PBIO 250 Lecture Notes

James L. Reveal

Norton-Brown Herbarium, University of Maryland

Selected Families of Angiosperms: Asteridae

Gentianales Lindl., 1833

Gentianaceae Juss., 1789: Herbs with opposite astipulate basally connate leaves and 4- or 5-merous flowers bearing 4-5 stamens alternating with the petals, and a bicarpellate superior ovary. CA(4-5) [CO(4-5) A4-5] GS(2). 80 genera, 1200 species. Cosmopolitan. Some are cultivated as ornamentals (Exacum, Gentiana, Sabatia), a few for their medicinal properties (Canscora, Centaurium, Gentiana). The family is divided into approximately four tribes: Potalieae Endl., Gentianeae Bercht. & J. Presl, Helieae Gilg and Voyrieae Gilg. The first is sometimes recognized as a distnict family, Potaliaceae Mart. (3 genera and 70 species) and in that case, the name Lisianthieae Griseb. becomes the correct name for the first tribe. Full description from Watson & Dallwitz; family synonymy from Reveal; list of genera from Kew; family synopses from the University of Hawaii and Utah State.

Apocynaceae Juss., 1789: Trees, shrubs or herbs mostly with milky sap, opposite or whorled astipulate leaves and 5-merous flowers producing a follicle with tufted seeds. CA(5) [CO(5) A5] GS(2). 168 genera, 2100 species. Mostly tropical. Many are cultivated as ornamentals and medicinals, a few are used for timber. Strophanthus is used in the treatment of heart disease and a precursor of cortisone. Vinca is used to treat childhood leukemia. Several are poisonous. Nerium is oleander. Two subfamilies are usually recognized: Rauvolfioideae Kostel. (including Plumerioideae Leurss.) and Apocynoideae Burnett. Some taxonomist distinguished as Plumeriaceae (e.g., Hutchinson) as a distinct family. Most current workers now include Asclepiadaceae in Apocynaceae. Full description from Watson & Dallwitz; family synonymy from Reveal; list of genera from Kew; family synopses from the University of Hawaii and Utah State.

Asclepiadaceae R. Br., 1810: Herbs, shrubs or rarely trees with milky sap, opposite or whorled astipulate leaves and 5-merous flowers with a distinct corona, pollinia, translators and corpuscula producing a follicle with tufted seeds. CA5 [CO(5) [A5] GS2)]. 315 genera, 2900 species. Cosmopolitan but mainly in the tropics. Many cultivated as ornamentals (Stapelia), many weedy. Divided into three subfamilies, Periplocoideae Kostel., Secamonoideae Endl. and Asclepiadoideae Burnett. The first is occasionally accepted as a distinct family Plocospermataceae. Most workers now place Asclepiadaceae in the Apocynaceae. Full description from Watson & Dallwitz; family synonymy from Reveal; list of genera from Kew; family synopses from the University of Hawaii and Utah State. See also Cardillo & Samuels.

Solanales Dumort., 1829

Solanaceae Juss., 1789: Herbs, shrubs, vines or trees with alternate, astipulate leaves, actinomorphic 5-merous flowers with 5 stamens and a 2-locular ovary producing a berry or capsule. CA(5) [CO(5)A5] GS(2). 96 genera, 2900 species. Cosmopolitan but especially in South America where 56 genera (25 endemic) are found. Many food, spice, medicinal, hallucinogenic, insecticidal and ornamental plants. Includes the tomato, potato, chili and bell peppers, atropa (atropine), datura (hyoscine) and tobacco (nicotine). The famous mandrake is a member of this family. Commonly divided into three subfamilies: Browallioideae Kostel. (including the Cestroideae Burnett), Solanoideae Kostel. and Nolanoideae Kostel. with the latter occasionally accepted as the Nolanaceae (Nolana). Goetzeaceae and Sclerophylacaceae are often removed although these were included by Cronquist. Full description from Watson & Dallwitz; family synonymy from Reveal; list of genera from Kew; family synopsis from the University of Hawaii and Utah State. See also Cardillo & Samuels.

Convolvulaceae Juss., 1789: Herbs, shrubs or trees with the sap somewhat milky, alternate leaves, 5-merous flowers and a contorted corolla with 5 epipetalous stamens and a 2-carpellate ovary with 1-2 ovules. CA5 [CO(5) A5] GS(2). 58 genera, 1650 species. Cosmopolitan. Ipomoea, Calystegia and Convolvulus are frequently cultivated as ornamentals. The sweet potato, Ipomoea batatas, is a member of this family. Also in Ipomoea (a genus of some 500 species) is I. violacea, known as "heavenly blue" which is hallucinogenic as it contains lysergic acid (LSD). Three subfamilies are recognized: Convolvuloideae Burnett, Humbertioideae and Dichondroideae Endl. The latter two are rather distinct from the bulk of the family, and therefore Dichondraceae Dumort. and Humbertiaceae Pichon are sometimes recognized. Full description from Watson & Dallwitz; family synonymy from Reveal; list of genera from Kew; family synopses from the University of Hawaii and Utah State.

