The families of flowering plants
Including Ardisiaceae Juss., Embelieae (Embeliaceae) J.G. Agardh, Maesaceae (A.DC.) Anderb, Ståhl & Källersjö, Ophiospermes (Ophiospermae) Vent., Ophiospermataceae Kuntze; excluding Aegicerataceae, Theophrastaceae.
Habit and leaf form. Trees and shrubs, or lianas (a few and a few sub-herbaceous); with coloured juice, or non-laticiferous; resinous. Climbing (a few), or self supporting (mostly). Mesophytic. Leaves alternate; spiral; petiolate; non-sheathing; gland-dotted (often), or not gland-dotted (then glandular-hairy); aromatic, or without marked odour; simple. Lamina entire; pinnately veined; cross-venulate. Leaves exstipulate. Lamina margins often entire. Leaf development not graminaceous. Domatia occurring in the family (seen in 3 genera); manifested as pockets, or hair tufts.
Leaf anatomy. The leaf lamina generally dorsiventral. Mucilaginous epidermis present, or absent. Hairs present; eglandular, or glandular; mostly multicellular (and mostly not simple, being variously forked, branched or peltate). Lamina with secretory cavities (usually), or without secretory cavities. Secretory cavities containing resin (yellow or reddish brown); schizogenous. The mesophyll often with hypodermal fibres; containing crystals. The crystals druses, or solitary-prismatic. Minor leaf veins without phloem transfer cells (Ardisia, Suttonia (= Rapanea)).
Axial (stem, wood) anatomy. Secretory cavities commonly present; with resin (yellow or reddish-brown). Cork cambium present; initially superficial. Nodes unilacunar. Primary vascular tissues in a cylinder, without separate bundles; collateral. Internal phloem absent. Cortical bundles absent. Medullary bundles absent. Secondary thickening developing from a conventional cambial ring. Primary medullary rays generally narrow (?).
The wood diffuse porous (mostly), or semi-ring porous. The vessels moderately small; in radial multiples (commonly, of 3–4 cells), or clustered (these irregular), or in tangential arcs. The vessel end-walls simple (usually, exclusively), or scalariform and simple (occasionally in some genera). The vessels without vestured pits; with spiral thickening (rarely), or without spiral thickening. The axial xylem without fibre tracheids; with libriform fibres; usually including septate fibres. The fibres without spiral thickening. The parenchyma usually sparse, paratracheal; wood partially storied, or not storied.
Reproductive type, pollination. Plants hermaphrodite, or monoecious, or polygamomonoecious, or dioecious. Female flowers with staminodes (often, large), or without staminodes.
Inflorescence, floral, fruit and seed morphology. Flowers aggregated in inflorescences. The ultimate inflorescence units racemose. Inflorescences axillary, or terminal. Flowers bracteolate (rarely, then usually bibracteolate, e.g. Maesa), or ebracteolate (mostly); small; regular; mostly 4–5 merous; cyclic; tetracyclic. Free hypanthium absent.
Perianth with distinct calyx and corolla; 6–12; 2 whorled; isomerous. Calyx (3–)4–5(–6); 1 whorled; polysepalous, or gamosepalous (often basally connate); regular; imbricate, or contorted, or valvate. Corolla (3–)4–5(–6); 1 whorled; gamopetalous (usually), or polypetalous (e.g. Embelia); imbricate, or contorted, or valvate; regular; not fleshy.
Androecium (3–)4–5(–6). Androecial members free of the perianth (rarely), or adnate (to the corolla tube); free of one another, or coherent (sometimes); when coherent, 1 adelphous (the filaments connate); 1 whorled. Androecium of male-fertile flowers exclusively of fertile stamens, or including staminodes (occasionally). Stamens (3–)4–5(–6); usually isomerous with the perianth; alternisepalous; opposite the corolla members. Anthers cohering (Amblyanthus), or separate from one another; dehiscing via longitudinal slits, or dehiscing via pores; introrse; tetrasporangiate. Endothecium not developing fibrous thickenings (in Badula). Pollen grains aperturate; 3(–5) aperturate; colpate, or colporate (col(por)oidate, sometimes 4-rupate); 2-celled (in Ardisia and Wallenia).
