Editorial Committee

P.I.Forster (editor)

P.D.Bostock (technical advisor)

G.P.Guymer (technical advisor)

Graphic Design

Will Smith

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Yvonne Smith

Austrobaileya

Vol. 1, No. 1 was published on 1 December 1977

Vol. 8, No. 2 was published on 7 December 2010

Vol. 8, No. 3 was published on c. 5 December 2011

Austrobaileya is published once per year.

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ISSN 0155-4131

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Austrobaileya is the journal of the Queensland Herbarium and publishes peer-reviewed research

on plants, algae, fungi and lichens (systematics, morphology, geography, anatomy, karyology,

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Austrobaileya 8 ( 3 ): 225-440 ( 2011 )

Contents

A taxonomic revision of Symplocos Jacq. (Symplocaceae) in Australia

L.W. Jessup . 225-251

A taxonomic revision of Pseudoweinmannia Engl. (Cunoniaceae: Geissoieae )

A.C.Rozefelds & B.Pellow . 252-266

A taxonomic revision of Nyssanthes R.Br. (Amaranthaceae)

A. R. Be an & C. II. Miller . 267-279

A taxonomic revision of Pterocaulon section Monenteles (Labill.) Kuntze

(Asteraceae: Inuleae-Plucheinae)

A. R. Bean . 280-334

Heliotropium microspermum E.J.Thomps. (Boraginaceae), a new species

from Queensland

E. J.Thompson . 335-339

Two new species of Pluchea Cass. (Asteraceae: Plucheinae) from

Queensland, Australia

A. R. Bean . 340-346

Eucalyptus melanophloia subsp. nana D.Nicolle & Kleinig, a new mallee

ironbark (E. series Siderophloiae ; Myrtaceae) from central Australia and

north western Queensland

D.Nicolle & D.A.Kleinig . 347-355

Cycas terryana P.I.Forst. (Cycadaceae), a new species from central Queensland

P.I.Forster . 356-363

A taxonomic revision of Trichosanthes L. (Cucurbitaceae) in Australia,

including one new species from the Northern Territory

W.E.Cooper & H.J.de Boer . 364-386

Five new species of Plectranthus L.Her. (Lamiaceae) from New South Wales

and Queensland

P.I.Forster . 387-404

Wilkiea kaarruana Zich & A.J.Ford (Monimiaceae), a new species from

north-east Queensland

F. Zich & A.J.Ford . 405-411

New and reinstated species of the Solanum ellipticum R.Br. (Solanaceae)

species group

A. R. Be an .412-430

Luvunga monophylla (DC.) Mabb. (Rutaceae): a new species for

Queensland

D.G.Fell & D.J.Stanton . 431-434

Hydrocharis dubia (Blume) Backer (Hydrocharitaceae) is an alien species

in Australia

A. R. Be an . 435-437

Reinstatement of Vigna suberecta Benth. (Fabaceae: Phaseoleae)

A. E. Holland . 438-440

A taxonomic revision of Symplocos Jacq.

(Symplocaceae) in Australia

L.W. Jessup

Summary

Jessup, L.W. (2011). A taxonomic revision of Symplocos Jacq. (Symplocaceae) in Australia.

Austrobaileya 8(3): 225-251. The Australian taxa in the genus Symplocos are revised. Nineteen

species are recognised and separate identification keys are provided for flowering and fruiting material.

Symplocos boonjee Jessup, S. bullata Jessup and S. oresbia Jessup are described as new species, four

taxa are raised in rank to species viz. S. gittinsii Jessup, S. glabra Jessup, S. puberula Jessup and

S. wooroonooran Jessup and one new variety, S. cyanocarpa var. pilosa Jessup is described.

Key Words: Symplocaceae, Symplocos , Australia flora, Queensland flora. New South Wales flora,

Symplocos boonjee, Symplocos bullata, Symplocos cyanocarpa var. pilosa, Symplocos gittinsii,

Symplocos glabra, Symplocos oresbia, Symplocos puberula, Symplocos wooroonooran, new species,

identification key

Laurence W. Jessup, Queensland Herbarium, Department of Environment and Resource Management,

Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email:

Laurence.Jessup@derm.qld.gov.au

Introduction

Symplocos Jacq. is a predominantly

pantropical genus of over 320 species with

highest species diversity in Asia, Malesia and

the New World (Wang et al. 2004; Fritsch et

al. 2006, 2008). Symplocos has been usually

regarded as the sole generic representative

of the Symplococeae with division into four

subgenera and various sections based upon

floral structure (Brand 1901). Molecular

phylogenies to date have not supported

this classification and have resulted in the

elevation of Symplocos section Cordyloblaste

to generic rank (as Cordyloblaste Hensch.

ex Moritzi) and the recognition of only two

subgenera and relatively few sections and

series in Symplocos (Fritsch et al. 2008). This

infrageneric classification of the genus is

based on DNA sequence data from five genic

regions (for 90 species) and 51 morphological

characters (Fritsch et al. 2008). For the

Australian region, all taxa can be classified

into Symplocos (subgenus Symplocos)

section Lodhra G.Don (Fritsch et al. 2008)

and represents a major departure from that

proposed by Nooteboom (1975).

Accepted for publication 2 September 2011

Mueller (1862) was the first to recognise

Symplocos in Australia ( S. thwaitesii F.Muell.)

and followed this with a second new species (S.

stawellii F.Muell.) (Mueller 1865). Bentham

(1869) recognised Symplocos thwaitesii but

applied the name S. spicata var. australis

Benth. to specimens of S. stawellii. Bailey

(1890) described Symplocos paucistaminea

F.Muell. & F.M.Bailey ex F.M.Bailey and in

Bailey (1900b) treated the other Australian

species in the same sense as Bentham

(1869). Brand (1901) reinstated the name

Symplocos stawellii and described the Lord

Howe Island species S. candelabrum Brand.

Nooteboom (1975) revised the Old World

species creating a large number of synonyms

and four subspecies and thirty-four varieties

under the apparently polymorphic species

Symplocos cochinchinensis (Lour.) S.Moore.

He followed with a modified treatment of

Australian Symplocos in Nooteboom (1981).

Smith (1978, 1982) chose not to follow

Nooteboom’s broad taxonomic concept in his

treatment of the Fijian species as did Green

(1994) in the Oceanic Islands volume of the

Flora of Australia.

Since 1981 new taxa from Australia have

been recognised and some taxa that were

placed in synonymy by Nooteboom have

continued to be recognised as distinct in

226

Austrobaileya 8(3): 225-251 (2011)

the field and in the herbarium particularly

as more collections have been made (Jessup

1993, 1994, 1997, 2002, 2007, 2010). Also

all of the Australian varieties described by

Nooteboom under Symplocos cochinchinensis

subsp. thwaitesii (F.Muell.) Noot. are now

well-collected throughout their natural range

and are represented in the herbarium by

quality specimens that exhibit several distinct

correlating character states sufficient to allow

recognition as species. In this paper four

previously described taxa are raised in rank

to species and three new species and one new

variety are described.

Materials and methods

This paper is based on herbarium collections in

BM, BRI, CANB, CNS (previously QRS), K,

L, MEL and NSW and some field observations

by the author. All specimens cited have been

seen by the author. Descriptions of flowers

were prepared from material preserved

in FAA or 70% alcohol and glycerol or

reconstituted by briefly boiling in water.

Descriptions of fruit were prepared from

both dried and alcohol preserved material.

Common abbreviations in the specimen

citations are F.R. (Forest Reserve), L.A.

(Logging Area), N.R (National Park), S.F.R./

S.F. (State Forest Reserve/State Forest), T.R.

(Timber Reserve). The abbreviation NCA

is for the Queensland Nature Conservation

Act 1992 and its associated regulations. The

abbreviation EPBC refers to the Environment

Protection and Biodiversity Conservation Act

1999

Taxonomy

Symplocos Jacq., Enum. PI. Carib. 5, 24

(1760). Type species: S. martinicensis Jacq.

Trees or shrubs, dioecious, polygamous

or hermaphroditic. Leaves mostly spirally

arranged, simple, penninerved, serrate,

dentate, or entire, exstipulate. Inflorescence

spicate, racemose, a thyrse or a panicle, or

flowers solitary or fasciculate, axillary, rarely

terminal. Flowers actinomorphic, bisexual or

often functionally unisexual, subtended by

a bract and 2 bracteoles. Ovary inferior or

rarely semi-inferior. Hypanthium completely

adnate to ovary. Calyx limb epigynous,

mostly 5-lobed; lobes imbricate, persistent.

Corolla gamopetalous but divided to near

base into 3-10 petals (5 in Australian species),

quincuncially imbricate, white, cream or pink.

Stamens few to many, connate only at base,

adnate to the base of the corolla, pentadelphous

in alternipetalous bundles or (not in Australia)

monadelphous; filaments terete. Anthers

bilocular, introrse, dehiscing longitudinally.

Disc inside staminal ring. Ovary 2-5-locular

(3-locular in Australia) with 2-4 anatropous

ovules per locule. Placentation axile but septa

separating distally. Style 1, stigma simple,

capitate or peltate. Fruit a monopyrenous

drupe, crowned by persistent calyx lobes,

frequently 1-celled and 1-seeded by abortion;

mesocarp fleshy or not; endocarp hard, smooth

or longitudinally ridged, with an apical

depression. Seed with copious endosperm;

embryo straight or curved.

The genus includes 300-350 species in

tropical, subtropical and temperate parts of

Asia, Malesia, North and South America, the

West Indies, the Pacific islands and Australia.

Nineteen species occur in Australia.

Jessup, Symplocos in Australia

227

Fig. 1. Symplocos ampulliformis. A. part of inflorescence with flowers and buds x 4, Gray 3362 (BRI). B. fruit x 2,

Hyland 13459 (BRI). C. leafabaxial surface x fl Volck 1389 (BRI). .S', boonjee. D. inflorescence with flowers x 3, Gray

5575 (CNS). E. flower x 6, Hyland 14345 (CNS). F. fruit x 2, Gray 5575 (CNS). S. bullata. G. inflorescence with flowers

x 3. H. leaf adaxial surface x 1 . 1 . leaf abaxial surface x 1 . G-I from Gray 2543 (BRI). Del. W.Smith.

228

Austrobaileya 8(3): 225-251 (2011)

Key to Australian Symplocos taxa using flowering specimens

1 Inflorescence racemose or a panicle of shortly pedicellate flowers.2

1. Inflorescence spicate or a panicle or fascicle of sessile flowers, or flowers

solitary, axillary.10

2 Lamina margins serrate or dentate, sometimes obscurely denticulate.3

2. Lamina margins denticulate or entire.7

3 Petiole up to 4 mm long.4

3. Petiole 5-17 mm long.5

4 Young branchlets appressed puberulous; petals 1.4—1.6 mm wide;

stamens mostly 15-20.2. S. baeuerlenii

4. Young branchlets sparsely pilose with erect hairs; petals 2-2.5 mm wide;

stamens 30-40 . 11. S. harroldii

5 Young branchlets, petiole and inflorescence glabrous or nearly so.6

5. Young branchlets, petiole and inflorescence appressed puberulous or

pubescent.16. S. puberula

6 Calyx limb 1.7-2 mm long; stamens more than 40; petiole mostly 5-15

mm long.18. S. thwaitesii

6. Calyx limb 0.7-0.8 mm long; stamens less than 40; petiole mostly 20-30

mm long.5. S. candelabrum

7 Bracts up to 1.2 mm long; bracteoles up to 0.6 mm long; petals up to 4 mm

long; stamens up to 20; style up to 2.5 mm long.8

7. Bracts 1-2.5 mm long; bracteoles 0.6-1.2 mm long; petals 4 mm long or

more; stamens 30-50; style 3 mm long or more.9

8 Secondary veins mostly 7-9 pairs; basal ones not more acute and longer

than distal ones; bracts caducous before anthesis; petals 2-2.5 mm long;

disc and style glabrous.

8. Secondary veins mostly 4-6 pairs; basal ones more acute and longer than

distal ones; bracts persistent to after anthesis; petals 3-3.7 mm long;

disc and base of style puberulous.1. S

9 Young branchlets glabrescent; lamina 2-6.5 cm long; inflorescence

appressed puberulous; bracts ovate; style 3-3.5 mm long.19. S. wooroonooran

9. Young branchlets glabrous; lamina 7-15 cm long; inflorescence glabrous;

bracts oblong; style 4.5-5 mm long.8. S. glabra

10 Young branchlets, petiole or inflorescence pilose, villous or pubescent

with erect hairs. 11

10. Young branchlets glabrous or puberulous; petiole glabrous; inflorescence

glabrous, puberulous, or shortly appressed pubescent.16

11 Calyx densely pubescent outside.12

11. Calyx glabrous or sparsely pubescent outside.13

12 Secondary veins 6-12 pairs; inflorescence a few-flowered fascicle

.7. S. cyanocarpa var. pilosa

12. Secondary veins mostly 15-20 pairs; inflorescence spicate with many

flowers.5. S. crassiramifera

. 14. S. oresbia

. ampulliformis

Jessup, Symplocos in Australia 229

13 Sterile shoots rarely with cataphylls; twig hairs, reddish brown; midvein

above pilose or villous.14

13. Sterile shoots bearing cataphylls; twig hairs white, hyaline or pale ginger

brown; midvein glabrous above.15

14 Lamina in mature foliage bullate, margins more or less recurved.4. S. bullata

14. Lamina not bullate, margins flat.15. S. paucistaminea

15 Lamina sometimes bullate between veins, base mostly acutely cuneate,

margins often recurved; bracts c. 2 mm long,; calyx lobes obtuse or

rounded, entire or minutely denticulate.13. S. hylandii

15. Lamina not bullate, base rounded or cordate, margins plane; bracts 3-9

mm long; calyx lobes acute, ciliate.12. S. hayesii

16 Inflorescence axes usually branching above the base;.17

16. Inflorescence axes solitary or clustered, unbranched above the base

or flowers fasciculate or solitary. 21

17 Lamina margins crenulate and serrulate.17. S. stawellii

17. Lamina margins entire, rarely some denticulations evident.18

18 Stamens (12—)15—20.19

18. Stamens 30-70 . 20

19 Secondary veins mostly 7-9 pairs; basal ones not more acute and longer

than distal ones; bracts caducous before anthesis; petals 2-2.5 mm long;

disc and style glabrous.14. S. oresbia

19. Secondary veins mostly 4-6 pairs; basal ones often more acute and

longer than distal ones; bracts persistent to after anthesis; petals 3-3.7

mm long; disc and base of style puberulous.1. S. ampulliformis

20 Petiole 3-10 mm long; inflorescence glabrous; bracts oblong; stamens

mostly 30-50; style 4.5- mm long.9. S. glabra

20. Petiole 10-35 mm long; inflorescence sparsely appressed puberulous;

bracts broadly ovate or depressed ovate; stamens mostly 60-70; style

3.5-4 mm long.8. S. gittinsii

21 Inflorescence usually several clustered axillary spikes to 1.5 cm long;

bracts 1.2-1.5 mm long, sometimes denticulate, pubescent; style 2-2.5

mm long.10. S. graniticola

21. Inflorescence a fascicle, or flowers solitary; bracts 1.5 mm long or

more; style 2.5-3.5 mm long.22

22 Calyx lobes 1-1.4 mm long; petals rounded; filaments pilose

.7. S. cyanocarpa var. cyanocarpa

22. Calyx lobes 4-5.5 mm long; petals acute; filaments glabrous.3. S. boonjee

230

Austrobaileya 8(3): 225-251 (2011)

Key to Australian Symplocos taxa using fruiting specimens

1 Dry fruit ampulliform (flask-shaped).2

1. Dry fruit globose, depressed globose, or ellipsoid to subcylindrical,

rarely ovoid or obovoid.10

2 Young branchlets and petiole pilose or villous; infructescence spicate,

pilose or villous.3

2. Young branchlets and petiole glabrous, puberulous, or appressed

pubescent; infructescence paniculate, or racemose, glabrous or

appressed pubescent.6

3 Dry fruit 5-6 mm long, 3.5-4 mm wide.15. S. paucistaminea

3. Dry fruit 7-10 mm long, 4-5 mm wide.4

4 Secondary veins 15-20 pairs.6. S. crassiramifera

4. Secondary veins 7-12 pairs.5

5 Midvein densely villous on upper (adaxial) surface of lamina.4. S. bullata

5. Midvein glabrous on upper (adaxial) surface of lamina.13. S. hylandii

6 Lamina margins not entire, sometimes teeth scarcely evident on recurved

margins.7

6. Lamina margins entire.8

7 Young branchlets and petiole glabrous; lamina margins flat.18. S. thwaitesii

7. Young branchlets puberulous or pubescent; petiole glabrescent; lamina

margins recurved.16. S. puberula

8 Secondary veins mostly 4-6 pairs; basal pair more acute and longer; style

pubescent at base.1. S. ampulliformis

8. Secondary veins 7-12 pairs, not more acute and longer near base; style

glabrous.9

9 Young branchlets puberulous or pubescent, glabrescent; fruit 5.8-6.3

mm long, 4-5.5 mm wide.19. S. wooroonooran

9. Young branchlets glabrous; fruit 10-15 mm long; 7-9 mm wide.14. S. oresbia

10 Infructescence a glomerule or fruit solitary at each node.11

10. Infructescence spicate or racemose, branched or unbranched.12

11 Fruit glabrous; calyx lobes 4-5.5 mm long.3. S. boonjee

11. Fruit usually not glabrous; calyx lobes 1-1.4 mm long.7. S. cyanocarpa

12 Lamina margins entire or sometimes obscurely denticulate.13

12. Lamina margins serrate or dentate.16

13 Fruit mostly obovoid (Lord Howe Island endemic).5. S. candelabrum

13. Fruit globose, ovoid or ellipsoid (Australian mainland).14

14 Lamina length usually 3-6 times width.10. S. graniticola

14. Lamina length usually less than 3 times width.15

15 Petioles 10-35 mm long. Pedicels 0-0.2 mm long; fruit 6-7 mm long;

4-5 mm wide.

15. Petioles 3-10 mm long. Pedicels 0.2-2 mm long; fruit 9-11 mm long;

8-10 mm wide.

8. S. gittinsii

. 9. S. glabra

231

Jessup, Symplocos in Australia

16 Secondary and tertiary veins raised above (adaxially) (dried leaves).17

16. Secondary and tertiary veins not raised above (adaxially).19

17 Young branchlets, bud scales and infructescence axis glabrous.18. S. thwaitesii

17. Young branchlets, bud scales and infructescence axis not glabrous.18

18 Young branchlets sparsely appressed puberulous; petiole 6-25 mm long;

fruit 3.5-4.5 mm wide.17. S. stawellii

18. Young branchlets sparsely pilose with erect hairs; petiole 3-4 mm long;

fruit 6-7 mm wide.11. S. harroldii

19 Shoots usually without cataphylls; lamina 0.7-2 cm wide, sparsely

appressed puberulous on underside (abaxially), base acutely cuneate;

fruit 4-9 mm long.2. S. baeuerlenii

19. Shoots usually bearing scattered cataphylls; lamina 2.5-5.5 cm wide,

sparsely pilose on underside (abaxially), base rounded or cordate; fruit

15-20 mm long.12. S. hayesii

1. Symplocos ampulliformis C.T.White,

Proc. Roy. Soc. Queensland 50: 81 (1939).

Type: Queensland. Cook District: Mt

Spurgeon, September 1936, C.T.White 10581

(holo: BRI; iso: A n.v .; BM).

Illustrations : Hyland etal. (2010).

Shrub or small tree to 10m. Shoots sparsely

appressed puberulous, glabrescent; young

branchlets glabrous. Leaves chartaceous;

lamina oblanceolate or elliptic, 4.5-12 cm

long, 2-5 cm wide, acuminate at apex,

acute or attenuate at base, glabrous; margins

recurved, entire or nearly so; secondary

veins mostly 4-6 pairs, the basal pair often

longer and more acute; petiole 4-10 mm

long, glabrous. Inflorescence usually a cluster

of racemose axes to 2 cm long, glabrous or

sparsely appressed puberulous, sometimes

shortly branched near base; bracts and

bracteoles ovate, puberulous or ciliate,

persistent to anthesis, bracts c. 0.5 mm long,

bracteoles c. 0.4 mm long. Calyx limb 0.6-0.8

mm long, sparsely puberulous, glabrescent;

lobes depressed ovate or rounded, 0.3-0.5

mm long, glabrescent, margins ciliate. Petals

oblong or narrowly obovate, 2.5-3.5 mm

long, white, glabrous. Stamens (12—)15, 1.5-4

mm long. Disc puberulous. Style c. 2.5 mm

long, puberulous at base. Fruit ampulliform

to narrowly ovoid, 10-15 mm long, 5-6 mm

wide, blue to blue-black or purple-black. Fig.

1A-C.

Additional selected specimens examined: Queensland.

Cook District: N.P.R. 133, summit of Mt Sorrow walking

track. Cape Tribulation, Nov 2000, Ford AF2524 et al.

(BRI); S.F.R.144, Parish of Whypalla, Bower Bird L.A.,

Dec 1987, Hyland 13459 (BRI, CNS); loc. cit., May 1989,

Gray 5046 (BRI, CNS); S.F.R.144, Dasyurid L.A., Apr

1980, Gray 1682 (BRI, CNS); S.F.R.144, Chowchilla

L.A., Jul 1982, Gray 2663 (BRI, CNS); Mt Lewis, SW

of Mossman, Aug 1957, Volck 1389 (BRI); S.F.R.143,

Parish of Riflemead, North Mary L.A., Dec 1984, Gray

3758 (BRI); S.F.R.143, South Mary L.A., Aug 1973,

Irvine 612 (BRI); S.F.R.143, Parish of Riflemead, South

Mary L.A., Gray 3362 (BRI, CNS); S.F.R.143, Parish of

Riflemead, Carbine L.A., Nov 1987, Hyland 25249RFK

(BRI); loc. cit., Jan 1985, Gray 3835 (BRI); S.F.R. 143,

Bushy L.A., Feb 1982, Hyland 11691( BRI); T.R.143,

Zarda L.A., Sep 1973, Hyland 2884RFK (BRI); Along

Falchetti Track to the Mitchell Bomber crash site on

the NE peak of Mt Bartle Frere, Jul 2001, Jago 6011

& Gandini (BRI); Upper E Mt Bartle Frere, Oct 1985,

Godwin C2900 (BRI).

Distribution and habitat: This species is

endemic to Queensland in the Wet Tropics and

occurs on Mt Sorrow near Cape Tribulation,

the Mt Windsor Tableland, Mt Carbine

Tableland, in the vicinity of Mount Lewis,

and on Mt Bartle Frere (Map 1), in upland and

montane notophyll and microphyll vineforest

on granite, granodiorite (all upper montane

localities) and phyllite substrates.

Phenology : Flowers are recorded from March

to May; fruit from January to July.

Notes: This species was treated as Symplocos

cochinchinensis var. montana (C.T.White)

Noot. (as S. cochinchinensis subsp. thwaitesii

var. montana) in Nooteboom (1982).

232

Conservation status : Symplocos ampulliformis

is currently listed as Near Threatened under the

NCA.

Map 1. Distribution of Symplocos ampulliformis

2. Symplocos baeuerlenii R.T.Baker, Proc.

Linn. Soc. New South Wales 27: 594, t. 28

(1902). Type: New South Wales. Tumbulgum,

Murwillumbah, s.dat ., W. Bauerlen s.n. (holo:

NSW; iso: MEL).

Illustrations : Floyd (1989: 396); Leiper et al.

(2008: 336).

Shrub or small tree to 10 m. Shoots and young

branchlets puberulous. Leaves coriaceous;

lamina narrowly elliptic or lanceolate, 1.5-8

cm long, 0.7-2 cm wide, acuminate at apex,

acute at base, glabrous or with a few hairs on

midvein and undersurface; margins mostly

serrate; secondary veins mostly 7-10 pairs;

petiole 2-4 mm long, puberulous, glabrescent.

Inflorescence racemose, to 0.6 cm or flowers

solitary, puberulous; pedicels to 1 mm long;

bracts and bracteoles puberulous, semi-

persistent, bracts acutely triangular, 0.8-1.2

mm long, bracteoles ovate, 0.3-0.5 mm

long. Calyx limb 0.7-1.1 mm long, glabrous;

lobes depressed ovate or rounded, 0.5-0.8

mm long, glabrous. Petals obovate, 2-3 mm

long, 1.4-1.6 mm wide, white, glabrous.

Stamens mostly 15-20, 1.5-3 mm long. Disc

pubescent. Style 1.5-2.5 mm long, pubescent

at base. Fruit ovoid, to 9 mm long, red.

Additional selected specimens examined : Queensland.

Moreton District: Lyrebird Ridge Road, Springbrook

Plateau, Dec 1993, Grimshaw G259 (BRI); Springbrook,

Aug 1945, Blake 15892 (BRI); loc. cit ., Sep 1929, Rudder

4022 (BRI, L); loc. cit., Sep 1929, White 6265 (BRI); loc.

Austrobaileya 8(3): 225-251 (2011)

cit., Aug 1930, White 7059 (BM, BRI); Track to ‘Best of

All’ Lookout, Springbrook, Jul 1977, McDonald 1946 &

Elsol (BRI); End of Currumbin Valley road, Mt Cougal

N.P., Nov 1992, Thomas 96 & Barry (BRI). New South

Wales. Mt Nardi, Goommbar S.F., Feb 1979, Floyd 1194

(BRI); Whian Whian S.F., Jun 1945, White 12758 (BRI);

loc. cit., Jun 1947, Hayes s.n. (BRI [AQ537353]); loc. cit.,

Nov 1949, White s.n. (BRI [AQ335709]); loc. cit., 1953-

1958, Webb & Tracey s.n. (BRI [AQ84368]); Minyon,

near Mullimbimby, Aug 1936, White 10480 (BRI);

Whian Whian, N of Lismore, Jun 1965, Smith 12303 et

al. (BRI); c. 6 km W of Minyon Falls, Whian Whian S.F.,

Jul 1975, Williams s.n. (BRI [AQ504389]).

Distribution and habitat : This species is

endemic to south-east Queensland and north¬

east New South Wales where it occurs from

the Macpherson Range near Springbrook,

south to Lismore (Map 2), in microphyll and

notophyll vineforests on substrates derived

from basalt and enriched rhyolite.

Map 2. Distribution of Symplocos baeuerlenii

Phenology : Flowers are recorded from July

to September; fruit in January and February.

Notes : This species was included under

Symplocos cochinchinensis var. montana

(as S. cochinchinensis subsp. thwaitesii var.

montana ) in Nooteboom (1982).

Conservation status : Symplocos baeuerlenii

is currently listed as Vulnerable under the

NCA.

3. Symplocos boonjee Jessup species nova

S. cyanocarpam simulans sed bracteis

longioribus (2-5 mm contra 1.5-1.7 mm

longis) bracteolis longioribus (2-3 mm contra

0.5-1 mm) sepalis longioribus (4-5.5 mm

contra 1-1.4 mm) petalis ovatis acutis et

filamentis glabris differt. Typus: Queensland.

Cook District: Wooroonooran National Park,

Stockwellia site, 10 July 2000, P.I.Forster

Jessup, Symplocos in Australia

233

PIF25915, R.Booth & R.Jensen (holo: BRI;

isotypi distribuendi).

Symplocos sp. (Boonjie B.P.Hyland 2753);

(Jessup 1994, 1997, 2002, 2007, 2010).

Illustrations : Cooper & Cooper (2004: 538),

under Symplocos sp. (Boonjee); Hyland et

al. (2010), under Symplocos sp. (Boonjie

B.P.Hyland 2753).

Shrub to 3 (-5) m. Shoots, cataphylls and

young branchlets glabrous; cataphylls usually

caducous. Leaves chartaceous; lamina

narrowly obovate or narrowly oblanceolate,

4.5-11 cm long, 1-3 cm wide, acuminate at

apex, acute or attenuate at base, glabrous;

margins usually entire, slightly recurved;

secondary veins mostly 5-7 pairs; petiole

3- 7 mm long, glabrous. Inflorescence a

solitary flower in axil of a leaf or cataphyll;

the axis c. 1.5 mm long with up to 9 bracts

or a fascicle with one open flower surrounded

by several buds; lower bracts broadly ovate to

depressed ovate, upper ones ovate to narrowly

ovate, 3.5-5 mm long, persistent, sparsely

appressed pubescent to nearly glabrous;

margins denticulate, ciliate; bracteoles 2-3

mm long, similar to upper bracts. Calyx limb

4- 5.5 mm long; lobes narrowly ovate, acute,

sparsely puberulous; margin denticulate,

ciliate; tip incurved. Petals narrowly ovate,

acute or obtuse, 3-3.5 mm long, 1.5-1.7 mm

wide, cream to white, glabrous. Stamens

35-40 in males, 25-30 in females, 0.8-3 mm

long, filaments glabrous. Disc pilose. Style

3-3.5 mm long (c. 0.5 mm in males), with a

few hairs at base, otherwise glabrous. Fruit

oblong or ellipsoidal, c. 17 mm long and 3.5

mm wide, glabrous, blue. Fig. 1D-F.

Additional selected specimens examined : Queensland.

Cook District: E side of Mulgrave River, Goldsborough

Valley, Jun 1995, Hunter JH3704 (BRI); Gadgarra S.F.,

S of Butchers Creek, Jun 1995, Hunter JH3904 (BRI);

S.F.R. 310, Swipers F.A., E of Malanda, Aug 1963,

Hyland AFO/2753 (BRI); Swipers Flat, Wooroonooran

N.P., Jul 1995, Forster PIF17138 & Figg (BRI); Boonjee

F.A., near Bartle Frere track, Nov 1988, Jessup GJM362

et al. (BRI); Russell River, Johnson s.n. (MEF46567);

Palmerston Track ridge, Feb 1995 Hunter JH1158 (BRI);

Breeden Property, Hughes Road, Topaz, Mar 1992,

Cooper & Cooper 221 (CNS); T.R. 1230, Bartle Frere,

Boonjee F.A., Sep 1992, Gray 5575 (CNS); S.F.R. 755,

Bartle Frere, Gosschalk F.A., Nov 1991, Hyland 14345

(CNS); R.926, Kauri Reserve, Dirran, Oct 1987, Dansie

20166 (CNS); N.P.R. 904, along track between Coolamon

Creek and Combo Creek, Oct 2000, FordAF2462 (BRI);

Wooroonooran N.P, Combo Creek, Nov 1998, Forster

PIF23990 et al. (BRI).

Distribution and habitat : This species is

endemic to the Wet Tropics of Queensland

where it is known from notophyll and

mesophyll vineforests mostly on the western

side of Wooroonooran N.P, and nearby forest

remnants (Map 3), on metamorphic or basalt

substrates.

Map 3. Distribution of Symplocos boonjee

Phenology: Flowers are recorded from June

to February; fruit in July and August.

Notes: Specimens of this species (e.g. Hyland

2753 ) were tentatively included in Symplocos

cyanocarpa C.T.White in Nooteboom (1982).

Etymology: The species epithet

commemorates a botanically rich but loosely

defined locality on the eastern edge of the

Atherton Tableland known as Boonjee or

Boonjie. For the sake of consistency the first

alternative spelling is used as it has been used

in the names Syzygium boonjee B.Hyland and

Bulbophyllum boonjee B.Gray & D.L. Jones (=

Adelopetalum boonjee (B.Gray & D.L.Jones)

D.L.Jones & M.A.Clem.).

Conservation status: Symplocos boonjee is

only known from Wooroonooran N.P. At a

few sites it has been recorded as common.

234

Additional collections and observations are

required to determine its area of occupancy

and population numbers.

4. Symplocos bullata Jessup species nova

5. paucistamineam simulans sed foliis

bullatis, marginibus recurvis, stylis longibus

(2-2.5 mm, contra 1.2-1.8 mm) et fructibus

majoribus (8-9 x 3.5-4 mm, contra 5-5.8 x

7.5-8 mm) differt. Typus: Queensland. Cook

District: State Forest Reserve 143, North

Mary Logging Area, 25 April 1982, B.Gray

2543 (holo: BRI; iso: BRI, CNS).

Symplocos sp. (North Mary B.Gray 2543);

(Jessup 2002, 2007, 2010).

Illustrations: Hyland et al. (2010), under

Symplocos sp. (North Mary B.Gray 2543).

Dioecious tree to 15 m. Shoots and young

branchlets yellowish brown or reddish-

brown to dark brown villous, cataphylls

rarely evident. Leaves thinly coriaceous

or chartaceous; lamina lanceolate, elliptic,

obovate or oblanceolate, 4-14 cm long, 1.2-5

cm wide, acuminate at apex, cuneate, acute

or scarcely obtusely so or obtuse, truncate

or rounded at base, mature leaves bullate

between main veins, glabrescent above, pilose

below; margins obscurely denticulate or

entire, recurved; secondary veins mostly 7-

12 pairs, usually depressed on upper surface;

petiole 4-11 mm long, pilose. Inflorescences

spicate, to 5 cm, pilose; bracts and bracteoles

ovate, densely reddish-brown pilose or

villous, persistent, bracts 1.8-2.5 mm long,

bracteoles 1.3-1.5 mm long. Calyx limb

1.2-1.5 mm long; lobes ovate, 1-1.2 mm long,

pubescent in middle near apex with reddish-

brown hairs, sometimes glabrescent. Petals

obovate, 3-3.5 mm long, 1—1.5 mm wide,

cream to white, mostly glabrous. Hypanthium

glabrous. Stamens (15-) 20-25 in males, 7-10

in females and apparently sterile, 1-2.5 mm

long. Disc glabrous. Style 2-2.5 mm long,

glabrous; females with peltate stigma. Fruit

broadly ovoid or ampulliform 8-9 mm long,

7.5-8 mm wide, colour unknown. Fig. 1G-I.