Cuscutaceae Dumort., 1829: Achlorophyllous threadlike, leafless twining parasitic herbs with small flowers in stalked or sessile bractless clusters producing a dry or fleshy capsule. CA4-5 or (4-5) CO(4-5) A4-5 GS(2). 1 genus, 170 species. Widespread. A plant with a short-lived root system, soon wholly parasitic and often brightly orange-colored with the stems attached to the host by haustoria. Many cause serious crop losses. Sometimes included in Colvolvulaceae as a distinct subfamily, Cuscutoideae Link. This family and Convolvulaceae are sometimes placed in their own order, Convolvulales Dumort. Full description from Watson & Dallwitz; family synonymy from Reveal; family synopses from the University of Hawaii and Utah State.

Polemoniaceae Juss., 1789: Herbs with alternate or opposite, astipulate leaves with 5-merous flowers consisting of a united calyx and a united corolla bearing 5 stamens attached to the corolla lobes at various levels, and 3 stigmas. CA(5) [CO(5) A5] GS(3). 21 genera, 275 species. Eurasia and temperate (especially western) and tropical America. Phlox is commonly cultivated as an ornamental. The South American, bat-pollinated genus Cobaea is frequently referred to the Cobaeaceae D. Don. There is growing evidence that this family is not a member of the Asteridae but rather should be assigned to the Dilleniidae and positioned near the Fouquieriaceae DC. (Fouquieriales Takht. ex Reveal) in the order Polemoniales Bromhead. Two subfamilies are recognized: Cobaeoideae (D. Don) Arn. and Polemonioideae (Juss.) Arn. Full description from Watson & Dallwitz; family synonymy from Reveal; list of genera from Kew; family synopses from the University of Hawaii and Utah State.

Hydrophyllaceae R. Br. ex Ker-Gawl., 1817: Herbs with alternate astipulate leaves and 5-merous flowers in helicoid cymes, fused sepals and petals and a bicarpellate, unilocular ovary. CA(5) [CO(5) A5] GS(1:2). 19 genera, 275 species. Widespread except for Australia; mainly in western North America. Several are cultivated: Hesperochiron, Hydrophyllum, Nemophila, Phacelia, Romanzoffia and Wigandia. Cronquist retains the family in his Solaniales near Polemoniaceae; more recently it has been associated with Boraginaceae (e.g., Thorne and Takhtajan), occasionally in the Boraginales Dumort. Three tribes are recognized: Hydrophylleae Horan., Romanzoffieae Dumort. (including Phacelieae Benth. & Hook.f.) and Hydroleeae Choisy. Full description from Watson & Dallwitz; family synonymy from Reveal; list of genera from Kew; family synopses from the University of Hawaii and Utah State.

Lamiales Bromhead, 1838 - There is broad, general agreement at the moment that the Lamiales must be expanded to include the Scrophulariales and most likely even the Oleales Lindl. There is also considerable evidence to suggest that Plantaginales and Callitrichales are not only synoymous with Lamiales but that both Plantaginaceae and Callitrichaceae are synonyms of Scrophulariaceae. At the moment, I am inclined to retain both families but willing reduce both orders.

Boraginaceae Juss., 1789: Herbs, shrubs, vines or trees with alternate, astipulate leaves and 5-merous flowers in helicoid cymes, usually actinomorphic and bearing a 4-lobed, bicarpellate ovary, with the style typically arising from among the lobes, resulting in 4 nutlets. CA5 [CO(5) A5] GS(2). 131 genera, 2500 species. Cosmopolitan but mainly of tropical and warm regions of the world. Some are used for timber, dyes, potherbs and cultivated ornamentals (Borago, Echium, Heliotropium, Myosotis etc.). Typically divided into five subfamilies: Ehretioideae (Mart. ex Lindl.) Arn., Cordioideae Link, Heliotropioideae (Schrad.) Arn., Boraginoideae Arn. and Wellstedioideae Pilg.. The Boraginoideae are divided into six tribes according to Takhtajan (1997): Trigonotideae, Rochelieae A. DC. (including Eritricheae), Cynoglosseae W.D.J. Koch, Cerintheae Bercht. & J. Presl (including Lithospermeae Dumort.), Boragineae Bercht. & J. Presl and Myosotideae Rchb.f. A few authors, even today, recognize the Ehretiaceae and Wellstediaceae. Full description from Watson & Dallwitz; family synonymy from Reveal; list of genera from Kew; family synopses from the University of Hawaii and Utah State. See also Cardillo & Samuels.

