sign in sign up
<<
Home , Allocasuarina, Allocasuarina campestris, Allocasuarina crassa, Allocasuarina defungens, Allocasuarina dielsiana, Allocasuarina diminuta

FLORA OF

HAMAMELIDACEAE

H.J.Hewson

Shrubs or , monoecious, rarely dioecious. simple, alternate, rarely opposite, simple to palmately lobed, petiolate; stipules usually present. usually a spike, sometimes a raceme or panicle, bracteate. bisexual or unisexual, usually regular. 4 or 5, free or connate, small, sometimes absent. Petals 4 or 5, ligulate, small, sometimes absent. 4 or 5, free, often in 2 whorls with inner whorl staminodal; anthers usually basifixed, the connective often produced into an appendage, each locule (in Australia) with 2 pollen sacs and 1 valve. usually inferior, sometimes superior, with 2, rarely 3, carpels; styles 2, rarely 3, free, usually persistent in ; 1 or 2, pendulous, or 5–many, anatropous. Fruit (in Australia) a loculicidal capsule, woody. sometimes winged; present. A family of 26 genera and c. 100 , of subtropical to warm temperate regions around the world but predominantly in eastern . Three monotypic genera endemic in Australia. All 3 genera are in the tribe Hamamelideae of the subfamily Hamamelidoideae. The family has a long record and many representatives may be relictual. Species of Hamamelis L. (Witch-hazel) and Liquidambar L. are important ornamental . Species of Distylium Sieber & Zucc. and Loropetalum R.Br. have also been cultivated in Australia. H.Harms, Hamamelidaceae, Nat. Pflanzenfam. 2nd edn, 18a: 303–345 (1930); L.S.Smith, Hamamelidaceae in New species of and notes on plants. III, Proc. Roy. Soc. Queensland 69: 43–48 (1958); W.Vink, Hamamelidaceae, Fl. Males. 5: 363–379 (1958).

KEY TO GENERA

1 Stipules bristle-like, less than 5 mm long, leaving an obscure scar; flowers in open spikes, cream or white; staminodes absent 2. NEOSTREARIA 1: Stipules not bristle-like, more than 5 mm long, leaving an obvious scar; flowers in dense closed spikes, purple, red or green; staminodes present or absent 2 Stipules lanceolate, c. 7 mm long, leaving a curvilinear scar; petals expanding, pale green or yellow-green; staminodes 5 1. OSTREARIA 2: Stipules asymmetrically ovate, to 2 cm long, leaving an elliptic scar; petals inrolled, purple or red; staminodes absent 3. NOAHDENDRON

1. OSTREARIA

Ostrearia Baillon, Adansonia 10: 131 (1871); from the Latin ostrea (oyster), in reference to the bivalve-like form of the . Type: O. australiana Baillon Trees. Leaves entire, penniveined; stipules small, caducous leaving curvilinear scars. Spikes terminal, pedunculate, short and dense. Calyx 5-lobed above a short tube. Petals 5, linear. Stamens 5; anthers broad, cordate-ovate; connective developed to cover back of anther sacs and form an upcurved appendage; staminodes 5. Ovary half-inferior, bilocular; styles 2; 1 per locule. Capsule sessile, woody, compressed, obovoid, 2-locular, free or adnate with calyx tube leaving upper 2/3 free. Seeds obovoid or ellipsoidal. A monotypic endemic in northern Qld.

1 1. Ostrearia HAMAMELIDACEAE

Ostrearia australiana Baillon, Adansonia 10: 131 (1871) T: Rockingham Bay, Qld, ?J.Dallachy (ex herb. F.Mueller); holo: P n.v., fide L.S.Smith, Proc. Roy. Soc. Queensland 69: 43 (1958). Illustrations: L.S.Smith, op. cit. fig. 1i–p (1958); B.D.Morley & H.R.Toelken, Fl. Pl. Australia fig. 28a, b (1983). to 25 m tall; indumentum stellate. Leaves elliptic to lanceolate; lamina to 17 cm long and 6 cm wide; 7–12 mm long; stipules lanceolate, to 7 mm long. Spike on to 1 cm long; rachis to 3 cm long; bracts to 2 mm long; bracteoles to 1.5 mm long. Calyx lobes ovate, 3 mm long. Petals 7–9 mm long, 1–1.5 mm wide, pale green or yellow-green. Stamens c. 2 mm long; anther appendage c. 0.2 mm long; staminodes incurved, longer than stamens. Styles c. 0.3 mm long. Capsule glabrescent, 1.2–1.7 cm long, 1.2–2.4 cm wide, adnate with calyx tube for 3–5 mm. Seeds ellipsoidal, 1–1.5 cm long. Fig. 25A–C. Occurs from near Cooktown S to near Tully, Qld, near streams in at altitudes to 900 m. Region: CYRK. Map 1. Qld: Upper Parrot Ck, Annan R., L.J.Brass 20266 (BRI, CANB); Zarda, 16°27'S, 145°15'E, B.Hyland 2887 (BRI); Mt Finnigan, J.M.Powell 684 & J.A.Armstrong (NSW); Gap Ck, SSE of Cooktown, L.S.Smith \11209 (BRI); Jarrah Ck, near Tully, L.S.Smith & L.J.Webb 4897 (BRI). This species is the most common of the 3 Australian members of the family. The vestigial petal lobes which are usually absent place it close to the SE Asian genus Embolanthera Merr.

2. NEOSTREARIA

Neostrearia L.S.Smith, Proc. Roy. Soc. Queensland 69: 46 (1958); from the latin neo- (new) and Ostrearia (a closely related genus). Type: N. fleckeri L.S.Smith Trees. Leaves simple, penniveined; stipules small, caducous leaving obscure scars. Spikes terminal, pedunculate, elongate and open. Calyx 3-lobed above a short tube. Petals 5, linear. Stamens 5; anthers broad, cordate-ovate; connective developed to cover back of anther sacs and form a short apical appendage; staminodes absent. Ovary half-inferior, bilocular; styles 2; ovules 3 per locule. Capsule sessile, woody, compressed, obovoid, 2-locular, free from calyx tube in upper half. Seeds obovoid. A monotypic genus endemic in northern Qld.

Neostrearia fleckeri L.S.Smith, Proc. Roy. Soc. Queensland 69: 46, fig. 1a–h (1958) T: Babinda Ck, Happy Valley, Qld, May 1949, H.Flecker 12775; holo: BRI. Illustrations: L.S.Smith, loc. cit.; B.D.Morley & H.R.Toelken, Fl. Pl. Australia fig. 28c (1983). Tree to 12 m tall; indumentum stellate or lepidote. Leaves lanceolate to oblong-lanceolate; lamina to 18 cm long and 7 cm wide; petiole 3–12 mm long; stipules bristle-like, to 4 mm long; petiole 3–12 mm long. Spikes on peduncle to 3.5 cm long; rachis to 10 cm long; bracts to 2.5 mm long. Calyx lobes ovate, c. 3.5 mm long. Petals 1–1.3 cm long, 1.5–2 mm wide, cream to white. Stamens c. 1.5 mm long; anther appendage c. 0.1 mm long. Styles c. 0.2 mm long. Capsule 1–1.5 cm long, 1 mm wide. Seeds c. 1 cm long. Fig. 25D. Occurs from near Daintree S to near Tully, Qld, near streams in lowland rainforest. Region: CYRK. Map 2. Qld: Whyanbeel, 16°20'S, 145°20'E, B.Hyland 3028 R.F.K. (BRI); Boonjie, 17°25'S, 145°45'E, B.Hyland 6679 (BRI); The Boulders, near Babinda, J.G.Tracey 6302 (BRI). This genus is close to the SE Asian genera Maingaya Oliver and Embolanthera Merr. but differs from Maingaya in its elongate inflorescence and lack of staminodes and from

2

Figure 25. A–C, Ostrearia australiana. A, fruit ×0.5; B, stipule ×5; C, stipule scar ×5 (A–C, L.S.Webb & L.J.Webb 4897, BRI). D, Neostrearia fleckeri, stipule ×5 (B.Hyland 6679, BRI). E–F, Noahdendron nicholasii. E, habit ×0.5; F, stipule ×5 (E–F, K.Williams 82132, BRI). Drawn by D.Boyer.

3 2. Neostrearia HAMAMELIDACEAE

Embolanthera in its simple leaves and 2-valved anthers.

3. NOAHDENDRON

Noahdendron Endress, B.Hyland & Tracey, Bot. Jahrb. Syst. 107: 372 (1985); from Noah Ck (the type locality) and the Greek dendron (a tree). Type: N. nicholasii Endress, B.Hyland & Tracey Trees. Leaves distichous, simple, penniveined; stipules large, some persistent, others caducous leaving elliptic scars. Spikes terminal, pedunculate, dense, shorter than those of Neostrearia. Sepals 5, free. Petals 5, narrowly oblong, tightly inrolled. Stamens 5; anthers cordate-ovate; connective developed to cover back of anther sacs and form an apical appendage; staminodes absent. Ovary superior to half-inferior, bilocular; styles 2; ovules 3 per locule, 1 fertile. Capsule sessile, woody, 2-locular. Seeds ovoid. A monotypic genus endemic in northern Qld.

Noahdendron nicholasii Endress, B.Hyland & Tracey, Bot. Jahrb. Syst. 107: 372 (1985) T: Noah Ck, Qld, 16°07'S, 145°26'E, J.G.Tracey 14945; holo: Z n.v., fide P.K.Endress et al., loc. cit. Illustrations: P.K.Endress et al., op. cit. figs 1–29 (1985). Tree to 10 m tall; indumentum stellate. Leaves oblong to elliptic, acuminate; lamina to 30 cm long and 10 cm wide, attenuate at base; petiole 0.8–1.5 cm long; stipules asymmetrically ovate, attenuate at base, to 2 cm long, 1 cm wide. Spikes on peduncle to 5 cm long, pendent; rachis to 7 cm long; inflorescence bracts to 1 cm long; bracts c. 4 mm long. Sepals ovate-triangular, c. 3 mm long, densely stellate-hairy. Petals glabrous, c. 5 mm long (unrolled), red or purple. Stamens c. equal to unrolled petals, red; anther appendage long-apiculate, c. 1 mm long. Styles c. 1 mm long; capitate. Capsule 1 cm long and wide. Seeds c. 7 mm long. Fig. 25E–F. Restricted to the Cape Tribulation area, Qld, near streams in rainforest at altitudes to 100 m. Region: CYRK. Map 3. Qld: Cape Tribulation, 16°05'S, 145°29'E, K.A.W.Williams 82132 (BRI). This species is unique in the family with its inrolled and bent petals. The large stipules are not characteristic of the subfamily Hamamelidoideae.

ULMACEAE

H.J.Hewson

Trees or , evergreen or , monoecious. Leaves simple, mostly alternate, often distichous, pinnately nerved or 3-nerved at base, entire or serrate, petiolate; base often oblique; stipules extra- or intrapetiolar, free or connate, usually caducous. Inflorescence axillary, solitary, racemose or cymose. Flowers small, inconspicuous, regular, hypogynous or perigynous, unisexual or bisexual. 1-whorled with 4–8 lobes, usually 5. Stamens usually as many as perianth lobes, free or adnate to perianth tube; staminodes present or absent in female flowers. Ovary superior; carpels 2 or 3, fused, usually unilocular; pistillode present or absent in male flowers. Styles free. Ovule 1 per locule, usually pendulous. Fruit a , drupe or samara. Seeds with little or no endosperm. A family of 18 genera and c. 200 species in tropical and temperate regions in both hemispheres; in Australia 3 genera native, 1 naturalised. Gironniera Gaudich. has been

4 ULMACEAE recorded for Australia by L.Phuphathanaphong in Thai Forest Bull. (Bot.) 6: 49–59 (1972) but there appears to be no basis for this. G.Bentham, Tribe Celtideae in Urticeae, Fl. Austral. 6: 155–160 (1873); E.Soepadmo, Ulmaceae, Fl. Males. ser. 1, 8: 31–76 (1977).

KEY TO GENERA

1 Fruit a samara; perianth 4–8-lobed; ovary stipitate 1. ULMUS 1: Fruit a drupe; perianth 4- or 5-lobed; ovary sessile 2 Leaves coarsely serrate; serrations mucronate, usually pungent; female flowers solitary 2. APHANANTHE 2: Leaves entire to serrulate; serrations not pungent; female flowers more than 1 per inflorescence, or flowers bisexual 3 Fruit less than 5 mm wide 3. TREMA 3: Fruit more than 5 mm wide 4.

1. ULMUS

Ulmus L., Sp. Pl. 1: 225 (1753), Gen. Pl. 5th edn, 106 (1754); from an old Latin name for elm trees, possibly derived from the Celtic ulm. Type: U. campestris L. Trees, deciduous or semideciduous, usually developing suckers; leaves of suckers frequently different from adult leaves. Leaves alternate, pinnately veined, serrate or crenate, usually oblique at base; stipules extrapetiolar, caducous. Inflorescence of 3–15 flowers in clusters or cymes, produced before the leaves on short lateral shoots. Flowers bisexual, sometimes functionally unisexual. Perianth 4–8-lobed. Stamens 5 or 6, usually exserted; anthers reniform, extrorse. Ovary compressed, sometimes stipitate; style short; ovule 1. Fruit a dry flattened winged nutlet or samara. Seeds with straight embryo; endosperm absent. A genus of c. 25 species from , Asia and ; 1 species established in Australia.

*Ulmus hollandica Miller, Gard. Dict. 8th edn, no. 5 (1768) T: none designated. Large, spreading tree to 40 m tall; suckers numerous, forming thickets. Leaves broadly ovate, oblique at base, acuminate, double-serrate, lamina 6–14 cm long, scabrous above, glabrescent; petiole 0.5–1.5 cm long; stipules linear or lanceolate, 0.5–1 cm long. Flowers not seen. Samara flat, orbicular, 1–2 cm wide, broadly winged. Dutch Elm. Plants established around habitation in Qld, N.S.W., A.C.T., Vic. and Tas. Specimens are rarely collected, consequently distribution data and specimen citation are an understatement of the extent of establishment. Regions: MCPH, HOWE, TASM. Map 4. Qld: Mt Marshall, 8 Nov. 1963, E.Smith (BRI). N.S.W.: Murray R., near Ournie, Jan. 1976, J.Meakins (NSW). A.C.T.: Lanyon, I.Olsen 1378 (NSW). Tas.: Shot Tower, Hobart, W.M.Curtis (HO). F.Mueller, Fragm. 8: 101 (1873) recorded U. parvifolia Jacq. from Wide Bay, Qld. No further collections have been made and it is not regarded as established. J.H.Willis, Handb. Pl. Victoria 2: 27–28 (1972) noted that U. procera Salisb. is widely cultivated but not apparently naturalised. T.D.Raphael, Pap. & Proc. Roy. Soc. Tasmania 89: 150 (1955), listed U. thomasii Sarg. as an escape but it does not seem to be established.

5 2. Aphananthe ULMACEAE

2. APHANANTHE

Aphananthe Planchon, Ann. Sci. Nat. Bot. ser. 3, 10: 265, 337 (1848), nom. cons.; from the Greek aphanes (invisible) and anthos (flowers) in reference to the small inconspicuous flowers. Type: A. philippinensis Planchon Trees or shrubs, deciduous or semideciduous, monoecious. Leaves alternate, pinnately veined (in Australia); stipules extrapetiolar, free. Inflorescence axillary, unisexual, rarely with male and female flowers; bracts minute, caducous. Male inflorescence a condensed, many-flowered raceme, in lower axils of new shoots. Male flowers: pedicels short; perianth lobes 4 or 5, membranous, imbricate in ; stamens 4 or 5, inflexed in bud, included or exserted at anthesis; anthers introrse; pistillode absent. Female inflorescence in upper axils of new shoots, 1-flowered, sometimes 2 or 3 flowers of both male and female. Female flowers: pedicels long; perianth lobes 4 or 5, imbricate in bud; staminodes absent; ovary sessile; stigmas tubular. Fruit a drupe, slightly 3–5-angular or terete. with curved embryo; endosperm absent. A genus of 5 species in , Mexico and from E to Japan and the Solomon Islands and S to Australia; 1 species native in Australia. J.-F.Leroy, Le Genre Aphananthe (Ulmacées). Révision Systématique et Distribution Géographique des espèces, Bull. Mus. Hist. Nat. (Paris) ser. 2, 18: 118–123 (1946).

Aphananthe philippinensis Planchon, Ann. Sci. Nat. Bot. ser. 3, 10: 337 (1848) T: Luçonia, , Cunning 1311; n.v. Sponia ilicifolia Kurz, Flora 55: 448 (1872). T: Clarence R., N.S.W., J.v.Müller; n.v.; Brisbane R., Qld, J.v.Müller; n.v.; Rockhampton, Qld, J.v.Müller; n.v. Taxotrophis rectinervis F.Muell., Fragm. 6: 192 (1868); Aphananthe rectinervis (F.Muell.) Planchon in A.P. de Candolle, Prodr. 17: 208 (1873). T: Clarence R., N.S.W., Wilcox; syn: MEL; Rockingham Bay, Qld, ?F.Mueller; syn: MEL. Illustrations: J.H.Maiden, Forest Fl. 2: t. 50 (1907), W.D.Francis, Austral. Rainforest Trees 4th edn, figs 22, 23 (1981), B.D.Morley & H.R.Toelken, Fl. Pl. Australia fig. 29a–c (1983). or tree to 30 m tall; trunk sometimes buttressed. Leaves elliptic to broadly ovate, mostly serrate, the teeth distant, mucronate to pungent; lamina 3–10 cm long, rough-asperate; venation prominent below; petiole 3–5 mm long. Male flowers: tepals ovate-lanceolate, 1–1.5 mm long; stamens with filaments c. 0.5 mm long; anthers c. 0.75 mm long. Female flowers solitary; to 5 mm long; tepals narrowly ovate to lanceolate, 0.5–1 mm long, persistent; stigmatic arms recurved, 2–3 mm long, persistent. Drupe ovoid to globose, somewhat acuminate, 6–8 mm long, slightly angular, glabrescent; pedicel to 1 cm long. Native Elm, Axehandle , Greyhandle Wood. Fig. 26A–D. Occurs in coastal districts and up to 100 km inland, from , Qld, S to the Manning R., N.S.W.; grows in rainforest; also in the Philippines, and the Solomon Islands. Regions: CYRK, BURD, DWSN, MCPH. Map 5. Qld: Gracemere Stn, E.R.Anderson 3802 (BRI); Tamborine Mtn, J.Shirley 88056 (BRI); Wongabel, G.Stocker 655 (BRI, CANB). N.S.W.: Woolgoolga, Oct. 1934, W.A.W. de Beuzeville (NSW); Moore Park, Old Grevillea, R.Coveny 9953 & L.Haegi (CANB, NSW). A shrub in depauperate rainforest thickets to a canopy tree in monsoon rainforest. The timber has been used for linings in houses but more commonly for tool handles as the common names imply.

6

Figure 26. A–D, Aphananthe philippinensis. A, fruiting branch ×0.5 (near Canungra, Qld, K.Williams, BRI); B, ♂flower ×5 (E.Anderson 3802, BRI); C, ♀ flower ×5 (Tamborine Mtn, Qld, J.Shirley, BRI) D, fruit ×2.5 (J.Simmonds 485, BRI), E–F, Trema tomentosa var. tomentosa. E, fruiting branch ×0.5; F, fruit ×10 (E–F, J.Maconochie 1712, DNA). G–H, Trema tomentosa var. viridis. G, ♂flower ×10 (M.Rankin 1742, DNA); H, ♀flower ×10 (C.Dunlop 4792, DNA). Drawn by D.Boyer.

7 3. Trema ULMACEAE

3. TREMA

Trema Lour., Fl. Cochinch. 2: 539, 562 (1790); from the Greek trema (a hole) in reference to the pitting on the fruit. Type: T. cannabina Lour. Sponia Lam., Encycl. 4: 138 (1797). T: not designated. Trees or shrubs, evergreen, monoecious, with simple hairs and sometimes with multicellular glandular hairs. Leaves alternate, penninerved; stipules free, extrapetiolar, caducous. Inflorescence an axillary panicle or thyrse, either unisexual or with both male and female flowers; bracts minute, caducous. Male flower globular; perianth 4- or 5-lobed, induplicate-valvate; stamens 4 or 5, included or exserted; anthers dorsifixed, introrse; pistillode present, hirsute. Female flowers ovoid; perianth 4- or 5-lobed; staminodes usually present; ovary sessile, with short style; stigmas spreading or incurved. Fruit a drupe. Seeds with or without endosperm; embryo curved. A genus of c. 15 species around the world in tropical, subtropical and temperate regions; 2 species native in Australia. The genus was partially revised by E.Soepadmo, Fl. Males. ser. 1, 8: 47–55 (1977). He concluded that a world monograph is required. While his work helps somewhat to resolve the taxa in Australia, it seems that a satisfactory resolution cannot be found in the absence of a monographic treatment. In this treatment, the placement of Australian representatives of Trema does not entirely follow Soepadmo. All Australian representatives have black fruits. Hence T. cannabina Lour., with orange to red fruit, does not occur in Australia. Trema tomentosa is interpreted more broadly and is regarded as having 2 varieties. underside velvety over a dense hoary indumentum; leaf surface not visible 1. T. orientalis Leaf almost glabrous, scabrous, or velvety without an underlying indumentum; leaf surface visible 2. T. tomentosa

1. Trema orientalis (L.) Blume, Mus. Bot. 2: 62 (1856) Celtis orientalis L., Sp. Pl. 2: 1044 (1753). T: India, collector unknown; n.v. Illustration: E.Soepadmo, Fl. Males. ser. 1, 8: figs 13, 16 (1977). Tree to 30 m tall. Leaves usually coriaceous, ovate to narrowly elliptic or lanceolate, serrate, attenuate, the base cordate or rounded, symmetrical or oblique; lamina 4–20 cm long, markedly discolorous, scabrous above, velvety over hoary indumentum below; nerves 4–8 pairs; petiole 2–20 mm long; stipules linear-lanceolate, 3–10 mm long. Inflorescence few- to many-flowered, to 4 cm long. Flowers globose, c. 2 mm diam.; perianth persistent. Drupe ovoid, c. 3 mm wide, black. Occurs in northern N.T. and from Cape York S to Bundaberg, Qld; grows on sand dunes near mangrove swamps, in marginal rainforest and in alluvial soils in rainforest; also in and Asia. Regions: ARNM, CYRK, BURD, DWSN. Map 6. Qld: Mowbray R., L.J.Brass 2013 (BRI); Kelsey Ck, near Proserpine, N.Michael 1135 (BRI); Upper East Funnel Ck, Sarina Ra., M.E.Ritchie 2 (BRI); Gorge Ck, Netherdale, A.N.Rodd 3776 (BRI, NSW); c. 14 km NW of , R.Schodde 4134 (BRI). This species is characterised by the very dense hoary abaxial indumentum causing the epidermis to be invisible and the leaves to be markedly discolorous. E.Soepadmo, op. cit. 52 (1977), discussed three entities in T. orientalis as he treated it. He regarded Australian material as being included in ‘so-called T. discolor’. Trema discolor (Brongn.) Blume has not been formally applied to Australian material and Soepadmo included it as a synonym of T. orientalis.

8 ULMACEAE 3. Trema

2. Trema tomentosa (Roxb.) Hara, Fl. East. Himalaya 2: 19 (1971) Celtis tomentosa Roxb., Fl. Indica 2: 66 (1832). T: Chittagong, India, collector unknown; n.v. Shrub to 8 m tall. Leaves chartaceous to thickly coriaceous, broadly ovate to lanceolate, serrate to serrulate, acuminate, the base cordate to rounded, symmetrical or asymmetrical; lamina 1–14 cm long, mostly discolorous, usually scabrous above, slightly to strongly scabrous to scabrous-villous to velvety below; nerves 3–8 pairs, the lowest pair frequently giving a 3-nerved appearance; petiole 2–11 mm long; stipules linear-lanceolate, c. 5 mm long. Inflorescence few- to many-flowered, to c. 4 cm long. Flowers globose, c. 2 mm diam.; perianth persistent. Drupe ovoid, c. 2 mm diam., black. Peach Leaf Poison Bush, Native Peach. Occurs in coastal to near-coastal W.A., N.T., Qld and N.S.W.; also in tropical Africa, India, SE Asia and tropical Pacific islands to Hawaii. It has 2 varieties. The species is variable in leaf morphology and indumentum density. The variation is continuous and the recognition of taxa at the rank of species does not seem justifiable until monographic revisionary work is done. Two varieties are recognised here which take account of extremes in indumentum density on mature adult leaves. The determination of intermediates is subjective. It is made the more so by variation in indumentum density between sun and shade leaves. In no case is the indumentum hoary or sufficiently dense so as to obscure the epidermis. There are reports of varying toxicity in specimens. This may indicate that chemotaxonomy could be useful. Mature adult leaf underside velvety 2a. var. tomentosa Mature adult leaf underside scabrous to scabrous-villous 2b. var. viridis 2a. Trema tomentosa (Roxb.) Hara var. tomentosa [Trema amboinensis auct. non (Willd.) Blume: G.Bentham, Fl. Austral. 6: 159 (1873)] Adult leaves with velvety abaxial indumentum. Fig. 26E–F. Occurs in N.T., and from Cape York S to the Bunya Mtns, Qld; grows on margins of coastal rainforest and vine thickets and in eucalypt woodland. Regions: KIMB, ARNM, CYRK, BURD, DWSN, MCPH. Map 7. N.T.: tributary of Deaf Adder Ck, Kakadu Natl Park, M.Lazarides 9062 (CANB, DNA). Qld: 7 km from beach on track from Starke Stn to McIvor R. mouth, J.R.Clarkson 5188B (BRI); Eungella Tableland, 11 Mar. 1963, R.Matthews (BRI); Baileys Ck, c. 13 km ENE of Daintree, L.S.Smith 11598 (BRI); Lizard Is., R.L.Specht 306 (BRI). 2b. Trema tomentosa var. viridis (Planchon) Hewson, Fl. Australia 3: 190 (1989) Sponia viridis Planchon, Ann. Sci. Nat. Bot. ser. 3, 10: 319 (1848); Trema viridis (Planchon) Blume, Mus. Bot. 2: 58 (1856); Trema aspera var. viridis (Planchon) Benth., Fl. Austral. 6: 158 (1873). T: Port Essington, N.T., Armstrong 384; n.v. Celtis aspera Brongn. in L.I.Duperrey, Voy. Monde (Phan.) 213, t. 48 (Atlas) (1834); Sponia aspera (Brongn.) Decne, Nouv. Ann. Mus. Hist. Nat. 3: 498 (1834); Trema aspera (Brongn.) Blume, Mus. Bot. 2: 58 (1856); [Trema aspera var. typica Domin, Biblioth. Bot. 89: 560 (1921), nom. inval.]. T: Blue Mountains, near Port Jackson, N.S.W., R.P.Lesson & J.S.C.D. D’Urville; n.v. Trema aspera var. xerophila Domin, Biblioth. Bot. 89: 6 (1921). T: Chillagoe, Qld, 1910, K.Domin; n.v. [Trema cannabina auct. non Lour.: F.Mueller, Proc. Roy. Soc. New South Wales 14: 84 (1881); E.Soepadmo, Fl. Males. ser. 1, 8: 50 (1977)] Illustrations: Mahood in G.M.Chippendale, Poisonous Pl. N. Terr. 2: fig. 11 (1958); T.D.Stanley & E.M.Ross, Fl. S.E. Queensland 1: fig. 6H (1983). Adult leaves with scabrous to scabrous-villous abaxial indumentum. Fig. 26G–H. Occurs in the Kimberley, W.A., northern N.T. and disjunctly in the Macdonnell Ranges, and from Cape York, Qld, S to Mallacoota, Vic.; grows on margins of rainforest and vine

9 3. Trema ULMACEAE thickets and in eucalypt woodland. Regions: KIMB, ARNM, MACD, CYRK, BURD, DWSN, MCPH, NEPN, HOWE. Map 8. W.A.: island in Prince Frederick Harbour, K.F.Kenneally 8956 (PERTH). N.T.: c. 32 km E of Creek, N.Byrnes NB1344 (DNA). Qld: Burnett R. c. 32 km W of Bundaberg, R.C.Jansen 70 (BRI). N.S.W.: near Ballina, K.L.Wilson 3295 (NSW); c. 70 km NW of Grafton, M.Tindale & D.R.Brooke (NSW). E.Soepadmo, op. cit. 50 (1977), treated this taxon as T. cannabina Lour. However, T. cannabina has orange to red fruits and does not seem to occur in Australia.

4. CELTIS

Celtis L., Sp. Pl. 2: 1043 (1753), Gen. Pl. 5th edn, 467 (1754); a Latin name for an African species of Lotus, transferred to this genus by Linnaeus for no known reason. Type: C. australis L. Solenostigma Endl., Prodr. Fl. Norfolkicae 41 (1833); Celtis subg. Solenostigma (Endl.) Planchon, Ann. Sci. Nat. Bot. ser. 3, 10: 263, 305 (1848). T: S. paniculatum Endl. Trees or shrubs, evergreen or semideciduous, monoecious, sometimes with some bisexual flowers. Leaves alternate, 3-nerved at base; stipules lateral, free, scarious, enclosing bud, caducous. Inflorescence axillary on new shoots, a raceme or panicle; bracts minute, caducous. Male flowers globular; perianth lobes 4 or 5, imbricate in bud, free, recurved at anthesis; stamens 4 or 5, exserted at anthesis; anthers dorsifixed, extrorse; pistillode present or absent. Bisexual flowers ovoid; perianth lobes 4 or 5, as for male flowers; stamens 4 or 5; ovary 1-locular, sessile; style short or absent; stigmas elongate, entire or bifid, divergent; ovule pendulous. Female flowers: as for bisexual flowers but stamens reduced to staminodes. Fruit a drupe, ovoid or globose. Seed without endosperm; embryo curved. A genus of c. 60 species in tropical and temperate regions of the world; 2 species native in Australia and 3 species naturalised; one mainland species also on Norfolk Is.; a subspecies of a New Caledonian species endemic on Lord Howe Is. 1 Leaves serrate; inflorescence 1–few-flowered 2 Leaves ovate-lanceolate, the longest more than 10 cm long, serrate 3 Leaves scabrous above 1. C. australis 3: Leaves glabrous above 2. C. occidentalis 2: Leaves ovate, the longest not more than 10 cm long, serrate in upper half only 3. C. sinensis 1: Leaves entire; inflorescence many-flowered 4 Leaves markedly tri-nerved; fruit orange or red 4. C. philippensis 4: Leaves not markedly tri-nerved; fruit blue to purple-black 5. C. paniculata

1. * L., Sp. Pl. 2: 1043 (1753) T: southern Europe, collector unknown; n.v.; northern Africa, collector unknown; n.v. Tree to 25 m tall. Leaves ovate-lanceolate, acuminate, serrate or double-serrate, rounded or cordate at base, oblique; lamina usually 4–15 cm long, not markedly trinerved, scabrous above, velvety below; petiole 1–2 cm long; stipules 0.5–2 cm long. Inflorescence few-flowered. Male flowers: pistillode present. Bisexual flowers: perianth lobes ovate, c. 3 mm long; stamens not exceeding perianth; ovary ovoid-cylindrical; stigmatic arms spreading to horizontal or recurved. Drupe globose, 9–12 mm wide, brown to black; pedicel to 3.5 cm long. Native in Mediterranean Europe and Africa; naturalised in the central and southern tablelands region of N.S.W. and A.C.T. Regions: NEPN, HOWE. Map 9.

10 ULMACEAE 4. Celtis

N.S.W.: Mt Wellesley Gorge, Wellington district, 1977, G.W.Althofer (NSW). A.C.T.: Black Mtn, Canberra, H.S.McKee 11730 (CANB); north bank of Lake Burley Griffin near Scrivener Dam, I.R.Telford 10601 (CBG).

2. * L., Sp. Pl. 2: 1044 (1753) T: Virginia, [U.S.A.], collector unknown; n.v. Shrub or tree to c. 25 m tall. Leaves ovate-lanceolate to broadly ovate, acuminate, serrate, rounded at base, oblique; lamina 5–12 cm long, glabrous above, pubescent below, somewhat 3-nerved; petiole c. 1 cm long; stipules lanceolate, c. 1 cm long. Inflorescence few-flowered. Bisexual flowers usually solitary; perianth lobes linear-oblong, c. 3 mm long; stamens equal to or slightly longer than perianth; ovary ovoid-cylindrical, c. 4 mm long; stigmatic arms recurved. Drupe ellipsoidal or globose, 7–13 mm long, red to black; pedicel 0.7–2 cm long. Native of North America; naturalised in a few localities from Toowoomba, Qld, S to near Albury, N.S.W. Regions: MCPH, NEPN, HOWE. Map 10. N.S.W.: Razor Back Mtn, Camden, E.McBarron 15091 (NSW); Camden, Nepean R., Jan. 1960, J.K.Brown (NSW); Cobbitty, Apr. 1968, A.N.Rodd (NSW).

3. *Celtis sinensis Pers., Syn. Pl. 1: 292 (1805) T: from China; n.v. Tree to 20 m tall. Leaves ovate; lamina 4–10 cm long, glabrous, acuminate, entire in lower half, serrate in upper half; base rounded, oblique; venation 3-nerved at base; lateral veins 6–8; petiole 0.5–1 cm long; stipules not seen. Inflorescence 1- to few-flowered. Bisexual flowers usually 1–3 together; perianth lobes broadly ovate, c. 2 mm long; stamens slightly exceeding perianth; ovary cylindrical to ovoid, 1.5–3 mm long; stigmatic arms mostly divergent. Drupe globose, 7–9 mm long, orange to black; pedicel 0.4–1 cm long. Chinese Celtis. This species is established in south-eastern Qld. Region: MCPH. Map 11. Qld: Barambah, c. 6 km NE of Gayndah, B.Lebler & L.Durrington 29 (BRI, CANB); Yellow Gully, 3 km NE of Kin Kin, A.N.Rodd 3034 & S.Jacobs (CANB, NSW); Eidsvold, Apr. 1912, T.L.Bancroft (BRI); Brian Pastures, 11 Nov. 1952, S.L.Everist (BRI); Indooroopilly, L.W.Jessup 837 (CBG).

4. Celtis philippensis Blanco, Fl. Filipinas 197 (1837) [Celtis philippinensis Benth., Fl. Austral. 6: 156 (1873), orth. var.] T: Malaitmo, collector unknown; n.v. Celtis strychnoides Planchon, Ann. Sci. Nat. Bot. ser. 3, 10: 306 (1848). T: Careening Bay, [W.A.], A.Cunningham; ?iso: MEL. Solenostigma brevinerve Blume, Mus. Bot. Lugduno-Batavum 2: 67 (1856); Celtis brevinervis (Blume) Planchon, in A.P. de Candolle, Prodr. 17: 183 (1873). T: New Guinea, collector unknown; n.v. Illustrations: E.Soepadmo, Fl. Males. ser. 1, 8: fig. 18a–k (1977); D.Levitt, Plants and People fig. 80 (1981); B.D.Morley & H.R.Toelken, Fl. Pl. Australia fig. 29f, g (1983). Shrub or tree, to 15 m tall. Adult leaves elliptic to suborbicular, acuminate, mostly entire, rounded at base, sometimes oblique; lamina 4–18 cm long, markedly 3-nerved, glabrous to slightly scabrous above, glabrous to sparsely puberulous below; petiole 2–14 mm long; stipules ovate, 6–10 mm long, peltate. Inflorescence a many-flowered panicle, either male or male and bisexual. Flowers 5-partite. Male flowers: perianth lobes ovate-elliptic, 1.5–2 mm long; stamens 1.5–2.5 mm long; anthers subreniform. Bisexual flowers: perianth lobes ovate-elliptic, 2–2.5 mm long; stamens 1.5–3 mm long; ovary ovoid-cylindrical, 2–2.5 mm long; stigmatic arms 1–1.5 mm long, bilobed or bifid. Drupe ovoid, 7–15 mm long, orange or red; pedicel 0.3–1.5 cm long. Fig. 27A. Occurs in the Kimberley, W.A., in northern N.T. and from Cape York Peninsula S to Broad Sound, Qld. Grows in a wide range of habitats and in a wide range of soil types including basalt, quartzite, sandstone, limestone, coastal sand and black peaty soil. Habitats include open sclerophyll forest, vine forest and coastal dunes. Also occurs in tropical Africa,

11

Figure 27. Celtis. A, C. philippensis, fruiting branch ×0.5 (J.Must 1330, DNA). B–C, C. .paniculata. B, ♀flowering branch ×0.5; C, ♀flower ×10 (B–C, K.Williams 82157, BRI). Drawn by D.Boyer.

12 ULMACEAE 4. Celtis through India, SE Asia and Malesia to the Solomon Islands. Regions: KIMB, ARNM, BKLY, CARP, GGNA, CYRK, BURD. Map 12. W.A.: Walsh Point, Port Warrender, K.F.Kenneally 7815 (PERTH). N.T.: W of mouth of Alligator R., L.J.Webb & J.G.Tracey 12248 (BRI); Black Jungle, G.Wightman 301 & C.R.Dunlop (DNA). Qld: Cardwell Gap, Oct. 1976, S.L.Everist (BRI); South Percy Is., G.Porter 34 (BRI). Celtis philippensis has flat cotyledons. The species is markedly xerophytic in drier habitats. In W.A. and adjacent N.T. the leaves are the smallest, apiculate, with more-persistent indumentum and often with toothed margins. In Australia the trees do not reach the sizes reported by E.Soepadmo in Fl. Males. ser. 1, 8: 62 (1977). Australian specimens are assignable to C. philippensis Blanco var. philippensis, although 2 collections from Lawn Hill, Qld — L.J.Tracey & J.G.Webb 10650 (BRI), and May 1974, T.Farrell (BRI) — both have leaves with 1 or 2 pairs of nerves and may be referable to C. philippensis var. wrightii (Planchon) Soep. The fruit is eaten by Aborigines. The Bardi Tribe of the Kimberley region name it gulyindji. The Groote Eylandt Aborigines call it angarrakaba.

5. Celtis paniculata (Endl.) Planchon, Ann. Sci. Nat. Bot. ser. 3, 10: 305 (1848) Solenostigma paniculatum Endl., Prodr. Fl. Norfolkicae 42 (1833). T: Norfolk Is., F.Bauer; n.v. C. ingens F.Muell., Fragm. 4: 88 (1864). T: Hastings R., N.S.W., H.Beckler; ?syn: MEL; Richmond R., N.S.W., C.Moore; ?syn: MEL; Clarence R., N.S.W., C.Moore; ?syn: MEL; Moreton Bay, Qld, F.Mueller; syn: n.v.; Edgecombe Bay, Qld, J.Dallachy; syn: MEL; Sweers Is., Qld, D.Henne; syn: MEL. Illustration: E.Soepadmo, Fl. Males. ser. 1, 8: 59, fig. 20a–d (1977). Tree to 12 m tall. Leaves elliptic to ovate-elliptic, bluntly acute, entire, sometimes undulate, rounded at base, sometimes oblique; lamina 5–15 cm long, not markedly 3-nerved, glabrous or sparsely puberulous; petiole 6–15 mm long; stipules broadly ovate, 4–5 mm long, peltate. Inflorescence a many-flowered panicle, either male or female. Flowers 5-partite. Male flowers: perianth lobes ovate, 1–1.5 mm long, membranous; stamens 1–2 mm long; anthers subreniform. Female flowers: perianth lobes ovate to broadly ovate, 0.5–1 mm long, membranous, sometimes emarginate or bifid; staminodes rudimentary; ovary ovoid, 1.5–2 mm long; stigmatic arms bifid. Drupe ovoid, 7–12 mm long, blue to purple-black; pedicel 2–6 mm long. Fig. 27B–C. Occurs from Cape York, Qld, S to Kiama, N.S.W. Grows in a wide range of soil types (basaltic, granitic, calcareous, sandy and humus-rich) and in a wide range of habitats (sandy beaches, vine forest and sclerophyll forest) being somewhat mesophytic in wetter habitats and xerophytic in drier habitats. Also found in , , Malesia and Norfolk Is. Regions: CYRK, BURD, DWSN, MCPH, NEPN. Map 13. Qld: Bunjurgen, May 1985, L.H.Bird (BRI); Cape York, K.J.White 1215 (BRI); Cape Tribulation, K.A.W.Williams 82157 (BRI). N.S.W.: Ballina, W.Bäuerlen 1132 (NSW). This species has folded cotyledons. A collection made on Sweers Is., 1901, J.F.Bailey (BRI) claims to be a collection from the ‘Investigator Tree’. D.J.Mabberley, Jupiter Botanicus 103 (1985) reported that ‘left “the name of ‘H.M.S. Investigator’, with the date 1802” carved on a tree, which, with “heaps of broken bottles”, was all that remained in 1880 of the thriving township of Carnarvon which grew up ... The tree blew down in 1887 bearing thirteen ships’ inscriptions including that of the Beagle (1841).’ If the Bailey collection is authentic then the tree must have persisted. The wood of this species has been used for making tools which require the timber to be pliable, e.g. whip handles.

13

CANNABACEAE

R.D.Pearce

Shrubs or herbs, erect or climbing, usually dioecious, annual or perennial, with aromatic granules. Leaves stipulate, opposite or alternate, simple to palmately-nerved or palmately-compound. Male inflorescence paniculate; female inflorescence a spike or spike-like, the bracteate flowers usually dense and in cone-like structures. Flowers apetalous. Male flowers pedicellate; perianth 5-partite, with imbricate sepals; stamens 5, erect in bud; filaments shorter than anthers. Female flowers sessile; perianth undivided; ovary solitary, unilocular; ovule 1, pendulous; style central; stigmas 2. Fruit a nut or achene, subtended by whole or part of perianth or bract. Endosperm oily. A family of 2 genera with c. 4 species native to northern temperate regions; 2 species are grown as crops of agriculture, and 1 as an ornamental. In the past the 2 genera have been treated under a number of different families: Artocarpaceae, , and . W.T.Stearn, ed. C.R.Joyce & S.M. & A.Curry, The and Chemistry of (1970).

KEY TO GENERA

Climbers; female inflorescence cone-like; leaves usually 3–5-lobed 1. HUMULUS Erect herbs; female inflorescence not cone-like; leaves divided into (3–9) leaflets 2. CANNABIS

1. HUMULUS

Humulus L., Sp. Pl. 1028 (1753); Gen. Pl. 5th edn, 453 (1754); from the Latin humus (the earth), the plants being normally prostrate. Type: H. lupulus L. Annual or perennial twining climbers, dioecious. Stems terete, glabrous, or scabrous when young. Leaves ovate-cordate, 3–5-lobed, opposite. pendent; male much-branched; female lax, cone-like. A genus of 2 species in the ; 1 species sparsely naturalised in Australia.

*Humulus lupulus L., Sp. Pl. 1028 (1753) T: herb. Linn; n.v. Illustration: S.Ross-Craig, Draw. Brit. Pl. 27: 7 (1970). Stems to 6 m long, scabrid with reflexed hairs. Leaves opposite, to 10–13 cm long, serrate; upper surface with appressed hairs with squamellate bases or only the bases remaining; lower surface paler, bearing yellow, resinous, bitter and aromatic granules; petiole to 9 cm long. Peduncle slender, scabrous. Female inflorescence cone-like, c. 3 cm long at maturity; flowers subtended by large, persistent bracts; bracts ovate-acute, pale green to light brown, with aromatic granules, yellow-amber at maturity; outer surface of bracts pubescent or with appressed hairs outside, prominently veined inside. Seed c. 3 mm long, 2 mm wide, acute, with a central rib, shiny, brown, enclosed by a single large perianth bract. Hops. Indigenous in Europe and Western Asia N of 32°; in Australia cultivated in cooler areas of S.A., N.S.W. and Tas., but few collections outside cultivation. Regions: EYRE, HOWE, TASM. Map 14.

14 CANNABACEAE 1. Humulus

S.A.: Hindmarsh Valley, R.M.Kain, Dept. of Agric. 532 (AD). N.S.W.: Sams Ck, Cobargo, Mar. 1962, M.Bellert (NSW). Tas.: 1.6 km S of Darlington, Maria Is., J.H.Hemsley 6737 (NSW). Has a long record of usage, mostly in the brewing industry, but also in the past in bread-making and for its herbal properties.

2. CANNABIS

Cannabis L., Sp. Pl. 1027 (1753); Gen. Pl. 5th edn, 453 (1754); from kannabis, the Greek name for hemp. Type: C. sativa L. Erect branching herbs, dioecious or occasionally both sexes intermixed on the same . Stems ribbed. Leaves with 3–9 leaflets, palmate, the lower opposite, upper alternate. Male inflorescence paniculate, erect; female inflorescence spicate, erect. A genus of 1 or 2 species in Central Asia; 1 species sparingly naturalised in Australia.

1. *Cannabis sativa L., Sp. Pl. 1027 (1753) T: hort. Cliff.; lecto: BM n.v., fide W.T.Stearn, Bot. Mus. Leaflet Harvard Uni. 23: 325–336 (1974). Illustration: B.D.Morley & H.R.Toelken, Fl. Pl. Australia fig. 32a–c (1983). Plant to 6 m high. Stems with dense appressed hairs. Leaflets 3–9, linear-lanceolate, serrate; margins slightly reflexed; upper surface with tubercle based hairs; lower surface with resinous granules; hairs dense, deflexed; stipules lanceolate; petiole to 9 cm long. Male flowers: stamens drooping. Female flowers: bracts glandular-hairy, ciliate; stigma 8–10 mm long, with unicellular hairs, exserted beyond bracts; bracts persistent. Achenes persistent on plant, 1.5 mm long, pale-brown with dark brown blotching and reticulate venation. Hemp. Native to central Asia; illegally cultivated throughout settled areas of Australia and probably sparingly naturalised in S.A., Qld, N.S.W. and Vic. Regions: MCPH, NEPN, HOWE, EYRE. Map 15. S.A.: Adelaide Plains, 26 Feb. 1972, A.G.Spooner (AD). N.S.W.: Narrabri, 18 Oct. 1982, G.Radunz (NSW); Lochinvar, Hunter R. Valley, Oct. 1969, G.Schroder (NSW); Penrith, 26 Dec. 1912, A.A.Hamilton (NSW). Has a long history of cultivation and now grown world-wide in temperate areas for the production of hemp fibre and illegally for narcotic resin (hashish, marijuana). Material from female plants is preferred for smoking. There is considerable literature on its pharmacology, social effects and legal aspects of its cultivation, e.g. F.Crowley & L.Cartwright, A Citizen’s Guide to Marihuana in Australia (1977); W.Emboden, Narcotic Plants (1972).

MORACEAE

W.-L.Chew

Shrubs, trees, climbers or herbaceous plants, monoecious or dioecious, usually with milky latex. Leaves alternate or opposite, simple, or (not in Australia) pinnate or palmate, usually with cystoliths, stipulate. Inflorescence axillary, unisexual or bisexual, paniculate, racemose, spicate, discoid, capitate or urceolate. Flowers small, unisexual, apetalous. Tepals usually 4, or up to 8, free or united, imbricate or valvate, persistent, or absent. Stamens opposite tepals, or reduced to 3, 2 or 1; filaments straight, free or connate, or inflexed; anthers large and mucronate, to small, bilobate and non-mucronate. Ovary superior, inferior or immersed in sockets in inflorescence, usually 1-locular; styles 1 or 2; stigma simple or bifid; ovule 1,

15

anatropous or campylotropous, generally apical. Pistillode present or absent in male flowers. Fruit drupaceous, free, or connate in fleshy syncarps or syconia (figs), or achenes. Seeds small to large, with endocarp; testa membranous or disintegrated; embryo curved or straight; cotyledons plicate, conduplicate, or flat. n = 12, 13, 14, with intrageneric polyploidy. A family of c. 53 genera and over 1400 species, widely pantropical with few extensions to temperate regions. Represented in Australia by 7 genera and 47 species. The family contains a number of valuable food plants such as the Fig ( carica L.), the Mulberry ( alba L.), the and the ( spp.). Over 40 species of Ficus are cultivated as indoor ornamentals and garden and roadside trees. toxicaria Leschen. produces a highly toxic latex. J.Lindley, Moraceae, The Vegetable Kingdom 2nd edn, 266–268 (1847); J.Lindley, Artocarpaceae, op. cit. 269–271; E.Bureau, Moraceae, in A.P. de Candolle, Prodr. 17: 211–279 (1873); E.Bureau, Artocarpaceae, op. cit. 280–288; G.Bentham, & , Fl. Austral. 6: 160–182 (1873); G.Bentham & J.D.Hooker, Moreae & Artocarpeae, Gen. Pl. 3: 357–378 (1880); E.Hurst, Poisonous Plants of New South Wales 74–77 (1942); E.J.H.Corner, The Classification of Moraceae, Gard. Bull. 19: 187–252 (1962); C.C.Berg, Some Remarks on the Classification and Differentiation of Moraceae, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 386: 1–10 (1973); C.C.Berg, Revisions of African Moraceae, Bull. Jard. Bot. Etat 47: 267–407 (1977); M.O.Rankin, The Family Moraceae in the , N. Terr. Bot. Bull. 5: 1–69 (1982).

KEY TO TRIBES

1 Flowers inside urceolate receptacles (figs); styles not exserted from figs; orifice of fig closed by bracts Trib. 3. FICEAE 1: Flowers not in urceolate receptacles, if so orifice of receptacle not closed by bracts and styles exserted beyond it 2 Inflorescence spicate to racemose or capitate or sometimes in shortly branched cymes, rarely discoid; stamens inflexed in bud; branches not self-pruning Trib. 1. MOREAE 2: Inflorescence discoid to turbinate, sometimes cyathiform or capitate; stamens straight in bud; branches spirally self-pruning Trib. 2.

KEY TO GENERA

1 Flowers crowded on turbinate or urceolate receptacle 2 Shrubs, climbers, stranglers or trees; flowers inside an urceolate receptacle (fig) 7. FICUS 2: Herbs; flowers on a linear to orbicular turbinate or flat receptacle † 1: Flowers not so 3 Herbs shorter than 1 m 3. 3: Trees, shrubs taller than 1 m, or woody climbers 4 Woody climbers with axillary spines 4. 4: Trees, shrubs or climbers without spines 5 Woody climbers 2. MALAISIA 5: Trees or shrubs 6 Male and female inflorescences capitate, globose to clavate; male flowers with 1 ; fruit a fleshy syncarp 5. ARTOCARPUS 6: Male inflorescences discoid or spicate, female inflorescences few- flowered, not capitate; fruit not a syncarp

16 MORACEAE Key to genera

7 Male inflorescence discoid; male flowers with 2–4 stamens 6. ANTIARIS 7: Male inflorescence a spike; male flowers with 4 stamens 8 Deciduous trees; leaves broadly ovate to cordate, the lamina thinly chartaceous; female tepals pulpy in fruit † MORUS 8: Evergreen shrubs or trees; leaves ovate, elliptic or oblong, the lamina coriaceous; female tepals not pulpy in fruit 1. † Dorstenia arifolia Lam., a native of , has been reported as a weed in Qld, e.g. F.M.Bailey, Queensland Fl. 5: 1464 (1902), but does not appear to be naturalised. Morus alba L. is widely cultivated in Australia for its edible fruit and sometimes escapes, but does not appear naturalised.

Trib. 1. MOREAE

Moraceae trib. Moreae Gaudich., Voy. Uranie 509 ‘1826’ (1830). Type: Morus L. Moraceae trib. Artocarpeae R.Br., Bot. Congo 454 (1818). T: Artocarpus Forster & G.Forster Moraceae trib. Strebleae Bureau in A.P. de Candolle, Prodr. 17: 215 (1873). T: Streblus Lour. Moraceae trib. Fatoueae Benth. & J.D.Hook., Gen. Pl. 3: 344 (1880). T: Fatoua Gaudich. Trees, shrubs or herbs, sometimes with thorns; latex present except in Malaisia and Fatoua. Leaves alternate, often distichous; stipules mostly free. Inflorescence usually unisexual, spicate, racemose or capitate, sometimes cymose, rarely discoid and involucrate. Perianth of female flower with free or connate tepals, sometimes absent. Stamens straight or inflexed in bud. Pistillodes generally present in male flower. Fruit free or adnate to perianth. Seeds small to very large, with or without endosperm. A tribe with 15 genera largely in the Old World tropics; 5 genera in Australia. Weak distinction between tribes Moreae and Artocarpeae and the lack of unifying attributes within each resulted in their amalgamation by Berg (1977). Generic delimitation is also a major problem in this group. The tribe needs to be investigated from a much broader range of attributes than gross morphology in order to obtain a more satisfactory classification.

1. STREBLUS

Streblus Lour., Fl. Cochinch. 614 (1790); from the Greek streblos (crooked), probably in reference to the crooked trunk of the type species. Type: S. asper Lour. Paratrophis Blume, Mus. Bot. Lugd. . 2: 81 (1856). T: P. heterophyllus Blume Pseudomorus Bureau, Ann. Sci. Nat. ser. 5, 11: 371 (1869). T: P. brunoniana (Endl.) Bureau Monoecious trees or shrubs. Leaves simple, distichous, petiolate, without glands; stipules small. Inflorescence unisexual, rarely bisexual, pedunculate. Male inflorescence a many-flowered spike, with a sterile often spiral groove on rachis; female a few-flowered spike shorter than the male. Male flowers: tepals 4, free, pubescent; stamens 4, with inflexed filaments; pistillode present. Female flowers: tepals 4, free or slightly connate; ovary superior; stigma bifid. Fruit usually an achene, ovoid to globular, sometimes fleshy; embryo diverse. A genus of some 25 species widely spread from Madagascar through S and SE Asia to Australia and the Pacific; 2 species in Australia; 1 species endemic on Norfolk Is. The Australian species are placed in sect. Paratrophis. E.J.H.Corner, Streblus, The Classification of Moraceae, Gard. Bull. Singapore 19: 215–222 (1962).

17 1. Streblus MORACEAE

Leaves often scabrid, dentate; lateral veins 8–12 pairs, oblique to midrib; petiole c. 5 mm long; male spikes densely flowered, sterile groove not readily visible; achene globular 1. S. brunonianus Leaves glabrous, faintly crenulate; lateral veins 15–20 pairs, ±perpendicular to midrib; petiole c. 15 mm long; male spikes not densely flowered, sterile groove readily visible; achene ovoid 2. S. glaber

Sect. Paratrophis

Streblus sect. Paratrophis (Blume) Corner, Gard. Bull. Singapore 19: 216 (1962). Paratrophis Blume, Mus. Bot. 2: 81 (1856). T: Paratrophis heterophyllus Blume, nom. illeg. = S. heterophyllus Corner Inflorescence unisexual. Male inflorescence elongated, unbranched, with a sterile groove. Male flowers sessile, 4-merous; pistillode quadrate-columnar. Female flowers sessile; stigma bifid.

1. Streblus brunonianus (Endl.) F.Muell., Fragm. 6: 192 (1868) Morus brunoniana Endl., Atakta Bot. t. 32 (1835); Pseudomorus brunoniana (Endl.) Bureau, Ann. Sci. Nat. ser. 5, 11: 373 (1869); P. brunoniana var. australiana Bureau, op. cit. 373; P. pendulina var. australiana (Bureau) Stearn, J. Arnold Arbor. 28: 427 (1947). T: t. 32 in Endl., loc. cit. Illustrations: S.Endlicher, loc. cit.; J.H.Maiden, For. Fl. New South Wales 8 (75): t. 286 (1924); W.D.Francis, Australian Rainforest Trees 3rd edn, 73, fig. 26 (1970). Tree or shrub to 15 m; branchlets lightly pubescent, glabrescent. Leaves elliptic, ovate to lanceolate, acute to acuminate, cuneate at base, dentate; lamina 5–8 cm long, 2.5–4 cm wide, scabrid, glossy on upper surface, finely pubescent to scabrid on lower surface; lateral veins 8–12 pairs, oblique, the intercostals prominent; petiole c. 5 mm long. Inflorescence shorter than leaves. Male spikes 2–5 cm long, densely flowered, the sterile groove obscured. Male flowers c. 1.5 mm long and wide; tepals pubescent; stamens c. 1.5 mm long; pistillode quadrate-columnar. Female inflorescence to 3 cm long, sometimes reduced to 1 flower. Female flowers c. 2 mm long; stigma branches c. 1.5 mm long, pubescent. Achenes globular, 3–5 mm diam. White Handlewood, Whale Bone Tree, Prickly Fig. Occurs from north-eastern Qld S to Milton, N.S.W.; grows in rainforest in wetter parts of ranges, tablelands, slopes and riverine habitats. Regions: CYRK, BURD, DWSN, MCPH, NEPN, HOWE. Map 16. Qld: S of Canungra, 19 Nov. 1970, K.Williams (BRI, NSW); Brisbane R., Dec. 1917, C.T.White (BRI, NSW). N.S.W.: Gloucester Tops, R.Coveny 727 (NSW); near Cobbity trig., R.Coveny et al. 7388 (NSW); Terrara, Shoalhaven, 15 Mar. 1939, F.A.Rodway (NSW). A highly variable species long considered conspecific with S. pendulinus (Endl.) F.Muell. of Norfolk Is., from which it is easily distinguished by the considerably shorter male inflorescence and the less hairy leaves. Superficially similar to Aphananthe philippinensis. Sap reported to be an irritant to human eyes.

2. Streblus glaber (Merr.) Corner, Gard. Bull. Singapore 19: 221 (1962) var. australianus (C.White) Corner, loc. cit. Paratrophis australiana C.White, Contr. Arnold Arbor. 4: 15 (1933). T: Herberton Ra., Qld, S.F.Kajewski 1378; holo: BRI; iso: NSW. Illustration: C.T.White, op. cit. t. II. Tree to 20 m; branchlets glabrous. Leaves narrowly ovate, elliptic to oblong, acuminate to caudate, cuneate to rounded at base, entire to crenulate; lamina 5–12 cm long, 1.5–3.5 cm wide, glabrous; lateral veins 15–20 pairs, ±perpendicular to midrib; petiole c. 15 mm long. Male spikes to 4 cm long, slender. Male flowers with tepals densely pubescent; stamens to 2.5 mm long; pistillode slender, columnar. Female inflorescence c. 1.5 cm long. Female

18 MORACEAE 1. Streblus flowers c. 2 mm long; stigma branches to 2 mm long. Achenes ovoid, 5 mm long, 3 mm wide. Fig. 28A–B. An endemic variety in north-eastern Qld.; grows in montane rainforest. Region: CYRK. Map 17. Qld: Davies Ck, L.S.Smith 12072 (BRI); Mt Hemmant, N of Noah Ck, L.J.Webb & J.G.Tracey 11970 (BRI); Koolmoon Ck, L.S.Smith 10466 (BRI); Herberton Ra., S.F.Kajewski 1378 & 1383 (BRI, NSW). The type variety occurs in Malesia, including New Guinea.

2. MALAISIA

Malaisia Blanco, Fl. Filip. 789 (1837); from Malaisis, the Philippine local name of the species. Type: M. tortuosa Blanco = M. scandens (Lour.) Planchon Dioecious deciduous scrambling shrubs or climbers with or without latex. Leaves alternate, simple, petiolate; stipules small. Inflorescences unisexual, pedunculate. Male inflorescence a spike, densely flowered. Male flowers 3- or 4-partite; filaments inflexed in bud; pistillode present. Female inflorescence capitate. Perfect female flowers: perianth urceolate with small dentate orifice enveloping ovary; style long; stigma bifid. Fruit an achene, ovoid, several in a shallow receptacle; embryo incurved, the longer cotyledon enveloping the smaller. A monotypic genus mainly in SE Asia but extending to Australia and the Pacific. G.Bentham, Malaisia, Fl. Austral. 6: 180–181 (1873); E.J.H.Corner, Malaisia, The Classification of Moraceae, Gard. Bull. Singapore 19: 240–241 (1962).

Malaisia scandens (Lour.) Planchon, Ann. Sci. Nat. ser. 4, 3: 293 (1855) Caturus scandens Lour., Fl. Cochinch. 612 (1790). T: from Indo-China, J. de Loureiro; BM n.v., fide E.D.Merrill, Trans. Amer. Phil. Soc. 24: 133 (1935). subsp. scandens M. viridescens Planchon, loc. cit.; M. tortuosa var. viridescens (Planchon) Bureau in A.P. de Candolle, Prodr. 17: 222 (1873); Caturus viridescens (Planchon) Seemann, Fl. Vitiensis 254 (1868). T: east coast of Australia, A.Cunningham; n.v. M. cunninghamii Planchon, op. cit. 294; Caturus cunninghamii (Planchon) Seemann, loc. cit. T: Brisbane R. and Moreton Bay, Qld, A.Cunningham; n.v. M. acuminata Planchon, op. cit. 294; M. tortuosa var. acuminata (Planchon) Bureau, loc. cit.; Caturus acuminatus (Planchon) Seemann, loc. cit. T: Mt Marshall, Illawarra District, N.S.W., J.Backhouse; holo: K. Illustrations: C.Gaudichaud, Voy. Bonite, Bot. t. 97 (1844–1851) as Dumartroya fagifolia Gaudich.; N.C.W.Beadle, Stud. Fl. N.E. New South Wales 4: 502, fig. 217C (1980). Scrambling or climbing shrub; stem densely lenticellate, glabrous. Leaves ovate, elliptic to oblong, acuminate, cuneate to rounded at base, often slightly asymmetrical, entire; lamina 8–10 cm long, 2.5–5 cm wide, lightly scabrid on lower surface, coriaceous; lateral veins 8–10 pairs, prominent; petiole 0.5–1 cm long. Male spikes 1–1.5 cm long. Male flowers sessile, c. 1.5 mm long and wide; tepals densely pubescent. Female capitulum to 5 mm diam., usually only 1 or 2 flowers developing. Female flowers sessile; orifice of perianth tube fringed with hairs; ovary c. 1 mm long; stigma bifid, to 5 mm long. Achenes c. 8 mm long, 5 mm wide, 1–4 in a shallow red receptacle. Crow Ash. Fig. 28C–D. Occurs from the Kimberley, W.A., through northern N.T., and from north-eastern Qld S to Batemans Bay, N.S.W.; grows in rainforest in diverse soil types, and reported to be frequent in disturbed forest. Regions: KIMB, ARNM, CYRK, BURD, DWSN, MCPH, NEPN, HOWE. Map 18.

19 2. Malaisia MORACEAE

W.A.: Walsh Point, Port Warrender, K.F.Kenneally 7826 (PERTH). N.T.: Daly R. at Claravale, M.O.Rankin 1672 (DNA); Whitestone Ck, M.O.Rankin 1582 (DNA, NSW). Qld: Mt Glorious, C.T.White 1960 (BRI, NSW). N.S.W.: Upper Williams R., F.A.Rodway 1884 (NSW). A distinctive species with fig-like leaves, but rarely confused with climbing species of Ficus. Malaisia scandens comprises 2 subspecies: the type subspecies in mainland Australia and subsp. megacarpa Green, an endemic of Lord Howe Is. (P.S.Green, J. Arnold Arbor. 67: 113, 1986). Subsp. scandens differs from subsp. megacarpa in having acute to acuminate (not long-acuminate) leaf lamina and considerably smaller achenes.

3. FATOUA

Fatoua Gaudich., Voy. Uranie 509 (1830); origin of generic name unknown. Type: F. pilosa Gaudich. Monoecious annual herbs without latex. Leaves alternate, simple, petiolate; stipules intra-petiolar, small. Inflorescence a bisexual capitate cyme. Perianth parts free or slightly connate, lightly villous. Male flowers: tepals 4, stamens 4; filaments inflexed in bud; pistillode present. Female flowers: tepals 4–6; ovary obovoid, often stipitate; style lateral, with a minute branch near base; stigma filiform. Fruit an achene, warted, asymmetrically globular to ovoid, slightly compressed, enclosed by persistent perianth; embryo curved with cotyledons equal. A genus of 2 species from Madagascar to E Asia and Australia; 1 species in Australia.

Fatoua pilosa Gaudich., Voy. Uranie 509 (1830) T: Moluccas, C.Gaudichaud; holo: P. Illustrations: C.Gaudichaud, Voy. Bonite, Bot. t. 84 (1844); C.L.Blume, Mus. Bot. 2: t. 38 (1857) as F. japonica Blume. Herb less than 1 m high; rootstock present. Leaves ovate, broadly ovate to triangular, acute to acuminate, rounded to cordate at base, dentate; lamina to 7 cm long, 4 cm wide, sparsely armed with short stiff hairs, chartaceous; lateral veins 6–8 pairs; petiole 1–3 cm long. Inflorescence c. 7.5 mm diam., shorter than leaves. Male flowers pedicellate; tepals 4; stamens 4; pistillode minute. Female flowers subsessile; tepals 4 or 5. Arzerarzer. Fig. 28E. Occurs in the Kimberley, W.A., northern N.T. and north-eastern Qld; also in Indo-Malesia. Regions: KIMB, ARNM, CYRK. Map 19. W.A.: Walcott Inlet, W Kimberley, A.V.Milewski 22 (PERTH); Osborne Is., Bonaparte Archipelago, P.G.Wilson 11065 (PERTH). N.T.: Waterhouse R. bridge, J.D.Briggs 883, 889 & 891 (CBG); Beswick Stn, N.Byrnes 2606 (DNA). Qld: Murray Is., M.Lawrie 53 (BRI). Superficially similar to Laportea interrupta (L.) Chew of the Urticaceae. They can be distinguished easily by touch as the latter species is heavily armed with irritant hairs.

4. MACLURA

Maclura Nutt., Gen. N. Amer. Pl. 2: 233 (1818), nom. cons.; after William Maclure, a noted American geologist. Type: M. aurantiaca Nutt. Cudrania Trecul, Ann. Sci. Nat. ser. 3, 8: 122 (1847), nom. cons. T: C. javanensis Trecul Dioecious scrambling shrubs or small trees with axillary spines. Leaves alternate, often distichous, simple, petiolate, without glands; stipules lateral, small. Inflorescence unisexual, pedunculate. Male inflorescence paniculate or capitate. Male flowers: tepals 4, rarely 5, free; stamens 4, rarely 5; filaments erect or reflexed in bud; pistillode small to ?absent. Female

20

Figure 28. A–B, Streblus glaber var. australianus. A, ♂flowering branchlet ×1 (S.Kajewski 1383, NSW); B, ♀flowering branchlet ×0.5 (S.Kajewski 1378, BRI). C–D, Malaisia scandens. C, flowering branchlet ×0.5; D, fruit and pedicel ×1 (C–D, not recorded). E, Fatoua pilosa, flowering branchlet ×0.5 (M.Lawrie 53, BRI). Drawn by L.Hoare.

21 4.. Maclura MORACEAE inflorescence capitate. Female flowers free or connate in lower half; tepals 4, free or joined in lower half, fleshy at maturity; ovary free or embedded in sockets in fleshy receptacle; style simple or bifid. Fruit an achene (?drupe), thinly pulpy, enclosed in fleshy perianth, often concrescent below into a fleshy syncarp; embryo curved; cotyledons equal. A genus of 12 species in America, Africa, Asia and Australia; 1 species native in Australia. Maclura pomifera (Raf.) Schreider (Osage Orange), cultivated for hedges and ornament in southern Australia may persist in old gardens. A genus of 4 sections. E.Bureau in A.P. de Candolle, Prodr. 17: 226–232 (1873); G.Bentham, Cudrania, Fl. Austral. 6: 178–179 (1873); E.J.H.Corner, Maclura, The Classification of Moraceae, Gard. Bull. Singapore 19: 235–240 (1962); C.C.Berg, The delimitation and subdivision of the genus Maclura (Moraceae), Proc. Kon. Nederl. Akad. Wetensch. 89: 241–247 (1986).

Sect. Cudrania

Maclura sect. Cudrania (Trecul) Corner, Gard. Bull. Singapore 19: 237 (1962). Cudrania Trecul, Ann. Sci. Nat. ser. 3, 8: 122 (1847). T: C. javenensis Trecul = M. cochinchinensis (Lour.) Corner Climber or scrambling shrub. Male and female inflorescences bracteate and capitate. Male flowers with filaments straight in bud; style short. Syncarp fleshy. A section with 2 series.

Ser. Connatae

Maclura ser. Connatae Corner, Gard. Bull. Singapore 19: 238 (1962). Type: M. cochinchinensis (Lour.) Corner Female flowers connate at base; ovaries in sockets of fleshy receptacle; style simple, rarely bifid. A series of 3 species.

1. Maclura cochinchinensis (Lour.) Corner, Gard. Bull. Singapore 19: 239 (1962) Vanieria cochinchinensis Lour., Fl. Cochinch. 564 (1790); Cudrania cochinchinensis (Trecul) Kudo & Mosamane, Ann. Rep. Taihoku Bot. Gard. 2: 272 (1932). T: from Indo-China, J. de Loureiro; n.v. Cudrania javanensis Trecul, Ann. Sci. Nat. ser. 3, 8: 123 (1847). T: from , collector unknown; n.v. Maclura cochinchinensis var. bancroftii (Bailey) Corner, loc. cit.; Cudrania javanensis var. bancroftii Bailey, Syn. Queensland Fl. 491 (1883). T: raised by Dr Bancroft, of Brisbane; n.v. Illustrations: A.Trecul, op. cit. t. 3, figs 76–85, as Cudrania javanensis; C.L.Blume, Mus. Bot. 2: t. 31 (1856) as M. javanica Blume; N.C.W.Beadle, Stud. Fl. N.E. New South Wales 4: 502, fig. 217B (1980). Woody climber; twigs lenticellate, glabrous; spines curved, to 1.5 cm long. Leaves elliptic, rarely ovate to obovate, acute to apiculate, cuneate at base, entire; lamina 4–8 cm long, 1.5–2.5 cm wide, glabrous; lateral veins 7–10 pairs; petiole 0.5–2 cm long; stipules to 2 mm long, fulvous. Inflorescences paired, shorter than leaves. Male inflorescences 5–10 mm diam.; peduncle 5–8 mm long; flowers c. 1.5 mm long, obpyramidal; tepals 4, spathulate, concave at apex, pubescent; stamens 4; pistillode subulate. Female inflorescence slightly larger than male; flowers sessile, ±obconical, c. 1.5 mm long and wide; tepals 4, rarely 3, lightly pubescent. Syncarp 1.5–2 cm wide. Achenes obovoid to nearly globular, ±compressed, to 8 mm long and 6 mm wide, partially immersed in receptacle. Cockspur Thorn. Fig. 29A. Occurs from Cape York Peninsula, Qld, S to Milton, N.S.W.; grows mainly in rainforest and near watercourses. Also SE Asia to W Pacific. Regions: CYRK, BURD, DWSN, MCPH, NEPN, HOWE. Map 20.

22 MORACEAE 4. Maclura

Qld: Atherton, M.M.J. van Balgooy 1533 (L, NSW); Ithaca Ck, L.A.S.Johnson 236 (NSW). N.S.W.: Church Ck, A.N.Rodd 512A (NSW); Yellow Rock Ck, R.Schodde 5171 (NSW); Little Forest, near Milton, Sept. 1960, L.A.S.Johnson & E.F.Constable (NSW). Var. bancroftii is a cultivated variant with variegated leaves.

5. ARTOCARPUS

Artocarpus Forster & G.Forster, Char. Gen. Pl. 101, t. 51, 51a (1776), nom. cons.; from the Greek artos (bread) and karpos (fruit). Type: A. communis Forster & G.Forster Monoecious trees with latex. Leaves alternate, distichous or spiral, simple to pinnatifid or pinnate; stipules paired. Inflorescence cauliflorous, unisexual, pedunculate, capitate; head globose, or clavate to cylindrical, without involucral bracts. Flowers numerous; perianth lobes fused to each other and between flowers to form a syncarp of several to many flowers. Male flower: stamen 1; filament straight in bud; pistillode absent. Female flower: perianth tubular, lower part thin-walled, enclosing ovary, upper part thick-walled, often connate; styles exserted through perianth lobes. Fruit a syncarp developed from 1 to many flowers. Seed large; embryo straight or slightly curved. A genus of c. 50 species indigenous in S and SE Asia through Malesia to the Solomon Islands and Australia; 1 species in Australia. Artocarpus communis (Breadfruit), cultivated for its edible fruit, was spread across the Pacific islands by the Polynesians. The genus comprises 2 subgenera. F.M.Jarrett, Studies in Artocarpus and Allied Genera, III. A Revision of Artocarpus subgenus Artocarpus, J. Arnold Arbor. 40: 113–155, 298–368 (1959); F.M.Jarrett, Studies in Artocarpus and Allied Genera, IV. A Revision of Artocarpus subgenus Pseudojaca, J. Arnold Arbor. 41: 73–140 (1960).

Subg. Pseudojaca

Artocarpus subg. Pseudojaca Trecul, Ann. Sci. Nat. ser. 3, 8: 117 (1847). Type: A. lakoocha Roxb. Leaves alternate and distichous, mostly entire; stipules small, non-amplexicaul. Inflorescence without sterile flowers. Male flowers with 2–4-lobed or -partite perianth, ciliate. Female flowers with tubular perianth covering ovary. Syncarp globose to slightly lobed, fleshy. A subgenus represented by 1 species in Australia.

Artocarpus glaucus Blume, Bijdr. 483 (1825) as glauca T: from Java, Zollinger 704; neo: P, fide F.M.Jarett, J. Arnold Arbor. 41: 118 (1960). Illustrations: G.King, Ann. Roy. Bot. Gard. (Calcutta) 2: t. 8B (1889) as A. denisoniana King; M.O.Rankin, N. Terr. Bot. Bull. 5: t. 6 (1982). Tree to 20 m. Leaves ovate to elliptic, attenuate to acuminate, cuneate to rounded at base, often slightly asymmetric, entire; lamina 10–18 cm long, 5–8 cm wide, glabrous above, appressed puberulous to glabrescent and often grey-glaucous below; lateral veins 10–13 pairs, arching, prominent below; petiole 1–2 cm long, puberulous; stipules to 1 cm long, pubescent. Inflorescence solitary or paired, pedunculate; interfloral bracts peltate. Male head clavate, 5–12 mm long, 3–6 mm diam.; bracts thinly stalked; peduncle 1–3 mm long. Male flowers 0.6 mm long; tepals 2 or 3; stamen 0.8 mm long, filament cylindrical. Female head globose. Syncarp subglobose, often lobed, to 3 cm diam., maturing orange-red; peduncle 3–4 mm long. Fig. 29B–C.

