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). With a basal aggregation of leaves, or with neither basal nor terminal aggregations of leaves, or with terminal aggregations of leaves. Mesophytic (in rainforest), or xerophytic (mainly). Heterophyllous, or not heterophyllous. Leaves evergreen; 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; not connate; 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); edgewise to the stem, or with ‘normal’ orientation; 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. Leaves without a persistent basal meristem.
Leaf anatomy. Stomata present; commonly paracytic.
Adaxial hypodermis present, or absent. Lamina dorsiventral, or isobilateral, or centric; with secretory cavities (rarely), or without secretory cavities. The mesophyll with sclerencymatous idioblasts (commonly), or without sclerenchymatous idioblasts. Minor leaf veins without phloem transfer cells (Grevillea, Lomatia, Telopea).
Stem anatomy. Secretory cavities absent. Cork cambium present; initially deep-seated (rarely), or superficial. Nodes tri-lacunar. Cortical bundles present, or absent. Secondary thickening developing from a conventional cambial ring. Xylem with tracheids, or without tracheids; with fibre tracheids, or without fibre tracheids. Vessel end-walls simple, or scalariform and simple. Vessels without vestured pits. Primary medullary rays typically mixed wide and narrow. Wood parenchyma paratracheal.
Reproductive type, pollination. Plants hermaphrodite (usually), or monoecious, or andromonoecious, or dioecious. Entomophilous, or ornithophilous, or cheiropterophilous (?), or pollinated by unusual means (by small marsupials and rodents). Pollination 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); in racemes, in spikes, in heads, and in umbels. The fruiting inflorescences conelike (sometimes large and spectacular, especially in some Banksia species), or not conelike. The terminal inflorescence unit 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.
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 (i.e.to ‘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). Fruit without fleshy investment. 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, biochemistry. Cyanogenic, or not cyanogenic. Cynogenic constituents tyrosine-derived. Alkaloids present (rarely), or absent. Iridoids not detected. 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). Arbutin present. Saponins/sapogenins present, or absent. Aluminium accumulation demonstrated (very commonly), or not found. Sugars transported as sucrose (in Grevillea robusta). C3. C3 physiology recorded directly in Grevillea, Hakea.
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 (1998) Eudicot; peripheral Eudicot (non-core Eudicots, ‘neither Rosid nor Asterid’); 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.
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. • Technical details (Banksia). • Technical details (Leucospermum). • Technical details (Grevillea, Stenocarpus).
Cite this publication as: ‘L. Watson and M. J. Dallwitz (1992 onwards). The Families of Flowering Plants: Descriptions, Illustrations, Identification, and Information Retrieval. Version: 14th December 2000. http://biodiversity.uno.edu/delta/’. Dallwitz (1980), Dallwitz, Paine and Zurcher (1993, 1995, 2000), and Watson and Dallwitz (1991) should also be cited (see References).
