The families of flowering plants
Habit and leaf form. Glutinous-villous herbs. Plants autotrophic. Annual, or perennial; often tuberous. Leaves alternate, or opposite; petiolate; non-sheathing; simple. Lamina cross-venulate. Leaves exstipulate.
Leaf anatomy. The leaf lamina dorsiventral. Stomata present; on both surfaces; anomocytic. Hairs present; eglandular and glandular (the former long-uniseriate, the latter comprising capitate, short-staked mucilage hairs and/or hairs with unicellular or uniseriate stalks and spherical or turbinate heads). The mesophyll containing crystals. The crystals small, druses, or solitary-prismatic (mainly?). Minor leaf veins without phloem transfer cells (Martynia).
Axial (stem, wood) anatomy. Primary vascular tissues in a cylinder, without separate bundles; collateral. Secondary thickening developing from a conventional cambial ring. Primary medullary rays mixed wide and narrow.
The vessel end-walls simple.
Reproductive type, pollination. Plants hermaphrodite.
Inflorescence, floral, fruit and seed morphology. Flowers aggregated in inflorescences; in racemes. The ultimate inflorescence units racemose. Inflorescences terminal; terminal racemes. Flowers very irregular; zygomorphic. The floral irregularity involving the perianth and involving the androecium. Flowers 5 merous; cyclic; tetracyclic. Free hypanthium absent. Hypogynous disk present; intrastaminal.
Perianth with distinct calyx and corolla; 10; 2 whorled; isomerous. Calyx 5; 1 whorled; gamosepalous (sometimes nearly free, split down one side and spathaceous in Craneolaria); blunt-lobed; with the median member posterior. Corolla 5; 1 whorled; gamopetalous; imbricate; unequal but not bilabiate, or bilabiate.
Androecium 3, or 5. Androecial members adnate (epipetalous); markedly unequal; free of one another; 1 whorled. Androecium including staminodes. Staminodes 1, or 3 (the posterior member always reduced); in the same series as the fertile stamens; representing the posterior median member, or the posterior median member and the posterior-lateral pair. Fertile stamens representing the anterior-lateral pair, or the posterior-lateral pair and the anterior-lateral pair. Stamens 2 (the shorter pair sometimes sterile, as well as the posterior, in Martynia), or 4; didynamous, or not didynamous, not tetradynamous; reduced in number relative to the adjacent perianth; oppositisepalous; alternating with the corolla members. Anthers connivent; dehiscing via longitudinal slits; tetrasporangiate. Endothecium developing fibrous thickenings. Anther epidermis persistent. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral, or decussate. Anther wall initially with one middle layer. Tapetum glandular. Pollen grains aperturate, or nonaperturate (synrugoidate?); 13–40 aperturate (? polyaperturate); foraminate, or rugate; 3-celled (in Proboscidea).
Gynoecium 2 carpelled. Carpels reduced in number relative to the perianth. The pistil 1 celled, or 4 celled. Gynoecium syncarpous; synstylovarious to eu-syncarpous; superior. Ovary unilocular; 1 locular (but rendered more or less 4-locular in the fruit, by union of the T-shaped placentae with one another and with the endocarp). Locules secondarily divided by false septa to without false septa. Gynoecium median. Styles 1; attenuate from the ovary; apical. Stigmas 1, or 2; 2 lobed. Placentation parietal. Ovules in the single cavity 3–100 (few to many); anatropous; 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 (large). Endosperm formation cellular. Endosperm haustoria present; chalazal (in Martynia). Embryogeny onagrad.
Fruit fleshy to non-fleshy; dehiscent; a capsule. Capsules loculicidal (the soft outer pericarp shed, the inner woody). Dispersal by animals, associated with usually hooked or curved spurs which develop from the tip of the midrib of each carpel. Seeds scantily endospermic. Cotyledons 2. Embryo straight.
Seedling. Germination phanerocotylar.
Physiology, phytochemistry. Not cyanogenic. Verbascosides detected (Martynia). Cornoside detected (Martynia). Iridoids detected; Route II type (+decarb.). Saponins/sapogenins absent. Proanthocyanidins absent. Flavonols present, or absent; kaempferol. Ellagic acid absent (2 genera, 2 species).
Geography, cytology. Neotropical. Sub-tropical to tropical. Tropical and subtropical South America, Mexico.
Taxonomy. Subclass Dicotyledonae; Tenuinucelli. Dahlgrens Superorder Lamiiflorae; Scrophulariales. Cronquists Subclass Asteridae; Scrophulariales. APG 3 core angiosperms; core eudicot; Superorder Asteranae; lamiid; Order Lamiales (as a synonym of Pedaliaceae).
Species 13. Genera 4; Craniolaria, Ibicella, Martynia, Proboscidea.
General remarks. Differing from Pedaliaceae (q.v.) in conspicuous features of inflorescence and gynoecium morphology, and the compiled data suggest further differences in characters relying on limited sampling (leaf anatomy, anther wall and pollen development, embryology).
Illustrations. • Le Maout and Decaisne: Craniolaria, Martynia. • Martynia diandra: Bot. Reg. 2001, 1837. • Martynia fragrans: Bot. Reg. 6, 1841. • Martynia lutea: Bot. Reg. 934, 1825.
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: 9th January 2018. delta-intkey.com/angio’.