23 5. Artocarpus MORACEAE

Occurs in the north-western N.T.; grows in monsoon forests along water courses; also distributed in and to the Lesser Sunda Islands. Region: ARNM. Map 21. N.T.: Woolner Stn, Whitestone Ck, C.R.Dunlop 5189 (BRI, CBG, DNA, NSW, NT); Wagait Reserve, C.R.Dunlop 3103 (DNA, NSW, NT); Berry Springs Nature Park, M.O.Rankin 2585 (DNA); Channel Point, C.R.Dunlop & R.Petherick 3674 (DNA, NT); Channel Point, B.Hyland 3110 (NSW). First recorded for Australia in 1973.

Trib. 2. CASTILLEAE

Moraceae trib. Castilleae Berg, Acta Bot. Neerl. 26: 78 (1977). Type: Sesse Moraceae trib. Olmedieae Trecul, Ann. Sci. Nat. ser. 3, 8: 126 (1847) p.p. T: Olmedia Ruíz Lopez & Pavón Trees or shrubs with spirally arranged self-pruning branches; latex present. Leaves alternate, distichous; stipules rarely connate. Inflorescences unisexual, discoid to turbinate, in groups on short shoots. Perianth if present usually with imbricate tepals. Stamens straight in bud. Pistillode usually absent. Fruit adnate to perianth. Seeds large, without endosperm. A tribe comprising 8 genera mainly in tropical America and Africa, also SE Asia to the W Pacific; 1 genus in Australia.

6. ANTIARIS

Antiaris Leschen., Ann. Mus. Natl Hist. Nat. 16: 478 (1810), nom. cons.; from the Malay name Antjar. Type: A. toxicaria Leschen. Monoecious trees with latex turning dark on exposure. Leaves alternate, often distichous, simple; stipules free. Male inflorescence many-flowered, with an involucre of bracts. Male flowers: tepals 2–7; stamens 2–4 with straight filaments; pistillode absent. Female inflorescence with 1 or 2 flowers, with involucre of bracts. Female flowers: tepals 4; ovary partly adnate to receptacle; stigmas 2. Fruit a drupe, ellipsoidal to pyriform. Embryo straight. A genus of 4 species in Africa, Indo-Malesia, Australia and SW Pacific; 1 species in Australia. E.J.H.Corner, Antiaris, The Classification of Moraceae, Gard. Bull. Singapore 19: 244–249 (1962); C.C.Berg, Revisions of African Moraceae, Bull. Jard. Bot. État 47: 308–323 (1977).

1. Antiaris toxicaria Leschen., Ann. Mus. Natl Hist. Nat. 16: 478, t. 22 (1810) var. macrophylla (R.Br.) Corner, Gard. Bull. Singapore 19: 248 (1962) A. macrophylla R.Br. in M.Flinders, Voy. Terra Australis 2: 602, t. 5 (1814). T: Company Is., Arnhem Land, [N.T.], Feb. 1803, R.Brown; holo: BM. Illustration: R.Brown, loc. cit. Tree 10–28 m, deciduous. Leaves ovate to oblong, acuminate, unequally cordate to rounded at base, entire; lamina 7–17 cm long, 3–8 cm wide, usually glabrous, lightly scabrid to smooth, coriaceous to membranous; lateral veins c. 10 pairs; intercostal veins prominent; petiole c. 1 cm long. Male inflorescence 10–12.5 mm diam., somewhat fleshy. Male flowers: tepals 4; stamens 4. Female flowers: tepals 4; stigmas 2, divided to base. Infructescence obovoid, to 4 cm long and 2 cm diam. Figs. 2, 29D–E.

24

Figure 29. A, Maclura cochinchinensis, stem and flowering branchlet ×0.5 (D.Blaxell & R.Coveny 3353, NSW). B–C, Artocarpus glaucus. B, ♂flowering branchlet ×0.25 (M.Rankin 2585, DNA); C, ♀flower ×0.5 (N.T., J.Tracey s.n., DNA). D–E, Antiaris toxicaria var. macrophylla. D, ♀flowering branchlet ×0.5 (Cusack 12, DNA); E, ♂flower ×2 (A.Dockrill 561, BRI). Drawn by L.Hoare.

25 6. Antiaris MORACEAE

Occurs in northern N.T. and Cape York Peninsula, Qld; also from eastern Malesia to Melanesia. Regions: ARNM, CYRK. Map 22. N.T.: East Point, Darwin, M.O.Rankin 1825 (BRI, DNA, NSW, NT); Woolner Stn, Whitestone Ck, C.R.Dunlop 5190 (DNA, NSW). Qld: Cape York Peninsula, Claudie R., A.Dockrill 561 (BRI); Pin Pin, A.K.Irvine 690 (BRI). In Cape York Peninsula, Qld, trees can attain a height of c. 28 m. Reported to be deciduous in N.T. early in the dry season. Though the latex is known to be highly poisonous if injected into the blood stream (which is used widely in SE Asia by hunting communities), if swallowed its toxic properties are apparently not effective.

Trib. 3. FICEAE

Moraceae Trib. Ficeae Gaudich., Voy. Freyc. 510 ‘1826’ (1830). Type: Ficus L. Monoecious or dioecious trees, shrubs, stranglers or climbers, with latex. Leaves alternate or opposite, often distichous; stipules free and paired or connate and amplexicaul at base. Flowers small, unisexual, borne inside urceolate receptacles (figs), of 3 or 4 types: male, female, sterile male (neuter) and sterile female inhabited by wasp (gall flower). Styles not exserted from orifice of receptacle; orifice enclosed by an involucre of bracts. A tribe comprising the single genus Ficus occurring throughout the tropics and warm temperate regions. The flowers are pollinated by symbiotic fig-wasps (Hymenoptera, Chalcidoidea).

7. FICUS

Ficus L., Sp. Pl. 1059 (1753); Gen. Pl. 482 (1754); an ancient Latin name for the Fig-tree. Type: F. carica L. Mastosuke Raf., Sylva Tellur. 59 (1838). T: M. rubiginosa (Desf. ex Vent.) Raf. Urostigma Gasp., Nov. Gen. Fic. 7 (1844). T: not designated. Covellia Gasp., op. cit. 10. T: C. ulmifolia (Lam.) Gasp. Sycomorus Gasp., Ric. Caprifico 85 (1845). T: Ficus sycomorus L. Pharmacosycea Miq., Lond. J. Bot. 6: 525 (1847). T: not designated. Trees or shrubs, sometimes epiphytic stranglers or -climbers. Leaves simple, entire or palmately lobed, usually petiolate; glands often present on underside of leaves, in axils of veins or apex of petiole. Inflorescence (fig) urceolate, axillary, cauliflorous or on underground stolons. Tepals 2–8, free or connate, entirely gamophyllous or absent. Male flowers: stamens 1–8; anthers mucronate, introrse; pistillode usually absent. Female flowers: ovary 1, unilocular; style subterminal to gynobasic; stigma bifid or simple; ovule 1, anatropous. Gall flower with a bloated sterile ovary containing a fig-wasp; style short; stigma funnel-shaped. Fruit a drupelet, the woody endosperm forming a pyrene. A large genus of c. 1,000 species throughout tropical and warm temperate regions, with greatest diversity in SE Asia, Malesia and tropical South America; 42 species in Australia. Ficus carica L. produces the edible Fig of commerce, and F. elastica Roxb. ex Hornem. was the source of the Indian-rubber before the introduction of Para-rubber (Hevea brasiliensis (A.L.Juss.) Muell. Arg.). The genus has a large number of indoor ornamental plants and garden and roadside trees such as F. elastica Roxb. ex Hornem., F. religiosa L., F. benjamina, F. macrophylla, F. microcarpa and F. rubiginosa. Ficus pumila is grown widely as a climber on walls. The fig is the single essential morphological attribute of the genus, without which Ficus would have long since been split into numerous genera. This characteristic, coupled with the unique fig-wasp symbiotic relationship, is so strongly

26 MORACEAE 7. Ficus entrenched in the concept of the genus that early attempts by Miquel (1861) and others to break it up into smaller more manageable taxa were unsuccessful. Yet the variation in floral and other attributes within Ficus far exceeds those in many angiosperm families. To express this diversity without sacrificing the convenience of having the figs in 1 genus, a large number of infrageneric taxa were established by Miquel and others which E.J.H.Corner (1960–1965) has refined for Asia and Australasia into a group comprising 4 subgenera, 14 sections, 14 subsections, 54 series and 38 subseries accounting for some 450 species. They have, however, overlooked a number of infrageneric names which G.Don had previously and quite legitimately established in Loudon’s Hortus Britannicus in 1830 and which ought to have been adopted for priority of publication. Don’s taxa are unfortunately inadequately described; since they are not typified, their identity is not clear. They are therefore not followed in this work to avoid premature changes to the existing nomenclature. In the treatment below, subsections and subseries are omitted for convenience. Readers needing the detailed classifications are referred especially to E.J.H.Corner (1960–1965) F.A.G.Miquel, Note sur les Figuiers de la Nouvelle-Hollande, J. Bot. Neerl. 1: 230–243 (1861); O.Warburg, Australische Ficus-arten, Repert. Spec. Nov. Regni Veg. 1: 72–78 (1905); E.J.H.Corner, Taxonomic Notes on Ficus Linn., Asia and Australasia. Sections 1–4, Gard. Bull. Singapore 17: 368–485 (1960); —Sections 5 & 6, Gard. Bull. Singapore 18: 1–69 (1960); —Addendum, Gard. Bull. Singapore 18: 83–97 (1961); —Addendum II, Gard. Bull. Singapore 19: 385–401 (1962); E.J.H.Corner, Check-List of Ficus in Asia and Australasia with Keys to Identification, Gard. Bull. Singapore 21: 1–186 (1965); E.J.H.Corner in M.D.Dassanayake & F.R.Fosberg, Rev. Handb. Fl. Ceylon 3: 230–279 (1981).

KEY TO INFRAGENERIC TAXA

1 Dioecious; interfloral bracts absent subg. 4. FICUS 2 Root climbers; leaves often distichous, with appressed bathyphylls; fig-body with a collar of 3 basal bracts, without lateral bracts; seeds oblong, usually compressed, keeled sect. 5. RHIZOCLADUS 3 Leaves distichous or alternate; figs ripen purple to black; filaments free or slightly joined ser. 8. PLAGIOSTIGMATICAE 3: Leaves mostly distichous; figs ripen red; filaments joined ser. 9. RAMENTACEAE 2: Not this combination of characters 4 Perianth gamophyllous, entire or absent; stamen 1, rarely 2; gall-stigma widely funnel-shaped; seeds not compressed, rarely keeled sect. 9. SYCOCARPUS ser. 16. TUBERCULIFASCICULATAE 4: Perianth of separate tepals, or if joined, then lobate; gall-stigma narrowly funnel-shaped to subclavate; seeds often compressed and keeled 5 Stamens 2 or more; figs pedunculate, mostly cauliflorous, with collar of 3 basal bracts; lateral bracts absent sect. 8. NEOMORPHE ser. 14. VARIEGATAE 5: Stamen 1; if 2, then fig without collar of basal bracts but with lateral bracts 6 Seeds compressed, often with double keel at base, smooth sect. 7. ADENOSPERMA ser. 13. AMPHIGENAE 6: Seeds lenticular to oblong, with a single keel, rarely with double keel sect. 6. SYCIDIUM 7 Leaves long-petiolate, usually symmetrical, scabrid; figs pedicellate with dispersed lateral bracts ser. 10. COPIOSAE 7: Not this combination of characters

27 7. Ficus MORACEAE

8 Leaves asymmetrical, scabrid; figs often pedunculate with a collar of basal bracts ser. 11. SCABRAE 8: Leaves often strongly asymmetrical, lightly scabrid; figs usually not pedunculate, or at most shortly so; often without lateral bracts ser. 12. PALLIDAE 1: Monoecious; interfloral bracts often present 9 Epiphytic stranglers; leaves entire, usually coriaceous, with glands at back of petiole apex; figs with male, female and gall-flowers; stamen usually 1; seeds smoothsubg. subg. 1. UROSTIGMA 10 Fig with bilabiate or trilabiate ostiole with inflexed apical and internal bracts sect. 3. MALVANTHERA ser. 6. MALVANTHERAE 10: Fig ostiole closed by overlapping apical and internal bracts 11 Cystoliths on lower surface of lamina; petiole usually articulated; male flowers usually ostiolar; ovary wholly red-brown or at least in distal half sect. 1. UROSTIGMA 12 Fig with abundant internal chaffy-vescular bristles; basal bracts persistent ser. 2. CAULOBOTRYAE 12: Fig without internal bristles, or few and minute; basal bracts caducous ser. 1. SUPERBAE 11: Cystoliths on both surfaces or upper only of lamina; petiole not articulated; male flowers dispersed; ovary white with a red mark at base, at least on stylar side or lower half red sect. 2. CONOSYCEA 13 Figs sessile or pedicellate, ripen yellow to red ser. 3. DRUPACEAE 13: Not this combination of characters 14 Leaves somewhat distichous, subcoriaceous; basal veins not elongated; lateral veins distinctly raised on both sides in dried material ser. 4. BENJAMINEAE 14: Leaves alternate or distichous, often coriaceous; basal veins often elongated; lateral veins rarely distinctly raised on both sides ser. 5. CALLOPHYLLEAE 9: Trees, rarely shrubs not epiphytic; leaf-glands in axils of the basal veins or absent; stamens 2 (3), or 1, often with a pistillode 15 Male flowers dispersed in fig or ostiolar, often pedicellate; interfloral bracts often present; stigma generally bifid subg. 2. PHARMACOSYCEA sect. 4. OREOSYCEA ser. 7. VASCULOSAE 15: Male flowers mainly ostiolar, sessile, compressed; interfloral bracts generally absent; stigma simple subg. 3. SYCOMORUS

KEY TO SPECIES

1 Root climbers 2 Leaves foveolate beneath, villous; figs pyriform, at least 3 cm long, 2 cm diam., sparsely villous 21. F. pumila 2: Leaves not foveolate beneath, glabrous; figs globular, less than 1.8 cm diam., glabrous 22. F. pantoniana 1: Trees or stranglers, not climbers 3 Leaves 3–5-lobed, palmately nerved † F. carica 3: Leaves entire, very rarely lobed, pinnately nerved 4 Leaves scabrid (sand-paper) or hispid 5 Leaves only lightly scabrid or hispid

28 MORACEAE 7. Ficus

6 Leaf lamina lanceolate, attenuate, often 5 or more times longer than broad, apex long-attenuate, the base rounded; fig lightly scabrid; ostiole raised in a crown of apical bracts 30. F. coronulata 6: Not this combination of characters 7 Leaf lamina usually 6–12 cm long; lateral veins usually 7–8 pairs 8 Leaves somewhat distichous; lamina 8–12 cm long, elliptic to narrowly ovate, acute to long-acuminate, the base cuneate 25. F. leptoclada 8: Leaves alternate; lamina 6–15 cm long, oblong or ovate-elliptic, often strongly asymmetrical, acute, the base subcordate, rounded to cuneate on one side, often decurrent 32. F .tinctoria 7: Leaf lamina usually 16–35 cm long; lateral veins 10 pairs, rarely fewer 9 Buttressed tree; leaves oblong, the base unequally rounded to cordate; stipules small, not amplexicaule; figs globular to obovoid, mostly 1 cm diam. 24. F. melinocarpa 9: Spreading tree often branching at base; leaves crenulate to denticulate 10 Leaves alternate, broadly elliptic to rhombic, base symmetrically cordate; stipules amplexicaul; figs pyriform to discoid, 2.5 cm long, 2.5 cm diam., sometimes geocarpic 38. F. congesta 10: Leaves opposite-decussate, sometimes alternate, pentagonal to oblong, base asymmetrically subcordate; figs depressed globose, 1–2 cm diam., cauliflorous in clusters to 1 m long, often trailing 39. F. hispida 5: Leaves strongly scabrid (sand-paper) 11 Leaves subsessile, often whorled, strongly scabrid; lamina linear to narrowly oblong, to 5 cm long, 0.5 cm wide, the apex obtuse, rounded; lateral veins short 28. F. podocarpifolia 11: Not this combination of characters 12 Leaves mostly opposite-decussate, rarely alternate; leaf margin sinuous; lateral veins 8–13 pairs 13 Leaves ovate, oblong to somewhat orbicular, often sinuous with small rigid hairs; basal pair of lateral veins often more distinct 26. F. opposita 13: Leaves obovate, rarely pentagonal, often faintly sinuous; basal pair of lateral veins not more distinct 27. F. scobina 12: Leaves mostly alternate, rarely opposite; leaf margin entire to dentate or denticulate; lateral veins 6–8 pairs 14 Plant strongly scabrid but otherwise glabrous; leaves generally entire, not asymmetrical at base 29. F. fraseri 14: Plant strongly scabrid; leaves usually densely brown-villous beneath, dentate or denticulate, usually asymmetrical at base 31. F. coronata 4: Leaves glabrous to variously hairy, not scabrid 15 Leaves glabrous 16 Deciduous banyan or strangler to 30 m; leaves with prominent zig-zag intercostals; basal bracts of figs ovate, persistent, often reflexed; figs globular; ostiole enclosed by a flat disc of 3 apical bracts 2. F. virens 16: Not this combination of characters 17 Stipules 4–10 cm long; ostiole often umbonate to long-attenuate

29 7. Ficus MORACEAE

18 Stipules to 10 cm long; leaves broadly obovate, rarely broadly elliptic or ovate, 7–13 cm wide; figs cylindrical, to 6 cm long,

2 cm diam., often 3-angular, pubescent, mottled red-brown; ostiole horn-like, to 2 cm long 18. F. crassipes 18: Stipules 4–7 cm long; leaves generally elliptic, less than 7 cm wide; figs subglobose, oblong or ovoid, to 4 cm long and 3 cm diam.; ostiole umbonate to beaked, less than 1 cm long 19 Lateral veins of leaves 5–10 pairs; figs subglobose to oblong, 1.5–2 cm long, 1.2–1.6 cm diam.; ostiole umbonate, triradiate 7. F. triradiata 19: Lateral veins of leaves 10–16 pairs; figs ovoid to oblong, 2–4 cm long, 1.5–3 cm diam.; ostiole umbonate to beaked, to 8 mm long 8. F. watkinsiana 17: Stipules less than 4 cm long; ostiole at most very shortly beaked, not umbonate or attenuate 20 Tree with plank buttresses to 5 m high; leaf lamina ovate to very broadly ovate, 15–25 cm long, 8–16 cm wide; lateral veins 6–8 pairs, widely spaced, arching, the basal pair reaching at least 1/3 length of lamina; petiole 5–12 cm long 21 Figs without internal interfloral bristles; tepals entire 36. F. variegata 21: Fig wall with inward processes bearing the flowers; tepals dentate-laciniate 37. F. nodosa 20: Not this combination of characters 22 Stipules 1–1.5 cm long 23 Figs paired, sessile 24 Leaf lamina 10–20 cm long, rounded to subcordate at base; lateral veins distantly spaced; figs oblong, to 2.5 cm long 3. F. drupacea 24: Leaf lamina 3–12 cm long, cuneate to somewhat rounded at base; lateral veins closely set; figs globose, ellipsoidal, ovoid to obovoid, to 1.2 cm long 25 Branchlets pendulous (willow-like); leaves somewhat distichous, rounded to cuneate at base; basal veins not elongated; figs ellipsoidal, ovoid or obovoid, ripening dark red 5. F. benjamina 25: Branchlets erect; leaves alternate, asymmetrically cuneate at base; basal veins elongated; figs globose, ripen purple-black 6. F. microcarpa 23: Figs pedunculate 26 Deciduous; petiole to 7 cm long; figs globular, to 2.5 cm diam., yellow-orange mottled white 1. F. superba 26: Evergreen; petiole less than 4 cm long; figs globular, usually less than 1.5 cm diam. 27 Lamina 6–10 cm long; petiole flattened dorsiventrally, to 4 cm long; figs glabrous, at most pustular; ostiole triradiate 16. F. platypoda 27: Lamina 10–20 cm long; petiole not flattened, to 2 cm long; figs scabridulous; ostiole beaked with few protruding apical bracts 33. F. virgata 22: Stipules 2–3 cm long

30 MORACEAE 7. Ficus

28 Leaves pendulous; outer bark of petiole generally not flaky; figs obovoid to depressed globose, often with 6–8 longitudinal ribs 40. F. septica 28: Leaves not pendulous; outer bark of petiole generally brown flaky; figs globose, not ribbed 35. F. adenosperma 15: Leaves variously hairy, not scabrid 29 Leaves ferruginous-pubescent, at least on lower side when young 30 Lower side of leaves ferruginous-pubescent; lateral veins 13–16 pairs; basal bracts 2 31 Leaf lamina broadly elliptic to oblong, 10–25 cm long, 7–12 cm wide; stipules to 15 cm long; figs 2–2.5 cm diam.; peduncle to 2.5 cm long, 0.6 cm diam. 9. F. macrophylla 31: Leaf lamina elliptic, 12–15 cm long, 6.5–7.5 cm wide; stipules to 5 cm long; figs to 2 cm long, 1.5 cm diam.; peduncle 1–1.5 cm long, 0.5 cm diam. 10. F. baileyana 30: Both sides of leaves ferruginous-pubescent, at least when young; lateral veins 10–13 pairs; basal bracts 3 32 Stipules to 2 cm long; figs densely ferruginous-pubescent, reddish brown; ostiole usually bilabiate, raised in a crown 3 mm diam. 11. F. destruens 32: Stipules 4–7 cm long; figs ferruginous-pubescent, soon glabrous, prominently warted; ostiole 3- to 4-radiate, 3 mm diam., slightly raised 12. F. rubiginosa 29: Leaves variously hairy, not ferruginous-pubescent 33 Stipules to 20 cm long, densely pubescent in lower half; figs long-ovoid, to 6 cm long, 2.5 cm diam.; ostiole raised in a crown 5 mm long, 3 mm wide; peduncle to 1.5 cm long, dilated at distal end to form a cupule 17. F. pleurocarpa 33: Stipules much less than 20 cm long; other characters not as above 34 Stipules persistent, often leafy 20. F. racemosa 34: Stipules not persistent, not leafy 35 Leaves broadly ovate, obtuse, cordate at base; lamina very coriaceous, 10–30 cm long, 7–20 cm wide; lateral veins 5–7 pairs, the basal pair prominent, reaching 1/3 length of lamina; stipules thick, 1–1.5 cm long and wide 4. F. benghalensis 35: Not this combination of characters 36 Plant lightly to densely muriculate; leaves alternate to whorled, obovate to obpentagonal, long-acute to acuminate; lamina 25–35 cm long 23. F. copiosa 36: Plant variously hairy, not muriculate; leaves alternate, ovate to elliptic; lamina less than 20 cm long 37 Plant puberulous, soon glabrous; lateral veins indistinct; figs globular, glabrous, 1.5 cm diam. 38 Leaves elliptic, obtuse to acute, cuneate at base; lateral veins 10–12 pairs; petiole 1–2 cm long; stipules to 6 cm long; figs with 2 basal bracts 14. F. obliqua 38: Leaves ovate, obtuse, cuneate to rounded; lateral veins 8–10 pairs; petiole 3–8 cm long; stipules to 2.5 cm long; figs with 3 basal bracts 15. F. subpuberula 37: Plant pubescent to villous or velvety; lateral veins distinct to prominent; figs pubescent to villous

31 7. Ficus MORACEAE

39 Lateral veins 9–11 pairs; petiole woolly, soon glabrous; figs often cauliflorous, obovoid to globular, villous, c. 1 cm diam.; stipules to 1.5 cm long 34. F. mollior 39: Lateral veins 10–13 pairs; petiole pubescent; figs axillary, globular, c. 2 cm diam., pubescent; stipules to 2.5 cm long 40 Leaf lamina 8–13 cm long, acute to obtuse, rarely acuminate; petiole 1–2 cm long; stipules to 5 cm long; figs with 2 basal bracts 13. F. leucotricha 40: Leaf lamina 15–18 cm long, long-acuminate; petiole 3–7 cm long; stipules to 2 cm long; figs with 3 basal bracts 19. F. albipila † Ficus carica L., a native of the Mediterranean and Middle East, is widely cultivated in Australia for its edible fruit. Although reported as an escape in S.A., e.g. H.Eichler, Suppl. Black’s Fl. S. Australia 94 (1965), it does not appear to be naturalised.

Subg. 1. Urostigma

Ficus subg. Urostigma (Gasp.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 285 (1867). Urostigma Gasp., Nov. Gen. Fic. 7 (1844). T: Ficus religiosa L.; lecto, fide E.J.H.Corner, Gard. Bull. Singapore 12: 370 (1960). Monoecious epiphytic stranglers. Leaves entire, usually coriaceous, with abaxial glands at petiole apex. Figs with male, female and gall flowers, with interfloral bracts, without external lateral bracts; fig-wall with an outer and nner layer of sclerotic cells; male flowers with 1 stamen; female flowers with simple style. Seeds smooth. Of the 5 sections in the subgenus, 3 occur in Australia.

Sect. 1. Urostigma

Ficus sect. Urostigma (Gasp.) Endl., Gen. Pl. 4: 35 (1848). Lamina with cystoliths only on lower surface; petiole usually articulated to lamina. Figs with overlapping apical bracts. Male flower usually ostiolar. Female flower with ovary red-brown throughout or at least in distal half. A section of 4 series; 2 in Australia.

Ser. 1. Superbae

Ficus ser. Superbae Corner, Gard. Bull. Singapore 17: 371 (1960). Type: F. superba Miq. Leaves without hypodermis. Figs pedunculate, rarely sessile; internal bristles few and minute or absent; basal bracts caducous.

1. Ficus superba (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 2: 200 (1867) var. henneana (Miq.) Corner, Gard. Bull. Singapore 17: 376 (1960) F. henneana Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 216 (1867). T: Booby Is., Qld, Henne; iso: NSW. F. gracilipes Bailey, Queensland Bot. Bull. 3: 16 (1891). T: Brookfield, Qld, A.Exley; holo: BRI. F. parkinsonii Hiern, J. Bot. 39: 1, t. 417 (1901). T: Booby Is., [Qld], 23 Aug. 1770, J.Banks; n.v.

32 MORACEAE 7. Ficus

F. pritzelii Warb., Repert. Spec. Nov. Regni Veg. 1: 74 (1905). T: Barron, Qld, 22 May 1902, L.Diels 8371; n.v. Illustrations: W.P.Hiern, loc. cit., as F. parkinsonii; J.H.Maiden, For. Fl. New South Wales 2: t. 56 & photo opposite 209 (1907). Large shrub or strangler to 35 m, deciduous, glabrous. Leaves elliptic to oblong, rarely ovate, shortly acuminate, the base rounded, rarely cuneate or ±cordate; lamina 7–13 cm long, 3–6 cm wide; lateral veins 9–11 pairs; petiole 3–7 cm long; stipules to 1 cm long. Figs smooth, globular, to 2.5 cm diam.; yellow to orange, mottled white; ostiole 3–4 mm diam., rarely sunken, bracts forming a raised crown; basal bracts caducous, leaving a collar-scar; peduncle 2–3.5 cm long. Male flowers usually ostiolar, edicellate; tepals 3. Female flowers subsessile, tepals 4 or 5; stigma simple, elongated, sublateral. Gall-stigma bifid. An endemic variety occurring in eastern Arnhem Land, N.T., and from Cape York Peninsula, Qld, S to Nowra, N.S.W.; grows in monsoon forest and rainforest in a variety of alluvial soils. Regions: ARNM, CYRK, BURD, DWSN, MCPH, NEPN. Map 23. N.T.: Little Lagoon, Groote Eylandt, , R.L.Specht 446 (CANB, NSW). Qld: Hibbett Mt, Gundiah, S.F.Kajewski 1507 (BRI, NSW); Black Mt Chain, c. 26 km SSW of Cooktown, L.S.Smith 10681 (BRI). N.S.W.: Rivertree, Upper Clarence, R.H.Cambage 2874 (NSW); c. 16 km NNE of Ravensworth, R.Story 7149 (NSW).

Ser. 2. Caulobotryae

Ficus ser. Caulobotryae (Miq.) Corner, Gard. Bull. Singapore 17: 371 (1960). Urostigma ser. Caulobotrya Miq., Fl. Ned. Ind., Eerste Bijr. 1(2): 334 (1859). T: F. caulocarpa Miq. Lamina without hypodermis. Figs pedunculate or sessile; internal bristles white, chaffy-vesicular, abundant; basal bracts persistent.

2. Ficus virens Aiton, Hort. Kew. 3: 451 (1789) T: cultivated at Kew, from West Indies; holo: BM. Urostigma nesophilum Miq., J. Bot. Neerl. 1: 237 (1862); Ficus nesophila (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 286 (1867); F. glabella var. nesophila (Miq.) Schumann in K.Schumann & C.A.G.Lauterbach, Fl. Schutzgeb. Südsee 273 (1901). T: Quail Island, Flood; holo: ?K. [F. infectoria auct. non Willd.: W.Roxburgh, Fl. Ind. 3: 551 (1832); F.Mueller, Second Syst. Census Austral. Pl. 38 (1889); K.Domin, Biblioth. Bot. 89: 9 (1921)] [Ficus lacor auct. non Buch.-Ham.: R.L.Specht, Rec. Amer.-Austral. Sci. Exped. Arnhem Land 3: 317 (1958)] Deciduous banyan with few aerial or strangler to over 30 m. Leaves ovate to elliptic, sometimes sinuous, bluntly acuminate with tip less than 1 cm long; base subcordate, truncate to cuneate; lamina 8–19 cm long, 3.5–9 cm wide, glabrous; lateral veins 7–12 pairs with 1–4 zig-zag intercostals; petiole 2.5–7 cm long; stipules to 1.5 cm long, finely puberulous. Figs paired, globose, 1–2 cm diam., ripening purple to black; ostiole 1–2 mm diam., enclosed by 3 flat apical bracts in a disc; basal bracts 3, ovate, 1.5–3 mm long, 2–4 mm wide. Male flowers ostiolar, sessile; tepals 2 or 3. Female and gall lowers sessile; tepals free, 3 or 4; ovary sessile or stalked. Widely distributed in S Asia, Malesia through Melanesia to . The species comprises 4 varieties of which 3 are native to Australia. 1 Figs sessile, glabrous 2b. var. sublanceolata 1: Figs pedunculate, glabrous or pubescent 2 Figs glabrous 2a. var. virens 2: Figs white-pubescent 2c. var. dasycarpa

33

Figure 30. Ficus. A–B, F. virens var. virens. A, branchlet ×0.5; B, fruit ×1 (A–B, K.Kenneally 6172, PERTH). C, F. drupacea var. drupacea, flowering branchlet ×0.5 (J.Clarkson 3853, BRI). D, F. watkinsiana, fruiting branchlet ×0.5 (C.White 12787, BRI). Drawn by L.Hoare.

34 MORACEAE 7. Ficus

2a. Ficus virens Aiton var. virens Figs pedunculate, glabrous. Figs 7, 30A–B. Occurs in the Kimberley, W.A., northern N.T. and north-eastern Qld.; grows in monsoon forest and rainforest; also in S Asia through Malesia and Melanesia. Regions: KIMB, ARNM, BKLY, CARP, CYRK. Map 24. W.A.: North Heywood Is., Bonaparte Archipelago, P.G.Wilson 10915 (NSW, PERTH). N.T.: Gardens Road, Darwin, J.Must 916 (DNA, NSW, NT); Oenpelli, Arnhem Land, R.L.Specht 1145 (NSW). Qld: Barron R., Jan. 1927, J.L.Tandent (BRI); Lake Barrine, Atherton Tableland, S.F.Kajewski 1132 (BRI). 2b. Ficus virens var. sublanceolata (Miq.) Corner, Gard. Bull. Singapore 17: 377 (1960) F. saxophila var. sublanceolata Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 260 (1867). T: from Sumatra, De Vriese; n.v. Urostigma cunninghamii Miq., London J. Bot. 6: 560 (1847); Ficus cunninghamii (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 286 (1867); F. infectoria var. cunninghamii (Miq.) Domin, Biblioth. Bot. 89: 8 (1921). T: Brisbane R., [Qld], A.Cunningham; holo: K. Urostigma fraseri Miq., London J. Bot. 6: 561 (1847); Ficus caulobotrya var. fraseri (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 287 (1867); F. fraseri (Miq.) F.Muell., Fragm. 6: 195 (1868); F. infectoria var. fraseri (Miq.) Domin, Biblioth. Bot. 89: 8 (1921). T: Bremer R., [Qld], 1829, C.Fraser 704; holo: K. Urostigma psychotriifolium Miq., London J. Bot. 6: 561 (1847); Ficus psychotriifolia (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 286 (1867); F. infectoria var. psychotriifolia (Miq.) Domin, Biblioth. Bot. 89: 8 (1921). T: Brisbane R., [Qld], 1829, C.Fraser 73; holo: K. Illustration: W.D.Francis, Austral. Rainforest Trees 3rd edn, 75, fig. 27 (1970). Figs sessile, glabrous. White Fig. Fig. 6. Occurs from the Kimberley, W.A., through northern N.T., and from Cape York Peninsula, Qld, S to the Macleay R., N.S.W.; also widely distributed in S and SE Asia. Grows in monsoon forest and rainforest. Regions: KIMB, ARNM, CYRK, BURD, DWSN, MCPH. Map 25. W.A.: near Cape Leveque, W.V.Fitzgerald 1698 (PERTH). N.T.: Daly River, C.R.Dunlop 3593 (DNA, NT). Qld: Lake Barrine, Atherton Tableland, S.F.Kajewski 1120 (NSW). N.S.W.: Ballina, Apr. 1896, W.Bäuerlen (NSW); Clarence Gorge, A.G.Floyd 1710 (NSW). 2c. Ficus virens var. dasycarpa Corner, Blumea 22: 299 (1975) T: Dale Gorge, Hamersley Ra., W.A, Sept.–Oct. 1964, J.Thompson s.n.; holo: PERTH. Figs pedunculate, densely white villous. An endemic variety. Occurs disjunctly from the Hamersley Ra., W.A., through northern N.T. to Cape York Peninsula, Qld. Grows usually in rocky sites. Regions: PILB, KIMB, ARNM, CYRK. Map 26. W.A.: Wittenoom Gorge, K.Hill 441 et al. (NSW, PERTH); Dale Gorge, A.S.George 1053 (PERTH). N.T.: Arnhem Land, D.Thomson 112 (BRI); Katherine, Cutta Cutta Caves, M.O.Parker 1144 (DNA, NSW, NT). Qld: Kowanyama-Topsy Ck, P.Black 671 (BRI).

Sect. 2. Conosycea

Ficus sect. Conosycea (Miq.) Corner, Gard. Bull. Singapore 17: 371 (1960). Urostigma sect. Conosycea Miq., Fl. Ned. Ind., Eerste Bijv. 1(2): 349 (1859). T: F. annulata Blume Leaf lamina usually with an upper hypodermis; cystoliths on both surfaces or upper surface only; petiole not articulated to lamina. Figs with overlapping apical bracts. Male flowers dispersed. Female flowers: ovary white with a red mark at base, at least n the stylar side, or lower half red.

35 7. Ficus MORACEAE

The section has 8 series, 3 in Australia.

Ser. 3. Drupaceae

Ficus ser. Drupaceae Corner, Gard. Bull. Singapore 17: 372 (1960). Type: F. drupacea Thunb. Figs sessile or pedicellate, maturing yellow to red.

3. Ficus drupacea Thunb., Diss. Ficus 6, 11 (1786) Urostigma drupaceum (Thunb.) Miq., London J. Bot. 6: 581 (1847). T: from southern Asia, C.Thunberg; n.v. Ficus pilosa Reinw. ex Blume, Bijdr. 446 (1825); Urostigma pilosum (Reinw. ex Blume) Miq., Fl. Ned. Ind., Eerste Bijv. 1(2): 351 (1854). T: Mt Salak, Java, C.G.C.Reinwardt herb.; holo: L n.v., fide C.L.Blume, op. cit. 446. Spreading strangler to 20 m, glabrous except branchlet apex. Leaves elliptic-oblong to pentagonal, bluntly acuminate or with tip to 7 mm long, the base rounded, slightly cordate; lamina 10–20 cm long, 4–8 cm wide; lateral veins 8–10 pairs, distant, prominent; petiole 1.5–3 cm long; stipules to 1.5 cm long, often brown-hairy, soon glabrous. Figs paired, sessile, glabrous, oblong, 1–2.5 cm long, 1.5 cm wide, maturing orange to red; ostiole slightly umbonate; basal bracts small, concealed under fig. Male flowers pedicellate; tepals 3; stamen 1; bracts numerous; internal bristles absent. emale flowers usually sessile; stigma simple, dilated at base. Widespread in S and SE Asia, Malesia, Melanesia to Australia. The species comprises 5 varieties of which 2 occur in Australia. Basal bracts brown-hairy 3a. var drupacea Basal bracts glabrous to white-pubescent 3b. var glabrata 3a. Ficus drupacea Thunb. var. drupacea Basal bracts of fig usually brown-hairy. Fig. 30C. Occurs on Cape York Peninsula and islands of Torres Strait, Qld; also widely distributed from Burma and the Indo-Chinese Peninsula through Malesia and Melanesia. Region: CYRK. Map 27. Qld: Somerset, Cape York Peninsula, June 1897, F.M.Bailey (BRI); Saibai Is., E.W.Bick 43 (BRI); Boigu Is., J.R.Clarkson 3853 (BRI); Meringa, A.P.Dodd (BRI). 3b. Ficus drupacea var. glabrata Corner, Gard. Bull. Singapore 17: 380 (1960) T: Timor Laut [Tanimbar Is.], Indonesia, F.O.Forbes 3371; holo: BM n.v., fide E.J.H.Corner, op. cit. 381. Basal bracts of fig glabrous, or white-pubescent. Occurs on Cape York Peninsula, Qld; also widely from eastern Malesia to the Solomon Islands. Region: CYRK. Map 28. Qld: Galloways Ck, Bamaga, tip of Cape York Peninsula, L.J.Webb & J.G.Tracey 7116A (BRI).

4. *Ficus benghalensis L., Sp. Pl. 1059 (1753) Urostigma benghalense (L.) Gasp., Ric. Caprifico 82, t. 7, fig. 14–21 (1845). T: from India; n.v. Illustrations: G.King, Ann. Roy. Bot. Gard. (Calcutta) 1: t. 13 (1888); E.J.H.Corner in M.D.Dassanayake & F.R.Fosberg, Rev. Hand. Fl. Ceylon 3: 252 fig. 13 (1981). Wide-spreading banyan with copious aerial roots. Leaves broadly ovate, obtuse, the base cordate; lamina 10–30 cm long, 7–20 cm wide, very coriaceous, puberulous beneath; lateral

36 MORACEAE 7. Ficus veins 5–7 pairs, the basal pair prominent, reaching 1/3 of lamina length; petiole 1.5–7 cm long, 5 mm wide, puberulous; stipules thick, 1–1.5 cm long and wide, puberulous. Figs paired, sessile, puberulous, depressed-globular, 1.5–2 cm diam., maturing orange to red; ostiole broadly umbonate, enclosed by 3 flat apical bracts; basal bracts 3, foliaceous, obtuse, 3–7 mm long, 10–15 mm wide, puberulous. Male flowers edicellate; tepals 2 or 3. Female flowers sessile; tepals 3 or 4. Gall flowers pedicellate; tepals 3 or 4. Banyan Tree. Native to India and Pakistan; reported to be naturalised in north-eastern and central Qld in mixed eucalypt woodland with monsoon scrub species. Regions: BURD, DWSN. Map 29. Qld: 1 km W of Rita Marda Point, G.N.Batianoff & T.J.McDonald 790 (BRI). Easily recognised by its leathery heart-shaped leaves with blunt to rounded apex and softly puberulous under-surface, and the large sessile orange to red figs. Has great religious significance for the Hindus in India, and is widely cultivated in Asia.

Ser. 4. Benjamineae

Ficus ser. Benjamineae Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 287 (1867). Type: F. benjamina L. Leaves somewhat distichous, subcoriaceous; basal veins not elongate; lateral veins distinctly raised on both sides in dried material. A series of 3 species from SE Asia to Australia, 1 of which occurs in Australia.

5. Ficus benjamina L., Mant. Pl. 129 (1767) T: from South Asia; n.v. var. benjamina Ficus nitida Thunb., Diss. Ficus 10 (1786); Urostigma nitidum (Thunb.) Miq., London J. Bot. 6: 582 (1847), non Gasp. (1844). T: not designated. Illustrations: G.King, Ann. Roy. Bot. Gard. (Calcutta) 1: t. 52 (1888); E.J.H.Corner in M.D.Dassanayake & F.R.Fosberg, Rev. Handb. Fl. Ceylon, 3: 256 fig. 15 (1981). Large banyan, glabrous; adventitious roots few to none. Leaves subdistichous, elliptic to ovate, rarely oblong, acute to acuminate, cuneate to rounded at base; lamina 3–12 cm long, 1.5–6 cm wide; lateral veins 6–13 pairs; petiole 4–16 mm long, 1–1.5 mm wide; stipules to 1.2 cm long. Figs usually sessile, ellipsoidal, ovoid, obovoid, or rarely subglobose, 8–12 mm diam., ripening dark red; pedicel if present thick; ostiole enclosed by 3 flat apical bracts in a disc 2 mm diam., often with a rim; basal bracts 2 or 3, to 1.5 mm long, concealed beneath fig-body. Male flowers abundant, pedicellate; tepals 3. Female flowers sessile; tepals free, 3 or 4. Gall lowers pedicellate; tepals 3 or 4. Occurs in northern N.T. and north-eastern Qld; also widespread from S and SE Asia to SW Melanesia. Regions: ARNM, CYRK. Map 30. N.T.: Yirrkala, Arnhem Land, R.L.Specht 853 (NSW); Elcho Is., C.R.Dunlop 3932 (DNA, NT). Qld: Bellenden Ker, F.M.Bailey (BRI). Three varieties are recognised; only var. benjamina occurs in Australia. Widely cultivated as an indoor ornamental.

37 7. Ficus MORACEAE

Ser. 5. Callophylleae

Ficus ser. Callophylleae Corner, Gard. Bull. Singapore 17: 374 (1960). Type: F. callophylla Blume Leaves alternate or subdistichous, often coriaceous; basal veins often elongated; lateral veins rarely distinctly raised on both sides.

6. Ficus microcarpa L.f., Suppl. Pl. 442 (1781) Urostigma microcarpum (L.f.) Miq., London J. Bot. 6: 583 (1847). T: from Java, C.P.Thunberg; holo: UPS n.v. Ficus cairnsii Warb., Repert. Spec. Nov. Regni Veg. 1: 73 (1905). T: Cairns, Qld, 30 May 1902, L.Diels 8425; n.v. [F. nitida auct. non L.: F.Mueller, Fragm. 6: 195 (1868)] [F. retusa auct. non L.: G.Bentham, Fl. Austral. 6: 166 (1873)] Illustration: E.J.H.Corner in M.D.Dassanayake & F.R.Fosberg, Rev. Handb. Fl. Ceylon 3: 258, fig. 16 (1981). Large banyan with copious aerial roots. Leaves usually alternate, often asymmetric at base; lamina 3–12 cm long, 1.5–9 cm wide, usually glabrous; lateral veins 5–9 pairs, usually at an acute angle to midrib, rarely prominent; petiole 3–20 mm long, often puberulous; stipules to 1.5 cm long, the margin puberulous. Figs paired, sessile, usually somewhat globose, 9–12 mm diam., ripening purple to black; ostiole enclosed by 2 or 3 flat apical bracts forming a disc 2–2.5 mm diam.; basal bracts 1.5–3 mm long and wide, puberulous. Flowers with 3 tepals; male and female sessile; gall flowers pedicellate. Occurs widely in S and SE Asia, Malesia, Melanesia, the SW Pacific and north-eastern Australia. Often a large buttressed tree with wide spreading branches from which aerial roots descend to the ground and eventually form secondary stems. The species comprises 7 varieties of which 3 occur in Australia. With such a wide distribution, the species has a very wide range of morphological variation, especially in leaf form. The varieties are not easy to distinguish and detailed field investigation is needed to determine their status. 1 Leaf lamina broadly elliptic to broadly obovate, to 12 cm long and 9 cm wide, obtuse to shortly and bluntly acuminate, rounded to widely cuneate at base; venation somewhat prominent beneath 6c. var. latifolia 1: Leaf lamina narrowly elliptic to narrowly obovate, to 11 cm long and 6 cm wide, often bluntly acuminate to somewhat acute, cuneate to narrowly cuneate at base; venation not prominent beneath 2 Basal veins usually elongated and prominent; petiole not articulated; fig with internal bristles few to many 6a. var. microcarpa 2: Basal veins not so elongated or prominent; petiole somewhat articulated; fig with internal bristles usually absent 6b. var. hillii 6a. Ficus microcarpa L.f. var. microcarpa Leaves narrowly elliptic to narrowly obovate, often bluntly acuminate to somewhat acute, cuneate to narrowly cuneate at base; lamina to 11 cm long and 6 cm wide; venation not prominent beneath; basal veins usually elongated; petiole not articulated. Figs with few to many internal bristles. Fig. 4. Occurs from Cape York S to near Rockhampton, Qld, including coastal islands; also throughout the range of the species. Regions: CYRK, BURD, DWSN. Map 31. Qld: Red Island Point, Cape York Peninsula, L.J.Brass 18926 (BRI); Middle Percy Is., H.Tryon (BRI); Palm Islands, T.L.Bancroft 91 (BRI); Mt Cook, L.S.Smith 10605 (BRI).

38 MORACEAE 7. Ficus

6b. Ficus microcarpa var. hillii (Bailey) Corner, Gard. Bull. Singapore 17: 398 (1960) F. hillii Bailey, Queensland Bot. Bull. 3: 16 (1891). T: tropical Qld, W.Hill; holo: BRI. Leaves narrowly elliptic to narrowly obovate, often bluntly acuminate to somewhat acute, cuneate at base; lamina to 11 cm long and 6 cm wide; venation not prominent beneath; basal veins not prominent beneath; petiole somewhat articulated. Figs usually without internal bristles. Occurs in north-eastern Qld S to Yeppoon; also from the Sunda Islands, Indonesia, through New Guinea to and the Loyalty Islands. Grows in strand forest. Regions: CYRK, BURD, DWSN. Map 32. Qld: Clump Point, L.S.Smith 4817 (BRI); between Clump Point and Bingil Bay, L.S.Smith 4861 (BRI). Commonly cultivated in Australia as roadside tree, especially in warmer regions near the coast. 6c. Ficus microcarpa var. latifolia (Miq.) Corner, Gard. Bull. Singapore 17: 398 (1960) Urostigma accedens var. latifolium Miq., Fl. Ned. Ind., Eerste Bijv. 1(2): 347 (1859). T: Halawahoea Is., Moluccas, herb. C.G.C.Reinwardt; holo: L n.v., fide E.J.H.Corner, loc. cit. F. dictyophleba F.Muell. ex Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 218 (1867). T: islands off Cape Flattery, Qld, herb. F.Mueller; n.v. F. thyneana Bailey, Queensland Agric. J. 1: 231 (1897); F. thyneana var. typica Domin, Biblioth. Bot. 89: 9 (1921). T: Cairns, Qld, E.Cowley; n.v. F. thyneana var. minor Domin, Biblioth. Bot. 89: 9 (1921). T: Yarraba, Qld, Jan. 1910, K.Domin; n.v. Leaves broadly elliptic to broadly obovate, obtuse to shortly bluntly acuminate, rounded to widely cuneate at base; lamina to 12 cm long and 9 cm wide; venation ±prominent beneath; basal veins ±prominent; petiole not articulated. Figs with internal bristles. Occurs in north-eastern Qld S to near Cairns; also in the Celebes, Moluccas, New Guinea, and the . Grows in strand and riverine forest. Region: CYRK Map 33. Qld: Trinity Bay, F.M.Bailey (BRI); Mowbray R., L.J.Brass 2016 (BRI); Cairns, Aug. 1901, E.Betche (NSW); Coen, July 1949, H.Flecker (NSW).

Sect. 3. Malvanthera

Ficus sect. Malvanthera Corner, Gard. Bull. Singapore 17: 374 (1960). Type: F. macrophylla Desf. ex Pers. Fig with bilabiate or triradiate ostiole; apical and internal bracts inflexed. Male flowers dispersed; anthers unilocular with 2 pollen sacs. Ovary partly immersed in the fig-wall with a red mark at base or with red apex. A section of 2 series occurring from Malesia to Australia.

Ser. 6. Malvantherae

Ficus ser. Malvantherae Corner, Gard. Bull. Singapore 17: 375 (1960). Type: F. macrophylla Desf. ex Pers. Anthers reniform, dehiscing longitudinally and crescentically. A series of 12 species, 11 endemic in Australia, the other widespread in eastern Malesia, Australia and the SW Pacific.

39 7. Ficus MORACEAE

7. Ficus triradiata Corner, Gard. Bull. Singapore 17: 401 (1960) T: Mt Spurgeon, Qld, C.T.White 10536; holo: BRI. Large strangler, glabrous. Leaves elliptic to narrowly obovate, subacute, the base cuneate; lamina 6–14 cm long, 2–6 cm wide; primary lateral veins 5–10 pairs, not prominent; secondary lateral veins faint, the basal pair slightly elongated; petiole 1.5–3 cm long; stipules to 7 cm long and 0.7 cm wide. Figs paired, subglobose to oblong, 1.5–2 cm long, 1.2–1.6 cm wide; ostiole mammillate, triradiate, to 4 mm diam.; basal bracts 3, ovate, 5–7 mm long, 7–10 mm wide, persistent. Male flowers pedicellate; tepals 4, free. Female and gall flowers essile to shortly pedicellate; tepals 4 or 5; ovary free. Endemic in north-eastern Qld. Two varieties have been named but these may represent different developmental stages. Leaf lamina 7–14 cm long, 3.5–6 cm wide; figs pedunculate 7a. var. triradiata Leaf lamina 6–12 cm long, 2–4 cm wide; figs subsessile to sessile 7b. var. sessilicarpa 7a. Ficus triradiata Corner var. triradiata Leaf lamina elliptic to narrowly obovate, 7–14 cm long, 3.5–6 cm wide; primary lateral veins 6–10 pairs. Figs pedunculate, usually oblong. Apparently endemic in the Mt Lewis area, SW of Mossman, Qld; grows in montane rainforest. Region: CYRK. Map 34. Qld: Brooklyn–Mt Lewis track, H.Flecker, Qld Naturalists’ Club 7260 (BRI); Mt Lewis, B.Hyland 1883 (BRI). A rainforest strangler with copious white latex. 7b. Ficus triradiata var. sessilicarpa Corner, Gard. Bull. Singapore 17: 401 (1960) T: Mt Finnegan, Qld, L.J.Brass 20052; holo: BRI. Leaf lamina narrowly elliptic, 6–12 cm long, 2–4 cm wide; primary lateral veins 5–9 pairs; figs subsessile to sessile, usually subglobose. Endemic in the mountains from near Cooktown S to near Mossman, Qld; grows in montane rainforest in soils derived from granodiorite. Region: CYRK. Map 35. Qld: Mt Lewis, V.K.Moriarty 835 (BRI).

8. Ficus watkinsiana Bailey, Queensland Bot. Bull. 2: 18 (1891) T: Mooloolah, Qld, Dec. 1890, Field Naturalists; syn: n.v.; top of Bunya Mtns, Qld, H.Tryon; syn: n.v.; top of Blackall Ra., Qld, A.Anderson; syn: n.v. F. bellingeri C.Moore in C.Moore & E.Betche, Handb. Fl. New South Wales 81 (1893). T: Bellinger R., N.S.W., ?E.Betche; n.v. F. simmondsii Bailey, Queensland Agric. J. 25: 234, t. 19 (1910). T: Tweed Heads, N.S.W., Oct. 1910, J.H.Simmonds; holo: BRI. Illustrations: V.S.Summerhayes, Hooker’s Icon. Pl. 32: t. 3187 (1933); W.D.Francis, Austral. Rainforest Trees 3rd edn, fig. 33 (1970). Large strangler to 60 m; trunk to 2 m diam. Leaves usually elliptic, shortly acuminate, the base cuneate; lamina 8–20 cm long, 2–7 cm wide, glabrous; primary lateral veins 10–16 pairs, not prominent; petiole 4–7 cm long; stipules 4–7 cm long, glabrous. Figs paired, lenticellate, ovoid to oblong, 2–4 cm long, 1.5–3 cm wide; ostiole strongly mammillate, often to 8 mm long; basal bracts leaving a prominent collar to 1.5 cm diam. at base; peduncle 1–2 cm long, 0.6 cm diam. Male flowers long-pedicellate; tepals 4. Female and gall flowers sessile; tepals 4, narrowly ovate; style subterminal; stigma obtuse. Watkin’s Fig. Fig. 30D.

40 MORACEAE 7. Ficus

Occurs mainly from Gympie, Qld, S to near Dungog, N.S.W., with a disjunct population between Cairns and Townsville; grows usually in clay-loam derived from basalt. Regions: CYRK, MCPH. Map 36. Qld: Springbrook, McPherson Ra., C.E.Hubbard 5413 (BRI); Springbrook, W.J.F.McDonald & L.W.Jessup 2079 (BRI); Bunya Mtns, C.T.White 9150 (BRI). N.S.W.: Whian Whian State Forest, C.T.White 12787 (BRI); Lismore, Jan. 1950, J.L.Green (NSW). The large hollow ‘trunks’ formed by the aerial roots after the supporting tree has decayed are often called Cathedral Trees in south-eastern Qld.

9. Desf. ex Pers., Syn. Pl. 2: 609 (1807) Urostigma macrophyllum (Desf. ex Pers.) Miq., London J. Bot. 6: 560 (1847). T: cult. Hort. Bot. Paris; n.v. subsp. macrophylla Ficus huegelii Kunth & Bouché, Ind. Sem. Hort. Berol. 15 (1846); Urostigma huegelii (Kunth & Bouché) Miq., London J. Bot. 6: 586 (1847). T: n.v. Urostigma squamellosum Miq., J. Bot. Neerl. 1: 239 (1862); Ficus squamellosa (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 287 (1867). T: Clarence R., N.S.W., F.Mueller; n.v. Urostigma platypodum f. majus Miq., J. Bot. Neerl. 1: 236 (1862). T: Brisbane R., [Qld], C.Fraser; syn: n.v.; Hastings R., N.S.W., C.Fraser; syn: n.v. F. platypoda var. petiolaris Benth., Fl. Austral. 6: 169 (1873) p.p. as to C.Fraser, Brisbane & Hastings Rivers, see E.J.H.Corner, Gard. Bull. Singapore 17: 402 (1960). F. macrophylla f. stenophylla Domin, Biblioth. Bot. 89: 13 (1921). T: Tamborine Mtn, Qld, K.Domin; n.v. Illustrations: W.D.Francis, Austral. Rainforest Trees 3rd edn, figs 31, 32 (1970); B.D.Morley & H.R.Toelken, Fl. Pl. Australia fig. 30 (1983). Tree to 55 m with wide-spreading crown; trunk to 2 m diam., strongly buttressed. Leaves broadly elliptic to oblong, often broadly ovate, acute to bluntly acuminate, the base broadly cuneate to subrotund; lamina 10–25 cm long, 7–12 cm wide, ferruginous-pubescent beneath, often becoming glabrous; primary lateral veins 13–16 pairs, slightly more prominent than the secondary ones; petiole 5–10 cm long; stipules to 15 cm long, lightly pubescent. Figs paired, globose to oblong, 2–2.5 cm diam.; ostiole minute, hardly raised; basal bracts 2, very broadly ovate; peduncle 2–2.5 cm long, 0.6 cm diam. Male flowers pedicellate; tepals 3 or 4. Female flowers shortly pedicellate; tepals usually 3; tyle short; stigma simple. Moreton Bay Fig. Fig. 9. Occurs from north-eastern Qld S to Nowra, N.S.W.; grows in montane or coastal rainforest, often in soils derived from volcanics or alluvium. Regions: CYRK, DWSN, MCPH, NEPN. Map 37. Qld: Roberts Plateau, C.T.White 6054 (BRI); Lamington Natl Park, C.T.White 11891 (BRI); Parish of Haly, H.F.Swain (BRI). N.S.W.: Mt Elaine, Glenugie Peak, June 1957, L.A.S.Johnson & E.F.Constable (NSW); Fig Tree, near Wollongong, July 1951, C.E.Chadwick (NSW). Sometimes confused with F. rubiginosa but readily distinguished by the larger leaves and the figs with much longer peduncles each with 2 (not 3) basal bracts. Widely cultivated as garden trees south as far as Victoria. Fallen leaves have been reported palatable to stock. Wood said to be useful for cases and crates. Ficus macrophylla comprises 2 subspecies: the type subspecies in mainland Australia characterised by the presence of a main trunk, and subsp. columnaris of Lord Howe Is. without such a character, see P.S.Green, J. Arnold Arbor. 67: 111 (1986).

10. Ficus baileyana Domin, Biblioth. Bot. 89: 567 (1921) F. macrophylla var. pubescens Bailey, Queensland Agric. J. 26: 316 (1911). T: southern Qld, F.M.Bailey; n.v. Illustration: F.M.Bailey, loc. cit. t. 33.

41 7. Ficus MORACEAE

Tree to 20 m with spreading crown. Leaves usually elliptic, rarely ovate, acute, rarely bluntly acuminate, the base subrotund; lamina 12–15 cm long, 6.5–7.5 cm wide, ferruginous-pubescent beneath; primary lateral veins 13–16 pairs, slightly more prominent than the secondary ones; petiole 5–7 cm long, pubescent; stipules to 5 cm long, densely ferruginous-pubescent. Figs paired, lenticellate, ferruginous-pubescent, oblong, to 2 cm long and 1.5 cm wide; ostiole minute, very slightly raised; basal bracts sometimes broadly ovate, ferruginous-pubescent; peduncle 1–1.5 cm long, c. 5 mm diam., pubescent. Male and female flowers dispersed, shortly pedicellate; tepals 4. Endemic in central eastern Qld. Region: DWSN. Map 38. Qld: along Calliope R., N of Gladstone, R.F.Thorne 20567a (BRI). A close relative of F. macrophylla from which distinguished by the reddish brown vestiture and the more elongated figs.

11. Ficus destruens F.Muell. ex C.White, Contr. Arnold Arbor. 4: 16 (1933) T: Gadgarra, Atherton Tableland, Qld, 8 June 1929, S.F.Kajewski 1087; holo: BRI. Illustration: C.T.White, loc. cit. t. 3. Strangler to 30 m; twigs densely ferruginous-pubescent. Leaves narrowly elliptic to lanceolate, ±obtuse, the base cuneate; lamina 9–18 cm long, 2–6 cm wide, densely ferruginous-pubescent beneath; primary lateral veins 11–13 pairs, prominent beneath; petiole 3–5 cm long, ferruginous-pubescent; stipules 2 cm long, often densely ferruginous-pubescent. Figs paired, globose, 1–1.7 cm diam., densely ferruginous-pubescent, often reddish brown, glabrescent; ostiole usually bilabiate, raised to a crown 3 mm diam.; basal bracts 3, caducous, leaving a collar at base of fig; peduncle to 5 mm long, densely pubescent. Male flower: tepals 4–6; pedicel 2.5 mm long. Female flower: tepals 4–6; pedicel to 3 mm long. Endemic in north-eastern Qld, disjunctly from the Atherton Tableland S to near to the Eungella Ra., W of Mackay; grows in montane and coastal rainforest. Regions: CYRK, BURD. Map 39. Qld: Windin Ck, State Forest Reserve 310, A.W.Dockrill 80 (BRI, NSW); Topaz, near Malanda, S.T.Blake 15203 (BRI); Fenby Gap, Mission Beach area, R.F.Thorne & J.G.Tracey 22629 (BRI); Cucania, Russell & Mulgrave Rivers junction, R.F.Thorne & J.G.Tracey 23152 (BRI); State Forest Reserve 194, B.Hyland 1975 (BRI). A distinctive species easily recognised by the reddish brown to dark indumentum of the twigs, leaves and figs, the leaf shape and the usually bilabiate ostiole of the fig. In the Eungella Ra. area the species grows as a strangler on eucalypts.

12. Ficus rubiginosa Desf. ex Vent., Jard. Malm. 114, t. 114 (1805) F. australis Willd., Sp. Pl. 4: 1138 (1806), nom. illeg. based on F. rubiginosa Desf. ex Vent.; Mastosuke rubiginosa (Desf. ex Vent.) Raf., Sylva Tellur. 59 (1838); Urostigma rubiginosum (Desf. ex Vent.) Gasp., Nov. Gen. Fic. 7 (1844). T: cult. Hort. Bot. Paris [Ventenat, op. cit. t. 114]. F. rubiginosa var. lucida Maiden, Forest Fl. New South Wales 1: 10 (1902). T: not designated. F. rubiginosa var. variegata Guilfoyle, Austral. Pl. 178 (1911). T: not designated. F. rubiginosa var. glabrescens Bailey, Compr. Cat. Queensland Pl. 486 (1913). T: Eidsvold, Qld, T.L.Bancroft; n.v. Illustrations: R.Ventenat, Jard. Malm. t. 114 (1805); W.J.Hooker, Bot. Mag. 56: t. 2939 (1829); N.C.W.Beadle, Stud. Fl. N.E. New South Wales 4: 499, fig. 215 B (1980). Tree to 50 m, with spreading crown; trunk to 2 m diam.; young twigs ferruginous-pubescent. Leaves alternate, elliptic to ovate, rarely slightly obovate, obtuse to slightly acute, the base cuneate to subrotund; lamina 7–11 cm long, 3.5–7.5 cm wide, ferruginous-pubescent beneath; lateral veins 10–13 pairs, prominent; petiole 2–4 cm long; stipules 4–7 cm long. Figs globular, to 1.5 cm diam., prominently warted; ostiole 3- or 4-radiate, to 3 mm diam., slightly raised; basal bracts 3, to 2 cm long, ferruginous-pubescent; peduncle 0.5–1 cm long,

42 MORACEAE 7. Ficus often dilated at apex, puberulous. Male and female flowers interspersed, pedicellate; pedicels c. 1 mm long; tepals 3. Port Jackson Fig, Rusty Fig. Endemic in eastern Australia, with a very wide ecological range. Regions: BURD, DWSN, MCPH, NEPN, HOWE. Map 40. Qld: Gloucester Park, Sinclair Bay, A.N.Rodd 3083 (NSW); c. 48 km S of Emerald Township, M.Lazarides & R.Story 62 (CANB, NSW). N.S.W.: Durras Mtn, Murramarang Ra., B.G.Briggs 3131 (NSW); Shoalhaven R., Wogamia, F.A.Rodway 12711 (NSW). This species exhibits a broad spectrum of morphological variation; inland forms often come close to Ficus platypoda.

13. Ficus leucotricha (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 285 (1867) Urostigma leucotrichum Miq., J. Bot. Neerl. 1: 234 (1862). T: upper Victoria R., [N.T.], Dec. 1855, F.Mueller; holo: K. Urostigma platypodum f. ellipticum Miq., J. Bot. Neerl. 1: 236 (1862). T: York Sound, W.A., A.Cunningham .s.n.; holo: K. Spreading tree to 10 m, sometimes deciduous; most parts pubescent to velutinous. Leaves alternate, ovate to elliptic, sometimes oblong to pentagonal, acute to obtuse, rarely acuminate, the base broadly cuneate to rounded, often cordate; lamina 8–13 cm long, 4–9 cm wide; lateral veins 10–13 pairs, prominent; petiole 1–2 cm long; stipules 5 cm long. Figs globular, 2 cm diam., pubescent to velutinous; ostiole slightly raised, 2 mm diam., often triradiate; basal bracts 2, 1 cm long and wide, rarely to 1.5 cm; peduncle absent or to 3.5 cm long. Male flowers mainly ostiolar, mixed with female lowers, subsessile; tepals 3. Female flowers subsessile; tepals 4. Endemic in Australia. There are 3 varieties. 1 Figs sessile 13c. var. sessilis 1: Figs pedunculate 2 Fig-body 10–12 mm diam.; basal bracts 4–6 mm long and wide 13a. var. leucotricha 2: Fig-body 14–16 mm diam.; basal bracts 8–12 mm long and wide 13b. var. megacarpa 13a. Ficus leucotricha (Miq.) Miq. var. leucotricha Twigs usually less than 5 mm diam. Fig-body 10–12 mm diam.; basal bracts 4–6 mm long and wide; peduncle 4–35 mm long. Occurs widely in W.A., N.T. and Qld where it thrives in rocky country. Regions: KIMB, ARNM, BKLY, CARP, GGNA. Map 41. W.A.: Grant Ra., West Kimberley, Aug. 1906, W.V.Fitzgerald (NSW). N.T.: Settlement Ck, L.J.Brass 162 (BRI); Overland Telegraph Stn, S.T.Blake 17675 (BRI). Qld: Mt Isa, T.Farrell TF676 (BRI); Mt Isa, A.Schmid AS489 (BRI). 13b. Ficus leucotricha var. megacarpa F.Muell. ex Corner, Gard. Bull. Singapore 17: 402 (1960) T: Sea Range, Qld, F.Mueller; holo: K n.v., fide E.J.H.Corner, op. cit. 402 (1960). Twigs more than 5 mm diam. Fig-body 14–16 mm diam.; basal bracts 8–12 mm long and wide; peduncle 6–14 mm long. Largely confined to northern W.A. and N.T. Frequent on rocky sites especially quartzite hills. Region: KIMB. Map 42. W.A.: Dillon Springs, E Kimberley, W.V.Fitzgerald 1646 (NSW, PERTH); Augustus Is., Bonaparte Archipelago, P.G.Wilson 10696 (PERTH); Kelly Knob Lookout, Kununurra, K.F.Kenneally 1922 (PERTH). N.T.: Jasper Gorge, M.O.Parker 444 (BRI, DNA, NT); c. 108 km NNE of Creswell Stn, R.A.Perry 1661 (NT).

43 7. Ficus MORACEAE

The type of this variety was cited as collected in Queensland, but as no other collections have been made in eastern Australia the type locality may be incorrect. 13c. Ficus leucotricha var. sessilis Corner, Gard. Bull. Singapore 17: 402 (1960) T: locality uncertain, collector unknown, ex J.E.Smith herbarium; holo: MANCH. Twigs less than 5 mm diam. Fig-body 10–12 mm diam., sessile; basal bracts to 10 mm long and wide. Reported to be frequent on sandstone in W.A. Region: KIMB. Map 43. W.A.: Edkin Ra., W.V.Fitzgerald 1423 (PERTH); Prince Regent River Reserve, NW Kimberley, K.F.Kenneally 2086 (PERTH); Fern Gully, Prince Regent River Reserve, NW Kimberley, A.S.George 12621 (PERTH); 0.6 km NE of Elgie Cliffs Stn, M.Lazarides 5087 (NSW). This variety does not occur in N.S.W.; J.E.Smith’s collection has probably been wrongly annotated as to place of origin.

14. Ficus obliqua G.Forster, Fl. Ins. Austral. 77 (1786) Urostigma obliquum (G.Forster) Miq., London J. Bot. 6: 563 (1847). T: Tanna [Tana] Is., , G.Forster; n.v. Urostigma eugenioides Miq., J. Bot. Neerl. 1: 238 (1862); Ficus eugenioides (Miq.) F.Muell. ex Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 286 (1867). T: Albany Is., Qld, ; n.v. Urostigma backhousei Miq., J. Bot. Neerl. 1: 240 (1862); Ficus backhousei (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 288 (1867). T: eastern Australia, J.Backhouse; n.v. F. virginea Banks & Sol. ex Hiern, J. Bot. 39: 2 (1901). T: Booby Is., [Qld], 1770, J.Banks; holo: BM. F. tryoni Bailey, Queensland Agric. J. 17: 103 (1906). T: Middle Percy Is., [Qld] 1770, Dec. 1905, R.Tryon; holo: BRI. Tree 10–15 m, rarely to 30 m, puberulous, soon glabrous. Leaves alternate, elliptic, rarely narrowly ovate, obtuse to acute, the base cuneate; lamina 6–10 cm long, 2–3 cm wide, glabrous; lateral veins 10–12 pairs, indistinct; petiole slightly flattened dorsiventrally, 1–2 cm long; stipules 6 cm long. Figs globular, to 1.5 cm diam., glabrous, often spotted; ostiole triradiate, slightly raised; basal bracts 2, to 4 mm long; peduncle absent or to 1 cm long. Male and female flowers mixed, the male slightly more numerous near ostiole; flowers subsessile to shortly pedicellate; tepals generally 3. Widely distributed from E Malesia through Melanesia and Australia to SW Pacific. Three varieties are recognised; all occur in Australia. 1 Fig 10–15 mm diam.; peduncle 2.5–10 mm long 14b. var. petiolaris 1: Fig less than 10 mm diam.; peduncle absent or to 4 mm long 2 Leaf lamina cuneate at base 14a. var. obliqua 2: Leaf lamina rounded at base 14c. var. puberula 14a. Ficus obliqua G.Forster var. obliqua Illustration: F.M.Bailey, Queensland Agric. J. 17: t. 10 (1906) as F. tryoni. Leaf lamina elliptic, cuneate at base; petiole 1–1.5 cm long. Fig 6–9 mm diam.; peduncle absent or to 4 mm long. Fig. 5. Occurs from Celebes in Indonesia through Melanesia to SW Pacific. In Australia, occurs in W.A., Qld and N.S.W. Regions: KIMB, CYRK, DWSN, MCPH, NEPN. Map 44. W.A.: Prince Regent River Reserve, K.F.Kenneally 2094 (PERTH). Qld: Danbulla, State Forest Reserve 185, J.O’Farrell 47 (BRI, NSW); Atherton Tableland, S.F.Kajewski 1179 & 1314 (BRI, NSW). N.S.W.: Tweed R., Guilfoyle (NSW); Shellharbour, J.L.Boorman (NSW).

44 MORACEAE 7. Ficus

14b. Ficus obliqua var. petiolaris (Benth.) Corner, Gard. Bull. Singapore 17: 402 (1960) F. platypoda var. petiolaris Benth., Fl. Austral. 6: 169 (1873). T: Narran R., L.Leichhardt; syn: n.v.; Rockhampton & Cape R., Qld, Bowman; syn: n.v.; Cape Cleveland, Qld, Burdekin Expedition; syn: n.v.; Port Denison, Qld, Fitzalan; syn: n.v. Leaf lamina elliptic, cuneate to rounded at base; petiole to 4 cm long. Fig 10–15 mm diam.; peduncle 2.5–10 mm long. Endemic in Qld and N.S.W. Regions: CYRK, DWSN, MCPH, NEPN. Map 45. Qld: Tallebudgera Ck, C.T.White 6508 (BRI); Belmont, near Brisbane, C.T.White 6665 (BRI); Bald Mtns, Darling Downs, M.S.Clemens (BRI). 14c. Ficus obliqua var. puberula (Benth.) Corner, Gard. Bull. Singapore 17: 402 (1960) F. eugenioides var. puberula Benth., Fl. Austral. 6: 167 (1873); based on Urostigma brachypodum Miq., London J. Bot. 6: 562 (1847); Ficus brachypoda (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 287 (1867). T: York Sound, [W.A.], A.Cunningham 304; holo: K n.v., fide E.J.H.Corner, loc. cit. Leaf lamina elliptic-lanceolate, cuneate, somewhat abruptly rounded at base; petiole 3–5 mm long. Fig 5–6 mm diam.; peduncle absent or to 4 mm long. Recorded from northern W.A. but no recent collections seen. Region: KIMB. Map 46. No specimens seen.

15. Ficus subpuberula Corner, Gard. Bull. Singapore 17: 403 (1960) Urostigma puberulum Miq., London J. Bot. 6: 562 (1847); Ficus puberula (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 287 (1867), nom. illeg. non Kunth & Bouché (1847). T: York Sound, [W.A.], A.Cunningham s.n.; holo: K n.v., fide F.A.W.Miquel, London J. Bot. 6: 562 (1847). Illustration: F.A.W.Miquel, London J. Bot. 6: t. 23A (1847). Much-branched tree to 6 m; twigs puberulous. Leaves alternate, ovate, obtuse, the base cuneate to subrotund; lamina 8–12 cm long, 3–6 cm wide, glabrous; lateral veins 8–10 pairs, indistinct; petiole 3–8 cm long, soon glabrous; stipules to 2.5 cm long, soon glabrous. Figs globular, to 1.5 cm diam., softly pubescent, soon glabrous; ostiole mammilate, 3 mm diam.; basal bracts 3, caducous; peduncle to 1 cm long, distal end dilated, softly pubescent. Flowers shortly pedicellate to sessile. Male flowers dispersed; tepals 3. Female flowers: tepals 3 or 4. Occurs in northern W.A. and N.T.; grows in crevices of sandstone rocks. Regions: KIMB, ARNM, BKLY. Map 47. W.A.: Kimberlite Pipe area, A.S.Weston 12294 (PERTH); Prince Regent River Reserve, K.F.Kenneally 2025 (NSW, PERTH). N.T.: Nabarlek Airstrip, M.O.Rankin 2187 (DNA, NSW); c. 4 km SW of Mt Gilruth, M.Lazarides 7944 (NSW, NT, PERTH). Closely related to F. platypoda with which it can sometimes be confused. It might well be a variant of F. platypoda with thicker, rigid leaves.

16. Ficus platypoda (Miq.) Cunn. ex Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 287 (1867) Urostigma platypodum Miq., London J. Bot. 6: 561 (1847). T: York Sound, W.A., A.Cunningham; holo: K n.v., fide E.J.H.Corner, Gard. Bull. Singapore 17: 403 (1960). Urostigma vittelinum Miq., J. Bot. Neerl. 1: 237 (1862); Ficus vittelina (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 268, 288 (1867). T: Fitzmaurice River, [N.T.], F.Mueller; n.v. Spreading shrub or tree to 6 m. Leaves alternate, broadly ovate to elliptic, acute, the base cuneate, rounded to cordate; lamina 6–10 cm long, 1–7 cm wide, usually glabrous; lateral veins 10–15 pairs, prominent to indistinct; petiole usually dorsiventrally flattened, 1–4 cm long; stipules small, soon glabrous. Figs globular, 1–1.5 cm diam., glabrous; ostiole often triradiate, slightly umbonate, diam. variable; basal bracts 3, small, soon glabrous; peduncle c.

