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The families of flowering plants

L. Watson and M.J. Dallwitz

Proteaceae Juss.

Including Lepidocarpicae (Lepidocarpaceae) Schultz-Schultzenst.

Habit and leaf form. Trees, or shrubs (or often lignotuberous subshrubs, sometimes geoflorous), or herbs (sometimes, in Stirlingia). The few herbs perennial (and woody); with a basal aggregation of leaves, or without conspicuous aggregations of leaves, or with terminal aggregations of leaves. Mesophytic (in rainforest), or xerophytic (mainly). Heterophyllous, or not heterophyllous. Leaves persistent; small to very large; alternate (mostly), or opposite, or whorled; usually spiral; flat, or terete; leathery, or fleshy, or modified into spines; petiolate to sessile; non-sheathing (usually), or sheathing (sometimes in Synaphea). Leaf sheaths when sheathing, not tubular; with free margins. Leaves aromatic (occasionally), or without marked odour (mostly); borne edgewise to the stem, or ‘normally orientated’; simple, or compound (especially in juvenile stages); epulvinate; when compound, or considered so, ternate, or pinnate, or bipinnate, or multiply compound, or palmate (sometimes digitate with pinnate segments). Lamina when simple, dissected, or entire; when entire, often acicular, or linear; when simple/dissected, pinnatifid, or palmatifid, or much-divided, or finely dichotomously dissected (variously dichotomously branched, bipinnately dissected, or digitately dissected with pinnately dissected segments — and in Synaphea lending the plants an amazingly ‘pteridophytic’ aspect); one-veined, or pinnately veined, or palmately veined, or parallel-veined. Leaves exstipulate. Lamina margins entire, or crenate, or serrate, or dentate. Leaf development not ‘graminaceous’.

Leaf anatomy. The leaf lamina variously dorsiventral, or bifacial, or centric. Stomata often sunken, mainly confined to one surface (commonly abaxial on dorsiventral leaves), or on both surfaces; commonly paracytic (but subsidiary patterns may be indeterminable when the stomata are sunken). Hairs present; seemingly exclusively eglandular; unicellular (mostly, usually with thick walls and narrow lumina, but occasionally forming a dense felt and sometimes deciduous), or multicellular (bicellular with short basal and long terminal cells recorded in Lambertia, and bicellular-medifixed hairs each with a long horizontal cell supported in the middle by short basal one recorded in Hakea and Grevillea). Adaxial hypodermis present (sometimes with an abaxial one as well), or absent. Lamina with secretory cavities (rarely), or without secretory cavities. The mesophyll with sclerenchymatous idioblasts (commonly, branched or unbranched), or without sclerenchymatous idioblasts; containing crystals, or without crystals (rather infrequent). The crystals when present, druses, or solitary-prismatic. Minor leaf veins without phloem transfer cells (Grevillea, Lomatia, Telopea).

Axial (stem, wood) anatomy. Secretory cavities absent. Cork cambium present; initially deep-seated (rarely), or initially superficial. Nodes tri-lacunar. Primary vascular tissues in sufficiently young stems comprising a ring of bundles (the circle of closely placed but individually distinct bundles more or less sinuous); collateral. Internal phloem absent. Cortical bundles present (e.g., in species of Banksia, Dryandra and Protea), or absent. Medullary bundles absent. Secondary thickening developing from a conventional cambial ring. Primary medullary rays typically mixed wide and narrow.

The wood diffuse porous (nearly always), or ring porous to semi-ring porous (uncommonly). The vessels small to medium (mostly), or large (rarely). The vessel end-walls simple (typically), or scalariform and simple (rarely exhibiting a few members with scalariform plates). The vessels without vestured pits; with spiral thickening, or without spiral thickening. The axial xylem with tracheids, or without tracheids; with vasicentric tracheids (often), or without vasicentric tracheids; with fibre tracheids, or without fibre tracheids (the apertures usually exserted, the pit borders sometimes distinct but usually small or indistinct and sometimes lacking); with libriform fibres, or without libriform fibres; without septate fibres. The fibres with spiral thickening (rarely?), or without spiral thickening. The parenchyma paratracheal. The secondary phloem not stratified. ‘Included’ phloem absent. The wood not storied.

Reproductive type, pollination. Plants hermaphrodite (usually), or monoecious, or andromonoecious, or dioecious. Pollination entomophilous, or ornithophilous, or cheiropterophilous (?), or by unusual means (notably by small marsupials and rodents); mechanism conspicuously specialized (usually, exhibiting ‘the greatest diversity of pollen presenter morphology in the flowering plants’ (Ladd 1994)), or unspecialized (Sphalmioideae).