Gynoecium 3–5(–6) carpelled. Carpels isomerous with the perianth, or reduced in number relative to the perianth, or increased in number relative to the perianth. The pistil 1 celled. Gynoecium syncarpous; eu-syncarpous; superior (nearly always), or partly inferior (Maesa). Ovary 1 locular. Gynoecium stylate. Styles 1; attenuate from the ovary; apical. Stigmas 1; dry type; papillate; Group II type. Placentation basal, or free central. Ovules in the single cavity 3–100 (few to many); sunken in the placenta; ascending; non-arillate; anatropous; bitegmic; tenuinucellate. Outer integument not contributing to the micropyle. Endothelium differentiated. Embryo-sac development Polygonum-type. Endosperm formation nuclear. Embryogeny onagrad (?).
Fruit fleshy; indehiscent; a berry, or a drupe; 1 seeded (usually), or 3–100 seeded (many only in Maesa). Seeds endospermic. Endosperm oily. Seeds with amyloid. Cotyledons 2. Embryo achlorophyllous (1/3); straight to curved. Polyembryony recorded (e.g. Ardisia).
Seedling. Germination phanerocotylar, or cryptocotylar.
Physiology, phytochemistry. Anatomy non-C4 type (Ardisia). Sugars transported as sucrose (in Ardisia). Not cyanogenic. Alkaloids present (rarely), or absent. Iridoids not detected. Saponins/sapogenins present, or absent. Proanthocyanidins present (usually), or absent; cyanidin, or delphinidin, or cyanidin and delphinidin. Flavonols present; kaempferol, or kaempferol and quercetin, or myricetin. Ellagic acid absent (5 genera, 5 species).
Geography, cytology. Temperate to tropical. Pantropical, subtropical and extending North to Japan and Florida, and South to New Zealand. X = 10–13, 23.
Taxonomy. Subclass Dicotyledonae; Tenuinucelli. Dahlgrens Superorder Primuliflorae; Primulales. Cronquists Subclass Dilleniidae; Primulales. APG 3 core angiosperms; core eudicot; Superorder Asteranae; Order Ericales (as a synonym of Primulaceae).
Species 1000. Genera about 35; Amblyanthopsis, Amblyanthus, Antistrophe, Ardisia, Badula, Conandrium, Ctenardisia, Cybianthus, Discocalyx, Elingamita, Embelia, Emblemantha, Fittingia, Geissanthus, Heberdenia, Hymenandra, Labisia, Loheria, Maesa, Monoporus, Myrsine, Oncostemum, Parathesis, Pleiomeris, Rapanea, Sadiria, Solonia, Stylogyne, Tapeinosperma, Tetrardisia, Vegaea, Wallenia.
General remarks. After analysing a combination of nucleic acid sequences from the chloroplast genes rbcL, ndhF andatpB, Källersjö et al (2000) supported earlier claims that Primulaceae and Myrsinaceae as traditionally circumscribed are paraphyletic, with (e.g.) Anagallis, Ardisiandra, Coris, Lysimachia and Trientalis belonging in myrsinaceous rather than primulaceous clades. Rather than merging all the genera into one supposedly monophyletc family, Anderberg et al (2000) proposed raising Maesa to family rank, and adjusting the contents of Myrsinaceae s. lat. and Primulaceae s. str. In the absence of any attempts by modern re-classifiers to prepare the requisite comparative descriptions, the classical family circumscriptions are largely retained here. See comments under Primulaceae and Theophrastaceae.
Illustrations. • Technical details: Maesa (Thonner). • Technical details (Maesa, Ardisia). • Ardisia chinensis: as A. odontophylla, Bot. Reg. 1892 (1836). • Myrsine dependens, as M. myrtoides: Hook. Ic. Pl. 9 (1852). • Rapanea salicina (as Suttonia): Hooker, Fl. Novae-Zelandiae (1853). • Rapanea nummularia (as Suttonia): Hooker, Fl. Novae-Zelandiae (1853). • Leaf hairs of Ardisia, with Aegicerataceae (Aegiceras) and Theophrastaceae (Clavija and Jacquinia): Solereder, 1908.
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’.