45 7. Ficus MORACEAE

1 cm long, glabrous to pubescent. Male and female flowers interspersed; tepals 3 or 4. Female lowers sessile. Male flowers shortly pedicellate to subsessile. Rock Fig. Widely distributed in the northern half of Australia, with an extension to the Lesser Sunda Islands in Indonesia. The species is highly variable with considerable intergration between 5 varieties which are consequently hard to distinguish. 1 Leaf lamina broadly ovate, the base cordate to rounded 16c. var. cordata 1: Leaf lamina ovate to elliptic, the base cuneate to rounded 2 Twigs and petioles glabrous or puberulous 3 Leaf lamina narrowly ovate to narrowly elliptic, 1–4 cm wide, rigidly coriaceous, glabrous 16e. var. minor 3: Leaf lamina elliptic, rarely ovate or broadly ovate, 3–7 cm wide, not rigidly coriaceous, glabrous or slightly puberulous 16a. var. platypoda 2: Twigs and petioles pubescent 4 Leaf lamina velvety both sides 16b. var. angustata 4: Leaf lamina glabrous or velvety beneath 16d. var. lachnocaula 16a. Ficus platypoda (Miq.) Cunn. ex Miq. var. platypoda Leaf lamina elliptic to broadly ovate, 3–7 cm wide, the base usually rounded; glabrous, or twigs and stipules lightly puberulous. Occurs from north-western W.A. through northern N.T. and eastern Qld to north-eastern N.S.W. Regions: KIMB, SNDY, PILB, MRCH, ARNM, BKLY, CYRK, BURD, DWSN, MCPH. Map 48. W.A.: Camp Ck, Mitchell Plateau, K.F.Kenneally 5019 (PERTH). N.T.: El Sharana, Pine Creek road, P.Martensz & R.Schodde AE 493 (NT); Mary River, near Black Fella Is., M.O.Rankin 2555 (DNA). Qld: Lamington Natl Park, R.F.Thorne 21711 (BRI). N.S.W.: Stanthorpe road, near Woodenbong, S.L.Everist & L.J.Webb 1399 (BRI). Inland populations in the N.T. tend to have smaller and more coriaceous leaves than those nearer the coast, and can be difficult to distinguish from var. minor. In eastern Australia var. platypoda approaches F. rubiginosa with which it is often confused, differing chiefly in the absence of brown scurfy hairs on the underside of the lamina. 16b. Ficus platypoda var. angustata (Miq.) Corner, Gard. Bull. Singapore 21: 27 (1965) F. leichhardtii var. angustata Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 268 (1867). T: Whitsunday Is., Qld, Henne; iso: NSW. Urostigma leichhardtii Miq., J. Bot. Neerl. 1: 235 (1862); Ficus leichhardtii (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 268, 287 (1867). T: Cape Cleveland, Qld, F.Mueller; n.v. F. platypoda var. subacuminata Benth., Fl. Austral. 6: 170 (1873). T: Whitsunday Island, Qld, Henne; iso: NSW. Urostigma muelleri Miq., J. Bot. Neerl. 1: 235 (1862); Ficus muelleri (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 268, 287 (1867). T: Hastings River, Qld, F.Mueller; n.v. F. platypoda var. mollis Benth., Fl. Austral. 6: 170 (1873). T: Rockingham Bay, Qld, J.Dallachy; iso: NSW. F. shirleyana Domin, Biblioth. Bot. 89: 12 (1921). T: Yungaburra, Qld, Feb. 1910, K.Domin; n.v. Leaf lamina ovate to elliptic, cuneate to rounded at base, coriaceous, velvety both sides; lateral veins distinct; petiole and twigs pubescent. Widely distributed in northern W.A., N.T. and Qld. Regions: KIMB, ARNM, CARP, ?GGNA, CYRK, BURD, DWSN. Map 49. W.A.: South-West Osborne Is., Bonaparte Archipelago, P.G.Wilson 11142 (PERTH). N.T.: Reynolds Crossing, M.O.Parker 848 (DNA, NT); El Sharana, Pine Creek road, P.Martensz & R.Schodde AE517 (NT). Qld: Eungella Ra., C.T.White 12995 (BRI); Atherton Tableland, S.F.Kajewski 1330 (BRI).

46 MORACEAE 7. Ficus

This variety is hardly distinguishable from var. lachnocaula. 16c. Ficus platypoda var. cordata Specht, Records Amer.-Austral. Sci. Exped. Arnhem Land 3: 217 (1958) T: South Bay, Bickerton Is., Arnhem Land, N.T., R.L.Specht 449; holo: BRI; iso: NSW, PERTH. Illustration: R.L.Specht, op. cit. fig. 1. Leaf lamina broadly ovate, cordate to rounded at base, glabrous to sparsely pubescent, often coriaceous; lateral veins distinct. Occurs in far northern W.A. and N.T.; also in Indonesia. Regions: KIMB, ARNM. Map 50. W.A.: Cape Anjo, N coast of W.A., P.G.Wilson 11300 (PERTH). N.T.: 0.6 km E of Top Springs, G.Chippendale 2368 (NT); c. 155 km S of Manangrida, J.R.Maconochie 1579 (NT). 16d. Ficus platypoda var. lachnocaula (Miq.) Benth., Fl. Austral. 6: 169 (1873) as lachnocaulon Urostigma lachnocaulon Miq., J. Bot. Neerl. 1: 238 (1862); Ficus lachnocaula (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 287 (1867). T: , ?Baudin’s Expedition; n.v. Illustration: K.Domin, Biblioth. Bot. 89: 11, fig. 118 (1921). Leaf lamina ovate to elliptic, cuneate to rounded at base, glabrous or velvety beneath, coriaceous; lateral veins distinct. Occurs widely from north-western W.A. to northern N.T., possibly in Qld. Regions: KIMB, MRCH, PILB, SNDY, ARNM, ?BURD. Map 51. W.A.: Charles Knife Rd, Cape Ra., A.S.George 1353 (PERTH); West Lewis Is., Dampier Archipelago, R.D.Royce 7431 (PERTH). N.T.: Beatrice Hill, S.T.Blake 16987 (BRI); Deaf Adder Gorge, R.E.Fox 2524 (DNA, NT). 16e. Ficus platypoda var. minor Benth., Fl. Austral. 6: 169 (1873) T: N.W. coast, W.A., W.Bynoe; syn: n.v.; Nicol Bay, W.A., Gregory’s and Ridley’s Expeditions; syn: n.v. Urostigma platypodum f. glabrior Miq., J. Bot. Neerl. 1: 236 (1862). T: north-western W.A., W.Bynoe; syn: K; Macdonnell Ra., [N.T.], J.M.Stuart; syn: n.v. Leaf lamina narrowly ovate to narrowly elliptic, 1–4 cm wide, cuneate at base, glabrous, stiffly coriaceous; lateral veins indistinct. Fig. 3. Occurs widely in the northern half of W.A., southern N.T., northern S.A. and northern Qld. Regions: KIMB, MRCH, PILB, SNDY, GBSN, TNMI, MACD, SIMP, CARP. Map 52. W.A.: S of upper Ruddall R., A.S.George 10747 (PERTH). N.T.: Ayers Rock, G.Chippendale 683 (NSW, NT). S.A.: Musgrave Park, Cave Hill, S.H.Yengoyan et al. 52 (NT). Qld: Emerald Township, M.Lazarides & R.Story 62 (NSW). The most distinctive variety in the species, although its differentiation from var. platypoda is still not clear-cut. The only variety to occur in central Australia.

17. Ficus pleurocarpa F.Muell., Fragm. 8: 246 (1874) T: Johnstone R., Qld, W.Hill; n.v. F. cylindrica Warb., Repert. Spec. Nov. Regni Veg. 1: 72 (1905). T: Barron R., Qld, 22 May 1902, L.Diels 8382; n.v. Tree to 15 m. Leaves alternate, ovate to oblong, attenuate, the base broadly cuneate to subrotund; lamina 15–26 cm long, 5–9 cm wide, glabrous below except pubescent midrib; primary lateral veins 18–23 pairs; petiole to 8 cm long, usually flattened, densely pubescent, soon glabrous; stipules to 20 cm long, densely pubescent in lower half. Figs narrowly ovoid,

47

Figure 31. Ficus. A–B, F. crassipes. A, branchlet ×0.5; B, fruit ×0.5 (A–B, A.Irvine 999, BRI). C, F. pleurocarpa, flowering branchlet ×0.5 (S.Blake 15184, BRI). Drawn by L.Hoare.

48 MORACEAE 7. Ficus to 6 cm long and 2.5 cm diam., soon glabrous; ostiole raised in a crown 5 mm long, 3 mm wide; basal bracts 3, small, fused to distal end of peduncle; peduncle to 1.5 cm long, distal end often dilated to form a cupule at base of fig. Male and female flowers interspersed, pedicellate; tepals 3–5. Fig. 31C. Occurs from S of Cooktown to the Atherton Tableland area, Qld; grows in montane rainforest. Region: CYRK. Map 53. Qld: Boonjie, near Malanda, S.T.Blake 15184 (BRI); Theresa Ck, Millaa Millaa, W.G.Trapnell 261 (BRI); Big Tableland, c. 43.5 km SE of Cooktown, L.S.Smith 11176 (BRI); State Forest Reserve Moomin, B.Hyland 1751 (BRI); Atherton, J.Bravery 3998 (BRI).

18. Ficus crassipes Bailey, Rep. Pl. Prelim. Gen. Rep. Bot. Meston’s Exped. Bellenden Ker 2 (1889); Bot. Bellenden Ker Exped. 60 (1889) T: Harvey Ck, Russell R., Qld, F.M.Bailey & E.Meston; syn: BRI. Tree to 30 m, often strangling. Leaves alternate, broadly obovate, rarely broadly elliptic or ovate, bluntly acuminate, the base cuneate to subrotund; lamina 10–17 cm long, 7–13 cm wide, glabrous; primary lateral veins 9–12 pairs; petiole 6–10 cm long, often flattened, glabrous; stipules to 10 cm long, lightly pubescent. Figs cylindrical, often 3-angular, 6 cm long, 2 cm diam., pubescent, mottled reddish brown; ostiole horn-like, to 2 cm long; basal bracts 3, 1 cm long and wide; peduncle 1.5 cm long, 3-angled, dilated at apex forming with bracts a cupule at base of fig. Male and female flowers dispersed, pedicellate; tepals 4. Fig. 31A–B. Restricted to the Atherton and Evelyn Tableland areas, Qld, in montane rainforest and also in disturbed habitats. Region: CYRK. Map 54. Qld: Topaz, near Malanda, S.T.Blake 15204 (BRI); Jaggan, S of Malanda, L.S.Smith & L.J.Webb 3340 (BRI); Gadgarra, B.Hyland 1862 (BRI); Gadgarra, S.F.Kajewski 1097 (BRI); Atherton, A.K.Irvine 999 (BRI). Closely related to F. pleurocarpa from which it differs mainly in the shape of the leaves and the form of the fig ostiole.

Subg. 2. Pharmacosycea

Ficus subg. Pharmacosycea (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 299 (1867). Pharmacosycea Miq., London J. Bot. 6: 525 (1847). T: F. maxima Miller = F. radula Willd. Monoecious trees, rarely shrubs, not epiphytic or climbing. Leaves usually entire with a gland in axil of each main basal vein. Figs without lateral bracts on body; interfloral bracts often present. Male flowers ostiolar or dispersed; stamens 1–3; pistillode sometimes present. Gall and female flowers similar, the female usually sessile with longer style; ovary white or with a red spot at base, sessile; style glabrous; stigma bifid. Seeds smooth, often keeled over apex. A subgenus of 2 sections, 1 of which occurs in Australia, the other being confined to the American tropics.

49 7. Ficus MORACEAE

Sect. 4. Oreosycea

Ficus sect. Oreosycea (Miq.) Corner, Gard. Bull. Singapore 17: 406 (1960). Urostigma sect. Oreosycea Miq., London J. Bot. 6: 525 (1847). T: F. nervosa Heyne ex Roth Leaf lamina with cystoliths mostly on the lower side. Figs usually paired; ostiole variable in shape, not crateriform; interfloral bracts and internal bristles present or absent. Male flowers: stamens 1 or 2, rarely 3. Ovary without a red spot. A section with some 45 species from Africa and Madagascar through Asia to Australia and SW Melanesia.

Ser. 7. Vasculosae

Ficus ser. Vasculosae Corner, Gard. Bull. Singapore 17: 406 (1960). Type: F. vasculosa Wallich ex Miq. Medium to large trees. Reticulation of leaf lamina often raised on one or both sides, rarely with basal glands. Fig peduncle well developed. Stamens usually 2 in ostiolar flowers. Occurs in Madagascar through SE Asia to New Guinea and Australia. This series has 1 species in Australia.

19. Ficus albipila (Miq.) King, Ann. Roy. Bot. Gard. (Calcutta) 1: 179 (1888) var. albipila Covellia albipila Miq., Fl. Ned. Ind., Eerste Bijr. Suppl. 434 (1860); Ficus mollis var. albipila (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 296 (1867). T: from Indonesia, collector unknown; n.v. F. colossea F.Muell. ex Benth., Fl. Austral. 6: 163 (1873). T: Herbert R., Rockingham Bay, Qld, J.Dallachy; holo: K. Tree to 35 m; bole to 20 m long, 1 m diam., with buttresses to 5 m high. Leaves alternate, ovate to pentagonal, long-acuminate, the base cordate; lamina 15–18 cm long, 7–10 cm wide, lightly pubescent to glabrous on upper side, densely pubescent on lower side; lateral veins 10–13 pairs, prominent; petiole 3–7 cm long, pubescent; stipules to 2 cm long, densely pubescent. Figs globular, 2 cm diam., pubescent; ostiole 2–3 mm diam., slightly raised; basal bracts 3, forming collar just below fig-body; peduncle to 1 cm long, pubescent. Male and female flowers interspersed, pedicellate, surrounded by abundant bristles; tepals 3 or 4. Abbey Tree. Fig. 32A. Widely distributed in SE Asia; in Australia occurs in Qld. Region: CYRK. Map 55. Qld: Rocky R., B.Hyland 6855 (BRI, NSW); L.J.Webb & J.G.Tracey 9331 (BRI); Herbert Vale, F.M.Bailey (BRI); Cardwell, R.Michael (BRI); Claudie R., L.J.Webb & J.G.Tracey 8578 (BRI). Ficus albipila comprises 2 varieties, of which only the type variety occurs in Australia.

50

Figure 32. Ficus. A, F. albipila var . albipila, fruiting branchlet ×0.5 (B.Hyland 6855, BRI). B–C, F. racemosa var. racemosa. B, branchlet ×0.5; C, fruiting branchlet ×0.5 (B–C, not recorded). D–E, F. copiosa. D, leaf ×0.4; E, fruit ×0.5 (E–F, S.Kajewski 1178, NSW). Drawn by L.Hoare.

51 7. Ficus MORACEAE

Subg. 3. Sycomorus

Ficus subg. Sycomorus (Gasp.) Mildbr. & Burret, Bot. Jahrb. Syst. 46: 174 (1812) Ficus sect. Sycomorus (Gasp.) Miq., Fl. Ind. Bot. 1(2): 319 (1859); Sycomorus Gasp., Giorn. Bot. Ital. 2: 219 (1844); Ric. Caprifico 85 (1845). T: F. sycomorus L. Monoecious, buttressed trees, usually cauliflorous. Leaves often dentate, ±plicate in bud. Male flowers ostiolar, sessile, compressed; stamens usually 2; filaments joined in lower part. Gall and female flowers distinct; ovary dark red; stigma simple. Tepals usually laciniate-denticulate or entire in male flowers. Interfloral bracts absent. Largely African with 1 species in Australia.

20. Ficus racemosa L., Sp. Pl. 1060 (1753) T: from southern Asia, collector unknown; n.v. var. racemosa F. glomerata Roxb., Pl. Corom. 2: 13, fig. 123 (1798); Covellia glomerata (Roxb.) Miq., London J. Bot. 7: 465 (1848). T: Coromandel Coast, India, W.Roxburgh; n.v. F. vesca F.Muell. ex Miq., J. Bot. Neerl. 1: 243 (1862); F. racemosa var. vesca (F.Muell. ex Miq.) Barrett, Bull. Torrey Bot. Club 73: 323 (1946). T: Fitzmaurice R., N.T., F.Mueller; n.v. F. semicostata Bailey, Queensland Agric. J. 26: 316, t. 32 (1911). T: cult. Bot. Gard. Brisbane, Qld, F.M.Bailey; holo: BRI. Illustrations: W.Roxburgh, loc. cit.; G.King, Ann. Roy. Bot. Gard. (Calcutta) 1(2): t. 218, 219 (1888) as Ficus glomerata; E.J.H.Corner in M.D.Dassanayake & F.R.Fosberg, Rev. Handb. Fl. Ceylon 3: 267, fig. 21 (1981). Buttressed tree to 35 m, often deciduous. Leaves alternate, ovate, rarely oblong, obtuse to subacute or subacuminate, sometimes sinuate, the base cuneate to subcordate; lamina 6–20 cm long, 4–9 cm wide, lightly pubescent, soon glabrous; lateral veins 4–8 pairs, with 5–7 prominent intercostals; petiole 4–7 cm long; stipules persistent, 10–13 mm long. Figs cauliflorous in clusters to 25 cm long, globular to pyriform, c. 3 cm diam., often lenticellate-verrucose; ostiole closed by 5 or 6 apical bracts in a disc 2–3 mm diam.; internal bristles absent; basal bracts 3, to 2 mm long; peduncle to 12 mm long. Flowers sessile, with 3 or 4 irregular lobes; male flowers ostiolar, in 1 ring; female and gall flowers interspersed. Fig. 32B–C. Widely distributed from southern Asia to New Guinea; also in northern Australia where it occurs along streams in various soil types. Regions: PILB, KIMB, ARNM, BKLY, CARP, CYRK, BURD. Map 56. W.A.: Elvira R., c. 11 km N of Turner River Stn, R.A.Perry 2408 (BRI, NSW). N.T.: East Alligator R., Oenpelli, R.L.Specht 1184 (BRI, NSW); Adelaide R., M.O.Rankin 1756 (DNA). Qld: Ck, Mossman, S.T.Blake 15015 (BRI); Georgetown, C.T.White 1450 (BRI, NSW). Four varieties are recognised in the species in S and SE Asia; only the type variety occurs in Australia.

Subg. 4. Ficus

Ficus L. subg. Ficus. Ficus subg. Eusyce Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 289 (1867); Ficus subg. Carica Mildbr. & Burret, Bot. Jahrb. Syst. 46: 174 (1912). T: F. carica L. Dioecious plants of various growth habit. Leaves often dentate. Gall plants with male and gall flowers in the figs, seeding plants with female and neuter flowers. Interfloral bracts absent. Lateral bracts often present on body of fig. Seeds often with distinctive form or marking.

52 MORACEAE 7. Ficus

A subgenus of 8 sections, 5 of which occur in Australia.

Sect. 5. Rhizocladus

Ficus sect. Rhizocladus Endl., Gen. Pl. 4(2): 34 (1847). Type: F. pumila L. Root-climbers. Leaves distichous. Figs with collar of 3 basal bracts, without lateral bracts. Male flowers 2-staminate, ostiolar. Female flowers usually with ifid stigma. Seeds oblong, generally compressed and keeled. A section of 5 series, 2 of which occur in Australia.

Ser. 8. Plagiostigmaticae

Ficus ser. Plagiostigmaticae Corner, Gard. Bull. Singapore 18: 3 (1960). Type: F. pumila L. Male and neuter flowers ostiolar, in 1 to several rows, the males mostly pedicellate. Anthers mucronate; filaments free or slightly joined. Figs ripening purple to black. Represented in Australia by 1 introduced species.

21. *Ficus pumila L., Sp. Pl. 1060 (1753) T: from eastern Asia, collector unknown; n.v. Ficus stipulata Thunb., Diss. Bot. Fic. 7 (1786). T: not designated. Illustration: G.King, Ann. Roy. Bot. Gard. (Calcutta) 1(2): t. 158 (1888). Root-climber, generally villous. Leaves distichous, broadly ovate, elliptic to oblong, obtuse to subacute, the base cordate; lamina 4–10 cm long, 2.5–6 cm wide, pitted beneath; lower juvenile leaves usually appressed on climbing twig; lateral veins 3–6 pairs; petiole 2–4 cm long; stipules to 1.5 cm long, appressed-villous. Figs solitary, pyriform, 3–6 cm long, 2.5– 3.5 cm diam., sparsely villous, ripening purple to black, truncate and umbonate at apex; internal bristles abundant; basal bracts 5–7 mm long. Male flowers at distal half of fig, in many rows, 10–17 mm long; stamens 2, rarely 3. Female flowers with sessile ovary; style simple, lateral. Widespread in Asia; naturalised in Qld and N.S.W. Regions: NEPN, MCPH. Map 57. Qld: Brisbane, G.N.Batianoff 6 (BRI); Brisbane, J.Gillieatt (BRI). N.S.W.: Dubbo, July 1913, L.V.Miller (NSW); Cheltenham, Dec. 1948, L.A.S.Johnson (NSW). Grown widely as a garden-wall climber in Qld and N.S.W.

Ser. 9. Ramentaceae

Ficus ser. Ramentaceae Corner, Gard. Bull. Singapore 18: 4 (1960). Type: F. villosa Blume Similar to ser. Plagiostigmaticae but differs in filaments being joined and figs ripening red. A series of 53 species in Asia, 1 of which occurs in Australia.

53 7. Ficus MORACEAE

22. Ficus pantoniana King, J. Asiat. Soc. Bengal 55(2): 407 (1887) T: New Guinea, collector unknown; n.v. var. pantoniana F. scandens var. australis Bailey, Queensland Agric. J. 1: 370 (1897); F. nugentii Domin, Biblioth. Bot. 89: 13 (1921). T: Kamerunga, Qld, L.J.Nugent; n.v. Illustration: F.M.Bailey, Compr. Cat. Queensland Pl. 496, fig. 488 (1913). Root-climber, glabrous. Leaves alternate, generally ovate, obtuse to shortly acuminate, the base rounded; lamina 8–13 cm long, 4–7 cm wide, not pitted beneath; lateral veins 5–8 pairs, prominent; petiole 1.5–3 cm long; stipules narrow, 1 cm long. Figs pedunculate, globular, 1–1.5 cm diam.; ostiole somewhat sunken; basal bracts 2 mm long, forming a small collar. Male flowers mainly ostiolar; anthers to 2 mm long. Female flowers with ovary to 1.5 mm long; style simple; pedicel c. 1.5 mm ong. Dispersed in Moluccas (Indonesia) through New Guinea and Bismarck Archipelago to Qld, Australia. Common root-climber in swampy palm rainforest in granitic alluvial soils. Regions: CYRK, BURD. Map 58. Qld: Innisfail, C.T.White 11753 (BRI); Daintree R., L.J.Brass 2351 (BRI); Fishery Falls, between Gordonvale and Babinda, L.J.Webb & J.G.Tracey 7458 (BRI); Atherton, Porn 62, Alexandra, B.Hyland 7038 (BRI, NSW). Three varieties are recognised; only the type variety occurs in Australia. Distinguished from F. pumila by the non-pitted leaves and the globular, considerably smaller figs which ripen red.

Sect. 6. Sycidium

Ficus sect. Sycidium Miq., London J. Bot. 7: 228 (1848). Type: F. aspera G.Forster Trees, shrubs or stranglers. Leaves alternate, rarely distichous. Figs with or without collar of basal bracts, often with lateral bracts. Flowers often pedicellate. Male flowers ostiolar; stamens 1 or 2. Female flowers with simple, subclavate, subterminal stigmas; ovary white. Seeds lenticular to oblong, with a single keel. A section of c. 90 species from Africa through Asia to the South Pacific, 11 species in Australia.

Ser. 10. Copiosae

Ficus ser. Copiosae Corner, Gard. Bull. Singapore 17: 445 (1960). Type: F. copiosa Steudel Trees, often scabrid. Leaves mostly long-petiolate, symmetrical. Figs pedicellate, with dispersed small to minute lateral bracts. A series of 11 species from India to the Solomon Islands; epresented in Australia by the type species.

23. Ficus copiosa Steudel, Nom. Bot. 2nd edn, 1: 635 (1840) F. polycarpa Roxb., Fl. Ind. 1832 edn, 3: 556 (1832), nom. illeg. non Jacq. T: cult. Bot. Gard. Calcutta, India, collector unknown; n.v. F. magnifolia F.Muell., Fragm. 4: 50 (1863). T: Mt Elliot, Qld, E.Fitzalan & J.Dallachy; n.v. F. subinflata Warb., Repert. Spec. Nov. Regni Veg. 1: 76 (1905). T: Barron R., Qld, L.Diels 8379; n.v. F. copiosa var. pubescens Corner, Gard. Bull. Singapore 17: 455 (1960). T: New Guinea, A.J.Kostermans 310; holo: L n.v., fide E.J.H.Corner, loc. cit.

54 MORACEAE 7. Ficus

Tree to 7 m, lightly to densely muriculate. Leaves alternate to whorled, obovate to obpentagonal, long-acute to acuminate, the base cordate; lamina 25–35 cm long, 8–15 cm wide; lateral veins 8–10 pairs, basal pair straighter than others, prominent; petiole 4–7 cm long; stipules narrowly elliptic, 1–2 cm long. Figs axillary to cauliflorous, slightly depressed-globular, c. 2.5 cm diam., often sparsely covered with minute lateral bracts, often densely muricate; ostiole slightly raised; basal bracts occasionally present forming a small collar on peduncle. Male lowers ostiolar; stamens 1 or 2. Female flowers long-pedicellate; style short. Fig. 32D–E. Widespread in the Celebes, Moluccas (Indonesia) through New Guinea, Bismarck Archipelago, Solomon Islands to the Pacific. In Australia, occurs on the coast and tablelands of central and northern Qld. Regions: CYRK, BURD, DWSN. Map 59. Qld: Tolga Reserve, near Atherton, R.F.Thorne 20870 (BRI); Lake Eacham, Atherton Tableland, S.F.Kajewski 1178 (BRI, NSW); c. 1 km W of Cedar Bay, Bloomfield R. area, L.J.Webb & J.G.Tracey 8968 (BRI); slopes of Mt Quincan, Peeramon, S.T.Blake 15248 (BRI, NSW); Timber Camp Rd between Daintree & Bloomfield R., L.J.Webb & J.G.Tracey 12059 (BRI). On Australian material, it is impracticable to distinguish var. pubescens due to continuity of the diagnostic characters.

Ser. 11. Scabrae

Ficus ser. Scabrae Miq., London J. Bot. 7: 228 (1848). Type: F. aspera G.Forster Leaves shortly petiolate, often asymmetrical and scabrid. Figs often with a collar of basal bracts.

24. Ficus melinocarpa Blume, Bijdr. 460 (1825) var. hololampra (Diels) Corner, Gard. Bull. Singapore 17: 462 (1960) F. hololampra Diels, Bot. Jahrb. Syst. 67: 201 (1935). T: New Guinea, collector unknown; n.v. Buttressed tree; trunk to 40 cm diam. Leaves alternate, oblong, acute to shortly acuminate, the base unequally rounded to cordate; lamina 16–20 cm long, 7–9 cm wide, lightly scabrid; lateral veins c. 10 pairs; petiole c. 2 cm long; stipules small. Figs globular to obovoid, c. 1 cm diam., lightly scabrid; ostiole slightly raised, apical bracts slightly protruding; basal bracts persistent. Male flowers shortly pedicellate. Female flowers subsessile. Widely distributed from the Philippines to New Guinea; recently recorded from Qld, Australia. Region: CYRK. Map 60. Qld: Rocky R., B.Hyland 2837 (BRI, NSW).

25. Ficus leptoclada Benth., Fl. Austral. 6: 172 (1873) T: Rockingham Bay, Qld, J.Dallachy s.n.; iso: NSW. Tree to 15 m. Leaves distichous, elliptic to narrowly ovate, acute to long-acuminate, the base cuneate; lamina 8–12 cm long, 3–4 cm wide, glabrous, rarely sparsely scabrid; lateral veins 7–10 pairs; petiole 1 cm long; stipules less than 1 cm long. Figs often cauliflorous, globular, to 1.2 cm diam., glabrous; ostiolar bracts forming a small beak 3 mm diam.; basal bracts small, alternate. Male flowers ostiolar; stamens 1 or 2. Female flowers shortly pedicellate; tepals 4 or 5. Fig. 33A. Endemic in northern Qld, growing in rainforest to 1000 m altitude. Regions: CYRK, BURD, DWSN. Map 61. Qld: Atherton Tableland, State Forest Reserve 185, Robson Logging Area, A.K.Irvine 1250 (BRI, NSW); Darymple Heights, Armstrong Ck, Nov. 1947, M.S.Clemens (BRI); Herberton Ra., Forest Reserve 99,

55 7. Ficus MORACEAE

R.Schodde 4181 (BRI); Daintree R., S.F.Kajewski 1455 (BRI); Tinaroo Ra., road from Downfall Ck, L.J.Webb & J.G.Tracey 5752 (BRI).

26. Ficus opposita Miq., London J. Bot. 7: 426 (1848) T: Bremer R., [Qld], 1829, C.Fraser 101; n.v. F. fitzalanii Miq., J. Bot. Neerl. 1: 242 (1862). T: Cape Cleveland, Qld, E.Fitzalan; n.v. F. xerophila Domin, Biblioth. Bot. 89: 14 (1921). T: Smelling Bluff, near Chillagoe, Qld, Feb. 1910, K.Domin; n.v. F. yarrabensis Domin, op. cit. 15. T: Yarraba, Qld, Jan. 1910, K.Domin; n.v. Tree to 8 m. Leaves opposite, often decussate, sometimes alternate, ovate, oblong to ±orbicular, often sinuate with small rigid hairs, acute to obtuse, the base rounded to cordate; lamina 4–14 cm long, 3–6 cm wide, variously hairy, often strongly scabrid; lateral veins 8–10 pairs, the basal pair often more distinct; petiole 5–30 mm long, usually pubescent to villous, rarely glabrescent; stipules to 1 cm long. Figs globular, 1–1.5 cm diam., glabrescent-scabrid to densely villous; ostiole slightly umbonate, with protruding apical bracts; lateral bracts often present; basal bracts variously arranged. Flowers with 5–7 tepals. Male flowers in a single row; stamens 1 or 2. Sandpaper Fig. Widely distributed from eastern Malesia to Australia. The species is highly variable and the 3 varieties recognised intergrade considerably and are difficult to distinguish. 1 Petiole 10–30 mm long; lamina 6–14 cm long, 4–6 cm wide, the areolae usually villous beneath; fig 10–15 mm diam. 26a. var. opposita 1: Petiole 2–15 mm long; lamina to 9 cm long and 6 cm wide; fig 8–10 mm diam., the basal bracts often spaced along peduncle 2 Leaf lamina scabrid, minutely foveolate beneath with puberulous stomatal pits 26b. var. indecora 2: Leaf lamina aculeate on upper side, the margin with short rigid hairs; lower side villous to glabrescent, smooth to lightly foveolate from the bulging reticulations 26c. var. micracantha 26a. Ficus opposita Miq. var. opposita Illustration: K.Domin, Biblioth. Bot. 89: 17, fig. 120 (1921) as F. xerophila. Leaf lamina 6–14 cm long, 4–6 cm wide; petiole 10–30 mm long. Figs 10–15 mm diam. Widely distributed in eastern Malesia and New Guinea; in Australia, recorded from Qld, but possibly also in N.T. and N.S.W. Regions: CYRK, BURD, DWSN. Map 62. Qld: Laglan Stn, L.S.Smith 10305 (BRI); Stannary Hills, J.L.Bancroft s.n. (BRI); Cooktown, S.T.Blake 23509 (BRI, NSW). 26b. Ficus opposita var. indecora (Cunn. ex Miq.) Corner, Gard. Bull. Singapore 17: 471 (1960) F. indecora Cunn. ex Miq., London J. Bot. 7: 426 (1848). T: Careening Bay, [W.A.], A.Cunningham s.n.; holo: K. Leaf lamina to 9 cm long and 6 cm wide, scabrid, minutely foveolate beneath with puberulous stomatal pits, not villous; petiole 2–12 mm long. Occurs in northern W.A. and N.T. Regions: PILB, SNDY, KIMB, ARNM. Map 63. W.A.: Mt Trafalgar, A.S.George 12700 (PERTH); Yathalla Ck, Hamersley Ra., C.A.Gardner 6391 (PERTH); entrance to Logue Springs, K.F.Kenneally 5691 (PERTH). N.T.: Hayes Ck, S.T.Blake 16168A (BRI); Adelaide River, S.T.Blake 16689 (BRI).

56 MORACEAE 7. Ficus

26c. Ficus opposita var. micracantha (Miq.) Corner, Gard. Bull. Singapore 17: 471 (1960) F. micracantha Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 221 (1867); F. aculeata var. micracantha (Miq.) Benth., Fl. Austral. 6: 175 (1873). T: Cairnscross Is., Qld, F.Mueller; holo: K. F. aculeata Miq., London J. Bot. 7: 426 (1848). T: not designated. F. orbicularis Miq., London J. Bot. 7: 426 (1848). T: not designated. ?F. beckleri Miq., J. Bot. Neerl. 1: 241 (1861). T: Fitzmaurice R., Arnhem Land, [N.T.], Beckler; n.v. Leaf lamina aculeate above, the margin with short rigid hairs; underside villous glabrescent, smooth to lightly foveolate from the bulging reticulations; petiole 2–15 mm long. Occurs in northern W.A., N.T. and Qld. Regions: PILB, KIMB, ARNM, BKLY, CARP, CYRK, BURD, DWSN. Map 64. W.A.: Dampier Archipelago, R.D.Royce 7147 (PERTH); Winjana Gorge, A.C.Beauglehole 11223 (PERTH). N.T.: Bickerton Is., R.L.Specht 636 (BRI); Oenpelli, R.L.Specht 1055 (BRI). Qld: Karumba, S.T.Blake 15134 (BRI).

27. Ficus scobina Benth., Fl. Austral. 6: 176 (1873) T: Port Darwin, N.T., Schultz 410; lecto: K, fide E.J.H.Corner, Gard. Bull. Singapore 17: 471 (1960). Spreading shrub or tree to 8 m, generally strongly scabrid. Leaves opposite to alternate, obovate, rarely pentagonal, often faintly sinuous, obtuse, rarely acute, the base rounded and often narrowed; lamina 5–16 cm long, 3–8 cm wide; lateral veins 8–13 pairs; petiole to 1 cm long; stipules to 5 mm long. Figs rarely cauliflorous, globular, 1 cm diam., scabrid; ostiole to 3 mm diam., slightly raised, with protruding apical bracts; basal bracts variously arranged; peduncle to 1 cm long. Male flowers with 4 or 5 tepals; stamens 1 or 2. Female and gall flowers pedicellate; tepals 5. Sandpaper Fig. Fig. 8 Occurs in northern W.A. and N.T.; grows along watercourses in monsoon forest. Regions: KIMB, ARNM. Map 65. W.A.: near Goose Hill, W.V.Fitzgerald 1598 (PERTH). N.T.: Yirrkala, R.L.Specht 873 (BRI, NSW); Pig Hill, M.O.Rankin 1444 (NSW); Katherine, H.S.McKee 8420 (NSW). Closely related to F. opposita from which it differs in leaf venation and habitat preference. When injured, adventitious shoots develop that tend to produce linear to narrowly oblong scabrid leaves similar to those in F. podocarpifolia. May be ±deciduous in N.T.

28. Ficus podocarpifolia Corner, Blumea 22: 305 (1975) T: Boongaree Is., Prince Frederick Harbour, W.A., 14 July 1973, P.G.Wilson 11347; holo: PERTH; iso: NSW. Illustration: E.J.H.Corner, op. cit. fig. 2. Spreading shrub to 2 m, generally scabrid. Leaves alternate, often subverticillate or opposite, shortly petiolate to subsessile, usually reflexed, linear to oblong, setose, obtuse to apiculate, the base rounded to somewhat truncate or cordate; lamina 2–6.5 cm long, 0.3–0.8 cm wide, villous and scabrid; margins recurved; lateral veins 16–30 pairs, at 90° to midrib. Figs sessile, globular, to 1 cm diam., lightly to densely scabrid; ostiole 2–3 mm diam., ostiolar bracts forming a small umbo; basal bracts minute; peduncle to 2 mm long. Male flowers in 1 ring, pedicellate; tepals 3–5; stamen 1. Female and gall flowers sessile to subsessile; tepals 3–5. Fig. 33B–D. Occurs in far northern W.A. and N.T.; grows in sandstone country often close to water-courses. Regions: KIMB, ARNM. Map 66. W.A.: South-East Osborne Is., Bonaparte Archipelago, P.G.Wilson 11096 (PERTH); Mitchell Falls, West Kimberley, K.F.Kenneally 5001 (PERTH); near Fern Gully, Prince Regent River Reserve, A.S.George 12816 (PERTH). N.T.: Deaf Adder Gorge, C.R.Dunlop 4400 (DNA); 2 km north of Nabarlek, M.O.Rankin 2207 (DNA).

57

Figure 33. Ficus. A, F. leptoclada, fruiting branchlet ×0.5 (Hartley & Hyland 14160, BRI). B–D, F. podocarpifolia. B, branchlet ×0.5 (A.George 12816, PERTH); C, leaf lamina ×1; D, fruit ×1 (C–D, P.Wilson 11096, PERTH). E, F. coronulata ×0.5 (not recorded). Drawn by L.Hoare.

58 MORACEAE 7. Ficus

Distinguished by shrubby habit and linear to oblong scabrid leaves. Probably a juvenile form of Ficus scobina (see M.O.Parker 342 & M.Lazarides 9105, DNA).

29. Ficus fraseri Miq., London J. Bot. 7: 235 (1848) T: Brisbane R., [Qld], C.Fraser 154; holo: K. F. stenocarpa F.Muell. ex Benth., Fl. Austral. 6: 174 (1873). T: Rockingham Bay, Qld, J.Dallachy; syn: NSW. F. aspera var. subglabra Benth., Fl. Austral. 6: 175 (1873); F. subglabra (Benth.) F.Muell., Fragm. 9: 152 (1875); F. stephanocarpa var. subglabra (Benth.) Maiden & E.Betche, Census New South Wales Pl. 55 (1916). T: Brisbane R., Qld, F.Mueller & F.M.Bailey; syn: NSW; Hunter R., N.S.W., R.Brown; syn: n.v. Tree to 10 m, strongly scabrid but otherwise glabrous. Leaves alternate, rarely opposite, ovate, elliptic or obovate, entire, obtuse to shortly acuminate, the base rounded to acute; lamina 8–13 cm long, 4–6 cm wide, strongly scabrid; lateral veins 6–8 pairs; petiole 1–2 cm long; stipules small. Figs ellipsoidal, ovoid to globular, 1 cm diam., strongly scabrid; ostiole umbonate with protruding apical bracts; basal bracts 3, forming a small collar; peduncle to 1.5 cm long. Male flowers often in ellipsoidal figs, with stamens 2. Female flowers with 3–5 tepals, often fringed. Occurs widely but disjunctly in eastern Qld and north-eastern N.S.W., rare in N.T.; also in New Caledonia and Vanuatu. Regions: ARNM, CYRK, DWSN, MCPH, NEPN. Map 67. N.T.: junction of Daly R. & Bradshaw Ck, C.S.Robinson R.482 (DNA). Qld: Scrubby Ck, Herberton Ra., S.F.Kajewski 1362 (BRI); Etty Bay, C.T.White 11752 (BRI). N.S.W.: Upper Eungella, Tweed R., C.T.White 3279 (BRI); Yabbra State Forest, S.Clark et al. 1712 (NSW). Figs similar to those of F. leptoclada, but the 2 species are otherwise different.

30. Ficus coronulata Miq., J. Bot. Neerl. 1: 242 (1862) T: Arnhem Land, [N.T.], Aug. 1855; n.v. F. salicina F.Muell., Fragm. 4: 49 (1863). T: between Victoria R. & Fitzmaurice R., [N.T.], F.Mueller; n.v. Tree to 15 m with dense crown; branches often pendulous. Leaves alternate, lanceolate, attenuate, the base cuneate; lamina 15–25 cm long, 4–5 cm wide, lightly scabrid; lateral veins 20–30 pairs; petiole 2–4 cm long; stipules 1 cm long. Figs globular, to 1.3 cm diam., lightly scabrid; ostiole protruded into a crown 3 mm diam., 2 mm high; ostiolar bracts triangular with fringed margin; basal bracts to 2 mm long; peduncle to 2 cm long. Male flowers with 4 or 5 tepals; stamens 1 or 2. Female and gall flowers pedicellate; tepals 5. Peach Leaf Fig, Crown Fig. Fig. 33E. Endemic in northern W.A. and N.T.; grows along creeks in a variety of soil types. Regions: KIMB, ARNM, TNMI, BKLY. Map 68. W.A.: Fitzroy Crossing, T.E.H.Aplin 4703 (PERTH); Fitzroy Crossing, Oct. 1906, W.V.Fitzgerald (NSW). N.T.: Dorisvale, M.O.Rankin 1659 (DNA, NSW); Katherine R.–Maud Ck junction, S.Jacobs 3794 (NSW); Deaf Adder Ck Basin, R.Schodde AE92 (BRI). Easily distinguished by attenuate lamina and crown-like ostiole of the fig.

31. Ficus coronata Spin, Cat. Hort. St. Sebast. 29 (1818) T: cultivated specimen: TO n.v., fide E.J.H.Corner, Gard. Bull. Singapore 17: 472 (1960). F. muntia Link, Enum. Pl. Berol. 2: 450 (1822). T: not designated. F. scabrifolia A.Rich., Voy. Astrolabe 2: 44, t. 17 (1834). T: Moreton Bay, [Qld], collector unknown; n.v. F. stephanocarpa Warb., Repert. Spec. Nov. Regni Veg. 1: 75 (1905). T: Toowoomba, Qld, O.Warburg 18478; n.v. [F. aspera auct. non G.Forster: G.Bentham, Fl. Austral. 6: 174 (1873)] Illustration: L.Costermans, Native Trees & Shrubs S.E. Australia 150, 151 (1981).

59 7. Ficus MORACEAE

Shrub or tree to 12 m, generally strongly scabrid. Leaves spiral, ovate to oblong, dentate to crenulate, acuminate, cuneate to rounded and often asymmetrical at base, often densely brown-villous beneath; lamina 6–15 cm long, 3–8 cm wide; lateral veins 6–8 pairs; petiole and stipules to 1 cm long. Figs sometimes cauliflorous, ellipsoidal to ovoid, 1.5 cm long, scabrid, often also villous; ostiole to 4 mm diam., raised with protruding apical bracts; lateral bracts present; basal bracts variously arranged; peduncle to 1 cm long. Male flowers with 4 or 5 tepals; stamens 1 or 2. Female and gall flowers with 4–6 tepals. Sandpaper Fig. Fig. 34A. Endemic in Australia; occurs widely in eastern Qld and N.S.W., rare in N.T. and far eastern Vic. Grows in a wide range of habitats. Regions: BURD, DWSN, MCPH, NEPN, HOWE. Map 69. N.T.: Roper R., W.Baldwin Spencer (AD). Qld: Mt Tamborine, C.T.White 3589 (BRI). N.S.W.: Allyn R., P.Hind 351 (NSW); Whispering Gallery, R.Coveny 9734 (NSW). Vic.: E Gippsland, N.A.Wakefield 4585 (NSW).

Ser. 12. Pallidae

Ficus ser. Pallidae Miq., London J. Bot. 7: 433 (1848). Type: F. tinctoria G.Forster Trees, often strangling. Leaves coriaceous. Figs without lateral bracts, with minute internal bristles. Tepals puberulous or lightly hispid.

32. Ficus tinctoria G.Forster, Ins. Fl. Austral. 76 (1786) T: Society Islands, G.Forster; n.v. subsp. tinctoria F. validinervis F.Muell. ex Benth., Fl. Austral. 6: 166 (1873). T: Rockingham Bay, Qld, J.Dallachy; holo: K. Illustration: F.A.W.Miquel, London J. Bot. 7: t. 6B (1848). Large tree or banyan. Leaves alternate, oblong or ovate-elliptic, often strongly asymmetrical, acute, the base subcordate, rounded or cuneate, often decurrent; lamina 6–15 cm long, 4–15 cm wide, lightly scabrid, otherwise glabrous, coriaceous; lateral veins 7–9 pairs; petiole less than 1 cm long; stipules small. Figs often below leaves, globular, 8–12 mm diam., scabrid; ostiole slightly umbonate, 2 mm diam., with protruding apical bracts; peduncle usually absent; pedicel to 1 cm long; basal bracts 3, at base of pedicel. Male flowers in 1 or 2 rows, sessile; stamen 1. Female and gall flowers subsessile to pedicellate. Widely distributed in eastern Asia, through Malesia, the South Pacific and to Australia, where it occurs in northern W.A. and Qld. Regions: KIMB, CYRK. Map 70. W.A.: Boongaree Is., P.G.Wilson 11424 (PERTH); Mt Trafalgar, A.S.George 12679 (PERTH); Devils Pass, Napier Ra., C.A.Gardner 10134 (PERTH); Red Rock Ck, near Bungle Bungle, K.F.Kenneally 9236 (PERTH). Four subspecies are recognised; only the type occurs in Australia. This species has been confused with F. virgata.

33. Ficus virgata Reinw. ex Blume, Bijdr. 454 (1825) T: from Moluccas, C.G.C.Reinwardt 1376; holo: L. var. virgata F. pinkiana F.Muell., S. Sci. Rec. 2: 273 (1882). T: Trinity Bay, Qld, J.Pink; n.v. F. esmeralda Bailey, Queensland Agric. J. 1: 452 (1897). T: Mourilyan Harbour, Qld, W.Mugford; holo: BRI. [F. philippinensis auct. non Miq.: G.Bentham, Fl. Austral. 6: 173 (1873)]

60

Figure 34. Ficus. A, F. coronata, fruiting branchlet ×0.5 (T.Hartley 14016, NSW). B, F. virgata, fruiting branch ×1 (S.Kajewski 1327, BRI, NSW). C–D, F. variegata. C, branchlet ×0.5 (V.Moriarty 1933, BRI); D, fruiting branchlet ×0.5 (S.Kajewski 1416, BRI). Drawn by L.Hoare.

61 7. Ficus MORACEAE

Tree to 10 m. Leaves distichous to alternate, elliptic to oblong, often slightly asymmetrical, acuminate, the base rounded to cuneate, often decurrent; lamina 10–20 cm long, 4–7 cm wide, glabrous, coriaceous; lateral veins 10–12 pairs; petiole 1–1.5 cm long; stipules small. Figs globular, 1 cm diam., finely scabrid; ostiole umbonate, 2 mm diam., a few apical bracts protruding; peduncle to 5 mm long; pedicel 5 mm long; basal bracts 3, often at base of pedicel. Male flowers subsessile; stamen 1. Female flowers subsessile; tepals 3 or 4. Fig. 34B. Widely distributed from Formosa through eastern Malesia and Melanesia to the SW Pacific; in Australia occurs in Qld. Grows in rainforest. Region: CYRK. Map 71. Qld: between Cooktown & Bloomfield R., L.S.Smith 10730 (BRI); Atherton Tableland, S.F.Kajewski 1327 (BRI); Daintree R., L.J.Brass 2333 (BRI); Etty Bay, C.T.White 11756 (BRI); Mossman Gorge, R.F.Thorne & J.G.Tracey 22866 (BRI). Three varieties are recognised of which the type occurs in Australia. Closely related to F. tinctoria, but distinguished by the lamina being less asymmetrical, elliptic to oblong instead of subrhombic or angled, and by the much smaller figs.

Sect. 7. Adenosperma

Ficus sect. Adenosperma Corner, Reinwardtia 4(3): 353 (1958) [pagination in some sets as 43]. Type: F. adenosperma Miq. Dioecious trees or shrubs. Leaves alternate, rarely distichous. Figs mostly pedunculate, with collar of 3 basal bracts, often with lateral bracts. Flowers sessile or subsessile. Male flowers ostiolar in 1 or 2 rows; stamen 1. Ovary dark red or red-spotted; style lateral to gynobasic, glabrous. Seeds compressed, keeled, the keel usually double at base of seed, smooth. Only 1 of the 2 series in this section occurs in Australia.

Ser. 13. Amphigenae

Ficus ser. Amphigenae Corner, Reinwardtia 4(3): 354 (1958) [pagination in some sets as 44]. Type: F. adenosperma Miq. Cystoliths present on both surfaces of leaf lamina.

34. Ficus mollior F.Muell. ex Benth., Fl. Austral. 6: 173 (1873) T: Rockingham Bay, Qld, J.Dallachy s.n.; iso: NSW. var. mollior F. dielsii Warb., Repert. Spec. Nov. Regni Veg. 1: 75 (1905). T: Barron R., Qld, 3 June 1902, L.Diels s.n.; n.v. Tree to 15 m, low spreading branches often knobbly. Upper branchlets villous. Leaves alternate, elliptic to ovate, acute to shortly acuminate, the base narrowed to cordate; lamina 10–18 cm long, 5–9 cm wide, glabrous above, villous below; lateral veins 9–11 pairs; petiole 2–4 cm long, woolly, soon glabrous; stipules c. 1.5 cm long. Figs often cauliflorous, obovoid to globular, to 1 cm diam., villous; ostiolar bracts forming a small protuberance; basal bracts to 2 mm long; peduncle 5 mm long. Male flowers: tepals 3. Female flowers with unequal tepals. Widespread in the Moluccas through New Guinea to north-eastern Qld. Grows in rainforest along creeks. Regions: CYRK, BURD, DWSN. Map 72.

62 MORACEAE 7. Ficus

Qld: Range Rd, Atherton Tableland, S.F.Kajewski 1306 (BRI, NSW); Gadgarra Reserve, S.F.Kajewski 1190 (BRI, NSW); Jarra Ck, W of Tully, L.S.Smith 4899 (BRI); Mossman River Gorge, L.J.Brass 2133 (BRI); Palm Islands, N of Townsville, T.L.Bancroft 9 (BRI). Three varieties and 1 form have been distinguished, of which only the type variety occurs in Australia.

35. Ficus adenosperma Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 233 (1867) T: Indonesia; Celebes, Teysmann; Moluccas, De Vriese; syn: n.v. F. depressa Benth., Fl. Austral. 6: 172 (1873). T: Mt Elliott, Qld, E.Fitzalan; n.v. F. chaetophora Warb., Schumann & Lauterb., Nachtr. Fl. Sudsée 246 (1905); F. adenosperma var. chaetophora (Warb.) Corner, Gard. Bull. Singapore 18: 28 (1960). T: New Guinea, R.Schlechter 13816; syn: n.v.; New Guinea, Lauterbach 694, 2074, 2225, 2653; syn: n.v.; New Guinea, Rodatz & Klink s.n.; syn: n.v. F. frutescens Bailey, Queensland Agric. J. n. ser. 1: 276 (1914). T: Bellenden-Ker Ck and Harvey Ck, Qld, Feb. 1914, E.W.Bick; holo: BRI. F. adenosperma var. glabra Corner, Gard. Bull. Singapore 18: 28 (1960). T: Guadalcanal Is., Solomon Islands, S.F.Kajewski 2480; holo: L. Illustrations: F.M.Bailey, op. cit. 277, t. 40; M.O.Rankin, N. Terr. Bot. Bull. 5: t. 5 (1982). Tree 8–10 m. Leaves alternate, narrowly ovate, ovate, elliptic or somewhat pentagonal, shortly acuminate to attenuate, the base cuneate to subcordate; lamina 6–17 cm long, 3–7 cm wide, glabrous; lateral veins 8–10 pairs; petiole 1.5–4 cm long, with brown flaky outer bark, pubescent, soon glabrous; stipules 2–3 cm long. Figs globose, 1–1.3 cm diam., smooth to pustular; ostiole 3 mm diam.; peduncle to 1 cm long; basal bracts forming a collar on peduncle or at base of fig-body; lateral bracts sometimes present. Male flowers in 1 row; tepals 4 or 5. Female and gall flowers sessile; tepals 3 or 4. Occurs in northern N.T. and Qld in Australia and in the Celebes, Moluccas, New Guinea and from Bismarck Archipelago to Vanuatu. Regions: ARNM, CYRK, BURD. Map 73. N.T.: Adelaide R., 1 km downstream from Daly River road, M.O.Rankin 1761 (BRI, DNA, NT); Adelaide R., c. 1 km downstream from Daly River road, M.O.Rankin 1774, 1775 (DNA). Qld: Deeral, near Babinda, S.T.Blake 14964 (BRI); Sareena Ra., R.F.Thorne 22610 (BRI); Mossman River Gorge, L.J.Brass 2114 (BRI). The Australian populations of this species are indistinguishable from the type variety of F. adenosperma, and var. chaetophora and var. glabra are not maintained here. This decision is based on a comparison of recent collections from N.T. and Qld with authentic material of var. adenosperma from Indonesia and Vanuatu.

Sect. 8. Neomorphe

Ficus sect. Neomorphe King, Ann. Roy. Bot. Gard. (Calcutta) 1(1): 2 (1887). Type: F. variegata Blume Dioecious buttressed trees. Leaves alternate, with cystoliths. Figs pedunculate, mostly cauliflorous, with collar of 3 basal bracts; lateral bracts absent. Flowers sessile or subsessile; tepals free or united, red to brownish. Male flowers ostiolar; stamens 2; filaments shortly joined. Gall and female flowers: stigma simple. Seeds lenticular, smooth, slightly keeled. A section of 2 series, represented in Australia by ser. Variegatae.

63 7. Ficus MORACEAE

Ser. 14. Variegatae

Ficus ser. Variegatae Corner, Gard. Bull. Singapore 18: 33 (1960). Type: F. variegata Blume Female and gall flowers sessile to shortly pedicellate; ovary brownish red; style short, glabrous.

36. Ficus variegata Blume, Bijdr. 459 (1825) T: from Java; n.v. var. variegata F. ehretioides F.Muell. ex Benth., Fl. Austral. 6: 171 (1873). T: Rockingham Bay, Qld, J.Dallachy; iso: NSW. Tree to 20 m; trunk to 3 m diam., with plank buttresses to 5 m high. Leaves alternate, ovate to very broadly ovate, acuminate, sometimes sinuate-dentate, the base cordate; lamina 15–25 cm long, 8–16 cm wide, glabrous, smooth; lateral veins 6–8 pairs, widely spaced, arching, basal pair reaching to at least 1/3 length of lamina; petiole 5–12 cm long, often with brown flaky epidermis; stipules to 2 cm long. Figs cauliflorous, obovoid to pyriform, 2–2.5 cm long and wide, glabrous, densely lenticellate; ostiole slightly raised, 3–4 mm diam.; peduncle to 4 cm long; basal bracts 1–2 mm long. Female flowers sessile to subsessile; style subterminal; stigma truncate, glabrous. Fig. 34C–D. Widely distributed from S Asia through Malesia and Melanesia to northern Qld, Australia. Region: CYRK. Map 74. Qld: Daintree R., S.F.Kajewski 1416 (BRI, NSW); Mossman River Gorge, L.J.Brass 2112 (BRI); Mulgrave R., Bellenden-Ker Expedition, F.M.Bailey (BRI). Five varieties are recognised, of which only the type variety occurs in Australia.

37. Ficus nodosa Teysmann & Binnend., Tijdschr. Ned.-Indie 29: 245 (1867) T: Moluccas, collector unknown; n.v. Large buttressed trees. Leaves alternate, broadly ovate-cordate, sinuate, acuminate, the base cordate; lamina 18–22 cm long, 14–18 cm wide, glabrous, smooth; lateral veins 5–7 pairs, widely spaced, arching, basal pair reaching at least 1/3 length of lamina; petiole 5–13 cm long, often with brown flaky epidermis. Figs cauliflorous, oblong to obovoid, to 2.5 cm long and wide, densely lenticellate; ostiole slightly raised, 3–4 mm diam.; peduncle 2–3 cm long; basal bracts 2–9 mm long. Occurs on Cape York Peninsula, Qld; also in eastern Malesia and Melanesia. Region: CYRK. Map 75. Qld: Claudie R., B.Hyland 2915 (BRI); Massey Ck, J.R.Clarkson 2589 (BRI); upper Massey Ck, L.S.Smith 11881 (BRI); Smithfield, L.J.Webb & J.G.Tracey 7262A (BRI); McIlwraith Ra., L.J.Webb & J.G.Tracey 9137 (BRI). Poorly collected in Australia; further collections are needed to elucidate the floral morphology of the Australian populations. From gross morphology, this species appears remarkably similar to F. variegata though they belong to different subseries.

Sect. 9. Sycocarpus

Ficus sect. Sycocarpus Miq., Ann. Sci. Nat. ser. 3, 1: 33 (1844). Type: F. hispida L.f. F. sect. Covellia (Gasp.) Benth., Fl. Austral. 6: 163, 176 (1873); Covellia Gasp., Nov. Gen. Fic. 10 (1844). T: C. ulmifolia (Lam.) Gasp.

64 MORACEAE 7. Ficus

Dioecious trees or shrubs. Leaves alternate; lamina mostly without basal glands. Figs often cauliflorous, with or without lateral bracts, mostly with collar of 3 basal bracts. Tepals entire, gamophyllous, or absent. Male flowers ostiolar; stamen 1, rarely 2. Female stigma simple. Gall stigma widely funnel-shaped. Seeds not compressed, rarely keeled.

Ser. 16. Tuberculifasciculatae

Ficus ser. Tuberculifasciculatae Sata, Contr. Hort. Inst. Taihoku Univ. 32: 298, 338 (1944). Type: F. repandifolia Elmer = F. fistulosa Reinw. ex Blume Cystoliths present on lower surface of leaf lamina.

38. Ficus congesta Roxb., Fl. Ind. 2nd edn, 3: 560 (1832) T: from Moluccas, introduced into Bot. Gard. Calcutta; n.v. var. congesta F. fasciculata F.Muell. ex Benth., Fl. Austral. 6: 177 (1873). T: Fitzroy R., Rockhampton, Qld, Bowman, Thozet & Dallachy; n.v. F. fasciculata var. ?subopposita Benth., Fl. Austral. 6: 177 (1873). T: Rockingham Bay & Mackay R., Qld, J.Dallachy; n.v. F. setistyla Warb., Repert. Spec. Nov. Regni Veg. 1: 77 (1905). T: below Russell R., Qld, 6 June 1902, L.Diels 8497; n.v. F. trichostyla Warb., Repert. Spec. Nov. Regni Veg. 1: 77 (1905). T: Cairns, Qld, O.Warburg 18500; n.v. Illustration: M.O.Rankin, N. Terr. Bot. Bull. 5: t. 4 (1982). Spreading tree, often branching at base; branchlets often pubescent. Leaves alternate, broadly elliptic to rhombic, rarely ovate, crenulate to denticulate, acute to shortly acuminate, the base rounded to cuneate, often cordate; lamina 12–25 cm long, 8–13 cm wide, often scabrid, the lowerside often densely pubescent, chartaceous to coriaceous; lateral veins 10 pairs; petiole 1.5–4 cm long, reddish-brown-hispid; stipules 1–2 cm long, amplexicaule, often leaving prominent scars. Figs axillary and cauliflorous, sometimes geocarpic, pyriform to discoid, ribbed, 2.5 cm long and wide, hispid; ostiole depressed, 2–3 mm diam., with a few protruding apical bracts; peduncle 1 cm long, with collar of 3 basal bracts. Male flowers in 1 row; tepals 2; stamen 1. Female and gall flowers similar, sessile; perianth enveloping ovary. Occurs in rainforest in Qld and N.T.; also widely distributed from the Philippines through eastern Malesia to Melanesia. Regions: ARNM, CYRK, BURD. Map 76. N.T.: Limestone Arch, Douglas R., M.O.Rankin 1230 (DNA, NT); Howard Springs, M.O.Rankin 1241 (DNA, NT). Qld: Daintree R., L.J.Brass 2229 (BRI); Garradunga, Cook district, C.T.White 11755 (BRI); Whyanbeel Ck, near Mossman, S.T.Blake 15012 (BRI). Three varieties are recognised, of which only var. congesta occurs in Australia. In Qld, trees of this variety have been encountered bearing figs on runners issuing from the base of the trunk and rooting in the ground, thus showing features of geocarpism (L.J.Brass 2229; C.T.White 11755).

39. Ficus hispida L.f., Suppl. Pl. 442 (1781) Coviella hispida (L.f.) Miq., London J. Bot. 7: 462 (1848). T: from Java, C.P.Thunberg; n.v. var. hispida F. oppositifolia Roxb., Pl. Corom. 2: 14, fig. 124 (1798); Coviella oppositifolia (Roxb.) Gasp., Ric. Caprifico 85 t. VIII (1845). T: Coromandel Coast, India, W.Roxburgh; n.v. Illustrations: G.King, Ann. Roy. Bot. Gard. (Calcutta) 1(2): t. 154, 155 (1888); E.J.H.Corner in M.D.Dassanayake & F.R.Fosberg, Rev. Handb. Fl. Ceylon 3: 278, fig. 27 (1981); M.O.Rankin, N. Terr. Bot. Bull. 5: t. 3 (1982).

65 7. Ficus MORACEAE

Spreading tree to 13 m. Leaves opposite-decussate, sometimes alternate, pentagonal to oblong, crenulate, acuminate, the base subcordate, asymmetrical; lamina 15–35 cm long, 6–20 cm wide, hispid with short white stiff hairs; lateral veins 7–9 pairs, with prominent intercostals; petiole 5–10 cm long. Figs often cauliflorous in clusters to 1 m long, often trailing, depressed-globose, 1–2 cm diam., densely brown-pubescent; ostiole closed by 5–10 bracts; lateral bracts often present; peduncle 5–15 mm long; basal bracts 3, to 1.5 mm long. Male flowers in 1 or 2 rows. Female and gall flowers subsessile to pedicellate; perianth enveloping ovary. Fig. 10. Occurs in northern W.A., N.T. and Qld; also widely distributed from southern Asia through Malesia to New Guinea. Grows in monsoon forest. Regions: KIMB, ARNM, CYRK, BURD. Map 77. W.A.: Mt Trafalgar, A.S.George 12722 (PERTH). N.T.: Gunn Point area, M.O.Rankin 1579 (DNA, NSW); Darwin, S.N.Cousins 66 (DNA). Qld: Cardwell, A. & M.Thorsborne 3 (BRI); Yungaburra, Atherton, A.W.Dockrill 22 (BRI). Considered to comprise 3 varieties; only the typical variety reaches Australia, and is distinguished by the figs not being geocarpic.

40. Ficus septica Burman f., Fl. Ind. 226 (1768) T: from Asia, probably SE Asia, collector unknown; n.v. F. casearia F.Muell. ex Benth., Fl. Austral. 6: 177 (1873). T: Rockingham Bay, Qld, J.Dallachy; syn: n.v.; Fitzroy Is. & Endeavour R., Qld, collector unknown; syn: n.v. Tree to 15 m, glabrous, branches often pendulous. Leaves alternate, elliptic, ovate or oblong, entire, acuminate, the base cuneate; lamina 12–20 cm long, 5–7 cm wide; lateral veins 9–11 pairs; petiole 2–5 cm long; stipules 3 cm long. Figs axillary, cauliflorous, pedunculate, obovoid to depressed-globular, to 1.5 cm diam., often with 6–8 longitudinal ribs; ostiole often sunken, ostiolar bracts sometimes forming a slightly raised crown 3 mm diam.; basal bracts 3, 1–1.5 mm long, forming a collar. Male flowers: stamen 1. Female flowers with united sepals. Widespread in SE Asia through New Guinea and Solomon Islands to Vanuatu. In Australia, confined to northern Qld. Three varieties recognised of which 2 occur in Australia. Figs axillary and on leafless branchlets 40a. var. septica Figs axillary and cauliflorous on trunk 40b. var. cauliflora 40a. Ficus septica Burman f. var. septica Figs mostly axillary but also on upper branchlets below the leaves. Distribution as for species. In Australia, occurs in Qld, mainly in rainforest and wetter edges of scrub to 1,000 m altitude, often as a pioneer in regrowth in various soil types, especially granitic alluvium. Region: CYRK. Map 78. Qld: Ghurka Pocket, Boonjie, Atherton Tableland, S.F.Kajewski 1226 (BRI, NSW); Mt Lewis, near Julatten, P.Hind 133 (NSW); Clump Point, R.F.Thorne & J.G.Tracey 22495 (BRI); Gadgarra, Atherton, S.F.Kajewski 1021 (BRI); Claudie R., L.J.Webb & J.G.Tracey 8520 (BRI). 40b. Ficus septica var. cauliflora Corner, Gard. Bull. Singapore 18: 61 (1960) T: San Cristobal, Solomon Islands, E.S.Brown 5482; holo: BM n.v., fide E.J.H.Corner, loc. cit. Figs cauliflorous on trunk, in branched clusters, also axillary. Widely distributed in Tanimbar Islands, Solomon Islands and Vanuatu. In Australia, known from northern Qld. Region: CYRK. Map 79. Qld: Davies Ck, Atherton, A.W.Dockrill 50 (BRI); Shipton Flat, Cooktown, V.K.Moriarty 1426 (BRI); Innisfail, S.T.Blake 15282 (BRI); Boonjee, near Malanda, S.T.Blake 15212 (BRI).

66 MORACEAE 7. Ficus

The character may be a developmental feature, and further study is needed to assess the validity of the variety.

Doubtful and excluded names

Ficus aspera var. abbreviata Miq., J. Bot. Neerl. 1: 241 (1862). T: Twofold Bay, N.S.W., Sept. 1860, F.Mueller; n.v. Application uncertain. Ficus caulobotrya (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 287 (1867). Urostigma caulobotryum Miq., London J. Bot. 6: 568 (1847). T: India, Wright; n.v. Recorded for Australia by Miquel, but E.J.H.Corner, Gard. Bull. Singapore 17: 378 (1960); Gard. Bull. Singapore 21: 97 (1965), recommended that this name be regarded as doubtful. The type is a mixture of F. tsjahela Burman f. and F. arnottiana Miq. Ficus virens var. glabella (Blume) Corner, Gard. Bull. Singapore 17: 377 (1960). F. glabella Blume, Bijdr. 452 (1825). T: Java, Indonesia, collector unknown; n.v. Listed by F.Mueller, Second Syst. Census Austral. Pl. 37 (1889), but probably a misdetermination; the taxon does not occur in Australia. Ficus mourilyanensis Bailey, Queensland Agric. J. 1: 452 (1897). T: Mourilyan harbour, near Esmeralda Plantation, Qld, W.Mugford; n.v. Probably conspecific with F. copiosa Steudel, see E.J.H.Corner, Gard. Bull. Singapore 21: 95 (1965).

Ficus microcarpa var. rigo (Bailey) Corner, Gard. Bull. Singapore 19: 385 (1962). F. rigo Bailey, Queensland Agric. J. 1(3): 235 (1897); F. retusa var. rigo (Bailey) Diels, Bot. Jahrb. 67: 183 (1935). T: not designated. Native of New Guinea; not in Australia.

Ficus scabra G.Forster, Fl. Ins. Austral. Prodr. 76 (1786). T: not designated. Native to the SW Pacific. Recorded by F.Mueller, Fragm. 6: 196 (1868) from Rockingham Bay, Qld, but probably a misdetermination.

Ficus stipulosa (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 287 (1867). Urostigma stipulosum Miq., London J. Bot. 6: 568 (1847). T: Philippines, Cuming 1978; n.v. This is F. caulocarpa Miq., a native of southern Asia and Malesia; not in Australia.

Ficus subgelderi var. rigida Corner, Gard. Bull. Singapore 17: 387 (1960). Urostigma rigidum Miq., London J. Bot. 6: 578 (1847); F. rigida (Miq.) Miq., Ann. Mus. Bot. Lugduno-Batavum 3: 286 (1867), nom. illeg. non Jack (1822). T: Penang, Malaysia, Herb. Hookerianum; holo: K. Recorded for Australia by Miquel (1867), but native of SE Asia and ; not in Australia.

Ficus subulata Blume, Bijdr. 461 (1825). T: Mt Paray, Indonesia, collector unknown; n.v. Native from India to New Guinea and the Solomon Islands. Recorded by F.Mueller, Second Syst. Census Austral. Pl. 38 (1889), but no collection seen from Australia.

67 7. Ficus MORACEAE

Taxotrophis rectinervis F.Muell., Fragm. 6: 192 (1868). T: between Clarence R. and Rockingham Bay, Qld, collector unknown; n.v. Possibly a species of Streblus.

Urostigma subglaucinum Miq., J. Bot. Neerl. 240 (1861). T: Rockhampton, Qld, F.Mueller; n.v. The application of this name is uncertain. Miquel was unsure if it belonged in Urostigma, and it was not interpreted by G.Bentham, Fl. Austral. 6: 160–178 (1873) or E.J.H.Corner, Gard. Bull. Singapore 21: 1–186 (1965).

URTICACEAE

W.-L.Chew

Herbs, sometimes epiphytic, shrubs, climbers or trees, occasionally with stinging hairs. Leaves simple, petiolate, stipulate, mostly with cystoliths. Inflorescence axillary, monoecious or dioecious, cymose, spicate or capitate. Flowers small, unisexual, rarely bisexual; perianth 1-whorled; tepals usually 3–5, occasionally 1, 2 or absent, free or connate, imbricate or valvate, persistent. Male flowers: stamens usually 3–5, opposite tepals, or reduced to 2 or 1; filaments inflexed in bud; anthers bilobed; pistillode usually present. Female flowers: ovary superior, rarely immersed in sockets in the inflorescence, sometimes adnate to calyx, usually 1-locular; ovule 1, basal, orthotropous; style 1; stigma 1, simple; staminodes sometimes present. Fruit an achene, rarely a fleshy drupe. Seed with endosperm; embryo straight. A family of about 52 genera and 1900 species, widely distributed, mostly at mid-altitudes on tropical mountains; 9 genera and 21 species in Australia representing all 5 tribes in the family. Species of Lindley have been widely cultivated as indoor plants. The Fibre of commerce comes from nivea Gaudich. Young shoots of the European Nettle, urens L. have been eaten in times of economic hardship. Species of the tribe Urticeae, especially the genera Urtica, Laportea and , are usually armed with irritant stinging hairs. C.Gaudichaud-Beaupré, Urticeae, Voy. Uranie 492–505 ‘1826’ (1830); H.A.Weddell, Revue de la Famile des Urticées, Ann. Sci. Nat. Bot. ser. 4, 1: 173–212 (1854); C.L.Blume, Urticaceae, Mus. Bot. 2: 137–170, 193–256 (1856); H.A.Weddell, Monographie de la Famile des Urticées, Arch. Mus. Hist. Nat. 9: 1–592 (1856–1857); H.A.Weddell, Urticaceae, in A.P. de Candolle, Prodr. 16(1): 32–235*64 (1869); G.Bentham, Urticeae trib. Eu-urticeae, Fl. Austral. 6: 182–192 (1873); F.M.Bailey, Urticaceae trib. Urticeae, Queensland Fl. 5: 1481–1488 (1902); K.Domin, Urticaceae, Beitr. Fl. Pflanzengeogr. Austral., Biblioth. Bot. 89: 572–577 (1921).

KEY TO TRIBES

1 Plants with irritant hairs (Urtica , Laportea , Dendrocnide) Trib. 1. URTICEAE 1: Plants without irritant hairs 2 Female perianth 3–5-partite (Elatostema , Procris) Trib. 2. ELATOSTEMEAE 2: Female perianth tubular, with a contracted orifice 3 Male perianth (Australina) bilabiate; stamen 1 Trib. 5. FORSSKAOLEAE 3: Not this combination of characters

68 URTICACEAE Key to tribes

4 Flowers unisexual; leaves opposite-decussate to alternate; herbs, climbers, shrubs or trees (Boehmeria , Pouzolzia , Pipturus , Nothocnide) Trib. 3. BOEHMERIEAE 4: Flowers often bisexual; leaves alternate; herbs (Parietaria) Trib. 4. PARIETARIEAE

KEY TO GENERA

1 Plants with irritant hairs, sometimes sparse 3. DENDROCNIDE 2 Trees; leaves alternate 2: Herbs; leaves opposite or alternate 3 Leaves opposite 1. URTICA 3: Leaves alternate 2. LAPORTEA 1: Plants without irritant hairs 4 Climbers, vines or lianes 9. NOTHOCNIDE 4: Herbs, sometimes creeping, sometimes epiphytic, or shrubs or trees 5 Herbs, sometimes creeping; stem often woody at base 6 Male perianth bilabiate; stamen 1; pistillode absent 11. AUSTRALINA 6: Male perianth 3–5-lobed; stamens 3–5; pistillode small, usually present 7 Female perianth 3- or 4-partite 8 Flowers in heads on flat or concave receptacles 4. ELATOSTEMA 8: Flowers in clusters, not in heads † PILEA 7: Female perianth tubular, the orifice 2–4-dentate 9 Flowers solitary, axillary, subtended by an involucre of 1 bract and 2 bracteoles † SOLEIROLIA 9: Flowers in clusters of 3 or more 10 Stipules present at least in young shoots; flowers unisexual; stigma filiform, unilaterally stigmatic; achenes black, smooth, shiny 7. POUZOLZIA 10: Stipules absent; flowers unisexual and bisexual; stigma sessile to long-styled, subcapitate to capitate with long stigmatic processes; achenes brown, smooth, shiny 10. PARIETARIA 5: Subshrubs, shrubs or trees 11 Female perianth of 3 or 4 (rarely 5) tepals; leaves opposite, 1 of each pair greatly reduced or some aborted so as to appear alternate; male inflorescence a loose cyme; female inflorescence capitate-globose, sessile to subsessile 5. PROCRIS 11: Female perianth tubular, free or adnate to ovary; leaves opposite with 1 of each pair greatly reduced, or alternate 12 Stipules entire; stigma persistent; female perianth tubular, free or only slightly attached to ovary, 2–4-dentate 6. BOEHMERIA 12: Stipules bifid; stigma caducous; female perianth adnate to ovary, minutely dentate 8. PIPTURUS † Soleirolia soleirolii (Req.) Dandy, based on Helxine soleirolii Req., has been recorded for S.A. and Vic. (e.g. J.P.Jessop, Fl. S. Australia 1: 118, 1986) but is not considered naturalised. † Pilea microphylla (L.) Liebm., a native of tropical America, has been recorded as a bush-house weed in Qld (F.M.Bailey, Queensland Fl. 5: 1484, 1902), but is not considered naturalised. It is a small tufted glabrous herb with small leaves in opposite unequal pairs.

69 Trib. !. Urticeae URTICACEAE

Trib. 1. URTICEAE

Urticaceae trib. Urticeae Type: Urtica L. Urticaceae Trib. Urereae Gaudich., Voy. Uranie 496 ‘1826’ (1830). T: Gaudich. Monoecious, rarely dioecious herbs or trees with irritant hairs. Leaves opposite-decussate or alternate. Flowers unisexual. Male flowers: perianth 4- or 5-partite; stamens 4 or 5; pistillode present. Female flowers: perianth 4-partite, unequal, often one pair larger than the other; pistil present; staminodes absent. About 10 genera, very widely dispersed throughout the world; 3 genera in Australia.