Additional selected specimens examined : Queensland.

Cook District: Daintree N.P., NW of Black Mountain,

May 1998, Forster PIF22903 et al. (BRI); Daintree

N.R, near end of Mt Lewis road, Nov 1988, Jessup et al.

GJM155 (BRI); S.F.R.143 Riflemead, North Mary L.A.,

Austrobaileya 8(3): 225-251 (2011)

Jun 1995, Gray 6226 (BRI); loc. c/7., Dec 1995, Gray

6479 (CNS photo); S.F.R. 652, Mt Fisher, May 1975,

Hyland3I75RFK (BRI, CNS).

Distribution and habitat: Endemic to the Wet

Tropics of Queensland where it occurs in the

Mt Lewis to Mt Carbine Tableland area west

of Mossman and with an outlier at Mt Fisher,

south of Millaa Millaa (Map 4). Grows in

montane microphyll and notophyll vineforests

on granite substrates.

Map 4. Distribution of Symplocos bullata

Phenology: Flowers are recorded from April

to June; fruit in December.

Etymology: The specific epithet is derived

from the Latin adjective bullatus (bullate,

blistered or puckered) and refers to the

appearance of the upper surface of the lamina

between the veins.

Conservation status: Symplocos bullata

is only known from the localities cited

above. Additional collections are required

to determine its area of occupancy and

population numbers.

235

Jessup, Symplocos in Australia

5. Symplocos candelabrum Brand in Engl.,

Pflanzenr. 6: 39 (1901); S. cochinchinensis

var. candelabrum (Brand) Noot., Brunonia

4: 323 (1982). Type: New South Wales. Lord

Howe Island, April 1898, J.H.Maiden s.n.

(holo: B [destroyed]; iso: BM, BRI).

Illustration : Green (1994).

Tree to 13 m, usually much smaller. Shoots

and young branchlets glabrous or glabrescent.

Leaves thinly coriaceous; lamina elliptic

or lanceolate, 6-16 cm long, 2-6 cm wide,

acute to acuminate at apex, attenuate at

base, glabrous; margins entire to remotely

serrulate-denticulate near apex, more or less

recurved near base; secondary veins mostly

8-11 pairs; petiole 20-30 mm long, glabrous.

Inflorescence predominately racemose to 6

cm, glabrous or with few very sparse hairs;

pedicels to 5 mm long; bracts and bracteoles

triangular, ciliate; bracts 1—1.5 mm long,

caducous before anthesis; bracteoles 0.8-1.5

mm long, often persisting a little longer

than the bracts. Hypanthium glabrous; calyx

limb 0.6-0.8 mm long; lobes broadly ovate,

glabrous with ciliate margin. Petals obovate,

mostly 6-7 mm long, 3-3.5 mm wide, white,

glabrous. Stamens mostly 30-40, 5-6 mm

long. Disc with a few hairs. Style 4-5 mm

long, pubescent at base. Lruit mostly obovoid,

10-16 mm long, 7-10 mm wide, blue.

Additional selected specimens examined : New South

Wales. Lord Howe Island: Mt Gower walking track,

Nov 2000, Conn 4320 (BRI); Halfway to saddle from

Erskines Creek on summit track, Nov 2000, Le Cussan

1056 (BRI); loc. eit., Feb 2004, Le Cussan 1375 (BRI);

Intermediate Hill track. Mar 2003, Le Cussan 1280

(BRI).

Distribution and habitat: Occurs only in

submontane forest at the southern part of

Lord Howe Island.

Phenology: Llowers are recorded from March

to June; Mature fruit in Lebruary.

Notes: This species was included under

Symplocos cochinchinensis var. candelabrum

(C.T.White)Noot. (as S. cochinchinensis subsp.

thwaitesii var. candelabrum ) in Nooteboom

(1982).

Conservation status: Symplocos candelabrum

is currently not listed under the EPBC Act

despite its highly restricted distribution.

6. Symplocos crassiramifera Noot., Leiden

Bot. Ser. 1: 186 (1975). Type: Queensland.

Cook District: Devil Devil Creek, a few

miles NE of Rumula and c. 8 miles [13.3km]

SSW of Mossman, 1 September 1948, L.S.

Smith 3965 (holo: BRI; iso: L).

Illustrations: Hyland et al. (2010).

Shrub or small tree to 8 m with few branches.

Shoots and young branchlets with caducous

cataphylls, pilose. Leaves coriaceous; lamina

narrowly elliptic to narrowly oblanceolate,

20-41 cm long, 3.5-8.5 cm wide, acuminate at

apex, acute or obtuse at base, glabrous above,

pilose below; margins revolute, remotely

denticulate to entire; secondary veins mostly

15-20 pairs; petiole 8-20 mm long, pilose.

Inflorescences comprising clustered spicate

axes to 15 cm, villous or pilose; bracts and

bracteoles ovate, brown pubescent, persistent,

bracts 2-2.5 mm long, bracteoles 1.7-2

mm long. Calyx limb 1.5-2 mm long, lobes

broadly ovate or rounded, pubescent. Petals

obovate, 4-4.5 mm long, 1.7-2 mm wide,

pink, glabrous. Stamens c. 25, 3-4 mm long.

Disc inconspicuous. Style 2.5-4 mm long,

pubescent at base. Lruit ampulliform, 7-10

mm long, 4-5 mm wide, pink.

Additional selected specimens examined : Queensland.

Cook District: Little Mossman L.A., S.F.143, Jul 1994,

Forster PIF15513 et al. (BRI); Rex Range, c. 1 km

from Mossman-Julatten Road, on forest track c. 9 km

NE of Julatten, Dec 1988, Jessup 880 et al. (BRI); Rex

Highway, near Julatten, Apr 1962, Webb & Tracey 5961

(BRI); The Gate, property of G. & S. Kurmond, 105

Salisbury Drive, Flin Creek, Julatten, Jan 2002, Ford

AF3260 (BRI); T.R. 66, Mt Lewis, May 1979, Moriarty

2603 (BRI, CNS); 7 km along Mt Lewis Road, Mt Lewis

F.R., Apr 2002, Booth 3092 & Jensen (BRI); T.R. 66, 8

km along Mt Lewis Road, Jun 1997, Forster P1F21233 et

al. (BRI); Mt Lewis forestry road, SW of Mossman, May

1972, Wrigley & Telford NQ140 (BRI, CANB); Rumula,

Apr 1962, McKee 9106 (BRI); Portion 203 Garioch, Jun

1973, Hyland 6758 (BRI, CNS).

Distribution and habitat: Endemic to the

Wet Tropics of Queensland where it is known

only from the vicinity of the headwaters of

the South Mossman River to Mt Lewis (Map

5), in notophyll and mesophyll vineforests on

granite and metamorphic substrates.

236

Map 5. Distribution of Symplocos crassiramifera

Phenology : Flowers are recorded from May

to July; fruit in September.

Conservation status : Symplocos crassiramifera

is currently listed as Vulnerable under the NCA.

7. Symplocos cyanocarpa C.T.White, Proc.

Roy. Soc. Queensland 47: 69 (1936). Type:

Queensland. Cook District: Mossman River,

4 February 1932, L.J.Brass 2050 (holo: A,

n.v.\ iso: BRI, MEL).

Shrub or small tree to 5 m. Shoots and young

branchlets pilose, glabrescent or glabrous,

sometimes with pubescent cataphylls.

Leaves chartaceous to coriaceous; lamina

Austrobaileya 8(3): 225-251 (2011)

oblanceolate, narrowly obovate or elliptic, 8-

23 cm long, 2.5-7 cm wide, mostly acuminate,

sometimes acute or obtuse at apex, acute or

attenuate at base, glabrous or with scattered

hairs on midvein below; margins remotely

serrate towards tip or entire; secondary

veins 7—12(—15) pairs; petiole 3-10 mm long,

glabrous or pilose. Inflorescence a fascicle

of mostly 2-10 flowers, axillary or ramal;

bracts and bracteoles ovate to triangular,

sometimes denticulate, pubescent, persistent,

bracts 1.5-1.7 mm long, bracteoles 1-1.2 mm

long. Calyx limb 1.5-2 mm long; lobes ovate,

often denticulate and ciliate, 1-1.4 mm long,

pubescent. Petals oblong to obovate, 2.5-4

mm long, 1.4-1.8 mm wide, white or cream-

pink, glabrous or with a few hairs, sometimes

ciliolate. Stamens 15-30, 2.5-3.5 mm long,

filaments pilose. Disc pilose. Style 2.5-3 mm

long, shorter and without stigma in males,

sparsely pilose. Fruit cylindrical to narrowly

ellipsoidal, 15-25 mm long, 6-10 mm wide,

glabrescent, blue to purple.

Distribution and habitat : Endemic to the

Wet Tropics of Queensland where it occurs

from Big Tableland, north of Mt Finnigan,

south to the Mossman River, in mesophyll

and notophyll vine forest.

Two varieties are recognised.

Key to varieties

Young branchlets, petiole and lamina glabrous.var. cyanocarpa

Young branchlets, petiole and midvein of lamina pilose below.var. pilosa

7a. Symplocos cyanocarpa var. cyanocarpa

Shrub or small tree to 5 m. Shoots, young

branchlets, petiole and lamina glabrous

or nearly so. Bracts, bracteoles, and calyx

appressed puberulous. Fig. 2A-C.

Illustrations : Hyland et al. (2010).

Additional selected specimens examined : Queensland.

Cook District: Mt Finnigan, W slopes, Sep 1948, Brass

20043 (BRI); Cedar Bay N.P., W slopes of Mt Finnigan,

Oct 1999, Forster PIF25014 & Booth (BRI); Mt Misery,

Jun 1992, Forster PIF10751 et al. (BRI); Daintree N.P.,

Cedar Bay section. Gap Creek, Nov 2000, Forster

PIF26464 et al. (BRI); Cedar Bay N.P, Jul 1994, Forster

PIF15585 et al. (BRI); N of Daintree River, on road to

Cape Tribulation, Nov 1985, Tucker 37 (BRI); Palm

Road, 3 km W of Alexander Bay, Dec 2001, Forster

PIF27907 et al. (BRI); Whyanbeel, Dec 1978, Gray 1181

(BRI, L); loc. cit ., Dec 1978, Gray 1187 (BRI); c. 2 km

along Whyanbeel - Daintree track, from Whyanbeel

Creek, Jan 1986, Guymer 2060 (BRI); Near Cooper

Creek, Aug 1987, Sankowsky 639 & Sankowsky (BRI);

Mossman Gorge, W of Mossman, Jun 1972, Wrigley &

TelfordNQ1137 (BRI).

Distribution and habitat : Endemic to the

Wet Tropics of Queensland where it occurs

from the vicinity of Mt Finnigan to south of

Mossman (Map 6) in mesophyll and notophyll

vineforest on granitic and metamorphic

derived soils.

Jessup, Symplocos in Australia 237

Fig. 2. Symplocos cyanocarpa var. cyanocarpa. A. portion of stem with inflorescences and leaf base x 3. B. flower x

8. C. fruit x 2. A-C from Forster PIF10751 et al. (BRI). S. cyanocarpa var. pilosa. D. inflorescence and leaf base x

4, Hyland 10582 (BRI). S. glabra. E. branchlet with fruit x 0.8, Hyland 11798 (BRI). S. hayesii. F. inflorescence with

flowers x 4, Hyland 14628 (CNS). G. fruit x 2, Smith 10839 (BRI). H. leaf abaxial surface x 1, Smith 10839 (BRI). Del.

W. Smith.

238

Map 6. Distribution of Symplocos cyanocarpa var.

cyanocarpa

Phenology : Flowers are recorded from July

to January; fruit from November to March.

Conservation status : Symplocos cyanocarpa

var. cyanocarpa occurs mostly in National

Parks and other reserves throughout its range

and is not considered to be threatened.

7b. Symplocos cyanocarpa var. pilosa Jessup

varietas nova, differt a var. cyanocarpa

ramulis juvenibus, petiolo et costa abaxialiter

pilosis. Typus: Queensland. Cook District:

Timber Reserve 146, Tableland Logging

Area, 11 September 1980, B.P.Hyland 10582

(holo: BRI; iso: CNS).

Symplocos sp. (Big Tableland B.P.Hyland

10582); (Jessup 1997, 2002, 2007, 2010).

Illustrations : Hyland et al. (2010), under

Symplocos sp. (Big Tableland B.P.Hyland

10582).

Shrub or small tree to 3 m. Shoots, young

branchlets, petiole and main veins of leaves

on underside pilose with scattered erect hairs,

glabrous above. Bracts, bracteoles and calyx

pilose with more or less erect hairs. Fig. 2D.

Additional specimens examined : Queensland. Cook

District: T.R. 146, Tableland L.A., Jul 1975, Hyland

8324 (BRI, CNS); Hartley Creek at end of Mt Hartley

track, Oct 1996, Ford 1774 (BRI); Along track from

Hartley Creek to Mt Hartley, Oct 1996, Ford 1782 (BRI);

Mt Hartley, western slopes, T.R. 165, Jul 1995, Forster

PIF17296 & Figg (BRI).

Austrobaileya 8(3): 225-251 (2011)

Distribution and habitat : Endemic to the

Wet Tropics of Queensland where it occurs on

Big Tableland and Mt Hartley, northwest of

Cedar Bay (Map 7) in notophyll vineforest on

granite substrates.

Map 7. Distribution of Symplocos cyanocarpa var.

pilosa

Phenology : Flowers are recorded in July,

September and October and very immature

fruit in October.

Etymology : The varietal epithet is formed

from the Latin word pilosus (covered with

short, weak, thin hairs) and refers to the

indumentum on the young branchlets, petiole

and adaxial surface of the midvein.

Conservation status : Symplocos cyanocarpa

var. pilosa is only known from the localities

cited above. Additional collections are

required to determine its area of occupancy

and population numbers.

8. Symplocos gittinsii Jessup nom. et

stat. nov.; S. cochinchinensis var. gittonsii

Noot., Leiden Bot. Ser. 1: 162 (1975). Type:

Queensland. North Kennedy District: Mt

Spec Range, May 1962, C.H.Gittins 494 (holo:

BRI; iso: L).

Illustrations : Nooteboom (1982: Fig. 11);

Cooper & Cooper (2004: 537); Hyland et al.

(2010), all under Symplocos cochinchinensis

var. gittonsii

Tree to 30 m. Shoots and young branchlets

glabrous; cataphylls glabrous and ciliate.

Leaves coriaceous; lamina obovate,

239

Jessup, Symplocos in Australia

sometimes elliptic, 8-18 cm long, 3.5-9.5

cm wide, acuminate, acute or obtuse at apex,

acute or attenuate at base, glabrous; margins

entire, revolute; secondary veins mostly 7-

11 pairs; petiole 10-35 mm long, glabrous.

Inflorescence axes branched, to 13 cm long,

spicate, sparsely appressed puberulous; bracts

and bracteoles broadly or depressed ovate,

glabrous with ciliate margin, persistent to

after anthesis, sometimes to fruiting, bracts

0.8-1.5 mm long, bracteoles 0.7-1 mm long.

Calyx limb 0.7-1.5 mm long, glabrous; lobes

depressed obovate, mostly glabrous with

ciliate margin. Petals broadly obovate, 4-6

mm long, 3.5-4.5 mm wide, white, glabrous.

Stamens 60-70, 2.5-6 mm long. Disc slightly

raised, 5-lobed, glabrous. Ovary 3-locular,

with c. 3 ovules per locule. Style 4 mm long,

glabrous. Fruit ellipsoidal or ovoid, 6-7 mm

long, 4-5 mm wide, blue to black.

Additional selected specimens examined : Queensland.

Cook District: T.R. 30, Mt Lewis area, Oct 1967,

Hyland 1048 (BRI); Ridge between McKinnon &

Wrights Creeks, southwest of Edmonton, Jun 1996,

Forster PIF19263 et al. (BRI); S.F.R. 310, Gadgarra,

Feb 1981, Bragg s.n. (BRI [AQ874875]); T.R. 1230,

Boonjie F.A., May 1977, Gray 533 (BRI, MEF); S.F.R.

310, Gadgarra, Windin F.A., May 1990, Hyland 14017

(BRI); S.F.R. 310, Windin F.A., Jan 1978, Gray 850

(BRI, MEF); Topaz to Boonjee Road, May 1987, Starkey

74 (BRI); On Topaz Road, Dec 1985, Sankowsky 459 &

Sankowsky (BRI); Topaz Road, Nov 1992, Jessup 905

(BRI); Palmerston Highway, 1 km south-east of Beatrice

River, Jan 1993, Bean 5401 (BRI); Wooroonooran N.P.,

Palmerston section, Dec 1999, Forster PIF25209 (BRI);

S.F. 756, Mt Father Clancy, May 2000, Forster PIF25726

& Booth (BRI); Portion 297, Parish of Dirran, Apr 1984,

Gray 3992 (BRI, CNS). North Kennedy District: 4.5

km south along Culpa Road, S.F. 605 Koombooloomba,

Culpa F.A., May 2002, Forster PIF28705 & Booth

(BRI); Mt Spec near Paluma village. May 1985,

Jackes 14 (BRI); Paluma, May 1980, Horton s.n. (BRI

[AQ343157]); Paluma, Sep 1990, Gumming 10650 (BRI);

Mt Spec N.P, Paluma, behind Rangers barracks. May

1989, Fell DF1851 (BRI). South Kennedy District:

S.F. 652 Cawley, near Mt Macartney, Jun 1995, Forster

PIF16714 & Tucker (BRI); Clarke Range, 13.7km from

Eungella township, Eungella N.P, Apr 1991, Forster

PIF8057 & McDonald (BRI); Roadside c. 15 km N of

Eungella township on Dalrymple Road, Nov 1989,

McDonald 4465 (BRI); Dalrymple Heights, Eungella

Range, Fredrickson’s Farm, May 1987, Champion 227

(BRI).

Distribution and habitat : Endemic to

Queensland where it occurs from Mt Lewis

south to Culpa L.A. in the Wet Tropics, then

disjunct to Paluma and the Clarke Range near

Eungella (Map 8). Grows in upland notophyll

or mesophyll vine forest on diverse substrates

(basalts, granites, metamorphics, rhyolites).

Map 8. Distribution of Symplocos gittinsii

Phenology: Flowers are recorded in April and

May; fruit from November to January.

Notes: This species was treated as Symplocos

cochinchinensis var. gittonsii Noot. (as

S. cochinchinensis subsp. thwaitesii var.

gittonsii ) in Nooteboom (1982).

Etymology: The name commemorates

Clifford Halliday Gittins (1904-1995),

an amateur botanical collector of mostly

Queensland plant specimens.

Conservation status: Symplocos gittinsii

occurs in several National Parks and other

reserves throughout its range and is not

considered to be threatened.

9. Symplocos glabra Jessup nom. et stat.

nov.; S. cochinchinensis var. glaberrima

Noot., Brunonia 4: 319-320 (1982). Type:

Queensland. Cook District: State Forest

Reserve 194, Compartment 54, 17 March

1971, G.C. Stocker 622 (holo: L; iso: BRI,

CNS, MEL).

Illustrations: Nooteboom (1982, Fig. 11);

Hyland et al. (2010), both under Symplocos

cochinchinensis var. glaberrima.

Small tree to 9 m. Shoots and young

branchlets glabrous. Leaves coriaceous;

lamina oblanceolate or narrowly obovate,

rarely elliptic, 7-15 cm long, 2-7 cm wide,

acuminate, acute or obtuse at apex, acute or

attenuate at base, glabrous; margins entire,

240

Austrobaileya 8(3): 225-251 (2011)

revolute; secondary veins mostly 8-12 pairs;

petiole 3-10 mm long, glabrous. Inflorescence

axes 2 or 3, racemose, to 3 cm long, glabrous,

branching near base, sometimes single;

pedicels 0.2-2 mm long; bracts and bracteoles

glabrous but ciliate, caducous or persistent to

anthesis, bracts broadly ovate or oblong, 1.2-

2.5 mm long, bracteoles ovate, 0.7-1.2 mm

long. Calyx limb 1.2-1.5 mm long, glabrous;

lobes depressed obovate, glabrous with ciliate

margin. Petals narrowly obovate, 4.5-6 mm

long, 2-3 mm wide, cream to white, glabrous.

Stamens mostly 30-50, 2-5 mm long. Disc 5-

lobed, sparsely puberulous. Style 4.5-5 mm

long, glabrous. Fruit subglobose or ellipsoidal,

8-11 mm long, (5-) 8-10 mm wide, blue-black

to purple-black. Fig. 2E.

Additional selected specimens examined : Queensland.

Cook District: V.C.L. Noah, Mt Hemmant, Oct 1975,

Hyland RFK3339 (BRI); Mt Windsor F.R, c. 5.7 km

from Spencer Creek Forestry barracks, Jun 2004, Ford

4341 & Hewett (BRI); S.F.R. 144 Whypalla, Bower Bird

L.A., Dec 1987, Hyland 13454 (BRI); Mt Spurgeon, W of

Mossman, Sep 1936, White 10557 (BRI); Mt Bellenden

Ker, Jun 1969, Smith 14682B (BRI); Wooroonooran

N.R, Mt Bellenden Ker summit area, Dec 2000, Forster

PIF26510 et al. (BRI); loc. cit ., Dec 2001, Forster

PIF27945 et al. (BRI); Mt Bellenden Ker, c. 0.5 miles

[0.8 km] SW of centre peak, Jun 1969, Smith 14638 &

14638a (BRI); Summit of Centre Peak, Bellenden Ker,

Nov 1972, Webb & Tracey 11920 (BRI); S.F.R. 194

Herberton, c. 10 miles [16 km] S of Atherton, May 1964,

Rudder s.n. (BRI [AQ84400], L); Bellenden Ker, Jan

1923, White s.n. (BRI [AQ84393]); S.F.R. 310 Bellenden

Ker Range, Jan 1980, Hyland 10205 (BRI); Mt Bellenden

Ker, Feb 1982, Irvine 2210 (BRI, CNS); Mt Bellenden

Ker, Dec 2000, Jago 5779 & Forster (BRI); Summit of

Mt Bellenden Ker, Aug 1971, Hyland5312 (BRI); S.F.R.

755 Gosschalk L.A., Jun 1982, Hyland 11798 (BRI); 100

m W of summit of NW peak of Bartle Frere, Jul 1995,

Hunter JH4461 (BRI); Wooroonooran N.P., summit of

South Peak, Oct 1997, Forster PIF21771 et al. (BRI);

Mt Bartle Frere, Jun 1961, Martin & Hyland AFO/1890

(BRI); S.F.R. 194 Barron, Jan 1983, Gray 2909 (BRI);

S.F.R. 251, Ravenshoe - Tully Falls Road, Aug 1968,

Hyland 1737 (BRI).

Distribution and habitat: Endemic to the Wet

Tropics of Queensland where it occurs at Mt

Hemmant, Mt Windsor Tableland, Mt Carbine

Tableland, on the Lamb Range, Bellenden Ker,

Cardwell and Herberton Ranges (Map 9) in

simple to complex microphyll and notophyll

vineforests on granites (upper montane

localities) and rhyolites.

Phenology: Flowers are recorded from

December to May; fruit from December to

March.

Note: The superlative epithet ‘glaberrima’ is

not available for use at the rank of species.

This species was treated as Symplocos

cochinchinensis var. glaberrima Noot. (as

S. cochinchinensis subsp. thwaitesii var.

glaberrima) in Nooteboom (1982).

Conservation status: Symplocos glabra

occurs in several National Parks and other

reserves in the Wet Tropics and is not

considered to be threatened.

Etymology: The specific epithet is derived

from the Latin adjective glaber (glabrous) and

refers to the lack of indumentum on all parts

of the plant.

10. Symplocos graniticola Jessup,

Austrobaileya 4: 9 (1993) Type: Queensland.

Cook District: Mt Lewis, 24 December 1986,

G. Sankowsky 598 & N. Sankowsky (holo:

BRI).

Symplocos sp. ‘Mt Lewis’; (Thomas &

McDonald 1989).

Illustrations: Jessup (1993: 10); Hyland et al.

( 2010 ).

Shrub to 3 m. Shoots and cataphylls with

very sparse appressed hairs, glabrescent;

cataphylls caducous. Twigs glabrous. Leaves

coriaceous; lamina narrowly obovate or

narrowly oblanceolate, obtuse or shortly

acuminate, 9-23 cm long, 2-7.5 cm wide,

glabrous; margins entire, recurved, sometimes

remotely serrulate or denticulate, acute or

241

Jessup, Symplocos in Australia

attenuate at base; secondary veins mostly

6-12 pairs; petiole 8-20 mm long, glabrous.

Inflorescence axes clustered, spicate, to 1.5

cm, puberulous; bracts and bracteoles ovate,

slightly keeled, sometimes denticulate,

pubescent, persistent, bracts 1.2-1.5 mm long,

bracteoles 1—1.3 mm long. Calyx limb tube 1-

1.3 mm long, glabrous; limb 1-1.2 mm long;

lobes ovate or depressed ovate, 0.8-1 mm

long, sparsely puberulous to nearly glabrous

and with a ciliate margin. Petals oblong to

obovate, 2.5-3.5 mm long, 1.5-2 mm wide,

white, glabrous. Stamens 25-30, 2.5-4 mm

long, filaments glabrous. Disc pilose. Style

2.5 mm long, pilose. Fruit ovoid or ellipsoid,

10-18 mm long, 6-12 mm wide, black.

Additional selected specimens examined : Queensland.

Cook District: Platypus Creek, Jan 1936, Flecker 1216a

(CNS); Pinnacle Rock track, Jun 1992, Forster PIF10716

etal. (BRI); North Mary L.A., S.F. 143,31.5 km along Mt

Lewis road, Jun 1995, Forster PIF16774A (BRI); S.F.R.

143, South Mary L.A., May 1976, Hyland 8773 (BRI,

CNS); 32.6 km along Mt Lewis Road from Mossman

- Mt Molloy Road, Dec 1989, Jessup GJD3359 et al.

(BRI). S.F.R. 143, Kanawarra, Carbine Tableland, Nov

1995, Gray 6400 (BRI); Mt Lewis F.R., off Mt Lewis

Road, Jan 2008, FordAF5188 et al. (BRI).

Distribution and habitat : This species is

endemic to the Wet Tropics of Queensland

where it is known from Mt Spurgeon to Mt

Lewis, Platypus Creek and Pinnacle Rock

near Mossman (Map 10). It grows in simple

microphyll and notophyll vineforests on

granite or granodiorite substrates.

Map 10. Distribution of Symplocos graniticola

Phenology : Flowers are recorded from

December to May; green fruit from June to

September, mature fruit in November.

Conservation status: Symplocos graniticola

is currently listed as Vulnerable under the

NCA.

11. Symplocos harroldii Jessup,

Austrobaileya 4: 7 (1993). Type: Queensland.

Moreton District: Moggill State Forest, just

downstream of road crossing Pullen Creek, 16

November 1980, L.W.Jessup 268 (holo: BRI;

iso: K, NSW).

Symplocos sp. 1.; Stanley & Ross (1986).

Illustrations: Jessup (1993: 8); Leiper et al.

(2008: 336).

Shrub or small tree to 6 m. Shoots and twigs

pilose. Leaves coriaceous; lamina narrowly

obovate or oblanceolate, 3-10 cm long, 0.8-

3.5 cm wide, acuminate at apex, acute at base,

glabrous or with a few hairs on underside,

particularly on midvein; margins denticulate

to serrate towards tip; secondary veins mostly

6-12 pairs; petiole 3-4 mm long, usually

with scattered hairs. Inflorescence racemose,

to 2 cm, puberulous; pedicels 0.2-2 mm

long; bracts and bracteoles ciliate, semi-

persistent, bracts broadly triangular, 0.7-1

mm long, bracteoles triangular, 0.3-0.4 mm

long. Hypanthium glabrous. Calyx lobes

broadly ovate, 0.5-0.7 mm long, glabrous,

often with a ciliate margin. Petals obovate to

oblong, 3-4 mm long, 2-2.5 mm wide, white,

glabrous. Stamens 30-40, 1.8-5 mm long.

Disc puberulous. Ovary 3-locular with up

to 4 ovules per locule. Style 3.5-4 mm long,

glabrous, dilated at stigma. Fruit narrowly

ovoid or ellipsoid, 8-10 mm long, 6-7 mm

wide, red.

Additional selected specimens examined: Queensland.

Wide Bay District: Tinana Creek, 7 km E Tiaro, Dec

1992, Forster PIF12503 & Smyrell (BRI); Tinana

Creek, 7 km ENE of Tiaro, Mar 1992, Smyrell s.n. (BRI

[AQ541786]); S.F. 502, Veteran L.A., Gympie, Sep 1992,

Thomas 8 & Barry (BRI); 1 km ENE of Timberwah

[Tinbeerwah] Hall, junction forestry road & tributary of

Wooroi Creek, S.F. 959, Sep 1992, Thomas 1 & Barry

(BRI); Gum Tree Drive off Cooroy - Tewantin Road,

Sep 1992, Thomas 5 & Barry (BRI); Noosa N.P., off

Viewland Drive, Jan 1993, Thomas & Gayle s.n. (BRI

[AQ636338]); Panorama Drive via Duke Road, 8 km SE

of Cooroy, Apr 1993, Bean 5928 (BRI); N of Ringtail

242

Creek Road, Noosa Shire, Portion 8V, near S.F. 997,

Oct 1994, Thomas 102 (BRI); Noosa Shire hinterland

N of Collwood Road, near Lake McDonald, Lot 3 Rp

800331 Parish ofTewantin, Sep 1995, Thomas 33 (BRI);

Compartment 1, Ringtail S.F., NE of Pomona, Sep

1997, Bean 12412 (BRI); Doonan Wetland Conservation

Area, Jan 2007, Thomas MS112 (BRI); Maroochy Shire

Bushland Conservation Reserve, Doonan Creek East

(BCR), Jul 2007, Thomas MS139 (BRI); Near Doonan

Creek, Feb 2002, Thomas 420M (BRI). Moreton

District: 650 Campbells Pocket Road, Caboolture, Jan

2006, Weber s.n. (BRI [AQ735727]); 0.5 km SW of

McAfees Lookout, S.F. 309, Feb 1991, Forster PIF7790

& Bird (BRI); Dillon Road, The Gap c. 0.5 km from

intersection with Payne Road, Jul 1999, Harris WKH124

& Thompson (BRI); Greenford Street, Chapel Hill, Sep

1999, Stephens s.n. (BRI [AQ664972]); Cubberla Creek,

Fig Tree Pocket, Sep 1998, Stephens 2 (BRI); Daisy Hill

S.F., Logan City, Dec 1993, Thompson MOR323 (BRI);

Hinze Dam, Mar 2007, Jinks s.n. (BRI [AQ784176]);

Hinze Dam, c. 3.5 km SW of Mudgeeraba, Apr 2007,

Jinks DBA { BRI).

Distribution and habitat : This species is

endemic to southeast Queensland. Collections

of this species made since it was described in

1993 have extended the known range from

Tinana Creek near Tiaro to the Hinze Dam

near Mudgeeraba (Map 11).

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Map 11. Distribution of Symplocos haroldii

Austrobaileya 8(3): 225-251 (2011)

Phenology : Flowers are recorded from

September to November; fruit in January.

Note : The phrase in the description in Jessup

(1993) stated ‘peduncles 0.2-2 mm long’; this

should have read ‘pedicels 0.2-2 mm long’.

Conservation status : Symplocos harroldii is

currently listed as Near Threatened under the

NCA.

12. Symplocos hayesii C.T.White & Francis,

Proc. Roy. Soc. Queensland 33: 158, t.2 (1921).

Type: Queensland. Cook District: Glenallan

[Glen Allyn], Malanda, s.dat., H.C.Hayes

(holo: BRI [AQ23234]).

Illustrations: Cooper & Cooper (2004: 538);

Hyland et al. (2010).

Shrub or small tree to 2 m. Shoots and young

branchlets with persistent cataphylls, pilose.

Leaves chartaceous; lamina narrowly ovate

or narrowly obovate or elliptic, 6-12 cm long,

2.5- 5.5 cm wide, acuminate at tip, rounded

or cordate at base, glabrous except midvein

on underside; margins serrate; secondary

veins mostly 7-10 pairs; petiole 3-7 mm long,

pilose. Inflorescence spicate, to 1.5 cm, or a

fascicle, pilose; bracts and bracteoles sparsely

pilose and ciliate, persistent, bracts ovate to

narrowly triangular, 3-9 mm long, bracteoles

ovate, 2-3 mm long. Calyx limb 1-1.5 mm

long, glabrous; lobes ovate, acute, glabrous or

with a few hairs, margin often ciliate. Petals

ovate or oblong, denticulate, 2.2-2.5 mm

long, white, glabrous. Stamens mostly 12-15,

as long as petals in males, shorter in females.

