1 PLANTAE kingdom 1 MAGNOLIOPHYTA division PLANTAE 100 fam. MAGNOLIOPHYTA Proteaceae Prostrate to erect shrubs, or short to tall trees, sometimes lignotuberous, usually with short, 3-celled hairs, occasionally with glandular hairs. Clusters of short, lateral roots (‘proteoid roots’) often produced. Leaves alternate, less commonly opposite or whorled, simple, pinnatifid, pinnate or bipinnate, rarely palmate, usually coriaceous, exstipulate. Inflorescence simple or compound, axillary or terminal, racemose, paniculate or condensed (corymbose, capitate or cone-like). Flowers zygomorphic or actinomorphic, usually bisexual, solitary or paired in the axils of bracts, rarely ebracteate. Perianth of 4 valvate, free or variously united tepals; each with a slightly expanded limb. Stamens 4, usually all fertile, opposite tepals; filaments partly or wholly adnate to tepals, rarely free. Hypogynous glands usually present, (2–) 4, scale-like or fleshy, free or fused. Gynoecium of 1 carpel. Ovary usually superior, rarely perigynous, sessile or stipitate, 1-locular; ovules l to many, variously inserted; style simple, often persistent, often with the apex expanded as a pollen presenter; stigma small, terminal or subterminal. Fruit dehiscent, tardily dehiscent or indehiscent, a woody or coriaceous follicle, an achene, nut or a succulent or non-succulent drupe. Seeds 1 to many, sometimes winged, usually endospermic. This essentially Southern Hemisphere family of 79 genera and c. 1700 species has Australia and southern Africa as its centres of greatest diversity. Smaller numbers of species are known from Mexico, Central and South America, tropical Africa, Madagascar, India, eastern and south-eastern Asia, Malesia, New Caledonia, New Zealand and Fiji. In Australia, the family is represented by 46 genera and almost 1100 species and is found in all but the most arid regions. Speciation is most pronounced in south-western W.A., where prostrate to erect, usually sclerophyllous shrubs of large, ±locally endemic genera such as Petrophile, Conospermum, Synaphea, Adenanthos and Dryandra grow in oligotrophic, sandy soils and lateritic gravels. Similarly, species of other largely extratropical genera such as Isopogon, Grevillea, Hakea and Banksia are numerous in the south-west. Diversity is also high in the eastern states where Proteaceae tend to inhabit siliceous soils. A small but mainly disjunct and relictual component comprising 16 small genera occupies the rainforests of north-eastern Qld. The family name comes from that of the Greek sea-god, Proteus, who could change form at will, and was used by Linnaeus because the first representatives of this family seen by him were very variable. Coincidentally, it is an appropriate allusion for the rest of this morphologically diverse family. In an apparent adaptation to growth in poor soils, many taxa exhibit a specialised form of seasonal root growth known as ‘proteoid roots’. These are short, very dense root masses, produced laterally on the normal roots and heavily invested with root hairs. They are formed mainly in the leaf litter layer during seasonal growth flushes, usually shrivelling at the end of the season to be replaced again next year. Soil bacteria appear to play a role in their initiation. Proteoid roots probably function in an analogous way to mycorrhiza in other families, providing a greatly increased absorption surface for scarce nutrients and water. Mycorrhiza are not found in Proteaceae, although common in other Australian plant families growing in nutrient-poor soils. The inflorescences of Proteaceae are complex, and (particularly in Grevilleoideae) represent a reduced compound structure, which many authors prefer to describe as a conflorescence. However, for simplicity in the key and introduction, we have used the more generic term inflorescence to refer to all types of structure. Many taxa contain poisonous substances, mainly cyanogenetic glycosides, and skin irritants, and have been implicated in stock illnesses and deaths. Very few cases of human poisoning have been reported (Everist, 1981). R.Brown, On the Proteaceae of Jussieu, Trans. Linn. Soc. London 10: 15–226 (1810); C.F.Meisner, Proteaceae, in A.L.P.P. de Candolle, Prodr. 14: 209–482 (1856); G.Bentham, Proteaceae, Fl. Austral. 5: 315–584 (1870); H.Sleumer, Studies in Old World Proteaceae, Blumea 8: 1–95 (1955); C.Venkato Rao, Cytotaxonomy of the Proteaceae, Proc. Linn. Soc. New South Wales 82: 257–271 (1957); L.A.S.Johnson & B.G.Briggs, Evolution in the Proteaceae, Austral. J. Bot. 11: 21–61 (1963); L.A.S.Johnson & B.G.Briggs, On the Proteaceae – the evolution and classification of a southern family, Bot. J. Linn. Soc. 70: 83–182 (1975); S.L.Everist, Poisonous Plants of Australia, 2nd edn, 589–596 (1981); L.A.S.Johnson & B.G.Briggs, Proteaceae, in B.D.Morley & H.R.Toelken, Flowering Plants of Australia 238–244 (1983); A.S.George, An Introduction to the Proteaceae of Western Australia (1984); J.W.Wrigley & M.Fagg, Banksias, Waratahs and Grevilleas and all other plants in the Australian Proteaceae family (1989); G.J.Harden (ed.), Proteaceae, Fl. New South Wales 2: 2–71 (1991); R.J.Carpenter, Cuticular morphology and aspects of the ecology and fossil history of North Queensland forest Proteaceae, Bot. J. Linn. Soc. 116: 249–303 (1994). KEY TO THE GENERA OF PROTEACEAE IN AUSTRALIA P.H.Weston National Herbarium of New South Wales, Royal Botanic Gardens, Mrs Macquaries Road, Sydney, New South Wales, 2000. 1 Leaves opposite or whorled or pseudo-whorled 2 1: Leaves alternate 8 2 Fruit falling at maturity, either indehiscent or a tardily dehiscent, leathery, globose follicle 3 2: Fruit persistent on plant for several to many years, follicular, woody or cartilaginous 6 3 Flowers borne in regular, sessile, or shortly pedunculate pairs on an inflorescence axis; style tip swollen, functioning as a pollen presenter 4 3: Flowers borne singly in the axils of scale leaves or leaves in racemose or capitulate inflorescences; style tip not swollen, not functioning as a pollen presenter 5 4 Perianth strongly zygomorphic; hypogynous glands free, 4 or 2; pericarp fleshy, red to purple to blue 33 gen. TRIUNIA 4: Perianth actinomorphic; hypogynous glands connate, forming a ring around ovary; perianth leathery, green to brown or greyish 39 gen. MACADAMIA 5 Hypogynous glands absent; staminal filaments free or slightly adnate to the base of tepals; fruit more than 2 cm diam.; pericarp parenchymatous; rainforest trees 5 gen. EIDOTHEA 5: Hypogynous glands present; staminal filaments largely or wholly adnate to tepals; fruit less than 2 cm diam.; pericarp succulent; shrubs and small trees, mostly of sclerophyllous communities 3 gen. PERSOONIA 6 Inflorescence a 7-flowered head, or reduced to a single flower, surrounded by a conspicuous involucre; fruit beaked, often bearing conspicuous horns or spines 40 gen. LAMBERTIA 6: Inflorescence not consistently 1- or 7-flowered, lacking an involucre; fruit not beaked nor horned nor spiny 7 7 Inflorescence raceme-like, with a non-woody axis; follicle symmetrical, ellipsoidal to pear-shaped 32 gen. XYLOMELUM 7: Inflorescence cone-like, with a woody axis; follicle