| # The Families of Angiosperms |

L. Watson and M.J. Dallwitz

### Plantaginaceae Juss. (sensu stricto)

Including Littorellaceae S.F. Gray, Psylliaceae Horan.; excluding Globulariaceae, Callitrichaceae, and the numerous genera of traditional Scrophulariaceae transferred to this family by APG (see comments below).

Habit and leaf form. Herbs (mostly), or shrubs. Plants succulent (e.g. Plantago maritima), or non-succulent. Annual to perennial; with a basal aggregation of leaves, or without conspicuous aggregations of leaves. Sometimes pachycaul. Helophytic to xerophytic. Conspicuously heterophyllous, or not conspicuously heterophyllous. Leaves alternate; 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. The leaf lamina dorsiventral, or bifacial (then isobilateral, or sometimes with the mesophyll consisting entirely of palisade cells). Hydathodes present (commonly), or absent. Stomata variously mainly confined to one surface (adaxial only, or abaxial only), or on both surfaces; anomocytic, or tetracytic, or diacytic, or cyclocytic. Hairs present; eglandular (only, in Littorella), or eglandular and glandular. Minor leaf veins with phloem transfer cells (Littorella, Plantago).

Axial (stem, wood) anatomy. Cork cambium present (in rhizomes), or absent; initially deep-seated, or initially superficial. Nodes unilacunar, or tri-lacunar, or multilacunar. Primary vascular tissues in a cylinder, without separate bundles, or comprising a ring of bundles. Medullary bundles present, or absent. Secondary thickening absent, or developing from a conventional cambial ring.

The vessels (at least in Plantago fernandezia) small; solitary, radially paired, and in radial multiples (the multiples in long radial lines). The vessel end-walls simple. The axial xylem with tracheids (according to Solereder); without fibre tracheids; with libriform fibres. The parenchyma absent or extremely sparse; wood partially storied (VI), or not storied.

Reproductive type, pollination. Unisexual flowers present, or absent. 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 units 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.

Perianth with 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 pistil 2 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, phytochemistry. C3, or CAM. C3 physiology recorded directly in Plantago. CAM recorded directly in Littorella (aquatic CAM only). Anatomy non-C4 type (Plantago). Not cyanogenic. Alkaloids present, or absent. Verbascosides detected. Arbutin absent. Iridoids detected; ‘Route II’ type (+decarb.). Saponins/sapogenins present, or absent. Proanthocyanidins absent. Ellagic acid absent (4 s ecies of Plantago). Ursolic acid present. Aluminium accumulation not found.

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 III core angiosperms; core eudicot; Superorder Asteranae; lamiid. APG IV Order Lamiales.

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

General remarks. No attempt is made here to incorporate the radical APG re-organization of generic assignments involving transfer to Plantaginaceae sensu stricto of about 90 genera and 1,500 species of traditional Scrophulariaceae, the practical worth of which in terms of character correlations remains to be demonstrated. For discussion of the long recognised 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 provided preliminary insights from chloroplast gene sequencing. Meanwhile, in view of the numerous differences between Plantaginaceae and Callitrichaceae (q.v.) involving characters of habit, vegetative morphology and anatomy, general floral morphology, gynoecium and fruit structure, and embryology, the latter are a bizarre inclusion here in recent cladistic classifications. Likewise Globulariaceae (q.v), which in terms of the present compilation of data differ from Plantaginaceae in 18 characters, involving general morphology, anatomy, pollen form, embryology and cytology.

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

Quotations.

No salve, sir, but a plantain
(‘Love’s Labour’s Lost’ iii., 1 - Plantago major, the leaf reputedly good for a broken shin)

And with a wabret leaf he made a wallet,
With scrip to beg his crumbs and pick his sallet
(Quoted ‘from an early poet’ by Ann Pratt in 1857, describing humourously(?) a ‘bee’s pilgimage’ — medieval ‘wabret’ = Plantago major, ‘scrip’ = satchel, ‘sallet’ = salad)

Illustrations. • Le Maout and Decaisne: Plantago major. • Plantago palmata: Thonner. • Le Maout and Decaisne: Littorella uniflora, as L. lacustris. • Le Maout and Decaisne: Littorella, embryo and germination). • Littorella uniflora (B. Ent.). • Littorella uniflora: Eng. Bot. 1169, 1867. • Plantago asiatica: R. Wight 2 (1850). • 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. • Leaf hairs of Littorella lacustris, Plantago argentea and P. cylindrica: Solereder, 1908.

We advise against extracting comparative information from the descriptions. This is much more easily achieved using the DELTA data files or the interactive key, which allows access to the character list, illustrations, full and partial descriptions, diagnostic descriptions, differences and similarities between taxa, lists of taxa exhibiting or lacking specified attributes, distributions of character states within any set of taxa, geographical distribution, genera included in each family, and classifications (Dahlgren; Dahlgren, Clifford, and Yeo; Cronquist; APG). See also Guidelines for using data taken from Web publications.

Cite this publication as: ‘Watson, L., and Dallwitz, M.J. 1992 onwards. The families of flowering plants: descriptions, illustrations, identification, and information retrieval. Version: 8th June 2026. delta-intkey.com’.

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