FLORA OF AUSTRALIA

HAMAMELIDACEAE
H.J.Hewson

Shrubs or trees, monoecious, rarely dioecious. Leaves simple, alternate, rarely opposite,
simple to palmately lobed, petiolate; stipules usually present. Inflorescence usually a spike,
sometimes a raceme or panicle, bracteate. Flowers bisexual or unisexual, usually regular.
Sepals 4 or 5, free or connate, small, sometimes absent. Petals 4 or 5, ligulate, small,
sometimes absent. Stamens 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. Ovary usually inferior, sometimes superior, with 2,
rarely 3, carpels; styles 2, rarely 3, free, usually persistent in fruit; ovules 1 or 2, pendulous,
or 5–many, anatropous. Fruit (in Australia) a loculicidal capsule, woody. Seeds sometimes
winged; endosperm present.
A family of 26 genera and c. 100 species, of subtropical to warm temperate regions
around the world but predominantly in eastern Asia. Three monotypic genera endemic in
Australia. All 3 genera are in the tribe Hamamelideae of the subfamily Hamamelidoideae.
The family has a long fossil record and many representatives may be relictual. Species of
Hamamelis L. (Witch-hazel) and Liquidambar L. are important ornamental plants. 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 Queensland 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 fruits.
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;
ovule 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 genus 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).
Tree to 25 m tall; indumentum stellate. Leaves elliptic to lanceolate; lamina to 17 cm long
and 6 cm wide; petiole 7–12 mm long; stipules lanceolate, to 7 mm long. Spike on peduncle
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 rainforest 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; flower 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; stigma 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 shrubs, evergreen or deciduous, 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. Perianth 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
nut, 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. CELTIS

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 Europe, Asia and North America; 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 bud; 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. Seed with curved embryo;
endosperm absent.
A genus of 5 species in Madagascar, Mexico and from India 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, Philippines, 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. New South Wales 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).
Shrub 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; pedicel 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 Wood, Greyhandle Wood. Fig. 26A–D.
Occurs in coastal districts and up to 100 km inland, from Cape York Peninsula, Qld, S to
the Manning R., N.S.W.; grows in rainforest; also in the Philippines, New Guinea 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.
Leaf 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
Africa 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 Cairns, 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 Pine
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. *Celtis australis 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. *Celtis occidentalis 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 Melanesia, Polynesia, 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 Matthew
Flinders ‘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, Cannabaceae, Moraceae and Urticaceae.
W.T.Stearn, ed. C.R.Joyce & S.M. & A.Curry, The Botany and Chemistry of Cannabis
(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. Inflorescences pendent; male
much-branched; female lax, cone-like.
A genus of 2 species in the northern hemisphere; 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 human 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 plant.
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 (Ficus carica L.), the
Mulberry (Morus alba L.), the Jackfruit and the Breadfruit (Artocarpus spp.). Over 40
species of Ficus are cultivated as indoor ornamentals and garden and roadside trees. Antiaris
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, Artocarpeae &
Moreae, 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. Singapore 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
Northern Territory, 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. CASTILLEAE

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 † DORSTENIA
1: Flowers not so
3 Herbs shorter than 1 m 3. FATOUA
3: Trees, shrubs taller than 1 m, or woody climbers
4 Woody climbers with axillary spines 4. MACLURA
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 stamen; 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. STREBLUS
† Dorstenia arifolia Lam., a native of Brazil, 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. Bat. 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
A ustralia, 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
a cuminatus (Planchon) Seemann, loc. cit. T: Mt Marshall, Illawarra District, N.S.W., J.Backhouse; holo:
K.
I llustrations: 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 Java, 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 Malaysia and Indonesia 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: Castilla 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 root-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, Gulf of Carpentaria, 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 roots 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 northern Australia. 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.

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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.

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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 New Caledonia 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 Caroline Islands. 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. Ficus macrophylla 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., Vanuatu,
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, Woods; 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: western Australia, ?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: Cassowary 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.

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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.