Disc pilose. Style 1.5-2 mm long in females,

glabrous, stigma peltate. Fruit narrowly ovoid

to ellipsoidal, usually curved, 15-20 mm long,

3.5- 5 mm wide; blue to black, mesocarp thin,

fleshy; endocarp smooth; seed 1. Fig. 2F-H.

Additional selected specimens examined : Queensland.

Cook District: S.F. 185 Danbulla, 1 km N of Hoop Pine

Triangle on C road, Jul 1994, Forster PIF15609 et al.

(BRI); Danbulla, 12 miles [19.3 km] NE of Atherton, Jun

1949, Smith 4181 (BRI); Yungaburra, near Lake Barrine,

Blake 9602 (BRI, CANB); near Lamond’s Hill, 7.5 miles

[12 km] SE of Malanda, Sep 1959, Smith 10839 (BRI);

S.F.R. 310, Gadgarra, Boonjee L.A., Jan 1993, Hyland

14628 (CNS); T.R. 1230, Boonjee L.A., Gray 1246 (BRI);

S.F. 756, Mt Father Clancy, May 2000, Forster PIF25720

& Booth (BRI); Vacant Crown Land, Bartle Frere, E of

Glen Allyn Trig., Webb & Tracey 5795 (BRI). North

Kennedy District: tributary of Chunum Creek, May

2005, Wannan 3986 & Sinclair (BRI); Tableland L.A.,

243

Jessup, Symplocos in Australia

0.5 km S of Koolmoon Creek near junction of Ebony

Road and Tully Falls Road, 6 km NW of Tully Falls, Oct

1998, Jessup GJM2670 et al. (BRI).

Distribution and habitat : Endemic to the

Wet Tropics of Queensland where it occurs

from Tinaroo Range to Koolmoon Creek,

southeast of Ravenshoe (Map 12). Grows

in mesophyll and notophyll vineforests on

substrates derived from basalts, granites and

metamorphics.

Phenology: Flowers are recorded from

December to February; fruit from May to

September.

Conservation status: Symplocos hayesii

occurs in several National Parks and other

reserves on the Atherton Tableland is not

considered to be threatened.

13. Symplocos hylandii Noot., Brunonia 4:

323 (1982). Type: Queensland. Cook District:

State Forest Reserve 755 Boonjee, 21 March

1968, B.HylandRFK1395 (holo: CNS).

Illustrations: Hyland et al. (2010).

Shrub or small tree to 3 m. Shoots and

young branchlets with persistent cataphylls,

pilose. Feaves coriaceous; lamina narrowly

obovate, 10-16 cm long, 1.7-3.5 cm wide,

often bullate and brittle, acuminate at apex,

acute at base, glabrous or with a few hairs

on underside, particularly midvein; margins

serrate; secondary veins mostly 5-12 pairs,

raised below, sunken above; petiole 3-5 mm

long, pilose. Inflorescence spicate, to 1.5 cm,

pilose; bracts and bracteoles ovate, sparsely

pilose and ciliate, persistent, bracts, c. 2 mm

long, bracteoles, c. 1 mm long. Calyx limb

1.3-1.7 mm long, glabrous; lobes ovate or

broadly oblong, puberulous or nearly glabrous,

margins entire or minutely denticulate. Petals

ovate or oblong, c. 3 mm long, white, glabrous.

Stamens 10-15. Disc pulvinate, glabrous. Fruit

ampulliform to ovoid, (7-) 10-12 mm long,

8-9 mm wide, blue, mesocarp thin, endocarp

shallowly grooved; seed 1. Fig. 3A,B-

Additional selected specimens examined : Queensland.

Cook District: Mt Bellenden Ker, c. 7/8 mile [1.5 km]

SE of centre peak, Jun 1969, Smith 14717 (BRI); East

Mulgrave River Falls, May 2002, Jago 6212 (BRI);

T.R. 1230, Stockwellia Track, 4.9 km from Lamins Hill

turnoff (headwaters of Russell River), Aug 1989, Bostock

912 & Guymer (BRI); Swipers Flat, WooroonooranN.R,

Jul 1995, ForsterP1F17136& F/gg/BRI); Wooroonooran

N.R, track to Chuck Lunga Creek, E of Boonjee, Jun 1996,

Forster PIF19284 & Tucker (BRI); Wooroonooran N.R,

lower reaches of Russell River Track, Jan 2007, Ford

AF4913 & Metcalfe (BRI); N.P.R. 904 Wooroonooran,

Coolamon Creek, Oct 2001, Ford AF3047 et al. (BRI);

Coolamon Creek, end of Towalla Road, Wooroonooran

N.R, Oct 2001, Cooper WWC1606 (BRI).

Distribution and habitat: This species is

endemic to the Wet Tropics of Queensland

where it occurs from Mt Bellenden Ker

to near Millaa Millaa (Map 13) mostly in

mesophyll vineforests on substrates derived

from granites and metamorphics.

Phenology: Flowers are recorded in May;

fruit from October to January.

Conservation status: Symplocos hylandii is

located mostly in Wooroonooran N.P and is

not considered to be threatened.

244

14. Symplocos oresbia Jessup species nova

S. wooroonooran simulans sed innovationibus

et axibus inflorescentiarum glabris, bracteis

et bracteolis ante anthesin caducis, petalis

2-2.5 mm longis (non 4-5 mm) et 1-1.3 mm

latis (non 1.4-2.5), staminibus paucioribus

(16-19 non 30-35), stylis brevioribus (1.5-1.8

mm non 3-3.5 mm) et fructibus amplissimus

(10-15 x 7-9 mm non 5.8-6.3 x 4-5.5 mm)

differt. Typus: Queensland. Cook District:

Summit area of Thornton Peak, 12 November

1973, TG.Hartley 14045 (holo: BRI; iso:

CANB).

Symplocos sp. (Mt Finnigan L. J.Brass 20129);

(Jessup 1994, 1997, 2002, 2007, 2010).

Illustrations : Hyland et al. (2010), under

Symplocos sp. (Mt Finnigan L. J.Brass

20129).

Shrub or small tree to 8 m. Shoots and young

branchlets glabrous. Leaves coriaceous;

lamina elliptic, 3-8 cm long, 1-3 cm wide,

acuminate at apex, acute or attenuate at base,

glabrous; margins entire, revolute; secondary

veins mostly 7-9 pairs; petiole 3-7(-12) mm

long, glabrous. Inflorescence axes 2 or 3,

racemose, to 3 cm long, glabrous, branching

Austrobaileya 8(3): 225-251 (2011)

near base, sometimes single; pedicels 0-0.2

mm long, bracts broadly ovate, 1-1.2 mm long,

bracteoles ovate, 0.5-0.6 mm long, glabrous

or ciliate, caducous before anthesis; pedicel

0.2-1.5 mm long. Calyx limb 1-1.5 mm long,

lobes depressed ovate or rounded, 0.5-0.7

mm long, glabrous with ciliate margin. Petals

obovate, 2-2.5 mm long, 1-1.3 mm wide,

rounded, ?white, glabrous. Stamens 16-19;

filaments glabrous. Disc glabrous. Style

1.5-1.8 mm long, glabrous; stigma indistinct.

Fruit ampulliform, 7.8-8.5 mm long, 4.6-5.5

mm wide, colour unknown. Fig. 3C,D.

Additional selected specimens examined : Queensland.

Cook District: Mt Finnigan, Apr 1948, Brass 20129

(BRI, CANB); Cedar Bay N.P., Mt Finnigan summit

area, campsite on Horans Creek, Oct 1999, Forster

PIF25035 & Booth (BRI); Summit of Devils Thumb,

Mossman Gorge N.P., Apr 1986, Godwin C3093 (BRI);

Pinnacle Rock Track, Whyanbeel, Feb 1996, Gray 6566

(BRI); Mossman Bluff, about 10 km W of Mossman,

Dec 1988, Fell DF1686 (BRI).

Distribution and habitat : This species is

endemic to the Wet Tropics of Queensland

where it occurs from Mt Finnigan to Mossman

Bluff (Map 14) in exposed montane rainforest

(microphyll to nanophyll vineforests and

vinethickets) on substrates derived from

granite.

Map 14. Distribution of Symplocos oresbia

Phenology : Flowers are recorded from

February to April; fruit from December to

January.

Notes: Specimens of this species (e.g. Brass

20129 ) were included under Symplocos

cochinchinensis subsp. thwaitesii var.

montana in Nooteboom (1982).

Conservation status: Symplocos oresbia (as

Symplocos sp. (Mt Finnigan L. J.Brass 20129))

is currently listed as Near Threatened under

the NCA.

Jessup, Symplocos in Australia

245

Fig. 3. Symplocos hylandii. A. node with leaf base and budding inflorescence x 4, Jago 6212 (BRI). B. fruit x 2, Ford

AF4913 & Metcalfe (BRI). S. oresbia. C. flower bud x 8, Godwin C3093 (BRI). D. leaf base and inflorescence with

fruit x 2, Hartley 14045 (BRI). S. wooroonooran. E. node with leaf base and inflorescence with buds and flowers x

3, Forster PIF21767 et al. (BRI). F. Flower x 6, Forster PIF21767 et al. (BRI). G. fruit x 3, Hyland 5775 (BRI). Del.

W. Smith.

15. Symplocos paucistaminea F.Muell. &

F.M.Bailey ex F.M.Bailey, Syn. Queensland

FI, Supp. 3: 46 (1890). Type: Queensland.

Cook District: Harvey’s Creek, in 1889

(Bellenden Ker Expedition), F.M. Bailey

s.n. (holo: BRI [AQ23235]; iso: BM, K, L,

MEL46497).

Illustrations : Cooper & Cooper (2004: 538);

Hyland et al. (2010).

Dioecious shrub or tree to 18 m. Shoots and

young branchlets reddish-brown to dark brown

pilose, cataphylls rarely evident. Leaves thinly

coriaceous or chartaceous; lamina narrowly

246

Austrobaileya 8(3): 225-251 (2011)

obovate, oblanceolate or elliptic, 5-22 cm

long, 2-8.5 cm wide, acuminate at apex,

acutely or obtusely cuneate or rounded at

base, glabrescent above, pilose below, slightly

glossy; margins serrulate and denticulate;

secondary veins mostly 7-12 pairs, depressed,

flush or indistinct on upper surface; petiole 4-

12 mm long, pilose. Inflorescences clustered

or not, spicate, to 5 cm, pilose; bracts and

bracteoles ovate, reddish-brown pilose or

villous, persistent, bracts 1.5-2.5 mm long,

bracteoles 1.2-1.5 mm long. Calyx limb 1.2-

1.5 mm long; lobes narrowly ovate or oblong,

1-1.2 mm long, glabrous or sparsely pubescent

towards apex. Petals obovate, 2.5-3.5 mm

long, 1.3-1.5 mm wide, cream to white,

mostly glabrous. Stamens 15-20 in males,

7-10 in females and apparently sterile, 1-2.5

mm long. Disc glabrous. Style 1.2-1.8 mm

long, glabrous; females with peltate stigma.

Fruit ampulliform (when dry), globular when

fresh, 5-5.8 mm long, 3.5-4 mm wide, dark

blue to purple.

Additional selected specimens examined : Queensland.

Cook District: Mt Cook, Aug 1985, Scarth-Johnson

1641A (BRI); Gap Creek, T.R. 146, Fritz L.A., Jul 1973,

Stocker 997 (BRI); Cooper Creek, Cape Tribulation

Area, Aug 1996, Telford 12207 (BRI); V.C.L. Noah, Jun

1975, Hyland 8291 (BRI); Portion 188, Alexandra, Palm

Road, Jun 1985, Gray 4074 (BRI); Baileys Creek area, c.

0.2 miles [0.3 km] E of sawmill, c .7.5 miles [12 km] ENE

of Daintree, Oct 1962, Smith 11516 (BRI); Mt Spurgeon,

Apr 1968, Hyland RFK1508 (BRI); Nissen Creek, Mt

Perseverance Road, Jul 1995, Forster PIF17169 & Figg

(BRI); Churchill Creek, Churchill L.A., S.F. 143, Jul

1995, Forster PIF17204 & Figg (BRI); Churchill Creek,

S.F. 143, Churchill L.A., Nov 1998, Forster PIF24008 et

al. (BRI); F.R. 185, Kalorama L.A. near Danbulla, Jun

1961, Hyland 2047 (BRI); T.R. 1230, Boonjie L.A., May

1977, Gray 526 (BRI); Wooroonooran N.P., Palmerston

section. South Johnston Forestry Track., May 2005,

Forster PIF30970 & Jensen (BRI); S.F. 756 Mt Father

Clancy, May, 2000, Forster PIF25723 & Booth (BRI);

5.6 km S of Mena Creek township, SSW of Innisfail, Jul

2007, Bean 26546 (BRI). North Kennedy District: 27.5

km south along Culpa Road, S.F. 605 Koombooloomba,

May 2002, Forster PIF28789 & Booth (BRI); Paluma

Range, Dotswood Holding, Jun 1974, Hyland 3001

(BRI). South Kennedy District: Mt Macartney, S.F. 652

Cawley, May 1992, Forster PIF9911 & Tucker (BRI);

Dalrymple Heights and vicinity, Jun-Nov 1947, Clemens

s.n. (BM, BRI [AQ84447]).

Distribution and habitat: This species occurs

from the northern end of the Wet Tropics at

Cooktown south to the Clarke Range near

Eungella (Map 15). It grows in upland and

Map 15. Distribution of Symplocospaucistaminea

lowland mesophyll and notophyll vineforests

on substrates derived from basalts and

metamorphics.

Phenology: Flowers are recorded from May to

August; fruit from September to December.

Conservation status: Symplocos

paucistaminea is widely distributed in north

and central eastern Queensland and is not

considered to be threatened.

16. Symplocos puberula Jessup nom. et stat.

nov.; S. cochinchinensis var. pilosiuscula

Noot., Brunonia 4: 320 (1982). Type:

Queensland. Cook District: Speewah,

Upper Clohesy River, 5 June 1966, W.Rijkers

NQNC14754 (holo: L; iso: BRI, CNS).

Illustrations: Nooteboom (1981: Fig. 11);

Hyland et al. (2010), both under Symplocos

cochinchinensis var. pilosiuscula.

Tree to 25 m. Shoots and young branchlets

appressed puberulous, young branchlets

glabrescent. Leaves coriaceous; lamina

oblanceolate or obovate, sometimes elliptic,

6-15 cm long, 2.5-6 cm wide, acuminate

at apex, acute at base, glabrous or with a

few hairs on midvein on underside; margins

shallowly recurved, denticulate, nearly entire;

secondary veins mostly 6-12 pairs; petiole 5-

17 mm long, sparsely appressed puberulous,

247

Jessup, Symplocos in Australia

glabrescent. Inflorescence paniculate or

racemose to 7 cm, puberulous; pedicels 0.6-2

mm long; bracts and bracteoles pubescent,

caducous, bracts ovate, c. 1.5 mm long,

bracteoles narrowly triangular, c. 0.8 mm long.

Calyx limb 1-1.5 mm long, glabrous; lobes

depressed ovate or rounded, glabrous with

ciliate or denticulate margin. Petals oblong

or narrowly obovate, 3.5-5 mm long, 1.7-3.5

mm wide, white, glabrous. Stamens mostly

50-70, 2-6 mm long. Disc annular, shallowly

lobed, glabrous or with a few hairs. Style 3-4

mm long, glabrous or with a few hairs at base,

dilated at stigma. Fruit ampulliform, 7-10

mm long, 4-6 mm wide, dark blue to black.

Additional selected specimens examined : Queensland.

Cook District: 33 km along Leo Creek road, Mcllwraith

Range, Jun 1995, Forster PIF16822 (BRI); T.R. 14,

Head of Peach Creek, Mcllwraith Range, 26.5 km NE

of Coen, Cape York Peninsula, Aug 1993, Fell DGF3543

& Jensen (BRI); Hopevale Aboriginal Reserve, 13.2

km ESE of Hopevale Community, Endeavour River

right arm, Nov 1993, Fell DGF3859 & Stanton (BRI);

Mt Poverty Mine Road, near Normanby Tin Mines, Jun

2003, McDonald KRM1447 (BRI); Gap Creek, Cedar

Bay N.P., Jun 1992, Forster PIF10732 et al. (BRI);

Daintree N.P., Adeline Creek headwaters, top of hill

929, May 1999, Forster PIF24538 & Booth (BRI); S.F.

144, Mt Windsor Tableland, May 1979, Moriarty 2614

(BRI); T.R. 66, 3 km along Mt Lewis Road, Jun 2000,

Forster PIF25821 & Booth (BRI); S.F.R. 607, Shoteel

L.A., Oct 1982, Gray 2779 (BRI); S.F.R. 933, Little Pine

L.A., Feb 1983, Gray 2963 (BRI); Longlands Gap, S.F.

Map 16. Distribution of Symplocos puberula

194, Sep 2001, Forster PIF27514 et al. (BRI). North

Kennedy District: S.F.R. 605, Hilton L.A., Sep 1979,

Dockrill 1615 (BRI); Goold Island [Goold Island N.P],

Aug 1997, Cumming 16252 (BRI); Conway S.F., 2 km E

of Impulse Creek, along Timber Road, Sep 1994, Perry 2

(BRI); S.F. 299 Conway, Brandy Creek Road, 8 km S of

Airlie Beach, Feb 2004, Forster PIF29974 et al. (BRI).

South Kennedy District: 1 km SW of Mt Macartney,

Kangaroo Creek, Cathu S.F., Aug 1989, Thompson 148

(BRI); St Helens Creek, Eungella N.P., Aug 1992, Bean

4949 (BRI); Sarina Range, East Funnel Creek Road,

Sarina Shire, May 2006, Von Fahland 2 (BRI).

Distribution and habitat: This species occurs

from the Mcllwraith Range on Cape York

Peninsula south to the Sarina Range south

of Sarina (Map 16). Grows in mesophyll and

notophyll vineforests on substrates derived

from granites and metamorphics.

Phenology: Flowers are recorded from May to

November; fruit from December to February.

Notes: This species was treated as Symplocos

cochinchinensis var. pilosiuscula Noot. (as

S. cochinchinensis subsp. thwaitesii var.

pilosiuscula Noot.) in Nooteboom (1982).

Conservation status: Symplocos puberula is

widely distributed from Cape York Peninsula

to central eastern Queensland and is not

considered to be threatened.

17. Symplocos stawellii F.Muell., Fragm.

5: 60 (1865); S. stawellii var. stawellii ,

C.T.White, Proc. Roy. Soc. Queensland 50:

82 (1939); S. cochinchinensis var. stawellii

(F.Muell.) Noot., Leiden Bot. Ser. 1: 161

(1975). Type: Queensland. Rockingham Bay,

s.dat., J.Dallachy s.n. (syn: MEL).

Symplocos spicata var. australis Benth., FI.

Austral. 4: 292 (1868). Type: Queensland.

Rockingham Bay, s.dat. J.Dallachy s.n. (syn:

MEL); New South Wales. Richmond River,

s.dat., C. Moore s.n. (syn: NSW).

Illustrations: Nooteboom (1982, Fig. 11);

Hyland et al. (2010), both under Symplocos

cochinchinensis var. stawellii ; Floyd (1989:

397); Leiper et al. (2008: 336) both as S.

stawellii.

Shrub or tree to 30 m. Shoots sparsely

appressed pubescent, glabrescent; young

branchlets glabrescent. Leaves coriaceous;

lamina elliptic to oblanceolate or obovate,

6-18 cm long, 2-6 cm wide, acuminate

at apex, mostly acute or attenuate at base,

248

glabrous or with a few hairs on midvein on

underside; margins serrulate and crenulate,

sometimes slightly recurved; secondary veins

mostly 7-10 pairs; petiole 6-25 mm long,

glabrescent or glabrous. Inflorescence axes

branched, spicate, to 6 cm long, pubescent;

bracts and bracteoles broadly or depressed

ovate, sparsely pubescent or glabrous, ciliate,

persistent to fruiting, bracts 0.8-1.5 mm long,

bracteoles 0.6-0.8 mm long. Calyx limb 0.3-

0.5 mm long, glabrous; lobes very shallowly

rounded or depressed ovate, 0.1-0.4 mm

long, glabrous or nearly so and with a ciliate

margin. Petals obovate, 3.2-3.5 mm long,

1.7-2.5 mm wide, white, glabrous. Stamens

mostly 30-50, 1.5-3 mm long. Disc 5-lobed,

glabrous. Style 2.5-3.5 mm long, glabrous.

Fruit mostly ovoid, 5-5.5 mm long, 3.5-4.5

mm wide, blue to black.

Additional selected specimens examined : Queensland.

Cook District: Lake Barrine, Atherton Tableland, Sep

1929, Kajewski 1237 (BRI, L, P); Malanda - Atherton

Road, May 1976, Stocker 1438 (BRI, CNS, L). North

Kennedy District: S.F.R. 251, Parish of Ravenshoe, Jun

1988, Gray 4844 (BRI, CNS). South Kennedy District:

Dalrymple Road, Eungella N.P, Apr 2000, Forster

• Q

on 00

T-:

4 Rockham

z u o

pton

i ~

. | Bri

25 S

sbane

sxC ■

•

/Sydney

30 S -

OC°Q

iJF V

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l • v-r

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_i_

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Map 17. Distribution of Symplocos stawellii

Austrobaileya 8(3): 225-251 (2011)

PIF25512 & Booth (BRI); S.F.R. 679, Range L.A., 1 km

S of Crediton, Clarke Range, Jun 2001, Ford AF2887

(BRI). Port Curtis District: Byfield N.P., 9 km ENE

of Byfield township, Sep 2000, Forster PIF26237 et al.

(BRI). Wide Bay District: Verrierdale Road, 8 km W of

Peregian Beach, May 1990, Bean 1589 (BRI). Moreton

District: Running Creek, c. 2 miles [3.2 km] NW of

Yandina, May 1969, Smith 14616 (BRI). New South

Wales. Coombadjha Creek Road, Apr 1980, Floyd 1477

(BRI); Johns River near Newcastle, May 1956, Webb &

Tracey 3691 (BRI).

Distribution and habitat : This species is

widespread in eastern Queensland and New

South Wales from the Hann Tableland,

northwest of Mareeba to near Windsor, New

South Wales (Map 17). It generally grows in

notophyll vineforests, although it sometimes

occurs as an understorey tree in moist eucalypt

forest. Substrates are diverse.

Phenology : Flowers are recorded from April

to August; fruit from November to January.

Conservation status : Symplocos stawellii is

widely distributed from north Queensland to

central eastern New South Wales and is not

considered to be threatened.

18. Symplocos thwaitesii F.Muell., Fragm.

3: 22 (1862); S. cochinchinensis subsp.

thwaitesii (F.Muell.) Noot., Leiden Bot. Ser. 1:

159 (1975); S. cochinchinensis var. thwaitesii

(F.Muell.) Noot., Leiden Bot. Ser. 1: 160

(1975). Type: New South Wales. Hastings

River, s.dat ., Beckler s.n. (holo: MEL46540,

46541; iso: BRI, P).

Illustrations : Nooteboom (1982, Fig. 11)

under S. cochinchinensis var. thwaitesii ;

Floyd (1989: 397) as S. thwaitesii.

Shrub or tree to 20 m. Shoots and young

branchlets glabrous. Leaves coriaceous;

lamina oblanceolate, narrowly obovate or

elliptic, mostly 6-16 cm long, 1.5-5 cm wide,

acuminate, acute or obtuse at apex, acute or

attenuate at base, glabrous; margins serrate;

secondary veins mostly 8-11 pairs; petiole

5-15 mm long, glabrous. Inflorescence

paniculate or racemose to 7 cm, glabrous;

pedicels 0-8 mm long; bracts and bracteoles

obovate to triangular, glabrous or scarcely

ciliate, caducous, bracts 1.5-4 mm long,

bracteoles 1-2 mm long. Calyx limb 1.7-2

mm long, glabrous; lobes depressed obovate,

1.4-1.6 mm long glabrous. Petals broadly

249

Jessup, Symplocos in Australia

obovate, mostly 4.5-7 mm long, 3.5-5 mm

wide, white, glabrous. Stamens 80-100,

2-5 mm long. Disc glabrous. Style 4-5 mm

long, glabrous. Fruit ovoid to ampulliform,

6-11 mm long, 4-8 mm wide, blue, purple or

black.

Additional selected specimens examined : Queensland.

Port Curtis District: Isopod, Mt Parnassus, Oct 1976,

Hyland 4314RFK (BRI); Bulburin S.F. 67, Apr 1980,

McDonald 3220 et al. (BRI). Wide Bay District: The

Missings, S.F. 915 Tuan, Dec 1994, Forster PIF16046

et al. (BRI); S.F. 639 Wrattens, Beauty Spot 50, end of

Black Snake Road, Jan 2003, Haskard & Smyrell s.n.

(BRI [AQ772885]); c. 2 miles [3.2 km] NE Maleny,

May 1959, Thorne 21263 et al. (BRI); Mary Cairncross

Park, Sep 2004 , Russell s.n. (BRI [AQ613832]). Burnett

District: Bunya Mountains, Oct 1919, White s.n. (BRI

[AQ84418]). Moreton District: Brolga N.P, W of

Woombye, Feb 1990, Bean 1349 (BRI); Walking track

between Lawnton Road & Joyners Ridge Road, c. 1 km

from Lawton Road, Mt Glorious, Sep 2002, Phillips 783

(BRI); Mt Glorious c. 50 km W of Brisbane, Sep 1978,

Jessup 147 & Reynolds (BRI); Mt Mistake, May 1948,

Smith 3689 & Webb (BRI); Cainbable Range, Oct 1952,

Lahey s.n. (BRI [AQ478660]); Burleigh Heads N.P,

Tallebudgera, Jul 1978, McDonald 2165 et al. (BRI).

Darling Downs District: Mt Cordeaux, just below the

summit, Dec 1981, Guymer 1661 & Jessup (BRI); The

Head, Main Range N.P, Jan 2002, Forster PIF28066

6 Leiper (BRI); Border of Lamington N.P., Sarabah

Range, Oct 1969, Schodde 5576 (BRI). New South

Wales. Unumgar near base of Mt Lindsay, Nov 1944,

Jones s.n. (BRI [AQ84429]); Booyong Reserve NE of

Lismore, Nov 1982, Floyd s.n. (BRI [AQ393589]); c. 1

mile [1.6 km] S of Port Macquarie, Oct 1966, Schodde

5152 (BRI, CANB, L); Robertson Nature Reserve, Jun

1994, Kodela 315 et al. (BRI). Victoria. Spur between

two gullies entering Snowy River on W bank of Wood

Point [13 km NW of Orbost], Mar 1976, Cameron 6132a

(MEL).

Distribution and habitat: Endemic to eastern

Australia from central Queensland (Mt

Parnassus near Rockhampton) south to the

Snowy River in Victoria (Map 18). Grows

mostly in notophyll vine forest on substrates

derived primarily from basalts and granites.

Phenology: Flowers are recorded from

September to November; fruit in February.

Conservation status: Symplocos thwaitesii is

widely distributed and is not considered to be

threatened.

19. Symplocos wooroonooran Jessup nom. et

stat. nov.; S. stawellii var. montana C.T.White,

Proc. Roy. Soc. Queensland 50:82 (1939);

S. cochinchinensis var. montana (C.T.White)

Noot., Leiden Bot. Ser. 1: 160 (1975). Type:

Queensland. Cook District: Mt Bartle Frere,

7 October 1938, H.Flecker 857 (holo: BRI).

Illustrations: Cooper & Cooper (2004: 539);

Hyland et al. (2010), both under Symplocos

stawellii var. montana.

Tree to 8 m. Shoots appressed pubescent,

glabrescent; young branchlets glabrescent.

Leaves chartaceous to thinly coriaceous;

lamina oblanceolate to elliptic, 2-6.5 cm

long, 0.9-2.4 cm wide, acuminate at apex,

mostly attenuate at base, glabrous or nearly

so; margins recurved, remotely denticulate or

sometimes entire; secondary veins mostly 7-

10 pairs; petiole 2.5-4.5 mm long, glabrescent

or glabrous. Inflorescence axes 2 or 3,

racemose, to 1.5 cm long, lightly appressed

puberulous, branching at base, sometimes

single; pedicels 0-1 mm long; bracts and

bracteoles ovate, sparsely puberulous, ciliate,

mostly persistent until after anthesis, bracts 1-

2 mm long, bracteoles 0.6-1 mm long. Calyx

limb 0.75-1 mm, glabrous; lobes depressed

ovate, triangular or shortly oblong, rounded,

250

0.5-0.8 mm long, glabrous. Petals oblong

or obovate, 4-5 mm long, white, glabrous.

Stamens mostly 30-35. Disc pubescent. Style

3-3.5 mm long, pubescent near base. Fruit

ampulliform, 5.8-6.3 mm long, 4-5.5 mm

wide, black. Fig. 3E-G.

Additional selected specimens examined: Queensland.

Cook District: S.F.R. 310, Upper Goldsborough L.A.,

Sep 1976, Dockrill 1278 (BRI); Bellenden Ker, s.dat.,

Johnson s.n. (BRI [AQ519814]; MEL46558, 46559,

46560,46561,46562); Summit of Mt Bellenden Ker, Aug

1971, Hyland 5304 & 5316 (BRI); N.P.R. 226, Bellenden

Ker, Jan 1972, Hyland 5775 (BRI); Bellenden Ker

Range, Oct 1974, Hyland 7769 (BRI); Centre Peak, near

TV tower, summit of Bellenden Ker, Nov 1972, Webb &

Tracey 10802 (BRI); Mt Bellenden Ker, Jun 1969, Smith

14630 (BRI); loc. cit.. May 2001, Cooper 1525 & Cooper

(BRI); Wooroonooran N.P, Mt Bellenden Ker summit

area, Dec 2000, Forster PIF26516 et al. (BRI); Mt Bartle

Frere, Oct 1935, Blake 9843 (BRI); loc. cit., Nov 1966,

Gilmour 4076 (BRI); S.F.R. 755, Gosschalk L.A., Gray

702 (BRI); Summit area of Mt Bartle Frere, Sep 1977,

Powell 833 et al. (BRI); Wooroonooran N.P, Bartle

Frere, first heath to SE of South Peak, Oct 1997, Forster

PIF21767 et al. (BRI).

Distribution and habitat : This species is

endemic to the Wet Tropics of Queensland

where it occurs only on the upper slopes and

summits of Mt Bartle Frere and Mt Bellenden

Austrobaileya 8(3): 225-251 (2011)

Ker in Wooroonooran N.P. (Map 19). It grows

in montane rainforest (simple microphyll

vine-fern thicket and shrubland) on granite

derived substrates.

Phenology : Flowers are recorded in October

and November; fruit from January to May.

Notes : This species was treated as Symplocos

cochinchinensis var. montana (C.T.White)

Noot. (as S. cochinchinensis subsp. thwaitesii

var. montana) in Nooteboom (1982).

Conservation status : Symplocos

wooroonooran (as Symplocos stawellii

var. montana) is currently listed as Near

Threatened under the NCA.

Etymology : The species epithet

commemorates the botanically diverse

Wooroonooran N.P. and is also the aboriginal

name for Mt Bellenden Ker (Bailey 1900a).

Acknowledgements

This work was partially funded by a grant

from the Australian Biological Resources

Study (ABRS) for the purposes of providing

an account of the family for the Flora of

Australia. I would like to thank the directors

and staff of BM, BRI, CANB, CNS (formerly

QRS), K, L, MEL, NSW and P for access to

their collections. Will Smith produced the

line drawings and Peter Bostock checked and

corrected my Latin diagnoses.

References

Bailey, F.M. (1890). A Synopsis of the Queensland

Flora, Supplement 3: 46. H.J.Diddams & Co.:

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- (1900a). Leptospermum wooroonooran

F.M.Bailey. In The Queensland Flora 2: 588.

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Brisbane.

- (1900b). Symplocos Linn. In The Queensland

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Bentham, G. (1868). Symplocos Linn. (Styracaceae). In

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Brand, A. (1901). Symplocaceae. In H.G.A.Engler (ed.),

Das Pflanzenreich Heft 6(1 V, 242): 1-111.

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Map 19. Distribution of Symplocos wooroonooran

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Jessup, Symplocos in Australia

Bostock, P.D. & Holland, A.E. (eds.) (2010). Census

of the Queensland Flora 2010. Queensland

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Cooper, W. & Cooper, W.T. (2004). Fruits of the

Australian Tropical Rainforest. Nokomis

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eastern Australia. Inkata Press: Melbourne/

Sydney.

Fritsch, P.W., Cruz, B.C., Almeda, F., Wang, Y. & Shi,

5. (2006). Phylogeny of Symplocos based on

DNA sequences of the chloroplast trnC-trnD

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Fritsch, P.W., Kelly, L.M., Wang, Y., Almeda, F. &

Kriebel, R. (2008). Revised infrafamilial

classification of Symplocaceae based on

phylogenetic data from DNA sequences and

morphology. Taxon 57: 823-852.