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’ (nearly always), or solitary (sometimes, more or less, in Adenanthos); when solitary, axillary (towards the ends of the shoots); mostly in racemes, or in spikes, or in heads, or in umbels. The fruiting inflorescences conelike (sometimes large and spectacular, especially in some Banksia species), or not conelike. The ultimate inflorescence units racemose (or two flowered). Inflorescences terminal, or axillary, or intercalary; bracteate heads, cones, spikes or racemes; with involucral bracts, or without involucral bracts; pseudanthial, or not pseudanthial. Flowers bracteate (perhaps always), or ebracteate (depending on interpretation of cone-scales, in Isopogon, Petrophile); small to large (often very showy); regular to very irregular; when irregular, zygomorphic. The floral irregularity when present, involving the perianth and involving the androecium. Flowers mainly 4 merous; cyclic; tetracyclic, or tricyclic. Floral receptacle developing a gynophore, or with neither androphore nor gynophore. Free hypanthium present (represented by a ‘calyx tube’ with stamens attached), or absent (at least sometimes, regardless of the hypanthium/calyx-tube conundrum, since the stamens are sometimes hypogynous (e.g. Bellendina)). Hypogynous disk present (representing the corolla?), or absent; extrastaminal; of separate members, or annular.

Perianth with distinct calyx and corolla (there sometimes being ‘glands’ or ‘scales’, perhaps representing petals, internal to and alternating with the conspicuous tepals), or sepaline (the conspicuous perianth component seemingly representing the calyx, though ‘petaloid’); 4, or (6–)8; 1 whorled, or 2 whorled; isomerous, or anisomerous. Calyx (the conspicuous tepals being so interpreted) 4; gamosepalous (with a basal tube), or partially gamosepalous, or polysepalous (or interpretable as such, when the stamens are interpreted in the throat of the perianth — i.e. at the top of the ‘hypanthium’). When not completely gamosepalous, 3 of the members joined (and one free). Calyx tubular; unequal but not bilabiate (the tube sometimes cleft down most of one side), or bilabiate, or regular (but then commonly with the tube bent up, or with the lobes rolling back); valvate (but variously split when open). Corolla when present, (2–)4 (then represented by ‘glands’ or ‘scales’).

Androecium 4 (nearly always), or (3–)4 (Grevillea). Androecial members free of the perianth, or adnate (often inserted on the perianth, with only the anthers free); all equal, or markedly unequal; free of one another (usually), or coherent (sometimes in Conospermum, Synaphea); 1 whorled. Androecium exclusively of fertile stamens (but then often exhibiting stamens with one theca sterile), or including staminodes (Persoonia). Staminodes when present, in Petrophile 1; in the same series as the fertile stamens. Stamens 4 (usually), or (3–)4 (Grevillea, Petrophile); reduced in number relative to the adjacent perianth, or isomerous with the perianth; oppositisepalous; filantherous (with broad filaments), or with sessile anthers. Anthers cohering (occasionally), or connivent, or separate from one another; more or less basifixed; non-versatile; dehiscing via longitudinal slits; strongly introrse (usually), or latrorse (rarely); tetrasporangiate; appendaged (via the elongated connective), or unappendaged. Endothecium developing fibrous thickenings. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or decussate. Anther wall initially with more than one middle layer (2 or 3). Tapetum glandular. Pollen grains aperturate; (2–)3(–8) aperturate; porate (nearly always, perhaps occasionally colpoidate); 2-celled (in Grevillea, Leucadendron and Macadamia).

Gynoecium 1 carpelled. The pistil 1 celled. Gynoecium monomerous; of one carpel; superior. Carpel fully closed, or incompletely closed; non-stylate (rarely), or stylate (usually, the style often long and slender); apically stigmatic; (1–)3–100 ovuled ( ‘many’). Placentation marginal (mostly), or apical. Ovary sessile to stipitate. Styles bearing an ‘indusium’ beneath the stigma, or without an indusium. Stigmas dry type; papillate; Group II type. Ovules funicled, or sessile (the ovule sometimes adnate to the inner ovary wall, e.g. Leucospermum cordifolium); non-arillate; orthotropous, or anatropous, or amphitropous, or hemianatropous (the micropyle always pointing down); bitegmic; crassinucellate. Outer integument not contributing to the micropyle. Embryo-sac development Polygonum-type. Polar nuclei fusing only after one has been fertilized, or fusing simultaneously with the male gamete (?). Antipodal cells formed; 3; not proliferating; ephemeral, or persistent. Synergids usually hooked (and with filiform apparatus). Endosperm formation nuclear. Endosperm haustoria present; chalazal. Embryogeny asterad.

Fruit fleshy, or non-fleshy (frequently woody and long persistent). The fruiting carpel dehiscent, or indehiscent; a follicle, or drupaceous, or nucular, or an achene (see Manning and Brits (1993), who provide evidence for interpreting seed coats of indehiscent forms as outer integument rather than pericarp — thus casting doubt on all previous phylogenetic speculation involving seed coats). Gynoecia of adjoining flowers combining to form a multiple fruit, or not forming a multiple fruit. The multiple fruits coalescing (often, sometimes incorporated in woody cones), or not coalescing. Seeds non-endospermic (nearly always), or endospermic (Bellendina); winged (often), or wingless. Embryo well differentiated. Cotyledons 2(–9). Embryo achlorophyllous (1/1); straight.