asymmetrical, laterally compressed 45 gen. BANKSIA 8 Leaves palmately compound with 5, or occasionally 3 radiating segments; some or all of the segments themselves often pinnately compound 17 gen. CARNARVONIA 8: Leaves simple or pinnately or bipinnately or (pseudo-)dichotomously compound 9 9 Leaves (pseudo-)dichotomously dissected or compound 10 9: Leaves either entire or with dentate margins or pinnately compound or divided 11 10 Leaves prominently glandular; perianth basally connate to form a slender tube; free distal perianth segments more than 9 mm long 14 gen. FRANKLANDIA 10: Leaves lacking glands; perianth segments free, less than 8 mm long 9 gen. STIRLINGIA 11 Perianth zygomorphic 12 11: Perianth actinomorphic 32 12 Flowers sessile in dense, cone-like, globose to cylindrical inflorescences; inflorescence axis woody 45 gen. BANKSIA 12: Flowers pedicellate to sessile, but not in cone-like inflorescences; inflorescence axis not woody 13 13 Flowers borne singly along the inflorescence axis, or inflorescence umbellate or reduced to a single flower 14 13: Flowers borne in regular, sessile or pedunculate pairs along inflorescence axis 22 14 One anther and 2 half-anthers abortive; loculi of adjacent anthers coherent in bud; each half-anther apparently 1-locular; hypogynous glands absent 15 14: Anthers not as above, either all developed and fully 4-locular or 1 or 3 infertile; hypogynous glands usually present 16 15 Perianth white, blue, grey or pink; lower anther abortive; leaves entire 12 gen. CONOSPERMUM 15: Perianth yellow; upper anther abortive; leaves usually dissected 13 gen. SYNAPHEA 16 Upper stamen fertile, the other 3 reduced to staminodes; plant andromonoecious, most flowers lacking a gynoecium; fruit a follicle in which the winged seeds are oriented transversely 1 gen. PLACOSPERMUM 16: All stamens fertile, or rarely 1 or all stamens infertile; flowers usually bisexual; fruit either not follicular or follicular but with longitudinally oriented seeds 17 17 Gynoecium about half length of perianth, hooked so that tip sits in pouch of ventral tepal below ventral anther; fruit a drupe 3 gen. PERSOONIA 17: Gynoecium about as long as perianth, exserted; fruit an achene or follicle 18 18 Inflorescence reduced to a single flower which is subtended at its base by several imbricate scale leaves; leaf glands present at leaf tip or at tips of leaf lobes or over leaf surface; ovule solitary; fruit an achene 15 gen. ADENANTHOS 18: Inflorescence usually multi-flowered or rarely reduced to a single flower (but then each flower subtended by a solitary scale leaf or completely lacking a floral bract); leaf glands absent; ovules 2 or more; fruit a follicle 19 19 Inflorescence umbel-like or reduced to a single flower; axis of symmetry of gynoecium passing through anterior and posterior tepals; hairs simple, seed enclosed within a membranous envelope 20 19: Inflorescence raceme-like and multi-flowered or rarely umbel-like or reduced to a single flower; axis of symmetry of gynoecium passing between tepals; hairs mostly bifid, seed not enclosed within a membranous envelope 21 20 Ovules 2; follicle woody, 1-seeded 23 gen. STRANGEA 20: Ovules more than 2; follicle leathery or cartilaginous, multi-seeded 24 gen. STENOCARPUS 21 Fruit usually leathery, rarely woody, lacking secondary thickening, usually not persisting on the plant for more than a year; leaves usually flat or angular in cross-section, usually with dissimilar surfaces, rarely truly terete or unifacial 41 gen. GREVILLEA 21: Fruit with woody secondary thickening, persistent on the plant for several to many years; leaves terete or flat with similar surfaces 42 gen. HAKEA 22 Hypogynous glands 2, 3 or 4, free 23 22: Hypogynous gland solitary, mostly crescentic or annular or occasionally bifid 26 23 Ovary densely hairy; ovules 2 24 23: Ovary glabrous; ovules more than 2 25 24 Leaves with entire to dentate margins; pollen presenter ±radially symmetrical; hypogynous glands alternating with tepals; fruit drupe-like 33 gen. TRIUNIA 24: Leaves pinnatisect; pollen presenter appearing to be lateral on style tip, not radially symmetrical; hypogynous glands opposite tepals; fruit a follicle 41 gen. GREVILLEA 25 Hypogynous glands 3; axis of symmetry of gynoecium passing between tepals; follicle leathery; seed wing terminal, diaphanous 27 gen. LOMATIA 25: Hypogynous glands 4; axis of symmetry of gynoecium passing through tepals; follicle woody; seed wing surrounding embryo, opaque 22 gen. CARDWELLIA 26 Ovules 2; fruit 1–2 27 26: Ovules more than 2; fruit multi-seeded 29 27 Pedicels absent, each flower pair borne on a common peduncle; axis of symmetry of gynoecium passing through tepals; hairs simple; fruit indehiscent; seed not winged 34 gen. GEVUINA 27: Pedicels present; common peduncle of flower pair absent; axis of symmetry of gynoecium passing between tepals; hairs mostly bifid; fruit dehiscent either spontaneously or on drying; seed usually winged 28 28 Fruit usually leathery, rarely woody, lacking secondary thickening, usually not persisting on plant for more than a year; leaves usually flat or angular in cross-section, usually with dissimilar surfaces, rarely truly terete or unifacial 41 gen. GREVILLEA 28: Fruit with woody secondary thickening, persistent on plant for several to many years; leaves terete or flat with similar surfaces 42 gen. HAKEA 29 Axis of symmetry of gynoecium passing through tepals, or flowers asymmetrical; flowers white to cream, sometimes appearing brownish due to hairs on outside of tepals 30 29: Axis of symmetry of gynoecium passing between tepals; flowers bilaterally symmetrical, red (very rare individuals can have white or yellow flowers) 31 30 Lower surface of leaves densely covered in persistent, appressed, shining hairs; intermediate leaves pinnate; perianth c. 3 mm long; hypogynous gland bifid 26 gen. OPISTHIOLEPIS 30: Lower surface of leaves glabrous; intermediate leaves pinnatisect; perianth 7–20 mm long; hypogynous gland crenulate 25 gen. BUCKINGHAMIA 31 Inflorescence not surrounded by involucral bracts; style gently curved 28 gen. ALLOXYLON 31: Inflorescence surrounded by an involucre of enlarged, sometimes brightly coloured bracts; style strongly curved or bent 29 gen. TELOPEA 32 Style tip not differentiated from the rest of the style as a pollen presenter, not swollen 33 32: Style tip differentiated from the rest of the style as a pollen presenter, usually swollen 42 33 Hypogynous glands 4 34 33: Hypogynous glands absent 38 34 Inflorescence racemose, the flowers borne singly in the axils of scale leaves or leaves, frequently growing on into a leafy shoot; fruit a drupe; tepals most frequently yellow, less commonly white, green or marked with red 35 34: Inflorescence a pseudo-raceme but the flowers borne on the inflorescence axis in lateral pairs subtended by prominent, often striped scale leaves, not growing on into a leafy shoot; fruit a follicle; tepals white to cream 37 35 Leaf margins dentate; posterior anther terminated by a much longer appendage than the lateral or anterior anthers; staminal filaments adnate to tepals only at the base 7 gen. CENARRHENES 35: Leaf margins entire; anthers