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Hyland, B.P.M., Whiffin, T. & Zich, F. (2010).

Australian Tropical Rainforest Plants. Edition

6. http://kevs.trin.org.au:8080/kev-server/data/

0e0f0504-0103-430d-8004-060d07080d04/

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Jessup, L.W. (1993). Two new species of Symplocos

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- (1994). Symplocaceae. In R.J.F. Henderson

(ed.), Queensland Vascular Plants: Names and

Distribution , p. 329. Queensland Herbarium:

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- (1997). Symplocaceae. InR. J.F. Henderson (ed.),

Queensland Plants: Names and Distribution ,

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- (1865). Fragmenta Phytographiae Australiae 5:

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- (1982). A revision of the Australian species of

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326.

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Flora of south-eastern Queensland 2: 272-273.

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Kelly, L.M. (2004). Phylogeny and infrageneric

classification of Symplocos (Symplocaceae)

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- (2002). Symplocaceae. In R.J.F. Henderson

(ed.), Names and Distribution of Queensland

Plants, Algae and Lichens , p. 193. Queensland

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- (2007). Symplocaceae. In P.D. Bostock & A.E.

Holland (eds.). Census of the Queensland Flora

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Management: Brisbane.

A taxonomic revision of Pseudoweinmannia

Engl. (Cunoniaceae: Geissoieae)

Andrew C. Rozefelds 1 & Belinda Pellow 2

Summary

Rozefelds, AC. & Pellow, B. (2011). A taxonomic revision of Pseudoweinmannia Engl. (Cunoniaceae:

Geissoieae). Austrobaileya 8(3): 252-266. The vegetative and reproductive morphology of

Pseudoweinmannia is described and illustrated. The genus is endemic to eastern Australia with two

species, Pseudoweinmannia apetala (F.M.Bailey) Engl, and P. lachnocarpa (F.Muell.) Engl., that

can be differentiated by their leaf serration, stipule morphology and flower and fruit characters.

New interpretations of floral and fruit morphology in Pseudoweinmania are discussed, and their

significance in better understanding the phylogenetic relationships within the tribe Geissoieae are

examined.

Key Words: Cunoniaceae, Geissoieae , Pseudoweinmannia , Pseudoweinmannia apetala ,

Pseudoweinmannia lachnocarpa , Australia flora, Queensland flora. New South Wales flora,

comparative morphology

‘Andrew C. Rozefelds, Queensland Museum, GPO Box 3300, South Brisbane, Queensland 4101,

Australia. Email: andrew.rozefelds@qm.qld.gov.au

2 Belinda Pellow, Janet Cosh Herbarium, Department of Biological Sciences, University of

Wollongong, Northfields Avenue, Wollongong, New South Wales 2522, Australia.

Introduction

The family Cunoniaceae includes 15 genera

in Australia, which are all primarily rainforest

trees and/or shrubs, and are currently being

revised for the Flora of Australia. Recent

revisions have been completed on the genera:

Anodopetalum A.Cunn ex Endl. (Barnes

& Rozefelds 2000), Ceratopetalum Sm.

(Rozefelds & Barnes 2002), Davidsonia

F.Muell. (Harden & Williams 2000), the

Australian species of Geissois Labill.

(Schimanski & Rozefelds 2002), Vesselowskya

Pamp. (Rozefelds et al. 2001) and a new

species of Gillbeea F.Muell. was described

from north eastern Australia (Rozefelds &

Pellow 2001). In this paper a revision of the

Australian endemic genus Pseudoweinmannia

Engl, is presented.

Pseudoweinmannia was proposed by

Engler (1930) for two species, Weinmannia

lachnocarpa F.Muell. and W. apetala

F.M.Bailey from eastern Australia, which he

referred to the tribe Cunonieae (R.Br.) Schrank

& Mart, in Cunoniaceae. Pseudoweinmannia

differs from Weinmannia, the major genus

Accepted for publication 8 August 2011

in this tribe, in lacking petals, having a

polyandrousandroeciumandtardily-dehiscent

fruits. Based upon an analysis of stipule

morphology Dickison & Rutishauser (1990)

concluded, that Pseudoweinmannia was more

closely related to Geissois and Lamanonia

Veil, in the Tribe Belangerae Lindl ex Pfeiff.

(= Tribe Geissoieae Endl. ex Meisn.), than to

the Tribe Cunonieae.

In a cladistic analysis using morphological

characters, Hufford & Dickison (1992) showed

that Pseudoweinmannia was sister to Geissois

(including Lamanonia ), as it shared with this

genus, lateral stipules (Dickison & Rutishauser

1990), racemose inflorescences and a

polyandrous androecium. Dickison (1984)

and Hufford & Dickison (1992) also identified

structures on the seeds as elaiosomes, which

they interpreted as a putative autapomorphy

for the genus.

Combined morphological and molecular

data based upon analysis ofrZ>cL, and intron and

spacer regions in the combined trnL-G region,

and chloroplast DNA data provide support for

interpreting Pseudoweinmannia and Geissois

sens. lat. as sister taxa (Bradford & Barnes

2001). These authors recircumscribed the

Rozefelds & Pellow, Revision of Pseudoweinmannia

253

Tribe Geissoieae (= Tribe Belangerae sensu

Dickison & Rutishauser 1990) to include the

Australian and New Caledonian species of

Geissois , Lamanonia from South America,

and Pseudoweinmannia.

Pseudoweinmannia is endemic to eastern

Australia where it is restricted to rainforests.

Pseudoweinmannia lachnocarpa (F.Muell.)

Engl. (Fig. 1) occurs in north eastern New

South Wales and south eastern Queensland

(Map 1), and was described by Mueller (1874)

from material collected from the Tweed

District in north eastern New South Wales. A

second species, Pseudoweinmannia apetala

(F.M.Bailey) Engl. (Fig. 2), was described by

Bailey (1893), from Kamerunga, near Cairns

in north eastern Queensland (Map 1). Bailey

(1893), in describing Pseudoweinmannia

apetala , noted that it “approaches near to

lachnocarpa but differs in its more lax slender

inflorescence, more membranous leaves,

which are not so prominently reticulate, its

longer stamens, as well as probably the flower

lobes to the calyx”. Since their description

there has not been a critical re-examination of

the morphology of both species and as a result

the status of Pseudoweinmannia apetala has

become unclear. Recent reviews in flora and

state censuses have either considered it a

synonym of Pseudoweinmannia lachnocarpa

(e.g. Henderson 1997; Hyland & Whiffin

1993; Jessup 2010), or a distinct species (e.g.

Stanley & Ross 1983; Harden 1990; Bradford

etal. 2004).

In this paper the vegetative and

reproductive morphology of the two species

is described using herbarium collections

(both dried and in spirit) and consistent

morphological differences are recognized

between Pseudoweinmannia lachnocarpa and

P. apetala. The vegetative and reproductive

morphology of some key characters in

Pseudoweinmannia, i.e. stipule morphology,

inflorescence architecture and floral and fruit

morphology, are also described and illustrated

to permit comparisons with other genera in

the Tribe.

Methods and materials

Herbarium collections examined include those

from BRI, CANB, CNS (formerly in QRS),

K, NSW and MEL. The principal collections

used are cited herein and specimens examined

with the scanning electron microscopy (SEM)

are indicated by *. Common abbreviations

used in specimen citation are N.R (National

Park), L A. (Logging Area), S.F./S.F.R. (State

Forest/State Forest Reserve).

Terminology follows Walsh & Entwisle

(1999) for leaf morphology and Hickey (1979)

for leaf architecture.

For SEM the material prepared included

floral organs, pollen, seeds, and stipules. The

specimens were either air-dried or critical

point dried, and placed onto aluminium stubs

with tape. They were then sputter coated with

gold and examined with a Philips Electroscan

Environmental Scanning Electron Microscope

2020, under high vacuum operated at 10-15

kV.

Results

Stipule morphology

The stipules in Pseudoweinmannia are keel¬

shaped, lanceolate structures. Developmental

studies in Pseudoweinmannia and the

Australian Geissois species by Dickison

& Rutishauser (1990) showed that four

stipular primordia arise laterally at each

node. In Pseudoweinmannia the two stipular

primordial, on either side of the node, coalesce

to form a bifid laterally-positioned compound

structure (Fig. 3B). The margins of the

stipules have glandular trichomes and simple

hairs in Pseudoweinmannia apetala while in

P. lachnocarpa only glandular trichomes are

present (Fig. 3A-C).

Inflorescence architecture

Schimanski & Rozefelds (2002) have

described the inflorescence morphology in

the Australian species of the related genus

Geissois. Their terminology is applied here

to describe the inflorescence morphology of

Pseudoweinmannia.

The flowers of Pseudoweinmannia are

arranged in indeterminate racemose units.

254

Austrobaileya 8(3): 252-266 (2011)

20 DEC 2001

Depth m

Alt. 960m

173 Cunoniaceae

AQ 567426

QUEENSLAND HERBARIUM (BRI)

Flora of Queensland Cook

Pseudoweinmannia lachnocarpa (F.Muell.) Engl.

QUEENSLAND HERBARIUM (BRI)

Brisbane Australia

557426

Coll. A, Ford

Jensen,R.;

<16d 17m 36s 145d 5m 18s)[AGD66]

(55,295750,8197650) 7965-957976

SFR144, Chovvchilla Logging Area, along old logging track, Windsor

Tableland.

Complex notophyll rainforest on soil derived from granite. Agathis

robusta very common.

Canopy of Aleurites, Argyrodendron poly end rum and

Pseudoweinmannia.

Cdfnmon small trees - Austromyrtus spp., Dinosperma, Croton

insularis and Elaeodendron.

Canopy tree 15m high. New growth reddish.

Fruit slightly immature, hairs creamish when fresh.

Dup. NSW MEL HO\LN^

* May be cited as computerised collection Number

(Archival Paper)

Fig. 1 . Representative specimen of Pseudoweinmannia apetala {Ford AF3202 & Jensen [BRI])

Rozefelds & Pellow, Revision of Pseudoweinmannia

255

QUEENSLAND HERBARIUM (BRI)

Flora of Queensland

Pseudoweinmannia lachnocarpa (F.Muell.) Engl.

Coll, P.l.Forster PIF9353

+P.Machim

Moreton

6 JAN 1992

27 b 25'S 152*49'E Alt> m .

2l State Forest 309 Enoggera. " Grid ref!

9443-o24o60.

creek 3 '" 13 ' 1 not ° phyl 1 vine forest on dark brown soil along

Tree to 5m, cream flowers.

Occasional In area.

QUEENSLAND HERBARIUM (BRI)

Brisbane Australia

520250

Fig. 2. Representative specimen of Pseudoweinmannia lachnocarpa {Forster 9353 & Machin [BRI])

256

Following Bradford (1998), we use the term

raceme to describe these elements, although

occasionally the racemes are weakly branched

(e.g. Webb & Tracey 11246). Bracts subtend

each flower, but are quickly deciduous. The

pedicels are shortly hirsute and have an

abscission point about 1-2 mm above the

point of attachment.

Variation in Psendoweinmannia , as in

the Australian species of Geissois , occurs

at the following hierarchical levels: (1) in

the varying development of the racemes

and supporting axes and buds that form the

inflorescence (Inflorescence Module sensu

Bradford 1998); and (2) in the position that

the inflorescences occupy in relation to the

main stem axis. In both Pseudoweinmannia

apetala and P. lachnocarpa the common

condition, particularly in younger shoots, is

for the raceme to occupy an axial position in

the leaf axils and a terminal bud is present.

The pattern is more complicated in

older branch units, where inflorescence

units may develop in a number of leaf axils,

and a compound inflorescence is formed;

subtending leaves, may or may not, be

present. The inflorescence could be described

as a compound homothetic inflorescence

{sensu Weberling 1989), and the arrangement

of raceme-like units on the flowering stem is

consistent with the decussate architecture of

the shoot. In some specimens the terminal

bud may also be replaced by a raceme

{Pseudoweinmannia lachnocarpa {Blake

22848)' P. apetala {Whiffin 5978 , Gray

5978)). Heterothetic compound inflorescences

{sensu Weberling 1989) therefore result if the

terminal bud is replaced by a raceme-like unit

and inflorescences are initiated in a number

of leaf axils. In both Pseudoweinmannia

apetala and P. lachnocarpa , heterothetic and

homothetic compound inflorescences may

occur on the same specimen, and no consistent

inflorescence architectural differences have

been identified between the two species.

The racemes are somewhat variable in

length, although those of Pseudoweinmannia

apetala are usually longer than that in

P. lachnocarpa.

Austrobaileya 8(3): 252-266 (2011)

Floral and fruit morphology

TheflowersinbothPsezzdowzzzzzzawzzaspecies

are (5-)6(-7)-merous and polystemonous

(Bradford & Barnes 2001; Bradford etal. 2004;

Schimanski & Rozefelds 2002) (Fig. 4B,E).

The sepals are valvate, ovate in shape and have

a strigose indumentum over their inner and

outer surfaces (Fig. 4A,D). Petals are lacking

in both species (Fig. 4B,E). Anthers are

introrse, dorsifixed at just below the midpoint

of the anther and are weakly versatile and

lack a sterile distal protrusion (Fig. 4A,D,E).

A prominent, lobed annular nectary occurs at

the base of the ovary (Fig. 4B,F). The ovary is

bicarpellate, rarely tricarpellate, syncarpous,

and is covered by a dense indumentum of

strigose hairs (Fig. 4A,B>C,F). The styles

are furrowed ventrally, are slightly coiled

in bud and in young flowers (Fig. 4A) and

straighten with flower development (Fig. 4B).

The styles are exserted before the anthers

and while studies on stigmatic receptivity

have not been undertaken, the flowers are

possibly proterogynous (Fig. 4D). The stigma

is papillate at maturity (Fig. 4C). The fruit

is a tardily dehiscent capsule. The ovary

in Pseudoweinmannia is divided into two

parts, rarely three, indicating origin from

two carpels; the two carpels are adnate and

free (Fig. 5A,B) The seeds are irregularly

rounded pyramidal in shape (Fig. 6A,C), with

weakly reticulate cell walls (Fig. 6B,D) and

lack a wing-like extension.

Comparative morphology in the Tribe

Geissoieae

The Tribe Geissoieae consists of

Pseudoweinmannia , Lamanonia and Geissois

sens, lat., and includes species that occur in

Australia, South America and New Caledonia

(Bradford & Barnes 2001). The adult leaves

in Pseudoweinmannia are trifoliolate and

have two caducous stipels at the base of the

lateral leaflets. Australian Geissois species

also have stipels, or rarely reduced leaflets,

that occupy the same position as those in

Pseudoweinmannia (Schimanski & Rozefelds

2002). The New Caledonian Geissois taxa

and Lamanonia are typically digitate or

trifoliolate, the most basal leaflets being

the smallest. Bradford & Barnes (2001)

Rozefelds & Pellow, Revision of Pseudoweinmannia

257

Fig. 3. Variation in stipule morphology in Pseudoweinmannia apetala and P. lachnocarpa. A. P. lachnocarpa,

detail of stipule margin showing glandular trichomes. B. P. apetala, the pair of stipules and the margins between the

two stipules are arrowed. C. P. apetala stipule, detail of stipule margin showing unicellular and glandular trichomes

(arrowed). Scale bars A,C = 300 pm; B = 1 mm. A from Floyd s.n. (CANB 289092); B-C from Gray 1272 (CNS).

interpreted the common leaf condition in the

tribe as being digitate (palmately compound),

which is logical if the stipels in Geissois

and Pseudoweinmannia are interpreted as

homologues to lateral leaflets.

Dickison (1975) described the stomata in

Pseudoweinmannia as anomocytic, although

recent studies by Barnes in Bradford &

Barnes (2001) interpreted the stomata as

brachyparacytic, which is the common

condition in the family and is present for all

genera of the Tribe Geissoieae. The secondary

leaf venation is semi-craspedodromous,

which is the common condition in most of

the Cunoniaceae and is typical of Geissoieae

(Hufford & Dickison 1992; Barnes pers.

comm. 2011).

The androecium is defined as polyandrous

in the Geissoieae (Bradford & Barnes 2001).

In the New Caledonian species of Geissois ,

developmental studies show that the stamens

are grouped together in fascicles, occupy

an episepalous position and development

proceeds from a centre of the fascicle

outwards in both directions (Matthews &

Endress 2002). In developing flowers the

inner stamens are typically larger than the

outer ones in the fascicle. This arrangement

of stamens is evident in older stages of flower

development in the Australian species of

Pseudoweinmannia (Rozefelds pers. obs .)

and Geissois (Schimanski & Rozefelds 2002,

figs 25, 27). The arrangement of stamens

in Pseudoweinmannia and the Australian

species of Geissois is considered to be weakly

polyandrous.

The common fruit type in the Cunoniaceae

is a bilocular, ventrally dehiscent capsule

(Dickison 1984). Geissois s.l. and Lamanonia

have ventrally dehiscent capsules, while those

of Pseudoweinmannia were interpreted by

Dickison (1984) as an ‘indehiscent capsule’

and Bradford et al. (2004) similarly referred to

the fruits in Pseudoweinmannia as indehiscent

258

Austrobaileya 8(3): 252-266 (2011)

Rozefelds & Pellow, Revision of Pseudoweinmannia

fruits. Other researchers have indicated that

the fruits are septicidally dehiscent (Harden

1990) and open into two valves (Floyd 1989).

The semantic discontinuities imposed by

terms, such as ‘indehiscent’ versus ‘dehiscent’,

obscure the morphological similarities that

link these genera together.

The fruits in Pseudoweinmannia are

typically derived from two carpels, although

they are obscured by the mass of fine hairs

on the outside of the fruit (Dickison 1984,

fig. 13). Sections through the fruit indicate

that the walls of the carpels are adnate, but

are not fused together (Fig. 5A,B) The

placental proliferations that Dickison (1984,

pi. 2, Fig. 21) illustrated were not evident in

the specimens we sectioned. The thickened

(sclerenchyma) walls of the capsules can

be convoluted in appearance as the oblique

section revealed (Fig. 5B), but near the mid¬

region of the fruit these walls are straight

(Fig. 5B). It seems likely that Dickison’s

interpretations were based upon either a

section through an immature fruit and/or

an oblique section through the fruit that cut

through the developing, convoluted walls of

the capsule.

InPseudoweinmanniathQ fruits are similar

to those of the other genera as the walls of the

two carpels are not fused together but stay

conjoined for some time after fruit fall. The

mid-section through the Pseudoweinmannia

fruits indicates that sclerenchyma occupies

a largely identical position to that seen in

Geissois stipularis A.C.Sm (Dickison 1984, pi

Fig. 4. (facing page) Floral morphology of

Pseudoweinmannia apetala and P. laclinocarpa. A-C.

P. apetala. A. Partially opened bud with some stamens

removed to show dense covering of hairs on ovary, and

curved styles. B. Mature flower with fully exserted

stamens, and the two style branches are arrowed, note

conspicuous disc (nectary = N) at base of ovary. C. Top

view of flower and the papillate surface of the stigma

is arrowed (white arrow). D-F. P. lachnocarpa. D.

partially opened bud, note that the style branches are

exserted (white arrows) and the anthers are dorsifixed

at just beiow anther midpoint (black arrow). E. Newly

open flower, note the variation in length of the stamen

filaments. F. Mature flower with stamen filaments

extended and basal disc (N). Scale bars A,C,D = 1 mm;

B,E,F = 2 mm. A from Gray 5978 (CNS); D-E from

Rozefelds s.n. (HO); F from Forster PIF9353 & Machin

259

2, fig. 22). All genera in the Tribe Geissoieae

therefore have bi-locular fruits that are

derived from capsules and are dehiscent or

tardily dehiscent.

Pseudoweinmannia differs from the other

genera in the Tribe by stipule morphology

(Schimanski & Rozefelds 2002) and in

seed and fruit morphology. The stipules

in Pseudoweinmannia are keel-shaped,

lanceolate structures (Fig. 3B), while those

in Geissois from Australia (Schimanski &

Rozefelds 2002) and Lamanonia from South

America are large and often foliose (Leite

1983; Zickel & Leitao Filho 1993; Engler

1930).

Pseudoweinmannia has rounded,

pyramidal seeds while the other genera in

the tribe have winged seeds. Dickison (1984)

described the seeds of Geissois s. lat. as

having distal wings. Lamanonia also has a

distal wing (Engler’s (1930) illustration of

L. tomentosa Camb. (= 1L. ternata Veil.)',

Zickel & Leitao Filho 1993; Leite 1983),

and Dickison (1984) interpreted the seed in

Lamanonia ternata as having a distal wing

and lateral extensions. Dickison (1984) drew

particular attention to an aril-like outgrowth

on the seeds in Pseudoweinmannia , which

he interpreted as an elaiosome. We observed

a few, thin-walled parenchyma cells around

the point of attachment of the seed, but we

interpret these cells as being derived from the

placental parenchyma (Fig. 5B,C). We are

unaware of any ecological or field studies that

support interpretation of these structures as

an elaiosome.

The seeds of Pseudoweinmannia were

described as having either a smooth (Dickison

1984) or papillate (Hufford & Dickison 1992)

surface, although the material we examined of

both species had a smooth, weakly reticulate

appearance due to the cell walls (Fig. 6A-D).

Dickison (1984) considered the epidermal

cells of the Australian species of Geissois to be

more ornamented and they could be identified

apart by differences in the epidermal cell

structures. The epidermal cells of the seed

coats of Lamanonia and Geissois stipularis

A.C.Sm. (New Caledonia) were described

as having irregular outlines and moderately

thick to thin lateral walls (Dickison 1984).

260

Austrobaileya 8(3): 252-266 (2011)

Fig. 5. A. Pseudoweinmamtia lachnocarpa, oblique

transverse section through a mature fruit. Note the

position of the seeds in the fruit and the thickened seed

coat (SC). B. Drawing of A showing that the walls of

the two carpels are adnate but free; the sclerenchyma

wall (sc) of the right hand carpel are not continuous and

remain free near the mid-section of the fruit. The more

oblique section through the left hand carpel shows that

the convoluted sclerenchyma (sc) wall of the carpel is

continuous. C. Simplified transverse section of fruit of

Geissois stipularis A.C.Sm., redrawn from Dickison

(1984), note that the sclerenchyma walls (sc) of the

capsule remain free. (S = seed; SC = seed coat; sc =

sclerenchyna). Scale bar = 1 mm. A from Williams s.n.

(MEL).

The tribe Geissoieae are a very cohesive

group, with all species having a similar

vegetative and reproductive morphology. The

species share digitate (palmately compound)

leaves, brachyparacytic stomates and semi-

craspedodromous secondary venation. The

floral plan in all genera consists of valvate

sepals, no petals, polyandrous androecium,

the anthers lack a distal connective protrusion,

and the bicarpellate ovary is covered in a thick

indumentum with a prominent disc (nectary)

at the base. Pseudoweinmannia also shares

with the other two genera dicolporate pollen,

raceme-like inflorescences and capsular

dehiscent fruits (Hufford & Dickison 1992;

Bradford & Barnes 2001; Schimanski &

Rozefelds 2002; herein).

Systematics

Family Cunoniaceae

Tribe Geissoieae Endl. ex Meisn.

Pseudoweinmannia Engl., Nat. Pflanzenfam.

2nd Ed., Vol. 18a: 249 (1930). Type species:

Pseudoweinmannia lachnocarpa (F.Muell.)

Engl. (= Weinmannia lachnocarpa F.Muell.)

Trees or shrubs. Leaves opposite, decussate,

trifoliolate and with a pair of stipels at base

of lateral leaflets (digitate); unifoliolate in

coppice growth or seedlings; leaflets sessile

or very shortly petiolulate, similar in size,

central leaflet sometimes up to one-third

larger, broadly lanceolate, elliptical to obovate,

apices typically acute, less commonly obtuse,

leaf bases are cuneate to attenuate, lateral

leaflets base slightly asymmetrical, margin

serrate to crenate, sinuses weakly to strongly

Rozefelds & Pellow, Revision of Pseudoweinmannia

developed, venation semi-craspedodromous,

midrib course straight in central leaflet and

slightly curved in lateral leaflets, secondary

venation thickness stout, alternate, uniformly

divergent, tertiary venation percurrent,

complete areolation is formed by higher-

order venation, teeth dark, glandular with

simple apical termination. Stipules 4 at each

node, keel-shaped, lanceolate, apex acute.

Inflorescences usually axillary, consisting of

raceme-like units, which typically occur in the

upper leaf axils, or occasionally terminate the

main axis. Flowers bisexual; sepals (5-)6(-7)

free; petals absent; androecium weakly

261

polystemonous, stamens 12-25, carpels 2 (or

3) fused, styles subulate, 2 (or 3) distally free,

ovary densely hairy, stigmas papillate, 2-4

ovules/carpel, disc (annular nectary) around

base of ovary. Fruit a dry, tardily dehiscent

capsule covered in a dense, woolly golden-

brown indumentum, stamen filaments and

styles persistent; carpels free; seed rounded-

pyramidal, not winged, seed walls weakly

reticulate.

Etymology: Name from the Latin pseudo

meaning (false), and Weinmannia a genus

named in honour of Johann Wilhelm

Weinmann, German pharmacist and botanist,

(1683-1741).

Key to Pseudoweinmannia species

Leaf margins crenulate-crenate, sinuses weakly developed; stipules

fringed with glandular and simple trichomes; stamens

12-20, disc < 2.5 mm high; fruit ovoid-globose. NE Qld. P. apetala

Leaf margins serrulate-serrate (rarely crenate), sinuses well developed;

stipules fringed with glandular trichomes; stamens (12-)20-24, disc

>3 mm high; fruit globose. SE Qld, NE NSW. P. lachnocarpa

Pseudoweinmannia apetala (F.M. Bailey)

Engl, in Engl. & Prantl., Nat. Pflanzenfam.

2nd Ed., Vol. 18a: 249 (1930); Weinmannia

apetala F.M.Bailey, Dept. Agric. Brisbane,

Bot. Bull. 8: 76 (1893). Type: Queensland.

Cook District: Kamerunga, January 1892,

E.Cowley 65, ‘the growing shoots from roots,

flower’ (holo: BRI [AQ341025]; iso: K n.v.

[photo at HO!]).

Pseudoweinmannia lachnocarpa auct. non

(F.Muell.) Engl.; Hyland & Whiffin (1993:

131); Cooper & Cooper (2004: 148).

Illustrations: Christophel & Hyland (1993:

pi. 25, fig. C); Cooper & Cooper (2004: 148);

all as P. lachnocarpa.

Tree up to 30 m tall, often prominently

buttressed in larger trees (Hyland & Whiffin

1993). Outer bark flaky ( Sanderson 1634).

Branchlets terete, bark grey with conspicuous

lenticels. Stipules 2-5 mm long, base to 2 mm

wide, margin entire with glandular trichomes

(colleters) and non-glandular strigose

trichomes, glabrous above, caducous. Leaves

opposite, decussate, petiolate, trifoliolate with

two caducous stipels; petiole 1.5-4.5 cm long,

broadly elliptic in cross-section, glabrous to

sparsely strigose; leaflets 5-12.5 cm long,

1.5-4.9 cm wide, sessile or occasionally with

short petiolule, broadly lanceolate, elliptical

to obovate, apex acute to acuminate, rarely

obtuse, bases cuneate to attenuate, margins

typically crenulate to crenate, sinuses weakly

developed; secondary veins 10-16 pairs,

60-80° to midrib and branching well inside

margin. Inflorescence usually axillary,

consisting of a simple raceme, 4.5-12 cm

long, occasionally with one or two racemose

lateral branches, or with some of the lower

flowers replaced by 3-flowered dichasia;

rachis terete, shortly hirsute with crispate

trichomes. Flowers bisexual; pedicels terete,

3-8 mm long, shortly hirsute with crispate

trichomes, abscission point 0.3-0.9 mm above

attachment; sepals valvate in bud, (5-)6(-7),

ovate, elliptical to narrowly ovate, 1.5-2.5

mm long, 0.6—1 mm wide, apex acute, densely

hirsute distally, trichomes more sparsely

distributed below; stamens 12-20; filaments

of unequal length, 2-4 mm long, glabrous,

anthers introrse, 0.4-0.5 mm long; disc

262

Austrobaileya 8(3): 252-266 (2011)

Fig. 6. Seed morphology in Pseudoweinmannia apetala andP. lachnocarpa. A & B. P. lachnocarpa. A. Seed showing

rounded, pyramidal shape and irregular surface and angular margins. B. Detail showing reticulate appearance of the

cell walls. C & D. P. apetala. C. seed showing rounded, pyramidal shape and angular margins. D. Detail showing

somewhat more wrinkled, reticulate appearance of the cell walls. Scale bars A = 1 mm; B,D = 200 pm; C = 0.5 mm. A

& B from Webb & Tracey 11246 (CANB); C&D from Risley 35 (NSW).

colour not noted, c. 0.2 mm high, glabrous,

slightly lobed; ovary subglobular-conical,

0.6-1 mm high, 0.9-1 mm wide, externally

woolly hirsute, 2 (or 3) celled, each cell with

two rows of 3 or 4 ovules, tapering into 2 (or

3) free styles; styles subulate, 1.8-2.8 mm

long, c. 0.2 mm wide at base, glabrous except

for a few trichomes near the base. Fruit ovoid-

globose, 3-4.5 mm high, 4-5 mm wide; seed

brown. Fig. 1.

Additional selected specimens examined : Queensland.

Cook District: T.R. 176, Monkhouse, Oct 1982, Hyland

12175 (CNS); Shiptons Flat on Tin Mine Road, May

1969, Smith 14341 (BRI); E.P 40, S.F.R. 144, Jun 1979,

Sanderson 1634 (CNS); S.F.R. 144, Dagmar, Chowchilla

L.A.,Feb 1995, Gray5978* (CNS); S.F.R. 144 Chowchilla

L.A., Windsor Tableland, Dec 2001, Ford AF3202 &

Jensen (BRI, CNS, HD); S.F.R. 1073, Dulanban, Hartley

L.A., Feb 1993, Hyland 14641* (CNS); near Kuranda,

Jan 1972, Hyland 5795 * (BRI, CNS); S.F.R. 933, Little

Pine L.A., Feb 1979, Gray 1272 (BRI, CNS); S.F.R. 185,

Downfall L.A., Mar 1973, Whiffin & Risley 586 (CNS);

S.F.R. 185, Downfall L.A., Jan 1985, Gray 3848* (CNS);

Downfall L.A., S.F.R. 185, Feb 1972, Risley 35* (BRI,

CANB, CNS, NSW); Atherton District, Dansie 2053

(BRI); Bellenden Ker Range near Babinda, Jan 1979,

Jago 270* (CNS); Downfall Creek Road, near Tinaroo

Dam, s.dat., Webb & Tracey 7646 (BRI); Babinda, May

1997, Jago 634 (CNS). North Kennedy District: Tully

River, rafting access point no. 9, SE of Cardstone, Jun

2001, FordAF2892 (BRI, CNS).

Distribution and habitat: Restricted to north

eastern Queensland from near Cardstone in

the south to Rossville near Cooktown in the

north (Map 1). Hyland & Whiffin (1993)

Rozefelds & Pellow, Revision of Pseudoweinmannia

noted that Pseudoweinmannia apetala (under

the name P. lachnocarpa) occurs from sea

level to 800 m, commonly in drier rain forest,

but also occasionally in wetter rainforest

communities. Cooper & Cooper (2004)

extended the altitudinal range to 950 m.

Phenology: Pseudoweinmannia apetala

flowers from January through to April, rarely

as late as June; with fruiting specimens

collected from December through to March.

Notes : The notes with the type specimen

i.e. “the growing shoots from roots, flower”

suggest that it may have been collected from

coppice growth at the base of the tree.

Etymology: The species name, apetala , comes

from the Greek a , not, and Latin petalum for

petals, which refers to the lack of petals in the

flowers.

Although Cooper & Cooper (2004)

list a common name as ‘Marara’, this

application is derived from populations of

Pseudoweinmannia lachnopetala and the

adoption of this aboriginal name by people

in southern Queensland and New South

Wales (Bailey 1900, 1913; Maiden 1922). No

common names were listed for this species by

Bailey (1900, 1913).

Pseudoweinmannia lachnocarpa (F.Muell.)

Engl, in Engl. & Prantl., Nat. Pflanzenfam.

2nd Ed., Vol. 18a: 249 (1930); Weinmannia

lachnocarpa F.Muell., Fragm. 8: 7 (1874);

Windmannia lachnocarpa (F.Muell.) Kuntze,

Revis. Gen. PI. 1: 228 (1891); Geissois

lachnocarpa (F.Muell.) Maiden, Forest FI.

New South Wales 7:6 (1922). Type: New South

Wales. Tweed, s.dat., [W]Guilfoyle s.n. (lecto

[here designated]: MEL104457; isolecto: K

n.v., photo at HO!; Tweed, s.dat., C.Moore s.n.

(lectopara: MEL104455, MEL104456).

