The families of flowering plants
~ Malvaceae sensu lato.
Including Sparmanniaceae J.G. Agardh; excluding Elaeocarpaceae, Muntingiaceae, Oceanopapaveraceae.
Habit and leaf form. Trees and shrubs, or herbs (rarely); non-laticiferous, without coloured juice; leptocaul. Mesophytic. Leaves alternate; spiral, or distichous (often, or at least two ranked on the upper half of the shoot); petiolate; non-sheathing; simple. Lamina dissected, or entire; conspicuously asymmetric (commonly), or not conspicuously asymmetric; when dissected, palmatifid; usually palmately veined. Leaves stipulate. Stipules intrapetiolar; free of one another; caducous (often), or persistent. Leaf development not graminaceous. Domatia occurring in the family (known from 9 genera); manifested as hair tufts (nearly always), or pockets (rarely).
Leaf anatomy. The leaf lamina dorsiventral (usually), or bifacial (sometimes consisting wholly of palisade). Mucilaginous epidermis present, or absent. Stomata present; usually anomocytic. Hairs present (with numerous kinds represented in the family), or absent; eglandular, or glandular; unicellular, or multicellular. Complex hairs present, or absent; peltate, or stellate, or capitate. Lamina with secretory cavities, or without secretory cavities. Secretory cavities containing mucilage (then with mucilaginous canals and/or cells); schizogenous, or lysigenous. The mesophyll usually containing mucilage cells. Minor leaf veins without phloem transfer cells (Entelea, Sparmannia).
Axial (stem, wood) anatomy. Secretory cavities present (usually, in pith and cortex); with mucilage. Cork cambium present; initially superficial. Nodes tri-lacunar. Primary vascular tissues in a cylinder, without separate bundles; collateral. Internal phloem absent. Secondary thickening developing from a conventional cambial ring. Primary medullary rays wide, mixed wide and narrow, and narrow (the xylem ring more interrrupted in some species than others).
The wood semi-ring porous, or diffuse porous. The vessels small to medium; solitary, or radially paired to in radial multiples (commonly), or clustered. The vessel end-walls simple. The vessels without vestured pits; with spiral thickening, or without spiral thickening. The axial xylem with tracheids, or without tracheids; without vasicentric tracheids; with fibre tracheids (with small bordered pits in Tilia), or without fibre tracheids (usually); with libriform fibres; without septate fibres. The fibres without spiral thickening. The parenchyma apotracheal, or paratracheal. The secondary phloem stratified into hard (fibrous) and soft (parenchymatous) zones. Included phloem absent. Tile cells present (Durio and Pterospermum types). The wood storied, or partially storied (VPI).
Reproductive type, pollination. Plants hermaphrodite, or monoecious, or polygamomonoecious.
Inflorescence, floral, fruit and seed morphology. Flowers solitary, or aggregated in inflorescences (and sometimes paired); when solitary, axillary; when aggregated, in cymes. The ultimate inflorescence units cymose (mostly), or racemose. Inflorescences axillary (or displaced-axillary, with the foliage leaf subtending both a vegetative and an inflorescence bud: see Rendle (1930) for interpretation); mostly cymes, often very complex. Flowers regular; (3–)5 merous; cyclic, or partially acyclic. Sometimes the androecium acyclic. Floral receptacle developing an androphore, or with neither androphore nor gynophore. Free hypanthium absent. Hypogynous disk absent.
Perianth with distinct calyx and corolla, or sepaline (corolla rarely lacking); (4–)5–10; 2 whorled (usually), or 1 whorled; isomerous (usually). Calyx (3–)5; 1 whorled; polysepalous, or gamosepalous (sometimes basally connate); regular; imbricate. Epicalyx present, or absent. Corolla normally (4–)5; 1 whorled; polypetalous; imbricate, or contorted; regular. Petals deeply bifid, or entire.
Androecium (10–)15–100 (usually many). Androecial members branched; maturing centrifugally; free of the perianth (inserted at the base of the petals, or on an androphore); free of one another, or coherent; when coherent, 1 adelphous, or 5 adelphous, or 10 adelphous; 1–10 whorled (or acyclic and covering an androphore). Androecium exclusively of fertile stamens, or including staminodes. Staminodes when present, 5–15 (?); non-petaloid. Stamens (10–)15–100 (usually many); diplostemonous (rarely), or triplostemonous to polystemonous. Anthers dehiscing via pores, or dehiscing via short slits, or dehiscing via longitudinal slits; bilocular (by contrast with Malvaceae); bisporangiate. Endothecium developing fibrous thickenings. Anther epidermis persistent. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral. Anther wall initially with one middle layer, or initially with more than one middle layer (1 or 2); of the basic type. Tapetum glandular. Pollen grains aperturate; (2–)3–4 aperturate, or 6 aperturate; porate (34), or colporate (most commonly tricolporate), or foraminate (oligo-), or rugate (6-); 2-celled (in 6 genera).