1. URTICA

Urtica L., Sp. Pl. 983 (1753); Gen. Pl. 5th edn, 423 (1754); the Latin name for the nettle. Type: U. dioica L. Herbs, monoecious, rarely dioecious, with irritant hairs. Leaves opposite-decussate, petiolate, simple, dentate to incised, very rarely subentire, with 3 or 5 nerves; cystoliths mainly punctiform; stipules free and lateral to connate and intrapetiolar. Inflorescences axillary, paired interrupted spikes or panicles, unisexual or bisexual. Male flowers: tepals 4, free, with irritant hairs; stamens 4; pistillode cupuliform, translucent. Female flowers: tepals 4, 1 opposite pair larger than other; pistil straight; style short or absent; stigma penicillate-capitate. Achene laterally compressed, ovoid, enclosed by tepals. A widespread genus of about 100 species throughout the world, mostly in temperate and warm temperate regions; 2 species in Australia, of which 1 is introduced. The common nettle, Urtica dioica L., has been recorded for Australia, e.g. F.Mueller, Fragm. 4: 87 (1864), R.H.Anderson, Contr. New South Wales Natl. Herb. 1: 19 (1939), but is apparently not established. It can be confused with U. urens but is a perennial dioecious herb, the lamina of the lower leaves longer than the petiole, and the inflorescence up to 10 cm long (J.H.Willis, Handb. Pl.Victoria 2: 31, 1972). Perennial herb; leaves broadly dentate; inflorescence unisexual, longer than petiole; larger tepals of female flowers glabrous 1. U. incisa Annual herb; leaves deeply, narrowly dentate; inflorescence bisexual, rarely longer than petiole; larger tepals of female flowers fringed with curved hairs 2. U. urens

1. Urtica incisa Poiret in J.B.A.P.M. de Lamarck, Encycl. Suppl. 4: 224 (1816) T: Australia, J.J.H. de Labillardière; n.v. Urtica lucifuga Hook., London J. Bot. 6: 285 (1847). T: ‘Hobart Town & Circular Head’, Tas., R.C.Gunn; n.v. Illustration: J.M.Black, Fl. S. Australia 2nd edn, 2: 261, fig. 312 (1963). Monoecious perennial herb. Leaves triangular to ovate, broadly dentate to incised, acute to caudate, truncate at base; lamina 5–12 cm long, 3–6 cm wide; lateral veins 4 or 5 pairs, 2 pairs from base, prominent, extending 2/3 the lamina length, the others short, near lamina apex; petiole 4–7 cm long; stipules 5–15 mm long. Inflorescences often paired, paniculate, unisexual, longer than petiole; male inflorescences sparsely branched, reduced to interrupted spikes. Flowers pedicellate, c. 1.5 mm long, 1 mm broad, glabrous. Male flowers: pistillode small. Female flowers: ovary ovoid. Achenes ovoid, smooth, c. 2 mm long. Scrub Nettle. Fig. 35A. Widespread in temperate W.A., Qld, N.S.W., Vic., S.A. and Tas.; frequent in wet cool places, especially near water courses and in wet highland areas in warmer latitudes. Regions:

70

Figure 35. A, Urtica incisa, habit ×0.5 (R.Pullen 4170, NSW). B–F, Laportea interrupta. B, ♂flower ×20; C, achene ×20; D, flowering branch ×0.75; E, ♀perianth ×22.5; F, ♀flower ×40. A drawn by D.Mackay. B–F reproduced by permission from Garden’s Bulletin, Singapore 25: 147, fig. 12 (1969).

71 1. Urtica URTICACEAE

EYRE, CYRK, BURD, DWSN, MCPH, NEPN, DARL, RIVR, OTWY, HOWE, TASM. Map 80. W.A.: Esperance, July 1937, R.F.Bredges (PERTH). S.A.: S of Blanchetown, Murray R., J.Z.Weber 3475 (NSW). Qld: between Tolga & Atherton, H.R.Thurston 444 (QRS). N.S.W.: The Native Wineyard, 5.5 km N of Cobbity, R.Coveny 7429 et al. (NSW). Vic.: W shore of Lake Wellington, H.Salasoo 5006 (NSW). Tas: Ocean Beach, Strahan, 20 Jan. 1949, L.A.S.Johnson (NSW).

2. *Urtica urens L., Sp. Pl. 984 (1753) T: Europe, collector unknown; n.v. Illustration: J.M.Black, Fl. S. Australia 2nd edn, 2: 261, fig. 311 (1963). Monoecious annual herb. Leaves ovate to elliptic, deeply incised, caudate, broadly cuneate at base; lamina 3–6 cm long, 1.5–4 cm wide; lateral veins 4 or 5 pairs, the second pair reaching nearly to lamina apex, the last two pairs short, near apex; petiole to 5 cm long; stipules small. Inflorescences paired, short spikes, bisexual, often shorter than petiole. Flowers subsessile, 1–2 mm long, c. 1 mm broad; males often at base of inflorescence. Male flowers: pistillode small. Female flowers: ovary ovoid. Achenes compressed-ovoid, finely warted, c. 2 mm long. Dwarf Nettle, Small Nettle. Fig. 11 Native of Europe; widely naturalised in Australia, in all States except N.T. Common in waste and cultivated land especially in the cooler latitudes and in the highlands of the warmer north. Regions: EYRE, BENC, LUWN, ESPR, MCPH, NEPN, DARL, RIVR, OTWY, HOWE, TASM. Map 81. W.A.: Kings Park, G.J.Keighery 2388 (PERTH). S.A.: 1.6 km NW of Myopolonga, Murray R., G.Barker 5 (NSW). N.S.W.: 19 km W of Kerrabee, R.Coveny 9588 (NSW); 16 km NNE of Wentworth, T.B.Muir 5816 (NSW). Tas.: Cascade Gully, April 1914, A.H.S.Lucas (NSW).

Doubtful Name

Urtica lucifuga var. linearifolia J.D.Hook., London J. Bot. 6: 285 (1847) T: Hobart and Circular Head, Tas., R.Gunn s.n.; n.v. Possibly U. incisa.

2. LAPORTEA

Laportea Gaudich., Voy. Uranie 498 ‘1826’ (1830), nom. cons.; after M.Laporte, a naval officer on the Uranie during its voyage around the world, 1817–1820. Type: L. canadensis (L.) Gaudich. Urticastrum Heist. ex Fabric., Enum. 204 (1759), nom. rej. T: not designated. Herbs or shrubs with irritant hairs, monoecious, rarely dioecious. Leaves alternate, petiolate, simple, chartaceous, variously dentate; stipules intrapetiolar, partially connate, bifid. Inflorescence axillary, paniculate, pedunculate; flowers in loose glomerules; pedicels winged. Male flowers: tepals 4 or 5; stamens 4 or 5; filaments reflexed; pistillode small. Female flowers: tepals 4, the dorsiventral ones unequal, smaller than the laterals; staminodes absent; ovary ovoid; style linear, rarely brush-like. Achenes compressed, ovoid to hemispherical, sessile or stipitate, to 4 mm long, reflexed. A genus of 22 species centred in Africa and Madagascar; 3 species are pantropical, one of which occurs in mainland Australia, another on Cocos (Keeling) and Christmas Islands (Indian Ocean). The genus comprises two sections: sect. Laportea and sect. Fleurya (Gaudich.) Chew; only the latter occurs in Australia.

72 URTICACEAE 2. Laportea

W.-L.Chew, Laportea and Allied Genera (Urticaceae), Gard. Bull. Singapore 21: 198–201 (1965); A Monograph of Laportea (Urticaceae), Gard. Bull. Singapore 25: 111–178 (1969).

Sect. Fleurya

Laportea sect. Fleurya (Gaudich.) Chew, Gard. Bull. Singapore 21: 199 (1965) Fleurya Gaudich., Voy. Uranie 497 ‘1826’ (1830); Urtica d. Fleurya (Gaudich.) Endl.; Gen. Pl. 283 (1837). T: F. spicata Gaudich. = L. interrupta (L.) Chew, lecto, fide W.-L.Chew, op. cit. 200. Herbs. Pedicels of female flowers usually winged dorsiventrally and asymmetrically. Achene not articulated on pedicel.

1. Laportea interrupta (L.) Chew, Gard. Bull. Singapore 21: 200 (1965) Urtica interrupta L., Sp. Pl. 985 (1753); Fleurya interrupta (L.) Wight, Icon. Pl. 6: 10, 1975 (1853). T: Ceylon [Sri Lanka], Hermann; holo: BM. Illustration: W.-L.Chew, Gard. Bull. Singapore 25: 147, fig. 12 (1969). Monoecious annual herb to 2 m, woody at base. Leaves ovate, serrate, acuminate, rounded at base; lamina 8–12 cm long, 5–7 cm wide; lateral veins 3–6 pairs, the basal pair 2/3 length of lamina; petiole 5–8 cm long; stipules 3–5 mm long, glabrous. Inflorescence bisexual, to 30 cm long, sometimes at base of stem; side branches solitary, often suppressed; flowers fasciculate on elongated peduncle, pedicellate; tepals 4, 1–1.5 mm long. Male flowers: pedicel c. 1 mm long; tepals corniculate; pistillode 0.5 mm long. Female flowers sessile; ovary c. 0.75 mm long; stigma trifid, reflexed. Achenes c. 1.25 mm long, sessile. Fig. 35B–F. Occurs in northern N.T. and Cape York Peninsula, Qld; also throughout the Old World tropics. Grows as transient elements of disturbed habitats in partial shade. Regions: ARNM, CYRK. Map 82. N.T.: Kapalga, C.R.Dunlop & Taylor 6087 (DNA, NSW); Mt Howship, C.R.Dunlop 6635 (DNA). Qld: Iron Ra., H.Flecker 8673 (NSW, QRS); Barron R., Feb. 1914, E.W.Bick (NSW); N of Palmer R., J.R.Clarkson 4668 (BRI, QRS).

3. DENDROCNIDE

Dendrocnide Miq., Pl. Jungh. 29 (1851); from the Greek dendron (tree) and knide (nettle). Type: D. costata Miq. = D. stimulans (L.f.) Chew Laportea Gaudich., Voy. Uranie 498 ‘1826’ (1830) p.p. but not as to lectotype. [Urticastrum Heist. ex Fabric. sensu O.Kuntze, Rev. Gen. Pl. 634 (1891) p.p. but not as to lectotype] Shrubs or trees with irritant hairs, dioecious or monoecious. Leaves alternate, simple, coriaceous, crenulate, undulate to smooth, petiolate; stipules entirely connate, intrapetiolar, coriaceous. Inflorescence axillary, pedunculate, racemose, bracteate. Flowers free, in small fascicles or on flabellate receptacles. Male flowers: tepals 4, rarely 5; stamens 4, rarely 5; pistillode present. Female flowers sessile to pedicellate, flabellately or distichously arranged, or in loose fascicles; tepals 4; ovary ovoid, unilocular; stigmas usually ligulate; ovule erect; staminodes absent; pedicels simple or swollen. Achenes compressed or ellipsoidal to ovoid, not chartaceous, usually strongly warted. Stinging Trees. A genus of 37 species native to SE Asia, Australia and Pacific islands; 5 species in Australia. The genus comprises 2 sections; Australian species belong to sect. Sarcopus. W.-L.Chew, Laportea and Allied Genera (Urticaceae), Gard. Bull. Singapore 21: 201–207 (1965); W.-L.Chew, A Monograph of Dendrocnide (Urticaceae), Gard. Bull. Singapore 25: 1–104 (1969).

73

Figure 36. Dendrocnide. A–D, D. corallodesme. A, cauliflorous ♀inflorescence ×0.8; B, ♂flower ×16; C, achene ×8; D, leaf ×0.6. E–G, D. moroides. E, flowering and fruiting branchlet ×0.5; F, achene ×13; G, ♀flower ×20. Reproduced by permission from Garden’s Bulletin, Singapore 25: 54, fig. 21, 57, fig. 22 (1969).

74 URTICACEAE 3. Dendrocnide

1 Leaves glabrous or almost so, the irritant hairs absent or confined to veins on underside; leaf base cuneate to rounded 2 Lamina narrowly elliptic, strongly cuneate at base, long acuminate to attenuate 1. D. corallodesme 2: Lamina elliptic to ovate, broadly cuneate to rounded at base, shortly acuminate 4. D. photinophylla 1: Leaves hairy, with irritant hairs; leaf base cordate to cordate-peltate 3 Lamina densely pubescent on both sides; base cordate-peltate; inflorescence bisexual 2. D. moroides 3: Lamina usually densely pubescent on underside, less so on upperside; base cordate, not peltate; inflorescence unisexual 4 Lamina densely woolly on underside; base deeply cordate, the two lobes often overlapping. 5. D. excelsa 4: Lamina not densely woolly beneath; base broadly cordate, the lobes not overlapping 3. D. cordata

Sect. Sarcopus

Dendrocnide sect. Sarcopus (Wedd.) Chew, Gard. Bull. Singapore 25: 7 (1969). Laportea § Sarcopus Wedd., Arch. Mus. Hist. Nat. 9: 129 (1856); Laportea sect. Sarcopus (Wedd.) Wedd. in DC., Prodr. 16(1): 82 (1869). Type: Laportea gigas Wedd. = D. excelsa (Wedd.) Chew Female flowers sessile or pedicellate, borne freely in loose fascicles or distichously at ends of peduncles; pedicel often fleshy. A section of 27 species in E Malesia, Australia and the South Pacific.

1. Dendrocnide corallodesme (Lauterb.) Chew, Gard. Bull. Singapore 21: 202 (1965) Laportea corallodesme Lauterb. in K.Schumann & C.A.G.Lauterbach, Nachtr. Fl. Schutzgeb. 252 (1905). T: Madang District, New Guinea, Jan. 1902, R.Schlechter 13840; holo: WRSL. Illustration: W.-L.Chew, Gard. Bull. Singapore 25: 54, fig. 21 (1969). Dioecious tree to 6 m; irritant hairs mainly on inflorescence and abaxial leaf midrib. Leaves elliptic, entire, acuminate, narrowly cuneate at base; lamina 12–20 cm long, 4–7 cm wide, glabrous except abaxial midrib; lateral veins 13–17 pairs, short, closely set; petiole 2–4 cm long; stipules 1–1.5 cm long. Inflorescence ramiflorous, unisexual, branched racemes, paired; peduncle pubescent with short irritant hairs; pedicels 1–1.5 mm long. Tepals 4, with irritant hairs. Male flowers: tepals keeled; pistillode small. Female flowers: ovary c. 1.5 mm long, 0.8 mm wide; stigma to 3 mm long. Achenes ellipsoidal to obovoid, 3 mm long, 2.5 mm wide, often warted; perianth persisting as small cupule at base. Mango-leafed Stinger. Fig. 36A–D. Occurs in the Iron Ra. and McIlwraith Ra. areas, Qld, along creeks in rainforest; also in New Guinea. Region: CYRK. Map 83. Qld: between Iron Ra. & Nundah, Mosquito Ck, Cape York Peninsula, B.Hyland 2921 (NSW, QRS); Claudie R., B.Hyland 3613 (QRS); Chester R., B.Hyland 3533 (QRS); Chester R., McIlwraith Ra., J.R.Clarkson 2439 (BRI). Qld plants differ from those of New Guinea in having irritant hairs on the abaxial midrib. The species can cause severe irritation to the skin on contact.

75 3. Dendrocnide URTICACEAE

2. (Wedd.) Chew, Gard. Bull. Singapore 21: 204 (1965) Laportea moroides Wedd., Arch. Mus. Hist. Nat. 9: 142 (1856); Urticastrum moroides (Wedd.) Kuntze, Revis. Gen. Pl. 2: 635 (1891) as morodes. T: Endeavour R., [Qld], A.Cunningham s.n.; holo: G. Illustrations: W.-L.Chew, Gard. Bull. Singapore 25: 57, fig. 22 (1969); K.A.W.Williams, Native Pl. Queensland 1: 91 (1979). Monoecious shrub or tree to 10 m, densely pubescent. Leaves peltate, broadly ovate, dentate, acuminate, cordate at base; lamina 12–22 cm long, 11–18 cm wide, densely pubescent; lateral veins 6–8 pairs, the basal pair prominent; petiole 10–18 cm long; stipules c. 1 cm long. Inflorescence a bisexual, branched raceme, to 15 cm long and 8 cm wide, often paired. Pedicels to 1.5 mm long. Tepals 4. Male flowers: pistillode c. 0.75 mm long. Female flowers: perianth c. 0.75 mm long; ovary c. 0.5 mm long; stigma c. 2 mm long. Achenes ovoid, c. 2 mm long, 1.5 mm wide, warted, enclosed by lateral tepals. Gympie, Mulberry-leaved Stinger Figs 12, 36E–G. Occurs in eastern Qld and N.S.W. growing in rainforest; also in the Moluccas and Indonesia. Regions: CYRK, BURD, DWSN, MCPH. Map 84. Qld: near Atherton, E.Volck & S.Dansie 1471 (BRI, NSW); Wongabel, L.S.Smith 10175 (BRI, NSW); Mowbray R., L.J.Brass 1981 (A, B, BRI). N.S.W.: Nimbiro, near Lismore, Sept. 1925, E.Cheel (NSW); Tweed R., W.Bäuerlen, NSW 77926 (NSW). Perhaps the most virulent stinger in the genus and much feared by susceptible foresters in the field. The species is unique in the genus in having bisexual inflorescences in which the few male flowers are surrounded by female flowers in each floral fascicle.

3. Dendrocnide cordata (Warb. ex H.J.Winkler) Chew, Gard. Bull. Singapore 21: 202 (1965) Laportea cordata Warb. ex H.J.Winkler, Bot. Jahrb. Syst. 57: 503 (1922). T: Sepik River, New Guinea, K.Hollrung 513; iso: BO, K, MEL. Laportea cordifolia L.S.Smith, Proc. Roy. Soc. Queensland 70: 31 (1959). T: c. 4 miles [6.4 km] SW of Atherton, Qld, 4 Sept 1957, L.S.Smith 10132; holo: BRI. Illustration: W.-L.Chew, Gard. Bull. Singapore 25: 59, fig. 23 (1969). Monoecious shrub or tree to 10 m, puberulous. Leaves very broadly ovate, irregularly dentate to crenate-sinuate, acuminate, cordate at base; lamina 18–25 cm long, 16–22 cm wide, pubescent becoming puberulous with irritant hairs; lateral veins 8–10 pairs, two basal pairs prominent; petiole 18–22 cm long; stipules 1.5–2 cm long. Inflorescence a unisexual, loosely branched raceme to 20 cm long, 10 cm wide, paired. Tepals 4. Male flowers: pedicel less than 1 mm long; pistillode c. 0.5 mm long. Female flowers: pedicel 1–1.5 mm long; perianth c. 1 mm long, 1 tepal larger than others; ovary to 1 mm long; stigma to 2 mm long. Achenes c. 2 mm long, 1.5 mm wide, bloated, warted, reflexed, covered by lateral tepals. Stinger. Fig. 37E–F. Occurs on Cape York Peninsula, Qld; grows in rainforest; also in the Moluccas, New Guinea and the Bismarck Archipelago. Region: CYRK. Map 85. Qld: near Atherton, E.Volck & S.Dansie 1470 (BRI, NSW); State Forest Reserve 194, G.C.Stocker 1796 (QRS). Easily distinguished from D. moroides through most of its geographical range, but in Australia is sometimes as hairy and virulent as that species, the 2 species then being difficult to separate.

4. Dendrocnide photinophylla (Kunth) Chew, Gard. Bull. Singapore 21: 205 (1965) Fleurya photinophylla Kunth, Ann. Sci. Nat. Bot. ser. 3, 7: 183 (1847); Laportea photinophylla (Kunth) Wedd., Arch. Mus. Hist. Nat. 9: 138 (1856); Urticastrum photinophyllum (Kunth) Kuntze, Revis. Gen. Pl. 2: 635 (1891). T: cult. Göttingen, Germany; holo: B, destroyed; Nudgee, Qld, C.T.White 5600; neo: A; isoneo: BRI, NY fide W.-L.Chew, Gard. Bull. Singapore 25: 62 (1969). Urera leichardiana Wedd., Ann. Sci. Nat. Bot. ser. 4, 1: 178 (1854). T: The Creek, Moreton District, Qld,

76

Figure 37. Dendrocnide. A–D, D. photinophylla. A, juvenile leaf ×0.5; B, flowering and fruiting branchlet ×0.5; C, achene ×8; D, ♀flower ×16. E–F, D. cordata. E, flowering branchlet ×0.3; F, ♂inflorescence ×0.3. Reproduced by permission from Garden’s Bulletin, Singapore 25: 59, fig. 23, 61, fig. 24 (1969).

77 3. Dendrocnide URTICACEAE

L.Leichhardt 26; holo: P. Illustrations: W.-L.Chew, Gard. Bull. Singapore 25: 61, fig. 24 (1969); W.D.Francis, Austral. Rainforest Trees 3rd edn, 88, 89 (1970). Dioecious tree to 25 m, puberulous, glabrescent; irritant hairs sparse. Leaves elliptic to ovate, entire, rarely dentate, obtusely acuminate, cuneate to rounded at base; lamina 8–12 cm long, 4–6 cm wide; lateral veins 6–8 pairs, the basal pair prominent; petiole 3–7 cm long; stipules c. 1 cm long. Inflorescence a unisexual, branched raceme, 6 mm long, solitary. Pedicels c. 1.5 mm long. Tepals 4. Male flowers: pedicel fleshy at anthesis; pistillode small. Female flowers: lateral tepals larger than dorsiventrals; stigma c. 1.5 mm long. Achenes 2 mm long, sigmoid, smooth, half covered by persistent tepals, reflexed over enlarged pedicel. Shining-leafed Stinger, Shiny leaf Stinging Tree. Fig. 37A–D. Occurs from Cape York, Qld, S to the Nepean region, N.S.W.; grows in dense rainforest, sometimes in deep sand. Regions: CYRK, BURD, DWSN, MCPH, NEPN. Map 86. Qld: McIvor R., B.P.M.Hyland 6266 (QRS); Mt Tamborine, C.T.White 12726 (BO, BRI, K). N.S.W.: Walcna, Crawford 16 (MEL); Tooloom Ra., Apr. 1953, J.W.Vickery s.n. (NSW).

5. (Wedd.) Chew, Gard. Bull. Singapore 21: 203 (1965) Urera excelsa Wedd., Ann. Sci. Nat. Bot. ser. 4, 1: 178 (1854). T: Australia, L.Leichhardt 17; holo: P. Urera rotundifolia Wedd., Ann. Sci. Nat. Bot. ser. 4, 1: 177 (1854). T: Australia, collector unknown; n.v. Laportea gigas Wedd., Arch. Mus. Hist. Nat. 9: 129 (1856), nom. illeg.; Urticastrum gigas (Wedd.) Kuntze, Revis. Gen. Pl. 2: 635 (1891), nom. illeg. T: ‘Five Islands Country’, N.S.W., A.Cunningham; syn: K. Illustrations: H.A.Weddell, Arch. Mus. Hist. Nat. Paris 9: t. 3 (1856) as Laportea gigas; W.-L.Chew, Gard. Bull. Singapore 25: 65, fig. 26 (1969); W.D.Francis, Austral. Rainforest Trees 3rd edn, 84, 85 (1970). Dioecious buttressed tree to 35 m, with irritant hairs. Leaves broadly ovate, broadly crenate or sinuate to entire, bluntly acuminate, cordate at base; lamina 12–22 cm long, 10–12 cm wide, juveniles considerably larger, pubescent above, villous below; lateral veins 6–8 pairs, the basal pair prominent; petiole 5–10 cm long, pubescent to villous; stipules 1–2 cm long. Inflorescences unisexual, to 12 cm long and 6 cm wide, usually paired. Flowers sessile or on pedicels to 1.6 mm long. Tepals 4. Male flowers: pistillode c. 0.4 mm long. Female flowers densely pubescent; stigma c. 1 mm long. Achenes ovoid, c. 2 mm long, 1.5 mm wide, tuberculate, on fleshy pedicels. Giant Stinger, Giant Stinging Tree. Fig. 38. Occurs from the Bunya Mtns, Qld, S to Kiama, N.S.W. Common in rainforest especially on slopes and in gullies, often in basaltic soil. Regions: DWSN, MCPH, NEPN, HOWE. Map 87. Qld: Bunya Mtns, C.T.White 9145 (A, BRI, NY); Mt Glorious, L.S.Smith 10540 (BRI); Imbil, Sept. 1922, Petrie (BRI). N.S.W.: Cairds Gap, Liverpool Ra., Oct. 1954, L.Johnson & E.Constable (NSW); Kiama, R.Pullen 4065 (L, NSW). Trees in excess of 35 metres with buttresses to over 10 metres high have been recorded in rainforest in Qld.

Trib. 2. ELATOSTEMEAE

Urticaceae trib. Elatostemeae Gaudich., Voy. Uranie 493 ‘1826’ (1830). Type: Elatostema Forster & G.Forster, nom. cons. Urticaceae trib. Lecantheae Wedd., Ann. Sci. Nat. Bot. ser. 4, 1: 174 (1854). T: Lecanthus Wedd. Urticaceae trib. Procrideae Wedd., Arch. Mus. Hist. Nat. 9: 170 (1856); Urticeae subtrib. Procrideae Benth., Fl. Austral. 6: 182 (1873). T: Procris Comm. ex A.L.Juss. Monoecious or dioecious herbs or shrubs, sometimes epiphytic, without irritant hairs. Leaves opposite-decussate, opposite with one reduced, or alternate, distichous; cystoliths elongate. Flowers unisexual. Male flowers: tepals 4 or 5; stamens 4 or 5; pistillode present, obconical, small to minute. Female flowers: tepals 3–5; staminodes present.

78

Figure 38. Dendrocnide excelsa. A, ♀flower ×10; B, achene ×10; C, flowering and fruiting branchlet ×0.7. Reproduced by permission from Garden’s Bulletin, Singapore 25: 65, fig. 26 (1969).

79 Trib. 2. Elatostemeae URTICACEAE

A tribe of 6 genera, widespread in tropical and subtropical regions, especially in the mountains; 2 genera in Australia.

4. ELATOSTEMA

Elatostema Forster & G.Forster, Char. Gen. Pl. 105 (1775), nom. cons.; from the Greek elatos (elastic) and stema (stamen), referring to the stamens being inflexed and then reflexing. [Elatostemma Endl., Prodr. Fl. Norfolk. 39 (1833), orth. var.]. Type: E. sessile Forster & G.Forster, typ. cons. Monoecious or dioecious herbs, rarely subshrubs, without irritant hairs. Leaves shortly petiolate, distichous, alternate or opposite, with one of the pair alternately greatly reduced; lamina often asymmetrical, dentate, sinuate to subentire; stipules intrapetiolar. Inflorescence unisexual, cymose-capitate; flowers in open, discoid or concave often fleshy receptacles surrounded by an involucre of bracts, or in dense axillary cymes. Male flowers: tepals 4 or 5; stamens 4 or 5; pistillode small to minute. Female flowers: tepals 3 or 5; stigma subsessile, brush-like; staminodes often present. Achenes ovoid, compressed, often warted. A genus of c. 250 species widespread in the Old World tropics, especially in moist lower- and mid-montane regions; 2 species in Australia, 1 also on Lord Howe Is. The extended genus comprises 4 subgenera: subg. Elatostema, subg. Pelliona, subg. Elatostematoides and subg. Weddellia; all the Australian species belong to subg. Elatostema. H.Schroeter & H.J.Winkler, Monographie der Gattung Elatostema sens. lat., Feddes Repert. Beih. 83(1): 1–55 (1935); 83(2): 1–174 (1936). Leaves more than 10 cm long, sharply dentate; female head subsessile or on short thick peduncle 1. E. reticulatum Leaves less than 10 cm along, the marginal teeth rounded; female head on long, thin peduncle 2. E. stipitatum

1. Elatostema reticulatum Wedd., Ann. Sci. Nat. Bot. ser. 4, 1: 188 (1854) T: Australia, L.Leichhardt; n.v. E. reticulatum. var. pubescenti-hirta Benth., Fl. Austral. 6: 183 (1873). T: Upper Hunter, Kiri Brush, N.S.W., Jan. 1843, L.Leichhardt s.n.; iso: NSW. E. reticulatum var. sessile Benth., Fl. Austral. 6: 184 (1873). T: Macleay R., N.S.W., H.Beckler; n.v. E. reticulatum var. minus Domin, Biblioth. Bot. 89: 28 (1921). T: Tamborine Mtn, Qld, Mar. 1910, K.Domin; n.v. [E. reticulatum var. glabrum Domin, Biblioth. Bot. 89: 20 (1921), nom. inval. based on type of species] Straggly herb to 1 m, rarely scandent, often with adventitious roots, sparsely pubescent, soon glabrescent. Leaves distichous, asymmetrically elliptic to obovoid, curved, sharply dentate, acuminate, asymmetrically cuneate and auriculate at base; lamina 10–20 cm long, 4–9 cm wide, glabrescent, dense with elongated cystoliths; lateral veins 8 or 9 pairs, prominent; petiole short or absent. Inflorescence unisexual, capitate, 1–2 cm diam., pedunculate to subsessile. Male inflorescence: peduncle c. 5 cm long; flowers c. 1.5 mm long; tepals 4; stamens 4; pistillode minute. Female inflorescence: peduncle c. 3 mm long; flowers 1.5 mm long; tepals 3, unequal; staminodes 3, reflexed. Achenes ellipsoidal, 1–2 mm long, 1 mm wide. Rainforest Spinach. Figs 13, 39A. Occurs from Mt Bartle Frere, Qld, S to Clyde Mtn, N.S.W.; grows on coastal mountains in wet sites in rainforest. Regions: CYRK, BURD, DWSN, MCPH, NEPN, HOWE. Map 88. Qld: North Mary Logging Area, State Forest Reserve 143, B.Hyland 7924 (NSW, QRS); Blackall Ra., Dec. 1916, C.T.White s.n. (NSW). N.S.W.: Night Cap Rd., R.Melville 3328 (NSW); Macquarie Pass, M.Gray 5886 (NSW); Doyles River State Forest, E.F.Constable 6310 (NSW).

80

Figure 39. A, Elatostema reticulatum, habit ×0.5 (BRI 305758, BRI). B, , habit ×0.5 (BRI 339988, BRI). Drawn by D.Mackay.

81 4. Elatostema URTICACEAE

As all manner of gradations have been seen in the collections studied, the varieties distinguished by Bentham (1873) and Domin (1921) are not maintained.

2. Elatostema stipitatum Wedd., Ann. Sci. Nat. Bot. ser. 4, 1: 190 (1854) T: Australia, J.P.Verreaux 145; n.v. Prostrate herb, usually strigose. Leaves distichous, asymmetrically obovate, curved, acute to shortly acuminate, dentate, the teeth often rounded; base asymmetrical, strongly auriculate; lamina 4–8 cm long, 2–2.5 cm wide, strigose, especially on veins on lowerside, dense with elongated cystoliths; lateral veins 4–6 pairs, not prominent; petiole often absent. Inflorescence unisexual, capitate, c. 1 cm diam., pedunculate. Male inflorescence: peduncle c. 5 cm long; flowers c. 1.5 mm long; tepals 3 or 4; stamens 3 or 4; pistillode minute. Female inflorescence: peduncle less than 1.5 cm long; flowers c. 1.5 mm long; tepals 3; staminodes minute. Achenes ellipsoidal, 1–2 mm long, 1 mm wide. Occurs from Maryborough, Qld, S to Mt Boss, N.S.W.; grows on coastal mountains in damp sites in rainforest, where it often forms continuous mats, sometimes as lithophytes. Region: MCPH. Map 89. Qld: Beechmont Ra., 0.5 km WSW of Binna Burra, I.R.Telford 9725 (NSW); Mt Roberts, McPherson Ra., L.A.S.Johnson 48 (NSW). N.S.W.: Mt Lindesay, E.F.Constable 10544 (NSW); Mt Boss, 43 km NW of Wauchope, R.Coveny 6010 et al. (NSW); Dorrigo, Jan. 1918, J.B.Cleland s.n. (NSW).

Excluded species

Elatostema lineolatum Wight, Icon. 1984 (1853).

Elatostema lineolatum var. majus Thwaites ex Wedd., in A.P. de Candolle, Prodr. 16(1): 182 (1869). Mentioned incidentally by T.S.Liu & W.D.Huang, Fl. 2: 181 (1976), that this species and variety occur in Australia but without citation of collections.

Elatostema sessile Forster & G.Forster var. brongniartianum (Wedd.) Wedd., in A.P. de Candolle, Prodr. 16: 173 (1869); E. brongniartianum Wedd., Ann. Sci. Nat. ser. 4, 1: 190 (1854). This Malesian species was claimed by Weddell to have been recorded for Australia by F.Mueller. No collections have been seen.

5. PROCRIS

Procris Comm. ex A.L.Juss, Gen. 403 (1789). Type: P. axillaris J.Gmelin, Syst. Nat. 2: 267 (1791). Monoecious or dioecious perennial subshrubs, sometimes epiphytic, without irritant hairs. Leaves alternate or opposite with one of pair small to minute, petiolate, simple, coriaceous, often asymmetrical, curved, entire, to crenulate-dentate in distal half; stipules small, often caducous. Inflorescence unisexual. Male inflorescence divaricate, cymose; flowers in small receptacles. Male flowers: tepals 4 or 5; pistillode present. Female inflorescence capitulate, fleshy, globose, subsessile to sessile at nodes, often solitary. Female flowers: tepals 3 or 4; stigma subsessile to sessile, shortly ligulate. Achene ovoid, often exserted from perianth. A genus of c. 16 species widespread in the Old World tropics; 1 species in Australia, also on Norfolk Is. H.Schroeter, Monographie der Gattung Procris. I, Feddes Repert. 45: 179–192 (1938); II, op. cit. 257–300 (1938).

82

Figure 40. Procris pedunculata, habit ×0.75 (QRS 054401, QRS). Drawn by D.Mackay.

83 5. Procris URTICACEAE

1. Procris pedunculata (Forster & G.Forster) Wedd. in A.P. de Candolle, Prodr. 16(1): 191 (1869) Elatostema pedunculatum Forster & G.Forster,. Char. Gen. Pl. 106, t. 53 (1776). T: not designated. Monoecious shrub, sometimes epiphytic, glabrous. Leaves opposite but one of pair minute, narrowly obovate, asymmetrical, acuminate, cuneate at base, shallowly crenulate in distal half; lamina 8–15 cm long, 2.5–4 cm wide, with crowded elongated cystoliths; lateral veins 6–11 pairs, faint; petiole to 1 cm long. Male inflorescence cymose, few-flowered; peduncle to 3 cm long; flowers pedicellate; tepals 5. Female inflorescence capitulate, sessile, many-flowered; flowers subsessile to sessile, 1 mm long; stigma brush-like, short. Achene ovoid, warted, 1.5 mm long. Fig. 40. Occurs on Cape York Peninsula, Qld, in rainforest; also from the Mascarenes through Malesia to the Pacific. Region: CYRK. Map 90. Qld: State Forest Reserve 755, Barong Logging Area, B.Hyland 8360 (QRS). A collection from North Johnstone Logging Area, near Atherton, Qld, B.J.Wallace 83110 (NSW), is doubtfully referred to this species. The species has been considered to comprise 3 varieties (H.Schroeter, op. cit.); only the typical variant occurs in Australia. Because the species varies considerably throughout its range in Malesia and Melanesia, the varieties are not maintained here.

Trib. 3. BOEHMERIEAE

Urticaceae trib. Boehmerieae Gaudich., Voy. Uranie 499 ‘1826’ (1830). Type: Boehmeria Jacq. Trees, shrubs, herbs or climbers without irritant hairs. Leaves opposite-decussate or alternate; cystoliths mainly punctiform. Flowers unisexual. Male flowers: tepals 4 or 5; stamens 4 or 5; pistillode small. Female flowers: perianth tubular, with contracted 2–4-toothed orifice; staminodes absent. A tribe of 8 genera, throughout the tropics; 4 genera in Australia.

6. BOEHMERIA

Boehmeria Jacq., Enum. Syst. Pl. 9 (1760); after G.R.Boehmer (1723–1803), Professor of Botany, Wittenberg, Germany. Type: B. ramiflora Jacq. Monoecious or dioecious shrubs or trees, often pubescent. Leaves opposite, those of each pair often unequal, or alternate, petiolate, simple, chartaceous, dentate, the lamina often asymmetrical, trinerved; stipules lateral, free, rarely connate. Inflorescences at nodes, spike-like or cymose; flowers in globular heads along rachis of inflorescence, unisexual. Male flowers: tepals 4; pistillode present. Female flowers: perianth tubular, the orifice often contracted, 2–4-dentate; ovary enclosed by perianth, sometimes partially adnate with it; stigma filiform, often unilaterally stigmatic. Achenes enclosed by often winged perianth. A genus of over 100 species widespread throughout the tropics; 2 taxa in mainland Australia, 1 native, 1 introduced; 1 species endemic on Lord Howe Is., 1 species endemic on Norfolk Is. Leaves opposite; lamina strigose below; inflorescence spicate, often pendulous 1. B. macrophylla Leaves alternate; lamina densely white woolly below; inflorescence a divaricate raceme 2. B. nivea

84 URTICACEAE 6. Boehmeria

1. Boehmeria macrophylla Hornem., Hort. Bot. Hafn. 2: 890 (1815); I.Friis & W.Marais, Kew Bull. 37: 164 (1982), non D.Don (1825). T: cult. Copenhagen Bot. Gard., Denmark; holo: C. B. platyphylla D.Don, Prodr. Fl. Nepal. 60 (1825). T: Narainhetty, Nepal, Hamilton s.n.; holo: BM n.v., fide W.T.Stearn, J. Arnold Arbor. 26: 168 (1945). B. macrostachya (Wight) Bailey, Synop. Queensland Fl. Suppl. 2: 55 (1888), nom. illeg.; Splitgerbera macrostachya Wight, Icon. Pl. Ind. Orient. t. 1977 (1853). T: not designated. B. platyphylla var. austroqueenslandica Domin, Biblioth. Bot. 89: 22 (1921). T: Tamborine & Mts [Beechmont], Qld, Mar. 1910, K.Domin; n.v. Illustration: K.A.W.Williams, Native Pl. Queensland 3: 43 (1987) as B. platyphylla var. austro- queenslandica. Monoecious shrub to 2 m, strigose throughout. Leaves opposite, one of each pair smaller, ovate, serrate, acuminate to attenuate, broadly cuneate at base; lamina 12–22 cm long, 5–7 cm wide; lateral veins 4 or 5 pairs, the basal pair often reaching nearly to apex, the others short; petiole c. 5 cm long; stipules 6–8 mm long. Inflorescence a unisexual or bisexual spike to 25 cm long, pendulous, single or paired; flowers spirally glomerulate, bracteate. Male flowers mainly at base of spike, subsessile, c. 1 mm long. Female flowers sessile, 1–2 mm long; orifice 2-dentate, attenuate; stigma c. 2 mm long. Achenes ovoid, slightly compressed, 2 mm long. Figs 16, 39B. Occurs in the mountains of south-eastern Qld and north-eastern N.S.W., particularly on the ranges of the Tweed caldera; grows along creeks in rainforest. Also from Africa through Asia to Pacific islands. Region: MCPH. Map 91. Qld: Mt Roberts, S.T.Blake 21512 (BRI, NSW); Curtis Fall, Jan. 1916, C.T.White s.n. (BRI); Mt Clunie, R.Melville 3628 (NSW). N.S.W.: Mt Gipps, Wiangarie State Forest, R.Coveny 10570 (NSW); Mt Warning, SW of Murwillumbah, S.Clarke 1583 et al. (NSW). Boehmeria macrophylla is found widely from Africa through Asia to the Pacific. Highly variable; numerous varieties have been established for the species (H.A.Weddell in A.P. de Candolle, Prodr. 16(1): 210 (1869).

2. *Boehmeria nivea (L.) Gaudich., Voy. Uranie 499 ‘1826’ (1830) Urtica nivea L., Sp. Pl. 985 (1753). T: China, collector unknown; n.v. Illustrations: R.Wight, Icon. 2: t. 688 (1843) as Urtica tenacissima Roxb.; W.J.Hooker, Hooker’s J. Bot. Kew Gard. Misc. 3: t. viii (1851); H.A.Weddell, Arch. Mus. Hist. Nat. 9: t. xi, fig. 10–17 (1857). C.L.Blume, Mus. Bot. 2: t. 56 (1856), as Boehmeria tenacissima (Roxb.) Blume. Monoecious shrub, densely rough-strigose throughout. Leaves alternate, ovate; lamina 15–25 cm long, 5–8 cm wide, the underside densely white-woolly. Inflorescence of divaricate racemes, paired to numerous at each axil. Ramie. This species, native of South and SE Asia, produces the Ramie of commerce. Reported by K.Domin (loc. cit.) to have been seen in the wild. It is likely that such populations are escapes from cultivation and doubtfully naturalised. No collections from the wild have been seen. Region: CYRK. Map 92.

7. POUZOLZIA

Pouzolzia Gaudich., Voy. Uranie 503 ‘1826’ (1830); after P.M.C. de Pouzolz (1785–1858), botanist of Nîmes, . Type: P. laevigata (Poiret) Decne. Monoecious, rarely dioecious herbs or shrubs, finely pubescent, without irritant hairs. Leaves usually alternate and opposite, simple, chartaceous, the upper leaves grading into bracts, petiolate, entire, often 3-nerved; stipules free. Inflorescence small axillary clusters, sometimes spike-like when leaves not developed. Male flowers: tepals 3–5; stamens 3–5; pistillode

85 7. Pouzolzia URTICACEAE present. Female flowers: perianth tubular; orifice contracted, 2–4-dentate; ovary enclosed by perianth; stigma filiform, unilaterally stigmatic. Achene enclosed by, but free from, persistent perianth. A genus of c. 70 species widely dispersed in the tropics; 2 species in Australia. There are 2 sections: sect. Pouzolzia and sect. Memorialis Bennett; 1 species of each section in Australia. Though it has been maintained that the sections perhaps deserve generic status, they are most difficult to separate satisfactorily. This group of highly variable species is in urgent need of a monographic treatment. Diffuse herb; stems not woody; leaves 3-nerved at base, the veins branched near apex; male perianth not inflexed, appearing globular; perianth enclosing achene not winged 1. P. zeylanica Erect herb; stems elongated, woody at base; leaves 3-nerved to apex of leaf; male perianth strongly inflexed, concave at apex; perianth enclosing achene strongly winged 2. P. hirta

Sect. 1. Pouzolzia

Pouzolzia Gaudich. sect. Pouzolzia. Leaves entire or serrate-dentate, 3-nerved at base, the lateral veins branched. Male tepals convex or gibbous at the back.

1. Pouzolzia zeylanica (L.) Bennett, Pl. Jav. Rar. 67 (1838) Parietaria zeylanica L., Sp. Pl. 1052 (1753). T: Ceylon [Sri Lanka], P.Hermann; holo: BM. Parietaria indica L., Mant. Pl. 1: 128 (1767); Pouzolzia indica (L.) Wight, Icon. 6: 43 (1853). T: Asia, collector unknown; n.v. Urtica alienata L., Syst. Nat. 13th edn, 2: 622 (1770), nom. superfl. (based on same type as Parietaria zeylanica L.); Pouzolzia indica var. alienata (L.) Wedd., Arch. Mus. Hist. Nat. Paris 8: 398 (1856). Pouzolzia arnhemica F.Muell., Fragm. 4: 87 (1864). T: Victoria R., N.T., F.Mueller & Wilson; n.v. Illustration: R.Wight, Icones Indiae Plantarum Orientalis 6: t. 1980, fig. 1; t. 2100, fig. 40 (1853) as P. indica (L.) Wight. Diffuse, monoecious perennial herb. Stems not woody, pubescent to strigose. Basal leaves opposite, upper ones alternate, ovate to broadly ovate, entire, acute to obtuse, rounded to broadly cuneate at base; lamina 10–17 mm long, 5–10 mm wide, 3-nerved at base, with 1 or 2 laterals arising near apex; petiole short; stipules free, persistent. Inflorescence bisexual, of few-flowered, sessile cymules. Male flowers pedicellate; tepals 4; pistillode small. Female flowers subsessile to sessile; perianth tubular with prominent ribs, the orifice 4- or 5-dentate; stigma caducous. Achene ovoid, black, shiny, enlosed by thickened perianth. Fig. 41A. Occurs commonly in the Kimberley, W.A., Arnhem Land, N.T., and on Cape York Peninsula, Qld. Widespread in India and SE Asia. Regions: KIMB, ARNM, CYRK. Map 93. W.A.: Devils Pass, Napier Ra., W.V.Fitzgerald 615 (PERTH); May R., W Kimberley, May 1905, W.V.Fitzgerald s.n. (NSW); King Sound, 1888, Frogatt s.n. (NSW). Qld: Saibai Is., J.R.Clarkson 3880 (QRS).

Sect. 2. Memorialis

Pouzolzia sect. Memorialis Bennett, Pl. Jav. Rar. 65 (1838). Memorialis (Bennett) Ham. ex Wedd., Arch. Mus. Hist. Nat. 9: 415 (1856). T: not designated. Leaves entire, 3-nerved usually to leaf apex; upper leaves grading into bracts. Male tepals inflexed at middle, the flowers appearing truncated, often with filiform processes.

86

Figure 41. A, Pouzolzia zeylanica, habit ×0.75 (BRI 036641, BRI). B–C, Pipturus argenteus. B, habit (♂) ×0.5 (QRS 042132, QRS); C, ♀inflorescence ×0.5 (QRS 042128, QRS). Drawn by D.Mackay.

87 7. Pouzolzia URTICACEAE

2. Pouzolzia hirta (Blume) Hassk., Cat. Hort. Bogor. 80 (1844) Urtica hirta Blume, Bijdr. 495 (1826); Gonostegia hirta (Blume) Miq., Ann. Mus. Bot. Lugduno-Batavum 4: 303 (1869); Hyrtanandra hirta (Blume) Miq., Pl. Jungh. 1: 26 (1851); Memorialis hirta (Blume) Wedd. in A.P. de Candolle, Prodr. 16(1): 235*6 (1869); [Pouzolzia hirta var. typica Domin, Biblioth. Bot. 89: 22 (1921) nom. inval.] T: Java, Indonesia, Junghuhn (?); n.v. Pouzolzia quinquenervis Bennett, Pl. Jav. Rar. 66 (1838); Pouzolzia hirta var. quinquenervis (Bennett) Domin, Biblioth. Bot. 89: 23 (1921). T: Nepal, Apr. 1808, Hamilton, Wallich Cat. 4601A, p.p. ex Bhotgong; syn: K. Hyrtanandra lythroides F.Muell., Fragm. 5: 194 (1866); Pouzolzia hirta var. lythroides (F.Muell.) Domin, Biblioth. Bot. 89: 23 (1921). T: Lynds R., Qld, L.Leichhardt; n.v. Erect monoecious herb. Stems elongated, woody at base, ribbed, lightly strigose on ribs. Leaves opposite, ovate, entire, acute, rounded to truncate at base; lamina c. 6 cm long and 1.5 cm wide, prominently 3-nerved to apex, lightly strigose, often sessile; stipules free, ovate, 1 mm long, persistent. Inflorescence bisexual, 3–7-flowered. Male flowers subsessile; tepals 5, strongly inflexed. Female flowers subsessile to sessile; perianth enclosing ovary. Achene black, shiny, enclosed by broadly winged perianth. Occurs in the Lynd R. and Mackay areas, Qld; grows in disturbed habitats. Widespread in India and SE Asia. Regions: CYRK, BURD. Map 94. Qld: Port Mackay, A.Dietrich 2487 (NSW).

8. PIPTURUS

Pipturus Wedd., Ann. Sci. Nat. Bot. ser. 4, 1: 196 (1854); from the Greek pipto (to fall) and oura (tail), alluding to the deciduous stigma. Type: P. velutinus (Decne.) Wedd., lecto, fide C.Skottsberg, Acta Horti Gothob. 8: 117 (1933). Dioecious, rarely monoecious, shrubs or soft-wooded trees. Leaves alternate, simple, coriaceous, serrulate to dentate, trinerved, closely villous beneath, petiolate; stipules connate, intrapetiolar, bifid. Inflorescence unisexual; flowers borne in globular heads sessile at nodes or alternating on interrupted spikes. Male flowers: tepals 4 or 5; pistillode present, densely villous. Female flowers: perianth tubular, the orifice contracted, 2–4-dentate; ovary enclosed by perianth; stigma filiform, unilaterally stigmatic. Achene adnate to enclosing fleshy perianth. A genus of c. 40 species in the Old World Tropics; 1 species in Australia. C.Skottsberg, Acta Horti Gothob. 7: 43–63 (1932).

Pipturus argenteus (G.Forster) Wedd. in A.P. de Candolle, Prodr. 16(1): 235*19 (1869) Urtica argentea G.Forster, Prodr. 65 (1786). T: Society Islands, G.Forster; n.v. Boehmeria propinqua Decne., Herb. Timor. 163 (1834); Pipturus propinquus (Decne.) Wedd., Ann. Sci. Nat. ser. 4, 1: 196 (1854). T: Timor, collector unknown; n.v. P. argenteus var. calcicola Domin, Biblioth. Bot. 89: 577 (1921). T: Chillagoe, Qld, Feb. 1910, K.Domin; n.v. Illustration: K.A.W.Williams, Native Pl. Queensland 2: 225 (1984). Dioecious shrub or tree to 10 m. Branchlets strigose. Leaves alternate, ovate, serrate, attenuate to acuminate, rounded at base; lamina 10–15 cm long, 5–8 cm wide, densely strigose, densely woolly beneath; cystoliths punctiform, dense; lateral veins 5–7 pairs, the basal prominent; petiole 5–10 cm long; stipules less than 3 mm long. Inflorescence single to few, spicate, as long as petiole; flowers clustered at intervals, subsessile to sessile, densely strigose. Male flowers: tepals 4; stamens 4; pistillode present. Female flowers: perianth tubular, the orifice 3–5-dentate; style long. Achenes enclosed by succulent perianth. Fig. 41B–C.

88 URTICACEAE 8. Pipturus

Occurs from Arnhem Land, N.T., through eastern Qld S to north-eastern N.S.W.; grows in open rainforest, especially as thickets in disturbed areas. Also in SE Asia and islands of the South Pacific. Regions: ARNM, CYRK, BURD, DWSN, MCPH. Map 95. N.T.: Glyde R., H.Reeves 323 (DNA); Black Jungle, 51 km SE of Darwin, G.Chippendale 4505 (NSW). Qld: Rockhampton, A.Dietrich 1499 (NSW); Hervey Holding, Paluma Ra., B.Hyland 7299 (QRS). N.S.W.: between Shannon & Terania Ck Forest, R.Coveny 10624 (NSW). A distinctive species but highly variable especially in the indumentum of the branchlets and leaves. The var. calcicola is not clearly distinguished.

9. NOTHOCNIDE

Nothocnide Blume, Mus. Bot. 2: t. XIV (1856); from the Greek nothos (false) and knide (nettle), reference unclear. Type: N. repanda (Blume) Blume Pseudopipturus Skottsb., Acta Horti Gothob. 8: 117 (1933). T: P. repandus (Blume) Skottsb. = Nothocnide repanda (Blume) Blume Dioecious climbers or lianes. Leaves alternate, simple, coriaceous, entire, distinctly 3-nerved, glabrous, usually petiolate; stipules intrapetiolar, bifid. Inflorescence spicate, axillary. Flowers subsessile to sessile, clustered at intervals along rachis, surrounded by dense long white hairs. Male flowers: tepals 4; pistillode surrounded by long white hairs. Female flowers: perianth tubular, minutely toothed at apex, adnate to ovary; stigma ligulate. Achene enclosed by succulent perianth. A genus of 5 species centred in Malesia; 1 species in Australia. W.-L.Chew, Nothocnide (Urticaceae) in Malesia, Gard. Bull. Singapore 24: 361–373 (1969).

1. Nothocnide repanda (Blume) Blume, Mus. Bot. 2: t. xiv (1856) Urtica repanda Blume, Bijdr. 501 (1825); Boehmeria repanda (Blume) Hassk., Cat. Hort. Bogor. 79 (1844); Pipturus repandus (Blume) Wedd., Ann. Sci. Nat. Bot. ser. 4, 1: 196 (1854); Pseudopipturus repandus (Blume) Skottsb., Acta Horti Gothob. 8: 117 (1933). T: from Indonesia, collector not designated; n.v. Illustrations: C.L.Blume, Mus. Bot. 2: t. xiv (1856). W.-L.Chew, Gard. Bull. Singapore 24: 363, fig. 1 (1969). Dioecious climber or liane, sparsely strigose to glabrous. Leaves alternate, broadly elliptic to obovate, entire to lightly crenulate, acute to acuminate, cuneate to rounded at base; lamina 12–15 cm long, 5–7 cm wide, with dense punctiform cystoliths, sparsely strigose on veins, 3-nerved, with 1 or 2 more pairs near apex; petiole 3–5 cm long, lightly strigose; stipules to 6 mm long. Inflorescences 2–4 per axil, to 10 cm long, the females slightly longer. Male flowers c. 1 mm long, strigose. Female flowers: to 1.5 mm long, 1 mm wide; perianth orifice protruded slightly, 4-dentate, strigose; stigma 3 mm long, caducous. Achenes obovoid, 1.5 mm long, 1 mm broad. Fig. 42A–E. Occurs on Cape York Peninsula, Qld, in rainforest; also in Malesia and the Solomon Islands. Region: CYRK. Map 96. Qld: Timber Reserve Massy, B.Hyland 21122V (QRS); Claudie R., B.Hyland 7008 (QRS); Garraway R., B.Hyland 9534 (QRS); Atherton, B.Hyland 8631 (QRS). Australian populations appear to differ slightly from the Malesian counterparts in a few characters: leaves more broadly elliptic and very sparsely strigose on veins below; orifice of female perianth more protruded.

89

Figure 42. A–E, Nothocnide repanda. A, stipule ×7.5; B, L.S. ♀flower ×20; C, ♀inflorescence ×7; D, ♀flower ×5; E, habit ×0.5 (D–E, QRS 023520, QRS). F–G, Parietaria cardiostegia. F, habit ×0.75; G, fruits ×5 (F–G, K.Newbey 5792, PERTH). H–I, Australina pusilla. H, habit ×0.75, Victorian form (NSW 25562, NSW); I, leaves ×0.75, Tasmanian form (T. & J.Whaite 2390, NSW). A–C reproduced by permission from Garden’s Bulletin, Singapore 24: 363, fig. 1 (1969). D–I drawn by D.Mackay.

90 URTICACEAE Trib. 4. Parietarieae

Trib. 4. PARIETARIEAE

Urticaceae trib. Parietarieae Gaudich., Voy. Uranie 501 ‘1826’ (1830). Type: Parietaria L. Herbs or (not in Australia) shrubs, without irritant hairs. Leaves alternate; cystoliths punctiform. Flowers unisexual or bisexual. Male flowers: tepals 3 or 4; stamens 3 or 4; pistillode present. Female flowers: perianth tubular, with contracted 2–4-dentate orifice; staminodes absent. A small tribe of 5 genera; 1 in Australia.

10. PARIETARIA

Parietaria L., Sp. Pl. 1052 (1753); Gen. Pl. 4th edn, 471 (1754); from the Latin parietarius (of walls) with which the plant is often associated, as observed by Pliny. Type: P. officinalis L. Freirea Gaudich., Voy. Uranie 502 ‘1826’ (1830). T: not designated. Monoecious or polygamous annual or perennial herbs, usually densely pubescent with hooked or curved hairs. Leaves alternate, chartaceous, entire, 3-nerved, petiolate; stipules absent. Flowers in clusters or cymules in axils, the lateral flowers with an involucre of leafy bracts. Male flowers: tepals 3 or 4; stamens 3 or 4. Female flowers: perianth 3- or 4-dentate enclosing the free ovary; stigma subcapitate to capitate, dense with long stigmatic processes. Bisexual flowers: perianth and stamens as in males, pistil as in females. Achenes ovoid, smooth to polished, brown, enclosed by persistent perianth subtended by foliaceous bracts. A small genus of c. 20 species chiefly in the Middle East with a few species in South America, Asia and Australia; 3 species in Australia, of which 1 is naturalised. 1 Leaf lamina rhombic, acute; bisexual flowers numerous; stigma on long thin style; bracts lanceolate, elliptic or ovate, connate at base 3. P. judaica 1: Leaf lamina ovate, obtuse; bisexual flowers few; stigma subsessile; bracts variable in shape, free 2 Bracts lanceolate to broadly elliptic, with simple or ±branched midvein 1. P. debilis 2: Bracts cordate, with a conspicuous radial reticulate venation 2. P. cardiostegia

1. Parietaria debilis G.Forster, Prodr. 73 (1786) T: New Zealand, G.Forster; n.v. Freirea australis Nees in J.G.C.Lehmann, Pl. Preiss. 1: 638 (1845); Parietaria australis (Nees) Blume, Mus. Bot. 2: 256 (1857); Parietaria debilis var. australis (Nees) J.Black, Fl. S. Australia 2: 157 (1924). T: Rottnest Is., W.A., Aug. 1839, L.Preiss 2399; n.v. Parietaria squalida J.D.Hook., London J. Bot. 6: 285 (1847); P. debilis var. squalida (J.D.Hook.) J.D.Hook., Fl. Tasman. 1: 344 (1857). T: Circular Head, Tas., R.C.Gunn; n.v. Illustration: J.M.Black, Fl. S. Australia 2: 157, fig. 36 (1922). Polygamous annual herb. Leaves ovate, entire, obtuse, rounded at base; lamina 1.5–2.5 cm long, 0.5–1.5 cm wide; lateral veins in 3 pairs, the basal pair prominent; petiole 1–2 cm long. Inflorescence a few-flowered sessile cymule or cluster, the lower axils mainly female, the upper mixed male, female and bisexual; lateral flowers subtended by involucre of 3 free lanceolate or ligulate bracts. Male flowers subsessile, c. 1 mm long. Female flowers c. 2 mm long; perianth 4-dentate; ovary ellipsoidal to ovoid; stigma subsessile, subcapitate, with long stigmatic processes. Lateral flowers long-pedicellate in fruit. Achenes ellipsoidal, c. 2 mm long, smooth, enclosing perianth, brown-ribbed. Native Pellitory. Occurs in temperate Australia, locally common in damp shaded places, preferring calcareous soils; widely distributed throughout the temperate regions of the world. Regions: PILB,

91 10. Parietaria URTICACEAE

MRCH, BENC, LUWN, LEON, ESPR, GBSN, MACD, VICD, NULL, SIMP, TRNS, EYRE, RIVR, OTWY, WRGO, DARL, DWSN, MCPH, NEPN, HOWE, TASM. Map 97. N.T.: 40 km S of Alice Springs, D.J.Nelson 475 (NSW). S.A.: Koonalda, D.E.Symon 4527 (NSW). N.S.W.: Wuuluman, R.Coveny 10328 & J.Armstrong (NSW). Vic.: Bridgewater Bay, R.Melville 1500 (NSW). Tas.: Shingle Beach, Western plains, R.C.Gunn 886 (NSW).

2. Parietaria cardiostegia Greuter, Fl. Australia 3: 190 (1989) T: ‘Mesa’ hills, c. 12 km NW of Fowler Gap, N.S.W., Sept. 1981, W.Greuter 18387; holo: B; iso: NSW. Similar to P. debilis but bracts of involucre of lateral flowers broadly triangular-cordate, constricted at base in a short stout claw continuous with peduncle, with a conspicuous radial reticulate venation; involucre concealing flower and shed with the fruit. Fig. 42F–G. Endemic in semi-arid and arid areas of southern W.A., southern N.T., S.A., western N.S.W., and in Tas. Regions: MRCH, LUWN, BENC, VICD, NULL, MACD, EYRE, TRNS, COOP, DARL, TASM. Map 183. W.A.: Niagara, A.S.George 2742 (PERTH). N.T.: Pinta Springs, 45 km NW of Alice Springs, Aug. 1932, J.B.Cleland (AD). S.A.: Yarlbrinda Well, Koondoolka, Gawler Ra., J.Z.Weber 3142B (AD, NSW). N.S.W.: jump-up S of Olive Downs, Sturt Natl Park, W.Greuter 18495 (B, NSW). Tas.: Big Chalky Is., Furneaux Group, Bass Strait, J.S.Whinray 690 (AD). Previously included with the widespread, variable P. debilis which occurs in the same regions but grows in separate pure stands.

3. *Parietaria judaica L., Sp. Pl. 2nd edn, 2: 1492 (1764) T: Middle East, Palestine, collector unknown; n.v. [Parietaria diffusa auct. non Mert. & Koch; N.C.Beadle et al., Fl. Sydney Region 302 (1962); J.H.Willis, Handb. Pl. Victoria 2: 31 (1972)] Perennial herb. Leaves rhombic, entire, acute, cuneate at base, often slightly decurrent on petiole; lamina 2–3 cm long, 1–1.5 cm wide; lateral veins 2 or 3 pairs, the basal pair more prominent, ±decurrent on midrib; petiole 1–1.5 cm long. Inflorescence a few-flowered sessile cymule or cluster of male, female and bisexual flowers; bracts leafy, often connate at base. Male flowers sessile, c. 1 mm long. Female flowers subsessile or on pedicel 1–2 mm long; perianth 4-dentate, 2 mm long; ovary ovoid; style thin, long; stigma capitate. Bisexual flowers numerous. Achenes ellipsoidal, smooth. Pellitory. Fig. 14. Naturalised in many areas of temperate Australia as a common roadside and garden weed. Native of the Middle East. Regions: LUWN, EYRE, OTWY, NEPN. Map 98. W.A.: Essex Rd, Fremantle, G.J.Keighery 5903 (PERTH). N.S.W.: Sydney City, E.McBarron 22393 (NSW); Paddington, Sydney, R.Coveny 7609 (NSW).

Trib. 5. FORSSKAOLEAE

Urticaceae trib. Forsskaoleae Gaudich., Voy. Uranie 504 ‘1826’ (1830), as Forskalieae. Type: Forsskaolia L. Monoecious herbs without irritant hairs. Leaves alternate, rarely opposite; cystoliths linear or punctiform. Flowers unisexual, with tubular perianth. Male flowers: perianth often bilabiate; stamen 1; pistillode absent. Female flowers: perianth with contracted 4-dentate orifice, sometimes absent; staminode absent. A tribe of 5 genera; 1 genus in Australia. I.Friis & C.M.Wilmot-Dear, A revision of the tribe Forsskaoleae (Urticaceae), Nordic. J. Bot. 8(1): 25–59 (1988).

92 URTICACEAE 11. Australina

11. AUSTRALINA

Australina Gaudich., Voy. Uranie 505 ‘1826’ (1830); from the Latin australis (southern or south), in reference to Australia, where the type was collected by Labillardière. Type: A. pusilla (Poiret) Gaudich. Monoecious diffuse perennial herbs, often strigose. Leaves alternate, the upper ones opposite, membranous, 3-nerved, petiolate; stipules small, free, caducous. Inflorescence unisexual, axillary, few-flowered, without involucre of bracts. Male inflorescence 2–4-flowered, pedunculate; flowers sessile at peduncle apex; perianth bilabiate, the larger lobe incurved. Female inflorescence 2–5-flowered, sessile, usually below the male; flowers subsessile; perianth attenuate, 2- or 3-dentate; stigma filiform. Achene ellipsoidal, compressed, enclosed by perianth. A genus of 2 species, 1 in south-eastern Australia and New Zealand, the other in Ethiopia and .

1. Australina pusilla (Poiret) Gaudich., Voy. Bonite, Bot. 3: t. 114 (1852) Urtica pusilla Poiret in J.B.A.P.M. de Lamarck, Encycl. Suppl. 4: 224 (1816). T: Tasmania, J.J.H. de Labillardière; n.v. A. muelleri Wedd., Arch. Mus. Hist. Nat. 9: 545 (1857); A. pusilla subsp. muelleri (Wedd.) Friis & Wilmot-Dear, Nordic J. Bot. 7: 126 (1988). T: Delatite R., Vic., F.Mueller; holo: K. A. pusilla ß major J.D.Hook., Fl. Tasman. 2: 371 (1859). T: Coal R. Tier, A.Oldfield; n.v. Illustrations: W.M.Curtis, Stud. Fl. Tasmania 3: 641, fig. 136 (1967); I.Friis & C.M.Wilmot-Dear, Nordic J. Bot. 8 (1): 54, fig. 19 (1988). Herb to 50 cm, lightly strigose to glabrescent. Leaves almost circular, elliptic to obovate, dentate, obtuse to acute, cuneate at base; lamina 7–60 mm long, 7–20 mm wide; lateral veins 3 or 4 pairs, the basal pair prominent; petiole 3–30 mm long. Male inflorescence: peduncle to 1.3 cm long, thin; flowers c. 2 mm long, densely strigose. Female inflorescence 2–5-flowered; perianth tubular, attenuate, 2-dentate; stigma filiform, unilaterally stigmatic, strigose. Achene asymmetrically ellipsoidal, shortly stipitate, smooth, brown. Small Shade Nettle. Fig. 42H–I. Occurs from the Macpherson Ra., N.S.W., to the Otway Ra., Vic., and in Tas.; grows in shaded areas along watercourses, especially banks of running streams; also in New Zealand. Regions: MCPH, NEPN, HOWE, OTWY, TASM. Map 99. N.S.W.: Barrington Tops, Jan. 1934, L.Fraser & J.Vickery (NSW); Macleay R., 1882, E.Betche (NSW). Vic.: Sherbrooke Forest, Feb. 1949, L.A.S.Johnson (NSW). Tas.: Mt Field Natl Park, P.H.Raven et al. R25912 (NSW); Mt Barrow, Jan. 1922, H.M.R.Rupp (NSW). A highly variable species ranging from a tiny frail herb with almost circular leaves to a fairly tall erect herb. It is not practicable to segregate the infraspecific taxa recognised by Hooker (loc. cit.), and Friis & Wilmot-Dear (loc. cit.).

BALANOPACEAE

S.Carlquist

Dioecious, small to large trees. White circular lenticels prominent on older bark. Young shoots with scale leaves near base. Leaves alternate or subverticillate, simple, glabrous at maturity; margins recurved, toothed; stipules minute. Inflorescences usually axillary to scale leaves. Male inflorescence a ; flowers shortly pedicellate, axillary to scales on catkin axis. Male flowers: perianth reduced, of 1 or more teeth often fewer than stamens; stamens

93

Figure 43. Balanops australiana. A, ♀flower ×3.5; B, L.S. ovary ×6.25; C, ♀flowering branch ×0.38 (A–C, S.Carlquist 15210, RSA); D, fruit ×1.13; E, L.S. fruit ×2.13 (D–E, A.Dockrill 1405, RSA); F, ♂flowering branch ×0.38; G, ♂catkin ×7.5; H, anther ×25 (F–H, B.Hyland 2159, RSA). Drawn by S.Summit.

94 BALANOPACEAE

1–12; anthers oblong-elliptic, dehiscing laterally; pistillodes small or lacking. Female inflorescences a single terminal flower subtended by numerous crowded, triangular bracts; peduncle elongate or condensed. Female flower: perianth and staminodes absent; ovary bicarpellate or tricarpellate, the locules imperfectly septate at anthesis, fully septate in fruit; ovules 2 per carpel; styles once or twice divaricate, filiform, the bases persistent. Fruit drupaceous, yellow, orange or brown; pyrene 1 per carpel; embryo green, straight, erect, with 2 succulent elliptic-oblong cotyledons filling pyrene. A monogeneric family (formerly known as Balanopsidaceae) with 9 species in Australia, New Caledonia, New Hebrides, and ; 1 species endemic in north-eastern Qld. The segregate genus Trilocularia Schltr. is not recognised by recent authors. The affinity of the family, formerly believed to lie with Fagaceae (Cronquist, 1981), appears to lie elsewhere (possibly near Hamamelidales or ). No economic uses are known. A.Guillaumin, Recherches sur l’anatomie et la classification des Balanopsidacées, Rev. Gén. Bot. 37: 433–449 (1925); H.Hjelmquist, Studies on the floral morphology and phylogeny of the Amentiferae, Bot. Not. Suppl. 2: 1–171 (1948); S.Carlquist, Anatomy and systematics of Balanopaceae, Allertonia 2: 191–246 (1980).

1. BALANOPS

Balanops Baillon, Adansonia 10: 117 (1871); from the Greek balanos () and ops (looks like), in reference to the nature of the fruit. Type: B. vieillardii Baillon (as ‘vieiardii’)

1. Balanops australiana F.Muell., Fragm. 10: 114 (1877) T: Rockingham Bay, Qld, J.Dallachy; holo: MEL; iso: K. B. montana C.White, Contr. Arnold Arbor. 4: 15 (1933). T: Mt Alexander [Thornton Peak], Qld, S.F.Kajewski 1485; holo: A; iso: BRI, K. Illustration: B.D.Morley & H.R.Toelken, Fl. Pl. Australia fig. 34e (1983). Tree to 35 m. Leaves scattered along upper portion of shoots, obovate-elliptic to elliptic to oblanceolate, acute to acuminate, thinly coriaceous, the veins not prominent in dried leaves; lamina 4.5–9 cm long, 1.5–5 cm wide; margins apparently entire, the teeth small or not visible; petiole 5–10 mm long. Male inflorescence dense, c. 5 mm long. Female inflorescence c. 6 mm diam. at anthesis, sessile; bracts hairy, orbicular-rhomboid, to 5 mm long in fruit. Female flower: carpels 2; styles twice bifurcate near base. Fruit oblong, c. 1.5 cm long, 8 mm diam., bright orange at maturity; pyrenes c. 1.3 cm long, the endocarp papery-fibrous. Fig. 43. Endemic in north-eastern Qld, from near Cooktown southwards to the Eungella Ra., at 500–1500 m elevation, in both granitic and volcanic soil in wet forest. Regions: CYRK, BURD. Map 100. Qld: Mt Dalrymple, N.Hansen 129 (QRS); Atherton and vicinity, S.Carlquist 15210 (RSA); Thornton Peak, B.Hyland 707 (RSA, QRS); Mt Finnegan, L.J.Brass 20328 (A, K). Plants on exposed summits usually are smaller and have smaller leaves than those in deep rainforest.

95

BETULACEAE

H.J.Hewson

Deciduous trees or shrubs, monoecious. Leaves alternate, simple, pinnately veined, petiolate; stipules caducous. Inflorescence a terminal or axillary, paniculate or solitary unisexual cymose catkin or spike, bracteate. Male inflorescence pendulous. Male flowers solitary or in 3-flowered dichasia; sepals 1–6, scale-like or obsolete; corolla absent; stamens usually equal in number to, and opposite, sepals; pistillode present or absent. Female inflorescence erect. Female flowers in 2- or 3-flowered dichasia; perianth usually absent; staminodes absent; ovary inferior; carpels 2 or 3, each with 1 or 2 axile, pendulous ovules; styles usually separate. Fruit a nut or a 2-winged samara enclosed by thickened bracts and bracteoles, usually in a cone-like infructescence. Seed usually solitary, with or without endosperm. A family of 6 genera and c. 120 species mainly in the temperate and cooler regions of the northern hemisphere; one species naturalised in Australia. Betula alba L. () is listed as an escape in Tasmania, T.D.Raphael, Proc. Roy. Soc. Tasmania 89: 150 (1955), but is not regarded as naturalised. H.Winkler, , Pflanzenr. 19: 1–149 (1904).

1. ALNUS

Alnus Miller, Gard. Dict. Abr. 4th edn (1754); the Latin name for the . Type: A. glutinosa (L.) Gaertner Trees or shrubs. Inflorescence composed of dichasia; dichasia each with 1 bract and 2 bracteoles. Male flowers 3 per dichasium; perianth usually 4-partite. Female flowers 2 per dichasium; perianth absent. Infructescence cone-like, ovoid or ellipsoidal; scales 5-lobed, thick, woody, persistent. A genus of 30 species in temperate regions of the northern hemisphere and the Andes in South America; one species naturalised in Australia.

1. *Alnus cordata (Lois.) Desf., Tabl. Hort. Par. 2nd edn, 244 (1815) Betula cordata Lois., Not. Fl. France 139 (1810). T: Corsica, M.G.Robert; n.v. Illustration: H.Winkler, Pflanzenr. 19: fig. 26 (1904). Tree to 15 m tall. Vegetative enclosed by stipules of the first foliage leaf, resinous from secretions from glands on buds and stipules. Leaves ovate to suborbicular, obtuse or acuminate, truncate or cordate at base, crenate, dentate or serrate; lamina 2–11 cm long, glabrescent; petiole 2–3 cm long. emerging before leaves. Male catkins 3–7 at ends of twigs, 2–2.5 cm long. Female catkins 1–3 together, ovoid-cylindrical in flower, 1.5–3 cm long, 1–2 cm wide, ovoid-oblong in fruit. Nutlet narrowly winged. Alder. Native of Italy and Corsica; recorded as sparingly naturalised near St Marys and New Norfolk in Tas. by W.M.Curtis, Stud. Fl. Tasmania 3: 645 (1967). Region: TASM. Map 101. Tas.: St Marys, W.M.Curtis (HO).

96

FAGACEAE

H.J.Hewson

Trees or shrubs, mostly monoecious, deciduous or evergreen. Leaves alternate (in Australia), simple, pinnately veined, stipulate; stipules caducous. Inflorescence axillary, cymose, usually with flowers in reduced dichasia and the dichasia organised in a catkin or spike, bracteate. Flowers small, inconspicuous, unisexual; perianth a single connate whorl, usually 6-lobed, sometimes obsolete. Male flowers solitary or in dichasial clusters; stamens 4–40, usually 6–12; anthers dehiscing by longitudinal slits; pistillode sometimes present. Female flowers solitary or in dichasial clusters subtended by an involucre of bracts which develop into a cupule; staminodes present or absent; ovary inferior, of 2–12 carpels each with a distinct style; ovules 2 per locule, pendulous; placentation axile. Fruit a nut (or nuts) subtended by a cupule. Seed 1, without endosperm. An almost cosmopolitan family (excluding tropical and southern Africa) of 6–8 genera and c. 800 species; represented in Australia by 1 genus and 3 endemic species. Seedlings of Quercus (Q. robur L. and Q. ilex L.) persist around planted trees, but neither species is regarded as naturalised. The Fagaceae are important economically particularly for timber, cork and ornament. Species of Quercus () and Fagus (Beech) from the Northern Hemisphere and (Antarctic Beech and Southern Beech) in the Southern Hemisphere are important timber trees. L. yields cork. Miller () produces edible nuts. G.Bentham, Cupuliferae, Fl. Austral. 6: 209–211 (1873); E.Soepadmo, Fagaceae, Fl. Males. ser. 1, 7: 265–403 (1973).

