Including Aparin(ac)eae Hoffmgg. & Link, Cinchonaceae Lindl., Coffeaceae J.G. Agardh, Gardeniaceae Dum., Lygodysodeaceae Bartl., Operculariaceae Dum.
Excluding Henriqueziaceae, Naucleaceae
Habit and leaf form. Trees and shrubs (mostly), or lianas, or herbs (then mostly with tetragonous, knotted stems). Self supporting, or epiphytic, or climbing. Helophytic, or mesophytic, or xerophytic, or hydrophytic (Limnosipanea). Leaves opposite (nearly always, and decussate), or whorled (or at least ostensibly so, seemingly representing paired leaves with enlarged, leaflike interpetiolar stipules - and rarely ostensibly alternate, through suppression of one member of each pair); petiolate to sessile; connate (often, via the stipules), or not connate; gland-dotted, or not gland-dotted; simple; epulvinate. Lamina entire; one-veined, or pinnately veined; cross-venulate. Leaves stipulate. Stipules interpetiolar (usually), or intrapetiolar. Lamina margins entire, or serrate. Leaves without a persistent basal meristem. Domatia recorded (frequently), or not recorded; represented by pits (mostly), or pockets, or hair tufts.
General anatomy. Plants with `crystal sand' (very commonly), or without `crystal sand'.
Leaf anatomy. Stomata typically paracytic.
Lamina dorsiventral (nearly always), or centric (rarely). The mesophyll with sclerencymatous idioblasts, or without sclerenchymatous idioblasts. Minor leaf veins with phloem transfer cells (5 genera - Asperula, Galium, Phuopsis, Rubia, Sherardia), or without phloem transfer cells (20 genera, e.g. Coprosma, Cinchona, Coffea, Gardenia, Hoffmannia, Ixora, Pavetta, Randia).
Stem anatomy. Young stems tetragonal. Cork cambium present, or absent (e.g. in Galieae); initially deep-seated, or superficial. Nodes unilacunar, or tri-lacunar. Internal phloem absent. Secondary thickening developing from a conventional cambial ring (nearly always), or anomalous (concentric cambia recorded only in Basanacantha, `Chiococca'); via concentric cambia. `Included' phloem present (Basanacantha), or absent. Xylem with tracheids; with fibre tracheids; with vessels. Vessel end-walls mostly simple. Vessels with vestured pits. Wood parenchyma apotracheal, or paratracheal.
Reproductive type, pollination. Hermaphrodite (usually), or monoecious, or polygamomonoecious, or dioecious (e.g. Coprosma); plants often heterostylous. Entomophilous.
Inflorescence, floral, fruit and seed morphology. Flowers aggregated in `inflorescences', or solitary (less often); in cymes, or in panicles, or in verticils, or in heads (rarely, e.g. Morindeae, Gardenieae). The terminal inflorescence unit cymose. Flowers small, or medium-sized; regular; mostly 4 merous, or 5 merous; cyclic; tetracyclic. Free hypanthium absent.
Perianth with distinct calyx and corolla, or petaline (the calyx often absent or almost so); 7-15; 2 whorled, or 1 whorled; the two whorls isomerous. Calyx when detectable 4, or 5; 1 whorled; polysepalous (but epigynous), or gamosepalous (the lobes varying from practically lacking to enlarged and brightly coloured); when gamosepalous, entire, or lobulate, or lobed, or toothed; regular; open in bud. Corolla (3-)4, or 5, or 8-10; 1 whorled; gamopetalous; imbricate, or valvate, or contorted; tubular, or hypocrateriform; regular, or bilabiate (rarely).
Androecium 4, or 5. Androecial members adnate (to the corolla tube, or attached at its very base in Coprosma); free of one another (usually), or coherent (occasionally, via the anthers); 1 - whorled. Androecium exclusively of fertile stamens. Stamens 4, or 5; isomerous with the perianth; oppositisepalous. Anthers cohering (occasionally), or connivent, or separate from one another; dehiscing via longitudinal slits; introrse; tetrasporangiate; slightly appendaged (rarely?), or unappendaged. The anther appendages observed in Coprosma apical (by slight extension of the connective). 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; of the `dicot' type. Tapetum glandular. Pollen shed in aggregates (in the Gardenieae), or shed as single grains (mostly); when aggregated, in tetrads. Pollen grains aperturate, or nonaperturate (rarely); 3(-4) - aperturate (mostly), or 4-12 - aperturate; colpate (Paederieae, Galieae, Spermacoceae p.p.), or porate (Gardenieae), or colporate (polycolporate in Richardsonia), or foraminate; 2-celled, or 3-celled.