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 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 cauliflory 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 Borneo; 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 Pilea Lindley have been widely cultivated as indoor plants. The Ramie Fibre of
commerce comes from Boehmeria nivea Gaudich. Young shoots of the European Nettle,
Urtica urens L. have been eaten in times of economic hardship. Species of the tribe Urticeae,
especially the genera Urtica, Laportea and Dendrocnide, 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: Urera 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. Dendrocnide moroides (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. Dendrocnide excelsa (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, Boehmeria
macrophylla, 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. Taiwan 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 & Beech 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, France.
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).

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 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 Kenya.

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 catkin; 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 Fiji; 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 Rosales). 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 (acorn) 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. (birch) 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, Betulaceae, Pflanzenr. 19: 1–149 (1904).

1. ALNUS

Alnus Miller, Gard. Dict. Abr. 4th edn (1754); the Latin name for the alder.
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 buds 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. Catkins 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).

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 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 (Oak) and Fagus (Beech) from the Northern Hemisphere and
Nothofagus (Antarctic Beech and Southern Beech) in the Southern Hemisphere are
important timber trees. Quercus suber L. yields cork. Castanea sativa Miller (Chestnut)
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,
ecology 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 temperate rainforest 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.

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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 Gymnostoma, 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 Allocasuarina, 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 Casuarina, 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 (Hemiptera) (Gullan, 1984), form distinctive
galls on Casuarinaceae; some of these may be mistaken for fruiting cones but they are
actually modified vegetative branchlets. Actinomycete bacteria (Frankia) occur in nodules on

100
Figure 45. Details of Casuarinaceae. A–H, samaras ×5. A, Gymnostoma australianum
(NSW 63049, NSW); B, Casuarina cristata (K.Wilson 5906, NSW); C, C. cunninghamiana
subsp. cunninghamiana (N.Hall H80/19, NSW); D, Allocasuarina paludosa (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.

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 CASUARINACEAE

the roots and are effective in fixation of atmospheric nitrogen. Vesicular-arbuscular

endotrophic mycorrhizae have also been reported.
Macrofossils from the Eocene 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 fossils 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 South Africa, 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 Casuarina glauca and Allocasuarina littoralis 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-oaks, 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. Casuarina equisetifolia 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 Vietnam, 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.

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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. Casuarina cunninghamiana 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. Casuarina obesa 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. Casuarina pauper 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.

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 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

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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

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 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)

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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

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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

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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

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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

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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

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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

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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

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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

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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

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 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

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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

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 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

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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

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 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.

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 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.

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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. Allocasuarina luehmannii (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.

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 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.

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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.

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 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).

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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. Allocasuarina globosa 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.

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 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. Allocasuarina campestris (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.

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 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.

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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.

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 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. Allocasuarina dielsiana (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. Allocasuarina helmsii (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.

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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. Allocasuarina torulosa (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

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 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. Allocasuarina fraseriana (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 (Eucalyptus marginata)
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).

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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.

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3. Allocasuarina CASUARINACEAE

23. Allocasuarina huegeliana (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);
♀E) . (
isolecto: BM, BR, C, FI, K, L, LE, MEL, P, S; syn: L
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.

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 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. Allocasuarina inophloia (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.

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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.

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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

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 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

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A. emuina and A. thalassoscopica it is distinguished by its longer articles with glabrous

furrows, 7–9 teeth, and usually smaller cones.

32. Allocasuarina portuensis 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.

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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.

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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. Allocasuarina distyla (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.

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37. Allocasuarina diminuta 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)
♂, (with cones
T: Bundanoon–Penrose, N.S.W., Apr. 1948, M.Tindale NSW 62492; holo: NSW -
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. Allocasuarina defungens 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. Allocasuarina grampiana 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. Allocasuarina monilifera (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. Allocasuarina crassa 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. Allocasuarina paradoxa (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. Allocasuarina pusilla (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.

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 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. Allocasuarina striata (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

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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.