similar, with or without appendages; staminal filaments largely or wholly adnate to tepals 36 36 Anthers and anther appendage gently incurved; endocarp subtransversely ribbed 2 gen. ACIDONIA 36: Anthers straight or recurved to revolute; anther appendage straight to recurved or absent; endocarp smooth 3 gen. PERSOONIA 37 Ovules 2; intermediate leaves simple, entire to dentate or pinnatisect 18 gen. ORITES 37: Ovules 6–8; intermediate leaves pinnate 19 gen. NEORITES 38 Flowers ebracteate 4 gen. BELLENDENA 38: Flowers each borne in the axil of a scale leaf 39 39 Fruit a follicle; intermediate leaves pinnate; some flowers lacking a gynoecium; rainforest trees [north-eastern Qld] 16 gen. SPHALMIUM 39: Fruit indehiscent; leaves simple, entire to pinnatisect; flowers bisexual; small trees or shrubs, usually in sclerophyllous communities [southern Australia] 40 40 Leaves pinnatisect 8 gen. SYMPHIONEMA 40: Leaves entire 41 41 Perianth tubular at base; fruit not winged 12 gen. CONOSPERMUM 41: Perianth not tubular at base; fruit prominently winged 6 gen. AGASTACHYS 42 Flowers borne singly in the axils of fleshy bracts in a dense cone-like inflorescence; fruit a nut; ovule solitary 43 42: Flowers borne in lateral pairs on the inflorescence axis; inflorescence raceme-like or dense and cone-like; fruit dehiscent or indehiscent but then usually with a succulent, outer mesocarp, rarely indehiscent and dry; ovules more than 1 44 43 Cone scales falling with the fruits; nut not strongly compressed 11 gen. ISOPOGON 43: Cone scales adhering firmly to the inflorescence axis and opening to release the flattened nuts 10 gen. PETROPHILE 44 Flowers sessile, densely packed in heads or cone-like, globose or cylindrical inflorescences 45 44: Flowers either pedicellate or sessile (but then not densely packed in heads or cone-like inflorescences) 46 45 Bracts surrounding base of inflorescence enlarged, imbricate, forming a prominent involucre 46 gen. DRYANDRA 45: Bracts surrounding base of inflorescence either absent or inconspicuous, not enlarged 45 gen. BANKSIA 46 Hypogynous glands fused to form a single, annular or horse shoe-shaped nectary; ovary usually stipitate 47 46: Hypogynous glands distinct, free or basally fused to form a 4-lobed ring; ovary sessile 49 47 Inflorescence a 2–8-flowered umbel, or reduced to a single flower; axis of symmetry of gynoecium passing through anterior and posterior tepals; hairs simple, seed enclosed within a membranous envelope 23 gen. STRANGEA 47: Inflorescence raceme-like and multi-flowered or rarely umbel-like or reduced to a single flower; axis of symmetry of gynoecium passing between tepals; hairs mostly bifid, seed not enclosed within a membranous envelope 48 48 Fruit usually leathery, rarely woody, lacking secondary thickening, usually not persisting on the plant for more than a year; leaves usually flat or angular in cross-section, usually with dissimilar surfaces, rarely truly terete or unifacial 41 gen. GREVILLEA 48: Fruit with woody secondary thickening, persistent on plant for several to many years; leaves terete or flat with similar surfaces 42 gen. HAKEA 49 Hypogynous glands 3; pollen grains 2-porate, curved-ellipsoidal 50 49: Hypogynous glands 4; pollen grains 3-porate, 3-angled 51 50 Mature, hardened leaves glabrous; inflorescence 30–90 cm long, pendulous; perianth c. 20 mm long; intermediate leaves pinnate 44 gen. AUSTROMUELLERA 50: Mature, hardened leaves densely and finely tomentose underneath; inflorescence to 20 cm long, ascending to erect; perianth c. 5 mm long; intermediate leaves simple although sometimes deeply lobed 43 gen. MUSGRAVEA 51 Adult leaves pinnate, often with a winged rachis 35 gen. HICKSBEACHIA 51: Adult leaves simple, entire to pinnatifid 52 52 Ovules more than 2; seeds 2 or more 53 52: Ovules 2; seeds 1 or 2 54 53 Inflorescence spreading to erect; tepals cream; ovules 4; seeds winged 21 gen. DARLINGIA 53: Inflorescence pendulous; tepals pink to mauve or red; ovules more than 4; seeds not winged 30 gen. HOLLANDAEA 54 Pollen presenter only slightly thicker than the more proximal part of the style; fruit a follicle; seeds 2, winged 55 54: Pollen presenter c. twice as thick as the more proximal part of the style; fruit indehiscent; seed solitary, not winged 56 55 Inflorescence ramiflorous, borne on the branches below the leaves; tepals 22–29 mm long; intermediate and some adult leaves with amplexicaul bases 20 gen. MEGAHERTZIA 55: Inflorescence axillary or terminal; tepals 4–8.5 mm long; leaf bases tapered 18 gen. ORITES 56 Leaves entire throughout life cycle; fruit brown or red when mature 57 56: Leaves with toothed margins present in juvenile phase and usually also in adults; sometimes adult leaves entire; fruit pinkish blue to deep blue to purple or black 58 57 Plant completely glabrous; tepals pink to red; fruit red; outer pericarp succulent; inner pericarp bony 37 gen. CATALEPIDIA 57: Young stems and outer surface of tepals hairy; tepals cream; fruit brown; pericarp bony 38 gen. FLOYDIA 58 Ovules orthotropous; intermediate leaves deeply pinnatifid; tepals 19–23 mm long; inflorescence 15–34 cm long; fruit lenticular, 3.6–4.1 cm long 36 gen. ATHERTONIA 58: Ovules anatropous; intermediate leaves not lobed; tepals 6–18 mm long; inflorescence 4–17 cm long; fruit ovoid to globose, 0.6–1.4 cm long 31 gen. HELICIA 4 Proteaceae subfam. Proteaceae Proteoideae Type: Protea L. Mostly sclerophyllous shrubs of infertile soils. Leaves simple, entire, toothed or variously divided. Inflorescence simple axillary racemes, compound terminal racemes, capitula (‘cones’) or reduced to single flowers. Flowers actinomorphic or zygomorphic, andromonoecious or bisexual, not in regular pairs; bracts present. Staminal filaments free or partly fused to tepals, or anthers sessile. Hypogynous glands present or absent. Pollen presenter present or absent. Fruit dry, indehiscent, usually a small nut or achene, rarely a drupe. Chromosomes usually small at mitotic metaphase, c. 0.5 µm long, but rarely to 5 µm. n = 14, sometimes 13, 11, 10, 24 or 28. A subfamily of 3 tribes, 26 genera and c. 637 species, with 2 major centres of diversity, southern Africa and Australia, but extending also to tropical and subtropical Africa, Madagascar and New Caledonia. In Australia, 2 tribes, 10 genera and 237 species, the majority subshrubs, shrubs or, rarely, small trees of sclerophyllous communities. Often in poor soils. 1 Proteaceae trib. PROTEOIDEAE Conospermeae Endl. Gen. Pl. 338 1837 Type: Conospermum Sm. Leaves entire or variously divided; glandular cavities absent. Inflorescence capitate, spicate, racemose or cone-like. Flowers actinomorphic or zygomorphic. Stamens free, or often variously fused and with reduced anthers. Hypogynous glands absent or, if present (in Cenarrhenes), short and thick. Fruit usually a nut, rarely a drupe. A tribe of 5 subtribes; 3 subtribes comprising 5 genera endemic in Australia, 1 (Dilobeiinae) confined to Madagascar, and 1 (Cenarrheninae) comprising 3 endemic Australian genera and 2 from New Caledonia. 