The families of flowering plants
Including Coptaceae (Gregory) Löve & Löve, Helleboraceae von Vest, Nigellaceae J.G. Agardh, Thalictraceae Rafin.; excluding Glaucidiaceae, Hydrastidaceae, Kingdoniaceae.
Habit and leaf form. Herbs (mostly), or shrubs (Xanthorhiza), or lianas (Clematis); without essential oils. Annual to perennial; with a basal aggregation of leaves, or without conspicuous aggregations of leaves; often rhizomatous, or tuberous. Self supporting, or climbing. Hydrophytic to mesophytic; when hydrophytic, rooted. Leaves of aquatics emergent, or submerged, or submerged and floating. Heterophyllous (commonly, when hydrophytic), or not heterophyllous. Leaves alternate (usually), or opposite (Clematis); usually spiral; petiolate; sheathing. Leaf sheaths with free margins. Leaves not gland-dotted; without marked odour, or foetid; simple, or compound; peltate (more or less, occasionally), or not peltate; epulvinate; when compound pinnate, or palmate, or bipinnate, or multiply compound. Lamina when simple dissected, or entire; when simple/dissected palmatifid, or much-divided; pinnately veined, or palmately veined; cross-venulate (usually), or without cross-venules. Leaves stipulate (commonly rather conspicuously so), or exstipulate. Stipules intrapetiolar. Lamina margins entire, or crenate, or serrate, or dentate. Leaf development not graminaceous.
Leaf anatomy. Hydathodes present (occasionally), or absent. Minor leaf veins without phloem transfer cells (Helleborus, Ranunculus).
Axial (stem, wood) anatomy. Young stems with solid internodes, or with hollow internodes. Pith homogeneous, or heterogeneous. Cork cambium present, or absent. Nodes unilacunar (rarely), or tri-lacunar to multilacunar. Primary vascular tissues comprising a ring of bundles, or comprising two or more rings of bundles, or consisting of scattered bundles; collateral. Secondary thickening absent (commonly), or developing from a conventional cambial ring. The axial xylem with vessels.
The vessels in Clematis, clustered, or in tangential arcs. The vessel end-walls simple (at least usually). The axial xylem with fibre tracheids; with libriform fibres, or without libriform fibres. The wood partially storied (VP), or not storied.
Reproductive type, pollination. Plants hermaphrodite, or dioecious (rarely, by suppression). Floral nectaries present (usually), or absent (e.g. Anemone, Clematis, Thalictrum). Nectar secretion from the perianth, or from the androecium (from the bases of the petals, considered staminodial in origin). Pollination entomophilous (usually attracting insects by either nectar or pollen), or anemophilous (e.g. Thalictrum).
Inflorescence, floral, fruit and seed morphology. Flowers solitary (often pedunculate), or aggregated in inflorescences; in cymes, or in racemes, or in panicles. The ultimate inflorescence units cymose. Inflorescences scapiflorous, or not scapiflorous; terminal, or axillary; various, but determinate. Flowers small to medium-sized; regular (usually), or somewhat irregular to very irregular (Aconitum etc.); cyclic (Aquilegia), or partially acyclic, or acyclic. When more or less acyclic the perianth acyclic, the androecium acyclic, and the gynoecium acyclic, or the androecium acyclic and the gynoecium acyclic, or the gynoecium acyclic. Floral receptacle not markedly hollowed (convex or elongated). Free hypanthium absent. Hypogynous disk absent.
Perianth with distinct calyx and corolla, or sequentially intergrading from sepals to petals, or sepaline, or petaline; 5–50 (to many); free, or joined. Calyx when definable (3–)5–8 (or more, often becoming petaloid); polysepalous, or partially gamosepalous, or gamosepalous; spurred, or neither appendaged nor spurred; persistent, or not persistent; imbricate, or valvate. Corolla when definable 3–50 (to many perhaps staminodal in origin); polypetalous, or partially gamopetalous, or gamopetalous; imbricate; green, or white, or yellow, or red (e.g. some Aquilegia species), or purple, or blue; spurred (occasionally), or not spurred (or with little tubular nectariferous petals). Petals clawed, or sessile.
Androecium 15–100 (usually many). Androecial members unbranched; maturing centripetally; free of the perianth; free of one another; 1–13 whorled (or spiralled). Androecium including staminodes (usually several or many, if nectaries of various kinds between perianth and stamens are interpreted as such), or exclusively of fertile stamens. Staminodes external to the fertile stamens; petaloid, or non-petaloid. Stamens (5–)10–100 (usually many). Anthers adnate; non-versatile; dehiscing via longitudinal slits, or dehiscing by longitudinal valves (e.g. Trautvetteria); extrorse, or latrorse; tetrasporangiate. Endothecium developing fibrous thickenings. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral, or decussate. Anther wall initially with one middle layer, or initially with more than one middle layer (1 or 2); of the dicot type. Tapetum glandular. Pollen grains aperturate (usually), or nonaperturate (Souliea); (2–)3 aperturate; variously colpate, or foraminate, or rugate, or spiraperturate (but not colporate, not even colporoidate); 2-celled (recorded in 9 genera).