1. NOTHOFAGUS

Nothofagus Blume, Ann. Mus. Bot. Lugduno-Batavum 1: 307 (1850), nom. cons.; from the Greek notho- (false) and Fagus (Beech), meaning false Beech or mongrel Beech. Type: N. antarctica (G.Forster) Oersted, typ. cons. Trees or shrubs, deciduous or evergreen. Leaves in bud appearing decussate, after expansion distichous, petiolate, gland-dotted; stipules peltate. Inflorescences in axils of leaves of current season’s growth. Male flowers sessile or shortly pedunculate in 1- or 3-flowered dichasia; stamens 6–18; anthers linear, apiculate, basifixed, latrorse. Female flowers above the male flowers in 1- or 3-flowered dichasia; cupule 4-valved (in Australia). Fruits with one 2-winged central nut and two 3-winged lateral nuts. A genus of c. 35 species occurring in the Southern Hemisphere in South America, New Zealand, Australia, New Caledonia and New Guinea; 3 species endemic in Australia. Following the classification proposed by van Steenis (1953), both N. moorei and N. cunninghamii belong in sect. Calusparassus subsect. Quadripartitae Steenis in having persistent leaves not plicate in bud, lateral female flowers 3-merous and a 4-partite cupule. Nothofagus gunnii belongs in sect. Calucechinus subsect. Antarcticae Steenis in having deciduous leaves plicate in bud, 3-flowered female inflorescences and a 4-partite cupule. C.G.G.J.van Steenis, Papuan Nothofagus, J. Arnold Arbor. 34: 301–347 (1953). C.G.G.J.van Steenis, Nothofagus, Key genus of plant geography in time and space, living and fossil, and phylogeny, Blumea 19: 65–98 (1971); R.S.Hill, Evolution of Nothofagus cunninghamii and its Relationship to N. moorei as Inferred from Tasmanian Macrofossils, Austral. J. Bot. 31: 453–465 (1983); J.R.Busby, Nothofagus cunninghamii (Southern Beech) vegetation in Australia, Austral. Fl. Fauna Ser. 1: 1–69 (1984).

97 Nothofagus FAGACEAE

1 Leaves ±ovate-elliptic, 30–80 mm long, acute, with serrulate margins 1. N. moorei 1: Leaves broadly ovate, rhomboidal, triangular or orbicular, 3–25 mm long, obtuse, with crenate or bluntly serrate margins 2 Plant evergreen; leaves not plicate in bud; leaf margins bluntly serrate 2. N. cunninghamii 2: Plant deciduous; leaves plicate in bud; leaf margins crenate 3. N. gunnii

1. Nothofagus moorei (F.Muell.) Krasser, Ann. K. K. Naturhist. Hofmus 11: 161 (1896) Fagus moorei F.Muell., Fragm. 5: 109 (1866). T: Bellinger R., N.S.W., C.Moore; syn: n.v.; Bielsdown Ck, N.S.W., C.Moore; syn: MEL; Macleay R., N.S.W., C.Moore; syn: MEL. Illustrations: J.H.Maiden, Forest Fl. New South Wales 9: t. 258 (1922); N.C.W.Beadle, Stud. Fl. N.E. New South Wales 4: fig. 213 (1980); D.J.Boland et al., Forest Trees Australia 4th edn, 135 (1984). Evergreen tree to 50 m tall, sometimes multi-trunked, frequently with adventitious shoots; trunk to 1.5 m diam. Leaves ovate to ovate-elliptic or ovate-lanceolate, acute, serrulate; lamina 30–80 mm long; stipules 1–1.5 cm long. Male inflorescence rounded or flattened, c. 1 cm diam. with 8–12-lobed involucre surrounding 1–3 flowers totalling 15–40 stamens. Female inflorescence ovoid, c. 5 mm long, with 3 flowers. Fruits in a cupule 8–10 mm long; valves 4, prickly. Negrohead Beech, Antarctic Beech. Fig. 44A. Occurs from the McPherson Ra., Qld, S to Barrington and Gloucester Tops, N.S.W.; grows in cool often in pure stands, or as an understorey in tall eucalypt forest at altitudes above 800 m, usually in fertile soils. Regions: MCPH, NEPN. Map 102. Qld: Nothofagus Mtn, A.G.Floyd 761 (CANB); Lamington Natl Park, C.T.White 11402 (CANB). N.S.W.: Cockerawombeena Reserve, Mt Boss State Forest, R.Coveny 10870 (NSW); Tyalgum Ridge, 28°18'S, 153°08'E, R.Coveny 9940 & L.Haegi (CANB, NSW); Wiangaree State Forest, Tweed Ra., R.Schodde 5621 (CANB, MEL, NSW). Great ages have been accredited to multi-trunked linked coppice clumps on McPherson Ra. summits.

2. Nothofagus cunninghamii (Hook.) Oersted, Kongel Danske Vidensk.-Selsk. Skr. ser. 5, 9: 355 (1871) = Bidr. Egefam. 25 (1871) Fagus cunninghamii Hook., Hooker’s J. Bot. Kew Gard. Misc. 2: 152, t. 7 (1840). T: Pine Cove, Macquarie Harbour, Tas., A.Cunningham; syn: n.v.; Emu Bay & Mt Wellington, Tas., J.Backhouse; syn: n.v.; western Tas., J.Milligan & R.C.Gunn; syn: MEL. Illustrations: W.J.Hooker, loc. cit.; D.J.Boland et al., Forest Trees Australia 4th edn, 131 (1984). Evergreen shrub or tree to 50 m tall, frequently with adventitious shoots, sometimes multi-stemmed; trunk to 2.5 m diam. Leaves ovate to triangular, rhomboidal or orbicular, obtuse, bluntly serrate; lamina 3–25 mm long, stipules 2–4 mm long. Male inflorescence with 1 or 3 flowers; perianth 6-lobed, 2–3 mm long; stamens 8–12. Female inflorescence ovoid, 2–4 mm long, with 3 flowers. Fruits in a cupule c. 6 mm long; valves with 4 or 5 rows of recurved teeth. Myrtle, Myrtle Beech, Southern Beech. Fig. 44B. Occurs in the Otway Ranges, the highlands E of Melbourne, Strzelecki Ranges and Wilsons Promontory, Vic., and in Tas. Grows in temperate rainforest, often in pure stands, or as an understorey in tall eucalypt forest. Regions: OTWY, HOWE, TASM. Map 103. Vic.: Baw Baw Plateau, L.A.Craven 1783 (CANB, NSW); Mt Donna Buang, R.Schodde 3188 (CANB, NSW); Thompson R., Apr. 1911, J.Staer (NSW). Tas.: c. 8 km N of Tarraleah, N.T.Burbidge 3318 (CANB); Mt Laperouse, Dec. 1898, F.A.Rodway (NSW). At higher altitudes, the habit may be shrubby.

98

Figure 44. Nothofagus. A, N. moorei, leaf (underside) ×1 (R.Story 7567, CANB). B, N. cunninghamii, leaf (underside) ×1 (R.Schodde 3188, CANB). C–F, N. gunnii. C, habit ×1; D, ♂flower ×2.5 (C–D, near Lake Fenton, Tas., J.Somerville, HO); E, fruit ×2.5 (T.Burns K327, HO); F, ♀flower ×2.5 (A.Dobson 77136, HO). Drawn by D.Boyer.

99 Nothofagus FAGACEAE

3. Nothofagus gunnii (J.D.Hook.) Oersted, Kongel Danske Vidensk.-Selsk. Skr. ser. 5, 9: 354 (1871) = Bidr. Egefam. 24 (1871) Fagus gunnii J.D.Hook., Icon. Pl. 9: t. 881 (1851). T: summit of Mount Olympus (4500–5000 ft [c. 1370–1520 m]), Tas., R.C.Gunn; iso: HO n.v., fide A.E.Orchard, pers. comm. Illustration: J.D.Hooker, loc. cit. Deciduous shrub or tree to 3 m tall, sometimes to 8 m. Leaves broadly ovate to orbicular, obtuse or emarginate, crenate; lamina 10–20 mm long; stipules c. 3 mm long, saccate, ±persistent. Male inflorescence with 1–3 flowers, on recurved pedicel; perianth 3–6-lobed, c. 4 mm long; stamens 6–12. Female inflorescence ovoid, 3–5 mm long, with 3 flowers. Fruits in a cupule; valves each with 4–6 sets of recurved scales; cupule 8–9 mm long. Tanglefoot, Fagus. Figs 15, 44C–F. Occurs on the western and central plateau mountains, Tas. Grows in subalpine shrub communities sometimes in pure stands. Region: TASM. Map 104. Tas.: near Lindon Tarns, Cradle Mtn, A.M.Buchanan 874 (HO); Moores Pimple, Jan. 1894, W.V.Fitzgerald (NSW); near Lake Fenton, Mt Field Natl Park, R.Melville 2301 (NSW); near Lake Fenton, Mt Field Natl Park, Dec. 1960, J.Somerville (HO). In exposed situations, often grows as a wiry, tangled shrub to 2.5 m tall forming impenetrable stands.

CASUARINACEAE

K.L.Wilson and L.A.S.Johnson

Dioecious or monoecious trees or shrubs. Branchlets articulate, slender, wiry, with several short basal articles and 1–numerous elongated articles; articles with as many ridges (phyllichnia) as there are teeth (reduced leaves), the phyllichnia separated by furrows (shallow and open in , deep and closed in other genera) containing the stomates. Leaves reduced to teeth in a whorl of 4–20 at apex of each article of assimilatory branchlets. Inflorescences of alternating whorls of tooth-like bracts; within each bract 2 lateral scale-like bracteoles, persistent, occasionally deciduous in male , and a single flower. Male inflorescence a short to elongated spike (Fig. 45J, K). Male flowers: tepals 1 or 2, hooded, scale-like, deciduous at anthesis; stamen 1; anther 2-locular, basifixed. Female inflorescence a globular or ovoid head (Fig. 45 I). Female flowers: perianth absent; carpels 2, fused; ovules 2, rarely 4; style 2-branched, reddish. Infructescence a ±woody ‘cone’, the 2 floral bracteoles enlarged as valves. Fruit a winged nut (samara) (Fig. 45A–H). Seed solitary in each samara; cotyledons large; endosperm absent; often with more than 1 embryo. A family of 4 genera and 90 species from Australia to the islands of the Pacific and SE Asia; 3 genera and 66 species in Australia. A number of species, especially of , are commonly planted in Australia and overseas as ornamentals, and for fuel or rough timber in developing countries. C. equisetifolia is also used in stabilising coastal sands. In the past the timber of various species of Casuarina and Allocasuarina was used for shingles, bullock yokes, etc., but the timber is now of little commercial importance. Various larger species are known as ‘Forest Oak’, ‘Bull Oak’ etc.; the general name ‘Sheoak’ was given by early settlers in allusion to the timber, which is oak-like in appearance but was considered inferior in strength to English Oak. Various species of insects, especially coccoids () (Gullan, 1984), form distinctive galls on ; some of these may be mistaken for fruiting cones but they are actually modified vegetative branchlets. Actinomycete () occur in nodules on

100

Figure 45. Details of Casuarinaceae. A–H, samaras ×5. A, (NSW 63049, NSW); B, (K.Wilson 5906, NSW); C, C. cunninghamiana subsp. cunninghamiana (N.Hall H80/19, NSW); D, (R.Melville 3785A, NSW); E, A. drummondiana (NSW 53232, NSW); F, A. microstachya (NSW 95258, NSW); G, A. distyla (A.Rodd & M.Watson 3537, NSW); H, A. grevilleoides (E.Bennett 1657, NSW). I–K, Allocasuarina inflorescences. I, A. paludosa (♀) ×5 (R.Melville 3785, NSW); J, A. dielsiana (♂) ×5 (K.Wilson 2655, NSW); K, A. verticillata (♂) ×2 (NSW 77451, NSW). Drawn by D.Mackay.

101 CASUARINACEAE the roots and are effective in fixation of atmospheric nitrogen. Vesicular-arbuscular endotrophic mycorrhizae have also been reported. Macrofossils from the and Miocene indicate a much wider distribution for the family in the past, especially Gymnostoma which occurred in south-eastern Australia, New Zealand and Patagonia. Allocasuarina have been found in the South Is. of New Zealand (Campbell & Holden, 1984) and possibly in Victoria. Fossil pollen of the family has been found in , Argentina and on Ninety East Ridge in the Indian Ocean, as well as being common in Australia. Cones open readily at maturity and seed is short-lived, except in Allocasuarina, where cones may remain unopened for several years and seed is correspondingly long-lived. In exposed maritime situations, species such as and become stunted and have very thick articles with more teeth than usual. Dried specimens show some important differences from fresh material, notably in the cones and branchlets. When fresh, the fruiting bracteoles of the cone are nearly always appressed to each other, enclosing the samara; when the cone is removed from the plant and dries out, the bracteoles separate. In this treatment, measurements of cone length do not include any sterile apex (mentioned separately if of significant size); cone body diam. does not include any portion of the bracteoles extending beyond the main body of the cone. Samaras are paler when immature. The softer tissues of branchlets tend to contract when dried, so that features such as angularity or roundness of phyllichnia are emphasised in dried specimens. The keys and descriptions are mostly based on dried specimens. F.A.W.Miquel, Revisio critica Casuarinarum (1848), apparently published simultaneously in Nieuwe Verh. Eerste Kl. Kon. Ned. Inst. Wetensch. Amsterdam ser. 2, 13: 267–350, t. 1–12; G.Bentham, Casuarinaceae, Fl. Austral. 6: 192–202 (1873); J.Poisson, Recherches sur les Casuarina et en particulier sur ceux de la Nouvelle-Calédonie, Nouv. Arch. Mus. Hist. Nat. ser. 1, 10: 59–111, t. 4–7 (1874); B.A.Barlow, Chromosome numbers in the Casuarinaceae, Austral. J. Bot. 7: 230–237 (1959); D.C.Christophel, Occurrence of Casuarina megafossils in the Tertiary of South-eastern Australia, Austral. J. Bot. 28: 249–259 (1980); E.M.Bennett, A guide to the Western Australian she-, W. Austral. Naturalist 15: 77–105 (1982); S.J.Midgley, J.W.Turnbull & R.D.Johnston (eds.), Casuarina Ecology, Management & Utilization (1983); J.D.Campbell & A.M.Holden, Miocene Casuarinacean fossils from Southland and Central Otago, New Zealand J. Bot. 22: 159–167 (1984); P.J.Gullan, A Revision of the Gall-Forming Coccoid Genus Cylindrococcus Maskell (Homoptera: Eriococcidae), Austral. J. Zool. 32: 677–690 (1984).

KEY TO GENERA

1 Furrows of branchlets shallow and open, exposing the stomates; teeth 4 per whorl; cone bracts broad and woody beneath each pair of bracteoles 1. GYMNOSTOMA 1: Furrows of branchlets deep and narrow, the stomates not visible; teeth 4–20 per whorl; cone bracts mostly thin at outer face, not conspicuous 2 Mature samaras grey or yellow-brown, dull; cone bracteoles thinly woody, prominent, extending well beyond cone body, without dorsal protuberance; teeth 5–20 per whorl 2. CASUARINA 2: Mature samaras red-brown to black, shining; cone bracteoles thickly woody and convex, mostly extending only slightly beyond cone body, mostly with a separate angular, divided or spiny dorsal protuberance; teeth 4–14 per whorl 3. ALLOCASUARINA

102 CASUARINACEAE 1. Gymnostoma

1. GYMNOSTOMA

Gymnostoma L.Johnson, Telopea 2: 83 (1980); from the Greek gymnos (naked) and stoma (mouth), in reference to the exposed stomates on the branchlets. Type: G. nodiflorum (Thunb.) L.Johnson Trees or tall shrubs, dioecious or monoecious. Young persistent branchlets similar to deciduous branchlets; all articles quadrangular; furrows shallow and open, exposing stomates. Leaves in whorls of 4. Male inflorescences on branchlets similar to vegetative branchlets, simple or appearing compound owing to condensation of branching; bracteoles persistent. Female inflorescences on short or elongated branchlets similar to vegetative branchlets. ‘Cones’ mostly borne amongst assimilatory branchlets; bracts much expanded laterally, broader than high; bracteoles protruding, rounded on back but neither split nor with a dorsal protuberance. Samara body striate, glabrous, yellow-brown or greyish, dull. 2n = 16 (3 spp. studied), B.A.Barlow, Austral. J. Bot. 7: 232 (1959), and B.A.Barlow pers. comm. A tropical genus of 18 species in Malesia, Fiji, New Caledonia and north-eastern Australia; 1 species endemic in Australia.

1. Gymnostoma australianum L.Johnson, Fl. Australia 3: 202 (1989) T: Mt Alexander [Thornton Peak], Qld, 17 Dec. 1929, S.F.Kajewski 1492; holo: NSW, with cones; iso: A, BO, BRI, K, MEL, NY, SING. Dioecious or monoecious tree 4–7 m high with some buttress roots, a broad flattish crown and rather candelabriform branching habit. Bark fissured, brown. Branchlets stiff, ascending, to 13 cm long; new shoots ferruginous- or white-pubescent; articles acutely angular, 2–4 mm long, 0.5–0.8 mm diam., smooth; teeth 0.4–0.7 mm long. Male inflorescences compound (so far as known); spikes 1.5–2 mm long; anther c. 0.5 mm long. Cones short-cylindrical, white- to ferruginous-pubescent especially when immature; peduncle to 5 mm long; cone body 7–10 mm long, 8–10 mm diam.; bracts not striate; bracteoles striate, acute. Samara 7–8 mm long, pale brown. Figs 45A, 46A–B. Apparently restricted to a few small populations on or near Thornton Peak, Qld; grows in rainforest and as an emergent in cloud forest, mostly near streams. Region: CYRK. Map 105. Qld: Roaring Meg Creek Falls near China Camp, D.F.Blaxell 1110 (NSW); Thornton Peak, T.Hartley 14074 (BRI); Thornton Peak, P.Hind 2416 (NSW); Noah Ck, Oct. 1962, W.Mason, K.Beck & A.Dockrill (K, NSW 147110). Differs from all other species of Gymnostoma in having the tips of young branchlets ferruginous- or white-pubescent. The mature cones are also generally pubescent, unlike other species, and the combination of size and shape of organs differs from those of other members of the genus.

2. CASUARINA

Casuarina L., Amoen. Acad. 143 (1759); from the neo-Latin casuarius (cassowary), from the resemblance of the drooping branchlets to the feathers of the cassowary. Type: C. equisetifolia L. Trees, dioecious, monoecious in C. equisetifolia. Young persistent branchlets differing from deciduous branchlets in shorter articles and shape or size of leaves (‘teeth’). Articles terete, smooth; furrows deep and closed, concealing stomates. Leaves in whorls of 5–20. Male inflorescences simple elongate spikes; bracteoles persistent. Female inflorescences on short lateral branchlets (‘peduncles’) differing in appearance from the vegetative branchlets. Cones borne among or below assimilatory branchlets, pedunculate, pubescent at least when

103 2. Casuarina CASUARINACEAE immature; bracts thin in exposed portion, not vertically expanded; bracteoles protruding from cone surface, never greatly thickened and always lacking a dorsal protuberance. Samara body glabrous, pale yellow-brown or greyish, dull. 2n = 18 (6 spp. studied), B.A.Barlow, Austral. J. Bot. 7: 230–237 (1959). A genus of 17 species in SE Asia, Malesia, Melanesia, Polynesia, New Caledonia and Australia; 6 species in Australia, widespread except for Tas. and part of south-western and central Australia. Not usually found in soils greatly deficient in nutrients (cf. Allocasuarina). 1 Phyllichnia narrow and prominently angular, occasionally flattish in older growth in C. equisetifolia but then densely pubescent on phyllichnia as well as in furrows; teeth 6–10 2 Branchlets and cones ±densely and obviously pubescent; teeth 6–8, not marcescent; cone body 10–24 mm long, 9–13 mm diam.; cone bracteoles acute; samara 6–8 mm long 1. C. equisetifolia 2: Branchlets and cones sparsely and minutely pubescent; teeth 6–10, marcescent or not; cone body 7–14 mm long, 4–6 mm diam.; cone bracteoles broadly acute to acute; samara 3–4 mm long 2. C. cunninghamiana 1: Phyllichnia broad, flat or slightly rounded or with a median groove; teeth 8–20, mostly marcescent 3 Teeth 12–20; phyllichnia smooth, flat or slightly rounded; cone body 7–10 mm diam.; cone bracteoles 2.0–2.5 mm wide, thin, broadly acute, with no more than 1 obvious striation; samara 3.5–5.0 mm long 4 Teeth on young permanent shoots long-recurved 3. C. glauca 4: Teeth on young permanent shoots appressed or slightly spreading 4. C. obesa 3: Teeth 8–13; phyllichnia slightly wrinkled, flat or with a yellowish median groove (often masked by wax); cone body 10–16 mm diam.; cone bracteoles 3.5–4.0 mm wide, rather thick, acute, with several obvious striations; samara 5.5–10.5 mm long 5 Teeth erect, appressed; articles 0.6–0.9 mm diam., somewhat waxy or occasionally sparsely pubescent 5. C. cristata 5: Teeth spreading to recurved; articles 1–1.8 mm diam., densely and very shortly pubescent, strongly waxy 6. C. pauper

1. L., Amoen. Acad. 143 (1759) T: Rumphius, Herbarium Amboinense 3: t. 57 (1743); holo. Illustration: D.J.Boland et al., Forest Trees Australia 4th edn, 105 (1984). Monoecious tree 6–35 m high. Bark scaly, grey-brown to black. New shoots with teeth erect. Branchlets drooping, to 30 cm long; articles 5–13 mm long, 0.5–1.0 mm diam.; furrows usually densely pubescent; phyllichnia angular or occasionally flat in older growth, glabrous or pubescent; teeth 7 or 8, occasionally 6, erect (rarely spreading in New Guinea), 0.3–0.8 mm long, not marcescent. Male spikes 0.7–4 cm long, 7–11.5 whorls per cm; anther 0.6–0.8 mm long. Cones sparsely pubescent to tomentose; peduncle 3–13 mm long; cone body 10–24 mm long, 9–13 mm diam.; bracteoles acute. Samara 6–8 mm long. Coast Sheoak. Tropical and subtropical coastlines of northern and north-eastern Australia; also from Madagascar (doubtfully native) and Burma to , Malesia, Melanesia and Polynesia. There are 2 subspecies, intergrading on the Qld coast between Cooktown and Sarina, and in Vanuatu. Articles 0.5–0.7 mm diam.; phyllichnia prominently angular, glabrous or glabrescent; tall erect tree 1a. subsp. equisetifolia Articles 0.7–1 mm diam.; phyllichnia prominently angular to flat in older growth, often on the same branchlet, usually densely pubescent (at least when immature); small tree with rounded crown 1b. subsp. incana

104

Figure 46. A–B, Gymnostoma australianum. A, cone ×1.5; B, branchlet ×7.5 (A–B, S.Kajewski 1492, NSW). Casuarina cones ×1.5, branchlets ×7.5, C–D, C. equisetifolia subsp. equisetifolia. C, cone (M.Koie & S.Olsen 1229, NSW); D, branchlet (NGF 25332, NSW). E–F, C. equisetifolia subsp. incana. E, cone; F, branchlet (E–F, NSW 50234, NSW). G–H, C. cunninghamiana subsp. cunninghamiana. G, cone; H, branchlet (G–H, S.Blake 17377, NSW). I, C. cunninghamiana subsp. miodon, branchlet (N.Speck 1656, NSW). Drawn by D.Mackay.

105 2. Casuarina CASUARINACEAE

1a. Casuarina equisetifolia L. subsp. equisetifolia Tree 7–35 m high; articles 5–8 mm long, 0.5–0.7 mm diam.; phyllichnia prominently angular, glabrous; furrows sparsely to densely pubescent; teeth 0.3–0.8 mm long, occasionally to 1.0 mm. Male spikes 0.7–4 cm long. Cone ±sparsely pubescent; peduncle 3–10 mm long; cone body 12–24 mm long, 9–11 mm diam. 2n = 18, B.A.Barlow, Austral. J. Bot. 7: 232 (1959). Fig. 46C–D. Occurs from near Darwin, N.T., E to about Cairns, Qld; on sandy and low rocky sea coasts; also from Burma through Malesia to Vietnam, Melanesia and Polynesia. Regions: ARNM, CARP, CYRK. Map 106. N.T.: 5 km S of Danger Point, Cobourg Peninsula, G.M.Chippendale 8234 (NSW, NT); Elcho Is., J.R.Maconochie 2117 (CANB, DNA, K, L, NSW, NT); Dripstone Park, Darwin, K.L.Wilson 5089, R.Barker & C.Scarlett (DNA, K, NSW). Qld: near mouth of Settlement Ck, Gulf of Carpentaria, R.A.Perry 1226 (CANB, K, NSW). 1b. Casuarina equisetifolia subsp. incana (Benth.) L.Johnson, J. Adelaide Bot. Gard. 6: 79 (1982) C. equisetifolia var. incana Benth., Fl. Austral. 6: 197 (1873). T: Port Macquarie, N.S.W., May 1819, A.Cunningham 45; lecto: K, with cones, fide L.A.S.Johnson, loc. cit.; isolecto: BM. Illustration: J.H.Maiden, Forest Fl. New South Wales 5: t. 182 (1913). Tree 6–12 m high. Articles 7–13 mm long, 0.7–1.0 mm diam., densely pubescent on phyllichnia when immature as well as in furrows; phyllichnia prominently angular to flat (both extremes may be found on the one branchlet), when flat often wrinkled; teeth c. 0.7 mm long. Male spikes 1.2–2.5 cm long. Cone densely white- to ferruginous-pubescent or tomentose; peduncle 3–13 mm long; cone body 10–20 mm long, 10–13 mm diam. 2n = 18, B.A.Barlow, Austral. J. Bot. 7: 232 (1959). Figs 21, 46E–F. Occurs from Rockhampton, Qld, S to Laurieton, N.S.W., on sea coasts, behind beaches and on rocky headlands; also New Caledonia and southern Vanuatu. Regions: DWSN, MCPH. Map 107. Qld: Bishop Is., mouth of Brisbane R., S.T.Blake 3342 & S.L.Everist (BRI, NSW); The Causeway, c. 8 km N of Emu Park, L.A.S.Johnson 8655 & K.L.Wilson (BRI, NSW). N.S.W.: Point Perpendicular, near Laurieton, H.S.McKee 5776 (NSW); 4 km N of Iluka, Back Beach, I.R.Telford 8977 & G.Butler (BISH, CBG, MEL, NSW).

2. Miq., Revis. Crit. Casuarinarum 56, t. 6A (1848) T: sandy shores of Moreton Bay and Glasshouse Bay [Qld], [collector cited by Miq. as Cunningham but probably not so]; lecto: K, with cones; fide L.A.S.Johnson, Fl. Australia 3: 200 (1989). Tree 15–35 m high. Bark finely fissured and scaly, grey-brown. Teeth on new shoots erect. Branchlets drooping in vigorous specimens, erect in depauperate specimens; articles 4–9 mm long, 0.4–0.7 mm diam., mostly glabrous; edges of furrows often marked (when dry) by a slight ridge; phyllichnia angular to flat with a median rib; teeth 6–10, erect, 0.3–0.5 mm long, marcescent or not. Male spikes 0.4–4 cm long, 11–13 whorls per cm; anther 0.4–0.7 mm long. Cones sparsely pubescent; peduncle 2–9 mm long; cone body 7–14 mm long, 4–6 mm diam.; bracteoles broadly acute to acute. Samara 3–4 mm long. River Oak, River Sheoak. Occurs from the Daly R., N.T., E around the Gulf of Carpentaria and S to Bega, N.S.W. Grows along permanent streams. There are 2 subspecies. Teeth 8–10, yellow at base, darker brown towards apex, marcescent; phyllichnia prominently angular; cone bracteoles broadly acute 2a. subsp. cunninghamiana Teeth 6 or 7, uniformly yellow, not marcescent; phyllichnia angular to nearly flat; cone bracteoles sharply acute 2b. subsp. miodon

106 CASUARINACEAE 2. Casuarina

2a. Casuarina cunninghamiana Miq. subsp. cunninghamiana C. equisetifolia var. microcarpa F.Muell., Fragm. 6: 17 (1867). T: Glendon, N.S.W., L.Leichhardt; lecto: MEL, with cones, fide L.A.S.Johnson, Fl. Australia 3: 201 (1989). Illustrations: J.H.Maiden, Forest Fl. New South Wales 2: t. 59 (1905); L.Fuller, Wollongong’s Native Trees 2nd edn, 103 (1982); D.J.Boland et al., Forest Trees Australia 4th edn, 97–99 (1984). Tree to 35 m high. Articles 6–9 mm long, 0.4–0.6 mm diam.; phyllichnia prominently angular; teeth 8–10, marcescent, yellow at base, darker brown towards apex. Cone bracteoles broadly acute. 2n = 18, B.A.Barlow, Austral. J. Bot. 7: 232 (1959). Figs 45C, 46G–H. Occurs from Laura, Qld, S to Bega, N.S.W., inland to E of Chillagoe and Augathella in Qld, to Condobolin and W of Narrandera, N.S.W., including A.C.T. Grows along permanent freshwater streams. Regions: CYRK, BURD, DWSN, MCPH, NEPN, HOWE. Map 108. Qld: Laura road, 22 km from Mulligan Hwy, C.H.Gittins 967 (NSW); Dawson R., Chauvel homestead, N of Theodore, N.H.Speck 1884 (CANB, K, NSW). N.S.W.: Baradine Ck, Kenebri, E.F.Constable NSW 20781 (K, L, NSW); Brogo R., c. 14 km N of Bega, L.A.S.Johnson NSW 50311 (NSW). Hybridises naturally with C. cristata and C. glauca. Other hybrids are recorded in cultivation. 2b. Casuarina cunninghamiana subsp. miodon L.Johnson, Fl. Australia 3: 200 (1989) T: 23 km NNE of Borroloola on Bing Bong road, N.T., 13 May 1983, K.L.Wilson 5361; holo: NSW, with cones; iso: BRI, CANB, DNA. Tree to 20 m high. Articles 4–7 mm long, 0.5–0.7 mm diam.; phyllichnia angular to nearly flat but still with a central rib; teeth 6 or 7, not marcescent, uniformly yellow. Cone bracteoles sharply acute. Fig. 46 I. Occurs from the Daly R., N.T., E to the Gulf of Carpentaria, Qld. Grows along fresh or brackish permanent streams. Regions: ARNM, CARP. Map 109. N.T.: Daly R., S.T.Blake 16255 (BRI, NSW); 50 km S of Katherine, N.H.Speck 1656 (CANB, NSW); Mainoru homestead, D.E.Symon 7666 (AD, NSW, NT). Qld: 50 km E of Doomadgee Mission, R.A.Perry 1388 (CANB, K, NSW). Subsp. miodon has articles of similar diameter to subsp. cunninghamii but has fewer teeth, so that the individual phyllichnia are broader in subsp. miodon. Specimens from the Burketown region are somewhat intermediate between the 2 subspecies.

3. Casuarina glauca Sieber ex Sprengel, Syst. Veg. 3: 803 (1826) T: New South Wales, F.W.Sieber 325; holo: B (♂); iso: BM, C, FI, K, L, LE, MEL, P, PR, S. Illustrations: L.Fuller, Wollongong’s Native Trees 2nd edn, 105 (1982); D.J.Boland et al., Forest Trees Australia 4th edn, 103 (1984). Tree 8–20 m high or rarely shrub to c. 2 m, frequently producing root suckers. Bark finely fissured and scaly, grey-brown. Teeth on new shoots long-recurved. Branchlets spreading to drooping, to 38 cm long; articles 8–20 mm long, 0.9–1.2 mm diam., glabrous, occasionally waxy; phyllichnia flat to slightly rounded; teeth 12–17, rarely to 20, erect, 0.6–0.9 mm long, usually marcescent. Male spikes 1.2–4 cm long, 7–10 whorls per cm; anther c. 0.8 mm long. Cones ferruginous- to white-pubescent, becoming glabrous; peduncle 3–12 mm long; cone body 9–18 mm long, 7–9 mm diam.; bracteoles broadly acute. Samara 3.5–5.0 mm long. 2n = 18, B.A.Barlow, Austral. J. Bot. 7: 232 (1959). Swamp Oak. Fig. 47A–C. Occurs from Yeppoon, Qld, S to Bermagui, N.S.W. Grows in brackish situations along estuaries and streams, usually near the coast but somewhat farther inland along the Clarence, Hunter and Hawkesbury R. valleys; often forming pure stands as an open forest or woodland. Regions: DWSN, MCPH, NEPN, HOWE. Map 110.

107 2. Casuarina CASUARINACEAE

Qld: Stradbroke Is., C.E.Hubbard 2326 (BRI, K, L). N.S.W.: Ballina, W.Bäuerlen 1631 (LE, NSW); Nandudga Lake, 3 km by road S of Narooma, E.F.Constable 6835 (NSW); Heathcote Rd at Deadmans Ck, R.G.Coveny 11173 & J.Thomas (K, MO, NSW); Singleton district, R.Story 7680 (CANB, K, NSW). Occurs as a shrub c. 2 m high, with coarse branchlets bearing up to 20 teeth, on exposed headlands. When dried, the articles are of noticeably greater diam. at the apex than towards the base of the article (cf. C. obesa). Hybridises occasionally with C. cunninghamiana where their ranges meet along the coastal rivers.

4. Miq. in J.G.C.Lehmann, Pl. Preiss. 1: 640 (1845) T: Swan R. above Perth, W.A., 27 June 1839, and near Leschenault Inlet, W.A., 30 Dec. 1839, L.Preiss 2002; lecto: U, with cones, fide L.A.S.Johnson, Fl. Australia 3: 201 (1989); isolecto: LE, MEL, NSW, P, S. Similar to C. glauca. Teeth on new shoots erect. Branchlets drooping or spreading, to 21 cm long; articles usually waxy, not or only slightly greater in diam. at apex when dried; phyllichnia flat; teeth 12–16, occasionally slightly spreading, 0.3–1 mm long. Cones sessile or on peduncle to 10 mm long; cone body 10–20 mm long, 8–10 mm diam. 2n = 18, B.A.Barlow, Austral. J. Bot. 7: 232 (1959). Fig. 47D–F. Widespread in south-western W.A., except the extreme SW corner, with a few occurrences in the Wimmera region, Vic., and now destroyed at the only locality known in south-western N.S.W., near Euston. A supposed occurrence near Lake Torrens, S.A., is now thought to be depauperate C. pauper. Grows in brackish or saline situations along rivers or near salt lakes. Regions: MRCH, LUWN, BENC, LEON, ESPR, VICD, RIVR, OTWY. Map 111. W.A.: Finlayson Ra., N.T.Burbidge 4802 (CANB, NSW); Murchison R. bridge, 613 km N of Perth, M.E.Phillips CBG 35672 (CBG, NSW); 25 km S of Borden, B.Rockel CO21 (FRI, NSW). N.S.W.: between Sturt Hwy and Lake Benanee, L.A.S.Johnson NSW 46618 (NSW). Vic.: c. 10 km SW of Mt Arapiles, J.McQueen & A.C.Beauglehole 56279 (MEL, NSW). This is the western and inland vicariant of C. glauca, most obviously differing from that species in the erect teeth at the apex of new shoots and in the articles, which are less variable in diam. over their length when dried. Occasionally hybridises with C. pauper in W.A.

5. Casuarina cristata Miq., Revis. Crit. Casuarinarum 70, t. 10 (1848) C. quadrivalvis var. cristata (Miq). Miq., Flora 48: 18 (1865). T: Lachlan R. [Condobolin area], N.S.W., A.Cunningham; holo: K, with cones; iso: U. C. lepidophloia F.Muell., Fragm. 10: 115 (1877). T: between the Bogan R. and the Lachlan R., N.S.W., L.Morton; lecto: MEL ♀),( fide L.A.S.Johnson, Fl. Australia 3: 201 (1989); isolecto: NSW 46689. C. cambagei R.Baker, Proc. Linn. Soc. New South Wales 24: 605 (1900). T: Grawlin, Forbes, N.S.W., 28 July 1899, R.H.Cambage NSW 46670; lecto: NSW, with cones, fide L.A.S.Johnson, Fl. Australia 3: 200 (1989). Illustrations: J.H.Maiden, Forest Fl. New South Wales 2: t. 51 (1905) as C. lepidophloia; G.M.Cunningham et al., Pl. W. New South Wales 207 (1982); D.J.Boland et al., Forest Trees Australia 4th edn, 94–95 (1984). Tree 10–20 m high, frequently producing suckers. Bark finely fissured or scaly, grey-brown. Teeth on new shoots erect to somewhat spreading. Branchlets drooping in vigorous specimens, spreading in depauperate specimens, to 25 cm long; articles usually slightly wrinkled, 8–17 mm long, 0.6–0.9 mm diam., somewhat waxy, occasionally sparsely pubescent, disarticulating readily; phyllichnia flat or with slight median groove, often masked by wax; teeth 8–12, erect, 0.5–0.7 mm long, marcescent. Male spikes 1.3–5 cm long, 6–10 whorls per cm; anther 0.8–1.1 mm long. Cones ferruginous-pubescent when young, nearly glabrous at maturity; peduncle 1–14 mm long; cone body 13–18 mm, occasionally to 25 mm long, 10–16 mm diam.; bracteoles acute. Samara 6–10.5 mm long. 2n = 18, B.A.Barlow, Austral. J. Bot. 7: 232 (1959). Belah. Figs 45B, 47G–H.

108

Figure 47. Casuarina cones ×1.5, branchlets ×7.5. A–C, C. glauca. A, cone; B, branchlet; C, branchlet apex ×5 (A–C, NSW 46556, NSW). D–F, C. obesa. D, cone; E, branchlet (D–E, P.Wilson 4140, NSW); F, branchlet apex ×5 (NSW 46603, NSW). G–H, C. cristata. G, cone (NSW 46655, NSW); H, branchlet (NSW 46625, NSW). I–J, C. pauper. I, cone (E.Bennett 3250, NSW); J, branchlet (NSW 14594, NSW). Drawn by D.Mackay.

109 2. Casuarina CASUARINACEAE

Occurs from Clermont, Qld, S to Temora, N.S.W.; usually in clayey grey or brown soils with calcareous nodules near the surface. May form pure woodland stands; clonal stands occur. Some intergradation with C. pauper occurs in the area from Bourke to W of Condobolin, N.S.W. Regions: DWSN, WRGO, MCPH, NEPN, DARL, RIVR. Map 112. Qld: Yelarbon, S.T.Blake 10471 (BRI, NSW); 8 km N of Nathan Rd and Leichhardt Hwy junction, N.H.Speck 1906 (CANB, K, NSW). N.S.W.: Lightning Ridge, E.F.Constable NSW 19232 (K, NSW); Gilgandra–Collie road, E.F.Constable NSW 20370 (K, NSW).

6. F.Muell. ex L.Johnson, Fl. Australia 3: 202 (1989) T: Flinders Ranges, S.A., Oct. 1851, F.Mueller; holo: MEL, (♀, with inflorescences and cones). [C. pauper Miq., Ned. Kruidk. Arch. 4: 100 (1856), nom. inval.; C. cristata subsp. pauper (Miq.) L.Johnson, Nuytsia 1: 265 (1972), nom. inval.] Similar to C. cristata. Tree 5–15 m high. Teeth on new shoots spreading to recurved; articles 1–1.8 mm diam., strongly waxy, shortly and densely pubescent; teeth 9–13, persistently ferruginous-pubescent. Samara 5.5–7.0 mm long. Black Oak. Fig. 47 I–J. Occurs from far south-western Qld through far western N.S.W., north-western Vic. and inland S.A. to southern inland W.A.; usually in red-brown soils with light-textured topsoil and calcareous subsoil. Regions: LEON, VICD, NULL, EYRE, TRNS, COOP, RIVR, DARL. Map 113. W.A.: 13 km N of Kalgoorlie, R.Melville 3996 (K, NSW). S.A.: Commonwealth Hill Stn, D.E.Symon 3389 (AD, CANB, K, NSW). Qld: 8 km E of Warri Gate, S end of Bygrave Ra., L.A.S.Johnson & E.F.Constable NSW 32033 (BRI, NSW). N.S.W.: Koonenberry Mtns, c. 99 km SSE of Milparinka, E.F.Constable 4612 (NSW). Vic.: Sunny Cliffs S of Mildura, A.C.Beauglehole 16037 (MEL, NSW). Trees are usually smaller and of poorer form than in C. cristata; also differing in the shorter cone bracteoles.

3. ALLOCASUARINA

Allocasuarina L.Johnson, J. Adelaide Bot. Gard. 6: 73 (1982); from Greek allos (other) and neo-Latin Casuarina, referring to the relationship with the genus Casuarina. Type: A. torulosa (Aiton) L.Johnson Shrubs or trees, dioecious or monoecious. Young persistent branchlets usually distinguishable from the deciduous branchlets. Branchlet articles with furrows deep, concealing the stomates. Leaves in whorls of 4–14. Male flowers in simple short to long spikes, on branchlets which in the flowering region are usually distinctly different from the vegetative branchlets. Female inflorescences on short lateral branches or ±sessile. Infructescences (‘cones’) pedunculate or sessile; bracts thin in exposed portion; bracteoles considerably thickened and often divided so that dorsal portion forms 1 or more distinct protuberances. Samara body brown to black, shining, glabrous or hairy. x = 10, 11, 12, 13 or 14, B.A.Barlow, Austral. J. Bot. 7: 230–237 (1959), and B.A.Barlow & B.G.Briggs pers. comm. An endemic genus of 59 species, chiefly in southern Australia but 4 species extending to north-eastern Qld and one, A. decaisneana, in tropical and subtropical parts of the eremean region. Usually grows in soils markedly deficient in nutrients. Polyploidy occurs in some species (tetraploids are fertile). Hybridisation occurs occasionally. Several species include apomictic plants. In section Cylindropitys there are many closely related species, often but not always of limited distribution. These may require particular care to distinguish. The small size of some parts requires a higher level of precision of measurements than is usual in a Flora treatment. Branchlet details reflect underlying anatomical differences, which have been helpful in defining the species. Those taxa that are sharply although finely distinctive are treated as species, reserving the category of subspecies for those that exhibit intergradation.

110 CASUARINACEAE 3. Allocasuarina

KEY TO SECTIONS

Note. This key requires full male and female material of a species for its use but is essentially a practical key for identification of the sections and therefore does not include all significant morphological and karyological features, which can be found in the English and Latin descriptions. 1 Branchlets 1–6 cm long, with 1–5 elongate articles each 0.6–5 cm long, mostly pungent sect. 2. OXYPITYS 1: Branchlets mostly 8–50 cm long, with numerous elongate articles each 0.1–6 cm long, not pungent (except A. decaisneana ), if branchlets less than 8 cm long and/or if fewer than 7 elongated articles then articles less than 1 cm long and penultimate branchlets mostly green 2 Male bracteoles deciduous 3 Trees; articles verruculose 4 Bark furrowed; cone bracteoles with 1 abaxial protuberance 5 Branchlets ascending; teeth erect; cones very shortly cylindrical, broader than long, with 1–4 fertile whorls sect. 3. PLATYPITYS 5: Branchlets drooping; teeth spreading; cones ovoid, with 6–16 fertile whorls sect. 8. OÖPITYS 4: Bark ribbony-fibrous; cone bracteoles subdivided into 2 or 3 bodies sect. 10. INOPITYS 3: Shrubs; articles smooth sect. 11. CYLINDROPITYS (p.p.) 2: Male bracteoles persistent 6 Cone bracteoles with conspicuous abaxial spine or awn 7 Branchlets with a strong waxy coating at maturity; teeth 8–11 sect. 4. ECHINOPITYS 7: Branchlets with no more than a faint waxy bloom; teeth 5–6 sect. 14. ACANTHOPITYS 6: Cone bracteoles without abaxial spine or awn 8 Branchlets with a strong waxy coating at maturity sect. 5. CEROPITYS 8: Branchlets with no more than a faint waxy bloom 9 Cone bracteoles divided into 4–20 small bodies 10 Trees; bark corky or scaly 11 Teeth 4 or 5 sect. 6. ALLOCASUARINA 11: Teeth 6–8 sect. 7. AMORPHOPITYS 10: Shrubs; bark smooth or finely striate 12 Cone bracteoles subdivided into 3–5 bodies sect. 12. NANNOPITYS 12: Cone bracteoles subdivided into 6–9 bodies sect. 13. TRACHYPITYS 9: Cone bracteoles with 1 protuberance (rarely superficially 2–3-lobed) or protuberance fused with bracteole body 13 Teeth pungent sect. 1. DOLICHOPITYS 13: Teeth not pungent 14 Cone bracteoles acuminate, mucronate sect. 9. TRICHOPITYS 14: Cone bracteoles obtuse to acute, not mucronate sect. 11. CYLINDROPITYS

111 3. Allocasuarina CASUARINACEAE

KEY TO SPECIES BASED ON FEMALE SPECIMENS Notes. Hairs are produced in the furrows of all articles, as can be seen in T.S. However, the constriction of the furrows often means that no hairs are visible externally. Article furrows are here referred to as ‘pubescent’ when the hairs in the furrows are obvious using at least a hand lens, and as ‘glabrous’ when the hairs are hidden. In the key below, ‘teeth’ refers to the reduced leaves at the apex of each branchlet article. Tooth measurements should be made on the elongated articles, not on new shoots where the teeth are often much longer. 1 Teeth 4–8 per whorl (to p. 117) 2 Branchlets 1–6 cm long with 1–5 elongate articles each 0.6–5 cm long, mostly pungent 3 Teeth 4, usually erect, occasionally scarious but without broad white margins 4 Teeth 0.3–0.6 mm long; samara with long ferruginous hairs, the wing absent or rudimentary; cone bracteoles without dorsal awn 4. A. grevilleoides 4: Teeth 0.8–5 mm long; samara glabrous, with well-developed wing; cone bracteoles with dorsal awn 1–9 mm long 5 Branchlets with 1 elongate article 15–50 mm long; cone bracteole protuberance with well-developed yellowish awn 1.3–9 mm long 6 Awn adherent to near apex of cone bracteole; slender plant with terete articles 0.6–0.9 mm diam. 2. A. acuaria 6: Awn separating from cone bracteole near base; coarse plant with quadrangular articles 1–1.2 mm diam. 3. A. pinaster 5: Branchlets with 2–4 elongate articles, each 8–16 mm long; cone bracteole protuberance with acuminate, awn-like apex c. 1 mm long (cone details obscured by tangled mass of greyish coarse hairs c. 1 cm long) 5. A. fibrosa 3: Teeth 5, slightly spreading, with deciduous broad white margins, 0.6–1.3 mm long 6. A. ramosissima 2: Branchlets mostly 8–50 cm long, with numerous elongate articles each 0.1–6 cm long, not pungent except A. decaisneana (if branchlets shorter than 8 cm long and/or if fewer than 7 elongated articles, then articles less than 1 cm long and penultimate branchlets green) 7 Penultimate as well as ultimate branchlets green; shrubs, intricate at maturity 8 Teeth 4; phyllichnia verruculose to serrate; articles 2–6 mm long; samara wingless, with hairs shorter than body of samara 58. A. microstachya 8: Teeth 6 or 7; phyllichnia smooth; articles 1.5–2.5 mm long; samara with small wing and hairs exceeding body of samara 57. A. drummondiana 7: Only ultimate branchlets green; penultimate branches slender but woody with brown to grey bark; trees or shrubs, rarely intricate 9 Cones warty or tessellated, with each cone bracteole divided into 3–20 small bodies 10 Articles quadrangular in T.S.; cone body diam. greater than or equal to length; pyramidal bodies 12–20 20. A. decussata 10: Articles ±terete in T.S. (occasionally quadrangular in A. torulosa); cone body length greater than or equal to diam.; pyramidal bodies 4–12 11 Teeth spreading [6–8] 21. A. fraseriana 11: Teeth erect to appressed 12 Teeth 8–10; articles 7–14 mm long 16. A. tessellata 12: Teeth 4–7; articles 3–7 mm long

112 CASUARINACEAE 3. Allocasuarina

13 Cone peduncle 8–30 mm long; tree with drooping branchlets 19. A. torulosa 13: Cone peduncle 0–4 mm long; shrub with stiff ascending branchlets 14 Phyllichnia angular to strongly rounded, usually with median ridge often minutely serrate; teeth overlapping at least at base when young 56. A. humilis 14: Phyllichnia flat to slightly rounded, often with smooth yellowish median line but no ridge; teeth not overlapping 15 Furrows pubescent; cone bracteole bodies acute to broadly acute; mature samara very dark red-brown to black (N.S.W., Vic.) 55. A. nana 15: Furrows glabrous; cone bracteole bodies truncate; mature samara pale red-brown (W.A., S.A.) 18. A. helmsii 9: Cones with single protuberance or with protuberance fused to bracteole body 16 Teeth pungent; articles 20–60 mm long; phyllichnia with median groove (eremean W.A., N.T., S.A.) 1. A. decaisneana 16: Teeth not pungent; articles 1–20 mm long, rarely with median groove, rarely to 25 mm long and then phyllichnia without median groove 17 Protuberance produced in a slender spine 1–8 mm long on at least some cones on a plant 18 Teeth 8–10 8. A. corniculata 18: Teeth 5 or 6 59. A. thuyoides 17: Protuberance acute to obtuse or fused with cone bracteole body or rarely both bracteole and protuberance acuminate and pungent 19 Phyllichnia verruculose or minutely serrate on median ridge 20 Phyllichnia rounded (strongly to slightly so), verruculose; trees 3–10 m high 21 Teeth spreading, 0.5–0.7 mm long; articles 8–13 mm long, 0.8–0.9 mm diam.; furrows pubescent; bark smooth or furrowed (W.A.) 23. A. huegeliana 21: Teeth erect, 0.3–0.5 mm long; articles 4–7 mm long, 0.5–0.6 mm diam.; furrows glabrous; bark on trunk extremely ribbony-fibrous, red-brown (Qld, N.S.W.) 25. A. inophloia 20: Phyllichnia strongly angular, minutely serrate on ridge; shrub 0.5–3 m high (S.A., Vic.) 54. A. muelleriana 19: Phyllichnia smooth, occasionally pubescent 22 Protuberance fused with bracteole body (often apex of protuberance discernible as a transverse line, occasionally with a tiny mucro) 23 Teeth erect; cones erect; mature samara black; shrub 1–3 m high 24 Phyllichnia rounded or nearly flat with small but definite median ridge; cones sessile or on peduncle to 5 mm long 14. A. campestris 24: Phyllichnia flat or slightly rounded without median ridge; cones on peduncle 4–15 mm long 15. A. eriochlamys 23: Teeth spreading; cones recurved relative to branch; mature samara mid-brown; tree 4–9 m high 17. A. dielsiana 22: Protuberance 1 (occasionally superficially 2- or 3-lobed in A. striata)

113 3. Allocasuarina CASUARINACEAE

25 Teeth 1.5–3.5 mm long 26 Teeth not overlapping at maturity 27 Teeth 7–10; cones sessile or on peduncle to 3 mm long 28 Articles 0.9–1 mm diam., glabrous; cone bracteoles pungent, with acute or acuminate apex and with dorsal protuberance of similar shape and size (W.A.) 24. A. trichodon 28: Articles 1.4–2 mm diam., with densely pubescent furrows; cone bracteoles obtuse to truncate, with dorsal protuberance thick-pyramidal and slightly shorter than bracteole body (Tas.) 43. A. crassa 27: Teeth 5 or 6; cones on peduncle 2–9 mm long (Qld) 28. A. filidens 26: Teeth overlapping 29 Teeth recurved to spreading, strongly marcescent 46. A. mackliniana 29: Teeth erect, marcescent only at apex 50. A. striata 25: Teeth less than 1.5 mm long 30 Phyllichnia with median groove along their length 31 Teeth overlapping (S.A., Vic.) 48. A. pusilla 31: Teeth not overlapping 32 Articles 2–5 mm long, 0.4–0.7 mm diam. (N.S.W.) 53. A. brachystachya 32: Articles 5–14 mm long, 0.7–1.2 mm diam. 33 Samara mid-brown, 5.5–6.0 mm long; teeth strongly marcescent (W.A.) 13. A. tortiramula 33: Samara very dark brown to black, 3.5–5.5 mm long; teeth not or occasionally marcescent 34 Furrows glabrous (W.A.) 44. A. lehmanniana 34: Furrows pubescent (S.A., N.S.W., Vic., Tas.) 52. A. paludosa 30: Phyllichnia without median groove 35 Teeth overlapping at least at base when young 36 Furrows obviously pubescent (pubescent and glabrous sometimes mixed on same plant in A. simulans) 37 Teeth recurved to spreading 46. A. mackliniana 37: Teeth erect 38 Phyllichnia only slightly rounded 51. A. gymnanthera 38: Phyllichnia angular to strongly rounded 39 Cones large, 11–22 mm diam., on stout peduncle 2–4 mm diam.; articles dark blue-green (Port Stephens–Cooma, N.S.W.) 35. A. distyla 39: Cones slender, 9–12 mm diam., on slender peduncle c. 2 mm diam.; articles pale yellow-green (Nabiac–Forster, N.S.W.) 36. A. simulans 36: Furrows glabrous 40 Phyllichnia and teeth flat or slightly rounded (occasionally more rounded towards base of article but always rather flattened near apex) 41 Usually monoecious 42 Cones on peduncle 2–10 mm long; cone bracteoles truncate to broadly acute (Tas.) 42. A. monilifera 42: Cones sessile or on peduncle to 3 mm long; cone bracteoles broadly acute to obtuse (S.A.) 49. A. robusta

114 CASUARINACEAE 3. Allocasuarina

41: Usually dioecious 43 Teeth recurved to spreading, strongly 46. A. mackliniana marcescent (S.A., Vic.) 43: Teeth erect to spreading, not marcescent or slightly so at apex 44 Mature samara red- to olive-brown, 3.0–3.5 mm long 45 Articles blue-green, 6–8 mm long; cone body 5–7 mm diam. (Nabiac, N.S.W.) 38. A. defungens 45: Articles yellow-green, 5–11 mm long; cone body 7–8 mm diam. (Penrith, N.S.W.) 39. A. glareicola 44: Mature samara very dark brown to black, 4–7 mm long 46 Cones shortly cylindrical, sessile; cone bracteoles broadly acute to obtuse (S.A., Vic.) 48. A. pusilla 46: Cones long-cylindrical, on peduncle 3–8 mm long; cone bracteoles truncate to obtuse (N.S.W.) 51. A. gymnanthera 40: Phyllichnia angular or strongly rounded, with ridge or convexity continued onto teeth 47 Mature samara red-brown to olive-brown 48 Teeth recurved to spreading, mostly marcescent; articles 10–25 mm long 49 Articles 0.7–1.5 mm diam.; phyllichnia angular to sometimes rounded 31. A. rigida 49: Articles 0.8–1 mm diam.; phyllichnia rounded to sometimes angular 32. A. portuensis 48: Teeth erect to slightly spreading, not marcescent; articles 5–12 mm long 50 Articles of greater diam. at apex than towards base 51 Teeth narrowly to broadly deltoid, ±straight-sided; articles often with a waxy bloom 37. A. diminuta 51: Teeth very broadly deltoid, very convex-sided; articles without waxy bloom 33. A. rupicola 50: Articles of same diam. throughout their length, not waxy 39. A. glareicola 47: Mature samara very dark brown to black 52 Phyllichnia rounded 53 Articles with a waxy bloom at least when dry; teeth erect, not marcescent; cones on peduncle 2–12 mm long 54 Articles of nearly same diam. throughout their length or slightly enlarged at apex 55 Dioecious; cone body 7–9 mm diam.; phyllichnia strongly rounded to angular (Vic.) 40. A. grampiana 55: Monoecious mostly; cone body 8–14 mm diam.; phyllichnia rounded to strongly rounded (Tas.)

42. A. monilifera

115 3. Allocasuarina CASUARINACEAE

54: Articles of markedly greater diam. at apex 50. A. striata 53: Articles without a waxy bloom; teeth recurved to erect, often marcescent; cones sessile or on peduncle to 5 mm long 56 Teeth 5–7, erect and appressed to occasionally spreading; articles 4–10 mm long, 0.3–0.8 mm diam. 47. A. misera 56: Teeth 7–11, recurved to spreading; articles 6–17 mm long, 0.6–1.2 mm diam. 57 Teeth spreading, mostly somewhat marcescent; furrows glabrous 45. A. paradoxa 57: Teeth recurved to spreading, strongly marcescent; furrows glabrous or densely pubescent 46. A. mackliniana 52: Phyllichnia angular 58 Articles with a strong waxy bloom (Vic.) 40. A. grampiana 58: Articles with a faint waxy bloom or none (Tas.) 41. A. zephyrea 35: Teeth not overlapping, mostly not marcescent 59 Furrows obviously pubescent (at least shortly) 60 Tree 5–15 m high 26. A. littoralis 60: Shrub or small tree 0.5–3 m high 61 Phyllichnia angular to strongly rounded, without median groove; dioecious 62 Phyllichnia strongly angular with white median ridge (often minutely serrate); teeth erect and appressed; articles of same diam. throughout their length (S.A., Vic.) 54. A. muelleriana 62: Phyllichnia rounded without well-defined white median ridge 63 Teeth spreading to recurved (Tas.) 43. A. crassa 63: Teeth erect but not appressed (Qld, N.S.W.) 64 Articles 10–22 mm long, 0.8–1.5 mm diam.; teeth thick-textured, 0.5–1.2 mm long 65 Cones 11–22 mm diam., on peduncle 2–4 mm diam.; articles dark blue-green 35. A. distyla 65: Cones 9–12 mm diam., on peduncle c. 2 mm diam.; articles pale yellow-green 36. A. simulans 64: Articles 4–12 mm long, 0.5–0.9 mm diam.; teeth thin-textured, 0.3–0.7 mm long 66 Articles 4–8 mm long 29. A. emuina 66: Articles 7–12 mm long 61: Phyllichnia flat or slightly rounded (often with a median groove towards base in A. brachystachya) 67 Cone body 14–40 mm long, 9–12 mm diam.; articles 5–16 mm long 51. A. gymnanthera 67: Cone body 7–14 mm long, 5–8 mm diam.; articles 2–5 mm long 53. A. brachystachya 59: Furrows glabrous 68 Mature samara red- to olive-brown

116 CASUARINACEAE 3. Allocasuarina

69 Articles of the same diam. throughout their length, not waxy; phyllichnia rounded (usually only slightly so near apex) 39. A. glareicola 69: Articles at least slightly enlarged in diam. at apex, often with a waxy bloom at least when dry; phyllichnia angular or strongly rounded 70 Articles without a waxy bloom; teeth often marcescent at apex 31. A. rigida 70: Articles often with a waxy bloom 71 Teeth marcescent; articles 13–20 mm long 32. A. portuensis 71: Teeth not marcescent; articles 5–14 mm long 72 Teeth narrowly deltoid; occurs on serpentinite outcrops 34. A. ophiolitica 72: Teeth broadly deltoid; occurs on siliceous soils 37. A. diminuta 68: Mature samara very dark brown to black 73 Teeth erect, appressed (occasionally some teeth spreading in A. misera) 74 Phyllichnia flat to rounded or broadly angular with pale median line; samara c. 4 mm long 75 Articles 0.3–0.8 mm diam. (Vic.) 47. A. misera 75: Articles 0.7–1.2 mm diam. 76 Articles 7–11 mm long; teeth 0.5–1.0 mm long; cones on peduncle 2–6 mm long; usually monoecious (Tas.) 42. A. monilifera 76: Articles 5–9 mm long; teeth 0.4–0.8 mm long; cones on peduncle 6–35 mm long or less commonly sessile; mostly dioecious (W.A.) 44. A. lehmanniana 74: Phyllichnia sharply angular with pale median ridge; samara 6–9 mm long (S.A., Vic.) 54. A. muelleriana 73: Teeth spreading, or erect but not appressed and often with some apices spreading 77 Monoecious usually 78 Cones on peduncle 2–10 mm long; articles often with a waxy bloom (Tas.) 42. A. monilifera 78: Cones sessile or on peduncle to 3 mm long; articles not waxy (S.A.) 49. A. robusta 77: Dioecious usually 79 Articles with a strong waxy bloom (Vic.) 40. A. grampiana 79: Articles with a faint waxy bloom or none 80 Articles 0.5–0.8 mm diam.; samara 5–8 mm long (Vic.) 27. A. media 80: Articles 0.6–1.3 mm diam.; samara 4–5.5 mm long (Tas.) 41. A. zephyrea 1: Teeth more than 8 per whorl 81 Articles verruculose 82 Articles 4–7 mm long, 0.5–0.6 mm diam.; bark very ribbony-fibrous (Qld, N.S.W.) 25. A. inophloia 82: Articles 8–40 mm long, 0.7–2 mm diam.; bark smooth to furrowed 83 Teeth spreading; phyllichnia rounded

117 3. Allocasuarina CASUARINACEAE

84 Articles 15–40 mm long, 0.7–1.5 mm diam.; branchlets strongly drooping (S.A., N.S.W., Vic., Tas.) 22. A. verticillata 84: Articles 8–13 mm long, 0.8–0.9 mm diam.; branchlets drooping to ascending (W.A.) 23. A. huegeliana 83: Teeth erect; phyllichnia flat or slightly rounded; branchlets ascending 85 Monoecious; cone bracteoles with protuberance produced in a long spine 4–11 mm long 9. A. spinosissima 85: Dioecious; cone bracteoles with protuberance obtuse or acuminate with mucro less than 1 mm long 86 Articles 1.0–2.0 mm diam.; cone diam. greater than length; cones 5–12 mm long; cone bracteoles with short obtuse dorsal protuberance (S.A., Qld, N.S.W., Vic.) 7. A. luehmannii 86: Articles 0.7–1.2 mm diam.; cone length greater than diam.; cones 15–35 mm long; cone bracteole and protuberance fused or divided into 2 or 3 acuminate or acute bodies (W.A.) 10. A. acutivalvis 81: Articles smooth 87 Dorsal protuberance fused with cone bracteole 88 Articles 20–52 mm long; teeth 1.3–2.7 mm long 11. A. scleroclada 88: Articles 6–13 mm long; teeth 0.6–1.2 mm long 14. A. campestris 87: Dorsal protuberance a separate pyramidal body, or cone bracteole divided into 3–5 bodies (dorsal protuberance not distinguishable) 89 Cone bracteole divided into 3–5 bodies; branchlets mostly with a thick waxy coating 90 Teeth 10–12; articles 17–28 mm long; furrows mostly shortly pubescent 12. A. globosa 90: Teeth 8–9; articles 7–14 mm long; furrows glabrous 16. A. tessellata 89: Dorsal protuberance a single body; branchlets occasionally with a waxy bloom (more obvious when dry) but not forming a thick waxy coating 91 Teeth 1.5–3.5 mm long 92 Phyllichnia angular; articles glabrous, 0.9–1 mm diam.; cone bracteoles acute to acuminate (W.A.) 24. A. trichodon 92: Phyllichnia rounded, occasionally only slightly so; cone bracteoles truncate to broadly acute 93 Phyllichnia moderately rounded (if only slightly so, then furrows densely pubescent); articles glabrous or pubescent in furrows, 0.7–1.3 mm diam. (S.A., Vic.) 46. A. mackliniana 93: Phyllichnia strongly rounded; articles pubescent in furrows, 1.2–2 mm diam. (Tas.) 43. A. crassa 91: Teeth less than 1.5 mm long 94 Protuberance produced in a spine 2–5 mm long (W.A.) 8. A. corniculata 94: Protuberance truncate to broadly acute, without spine 95 Furrows pubescent 96 Phyllichnia nearly flat to slightly rounded (S.A., Vic.) 46. A. mackliniana 96: Phyllichnia strongly rounded 43. A. crassa 95: Furrows glabrous 97 Mature samara red- to olive-brown

118 CASUARINACEAE 3. Allocasuarina

98 Articles without a waxy bloom; teeth often marcescent at apex (Qld, N.S.W.) 31. A. rigida 98: Articles with a waxy bloom at least when dry; teeth usually not marcescent 99 Teeth narrowly deltoid; occurs on serpentinite outcrops. 34. A. ophiolitica 99: Teeth broadly deltoid; occurs on siliceous soils 37. A. diminuta 97: Mature samara very dark brown to black 100 Cones on peduncle 2–15 mm long; teeth erect to spreading (Tas.) 101 Dioecious; phyllichnia angular to rounded 41. A. zephyrea 101: Monoecious; phyllichnia strongly to slightly rounded 42. A. monilifera 100: Cones sessile or on peduncle to 3 mm long; teeth recurved to spreading (S.A., Vic.) 102 Teeth spreading, mostly somewhat marcescent; furrows glabrous 45. A. paradoxa 102: Teeth recurved to spreading, strongly marcescent; furrows glabrous or densely pubescent 46. A. mackliniana

KEY TO SPECIES BASED ON MALE SPECIMENS

Male inflorescences are said to be moniliform if this state is reached by the time of anthesis (all male spikes may eventually elongate and become more or less moniliform with age). In the key below, ‘teeth’ refers to the reduced leaves at the apex of each branchlet article. 1 Teeth 4–8 per whorl (to p. 125) 2 Branchlets 1–6 cm long with 1–5 elongate articles, each 0.6–5 cm long, mostly pungent 3 Teeth 4, erect usually, occasionally scarious but without broad white margins 4 Teeth 0.3–0.6 mm long 4. A. grevilleoides 4: Teeth 0.8–5 mm long 5 Branchlets with 1 elongate article 15–50 mm long 6 Slender plant with articles 0.6–0.9 mm diam.; spikes elongate, moniliform, 1–4 cm long; anther 0.7–0.9 mm long 2. A. acuaria 6: Coarse plant with articles 1–1.2 mm diam.; spikes short, dense, 0.5–1 cm long; anther 1–1.4 mm long 3. A. pinaster 5: Branchlets with 2–4 elongate articles, each 8–16 mm long; spikes dense, 0.4–0.7 cm long; anther 0.5–0.8 mm long 5. A. fibrosa 3: Teeth 5, slightly spreading, with deciduous broad white margins, 0.6–1.3 mm long 6. A. ramosissima 2: Branchlets mostly 8–50 cm long with numerous elongate articles 0.1–6 cm long, not pungent except A. decaisneana (if branchlets less than 8 cm long and/or if fewer than 7 elongated articles, then articles less than 1 cm long and penultimate branchlets mostly green) 7 Penultimate as well as ultimate branchlets green; shrubs, intricate at maturity 8 Teeth 4; phyllichnia verruculose to serrate; articles 2–6 mm long 58. A. microstachya 8: Teeth 6–7; phyllichnia smooth; articles 1.5–2.5 mm long 57. A. drummondiana 7: Only ultimate branchlets green; penultimate branches slender but woody with brown to grey bark; trees or shrubs, rarely intricate 9 Phyllichnia verruculose or with minutely serrate median ridge

119 3. Allocasuarina CASUARINACEAE

10 Phyllichnia verruculose, strongly to slightly rounded; tree 3–15 m high 11 Teeth spreading; bark scaly or strongly furrowed (W.A.) 12 Male bracteoles persistent; teeth 8–10, 0.7–1.2 mm long 21. A. fraseriana 12: Male bracteoles deciduous, usually falling as a unit with the 2 perianth segments; teeth 6–8, 0.5–0.7 mm long 23. A. huegeliana 11: Teeth erect; bark on trunk extremely ribbony-fibrous, red-brown; male bracteoles deciduous (Qld, N.S.W.) 25. A. inophloia 10: Phyllichnia angular to strongly rounded with minutely serrate ridge; shrub 0.2–3 m high; teeth erect; bark smooth or finely fissured 54. A. muelleriana 9: Phyllichnia smooth 13 Male spikes ±moniliform, 6–8 whorls per cm; teeth not overlapping 54. A. muelleriana 13: Male spikes dense, 14–16 whorls per cm; teeth overlapping at base at least when young 56. A. humilis 14 Teeth 1.5–3.5 mm long 15 Teeth pungent; articles 20–60 mm long; phyllichnia with median groove 1. A. decaisneana 15: Teeth not pungent; articles 1–20 mm long; phyllichnia without median groove 16 Teeth not overlapping at maturity (may overlap on young shoots) 17 Teeth erect; articles mostly with a waxy coating 15. A. eriochlamys 17: Teeth spreading to recurved; articles not waxy 18 Teeth 7–10 19 Furrows glabrous; articles 12–15 mm long, 0.5–0.8 mm diam. (W.A.) 24. A. trichodon 19: Furrows pubescent; articles 10–20 mm long, 1.2–2 mm diam. (Tas.) 43. A. crassa 18: Teeth 5–6; articles 12–15 mm long, 0.5–0.8 mm diam. (Qld) 28. A. filidens 16: Teeth overlapping 20 Teeth recurved to spreading, strongly marcescent 46. A. mackliniana 20: Teeth erect, marcescent only at apex 50. A. striata 14: Teeth less than 1.5 mm long 21 Phyllichnia with median groove along their whole length 22 Teeth overlapping; anther 0.3–0.6 mm long (S.A., Vic.) 48. A. pusilla 22: Teeth not overlapping; anther 0.6–1.2 mm long 23 Articles 2–5 mm long, 0.4–0.7 mm diam. (N.S.W.) 53. A. brachystachya 23: Articles 5–14 mm long, 0.7–1.2 mm diam. 24 Male spikes less than 1 cm long; teeth strongly marcescent (W.A.) 13. A. tortiramula 24: Male spikes 1–6 cm long; teeth not or occasionally marcescent 25 Furrows glabrous (W.A.) 44. A. lehmanniana 25: Furrows pubescent (S.A., N.S.W., Vic., Tas.) 52. A. paludosa 21: Phyllichnia without median groove 26 Teeth overlapping at least at base when young

120 CASUARINACEAE 3. Allocasuarina

27 Furrows pubescent (pubescent and glabrous sometimes mixed on same individual in A. simulans ) 28 Teeth recurved to spreading (S.A., Vic.) 46. A. mackliniana 28: Teeth erect (W.A., N.S.W.) 29 Phyllichnia only slightly rounded 51. A. gymnanthera 29: Phyllichnia angular to strongly rounded 30 Articles 10–22 mm long 31 Articles dark blue-green; phyllichnia angular to rounded; generally parts coarser (Port Stephens–Cooma, N.S.W.) 35. A. distyla 31: Articles pale yellow-green; phyllichnia rounded to angular; generally parts more slender (Nabiac–Forster, N.S.W.) 36. A. simulans 30: Articles 3–6 mm long (W.A.) 56. A. humilis 27: Furrows glabrous 32 Phyllichnia and teeth flat or slightly rounded, occasionally more rounded towards base of article but always rather flattened near apex 33 Usually monoecious 34 Male spikes mostly moniliform (Tas.) 42. A. monilifera 34: Male spikes mostly not moniliform (S.A.) 49. A. robusta 33: Usually dioecious 35 Teeth recurved to spreading, strongly marcescent 46. A. mackliniana 35: Teeth erect to spreading, not marcescent or slightly so at apex 36 Male spikes moniliform, 3–5 whorls per cm 37 Male bracteoles persistent; male spikes 0.2–2.5 cm long 38 Articles blue-green, 6–8 mm long; spikes 0.4–1 cm long (Nabiac, N.S.W.) 38. A. defungens 38: Articles yellow-green, 5–11 mm long; spikes 0.2–2.5 cm long (Penrith, N.S.W.) 39. A. glareicola 37: Male bracteoles deciduous; male spikes 2.5–6 cm long 51. A. gymnanthera 36: Male spikes not moniliform, 8–11 whorls per cm 39 Articles without a waxy bloom; teeth 0.3–0.8 mm long, mostly brown 47. A. misera 39: Articles with a waxy bloom; teeth 0.3–0.5 mm long, mostly pale yellow 48. A. pusilla 32: Phyllichnia angular or strongly rounded, with ridge or convexity continued onto teeth 40 Mature male spikes moniliform 41 Monoecious 42 Phyllichnia rounded to angular (N.S.W.) 37. A. diminuta 42: Phyllichnia angular (Tas.) 42. A. monilifera 41: Dioecious 43 Articles not waxy; teeth erect 44 Male spike with 4–6.5 whorls per cm 31. A. rigida 44: Male spike with 7.5–8 whorls per cm 33. A. rupicola

121 3. Allocasuarina CASUARINACEAE

43: Articles with at least a faint waxy bloom; teeth spreading to erect or recurved 45 Teeth 6–7; articles with a waxy bloom 40. A. grampiana 45: Teeth 7–9; articles with a faint waxy bloom 46 Male spikes 1–3.5 cm long, 5–7 whorls per cm (Tas.) 41. A. zephyrea 46: Male spikes 5–10 cm long, 3.5–4.5 whorls per cm (N.S.W.) 32. A. portuensis 40: Mature male spikes dense, not moniliform 47 Male bracteoles deciduous 50. A. striata 47: Male bracteoles persistent 48 Male spikes very dense, 14–16 whorls per cm 56. A. humilis 48: Male spikes less dense, 3.5–11 whorls per cm 49 Teeth strongly marcescent, spreading or recurved, rarely erect 50 Articles 10–25 mm long; phyllichnia angular to rounded (Qld, N.S.W.) 31. A. rigida 50: Articles 6–7 mm long; phyllichnia rounded to angular (S.A., Vic.) 51 Male spikes with 3.5–7 whorls per cm, not appearing much thicker than vegetative branchlets 45. A. paradoxa 51: Male spikes with 6–9 whorls per cm, the recurved teeth making spikes appear about twice as thick as branchlets 46. A. mackliniana 49: Teeth not marcescent or occasionally so at very apex, erect or spreading 52 Anther 0.8–1.2 mm long; teeth 0.6–1 mm long, spreading to suberect 49. A. robusta 52: Anther 0.5–0.8 mm long; teeth 0.3–0.8 mm long, erect to spreading 53 Articles angular to sharply rounded, often with a waxy bloom, 0.6–1.1 mm diam.; teeth 6–9 (N.S.W.) 37. A. diminuta 53: Articles broadly rounded, not waxy, 0.3–0.8 mm diam.; teeth 5–7 (Vic.) 47. A. misera 26: Teeth not overlapping, mostly not marcescent 54 Phyllichnia and teeth flat to slightly rounded at least near apex 55 Tree 5–20 m high; bark corky or furrowed 56 Teeth 4 or 5; phyllichnia with pale median line 19. A. torulosa 56: Teeth 6–8; phyllichnia with small but definite pale median ridge 26. A. littoralis 55: Shrub 0.5–5 m high; bark smooth to finely fissured 57 Teeth 7–10, marcescent 58 Male spikes with 10–16 whorls per cm 8. A. corniculata 58: Male spikes with 7–11 whorls per cm 59 Articles with small but definite median ridge on phyllichnia 14. A. campestris 59: Articles without ridge on phyllichnia 15. A. eriochlamys