Weinmannia lachnocarpa var. parvifolia

F.M.Bailey, Queensland Agric. J. 28:

196 (1912). Type: Queensland. Moreton

District: Macpherson Range, February 1912,

C.T.White s.n. (holo: BRI [AQ321400]; iso:

NSW104762).

Illustrations: Maiden (1922, t. 229); Francis

(1951: figs 11, 87, 88); Beadle (1976: fig. 150C);

Stanley & Ross (1983: Fig. 36D1,D2); Harden

(1990: 518).

263

Tree up to 40 m tall, often prominently

buttressed in larger trees (Francis 1951;

Beadle 1976). Outer bark grey or fawn pink

usually rough with small scales and pustules

(Floyd 1989). Branchlets terete, bark grey

with conspicuous lenticels. Stipules 2-3.5

mm long, base to 1 mm wide, margin entire

with glandular trichomes (colleters), glabrous

above, caducous. Leaves opposite, decussate,

petiolate, trifoliolate with two caducous

stipels; petiole 1.5-3.5 cm long, broadly

elliptic in cross-section, glabrescent, leaflets

2-13.3 cm long, 0.5-5.8 cm wide, sessile or

occasionally with a short petiolule, broadly

lanceolate, elliptical to obovate, apex acute

to obtuse and ± acuminate, base cuneate to

attenuate, margins serrulate to serrate, rarely

crenate, sinuses typically strongly developed;

secondary veins 11—16(—20) pairs, at 60-80°

to midrib and branching well inside margin.

Inflorescence typically consisting of a simple

raceme, 3-10 cm long, occasionally with one

or two racemose lateral branches, or with some

of the lower flowers replaced by 3-flowered

dichasia, rachis terete, shortly hirsute, with

crispate trichomes. Flowers bisexual; pedicels

terete, 3-8 mm long, with shortly hirsute

crispate trichomes on rachis, abscission point

0.3-1 mm above attachment; sepals valvate

in bud, (5-)6(-7), ovate-elliptical to narrowly

ovate, 2-3 mm long, 0.9-1.2 mm wide, apex

acute, densely hirsute distally and proximally,

with sparse glands on margins; stamens

(12-)20-24; filaments of unequal length 3-5

mm long, glabrous, anthers introrse, 0.5-0.8

mm long; disc red-brown in colour, c. 0.4 mm

high, distinctly lobed; ovary globular, 0.8-1

mm high, 0.9-1 mm wide, externally woolly

hirsute, 2(-3) celled, each cell with two rows

of 2, 3 or 4 ovules, tapering into 2(-3) free

styles, styles subulate, 2-3 mm long, c. 0.35

mm wide at base, glabrous except for a few

trichomes near the base. Fruit globose, 3-5

mm high, 4-7 mm wide; seed light brown.

Fig. 2.

Additional selected specimens examined : Queensland.

Wide Bay District: Tinana Creek, 7 km ENE of Tiaro,

Jan 2005, Forster PIF30579 et al. (BRI); NW slopes of

Mt Glastonbury, Dec 1991, Forster PIF9273 & Sharpe

(BRI, CNS, MEL); S of Anderson Road, 10 km W of

Cooroy, Nov 1993, Bean 7065 (BRI, NSW; AD, DNA

n.v.)- Imbil, Dec 1943, Clemens s.n. (BRI [AQ321401]);

near Imbil, Jun 1947, Smith & Webb 31162 (BRI).

264

Moreton District: Scientific Area 2, S.F. 309, Enoggera,

Jan 1992, Forster PIF9353 & Machin* (BRI, CNS,

MEL, NSW; A, B, K, L, NY, MO n.v); Laceys Creek, Mt

Glorious Road, Taylor Range, Feb 1972, Webb & Tracey

11246 (BRI, CANB); The Rafting Ground, Kenmore, Dec

1981, Dickison 294 & Jessup (BRI); Pimpama River, 3.7

km along Ormeau Road from Pacific Highway, c. 11 km

SE of Beenleigh, Jul 1986, Beesley 8497 & Ollerenshaw

(CANB); c. 2 miles [3.2 km] SW of Canungra, Jan 1971,

Williams s.n.* (BRI [AQ310510], CANB, MEL); Mt

Roberts, McPherson Range, Oct 1967, Blake 22848*

(BRI); Roberts Plateau, McPherson Range, Mar 1920,

White s.n.* (BRI [AQ321398]); O’Reilly’s, Lamington

N.P, Nov 1999, Rozefelds s.n.* (HO); Morans Falls,

Lamington N.P, Aug 1978, Floyd s.n.* (CANB289092).

New South Wales. Pacific Highway - Byron Bay Road,

May 1964, Williams s.n. (NSW); Casino, Dec 1908,

Pope s.n. (NSW).

Distribution andhabitat : Pseudoweinmannia

lachnocarpa extends from near Tinana in

south eastern Queensland to Casino in New

South Wales (Map 1). It occurs in riverine,

coastal or montane rainforest from near sea

level to 1000 m.

Phenology : Pseudoweinmannia lachnocarpa

flowers from October through to January;

with fruiting specimens collected between

November through to March.

Typification : Mueller’s (1874) description of

Weinmannia lachnocarpa cited collections

by both C.Moore and W.Guilfoyle from the

Tweed District of northern New South Wales.

The Guilfoyle specimen was initially selected

by Hoogland in 1984 as the lectotype, and

is an appropriate choice as it is the only

fruiting specimen. A specimen in K, which

is also fruiting, was sent by Mueller and it

matches the colouration and appearance of

the lectotype and is interpreted as a putative

isolectotype.

Notes : The type of Pseudoweinmannia

lachnocarpa var. parvifolia F.M. Bailey

differs from the more typical form of the plant

in having smaller leaves and fewer stamens.

More comprehensive collections, particularly

of flowering material, from throughout

the range of the species are needed to fully

assess this variation although there seems

little justification, at present, to recognise a

separate variety.

Etymology : The species name, lachnocarpa ,

comes from the Greek lachnos , wool and

Austrobaileya 8(3): 252-266 (2011)

karpos , fruit, alludes to the dense indumentum

covering the fruits.

Common names applied to this species

include ‘Marara’, ‘Rose Marara’, ‘Mararie’,

‘Merrany’, ‘ScrubRosewood’, ‘RedCarabeen’

(Bailey 1900, 1913; Maiden 1922; Francis

1951; Floyd 1989).

Acknowledgments

This study has benefited from the detailed

notes compiled by the late Ru Hoogland,

although our conclusions regarding the number

of species occurring in Australia differs from

his. We are grateful to an anonymous referee

and Paul Forster for their helpful advice

and comments on an initial draft of this ms.

We thank the Australian Botanical Liaison

Officer, Ken Hill, for arranging cibachromes

of material of both species in Kew. Andrew

Ford and Bernie Hyland, and Robert

Kooyman assisted with fieldwork in 1998 and

1999 respectively. We thank Phillip Brown,

Department of Agriculture, University of

Tasmania for access to a critical point drier

and David Steele, Central Science Laboratory,

University of Tasmania for assistance with

SEM studies.

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266

Austrobaileya 8(3): 252-266 (2011)

Map 1. Distribution of Pseudoweinmannialachnocarpa

and P. apetala in eastern Australia.

A taxonomic revision of Nyssanthes R.Br. (Amaranthaceae)

A.R.Bean 1 & C.H.Miller 2

Summary

Bean, A.R. & Miller, C.H. (2011). A taxonomic revision of Nyssanthes R.Br. (Amaranthaceae).

Austrobaileya 8(3): 267-279. The endemic Australian genus Nyssanthes is revised, with the

recognition of four species, N. erecta R.Br., N. diffusa R.Br., N. impervia A.R.Bean sp. nov. and N.

longistyla C.H.Mill. sp. nov. All species are illustrated, and distribution maps provided. Notes for

each species are given on conservation status, distribution and habitat, and an identification key is

presented.

Key Words: Amaranthaceae, Nyssanthes, Nyssanthes impervia , Nyssanthes longistyla , Australia

flora, Queensland flora, taxonomy, identification key, new species

'A.R. Bean, Queensland Herbarium, Department of Environment and Resource Management,

Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong 4066, Queensland, Australia. E-mail: tony.

bean@der m. qld. gov. au

2 C.H. Miller, Australian National Herbarium, Centre for Australian National Biodiversity Research,

G.RO. Box 1600, Canberra 2601, Australian Capital Territory, Australia. E-mail: cathy.miller@csiro.au

Introduction

Nyssanthes R.Br. is a small genus endemic to

eastern Australia. All species are perennial

shrubs or subshrubs characterised by their

opposite leaves, 2-locular anthers, 4-merous

flowers, spreading spinose bracteoles, and

tepals of unequal length. Nyssanthes belongs to

Amaranthaceae Subfamily Amaranthoideae,

but its closest affinities are unclear. It shows

superficial similarities to Achyranthes L.,

but pollen morphology suggests an affinity

with Deeringia R.Br. and Celosia L. (Borsch

1998).

Nyssanthes was named by Brown (1810),

from specimens he collected while in

Australia. Brown described three species,

Nyssanthes diffusa, N. erecta and N. media.

Moquin (1849) maintained all of these, but

Bentham (1870) reduced Nyssanthes media

to synonymy with N. diffusa. Little has been

written about the taxonomy of Nyssanthes

since the work of Bentham. Townsend

(1993) included a description of Nyssanthes

in his comprehensive synopsis of the

Amaranthaceae, although this did not expand

beyond that written by Bentham.

The two common species, Nyssanthes

diffusa and N. erecta , have been treated in

state or regional floras (Ross 1983; Jacobs &

Accepted for publication 28 June 2011

Lapinpuro 1990). They are not infrequent as

a component of the understorey in woodlands

and forests of eastern Queensland and New

South Wales. Despite being indigenous,

they are often termed “weeds”, because

they are prickly and may proliferate under

optimal conditions. Two additional species

have recently been recognised from western

Queensland, and they are newly described

here as Nyssanthes impervia A.R.Bean and

N. longistyla C.H.Mill. They inhabit shallow

or skeletal soils in areas that receive low

annual rainfall (280-445 mm).

Discussion of some morphological

characters

Due to the lack of basic knowledge of

Nyssanthes morphology, a number of

misconceptions have been propagated in the

literature. Townsend (1993) mentioned that

all flowers are hermaphroditic in Nyssanthes.

However, the basal flowers formed in the leaf

axils of Nyssanthes diffusa are sterile. They

usually comprise a bract, two bracteoles,

and one tepal. All of the other tepals are

rudimentary or absent, and no male or female

parts can be observed. These sterile flowers

seem to develop in all leaf axils and persist

year-round. Subsequently (and perhaps at

certain times of the year) fully fertile and

bisexual flowers develop. As far as is known,

sterile flowers do not occur in the other

species.

268

Austrobaileya 8(3): 267-279 (2011)

The length of the bract, bracteoles and the

longest tepal vary greatly within each species

of this genus, as Bentham (1870) observed.

However, the variation is not random. The

sterile flowers of Nyssanthes diffusa have

very long spinose bracts and bracteoles, and

sometimes the longest tepal is also of similar

length and is spinose. Subsequent flowers in

an inflorescence display a gradual decrease

in the lengths of the bracts and bracteoles,

even though the change from sterile to fertile

flowers is apparently rapid and without

intermediate forms.

Jacobs & Lapinpuro (1990) stated that

stamen number varies within species of this

genus. We have not found this to be the case.

All flowers of Nyssanthes erecta examined,

have possessed four fertile stamens, and for

N. diffusa consistently two. However, the

anthers are very easily shed, and an erroneous

stamen count can easily occur if the flower is

not opened very carefully.

Materials and methods

This revision is based upon field examination

of all species, and herbarium holdings at AD,

BRI, CANB, CNS, HO, MEL, NSW and

NT, and type specimens from BM and K.

Measurements of leaves, bracts and bracteoles

were made from dried herbarium specimens;

measurements of tepals, stamens and styles

were made from material preserved in spirit

or reconstituted with boiling water.

Common abbreviations in the specimen

citations are N.R (National Park) and S.F.

(State Forest).

Taxonomy

Nyssanthes R.Br., Prodr. 418 (1810).

Type species: N. diffusa R.Br. (lecto: here

designated).

Perennial herbs or shrubs. Leaves opposite,

entire, exstipulate. Inflorescence spicate, in

leaf axils. Rachis present or absent. Flowers

sessile. Bract 1 per flower, bracteoles 2 per

flower, spinose; basal parts cymbiform, more

or less appressed. Sterile flowers sometimes

present. Fertile flowers all bisexual, sessile.

Perianth 4-merous, tepals cymbiform,

heteromorphic, the outermost (posterior) tepal

being the longest, the next (anterior) tepal

being shorter; the two inner tepals shorter

again, but about the same size and shape as

each other. Stamens 2 or 4; anthers bilocular.

Pseudostaminodes present, 4. Ovary 1-locular,

1-ovular. Style not lobed, capitate. Fruit an

indehiscent utricle, dispersed as a unit with

perianth and bracteoles. Seeds 1, ellipsoid to

ovoid.

Etymology : From the Greek nyssa (a prickle),

and anthos (flower), in reference to the prickly

bracteoles and bracts found in all species of

the genus.

Typification: In Brown’s diary the description

of Nyssanthes diffusa has been annotated with

“nov: gen Amaranthor: Infs:”. Additionally the

name was written as “Nyssanthos axillaris”

but this has been crossed out and replaced by

“Nyssanthes diffusa” while the other species

names had not been altered. This would

indicate that Brown had determined the name

of this taxon before he named the other two

taxa (J.G.West pers. comm). For these reasons

Nyssanthes diffusa R.Br. has been selected as

the lectotype species for the genus.

A genus of four species endemic to eastern

Australia.

Bean & Miller, Revision of Nyssanthes

269

Key to the species of Nyssanthes

1 Leaf lamina 2.5-5.5 times longer than broad; leaf apical mucro absent or

up to 0.7 mm long; longest tepal recurved; central Queensland, eastern

New South Wales.1. N. erecta

1. Leaf lamina 1.5-2.5 times longer than broad; leaf apical mucro 0.5-2.1

mm long; longest tepal straight.2

2 Fully expanded leaves sparsely tomentose; inflorescence up to 22-flowered;

stamens two; eastern New South Wales & Queensland.2. N. diffusa

2. Fully expanded leaves moderately to densely tomentose; inflorescence

2-5(-8) flowered; stamens four.3

3 Style 0.6-0.8 mm long; pseudostaminodes without dorsal appendage;

bracts 0.8-1.2 mm wide; southwest Queensland.3. N. impervia

3. Style 27-3.5 mm long; pseudostaminodes with trifid dorsal appendage;

bracts 1.4-2 mm wide; southwest Queensland.4. N. longistyla

1. Nyssanthes erecta R.Br., Prodr. 418

(1810). Type citation: “(J.) v.v.” Type: New

South Wales. Banks of the Nepean [River] on

Badgery’s farm, [December 1804 or January

1805], R. Brown [Bennett No. 3070] (lecto:

BM000795238, here designated; isolecto:

BM000795214).

Dichasially branched perennial forb or

shrub to 0.8 m high. Stems striate, grooved,

sparsely tomentose; hairs antrorse, appressed

to spreading, 0.1-0.5 mm long. Petioles

1-5 mm long; lamina narrowly-elliptic

to narrowly ovate, 14-26 mm long, 3.5-8

mm wide, 2.5-5.5 times longer than broad,

sparsely tomentose when fully expanded;

hairs unicellular, appressed to spreading,

0.15-0.5 mm long; base attenuate, apex

acute or mucronate (mucro 0-0.7 mm long).

Inflorescence up to 16-flowered. Rachis to

8 mm long, sparsely to densely tomentose.

All flowers fertile. Bract 1.3-1.75 mm long,

1-1.3 mm wide; apex acuminate, spinulose,

acumen 0.2-1.5 mm long; base broadly ovate,

hyaline; remaining firmly attached to rachis

upon disarticulation of flower. Bracteoles

2.5-4.5 mm long, 0.9-1.7 mm wide; basal

part ovate to broadly ovate, moderately

tomentose, with thick, yellowish-brown

midrib, otherwise hyaline; acumen slender,

patent, yellowish-brown, spinose; remaining

attached to perianth upon disarticulation of

flower. Perianth 1.4-2 mm wide. Longest

tepal lanceolate-acuminate, 5.5-77 mm

long, recurved, basal half very sparsely to

densely tomentose; midrib prominent, with

1-2 pairs of lateral longitudinal ribs visible;

apex spinose. Next longest (anterior) tepal

slightly shorter but otherwise similar, 4.2-

6.8 mm long. Inner 2 tepals narrowly-ovate,

2.8-3.8 mm long, glabrous; apex acuminate,

spinulose. Fertile stamens 4, filaments

slender, 0.4-07 mm long; anthers 0.35-0.5

mm long, white to yellow. Staminodes absent.

Pseudostaminodes quadrate, glabrous, almost

filling space between filaments, 0.3-0.8

mm high, translucent; distal margin entire

or ciliate; dorsal appendage absent. Ovary

oblate, c. 0.8 mm long, glabrous. Style simple,

0.4-0.6 mm long, stigma capitate. Utricle

with membranous covering, irregularly

fragmenting. Seed obloid, 0.75-1.6 mm long,

0.65-1.1 mm wide, brown. Fig. 1.

Additional selected specimens examined: Queensland.

South Kennedy District: ‘Logan Downs’, Clermont,

Jun 1942, Fairbairn s.n. (BRI [AQ683973]). Mitchell

District: Jericho, Apr 1946, Clemens s.n. (BRI

[AQ683970]); ‘Gartmore’ on Alpha - Tambo road, c.

11 km NNE of Tambo, Apr 1961, Johnson 2185 (BRI).

Leichhardt District: ‘Minerva’, Mar 1935, Blake

7946 (BRI, CANB); ‘Cluden’, south of Taroom, Oct

1954, Johnson & Pedley 17 (BRI); ‘Nirvana’, c. 15

km WNW of Banana, Apr 2003, Bean 20152 (BRI);

SE of ‘Redcliffe’ via Baralaba, Mar 2005, Bean 23609

(BRI). Port Curtis District: Charons Ferry Station

near Styx, Jun 1942, Blake 14606 (BRI); Rosedale,

N.C. [North Coast] Line, Nov 1932, Dovey 164 (BRI);

Experimental Farm, Biloela, Oct 1947, Smith 3550

(BRI). Burnett District: on property of H.J. Brunje,

West Goomeri, Mar 1962, Saint-Smith s.n. (BRI

[AQ178268]). Warrego District: 51 kmN of Charleville

along road to Augathella, W side of road. Mar 1976,

Purdie & Boyland 163 (BRI). Darling Downs District:

270 Austrobaileya 8(3): 267-279 (2011)

Fig. 1. Nyssanthes erecta. A. upper part of flowering plant x l.B. leafy branchlet x 1.5. C. complete flower x 8. D. bract

x 12. E. bracteole x 12. F. bracteole x 12. G. outermost tepal x 12. H. second tepal x 12.1. third tepal x 12. J. innermost

tepal x 12. K. ovary, style, stamens and pseudostaminodes x 16. L. seed x 16. A, C from Bean 20152 (BRI); B from

Bean 23609 (BRI); L from Clemens s.n. (BRI [AQ683970], Del. W.Smith.

Bean & Miller, Revision of Nyssanthes

Chinchilla, Portion 100, ‘Mylo’, Chinchilla Shire, Dec

1989, Mann s.n. (BRI [AQ467453]); Kindon Station,

about 54 miles [90 km] NNE of Goondiwindi, Dec 1938,

Smith 610 (BRI); ‘Woodside’, Pratten, Oct 1962, Walsh

s.n. (BRI [AQ169899]); Bybera, Jan 1934, White 9700

(BRI); Wallumbilla, May 1916, White 9984 (BRI); c. 0.5

km S of Pelican Hall on Chinchilla - Burra Burri Road,

Apr 1981, Wilson 3395 (BRI, NSW). Moreton District:

Crows Nest, Mar 1948, Clydesdale 87 (BRI). New South

Wales. ‘Roma’, near Moree, May 1977, Bradbrook

s.n. (NSW); Stoney Creek Road at South Creek, south

of ‘Berkshire Park’, Dec 1982, Coveny et al. 11473

(NSW); Tenterfield, s.dat., s.coll. (MEL 59864); Bank

of Shoalhaven River, Nowra, Feb 1933, Menaghan s.n.

(NSW); Bullen Buttress, Shoalhaven Gorge, c. 8 km

upstream from the Tallowa Dam site, Feb 1974, Pullen

8775 (AD, CANB, HO, NSW); New England, s.dat.,

Stuart s.n. (MEL59863).

Distribution and habitat : Nyssanthes erecta

is widespread from the Mt Coolon area (west

of Mackay) in Queensland south to Nowra in

New South Wales (Map 1). It grows on heavy

clay soils, often in association with brigalow

(Acacia harpophylla F.Muell. ex Benth.) or

poplar box (Eucalyptus populnea F.Muell.),

but also in a range of other eucalypt-dominated

communities.

Phenology: Flowers and fruits may be found

throughout the year.

Typification: Brown’s diary does not specify

any locality for Nyssanthes erecta. However,

one of the sheets at BM (000795238) has

the locality written in Brown’s handwriting.

Also written next to the name is “Prod. 418”

indicating that Brown probably saw this

specimen. This sheet has been designated

as the lectotype. There is another sheet at

BM (000795214) that matches the lectotype

material but does not have a label in Brown’s

handwriting and with a general location of

Port Jackson. This was probably from the

same collection but has been separated as

there is no Bennett number (J.G. West pers.

comm). This specimen is here designated as

an isolectotype.

Notes: The conspicuously recurved perianth

exhibited by this species is not found in any

other species of Nyssanthes. In all of the other

species, the perianth is straight or slightly

incurved.

Conservation status: A widespread and

relatively common species.

271

2. Nyssanthes diffusa R.Br., Prodr. 418 (1810).

Type: [Queensland. Port Curtis District:].

Pine Mountain, Shoalwater Bay, 3 September

1802, R.Br own [Bennett No. 3069] (holo: BM;

iso: E n.v., K000357277 & 000357276).

N. media R.Br., Prodr. 418 (1810). Type

citation: “(J.) v.v” Type: New South Wales.

Port Jackson, banks of the Hawkesbury,

R.Brown s.n. (lecto: BM000795233, specimen

on right hand side of sheet, here designated;

isolecto: BM000795234 & 000795235).

Divaricately branched herb or shrub to 0.7

m high. Stems striate, grooved, sparsely

tomentose, glabrescent; hairs antrorse,

appressed to spreading, 0.2-0.4 mm long.

Petioles 0.5-3 mm long; lamina broadly

elliptical, 8-35 mm long, 4.5-18 mm wide,

1.5-2.5 times longer than broad, sparsely

tomentose when fully expanded; hairs

unicellular, appressed, 0.2-0.5 mm long; base

cuneate, apex mucronate (mucro 0.5-1.7 mm

long). Inflorescence up to 22-flowered. Rachis

to 10 mm long, with simple non-ciliolate

hairs. Up to 4 basal flowers sterile, subsequent

flowers fertile. Sterile flowers: Bract subulate

throughout and spinose, remaining attached

to perianth upon disarticulation of the

flower, 3.5-7 mm long; bracteoles subulate

throughout and spinose, 5-9 mm long;

longest tepal somewhat expanded at the base,

3-7mm long; all other tepals rudimentary

or absent; stamens and ovary absent. Fertile

flowers: Bract subulate, 1.3-4.5 mm long,

O. 5-1.2 mm wide; basal part ovate to broadly

ovate, hyaline, 0.6-1.5 mm long; acumen 1-

3.5 mm long, spreading, spinose; remaining

attached to rachis upon disarticulation of

flower. Bracteoles 2.1-4.6 mm long, 0.6-1.2

mm wide; basal part ovate to broadly ovate,

with thick, yellow-brown midrib, otherwise

hyaline, sparsely tomentose; acumen slender,

patent, 1-5 mm long, yellowish-brown,

spinose; remaining attached to perianth upon

disarticulation of flower. Perianth 0.9-1.4 mm

wide. Longest (posterior) tepal lanceolate-

acuminate, 5-8.7 mm long, straight, basal half

tomentose on outer surface; midrib prominent,

with 2 or 3 pairs of lateral longitudinal ribs;

apex subulate, spinose. Next longest (anterior)

tepal 2.6-4.9 mm long, straight, sparsely

tomentose on outer surface; apex acute,

272

spinulose. Inner 2 tepals narrowly-ovate, 2.1-

2.9 mm long, sparsely tomentose or glabrous

on outer surface; apex acute, not spinose.

Fertile stamens 2, filaments slender, 0.7-1.1

mm long, often differing in length in the same

flower; anthers 0.25-0.4 mm long, white to

yellow. Staminodes 0-2, 0.5-0.8 mm long,

shorter and thinner than stamen filaments.

Pseudostaminodes quadrate, translucent,

0.4-0.8 mm high, almost filling the space

between filaments, glabrous, distal margin

entire or denticulate, dorsal appendage absent.

Ovary lenticular, 0.3-0.6 mm long, glabrous.

Style simple, 0.3-0.5 mm long, stigma

capitate. Utricle with membranous covering,

irregularly fragmenting. Seed ovoid, 1-1.5

mm long, 0.9-1.1 mm wide, 0.7-0.8 mm deep,

brown. Fig. 2.

Additional selected specimens examined : Queensland.

North Kennedy District: 3 km along Tumoulin Road

from Kennedy Highway Junction, Jan 2004, McDonald

1692 (BRI). South Kennedy District: Eungella Range,

Sep 1938, White 12979 (BRI). Leichhardt District:

Crows Apple Scrub, ‘Rookwood’, Apr 1991, Forster

P1F7977&McDonald(BRiy, Rosedale near Baralaba, Sep

1959, Johnson 920 (BRI). Brigalow Research Station, 20

miles [33 km] NW of Theodore, Mar 1963, Johnson 2632

(BRI); Auburn Range S.F., about 50 km S of Thangool,

Jun 1996, Bean 10363 (BRI); road to Sandstone Bore,

S.F. 46, NW of Taroom, Jan 2003, Bean 19921 (BRI).

Port Curtis District: Bulburin S.F. 67, vicinity of Forest

station, Apr 1980, McDonald et al. 3228 (BRI, CANB).

Burnett District: Bunya Mountains. Near Mowbullan

guest house, Jun 1961, Willis s.n. (MEL). Wide Bay

District: Stony Creek, 4 km E of Didcot, Biggenden

Shire, Dec 1984, Forster P1F1969 (BRI); Glastonbury

district; c. 24 km WSW of Gympie, Apr 1978, Henderson

2623 (BRI). Darling Downs District: Teviot Range, 35

1cm NE of Killarney (SW of Wilsons Peak), May 1971,

Briggs 4210 (NSW). Moreton District: 10 km E of Mt

Nebo, Nov 1980, Dillewaard& Olsen 241 (BRI, CANB);

Table Top, Main Range, Toowoomba, May 1937, Roe

A27 (CANB); Lamington N.P, near turnoff to Moran’s

Falls from O’Reilly’s guest house. May 1981, Willis s.n.

(MEL). New South Wales. 0.5 km along Tucker Box

Road, Breury S.F., 20 km NNW of Urbenville, Nov

1987, Coveny et al. 12810 (CANB, NSW); ‘The Native

Vineyard’ (near Cobbity Trig), 5.5 km N of Cobbity,

Mar 1976, Coveny et al. 7445 (NSW); Waa Gorge,

Mt Kaputar N.P, 68 km NE of Narrabri by road, Nov

1976, Coveny & Roy 9019 (NSW); Yellow-Rock Road,

about 4 miles [6.6 km] SW of ‘Albion Park’, Feb 1967,

Evans 2584 (BRI, CANB, MEL, NSW); Berry, Jan 1929,

George s.n. (NSW); Acacia Creek, Jan 1956, Gray 3787

(CANB); ‘Taroona’, south of Bowling Alley Point, Mar

1995, Hosking 1096 (CANB); Scone, May 1902, Maiden

& Boorman s.n. (NSW); ‘Bajoloma’, Cowal Creek Road,

Bellangry, Sep 1987, Miller & Whaite 458 (CANB);

Between Kiama and Jamberoo, South Coast, Mar 1964,

Austrobaileya 8 ( 3 ): 267-279 ( 2011 )

Pullen 4028 (CANB, NSW); c. 8 km SW of Beelbrook

on Mackenzies Creek Road, Apr 1976, Wilson 1564

(NSW); 5.8 km W of Bingara on Narrabri Road, May

1985, Wilson 6232 (BRI, NSW).

Distribution and habitat : Nyssanthes diffusa

extends from the southern coast of New South

Wales to Ravenshoe in north Queensland.

While most occurrences are within 200 km

of the coast, it does extend as far inland as

Carnarvon Station, south-west of Springsure

(Map 2). It inhabits sunny spots (e.g. along

tracks) in rainforest (especially the Araucarian

vine forests), and sometimes in eucalypt

open forest. When it occurs in eucalypt

communities, it is usually found in a shady

microhabitat.

Phenology : Flowers and fruits may be found

at any time of the year.

Typification : There are three sheets of

Nyssanthes media located at BM. The sheet

BM000795233 has a reference to Brown’s

Prodromus written on the label in Brown’s

handwriting attached to the specimen on

the right hand side of the sheet and is here

designated as the lectotype. The specimen

on the left hand side of the sheet has a

different location (Mount Hunter) and is a

lusher specimen than normal and has been

designated as a residual syntype. The other

two BM sheets (000795234 and 000795235)

do not have labels in Brown’s handwriting

but have other labels indicating Port Jackson

and match the lectotype material (J.G. West

pers. comm.). These are here designated as

isolectotypes.

Notes : The basal (first-formed) flowers of the

inflorescence in this species are usually sterile.

The bract and bracteoles and the posterior

tepal are all long and rigid, and the latter

has only a small hyaline swelling at its very

base. The other tepals are much reduced, and

there are no male or female parts. Bentham

(1870) wrote “sometimes all three bracts are

subulate almost from the base and 3 to 4 lines

long”. This is a reference to the sterile flowers,

although he did not record them as such.

Nyssanthes media has been maintained in

synonymy here following Bentham (1870) as

there are not enough characters to distinguish

it from N. diffusa. Brown (1810) separated

Bean & Miller, Revision of Nyssanthes

273

Fig. 2. Nyssanthes diffusa. A. branchlet with sterile flowers x 2. B. sterile flower x 6. C. branchlet with fertile flowers

x 1.5. D. inflorescence with fertile flowers x 3. E. fertile flower x 12. F. bract x 12. G. bracteole x 12. H. outermost tepal

x 12.1. second tepal x 12. J. third tepal x 12. K. innermost tepal x 12. L. style, stamens and pseudostaminodes x 24. M.

seed x 24. A, B from Bean 19921 (BRI); C-L from Bean 10363 (BRI); M from Johnson 920 (BRI). Del. W.Smith.

274

the two taxa by the number of nerves on the

tepals, the level of pubescence of the plant

and the awn to leaflet ratio. Examination of

the type specimens found that the first two

characters varied considerably and could not

be used to consistently separate the taxa. The

third character is not useful as it is not clear

from his descriptions as to his definition of an

awn and several interpretations are possible.

Bentham (1870) stated that ‘W. media appears

to me to be rather a luxuriant state than a

variety of N. diffusa ”.

Conservation status: A widespread and

relatively common species.

3. Nyssanthes impervia A.R.Bean species

nova afinis N. diffusae , sed foliis hirsutioribus,

inflorescentiis 3-floris, et staminibus fertilibus

4 antheris 0.2-0.3 mm longis, distinguenda.

Typus: Queensland. Gregory South District:

4 km (direct) NNW of‘Trinidad’ Homestead,

E ofWindorah, 1 September 2010, A.R. Bean

30219 (holo: BRI; iso: CANB, K, MEL,

NSW).

Divaricately branched, woody shrub up

to 1 m high. Stems obscurely grooved or

terete, young growth moderately to densely

tomentose; hairs antrorse, appressed to

spreading, 0.15-0.4 mm long. Petioles 0-2

mm long. Lamina broadly elliptical, 5-11 mm

long, 2.5-6 mm wide, 1.8—2.2 times longer

than broad, with dense hairs on both surfaces

when fully expanded; hairs unicellular,

appressed, 0.25-0.6 mm long; base cuneate,

apex mucronate (mucro 0.7-2.1 mm long).

Inflorescences 3-flowered; rachis absent.