1 Proteaceae subtrib. CONOSPERMEAE Cenarrheninae L.A.S.Johnson & B.G.Briggs Bot. J. Linn. Soc. 70: 171 1975 Type: Cenarrhenes Labill. Leaves various, not dichotomous. Inflorescence a spike or raceme. Flowers actinomorphic, bisexual. Filaments often incurved beneath anthers. Hypogynous glands 4 or absent. Pollen presenter absent. Fruit drupaceous or dry, glabrous or almost so; endocarp woody, crystalliferous or absent. n = 10, 11, 13 or 14. Includes the Australian endemic genera Agastachys, Cenarrhenes and Symphionema as well as Beauprea Brongn. & Gris and Beaupreopsis Virot from New Caledonia. 8 I.R.H.Telford "Invergowrie", via Armidale, New South Wales, 2350. gen. CENARRHENINAE PROTEACEAE Symphionema R.Br. Trans. Linn. Soc. London 10: 157 1810 from the Greek symphio (to unite) and nema (a thread), in reference to the staminal filaments being connate towards their apices Type: not designated. Symphyonema Spreng., Anleit. Kenntn. Gew. 2nd edn, 2: 333 (1817), orth. var. Shrubs or undershrubs; stems erect. Leaves simple, usually 1–3-pinnatisect; upper leaves sometimes less dissected, linear or trifid, acute. Inflorescence a spike or panicle, terminal, or in upper leaf axils; bracts small, deltoid. Flowers ±actinomorphic. Perianth segments free from base, ±equal, linear, spreading. Stamens epipetalous; filaments inserted towards bases of perianth segments, incurved and cohering around style near their apices; anthers erect, 2 locular, dehiscing longitudinally. Hypogynous glands absent. Ovary sessile, ellipsoidal, 1 locular; ovule solitary; style length equalling perianth; stigma capitate or slightly dilated. Fruit an ellipsoidal achene. A genus of 2 species endemic in south-eastern Australia. J.W.Wrigley & M.Fagg, Banksias, Waratahs & Grevilleas 530–531 (1989). Leaf segments flat, 2–4 mm wide 1 sp. montanum Leaf segments ±terete, 0.5–1.5 mm wide 2 sp. paludosum 1 sp. Symphionema Symphionema montanum R.Br. Trans. Linn. Soc. London 10: 157 1810 T: near Port Jackson [Grose R., N.S.W.], R.Brown s.n.; holo: BM? n.v. Illustrations: N.C.W.Beadle et al., Fl. Sydney Reg. 3rd edn, t. 10 (1982), as Symphionema paludosa; E.R.Rotherham et al., Fl. & Pl. New South Wales & Southern Queensland 44, fig. 99 (1975). Shrub to 80 cm tall. Leaves 2-pinnatisect, 2–5 cm long; segments flat, linear to linear-lanceolate, 2–4 mm wide. Inflorescence 2–6 cm long. Perianth 4–5 mm long, pale yellow. Achene c. 2 mm long. 48F–H v16_f48.jpg Occurs on the Central Tablelands and Woronora Plateau, N.S.W., from near Putty south through the Blue Mountains to near Macquarie Pass. Grows in heath or open forest, usually in moist sites in sandy soil on sandstone at altitudes of 400–1200 m. Flowers Oct.–Dec. 122 v16_m122.jpg N.S.W.: c. 8 km W of Mt Cameron, c. 30 km N of Bell, J.Pickard 1242 (NSW); Cliff Drive, Katoomba, 19 Nov. 1970, C.Burgess (CBG, NSW); Blue Mtns Natl Park, Ingar Picnic Area, I.R.Telford 2984 (BISH, CBG, MEL); Mt Werong, 26 Oct. 1940, W.F.Blakely & F.L.Ludowier (NSW); West Dapto, May 1901, W.Cambage (NSW). Mature stamens may separate and the anthers dehisce almost explosively when triggered. 2 sp. Symphionema Symphionema paludosum R.Br. Trans. Linn. Soc. London 10: 157 1810 T: near Port Jackson [N.S.W.], R.Brown s.n.; holo: BM n.v. Illustration: N.C.W.Beadle, Students Fl. NE New South Wales 231, fig. 105C (1972). Shrub to 60 cm tall. Leaves mostly trifid or 1- or 2-pinnatisect; upper leaves sometimes undissected, linear; segments ±terete, 0.5–1.5 mm wide. Inflorescence 1–8 cm long. Perianth segments 4–5 mm long, pale yellow. Achene c. 2 mm long. 48A–E v16_f48.jpg Occurs mainly in coastal N.S.W., including near-coastal and escarpment plateaus, rarely in the higher Central Tablelands, from the Port Macquarie area south almost to the Victorian border and inland to Kanangra Tops. Grows usually in wet heath, or in open woodland or open forest, usually in sandy soils, at altitudes mostly below 800 m, rarely to 1200 m. Flowers Aug.–Nov. 123 v16_m123.jpg N.S.W.: 2 km W of Point Plomer, 16 km N of Port Macquarie, C.O.Boyd & D.J.McGillivray 1874 (NSW); Gosford, 6 Dec. 1922, J.S.Fletcher (NSW); 13 km E of Robertson, by Robertson–Jamberoo road, M.Evans 2677 (CANB, NSW); northern Budawang Ra., below Corang trig., I.R.Telford 10741 (AD, BRI, CBG, HO, MEL, NSW); Nadgee State Forest, 3.4 km N of Nagha, M.D.Crisp 4615 & I.R.Telford (BM, CBG, MEL, US). One of the least woody members of the family with plants sometimes appearing annual-like and flowering when only c. 10 cm tall. 2 subtrib. Conospermeae Proteaceae Stirlingiinae L.A.S.Johnson & B.G.Briggs Bot. J. Linn. Soc. 70: 171 1975 Type: Stirlingia Endl. Leaves ±dichotomous; subdivisions pinnate. Inflorescence a small head or short spike. Flowers actinomorphic, andromonoecious. Anthers fully developed; loculi 4, contiguous but not connate. Hypogynous glands absent. Pollen presenter absent. Fruit a small nut with long hairs. n = 13. One genus endemic in south-western W.A. 9 A.S.George 'Four Gables', 18 Barclay Road, Kardinya, Western Australia 6163 gen. STIRLINGIINAE PROTEACEAE Stirlingia Endl. Gen. Pl. 339 1837 named after James Stirling (1791–1865), who explored the Swan River in 1827 and was the first Governor of Western Australia Type: Stirlingia anethifolia (R.Br.) Endl.; lecto, fide A.S.George, Fl. Australia 16: 473 (1995). Simsia R.Br., Trans. Linn. Soc. London 10: 152 (1810), nom. illeg. non Pers. T: Simsia anethifolia R.Br.; lecto, fide A.S.George, Fl. Australia 16: 473 (1995). Shrubs or perennial herbs with woody rootstock, or with taproot, glabrous except bracts and fruit. Leaves mostly on lower part of stem, ±dichotomously divided, leathery or soft. Inflorescence scapose, paniculately branched or simple. Flowers in heads or very short spikes; each flower subtended by small bract, pale yellow, often tinged red-brown, turning black. Perianth straight, actinomorphic; lobes of limb separating and reflexed at anthesis. Anthers on short filaments, 2-locular. Hypogynous scales absent. Gynoecium straight; ovary sessile; ovule 1; style end broadly cupular. Fruit an obconical nut, densely silky, falling when mature. n = 13, fide L.A.S.Johnson & B.G.Briggs, Bot. J. Linn. Soc. 70: 103 (1975). A genus of 7 species endemic in south-western W.A. Distinguished by the straight, regular perianth with lobes sharply reflexed after anthesis, the expanded stigma, the broad, hirsute nut and ±dichotomously divided leaves. Floral morphology is relatively uniform, species being delimited largely on the basis of habit, leaves and inflorescence. In the descriptions below, the relative width of the perianth limb and tube should be noted at the late bud stage. The leaves of the last 4 species are remarkably soft but persist through the typically dry, hot summer. A.S.George, Intr. Proteaceae W. Australia 101–102 (1984); J.W.Wrigley & M.Fagg, Banksias, Waratahs & Grevilleas 524–526 (1989); P.G.Ladd & S.W.Connell, Andromonoecy and fruit set in three genere of the Proteaceae, Bot. J. Linn. Soc. 116: 77–88 (1994). 