The families of flowering plants
Including Gustaviaceae; excluding Asteranthaceae, Barringtoniaceae, Foetidiaceae, Napoleonaeaceae.
Habit and leaf form. Trees; without essential oils. Leaves medium-sized, or large; alternate; spiral (aggregated at the tips of the twigs); herbaceous, or leathery; petiolate; non-sheathing; simple. Lamina entire; pinnately veined; cross-venulate. Leaves exstipulate. Domatia occurring in the family (Combretodendron); manifested as pits.
General anatomy. Plants with silica bodies.
Leaf anatomy. The leaf lamina dorsiventral. Stomata mainly confined to one surface (abaxial, usually), or on both surfaces; usually anisocytic. Hairs present; eglandular; simple, unicellular, or multicellular. Unicellular hairs simple (occasionally tufted). Adaxial hypodermis usually absent. Lamina without secretory cavities. The mesophyll containing crystals. The crystals druses and solitary-prismatic.
Axial (stem, wood) anatomy. Cork cambium present; initially superficial. Nodes unilacunar. Primary vascular tissues in a cylinder, without separate bundles; collateral. Internal phloem absent. Cortical bundles present (these usually conventionally orientated). Medullary bundles absent. Secondary thickening developing from a conventional cambial ring.
The wood diffuse porous. The vessels very small to medium (but mostly medium sized); solitary, radially paired, and in radial multiples, or clustered. The vessel end-walls simple, or scalariform and simple (rarely). The vessels without vestured pits. The axial xylem with tracheids (Allantoma), or without tracheids (usually); without vasicentric tracheids; with fibre tracheids, or without fibre tracheids; with libriform fibres, or without libriform fibres; including septate fibres (rarely), or without septate fibres. The parenchyma usually abundant, apotracheal, or apotracheal and paratracheal (typically mostly in apotracheal bands). The secondary phloem usually stratified into hard (fibrous) and soft (parenchymatous) zones. Included phloem absent. The wood not storied. Tyloses commonly present.
Reproductive type, pollination. Plants hermaphrodite. Floral nectaries present. Nectar secretion from the disk. Pollination ornithophilous, or cheiropterophilous.
Inflorescence, floral, fruit and seed morphology. Flowers solitary, or aggregated in inflorescences. The ultimate inflorescence units when flowers aggregated, racemose. Flowers regular to very irregular; when irregular, zygomorphic (sometimes the androecium spectacularly so). The floral irregularity involving the perianth and involving the androecium. Flowers cyclic; polycyclic. Free hypanthium absent (always with complete fusion of receptacle and ovary).
Perianth with distinct calyx and corolla; 8–12; 2 whorled; isomerous. Calyx usually 4–6; 1 whorled; polysepalous; not persistent (on the fruit); valvate. Corolla 4–6; 1 whorled; polypetalous (usually), or gamopetalous; imbricate; unequal but not bilabiate to regular; deciduous (with the stamens).
Androecium 40–1000 (many). Androecial members maturing centrifugally; free of the perianth; coherent (more or less united below, the connate parts sometimes produced on one side of the androecium to form a flat ligule or a hooding structure); 3–5 whorled (? in several whorls). Androecium exclusively of fertile stamens, or including staminodes (by abortion of some anthers, in association with one-sided development of the union). Stamens 40–1000 (many); polystemonous; inflexed in bud. Anthers usually versatile; dehiscing via longitudinal slits; tetrasporangiate. Endothecium developing fibrous thickenings. Anther epidermis persistent. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral. Anther wall initially with more than one middle layer; of the basic type. Tapetum amoeboid. Pollen grains aperturate; 3 aperturate; colporate (or colporoidate, not synocolpate); 2-celled (in Gustavia).
Gynoecium 2–6(–10) carpelled. Carpels reduced in number relative to the perianth to increased in number relative to the perianth. The pistil 2–6(–10) celled. Gynoecium syncarpous; synstylovarious, or eu-syncarpous (?); partly inferior, or inferior. Ovary 2–6(–10) locular. Epigynous disk (disks) present (usually with an intrastaminal disk, as well as one under C and A: Airy Shaw 1973). Gynoecium stylate. Styles 1; apical. Placentation basal (Eschweilera), or basal to axile, or axile to apical, or apical. Ovules 1–50 per locule (to many); pendulous, or horizontal, or ascending; arillate (often, with a funicular aril), or non-arillate; anatropous; bitegmic; tenuinucellate. Outer integument not contributing to the micropyle. Endothelium differentiated. Embryo-sac development Polygonum-type. Antipodal cells formed; 3; not proliferating; ephemeral. Synergids pear-shaped. Endosperm formation nuclear.
Fruit often large, fleshy, or non-fleshy; dehiscent, or indehiscent; woody, a capsule, or capsular-indehiscent, or a berry. Capsules when dehiscent, circumscissile (monkey pots). Seeds non-endospermic. Embryo large, rudimentary at the time of seed release to weakly differentiated (as in Bertholletia (the Brazil nut), where it consists mainly of the much thickened hypocotyl), or well differentiated. Cotyledons 2.
Seedling. Germination phanerocotylar, or cryptocotylar.
Physiology, phytochemistry. Sugars transported as sucrose (Lecythis). Cyanogenic, or not cyanogenic. Alkaloids absent (2 species). Iridoids not detected. Saponins/sapogenins present. Proanthocyanidins present, or absent; when present, cyanidin and delphinidin. Flavonols present, or absent; kaempferol, or quercetin. Ellagic acid present (3 genera, 3 species).
Geography, cytology. Neotropical. Sub-tropical to tropical. Tropical America. X = 17.
Taxonomy. Subclass Dicotyledonae; Tenuinucelli. Dahlgrens Superorder Theiflorae; Theales. Cronquists Subclass Dilleniidae; Lecythidales. APG III core angiosperms; core eudicot; Superorder Asteranae. APG IV Order Ericales.
Species 325. Genera 10; Allontoma, Bertholletia, Cariniana, Corythophora, Couratari, Couroupita, Eschweilera, Grias, Gustavia, Lecythis.
General remarks. Morton et al. (1998) interpret these genera as subfamily Lecythidoideae of their expanded Lecythidaceae.
Illustrations. • Lecythis ovata: Saint-Hilaire et al., Fl. Brasil. (1829). • Le Maout and Decaisne: Bertholletia, Couroupita, Lecythis. • Lecythis ovata: Lindley. • Gustavia gracillima: Bot. Mag. 101 (1875).
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: 5th March 2018. delta-intkey.com/angio’.