All flowers fertile, lateral flowers maturing

before central flower. Bract 6-8.5 mm long,

0.8-1.2 mm wide, remaining attached to

stem upon disarticulation of fruit; basal part

ovate, hyaline, 1.2-1.6 mm long, sparsely

tomentose; upper part subulate, 4.5-7 mm

long. Bracteoles 5.2-7.8 mm long, 1.5-2 mm

wide; basal part ovate, hyaline, 1.3-1.8 mm

long, acumen 4.2-6.5 mm long, spreading,

spinose. Perianth 1.2-1.8 mm wide. Longest

tepal lanceolate-acuminate, 6.5-9.5 mm long,

straight, glabrous, midrib prominent and 2 or 3

pairs of lateral ribs visible; apex spinose. Next

longest tepal lanceolate-acuminate, 5-6.8

mm long, with dense, appressed to spreading

Austrobaileya 8(3): 267-279 (2011)

tomentum on outer surface, apex acute, not

spinose. Inner 2 tepals deflate, 3.5-6.2 mm

long, with moderately dense, appressed to

spreading tomentum on outer surface, apex

acute, not spinose. Fertile stamens 4, filaments

0.2-0.45 mm long; anthers 0.2-0.3 mm long,

white. Staminodes absent. Pseudostaminodes

4, quadrate, c. 0.3 mm long, distal margin

irregular, dorsal appendage absent. Ovary

oblate, c. 0.25 mm long, glabrous. Style

simple, 0.6-0.8 mm long, stigma capitate.

Fruits not seen. Fig. 3.

Additional specimens examined: Queensland. Gregory

South District: 4 km (direct) NNW of ‘Trinidad’

Homestead, E of Windorah, Sep 2010, Bean 30228

(AD, BRI, US); Cheviot Range, c. two-thirds of the

way between ‘Budgerygar’ Homestead and ‘Trinidad’

Homestead (towards Trinidad), Aug 1997, Purdie 4598

(BRI, CANB).

Distribution and habitat: Nyssanthes

impervia is currently known only from the

type location in south-western Queensland

(Map 3). It inhabits shallow sandy soils at the

base of rock outcrops or cliffs, in low open

woodland dominated by Acacia catenulata

C.T.White, A. shirleyi Maiden or Eucalyptus

thozetiana F.Muell. ex R.T.Baker. Other

associated shrubs include Eremophila spp.,

Ptilotus obovatus and Sida spp.

Phenology: Flowers are recorded in August

and September; fruits have not been

collected.

Notes: Nyssanthes impervia is similar to N.

diffusa , but differs by the denser indumentum

of the leaves, the 3-flowered inflorescences

(up to 22-flowered for N. diffusa ), the 4 fertile

stamens in each flower (2 fertile stamens

for N. diffusa), and the anthers 0.2-0.3

mm long (0.25-0.4 mm for N. diffusa). It is

similar stature and appearance to Nyssanthes

longistyla. However, Nyssanthes impervia

differs by the much shorter style (0.6-0.8 mm

long), and the pseudostaminodes that lack a

dorsal appendage.

Conservation status: There are 300-400

plants of this species currently known at the

type locality. Feral goats are present in the

area (J. Silcock pers. comm. Nov 2010) and

the effects of their grazing were evident on

most plants in September 2010. Under the

Bean & Miller, Revision of Nyssanthes

275

Fig. 3. Nyssanthes impervia. A. flowering branchlet x 3. B. immature flower x 9. C. bract x 12. D. bracteole x 12.

E. bracteole x 12. F. outermost tepal x 12. G. second tepal x 12. H. style, stamens, pseudostaminodes and innermost

tepal x 24.1. third tepal x 12. J. innermost tepal x 12. A-E from Purdie 4598 (BRI); F-J from Bean 30219 (BRI). Del.

W. Smith.

276

criteria of the IUCN Red List (IUCN 2001), a

conservation status of Vulnerable is suggested

[VU Blab(v) + 2ab(v); Cl+2a(i,ii); Dl+2],

Etymology : From the Latin impervius

meaning impassable. This is in reference to

the very spiny nature of the plant, making it

impervious to human touch and impassable to

human traffic.

4. Nyssanthes longistyla C.H.Miller species

nova distinguenda a N. erecta habitu lignea;

tepalis duobus exterioribus pilis extus dense

obtectis; tepalis interioribus, filamentis

staminalibus, antheris et stylo longioribus;

pseudostaminodiis multo longioribus,

laciniatis et fimbriatis, dorsaliter squama

trifida praeditis; fructibus multo latioribus.

Typus: Queensland. Gregory North District:

Mt Booka Booka, Elderslie Station, c. 6 km E

of Diamantina River, Boulia - Winton road,

10 June 1997, L.A.Craven, C.L.Brubaker &

J.P. Grace 9915 (holo: CANB; iso: AD, BRI,

DNA, K, MEL, MO, NSW).

Dichasially branched woody shrub up to 1.2

m tall. Stems striate, grooved, on older stems,

young growth densely tomentose; hairs

antrorse, appressed to spreading, 0.3-0.6

mm long. Petioles 0.5-3.5 mm long. Lamina

elliptical, 6-15 mm long, 3-8.5 mm wide,

2-2.5 times longer than broad, moderately

tomentose on both surfaces when fully

expanded; hairs unicellular, appressed to

spreading, 0.5-1 mm long; base attenuate to

cuneate, apex mucronate, (mucro 0.5-1.3 mm

long). Inflorescence with (2-) 5-8 flowers;

rachis to 1 mm long. All flowers fertile. Bract

3.2-6.5 mm long, 1.4-2 mm wide, remaining

firmly attached to rachis upon disarticulation

of fruit; basal part ovate to broadly ovate,

hyaline, 1.2-2.5 mm long, becoming recurved

with maturity, glabrous or with a few hairs on

back near base; upper part subulate, 2.5-5.8

mm long. Bracteoles 4-7.2 mm long, 1.7—

2.3 mm wide; base ovate to broadly ovate,

hyaline, 1.5-2.8 mm long, acumen 2.5-6.5

mm long, spinose; moderately tomentose

along edge of nerve, remaining attached

to perianth upon disarticulation of flower.

Perianth 1.5-2.2 mm wide. Longest (posterior)

tepal lanceolate-acuminate, 6.5-9 mm long,

straight, sparsely tomentose along edge of

Austrobaileya 8(3): 267-279 (2011)

nerve of top half; midrib prominent and 3-5

pairs of lateral longitudinal ribs visible; apex

spinose. Next longest (anterior) tepal similar,

5.5-7.5 mm long, densely tomentose. Inner

2 tepals narrowly-ovate, 5-6.5 mm long,

glabrous; apex acuminate. Fertile stamens 4,

filaments slender, 2.6-3.3 mm long; anthers

1.2-1.7 mm long, cream. Staminodes absent.

Pseudostaminodes 4, quadrate, 1.5-2 mmhigh,

distal margin laciniate, fimbriate translucent,

dorsal appendage trifid with ciliate margins,

0.6-1.1 mm long. Ovary obconical, c. 0.8 mm

long, glabrous. Style simple, 27-3.5 mm long,

stigma capitate. Utricle with membranous

covering, irregularly fragmenting. Seed

ellipsoid, 1.8-2.1 mm long, 1.4-1.5 mm wide,

brown. Fig. 4.

Additional specimens examined : Queensland:

Mitchell District: Mt Felix, c. 90 km N of Jundah on

road to Winton, May 2010, Bean 29779 & Emmott (BRI,

CANB, MEL, MO). Gregory North District: On upper

slopes of Mt Booka Booka, ‘Elderslie’, Oct 1935, Blake

10056 (BRI, NT); Mt Booka Booka, ‘Elderslie’, c. 70 km

W of Winton, Aug 2009, Silcocks.n. (BRI [AQ785536]).

Distribution and habitat : Endemic to

Queensland, where it is known from two

locations in the west of the state (Map 3).

It grows on the upper slopes of mountains

in gravelly shaly soil in association with

other shrubby genera including Senna and

Abutilon.

Phenology : Flowers and fruits have been

recorded in May, August and October.

Notes : This species varies considerably from

the other species of Nyssanthes. It differs from

Nyssanthes erecta by being a woody shrub

sometimes more than a metre tall, densely

covered in hairs all over; leaves are small

and elliptic; the tepals are usually densely

covered with hairs on the outside; the fruits

are much broader; style, staminal filaments,

anthers and pseudostaminodes are much

longer. It is the only Nyssanthes species to

have a trifid appendage on the dorsal surface

of the pseudostaminode. It and Nyssanthes

impervia have a scattered distribution in

inland Queensland, compared to the more

extensive distribution of N. erecta and

N. diffusa throughout large areas of eastern

Queensland and New South Wales.

Bean & Miller, Revision of Nyssanthes

277

Fig. 4. Nyssanthes longistyla. A. flowering branchlet x 4. B. flower at anthesis x 8. C. bracteole x 12. D. bract x 12.

E. outermost tepal x 12. F. second tepal x 12. G. third tepal x 12. H. innermost tepal x 12.1. ovary, style, stamens and

pseudostaminodes x 12. J. pseudostaminode x 16. A-C, E-J from Bean 29779 & Emmott ( BRI); D from Silcock s.n.

(BRI [AQ785536]). Del. W.Smith.

278

Conservation status : There are between

100 and 200 plants present of this species

on Mt Booka Booka (J.Silcock pers. comm.

Aug 2009), and all plants are very heavily

grazed by feral goats. At Mt Felix, there are

more than 1000 plants; here all plants are

in good condition and feral goats are not

present. Applying the criteria of the IUCN

Red List (IUCN 2001), a conservation status

of Vulnerable is proposed. [VU Blab(v) +

2ab(v); Cl; D2],

Etymology : From the Latin longus meaning

long and stylus meaning style in reference to

its long style. The other Nyssanthes taxa have

styles that are 0.3-0.8 mm long, while in N.

longistyla the style is 27-3.3 mm long.

Acknowledgements

We are grateful to Will Smith (BRI) for the

excellent illustrations, Laurie Adams (CANB)

and Peter Bostock (BRI) for the Latin

diagnoses and Nunzio Knerr (CANB) for the

maps. The first author thanks Jenny Silcock

for surveying the Nyssanthes population

on Mt Booka Booka, and Angus Emmott

for assisting with the visit to Mt Felix. The

second author thanks Lyn Craven (CANB) for

visiting Mt Booka Booka and bringing back

specimens to study and Judy West (CANB)

for assistance with nomenclatural matters.

References

Bentham, G. (1870). Nyssanthes. In Flora Australiensis

5: 246-248. L. Reeve & Co.: London.

Borsch, T. (1998). Pollen types in the Amaranthaceae:

morphology and evolutionary significance.

Grana 37: 129-142.

Brown, R. (1810) [1963 fascimile], Nyssanthes. In

Prodromus florae Novae Hollandiae et Insulae

van-Diemen, p. 418. J.Cramer: New York.

Iucn (2001). IUCN Red List Categories: version 3.1.

IUCN Species Survival Commission. IUCN:

Gland, Switzerland.

Jacobs, S.W.L. & Lapinpuro, L. (1990). Amaranthaceae.

In G. Harden (ed.), Flora of New South Wales 1:

248-60. University of New South Wales Press:

Sydney.

Moquin, C.H. (1849). Amaranthaceae. In A.P de

Candolle (ed.), Prodromus Systematis Naturalis

Regni Vegetabilis 13(2): 231-424.

Austrobaileya 8(3): 267-279 (2011)

Ross, E.M. (1983). Amaranthaceae. In T.D. Stanley

& E.M. Ross (eds.). Flora of south-eastern

Queensland 1: 135. Department of Primary

Industries: Brisbane.

Townsend, C.C. (1993). Amaranthaceae. In K. Kubitzki

(ed ). Families and Genera of Vascular Plants 2:

70-91. Springer-Verlag: Berlin.

Map 1. Distribution of Nyssanthes erect a

Bean & Miller, Revision of Nyssanthes

279

Map 2. Distribution of Nyssanthes diffusa

Map 3. Distribution of Nyssanthes impervia m and

N. longistyla ▲

A taxonomic revision of Pterocaulon section Monenteles

(Labill.) Kuntze (Asteraceae: Inuleae-Plucheinae)

A.R. Bean

Summary

Bean, A.R. (2011). A taxonomic revision of Pterocaulon section Monenteles (Labill.) Kuntze

(Asteraceae: Inuleae-Plucheinae). Austrobaileya 8(3): 280-334. Pterocaulon section Monenteles,

comprising all species of the genus from Asia, Malesia, Australia and Melanesia, is revised. 14

species are recognised, including the six new species, P. brachyanthum A.R.Bean, P. ciliosum

A.R.Bean, P. discolor A.R.Bean, P. paradoxum A.R.Bean, P. tricholobum A.R.Bean and P. xenicum

A.R.Bean, and one new combination, P. intermedium (DC.) A.R.Bean. Descriptions, illustrations,

and distribution maps are provided for all taxa. Monenteles sphacelatus and M. spicatus are

lectotypified. The ecology, chemistry and diagnostic morphological characters of members of the

section are discussed, and an identification key to species is provided.

Key Words: Asteraceae, Pterocaulon, Pterocaulon brachyanthum, Pterocaulon ciliosum, Pterocaulon

discolor , Pterocaulon intermedium, Pterocaulon paradoxum, Pterocaulon tricholobum, Pterocaulon

xenicum, taxonomy, Australia flora. New Caledonia flora, Malesia flora, coumarins, identification

key, taxonomy, new species

A.R.Bean, Queensland Herbarium, Department of Environment and Resource Management,

Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. E-mail: tony.

bean@der m. qld. gov. au

Introduction

Pterocaulon Elliott comprises 26 species

distributed in North America, South America,

south-east Asia, Malesia, Melanesia and

Australia. In a revision of the genus by Cabrera

& Ragonese (1978), four taxonomic sections

were recognised. Three of these sections

consist wholly of American species while

the fourth, Pterocaulon section Monenteles

(Labill.) Kuntze, comprises all of the Old

World taxa.

Monenteles Labill. was described at

generic rank by Labillardiere (1825), with two

species, M. sphacelatus Labi 11. andM spicatus

Labill, both named from specimens collected

in New Caledonia. De Candolle (1836) and

Bentham (1867) maintained Monenteles at

generic rank, but the latter commented, “From

the American genus Pterocaulon it scarcely

differs in the hermaphrodite flowers usually

reduced to a single one”.

The reduction of Monenteles to synonymy

with Pterocaulon was effected by Bentham

(1873), and species combinations were made

by Mueller (1882). Monenteles was reinstated

as a section of Pterocaulon by Kuntze (1903).

Anderberg (1989) suggested that the genus

Monenteles could be resurrected, but later

(Anderberg 1991) he abandoned the idea.

The differences between Pterocaulon

section Pterocaulon and P. section Monenteles

are minimal. Nesom (2006) stated that species

in Pterocaulon section Pterocaulon have 1-17

functionally staminate florets per capitulum,

a 1 or 2-seriate pappus, and the hairs have a

relatively long aseptate portion arising from

a cluster of basal cells; while species in P.

section Monenteles have a single staminate

floret per capitulum, a uniseriate pappus, and

the hairs are more-or-less uniformly septate

throughout.

Pterocaulon has been consistently placed

in the Pluchea-growp of genera. The rank of

this group was originally a Subtribe (Subtribe

Plucheinae Benth. & Hook.f. of the Tribe

Inuleae Cass.; Bentham 1873); then it was

deemed a Tribe (Tribe Plucheeae A.Anderb.;

Anderberg 1991, 1994). Evidence from

molecular data now favours subtribal rank

(Anderberg et al. 2005). Within the subtribe,

Anderberg (1991) grouped Pterocaulon

alongside Neojeffreya Cabrera and

Stenachaenium Benth. in the “Pterocaulon

clade”

Accepted for publication 23 August 2011

Bean, Pterocaulon section Monenteles

281

Pterocaulon species are readily identified

by the winged stems formed by decurrent

leaf bases, the sessile capitula grouped into

spiciform, cylindrical or spherical secondary

heads (capitulescences), and the pappus of

capillary bristles. In Pterocaulon section

Monenteles , the secondary heads are usually

aggregated to form a globose or ellipsoidal

capitulescence, and only two or three species

have a spicate or cylindrical capitulescence.

Pterocaulon is sometimes confused with the

related genus Sphaeranthus L., as the latter

also has globose capitulescences and winged

stems. However, Sphaeranthus differs by the

much-reduced capitula and the florets that

lack a pappus.

Cabrera & Ragonese (1978) recognised six

species in Pterocaulon section Monenteles ,

including one species newly described

therein. Wilson (1992) reinstated Pterocaulon

globuliflorum W.Fitzg., so that there were

seven accepted species prior to this study,

viz. P. globuliflorum , P. niveum Cabrera &

A.M.Ragonese, P. redolens (Willd.) Fern.-

Vill., P. serrulatum (Montrouz.) Guillaumin,

P. sphacelatum (Labill.) F.Muell.,

P. sphaeranthoides (DC.) F.Muell. and

P. verbascifolium (Benth.) F.Muell.

Fourteen species are recognised in this

paper. The following species are described as

new; Pterocaulon brachyanthum A.R.Bean,

P. ciliosum A.R.Bean, P. discolor A.R.Bean,

P. paradoxum A.R.Bean, P. tricholobum

A.R.Bean and P. xenicum A.R.Bean.

Monenteles intermedins DC. is reinstated

and transferred to Pterocaulon. Pterocaulon

serrulatum is maintained with two varieties.

Materials and methods

This revision is based on morphological

examination of herbarium specimens from A,

AMES, BRI, CANB, CNS, DNA, GH, K, L,

MEL, NOU and PERTH, and images of type

specimens from B, FI, G, K and P. Specimens

from NSW and P were not available for loan

when this paper was being prepared.

All measurements were made from dried

material. Leaf measurements are of cauline

leaves only, and do not include leaves from

the basal rosette (if present). Leaf length is the

distance from the apex to where the midrib

joins the stem. The width of the stem wings

was measured on the main stem at 20-25 cm

from the apex of the specimen.

Measurements of the dimensions of

capitulescences (secondary heads) reported in

this paper were made at or near anthesis, on

capitulescences where the corolla lobes of the

hermaphrodite floret were open, and where

the inner involucral bracts and the florets were

still attached to the receptacle. After anthesis,

and especially in fruiting plants, the inner

bracts and florets become detached from the

receptacle and form expanded “fluffy” heads,

often two or more times the diameter of the

intact heads. Pappus length was measured on

female florets.

Notes on conservation status have

not been given. Most species are common

and widespread; for the species that may

perhaps be threatened (Pterocaulon discolor ,

P. globuliflorum , P. xenicum ), I have no

knowledge of their extent or population sizes.

The catalogue of Malesian plant collectors

(van Steenis-Kruseman 2006-onwards) was

used for checking the interpretation of label

data of Malesian specimens.

Species treatments are arranged in

alphabetical order. Common abbreviations

used in the specimen citations are N.P

(National Park) and S.F. (State Forest).

Chemistry

Species in the genus Pterocaulon are rich in

chemical compounds, particularly coumarins

and flavonoids. These are thought to have a

biocidal effect against bacteria and fungi,

and to protect against herbivory (Daniel et al.

1999).

Johns et al. (1968) identified 6,7-

dimethoxycoumarin as a major alkaloid

constituent of a Pterocaulon from Inkerman,

Queensland, which they identified as

P. sphacelatum. I have located the specimen

on which this is based. It is held at CANB

(Inkerman, 8 June 1960, W.T.Jones 1531),

and it bears a stamp saying “Voucher for

identity of Bulk Sample No. 6489, Aust.

Phytochemical Survey”. The specimen is

Pterocaulon intermedium.

282

MacLeod & Rasmussen (1999) reported

the identification of a new caryophyllene

from Pterocaulon serrulatum, as well

as three coumarins and a flavanone.

Semple et al. (1998) found that a flavonoid

(chrysosplenol C) they extracted from the

leaves of Pterocaulon sphacelatum has an

antiviral effect, by inhibiting the replication

of rhinoviruses, the most frequent causative

agent of the Common Cold. Pterocaulon

species, especially P. sphacelatum and P.

serrulatum have traditionally been used by

Aboriginal peoples as a medicine, helpful in

the treatment of respiratory infections, colds

and fevers (Barr et al. 1993; Latz 1995).

In Asteraceae Subtribe Plucheinae,

Pterocaulon is the only genus known to

contain coumarins, and around 40 different

coumarins have been found in the 15

investigated species (Stein et al. 2007). These

authors stated that although the species

of Pterocaulon exhibit other classes of

compounds, coumarins are characteristic of

the genus and they postulated that they could

be considered chemotaxonomic markers for

the genus.

Ecology and distribution

Taxa belonging to Pterocaulon section

Monenteles are forbs and shrubs. They do not

occur in rainforest or other densely shaded

areas and are primarily found in sunny

shrublands, woodlands and open forests.

Most species are confined to areas where the

climate is frost-free and the rainfall exceeds

c. 600 mm per year. All species show a

preference for coarse-grained soils, probably

because of the associated good drainage, but

can sometimes be found on finer grained clay-

loams. None however, occurs on cracking

clays.

Many species of Pterocaulon clearly

tolerate salt-bearing wind, as a few species are

found in littoral habitats, not only in Australia,

but also in Indonesia and New Caledonia. In

coastal areas, they can be found close to salt

marsh or mangrove communities, and in the

arid areas close to salt lakes. The label for

Henry 952 (NT), collected from the Simpson

Desert, says that Pterocaulon sphacelatum is

“common in saline soil on surrounding edges

of salt lake”.

Austrobaileya 8(3): 280-334 (2011)

It is presumed that Pterocaulons do not

have strict requirements with regard to soil

acidity or soil chemistry. While they frequently

occur on soils derived from sandstone, where

the pH would be quite low, some species

(e.g. Pterocaulon globuliflorum, P. niveum )

occur on alkaline limestone substrates. In

New Caledonia and near Rockhampton in

Queensland, Pterocaulon spp. occur on

serpentinite, and the soils that develop there

are toxic to most plant species.

As with numerous other species in the

Asteraceae, Pterocaulon species are pioneer

plants. Accordingly, they favour places that

have been disturbed by fire or machinery,

and may occur in large numbers at such sites.

For this reason, they are sometimes regarded

as weeds, but they are readily displaced by

alien species that have a similar ecological

preference.

The life cycle is not known in detail

for any species of Pterocaulon section

Monenteles. All species appear to develop a

rootstock of some kind, and in some species

(e.g. P. intermedium ) it is distinctly woody.

Some species are clearly perennial, living for

three or four years at least; while others are

probably annual resprouters (above-ground

parts dying each year, but resprouting for one

or more years).

Pterocaulon section Monenteles is

distributed in southern China, Burma,

Thailand, Indochina, Malesia, much of

Australia (north of about 30°S), New Caledonia

and Vanuatu. The greatest diversity of

species is in the Kimberley region of Western

Australia and the ‘Top End’ of the Northern

Territory. Only one species {Pterocaulon

brachyanthum) does not occur in Australia.

Discussion of characters

1. Winged stems

Winged stems formed from the decurrent leaf

bases are a constant feature of all species in

the genus, and the generic name is derived

from this attribute. The width of the stem

wings is useful in distinguishing some

species in Pterocaulon section Monenteles.

For example, the stem wings of Pterocaulon

niveum are invariably narrow (usually 0.5-

Bean, Pterocaulon section Monenteles

283

1.5 mm wide), while the stem wings of the

vegetatively similar P. verbascifolium are

broad (usually 2.5-6 mm wide). In Pterocaulon

serrulatum and P. sphaeranthoides the stem

wings are serrated; in other species they are

nearly always entire ( P. sphacelatum and

P. brachyanthum sometimes have sparsely

toothed stem wings).

2. Leaves

Some species, and perhaps all species, begin as

a rosette of leaves at ground level. Knowledge

about this stage of the plant’s life cycle is

poor, as these leaves are rarely represented

on herbarium specimens. In some species, the

cauline leaves are obovate or oblanceolate,

i.e. broadest above the middle, while in

others, they are elliptical, i.e. broadest at

around the midpoint. The upper leaf surface

of most species is bullate to a greater or lesser

extent. The leaf margins are always dentate

to denticulate, though sometimes appearing

entire as the teeth can be obscured by dense

indumentum or by recurved margins.

3. Leaf indumentum

Lanate hairs (uniseriate, multicellular)

(Fig. 1A) are found in all species, but their

density varies greatly, with some taxonomic

correlation. In Pterocaulon verbascifolium

and P. niveum , the lanate hairs are dense

enough to obscure the lamina surface even

under magnification; in P. sphaeranthoides

and P. serrulatum var. serrulatum , the lanate

hairs may be very sparse. The upper leaf

surface often has a sparser indumentum

than the lower, and this is particularly the

case in Pterocaulon discolor. In Pterocaulon

intermedium , the lanate hairs on the upper

surface are as dense as or denser than those

on the lower surface.

Two taxa have broad-based stalked

glandular hairs on the leaves and stem wings

(Fig. IB). Globose sessile glands (often

shining-yellow) are found on the lower leaf

surface of all species, and their density can be

diagnostic for some species (Fig. 1A).

4. Capitulescence (secondary head) shape

and diameter

The capitulescences of Pterocaulon

globuliflorum and P. brachyanthum are

consistently narrower in diameter than other

species in the section.

While head shape varies within species,

it is still quite diagnostic. Three species

{Pterocaulon serrulatum, P. redolens and

P. verbascifolium ) have heads usually

described as cylindrical or spicate. In

Pterocaulon paradoxum, capitulescences

are usually markedly ellipsoidal, though

sometimes approaching globose. Similarly,

the heads of Pterocaulon serrulatum are

sometimes almost globose.

For the remainder, heads are globose or

ellipsoidal; in some species (e.g. Pterocaulon

intermedium) the capitulescences are

always close to globose while in others (e.g.

P. sphaeranthoides , P. sphacelatum) they

may vary from globose to broadly-ellipsoidal,

sometimes on the same plant.

5. Branching patterns and peduncles

Capitulescences are in some species

predominantly sessile, while in others

consistently pedunculate. Peduncle length can

be reliably determined only for overtopped

capitulescences, i.e. those where a side branch

has arisen below it, exceeding it and producing

a further (younger) capitulescence. The

peduncle length is the distance between the

base of the overtopped capitulescence and the

insertion of the lateral branchlet. Pterocaulon

intermedium is notable for its consistently

pedunculate heads. The lateral branchlets

usually protrude at 30-60° and sometimes

at right angles to the subtending branch. In

a few species with sessile capitulescences,

e.g. Pterocaulon tricholobum , they may

continue in more-or-less the same direction

(0-30°), here termed ‘continuous’ branchlets

(e.g. Fig. 19). Where this happens, the sessile

capitulescence appears to be borne laterally

and its shape can be globose-truncate.

284

Fig. 1. Leaf hairs in Pterocaulon. A. P. serrulatum var.

velutinum, showing lanate hairs and sessile glands x 40.

B. P. serrulatum var. serrulatum, showing broad-based

glandular trichomes and sessile glands * 40. A from

Bean 22246 (BRI); B from Fox 25 & Bean (BRI). Del.

W. Smith.

6. Inner involucral bracts

The inner involucral bracts offer some

diagnostic characters. Firstly, the bract length

(only the longest (innermost) bracts are

documented in this paper) may be diagnostic

for some species. Their colour may be white,

straw-coloured or pink, or pink with violet

speckles. The colour is not totally fixed for each

species, but some species have predominantly

Austrobaileya 8(3): 280-334 (2011)

pink bracts, some have predominantly white,

and in others the bracts are straw-coloured.

The upper margins of the inner bracts may

be entire, slightly toothed, strongly toothed or

lacerate, and this is strongly correlated to the

species. For example, in Pterocaulon redolens

the upper margins are entire, tapering to an

attenuate apex; in P. sphacelatum, the upper

margins are often lacerate, and there is no

single bract apex.

7. Cilia

The margins of the inner bracts of every

species have ± straight, hyaline, antrorse

cilia, up to 75% of the bract length. The cilia

of different species may vary in number,

position and length.

This character may be usefully employed

on intact or in-situ heads, i.e. where the

capitula are still attached to the receptacle.

The cilia protrude beyond the outer bracts

and may be seen alongside the inner bracts

and the pappus. The cilia of the bracts are

distinguishable from the bristles of the pappus

because the cilia are not barbellate. The

prominence of cilia varies greatly between

species. In some species e.g. Pterocaulon

tricholobum , P. ciliosum , numerous cilia

are clearly visible on the protruding part of

inner bracts, and in P. ciliosum only, these

cilia can exceed the bract apices; in other

species e.g. P. intermedium , P. sphacelatum ,

P. brachyanthum , very few if any cilia can be

seen on the protruding part of inner bracts

(Fig. 2).

8. Number of female florets per capitulum

The number of female florets is difficult

to determine by casual observation of

the capitulum, and an accurate count is

rather tedious, involving removal of an

intact capitulum and spreading out the

contents so that each element is clearly

visible. However, it can be very useful in

distinguishing between some species in

the section. Three groups of species can be

distinguished using this character; Pterocaulon

redolens , P. serrulatum, P. verbascifolium and

P. xenicum with 26-58 florets; P. ciliosum ,

P. globuliflorum,P. paradoxum,P. sphacelatum ,

P. sphaeranthoides and P. tricholobum with

Bean, Pterocaulon section Monenteles

285

Fig. 2. Single capitula in Pterocaulon. A. P. sphacelatum

x 16. B. P. ciliosum x 12. A from Piercey 51 (DNA); B

from Bean 19221 (BRI). Del. W.Smith.

14-29 florets; P. brachyanthum, P. discolor ,

P. intermedium and P. niveum with 7-17

florets.

9. Florets

There is a single hermaphrodite floret in each

capitulum of every species, except Pterocaulon

verbascifolium and P. sphaeranthoides , which

can sometimes have two.

On this floret, a cluster of globose sessile

glands is consistently present on the outer

surface of each corolla lobe, for all species

except Pterocaulon verbascifolium , in which

these glands are lacking. In some species (e.g.

Pterocaulon tricholobum) the corolla lobes

of the hermaphrodite floret consistently bear

numerous conspicuous eglandular unicellular

trichomes on their outer surface. In other

species (e.g. Pterocaulon intermedium) the

lobes are always glabrous. In some species

(e.g. Pterocaulon ciliosum) the trichomes are

absent on some collections, but present on

others.

The length of the corolla lobes of the

hermaphrodite floret is taxonomically

informative. In one group of species, the

corolla lobes are 0.7-1.2 mm long; in another

group, they are 0.3-0.6 mm long. In only

two species (Pterocaulon intermedium and

P. paradoxum) are the lobes of intermediate

length.

The female florets are very slender, with

an inconspicuously lobed corolla and a 2-fid

style (Fig. 20L). The female corolla length

can be diagnostic for species determination.

10. Achenes and pappus

The length of achenes varies from 0.5-1.2 mm

within the section, with each species showing

a small range of lengths. Twin hairs are found

on the achenes of all species, but the number

of hairs varies between species, and is hence

taxonomically useful.

In a few species, it is possible to find a ring

of transparent globose glands at the junction

of the achene and the pappus (Fig. 11L).

These glands are not consistently present in

any species, but can most frequently be seen

in Pterocaulon redolens. The pappus length

varies between species and is often diagnostic,

286

Austrobaileya 8(3): 280-334 (2011)

moz

HERBARIUM BOGORiENSE

Rojor, lodoaeti

Coll.

No.

K isternan*

dd ?5

Oct.1961

Fain-

Cortpoai..'ia

Gen.

Ptorocnulan

Spec.

Imeel fcu ' .

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Pet.

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v. Suribawn

Lee.

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m. alt.

Note*

On sunny deser

ted field.

. Ike only

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TNi^is. I.nk, Gcn^yc, Sarawak.-Jeha^a, Svtjncy. New YVk. Br. Mus

HnWali. Calcutta. Kepony, Briitbanc, Canberra. Sandakan, China

Smimnon, For, Res Inst Honor.

Please notify Bog or Herbarium of new identification oF this specimen

HbLtrfyp^ Queensland Herbarium (8RI)

f-f&WOuJo*

0et /f.A.de*-' ™*/0/SdZoN

QUEENSLAND

herbarium

354655

BRISBANE

Fig. 3. Holotype of Pterocaulon brachyanthum (Kostermans 19111 [BRI]).

Bean, Pterocaulon section Monenteles

287

but no other pappus characteristics appear to

be of taxonomic utility.

Taxonomy

Pterocaulon section Monenteles (Labill.)

Kuntze, Lex. Gen. Phan. 468 (1903);

Monenteles Labill., Serf. Austro-Caledon. 42,

t. 43, 44 (1825). Type species: Monenteles

spicatus Labill. (= Pterocaulon redolens

(Willd.) Fern.-Vill.,/^ Cabrera & Ragonese

[1978: 202]).

Annually resprouting or perennial forbs or

shrubs, often aromatic. Basal rosette present

in some species. Stems conspicuously

winged due to decurrent leaf margins, wings

extending for 1 and 2 nodes. Leaves alternate,

sessile, decurrent; indumentum of uniseriate,

multicellular hairs; hairs of the upper surface

with short basal cells and relatively long upper

cells; hairs of the lower surface with cell

length relatively uniform throughout; broad-

based glandular hairs sometimes present;

lower surface with yellow, shining, globose

sessile glands. Capitulescences terminal,

monochasially cymose; globose, ellipsoidal,

cylindrical or spiciform; partial clusters

subtended by a bract. Capitula cylindrical,

with involucral bracts in two series; the outer

bracts persistent, lanate-hairy and sometimes

glandular-hairy, all similar in size; the inner

bracts caducous, margins with ± straight,

spreading to antrorse cilia, dorsal surface

glabrous or with tiny translucent sessile

glands, inner whorls slender, outer whorls

shorter and broader. Receptacle epaleate,

glabrous. Outer florets numerous, filiform,

female; corolla three-lobed, style bifid. Disc

floret hermaphrodite, solitary (rarely 2),

tubular, functionally male; corolla broad, 5-

lobed; anthers tailed; style bifid; style branches

with acute sweeping-hairs reaching beyond

the furcation. Achenes narrowly-ellipsoid

to obovoid, with a conspicuous white ring-

shaped carpopodium; twin hairs antrorse,

appressed. Pappus comprising a single row

of barbellate capillary bristles, united at the

base, persistent.