1 Leaf segments flat; scape robust, to 1.5 m tall 1 sp. latifolia 1: Leaf segments terete; scape slender, not more than 1 m tall 2 2 Leaves less than 3 cm long, borne along more than half the stem 6 sp. abrotanoides 2: Leaves more than 5 cm long, mostly confined to lower part of stem 3 3 Heads 6–10 mm diam.; perianth 3–3.5 mm long; leaves leathery 2 sp. anethifolia 3: Heads 9–15 mm diam.; perianth 4.5–7 mm long; leaves ±soft 4 4 Scape simple or sparsely branched 7 sp. simplex 4: Scape openly branched 5 5 Petiole 12–14 cm long; perianth limb broader than tube 5 sp. divaricatissima 5: Petiole 1.5–6.5 cm long; perianth limb narrower than tube 6 6 Leaves divided up to 10 times into curved, crowded segments; ultimate lobes 1–4 mm long; bracts hoary 4 sp. tenuifolia 6: Leaves divided up to 6 times into straight, open segments; ultimate lobes 2–10 mm long; bracts glabrous 3 sp. seselifolia 1 sp. Stirlingia Stirlingia latifolia (R.Br.) Steud. Nomencl. Bot. 2nd edn, 2: 644 1841 Simsia latifolia R.Br., Suppl. Prodr. Fl. Nov. Holl. 9 (1830). T: Mt Manypeak[s], near King George Sound [W.A.], 1823, W.Baxter; holo: BM. Stirlingia paniculata Lindl., Sketch Veg. Swan R. xxx (1839). T: Swan River district, W.A., 183–, J.Mangles; holo: CGE. Simsia latifolia var. gracilis Ostenf., Biol. Meddel. Kongel. Danske Vidensk. Selsk. 3: 50 (1921). T: Kings Park, Perth, W.A., 13 Oct. 1914, C.H.Ostenfeld 681; iso: PERTH. Illustrations: R.Erickson et al., Fl. & Pl. W. Australia 27, pl. 39 (1973); A.S.George, Intr. Proteaceae W. Australia 102, figs 151–153 (1984); J.W.Wrigley & M.Fagg, Banksias, Waratahs & Grevilleas 525 (1989). Shrub with woody rootstock. Stems many, to 70 cm long. Leaves extending well up stem, leathery; petiole 3–10 cm long, ±terete at base, flattened upwards; lamina to 10 cm long, divided up to 3 times into flat, obtuse, often shallowly hooded lobes 1–15 mm wide. Scape to 1.5 m tall; panicle much-branched; heads 9–15 mm diam.; bracts ovate, obtuse to acute, 0.5–1 mm long, glabrous. Perianth 3–7 mm long; limb narrower than tube. Nut 4–5.5 mm diam., silky, hairy on top. Blueboy Widespread and often common in south-western W.A. from Kalbarri to Albany, mostly within 50 km of the coast, but extending inland to the Stirling Range. Grows in deep sand, in kwongan, shrubland and woodland. Flowers Sept.–Oct.; fruits Oct.–Jan. 124 v16_m124.jpg W.A.: Cockleshell Gully, W.E.Blackall 3587 (PERTH); South Perth, R.J.Cranfield 491 (PERTH); Yelverton State Forest, ENE of Margaret River, G.J.Keighery 11515 (PERTH); 25 km E of Cranbrook, R.H.Kuchel 1909 (AD, PERTH); 19 km E of Green Head, A.E.Orchard 423 (AD, PERTH). Easily distinguished by the robust habit, flat leaf segments and large nut. Variable in the width of leaf lobes and in the size and density of the panicle. Flowers pungently scented. The common name refers to the fact that wall plaster made using sand from where the species occurs turns blue. Flowering is enhanced the first season after fire. 2 sp. Stirlingia Stirlingia anethifolia (R.Br.) Endl. Iconogr. Gen. Pl. t. 23 1837 Simsia anethifolia R.Br., Trans. Linn. Soc. London 10: 153 (1810); Stirlingia tenuifolia var. anethifolia (R.Br.) Benth., Fl. Austral. 5: 358 (1870). T: Lucky Bay [W.A.], Jan. 1802, R.Brown s.n.; holo: BM. Stirlingia teretifolia Meisn. in J.G.C.Lehmann, Pl. Preiss. 1: 515 (1845); Simsia teretifolia (Meisn.) F.Muell., Syst. Census Austral. Pl. 66 (1882). T: interior of south-western W.A., Feb. 1841, L.Preiss 768; iso: B. Stirlingia affinis Meisn. in J.G.C.Lehmann, Pl. Preiss. 1: 516 (1845). T: Mt Wuljenup [= Mt Willyung], W.A., 14 Oct. 1840, L.Preiss 770; iso: B, MEL (2 sheets), P (2 sheets). Stirlingia acutifolia Endl., Gen. Pl. Suppl. 4(2): 81 (1848), apparently an error for Stirlingia anethifolia (R.Br.) Endl. Stirlingia intricata Meisn. in A.L.P.P. de Candolle, Prodr. 14: 325 (1856). T: Swan River Colony, W.A., J.Drummond 268 (or herb. Shuttleworth 286); iso: BM (2 sheets), CGE, K (2 sheets). Shrub with perennial rootstock. Stems several, to 10 cm long. Leaves mainly basal or extending a short way up the stem, leathery; petiole terete, 2–12 cm long; lamina divided to 6 times into terete, curved lobes; ultimate lobes divergent, 3–20 (–50) mm long. Scape 4–25 cm tall, often scarcely exceeding leaves, openly branched; stems ±curved; heads 6–10 mm diam.; bracts ovate, obtuse to almost acute, 1–1.5 mm long, glabrous, ciliate or hoary. Perianth 3–3.5 mm long; limb wider than tube. Nut 2–2.5 mm diam., densely silky. 49F–H v16_f49.jpg Widespread in near-coastal regions of south-western W.A. from near Albany to Israelite Bay. Grows in sand in kwongan. Flowers Sept.–Nov.; fruits Dec.–Jan. 125 v16_m125.jpg W.A.: 22.5 km [S]E of Mt Ragged road junction towards Israelite Bay, D.B.Foreman 1312 (AD, CANB, MEL, PERTH); Betty Beach, E of Albany, G.J.Keighery 8721 (PERTH); c. 8.5 km WSW of Howick Hill, A.E.Orchard 1081 (AD, PERTH); Cape Le Grand Natl Park, R.D.Royce 8738 (PERTH); Cape Riche, 25 Oct. 1968, J.W.Wrigley (CBG, PERTH). Very variable in leaf size and scape height, but without recognisable infraspecific taxa. Leaf lobes usually divaricate and curved, but also often erect and ±straight. The most xeromorphic of the terete-leaved species. For many years it has been known as S. teretifolia. 3 sp. Stirlingia Stirlingia seselifolia Domin V?stn. Král. Ceské Spole?n. Nauk, T?. Mat. P?ír. 1921–22(2): 7 1923 T: ‘Mallet’, W.A., 1910, A.A.Dorrien-Smith; holo: K. Perennial herb with verrucose taproot. Stems 1–several, to 15 cm long. Leaves at or near base of stem, rather soft; petiole terete, 1–3.5 cm long; lamina divided up to 6 times into terete, ±straight lobes; ultimate lobes divergent, 2–10 mm long. Scape 35–70 cm tall, openly branched; stems ±straight; heads 10–12 mm diam.; bracts broadly ovate, acute to obtuse, 1–1.5 mm long, glabrous. Perianth 5–6 mm long; limb narrower than tube. Nut 3 mm diam., silky. 49D v16_f49.jpg Recorded from scattered localities between Bunbury and Frankland, south-western W.A. Grows in low-lying areas, in sand, sometimes over laterite in open Jarrah (Eucalyptus marginata) forest. Flowers Oct.; fruits Nov.–Dec. 126 v16_m126.jpg W.A.: near Moorinup L., N of L. Muir, A.S.George 11128 (PERTH); 49 km SW of Kojonup, A.S.George 15240 (MEL, NSW, PERTH); Milyeannup Coast Rd, 1 km SW of Brockman Hwy, A.S.George 17122 (CANB, MEL, NSW, PERTH); Lowden, M.Koch 2052 (K, NSW). Closely related to S. tenuifolia but of stiffer habit, the leaf lobes straighter and longer, the scape less branched and the bracts glabrous. Flowers not scented. Epithet was given wrongly as sessilifolia in A.D.Chapman, Austral. Pl. Name Index Q–Z: 2747 (1991). 4 sp. Stirlingia Stirlingia tenuifolia (R.Br.) Steud. Nomencl. Bot. 2nd edn, 2: 644 1841 Simsia tenuifolia R.Br., Trans. Linn. Soc. London 10: 152 (1810). T: Princess Royal Harbour [Albany, W.A.], 22 Dec. 1801, R.Brown s.n.; holo: BM. Illustrations: A.S.George, Intr. Proteaceae W. Australia 101, pl. 150 (1984); J.W.Wrigley & M.Fagg, Banksias, Waratahs & Grevilleas 517 (1989). Perennial herb with verrucose taproot, probably fire-sensitive. Stems 1–several, 5–25 cm long. Leaves soft; petiole 1.5–6.5 cm long; lamina divided up to 10 times into terete, curved lobes; ultimate lobes erect, 1–4 mm long. Scape 10–100 cm tall, openly branched; stems ±curved; heads 10–12 mm diam.; bracts ovate-triangular, obtuse, 0.8–1.2 mm long, tomentose. Perianth 5–6 mm long; limb narrower than tube. Nut 2 mm diam., densely silky. 