Gynoecium (1–)3–100 carpelled (i.e. to many). The pistil when other than apocarpous, 1 celled, or 3–5 celled. Gynoecium monomerous, or apocarpous, or syncarpous; of one carpel (Actaea), or eu-apocarpous (nearly always), or semicarpous (carpels sometimes more or less connate, e.g. Aquilegia), or synovarious (e.g. Nigella); superior. Carpel non-stylate, or stylate; apically stigmatic; (when monomeric or apocarpous) 1 ovuled, or 2–100 ovuled (several to many). Placentation when monomeric or apocarpous marginal, or basal. Ovary when syncarpous, 3–5 locular. Styles when syncarpous, 3–5; apical. Stigmas dry type; papillate, or non-papillate; Group II type. Placentation when syncarpous, axile. Ovules when syncarpous, 3–15 per locule (?); pendulous, or horizontal, or ascending; with ventral raphe to with dorsal raphe; non-arillate; hemianatropous, or anatropous; unitegmic, or bitegmic; crassinucellate (when bitegmic), or pseudocrassinucellate (when unitegmic). Embryo-sac development Polygonum-type, or Allium-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3 (sometimes multinucleate); proliferating (rarely), or not proliferating. Synergids pear-shaped, or hooked (sometimes with filiform apparatus). Endosperm formation nuclear. Embryogeny onagrad (or undifferentiated).
Fruit non-fleshy (usually), or fleshy (rarely); an aggregate (usually), or not an aggregate. The fruiting carpel dehiscent, or indehiscent; a follicle, or an achene, or baccate (Actaea). Fruit when syncarpous, dehiscent; when syncarpous a capsule (e.g. Nigella). Seeds copiously endospermic. Endosperm oily. Seeds without amyloid. Embryo rudimentary at the time of seed release to weakly differentiated, or well differentiated. Cotyledons 1 (occasionally), or 2 (often connate). Embryo achlorophyllous (14/25); straight.
Seedling. Germination phanerocotylar, or cryptocotylar.
Physiology, phytochemistry. C3. C3 physiology recorded directly in Anemone, Clematis, Delphinium, Ranunculus. Anatomy non-C4 type (Anemone, Delphinium, Ranunculus). Cyanogenic, or not cyanogenic. Cynogenic constituents tyrosine-derived (including triglochinin). Alkaloids present (mostly), or absent. Berberine present (in the rhizomes of Coptis and Xanthorhiza), or absent. Arbutin absent. Iridoids not detected. Saponins/sapogenins present, or absent. Proanthocyanidins absent (usually), or present (e.g. Clematis); when present, cyanidin. Flavonols present (mostly), or absent; kaempferol, or quercetin, or kaempferol and quercetin (nearly always both). Ellagic acid absent (13 species, 9 genera). Aluminium accumulation not found. Sieve-tube plastids S-type.
Geography, cytology. Frigid zone to tropical. Cosmopolitan, concentrated in the North temperate.
Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgrens Superorder Ranunculiflorae; Ranunculales. Cronquists Subclass Magnoliidae; Ranunculales. APG 3 core angiosperms; peripheral eudicot; Superorder Ranunculanae; Order Ranunculales.
Species 1500. Genera about 50; Aconitum, Actaea, Adonis, Anemone, Anemonopsis, Aquilegia, Archiclematis, Asteropyrum, Barneoudia, Beesia, Calathodes, Callianthemum, Caltha, Ceratocephala, Cimicifuga, Clematis, Clematopsis, Consolida, Coptis, Delphinium, Dichocarpum, Enemion, Eranthis, Hamadryas, Helleborus, Hepatica, Isopyrum, Knowltonia, Komaroffia, Krapfia, Kumlienia, Laccopetalum, Leptopyrum, Megaleranthis, Metanomone, Miyakea, Myosurus, Naravelia, Nigella, Oreithales, Paraquilegia, Paroxygraphis, Pulsatilla, Ranunculus, Semiaquilegia, Souliea, Thalictrum, Trautvetteria, Trollius, Urophysa, Xanthorhiza.
Economic uses, etc. Many cultivated ornamentals (Ranunculus, Anemone, Helleborus, Trollius, Delphinium, Aconitum, Aquilegia, some (e.g. Aconitum) supply poisonous narcotic drugs.
When daisies pied, and
And cuckoo-buds of yellow hue,
And Lady-smocks all silver-white,
Do paint the meadows with delight
(Love Labours Lost, v., 2 - cuckoo-buds = buttercups)
buttercups, the little childrens dower
- Far brighter than this gaudy melon-flower!
