The families of flowering plants
Including Chironiaceae Horan.Excluding Saccifoliaceae
Habit and leaf form. Herbs (mostly, often with dichotomous branching), or shrubs (a few); non-laticiferous and without coloured juice. Leaves well developed, or much reduced. Plants autotrophic, or saprophytic. Annual, or biennial; with a basal aggregation of leaves, or with neither basal nor terminal aggregations of leaves. Climbing (Crawfurdia ~ Gentiana p.p.), or self supporting (usually). Helophytic, or mesophytic (often alpine). Leaves opposite (and decussate, usually), or whorled (rarely Swertia), or alternate (seldom); rarely (i.e. when alternate) spiral; herbaceous, or membranous (in mycotrophic species); petiolate to sessile; connate (often), or not connate (but then often connected by a transverse line); simple; epulvinate. Lamina entire; pinnately veined, or palmately veined. Leaves exstipulate (but the opposing leaf bases sometimes connected by a transverse line). Lamina margins usually entire. Leaves without a persistent basal meristem.
Leaf anatomy. Abaxial epidermis commonly papillose. Mucilaginous epidermis present, or absent. Stomata present; anomocytic (commonly), or anisocytic.
Adaxial hypodermis present (sometimes mucilaginous), or absent. Lamina dorsiventral, or isobilateral; without secretory cavities. The mesophyll containing mucilage cells (or even consisting of mucilaginous cells), or not containing mucilage cells. Minor leaf veins without phloem transfer cells (4 genera).
Stem anatomy. Young stems cylindrical, or oval in section. Nodes unilacunar, or tri-lacunar, or multilacunar. Primary vascular tissue centrifugal, or bicollateral (often). Cortical bundles absent. Medullary bundles present (commonly), or absent. Internal phloem present. Secondary thickening developing from a conventional cambial ring, or anomalous (Gentianoideae); from a single cambial ring. Included phloem present (commonly), or absent. Xylem with libriform fibres; with vessels. Vessel end-walls simple. Sieve-tube plastids S-type.
Reproductive type, pollination. Plants hermaphrodite (usually), or polygamomonoecious (rarely); homostylous, or heterostylous. Pollination entomophilous.
Inflorescence, floral, fruit and seed morphology. Flowers solitary, or aggregated in inflorescences; in cymes. The ultimate inflorescence unit (when flowers aggregated) nearly always cymose. Inflorescences terminal, or axillary; usually a simple or compound dichasium. Flowers bracteate, or ebracteate; bracteolate, or ebracteolate; small, or medium-sized; regular, or somewhat irregular to very irregular. The floral irregularity (when manifest) involving the perianth (K only), or involving the perianth and involving the androecium. Flowers 4–5(–12) merous; cyclic; tetracyclic. Floral receptacle developing a gynophore, or with neither androphore nor gynophore. Free hypanthium absent. Hypogynous disk present, or absent; when present, of separate members, or annular.
Perianth with distinct calyx and corolla; 8–10(–24); 2 whorled; isomerous. Calyx 4–5(–12); 1 whorled; polysepalous (rarely), or gamosepalous; entire (occasionally), or lobulate, or blunt-lobed (or polysepalous); regular, or bilabiate (rarely); imbricate (usually), or open in bud (sometimes). Epicalyx present, or absent. Corolla 4–5(–12); 1 whorled; appendiculate (often with scales or nectary pits inside), or not appendiculate; gamopetalous; contorted (commonly), or imbricate; campanulate, or funnel-shaped, or cyathiform, or rotate, or hypocrateriform; regular; white, or pink, or purple, or blue (often showy); not fleshy.
Androecium 4–5(–12) (as many as C). Androecial members adnate (to the tube); all equal, or markedly unequal (sometimes declinate); free of one another (usually), or coherent (forming a tube in some saprophytic genera); 1 whorled. Androecium exclusively of fertile stamens (nearly always), or including staminodes. Staminodes when present, 1–4; in the same series as the fertile stamens. Stamens (1–)4–5(–12); inserted near the base of the corolla tube, or midway down the corolla tube, or in the throat of the corolla tube; isomerous with the perianth (nearly always), or reduced in number relative to the adjacent perianth (very occasionally with some stamens staminodal or wanting); oppositisepalous. Anthers dorsifixed (usually), or basifixed; versatile (when dorsifixed), or non-versatile; dehiscing via longitudinal slits, or dehiscing via pores (rarely via apical pores); usually introrse; tetrasporangiate; appendaged (sometimes, with glands), or unappendaged. Endothecium developing fibrous thickenings (usually), or not developing fibrous thickenings (rarely, e.g. Cotylanthera). Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral. Anther wall of the dicot type. Tapetum amoeboid, or glandular. Pollen polysiphonous, or monosiphonous; shed in aggregates (Helieae), or shed as single grains; when aggregated, in tetrads. Pollen grains aperturate; (1–)2 aperturate, or 3(–4) aperturate; porate (and sometimes ruporate), or colporate; 2-celled (4 genera), or 3-celled (4 genera).