49A–C v16_f49.jpg Occurs in south-western W.A. from the Frankland River to Middle Mt Barren. Grows in sand and shale, in kwongan or woodland on hillsides and low-lying areas. Flowers Sept.–Oct.; fruits Nov.–Dec. 127 v16_m127.jpg W.A.: Red Gum Spring, Stirling Ra., E.M.Canning CBG 28444 (CBG, NSW); Middle Mt Barren, C.A.Gardner 9172 (PERTH); Grasmere L. (L. Powell), G.L.Webster 18770 (NSW); 23.7 km S of Muir Hwy on Nornalup Rd, A.S.George 17140 (CANB, PERTH). Variable in size of plant, sometimes flowering when c. 10 cm tall, possibly in the first year. 5 sp. Stirlingia Stirlingia divaricatissima A.S.George Fl. Australia 16: 473 1995 T: 20 miles [32 km] N of Bow Bridge, N of Peaceful Bay, W.A., 22 Oct. 1971, J.Boyd 33; holo: PERTH. Shrub to 1.7 m tall. Branching habit unknown. Leaves soft; petiole terete, 12–14 cm long; lamina divaricately and intricately divided up to 10 times; ultimate segments straight to slightly curved, very slender, 2–4 mm long. Scape at least 45 cm long (not seen complete), sparsely branched; stems ±straight; heads 9 mm diam.; bracts ovate, obtuse to ±acute, 1.5 mm long, closely tomentose. Perianth 4.5–5 mm long; limb broader than tube. Nut not seen. 49E v16_f49.jpg Known from only two collections from north of Bow River, W.A. Grows in loam (habitat unknown) and in sand in shrubland among Jarrah–Marri forest. Flowers Oct. 128 v16_m128.jpg W.A.: N of Walpole, A.S.Weston 95.3.11 (PERTH). The tall habit and very finely divided leaves characterise the species. 6 sp. Stirlingia Stirlingia abrotanoides Meisn. J.G.C.Lehmann Pl. Preiss. 1: 517 1845 Simsia abrotanoides (Meisn.) F.Muell., Syst. Census Austral. Pl. 66 (1882). T: near Swan River, W.A., 30 Sept. 1839, L.Preiss 2622; n.v.; Swan River, W.A., J.Drummond 587; syn: CGE, MEL, P (3 sheets). Shrub or perennial herb with fire-tolerant rootstock. Stems several, to 15 cm long. Leaves borne along more than half the stem, soft; petiole terete, 4–10 mm long; lamina divided 4 or 5 times into terete, ±straight lobes; ultimate lobes ±erect, 1–5 mm long. Scape 14–16 cm tall, simple or sparingly branched; branches erect; stems ±curved; heads 12–14 mm diam.; bracts lanceolate, acute, 1.5–3 mm long, glabrous. Perianth 5 mm long; limb c. as wide as tube. Fruit 2 mm diam., silky. Occurs between Eneabba and Mogumber and further inland near Wongan Hills, Northam, Tammin and Narembeen, south-western W.A. Grows in sand in kwongan. Flowers Sept.–Oct.; fruits Nov.–Dec. 129 v16_m129.jpg W.A.: Hill R., J.S.Beard 1878 (PERTH); 10 km E of Eneabba, D.B.Foreman 516 (NSW); Coomallo, E.A.Griffin 5459 (PERTH); Tammin, Sept. 1909, J.H.Maiden (NSW); W of Wongan Hills, 2 Oct. 1903, A.Morrison (PERTH). Distinguished especially by the small leaves that are less divided and extend further up the stem than in the other terete-leaved species. Inland specimens have more woody stems. It has often been included in the synonymy of S. simplex. 7 sp. Stirlingia Stirlingia simplex Lindl. Sketch Veg. Swan R. xxx 1839 Simsia simplex (Lindl.) F.Muell., Syst. Cens. Austral. Pl. 66 (1882). T: Swan River district, W.A., before 1839, J.Drummond s.n.; lecto: CGE, fide A.S.George, Fl. Australia 16: 474 (1995); Swan River, W.A., D.Toward; syn: CGE. Stirlingia capillifolia Meisn., Hooker’s J. Bot. Kew Gard. Misc. 7: 70 (1855). T: south-western W.A., 1850–1851, J.Drummond 6: 173; iso: CGE, K, NSW, PERTH. Illustration: J.W.Wrigley & M.Fagg, Banksias, Waratahs & Grevilleas 517 (1989). Herb with perennial rootstock. Stems few to many, short-lived, to 10 cm long. Leaves mainly basal, crowded, soft; petiole terete, 3–13 mm long; lamina divided up to 7 times into terete, slightly curved lobes; ultimate lobes erect to spreading, 2–20 mm long. Scape 10–60 cm tall, simple or sparingly branched; heads 10–15 mm diam., sometimes slightly elongated; bracts lanceolate, acute, 1.5–2 mm long, glabrous. Perianth 5.5–7 mm long; limb narrower than tube. Nut 2 mm diam., densely silky. Occurs in scattered localities from Eneabba to Waroona and east to Hyden, south-western W.A. Grows in sand, gravelly sand, gravelly clay and gravel in kwongan and eucalypt woodland. Flowers late Sept.–early Nov.; fruits Nov.–Dec. 130 v16_m130.jpg W.A.: Bendering, C.A.Gardner 1830 (PERTH); Yandanooka West Rd, S of Mingenew, A.S.George 16863B (CANB, MEL, NSW, PERTH); Greenmount, Darling Ra., 21 Oct. 1908, A.Morrison (PERTH); Waroona, R.D.Royce 3135 (PERTH); 10 km from Pingaring towards Hyden, 10 Nov. 1968, J.W.Wrigley (CBG, NSW). Plants from the Darling Range near Perth have larger flowers than those from elsewhere. Flowers sweetly scented. 3 Proteaceae subtrib. Conospermeae Petrophilinae L.A.S.Johnson & B.G.Briggs Bot. J. Linn. Soc. 70: 171 1975 Type: Petrophile R.Br. ex Knight Leaves various, not dichotomous. Inflorescence a dense cone-like head with imbricate, scale-like floral bracts. Flowers actinomorphic, bisexual; bases of tepals connate. Anthers 4, fully developed; loculi of adjacent anthers not closely appressed. Hypogynous glands absent. Style modified as pollen presenter, often with a brush of hairs. Fruit a nut, often with long hairs; endocarp frequently crystalliferous. n = 13. Comprises the extra-tropical Australian endemic genera Petrophile and Isopogon. 11 D.B.Foreman 50 Benjamin Drive, Lara, Victoria, 3212. gen. PETROPHILINAE PROTEACEAE Isopogon R.Br. ex Knight Cult. Prot. 93 1809 nom. cons. from the Greek isos (equal) and pogon (a beard), referring to the tufts of hairs at the apex of the perianth segments of some species or, possibly, referring to the hairs of ±equal length which cover the nut on all sides Type: Protea anemonifolia Salisb. = Isopogon anemonifolius (Salisb.) Knight Atylus Salisb., Parad. Lond. 66 (1807) p.p. [Description incorporates features of both Isopogon and Petrophile]. Atylus sect. Isopogon (R.Br. ex Knight) Kuntze in T.E. von Post & C.E.O.Kuntze, Lex. Gen. Phan. 54 (1903). Shrubs or small trees. Leaves simple or compound, terete or flattened, rigid, sometimes broad. Inflorescence a dense spike or ‘cone’, terminal or axillary, ±globose or ovoid, sessile or pedunculate, solitary or several clustered; receptacle cylindrical to concave; involucral bracts usually present, somewhat more persistent than cone scales; cone scales tomentose to villous outside, glabrous inside, deciduous after flowering or shed with the fruit; floral bracts absent. Flowers actinomorphic, bisexual, sessile. Tepals spreading at anthesis; upper part decidous; basal part persisting until the fruit expands on ripening. Stamens 4, sessile, with a short connective. Hypogynous glands absent. Ovary sessile, 1-locular; ovules 1 (–2), pendulous; style filiform, straight; pollen presenter usually dilated or clavate, often separated from the brush by a constriction, with various parts glabrous or hairy, rarely fusiform, with a very small stigma, otherwise the tip slightly enlarged to form a stigmatic cup. Fruiting cones not persisting; scales separating readily. Fruit a small, hairy nut, not compressed or winged. Conesticks, Conebushes, Coneflowers n = 13, H.P.Ramsay, Austral. J. Bot. 11: 4 (1963). A genus of 35 species, including 7 subspecies and 4 varieties, endemic in extra-tropical Australia. Most species (27) are