Family Phrymataceae

Phrymataceae Schauer

Alternatively Phrymaceae~ Verbenaceae p.p.

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.

Stem anatomy. Young stems tetragonal. Primary vascular tissue comprising a ring of bundles (one bundle at each angle of the young stem); centrifugal. Cortical bundles absent. Medullary bundles absent. Internal phloem absent. Secondary thickening developing from a conventional cambial ring. ‘Included’ phloem absent.

Reproductive type, pollination. Plants hermaphrodite.

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’; in spikes. The ultimate inflorescence unit 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, biochemistry. Not cyanogenic. Alkaloids absent (one species). Iridoids detected. Proanthocyanidins absent. Saponins/sapogenins 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 3 core angiosperms; core eudicot; Superorder Asteranae; lamiid; Order Lamiales.

Species 2. Genera 1; only genus, Phryma.

This is part of the Labiatae/Verbenaceae problem: see remarks under Labiatae.