| # The Families of Angiosperms |

L. Watson and M.J. Dallwitz

### Phrymataceae Schauer

Alternatively Phrymaceae; ~ Verbenaceae p.p.

Excluding Mazaceae Reveal

Habit and leaf form. Herbs. Erect, perennial; rhizomatous. Leaves opposite; petiolate; simple. Lamina entire; obovate; pinnately veined; cross-venulate. Leaves exstipulate. Lamina margins coarsely dentate.

Leaf anatomy. Stomata present; mainly confined to one surface (abaxial); anomocytic.

Axial (stem, wood) anatomy. Young stems tetragonal in section. Primary vascular tissues comprising a ring of bundles (one bundle at each angle of the young stem); collateral. Internal phloem absent. Cortical bundles absent. Medullary bundles absent. Secondary thickening developing from a conventional cambial ring.

‘Included’ phloem absent.

Reproductive type, pollination. Unisexual flowers absent. Plants hermaphrodite.

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’; in spikes. The ultimate inflorescence units racemose. Inflorescences terminal and axillary; slender spikes, the flowers erect in bud but later spreading to deflexed. Flowers bracteate; small; very irregular; zygomorphic; cyclic; tetracyclic. Free hypanthium absent.

Perianth with distinct calyx and corolla; 10; isomerous. Calyx 5; 1 whorled; gamosepalous; five blunt-lobed to toothed; tubular; bilabiate (prominently five nerved, the posterior three lobes subulate and hooked at the apex, the anterior two short and muticous); persistent (appressed to the axis in the fruit); with the median member posterior. Corolla 5; 1 whorled; gamopetalous; tubular; bilabiate (the posterior lip erect and shortly bilobed, the anterior longer, spreading, broadly trifid); purple, or blue.

Androecium 4. Androecial members adnate; markedly unequal; free of one another; 1 whorled. Androecium exclusively of fertile stamens. Stamens 4; inserted midway down the corolla tube to in the throat of the corolla tube; didynamous; reduced in number relative to the adjacent perianth; oppositisepalous; alternating with the corolla members; filantherous (the filaments slender). Anthers subreniform; dehiscing via longitudinal slits; tetrasporangiate. Pollen grains aperturate; 3 aperturate; colpate; 3-celled.

Gynoecium theoretically 2 carpelled, or 1 carpelled (ostensibly). Carpels reduced in number relative to the perianth. The pistil 1 celled. Gynoecium syncarpous (but pseudomonomerous); synstylovarious (detectable from monomery by the apically bifid style); superior. Ovary 1 locular (i.e. pseudomonomerous); sessile. Gynoecium stylate. Styles 1; apical to lateral. Stigmas 2. Placentation basal. Ovules in the single cavity 1; ascending; orthotropous; unitegmic; tenuinucellate. Embryo-sac development Polygonum-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; not proliferating; persistent. Synergids with filiform apparatus. Hypostase present. Endosperm formation cellular. Endosperm haustoria present; chalazal. Embryogeny solanad.

Fruit non-fleshy; indehiscent; a nut (enclosed by the persistent calyx, the pericarp thin); 1 seeded. Seeds non-endospermic. Embryo well differentiated. Cotyledons 2 (folded); rolled. Embryo oblong. Testa membranous, loosely applied to the pericarp.

Physiology, phytochemistry. Not cyanogenic. Alkaloids absent (one species). Iridoids detected. Saponins/sapogenins absent. Proanthocyanidins absent.

Geography, cytology. Holarctic and Paleotropical. Temperate to sub-tropical. India to Japan, Eastern North America.

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

Species 1 (Phryma leptostachya). Genera 1; only genus, Phryma.

General remarks. This is part of the Labiatae/Verbenaceae imbroglio, see remarks under Labiatae; extended by APG to include taxa referred here to Scrophulariaceae - Elacholoma, Erythranthe, Glossostigma, Hemichaena, Lancea, Mazus, Dodartia, Leacocarpus, Mimulus, as well as verbenaceous Phryma).

Illustrations. • Erythranthe cuprea (as Mimulus luteus var. cuprea): Bot. Mag. 90 (1864). • Phryma leptostachya: Nat. Pflanzenfam. IV (1895). • Phryma leptostachya: Hutchinson.

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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