concentrated in south-western W.A. Preliminary observations suggest that at least some of the taxa presently included in Isopogon, e.g. I. buxifolius s. lat., may have as their closest relatives some of the South African Proteoideae. Isopogon is generally distinguishable from Petrophile by the dried cone scales that fall with the fruits and by the fruits, themselves, which are hairy and not compressed. Less clearly defined are differences in the morphology of the pollen presenters of the two genera, with taxa in both Isopogon and Petrophile having simple, ±fusiform structures or more ornate pollen presenters. Studies presently underway, including a cladistic analysis of the family, may shed more light on the relationships between Isopogon, Petrophile and other Proteoideae. In the following descriptions the inflorescence measurement (usually its diameter) is inclusive of the flowers. If the flowers tend to be held more erect rather than spreading the overall length of the inflorescence is given. The morphology of the receptacle is recorded only if it might be a useful diagnostic character. B.L.Rye, Isopogon, in N.G.Marchant et al., Fl. Perth Reg. 1: 345–347 (1987); R.M.Sainsbury, Field Guide Isopogons & Petrophiles 1–55 (1987); W.E.Blackall & B.J.Grieve, How to Know W. Austral. Wildflowers part 1, 2nd edn, 143–152 (1988); J.W.Wrigley & M.Fagg, Banksias, Waratahs & Grevilleas 425–440 (1989). 1 Leaves terete 2 1: Leaves flat or slightly concave 14 2 Flowers creamy yellow or yellow 3 2: Flowers dirty white tinged with pink, creamy pink, reddish, purple or mauve 6 3 Leaves simple 11 sp. sp. A 3: Leaves pinnate, bipinnate or divaricately divided 4 4 Leaves pinnate or bipinnate; shrub to 2 m tall [N.S.W.] 2 sp. anethifolius 4: Leaves divaricately divided; shrub to 1.6 m tall [W.A.] 5 5 Shrub to 0.6 m tall; petiole to c. 13 cm long; lamina to c. 12 cm long; flowers to 20 mm long 12 sp. villosus 5: Shrub to 1.6 m tall; petiole to c. 25 mm long; lamina to c. 4 cm long; flowers to 30 mm long 15 sp. gardneri 6 Perianth completely glabrous 20 sp. asper 6: Perianth hirsute, villous, densely pubescent or largely glabrous but with a tuft of hairs at the apex of each tepal 7 7 Perianth glabrous but with a tuft of hairs at the apex of each tepal; flowers pale to dark pink, reddish or mauve 8 7: Perianth hirsute, villous or densley pubescent; flowers creamy pink, pink, or dirty white tinged with pink or purple 12 8 Mature leaves 1.2–2.6 cm long, with long, fine, spreading hairs, trifid or occasionally simple 23 sp. adenanthoides 8: Mature leaves to c. 5.5–18 cm long, ±glabrous, variously divided but not trifid 9 9 Leaves with a shallow longitudinal groove 10 9: Leaves lacking a shallow longitudinal groove 11 10 Flowers to c. 25 mm long; leaves to 5.5 cm long, smooth 16 sp. formosus 10: Flowers to c. 30 mm long; leaves to 18 cm long, longitudinally wrinkled, thicker than above 13 sp. heterophyllus 11 Leaves usually pinnate, occasionally simple; branchlets glabrous; flowers to 25 mm long 17 sp. divergens 11: Leaves simple or forked; branchlets minutely pubescent; flowers to c. 18 mm long 18 sp. scabriusculus 12 Flowers drooping, grey-hirsute; low shrub to 0.8 m tall 22 sp. inconspicuus 12: Flowers not drooping, villous or densely pubescent; shrub to 2 m tall 13 13 Flowers densely pubescent, to c. 15 mm long; shrub to 2 m tall 14 sp. teretifolius 13: Flowers villous, to c. 30 mm long; shrub to 1.6 m tall 15 sp. gardneri 14 Leaves undivided; margins entire 15 14: Leaves variously divided or lobed 26 15 Flowers pink to mauve to purplish pink 16 15: Flowers white, creamy white, greenish, creamy yellow, yellow or creamy green 20 16 Inflorescence axillary; flowers villous-plumose towards the apex 34 sp. axillaris 16: Inflorescence terminal, sometimes clustered towards the end of the branchlets; flowers glabrous or with a tuft of hairs at the apex of each tepal 17 17 Perianth glabrous 25 sp. linearis 17: Perianth with a tuft of hairs at the apex of each tepal 18 18 Inflorescence terminal or clustered towards the ends of the branchlets, to 2 cm long; flowers 10–15 mm long 35 sp. buxifolius 18: Inflorescence terminal, solitary, to 8 cm diam.; flowers to c. 35 mm long 19 19 Lamina narrowly obovate to oblong-obovate; apex obtuse; flowers c. 25 mm long 30 sp. cuneatus 19: Lamina obovate to broadly elliptic; apex acute; flowers c. 35 mm long 31 sp. latifolius 20 Flowers glabrous 21 20: Flowers silky-villous, or with a tuft of hairs at the apex of each tepal or or the whole limb and sometimes part of the tube villous 22 21 Inflorescences often densely clustered; flowers whitish to yellow or cream; spreading shrub to 1 m tall [W.A.] 26 sp. polycephalus 21: Inflorescences solitary; flowers creamy yellow to creamy green; erect bushy shrub to c. 1.5 m tall [N.S.W.] 4 sp. fletcheri 22 Tufted shrub to 0.4 m tall; lamina apex usually uncinate 28 sp. uncinatus 22: Erect bushy shrub, 1–2.5 m tall; lamina apex never uncinate 23 23 Flowers with a tuft of hairs on the apex of each tepal; lamina to 3.5 cm long 35 sp. buxifolius 23: Flowers silky-villous or the limb and sometimes part of the tube villous; lamina to 16 cm long 24 24 Flowers silky-villous; shrub to 2.5 m tall; inflorescence solitary 10 sp. longifolius 24: Flowers with the limb and sometimes part of the tube villous 25 25 Leaves glabrous, oblong-spathulate to ±linear; lamina margins not recurved; petiole to c. 5 cm long 27 sp. attenuatus 25: Leaves often remaining hirsute, linear to narrowly obovate; lamina margins often recurved; petiole not discernible 29 sp. sphaerocephalus 26 Flowers white, silvery white, creamy white, pale grey, pink or purple 27 26: Flowers creamy yellow or yellow 33 27 Erect shrub or small tree, to c. 5 m tall [N.S.W.] 3 sp. dawsonii 27: Low to medium, erect shrub, 0.4–1.5 (–2) m tall [W.A.] 28 28 Lamina canaliculate 32 sp. dubius 28: Lamina not canaliculate 29 29 Flowers to 35 mm long, greyish-villous 21 sp. baxteri 29: Flowers to 30 mm long, glabrous except for a tuft of hairs on the apex of each tepal, or pubescent or silky 30 30 Lamina to 6.5 cm long 31 30: Lamina to c. 18 cm long 32 31 Leaves simple, 3-toothed; lamina, to 6.5 cm long; flowers mostly white 24 sp. tridens 31: Leaves once or twice ternately divided; lamina to 4 cm long; flowers reddish pink 33 sp. crithmifolius 32 Low shrub, to 0.45 m tall; flowers yellowish, densely silky towards the top 19 sp. alcicornis 32: Shrub to 2 m tall; flowers pink or red, with a tuft of hairs at the apex of each tepal, the tube glabrous or pubescent 18 sp. scabriusculus 33 Perianth glabrous except for a tuft of hairs at the apex of each tepal, or glabrous at the base and becoming villous towards the apex [eastern Australia] 34 33: Perianth ±uniformly silky-villous or silky-pubescent [W.A.] 38 34 Perianth creamy yellow, glabrous at the base and becoming villous towards the apex, 17–20 mm long 5 sp. mnoraifolius 34: Perianth yellow, glabrous except for a tuft of hairs at the apex of each tepal, to c. 10–15 mm long 35 35 Erect shrub to 2 m tall 6 sp. anemonifolius 35: Low-growing or prostrate shrub to 1 m tall, often less 36 36 Petiole to c. 12 cm long; involucral bracts pubescent 1 