The families of flowering plants.
IncludingArdisiaceae Juss., Embelieae (Embeliaceae) J.G. Agardh, Maesaceae (A.DC.) Anderb, Ståhl & Källersjö, Ophiospermes (Ophiospermae) Vent., Ophiospermataceae KuntzeExcluding Aegicerataceae
Habit and leaf form. Trees and shrubs, or lianas (a few — and a few sub-herbaceous); with coloured juice, or non-laticiferous and without coloured juice; 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. Leaves without a persistent basal meristem. Domatia occurring in the family (3 genera); manifested as pockets, or hair tufts.
Leaf anatomy. Mucilaginous epidermis present, or absent.
Lamina with secretory cavities (usually), or without secretory cavities. Secretory cavities containing resin (yellow or reddish brown); schizogenous. The mesophyll often with hypodermal fibres. Minor leaf veins without phloem transfer cells (Ardisia, Suttonia (= Rapanea)).
Stem anatomy. Secretory cavities commonly present; with resin (yellow or reddish-brown). Cork cambium present; initially superficial. Nodes unilacunar. Secondary thickening developing from a conventional cambial ring. Xylem with fibre tracheids, or without fibre tracheids; with libriform fibres; with vessels. Vessel end-walls simple, or scalariform and simple. Vessels without vestured pits. Wood parenchyma paratracheal.
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 unit 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.
Perianthwith 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). Ovary1 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 (?).
Fruitfleshy; indehiscent; a berry, or a drupe; 1 seeded (usually), or 3–100 seeded (‘many’ only in Maesa). Seeds endospermic. Endosperm oily. Seedswith amyloid. Cotyledons 2. Embryo achlorophyllous (1/3); straight to curved. Polyembryony recorded (e.g. Ardisia).
Seedling.Germination phanerocotylar, or cryptocotylar.
Physiology, biochemistry. Not cyanogenic. Alkaloids present (rarely), or absent. Iridoids not detected. 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). Saponins/sapogenins present, or absent. Sugars transported as sucrose (in Ardisia). Anatomy non-C4 type (Ardisia).
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. Dahlgren’s Superorder Primuliflorae; Primulales. Cronquist’s 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. See comments under Primulaceae and Theophrastaceae.