Gynoecium 2 carpelled. The pistil 1 celled, or 2 celled (rarely). Gynoecium syncarpous; synstylovarious to eu-syncarpous; superior. Ovary 1 locular (usually), or 2 locular (rarely). Gynoecium median, or transverse; stylate, or non-stylate (Lomatogonium, with stigmas decurrent along the sides of the ovary). Styles usually 1; attenuate from the ovary; apical. Stigmas 1, or 2; 1 lobed, or 2 lobed; wet type; papillate; Group III type. Placentation parietal (usually), or free central (rarely); rarely, when bilocular axile. Ovules in the single cavity when unilocular, 15–100 (i.e. many); when ovary bilocular, 15–50 per locule (i.e. many); horizontal; non-arillate; anatropous (usually), or orthotropous (e.g. Leiphaimos); unitegmic; tenuinucellate. Endothelium differentiated. Embryo-sac development Polygonum-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; proliferating (e.g. Swertia), or not proliferating; usually ephemeral. Synergids pear-shaped. Endosperm formation cellular, or nuclear. Embryogeny solanad.
Fruit fleshy (rarely), or non-fleshy; dehiscent (usually), or indehiscent (rarely); a capsule (usually), or a berry (rarely). Capsules septicidal. Seeds usually copiously endospermic (but scantily so in saprophytic forms). Endosperm oily. Seeds winged, or wingless. Embryo rudimentary at the time of seed release to weakly differentiated (in saprophytes), or well differentiated. Cotyledons when developed, 2. Embryo chlorophyllous (Swertia iberica), or achlorophyllous (2/2); straight.
Seedling. Germination phanerocotylar.
Physiology, biochemistry. Cyanogenic (sometimes), or not cyanogenic. Iridoids detected; Route I type (normal and seco). Proanthocyanidins absent. Ellagic acid absent (2 genera, 3 species). Saponins/sapogenins present (doubtfully), or absent. Aluminium accumulation demonstrated (in a few genera). Anatomy non-C4 type (Blackstonia).
Peculiar feature. Lamina tip not abaxially pouched.
Geography, cytology. Frigid zone to tropical. Cosmopolitan. X = 5–13(+).
Taxonomy. Subclass Dicotyledonae; Tenuinucelli. Dahlgrens Superorder Gentianiflorae; Gentianales. Cronquists Subclass Asteridae; Gentianales. APG 3 core angiosperms; core eudicot; Superorder Asteranae; lamiid; Order Gentianales.
Species 900. Genera about 80; Bartonia, Belmontia, Bisgoeppertia, Blackstonia, Canscora, Celiantha, Centaureum, Chironia, Chorisepalum, Cicendia, Comastoma, Congolanthus, Cotylanthera, Coutoubea, Cracosna, Crawfurdia, Curtia, Deianira, Djaloniella, Enicostema, Eustoma, Exaculum, Exacum, Faroa, Frasera, Gentiana, Gentianella, Gentianopsis, Gentianothamnus, Halenia, Hockinia, Hoppea, Irlbachia, Ixanthus, Jaeschkea, Karina, Latouchea, Lehmanniella, Lisianthius, Lomatogoniopsis, Lomatogonium, Macrocarpaea, Megacodon, Microrphium, Monodiella, Neblinantha, Neurotheca, Obolaria, Oreonesion, Ornichia, Orphium, Parajaeschkea, Prepusa, Pterygocalyx, Pycnosphaera, Rogersonanthus, Sabatia, Schinziella, Schultesia, Sebaea, Senaea, Sipapoantha, Swertia, Symbolanthus, Symphyllophyton, Tachia, Tachiadenus, Tapeinostemon, Tetrapollinia, Tripterospermum, Urogentias, Veratrilla, Voyria, Voyriella, Wurdackanthus, Zonanthus, Zygostigma.
Illustrations. • Technical details: Gentiana, Erythraea = Centaurium. • Centaurium, Gentiana, Gentianella, Blackstonia (B. Ent. compilation). • Blackstonia perfoliata (as Chlora perfoliata): Eng. Bot. 913 (1866). • Centaurium erythraea (as Erythraea centaurium: Eng. Bot. 909 (1866). • Chironia peduncularis: Bot. Reg. 1803, 1836. • Gentiana acaulis: Bot. Mag. 2, 1788. • Gentiana pneumonanthe: Eng. Bot. 914 (1866). • Gentiana verna: Eng. Bot. 915 (1866). • Orphium frutescens: Bot. Mag. 2, 1788.
Who bade the sun
Clothe you with rainbows? Who, with living flowers
Of loveliest blue, spread garlands at your feet?
(Coleridge, To Chamouni - alluding to gentians)
This description is offered for casual browsing only. We strongly advise against extracting comparative information from it. This is much more easily achieved using 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).
Cite this publication as: ‘Watson, L., and Dallwitz, M.J. 1992 onwards. The families of flowering plants: descriptions, illustrations, identification, and information retrieval. Version: 19th December 2012. http://delta-intkey.com’.