sp. petiolaris 36: Petiole to 6 cm long; involucral bracts glabrous or sparsely hairy, often only the margins ciliate 37 37 Prostrate, spreading shrub; lamina to c. 4.5 cm long, variously divided into linear lobes; apices acute; petiole to c. 6 cm long 7 sp. prostratus 37: Compact shrub; lamina 1.8–3.6 cm long, pinnatisect; apices pungent-pointed; petiole 1.2–5.6 cm long 8 sp. ceratophyllus 38 Low shrub to 45 cm tall; lamina 10–40 cm long; flowers white or pink, 15–20 mm long 19 sp. alcicornis 38: Shrub to 2–2.5 m tall; lamina to 15.5 cm long; flowers cream to yellow, 8–15 mm long 39 39 Flowers c. 8–10 mm long, silky-pubescent; lamina 3–9-toothed to deeply 3–5-lobed, 2–5.5 cm long 9 sp. trilobus 39: Flowers to c. 15 mm long, silky-villous; lamina simple or deeply 2–3-lobed, c. 15.5 cm long 10 sp. longifolius 0 Published since the Flora of Australia treatment 36 sp. robustus 1 sp. Isopogon Isopogon petiolaris A.Cunn. ex R.Br. Suppl. Prodr. Fl. Nov. Holl. 8 1830 Atylus petiolaris (A.Cunn. ex R.Br.) Kuntze, Revis. Gen. Pl. 2: 577 (1891). T: ora orient., Moreton Bay [Qld], 1827, A.Cunningham; syn: BM. Isopogon anemonifolius var. ceratophylloides Cheel, Proc. Linn. Soc. New South Wales 48: 682 (1923). T: Warialda, N.S.W., Oct. 1904, N.W.Garrand s.n.; syn: NSW. Illustration: G.J.Harden in G.J.Harden (ed.), Fl. New South Wales 2: 21 (1991). Low, spreading shrub, mostly less than 1 m tall. Branchlets reddish brown, pubescent. Leaves ternately to pinnately divided, erect; petiole to c. 3–12 cm long; lamina flat, to c. 1.5–7.5 cm long, striate, acute to pungent-pointed, glabrescent; lateral pinnae often divaricate. Inflorescence globose, terminal, sessile, solitary, c. 15–20 mm diam., surrounded by leaves; involucral bracts broad, acuminate, pubescent; cone scales broadly cuneate, villous to very woolly on outside, except for acuminate apex. Flowers to c. 10 mm long, yellow, glabrous except for a short tuft of apical hairs on each tepal. Pollen presenter to c. 2–3 mm long; basal part minutely papillose, scarcely swollen, constricted near mid-point then slightly dilated; apical part glabrous, scarcely swollen except for stigmatic cup. Cones ovoid to globose, to c. 20 mm diam. Nuts ovoid, beaked, to c. 4.5 mm long, villous. 99K v16_f99.jpg Widely distributed from the Darling Downs, south-eastern Qld, to the New England Tableland, extending south to near Parramatta and west to the Pilliga State Forest and Bumberry, near Parkes, N.S.W. Grows in heath or dry sclerophyll forest or woodland, often in sandy soils of granite origin. Flowers July–Nov. 185 v16_m185.jpg Qld: Bald Rock Ck, 10 km N of Wallangara, I.R.Telford 3185 (CBG); Tobacco Rd, 17 km ESE of Inglewood, A.N.Rodd 4123 (MEL). N.S.W.: 40 km NNE of Coonabarabran on Narrabri road, H.Streimann 649 (CBG); Parramatta, July 1855, F.Mueller (MEL); Pilliga State Forest, 0.4 km W along Burma Rd from Newell Hwy, R.G.Coveny 12740, P.Cuneo & B.Wiecek (MEL, NSW). Reported as having several stems arising from a lignotuber. 2 sp. Isopogon Isopogon anethifolius (Salisb.) Knight Cult. Prot. 94 1809 Protea anethifolia Salisb., Prodr. Stirp. Chap. Allerton 48 (1796); Atylus anethifolius (Salisb.) Kuntze, Revis. Gen. Pl. 2: 577 (1891). T: ex Port Jackson [N.S.W.], auct. Jac. Lee; not located. Isopogon eriophorus Gand., Bull. Soc. Bot. France 66: 229 (1919). T: Berowra, N.S.W., July 1899, J.H.Maiden; holo: LY, fide D.J.McGillivray, Contr. New South Wales Natl. Herb. 4: 344 (1973). Isopogon confertus Gand., Bull. Soc. Bot. France 66: 229 (1919). T: Never Never, Rylstone, N.S.W., Oct. 1897, R.T.Baker s.n.; holo: LY, fide D.J.McGillivray, Contr. New South Wales Natl. Herb. 4: 344 (1973). Isopogon globosus Gand., Bull. Soc. Bot. France 66: 229 (1919). T: Port Jackson, N.S.W., Aug. 1897, C.Walter s.n.; lecto: LY, fide D.J.McGillivray, Contr. New South Wales Natl. Herb. 4: 344 (1973); Port Jackson, N.S.W., 1910, J.R.Tovey s.n.; syn: LY. Isopogon virgulatus Gand., Bull. Soc. Bot. France 66: 229 (1919). T: (Australia occid.), N.S.W., 1902 (ex Herb. C.Walter); holo: LY, fide D.J.McGillivray, Contr. New South Wales Natl. Herb. 4: 344 (1973). Illustrations: W.R.Elliot & D.L.Jones, Encycl. Austral. Pl. 5: 441 (1990); G.J.Harden in G.J.Harden (ed.), Fl. New South Wales 2: 21 (1991). Erect, bushy shrub to c. 2 m tall. Branchlets reddish brown, glabrous. Leaves pinnate or bipinnate; petiole c. 3.5–5.5 cm long; lamina terete, 6–10.5 cm long, acute, glabrous. Inflorescence globose, terminal or axillary, sessile or shortly pedunculate, solitary, to c. 40 mm diam.; involucral bracts small and narrow, ±acuminate, glabrous or with a short marginal fringe of hairs; inner ones broader; cone scales numerous, imbricate, broad, truncate, tomentose to villous except for deciduous apex. Flowers to c. 12 mm long, yellow, shortly silky, with longer tufts of hairs on apex of each tepal. Pollen presenter to c. 3.5 mm long; basal part swollen, minutely pubescent, constricted near mid-point then dilated into a globose part; apical part scarcely swollen, glabrous. Cones ovoid to globose, to c. 25 mm diam. Nuts ovoid, beaked, to c. 4 mm long, villous. Narrowleaf Drumsticks 99J v16_f99.jpg Restricted to coastal and nearby tablelands of central and southern N.S.W. Grows in dry sclerophyll forest and heathland on sandstone. Flower Sept.–Nov. 186 v16_m186.jpg N.S.W.: Bulee Gap, 7.1 km from Nerriga towards Nowra, H.Thompson 814 & S.Donaldson (CBG, MEL, NSW); Green Gully, Glen Davis, c. 40 km N of Lithgow, E.F.Constable 5155 (NSW); Wattamolla Rd, Royal Natl Park, D.J.McGillivray & E.F.Constable 5699 (NSW); Deanes siding (abandoned), c. 20 km NNE of Lithgow, B.G.Briggs 7177 & L.A.S.Johnson (NSW). 1 doubtful doubtful sp. Isopogon Isopogon divaricatus Roem. & Schult. Syst. Veg. 44 1817 T: n.v. According to A.D.Chapman, Austral. Pl. Name Index D–J: 1680 (1991), this taxon is based on the illegitimate name Protea divaricata Andrews, which is possibly based on cultivated material from the Cape of Good Hope. sp. doubtful Isopogon Isopogon pedunculatus R.Br. Suppl. Prodr. Fl. Nov. Holl. 5 1830 T: ora merid.-occid., Swan River [W.A.], 1827, C.Fraser 8; syn: BM. This is Petrophile seminuda Lindl. 2 doubtful doubtful sp. Isopogon Isopogon drummondii Jacques Ann. Fl. Pomone 216 1843 T: Nouvelle-Hollande, date and coll. unknown, fide A.D.Chapman, Austral. Pl. Name Index 2: 1680 (1991); not located. doubtful sp. Isopogon Isopogon eugellii Jacques Ann. Fl. Pomone 216 1843 T: Nouvelle-Hollande, date and coll. unknown, fide A.D.Chapman, Austral. Pl. Name Index 2: 1680 (1991); not located. sp. Isopogon Isopogon doubtful propinquus A.Cunn. ex Sweet Hort. Brit. 490 1827 T: New Holland, 1824, coll. unknown, fide A.D.Chapman, Austral. Pl. Name Index 2: 1681 (1991), not located, nom. inval., nom. nud. doubtful sp. Isopogon Isopogon diversifolius Ettingsh. Blatt-Skel. Dikot. 284, t. 12, fig. 2 1861 T: cultivirt im k.k. Hofgarten zu Schönbrunn, fide A.D.Chapman, Austral. Pl. Name Index 2: 1680 (1991), not located, nom. inval. Apparently based only on an illustration without a diagnosis. Data derived from Flora of Australia Volumes 16 (1995), 17A (2000) and 17B (1999), products of ABRS, ©Commonwealth of Australia