Verbenaceae J. St.-Hil., 1805: Herbs, shrubs, or trees with 4-angled stems, opposite leaves, 5-merous, zygomorphic flowers with a single terminal style. CA(5) [COz(5) A2+2] GS(2). 91 genera, 1900 species. Mostly tropical. Used for timber, flavoring, tea and medicines. Several genera are cultivated (Clerodendrum, Duranta, Lantana, Verbena) and some are widespread weeds (Lantana). As defined by Cronquist the family cannot be distinguished from Lamiaceae. One workable solution is to recognize eight additional families: Avicenniaceae Endl. (1/14, Avicennia, a mangrove); Chloanthaceae Hutch. (incorrectly given as "Dicrastylidaceae", a nom. nud. - 5/65); Cyclocheilaceae Marais (2/17); Nesogenaceae Marais (1/6); Phrymaceae Schauer (note the correct orthography of this name - 1/2, 1 eastern North America, 1 southeast Asia); Stilbaceae Kunth (5/12 species, southern Africa), Symphoremataceae (Meisn.) Mold. ex Reveal & Hoogland (3/34, tropical shrubs with free-central placentation) and Viticaceae Juss. (1/250, temperate and tropical trees and shrubs, many ornamental with palmately compound leaves) as suggested by Takhtajan (1997). More recent data suggests recognition of Phrymaceae, Cyclocheilaceae (including Nesogenaceae), Stilbaceae and Avicenniaceae as distinct from Verbenaceae, with Chloanthaceae, Symphoremataceae and Viticaceae referred to Lamiaceae; likewise, Tetrachondraceae Wettst. is removed from Cronquist's Lamiaceae (fide Bremer et al.). Newer data, some still formally unpublished, suggests, however, that Symphoremataceae merits recognition. Full description from Watson & Dallwitz; family synonymy from Reveal; list of genera from Kew; family synopses from the University of Hawaii and Utah State.

Lamiaceae Lindl., 1836: Herbs or shrubs often with 4-angled stems, opposite leaves, 5-merous, zygomorphic flowers with a deeply 4-lobed ovary bearing a basally attached style. CA(5) [COz(5) A2 or 4] GS(2). 200 genera, 3200 species as defined by Cronquist. Cosmopolitan. Many medicinal and culinary plants: lavender, mint, basil, oregano, marjoram, rosemary, sage, savory, thyme. Many used as cultivated ornamentals: Ajuga, Calamintha, Horminum, Moluccella, Monarda, Nepeta, Physostegia, Salvia, Scutellaria, Solenstemon and Westringia. As now defined, the family is expanded to include numerous genera of Cronquist's Verbenaceae so that the Lamiaceae consists of some 290 genera and some 7700 species. In this sense, the family is divided into seven (perhaps eight) subfamilies: Caryopteridoideae Briq., Ajugoideae Kostel. (including Teucrioideae Caruel), Prostantheroideae Leurss. (including Chloanthoideae Briq.), Wenchengioideae, Stachydoideae Prantl, Lamioideae (including Pogostemonoideae) and Nepetoideae Kostel. (including Ocimoideae Kostel.). The Viticoideae Briq. is probably distinct. If Viticoideae and Carypteridoideae are combined with the Teucrioideae, and these considered distinct from the Ajugoideae, as proposed by Thorne, the correct name for the taxon is Teucrioideae. The genus Tetrachondra is now placed in Tetrachondraceae Wettst. Full description from Watson & Dallwitz; family synonymy from Reveal; list of genera from Kew; family synopses from the University of Hawaii and Utah State. See also Cardillo & Samuels.

Plantaginales Lindl., 1833 - not distinct from Lamiales.

Plantaginaceae Juss., 1789: Herbs with prominently parallel-veined leaves in basal rosettes, spicate or capitate inflorescences bearing 4-merous flowers with a membranous corolla and generally exserted stamens. CA(4) [CO(4) A4] GS(2). 3 genera, 255 species. Cosmopolitan, with Plantago often a major weed. Psyllium, a laxative, has become popular. Current molecular data suggests this family should be submerged with a family split from Cronquist's Scrophulariaceae. The name Antirrhinaceae Pers. has been proposed as a conserved name with Plantaginaceae rejected in favor of Antirrhinaceae.Full description from Watson & Dallwitz; family synonymy from Reveal; list of genera from Kew; family synopses from the University of Hawaii and Utah State.

Scrophulariales Lindl., 1833 - not distinct from Lamiales by some; retained by Takhtajan (1997). If included in the Lamiales, then the Oleaceae are the basal group in the order.