Gynoecium 2–100 carpelled (to many). 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, or 2–100 celled (to many). Gynoecium syncarpous; eu-syncarpous; superior, or inferior (Neotessmannia). Ovary 1 locular (the septa incomplete), or 2–100 locular (to many); sessile. Gynoecium stylate. Styles 1; apical. Stigmas 1; capitate (or lobed); dry type; papillate; Group II type. Placentation when unilocular (i.e. rarely), free central; usually axile. Ovules in the single cavity when unilocular, 2–100 (to many); (1–)2–50 per locule (to many); ascending (usually, or always with Neotessmannia excluded?); more or less apotropous (?); with ventral raphe, or with lateral raphe; arillate (sometimes), or non-arillate; hemianatropous to anatropous; bitegmic; crassinucellate. Outer integument contributing to the micropyle, or not contributing to the micropyle. Endothelium differentiated. Embryo-sac development Polygonum-type. Antipodal cells formed; 3; not proliferating; ephemeral. Synergids hooked (or very elongated). Endosperm formation nuclear. Embryogeny onagrad, or asterad.
Fruit fleshy, or non-fleshy; dehiscent, or indehiscent, or a schizocarp. Mericarps when schizocarpic, 2–100 (to many); comprising nutlets, or samaroid, or comprising drupelets (or other?). Fruit when non-schizocarpic, a capsule (usually), or capsular-indehiscent, or a drupe, or a nut (or other?). Capsules denticidal, or poricidal, or loculicidal (or other?). Seeds endospermic. Endosperm oily. Cotyledons 2; flat. Embryo chlorophyllous (4/8); curved, or bent. Micropyle zigzag.
Seedling. Germination phanerocotylar.
Physiology, phytochemistry. C3. C3 physiology recorded directly in Corchorus, Tilia. Sugars transported as oligosaccharides + sucrose (predominantly, but some myoinositol usually detected as well). Not cyanogenic. Alkaloids present (rarely), or absent. Arbutin absent. Iridoids not detected. Proanthocyanidins present; cyanidin. Flavonols present, or absent; kaempferol, or kaempferol and quercetin. Ellagic acid absent (4 species, 4 genera). Aluminium accumulation not found. Sieve-tube plastids S-type.
Geography, cytology. Temperate to tropical. Cosmopolitan, except for frigid regions. X = 7–41.
Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgrens Superorder Malviflorae; Malvales. Cronquists Subclass Dilleniidae; Malvales. APG III core angiosperms; core eudicot; Superorder Rosanae; malvid. APG IV Order Malvales (as a synonym of Malvaceae).
Species 450. Genera about 50; Ancistrocarpus, Apeiba, Asterophorum, Berrya, Brownlowia, Burretiodendron, Christiana, Clappertonia, Colona, Corchorus, Craigia, Desplatsia, Diplodiscus, Duboscia, Eleutherostylis, Entelea, Erinocarpus, Glyphaea, Goethalsia, Grewia, Hainania, Heliocarpus, Hydrogaster, Jarandersonia, Luehea, Lueheopsis, Microcos, Mollia, Mortoniodendron, Neotessmannia (perhaps bettert referred to Muntingiaceae, q.v.), Pentace, Pentaplaris, Pseudocorchorus, Schoutenia, Sicrea, Sparmannia, Tahitia, Tetralix, Tilia, Trichospermum, Triumfetta, Vasivaea, Vinticena.
General remarks. Bayer et al.(1999) expanded Malvaceae to include Bombacaceae, Sterculiaceae and Tiliaceae, consequent on a combined analysis of plastid atpB and rbcL DNA sequences. However, the descriptions compiled for the present package depict Tiliaceae differing from Malvaceae sensu stricto (q.v.) in the imbricate calyx, androecial members free of the perianth, bilocular anthers (differing also in the glandular tapetum and the type of wall formation), and in the eusyncarpous gynoecium; also in the record of transported sugars containing oligosaccharides.
Economic uses, etc. Tilia supplies lumber (basswood, whitewood), also ornamental and shade trees popular for street plantings.
In the line-grove
which weather-fends your cell
(Tempest, v., 1. Line deriving from linden, subsequently corrupted to lime re. Tilia)
Illustrations. • Le Maout and Decaisne: Tilia. • Apeiba, Berrya, Triumfetta: Lindley. • Clappertonia polyandra, as Cephalonema: Hook. Ic. Pl. 31 (1915). • Grewia occidentalis: Bot. Mag. 422, 1798. • Mollia speciosa: Martius, Nova Gen. et Spec. Pl. Brasiliensium (1824). • Tilia platyphyllos (as T. grandifolia), T. x europaea (as T. intermedia), T. cordata (as T. parvifolia): Eng. Bot. 295–297, 1864. • Tilia x europaea (B. Ent.). • Triumfetta rhomboidea, as T. angulata: Wight’s Figs. of Indian Plants 2 (1843). • Triumfetta digitata, as Ceratosepalum: Hook. Ic. Pl. 24 (1894).
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: 15th April 2018. delta-intkey.com/angio’.