Gynoecium 2(-9). Carpels usually reduced in number relative to the perianth. Gynoecium syncarpous; synovarious to eu-syncarpous; inferior (nearly always), or superior (only Gaertnera and Pagamea). Ovary 1 locular (rarely e.g. Gardenia), or 2(-9) locular. Gynoecium when bilocular (i.e. usually), transverse. Epigynous disk often present. Gynoecium stylate. Styles 1 (often simple), or 2(-5); free, or partially joined; attenuate from the ovary, or from a depression at the top of the ovary; apical; shorter than the mature ovary to much longer than the mature ovary. Stigmas wet type, or dry type; papillate, or non-papillate; Group II type and Group IV type. Placentation when unilocular, parietal; when two or more locular, axile. Ovules in the single cavity when unilocular, 6-100 (`many'); 1-50 per locule (commonly one (Rubioideae), to `many'); pendulous, or horizontal, or ascending; anatropous, or hemianatropous; unitegmic; tenuinucellate. Endothelium not differentiated. Embryo-sac development Polygonum-type. Antipodal cells formed; 3; proliferating (occasionally), or not proliferating; ephemeral, or persistent. Endosperm formation nuclear. Embryogeny solanad.
Fruit fleshy, or non-fleshy; dehiscent, or indehiscent, or a schizocarp; when schizocarpic comprising achenes, or comprising mericarps, or comprising nutlets, or comprising drupelets (?); when non-schizocarpic a capsule, or a berry, or a drupe. Capsules septicidal, or loculicidal. Fruit 1-30 seeded. Seeds endospermic (usually), or non-endospermic (Guettardinae). Endosperm ruminate, or not ruminate; when present, oily. Seeds winged (rarely), or wingless. Cotyledons 2. Embryo achlorophyllous (8/11); straight (usually), or curved.
Seedling. Germination phanerocotylar, or cryptocotylar.
Physiology, biochemistry. Cyanogenic, or not cyanogenic. Cynogenic constituents phenylalanine-derived (?). Alkaloids present (commonly), or absent. Iridoids recorded; carbocyclic and seco-compounds. Proanthocyanidins present, or absent. Flavonols present (mostly), or absent; kaempferol and quercetin (mostly), or kaempferol, or quercetin. Ellagic acid absent (9 species, 9 gebera). Arbutin present. Ursolic acid present. Saponins/sapogenins present (occasionally), or absent. Aluminium accumulation demonstrated, or not found. Inulin recorded (Cinchona, Gibbs 1974). C3. C3 recorded in Borreria, Coprosma, Crucianella, Galium, Richardia. Anatomy non-C4 type (Cephalanthus, Crucianella, Diodia, Galium).
Geography, cytology. Frigid zone to tropical. Cosmopolitan. X = (6-)9/11(-17).
Taxonomy. Subclass Dicotyledonae; Tenuinucelli. Dahlgren's Superorder Gentianiflorae; Gentianales. Cronquist's Subclass Asteridae; Rubiales. Takhtajan's Subclass Asteridae; Gentiananae; Gentianales. Species 6000. Genera about 600; Asperula, Bouvardia, Chiococca, Cinchona, Coffea, Condaminea, Coprosma, Cosmibuena, Crucianella, Cuviera, Duroia, Faramea, Galium, Gardenia, Guettarda, Houstonia, Ixora, Lasianthus, Mitchella, Morinda, Myrmecodia, Mussaenda, Nertera, Oldenlandia, Paederia, Pavetta, Pentas, Plectronia, Posoqueria, Psychotria, Rondeletia, Rubia, Rudgea, Sherardia, Uragoga, etc.
Economic uses, etc. Sources of coffee (Coffea), quinine (Cinchona, ipecac (Cephaelis), and many ornamentals (e.g. Gardenia, Rubia, Mitchella, Coprosma).
Illustrations. rubia483.gif rubia761.gif rubia517.gif rubia484.gif
Additional, to be intercalated. Stipules with colleters (secreting mucilage) (typically).