Oleaceae Hofmannss. & Link, 1813-1820: Trees or shrubs with opposite leaves and 4-merous flowers bearing 2 stamens and a 2-locular ovary with 2 seeds in each locule. CA(4) CO(4) A2 GS(2). 24 genera, 900 species. Widespread but mainly in the warm regions of Asia. Many are used for timber, (Fraxinus), olive (Olea), scents (Jasminum), and cultivated ornamentals (Chionanthus, Forsythia, Fraxinus, Jasminum, Ligustrum, Osmanthus, Syringa). Sometimes placed in Oleales. Divided into two subfamilies, Jasminoideae (Adans.) Arn. and Oleoideae (Hoffmanns. & Link) Arn.. Full description from Watson & Dallwitz; family synonymy from Reveal; list of genera from Kew; family synopses from the University of Hawaii and Utah State.

Scrophulariaceae Lindl., 1833: Herbs or shrubs with opposite or whorled leaves and 5-merous, zygomorphic, bilabiate flowers bearing 2 or 4 functional stamens and a staminode, a 2-locular ovary with a terminal style and numerous ovules, and a dehiscent capsule. CA(5) [COz(5) A2 or 4] GS(2). As defined by Cronquist, a family of some 190 genera and 4000 species; Takhtajan lists 300 genera and 500 species while Thorne suggests 220 genera and 3000 species. In this sense, the probable numers are 275 genera and 4500 species. Cosmopolitan but mainly in temperate regions. Many cultivated as ornamentals (Calceolaria, Hebe), some are medicinal (Digitalis), and several are weedy Veronica). Traditionally divided into two subfamilies, Scrophularioideae Link and Rhinanthoideae Link. Recent work shows Scrophulariaceae to be paraphyletic and several genera within the traditional Scrophulariaceae must be placed elsewhere. Olmstead defines Scrophulariaceae to include the Buddlejaceae K. Wilh. Myoporaceae R. Br., Oftiaceae Takht. & Reveal and Selaginaceae Choisy, or about 35 genera, the more notable being Hemimeris, Leucophyllum, Verbascum and Scrophularia. I prefer to retain Buddlejaceae, Myoporaceae and Oftiaceae. The Antirrhinaceae are defined by Olmstead to include three highly reduced and modified families, Callitriche Hippuridaceae and Plantaginaceae in addition to Globulariaceae. I prefer to remove each of these to form a more narrowly defined family with such representatives as Chelone, Collinsia, Digitalis, Gratiola, Penstemon and Veronica. The hemiparasitic members of the Scrophulariaceae (most of the Rhinanthoideae) are properly placed by Olmstead in a greatly expanded Orobanchaceae Vent. traditionally characterized as those members that are wholly parasitic. As now defined, the Orobanchaceae includes such genera as Castilleja, Melampyrum, Pedicularis and Seymeria. The largely tropical genus Calceolaria and their relatives will be placed in a new family by Olmstead. In addition, two families not recognized by Cronquist are adopted: Paulowniaceae Nakai and Schlegeliaceae (A.H. Gentry) Reveal. My own views are summaries elsewhere. Full description from Watson & Dallwitz; family synonymy from Reveal; list of genera from Kew; family synopses from the University of Hawaii and Utah State.

Acanthaceae Juss., 1789: Herbs or shrubs with opposite leaves and 5-merous, zygomorphic, bilabiate flowers bearing 2 or 4 functional stamens and a staminode, a 2-locular ovary with a terminal style and 2 ovules, and an explosively dehiscent capsule. CA(5) [COz(5) A2 or 4] GS(2). 357 genera, 4350 species. Widespread but mainly tropical. The generic lines are much debated (some accepting only 250 genera). Many elegant cultivated ornamentals for the hothouse (Aphelandra, Eranthemum, Fittonia, Justicia, Ruellia, Strobilanthes, Thunbergia). Divided into five subfamilies: Nelsonioideae Lindl. ex Pfeiff., Thunbergioideae Kostel., Mendoncioideae Lindau, Acanthoideae Link and Ruellioideae T. Anders. The first three subfamilies are sometimes recognized as Nelsoniaceae (Nees) Sreem., Thunbergiaceae (Dumort.) Lilja and Mendonciaceae Bremek.. Full description from Watson & Dallwitz; family synonymy from Reveal; list of genera from Kew; family synopses from the University of Hawaii and Utah State.

Bignoniaceae Juss., 1789: Woody vines or trees with opposite, often compound leaves, more or less zygomorphic, 5-merous flowers producing a woody capsule typically with conspicuously winged seeds CA(5) [COz(5) A4] GS(2). 113 genera, 800 species. Mainly of tropical regions. Some are used for timber and ornamental trees and vines. Divided into seven tribes: Tecomeae Endl., Oroxyleae, Bignonieae Dumort., Eccremocarpeae Hogg, Tourrettieae G. Don, Crescentieae G. Don and Coleeae. Full description from Watson & Dallwitz; family synonymy from Reveal; list of genera from Kew; family synopsse from the University of Hawaii and Utah State.

Last revised: 27 Jan 1999