Cytology

The only published cytological record for

Pterocaulon section Monenteles is n = 10 for

Pterocaulon sphacelatum (Watanabe et al.

1999).

Etymology

Monenteles is derived from the Greek

monos meaning ‘single’ and enteles meaning

‘complete, entire or perfect’. This refers to the

single hermaphrodite floret in each capitulum,

surrounded by numerous female florets.

Fourteen species; distributed in Australia,

Melanesia, Malesia and South-east Asia.

Key to the species of Pterocaulon section Monenteles

1 Capitulescences globose-ovoid, spherical to ellipsoidal, 0.7-2.1 times

longer than wide.2

1. Capitulescences cylindrical or spicate, 2.2-5.8 times longer than wide.13

2 Some capitulescences on each plant laterally placed on continuous

branchlets, side branches erect or diverging at less than 30 degrees

(Fig-7).3

2. All capitulescences terminal, side branchlets diverging at 30-90 degrees

(Fig. 5).6

3 Stem wings broad, 2.8-5.5 mm wide .12. P. tricholobum

3. Stem wings narrow, 0.3-2.5 mm wide.4

4 Upper side of leaves with dense to very dense lanate indumentum; inner

involucral bracts pink to violet in dried specimens

6. P. niveum

288 Austrobaileya 8(3): 280-334 (2011)

4. Upper side of leaves with sparse lanate indumentum; inner involucral

bracts greenish-white to white in dried specimens.5

5 Capitulescences 8.5-10 mm wide; pappus 1.3-1.8 mm long .4. P. globuliflorum

5. Capitulescences 11-13 mm wide; pappus 2.6-3.4 mm long .3. P. discolor

6 Stem wings serrate or dentate. 7

6. Stem wings entire or sparsely toothed.8

7 Leaf upper surface conspicuously bullate; few or no cilia visible on

protruding part of inner bracts (Fig. 2A); inner bracts pink, violet or

white.11. P. sphaeranthoides

7. Leaf upper surface not or slightly bullate; many cilia visible on protruding

part of inner bracts (Fig. 2B); inner bracts white or greenish-white . . 9. P. serrulatum

8 Capitulescences 8.5-10 mm wide at anthesis; longest inner bracts 2.8-3.5

mm long; sessile glands on lower leaf surface very sparse.1. P. brachyanthum

8. Capitulescences 10-15 mm wide at anthesis; longest inner bracts 3.4-5.6

mm long; sessile glands on lower leaf surface sparse to dense.9

9 Female florets 8-17 per capitulum; achenes hairs 0-25; leaves elliptic to

lanceolate, broadest near the middle .5. P. intermedium

9. Female florets 16-44 per capitulum; achenes hairs 25-50 or more than

50; leaves oblanceolate to obovate, broadest towards apex.10

10 Corolla lobes of hermaphrodite floret 0.5-0.8 mm long; numerous cilia

visible on protruding part of inner bracts, but none exceeding apices . 7. P. paradoxum

10. Corolla lobes of hermaphrodite floret 0.35-0.6 mm long; cilia on

protruding part of inner bracts either few or absent, or cilia abundant

with some exceeding bract apices.11

11 Stem wings 0.3-0.5 mm wide; longest inner bracts 4.8-5.6 mm long . . . . 14. P. xenicum

11. Stem wings 0.7-3 mm wide; longest inner bracts 3.4-4.6 mm long.12

12 Protrudingpartofinnerbractswithabundantcilia(Fig.2A), some exceeding

bract apices (Fig. 2B); inner involucral bracts straw to white coloured;

leaves 2.1-3.5 times longer than wide.2. P. ciliosum

12. Protruding part of inner bracts with few or no cilia; inner involucral

bracts, white or pink to violet; leaves 3.3-5.8 times longer than wide 10. P. sphacelation

13 Stem wings serrate; achenes 0.7-1 mm long; outer involucral bracts with

broad-based glandular hairs and lanate trichomes outside.9. P. serrulatum

13. Stem wings entire; achenes 0.5-0.7 mm or 1.1-1.3 mm long; outer

involucral bracts with lanate hairs only outside.14

14 Capitulescence 17-30 mm wide at anthesis; hermaphrodite corolla 4-6.5

mm long; achenes 1.1-1.3 mm long.13. P. verbascifolium

14. Capitulescence 8-14 mm wide at anthesis; hermaphrodite corolla 2.5-3

mm long; achenes 0.5-0.7 mm long.8. P. redolens

Bean, Pterocaulon section Monenteles

289

1. Pterocaulon brachyanthum A.R.Bean

species nova affinis P. sphacelato sed

capitulescentiis minoribus, bracteis

interioribus brevioribus, dentibus minoribus

in margine superiore illarum et flosculis

femineis paucioribus in quoque capitulo

differens. Typus: Indonesia. West Nusa

Tenggara Province: Trail from Batudulang

to Punik, Mt. Batulante, west Sumbawa, 25

October 1961, A.J.G.Kostermans 19111 (holo:

BRI; iso: CANB, L).

Shrub 40-70 cm high; stems densely to very

densely lanate; stem wings entire or with

scattered teeth, 0.9-3 mm wide. Leaves

narrowly-elliptic to oblanceolate, 36-55 x

10-22 mm, 2.5-3.6 times longer than wide,

apex acute; margins recurved, denticulate,

with 2-14 pairs of blunt teeth, sometimes

obscure. Indumentum comprising lanate

multi-cellular hairs, glandular hairs absent.

Upper surface with hairs erect to spreading,

sparse to moderately dense, basal cells

short; globose sessile glands absent. Lower

surface with lanate hairs spreading, dense to

very dense, extending 0.4-0.7 mm from the

surface, basal cells not very short; sessile

glands globose, yellow and shining, very

sparse, 8-30 diameters apart. Capitulescence

globose to ellipsoidal, 9—14 mm long and 8.5-

10 mm wide at anthesis, 1-1.5 times longer

than wide; side branchlets never continuous,

at 20-60°; peduncles 2-17 mm long. Outer

involucral bracts 1.5-1.8 mm long, oblong

to oblong-spathulate, with dense spreading

lanate hairs on outer surfaces, inner surface

glabrous or with hairs on the distal end of

inner surface; glands absent; apex obtuse,

erose or truncate. Inner bracts straw-coloured

or white on dorsal surface; longest ones linear,

2.8-3.5 x 0.25-0.5 mm, apex acute, upper

margins with numerous small teeth; cilia

3-9, up to 1.7 mm long, attached along the

proximal half, few extending to the distal one

third, none exceeding the bract apex. Filiform

(female) florets 7-17, colour unknown, corolla

2-2.4 mm long, styles extending 0.3-0.5 mm

beyond corolla. Hermaphrodite floret solitary,

corolla colour unknown, 2.4-2.8 mm long;

corolla lobes 0.3-0.5 mm long, globose sessile

glands present, eglandular trichomes absent.

Achenes narrowly ellipsoidal, 0.5-0.8 mm

long, dark brown, with 0-25 or 25-50 twin

hairs each c. 0.05 mm long; globose sessile

glands at junction between achene and pappus

sometimes present; pappus 2.1-2.5 mm long.

Figs. 3, 4A-D.

Additional specimens examined : Indonesia. Likoeatang

- Taramana, Alor Island, May 1938, Jaag853 (L); Saseel

Island, Kangean Archipelago, Apr 1919, Backer 288635

(L); Karimunjawa Island, north-central Java, Nov 1955,

Hoogerwerf 172 (L); Flores, Lesser Sunda Islands, Sep

1966, Schmutz 355 (L); Soembawa, Nov 1879, Coifs 129

(L); Kambaniroe, Soemba [Sumba] Island, s.dat. [Sep

1873], Teysmann 8824 (A, L); Soemba [Sumba Island],

Lesser Sunda Islands, Nov 1932, Posthumus 3026 (L);

Sumba, Lesser Sunda Islands, Jul 1974, Verheijen 4147

(L); Banyupoh, Bali, Oct 1929, Demandt & VanDillewijn

s.n. (L); Timor, s.dat., Spanoghe s.n. (L); Timor, s.dat.,

Zippelius s.n. (L); Buton Island, south-east Celebes, Sep

1909, Elbert 2598 (L); Timor, Aug 1973, Kooy 957 (K,

L). Timor-Leste, s.loc., 1962-63, Cinatti 210 (L); in

Tasitolu area, just west of Dili, Oct 2006, Cowie 11213

(DNA, L).

Distribution and habitat : Pterocaulon

brachyanthum is endemic to Indonesia and

Timor-Leste. Most collections are from the

Lesser Sunda Islands, east of Java, and from

Timor, but there are two records from islands

north of Java, and one from the south-eastern

end of Sulawesi (Map 1). It inhabits grassy

areas near the coast. One specimen label states

“from the upper margin of mangroves and

salt flat, at the base of a steep shale hilL. The

collections seen were made between altitudes

of 0-200 metres, except the type, which was

from 700 metres.

Phenology : Flowers and fruits are recorded

from April to November.

Affinities : Pterocaulon brachyanthum is

similar to P. sphacelatum, but differs by

the 8.5-10 mm wide capitulescences (10-14

mm wide for P. sphacelatum ); longest inner

bracts 2.8-3.5 mm long (3.5-4.5 mm long for

P. sphacelatum ); upper margins of inner bracts

with small teeth (lacerate or conspicuously

toothed for P. sphacelatum ); female florets

7-17 (17-29 for P. sphacelatum)', and globose

sessile glands very sparse on the lower leaf

surface (dense for P. sphacelatum).

Notes: Cabrera & Ragonese (1978) cited

specimens of the type ( Kostermans 19111 )

from P and NY.

290

Austrobaileya 8(3): 280-334 (2011)

Fig. 4. Pterocaulon brachyantlium. A. hermaphrodite floret x 32. B. inner involucral bract (cilia not shown) x 16.

C. upper portion of inner involucral bract x 32. D. mature achene and basal portion of pappus x 32 . P. ciliosum. E.

hermaphrodite floret x 32. F. inner involucral bract (cilia not shown) x 16. G. upper portion of inner involucral bract

x 32. H. mature achene and basal portion of pappus x 32. P. discolor. I. hermaphrodite floret x 32. J. inner involucral

bract (cilia not shown) x 16. K. upper portion of inner involucral bract x 32. L. mature achene and basal portion of

pappus x 32. A-D from Cowie 11213 (DNA); E,H from Bean 12069 (BRI); F,G from Bean 27963 (BRI); I-L from

Kerrigan 579 (DNA).

Bean, Pterocaulon section Monenteles

291

Distinctive features of this species are

the capitulescences 8.5-10 mm wide; stem

wings often toothed; glands very sparse on

lower leaf surface; female florets few in each

capitulum.

Etymology : From the Greek brachys meaning

short and anthos meaning flower. This is

in reference to both the hermaphrodite and

female florets, which are shorter than in most

other species of the section.

2. Pterocaulon ciliosum A.R.Bean species

nova affinis P. sphacelato sed foliis latioribus,

bracteis interioribus stramineis, ciliis

abundantibus in capitulis in situ visibilibus,

dentibus minoribus in margine superiore in

bracteis interioribus et pedunculis brevioribus

differens. Typus: Australia: Queensland.

Port Curtis District: 7.1 km by road NNW

of Kalpowar Railway Station towards Many

Peaks, 30 August 1975, R.Coveny 6847 &

P.Hind{ holo: BRI; iso: L, NSW).

Illustration : Stanley & Ross (1986: 532), as

P. sphacelatum

Aromatic shrub 40-100 cm high; stems one to

several from non-woody rootstock, sparsely

to densely lanate and sometimes with a few

globose sessile glands; stem wings entire, 1.2-

3 mm wide. Leaves oblanceolate to obovate,

32-56 x 11-21 mm, 2.1-3.5 times longer

than wide, apex acute; margins recurved,

denticulate, with 8-18 pairs of blunt teeth.

Indumentum comprising lanate multi-cellular

hairs, glandular hairs absent. Upper surface

somewhat bullate; hairs erect to spreading,

sparse to moderately dense, basal cells

short; globose sessile glands absent. Lower

surface with lanate hairs erect to spreading,

moderately dense to dense, extending 0.5-1

mm from the surface, basal cells not very

short; globose sessile glands yellow and

shining, dense, touching or up to 3 diameters

apart. Capitulescence globose, 10-16 mm

long and 10-13 mm wide at anthesis, 0.9-

1.2 times longer than wide; side branchlets

never continuous, at 30-90°; peduncles to

7 mm long. Outer involucral bracts 1.8-2.4

mm long, oblong to oblong-spathulate, with

dense spreading lanate hairs on outer surface,

glabrous or with hairs on the distal end of inner

surface; glands absent; apex obtuse. Inner

bracts white to straw coloured (sometimes

tinged pink) on dorsal surface; longest ones

linear, 3.4-4.3 x 0.2-0.4 mm, apex acute,

upper margins with some small teeth; cilia

7-15, up to 2.9 mm long, attached along the

proximal half and consistently extending to

the distal one third, and some exceeding the

bract apex. Filiform (female) florets 16-23,

pink or straw-coloured, corolla 2.2-3 mm

long, styles extending 0.2-0.5 mm beyond

corolla. Hermaphrodite floret solitary, corolla

pink, 2.9-3.5 mm long; corolla lobes 0.35-0.5

mm long, globose sessile glands present,

eglandular trichomes absent or present.

Achenes narrowly ellipsoidal, 0.7-0.9 mm

long, obscurely quadrangular, dark brown,

with 25-50 twin hairs each <0.1 mm long,

evenly distributed; globose sessile glands at

junction between achene and pappus absent;

pappus 2.3-3.1 mm long. Figs. 2B, 4E-H, 5.

Additional selected specimens examined : Indonesia.

Flores, May 1967, Schmutz 1545 (L); Manggarai, Flores,

Jul 1983, Schmutz 5996 (L). Australia: Northern

Territory. Blue Mud Bay, 10 km N of Harris Creek

crossing, Sep 1987, Russeli-Smith 3117 & Lucas (DNA);

Tanumbirini Station, Apr 1991, Wilson 522 (BRI, DNA);

50 miles [80 km] S of Borroloola Road - Anthonys

Lagoon Road junction, Jun 1971, Latz 1539 (DNA); c. 21

km W of ‘Wol logorang’ on the Calvert Hills Road, May

1974, Pullen 9214 (CANB, DNA); 8.7 miles [14.0 km] S of

‘Dunmurra’, Sep 1957, Chippendale 3751 (BRI, CANB,

DNA); Goulburn Island, Anyiminali Point, May 1992,

Coleman 280 (DNA); Caledon Bay, Jun 1972, Latz 2691

(DNA). Queensland. Burke District: 38.5 km NW of

‘Bowthorn’ Homestead on Bowthorn Station, Jun 2006,

Thompson WES745 & Hogan (BRI). Cook District: 4.4

km S of ‘Fairlight’ on the Palmerville Road, Jun 1992,

Clarkson 9607 & Neldner (BRI, CNS, K, MEL); Palm

Cove, Cairns, Jul 1995, Jago 3526 (BRI, DNA); Spear

Creek, c. 1 mile [1.6 km] north of Mt Molloy, Aug 1963,

Schodde 3355 (A, AD, B, BRI, E, L). North Kennedy

District: Hook Island, Whitsunday Group, Jul 1985,

Warrian CW712 (BRI); Mt Abbot, 50 km W of Bowen,

Aug 1992, Bean 4893 (BRI); May’s firebreak, W of

Ravenshoe, Aug 2010, Bean 29858 & McDonald (AD,

BRI, L). South Kennedy District: Brampton Island,

Sep 1988, Batianoff 5114 (BRI); 5.4 km from ‘Redcliffe

Vale’ Homestead on road to Turrawallah, May 1997,

Thompson 266 (BRI). Leichhardt District: Grey Rock,

Glenlea Road, WNW of Springsure, Jul 1989, O ’Keeffe

894 (BRI); 10 km S of Isla Gorge lookout, 37 km S of

Theodore, Jun 1971, Johnson 7169 & Briggs (BRI,

NSW). Port Curtis District: Hill W of Mt Couti-Outi,

Stanage Bay area, N of Rockhampton, Jul 1997, Bean

12069 (BRI); 4 km SW of Mt Castletower, c. 40 km SW of

Gladstone, Jun 1977, Crisp 2739 (BRI, CANB). Burnett

District: Near junction of Delubra and Cadarga Creeks,

35 km SW of Mundubbera, Sep 2008, Bean 27963 (BRI).

292

Austrobaileya 8(3): 280-334 (2011)

Ex- NATIONAL HERBARIUM OF NEW SOUTH WALES

ROYAL BOTANIC GARDENS, SYDNEY

313281

Pterocaulon sphacelatum (Labi11.) F.Muell,

Lot. 24 * 38 'S Long. 151 0 17 ' E

LocJ.lka (4.4mls) by toad NNW of Kalpower

railway atn towards Many Fe-'ks, Old .

Coll. H. Coveny 6847 & P. Bind D °t e 30.Tiii.191

Notes Herb 50 —6 Oct high with mauve flowers,

scattered in sandy creek bed beneath Caauarina

cumin j&amlana . Melaleuca , etc.

Ho£ja~rf P ^ ef C[ueeT ' slan(1 n erbarium ( BR| )

Pfcrocjzuio/L- CthofUmr A. A. &4r\

///^

247327

Brisbane

Fig. 5. Holotype of Pterocaulon ciliosum (Coveny 6847 & Hind [BRI]).

Bean, Pterocaulon section Monenteles

293

Wide Bay District: Fairlies Knob, Seaview Range, May

2000, Phillips 389 & Phillips (BRI); Western edge of

Mt Walsh N.R, near Biggenden, Sep 2002, Bean 19221

(BRI). Moreton District: Upper Ithaca Creek, Aug 1887,

Simmonds s.n. (BRI); Mermaid Mt, NW of Brookfield,

May 1970, Telford 1590 (CANB).

Distribution and habitat: Pterocaulon

ciliosum is a widely distributed species. It

has been collected on the island of Flores

in Indonesia; it occurs in the north-east of

Northern Territory (extending to far north¬

west Queensland), and is widespread in

eastern Queensland from Murray Island in

the Torres Strait to Brisbane (Map 2). It is

found in eucalypt woodland, mainly on well-

drained sandy to loamy soils, including sites

near the ocean.

Phenology : Flowers and fruits have been

recorded from April to October.

Affinities: Pterocaulon ciliosum is similar

to P. sphacelatum , but differs by the leaves

2.1-3.4 times longer than wide (3.3-5 times

for P. sphacelatum ); the straw-coloured

inner bracts on dried material (white or

pink for P. sphacelatum ); abundant cilia

visible on the protruding part of inner bracts

(few or no visible cilia for P. sphacelatum );

the upper margins of the inner bracts with

small teeth (strongly toothed or lacerate for

P. sphacelatum)., and the peduncles to 7 mm

long (to 22 mm long for P. sphacelatum).

Notes: Distinctive features for this species are

the abundant cilia visible on the protruding

part of inner bracts, straw-coloured inner

bracts on dried material and the short corolla

lobes of the hermaphrodite floret.

Etymology: The specific epithet is from the

Latin cilia meaning ‘full of cilia’ and refers

to the numerous cilia on the margins of the

inner bracts.

3. Pterocaulon discolor A.R.Bean species

nova affinis P. tricholobae sed alis caulinis

angustioribus, foliis valde discoloribus,

foliorum dentibus paucioribus, flosculis

femineis paucioribus in capitulo quoque

numeris majoribus pilorum in achenis

differens. Typus: Australia: Northern

Territory. 3 km N of Adelaide River bridge on

Stuart Highway, 3 May 1983, J.D. Briggs 819

(holo: CANB; iso: MEL).

Forb 50-80 cm high, aromatic properties

unknown; stems one to several from woody

rootstock, densely lanate but without globose

sessile glands; stem wings entire, 1-2.5 mm

wide. Leaves elliptical, 35-68 x 13-31 mm,

2-2.7 times longer than wide, apex obtuse or

mucronate; margins not recurved, denticulate

to crenulate, with 10-16 pairs of small teeth.

Indumentum comprising lanate multi-cellular

hairs, glandular hairs absent. Upper surface

not or slightly bullate; hairs erect to spreading,

sparse, basal cell short; globose sessile glands

absent. Lower surface with lanate hairs erect

to spreading, very dense, obscuring surface,

extending 0.2-0.3 mm from the surface, basal

cells not very short; globose sessile glands not

visible. Capitulescence globose to ellipsoidal,

12—15(—19) mm long and 11-13 mm wide at

anthesis, 1—1.2(—1.5) times longer than wide;

side branchlets sometimes continuous, at 0-

60°; peduncles absent or up to 12 mm long.

Outer involucral bracts oblong-spathulate,

2.1-2.4 mm long, with dense spreading lanate

hairs on outer surface, sparse to moderately

dense hairs on the distal end of inner surface,

glands few, apex obtuse. Inner bracts white or

greenish-white on dorsal surface; longest ones

linear, 4.5-5 x 0.3-0.4 mm, apex acute, upper

margins entire or with some small teeth; cilia

8-12, up to 3.3 mm long, attached along the

proximal half and consistently extending to

the distal one third, not exceeding the bract

apex. Filiform (female) florets 7-14, white

or straw-coloured, corolla 27-3.2 mm long,

styles extending 0.3-07 mm beyond corolla.

Hermaphrodite floret solitary, corolla pink,

3.3-4 mm long; corolla lobes 07-1.1 mm long,

globose sessile glands present, eglandular

trichomes 1-5 per lobe. Achenes narrowly

ellipsoidal, 0.8-0.9 mm long, faintly ribbed,

dark brown, with 25-50 twin hairs each

0.1-0.2 mm long, evenly distributed; globose

sessile glands at junction between achene and

pappus absent; pappus 2.6-3.4 mm long. Figs.

4I-L, 6.

Additional specimens examined : Australia: Northern

Territory. Tipperary, May 1963, Muspratt 575 (DNA);

0.5 miles [0.8 km] E of Stuart Highway, 60 miles [98

km] [from Darwin?], Apr 1964, Robinson R343 (DNA);

Nitmiluk N.R, eastern boundary. May 2002, Michell

3875 (DNA); Nitmiluk N.R, Apr 2001, Michell 3329

(DNA); Nitmiluk N.R, Edith Falls, May 2002, Kerrigan

294

Austrobaileya 8(3): 280-334 (2011)

AUSTRALIAN NATIONAL HERBARIUM

mi >iiinii!i ;ihmi mi ii

CBG 8307528

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HERBARIUM. CANBERRA BOTANIC GARDENS

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Fig. 6. Holotype of Pterocaulon discolor (Briggs 819 [CANB])

Bean, Pterocaulon section Monenteles

295

579 (DNA); Tin Camp Creek, c. 20 miles [32 km] S

of Nabarlek mining camp. May 1973, Hartley 13824

(CANB [2 nd sheet only], DNA, L); 11 km E of Katherine

Gorge ranger station. May 2009, Latz 24367 & Quarmby

(DNA, K, NT).

Distribution and habitat : Pterocaulon

discolor is known from a few scattered

locations in the ‘Top End’ of the Northern

Territory, including Nitmiluk N.P. (Map 3).

It grows on sandstone and quartzite hills and

ridges in mixed open forest or open woodland,

in sandy soil.

Phenology : Flowers and fruits have been

collected in April and May.

Affinities: Pterocaulon discolor is similar

to P. tricholobum, but differs by the stem

wings 1-2.5 mm wide (2.8-5.5 mm for

P. tricholobum ); the lanate indumentum of the

lower leaf surface 0.2-0.3 mm high (0.4-0.8

mm high for P. tricholobum ); the 10-16 pairs

of readily observed leaf teeth (15-30 pairs

of obscure leaf teeth for P. tricholobum)., the

7-14 female florets per capitulum (15-21 for

P. tricholobum ); and the 25-50 hairs on the

achene (0-25 hairs for P. tricholobum).

Notes : This species is distinctive in the

strongly discolorous leaves, few female florets

per capitulum, conspicuous leaf teeth, white or

greenish-white inner bracts and the trichomes

on the corolla lobes of hermaphrodite floret.

Etymology : From the Fatin discolor , meaning

‘of different colours’. This refers to the leaves,

which in this species are strikingly green on

the upper surface and snowy white on the

lower surface.

4. Pterocaulon globuliflorum W.Fitzg., J.

& Proc. Roy. Soc. Western Australia 3: 223

(1918), (as “ globuliflorus ”). Type: Australia:

Western Australia. Devil’s Pass, Napier

Range, May 1905, W.VFitzgerald 607 (holo:

PERTH; iso: K373285; NSW n.v).

Shrub to 60 cm high; aromatic properties

unknown; stems sparsely to densely lanate and

with a few globose sessile glands; stem wings

entire, 0.3-1.5 mm wide. Feaves elliptical,

18-51 x 7-24 mm, 2-2.6 times longer than

wide, apex acute or obtuse; margins not

recurved, denticulate, with 7-15 pairs of blunt

teeth. Indumentum comprising lanate multi¬

cellular hairs, glandular hairs absent. Upper

surface not or slightly bullate, hairs sparse,

flexuose, basal cells not short; globose sessile

glands absent. Fower surface with lanate

hairs flexuose, dense to very dense, extending

0.15-0.25 mm from the surface, basal cells not

very short; globose sessile glands yellow and

shining, dense, touching or up to 3 diameters

apart, but usually obscured by indumentum.

Capitulescence globose or globose-truncate,

6—10 mm long, 8.5-10 mm wide at anthesis,

0.7-1.1 times longer than broad; sidebranchlets

0-50°, often continuous, peduncles 0-4 mm

long. Outer involucral bracts oblong, 0.8-1.9

mm long, with dense spreading lanate hairs

on the distal one-half of outer surface, and

glabrous on the inner surface; apex obtuse or

truncate. Inner bracts white on dorsal surface,

sessile glands absent; longest ones linear, 3.1-

3.5 x 0.2-0.25 mm, apex acute, upper margins

with small teeth; cilia 5-8, up to 1.6 mm long,

mostly attached proximally and extending to

the distal one third. Filiform (female) florets

16-20, colour unknown, corolla 1.7-2 mm

long, styles extending 0.3-0.5 mm beyond

corolla. Hermaphrodite floret solitary, corolla

pink, 2.5-2.8 mm long; corolla lobes 0.7-1.2

mm long, globose sessile glands present,

eglandular trichomes absent or with 1 or 2 per

lobe. Achenes narrowly ellipsoidal, 0.7-0.8

mm long, slightly longitudinally ribbed, dark

brown, with 25-50 twin hairs each c. 0.05

mm long, evenly distributed; globose sessile

glands at junction between achene and pappus

absent; pappus 1.3-1.8 mm long. Figs. 7, 8A-

Additional specimens examined : Australia: Western

Australia. Windjana Gorge, Napier Range, Jul 1974,

Carr 3882 & Beanglehole 47660 (PERTH); Windjana

Gorge, Napier Range, Jul 1974, Carr 4036 & Beanglehole

47814 (PERTH); Yammera Gap, Napier Range, 9 km NE

of Lennard River crossing on Gibb River Road, May

1988, Streimann 8269 (A, B, CANB, L); Old Fossil

Downs Station, May 1970, Wolfe & Martin 72 (CANB).

Distribution and habitat : Endemic to the

Napier and Geikie Ranges, Kimberley region,

Western Australia (Map 3). It is confined

to limestone ranges and hills, growing in

grassland or low open woodland.

Phenology: Flowers and fruits have been

recorded in May and July.

296

Austrobaileya 8(3): 280-334 (2011)

Flora of Australia

AUSTRALIAN NATIONAL BOTANIC GARDENS

HERBARIUM (CBG)

Notificatjon of change of determination would be appredated by CBG

AMARANTHACEAE

AUSTRALIA. WESTERN AUSTRALIA. Yammera Gap.

Napier Range, 9 Km NE of Lennard River crossing on Gibb

River Road.

c6A6

PUfocavalon ntoeuifn C a W<

Queensland Herbarium (BRI)

j/oSaJ/f/ort*, &

Det 20 //

17° 20' S 124" SITE 70 m alt.

Grasslands at base of limestone range with scattered small

shrubs.

Open spreading herb to 40 cm tall. Inflorescence white;

leaves dull dark green above, grey-green below.

H. Strelmann 8269 IS May. 1988

Dupl.t (6) L, DIVA, A. PERTH. l^E. B

Fig. 7. Representative specimen of Pterocaulon globuliflorum (Streimann 8269 [CANB]).

Bean, Pterocaulon section Monenteles 297

Fig. 8. Pterocaulon globuliflorum. A. hermaphrodite floret * 32. B. inner involucral bract (cilia not shown) x 16. C.

upper portion of inner involucral bract x 32. D. mature achene and basal portion of pappus x 32. P. intermedium. E.

hermaphrodite floret x 32. F. inner involucral bract (cilia shown) x 16. G. upper portion of inner involucral bract x 32.

H. mature achene and basal portion of pappus x 32. A-D from Carr 4036 & Beauglehole 47814 (PERTH); E-G from

Bean 12186 (BRI); H from Clarkson 4854 (BRI).

Affinities : Pterocaulon globuliflorum appears

to be allied to P. tricholobum and P. niveum ,

on the basis of the long corolla lobes of

the hermaphrodite floret, and the often

‘continuous’ branchlets giving rise to lateral

capitulescences.

Typification: Fitzgerald wrote the location

‘Devil’s Pass’ on the label of his specimen. In

the protologue, he used the locality ‘Wingrah

Pass’. According to an annotation by Annette

Wilson on the holotype, both of these terms

are equivalent to Windjana Pass or Windjana

Gorge.

Notes : Distinctive features for this species

include the small capitulescences (8.5-10 mm

diameter) that are often lateral; leaves short

and broad, 2-2.6 times longer than wide,

strongly discolorous; stem wings very narrow

and the very short pappus.

298

5. Pterocaulon intermedium (DC.) A.R.Bean

comb, nov.; Monenteles intermedins DC.,

Prodr. 5: 456 (1836). Type: [Australia:

Queensland. North Kennedy:] Sandy shores of

Cape Cleveland, 14 June 1819 , A.Cunningham

52 (holo: G-DC [G214737]; iso: K373284).

Monenteles globiferus DC., Prodr. 5: 455

(1836). Type: [Australia: Queensland.

Moreton District:] Lockyer’s Creek, Brisbane

River, Moreton Bay, July 1829, A. Cunningham

51 (holo: G-DC [G214749]; iso: K373282).

Pterocaulon sp. A; Wilson (1992: 954).

Pterocaulon sp. A Kimberley Flora (B.J.Carter

599); Paczkowska & Chapman (2000: 172).

Pterocaulon sp. (Yarrowmere Station E.J.

Thompson+ BUC340); Bostock & Holland

(2007: 30); Bostock & Holland (2010: 25).

Illustration : Kenneally et al. (1996: 67), as

‘Pterocaulon sp. A, Kimb. Flora’

Aromatic perennial shrub 20-60 cm high;

stems sprouting from a woody rootstock,

sparsely to densely lanate and sometimes

with a few globose sessile glands; stem wings

entire, 0.2-1.4 mm wide. Leaves lanceolate,

narrowly-elliptic or elliptic, 12-52 x 3.5-

18 mm, 2.6-4.6 times longer than wide,

apex acute; margins not recurved, entire or

denticulate, with 2-20 pairs of blunt teeth.

Indumentum comprising lanate multi-cellular

hairs, glandular hairs absent. Upper surface

not bullate, hairs moderately dense to very

dense, erect near base then antrorse, basal cells

short; globose sessile glands absent. Lower

surface with lanate hairs erect to spreading,

moderately dense to dense, extending 0.5-1

mm from the surface, basal cells not very

short; globose sessile glands yellow and

shining, dense, touching or up to 3 diameters

apart. Capitulescence globose or almost so,

10-15 mm long, 10-14 mm wide at anthesis,

1-1.2 times longer than broad; side branchlets

never continuous, at 30-90°; peduncles (5-)7-

25 mm long. Outer involucral bracts oblong

to spathulate, 1.7-2.4 mm long, with dense

spreading lanate hairs on the distal one-third

of outer surface, and glabrous or with a few

hairs distally on the inner surface, glands

absent; apex acute, obtuse or truncate. Inner

Austrobaileya 8(3): 280-334 (2011)

bracts pink to violet on dorsal surface; longest

ones linear, 4-4.8 x 0.2-0.4 mm, apex acute,

upper margins with small teeth; cilia 1-7, up

to 2.8 mm long, mostly attached proximally

and not extending to the distal one third.

