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The families of flowering plants.                                                                                                                                                                

Plantaginaceae Juss.

                        

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IncludingLittorellaceae S.F. Gray, Psylliaceae Horan.Excluding Globulariaceae

Habit and leaf form. Herbs (mostly), or shrubs. ‘Normal’ plants. Plants succulent (e.g. Plantago maritima), or non-succulent. Annual to perennial; with a basal aggregation of leaves, or with neither basal nor terminal aggregations of leaves. Sometimes pachycaul. Helophytic to xerophytic. Heterophyllous, or not heterophyllous. Leavesalternate; spiral; ‘herbaceous’, or leathery, or fleshy (occasionally); petiolate to sessile (the blade/petiole distinction never clear); sheathing. Leaf sheaths not tubular; with free margins. Leaves simple; epulvinate. Lamina entire (usually), or dissected; linear (sometimes ericoid), or lanceolate, or oblanceolate, or ovate, or orbicular; when dissected, more or less pinnatifid; parallel-veined; cross-venulate. Leaves exstipulate.

Leaf anatomy. Hydathodes present (commonly), or absent. Stomata mainly confined to one surface, or on both surfaces; anomocytic, or tetracytic, or diacytic, or cyclocytic.

Lamina dorsiventral, or isobilateral. Minor leaf veins with phloem transfer cells (Littorella,Plantago).

Stem anatomy. Cork cambium present (in rhizomes), or absent; initially deep-seated, or superficial. Nodes unilacunar, or tri-lacunar, or multilacunar. Medullary bundles present, or absent. Secondary thickening absent, or developing from a conventional cambial ring. Xylem with libriform fibres; with vessels. Vessel end-walls simple. Wood partially storied (VI), or not storied; parenchyma absent or extremely sparse.

Reproductive type, pollination. Plants hermaphrodite, or gynomonoecious, or monoecious. Pollination anemophilous, or anemophilous and entomophilous.

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’; in spikes, or in heads. The ultimate inflorescence unit racemose. Inflorescences scapiflorous (at least, pedunculate); terminal; heads or spikes. Flowers bracteate; ebracteolate; small; regular; (3–)4 merous; cyclic; tetracyclic. Free hypanthium absent. Hypogynous disk absent.

Perianthwith distinct calyx and corolla; (6–)8; 2 whorled; isomerous. Calyx (3–)4 (the posterior sepal seemingly lacking); 1 whorled; gamosepalous, or partially gamosepalous. When ostensibly only partially gamosepalous, 2 of the members joined (the abaxial pair). Calyx regular (diagonal); persistent; imbricate. Corolla (3–)4 (the two posterior petals fused?); 1 whorled; gamopetalous; imbricate (membranous); regular, or unequal but not bilabiate (the lobes unequal in Bougueria).

Androecium (1–)4. Androecial members adnate (to the corolla); all equal; free of one another; 1 whorled. Androecium exclusively of fertile stamens. Stamens(1–)4; reduced in number relative to the adjacent perianth to isomerous with the perianth; oppositisepalous. Anthers dorsifixed; versatile; dehiscing via longitudinal slits; introrse. Microsporogenesis simultaneous. Tapetum glandular. Pollen grains aperturate; (3–)4–14 aperturate; porate (occasionally), or foraminate; 2-celled (Littorella), or 2-celled and 3-celled (with both states recordedin Plantago).

Gynoecium 2 carpelled. Carpels reduced in number relative to the perianth. The pistil2 celled. Gynoecium syncarpous; synstylovarious to eu-syncarpous; superior. Ovary 2 locular. Gynoecium median; stylate. Styles 1; attenuate from the ovary; apical. Stigmas usually 2 lobed; dry type; non-papillate; Group II type. Placentation axile (Plantago), or basal. Ovules 5–50 per locule (i.e. to many, in Plantago), or 1 per locule; peltate; non-arillate; hemianatropous, or anatropous, or campylotropous; unitegmic; tenuinucellate. Endothelium differentiated. Embryo-sac development Polygonum-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; not proliferating; persistent. Synergids pear-shaped. Endosperm formation cellular. Endosperm haustoria present; chalazal and micropylar (aggressive). Embryogeny onagrad.

Fruit non-fleshy; dehiscent, or indehiscent; a capsule (membranous), or a nut.Capsules circumscissile. Seeds copiously endospermic. Cotyledons 2. Embryo achlorophyllous (6 species of Plantago); straight (usually), or curved.

Seedling.Germination phanerocotylar.

Physiology, biochemistry. Not cyanogenic. Alkaloids present, or absent. Iridoids detected; ‘Route II’ type (+decarb.). Verbascosides detected. Proanthocyanidins absent. Ellagic acid absent (4 s ecies of Plantago). Arbutin absent. Ursolic acid present. Saponins/sapogenins present, or absent. Aluminium accumulation not found. C3 and CAM. C3 physiology recorded directly in Plantago. CAM recorded directly in Littorella (aquatic CAM only). Anatomy non-C4 type (Plantago).

Geography, cytology. Frigid zone to tropical. Cosmopolitan. X = 4–12(+).

Taxonomy.Subclass Dicotyledonae; Tenuinucelli. Dahlgren’s Superorder Lamiiflorae; Scrophulariales. Cronquist’s Subclass Asteridae; Plantaginales. APG 3 core angiosperms; core eudicot; Superorder Asteranae; lamiid; Order Lamiales.

Species 270. Genera 3; Bougueria, Littorella, Plantago.

General remarks. For discussion of classificatory problems posed by Scrophulariaceae, impinging on Bignoniaceae, Buddlejaceae, Callitrichaceae, Plantaginaceae,Hippuridaceae, Lentibulariaceae, and Hydrostachydaceae, and such problem genera as Paulownia and Schlegelia, see Olmstead and Reeves (1995), who provide preliminary insights from chloroplast gene sequencing.

Economic uses, etc. Widespread weeds, and a laxative from P. psyllium seed.

 Illustrations:

  • Technical details: Plantago.
  • Technical details: Plantago (Thonner).
  • Technical details: Littorella.
  • Technical details (embryo, germination): Littorella.
  • Littorella uniflora (B. Ent.).
  • Littorella uniflora: Eng. Bot. 1169, 1867.
  • Plantago coronopus: Eng. Bot. 1168, 1867.
  • Plantago lanceolata (B. Ent.).
  • Plantago lanceolata: Eng. Bot. 1164, 1867. 
  • Plantago major: Eng. Bot. 1162, 1867.
  • Plantago maritima: Eng. Bot. 1166, 1867.
  • Plantago media (B. Ent.).
  • Plantago media: Eng. Bot. 1163, 1867.
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