122 CASUARINACEAE 3. Allocasuarina

57: Teeth 4–6, mostly not marcescent, rarely to 8 and then not marcescent 60 Furrows glabrous 61 Anther 0.7–1.2 mm long; articles 5–14 mm long 62 Male spikes usually moniliform; anther 0.6–0.9 mm long; teeth 0.2–0.7 mm long 63 Teeth 0.2–0.5 mm long, erect to slightly spreading (N.S.W.) 39. A. glareicola 63: Teeth 0.4–0.7 mm long, appressed to erect (W.A.) 44. A. lehmanniana 62: Male spikes usually not moniliform; anther 0.8–1.2 mm long; teeth 0.6–1 mm long (S.A.) 49. A. robusta 61: Anther 0.4–0.7 mm long; articles 1–7 mm long; teeth 0.3–0.5 mm long; male spikes not moniliform 64 Male spikes 5–25 mm long, 9–12 whorls per cm; articles 3–7 mm long 18. A. helmsii 64: Male spikes 1–5 mm long, of only 1–4 whorls; articles 1–3 mm long 59. A. huyoides 60: Furrows pubescent 65 Male spikes 2.5–6 cm long, moniliform, 3–5 whorls per cm; articles 5–12 mm long 51. A. gymnanthera 65: Male spikes 0.5–1.7 cm long, not moniliform, 9–20 whorls per cm 66 Usually monoecious; articles 2–5 mm long, not waxy; phyllichnia often with median groove towards base 53. A. brachystachya 66: Usually dioecious; articles 5–6 mm long, often waxy; phyllichnia without median groove 55. A. nana 54: Phyllichnia angular to strongly rounded, with ridge or convexity continuing onto teeth 67 Male inflorescence very short, head-like, c. 0.5–0.6 cm diam.; tree or tall shrub 17. A. dielsiana 67: Male inflorescence an elongate spike, 1–9 cm long, rarely less than 1 cm long and then a shrub 68 Tree 5–15 m high; bark corky or strongly fissured 69 Articles quadrangular; teeth 4; monoecious 20. A. decussata 69: Articles terete; teeth 6–8; usually dioecious 70 Teeth spreading, 0.7–1.2 mm long (W.A.) 21. A. fraseriana 70: Teeth usually erect, 0.3–0.9 mm long (Qld to Tas.) 26. A. littoralis 68: Shrub or small tree 0.5–5 m high; bark smooth or slightly fissured 71 Male spikes with 12–16 whorls per cm, 6–18 mm long 56. A. humilis 71: Male spikes with 4–11 whorls per cm, 1–9 cm long 72 Furrows glabrous 73 Male spikes moniliform 74 Teeth erect, appressed

123 3. Allocasuarina CASUARINACEAE

75 Phyllichnia rounded with pale median line (W.A.) 44. A. lehmanniana 75: Phyllichnia angular with well-defined white median ridge (S.A., Vic.) 54. A. muelleriana 74: Teeth spreading or erect but not appressed and often with some apices spreading 76 Usually monoecious 42. A. monilifera 76: Usually dioecious 77 Articles 0.5–0.7 mm diam., with phyllichnia rounded, without a waxy bloom; teeth 0.2–0.5 mm long (N.S.W.) 39. A. glareicola 77: Articles 0.6–1.3 mm diam., with phyllichnia angular or occasionally rounded; teeth 0.4–1.2 mm long 78 Articles with a waxy bloom (Vic.) 40. A. grampiana 78: Articles with only a faint waxy bloom or none 79 Teeth recurved to erect, mostly marcescent (Qld, N.S.W.) 80 Articles 10–14 mm long; teeth erect to slightly spreading 31. A. rigida 80: Articles 13–20 mm long; teeth recurved to spreading 32. A. portuensis 79: Teeth erect to spreading, mostly not marcescent (Tas.) 41. A. zephyrea 73: Male spikes dense, not moniliform 81 Teeth marcescent (W.A.) 16. A. tessellata 81: Teeth not marcescent or occasionally so at very apex 82 Teeth spreading to erect, mostly at least the apex spreading; phyllichnia strongly rounded to angular without well-defined median ridge 83 Articles 0.5–0.8 mm diam., of nearly same diam. throughout length, without a waxy bloom (Vic.) 27. A. media 83: Articles 0.6–1.3 mm diam., mostly enlarged at apex, often with a waxy bloom 84 Anther 0.7–1.2 mm long; teeth narrowly deltoid 85 Anther 0.8–1.2 mm long (N.S.W.) 34. A. ophiolitica 85: Anther 0.7–0.9 mm long (Tas.) 41. A. zephyrea 84: Anther 0.5–0.8 mm long; teeth broadly deltoid (N.S.W.) 37. A. diminuta 82: Teeth erect, appressed to somewhat incurved 86 Phyllichnia rounded with pale median line 87 Male spikes with 5–8 whorls per cm; articles 0.7–1.2 mm diam., occasionally with a waxy bloom (W.A.) 44. A. lehmanniana 87: Male spikes with 9–11 whorls per cm; articles 0.3–0.8 mm diam., without a waxy bloom (Vic.) 47. A. misera

124 CASUARINACEAE 3. Allocasuarina

86: Phyllichnia strongly angular with well-defined median ridge (S.A., Vic.) 54. A. muelleriana 72: Furrows pubescent 88 Teeth marcescent; articles usually with a strong waxy coating 14. A. campestris 88: Teeth not marcescent; articles sometimes somewhat waxy 89 Male whorls 3.5–6.5 per cm; articles 10–22 mm long 90 Articles dark blue-green; phyllichnia angular to rounded 35. A. distyla 90: Articles pale yellow-green; phyllichnia rounded to angular 91 Articles 0.9–1.3 mm diam.; teeth 6, 0.5–1.1 mm long (N.S.W.) 36. A. simulans 91: Articles 1.2–2 mm diam,; teeth 7–10, 1.1–3 mm long (Tas.) 43. A. crassa 89: Male whorls 7–9.5 per cm, rarely 6 and then phyllichnia strongly angular; articles 3–12 mm long 92 Phyllichnia and male bracts strongly angular (S.A., Vic.) 54. A. muelleriana 92: Phyllichnia and male bracts angular to strongly rounded (Qld) 93 Articles 4–8 mm long; anther 0.8–0.9 mm long 29. A. emuina 93: Articles 7–12 mm long; anther 0.6–0.8 mm long 1: Teeth more than 8 per whorl 94 Articles verruculose 95 Articles short and slender, 4–7 mm long, 0.5–0.6 mm diam.; bark very ribbony-fibrous 25. A. inophloia 95: Articles 8–40 mm long, 0.7–2 mm diam.; bark smooth to furrowed 96 Teeth spreading; phyllichnia rounded; branchlets drooping; male bracteoles usually falling as a unit with the 1 or 2 perianth segments 97 Articles 15–40 mm long, 0.7–1.5 mm diam.; anther 1.2–2.5 mm long (S.A., N.S.W. to Tas.) 22. A. verticillata 97: Articles 8–13 mm long, 0.8–0.9 mm diam.; anther 0.8–1.2 mm long (W.A.) 23. A. huegeliana 96: Teeth erect; phyllichnia flat or slightly rounded; branchlets ascending; male bracteoles falling individually 98 Monoecious; male bracteoles persistent; male spikes dense, 7 or 8 whorls per cm; articles 1–1.8 mm diam. 9. A. spinosissima 98: Dioecious 99 Male bracteoles deciduous; male spikes with 5–8 whorls per cm; articles 1.0–2.0 mm diam. (S.A., Qld to Vic.) 7. A. luehmannii 99: Male bracteoles persistent; male spikes usually less dense, with 3.5–6 whorls per cm; articles 0.7–1.2 mm diam. (W.A.) 10. A. acutivalvis 94: Articles smooth 100 Articles 20–52 mm long

125 3. Allocasuarina CASUARINACEAE

101 Teeth not overlapping 102 Teeth 1.1–3 mm long 103 Teeth 10–12; phyllichnia flat to slightly rounded (W.A.) 11. A. scleroclada 103: Teeth 7–10; phyllichnia strongly rounded (Tas.) 43. A. crassa 102: Teeth 0.6–1 mm long, 10–12 (W.A.) 12. A. globosa 101: Teeth overlapping, 7–10 (Qld, N.S.W.) 31. A. rigida 100: Articles less than 20 mm long 104 Teeth 1.5–3.5 mm long, not overlapping 105 Phyllichnia angular; articles glabrous, 0.9–1.1 mm diam. (W.A.) 24. A. trichodon 105: Phyllichnia rounded, occasionally only slightly so 106 Phyllichnia rounded, if only slightly so then furrows densely pubescent; articles 0.7–1.3 mm diam. (S.A., Vic.) 46. A. mackliniana 106: Phyllichnia strongly rounded; articles pubescent in furrows, 1.2–2 mm diam. (Tas.) 43. A. crassa 104: Teeth less than 1.5 mm long 107 Phyllichnia flat or slightly broadly rounded 108 Teeth not overlapping; articles mostly with a waxy coating (W.A.) 109 Articles 6–8 mm long; teeth 8–10 110 Male spikes with 10–16 whorls per cm 8. A. corniculata 110: Male spikes with 7–11 whorls per cm 111 Articles with a small but definite median ridge on phyllichnia 14. A. campestris 111: Articles without a ridge on phyllichnia 15. A. eriochlamys 109: Articles 17–28 mm long; teeth 10–12 12. A. globosa 108: Teeth overlapping; articles without a waxy coating or bloom (S.A., Vic.) 46. A. mackliniana 107: Phyllichnia angular or strongly rounded 112 Male spikes moniliform; teeth erect 113 Teeth recurved to spreading, mostly marcescent (Qld, N.S.W.) 31. A. rigida 113: Teeth erect to spreading, mostly not marcescent (Tas.) 114 Usually dioecious; phyllichnia angular 41. A. zephyrea 114: Usually monoecious; phyllichnia rounded (strongly to slightly so) 42. A. monilifera 112: Male spikes usually dense, occasionally moniliform but then teeth recurved to spreading 115 Teeth marcescent 116 Teeth erect, not overlapping (W.A.) 117 Anther broader than long; male spikes with 7–8 whorls per cm 16. A. tessellata 117: Anther longer than broad; male spikes with 8.5–11 whorls per cm 14. A. campestris 116: Teeth recurved to spreading, usually overlapping 118 Male spikes with 6–9 whorls per cm (S.A., Vic.) 46. A. mackliniana 118: Male spikes with 4–6.5 whorls per cm 119 Articles 10–25 mm long (Qld, N.S.W.) 31. A. rigida 119: Articles 6–14 mm long (Vic.) 45. A. paradoxa

126 CASUARINACEAE 3. Allocasuarina

115: Teeth generally not marcescent 120 Anther 0.7–1.2 mm long; teeth narrowly deltoid 121 Anther 0.8–1.2 mm long (N.S.W.) 34. A. ophiolitica 121: Anther 0.7–0.9 mm long (Tas.) 41. A. zephyrea 120: Anther 0.5–0.8 mm long; teeth broadly to narrowly deltoid 37. A. diminuta

Sect. 1. Dolichopitys

Allocasuarina sect. Dolichopitys L.Johnson, Fl. Australia 3: 192 (1989). Type: A. decaisneana (F.Muell.) L.Johnson Dioecious trees; penultimate branchlets woody. Bark thickly furrowed. Branchlets not waxy; elongate articles numerous, terete or quadrangular, smooth; teeth usually not overlapping. Male spikes elongate; bracteoles persistent. Cone bracts prominent, splitting at maturity so as to appear as isolated crescent-shaped bodies below bracteoles; bracteoles with protuberance entire. Samaras glabrous, dark brown to black. A section of 1 species in eremean W.A., N.T. and S.A.

1. Allocasuarina decaisneana (F.Muell.) L.Johnson, J. Adelaide Bot. Gard. 6: 74 (1982) Casuarina decaisneana F.Muell., Fragm. 1: 61 (1858). T: near Mt Mueller, W.A., F.Mueller; holo: MEL (♀), not found; iso: K, vegetative. Illustrations: J.P.Jessop (ed.), Fl. Central Australia 15 (1981); D.J.Boland et al., Forest Trees Australia 4th edn, 104 (1984). Tree 10–16 m high. Branchlets drooping, to 50 cm long; articles 2–6 cm long, 0.7–1.5 mm diam., pubescent becoming glabrous; phyllichnia flat with broad shallow median groove; teeth 4, pungent, usually erect, 1.7–3.2 mm long. Male spikes 2–4 cm long, c. 10 whorls per cm; anther c. 0.7–0.8 mm long. Cones very woody, long-cylindrical, often pubescent when young, becoming glabrous; peduncle 5–15 mm long; cone body 28–95 mm long, 20–35 mm diam.; bracteoles acute, protuberance acute, nearly as long as bracteole body. Samara 8.5–17 mm long. 2n = 28, B.A.Barlow, Autral. J. Bot. 7: 232 (1959). Desert Oak, Desert Sheoak. Figs 24, 48A–B. Occurs sporadically from Billiluna, W.A., south-east to the Mann and Musgrave Ranges, S.A.; grows chiefly in swales between sand dunes. Regions: SNDY, TNMI, GBSN, MACD. Map 114. W.A.: 75 km S of Giles, D.E.Symon 2541 (AD, NSW); 11 km N of No. 11 Well, Canning Stock Route, H.M.Wilson 18 (PERTH). N.T.: 56 km S of Central Mount Wedge, H.S.McKee 8574 (K, NSW); 1 km S of Mount Santa, J.R.Maconochie 2281 (AD, B, CBG, K, NT, NY). S.A.: Amata, Musgrave Ra., P.L.Milthorpe 3087 (NSW).

Sect. 2. Oxypitys

Allocasuarina sect. Oxypitys L.Johnson, Fl. Australia 3: 193 (1989). Type: A. acuaria (F.Muell.) L.Johnson Dioecious shrubs; penultimate branchlets green or woody. Bark smooth, flaky or striate. Branchlets usually pungent, not waxy; elongate articles 1–5, terete or quadrangular, smooth; teeth overlapping or not, marcescent or not. Male spikes often only shortly elongate; bracteoles persistent. Cone bracts prominent, thickened; bracteoles with protuberance entire and awned, or divided. Samaras glabrous or hairy, dark brown to black. A section of 5 species in south-western W.A.

127

Figure 48. Allocasuarina cones ×1.5. A–B, A. decaisneana. A, cone (B.Briggs 3553, NSW); B, branchlet ×7.5 (D.Symon 10388, NSW). C–E, A. acuaria. C, branching habit ×1.5; D, cone; E, branchlet ×3 (C–E, E.Bennett 1578, NSW). F–H, A. pinaster. F, branching habit × 1.5 (F & H, R.Saffrey 429, NSW); G, cone (NSW 105051, NSW); H, branchlet ×3. Drawn by D.Mackay.

128 CASUARINACEAE 3. Allocasuarina

2. Allocasuarina acuaria (F.Muell.) L.Johnson, J. Adelaide Bot. Gard. 6: 74 (1982) Casuarina acuaria F.Muell., Fragm. 6: 16, t. 54 (1867); C. oxyclada Miq. in DC., Prodr. 16(2): 344 (1868), nom. illeg. based on C. acuaria Endl. T: south-western W.A., J.Drummond 241; lecto: MEL 540117, with cones, fide L.A.S.Johnson, J. Adelaide Bot. Gard. 6: 74 (1982); isolecto: BM, CGE, K, LE, P. Shrub 1–3 m high; penultimate branchlets woody. Branchlets ascending, pungent, 1.5–3 cm long, with 1 elongate article; article terete, 1.5–3 cm long, 0.6–0.9 mm diam., pubescent when young; phyllichnia rounded with broad median groove; teeth 4, erect, or spreading to recurved on young plants and new shoots, slightly or not overlapping, 0.8–1.3 mm long, ±marcescent. Male spikes moniliform, 1–4 cm long, 4.5–5.5 whorls per cm; anther 0.7–0.9 mm long. Cones ovoid to subglobose; peduncle 4–10 mm long; cone body 15–19 mm long (20–26 mm long including bracteoles), 13–18 mm diam., shortly and densely hairy; bracts glabrous; bracteoles glabrous, acute, protuberance forming a yellowish ridge on bracteole body adnate to near apex, extending as a pungent eccentric awn 4–9 mm long, often with hooked tip. Samara c. 6 mm long, black. 2n = 28, B.G.Briggs (ex Kings Park, W.A., s.n.). Fig. 48C–E. Occurs in the Tambellup–Ravensthorpe area, W.A.; in heath in sand. Regions: BENC, ESPR. Map 115. W.A.: just S of Tambellup, E.M.Bennett 1578 (NSW, PERTH); Pingrup, 24 Sept. 1932, W.E.Blackall (NSW, PERTH); 38 km NE of Ongerup, K.Newbey 3029 (NSW, PERTH). The shape of the cone is obscured by the elongate bracteoles. The species is more slender than A. pinaster.

3. Allocasuarina pinaster (C.Gardner) L.Johnson, J. Adelaide Bot. Gard. 6: 77 (1982) Casuarina pinaster C.Gardner, J. Proc. Roy. Soc. W. Australia 27: 166 (1942). T: Kukerin, W.A., 13 Sept. 1925, C.A.Gardner 1732; lecto: PERTH♀, ( with cones); fide L.A.S.Johnson, J. Adelaide Bot. Gard. 6: 77 (1982). Similar to A. acuaria. Shrub 1–3 m high. Branchlets 2–6 cm long with 1 elongate article; article quadrangular, 2–5 cm long, 1–1.2 mm diam., usually pubescent; phyllichnia flat or with shallow median groove; teeth 4, erect, 4.5–5.0 mm long. Male spikes 5–10 mm long, 8–10 whorls per cm; anther 1–1.4 mm long. Cones sessile or on peduncle to 5 mm long; cone body 14–25 mm long, 12–16 mm diam.; bracteole apex with small deciduous mucro, protuberance separating from bracteole body near base, extending into yellowish awn 3–7 mm long and about as long as or exceeding bracteole apex. Samara 10–11 mm long, dark brown to black. 2n = 28, B.A.Barlow, Austral. J. Bot. 7: 232 (1959); B.G.Briggs pers. comm. (ex Kings Park, s.n.). Compass Bush. Fig. 48F–H. Restricted to the Hyden–Dumbleyung area, W.A.; in tall shrubland and heathland in lateritic soils. Region: ESPR. Map 116. W.A.: 28 km W of Lake Grace, A.S.George 349 (NSW, PERTH); 21–22 km E of Newdegate, A.S.George 522 (PERTH); 13 km N of ‘The Humps’, NE of Hyden, B.R.Maslin 568 (MEL, PERTH); Tarin Rock, B.Rockel CPIN2, CPIN3A (FRI, NSW); near Commander Rocks, R.A.Saffrey 429 (K, NSW, PERTH). Similar to A. acuaria, but generally larger. Female plants have a distinct main stem and a number of main branches aligned parallel to it, the whole axis thus formed being inclined at 30–40° from the vertical and in a consistent southerly direction; male plants are smaller and are more spreading with no main stem and branchlets less clustered (J.S.Beard, W. Austral. Nat. 11: 73–75, 1969).

4. Allocasuarina grevilleoides (Diels) L.Johnson, J. Adelaide Bot. Gard. 6: 75 (1982) Casuarina grevilleoides Diels in A.Engler, Bot. Jahrb. Syst. 35: 130, figs 11J–M (1905). T: Babilion Ra. [Babilion Hills], W.A., 30 Aug. 1901, L.Diels 4019; holo: B (♂, ♀ - monoecious). Spreading, lignotuberous shrub, 15–30 cm high; penultimate branchlets green. Branchlets ascending, ±pungent, 1–3 cm long; elongate article(s) 1 or 2, terete to subquadrangular, 10–24 mm long, 0.5–0.8 mm diam., often pubescent when young; phyllichnia flat or with

129 3. Allocasuarina CASUARINACEAE broad shallow median groove; teeth 4, erect to slightly spreading, overlapping, usually shortly fimbriate, 0.3–0.6 mm long, marcescent. Male spikes 2–5 mm long, dense; anther 0.6–0.7 mm long. Cones inconspicuous, ovoid, irregular, sessile, densely and minutely pubescent, becoming glabrous; cone body 9–14 mm long, 6–9 mm diam.; bracts with deciduous awn c. 1.5 mm long; bracteoles greatly thickened but relatively small, acute, protuberance much shorter than bracteole body, divided into 4–6 small pyramidal bodies. Samara 5.5–7 mm long, very dark brown, with long ferruginous hairs. Figs 45H, 49A–C. Restricted to the Jurien Bay–Mogumber area, W.A.; in heath. Regions: LUWN, BENC. Map 117. W.A.: 1.6 km S of Watheroo, E.M.Bennett 1657 (NSW, PERTH); Moore R., L.Diels & E.Pritzel 527 (NSW, PERTH); between Dandaragan and Hill R., C.A.Gardner 9016 (NSW, PERTH); Mogumber, C.A.Gardner 1890 (MEL, PERTH); Mogumber, Jan. 1936, C.A.Gardner (NSW, PERTH). This species has an unusual samara that is usually wingless and very plump, with a dense mass of long (5–7 mm) ferruginous septate hairs (readily deciduous).

5. Allocasuarina fibrosa (C.Gardner) L.Johnson, J. Adelaide Bot. Gard. 6: 75 (1982) Casuarina fibrosa C.Gardner, J. Proc. Roy. Soc. W. Australia 13: 61, t. 24A–F (1927). T: Tammin, W.A., Aug. [cited as Sept.] 1926, C.A.Gardner s.n.; holo: PERTH ♀,( with inflorescences and cones); iso: MEL. Similar to A. grevilleoides. Shrub 0.5–1.5 m high; penultimate branchlets woody. Branchlets 2–5 cm long; elongate articles 2–4, terete, 8–16 mm long, glabrous; teeth 4, erect, with broad whitish translucent margins at least towards apex, 1.5–2.0 mm long. Male spikes 4–7 mm long; anther 0.5–0.6 mm long. Cones subcylindrical to globose, with long coarse hairs; cone body 11–25 mm long, 9–11 mm diam.; bracteoles thin, long-acuminate, often awn-like (c. 1 mm long) and hooked, with thick pyramidal protuberance separating from bracteole near base, with short curved awn 1–2 mm long. Samara 6–7 mm long, glabrous, brown to black. Fig. 49D–F. Only one population known, near Tammin, W.A.; in tall open-heath in sand over laterite on a low ridge. Region: BENC. Map 118. W.A.: Tammin, C.A.Gardner 12117 (NSW, PERTH); Tammin, July 1949, C.A.Gardner (K, NSW, PERTH); Charles Gardner Natl Park, A.S.George 10891 p.p. (NSW, PERTH). This species has a cone with long (10–20 mm) tangled coarse hairs on the body, protruding from between the bracts and bracteoles. Listed as an endangered species.

6. Allocasuarina ramosissima (C.Gardner) L.Johnson, J. Adelaide Bot. Gard. 6: 78 (1982) Casuarina ramosissima C.Gardner, J. Proc. Roy. Soc. W. Australia 47: 54 (1964). T: c. 14 km N of Dandaragan, W.A., 23 Aug. 1948, C.A.Gardner 9013; holo: PERTH ♀,( with inflorescences & cones). Somewhat divaricate shrub c. 1 m high; penultimate branchlets green. Branchlets ascending, verticillate, 1–6 cm long; elongate articles 1–5, terete to subangular, 6–20 mm long, 0.6–1 mm diam., glabrous or pubescent especially in furrows; phyllichnia rounded, lacking median groove; teeth 5, ascending to spreading, markedly overlapping, 0.6–1.3 mm long, marcescent, dark brown with broad white erose margins. Male spikes head-like, 1.5–2.0 mm long, with prominent bracts; anther 0.6–0.7 mm long. Cones ovoid to cylindrical, sessile, with short coarse hairs 1–2 mm long around bracts and bracteoles; cone body 9–13 mm long, 7–9 mm diam.; bracts densely pubescent when young, with deciduous dark brown broadly acute apex with broad erose margins; bracteoles inconspicuous, truncate, with small protuberance apparently superficially divided. Samara 4.5–5.0 mm long, dark brown (including short wing), shortly pubescent on sides of body at least towards base. Fig. 49G–I. Restricted to Badgingarra–Dandaragan area, W.A.; in heath in sand. Region: LUWN. Map 119. W.A.: Tootbardi Rd, NE of Coomallo, M.G.Corrick 8305 (MEL, NSW); 11 km SE of Badgingarra, K.L.Wilson 2707, 2708 (NSW, PERTH).

130

Figure 49. Allocasuarina cones ×1.5. A–C, A. grevilleoides. A, branching habit ×1.5; B, cone; C, branchlet ×3 (A–C, C.A.Gardner, Jan. 1936, NSW). D–F, A. fibrosa. D, branching habit ×1.5; E, cone; F, branchlet ×3 (D–F, C.A.Gardner, July 1949, NSW). G–I, A. ramosissima. G, branching habit ×1.5; H, cone; I, branchlet ×5 (G–I, A.George 6428, NSW). J–K, A. luehmannii. J, cone (NSW 50717, NSW); K, branchlet ×7.5 (NSW 16401, NSW). Drawn by D.Mackay.

131 3. Allocasuarina CASUARINACEAE

Cone features are obscured by the very prominent cone bracts and ferruginous hairs and deciduous coarse grey hairs or hair-like appendages.

Sect. 3. Platypitys

Allocasuarina sect. Platypitys L.Johnson, Fl. Australia 3: 193 (1989). Type: A. luehmannii (R.Baker) L.Johnson Dioecious trees; penultimate branchlets woody. Bark furrowed. Branchlets often waxy; elongate articles numerous, terete, verruculose; teeth slightly overlapping, usually marcescent. Male spikes elongate; bracteoles deciduous individually. Cone bracts inconspicuous; bracteole protuberance entire. Samaras glabrous, red-brown. A section of 1 species in inland eastern Australia.

7. (R.Baker) L.Johnson, J. Adelaide Bot. Gard. 7: 314 (1985) Casuarina luehmannii R.Baker, Proc. Linn. Soc. New South Wales 24: 608, t. 47 (1900). T: Grenfell, N.S.W., [no date], R.H.Cambage NSW 50724; lecto: NSW ♂,( with cones), fide L.A.S.Johnson, Fl. Australia 3: 201 (1989). Illustrations: J.H.Maiden, Forest Fl. New South Wales 2: t. 76 (1905); L.Costermans, Native Trees & Shrubs S.E. Australia 146–147 (1981); G.M.Cunningham et al., Pl. W. New South Wales 208 (1982). Tree 5–15 m high. Branchlets ascending, to 40 cm long; articles 8–22 mm long, 1–2 mm diam., finely pubescent especially in furrows when immature; phyllichnia flat to slightly rounded, verruculose; teeth 10–14, erect, 0.5–1.0 mm long. Male spikes 1.5–4.5 cm long, 5–8 whorls per cm; anther 1–1.3 mm long. Cones very shortly cylindrical (broader than long), pubescent at least when young, sessile or on peduncle to 5 mm long; cone body 5–12 mm long, 8–14 mm diam.; bracteoles acute to obtuse, protuberance pyramidal or flattened, obtuse, much shorter than bracteole body and diverging from it near its base. Samara 4.5–5.0 mm long. 2n = 56, B.A.Barlow, Austral. J. Bot. 7: 232 (1959). Bull Oak. Fig. 49J–K. Occurs from Mareeba, Qld, S through central Qld and N.S.W. to north-western Vic. and adjoining areas of S.A. to S of Bordertown; rarely towards the coast, as in the Hunter Valley, N.S.W., and near Rockhampton, Qld; also in A.C.T. Scattered in woodland in non-calcareous soils. Regions: CYRK, BURD, DWSN, WRGO, MCPH, DARL, NEPN, RIVR, HOWE. Map 120. S.A.: near Bordertown towards Wolseley, I.Ridway ADW 42277 (AD, CANB, NSW). Qld: 20 km N of Mareeba on road to Mt Molloy, J.R.Clarkson 5704 (BRI, QRS, NSW). N.S.W.: 4 km SE of Mt Wambo, 22 km WSW of Singleton, R.G.Coveny 5615 & S.W.L.Jacobs (A, BO, FI, K, L, MEL, NSW, RSA, US); Bebo State Forest, E of Yetman, R.G.Coveny 11693 & P.G.Wilson (BRI, K, NSW). Vic.: 10 km W of Echuca, R.Melville 3905 (K, NSW).

Sect. 4. Echinopitys

Allocasuarina sect. Echinopitys L.Johnson, Fl. Australia 3: 192 (1989). Type: A. corniculata (F.Muell.) L.Johnson Dioecious or monoecious shrubs; penultimate branchlets woody. Bark smooth to striate or flaky. Branchlets waxy; elongate articles numerous, terete, smooth or verruculose; teeth not or slightly overlapping, marcescent. Male spikes elongate; bracteoles persistent. Cone bracts inconspicuous; bracteoles with spine-like entire protuberance. Samaras glabrous, red-brown to black. A section of 2 species in south-western Australia.

132 CASUARINACEAE 3. Allocasuarina

8. Allocasuarina corniculata (F.Muell.) L.Johnson, J. Adelaide Bot. Gard. 6: 74 (1982) Casuarina corniculata F.Muell., Fragm. 10: 62 (1876). T: near Mt Churchman, W.A., 1875, D.Young; holo: MEL, with cones; iso: B. Casuarina horrida D.A.Herbert, J. Proc. Roy. Soc. W. Australia 7: 87 (1921). T: Burracoppin [published as Merredin], W.A., Nov. 1920, E.Wilson [99] & D.A.Herbert; holo: PERTH, with cones; iso: A, UC. Dioecious shrub, erect to spreading, 2–3 m high. Branchlets ascending, to 26 cm long; articles 6–11 mm long, rarely to 16 mm, 0.9–1.4 mm diam., smooth, glabrous; phyllichnia flat or slightly rounded; teeth 8–10, erect to slightly spreading, not overlapping, 0.3–0.6 mm long. Male spikes 0.5–2 cm long, 10–16 whorls per cm; anther 0.5–0.9 mm long. Cones cylindrical, occasionally irregular, sparsely pubescent to glabrous, sessile or on peduncle to 3 mm long; cone body 8–15 mm long, 7–9 mm diam.; bracteoles broadly acute to obtuse, pyramidal protuberance separating near base of bracteole body and with a deciduous slender yellowish awn, usually pungent, 2–5 mm long, with hooked or curved apex. Samara 3–4 mm long, red-brown. Fig. 50A. Occurs from Wubin SE to Norseman, W.A.; in tall heath on sandplain. Regions: BENC, LEON, ESPR. Map 121. W.A.: 62 km NE of Wubin, M.G.Corrick 9311 (MEL, NSW); 1 km E of Cleary on road to Beacon, D.B.Foreman 678 (K, MEL, NSW, PERTH); Queen Victoria Rock, SW of Coolgardie, A.S.George 8038 (NSW, PERTH); W of Burracoppin, R.Melville 123B & C.A.Gardner (K, NSW); 90 Mile Tank, c. 80 km W of Daniell, P.G.Wilson 3198 (AD, NSW).

9. Allocasuarina spinosissima (C.Gardner) L.Johnson, Fl. Australia 3: 199 (1989) Casuarina spinosissima C.Gardner, J. Proc. Roy. Soc. W. Australia 9: 38 (1923). T: Carrabin, W.A., 6 Oct. 1922, C.A.Gardner 1773 [published as 1273 (original collecting no.)]; holo: PERTH, with cones; iso: MEL. Similar to A. corniculata. Monoecious shrub 2–4 m high. Branchlets with articles 8–13 mm long, 1–1.8 mm diam., verruculose; teeth 9–11, erect, slightly overlapping at bases, 0.5–1.0 mm long. Male spikes 1.5–3 cm long, 7 or 8 whorls per cm; anther 0.8–1.3 mm long. Cones ovoid to cylindrical, sessile or on peduncle to 6 mm long; cone body 10–23 mm long, 8–12 mm diam.; protuberance usually extending into a stout, often deciduous awn 4–11 mm long. Samara 5.5–7 mm long, very dark purple-brown to black. Fig. 50B–C. Widespread from Southern Cross to Norseman and Queen Victoria Spring, W.A.; in tall heath on sandplain. Regions: BENC, LEON, ESPR. Map 122. W.A.: 91 km from Southern Cross towards Coolgardie, E.M.Canning CBG 46357 (CBG, L, NSW); Queen Victoria Spring, 25 Sept. 1891, R.Helms NSW 50159 (AD, K, MEL, NSW); Comet Vale, J.H.Maiden NSW 18843 (A, K, L, NSW); Bronti, R.Melville 154 & C.A.Gardner (K, NSW); 18 km NE of Hyden, K.Newbey 1079 (PERTH). Previously regarded as a large-coned form of A. corniculata, which it resembles in habit and overlaps in range, but differing in the larger cones; larger black samara; verruculose and somewhat stouter articles; teeth 9–11, slightly overlapping and broader; and in being monoecious. A collection from the N margin of the Nullarbor Plain (D.L.Serventy & A.R.Main s.n., PERTH) lacks the long spines on the cone bracteole protuberance (but has a long-acuminate apex c. 1 mm long). This may represent a population which could then be given subspecific recognition.

133

Figure 50. Allocasuarina cones ×1.5. A, A. corniculata, branchlet ×7.5 (R.Melville 123B, NSW). B–C, A. spinosissima. B, cone; C, branchlet ×7.5 (B–C, E.Bennett 2146, NSW). D–E, A. acutivalvis subsp. acutivalvis. D, cone; E, branchlet ×15 (D–E, R.Coveny 8359, NSW). F, A. acutivalvis subsp. prinsepiana, cone (NE of Yuna, A.McCusker, NSW). G–H, A. scleroclada. G, cone; H, branchlet ×7.5 (G–H, NSW 94946, NSW). I–J, A. globosa. I, cone; J, branchlet ×7.5 (I–J, J.Beard 3838, PERTH). Drawn by D.Mackay.

134 CASUARINACEAE 3. Allocasuarina

Sect. 5. Ceropitys

Allocasuarina sect. Ceropitys L.Johnson, Fl. Australia 3: 191 (1989). Type: A. acutivalvis (F.Muell.) L.Johnson Dioecious or monoecious shrubs or trees; penultimate branchlets woody. Bark smooth or fissured. Branchlets usually with a strong waxy coating at maturity; elongate articles numerous, terete, smooth or verruculose; teeth not overlapping, mostly marcescent. Male spikes elongate or head-like; bracteoles persistent. Cone bracts occasionally prominent and thickened; bracteoles with protuberance entire or fused with bracteole body, or whole bracteole divided into 2–7 bodies. Samaras glabrous, mid-brown to black. A section of 9 species in south-western and southern Australia.

10. Allocasuarina acutivalvis (F.Muell.) L.Johnson, J. Adelaide Bot. Gard. 6: 74 (1982) Casuarina acutivalvis F.Muell., Fragm. 10: 61 (1876). T: Queen Victoria Spring, W.A., 30 Sept. 1875, D.Young; holo: MEL 540129, with cones. Dioecious shrub to small tree, 3–8 m high. Branchlets ascending, to 20 cm long; articles 10–25 mm long, 0.7–1.2 mm diam., verruculose, occasionally pubescent; phyllichnia flat, slightly verruculose; teeth 10–14, erect, 0.3–1.3 mm long, rarely to 1.8 mm long, marcescent. Male spikes 1–8 cm long, 3.5–6 whorls per cm; anther 1.2–2.1 mm long. Cones cylindrical to ovoid, finely white-pubescent to glabrous, often with longer ferruginous hairs when young, sessile or on peduncle to 10 mm long; cone body 15–35 mm long, 15–28 mm diam.; bracts prominent; bracteole and protuberance indistinguishable, very thick, pyramidal, acuminate with rather pungent mucro to 1.5 mm long or occasionally shortly bifid, or bracteole plus protuberance divided into 2 or 3 acuminate to acute bodies. Samara 6–12 mm long, black or very dark brown. 2n = 24, B.A.Barlow, Austral. J. Bot. 7: 232 (1959). Fig. 18. Widespread from Nerren Nerren Stn, N of the Murchison R., SE to Queen Victoria Spring and Zanthus, W.A.; in tall heath and open woodland and on rocky hillsides, in lateritic and sandy soils. Cone bracts more prominent than in many other species. There are 2 subspecies, chiefly distinguishable on whether the cone bracteoles are entire or deeply divided, and on the degree of spreading of teeth on new shoots. Some intergrading individuals are found with the bracteoles variously entire or divided even on the one cone. There is a zone of intergradation from N of the Murchison R. to near Merredin. Bracteole and protuberance fused, pyramidal, acuminate, occasionally shortly bifid; teeth on new shoots spreading to recurved 10a. subsp. acutivalvis Bracteole and protuberance divided into 2 or 3 bodies, each acuminate to acute; teeth on new shoots erect to spreading 10b. subsp. prinsepiana 10a. Allocasuarina acutivalvis (F.Muell.) L.Johnson subsp. acutivalvis Casuarina stowardii S.Moore, J. Linn. Soc., Bot. 45: 193 (1920). T: Nungarin, W.A., 1916, F.Stoward 743; holo: BM, with cones. Articles 10–25 mm long. Teeth 10–14, rarely 9, erect or occasionally spreading, 0.3–1.3 (rarely to 1.8) mm long. Cone body 15–35 mm long, 15–20 (rarely to 28) mm diam.; bracteole and protuberance indistinguishable, acuminate with rather pungent mucro to 1.5 mm long, occasionally the apex shortly bifid. Fig. 50D–E. From N of the lower Murchison R. to near Queen Victoria Spring and S almost to Ravensthorpe, W.A. Regions: MRCH, BENC, ESPR, LEON. Map 123. W.A.: 40 km S of Norseman, E.D.Briggs NSW 52364, 52365 (K, NSW); 16 km N of Cundeelee, M.I.H.Brooker 2588 (MEL, NSW, PERTH); 6.8 km E of Carrabin by road, R.G.Coveny 8359 & B.Haberley (NSW, PERTH); 1.5 km E of the Lake King–Ravensthorpe road towards Mt Madden, R.A.Saffrey 402 (PERTH).

135 3. Allocasuarina CASUARINACEAE

10b. Allocasuarina acutivalvis subsp. prinsepiana (C.Andrews) L.Johnson, J. Adelaide Bot. Gard. 6: 74 (1982) Casuarina prinsepiana C.Andrews, J. W. Austral. Nat. Hist. Soc. 1: 43 (1904). T: Mullewa, W.A., July 1903, C.R.P.Andrews; lecto: PERTH♀, ( with inflorescences; also ♂), fide L.A.S.Johnson, Fl. Australia 3: 202 (1989). Articles 12–20 mm long. Teeth 11–13, erect, 0.4–1 mm long. Cone body 25–32 mm long, 18–22 mm diam.; bracteole and protuberance divided into 2 or 3 bodies, acuminate to acute, with rather pungent mucro c. 1 mm long. Fig. 50F. Occurs from near Mullewa to N of Merredin and somewhat eastwards, flanked to the W and S by subsp. acutivalvis. Regions: MRCH, BENC. Map 124. W.A.: 16 km WSW of Paynes Find, A.C.Beauglehole 49169 (MEL, NSW, PERTH); 5.1 km from Bunna Warra turn-off towards Yalgoo, K.Hill 2581 & L.A.S.Johnson (NSW, PERTH); c. 6.9 km N of Morawa, R.J.Hnatiuk 760615 (PERTH); E of Pithara, R.D.Royce 1446 (PERTH); 19 km S of Paynes Find on Great Northern Hwy, R.A.Saffrey 834 (PERTH).

11. Allocasuarina scleroclada (L.Johnson) L.Johnson, J. Adelaide Bot. Gard. 6: 78 (1982) Casuarina scleroclada L.Johnson, Nuytsia 1: 261 (1972). T: c. 40 km SSE of Caiguna (c. 3.2 km from sea), W.A., 1 Sept. 1967, L.A.S.Johnson 2155; holo: NSW, with cones; iso: PERTH. Straggling dioecious shrub, 1–3 m high. Branchlets spreading or drooping, to 23 cm long; articles 20–52 mm long, 1.0–1.5 mm diam., smooth, occasionally pubescent in furrows; phyllichnia flat to slightly rounded; teeth 10–11, erect, 1.3–2.7 mm long, marcescent. Male spikes 6–16 mm long, dense; anther 0.5–0.8 mm long. Cones subcylindrical, sessile, glabrous except scattered hairs between bracteoles; cone body 18–25 mm long, 13–19 mm diam.; bracts inconspicuous; bracteole and protuberance fused, greatly thickened, broadly acute when young, becoming thickly obtuse, with or without a tiny deciduous mucro. Samara 5–8 mm long, black. 2n = c. 48, B.G.Briggs pers. comm. (B.G.Briggs 442). Fig. 50G–H. Occurs sporadically along the south-western coast of W.A., from Borden to the western part of the Great Australian Bight; in scrub and low woodland, in lateritic soils on rocky hillsides and on limestone pavements near the sea. Regions: ESPR, NULL. Map 125. W.A.: c. 495 km from Perth, Lake King–Ravensthorpe road, E.M.Bennett 3120 (NSW, PERTH); 46 km W of Phillips R., R.G.Coveny 7311 (NSW, PERTH); Salmon Gums, May 1924, C.A.Gardner (NSW, PERTH); Mt Ragged, A.S.George 2110 (PERTH); Mt Desmond, S of Ravensthorpe, L.A.S.Johnson 8727 (NSW). This species has a very straggling and open habit, often with a single main stem with arching drooping branches, especially in the female.

12. L.Johnson, Fl. Australia 3: 196 (1989) T: Bremer Ra., W.A., 25 Oct. 1964, J.S.Beard 3838; holo: PERTH, with cones. Dioecious shrub c. 1.5 m high. Branchlets ascending, to 12 cm long; articles 17–28 mm long, 0.9–1.2 mm diam., smooth, shortly pubescent in furrows; phyllichnia slightly rounded; teeth 10–12, erect to slightly spreading, 0.6–1 mm long, marcescent. Males unknown. Cones subglobose, tessellated, sessile; cone body 15–17 mm long, 13–15 mm diam.; bracts inconspicuous; bracteoles with obtuse apex, divided into 3 obtuse bodies surmounted by a tiny mucro. Samara 6.0–6.5 mm long, mid-brown. Fig. 50 I–J. Only known from Mt Day at the NW end of the Bremer Ra., W of Norseman, W.A.; in dense scrub with A. campestris, at the foot of a steep slope (J.S.Beard, Proc. Linn. Soc. New South Wales 93: 264, 1969). Region: ESPR. Map 126. This species differs from A. scleroclada in the shorter teeth, slender articles, paler samara and divided bracteoles; from A. campestris and A. tessellata in the longer articles, inconspicuous cone bracts, shorter cone, paler samara and the form of the divided cone bracteoles.

136 CASUARINACEAE 3. Allocasuarina

13. Allocasuarina tortiramula E.Bennett, Fl. Australia 3: 190 (1989) T: Nature Reserve 39422, W of Lake King, W.A., 22 July 1987, M.Graham 1127; holo: PERTH; iso: CANB, K, NSW. Dioecious shrub c. 1.7 m high. Branchlets spreading, twisted, to 10 cm long; articles 6–10 mm long, 0.8–1 mm diam., smooth, glabrous or shortly pubescent in furrows; phyllichnia rounded with median groove; teeth 7, spreading to erect, not overlapping, 0.5–0.8 mm long, marcescent. Male spikes 5–9 mm long, dense; anther 1.1–1.2 mm long. Cones cylindrical, sessile; cone body 10–15 mm long, c. 10 mm diam.; bracts inconspicuous; bracteoles obtuse, protuberance obtuse, shorter than bracteole body. Samara 5.5–6 mm long, mid-brown. Fig. 51. Known only from two populations in and near a Nature Reserve W of Lake King, W.A.; in dense tall heath in loam soil on granite. Region: ESPR. Map 127. Similar to A. globosa, but readily distinguished by the shorter articles with median furrow down the phyllichnia and the fewer, broader, thicker-textured teeth. It further differs from that species in the undivided bracteole protuberance on the cone.

14. (Diels) L.Johnson, J. Adelaide Bot. Gard. 6: 74 (1982) Casuarina campestris Diels in A.Engler, Bot. Jahrb. Syst. 35: 126, figs 10A–E (1905). T: Watheroo, W.A., 31 Dec. 1900, L.Diels 2038; lecto: B, with cones, fide L.A.S.Johnson, Nuytsia 1: 263 (1972); isolecto: NSW. Dioecious or monoecious shrub, dense, erect, 1–3 m high. Branchlets ascending, to 20 cm long; articles smooth, often pubescent in furrows, 6–13 mm long, 0.6–1.2 mm diam.; phyllichnia rounded or flat with a small but definite ridge; teeth 7–9, erect, 0.3–1.2 mm long, marcescent. Male spikes 0.4–2.8 cm long, 8.5–11 whorls per cm; anther 0.6–1 mm long. Cones long-cylindrical, shortly white-pubescent at least between bracts and bracteoles, sessile or sometimes on peduncle to 5 mm long; cone body 19–42 mm long, 10–17 mm diam.; bracts inconspicuous; bracteoles prominent, obtuse to broadly acute, protuberance fused to bracteole body except apex of protuberance which is marked by a curved or straight line or indentation close to body apex or rarely by a tiny mucro on surface of body. Samara 4.7–10 mm long, black. 2n = 24, B.A.Barlow, Austral. J. Bot. 7: 232 (1959). Fig. 52A–B. Widespread in the wheatbelt of W.A., extending from N of the Murchison R. almost to the S coast near Ravensthorpe and E of Esperance; on sandplain and laterite. Regions: MRCH, BENC, LEON, ESPR. Map 128. W.A.: 45 km SE of Coolgardie on Norseman road, A.C.Beauglehole 12283 (MEL, NSW, PERTH); 70 km S of Wannoo, D.F.Blaxell W75/115 (NSW, PERTH); 28.8 km ESE along Mt Gibson homestead road off Wubin–Paynes Find road, R.G.Coveny 7892 & B.R.Maslin (K, NSW, PERTH); Julimar Reserve, 80 km NE of Perth, D.Lewis NSW 150427 (K, MO, NSW, PERTH); 10 km S of Toompup, B.Rockel CS4 (FRI, NSW). Sometimes spreading clonally from roots or underground stems, at least in southern near-coastal populations. Some plants near the S coast produce large cones but these are usually sessile, not pedunculate as in A. eriochlamys, nor do they have the more prominent cone bracts of that species.

15. Allocasuarina eriochlamys (L.Johnson) L.Johnson, Fl. Australia 3: 195 (1989) Casuarina campestris subsp. eriochlamys L.Johnson, Nuytsia 1: 264 (1972); Allocasuarina campestris subsp. eriochlamys (L.Johnson) L.Johnson, J. Adelaide Bot. Gard. 6: 74 (1982). T: Comet Vale, W.A., Aug. 1917, J.T.Jutson 255; holo: NSW (♀). Dioecious or monoecious shrub, dense, erect, 1–3 m high. Branchlets ascending, to 23 cm long; articles smooth, often pubescent in furrows, 5–18 mm long, 0.5–1.1 mm diam.; phyllichnia flat or slightly rounded, not ridged; teeth 8–10, erect or rarely slightly spreading, 0.3–1 (rarely to 1.6) mm long, marcescent. Male spikes 1–3.5 cm long, 7–11 whorls per cm; anther 0.6–0.8 mm long. Cones long-cylindrical, shortly white-pubescent at least between bracts and bracteoles with some longer ferruginous hairs; peduncle 4–15 mm long; cone

137

Figure 51. Allocasuarina tortiramula. A, cone ×2; B, ♀inflorescence ×5; C, samara ×5; D, stamens ×12.5; E, ♂inflorescence ×2.5; F, branching habit (♂) ×1; G, habit ×0.75. (A–G, M.Graham 1127, PERTH). Drawn by C.Vasiliu.

138 CASUARINACEAE 3. Allocasuarina body 20–45 mm long, 13–21 mm diam.; bracts often inconspicuous; bracteoles with obtuse to acute apex, protuberance completely fused to bracteole body except apex of protuberance which is marked by a curved or straight line or indentation (rarely a tiny mucro) across bracteole body. Samara 5–10.5 mm long, black. Occurs from Comet Vale to Norseman, W.A., with an apparent outlier SSE of Zanthus; usually on or near rocky outcrops. Related to A. campestris, but differing morphologically in having pedunculate cones with more persistent indumentum, and phyllichnia without a small ridge; they differ also in habitat preferences. There are 2 subspecies. Cones with bract slightly thickened and bracteoles scarcely exceeding cone body; articles 0.5–0.9 mm diam.; cone body 13–18 mm diam. 15a. subsp. eriochlamys Cones with bract strongly thickened and bracteoles protruding prominently beyond cone body; articles 0.8–1.1 mm diam.; cone body 14–21 mm diam. 15b. subsp. grossa 15a. Allocasuarina eriochlamys (L.Johnson) L.Johnson subsp. eriochlamys Articles 5–14 mm long, 0.5–0.9 mm diam.; phyllichnia flat or slightly rounded, not ridged; teeth 8 or 9, 0.3–1 mm long. Male spikes 10–34 mm long, c. 7 whorls per cm. Cones on peduncle 4–13 mm long; cone body 23–39 mm long, 13–18 mm diam.; bracteoles not protruding beyond cone body, with obtuse to truncate apex, indentation marking apex of protuberance close to body apex. Fig. 52C–D. Occurs in the Comet Vale–Kalgoorlie area, W.A.; in mallee woodland on stony and gravelly slopes, probably usually near subdued rocky outcrops. Region: LEON. Map 129. W.A.: 3 km W of Musson Soak, E of Goongarrie, J.S.Beard 6257 (NSW); 11 km E of Riverina homestead on road to Menzies, M.G.Corrick 9125 (MEL, NSW); Seven Mile Hill, W of Kalgoorlie, R.Melville 3989, 3990 (K, NSW); Comet Vale, B.Rockel CCE3, CCE6 (FRI, NSW). 15b. Allocasuarina eriochlamys subsp. grossa (L.Johnson) L.Johnson, Fl. Australia 3: 195 (1989) Casuarina campestris subsp. grossa L.Johnson, Nuytsia 1: 264 (1972); Allocasuarina campestris subsp. grossa (L.Johnson) L.Johnson, J. Adelaide Bot. Gard. 6: 74 (1982). T: 9 km N of Norseman, W.A., 18 Dec. 1960, L.A.S.Johnson W177; holo: NSW, with cones; iso: PERTH. Articles 9–18 mm long, 0.8–1.1 mm diam.; phyllichnia flat or slightly rounded, not ridged; teeth 8–11, 0.5–0.9 (rarely to 1.6) mm long. Male spikes 16–35 mm long, 10–11 whorls per cm. Cones on peduncle 4–15 mm long; cone body 20–38 mm long, 14–21 mm diam.; bracts thickened laterally, so base appears 2-lobed; bracteoles protruding prominently beyond cone body, with acute to broadly acute apex, indentation marking apex of protuberance below body apex. 2n = 24, B.G.Briggs pers. comm. (B.G.Briggs 290). Fig. 52E–F. Occurs in the Norseman district, W.A., with one record SSE of Zanthus; around granite outcrops. Regions: LEON. Map 130. W.A.: 5 km NE of Norseman, A.C.Beauglehole 49399 (MEL, NSW, PERTH); between Zanthus and Cocklebiddy, Oct. 1964, P.Garstone (PERTH); Lake Lefroy, R.J.Larsen NSW 61828 (NSW); Beacon Hill, Norseman, M.E.Phillips CBG 44193 (CBG, NSW); SW arm of Lake Cowan, B.Rockel CCG5 (FRI, NSW). Cone bracts are more prominent and thicker than in A. campestris; bracteole has indentation below apex (cf. subsp. eriochlamys and A. campestris) and is slightly less prominent than in A. campestris.

16. Allocasuarina tessellata (C.Gardner) L.Johnson, J. Adelaide Bot. Gard. 6: 78 (1982) Casuarina tessellata C.Gardner, J. Proc. Roy. Soc. W. Australia 22: 119 (1936); C. campestris subsp. tessellata (C.Gardner) L.Johnson, Nuytsia 1: 263 (1972). T: summit of Mt Singleton, W.A., 9 July 1931, C.A.Gardner 2217; holo: PERTH, with cones; iso: B, K.

139

Figure 52. Allocasuarina cones ×1.5. A–B, A. campestris. A, cone; B, branchlet ×15 (A–B, R.Coveny 8324, NSW). C–D, A. eriochlamys subsp. eriochlamys. C, cone; D, branchlet × 15 (C–D, NSW 61832, NSW). E–F, A. eriochlamys subsp. grossa. E, cone (B.Briggs 289, NSW); F, branchlet ×15 (NSW 58480, NSW). G–H, A. tessellata. G, cone; H, branchlet ×15 (G–H, P.Wilson 8616, NSW). I–J, A. dielsiana. I, cone; J, branchlet ×7.5 (I–J, E.Bennett 1461, NSW). K–L, A. helmsii. K, cone; L, branchlet ×15 (K–L, D.Symon 2627, NSW). Drawn by D.Mackay.

140 CASUARINACEAE 3. Allocasuarina

Similar to A. campestris. Dioecious shrub or tree, 3–5 m high. Branchlets ascending; articles 7–14 mm long, 0.7–1 mm diam., glabrous; phyllichnia rounded; teeth 8 or 9. Male spikes 2–4 cm long, 7 or 8 whorls per cm; anther c. 0.8 mm long, much broader than long. Cones occasionally with sterile apex to 5 mm long; peduncle 7–13 mm long; cone body 26–55 mm long, 14–18 mm diam.; bracts thickened; bracteoles obtuse, body and protuberance not distinguishable, the whole divided into 3–5 tightly packed, thickened bodies. Samara 5–7.5 mm long. Fig. 52G–H. Occurs on Mt Singleton, W.A.; on rocky hillsides. Region: BENC. Map 131. W.A.: Mt Singleton, R.G.Coveny 7915 & B.R.Maslin (K, NSW, PERTH); Ninghan Stn, c. 40 km SW of Paynes Find, B.Rockel CCT10 (FRI, NSW); 13 km from main road, through Ninghan to Mt Singleton (N edge), E.M.Scrymgeour 2111 (NSW, PERTH). More tree-like than A. campestris.

17. (C.Gardner) L.Johnson, J. Adelaide Bot. Gard. 6: 75 (1982) Casuarina dielsiana C.Gardner, J. Proc. Roy. Soc. W. Australia 22: 119 (1936). T: summit of Mt Singleton, W.A., 9 July 1931, C.A.Gardner 2218; holo: PERTH, with cones; iso: B. Usually dioecious tree, 4–9 m high. Branchlets ascending, to 20 cm long; articles 6–10 mm long, 0.8–1 mm diam., smooth, glabrous; phyllichnia angular; teeth 6–8, slightly spreading, 0.5–0.7 mm long, the tips breaking off readily, marcescent. Male spikes head-like or slightly elongate, 5–6 mm long; anther 0.8–1.1 mm long. Cones cylindrical, very finely pubescent becoming glabrous, sessile or on peduncle to 7 mm long, spreading or recurved relative to branch; cone body 14–30 mm long, 12–17 mm diam.; bracts thickened; bracteoles and protuberances indistinguishable, acute to broadly acute. Samara 8–10 mm long, mid-brown. 2n = 28, B.A.Barlow, Austral. J. Bot. 7: 232 (1959). Northern Sheoak. Figs 45J, 52 I–J. Occurs from the Murchison R. area to Paynes Find and the Die Hardy Ra.; in lateritic hilly country. Regions: BENC, LEON. Map 132. W.A.: between Mullewa and Pindar, 10 Sept. 1928, C.A.Gardner (K, PERTH); Die Hardy Ra., A.S.George 8682 (K, PERTH); Rothsay townsite, 70 km E of Perenjori, B.Rockel CDL10, CDL12 (FRI, NSW); 104 km NE of Wubin on Paynes Find road, K.L.Wilson 2617–2619 (K, L, NSW, PERTH, RSA); 10 km N of Murchison R. on North West Coastal Hwy, K.L.Wilson 2655, 2656 (K, NSW, PERTH). The globular male spikes are distinctive in the genus.

18. (Ewart & M.Gordon) L.Johnson, J. Adelaide Bot. Gard. 6: 75 (1982) Casuarina helmsii Ewart & M.Gordon, Proc. Roy. Soc. Victoria n. ser. 32: 192, t. 12 (1920). T: Gnarlbine, W.A., 12 Nov. 1891, R.Helms; lecto: MEL, with cones, fide L.A.S.Johnson, J. Adelaide Bot. Gard. 6: 75 (1982); isolecto: BM, K, LE, MEL. Casuarina leptotrema S.Moore, J. Linn. Soc., Bot. 45: 193 (1920). T: Nungarin, W.A., 1916, F.Stoward 742; holo: BM, with cones; iso: MEL. [C. humilis auct. non Otto & Dietr.: J.M.Black, Fl. S. Australia 2nd edn, 2: 259 (1948)] Dioecious shrub, 1–5 m high. Branchlets erect, to 16 cm long; articles 3–7 mm long, 0.7–0.9 mm diam., smooth, glabrous; phyllichnia flat to slightly rounded, often with faint median groove more obvious towards base of article; teeth 5 or 6, erect, 0.3–0.5 mm long, not marcescent. Male spikes 0.5–2.5 cm long, 9–12 whorls per cm; anther 0.5–0.7 mm long. Cone usually long-cylindrical, of smoothish outline and tessellated surface, shortly white-pubescent, sometimes ferruginous-pubescent when young, sessile or on peduncle to 4 mm long; cone body 15–33 mm long, 8–13 mm diam.; bracts inconspicuous; bracteoles thick, body and protuberance not distinguishable, the whole divided into 4–7 obtuse or truncate bodies. Samara 4–5 mm long, mid-brown. 2n = 24, B.A.Barlow, Austral. J. Bot. 7: 232 (1959). Fig. 52K–L.

141 3. Allocasuarina CASUARINACEAE

Occurs from Nungarin, W.A., to north-western S.A., with outliers on the northern Eyre Peninsula, S.A., and Dirk Hartog Is., W.A.; in mallee woodland and tall heath, in a wide range of soils. Regions: MRCH, BENC, LEON, NULL, EYRE. Map 133. W.A.: Bullabulling, C.A.Gardner 9273 (PERTH); Dirk Hartog Is., A.S.George 11588 (NSW, PERTH); near Point Dover, Great Australian Bight, P.G.Wilson 5942 (K). S.A.: Great Victoria Desert, D.E.Symon 12369 (AD, CANB, NSW); Buckleboo railway siding, P.G.Wilson 607, 608 (AD, CANB, K, L, NE, NSW, RSA, UC, W).

Sect. 6. Allocasuarina

Allocasuarina L.Johnson sect. Allocasuarina Type: A. torulosa (Aiton) L.Johnson Dioecious or monoecious trees; penultimate branchlets woody. Bark thick, corky. Branchlets not waxy; elongate articles numerous, terete or quadrangular, smooth; teeth not overlapping or marcescent. Male spikes elongate; bracteoles persistent. Cone bracts inconspicuous; bracteoles with protuberance 8–20-parted. Samaras glabrous, mid-brown to black. A section of 2 species; 1 in eastern Australia, the other in south-western W.A.

19. (Aiton) L.Johnson, J. Adelaide Bot. Gard. 6: 79 (1982) Casuarina torulosa Aiton, Hort. Kew. 3: 320 (1789); C. lugubris Salisb., Prodr. 2 (1796), nom. illeg. based on C. torulosa Aiton. T: Bay of Inlets, [Qld], 1770, J.Banks & D.Solander; neo: BM, with cones, fide L.A.S.Johnson, J. Adelaide Bot. Gard. 6: 79 (1982); isoneo: MEL, NSW, P, W. Casuarina tenuissima Sieber ex Sprengel, Syst. Veg. 3: 804 (1826). T: New South Wales, F.W.Sieber 327; holo: B (♂) n.v.; iso: BM, BR, C, FI, K, L, MEL, P, PR, S, W. Illustrations: J.H.Maiden, Forest Fl. New South Wales 2: t. 63 (1905); L.Fuller, Wollongong’s Native Trees 2nd edn, 73 (1982); D.J.Boland et al., Forest Trees Australia 4th edn, 107 (1984). Usually dioecious tree 5–20 m high. Branchlets drooping, to 14 cm long; articles terete, quadrangular on young growth, 5–6 mm long, 0.4–0.5 mm diam., minutely pubescent in furrows; phyllichnia slightly rounded; teeth 4 or 5, erect, 0.3–0.8 mm long. Male spikes 0.5–3 cm long, 7–12 whorls per cm; anther 0.5–0.6 mm long. Cones shortly cylindrical or barrel-shaped, warty, occasionally densely pubescent, pendent or spreading; peduncle 8–30 mm long; cone body 15–33 mm long, 12–25 mm diam.; bracts inconspicuous; bracteoles acute, protuberance divided into 8–12 small tubercles slightly shorter than or as long as bracteole body. Samara 7–10 mm long, mid to dark brown. 2n = 24, B.A.Barlow, Austral. J. Bot. 7: 233 (1959); B.G.Briggs pers. comm. (K.Thurtell & R.G.Coveny 3838). Forest Oak, Rose Sheoak. Fig. 53A–B. Occurs from the McIlwraith Ra., Qld, S to Macquarie Pass and Jenolan Caves, N.S.W.; grows on coastal hills and ranges as understorey in open forest to tall open-forest, in a wide range of soils. Regions: CYRK, BURD, DWSN, MCPH, NEPN. Map 134. Qld: near Kenilworth, S.T.Blake 13944 (BRI, NSW); Davies Ck Forestry Road c. 16 km NNE of Mareeba, R.Hoogland 8504 (CANB, NSW). N.S.W.: Toonumbar, E.F.Constable NSW 10758 (NSW); Macquarie Pass, L.A.S.Johnson NSW 61928 (NSW); Mt Royal, 40 km E of Aberdeen, R.Story 7182 (CANB, NSW). In north Qld material, phyllichnia are often more angular. Generally grows in higher-nutrient soils and in moister forest than A. littoralis.

20. Allocasuarina decussata (Benth.) L.Johnson, J. Adelaide Bot. Gard. 6: 75 (1982) Casuarina decussata Benth., Fl. Austral. 6: 200 (1873). T: towards Cape Riche, W.A., J.Drummond 5: 434; holo: K (♂, with cones - monoecious); iso: BM, FI, LE, MEL 545419. Monoecious tree 8–15 m high, or more rarely shrub 1–3 m. Branchlets ascending, to 14 cm long; articles quadrangular, 3–7 mm long, 0.6–0.8 mm diam., occasionally finely pubescent when young; phyllichnia sharply angular; teeth 4, erect, 0.6–1 mm long. Male spikes 1–3.5

142 CASUARINACEAE 3. Allocasuarina cm long, 8–10 whorls per cm; anther 0.9–1 mm long. Cones shortly cylindrical, diam. greater than or equal to length, warty, sessile or on peduncle to 2 mm long; cone body 10–20 mm long, 15–24 mm diam., rarely to 40 mm long and 30 mm diam.; bracts inconspicuous; bracteoles obtuse, 3-lobed with a tiny mucro, protuberance longer than bracteole body and divided into 12–20 small bodies, pungent or with rounded apex. Samara 7–9 mm long, blackish. 2n = 20, B.A.Barlow, Austral. J. Bot. 7: 232 (1959); B.G.Briggs pers. comm. (NSW 95218). Karri Oak. Fig. 53C–D. Occurs from Blackwood R. south to Denmark, W.A.; grows in loam in Karri (Eucalyptus diversicolor) forest; also on Bluff Knoll, Stirling Ra., where it occurs as a shrub or depauperate tree in shrubland. Regions: LUWN, ESPR. Map 135. W.A.: c. 6 km along Rainbow Trail from Pemberton, A.C.Beauglehole 12628 (MEL, NSW, PERTH); Mt Chudalup, A.S.George 3215 (PERTH); Deep R., Dec. 1912, S.W.Jackson (BO, FI, K, L, NSW, US); Bluff Knoll, A.Strid 21566 (NSW, PERTH); Denmark, C.T.White 5367 (B, BRI, K, NY, S).

Sect. 7. Amorphopitys

Allocasuarina sect. Amorphopitys L.Johnson, Fl. Australia 3: 191 (1989). Type: A. fraseriana (Miq.) L.Johnson Dioecious trees; penultimate branchlets woody. Bark scaly or corky. Branchlets occasionally waxy; elongate articles numerous, terete, verruculose or smooth; teeth not overlapping or marcescent. Male spikes elongate; bracteoles persistent. Cone bracts somewhat thickened but inconspicuous; bracteoles with protuberance 4–8-parted. Samaras glabrous, very dark brown to black. A section of 1 species in south-western Australia.

21. (Miq.) L.Johnson, J. Adelaide Bot. Gard. 6: 75 (1982) Casuarina fraseriana Miq., Rev. Crit. Casuarineae 59, t. 6D (1848); C. stricta var. fraseriana (Miq.) Miq., Flora 48: 20 (1865). T: near Perth, W.A., 9 Apr. 1840, L.Preiss 2000; lecto: U, with cone, fide L.A.S.Johnson, J. Adelaide Bot. Gard. 6: 75 (1982); isolecto: B, LE, MEL, W. Casuarina torulosa f. gracilior? Miq. in J.G.C.Lehmann, Pl. Preiss. 1: 640 (1845). T: Mt Wuljenup [Mt Willyung], W.A., 14 Oct. 1840, L.Preiss 2007; holo: U (vegetative). Illustration: D.J.Boland et al., Forest Trees Australia 4th edn, 101 (1984). Tree 5–15 m high. Branchlets ascending, to 30 cm long; articles 7–15 mm long, 0.8–1.3 mm diam., occasionally strongly waxy; phyllichnia rounded, verruculose or smooth; teeth 6–8, somewhat spreading, 0.7–1.2 mm long. Male spikes 3–8 cm long, 5 or 6 whorls per cm; anther 0.7–1.2 mm long. Cones coarse, usually shortly cylindrical, warty, pubescent at least when young, sessile or on peduncle to 32 mm long; cone body 15–40 mm long, 15–22 mm diam.; bracteoles broadly acute to obtuse, protuberance slightly shorter than bracteole body and divided into 4–8 bodies, pungent or obtuse. Samara 9–10 mm long. 2n = 26, B.A.Barlow, Austral. J. Bot. 7: 232 (1959). Western Sheoak. Fig. 53E–F. Occurs from Perth to Albany, W.A., with an isolated population between Moora and Jurien Bay, W.A.; on the Darling Ra. grows in lateritic soil in Jarrah () forest; near the coast grows in sand in woodland and open forest. Regions: BENC, LUWN. Map 136. W.A.: 2.2 km S of Napier Ck, E of Albany, E.M.Bennett 1049 (K, PERTH); Red Hill, Upper Swan, R.J.Cranfield 418 (NSW, PERTH); between Boggy Lake and Deep R., J.W.Green 1117 (PERTH); Bayswater, 22 June 1907, A.Morrison (K, P, US).

143 3. Allocasuarina CASUARINACEAE

Sect. 8. Oöpitys

Allocasuarina sect. Oöpitys L.Johnson, Fl. Australia 3: 192 (1989). Type: A. verticillata (Lam.) L.Johnson Dioecious trees; penultimate branchlets woody. Bark fissured. Branchlets not waxy; elongate articles numerous, terete, verruculose; teeth not overlapping, marcescent. Male spikes elongate; bracteoles deciduous usually as a pair or united with tepals. Cone bracts inconspicuous; bracteoles with protuberance very small or fused with bracteole body. Samaras glabrous, dark brown to black. A section of 2 species, 1 in south-eastern Australia, the other in south-western W.A. The male bracteoles tend to fall as a pair or together with the 1 or 2 tepals when the fimbriate apices become entangled.

22. Allocasuarina verticillata (Lam.) L.Johnson, J. Adelaide Bot. Gard. 6: 79 (1982) Casuarina verticillata Lam., Encycl. 2: 501 (1788). T: cult. Jardin du Roi, Paris, France [probably coll. in Tasmania], herb. Lamarck; lecto: P-LA, vegetative, fide L.A.S.Johnson, J. Adelaide Bot. Gard. 6: 79 (1982). Casuarina stricta Aiton, Hort. Kew. 3: 320 (1789); C. excelsa Salisb., Prodr. 2 (1796), nom. illeg. based on C. stricta Aiton. T: Hort. Kew, 1778, W.Aiton; holo: BM ♂).( Casuarina quadrivalvis Labill., Nov. Holl. Pl. 2: 67, t. 218 (1806). T: Australia, probably Tas., J.J.H. de Labillardière; lecto: FI ♀(, with inflorescences and cones), fide L.A.S.Johnson, Fl. Australia 3: 202 (1989); probable isolecto: BM, P. Casuarina quadrivalvis Labill. var. macrocarpa Miq., Rev. Crit. Casuarineae 73, t. 10B (1848). T: Lachlan R., N.S.W., 1817, A.Cunningham 361; holo: K (♀, with inflorescences and cones). Casuarina quadrivalvis Labill. var. spectabilis Miq., Rev. Crit. Casuarineae 73, t. 10C (1848); C. gunnii J.D.Hook. ex Miq., Ned. Kruidk. Arch. 4: 100 (1859). T: near George Town, Tas., 17 Jan. 1843, R.Gunn 1247 [cited as Hooker 1247]; holo: K ♂);( iso: BM, U. Illustrations: J.H.Maiden, Forest Fl. New South Wales 2, t. 65 (1905); L.Costermans, Native Trees & Shrubs S.E. Australia 146–147 (1981); L.Fuller, Wollongong’s Native Trees 2nd edn, 75 (1982); G.M.Cunningham et al., Pl. W. New South Wales 209 (1982). Tree 4–10 m high. Branchlets drooping, to 40 cm long; articles 15–40 mm long, 0.7–1.5 mm diam., mostly densely pubescent in furrows; phyllichnia slightly rounded with a verruculose ridge; teeth 9–13, spreading, 0.7–1.2 mm long. Male spikes 3–12 cm long, 2.5–4 whorls per cm; anther 1.2–2.5 mm long. Cones cylindrical to barrel-shaped, sessile or on peduncle to 10 mm long; cone body 20–50 mm long, 17–30 mm diam.; bracts with long slender mucro; bracteoles broadly acute to acute, often pungent, protuberance fused with body or inconspicuous and only about as long as bracts. Samara 7–12 mm long, very dark brown. 2n = 26, B.A.Barlow, Austral. J. Bot. 7: 232 (1959). Drooping Sheoak. Figs 17, 45K, 53G–H. Occurs from near Cobar, N.S.W., southwards and extending to the coast on shale patches near Sydney and on the S coast of N.S.W., widespread through Vic. except in the NW, W to the Flinders and Gawler Ranges, western Eyre Peninsula and Kangaroo Is., S.A., and in Tas. from Launceston down the E coast to Kingston and the Tasman Peninsula. Grows in usually grassy woodland, forming pure stands or amongst eucalypts; also on rocky sea-coasts and dry rocky hills and ridges inland. Regions: DARL, NEPN, RIVR, HOWE, OTWY, TRNS, EYRE, TASM. Map 137. S.A.: Miltalie, c. 16 km NW of Cowell, R.Pearce ADW 29661 (AD, CANB, NSW). N.S.W.: Newport, R.H.Cambage NSW 61957, 61959 (NSW); Merrimerriwa Ra., Yathong Nature Reserve, K.Hill 117 & L.A.S.Johnson (K, NSW). Vic.: Lawloit Ra., c. 19 km E of Kaniva, R.Melville 1189, 1192 (K, NSW). Tas.: Blowhole, Eaglehawk Neck, M.E.Phillips CBG 13872 (CBG, NSW).

144

Figure 53. Allocasuarina cones ×1.5. A–B, A. torulosa. A, cone (NSW 61908, NSW); B, branchlet ×15 (NSW 61923, NSW). C–D, A. decussata. C, cone (A.Beauglehole 12628, NSW); D, branchlet ×15 (NSW 95218, NSW). E–F, A. fraseriana. E, cone (NSW 62076, NSW); F, branchlet ×7.5 (E.Bennett 1114, NSW). G–H, A. verticillata. G, cone (R.Coveny 7301, NSW); H, branchlet ×7.5 (G.M.Cunningham 3799, NSW). I–J, A. huegeliana. I, cone; J, branchlet ×15 (I–J, E.Bennett 588, NSW). K–L, A. trichodon. K, cone; L, branchlet ×7.5 (K–L, NSW 106388, NSW). Drawn by D.Mackay.

145 3. Allocasuarina CASUARINACEAE

23. (Miq.) L.Johnson, J. Adelaide Bot. Gard. 6: 76 (1982) Casuarina huegeliana Miq. in J.G.C.Lehmann, Pl. Preiss. 1: 640 (1845). T: Mt Brown, near York, W.A., 5 Sept. 1839, L.Preiss 2006 p.p.; lecto: U♂ ),( fide L.A.S.Johnson, J. Adelaide Bot. Gard. 6: 76 (1982); isolecto: BM, BR, C, FI, K, L, LE, MEL, P, S; syn: L♀)E. ( Casuarina dorrienii Domin, J. Linn. Soc., Bot. 41: 246, figs 25–27 (1912). T: Warrungup Hill, Stirling Ra., W.A., 1910, A.Dorrien-Smith; holo: K (♂, ♀). Illustrations: L.Diels in A.Engler, Bot. Jahrb. Syst. 35: fig. 9C–E (1905). Similar to A. verticillata. Tree 4–10 m high. Branchlets drooping to ascending; articles not or scarcely greater in diam. at apex when dried, 8–13 mm long, 0.8–0.9 mm diam.; teeth 8–10, 0.5–0.7 mm long. Male spikes 2–10 cm long, 4.5–7 whorls per cm; anther 0.8–1.2 mm long. Cones on peduncle 3–5 mm long; cone body 14–35 mm long, 10–14 mm diam.; bracteoles obtuse to acute. Samara 5–7 mm long, dark brown to black. 2n = 26, B.A.Barlow, Austral. J. Bot. 7: 232 (1959). Rock Sheoak. Fig. 53 I–J. Occurs from Murchison R. and Mingenew S to the S coast and E to Newman Rock, W of Balladonia, W.A.; associated with granite.. Regions: BENC, ESPR, NULL. Map 138. W.A.: Merredin, M.Koch 2978 (NSW); Mingenew, J.H.Maiden NSW 62094 (NSW); Beaufort Inlet, K.Newbey 1246 (PERTH); Newman Rock, 51 km W of Balladonia, T. & J.Whaite 4036 (K, NSW, PERTH); c. 8 km NW of Bullaring on Pingelly road, K.L.Wilson 2607 (K, NSW, PERTH). Close to A. verticillata, but A. huegeliana is more slender and has cones with more rounded bracteoles.