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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. Allocasuarina nana (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. Allocasuarina humilis (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.,
( ; iso: U.
J.Drummond in Herb. Hooker; holo: K♂)
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. Allocasuarina thuyoides (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♀
eets peci
as 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)
4. Ulmus hollandica (5)
7. Trema tomentosa
var. tomentosa (9)
10. Celtis occidentalis (11)
13. Celtis paniculata (13)
2. Neostrearia fleckeri (2)
5. Aphananthe philippinensis (5)
8. Trema tomentosa
var. viridis (9)
11. Celtis sinensis (11)
14. Humulus lupulus (14)
3. Noahdendron nicholasii (4)
6. Trema orientalis (8)
9. Celtis australis (10)
12. Celtis philippensis (11)
15. Cannabis sativa (15)
177
16. Streblus brunonianus (18)
19. Fatoua pilosa (20)
22. Antiaris toxicaria
var. macrophylla (24)
25. Ficus virens
var. sublanceolata (35)
28. Ficus drupacea
var. glabrata (36)
178
17. Streblus glaber
var. australianus (18)
20. Maclura cochinchinensis (22)
23. Ficus superba
var. henneana (32)
26. Ficus virens
var. dasycarpa (35)
29. Ficus benghalensis (36)
18. Malaisia scandens
var. scandens (19)
21. Artocarpus glaucus (23)
24. Ficus virens
var. virens (35)
27. Ficus drupacea
var. drupacea (36)
30. Ficus benjamina
var. benjamina (37)
31. Ficus microcarpa
var. microcarpa (38)
34. Ficus triradiata
var. triradiata (40)
37. Ficus macrophylla
subsp. macrophylla (41)
40. Ficus rubiginosa (42)
43. Ficus leucotricha
var. sessilis (44)
32. Ficus microcarpa
var. hillii (39)
35. Ficus triradiata
var. sessilicarpa (40)
38. Ficus baileyana (41)
41. Ficus leucotricha
var. leucotricha (43)
44. Ficus obliqua
var. obliqua (44)
32. Ficus microcarpa
var. latifolia (39)
36. Ficus watkinsiana (40)
39. Ficus destruens (42)
42. Ficus leucotricha
var. megacarpa (43)
45. Ficus obliqua
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)
64. Ficus opposita
var. micracantha (57)
67. Ficus fraseri (59)
70. Ficus tinctoria
subsp. tinctoria (60)
73. Ficus adenosperma (63)
62. Ficus opposita 63. Ficus opposita
var. opposita (56) var. indecora (56)
65. Ficus scobina (57) 66. Ficus podocarpifolia (57)
68. Ficus coronulata (59) 69. Ficus coronata (59)
71. Ficus virgata 72. Ficus mollior
var. virgata (60) var. mollior (62)
74. Ficus variegata 75. Ficus nodosa (64)
var. variegata (64)
181
76. Ficus congesta
var. congesta (65)
79. Ficus septica
var. cauliflora (66)
82. Laportea interrupta (73)
85. Dendrocnide cordata (76)
88. Elatostema reticulatum (80)
182
77. Ficus hispida
var.hispida (65)
80. Urtica incisa (70)
83. Dendrocnide
corallodesme (75)
86. Dendrocnide
photinophylla (76)
89. Elatostema stipitatum (82)
78. Ficus septica
var. septica (66)
81. Urtica urens (72)
84. Dendrocnide moroides (76)
87. Dendrocnide excelsa (78)
90. Procris pedunculata (84)
91. Boehmeria macrophylla (85)
94. Pouzolzia hirta (88)
97. Parietaria debilis (91)
100. Balanops australiana (95)
103. Nothofagus
cunninghamii (98)
92. Boehmeria nivea (85)
95. Pipturus argenteus (88)
98. Parietaria judaica (92)
101. Alnus cordata (96)
104. Nothofagus gunnii (100)
93. Pouzolzia zeylanica (86)
96. Nothocnide repanda (89)
99. Australina pusilla (93)
102. Nothofagus moorei (98)
105. Gymnostoma
australianum (103)
183
106. Casuarina equisetifolia
subsp. equisetifolia (106)
109. Casuarina cunninghamiana
subsp. miodon (107)
112. Casuarina cristata (108)
115. Allocasuarina acuaria (129)
118. Allocasuarina fibrosa (130)
184
107. Casuarina equisetifolia
subsp. incana (106)
110. Casuarina glauca (107)
113. Casuarina pauper (110)
116. Allocasuarina pinaster (129)
119. Allocasuarina
ramosissima (130)
108. Casuarina cunninghamiana
subsp. cunninghamiana
(107)