Seedling. Germination phanerocotylar, or cryptocotylar.

Physiology, phytochemistry. C3. C3 physiology recorded directly in Grevillea, Hakea. Sugars transported as sucrose (in Grevillea robusta). Cyanogenic, or not cyanogenic. Cynogenic constituents tyrosine-derived. Alkaloids present (rarely), or absent. Arbutin present. Iridoids not detected. Saponins/sapogenins present, or absent. Proanthocyanidins present; delphinidin, or cyanidin and delphinidin. Flavonols present; kaempferol, or kaempferol, or kaempferol and quercetin, or quercetin and myricetin. Ellagic acid absent (7 species, 6 genera). Aluminium accumulation demonstrated (very commonly), or not found.

Geography, cytology. Temperate to tropical. Pantropical and warm, mainly requiring a long dry season. X = 5, 7, 10–13 (the chromosomes sometimes very large). Supposed basic chromosome number of family: 7.

Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgren’s Superorder Proteiflorae; Proteales. Cronquist’s Subclass Rosidae; Proteales. APG III core angiosperms; peripheral eudicot; Superorder Proteanae. APG IV Order Proteales.

Species 1050. Genera about 75; Acidonia, Adenanthos, Agastachys, Alloxylon, Athertonia, Aulax, Austromuellera, Banksia, Beauprea, Beapreopsis, Bellendina, Brabejum, Buckinghamia, Cardwellia, Carnarvonia, Cenarrhenes, Conospermum, Darlingia, Diastella, Dilobeia, Dryandra, Embothrium, Eucarpha, Euplassa, Faurea, Finschia, Floydia, Franklandia, Garnieria, Gevuina, Grevillea, Hakea, Helicia, Heliciopsis, Hicksbeachia, Hollandaea, Isopogon, Kermadecia, Knightia, Lambertia, Leucadendron, Leucospermum, Lomatia, Macadamia, Malagasia, Mimetes, Musgravea, Neorites, Opisthiolepis, Oreocallis, Orites, Orothamnus, Panopsis, Paranomus, Persoonia, Petrophile, Placospermum, Protea, Roupala, Serruria, Sleumerodendron, Sorocephalus, Spatalla, Sphalmium, Stenocarpus, Stirlingia, Strangea, Symphionema, Synaphea, Telopea, Toronia, Triunia, Turrillia, Vexatorella, Virotia, Xylomelum.

General remarks. Discussed in detail by Johnson and Briggs (1975).

Economic uses, etc. Many genera and species are cultivated as ornamentals and barrier plants, and Macadamia produces excellent edible nuts.

Illustrations. • Le Maout and Decaisne: Banksia. • Le Maout and Decaisne: Grevillea, Stenocarpus. • Leucospermum conocarpum: Thonner. • Banksia coccinea: inflorescences (photo). • Banksia coccinea: habitat (photo). • Banksia grandis: inflorescences (photo). • Banksia grandis: young fruiting inflorescence (photo). • Banksia aemula: Bot. Reg. 688, 1822. • Banksia marginata: Bot. Reg. 787, 1824. • Banksia paludosa: Bot. Reg. 697, 1823. • Banksia quercifolia: Bot. Reg. 1430, 1831. • Banksia speciosa (immature inflorescence): Bot. Reg. 1728, 1835. • Dryandra (Banksia) falcata: as Hemiclidia baxteri, Bot. Reg. 1455 (1831). • Dryandra (Banksia) formosa: inflorescence (photo). • Dryandra (Banksia) formosa: habitat (photo). • Grevillea concinna: Bot. Reg. 1383, 1830. • Grevillea manglesii, as Anadenia: Hook. Ic. Pl. 4 (1841). • Grevillea punicea: Bot. Mag. 109 (1883). • Grevillea punicea: Bot. Reg. 1319, 1830. • Hakea cucullata: Bot. Mag. 76 (1850). • Hakea microcarpa: Bot. Reg. 475, 1820. • Helicia grandis: Hook. Ic. Pl. 27 (1900). • Isopogon formosus: Bot. Reg. 1288, 1829. • Isopogon longifolius: Bot. Reg. 900, 1825. • Lambertia inermis (photo). • Lambertia inermis var. drummondii (photo). • Lambertia formosa: Bot. Reg. 528, 1821. • Lomatia longifolia, = L. myricoides: Bot. Reg. 442, 1820. • Protea mellifera, = P. repens: Bot. Mag. 346, 1796. • Stenocarpus salignus: Bot. Reg. 441, 1820. • Grevillea buxifolia: Bot. Reg. 443, 1820. • TS leaves of Franklandia, Hakea and Isopogon, and foliar hairs of Grevillea and Lambertia: Solereder, 1908.

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.’.