Sect. 9. Trichopitys

Allocasuarina sect. Trichopitys L.Johnson, Fl. Australia 3: 193 (1989). Type: A. trichodon (Miq.) L.Johnson Dioecious or rarely monoecious shrub; penultimate branchlets woody. Bark finely striate. Branchlets not waxy; elongate articles numerous, terete, smooth; teeth not overlapping, long and marcescent. Male spikes elongate; bracteoles persistent. Cone bracts small but often prominent because of dense white pubescence and long slender mucro to 3 mm long; bracteoles with protuberance entire. Samaras glabrous, dark brown to black. A section of 1 species in south-western Australia.

24. Allocasuarina trichodon (Miq.) L.Johnson, J. Adelaide Bot. Gard. 6: 79 (1982) Casuarina trichodon Miq. in J.G.C.Lehmann, Pl. Preiss. 1: 641 (1845). T: Konkoberup Hills [near Cape Riche], W.A., 19 Nov. 1840, L.Preiss 2001; holo: U (♂); iso: L, LE, MEL, NSW, P. Casuarina bicuspidata Benth., Fl. Austral. 6: 202 (1873). T: probably King George Sound or Lucky Bay, [W.A.], (cited as Flinders Is., S.A., but the species occurs only in W.A.), R.Brown; lecto: K♀, ( with inflorescences and cones), fide L.A.S.Johnson, Fl. Australia 3: 200 (1989); isolecto: B, BM. Erect to spreading shrub, 0.5–3 m high. Branchlets ascending, to 30 cm long; articles 9–15 mm long, 0.9–1 mm diam., glabrous; phyllichnia markedly angular with median ridge; teeth 8–10, spreading, 1.8–3.5 mm long, often only c. 1 mm remaining on older articles. Male spikes 3–8 cm long, 8 whorls per cm; anther 1–1.2 mm long. Cones cylindrical to barrel-shaped, pubescent, sessile; cone body 15–50 mm long, 14–20 mm diam.; bracteoles slender, acute or acuminate with small mucro, usually pungent, protuberance nearly as long as bracteole body and very similar in size and shape. Samara 8–10 mm long. 2n = 20, B.A.Barlow, Austral. J. Bot. 7: 233 (1959). Fig. 53K–L. Occurs from Albany E to beyond Esperance and in the Stirling Ra., W.A.; grows in tall heathland, often in skeletal soils. Regions: LUWN, ESPR. Map 139.

146 CASUARINACEAE 3. Allocasuarina

W.A.: Stirling Ra. Natl Park, E.M.Canning CBG 32116 (CBG, NSW); East Mt Barren, 27 Jan. 1935, C.A.Gardner (PERTH); Mt Gardner, E of Albany, A.S.George 6320 (NSW, PERTH); Mt Le Grand, A.S.George 2222 (PERTH). The long spreading leaf teeth are distinctive amongst Western Australian species.

Sect. 10. Inopitys

Allocasuarina sect. Inopitys L.Johnson, Fl. Australia 3: 192 (1989). Type: A. inophloia (F.Muell. & Bailey) L.Johnson Dioecious trees; penultimate branchlets woody. Bark very ribbony-fibrous. Branchlets not waxy; elongate articles numerous, terete, verruculose; teeth not overlapping or marcescent. Male spikes elongate; bracteoles deciduous individually. Cone bracts usually split down middle and not visible; bracteoles with protuberance 2- or 3-parted. Samaras glabrous. A section of 1 species in eastern Australia.

25. (F.Muell. & Bailey) L.Johnson, J. Adelaide Bot. Gard. 6: 76 (1982) Casuarina inophloia F.Muell. & Bailey, Chem. & Druggist 4: Australas. Suppl. 92 (1882). T: Queensland, collector unknown; lecto: MEL, sterile, fide L.A.S.Johnson, J. Adelaide Bot. Gard. 6: 76 (1982). Illustration: J.H.Maiden, Forest Fl. New South Wales 3: t. 80 (1906). Open tree 3–10 m high. Branchlets drooping or ascending, to 21 cm long; articles 4–7 mm long, 0.5–0.6 mm diam., glabrous; phyllichnia slightly rounded with verruculose median ridge or line; teeth 7–9, erect, 0.3–0.5 mm long. Male spikes 2–4 cm long, 7–14 whorls per cm; anther 0.5–0.8 mm long. Cones cylindrical, densely pubescent; peduncle 3–8 mm long; cone body 10–20 mm long, 9–12 mm diam.; bracteoles acute to obtuse, protuberance usually divided near apex into 2 or 3 bodies, acute to obtuse, rarely only 1 body. Samara 5–6 mm long. 2n = 24, B.A.Barlow, Austral. J. Bot. 7: 232 (1959). Stringybark Oak. Figs 22, 54A–B. Occurs from the Herberton area, Qld, S to Warialda, N.S.W.; grows in woodland on sandstone or laterite ridges. Regions: CYRK, BURD, DWSN, MCPH. Map 140. Qld: Proston, S.T.Blake 14251 (BRI, NSW); E of Baal Gammon Mine, c. 7 km W of Herberton, B.J.Conn & J. de Campo 1295 (BRI, MEL, NSW); Alton, Moonie Hwy, T.J.McDonald 411 (BRI, MEL, NSW). N.S.W.: Warialda, J.L.Boorman NSW 19485 (BO, BR, BRI, K, NSW, U, W); Mulvaine Hill, 5 km S of Torrington, E.F.Constable 2028 (NSW). The ribbony-fibrous bark of this species is unique in the family.

Sect. 11. Cylindropitys

Allocasuarina sect. Cylindropitys L.Johnson, Fl. Australia 3: 191 (1989). Type: A. littoralis (Salisb.) L.Johnson Dioecious or monoecious shrubs or trees; penultimate branchlets woody. Bark smooth or fissured. Branchlets occasionally with a waxy bloom; elongate articles numerous, terete, smooth or occasionally verruculose; teeth may be overlapping and marcescent. Male spikes elongate, occasionally only shortly so; bracteoles persistent or rarely deciduous individually. Cone bracts inconspicuous; bracteoles with protuberance entire or rarely 2- or 3-parted. Samaras glabrous, red-brown to black. A section of 29 species, mainly in eastern and south-eastern Australia, 1 in south-western W.A.

147 3. Allocasuarina CASUARINACEAE

26. Allocasuarina littoralis (Salisb.) L.Johnson, J. Adelaide Bot. Gard. 6: 76 (1982) Casuarina littoralis Salisb., Prodr. 2 (1796). T: Botany Bay, [N.S.W.], 1770, J.Banks & D.Solander; holo: BM, with cones; iso: NSW, P. Casuarina suberosa Otto & Dietr., Allg. Gartenzeitung 9: 155 (1841). T: cult. Berlin bot. gard., collector unknown; holo: B (♂, ♀ - monoecious); probable iso: L, MEL. Casuarina leptoclada Miq., Rev. Crit. Casuarineae 41, t. 4C (1848). T: Sidney [Sydney], N.S.W., Clowes; lecto: K (♂), fide L.A.S.Johnson, Fl. Australia 3: 201 (1989); iso: U. Casuarina moesta F.Muell. ex Miq., Ned. Kruidk. Arch. 4: 98 (1856). T: near Melbourne, Vic., F.Mueller; lecto: MEL♀, ( with inflorescences), fide L.A.S.Johnson, Fl. Australia 3: 201 (1989); isolecto: B, LE, P. Illustrations: J.H.Maiden, Forest Fl. New South Wales 2: 72 (1905); L.Fuller, Wollongong’s Native Trees 2nd edn, 71 (1982). Usually dioecious tree 5–15 m high or rarely shrub. Bark fissured. Branchlets ascending or drooping, to 20 (rarely to 35) cm long; articles 4–10 mm long, 0.4–1 mm diam., smooth, pubescent in furrows; phyllichnia angular or rounded with median ridge; teeth 6–8, rarely 5 or 9, erect or rarely spreading, not overlapping, 0.3–0.9 mm long, usually not marcescent. Male spikes 0.5–5 cm long, 6–12.5 whorls per cm; bracteoles persistent; anther 0.4–0.8 mm long. Cones cylindrical, rarely broader than long, pubescent at least when young; peduncle 4–23 mm long; cone body 10–30 (rarely to 45) mm long, 8–21 mm diam.; bracteoles thin, broadly acute to obtuse, thick pyramidal protuberance shorter than bracteole body, occasionally with 2 lateral bodies. Samara 4–10 mm long, dark brown to black. 2n = 22, 44, B.A.Barlow, Austral. J. Bot. 6: 38–46 (1958). Black Sheoak. Figs 20, 54C–D. Occurs from Cape York, Qld, S along coast and adjoining tablelands to the Otway Ra., Vic., and down the E coast of Tas. to S of Hobart; grows in woodland and occasionally tall heath, in sandy and other poor soils (cf. A. torulosa). Regions: CYRK, BURD, DWSN, MCPH, NEPN, HOWE, OTWY, TASM. Map 141. Qld: Moreton Is., L.Durrington 374 (BRI, NSW); Beatie Ck, 72 km N of Wenlock R., P.Hind 536 (BRI, K, MO, NSW). N.S.W.: Wentworth Falls, E.F.Constable NSW 62282 (CHR, NSW). Vic.: Agahook Forest Park, Anglesea–Bambra, C.W.Huggins 23 (MEL, NSW). Tas.: 27 km from Ross on Cressy road, N.T.Burbidge 3548, 3529 (CANB, K, NSW). Very variable in size of articles and cones. Occurs as a coarse shrub in maritime exposed situations. Variation is not simply linked to chromosome number or distribution, and there appears to be complete intergradation between forms.

27. Allocasuarina media L.Johnson, Fl. Australia 3: 197 (1989) T: c. 3 km along Five Mile Rd from Tidal River Rd, Wilsons Promontory Natl Park, Vic., 4 Mar. 1986, K.L.Wilson 6680d & L.A.S.Johnson; holo: NSW ♀,( with cones); iso: MEL. Dioecious or rarely monoecious shrub 1–3 m high. Bark smooth. Branchlets ascending, to 19 cm long; articles terete, 5–12 mm long, 0.5–0.8 mm diam., mostly glabrous in furrows; phyllichnia rounded to angular; teeth 6–8, erect or slightly spreading, rarely overlapping at base, 0.3–0.8 mm long, not marcescent. Male spikes occasionally moniliform, 1–4.5 cm long, 5–7 whorls per cm; bracteoles persistent; anther 0.6–1 mm long. Cones cylindrical, sessile or on peduncle to 12 mm long; cone body 14–27 (rarely to 45) mm long, 8–15 mm diam.; bracteoles obtuse to broadly acute, pyramidal protuberance slightly shorter than bracteole body. Samara 5–8 mm long, very dark red-brown to black. Fig. 54E–F. Only at the northern end of Wilsons Promontory, Vic.; in low woodland in sand. Region: HOWE. Map 142. Vic.: Wilsons Promontory, A.Andersen NSW 154628 (NSW); 6 km S of Yanakie, 1 km N of entrance to Wilsons Promontory Natl Park, K.L.Wilson 6678, 6679 & L.A.S.Johnson (MEL, NSW); walking track to Five Mile Beach, between Barry Ck and Chinaman Ck crossings, D.E.Albrecht 2609 (NSW). Probably a taxon of recent hybrid origin from A. littoralis and A. paradoxa (which are in the region but not in the particular localities), but fairly well stabilised and self-reproducing. Differs from A. littoralis in being a smooth-barked shrub, with branchlets usually with

148

Figure 54. Allocasuarina cones ×1.5, branchlets ×15. A–B, A. inophloia. A, cone (NSW 62135, NSW); B, branchlet (NSW 62138, NSW). C–D, A. littoralis. C, cone (C.Moore 2671, NSW); D, branchlet (Tully Falls, Qld, A.Fielding, NSW). E–F, A. media. E, cone; F, branchlet (E–F, K.Wilson 6678b, NSW). G–H, A. filidens. G, cone; H, branchlet (G–H, S.Blake 22762, NSW). I, A. emuina, branchlet (P.Sharpe 2650, NSW). J, A. thalassoscopica, branchlet (P.Sharpe 2651, NSW). Drawn by D.Mackay.

149 3. Allocasuarina CASUARINACEAE glabrous furrows, with male spikes mostly moniliform, and with cones on generally shorter peduncles. Differs from A. paradoxa in the often more slender branchlets with erect to only slightly spreading teeth, which are more slender and often shorter than in A. paradoxa and are not marcescent.

28. Allocasuarina filidens L.Johnson, Fl. Australia 3: 195 (1989) T: Mt Beerwah, Qld, 13 Aug. 1966, S.T.Blake 22762; holo: NSW, with cones; iso: BRI. Dioecious shrub 1.5–3 m high. Bark becoming rough. Branchlets ascending, to 20 cm long; articles 12–15 mm long, 0.5–0.8 mm diam., smooth, when young with hairs protruding up to 2 mm from furrows but becoming glabrous; phyllichnia angular, with median ridge; teeth 5 or 6, spreading to recurved, not overlapping, 1.5–2.0 mm long, occasionally marcescent and c. 1 mm. Male spikes elongate but too young to describe details. Cones cylindrical to barrel-shaped, sparsely pubescent; peduncle 2–9 mm long; cone body 14–30 mm long, 11–18 mm diam.; bracteoles acute to broadly acute, often with small mucro, pyramidal protuberance as long as or exceeding bracteole body, with or without little mucro. Samara 6.5–9 mm long, dark brown to black. 2n = 44, B.A.Barlow pers. comm. (NSW 62406). Fig. 54G–H. Restricted to the Glass House Mtns, Qld; grows on summits and exposed upper slopes, in crevices of trachyte rocks with other shrubs. Region: MCPH. Map 143. Qld: Mt Beerwah, 12 June 1961, B.A.Barlow (BRIU, NSW 62406); Cruickneck, Glass House Mtns, 24 May 1935, D.A.Goy (BRI 246880); Mt Beerwah, K.Hill 1249 & L.A.S.Johnson (NSW). Differs from A. littoralis in being a shrub and in having longer articles and longer teeth which are recurved to spreading and only 5 or 6 in number.

29. Allocasuarina emuina L.Johnson, Fl. Australia 3: 195 (1989) T: Mt Emu, Qld, 28 Feb. 1981, P.R.Sharpe 2650; holo: NSW, with cones; iso: BRI. Dioecious spreading shrub 0.5–1.5 m high. Bark smooth. Branchlets ascending, to 12 cm long; articles 4–8 mm long, 0.5–0.9 mm diam., smooth, pubescent in furrows; phyllichnia angular to rounded; teeth 6 or 7, erect to slightly spreading, not overlapping, 0.3–0.7 mm long, not marcescent. Male spikes 1–3 cm long, 8.5–9.5 whorls per cm; bracteoles persistent; anther 0.8–0.9 mm long. Cones cylindrical, occasionally irregular and with a sterile apex to c. 5 mm long; peduncle 3–13 mm long, slender; cone body 12–28 mm long, 6–15 mm diam.; bracteoles broadly acute to obtuse, pyramidal protuberance shorter than bracteole body and diverging from it low down. Samara 4.5–7.5 mm long, dark brown to black. Fig. 54 I. Occurs from northerly slopes of Mt Emu, N of Coolum, S to Caloundra, Qld; grows in low heath in acid volcanic soil and in nearby wallum areas. Regions: DWSN, MCPH. Map 144. Qld: Caloundra, S.T.Blake 4105 (BRI, NSW); Sippy Ck, 16 km S of Nambour, J.E.Coaldrake QCC 356 (NSW); Mt Emu, L.A.S.Johnson 8560, P.R.Sharpe & K.L.Wilson (NSW), P.R.Sharpe 2650 (BRI, NSW), P.R.Sharpe 2723 (BRI, NSW); Emu Mtn near Coolum, C.T.White 11432 (BRI). The populations are slightly different. Closely related to A. thalassoscopica but can be distinguished by its shorter articles, longer anther, usually more rounded phyllichnia and less appressed teeth.

30. Allocasuarina thalassoscopica L.Johnson, Fl. Australia 3: 199 (1989) T: Mt Coolum, Qld, 23 Apr. 1981, L.A.S.Johnson 8562, P.R.Sharpe & K.L.Wilson; holo: NSW, with cones; iso: BRI. Similar to A. emuina. Shrub c. 1 m high, rarely to 2.5 m. Branchlet articles 7–12 mm long, 0.7–0.8 mm diam.; phyllichnia angular; teeth erect, 0.3–0.6 mm long, occasionally marcescent. Male spikes 0.5–4.5 cm long; anther 0.6–0.8 mm long. Cones on peduncle 8–14

150 CASUARINACEAE 3. Allocasuarina mm long; cone body 10–26 mm long, 9–15 mm diam.; bracteoles obtuse. Samara dark brown. Fig. 54J. Known only from the windswept S-facing upper slopes of Mt Coolum, Qld; forms a dense low closed heath. Region: MCPH. Map 145. Qld: Mt Coolum, P.R.Sharpe 2651, 2724 (BRI, NSW).

31. Allocasuarina rigida (Miq.) L.Johnson, J. Adelaide Bot. Gard. 6: 78 (1982) Casuarina rigida Miq., Rev. Crit. Casuarineae 61, t. 7D (1848); C. distyla var. rigida (Miq.) J.D.Hook., Fl. Tasman. 1: 348 (1857); C. stricta var. rigida (Miq.) Miq., Flora 48: 20 (1865). T: Moreton Bay [actually from or near A.Cunningham’s Mt Lindesay, now Mt Barney, Qld], C.Fraser 189; lecto: K, with cones, fide L.A.S.Johnson in W.M.Curtis, Stud. Fl. Tasmania 3: 653 (1967); isolecto: U. Dioecious shrub 0.5–4 m high. Bark smooth. Branchlets ascending, to 33 cm long; articles 10–25 mm long, 0.7–1.5 mm diam., smooth, usually glabrous; phyllichnia angular to sometimes rounded, with median ridge; teeth 7–10, spreading or recurved to erect, overlapping at least slightly when young, 0.4–1.3 mm long, usually marcescent. Male spikes occasionally moniliform, 1–7 cm long, rarely to 9 cm, 4–6.5 whorls per cm; bracteoles persistent; anther 0.7–1.2 mm long. Cones cylindrical to ovoid, pubescent; peduncle 2–9 mm long; cone body 8–27 mm long, 7–14 mm diam.; bracteoles truncate to obtuse, pyramidal protuberance shorter than bracteole body. Samara 3–7.5 mm long, mid to dark brown. Occurs from the McPherson Ra., Qld, S to the Ebor area, N.S.W., with isolated occurrences on Mt Cooroora, Qld, and in N.S.W. on the Koonyum Ra. and Big Nellie N of Taree; grows in poor sandy soils on acid granite, rhyolite or trachyte, in exposed situations. There are 2 subspecies. Articles 0.8–1.5 mm diam., 10–25 mm long; teeth recurved to spreading, overlapping 31a. subsp. rigida Articles 0.7–0.9 mm diam., 10–14 mm long; teeth erect to slightly spreading, overlapping slightly only when young 31b. subsp. exsul 31a. Allocasuarina rigida (Miq.) L.Johnson subsp. rigida Articles 10–25 mm long, 0.8–1.5 mm diam.; teeth recurved to suberect, overlapping, 0.4–1.3 mm long, marcescent. Male spikes not or shortly moniliform. Cone body 8–27 mm long, 7–14 mm diam. Samara 3–7.3 mm long. 2n = 22, B.A.Barlow, Austral. J. Bot. 7: 240 (1959). Fig. 55A–B. Throughout range of species except Mt Cooroora, Qld. Region: MCPH. Map 146. Qld: Mt Maroon, S.T.Blake 18244 (BRI, NSW). N.S.W.: Big Nellie, Landsdowne State Forest, D.Binns & W.Chapman DLB67 (NSW); Gibraltar Ra. Natl Park, R.G.Coveny 5687, 5688 & N.Lander (BRI, K, NSW); Barren Mtn, SE of Ebor, I.R.Telford (CBG 29732, 29736, NSW); Koonyum Ra. road, K.L.Wilson 5782 (BRI, K, L, MO, NSW). 31b. Allocasuarina rigida subsp. exsul L.Johnson, Fl. Australia 3: 198 (1989) T: Mt Cooroora, Qld, 12 Apr. 1984, P.R.Sharpe 3544; holo: NSW 179094, with cones; iso: BRI. Articles 10–14 mm long, 0.7–0.9 mm diam.; teeth erect to slightly spreading, overlapping slightly only when young, slightly narrower than in subsp. rigida, 0.4–0.9 mm long, somewhat marcescent. Male spikes strongly moniliform. Cone body 9–19 mm long, 6–11 mm diam. Samara 3.5–4.5 mm long. Fig. 55C. Restricted to Mt Cooroora near Pomona, Qld; grows on rocky (trachyte) hillside. Region: MCPH. Map 147. Qld: Mt Cooroora, P.R.Sharpe 3554 (BRI, NSW); Mt Cooroora, I.R.Telford 3442 (CBG, NSW). Differs from subsp. rigida (from which it is geographically isolated) in the less overlapping and spreading and slightly narrower teeth and in the strongly moniliform male spikes. From

151 3. Allocasuarina CASUARINACEAE

A. emuina and A. thalassoscopica it is distinguished by its longer articles with glabrous furrows, 7–9 teeth, and usually smaller cones.

32. L.Johnson, Fl. Australia 3: 198 (1989) T: Sydney Harbour Natl Park, N.S.W., 3 Apr. 1986, K.L.Wilson 6843 & L.A.S.Johnson; holo: NSW, with cones; iso: CANB, K, MEL. Slender dioecious shrub, 3–5 m high. Bark smooth. Branchlets drooping to spreading, to 27 cm long; articles terete, 13–20 mm long, 0.8–1 mm diam., usually with a faint waxy bloom, glabrous; phyllichnia rounded to sometimes angular; teeth 7 or 8, spreading to recurved, often slightly overlapping, 0.7–1.1 mm long, often marcescent. Male spikes moniliform, 5–10 cm long, 3.5–4.5 whorls per cm; bracteoles persistent; anther 0.8–1 mm long. Cones cylindrical; peduncle 2–15 mm long; cone body 12–15 mm long, 8–10 mm diam.; bracteoles obtuse, pyramidal, protuberance shorter than bracteole body. Samara 4–5 mm long, dark brown. Fig. 55D–E. Known only from 10 individuals in Sydney Harbour Natl Park, c. 6 km ENE of Sydney, N.S.W.; in tall shrubland on slope of sandstone headland. Region: NEPN. Map 148. N.S.W.: Sydney Harbour Natl Park P.Brookhouse NSW 181689–181691, 183659, K.L.Wilson 6840–6842 & L.A.S.Johnson (all in NSW). Differs from typical A. rigida and A. diminuta in the strongly moniliform male spikes. From A. rigida subsp. exsul it differs in its generally longer articles and often longer, broader teeth. It further differs from A. rigida in its more rounded phyllichnia and generally more slender articles; from A. diminuta in its spreading and mostly slightly overlapping teeth, its longer articles and its longer, less dense male spikes.

33. Allocasuarina rupicola L.Johnson, Fl. Australia 3: 199 (1989) T: 6.4 km on Boonoo Boonoo Falls road from Boonoo Boonoo, N.S.W., 25 Mar. 1981, L.A.S.Johnson 8539; holo: NSW, with cones; iso: BRI, CANB, K, MO. Similar to A. rigida. Shrub 1–3 m high. Branchlet to 18 cm long; articles 8–11 mm long, 0.7–0.9 mm diam.; teeth 7 or 8, erect to slightly spreading when old, 0.2–0.6 mm long, occasionally marcescent. Male spikes moniliform, 1–2.5 cm long, 7.5–8 whorls per cm; anther 0.7–0.8 mm long. Cones shortly cylindrical; peduncle 2–14 mm long; cone body 6–19 mm long, 6–10 mm diam. Samara 2.8–5 mm long. Fig. 55F–G. Restricted to the area between Wyberba, Qld, and Boonoo Boonoo Falls, N.S.W.; in clefts in granite on mountain slopes and along rocky creeks. Region: MCPH. Map 149. Qld: Wyberba, bank of Bald Rock Ck, S.T.Blake 4656 (BRI, NSW); Mt Norman, K.Hill 1257 & L.A.S.Johnson (NSW). N.S.W.: 6 km NNE of Boonoo Boonoo, K.Thurtell & R.G.Coveny 3894 (NSW). Close to A. rigida, but differs in teeth not spreading when fresh, and forming quite distinct populations.

34. Allocasuarina ophiolitica L.Johnson, Fl. Australia 3: 198 (1989) T: Bralga Tops, Glenrock Stn, N.S.W., 24 Mar. 1981, L.A.S.Johnson 8537; holo: NSW, with cones; iso: BRI, CANB, K, MO. Similar to A. rigida. Shrub 1–3 m high. Branchlets to 19 cm long; articles often with a waxy bloom, 7–14 mm long, 0.6–1 mm diam.; phyllichnia rounded to angular; teeth 7–9, erect to slightly spreading, not overlapping, 0.5–1.3 mm long, not marcescent. Male spikes not moniliform, 1–2.5 cm long, c. 6 whorls per cm; anther 0.8–1.2 mm long. Cones on peduncle 3–15 mm long; cone body 9–20 mm long, 7–12 mm diam.; bracteoles obtuse to very broadly acute. Samara 3.5–6 mm long, mid-brown. Fig. 55H–I. Restricted to the southern end of the New England region of N.S.W. and adjoining coastal ranges from Bralga Tops to Curricabark and Glenrock, N.S.W.; grows on serpentinite outcrops in tall heath and low open woodland. Region: MCPH. Map 150.

152

Figure 55. Allocasuarina cones ×1.5. A–B, A. rigida subsp. rigida. A, cone; B, branchlet ×15 (A–B, NSW 23723, NSW). C, A. rigida subsp. exsul, branchlet ×15 (P.Sharpe 3544, NSW). D–E, A. portuensis. D, cone; E, branchlet ×10 (D–E, K.Wilson 6843, NSW). F–G, A. rupicola. F, cone; G, branchlet ×15 (F–G, L.Johnson 8541, NSW). H–I, A. ophiolitica. H, cone; I, branchlet ×15 (H–I, NSW 26343, NSW). Drawn by D.Mackay.

153 3. Allocasuarina CASUARINACEAE

N.S.W.: Watchimbark Ck, NW of Gloucester, D.F.Blaxell 11 & R.G.Coveny (NSW); between Manning R. and Pigna Barney R., c. 1.5 km WSW of ‘Boonara’, L.A.S.Johnson NSW 103927 (NSW); Curricabark Ck, Curricabark, L.A.S.Johnson NSW 26343, 62396 (NSW). Similar to A. rigida, differing in its generally shorter and more slender articles with more rounded phyllichnia and more erect teeth (which are not overlapping or marcescent), its cone bracteoles (which are often more acute) and its paler samara. Plants small in more exposed situations, tree-like in more sheltered situations.

35. (Vent.) L.Johnson, J. Adelaide Bot. Gard. 6: 75 (1982) Casuarina distyla Vent., Jard. Cels 62, t. 62 (1802). T: Cap de Diemen [but presumably from N.S.W.], cult. Jardin de Cels, Paris, France, herb. Ventenat; lecto: G, with cones, n.v., fide L.A.S.Johnson, J. Adelaide Bot. Gard. 6: 75 (1982); isolecto: K, P, NSW (fragment ex G). Dioecious shrub 1–3 m high. Bark mainly smooth. Branchlets ascending, to 35 cm long; articles smooth, 10–20 mm long, 0.8–1.5 mm diam. occasionally waxy, pubescent in furrows; phyllichnia angular to rounded, occasionally sparsely and minutely pubescent; teeth 6–8, erect, slightly overlapping at least when young, 0.5–1.2 mm long, occasionally marcescent. Male spikes occasionally moniliform, 1.5–5 (rarely to 9) cm long, 4.5–6.5 whorls per cm; bracteoles persistent; anther 0.8–1.3 mm long. Cones long-cylindrical, often with sterile apex to 12 mm long; peduncle 2–15 mm long, rarely to 32 mm long and 2–4 mm diam.; cone body 13–35 (rarely to 50) mm long, 11–22 mm diam.; bracteoles obtuse to truncate, pyramidal protuberance shorter than bracteole body. Samara 4–8 mm long, very dark brown to black. 2n = 22, 33, B.A.Barlow, Austral. J. Bot. 7: 241 (1959). Figs 45G, 56A–B. Occurs from the S side of Port Stephens and Gospers Mtn S to the ranges E of Cooma, N.S.W., and perhaps on the upper Genoa R., Vic.; grows in tall heath on sandstone ridge tops. Regions: NEPN, HOWE. Map 151. N.S.W.: Mt Jillacambra, D.E.Albrecht 1680 (MEL, NSW); 37 km N of Clarence, E.F.Constable NSW 52370 (NSW); McCarrs Ck, R.Melville 526B (K, MEL, NSW); c. 6 km E of Nerriga, Braidwood–Nowra road, R.Pullen 4107 (AD, BRI, CANB, K, L, MEL, NSW); Fingal Bay, K.L.Wilson 4409, 4410 (MO, NSW). Hybrids between this species and A. littoralis are quite common in disturbed areas between Broken Bay and Port Hacking.

36. Allocasuarina simulans L.Johnson, Fl. Australia 3: 199 (1989) T: 6.4 km SE of Nabiac at old landing site, N.S.W., 16 Feb. 1975, R.G.Coveny 6031, P.Hind & R.Hancock; holo: NSW, with cones; iso: A, K, L, MO, RSA. Similar to A. distyla. Branchlets to 19 cm long; articles 13–22 mm long, 0.9–1.3 mm diam., with furrows pubescent or glabrous (often both conditions on same plant); phyllichnia rounded to angular; teeth 6, 0.5–1.1 mm long. Male spikes 1.5–4.5 cm long, c. 4 whorls per cm; anther c. 1.3 mm long. Cones often irregular with sterile apex to 12 mm long; peduncle 3–14 mm long and c. 2 mm diam.; cone body 14–33 mm long, 9–12 mm diam.; bracteoles broadly acute to obtuse. Samara 4.5–6 mm long. Fig. 56C. Only known from Myall Lakes area between Booti Booti and Nabiac, N.S.W.; grows in heath in sand. Region: NEPN. Map 152. N.S.W.: old Nabiac aerodrome, L.A.S.Johnson 8531 (NSW); Booti Booti State Recreation Area, K.L.Wilson 2561, 2563 & J.T.Waterhouse (NSW). Similar to A. distyla but generally more slender, and with the phyllichnia less angular when dry. The cones have smaller parts.

154 CASUARINACEAE 3. Allocasuarina

37. L.Johnson, Fl. Australia 3: 194 (1989) T: c. 3 km E of ‘Wattle Valley’ turn-off on Kangarooby Rd, N.S.W., 29 Apr. 1982, K.L.Wilson 4451; holo: NSW, with cones; iso: CANB, K. Dioecious or monoecious shrub or small tree, 1–5 m high. Bark smooth. Branchlets ascending, to 23 cm long; articles terete, 5–12 mm long, 0.6–1.1 mm diam., often with a waxy bloom, glabrous; phyllichnia rounded to angular; teeth 6–9, erect to spreading, mostly not overlapping, 0.3–0.8 mm long, not marcescent. Male spikes occasionally moniliform, 0.5–5 cm long, 5–10 whorls per cm; bracteoles persistent; anther 0.5–0.8 mm long. Cones shortly to long-cylindrical, occasionally irregular, pubescent at least when young; peduncle 2–10 mm long; cone body 5–20 mm long, 5–12 mm diam.; bracteoles truncate to broadly acute, pyramidal protuberance shorter than bracteole body. Samara 3.5–5 mm long, dark brown. Occurs in N.S.W. in several disjunct areas, from the Pilliga Scrub S to Temora, from Capertee to Bathurst and from Blackheath S to Braidwood, and isolated occurrences in the Sydney area and near Eden; on tablelands, slopes and coastal ranges, on sandstone ridges and hillsides in heath and low open woodland. Differs from A. rigida in its mostly shorter and more slender articles, with teeth often shorter and mostly not overlapping or strongly spreading or marcescent, and in its often smaller cones and in often being monoecious. Differs from A. gymnanthera, which has a partly overlapping range, in its glabrous articles with more strongly rounded phyllichnia and mostly non-overlapping teeth, its rarely moniliform male spikes with persistent bracteoles and smaller anthers, and its smaller cones. There are 3 subspecies. 1 Phyllichnia rounded (occasionally only slightly so); teeth straight-sided 37a. subsp. diminuta 1: Phyllichnia angular to rounded 2 Teeth broadly deltoid, convex-sided; articles often with a waxy bloom 37b. subsp. mimica 2: Teeth narrowly or broadly deltoid, straight- or convex-sided; articles not waxy 37c. subsp. annectens 37a. Allocasuarina diminuta L.Johnson subsp. diminuta Shrub 2–5 m high. Articles often with a waxy bloom; phyllichnia rounded, occasionally only slightly so; teeth 6 or 7, narrowly deltoid, straight-sided, 0.3–0.5 mm long. Anther 0.5–0.7 mm long. Cone bracteoles obtuse to truncate. 2n = 22, B.A.Barlow, Austral. J. Bot. 7: 240 (1959), as Casuarina sp. 2 p.p. Fig. 56D–E. Occurs from the Pilliga Scrub S to Temora, and in the Capertee to Bathurst area, N.S.W.; on tablelands and western slopes. Regions: DARL, NEPN. Map 153. N.S.W: Yetholme trig, R.G.Coveny 9659 (K, L, NSW); 42 km from Cowra along road to Boorowa, M.D.Crisp 7260 & I.R.Telford (CBG, MEL, NSW); 17.5 km S of Salt Caves on Gibbican Forest Rd, Timmallallie State Forest, K.L.Wilson 2460, 2461 & J.Waterhouse (K, NSW). 37b. Allocasuarina diminuta subsp. mimica L.Johnson, Fl. Australia 3: 195 (1989) T: Bundanoon–Penrose, N.S.W., Apr. 1948, M.Tindale NSW 62492; holo: NSW♂, ( with cones - monoecious). Shrub 1–2.5 m high. Articles often with a waxy bloom; phyllichnia angular to rounded; teeth 6–10, broadly deltoid, convex-sided, 0.3–0.6 mm long. Anther 0.5–0.7 mm long. Cone bracteoles obtuse to broadly acute. 2n = 22, 44, B.A.Barlow, Austral. J. Bot. 7: 240 (1959), as Casuarina sp. 2 p.p. Fig. 56F. Occurs in the Sydney area from Kingsford to Little Bay and NW of Heathcote and also in the Central Tablelands from Blackheath to Taralga and Bundanoon. Region: NEPN. Map 154.

155

Figure 56. Allocasuarina cones ×1.5. A–B, A. distyla. A, cone; B, branchlet ×7.5 (A–B, NSW 62460, NSW). C, A. simulans, branchlet ×10 (L.Johnson 8529, NSW). D–E, A. diminuta subsp. diminuta. D, cone; E, branchlet ×15 (D–E, NSW 62473, NSW). F, A. diminuta subsp. mimica, branchlet ×15 (J.Waterhouse 5793, NSW). G, A. diminuta subsp. annectens, branchlet ×15 (A.Orchard 4487, NSW). H, A. defungens, branchlet ×15 (L.Johnson 8505, NSW). I, A. glareicola, branchlet ×15 (NSW 155948, NSW). J, A. grampiana, branchlet ×15 (NSW 71533, NSW). Drawn by D.Mackay.

156 CASUARINACEAE 3. Allocasuarina

N.S.W.: Colong–Yerranderie, R.H.Cambage 2281 (NSW); 5.1 km N of Menai turn-off on Heathcote Rd, R.G.Coveny 11166 & J.Thomas (K, NSW); Long Bay, L.A.S.Johnson & E.F.Constable NSW 47620 (NSW); Berrima, K.L.Wilson 5924, 5925 (CANB, K, NSW). 37c. Allocasuarina diminuta subsp. annectens L.Johnson, Fl. Australia 3: 194 (1989) T: Corang R., Braidwood–Nerriga road, N.S.W., 24 Sept. 1960, R.Pullen 2207; holo: NSW ♂,( with cones - monoecious); iso: CANB, MEL. Shrub 1–2.5 m high. Articles not waxy; phyllichnia angular to rounded; teeth 6–8, broadly to narrowly deltoid, convex- or straight-sided, 0.4–0.8 mm long. Anther 0.6–0.8 mm long. Cone bracteoles obtuse to truncate. Fig. 56G. Occurs from Sassafras to Lake Bathurst and Braidwood, and with an apparently isolated occurrence SW of Eden, N.S.W.; on coastal ranges and adjacent tablelands. Region: HOWE. Map 155. N.S.W.: W of Sassafras, L.A.S.Johnson NSW 62491 (NSW); Wallagaraugh R.–Imlay Ck junction, 31 km SW of Eden, I.R.Telford 6833, 6834 (BISH, CBG, NSW); Oak Park, 3.5 km S of Sandy Point junction on Cullulla road, K.L.Wilson 5928–5934 (CANB, K, NSW). Morphologically intermediate between the other 2 subspecies, but with a more southern distribution.

38. L.Johnson, Fl. Australia 3: 194 (1989) T: Nabiac airstrip, N.S.W., 2 Sept. 1967, R.G.Coveny 73; holo: NSW ♂,( ♀ - monoecious). Dioecious or monoecious, lignotuberous, depauperate, erect or suberect shrub 0.5–2 m high. Bark smooth. Branchlets ascending, to 12 cm long; articles 6–8 mm long, 0.5–0.6 mm diam., smooth, glabrous; phyllichnia rounded (often only slightly so); teeth 5–7, erect, slightly overlapping when young, 0.2–0.5 mm long, often marcescent at apex. Male spikes shortly elongate, moniliform, 4–10 mm long; bracteoles persistent; anther 0.7–0.9 mm long. Cones very irregular, shortly cylindrical, on slender peduncle 3–7 mm long; cone body 8–11 mm long, 5–7 mm diam.; bracteoles smooth, obtuse to truncate with tiny mucro, pyramidal protuberance shorter than bracteole body. Samara c. 3 mm long, dark brown. Fig. 56H. Apparently restricted to Nabiac area, N.S.W.; in tall heath in sand. Region: NEPN. Map 156. N.S.W.: 7 km ESE of Nabiac, 3.5 km by road E of Pacific Hwy, L.A.S.Johnson 8510 (NSW). Distinguished from A. diminuta and A. rigida by its slender articles with only slightly rounded phyllichnia and 5–7 teeth, and its usually short moniliform male spikes. It may hybridise with A. littoralis. Differs from A. glareicola in its blue-green articles. Endangered or possibly now destroyed by sand-mining.

39. Allocasuarina glareicola L.Johnson, Fl. Australia 3: 195 (1989) T: Castlereagh State Forest, N.S.W., 11 Oct. 1983, D.Benson & D.Keith NSW 155948; holo: NSW♂, ( ♀ - monoecious). Dioecious or monoecious slender, erect shrub 1–2 m high. Bark smooth. Branchlets ascending, to 20 cm long; articles 5–11 mm long, 0.5–0.7 mm diam., smooth, glabrous; phyllichnia rounded; teeth 5–7, erect to slightly spreading, not or slightly overlapping when young, 0.2–0.5 mm long, not marcescent. Male spikes moniliform, 0.2–2.5 cm long, c. 5 whorls per cm; bracteoles persistent; anther 0.7–0.8 mm long. Cones cylindrical, on peduncle 4–7 mm long; cone body 10–13 mm long, 7–8 mm diam.; bracteoles broadly acute to obtuse with tiny mucro, pyramidal protuberance much shorter than bracteole body, with obtuse apex. Samara 3.0–3.5 mm long, mid-brown. Fig. 56 I. Restricted to a few small populations in and beside Castlereagh State Forest NE of Penrith on Cumberland Plain, N.S.W.; in open forest in Tertiary alluvial gravels with clayey subsoil. Region: NEPN. Map 157.

157 3. Allocasuarina CASUARINACEAE

N.S.W.: just E of Castlereagh State Forest, K.L.Wilson 5919–5922, L.A.S.Johnson & D.Benson (NSW); Castlereagh State Forest, K.L.Wilson 5911–5915, L.A.S.Johnson & D.Benson (NSW). Differs from A. rigida in its shorter, more slender articles (which often have only slightly rounded phyllichnia), 5–7 teeth which are usually shorter, not marcescent and much less spreading and overlapping, and its smaller cones. Differs from A. defungens in its yellow-green articles, phyllichnia that are occasionally strongly rounded, and teeth that are straighter-sided and often somewhat spreading.

40. L.Johnson, Fl. Australia 3: 196 (1989) T: upper slopes of Mt Rosea, The Grampians, Vic., 26 Aug. 1965, D.E.Symon 3507; holo: NSW, with cones; iso: AD, MEL. Dioecious shrub 1.5–4 m high. Bark smooth. Branchlets ascending, to 15 cm long; articles 5–12 mm long, 0.7–1 mm diam., smooth, glabrous, with a waxy bloom; phyllichnia angular to rounded; teeth 6 or 7, erect or rarely slightly spreading, occasionally overlapping when young, 0.4–0.8 mm long, not marcescent. Male spikes moniliform, 1–4 cm long, 5 or 6 whorls per cm; bracteoles persistent; anther 0.7–0.8 mm long. Cones long-cylindrical, with short sterile apex to 3 mm long; peduncle 2–6 mm long; cone body 13–35 mm long, 7–9 mm diam.; bracteoles broadly acute to obtuse, pyramidal protuberance shorter than bracteole body. Samara 4.5–5.5 mm long, black. 2n = 22, 33, 44, B.G.Briggs pers. comm. (NSW 89356, 76102). Fig. 56J. Restricted to The Grampians, Vic.; associated with sandstone outcrops. Region: OTWY. Map 158. Vic.: Boronia Peak, near Halls Gap, A.C.Beauglehole 30171 (MEL, NSW); below the Mt Military walk turn-off, The Grampians, D.E.Symon 3500 (AD, NSW); Mt William, H.Streimann 2991 (A, CBG, L, NSW); Mt Thackeray, Victoria Ra., H.Streimann 2826, 2828 (A, CBG, L, NSW). Close to A. monilifera and A. zephyrea of Tas. but distinguished by a combination of the phyllichnia being generally more angular and the articles having a stronger waxy bloom.

41. Allocasuarina zephyrea L.Johnson, Fl. Australia 3: 199 (1989) T: Ocean Beach, Strahan, Tas., 20 Jan. 1949, L.A.S.Johnson NSW 62623; holo: NSW, with cones. Similar to A. grampiana. Shrub 0.5–2 m high. Branchlets to 19 cm long; articles 4–15 mm long, 0.6–1.3 mm diam., occasionally slightly waxy; phyllichnia angular or rounded; teeth 7–9, rarely 10, spreading to erect, often slightly overlapping, 0.4–1.2 mm long, the apex eventually marcescent. Male spikes often moniliform, 1–3.5 cm long, 5–7 whorls per cm; anther 0.7–0.9 mm long. Cones on peduncle 2–15 mm long; cone body 10–25 mm long, 6–11 mm diam. Samara 4.0–5.5 mm long, black. 2n = 22, B.A.Barlow, Austral. J. Bot. 7: 240 (1959), as Casuarina sp. 4 p.p. Fig. 57A–C. Occurs from the western lowlands to central and south-eastern highlands, Tas., also on King Is.; grows in heath and on rocky outcrops. Region: TASM. Map 159. Tas.: Seal Rocks track, King Is., M.I.H.Brooker 5854 (FRI, NSW); Smithton, N.T.Burbidge 3469 (CANB, HO, K, NSW); Cradle Mtn, N.T.Burbidge 3519 (CANB, HO, K, NSW); Port Davey to New Harbour, C.Davis NSW 62625 (NSW); West Point, SW of Marrawah, K.L.Wilson 6387, 6388 (HO, NSW). There is variation between populations in length of articles and in shape and amount of overlapping of teeth, and this may prove worthy of taxonomic recognition. Forms close to the sea are coarser and there is a fairly well-defined form with short slender articles on the highlands from Cradle Mtn to Lake St Clair and Lake Margaret.

158 CASUARINACEAE 3. Allocasuarina

42. (L.Johnson) L.Johnson, J. Adelaide Bot. Gard. 6: 76 (1982) Casuarina monilifera L.Johnson in W.Curtis, Stud. Fl. Tasmania 3: 651 (1967). T: Tasman Arch, Eaglehawk Neck, Tas., 15 Jan. 1949, L.A.S.Johnson NSW 62593; holo: NSW ♂);( iso: HO. Similar to A. grampiana. Usually monoecious shrub 1.5–4 m high. Branchlet articles 6–11 mm long, 0.6–1.2 mm diam., at least slightly waxy; phyllichnia strongly to slightly rounded; teeth 6–9, erect and appressed to slightly spreading, often overlapping, eventually the apex marcescent, 0.5–1 mm long. Male spikes 1–3.5 cm long, with 3.5–7 whorls per cm; anther 0.7–1.2 mm long. Cones on peduncle 2–10 mm long; cone body 15–30 mm long, 8–14 mm diam.; bracteoles truncate to broadly acute; bracteole protuberance shorter than or occasionally equalling body. Samara 5–6 mm long, very dark brown to black. 2n = 44, B.A.Barlow, Austral. J. Bot. 7: 240 (1959), as Casuarina sp. 4 p.p. Fig. 57D–E. Occurs in northern and eastern Tas., also Flinders Is. and Kent Group in Bass Strait; grows in lowland heath and open woodland in sandy soil. Region: TASM. Map 160. Tas.: Kent Group, Allen NSW 62590 (NSW); Cape Tourville, J.Armstrong 878 & J.Powell (HO, NSW); Boat Harbour, N.T.Burbidge 3093, 3096 (CANB, HO, K, NSW); Five Mile Lagoon, Flinders Is., P.B.Nye 10 (A, HO, K); Dover, M.E.Phillips CBG 13903 (CBG, NSW). Similar to A. grampiana. Also close to A. zephyrea but differing in being tetraploid and mostly monoecious and in having slightly broader teeth that are mostly erect and appressed (occasionally teeth spreading) and in typically having articles more waxy. Articles vary in coarseness, while phyllichnium shape varies from strongly and narrowly rounded with a prominent rounded ridge to nearly flat, and teeth vary from appressed to spreading. However, there is no clear pattern to this variation and hence no formal subspecific taxa are here recognised.

43. L.Johnson, Fl. Australia 3: 194 (1989) T: Cape Pillar, Tasman Peninsula, Tas., Mar. 1976, T.Moscal NSW 145870; holo: NSW, with cones. Dioecious or monoecious shrub, 1–2 m high. Bark smooth. Branchlets spreading to ascending, to 17 cm long; articles 10–20 mm long, 1.2–2 mm diam., smooth, with densely pubescent furrows; phyllichnia strongly rounded; teeth 7–10, slender, spreading to slightly recurved, usually not overlapping, 1.1–3 mm long, not marcescent. Male spikes rarely moniliform, c. 2 cm long, 3.5–4 whorls per cm; bracteoles persistent; anther 0.8–1 mm long. Cones long-cylindrical, sessile or on peduncle to 3 mm long; cone body 15–34 mm long, 12–15 mm diam.; bracteoles obtuse to truncate, pyramidal protuberance slightly shorter than bracteole body. Samara 5–8 mm long, black. Fig. 57F. Only known from Tasman Is. and the Cape Pillar area on Tasman Peninsula, Tas.; grows on exposed cliff-tops. Region: TASM. Map 161. Tas.: between The Blade and Cape Pillar, 4 Dec. 1984, P.M.Buchanan (HO); Tasman Is., 26 Sept. 1965, J.Thwaites (HO, NSW); c. 9 km along track to Cape Pillar, just S of Corruption Gully, K.L.Wilson 6527–6541 (HO, NSW). Differs from A. monilifera, which also grows on the Tasman Peninsula, in its larger and thicker articles with strongly pubescent furrows and with more marcescent, longer teeth, and its larger samara.

44. Allocasuarina lehmanniana (Miq.) L.Johnson, J. Adelaide Bot. Gard. 6: 76 (1982) Casuarina lehmanniana Miq. in J.G.C.Lehmann, Pl. Preiss. 1: 639 (1845). T: Hay District, W.A., 7 Nov. 1840, L.Preiss 2005; lecto: U♀ (), fide L.A.S.Johnson, J. Adelaide Bot. Gard. 6: 76 (1982); isolecto: LE, MEL 538293. Illustration: L.Diels in A.Engler, Bot. Jahrb. Syst. 35: fig 10F–H (1905) as C. distyla. Similar to A. grampiana. Dioecious or less commonly monoecious shrub 1.5–4 m high. Branchlet articles 5–9 mm long, 0.7–1.2 mm diam., occasionally with a waxy bloom; phyllichnia rounded to angular; teeth 7 or 8, not overlapping, 0.4–0.8 mm long, not marcescent. Male spikes often moniliform, 1–3 (rarely to 6) cm long, 5–8 whorls per cm;

159

Figure 57. Allocasuarina cones ×1.5. A–C, A. zephyrea. A, cone; B, branchlet ×7.5 (A–B, P.Darbyshire 1107, NSW); C, branchlet ×15 (NSW 62622, NSW). D–E, A. monilifera. D, branchlet ×10 (NSW 62570, NSW); E, branchlet ×10 (J.Armstrong 8788, NSW). F, A. crassa, branchlet ×7.5 (NSW 168721, NSW). G–H, A. lehmanniana subsp. lehmanniana. G, cone (B.Briggs 565, NSW); H, branchlet ×15 (B.Briggs 564, NSW). I, A. lehmanniana subsp. ecarinata, branchlet ×10 (K.Newbey 1253, NSW). J–K, A. paradoxa. J, cone (NSW 62555, NSW); K, branchlet ×10 (NSW 62552, NSW). Drawn by D.Mackay.

160 CASUARINACEAE 3. Allocasuarina anther 0.6–0.9 mm long. Cones on peduncle 2–38 mm long, rarely sessile; cone body 12–35 mm long, 7–12 mm diam. Samara 4–5.5 mm long, black. 2n = 22, B.A.Barlow, Austral. J. Bot. 7: 240 (1959). Occurs from Jurien Bay S to near Albany and E to Ravensthorpe, W.A.; grows in woodland and tall scrub, often near the sea, in sandy-loam and sandy soils. There are 2 subspecies. Phyllichnia strongly rounded to angular, without median groove; teeth 7 or 8; cones on peduncle 4–38 (rarely 2) mm long 44a. subsp. lehmanniana Phyllichnia flat to slightly rounded, often with median groove; teeth 6 or 7; cones sessile or on peduncle to 4 mm long 44b. subsp. ecarinata 44a. Allocasuarina lehmanniana (Miq.) L.Johnson subsp. lehmanniana Casuarina baxteriana Miq., Rev. Crit. Casuarineae 37, t. 3C (1848); C. suberosa var. baxteriana (Miq.) Miq., Flora 48: 21 (1865). T: King George Sound, W.A., herb. Hooker; holo: K, with cones; iso: U. Casuarina microstrobilus C.Gardner, J. Proc. Roy. Soc. W. Australia 9: 37 (1923). T: Cranbrook, W extremity of Stirling Ra., W.A., 7 Mar. 1922, C.A.Gardner 1774; holo: PERTH, with cones. Dioecious or rarely monoecious shrub 1–4 m high. Branchlets to 20 cm long; articles with phyllichnia strongly rounded to angular, without median groove; teeth 7 or 8. Cones on peduncle 4–38 (rarely to 2) mm long; cone body 13–35 mm long, 9–12 mm diam. Fig. 57G–H. Occurs from Jurien Bay S to E of Albany, W.A. Regions: LUWN, ESPR. Map 162. W.A.: 26 km S of Kojonup, J.S.Beard 7415 (K, NSW, PERTH); 1.5 km S of Lancelin, E.M.Bennett 1334 (NSW, PERTH); Dempster Inlet, Middle Mt Barren, C.A.Gardner 9202 (PERTH); between Jurien Bay and Green Head, M.E.Phillips CBG 25646 (CBG, NSW). 44b. Allocasuarina lehmanniana subsp. ecarinata L.Johnson, Fl. Australia 3: 196 (1989) T: creek behind S end of Mylies Beach, near East Mt Barren, 19 Nov. 1985, A.N.Rodd 5089 & J.McCarthy; holo: NSW ♂,( ♀, with cones - monoecious); iso: K, PERTH. Dioecious or monoecious shrub 0.5–1 m high. Branchlets to 12 cm long; articles with phyllichnia flat to slightly rounded, often with median groove; teeth 6 or 7. Cones sessile or on peduncle to 4 mm long; cone body 12–24 mm long, 7–11 mm diam. Fig. 57 I. Occurs from near Ongerup to E of Esperance, W.A. Region: ESPR. Map 163. W.A.: 13 km NE of Ongerup, K.Newbey 1253 (NSW, PERTH); c. 3 km NW of Young R. crossing on Ravensthorpe–Esperance road, N.N.Donner 2765 (AD, NSW); Lynburn, Alexander R., H.P.Turnbull NSW 62835 (NSW).

45. (Macklin) L.Johnson, J. Adelaide Bot. Gard. 6: 77 (1982) Casuarina paradoxa Macklin, Bull. Misc. Inform., Kew 150 (1931). T: Cheltenham, Vic., May 1925, W.Audas; lecto: AD, with cones, fide L.A.S.Johnson in W.M.Curtis, Stud. Fl. Tasmania 3: 653 (1967); isolecto K. Dioecious or less often monoecious shrub 0.5–2 m high. Bark smooth. Branchlets ascending, to 15 cm long; articles 6–14 mm long, 0.6–1.2 mm diam., smooth, glabrous; phyllichnia rounded; teeth 7–11, spreading to recurved, usually somewhat overlapping, 0.3–2 mm long, marcescent. Male spikes rarely moniliform, 1–3 cm long, 3.5–7 whorls per cm; bracteoles persistent; anther 0.7–1.2 mm long. Cones cylindrical, sessile or on peduncle to 3 mm long; cone body 13–25 mm long, 7–13 mm diam.; bracteoles obtuse to broadly acute, pyramidal protuberance slightly shorter than bracteole body. Samara 4–8 mm long, very dark red-brown to black. 2n = 44, B.A.Barlow, Austral. J. Bot. 7: 240 (1959) as C. pusilla subsp. 5. Fig. 57J–K. Occurs in The Grampians and from the Melbourne area to Wilsons Promontory, Vic.; often grows in tall heath in sandy soil. Regions: OTWY, HOWE. Map 164.

161 3. Allocasuarina CASUARINACEAE

Vic.: Sandringham, A.Meebold 21533 (NSW); Cranbourne, Royal Botanic Gardens Annexe, J.H.Ross 2597, 2602 & M.G.Corrick (MEL, NSW); 9.5 km by road N of Tidal R., 2 km N of Darby Saddle, Wilsons Promontory Natl Park, K.L.Wilson 6685–6687 & L.A.S.Johnson (MEL, NSW); Halls Gap–Dunkeld road, at turn-off to Teddy Bears Gap (Serra Rd), K.L.Wilson 6725, 6726 & L.A.S.Johnson (MEL, NSW). Allocasuarina paradoxa and A. mackliniana are closely related. They are more or less sympatric in The Grampians, but have separate ranges elsewhere. A. mackliniana is typically coarser than A. paradoxa (although the Wilsons Promontory form of A. paradoxa may be coarse), with phyllichnia less strongly rounded and typically with teeth strongly marcescent, more overlapping and recurved especially on the very dense, thick male spikes. The two species are closely related to A. misera and A. pusilla.

46. Allocasuarina mackliniana L.Johnson, Fl. Australia 3: 196 (1989) T: 5 km E of Lucindale on Lochaber road, S.A., 8 Mar. 1986, K.L.Wilson 6787 & L.A.S.Johnson; holo: NSW (♀, with cones); iso: AD, K. Dioecious or rarely monoecious shrub 0.5–3 m high. Bark smooth. Branchlets ascending, to 20 cm long; articles 7–17 mm long, 0.8–1.4 mm diam., smooth, pubescent or glabrous in furrows; phyllichnia slightly rounded; teeth 7–10, recurved to spreading, usually overlapping at least at bases, 0.7–2 mm long, strongly marcescent. Male spikes very dense and thick, 1–4 cm long, 4.5–10 whorls per cm; bracteoles persistent; anther 0.8–1.5 mm long. Cones cylindrical, occasionally broader than long, sessile or on peduncle to 3 mm long; cone body 12–22 (rarely to 30) mm long, 8–14 mm diam.; bracteoles obtuse to broadly acute, pyramidal protuberance slightly shorter than bracteole body. Samara 5–7.5 mm long, dark red-brown to black. Occurs from the Southern Lofty Ranges, S.A., east to western Vic. Closely related to A. paradoxa. There are 3 subspecies. 1 Branchlets with phyllichnia rounded and furrows glabrous 46a. subsp. mackliniana 1: Branchlets with phyllichnia nearly flat to slightly rounded 2 Branchlets with furrows pubescent, often strongly so; phyllichnia slightly rounded 46b. subsp. hirtilinea 2: Branchlets with furrows pubescent usually only when young; phyllichnia nearly flat to slightly rounded 46c. subsp. xerophila 46a. Allocasuarina mackliniana L.Johnson subsp. mackliniana Articles 9–17 mm long, 0.8–1.2 mm diam., with furrows usually glabrous; phyllichnia rounded; teeth 7–10, 0.7–2 mm long. Male spikes with 6–9 whorls per cm; anther 0.9–1.1 mm long. 2n = 44, B.A.Barlow, Austral. J. Bot. 7: 240 (1959) as C. pusilla subsp. 5. Fig. 58A–B. Occurs from the southern Lofty Ranges, S.A., to near Portland, Vic.; in heath in sandy soils. Regions: EYRE, OTWY. Map 165. S.A.: Goolwa road near Square Waterhole, c. 2 km S of Mt Compass, K.Czornij 687 (AD); c. 32 km S of Bordertown on Naracoorte road, L.A.S.Johnson 7925 & K.L.Wilson (AD, NSW); Mt Magnificent Conservation Park, L.A.S.Johnson 8672 & K.Horsell (AD, K, NSW). Vic.: 18 km SW of Edenhope post office, A.C.Beauglehole 49748 (MEL, NSW); Mount Clay, near Portland, R.Melville 1640, 1641 (K, NSW). 46b. Allocasuarina mackliniana L.Johnson subsp. hirtilinea L.Johnson, Fl. Australia 3: 197 (1989) T: NE side of Mt Zero, Vic., 21 Aug. 1968, A.C.Beauglehole 28185 (♀, with cones); holo: NSW; iso: MEL. Articles 11–15 mm long, 1–1.3 mm diam., with furrows pubescent, often strongly so; phyllichnia slightly rounded; teeth 8 or 9, 1.5–2 mm long. Male spikes with c. 7 whorls per cm; anther 1.1–1.5 mm long. Fig. 58C.

162 CASUARINACEAE 3. Allocasuarina

Known only from the western parts of The Grampians, Vic. Grows in woodland in sandy soils. Region: OTWY. Map 166. Vic: Mt Zero, 24 km SE of Horsham, H.Streimann 2644 (CBG, NSW); Victoria Ranges, collector unknown (NSW 62580). 46c. Allocasuarina mackliniana L.Johnson subsp. xerophila L.Johnson, Fl. Australia 3: 197 (1989) T: Scorpion Springs Conservation Park, S of Pinnaroo, S.A., 21 Oct. 1973, D.E.Symon 8610 (♂); holo: NSW; iso: AD n.v. Articles 7–13 mm long, 0.8–1.4 mm diam., with furrows pubescent when young, usually becoming glabrous; phyllichnia nearly flat to slightly rounded; teeth 7 or 8, 0.7–1.5 mm long. Male spikes with 4.5–5 whorls per cm; anther 0.8–1.5 mm long. 2n = 44, A.H.Burbidge, B.Sc. Hons. thesis, Flinders University, as ‘C. robusta’. Fig. 58D–E. Occurs from near Pinnaroo and Keith, S.A., to Wyperfeld Natl Park and the Little Desert, Vic.; in heath in sandy soils. Regions: RIVR, OTWY. Map 167. S.A.: 6 km NW of Keith, L.A.S.Johnson NSW 47220 (NSW); Mt Shaugh Conservation Park, D.E.Symon 10762 (AD, K, MO, NSW). Vic.: Quandong Hill, Wyperfeld Natl Park, A.C.Beauglehole 29349 (MEL, NSW); 16 km S of Nhill on Goroke–Gymbowen road, K.L.Wilson 6765, 6766 & L.A.S.Johnson (K, MEL, NSW). Some forms from the Little Desert may approach subsp. hirtilinea; there is also intergradation to subsp. mackliniana generally in the southern part of the range.

47. Allocasuarina misera L.Johnson, Fl. Australia 3: 197 (1989) T: Anglesea, Vic., 10 Mar. 1986, K.L.Wilson 6818 & L.A.S.Johnson; holo: NSW♀, ( with cones); iso: MEL. Dioecious or monoecious shrub 0.5–2 m high. Bark smooth. Branchlets ascending, to 10 cm long; articles 4–10 mm long, 0.3–0.8 mm diam., smooth, usually glabrous in furrows; phyllichnia rounded to nearly flat; teeth 5–7, erect and appressed to occasionally spreading, slightly overlapping, 0.3–0.8 mm long, occasionally marcescent. Male spikes rarely moniliform, 0.5–1.5 cm long, 9–11 whorls per cm; bracteoles persistent; anther 0.5–0.8 mm long. Cones cylindrical, sessile or on peduncle to 5 mm long; cone body 9–16 mm long, 7–13 mm diam.; bracteoles obtuse to acute, pyramidal protuberance usually slightly shorter than bracteole body. Samara 4–6 mm long, very dark red-brown to black. Fig. 58F. Occurs from The Grampians to near Bairnsdale, Vic.; grows in heath or open woodland in sandy soil. Regions: OTWY, HOWE. Map 168. Vic.: Providence Ponds Flora and Fauna Reserve, A.C.Beauglehole 77472 (MEL, NSW); near Bellbrae (Jan Juc) 11 km N of Anglesea, L.A.S.Johnson NSW 77843 (NSW); Three Jacks Wildflower Sanctuary, northern edge of Stawell, K.L.Wilson 6709, 6710 & L.A.S.Johnson (MEL, NSW); 9.5 km W of Cherrypool turn-off on Black Ra. Rd, K.L.Wilson 6756–6758 & L.A.S.Johnson (MEL, NSW). Occurs as isolated populations, showing some variation; in particular, some specimens from eastern Victoria have coarser branchlets (though still short-toothed). Allocasuarina misera is generally more slender and has more erect teeth than the related species A. mackliniana and A. paradoxa. It further differs from A. mackliniana in its slender male spikes. It differs from A. pusilla in lacking a waxy bloom and median groove on the phyllichnia and in often having longer teeth (0.3–0.8 mm long), which are brown rather than pale yellow.

48. (Macklin) L.Johnson, J. Adelaide Bot. Gard. 6: 77 (1982) Casuarina pusilla Macklin, Trans. & Proc. Roy. Soc. S. Australia 51: 272, figs 22–26, t. 13 (1927). T: Encounter Bay, S.A., 1 July 1927, E.Macklin 020; lecto: AD, with cones, fide L.A.S.Johnson, J. Adelaide |Bot. Gard. 6: 77 (1982); isolecto: BRI, K, MEL, NSW. Illustration: L.Costermans, Native Trees & Shrubs S.E. Australia 148 (1981).

163

Figure 58. Allocasuarina cones ×1.5. A–B, A. mackliniana subsp. mackliniana. A, cone; B, branchlet ×10 (A–B, NSW 146700, NSW). C, A. mackliniana subsp. hirtilinea, branchlet ×10 (C, NSW 146702, NSW). D–E, A. mackliniana subsp. xerophila. D, cone; E, branchlet ×10 (D–E, NSW 76100, NSW). F, A. misera, branchlet ×15 (NSW 62576, NSW). G–H, A. pusilla. G, cone; H, branchlet ×15 (G–H, A.Beauglehole 49746, NSW). I–J, A. robusta. I, cone; J, branchlet ×12 (I–J, AD 95812020, NSW). K–L, A. striata. K, cone; L, branchlet ×15 (K–L, NSW 62602, NSW). Drawn by D.Mackay.

164 CASUARINACEAE 3. Allocasuarina

Dioecious spreading shrub 0.2–2 m high. Bark smooth. Branchlets ascending or spreading, to 12 cm long; articles 3–9 mm long, 0.4–1 mm diam., smooth, glabrous, usually with a waxy bloom; phyllichnia flat or slightly rounded, with faint median groove or line; teeth 5–7, erect, overlapping, 0.3–0.5 mm long, only marcescent at apex. Male spikes 0.3–2 cm long, 8–11 whorls per cm; bracteoles persistent; anther 0.6–1 mm long. Cones subglobose or shortly cylindrical, occasionally as broad as long, sessile; cone body 10–15 mm long, 8–11 (rarely to 13) mm diam.; bracteoles broadly acute to obtuse, pyramidal protuberance shorter than bracteole body. Samara 4.5–5.0 mm long, very dark brown to black. 2n = 22, 33, 44, 55, B.A.Barlow, Austral. J. Bot. 7: 240 (1959). Dwarf Sheoak. Fig. 58G–H. Occurs from Yorke Peninsula, S.A., E to Big and Little Deserts, Vic.; grows in heath in sandy soil. Regions: EYRE, RIVR, OTWY. Map 169. S.A.: reserve at Chaunceys Line, S of Monarto, Hj.Eichler 15581, 15582 (AD, NSW); c. 32 km S of Bordertown on Naracoorte road, L.A.S.Johnson 7926, 7927 & K.L.Wilson (AD, NSW); Dark Island, 16 km NE of Keith, R.Melville 458 (K, MEL, NSW). Vic.: c. 18 km direct SE of Cowangie, Big Desert, M.G.Corrick 6735 & P.Short (MEL, NSW); c. 5 km S of Winiam on road to Goroke, D.J.McGillivray 3246 & C.Bartlett (K, MEL, NSW). Differs from A. mackliniana and A. paradoxa in its slender glabrous articles usually with a waxy bloom and flat to rounded phyllichnia (with median line or groove) and short, erect teeth, and its slender male spikes.

49. Allocasuarina robusta (Macklin) L.Johnson, J. Adelaide Bot. Gard. 6: 78 (1982) Casuarina paludosa var. robusta Macklin, Trans. & Proc. Roy. Soc. S. Australia 51: 271, figs 20, 21, t. 13 (1927). T: upper Hindmarsh Valley, S.A., 1 July 1927, E.Macklin 014; lecto: AD, with cones, fide L.A.S.Johnson, J. Adelaide Bot. Gard. 6: 78 (1982); isolecto: BRI, K, MEL, NSW. Monoecious or more rarely dioecious shrub 0.2–3 m high. Bark smooth. Branchlets ascending, to 20 cm long; articles 7–14 mm long, 0.7–1.1 mm diam., smooth, glabrous; phyllichnia rounded, occasionally only slightly so; teeth 5–7, suberect to spreading, occasionally slightly overlapping when young, often marcescent at apex, 0.6–1 mm long. Male spikes rarely moniliform, 0.5–4.5 cm long, 5–9 whorls per cm; bracteoles persistent; anther 0.8–1.2 mm long. Cones cylindrical, often irregular, sparsely pubescent, sessile or on peduncle to 3 mm long; cone body 12–20 mm long, 7–12 mm diam.; bracteoles broadly acute to obtuse, pyramidal protuberance shorter than bracteole body, acute or obtuse with mucro. Samara 5.5–6.0 mm long, black. Fig. 58 I–J. Occurs in the southern Lofty Ranges, S.A.; grows in upland heath and open woodland with heathy understorey. Region: EYRE. Map 170. S.A.: Myponga, Jan. 1929, J.B.Cleland (AD); Inman Hills, 23 Jan. 1925, J.B.Cleland (AD, NSW 98538); 3 km S of Mt Compass on Victor Harbor road, just S of ‘Square Waterhole’, L.A.S.Johnson 8680 & K.Horsell (NSW). Differs from A. paradoxa in its always glabrous articles with more rounded phyllichnia and less spreading, mostly non-overlapping teeth, and in being usually monoecious. Differs from A. striata in its often shorter articles and usually shorter and more slender teeth, its persistent male bracteoles, and its shorter cone peduncles. It is uncommon and could be considered endangered.

50. (Macklin) L.Johnson, J. Adelaide Bot. Gard. 6: 78 (1982) Casuarina striata Macklin, Trans. & Proc. Roy. Soc. S. Australia 51: 267, figs 14–16, t. 13 (1927). T: Belair, S.A., 18 Aug. 1927, E.Macklin 036; lecto: A♀D, ( with inflorescences and cones), fide L.A.S.Johnson, J. Adelaide Bot. Gard. 6: 78 (1982); isolecto: BRI, K, MEL, NSW. Illustration: L.Costermans, Native Trees & Shrubs S.E. Australia 148 (1981). Dioecious or monoecious shrub or small tree 1–4 m high. Bark smooth or somewhat fissured in older plants. Branchlets ascending, to 10 cm long; articles 8–20 mm long, 0.8–1.4 mm diam., smooth, usually glabrous, often with a waxy bloom; phyllichnia rounded occasionally only slightly so; teeth 5–7, erect, overlapping, 0.6–1.8 mm long, often apex

165 3. Allocasuarina CASUARINACEAE marcescent. Male spikes rarely moniliform, 1–3 cm long, 5–7 whorls per cm; bracteoles deciduous; anther 0.7–1.3 mm long. Cones cylindrical; peduncle 3–12 mm long; cone body 16–33 mm long, 10–13 mm diam.; bracteoles obtuse, pyramidal protuberance shorter than bracteole body, rarely superficially divided into 2 or 3 parts. Samara 6–7 mm long, very dark brown to black. 2n = 22, 33, 55, B.A.Barlow, Austral. J. Bot. 7: 241 (1959). Small Bull Oak. Fig. 58K–L. Occurs in the Southern Lofty Ranges and on Kangaroo Is., S.A.; grows in heath in lateritic or sandy soils. Region: EYRE. Map 171. S.A.: 16 km from Victor Harbor, E.J.Carroll CBG 23786 (CBG, NSW); Clarendon, 26 Feb. 1927, J.B.Cleland (AD, NSW); c. 13 km E of Kelly Hill, Kangaroo Is., Hj.Eichler 15419, 15420 (AD, NSW); Prospect Hill, c. 5 km SW of Meadows, L.A.S.Johnson NSW 42273 (NSW); Fleurieu Peninsula, Second Valley Forest area, D.E.Symon 13102 (AD, B, L, MO, NSW, US). Of variable habit, which may be correlated with ploidy level.

51. Allocasuarina gymnanthera L.Johnson, Fl. Australia 3: 196 (1989) T: Lees Pinch, Wollar–Merriwa road, N.S.W., 19 Sept. 1951, L.A.S.Johnson NSW 62526; holo: NSW, with cones; iso: CANB, K. Dioecious or rarely monoecious shrub or tree, 2–5 m high. Bark smooth or ±fissured. Branchlets ascending, to 20 cm long; articles 5–12 mm long, 0.5–1.0 mm diam., smooth, mostly with pubescent furrows; phyllichnia slightly rounded, often sparsely and minutely pubescent; teeth 6–8, rarely 9, erect, slightly overlapping at bases at least when young, 0.4–0.7 mm long, not marcescent. Male spikes moniliform, 2.5–6 cm long, 3–5 whorls per cm; bracteoles deciduous; anther 0.8–1.4 mm long. Cones usually long-cylindrical, often with sterile apex to 3 mm long, often sparsely pubescent; peduncle 3–8 mm long; cone body 14–40 mm long, 9–12 mm diam.; bracteoles obtuse or truncate, pyramidal protuberance slightly shorter than bracteole body, obtuse. Samara 4.5–7 mm long, very dark brown to black. 2n = 22, 44, B.A.Barlow, Austral. J. Bot. 7: 240 (1959), as Casuarina sp. 3. Fig. 59A–B. Occurs from the southern Pilliga S to the lower Goulburn R. Valley (Ulan and Baerami) and to Glen Davis, N.S.W.; grows in low open woodland in sandy soil on sandstone ridges. Regions: MCPH, DARL, NEPN. Map 172. N.S.W.: Coxs Gap, E.F.Constable NSW 16283 (A, AD, BRI, CANB, CBG, K, L, LE, MEL, MO, PE, PERTH, PRE, RSA); 1.5 km SW of Glen Davis, Green Gully, M.D.Crisp 2208 & I.R.Telford (CBG, NSW); Goonoo State Forest, c. 37 km by road NE of Dubbo on Mendooran road, L.A.S.Johnson 8344, 8345 (K, NSW); Warrumbungle Natl Park, E end of Dows High Tops, 13 May 1980, J.H.Willis (MEL, NSW). Differs from most species in section Cylindropitys in its long, strongly moniliform male spikes with deciduous bracteoles. Differs from A. striata in its shorter and more slender articles with usually pubescent furrows and 6–9 shortish teeth. Its geographical range overlaps that of A. diminuta.

52. Allocasuarina paludosa (Sieber ex Sprengel) L.Johnson, J. Adelaide Bot. Gard. 6: 77 (1982) Casuarina paludosa Sieber ex Sprengel, Syst. Veg. 3: 803 (1826); C. distyla var. paludosa (Sieber ex Sprengel) Benth., Fl. Austral. 6: 199 (1873). T: New South Wales, F.Sieber 329; holo: B♂, ( ♀); iso: BM, BR, C, FI, K, L, LE, MEL, NSW, PR, S. Casuarina pumila Otto & A.Dietr., Allg. Gartenzeitung 9: 162 (1841). T: cult. Berlin bot. gard., collector unknown; holo: B (♂) n.v.; iso: S. Casuarina pumila var. hirtella F.Muell. ex Miq., Ned. Kruidk. Arch. 4: 100 (1856). T: Alberts R., Gippsland, Vic., June 1853, F.Mueller; holo: U (♂); iso: MEL. Casuarina distyla var. prostrata Maiden & E.Betche, Proc. Linn. Soc. New South Wales 30: 371 (1905). T: Narrabeen to Barrenjoey, N.S.W., July 1905, R.H.Cambage & J.H.Maiden NSW 62648; lecto: NSW,

166 Figure 59. Allocasuarina cones ×1.5. A–B, A. gymnanthera. A, cone; B, branchlet ×15 (A–B, NSW 62485, NSW). C–D, A. paludosa. C, cone; D, branchlet ×7.5 (C–D, R.Coveny 5764, NSW). E–F, A. brachystachya. E, cone; F, branchlet ×15 (E–F, NSW 65602, NSW). G–H, A. muelleriana subsp. muelleriana. G, cone; H, branchlet ×15 (G–H, L.Johnson 7928, NSW). I, A. muelleriana subsp. notocolpica, cone (E.Jackson 4357, NSW). J, A. muelleriana subsp. alticola, branchlet ×10 (NSW 91540, NSW). Drawn by D.Mackay.

