,
Thanne love I most those floures white and rede,
Such as men callen daysyes.
Chaucer, ‘Legend of Good Women’, Prol. 43 (c. 1385))

Depeinted wonderly,
With many a thousand daisies, red as rose,
And white also.
(Chaucer, ‘The Court of Love’, xv (c.1450))

Let us Finde out the prettiest Daisied Plot we can.
(‘Cymbeline’ IV. ii. 398 (1611))

‘Pushing up daisies’ is their creed, you know.
(Wilfred Owen, ‘Poems’ 65 (1963))

I think she’s drinking herself under the daisies, so to speak.
(S. Vines, ‘Humours Unreconciled’ xxi, 268 (1928))

Where is the wife now? . . .
Pushing up daisies . . .
(G. Heyer, ‘Blunt Instrument’ xiii. 252 (1938))

Daisy roots.
Rhyming slang, boots.
(‘Oxford English Dictionary, Supplement’)

While waiting for my pal I had my daisies cleaned.
(‘Macm. Magazine’ XL, 503/1 (1879))

The umpires called play, Grange being again on the defensive to the under-hand ‘daisy cutters’ of Sadler.
(‘Bell’s Life’ I, Nov. 7/1 (1858))

The ball that kept low had to be a daisy-cutter although a first-class cricket pitch has surely been sufficiently mown to lose all likeness to a flowery meadow.
(‘Times’, 13 June 13/3 (1963))

So courteous, so kind,
As Merry Margaret, the midsummer flower,
Gentle as falcon or hawk of the tower.
(John Skelton (1460?-1529, ‘To Mistress Margaret Hussey’ - Margaret = marguerite)

‘Kiss-me-and-I’ll-tell-you’ replied an attractive native of the Southern States when asked the name of that plant which people of colder climes know as the daisy fleabane.
(W.N Clute, ‘Common Names of Plants’ 131 (1931))

Wormwood. It is said to have been so called because this plant, according to legend, sprang up in the track of the serpent as it writhed along the ground when driven out of Paradise. The word is also used figuratively to dentote bitterness or its cause.
(Ebenezer Cobham Brewer, revised by Ivor H. Evans, ‘Brewer’s Dictionary of Phrase and Fable’, Centenary Edn (1970) - Artemesia)

And the third angel sounded and there fell a great star from heaven, burning as it were a lamp, and it fell upon the third part of the rivers, and upon the fountains of waters.

And the name of the star is called Wormwood: and the third part of the waters became wormwood: and many men died of the waters, because they were made bitter.
(Bible, ‘Revelation’, ch. 8, vv. 10–11 - Artemisia

This riddle, Cuddy, if thou can’st explain . . .
What flower is that which bears the Virgin’s name,
The richest metal joined with the same?
(John Gay, ‘The Shepherd’s Week’, Monday - Calendula officinalis; 17th-century ‘Marygold’ (marigold) = a sovereign.)

I’ll . . . presently go put five hundred Marygolds in a Purse for you.
(Cowley, ‘Cutter of Coleman-st’ iII. iii (1663) - Calendula officinalis; 17th-century ‘Marygold’ (marigold) = a sovereign.)

Thistle. The heraldic emblem of Scotland which seems to have been adopted by James II (1451, 1460–1488), possibly as a symbol of defence. The motto ‘Nemo me impune lacessit, ‘Nobody touiches (or provokes) me with impunity’, first appeared on the coinage of James VI (1566, 1567–1603).

Thistles, especially ‘Our Lady’s Thistle’, are said to be a cure for stich in the side. According to the doctrine of signatures, Nature has labelled every plant, and the prickles of the thistle tell us that the plant is efficacious for ‘prickles’ or the stich. The species called Silybum marianum, we are told, owes the white markings on its leaves to milk from Our Lady’s breast, some of which fell thereon and left a white mark behind.
(Ebenezer Cobham Brewer, revised by Ivor H. Evans, ‘Brewer’s Dictionary of Phrase and Fable’, Centenary Edn (1970))

You can keep you litter and your hay and your oats. Long live the
thistles of the field, for there you can play the stallion to your
heart’s content.
(François Rabelais (1494?-1553?), ‘Pantagruel’, IV. 7)

He that hath a good harvest may be content with some thistles.
(Proverb, J. Clarke, ‘Paroemiologia Anglo-Latina’ (1639))

Doubts are like the thistle, a bad weed, but growing in good ground.
Baxter, ‘Saint’s R.’, L. vii. (1650))

If the brain sows not corn, it plants thistles.
(Proverb, G. Herbert, ‘Outlandish Proverbs’ (1640), 2nd edn entitled ‘Jacula Prudentum’ (1651))

Get you some of this distilled Carduus Benedictus and lay it to your heart; it is the only thing for a qualm. . . . I mean plain Holy Thistle.
(‘Much Ado about Nothing’ - Cnicus benedictus)

Like a yonge artichocke that always carries pepper and salt, in itselfe.
(Ben Jonson, ‘Ev. blan. in Humm.’ iv. ii. (1598))

Ah, Sun-flower! weary of time,
Who counteth the steps of the Sun,
Seeking after that sweet golden clime,
Where the traveller’s journey is done;
(William Blake (1757–1827), ‘Ah, Sun-Flower!’ - Helianthus)

Cite this publication as: ‘L. Watson and M. J. Dallwitz (1992 onwards). The Families of Flowering Plants: Descriptions, Illustrations, Identification, and Information Retrieval. Version: 14th December 2000. http://biodiversity.uno.edu/delta/’. Dallwitz (1980), Dallwitz, Paine and Zurcher (1993, 1995, 2000), and Watson and Dallwitz (1991) should also be cited (see References).

Index