(Rober Browning, Home Thoughts, from Abroad)
Both milkmaids shouts and herdsmans call
Have vanishd with the green
The king kups yellow shades and all
Shall never more be seen
For all the cropping that does grow
Will so efface the scene
(John Clare 1821, Helpston Green a moist part, if this really refers to Caltha palustris)
Whatsoever man fasting eats this
wort, leaves his life laughing
(Quoted by Gilmour (British Botanists, 1956), from a medieval manuscript - of Ranunculus sceleratus, with reference to facial strictus induced by its toxin)
Illustrations. • Technical details: Aconitum. • Technical details: Actaea. • Technical details: Adonis, Myosurus. • Technical details: Anemone pulsatilla. • Technical details: Anemone vesicatoria. • Technical details: Aquilegia, Nigella. • Technical details: Caltha. • Technical details: Clematis. • Technical details: Delphinium. • Technical details: Helleborus. • Technical details: Nigella. • Technical details: Ranunculus, Ceratocephala. • Technical details: Ranunculus. • Technical details: Thalictrum. • Technical details: Trollius. • Adonis annua (as A. autumnalis): Eng. Bot. 14, 1863. • Aconitum anglicum (J.E. Sowerby, 1861). • Aconitum napellus: Eng. Bot. 48, 1863. • Actaea spicata: Eng. Bot. 49, 1863. • Anemone appenina, A. nemorosa and Pulsatilla vulgaris (as Anemone pulsatilla): Eng. Bot. 9–11, 1863. • Anemone ranunculoides: Eng. Bot. 12, 1863. • Anemone sylvestris: Bot. Mag. 2, 1788. • Anemone vitifolia: Bot. Reg. 1385, 1830. • Aquilegia glauca: Bot. Reg. xxvi, 46 (1840). • Aquilegia vulgaris: Eng. Bot. 46, 1863. • Caltha palustris (with var. radicans as C. radicans): Eng. Bot. 40 and 41, 1863. • Clematis chlorantha: Bot. Reg. 1234, 1829. • Clematis integrifolia: Bot. Mag. 2, 1788. • Clematis pubescens (photos). • Clematis vitalba: Eng. Bot. 1, 1863. • Delphinium decorum: Bot. Reg. xxvi, 64 (1840). • cf. Delphinium elatum subsp. montanum: as Delphinium montanum, Bot. Reg. 1936, 1837. • cf. Delphinium elatum subsp. elatum: as D. intermedium, Bot. Reg. 1963, 1837. • Delphinium villosum var. laxiflorum: as D. laxiflorum, Bot. Reg. XXIV, 30 (1838). • Eranthis hyemalis: Eng. Bot. 43, 1863. • Helleborus corsicus: as H. lividus, Bot. Reg. XXIV, 54 (1838). • Helleborus foetidus (J. E. Sowerby, 1861). • Helleborus lividus: Bot. Mag. 2, 1788. • Helleborus cf. orientalis: Bot. Reg. 34, 1842. • Helleborus viridis and H. foetidus: Eng. Bot. 44 and 45, 1863. • Helleborus viridis (J. E. Sowerby, 1861). • Ranunculus acris and R. repens: Eng. Bot. 33 and 34, 1863. • Ranunculus aquatilis (as R. heterophyllus) and R. trichophyllus: Eng. Bot. 19 and 21, 1863. • Ranunculus baudotii and R. tripartitus: Eng. Bot. 22 abd 24, 1863. • Ranunculus circinatus and R. fluitans: Eng. Bot. 15–16, 1863. • Ranunculus flammula and R. auricomus: Eng. Bot. 29 and 32, 1863. • Ranunculus hirsutus and R. lingua: Eng. Bot. 36 and 31, 1863. • Ranunculus omiophyllus (as R. lenormandi): Eng. Bot. 25, 1863. • Ranunculus peltatus: Eng. Bot. 17, 1863. • Ranunculus sceleratus and R. ophioglossifolius: Eng. Bot. 27 and 28, 1863. • Thalictrum alpinum and T. flavum: Eng. Bot. 2 and 8, 1863. • Trollius acaulis: Bot. Reg. 29, 32 (1843). • Trollius europaeus: Eng. Bot. 42, 1863. • British: Myosurus minimus (B. Ent.). • British: Anemone, Adonis (B. Ent. compilation). • British: Clematis, Delphinium, Aquilegia (B. Ent. compilation). • British: Trollius, Helleborus, Thalictrum (B. Ent. compilation). • British: Ranunculus species (B. Ent. compilation). • British: Ranunculus species, Caltha (B. Ent. compilation).
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: 19th October 2016. delta-intkey.com’.