167 3. Allocasuarina CASUARINACEAE with cones, fide L.A.S.Johnson, Fl. Australia 3: 200 (1989); isolecto: BM, MEL. Illustrations: L.Costermans, Native Trees & Shrubs S.E. Australia 148–149 (1981). Monoecious or dioecious shrub, spreading, 0.3–3 m high. Bark smooth. Branchlets ascending or curved, to 20 cm long; articles 5–14 mm long, 0.7–1 mm diam., smooth, usually densely pubescent in furrows; phyllichnia flat or slightly rounded, usually with median groove, occasionally pubescent (not in groove); teeth 6–8, erect or spreading, not overlapping, 0.5–0.9 mm long, often marcescent. Male spikes 1–2.5 cm long, 7–9 whorls per cm; bracteoles persistent; anther 0.7–1.1 mm long. Cones cylindrical to ovoid, sessile or on peduncle to 2 mm long; cone body 10–18 mm long, 7–13 mm diam.; bracteoles truncate to obtuse, pyramidal protuberance shorter than bracteole body, truncate, often with minute deciduous mucro. Samara 3.5–5 mm long, dark brown to black. 2n = 22, 44, B.A.Barlow, Austral. J. Bot. 7: 240 (1959). Figs 45D, I, 59C–D. Occurs from Broken Bay, N.S.W., S along coast and adjoining tablelands to north-eastern Tas. (including Flinders Is.) and W to Lucindale and Mt Burr, S.A.; grows in heath and in poorly drained soils above swamps at edge of woodland. Regions: OTWY, HOWE, NEPN. Map 173. S.A.: 16 km S of Penola–Clay Wells road on Mt Burr road, K.L.Wilson 1216 & L.A.S.Johnson (AD, NSW). N.S.W.: 18.5 km NW of Mittagong on Wombeyan Caves road, R.G.Coveny 6065, P.Hind & R.Hancock (A, K, L, NSW); Mt Budawang, L.A.Craven 652, 653 (CANB, CHR, K, MEL, NSW, US). Vic.: Glenelg R., 35 km N of Dunkeld, The Grampians, H.Streimann 2767 (A, BRI, CBG, L, NSW, US). Tas.: Waterhouse, H.N.Barber NSW 62718 (K, NSW). Individuals in a population may appear green or grey depending on hairiness of articles. Variable in size.

53. Allocasuarina brachystachya L.Johnson, Fl. Australia 3: 193 (1989) T: 19.5 km SE of Tingha on Tenterden–Moredun Dams road, N.S.W., 9 Oct. 1980, K.L.Wilson 3191; holo: NSW, with cones; iso: BRI, CANB, K, MO. Similar to A. paludosa. Usually monoecious shrub to 3 m high. Branchlets ascending, to 7 cm long; articles 2–5 mm long, 0.4–0.7 mm diam., usually minutely pubescent in furrows; phyllichnia often with median groove near base of article, occasionally along whole length; teeth 5–7, erect, 0.2–0.5 mm long. Male spikes occasionally moniliform, 5–17 mm long, 9–16 whorls per cm; anther 0.3–0.6 mm long. Cones on peduncle 2–4 mm long; cone body 7–14 mm long, 5–8 mm diam. Samara 2.5–4 mm long. 2n = 44, B.G.Briggs pers. comm. (NSW 65602, 65603). Fig. 59E–F. Occurs on the W edge of the New England Tableland from Emmaville S to Guyra and Moredun areas, N.S.W.; grows in low open woodland in low-nutrient sandy soil. Region: MCPH. Map 174. N.S.W.: Limestone Ck, Guyra district, W.F.Blakely, E.N.McKie & T.Youman NSW 62762 (NSW); Emmaville, J.L.Boorman NSW 62759 (NSW); Taabinga, 37 km NW of Guyra, E.N.McKie 9 (BRI). Related to A. paludosa but generally smaller in its parts.

54. Allocasuarina muelleriana (Miq.) L.Johnson, J. Adelaide Bot. Gard. 6: 77 (1982) Casuarina muelleriana Miq., Ned. Kruidk. Arch. 4: 99 (1856); C. suberosa var. muelleriana (Miq.) Miq., Flora 48: 21 (1865). T: Mt Torrens, Lofty Ra., S.A., Aug. 1850, F.Mueller; holo: U, with cone; iso: MEL. Illustrations: E.Macklin, Trans. & Proc. Roy. Soc. S. Australia 51: figs 30–33, t. 13 (1927); L.Costermans, Native Trees & Shrubs S.E. Australia 149 (1981). Dioecious or rarely monoecious shrub 0.5–3 m high. Bark smooth. Branchlets ascending, to 12 cm long; articles 3–11 mm long, 0.6–1.1 mm diam., angular to terete, often somewhat verruculose, often waxy, often pubescent in furrows; phyllichnia strongly angular with white central rib (often minutely serrate); teeth 5–8, erect, not overlapping, 0.3–0.6 mm long, not marcescent. Male spikes often moniliform, 1–4 cm long, 6–8 whorls per cm; bracteoles

168 CASUARINACEAE 3. Allocasuarina persistent; anther 0.5–1.0 mm long. Cones cylindrical, often with short sterile apex to 5 mm long, sessile or on peduncle to 16 mm long; cone body 14–30 (rarely to 35) mm long, 9–18 mm diam.; bracteoles not protruding far from cone body, obtuse to truncate, thick pyramidal protuberance shorter than bracteole body. Samara 6–9 (rarely 5) mm long, black. Slaty Sheoak. Occurs from Ceduna and the Flinders Ranges, S.A., E to Bendigo, Vic., also on Kangaroo Is., S.A. Grows in scrub and heath, in rocky siliceous soils. There are 3 subspecies. 1 Articles 0.6–0.8 mm diam.; phyllichnium margins (when dry) often raised along furrows; cone body usually 9–12 mm diam.; peduncle usually 1–8 mm long 54a. subsp. muelleriana 1: Articles 0.7–1.1 mm diam.; phyllichnium margins flat; cone body 12–18 mm diam. 2 Cones on stout peduncle 8–17 mm long, 3–4.5 mm diam. 54b. subsp. notocolpica 2: Cones sessile or on peduncle to 3 mm long, 2.5 mm diam. 54c. subsp. alticola 54a. Allocasuarina muelleriana (Miq.) L.Johnson subsp. muelleriana Shrub 0.5–3 m high. Articles 3–8 mm long, 0.6–0.8 mm diam.; phyllichnium margins adjacent to furrows often raised (when dry); teeth 5 or 6, rarely 7, 0.3–0.5 mm long. Cones on peduncle 1–8 (rarely to 16) mm long; cone body 14–28 (rarely to 35) mm long, 9–12 (rarely to 15) mm diam. 2n = 22, B.A.Barlow, Austral. J. Bot. 7: 240 (1959). Fig. 59G–H. Occurs throughout range of species, except the Northern Flinders Ranges and Kangaroo Is. Regions: EYRE, RIVR, OTWY. Map 175. S.A.: Fairview Conservation Park, E.N.S.Jackson 4580 (AD, NSW); Second Valley Forest, R.Melville 3958, 3959 (K, NSW); c. 22 km SSE of Kimba, P.G.Wilson 202, 227 (AD, NSW). Vic.: Cave of Hands, Billwing area, W side of Victoria Ra., A.C.Beauglehole 4640, M.McGarvie & P.Finck (MEL, NSW); SW of Jacksons Flat Reservoir, 1.5 km N of Bendigo, R.Melville 1375, 1376 (K, MEL, NSW). 54b. Allocasuarina muelleriana subsp. notocolpica L.Johnson, Fl. Australia 3: 198 (1989) T: near ‘Meadow Lea’ homestead, Kangaroo Is., S.A., 20 Aug. 1982, E.N.S.Jackson 4357; holo: NSW, with cones; iso: AD. Shrub 2–3 m high. Articles 5–11 mm long, 0.9–1.1 mm diam.; phyllichnium margins not raised; teeth 6 or 7, rarely 8, 0.4–0.6 mm long. Cones on stout peduncle 8–17 mm long, 3–4.5 mm diam.; cone body 18–30 mm long, 12–18 mm diam. Fig. 59 I. Restricted to Kangaroo Is., S.A. Region: EYRE. Map 176. S.A.: Hog Bay, Clarke AD 97242307 (AD); Lake Ada, just W of Murray Lagoon, 2 Mar. 1926, J.B.Cleland (AD); Cape Borda, M.E.Phillips CBG 6644 (AD, CBG); 0.8 km E of Karatta, Kangaroo Is., M.E.Phillips CBG 23784 (AD, CBG, NSW). In eastern and northern parts of the island there may be some gradation towards subsp. muelleriana. 54c. Allocasuarina muelleriana subsp. alticola L.Johnson, Fl. Australia 3: 197 (1989) T: upper slopes of Mt McKinley, S.A., 4 Mar. 1966, D.E.Symon 4016; holo: NSW, with cones; iso: AD, K. Shrub 1.5–3 m high. Articles 5–7 mm long, 0.7–1 mm diam.; phyllichnium margins not raised; teeth 7 or 8, rarely 6, 0.3–0.5 mm long. Cones sessile or on short peduncle to 3 mm long; cone body 18–30 mm long, 12–17 mm diam. 2n = 22, B.A.Barlow, Austral. J. Bot. 7: 240 (1959). Fig. 59J.

169 3. Allocasuarina CASUARINACEAE

Occurs in the northern Flinders Ranges from Freeling Heights S to Wilpena Pound and Bibliando, S.A. Region: TRNS. Map 177. S.A.: Bibliando Stn, M.D.Crisp 754, 755 (AD); Wilpena Pound, D.E.Symon 1424, 1426 (AD, NSW); Wilpena Pound, E.N.S.Jackson 79 (AD, NSW); Mt Serle, E of Copley, D.E.Symon 3996, 3997 (AD, CANB, K, NSW).

Sect. 12. Nannopitys

Allocasuarina sect. Nannopitys L.Johnson, Fl. Australia 3: 192 (1989). Type: A. nana (Sieber ex Sprengel) L.Johnson Dioecious or rarely monoecious shrubs; penultimate branchlets woody. Bark smooth. Branchlets often with a waxy bloom; elongate articles numerous, terete, smooth; teeth not overlapping or marcescent. Male spikes shortly elongate; bracteoles persistent. Cone bracts inconspicuous; bracteoles with protuberance 3–5-parted. Samaras glabrous, red-brown to black. A section of 1 species in south-eastern Australia.

55. (Sieber ex Sprengel) L.Johnson, J. Adelaide Bot. Gard. 6: 77 (1982) Casuarina nana Sieber ex Sprengel, Syst. Veg. 3: 804 (1826). T: New South Wales, F.Sieber 328; holo: B (♂); iso: BM, BR, C, FI, K, L, LE, MEL, P, PR, S. Spreading shrub, 0.2–2 m high. Branchlets ascending, to 8 cm long; articles 5–6 mm long, 0.5–0.8 mm diam., with pubescent furrows; phyllichnia slightly rounded; teeth 5 or 6, rarely 4, erect, 0.3–0.6 mm long. Male spikes dense, 5–10 mm long, 16–20 whorls per cm; anther 0.5–0.6 mm long. Cones cylindrical to barrel-shaped, sessile or on peduncle to 3 mm long; cone body 14–24 mm long, 10–15 mm diam.; bracteoles thick, body and protuberance not distinguishable, acute to broadly acute, divided into 2 larger and 3 (rarely to 5) smaller bodies. Samara 4–6 mm long. 2n = 22, 33, 44, B.A.Barlow, Austral. J. Bot. 6: 205–207, 217 (1958). Fig. 60A–B. Occurs from Cowan and Glen Davis, N.S.W., S to the upper Genoa R., Vic.; grows on the coast and tablelands in heath on sandstone, in exposed situations such as ridges on the ranges of the eastern highlands. Regions: NEPN, OTWY. Map 178. N.S.W.: between Berowra and Cowan, W.F.Blakely & D.W.C.Shiress NSW 62793 (MEL, MO, NSW); Crafts Wall, Kanangra, E.F.Constable 5851 (NSW); 10 km N of Clarence, R.G.Coveny 3591 (CHR, K, NSW); between Cooma and Kydra, R.Pullen 3982, 3983 (CANB, NSW). Vic.: Yambulla Ck (upper Genoa R.), N.A.Wakefield 2847 (NSW). This is the only eastern Australian species with cones that are of smoothish outline with a tessellated surface because of the thick, divided bracteoles and protuberances. There are considerable disjunctions in its distribution.

Sect. 13. Trachypitys

Allocasuarina sect. Trachypitys (Benth.) L.Johnson, Fl. Australia 3: 193 (1989). Casuarina sect. Trachypitys Benth., Fl. Austral. 6: 194 (1873). T: A. humilis (Otto & A.Dietr.) L.Johnson Dioecious or more rarely monoecious shrubs; penultimate branchlets green or woody. Bark smooth or finely striate. Branchlets occasionally with a waxy bloom; elongate articles numerous, terete or quadrangular, smooth or minutely serrate on the angles; teeth occasionally overlapping, occasionally marcescent. Male spikes very short; bracteoles persistent. Cone bracts inconspicuous; bracteoles and protuberance indistinguishable, 6–9-parted. Samaras glabrous or hairy, black.

170

Figure 60. Allocasuarina cones ×1.5. A–B, A. nana. A, cone; B, branchlet ×15 (A–B, NSW 19376, NSW). C–D, A. humilis. C, cone; D, branchlet ×15 (C–D, NSW 62790, NSW). E–F, A. drummondiana. E, cone; F, branching habit ×7.5 (E–F, T. & J.Whaite 4125, NSW). G–H, A. microstachya. G, cone; H, branching habit ×7.5 (G–H, NSW 95258, NSW). I–J, A. thuyoides. I, cone; J, branchlet ×7.5 (I–J, NSW 62800, NSW). Drawn by D.Mackay.

171 3. Allocasuarina CASUARINACEAE

A section of 3 species in south-western Australia.

56. (Otto & A.Dietr.) L.Johnson, J. Adelaide Bot. Gard. 6: 76 (1982) Casuarina humilis Otto & A.Dietr., Allg. Gartenzeitung 9: 163 (1841). T: cult. Berlin bot. gard., collector unknown; holo: B (♂, ♀); iso: MEL, NSW. Casuarina preissiana Miq. in J.G.C.Lehmann, Pl. Preiss. 1: 640 (1845). T: Mt Eliza [Kings Park], W.A., 19 May 1839, L.Preiss 2008 p.p.; lecto: U♂ (), fide L.A.S.Johnson, Fl. Australia 3: 202 (1989); isolecto: ?BR, C, K, L, LE, MEL, P, ?S, UPS, W. Casuarina selaginoides Miq. in J.G.C.Lehmann, Pl. Preiss. 1: 643 (1845). T: Bald Head [King George Sound], W.A., 16 Oct. 1840, L.Preiss 2003; holo: U (♂); iso: LE. Casuarina tephrosperma Miq., Revis. Crit. Casuarinarum 31 (1848). T: cult. Hamburg, from Australia, collector unknown; holo: U, sterile. Casuarina humilis var. macrocarpa Miq. in A.P. de Candolle, Prodr. 16(2): 340 (1868). T: Swan R., W.A., J.Drummond 238 in herb. Boissier; holo: G n.v.; iso: B, CGE, FI (with cones), K (with cones), LE, P (with cones), W. Dioecious or monoecious shrub, erect to spreading, 0.2–2 m high; penultimate branchlets woody. Branchlets ascending, to 12 cm long; articles terete, 3–6 mm long, 0.8–1.2 mm diam., smooth, sometimes waxy, glabrous, or finely pubescent in furrows; phyllichnia angular to markedly rounded, often with minutely serrate median ridge; teeth 5–7, erect, overlapping at bases at least when young, 0.4–0.5 mm long, not marcescent. Male spikes 6–18 mm long, 12–16 whorls per cm; anther 0.7–0.8 mm long. Cones cylindrical, slightly longer than broad, of smoothish outline and tessellated surface, sessile; cone body 12–22 (rarely to 33) mm long, 10–17 mm diam.; bracteoles thick, flat-topped and triangular, divided into 6–8 obtuse or ± acute bodies. Samara 5–6 mm long, glabrous; wing truncate, shorter than body. 2n = 20, B.A.Barlow, Austral. J. Bot. 7: 232 (1959). Figs 19, 60C–D. Widespread from the Murchison R. south to the S coast and E to Israelite Bay, W.A.; grows in heath in sand. Regions: BENC, LUWN, ESPR. Map 179. W.A.: 38 km NW of Kalbarri turn-off from North West Coastal Hwy, A.C.Beauglehole 11966 (NSW); N of Frenchmans Cap, near Cape le Grand, B.G.Briggs 407 (NSW, PERTH); 6 km W of Tammin on Great Eastern Hwy, L.Haegi 1785, 1786 (NSW, PERTH); c. 1.8 km W of Mt Hopkins, Walpole–Nornalup Natl Park, L.Haegi 1809, 1810 & J.Powell (NSW, PERTH); Hassell Beach area, L.Haegi 1854, 1855 (NSW, PERTH). Exhibits considerable morphological variation.

57. Allocasuarina drummondiana (Miq.) L.Johnson, J. Adelaide Bot. Gard. 6: 75 (1982) Casuarina drummondiana Miq., Revis. Crit. Casuarinarum 26, t. 1 (1848). T: Swan R., W.A., J.Drummond in Herb. Hooker; holo: K♂) ( ; iso: U. Illustrations: L.Diels in A.Engler, Bot. Jahrb. Syst. 35: figs 11C, D (1905). Dioecious intricate shrub 0.5–3 m high; penultimate branchlets green. Branchlets to 2 cm long; articles terete or angular, 1.5–2.5 mm long, 0.9–1.2 mm diam., smooth, pubescent in furrows; phyllichnia angular to rounded, with central ridge; teeth 6 or 7, not overlapping, erect or apex slightly spreading, apical portion very thin-textured and marcescent leaving only rounded base, 0.5–0.9 mm long (basal region c. 0.2–0.3 mm long). Male spikes 4–10 mm long, c. 20 whorls per cm; anther c. 0.7 mm long. Cones ovoid to cylindrical, sessile on woody branches; cone body 8–15 mm long, 7–8 mm diam.; bracteoles thick, flat-topped and triangular, divided into 5–8 convex bodies. Samara 3.0–4.0 mm long, sparsely hairy. 2n = 20, B.A.Barlow (no voucher). Figs 23, 45E, 60E–F. Restricted to Three Springs–Wongan Hills area, W.A.; grows in tall heath on lateritic ridges and on sandplain. Region: BENC. Map 180. W.A.: 4.2 km NW of Wongan Hills towards Piawaning, R.G.Coveny 7790 & B.R.Maslin (K, NSW, PERTH); c. 5 km S of Waddington, L.A.S.Johnson NSW 53232, 53233 (NSW); 11 km NW of Carnamah on Geraldton Hwy, R.Melville 4128, 4129 (K, NSW).

172 CASUARINACEAE 3. Allocasuarina

The samara of this species has a very short wing and sparse short (1–2 mm long) white hairs on the sides of the body.

58. Allocasuarina microstachya (Miq.) L.Johnson, J. Adelaide Bot. Gard. 6: 76 (1982) Casuarina microstachya Miq. in J.G.C.Lehmann, Pl. Preiss. 1: 642 (1845). T: near Maddington, Perth, W.A., 2 Nov. 1839, L.Preiss 1999; lecto: U♂ (), fide L.A.S.Johnson, J. Adelaide Bot. Gard. 6: 76 (1982); isolecto: B, BR, L, MEL, P, PR, S. Illustration: L.Diels in A.Engler, Bot. Jahrb. Syst. 35: fig. 11E (1905). Dioecious intricate shrub 0.1–1 m high; penultimate branchlets green. Branchlets to 5 cm long; articles quadrangular, 2–6 mm long, 0.6–0.8 mm diam., smooth on faces, verruculose on angles with furrows often becoming open and obvious; phyllichnia angular, very finely verruculose to serrate on angle; teeth 4, not overlapping, erect but apex eventually spreading and marcescent, leaving only rounded base, 0.4–1.3 mm long (dark apex c. 0.3–1 mm long). Male spikes 1.5–3 mm long; anther 0.5–0.6 mm long. Cones shortly cylindrical to ovoid, irregular, sessile; cone body 8–12 mm long, 5–10 mm diam.; bracteoles thick, divided into 6–9 tubercles, rounded or flat-topped with deciduous mucro 0.5–3 mm long. Samara 2.5–5 mm long, hairy. 2n = 20, B.A.Barlow, Austral. J. Bot. 7: 232 (1959). Figs 45F, 60G–H. Occurs from Strawberry, near Geraldton, SE to Albany and the Munglinup R., E of Ravensthorpe, W.A.; grows in heath in gravelly sand over laterite. Regions: BENC, LUWN, ESPR. Map 181. W.A.: Pithara–Miling, W.E.Blackall 2882 (PERTH); 11 km N of Bullaring, B.G.Briggs 190 (MO, NSW, PERTH); 1.5 km W of Munglinup R., Ravensthorpe to Esperance, B.G.Briggs 434, 435 (NSW); near 359 mile peg [c. 574 km E of Perth], W of Coolgardie, Great Eastern Hwy, B.G.Briggs & E.M.Scrymgeour 748 (NSW, PERTH); E of Strawberry, L.A.S.Johnson NSW 58485 (NSW, PERTH). Variable in size characters. The samara is most striking, being wingless and having a mass of spreading, long, septate, usually ferruginous hairs c. 3–5 mm long.

Sect. 14. Acanthopitys

Allocasuarina sect. Acanthopitys (Miq.) L.Johnson, Fl. Australia 3: 191 (1989). Casuarina sect. Acanthopitys Miq. in J.G.C.Lehmann, Pl. Preiss. 1: 641 (1845). T: A. thuyoides (Miq.) L.Johnson Monoecious or dioecious shrubs; penultimate branchlets woody. Bark smooth. Branchlets not or slightly waxy; elongate articles numerous, terete, smooth; teeth not overlapping or marcescent. Male spikes very short; bracteoles persistent. Cone bracts inconspicuous; bracteoles with protuberance entire, awned. Samaras glabrous, black. A section of 1 species in south-western Australia.

59. (Miq.) L.Johnson, J. Adelaide Bot. Gard. 6: 78 (1982) Casuarina thuyoides Miq. in J.G.C.Lehmann, Pl. Preiss. 1: 641 (1845). T: Quangen, Victoria District, and below Gordon R., Hay District, W.A., 20 Mar. 1840 & 7 Nov. 1840, L.Preiss 2004; lecto: U ♂)( [impossible to separate localities and dates; on same sh♀eet s pecias men], fide L.A.S.Johnson, J. Adelaide Bot. Gard. 6: 78 (1982); isolecto: B, LE, MEL. Illustrations: L.Diels in A.Engler, Bot. Jahrb. Syst. 35: figs 11F–H (1905). Intricate shrub 0.3–2 m high. Branchlets to 3 cm long; articles 1–3 mm long, 0.4–0.6 mm diam., glabrous; phyllichnia slightly rounded or flat, often with faint median groove; teeth 5 or 6, erect, 0.3–0.5 mm long. Male spikes of 1–4 whorls at apex of branchlet, 1–5 mm long; anther 0.4–0.6 mm long. Cones shortly cylindrical to globose, as long as broad, spiny; peduncle slender, 2–7 mm long; cone body 8–20 mm long, 8–15 mm diam.; bracteoles thin, acuminate, occasionally with hooked tip, protuberance diverging low on bracteole body, with pyramidal portion nearly as long as bracteole body and with 2–8 mm long coarse yellowish

173 3. Allocasuarina CASUARINACEAE awn with a deciduous hooked tip. Samara 5.0–6.0 (rarely to 10) mm long. 2n = 44, B.A.Barlow, Austral. J. Bot. 7: 233 (1959.. Fig. 60 I–J. Widespread from the Murchison R. south to near Albany and E to Esperance, W.A.; in heath on lateritic foothills and sandplain. Regions: BENC, LUWN, ESPR. Map 182. W.A.: Lucky Bay, E of Esperance, B.G.Briggs 408 (NSW); 29 km from Geraldton–Carnarvon hwy, towards Kalbarri, E.M.Canning CBG 43465 (CBG, NSW); 1.5 km N of Gibson Soak, L.A.S.Johnson NSW 58471 (NSW); 29 km S of Cranbrook on Albany road, R.Melville 4364 (K, NSW); 18 km E of Newdegate on road to Lake King, M.D.Tindale 218 & B.R.Maslin (CANB, K, L, NSW, PERTH).

Doubtful names

Casuarina excelsa Dehnh. ex Miq., Revis. Crit. Casuarinarum 23 (1848), nom. illeg. non Salisb. (1796). T: cult. Hort. bot. Berolinensi, ex Hort. Neapolitano, probably from Australia; n.v. Insufficiently described.

Casuarina lucida Dehnh., Revista Napolitana 1(3): 175 (1839). T: ex Nov. Holl. cult. Hort. Camald. No specimens of this have been found. The description is inadequate.

Casuarina ramuliflora Otto & A.Dietr., Allg. Gartenzeitung 9: 163 (1841). T: cult. Berlin bot. gard. Probable isotype in U is vegetative and may be abnormal (galled) A. littoralis. The description is also consistent with this.

Casuarina sparsa Tausch, Flora 22: 480 (1839). T: cult. Hort. Exc. Comitis Salm. No specimens of this have been found. The description is inadequate.

174

175

MAPS

Number in brackets refers to the page on which the taxon is described.

176

1. Ostrearia australiana (2) 2. Neostrearia fleckeri (2) 3. Noahdendron nicholasii (4) 4. Ulmus hollandica (5) 5. Aphananthe philippinensis (5) 6. Trema orientalis (8) 7. Trema tomentosa 8. Trema tomentosa 9. Celtis australis (10) var. tomentosa (9) var. viridis (9) 10. Celtis occidentalis (11) 11. Celtis sinensis (11) 12. Celtis philippensis (11) 13. Celtis paniculata (13) 14. Humulus lupulus (14) 15. Cannabis sativa (15)

177

16. Streblus brunonianus (18) 17. Streblus glaber 18. Malaisia scandens var. australianus (18) var. scandens (19) 19. Fatoua pilosa (20) 20. Maclura cochinchinensis (22) 21. Artocarpus glaucus (23) 22. Antiaris toxicaria 23. Ficus superba 24. Ficus virens var. macrophylla (24) var. henneana (32) var. virens (35) 25. Ficus virens 26. Ficus virens 27. Ficus drupacea var. sublanceolata (35) var. dasycarpa (35) var. drupacea (36) 28. Ficus drupacea 29. Ficus benghalensis (36) 30. Ficus benjamina var. glabrata (36) var. benjamina (37)

178

31. Ficus microcarpa 32. Ficus microcarpa 32. Ficus microcarpa var. microcarpa (38) var. hillii (39) var. latifolia (39) 34. Ficus triradiata 35. Ficus triradiata 36. Ficus watkinsiana (40) var. triradiata (40) var. sessilicarpa (40) 37. Ficus macrophylla 38. Ficus baileyana (41) 39. Ficus destruens (42) subsp. macrophylla (41) 40. Ficus rubiginosa (42) 41. Ficus leucotricha 42. Ficus leucotricha var. leucotricha (43) var. megacarpa (43) 43. Ficus leucotricha 44. Ficus obliqua 45. Ficus obliqua var. sessilis (44) var. obliqua (44) var. petiolaris (45)

179

46. Ficus obliqua 47. Ficus subpuberula 48. Ficus platypoda var. puberula (45) var. platypoda (46) 49. Ficus platypoda 50. Ficus platypoda 51. Ficus platypoda var. angustata (46) var. cordata (47) var. lachnocaula (47) 52. Ficus platypoda 53. Ficus pleurocarpa (47) 54. Ficus crassipes (49) var. minor (47) 55. Ficus albipila 56. Ficus racemosa 57. Ficus pumila (53) var. albipila (50) var. racemosa (52) 58. Ficus pantoniana 59. Ficus copiosa (54) 60. Ficus melinocarpa var. pantoniana (54) var. hololampra (55)

180

61. Ficus leptoclada (55) 62. Ficus opposita 63. Ficus opposita var. opposita (56) var. indecora (56) 64. Ficus opposita 65. Ficus scobina (57) 66. Ficus podocarpifolia (57) var. micracantha (57) 67. Ficus fraseri (59) 68. Ficus coronulata (59) 69. Ficus coronata (59) 70. Ficus tinctoria 71. Ficus virgata 72. Ficus mollior subsp. tinctoria (60) var. virgata (60) var. mollior (62) 73. Ficus adenosperma (63) 74. Ficus variegata 75. Ficus nodosa (64) var. variegata (64)

181

76. Ficus congesta 77. Ficus hispida 78. Ficus septica var. congesta (65) var.hispida (65) var. septica (66) 79. Ficus septica 80. Urtica incisa (70) 81. Urtica urens (72) var. cauliflora (66) 82. Laportea interrupta (73) 83. Dendrocnide 84. Dendrocnide moroides (76) corallodesme (75) 85. Dendrocnide cordata (76) 86. Dendrocnide 87. Dendrocnide excelsa (78) photinophylla (76) 88. Elatostema reticulatum (80) 89. Elatostema stipitatum (82) 90. Procris pedunculata (84)

182

91. Boehmeria macrophylla (85) 92. Boehmeria nivea (85) 93. Pouzolzia zeylanica (86) 94. Pouzolzia hirta (88) 95. Pipturus argenteus (88) 96. Nothocnide repanda (89) 97. Parietaria debilis (91) 98. Parietaria judaica (92) 99. Australina pusilla (93) 100. Balanops australiana (95) 101. Alnus cordata (96) 102. Nothofagus moorei (98) 103. Nothofagus 104. Nothofagus gunnii (100) 105. Gymnostoma cunninghamii (98) australianum (103)

183

106. Casuarina equisetifolia 107. Casuarina equisetifolia 108. Casuarina cunninghamiana subsp. equisetifolia (106) subsp. incana (106) subsp. cunninghamiana (107) 109. Casuarina cunninghamiana 110. Casuarina glauca (107) 111. Casuarina obesa (108) subsp. miodon (107) 112. Casuarina cristata (108) 113. Casuarina pauper (110) 114. Allocasuarina decaisneana (127) 115. Allocasuarina acuaria (129) 116. Allocasuarina pinaster (129) 117. Allocasuarina grevilleoides (129) 118. Allocasuarina fibrosa (130) 119. Allocasuarina 120. Allocasuarina ramosissima (130) luehmannii (132)

184

121. Allocasuarina 122. Allocasuarina 123. Allocasuarina acutivalvis corniculata (133) spinosissima (133) subsp. acutivalvis (135) 124. Allocasuarina acutivalvis 125. Allocasuarina 126. Allocasuarina globosa (136) subsp. prinsepiana (136) scleroclada (136) 127. Allocasuarina 128. Allocasuarina 129. Allocasuarina eriochlamys tortiramula (137) campestris (128) subsp. eriochlamys (139) 130. Allocasuarina eriochlamys 131. Allocasuarina 132. Allocasuarina subsp. grossa (139) tessellata (139) dielsiana (141) 133. Allocasuarina helmsii (141) 134. Allocasuarina torulosa 135. Allocasuarina (142) decussata (142)

185

136. Allocasuarina 137. Allocasuarina 138. Allocasuarina fraseriana (143) verticillata (144) huegeliana (146) 139. Allocasuarina 140. Allocasuarina 141. Allocasuarina littoralis (148) trichodon (146) inophloia (147) 142. Allocasuarina media (148) 143. Allocasuarina filidens (150) 144. Allocasuarina emuina (150) 145. Allocasuarina 146. Allocasuarina rigida 147. Allocasuarina rigida thalassoscopica (150) subsp. rigida (151) subsp. exsul (151) 148. Allocasuarina 149. Allocasuarina rupicola (152) 150. Allocasuarina portuensis (152) ophiolitica (152)

186

151. Allocasuarina distyla (154) 152. Allocasuarina simulans (154) 153. Allocasuarina diminuta subsp. diminuta (155) 154. Allocasuarina diminuta 155. Allocasuarina diminuta 156. Allocasuarina subsp. mimica (155) subsp. annectens (157) defungens (157) 157. Allocasuarina 158. Allocasuarina 159. Allocasuarina glareicola (157) grampiana (158) zephyrea (158) 160. Allocasuarina 161. Allocasuarina crassa (159) 162. Allocasuarina lehmanniana monilifera (159) subsp. lehmanniana (161) 163. Allocasuarina lehmanniana 164. Allocasuarina 165. Allocasuarina mackliniana subsp. ecarinata (161) paradoxa (161) subsp. mackliniana (165)

187

166. Allocasuarina mackliniana 167. Allocasuarina mackliniana 168. Allocasuarina misera (163) subsp. hirtilinea (162) subsp. xerophila (163) 169. Allocasuarina pusilla (163) 170. Allocasuarina robusta (165) 171. Allocasuarina striata (165) 172. Allocasuarina 173. Allocasuarina 174. Allocasuarina gymnanthera (166) paludosa (166) brachystachya (168) 175. Allocasuarina muelleriana 176. Allocasuarina muelleriana 177. Allocasuarina muelleriana subsp. muelleriana (169) subsp. notocolpica (169) subsp. alticola (169) 178. Allocasuarina nana (170) 179. Allocasuarina humilis (172) 180. Allocasuarina drummondiana (172)

188

181. Allocasuarina 182. Allocasuarina 183. Parietaria cardiostegia (92) microstachya (173) thuyoides (173)

189

APPENDIX

New taxa, combinations and lectotypifications

New taxa, combinations and lectotypifications occurring in this Volume of the are formally published below. For economy the entries are brief; the treatment of new taxa is more comprehensive in the main text. The date of publication of this Volume will be given in Volume 18.

ULMACEAE

H.J.Hewson

Trema tomentosa var. viridis (Planchon) Hewson, stat. et comb. nov. Sponia viridis Planchon, Ann. Sci. Nat. Bot. ser. 3, 10: 319 (1848); Trema aspera var. viridis (Planchon) Benth., Fl. Austral. 6: 158 (1873). T: Port Essington, N.T., Armstrong 384; n.v. This is the earliest epithet available for the taxon at the rank of variety.

URTICACEAE

W.Greuter

Parietaria cardiostegia Greuter, sp. nov. Ab affini Parietaria debili, quacum adhuc confundebatur, praecipue differt bractearum involucri florum foemineorum forma et venatione: nam in nostra bracteae late cordato-triangulares basi in stipitem brevem crassum contractae prominule radiatim reticulato-nervosae sunt, dum in illa a forma lanceolata ad late ellipticam variant et nervo mediano debili simplici vel aliquanto ramoso percurruntur. T: `Mesa' hills c. 12 km NW of Fowlers Gap, N.S.W., Sept. 1981, W.Greuter 18387; holo: B; iso: NSW. Named from the Greek kardia (heart) and stege (shelter), in reference to the involucral bracts of the female flowers.

CASUARINACEAE

ALLOCASUARINA

E.M.Bennett

Allocasuarina tortiramula E.Bennett Frutex dioica ad 1.75 m alta. Rami tortiles. Ramuli assimilativi ad 10 cm longi, virides sed cinerascentes; folia 7 in verticillo quoque. Inflorescentia mascula terminalis, 12–15 mm longa, 1–4 mm lata. Infructescentia sessilis, ellipsoidalis, 5–9 mm longa, c. 14 mm lata; bracteoli transversaliter fissurati, extus irregulariter obtrullati ad mucronem centralem errigentes, intus velutini. Samara 5.5–6 mm longa; corpus rufo-brunneum albo-notatum, glabrum, costa elevata non centrali; ala hyalina, obtusa, rufo-brunnea, costa ad apicem centrali.

190 APPENDIX Casuarinaceae

T: 16.5 km W of Lake King crossroads, Nature Reserve 39422, W.A., 22 July 1987, M.Graham 1127; holo: PERTH; iso: CANB, K, MEL, NSW. Occurs at 2 localities W of Lake King, south-western W.A. Named from the Latin torti- (twisted) and ramulus (branchlet), in reference to the twisted branchlets.

ALLOCASUARINA

L.A.S.Johnson The sectional names are here all formed on the model of Casuarina sect. Acanthopitys Miq. and the term pitys (a pine) is taken as referring to Casuarinaceae in general, because of the pine-like appearance of most species. The sectional names may thus refer to characters of various parts of the plant, e.g. Dolichopitys refers to the long cones, while Ceropitys is chosen because of the waxy branchlets.

Allocasuarina sect. Acanthopitys (Miq.) L.Johnson, comb. nov. Casuarina sect. Acanthopitys Miq. in J.G.C.Lehmann, Pl. Preiss. 1: 641 (1845). Lecto (here chosen): A. thuyoides (Miq.) L.Johnson

Allocasuarina sect. Amorphopitys L.Johnson, sect. nov. Arbores; ramuli elongati articulis numerosis; folia 6–8 in verticillo unoquoque, laminis (`dentibus') elongato-triangularibus plus minusve patentibus pallidis vel brunnescentibus sed non marcescentibus; phyllichnia prominenter convexa sed non acute angulata, in sicco pallescentia aliquando glaucescentia; flores masculi bracteolis anthesi deciduis; infructescentiae cylindricae sed maturitate nonnunquam aliquanto irregulares c. 14-stichae bracteolis protuberatione aequilonga crassa multituberculata instructis. Chromosomata x = 13. Type: A. fraseriana (Miq.) L.Johnson. The name is from the Greek amorphos (formless) and pitys, referring to the rather irregularly shaped cones.

Allocasuarina sect. Ceropitys L.Johnson, sect. nov. Arbusculae vel frutices; ramuli elongati articulis pluribus vel numerosis; folia 5–13 in verticillo unoquoque, laminis breviter usque ad longissime triangularibus plerumque nigrescentibus marcescentibusque; phyllichnia plana vel convexa sed vix carinata, plus minusve incerata; flores masculi bracteolis anthesi persistentibus; infructescentiae cylindricae vel ellipsoidales, 10–26-stichae, bracteolis crassis, integris vel varie fissis sed partibus (`protuberationibus') aequilongis non elongato-spinescentibus. Chromosomata x = 12, raro 14. Type: A. acutivalvis (F.Muell.) L.Johnson The name is from the Greek keros (wax) and pitys, referring to the waxy coating of the branchlets.

Allocasuarina sect. Cylindropitys L.Johnson, sect. nov. Arbores vel frutices; ramuli plus minusve elongati articulis numerosis; folia plerumque 5–9 in verticillo unoquoque, laminis triangularibus erectis vel patentibus aliquando sed saepissime non marcescentibus; phyllichnia varia (angulata vel convexa vel plana vel concava) semper laevia; flores masculi bracteolis persistentibus vel rarius deciduis; infructescentiae cylindricae, plerumque 10–18-stichae, bracteolis protuberatione dorsali quam bracteola breviori vel aequilonga saepissime integra rarius fissa instructis. Chromosomata x= 11.

191 Casuarinaceae APPENDIX

Type: A. littoralis (Salisb.) L.Johnson The name is from the Greek kylindros (a cylinder) and pitys, referring to the cylindrical shape of the cones.

Allocasuarina sect. Dolichopitys L.Johnson, sect. nov. Arbores; ramuli elongati articulis numerosis; folia 4 in verticillo unoquoque, laminis elongatissime triangularibus acutissimisque primum flavidis; phyllichnia plana (non carinata) vel paullo sulcata; flores masculi bracteolis post anthesin persistentibus; infructescentiae elongatae cylindraceae 8-stichae bracteolis maturis crassis protuberatione dorsali trigono-pyramidali bracteolam aequanti instructis. Chromosomata x = 14. Type: A. decaisneana (F.Muell.) L.Johnson The name is from the Greek dolichos (long) and pitys, referring to the long cones.

Allocasuarina sect. Echinopitys L.Johnson, sect. nov. Arbusculae vel frutices magni; ramuli elongati articulis numerosis; folia 8–11 in verticillo unoquoque, laminis plus minusve elongato-triangularibus mox nigrescentibus marcescentibusque; phyllichnia plana; flores masculi bracteolis anthesi deciduis; infructescentiae globulares usque ad breviter ellipsoidales, c. 7–20-verticillatae, c. 14–20-stichae, bracteolis convexis obtusisque protuberatione dorsali longissime spiniformi instructis. Chromosomata x = 10. Type: A. corniculata (F.Muell.) L.Johnson The name is from the Greek echinos (a hedgehog or a sea-urchin) and pitys, referring to the spiny cones.

Allocasuarina sect. Inopitys L.Johnson, sect. nov. Arbores cortice fibroso; ramuli elongati articulis numerosis; folia 7–9 in verticillo unoquoque, laminis anguste triangularibus erectis pallidis non marcescentibus; phyllichnia subangulata trichomatibus brevibus unicellularibus instructa; flores masculi bracteolis deciduis; infructescentiae cylindricae, plerumque 12–16-stichae, bracteolis protuberatione subaequilonga pyramidali vel in partibus 2 vel 3 pyramidalibus fissa instructis. Chromosomata x = 12. Type: A. inophloia (F.Muell. & Bailey) L.Johnson. The name is from the Greek is, inos (fibre or sinew) and pitys, referring to the division of the thick bark into threads. The sole species is known as ‘Stringybark Oak'.

Allocasuarina sect. Nannopitys L.Johnson, sect. nov. Frutices; ramuli penultimi articulis numerosis; folia 4–6 in verticillo unoquoque, laminis breviter triangularibus, non marcescentibus; phyllichnia leviter convexa; flores masculi bracteolis persistentibus; infructescentiae cylindricae, c. 8–12-stichae, bracteolis maturis crassis fissis 3–5-partibus. Chromosomata x = 11. Type: A. nana (Sieber ex Sprengel) L.Johnson The name is from the Greek nannos (dwarf) and pitys, referring to the often low stature of A. nana.

Allocasuarina sect. Oöpitys L.Johnson, sect. nov. Arbores; ramuli elongati articulis numerosis; folia 8–13 in verticillo unoquoque, laminis elongato-triangularibus praesertim ad basin rufobrunneis tandem marcescentibus sed non nigrescentibus; phyllichnia convexa sed non carinata superficie prominentibus minutis instructa; flores masculi bracteolis anthesi deciduis; infructescentiae plus minusve

192 APPENDIX Casuarinaceae truncato-ellipsoidales, c. 18–24-stichae, bracteolis crassis dorso convexis acutis obtusisve prominentibus haud in protuberationibus divisis. Chromosomata x = 13. Type: A. verticillata (Lam.) L.Johnson The name is derived from the Greek oön (an egg) and pitys, referring to the shape of the cones. It should be pronounced as four syllables with the stress on `ö'.

Allocasuarina sect. Oxypitys L.Johnson, sect. nov. Frutices; ramuli penultimi interdum virides; ramuli breves articulis elongatis non nisi 1–5, saepe pungentes; folia 4 vel 5 in verticillo unoquoque, laminis breviter ad longissime triangularibus, plus minusve marcescentibus; phyllichnia plana vel convexa, interdum sulcata; flores masculi bracteolis persistentibus; infructescentiae globulares usque ad subcylindraceae, c. 10–12-stichae, bracteolis convexis protuberatione dorsali longissime spiniformi vel 4–6-partito instructis. Chromosomata x = 14. Type: A. acuaria (F.Muell.) L.Johnson The name is from the Greek oxys (sharp, pointed) and pitys, referring to the spiny cones and pungent branchlets of some species.

Allocasuarina sect. Platypitys L. Johnson, sect. nov. Arbores; ramuli elongati articulis numerosis; folia 10–14 in verticillo unoquoque, laminis plus minusve elongato-triangularibus mox nigrescentibus et tandem marcescentibus; phyllichnia plana; flores masculi bracteolis anthesi deciduis; infructescentiae perbreves, plerumque 1–3 verticillatae, c. 20-stichae, bracteolis protuberatione dorsali obtusiuscula quam bracteola breviore instructis. Chromosomata x = 14 (species cognita unica tetraploidea). Type: A. luehmannii (R.Baker) L.Johnson The name is from the Greek platys (broad) and pitys, referring to the short, flattened cones.

Allocasuarina sect. Trachypitys (Benth.) L.Johnson, comb. nov. Casuarina sect. Trachypitys Benth., Fl. Austral. 6: 194 (1873). Lecto (here chosen): A. humilis (Otto & Dietr.) L.Johnson

Allocasuarina sect. Trichopitys L.Johnson, sect. nov. Frutices; ramuli elongati articulis numerosis; folia 8–10 in verticillo unoquoque, laminis valde elongatis in acuminem setiformem attenuatis rufobrunneis patentibus tandem marcescentibus sed non nigrescentibus; phyllichnia laeves et prominenter carinata; flores masculi bracteolis persistentibus; infructescentiae cylindricae vel truncato-ellipsoidales, c. 16–20-stichae, bracteolis prominentibus acutis protuberatione convexa aequilonga instructis. Chromosomata x = 10. Type: A. trichodon (Miq.) L.Johnson The name is from the Greek thrix, trichos (hair) and pitys, in reference to the long hair-like ends of the leaves (‘teeth’) and also the bracts of the male and female flowers.

Allocasuarina brachystachya L.Johnson, sp. nov. A. paludosae affinis sed in multibus partibus gracilior est: articuli 2–5 mm longi, 0.4–0.7 mm diam.; laminae 0.2–0.5 mm longae; inflorescentiae masculae 5–17 mm longae; corpus infructescentiae 5–8 mm diam. T: 19.5 km SE of Tingha on Tenterden–Moredun Dams road, N.S.W., 9 Oct. 1980, K.L.Wilson 3191; holo: NSW, with cones; iso: BRI, CANB, K, MO. Occurs on the western edge of the New England Tableland, N.S.W.

193 Casuarinaceae APPENDIX

Named from the Greek brachys (short) and stachys (a spike), referring to the short male inflorescences.

Allocasuarina crassa L.Johnson, sp. nov. Inter species sectionis Cylindropityos combinatione distinguitur: dioica; articuli 10–20 mm longi, 1.2–2 mm diam., sulcis pubescentibus, phyllichniis valde rotundato-convexis; laminae 7–10, 1.1–3 mm longae, maturitate non imbricatae, marcescentes; samara atra. T: Cape Pillar, Tasman Peninsula, Tas., Mar. 1976, T.Moscal NSW 145870; holo: NSW, with cones. Known only from Tasman Is. and Cape Pillar area of Tasman Peninsula, Tas. Named from the Latin crassus (thick, stout), referring to its large articles, which are longer and thicker than in the related species A. monilifera and A. zephyrea.

Allocasuarina defungens L.Johnson, sp. nov. Inter species sectionis Cylindropityos combinatione distinguitur: dioica vel monoica; articuli glaucescentes, sulcis glabratis, phyllichniis parum rotundato-convexis; laminae 0.2–0.5 mm longae, parum imbricatae, ad apicem marcescentes; inflorescentiae masculae moniliformes; corpus infructescentiae 5–7 mm diam. T: Nabiac airstrip, N.S.W., 2 Sept. 1967, R.G.Coveny 73; holo: NSW (♂, ♀ - monoecious). Apparently restricted to Nabiac area, N.S.W.; endangered and perhaps now destroyed by sand-mining. Named from present participle of the Latin defungor (to finish, die), referring to its depauperate habit, general lack of vegetative and reproductive vigour and probable impending .

Allocasuarina diminuta L.Johnson, sp. nov. Inter species sectionis Cylindropityos combinatione distinguitur: dioica vel monoica; articuli saepe leviter cerei, sulcis glabratis, phyllichniis angulatis rotundato-convexisve; laminae plerumque late-triangulares, 0.3–0.8 mm longae, nec plerumque imbricatae nec marcescentes; inflorescentiae masculae raro moniliformes; samara atrorufa. T: c. 3 km E of ‘Wattle Valley’ turn-off on Kangarooby road, N.S.W., 29 Apr. 1982, K.L.Wilson 4451; holo: NSW, with cones; iso: CANB, K. Occurs in N.S.W. in several disjunct regions. Named from the Latin diminutus (made small, diminished), referring to its being smaller in all parts than the allied A. distyla.

Allocasuarina diminuta subsp. annectens L.Johnson, subsp. nov. Inter subspecies diminutam et mimicam morphologia intermedia est: articuli non cerei; laminae late angusteve triangulares, marginibus plus minusve convexis. T: Corang R., Braidwood–Nerriga road, N.S.W., 24 Sept. 1960, R.Pullen 2207; holo: NSW ( ♂, with cones - monoecious); iso: CANB, MEL. Occurs in south-eastern N.S.W. Named from the Latin annectens (binding, connecting), referring to this being morphologically intermediate between the other 2 subspecies.

194 APPENDIX Casuarinaceae

Allocasuarina diminuta subsp. mimica L.Johnson, subsp. nov. A subspecie diminuta laminis late-triangularibus, marginibus convexis; et articulis saepe cereis, differt. T: Bundanoon–Penrose, N.S.W., Apr. 1948, M.D.Tindale NSW 62492; holo: NSW (♂, with cones - monoecious). Occurs near Sydney and in the Central Tablelands of N.S.W. from Blackheath to Taralga and Bundanoon. Named from the Latin mimicus (mimicking), referring to the subspecies' appearance of being a little replica of A. distyla.

Allocasuarina emuina L.Johnson, sp. nov. Inter species sectionis Cylindropityos combinatione distinguitur: dioica; articuli 4–8 mm longi, sulcis pubescentibus, phyllichniis angulatis rotundato-convexisve; laminae 0.3–0.7 mm longae, erectae parum patentesve, nec imbricatae nec marcescentes; inflorescentiae masculae non moniliformes. T: Mt Emu, Qld, 28 Feb. 1981, P.R.Sharpe 2650; holo: NSW, with cones; iso: BRI. Occurs from near Coolum Beach S to Caloundra, Qld. Named from its occurrence on Mt Emu.

Allocasuarina eriochlamys (L.Johnson) L.Johnson, comb. et stat. nov. Casuarina campestris subsp. eriochlamys L.Johnson, Nuytsia 1: 264 (1972); Allocasuarina campestris subsp. eriochlamys (L.Johnson) L.Johnson, J. Adelaide Bot. Gard. 6: 74 (1982). T: Comet Vale, W.A., Aug. 1917, J.T.Jutson 255; holo: NSW (♀).

Allocasuarina eriochlamys subsp. grossa (L.Johnson) L.Johnson, comb. nov. Casuarina campestris subsp. grossa L.Johnson, Nuytsia 1: 264 (1972); Allocasuarina campestris subsp. grossa (L.Johnson) L.Johnson, J. Adelaide Bot. Gard. 6: 74 (1982). T: 5.5 miles [9 km] N of Norseman, W.A., 18 Dec. 1960, L.Johnson W177; holo: NSW, with cones; iso: PERTH.

Allocasuarina filidens L.Johnson, sp. nov. A. littorali affinis sed laminis 5 vel 6, longioribus (1.5–2.0 mm longis); et articulis longioribus, 12–15 mm longis, differt. T: Mt Beerwah, Qld, 13 Aug. 1966, S.T.Blake 22672; holo: NSW, with cones; iso: BRI. Restricted to Glass House Mtns, Qld. Named from the Latin filum (a thread) and dens (a tooth), referring to the very long, slender leaves (‘teeth’).

Allocasuarina glareicola L.Johnson, sp. nov. Inter species sectionis Cylindropityos combinatione distinguitur: dioica vel monoica; articuli virides, sulcis glabratis, phyllichniis parum rotundato-convexis; laminae 0.2–0.5 mm longae, saepissime non imbricatae, erectae parum patentesve, non marcescentes; inflorescentiae masculae moniliformes; corpus infructescentiae 5–7 mm diam.; samara spadicea. T: Castlereagh State Forest, N.S.W., 11 Oct. 1983, D.Benson & D.Keith NSW 155948; holo: NSW (♂, ♀ - monoecious). Restricted to Castlereagh State Forest area, NE of Penrith, N.S.W. Named from the Latin glarea (gravel) and –cola (a dweller), referring to its occurrence in gravelly, lateritic soils.

195 Casuarinaceae APPENDIX

Allocasuarina globosa L.Johnson, sp. nov. A. sclerocladae affinis sed laminis brevioribus (0.6–1 mm longis); articulis plerumque tenuioribus (0.9–1.2 mm diam.); samara ferruginea; bracteolis infructescentiae 3-partitis; differt. T: Bremer Ra., W.A., 25 Oct. 1964, J.S.Beard 3838; holo: PERTH, with cones. Known only from the Bremer Ra., W.A. Named from the Latin globosus (spherical), referring to the subglobose or shortly cylindrical infructescences (‘cones’). This epithet was pencilled on the type sheet by C.A.Gardner but, since he was intending to recognise it as a species of Casuarina, it would not be appropriate to attach his name to this usage.

Allocasuarina grampiana L.Johnson, sp. nov. Inter species sectionis Cylindropityos combinatione distinguitur: dioica; articuli leviter cerei, sulcis glabratis, phyllichniis rotundato-convexis angulatisve; laminae 0.4–0.8 mm longae, nec plerumque imbricatae nec marcescentes; inflorescentiae masculae moniliformes; corpus infructescentiae 7–9 mm diam.; samara atra. T: upper slopes of Mt Rosea, The Grampians, Vic., 26 Aug. 1965, D.E.Symon 3507; holo: NSW, with cones; iso: AD, MEL. Restricted to The Grampians, Vic., and named from its occurrence there, the combination being taken as grampi- with the Latin adjectival ending -anus.

Allocasuarina gymnanthera L.Johnson, sp. nov. Inter species sectionis Cylindropityos combinatione distinguitur: dioica vel raro monoica; articuli 5–12 mm longi, sulcis plerumque pubescentibus, phyllichniis parum rotundato-convexis; laminae 0.4–0.7 mm longae, plerumque parum imbricatae, non marcescentes; inflorescentiae masculae moniliformes, bracteolis masculis caducis; samara atrobrunnea vel atra. T: Lees Pinch, WollarMerriwa road, N.S.W., 19 Sept. 1951, L.A.S.Johnson NSW 62526; holo: NSW, with cones; iso: CANB, K. Occurs from the Pilliga Scrub S to Goulburn R. valley and Glen Davis, N.S.W. Named from the Greek gymnos (bare, naked) and anthera (an anther), referring to the male flowers, the bracteoles of which fall early, exposing the anther (a condition uncommon in species of section Cylindropitys). The epithet is pronounced with a long ‘e’.

Allocasuarina lehmanniana subsp. ecarinata L.Johnson, subsp. nov. T: creek behind S end of Mylies Beach, near East Mt Barren, W.A., 19 Nov. 1985, A.N.Rodd 5089 & J.McCarthy; holo: NSW (♂,♀, with cones - monoecious); iso: K, PERTH. A subspecie lehmanniana phyllichniis planis vel parum rotundato-convexis plusminusve sulcatis, infructescentiis sessilibus vel in pedunculis ad 4 mm longis, differt. Occurs from near Ongerup to E of Esperance, W.A. The name is from the Latin ecarinatus (without a keel) referring to the more or less flattened phyllichnia.

Allocasuarina mackliniana L.Johnson, sp. nov. A. paradoxae affinis, sed spicis masculis ad anthesin densis, sulcis articulorum saepe pubescentibus, differt. T: 5 km E of Lucindale on Lochaber road, S.A., 8 Mar. 1986, K.L.Wilson 6787 & L.A.S.Johnson; holo: NSW (♀, with cones); iso: AD, K.

196 APPENDIX Casuarinaceae

Occurs from the southern Lofty Ranges, S.A., to western Vic. Named in honour of Miss Ellen D. Macklin, of Adelaide, who published very useful pioneering work on the perplexing ‘distyla group' (section Cylindropitys) in the 1920s and 1930s.

Allocasuarina mackliniana L.Johnson subsp. hirtilinea L.Johnson, subsp. nov. A subspeciebus mackliniana xerophilaque sulcis articulorum valde pubescentibus, phyllichniis leviter rotundato-convexis, differt. T: NE side of Mt Zero, Vic., 21 Aug. 1968, A.C.Beauglehole 28185, ♀ with cones; holo: NSW; iso: MEL. Occurs in western parts of The Grampians, Vic. Named from the Latin hirtus (hairy) and linea (line), referring to the strongly pubescent furrows on the articles.

Allocasuarina mackliniana L.Johnson subsp. xerophila L.Johnson, subsp. nov. A subspeciebus mackliniana hirtilineaque sulcis articulorum initio pubescentibus demum glabrescentibus, phyllichniis fere planis vel vix rotundato-convexis, differt. T: Scorpion Springs Conservation Park, S of Pinnaroo, S.A., 21 Oct. 1973, D.E.Symon 8610 (♂); holo: NSW; iso: AD n.v. Occurs from near Pinnaroo and Keith, S.A., to Wyperfeld Natl Park and the Little Desert, Vic.; in heath in sandy soils. Named from the Greek xeros (dry) and –philos (loving), referring to its occurrence in rather dry habitats.

Allocasuarina media L.Johnson, sp. nov. A. littorali A. paradoxaeque affinis, sed ab hoc habitu fruticosi, cortice laevi, sulcis articulorum plerumque glabratis, inflorescentiis masculis plerumque moniliformibus, differt; ab illo articulis plerumque gracilioribus, dentibus gracilioribus, erectis vel parum patentibus non marcescentibus, differt. T: c. 3 km along Five Mile Road from Tidal River Road, Wilsons Promontory Natl Park, 4 Mar. 1986, K.L.Wilson 6680d & L.A.S.Johnson; holo: NSW (♀, with cones); iso: MEL. Found only at the northern end of Wilsons Promontory, Vic. Named from the Latin medius (middling) referring to its apparently intermediate position between A. littoralis and A. paradoxa. Probably a stabilised taxon of relatively recent hybrid origin from those 2 species (which are in the region but not growing with A. media).

Allocasuarina misera L.Johnson, sp. nov. A. paradoxae affinis sed plerumque tenuior; dentibus 5–7, erectis vel parum patentibus. T: Anglesea, Vic., 10 Mar. 1986, K.L.Wilson 6818 & L.A.S.Johnson; holo: NSW (♀, with cones); iso: MEL. Occurs on the periphery of The Grampians, also near Anglesea and Bairnsdale, Vic. Named from the Latin miser (wretched) for its depauperate appearance as compared with that of the related species A. paradoxa.

Allocasuarina muelleriana subsp. alticola L.Johnson, subsp. nov. A subspecie muelleriana marginibus phyllichniorum nunquam elevatis; articulis validioribus, 0.7–1 mm diam.; et infructescentiis grandioribus, corpore 12–17 mm diam., differt. A subspecie notocolpica infructescentiis sessilis vel peduncula ad 3 mm longa, differt.

197 Casuarinaceae APPENDIX

T: upper slopes of Mt McKinley, S.A., 4 Mar. 1966, D.E.Symon 4016; holo: NSW, with cones; iso: AD, K. Occurs in the northern Flinders Ranges, S.A. Named from the Latin altum (a height) and –cola (a dweller), referring to its occurrence in the Flinders Ranges.

Allocasuarina muelleriana subsp. notocolpica L.Johnson, subsp. nov. A subspecie muelleriana marginibus phyllichniorum nunquam elevatis; articulis validioribus, 0.9–1.1 mm diam.; et infructescentiis grandioribus, corpore 12–18 mm diam., differt. A subspecie alticola peduncula infructescentiae robustiore, 8–17 mm longa, differt. T: road running E–W near ‘Meadow Lea’ homestead joining the South Coast road N of Lake Ada, c. 1 km NW by road from Eleanor R. crossing, Kangaroo Is., S.A., 20 Aug. 1982, E.N.S.Jackson 4357; holo: NSW, with cones; iso: AD. Restricted to Kangaroo Is., S.A. Named from the Greek notos (south) and kolpos (a gulf), referring to its occurrence S of Gulf St Vincent and Spencer Gulf.

Allocasuarina ophiolitica L.Johnson, sp. nov. Inter species sectionis Cylindropityos combinatione distinguitur: dioica; articuli saepe cerei, sulcis plerumque glabratis, phyllichniis rotundato-convexis angulatisve; laminae 0.5–1.3 mm longae, nec imbricatae nec marcescentes; inflorescentiae masculae moniliformes. T: Bralga Tops, Glenrock Stn, N.S.W., 24 Mar. 1981, L.A.S.Johnson 8537; holo: NSW, with cones; iso: BRI, CANB, K, MO. Occurs only from Curricabark to Bralga Tops, N.S.W. Named from the Greek ophis, ophios (a serpent, snake) and lithos (a stone, rock), in the modified form –liticus customarily used in geology, referring to the species' occurrence only on serpentinite rocks.

Allocasuarina portuensis L.Johnson, sp. nov. Inter species sectionis Cylindropityos combinatione distinguitur: dioica; articuli 13–20 mm longi, leviter cerei, sulcis glabratis, phyllichniis angulatis rotundato-convexisve; laminae 0.7–1.1 mm longae, parum imbricatae, patentes; inflorescentiae masculae moniliformes. T: Sydney Harbour Natl Park, 3 Apr. 1986, K.L.Wilson 6843 & L.A.S.Johnson; holo: NSW, with cones; iso: CANB, K, MEL. Known only from Sydney Harbour Natl Park, c. 6 km ENE of Sydney, N.S.W. Named from the Latin portuensis (inhabiting a port) referring to its locality beside Sydney Harbour (Port Jackson).

Allocasuarina rigida subsp. exsul L.Johnson, subsp. nov. A subspecie rigida articulis brevioribus tenuioribusque, 10–14 mm longis, 0.7–0.9 mm diam.; laminis erectis vel parum patentibus, maturitate non vel parum imbricatis. T: Mt Cooroora, Qld, 12 Apr. 1984, P.R.Sharpe 3544; holo: NSW 179094, with cones; iso: BRI. Known only from Mt Cooroora, near Pomona, Qld. Named from the Latin exsul (an exile), referring to its occurrence in isolation from the type subspecies.

198 APPENDIX Casuarinaceae

Allocasuarina rupicola L.Johnson, sp. nov. Inter species sectionis Cylindropityos combinatione distinguitur: dioica; articuli non cerei, sulcis glabratis, phyllichniis angulatis rotundato-convexisve; laminae latissime triangulares, 0.2–0.6 mm longae, imbricatae, non marcescentes; inflorescentiae masculae moniliformes. T: 6.4 km on Boonoo Boonoo Falls road from Boonoo Boonoo, N.S.W., 25 Mar. 1981, L.A.S.Johnson 8539; holo: NSW, with cones; iso: BRI, CANB, K, MO. Occurs about the Qld–N.S.W. border, between Wallangarra and Boonoo Boonoo Falls. Named from the Latin rupes (a rock) and –cola (a dweller), referring to its occurrence amongst rocks.

Allocasuarina simulans L.Johnson, sp. nov. A. distylae affinis sed in omnes partes plerumque tenuior: corpore infructescentiae 9–12 mm diam., pedunculo gracili c. 2 mm diam.; et articulis viridibus, phyllichniis rotundato-convexis, differt. T: 6.4 km SE of Nabiac at old landing site, N.S.W., 16 Feb. 1975, R.G.Coveny 6031, P.Hind & R.Hancock; holo: NSW, with cones; iso: A, K, L, MO, RSA. Occurs from Nabiac to Booti Booti, N.S.W. Named from the Latin simulans (imitating, resembling), referring to its morphological similarity to A. distyla.

Allocasuarina spinosissima (C.Gardner) L.Johnson, comb. nov. Casuarina spinosissima C.Gardner, J. Proc. Roy. Soc. W. Australia 9: 38 (1923). T: Carrabin, W.A., 6 Oct. 1922, C.Gardner 1773 [published as 1273 (original collector's no.)]; holo: PERTH, with cones; iso: MEL.

Allocasuarina thalassoscopica L.Johnson, sp. nov. Inter species sectionis Cylindropityos combinatione distinguitur: dioica; articuli 7–12 mm longi, sulcis pubescentibus, phyllichniis angulatis; laminae 0.3–0.6 mm longae, erectae, nec imbricatae nec saepissime marcescentes; inflorescentiae masculae non moniliformes. T: Mt Coolum, Qld, 23 Apr. 1981, L.A.S.Johnson 8562, P.Sharpe & K.Wilson; holo: NSW, with cones; iso: BRI. Known only from Mt Coolum, Qld. Named from the Greek thalassa (the sea) and skopos (a watcher), referring to its situation on the mountain slope facing the sea.

Allocasuarina zephyrea L.Johnson, sp. nov. Inter species sectionis Cylindropityos combinatione distinguitur: dioica; articuli leviter cerei, sulcis glabratis, phyllichniis angulatis; laminae 0.4–1 mm longae, plerumque parum imbricatae, erectae patentesve, raro marcescentes; inflorescentiae masculae moniliformes; samara atra. T: Ocean Beach, Strahan, Tas., 20 Jan. 1949, L.A.S.Johnson NSW 62623; holo: NSW, with cones. Occurs in western and central Tas. Named from the Greek zephyros (the west wind), referring to its occurrence on the western side of Tas.

199 Casuarinaceae APPENDIX

CASUARINA

L.A.S.Johnson

Casuarina bicuspidata Benth., Fl. Austral. 6: 202 (1873). T: South Coast Island VII, 13 Mar. 1802, R.Brown; lecto (here chosen): K (♀, with inflorescences and cones); isolecto: B, BM. The locality was cited by Bentham as Flinders Is. but the sheet at BM is labelled Anchorage VII, which was Waldgrave Is., S.A., from which Flinders Is. was indeed visited by Robert Brown. However, there must have been confusion in labelling, since this species (=Allocasuarina trichodon) occurs only in W.A. It does grow in the areas in W.A. in which Brown collected, and it must be assumed that he collected it at one of those sites — probably King George Sound or Lucky Bay.

Casuarina cambagei R.Baker, Proc. Linn. Soc. New South Wales 24: 605 (1900). T: Grawlin, Forbes, N.S.W., 28 July 1899, R.H.Cambage NSW 46670; lecto (here chosen): NSW, with cones. Chosen as the only specimen found that was annotated by Baker. This is a synonym of C. cristata.

Casuarina cunninghamiana Miq., Rev. Crit. Casuarineae 56 (1848). T: sandy shores of Moreton Bay and Glasshouse Bay [Qld], collector unknown; lecto (here chosen): K, with cones. Miquel cited the collector as Allan Cunningham but there is no indication on the sheet that this is so (the label is not in Cunningham's script). The specimen can hardly have been collected on the bay shores: it occurs in the Moreton region but along freshwater streams only.

Casuarina cunninghamiana subsp. miodon L.Johnson, subsp. nov. A subspecie cunninghamiana laminis 6 vel 7, flavidis, non marcescentibus; phyllichniis latioribus et minus angulatis; et bracteolis infructescentiae acutioribus, differt. T: 23 km NNE of Borroloola on Bing Bong road, N.T., 13 May 1983, K.L.Wilson 5361; holo: NSW, with cones; iso: BRI, CANB, DNA. Occurs from the Daly R., N.T., to the Gulf of Carpentaria, north-western Qld. Named from the Greek meion (few) [Latinised as mio- in combinations] and odous, odontis (a tooth) [Latinised as -odon in compound substantives], referring to the relatively fewer teeth.

Casuarina distyla var. prostrata Maiden & Betche, Proc. Linn. Soc. New South Wales 30: 371 (1905). T: Narrabeen to Barrenjoey, N.S.W., July 1905, R.Cambage & J.Maiden NSW 62648; lecto (here chosen): NSW, with cones; isolecto: BM, MEL. The lectotype sheet was annotated by Betche and has a copy of the description attached. This is a synonym of Allocasuarina paludosa.

200 APPENDIX Casuarinaceae

Casuarina equisetifolia L., Amoen. Acad. 4: 143 (1759) as equisefolia. T: Rumphius, Herbarium Amboinense 3: pl. 57 (1743) as C. litorea; holo. There are no extant Linnaean specimens of this species: Linnaeus based the species solely on Rumphius's publication and illustration, which refer unambiguously to this species. The validity of Linnaeus's publication of this name is arguable though defended by Friis (Taxon 29: 499–501, 1980). The next valid publication is by Forster & G.Forster, Char. Gen. Pl. 104 (1775). If Linnaeus's publication were regarded as invalid, the name of this species and that of the genus would not be affected, though the authors and dates of publication would be changed.

Casuarina equisetifolia var. microcarpa F.Muell., Fragm. 6: 17 (1867). T: Glendon, N.S.W., L.Leichhardt; lecto (here chosen): MEL, with cones. This lectotypification makes the variety a synonym of C. cunninghamiana subsp. cunninghamiana. Besides specimens of C. cunninghamiana, Mueller also included specimens from the Barrier Ra., which are C. pauper.

Casuarina lepidophloia F.Muell., Fragm. 10: 115 (1877). T: between the Bogan R. and the Lachlan R., N.S.W., L.Morton; lecto (here chosen): MEL (♀, with cones and galls); isolecto: NSW 46689. There are several syntypes in MEL, from which the lectotype is chosen as bearing cones (Mueller saw only fruiting material). This name is regarded as a synonym of C. cristata. However, the material on which C. lepidophloia was based is somewhat inadequate and it may rather represent an intergrade or hybrid between C. cristata and C. pauper. It is not C. pauper itself.

Casuarina leptoclada Miq., Rev. Crit. Casuarineae 41 (1848). T: Sidney [Sydney], N.S.W., Clowes; lecto (here chosen): K (♂); iso: U. This is a synonym of Allocasuarina littoralis, one of the slender forms of that species.

Casuarina luehmannii R.Baker, Proc. Linn. Soc. New South Wales 24: 608 (1900). T: Grenfell, N.S.W., no date, R.H.Cambage NSW 50724; lecto (here chosen): NSW, with cones. This is Allocasuarina luehmannii. The type sheet also bears a specimen with male inflorescences.

Casuarina moesta F.Muell. ex Miq., Ned. Kruidk. Arch. 4: 98 (1859). T: near Melbourne, Vic., F.Mueller; lecto (here chosen): MEL (♀, with inflorescences); isolecto: B, LE, P. A MEL specimen is chosen since no potential type material was found in U. This is a synonym of Allocasuarina littoralis, representing one of the slender forms of that species.

Casuarina obesa Miq. in J.G.C.Lehmann, Pl. Preiss. 1: 640 (1845). T: towards Swan R. above Perth, W.A., 27 June 1839, and towards Port Leschenault, W.A., 30 Dec. 1839, L.Preiss 2002; lecto (here chosen): U (♀, with cones); isolecto: LE, MEL, NSW, P, S. It is impossible to separate localities and dates on the sheets seen. The material seen in U is female; however, the material in P includes male as well as female.

201 Casuarinaceae APPENDIX

Casuarina pauper F.Muell. ex L.Johnson, sp. nov. C. cristata affinis, sed laminis patentibus recurvatisve; articulis crassioribus, 1–1.8 mm diam., dense pubescentibus cereisque; differt. T: Flinders Ra., S.A., Oct. 1851, F.Mueller; holo: MEL (♀, with inflorescences and cones). Occurs in W.A., S.A., N.S.W. and Vic. Named from the Latin pauper (poor), referring to its depauperate habit compared to C. cristata. This is the epithet used by F.Mueller and it is written in his hand on the holotype. This was published as a new combination in Nuytsia 1: 265 (1972), based on C. pauper F.Muell. ex Miq., Ned. Kruidk. Arch. 4: 100 (1859). However, Miquel did not accept Mueller's species: he regarded it as not differing from C. glauca. Thus the previous publication of the subspecies as a new combination was invalid.

Casuarina preissiana Miq. in J.G.C.Lehmann, Pl. Preiss. 1: 640 (1845). T: Mt Eliza [Kings Park, Perth], W.A., 19 May 1839, L.Preiss 2008 p.p.; lecto (here chosen): U (♂); isolecto: ?BR, C, K, L, LE, MEL, P, ?S, UPS, W. The lectotype is the specimen in the best condition of those seen at U. This is a synonym of Allocasuarina humilis.

Casuarina prinsepiana C.Andrews, J. W. Austral. Nat. Hist. Soc. 1: 43 (1904). T: Mullewa, W.A., July 1903, C.R.P.Andrews; lecto (here chosen): PERTH (♀, with inflorescences). There is also a specimen bearing male inflorescences on what was previously regarded as the holotype sheet. This is Allocasuarina acutivalvis subsp. prinsepiana.

Casuarina quadrivalvis Labill., Nov. Holl. Pl. 2: 67 (1806). T: New Holland [Tas.], J.J.H. de Labillardi~hre; lecto (here chosen): FI (♀, with inflorescences and cones); probable isolecto: P. The lectotype is one of 3 sheets at FI and bears Labillardi~hre's manuscript description and notes. This is a synonym of Allocasuarina verticillata.

GYMNOSTOMA

L.A.S.Johnson

Gymnostoma australianum L.Johnson, sp. nov. Inter species Gymnostomatis combinatione distinguitur: ramuli rigidiusculi non laxe pendentes, apicibus novellorum ferrugineo-pubescentibus vel canis, articulis 2.5–4 mm longis, 0.5–0.8 mm diam., phyllichniis acutiuscule angulatis, laminis foliorum 0.4–0.7 mm longis anguste deltoideis; infructescentiae canae vel ferrugineo-pubescentes, corpore maturae 7–10 mm longo, c. 10 mm diam., bracteis non striatis, bracteolis plus minusve striatis. T: Mt Alexander, Daintree R., alt. 1300 m [but see below], Qld, 17 Dec. 1917, S.F.Kajewski 1492; holo: NSW, with cones; iso: A, BO, BRI, K, MEL, NY, SING. Evidently restricted to a very few small populations near Thornton Peak, Qld. According to botanists at the Queensland Herbarium (BRI), Kajewski mistook Thornton Peak for Mt Alexander and his indicated locality is therefore in error. Named in reference to its occurrence in Australia (the only extant species of the genus in Australia).

202

SUPPLEMENTARY GLOSSARY bathyphyll: a leaf at the base of a stem with the function of attachment to a substrate. hypodermis: a layer of cells below the epidermis. marcescent: withering without falling off. moniliform: contracted at short regular intervals like a string of beads. ostiole: an opening or pore, e.g. (in Moraceae) at the apex of a fig, or (in fungi and lichens) at the apex of a perithecium. adj. ostiolar, ostiolate. phyllichnium: in Casuarinaceae, the ridge of a branchlet article; pl. phyllichnia.

203