111. Casuarina obesa (108)
114. Allocasuarina
decaisneana (127)
117. Allocasuarina
grevilleoides (129)
120. Allocasuarina
luehmannii (132)
121. Allocasuarina
corniculata (133)
124. Allocasuarina acutivalvis
subsp. prinsepiana (136)
127. Allocasuarina
tortiramula (137)
130. Allocasuarina eriochlamys
subsp. grossa (139)
133. Allocasuarina helmsii (141)
122. Allocasuarina 123. Allocasuarina acutivalvis
spinosissima (133) subsp. acutivalvis (135)
125. Allocasuarina 126. Allocasuarina globosa (136)
scleroclada (136)
128. Allocasuarina 129. Allocasuarina eriochlamys
campestris (128) subsp. eriochlamys (139)
131. Allocasuarina 132. Allocasuarina
tessellata (139) dielsiana (141)
134. Allocasuarina torulosa 135. Allocasuarina
(142) decussata (142)
185
136. Allocasuarina
fraseriana (143)
139. Allocasuarina
trichodon (146)
142. Allocasuarina media (148)
145. Allocasuarina
thalassoscopica (150)
148. Allocasuarina
portuensis (152)
186
137. Allocasuarina
verticillata (144)
140. Allocasuarina
inophloia (147)
143. Allocasuarina filidens (150)
146. Allocasuarina rigida
subsp. rigida (151)
149. Allocasuarina rupicola (152)
138. Allocasuarina
huegeliana (146)
141. Allocasuarina littoralis (148)
144. Allocasuarina emuina (150)
147. Allocasuarina rigida
subsp. exsul (151)
150. Allocasuarina
ophiolitica (152)
151. Allocasuarina distyla (154)
154. Allocasuarina diminuta
subsp. mimica (155)
157. Allocasuarina
glareicola (157)
160. Allocasuarina
monilifera (159)
163. Allocasuarina lehmanniana
subsp. ecarinata (161)
152. Allocasuarina simulans (154)
155. Allocasuarina diminuta
subsp. annectens (157)
158. Allocasuarina
grampiana (158)
161. Allocasuarina crassa (159)
164. Allocasuarina
paradoxa (161)
153. Allocasuarina diminuta
subsp. diminuta (155)
156. Allocasuarina
defungens (157)
159. Allocasuarina
zephyrea (158)
162. Allocasuarina lehmanniana
subsp. lehmanniana (161)
165. Allocasuarina mackliniana
subsp. mackliniana (165)
187
166. Allocasuarina mackliniana
subsp. hirtilinea (162)
169. Allocasuarina pusilla (163)
172. Allocasuarina
gymnanthera (166)
175. Allocasuarina muelleriana
subsp. muelleriana (169)
178. Allocasuarina nana (170)
188
167. Allocasuarina mackliniana
subsp. xerophila (163)
170. Allocasuarina robusta (165)
173. Allocasuarina
paludosa (166)
176. Allocasuarina muelleriana
subsp. notocolpica (169)
179. Allocasuarina humilis (172)
168. Allocasuarina misera (163)
171. Allocasuarina striata (165)
174. Allocasuarina
brachystachya (168)
177. Allocasuarina muelleriana
subsp. alticola (169)
180. Allocasuarina
drummondiana (172)
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 Flora of
Australia 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.

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 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.

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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.

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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 extinction.

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.

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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.

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 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.

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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.

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 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.

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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.

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 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.

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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