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The families of flowering plants

L. Watson and M.J. Dallwitz

Morinaceae J.G. Agardh

~ Caprifoliaceae sensu lato, Dipsacaceae.

Habit and leaf form. Thistlelike herbs. Perennial (with simple stems). Leaves opposite, or whorled; simple. Lamina dissected (usually), or entire (rarely); pinnatifid, or pinnatifid and spinose (or spinose-dentate). Leaves exstipulate. Lamina margins entire (rarely), or serrate, or dentate (or spinose-dentate).

Leaf anatomy. The leaf lamina dorsiventral. Mucilaginous epidermis present, or absent. Minor leaf veins without phloem transfer cells.

Axial (stem, wood) anatomy. Primary vascular tissues in a cylinder, without separate bundles (the initial bundles soon becoming linked); collateral (?). Internal phloem absent (?). Secondary thickening developing from a conventional cambial ring.

The axial xylem with libriform fibres.

Reproductive type, pollination. Plants hermaphrodite. Floral nectaries present. Nectar secretion from the disk.

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’; in verticils (cf. Labiatae). Inflorescences with involucral bracts (each flower enclosed in a tubular-campanulate, spinose-margined involucel, cf. Dipsacaceae). Flowers bracteolate (if the involucel is interpreted as fused bracteoles), or ebracteolate; very irregular; zygomorphic. The floral irregularity involving the perianth and involving the androecium. Flowers cyclic; tetracyclic. Free hypanthium absent.

Perianth with distinct calyx and corolla; 7–9 (depending on interpretation of the calyx); 2 whorled; anisomerous. Calyx 2, or 4 (sometimes ‘four lobed’); 1 whorled; gamosepalous; blunt-lobed; bilabiate (the lips entire or two-lobed); persistent. Corolla 5; 1 whorled; gamopetalous; imbricate; tubular (tube usually slender and exserted, the throat gaping); bilabiate; yellow, or red, or pink, or white.

Androecium 4 (the posterior member missing). Androecial members adnate; markedly unequal; free of one another; 1 whorled. Androecium including staminodes. Staminodes 2 (the two anterior members); in the same series as the fertile stamens; representing the anterior-lateral pair. Fertile stamens representing the posterior-lateral pair. Stamens 2; inserted in the throat of the corolla tube; reduced in number relative to the adjacent perianth; oppositisepalous; alternating with the corolla members. Anthers dehiscing via longitudinal slits; introrse; tetrasporangiate. Endothecium developing fibrous thickenings. Microsporogenesis simultaneous. Anther wall initially with one middle layer; of the ‘dicot’ type. Tapetum amoeboid. Pollen shed as single grains. Pollen grains aperturate; 3(–4) aperturate; porate, or colporate (to zonorate); 2-celled.

Gynoecium basically probably 2 carpelled (but one obsolete). Carpels reduced in number relative to the perianth. The pistil 1 celled. Gynoecium syncarpous; eu-syncarpous; inferior. Ovary 1 locular. Epigynous disk present. Gynoecium stylate. Styles 1 (slender); apical. Stigmas 1; capitate. Placentation apical. Ovules in the single cavity 1; pendulous; anatropous; unitegmic; tenuinucellate. Endothelium differentiated. Embryo-sac development Polygonum-type. Antipodal cells formed; proliferating. Endosperm formation cellular.

Fruit non-fleshy; indehiscent; achene-like (the pericarp thickened, more or less rugose); 1 seeded. Seeds endospermic. Endosperm oily. Cotyledons 2. Embryo straight.

Physiology, phytochemistry. Not cyanogenic. Alkaloids absent. Verbascosides not detected. Iridoids not detected. Proanthocyanidins absent. Ellagic acid absent.

Geography, cytology. Holarctic. Temperate. Southeast Europe to Himalayas and Southwest China. X = 17.

Taxonomy. Subclass Dicotyledonae; Tenuinucelli. Dahlgren’s Superorder Corniflorae; Dipsacales. Cronquist’s Subclass Asteridae; Dipsacales. APG 3 core angiosperms; core eudicot; Superorder Asteranae; campanulid; Order Dipsacales (as a synonym of Caprifoliaceae?).

Species 17. Genera 3; Acanthocalyx, Cryptothladia, Morina.

General remarks. Comparing this description with Caprifoliaceae sensu lato (q.v.) shows differences in numerous characters invoving inflorescence, perianth, androecium, and gynoecium morphology; also in wood-anatomical and embryological characters relying on limited sampling.

Illustrations. • Morina longifolia: Bot. Reg. xxvi, 36 (1840). • Morina nepalensis (as M. betonicoides): Hook. Ic. Pl. 12 (1876).


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: 13th March 2017. delta-intkey.com/angio’.

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