Filiform (female) florets 8-17, pink, corolla

27-3.6 mm long, styles extending 0.3-0.8

mm beyond corolla. Hermaphrodite floret

solitary, corolla pink, 3.1-3.8 mm long; corolla

lobes 0.6-0.9 mm long, globose sessile glands

present, eglandulartrichomes absent. Achenes

narrowly ellipsoidal, 0.6-0.75 mm long,

longitudinally ribbed, dark brown, glabrous

or with 1-25 twin hairs each <0.1 mm long,

evenly distributed; globose sessile glands at

junction between achene and pappus absent;

pappus 2.5-3.1 mm long. Figs. 8E-H, 9.

Additional selected specimens examined : Indonesia.

Kisar Island [just N of Timor], Apr 1939, Bloembergen

3852 (A, L). Papua New Guinea. Morobe Province:

Sialum Station, May 1975, Henty & Katik NGF49770

(BRI, CANB, L). Australia: Western Australia. Fence

line at junction of Pindan, Calanjadie and Cardingy

paddocks, Anna Plains Homestead, Sep 2004, Byrne 1266

(PERTH); Sir Graham Moore Island, Jul 1973, Wilson

11263 (PERTH); adjacent to Telecom tower, c. 5 km E

of Broome, Oct 1993, Dureau DD150 (BRI, PERTH);

4 km SSE of Cape Bertholet, Dampier Peninsula, Aug

1992, Carter BJC599 (PERTH); Pender Bay at Weedong

Lake, Dampier Peninsula, Jun 1984, Forbes 2435 &

Kenneally (L, MEL). Northern Territory. Elcho Island,

Warangaya, Sep 1987, Russell-Smith 3282 & Lucas (BRI,

DNA); Elcho Island, Refuge Bay, Aug 1995, Cowie 6002

(BRI, CANB, DNA, MEL); Howard Island, Jun 1996,

Booth 1789 (DNA); Maria Island, Gulf of Carpentaria,

Jul 1972, Dunlop 2803 (BRI, DNA); Homestead Creek,

Bing Bong Station, Jun 1971, Dunlop 2245 (DNA).

Queensland. Burke District: Karumba, Jun 1901,

Bailey s.n. (BRI [AQ270420]); Pandanus Ridge, 20 km

NE of Burketown, May 2005, Booth 4281 & Thompson

(BRI); Settlement Creek, 2 miles [3 km] from coast.

Gulf of Carpentaria, Jun 1948, Perry 1217 (BRI, CANB,

DNA); Sweers Island, South Wellesley group. Gulf of

Carpentaria, Nov 2002, Thomas SWI111 (BRI). Cook

District: Lakefield N.P, Jane Table Hill, Jun 1983,

Clarkson 4854 (BRI); near California Creek, NW of Mt

Garnet, Aug 1997, Bean 12186 (BRI); 1.5 km N of the

North Kennedy River on the Laura - Musgrave Road,

May 1989, Clarkson 7996 & Neldner (BRI, DNA, K,

L); on southern bank of Nassau River 0.5 km upstream

from junction with Rocky Creek, Jun 1990, Neldner

3001 (BRI, L). North Kennedy District: 15 miles [25

km] NW of Bowen towards Townsville, May 1970, Fagg

698 (CANB, L). South Kennedy District: 22.5 km

5 of ‘Yarrowmere’ Homestead, Apr 1992, Thompson

BUC340 & Simon (AD, BRI). Port Curtis District:

‘Galloway Plains’, c. 28 km SW of Calliope, Mar 1989,

Anderson 4623 (BRI); S.F.60 Rundle Range, 37 km NW

of Gladstone, Aug 1989, Gibson TOI683 (BRI). Burnett

Bean, Pterocaulon section Monenteles

299

yueensiana neroanum idkij

QUEENSLAND HERBARIUM (BRI)

Fkxa of Queensland Burt<e

Pterocaulon sphacelatum (Labill.) F.Muell.

Pfetioctud*^ A A, Scan

Dei - A*. 6e^ Dale to 61 2 * 1 ,

QUEENSLAND HERBARIUM (BRI)

Brisbane Australia

7323

Fig. 9. Representative specimen of Pterocaulon intermedium (Booth 4281 & Thompson [BRI]).

300

District: Brian Pastures near Gayndah, Mar 1952, Blake

18894 (BRI); Wetheron, Apr 1980, Forster PIF739 (BRI).

Darling Downs District: ‘Rockwood’, 60 miles [98 km]

W of Dalby, Apr 1969, Dunlop 337 (CANB). Moreton

District: Ipswich, 1909, Hall s.n. (BRI [AQ270443]).

Distribution and habitat : Pterocaulon

intermedium is widely distributed in northern

Australia from Port Hedland (Western

Australia) to Ipswich (Queensland). It is often

found close to the coast, but it may occur up to

250 km inland. Outside Australia, it is known

from the island of Kisar in Indonesia, and

from one location in Papua New Guinea (Map

4). In coastal areas it favours sandy swales or

dunes, in shrubland or low woodland, and is

sometimes adjacent to saline tidal areas. In

the more inland areas, it grows in eucalypt

woodland with sandy soil.

Phenology: Flowers and fruits are recorded

from April to October.

Affinities: Pterocaulon intermedium is

perhaps closest to P. paradoxum, but differs

by the leaves being broadest near the midpoint,

the upper surface not bullate, the indumentum

of the upper surface as dense as or denser than

the lower, the shorter achenes and the fewer

female florets per capitulum.

Typification: The original label of the

holotype of Monenteles intermedius gives the

date of collection as “June 14 1829”. This is

incorrect. The isotype at K bears the correct

date of 14 June 1819, as confirmed by Curry

& Maslin (1990).

Notes: Distinctive features for Pterocaulon

intermedium are the narrow stem-wings;

consistently long peduncles; a consistently

± globose capitulescence, inner bracts pink

or violet; relatively few female florets per

capitulum; elliptical leaves; upper leaf surface

indumentum as dense as or denser than lower

surface; and few hairs on the achene.

One specimen label reports that Pterocaulon

intermedium has an “apple juice smell”.

6. Pterocaulon niveum Cabrera &

A.M.Ragonese, Darwiniana 21: 249 (1978).

Type: Australia: Western Australia. Mt

Amherst, 13 May 1951, C.A.Gardner 10206

(holo: PERTH; iso: SI).

Austrobaileya 8(3): 280-334 (2011)

Illustration: Cabrera & Ragonese (1978:

248).

Spreading shrub 40-100 cm high; stems very

densely lanate; stem wings entire, 0.5-1.5

(-2.5) mm wide. Leaves elliptical or obovate,

39-70 x 21-38 mm, 2.1-2.6 times longer

than wide, apex acute or obtuse; margins not

recurved, denticulate, with 9-20 pairs of blunt

teeth. Indumentum comprising lanate multi¬

cellular hairs, glandular hairs absent. Upper

surface with hairs erect to spreading, very

dense; globose sessile glands absent. Lower

surface with lanate hairs erect to spreading,

very dense, extending 0.5-1 mm from the

surface; globose sessile glands sparsely

present, but hidden by hairs. Capitulescence

globose to globose-truncate, 10-15 mm long

and 10-15 mm wide at anthesis, 0.7-1.1 times

longer than wide; side branchlets sometimes

continuous, at 0-70°; peduncles 0-8 mm

long. Outer involucral bracts 2.2-3.2 mm

long, oblong to oblong-spathulate, with dense

spreading lanate hairs on outer surface, inner

surface glabrous; glands absent; apex acute or

obtuse. Inner bracts pink to violet on dorsal

surface; longest ones linear, 4.5-5.4 x 0.2-

0.35 mm, apex acute; upper margins entire or

with a few small teeth; cilia 8-14, up to 3 mm

long, attached mainly along the proximal half

and consistently extending to the distal one

third, not exceeding the bract apex. Filiform

(female) florets 11-17, pink, corolla 3-3.7 mm

long, styles extending 0.4-1.2 mm beyond

corolla. Hermaphrodite floret solitary, corolla

pink, 3.3-4 mm long; corolla lobes 0.7-1.1

mm long, globose sessile glands present,

eglandular trichomes present, 2-8 per lobe.

Achenes narrowly ellipsoidal, 0.9-1 mm long,

terete or slightly angular, dark brown, with

25-50 or >50 twin hairs each c. 0.1 mm long,

evenly distributed; globose sessile glands at

junction between achene and pappus absent;

pappus 2-3.1 mm long. Figs. 10,11A-D.

Additional specimens examined: Australia: Western

Australia. 19 miles [31 km] NW of Turner River Station,

Jul 1949, Perry & Lazarides 2410 (CANB, DNA); China

Wall, Jun 1983, MacDonald 234 (BRI); Mt Amherst

Range, 1 mile [1.6 km] E of Mt Amherst Station, July

1959, Lazarides 6341 (BRI); 9 km SSW of Bungle

Bungle outcamp. East Kimberley, Jul 1984, Kenneally

9264 (BRI, CANB, PERTH); S slopes of Mt North, May

1988, Goble-Garratt 525 (BRI, PERTH); Mt House

Bean, Pterocaulon section Monenteles

301

Spreading, dense sub-shrub 3-4 ft high with

purple, globular flower*. Stems and leaves

woolly and white.

Thi* specimen is presented on the condition that new

identifications will be communicated to CS.T.R.O,, Canberra.

Coll. M. Lazarides 6341 Date 15 July, 1959

Loc. Mt. Amherst Range, 1 mile E. of Mt. Amherst

Station, Kimberley*, W.A.

Occasional on rocky granitic slopes with Triodia spp.

139800

270496

EX HERBARIO A USTR ALIENS!

CS.I.RXX/ Canboira

Collected and distributed by Division of Land Research and

Regional Survey, GSJ.R.O., Canberra

Name Pterocaulon sphaceLatum (Lablll.)

Be nth. et Hook.

M. Lazarides 30 June, i960

det - L LL

Ai l.yq Herb. gR/

Fig. 10. Representative specimen of Pterocaulon niveum {Lazarides 6341 [BRI]).

302

Austrobaileya 8(3): 280-334 (2011)

Fig. 11. Pterocaulon niveum. A. hermaphrodite floret x 32. B. inner involucral bract (cilia not shown) x 16. C.

upper portion of inner involucral bract x 32. D. mature achene and basal portion of pappus x 32. P. paradoxum. E.

hermaphrodite floret x 32. F. inner involucral bract (cilia not shown) x 16. G. upper portion of inner involucral bract

x 32. H. mature achene and basal portion of pappus x 32. P. redolens. I. hermaphrodite floret x 32. J. inner involucral

bract (cilia not shown) x 16. K. upper portion of inner involucral bract x 32. L. mature achene and basal portion of

pappus x 32. A-D from Brocklehurst 372 (DNA); E-G from Keighery 9018 (PERTH); H from George 13582 (PERTH);

I-K from Rodd & Hardie 4378 (BRI); L from McKee 10198 (CANB).

Bean, Pterocaulon section Monenteles

303

Station, Aug 1950, Royce 3304 (PERTH); Caroline Pool,

8 km E of Halls Creek, Jun 1976, Beauglehole ACB53312

(PERTH); 48.5 km N of ‘Springvale’ Homestead, 86

km N of Halls Creek, Jun 1976, Beauglehole ACB53527

(BRI, PERTH); Palm Springs, S of Halls Creek, Aug

2000, Handasyde & Start TH00287 (PERTH); McSpeery

Gap, Napier Range, Jun 1971, Maconochie 1298 (BRI,

DNA); between Njitparriya and Djimbitjba, 3 km SE of

Bungle Bungle Outcamp, Jul 1984, Forbes 2597 (BRI,

CANB, DNA, MEL, PERTH); Mt Broome, May 1971,

Byrnes 2253 (K). Northern Territory. Calcite flow stop.

Limestone Gorge, Gregory N.P, Jun 2000, Kerrigan &

Risler 171 (DNA); Mt Napier area. May 1974, Dunlop

4066 (DNA); Gregory N.P, Fire plot 41, May 1999,

Michell & Johnson 2449 (DNA, NT); Gregory N.P,

28 km SW of Bullita outstation, Apr 1996, Coles 28

& Barritt (DNA, MEL); Gregory N.P, Humbert River

road, Aug 1991, Brocklehurst 372 (DNA).

Distribution and habitat: Pterocaulon

niveum is widespread in the Kimberley region

of Western Australia, occurring in the Napier

Range (western Kimberley) and from Mt

Amherst (south-eastern Kimberley) eastwards

to the Gregory N.R in the Northern Territory

(Map 2). It grows in Triodia grassland or

low open eucalypt woodland, on rocky

slopes or in gullies with shallow sandy soils.

The geological substrate may be sandstone,

quartzite or limestone.

Phenology: Flowers and fruits have been

collected from April to August.

Affinities: Pterocaulon niveum, P.

tricholobum, P. discolor and P. globuliflorum

appear to be related, based on similarity in

morphology of the hermaphrodite floret,

i.e. corolla lobes long (07-1.2 mm long)

with eglandular trichomes present, and the

presence of lateral capitulescences.

Sterile specimens of Pterocaulon niveum

and P. verbascifolium may be confused, as

they have leaves of similar size, shape and

toothing, and the indumentum is extremely

dense for both. However, the much broader

stem wings of Pterocaulon verbascifolium

allow them to be easily distinguished.

Notes: Distinctive features of Pterocaulon

niveum are the indumentum of the lower leaf

surface being extremely dense and obscuring

the surface, the continuous branchlets with

lateral capitulescences present, narrow

stem wings and long corolla lobes on the

hermaphrodite floret.

7. Pterocaulon paradoxum A.R.Bean

species nova affinis P. sphacelato sed alis

caulinis latioribus, glandibus sparsius in

pagina inferiore folii distributis, dentibus

parvis in marginibus superioribus bractearum

interiorum, capitulescentiarum plus

elongatarum et lobis longioribus corollae

flosculi hermaphroditi differens. Typus:

Australia: Western Australia. 15 km ESE of

Winnama yards by Samin mining exploration

track, c. 8 km E of Palms yard, SE Kimberley,

15 May 1984, S.J.Forbes 2017 (holo: DNA;

iso: CANB, MEL).

Illustration: Kenneally et al. (1996), as

P. sphacelatum.

Shrub 60-120 cm high, aromatic properties

unknown; stems one or more from non-woody

rootstock, sparsely to densely lanate but

without globose sessile glands; stem wings

entire, 1.5-4.5 mm wide. Leaves elliptical

to oblanceolate, 30-65 x 10-20 mm, 2.6-4.6

times longer than wide, apex obtuse; margins

recurved, denticulate, with 13-23 pairs of very

small teeth. Indumentum comprising lanate

multi-cellular hairs, glandular hairs absent.

Upper surface slightly bullate; hairs erect to

spreading, moderate to very dense, basal cell

short; globose sessile glands absent. Lower

surface with lanate hairs erect to spreading,

dense to very dense, extending 0.4-0.7 mm

from the surface, basal cells not very short;

globose sessile glands yellow and shining,

moderately dense, 3-6 diameters apart.

Capitulescence ovoid to ellipsoidal, 14-28

mm long and 12-15 mm wide at anthesis, 1.2-

2.1 times longer than wide; side branchlets

never continuous, at 20-50°; peduncles 0-20

mm long. Outer involucral bracts spathulate,

2.4-27 mm long, with dense spreading lanate

hairs on outer surface, sparse hairs on the

distal end of inner surface; glands absent;

apex obtuse or truncate. Inner bracts white to

greenish (occasionally tinged pink) on dorsal

surface; longest ones linear, 3.5-4.8 mm long

and 0.2-0.4 mm wide, apex acute, upper

margins entire or with some small teeth; cilia

6-12, up to 3.1 mm long, attached along the

proximal half and consistently extending to

the distal one third, not exceeding the bract

apex. Filiform (female) florets 18-24, pink,

304

Austrobaileya 8(3): 280-334 (2011)

Lat. 17°14 1 S Long. 128°20*E Alt.: 380 m

Coll.: S-J. Forbes, no.2017, 15.v. 1984

Freq.: occasional

Dei.:

Notes: sandy flat with Eucalyptus confertiflora

Crotalaria novae-hollandiae , Heteropogon

contortus , Lysiphyllum cunninghamii

Habit; erect herb

Flowers white

-Notes: winged stems, weakly aromatic

HoioTyfiE' Queensland Herbarium (BRI)

Pferoca u/on pa,ra.i{e>xet/n d.A.8ca.n

Compositae

* A *4-fci20//

NATIONAL HERBARIUM OF VICTORIA (MEL)

MELBOURNE, AUSTRALIA

Pterocaulon sphacelatum (Labill.) Benth &

Hook.f» ex F.Muell.

Loc.: W.A, - SE Kimberley, (15 km ESE Winnama

yards by Samim mining exploration track; c. 8 kir

E Palms Yard

Fig. 12. Holotype of Pterocaulon paradoxum {Forbes 2017 [DNA]).

Bean, Pterocaulon section Monenteles

305

corolla 2.5-3.4 mm long, styles extending

0.3-07 mm beyond corolla. Hermaphrodite

floret solitary, corolla pink, 2.9-37 mm

long; corolla lobes 0.5-0.8 mm long, globose

sessile glands present, eglandular trichomes

absent. Achenes narrowly ellipsoidal, 0.9-

1.2 mm long, obscurely quadrangular, dark

brown, with 25-50 or >50 twin hairs each

c. 0.1 mm long, evenly distributed; globose

sessile glands at junction between achene and

pappus occasionally present; pappus 2.5-3.5

mm long. Figs. 11E-H, 12.

Additional selected specimens examined : Papua New

Guinea. Central Province: Rogers Airstrip, c. 8 miles

[13 km] W of Kanosia Plantation, Kairuku subdistrict,

July 1962, Darbyshire 647 (A, BRI, CANB, L). Australia:

Western Australia. Adjacent to King Edward River, 8.7

km NW of Gibb River - Kalumburu road intersection,

along old Mitchell River Station Road, Jun 1987, Koch

506 (PERTH); S side of Cockburn Range, c. 13 km W

of King River, Jul 1974, Carr 3338 & Beauglehole 47116

(CANB); King Edward River, 100 km S of Kalumburu,

Jun 1987, Keighery 9018 (PERTH); King Edward River,

Mitchell Plateau road, c. 200 km W of Wyndham, Jun

1976, Beauglehole ACB51933 (PERTH); ‘Beverley

Springs’ Homestead, May 1979, Muir et al. 757

(PERTH); near Dromaius Creek, near southern end of

Ashton Range, Drysdale River N.P, Aug 1975, George

13191 (PERTH); Mogurnda Creek, near Drysdale River,

Drysdale River N.P, Aug 1975, George 13582 (PERTH);

upper reaches of Barker River, 2 km N of ‘Mount Hart’

Homestead, Jun 1987, Edinger 423 (BRI, PERTH);

Oobraguma Road, 5 km E of Stuart River, 74 km NNE

of Derby, Jun 1976, Beauglehole ACB52918 (PERTH);

c. 100 m W of Taylor’s Lagoon, Sep 2005, Byrne 1639

(PERTH); One Arm Point, Dampier Peninsula, Jul 1989,

Carter 410 (PERTH); Mornington Station, Trapline

1, May 2004, Handasyde 2099 (PERTH). Northern

Territory. Middle Beach swamp. West Alligator Head,

Jul 1990, Brennan Bre646 (DNA); Near Maningrida,

Anamayirra Creek, Aug 1995, Cowie 5909 (CANB,

DNA, MEL); Danger Point, Coburg Peninsula, Jul 1961,

Chippendale 8244 (BRI, CANB, DNA); Grant Island,

north side. May 1987, Clark s.n. (DNA); Ramingining

area, W of Dhabla Road, Aug 1998, Cowie & Dunlop

7922 (DNA); Wunyu Beach in Aurari Bay, c. 30 km SE

of Murgenella, Jun 1988, Weber 10102 (AD, DNA).

Distribution and habitat: Pterocaulon

paradoxum is relatively widespread in the

Kimberley region of Western Australia, is

scattered along the north coast of Northern

Territory, and there is one record from Papua

New Guinea (Map 5). It inhabits eucalypt

woodland on alluvial flats (species including

Eucalyptus miniata A.Cunn. ex Schauer,

E. tetrodonta F. MuQ\\.,Corymbiaconfertiflora

(F.Muell.) K.D.Hill & L.A.S.Johnson), on

sheltered sandstone scree slopes, or on coastal

dunes. Soils include red-brown clayey sand,

red loam, and grey sand.

Phenology: Flowers and fruits are recorded

from April to September.

Affinities: Pterocaulon paradoxum appears

to be most closely allied to P. sphacelatum.

It differs by the stem wings 1.5-4.5 mm

wide (0.7-2 mm wide for P. sphacelatum).,

the corolla lobes of the hermaphrodite floret

(0.5-)0.6-0.8 mm long (0.4-0.6 mm long for

P. sphacelatum ); the globose sessile glands

on the lower leaf surface moderately dense,

3-8 diameters apart (dense, 0-3 diameters

apart for P. sphacelatum ); the upper margins

of the inner bracts with small teeth (strongly

toothed or lacerate for P. sphacelatum ); and

the capitulescences usually more elongated.

The capitulescences of this species

provide a link between the ‘spicate’ group

{Pterocaulon redolens, P. verbascifolium ,

P. serrulatum ) and the ‘globose to

ovoid’ group (P. sphacelatum, P. ciliosum,

P. sphaeranthoides, P. intermedium). The

capitulescences are more elongated that other

species in the ‘globose to ovoid’ group, but

not clearly cylindrical as in the ‘spicate’

group. One or more separate clusters of

capitula can usually been seen at the base

of the capitulescence, as in the spicate

species Pterocaulon redolens. In some areas,

particularly the Dampier Peninsula (Western

Australia), the capitulescences appear

consistently almost globose, although the

floret dimensions and other features appear to

be the same as in the type.

Notes: Distinctive features of Pterocaulon

paradoxum are the capitulescences often

markedly ellipsoidal, the medium length

corolla lobes on the hermaphrodite floret and

the long achenes.

Two forms of this species exist. The typical

form is endemic to the Kimberley, mainly

away from the coast, and is a densely hairy

plant with corolla lobes on the hermaphrodite

floret 0.6-0.8 mm long. The second form (e.g.

Cowie 5909) is found along the northern coast

of the Northern Territory, often in littoral

areas, and one specimen from Papua New

306

Guinea is rather doubtfully included here.

The indumentum on vegetative parts is much

sparser in this form than the typical one, there

are few cilia visible on protruding part of inner

bracts, the corolla lobes on the hermaphrodite

floret are shorter (0.5-0.6 mm long) and the

capitulescences are narrower. Both forms

can have ellipsoidal capitulescences, where

separate clusters of capitula are discernable.

Etymology : From the Latin paradoxus ,

meaning ‘contrary to expectations,

puzzling’. This species is puzzling because

of its sometimes ± globular and sometimes

markedly ellipsoidal capitulescences, and the

different forms within the species.

8. Pterocaulon redolens (Willd.) Fern.-Vill.,

Nov. App. ed. 3, 4(3): 116 (1880); Conyza

redolens Willd., Sp. PI. ed. 3(3), 1951 (1804);

Tessaria redolens (Willd.) Less., Linnaea 6:

151 (1831); Monenteles forsteri Endl., Ann.

Wiener Mus. Naturgesch. 1: 168 (1836), nom.

illeg:, Monenteles redolens (Willd.) DC.,

Prodr. 5: 455 (1836). Type: New Caledonia, in

1774, J.R.Forster & J.G.A.Forster (holo: B-W

No. 15639; iso: K373288, 1 st , 2 nd and 4 th pieces

from the left).

Monenteles spicatus Labill., Sert. Austro-

Caledon. 43 (1825); Pterocaulon billardierei

F.Muell., Descr. Notes Papuan PI. 8: 43 (1886),

nom. nov.; Pterocaulon spicatum (Labill.)

Domin, Biblioth. Bot. 89: 664 (1930), nom.

illeg. non DC. (1836). Type: New Caledonia.

[June 1792], J.J.H. Labillardiere (lecto: FI

[FI-W92604], here designated; isolecto: FI

[FI-W92605], right hand specimen only, FI

[FI-W92611], P537803).

Sphaeranthus elongatus Blanco, FI. Filip. 636

(1837), fide Merrill (1918: 379).

Pterocaulon cylindrostachyum C.B.Clarke,

Compos. Ind. 98 (1878), nom. illeg. {Monenteles

spicatus is listed as a synonym).

Gnaphalium cylindrostachyum Wallich, Num.

List [Wallich] 2931 (1831), nomen nudum.

Illustrations : Cabrera & Ragonese (1978:

240); Porteners (1992: 204); Stanley & Ross

(1986: 532).

Weakly aromatic shrub 40-100 cm high; stems

Austrobaileya 8(3): 280-334 (2011)

one to several from non-woody rootstock,

sparsely to densely lanate and sometimes

with a few globose sessile glands; stem wings

entire, 1.4-3.3 mm wide. Leaves oblanceolate

to obovate, 16-45 x 6.5-16 mm, 2.1-3.4

times longer than wide, apex acute or obtuse;

margins recurved, denticulate, with 7-16

pairs of blunt teeth. Indumentum comprising

lanate multi-cellular hairs, glandular hairs

absent. Upper surface somewhat bullate;

hairs erect to spreading, sparse to moderately

dense, basal cells short; globose sessile

glands absent. Lower surface with lanate

hairs erect to spreading, moderately dense to

dense, extending 0.5-1 mm from the surface,

basal cells not very short; globose sessile

glands yellow and shining, dense, touching

or up to 3 diameters apart. Capitulescence

spiciform, 33-75 mm long and 8-14 mm wide

at anthesis, 2.8-5.8 times longer than wide;

side branchlets never continuous, at 20-45°;

peduncles 5-25 mm long. Outer involucral

bracts 1.6-2 mm long, oblong to oblong-

spathulate, with dense spreading lanate hairs

on outer surface, glabrous on inner surface;

glands absent; apex obtuse. Inner bracts white

to straw coloured (sometimes tinged pink) on

dorsal surface; longest ones linear, 3-3.6 x 0.1-

0.25 mm wide, apex attenuate, upper margins

entire or with an occasional small tooth; cilia

5-9, up to 2.5 mm long, attached along the

proximal half and consistently extending to

the distal one third, not exceeding the bract

apex. Filiform (female) florets 27-45, white

or straw-coloured, corolla 2.3-2.8 mm long,

styles extending 0.1-0.7 mm beyond corolla.

Hermaphrodite floret solitary, corolla pink,

2.5-3 mm long; corolla lobes 0.25-0.5 mm

long, globose sessile glands always present,

eglandular trichomes absent. Achenes

narrowly ellipsoidal, 0.5-0.7 mm long, terete

or slightly angular, dark brown, with 0-25 or

25-50 twin hairs each c. 0.05 mm long, evenly

distributed; globose sessile glands at junction

between achene and pappus usually present;

pappus 2.2-3 mm long. Figs. 11I-L, 13.

Additional selected specimens examined : China.

Ch’ang-kiang District, Hainan Island, Feb 1934, Lau

3261 (GH); Taichow, Hainan Island, Jan 1938, How &

Chun 70001 (GH). India. Ghariajhor, 11 miles [18 km]

from Sundargarh, Orissa, Mar 1950, Mooney 3748 (K).

Myanmar. Pegu, s.dat., Kurz 2263 (K); Prome, Ava,

Bean, Pterocaulon section Monenteles

307

QUEENSLAND HERBARIUM (BRI)

Flora of Queensland Maranoa

Pterocaulon reddens (Willd.) Fern.-Vill.

QUEENSLAND HERBARIUM (BRI)

Brisbane Australia

747G83

Fig. 13. Representative specimen of Pterocaulon redolens (Eddie CPE1064 [BRI]).

308

s.datHance 17401 (K). Thailand. Ban Huay Nam

Kow, Chiang Mai Province, Feb 1993, Radanachalesa

1 (A). Indonesia, near Palu, Celebes, Apr 1975, Meijer

9173 (A, L). Philippines. Bataan Province: Luzon, Jul

1913, Merrill 341 (AMES, L). Papua New Guinea.

Morobe Province: Erap, Jan 1958, Henty NGF9481

(BRI, CANB, K). Central Province: Tupuseleia, Port

Moresby subdistrict, Aug 1967, Kairo & Streimann

NGF30803 (BRI, CANB, L). New Caledonia. lie Pott,

Mouane, Aug 1968, McKee 19366 (NOU); Plateau

central, lie des Pins, Jul 1970, McKee 22304 (NOU);

Noumea, 1868-1870, Balansa (A). Vanuatu. Tableland,

Eromanga, Jul 1896, Morrison s.n. (K). Australia:

Queensland. Cook District: Dimbulah, Apr 1935,

Flecker 398 (CNS); Ravenshoe - Hughenden Road, c.

150 km N of Hughenden, Apr 1975, Halliday 418 (BRI);

Bulleringa N.P., 80 km NW of Mt Surprise, Apr 1998,

Forster PIF22541 & Booth (BRI, DNA, MEL). North

Kennedy District: Barrabas Scrub, May 1972, Hyland

6085 (CNS); 13.5 km W of the Dry Swamp, May 1991,

Neldner 3268 & Thompson (BRI, K); 6 miles [10 km] N of

Craigie Station, Jul 1953, Lazarides 3718 (BRI, CANB);

Townsville, Aug 1916, White s.n. (BRI [AQ270344]).

Leichhardt District: Springsure, Apr 1961, Jones 1870

(BRI, CANB); Twenty Mile Yards, ‘Rookwood’, Apr

1991, Forster PIF7881 & McDonald (BRI, MEL). Port

Curtis District: Rundle Range, S.F. 60, 37 km NW of

Gladstone, Aug 1989, Gibson TOI698 (BRI). Burnett

District: 9 km SE of Mt Perry (town), Apr 1985, Rodd

& Hardie 4378 (BRI, CANB, NSW); Windera, Apr

1963, McKee 10198 (BRI, CANB). Maranoa District:

Boatman Station, Mar 1947, Everist 2813 (BRI); ‘Six

Mile’, Bulloak Paddock, NNW of Roma, Mar 2007,

Eddie CPE1064 (BRI). Warrego District: Yanna

Siding, via Charleville, May 1949, Heinemann s.n. (BRI

[AQ270369]). Moreton District: Rifle Range Reserve,

Goodna, E of Ipswich, Apr 1990, Bean 1512 (BRI).

Distribution and habitat : Pterocaulon

redolens is widespread, with occurrences in

China (Hainan), India, Myanmar, Thailand,

Philippines, Indonesia, Papua New Guinea,

Australia, New Caledonia and Vanuatu.

Cabrera & Ragonese (1978) reported seeing

specimens also from Laos, Cambodia and

Vietnam. In Australia, it is widely distributed

in eastern Queensland, south from Lakefield

National Park (Map 6), and was reported by

Porteners (1992) to occur in the far north¬

east of New South Wales. It inhabits grassy

areas at low altitudes in hilly or flat terrain,

with sandy to clay-loam soil. In Australia, it

is found in various eucalypt woodlands or

on the edge of vine thickets in shallow or

deep soils, sometimes behind coastal dunes.

The geological substrate may be granite,

sandstone, basalt or serpentinite.

Austrobaileya 8(3): 280-334 (2011)

Phenology : Flowers and fruits are recorded

for every month of the year, with most records

between April and September.

Affinities: Pterocaulon redolens is not

obviously allied to any other species. However,

sterile specimens of Pterocaulon redolens

are indistinguishable from the sympatric

P. ciliosum.

Typification: The isotype of Conyza redolens

at K [K000373288] has a blue hand-written

label stating “Gnaphalium redolens (Forster)/

Habitat in New Caladonia”, and next to that, a

printed label saying “The Forster Herbarium.

Presented by the Corporation of Liverpool,

August 1885”. There are five separate branches

mounted on the sheet; three are Pterocaulon

redolens and two are P. sphacelatum.

Notes: This species is distinctive in the

spicate, interrupted capitulescence; ring of

glands often present at apex of achene; upper

margins of inner bracts entire and many

female florets (27-45) per capitulum.

The author has seen a single specimen

of this species from India, as cited above.

Pterocaulon redolens must be a relatively

recent arrival in India, as Hooker (1882)

recorded it only from “Burma and the Eastern

Peninsula”. It was not recorded by Haines

(1961) as present in Bihar or Orissa. Rath &

Priyadarshini (2005) reported Pterocaulon

redolens from an abandoned manganese

quarry, a highly modified habitat. I conclude

that this species is very likely not native to

India, but naturalised.

9. Pterocaulon serrulatum (Montrouz.)

Guillaumin, Bull. Soc. Bot. France 84: 56

(1937); Monenteles serrulatus Montrouz.,

Mem. Acad. Roy. Sci. Lyon , Sect. Sci. 10: 225

(1860). Type: New Caledonia. Art Island,

s.dat., J.X.Montrouzier 125 (holo: LY,

destroyed).

Monenteles glandulosus Benth., FI. Austral.

3: 523 (1867); Pterocaulon glandulosum

(Benth.) F.Muell, Syst. Census Austral. PI.

79 (1883); P. glandulosum var. glandulosum ,

Ewart & Davies, FI. N. Territory 277 (1917).

Type: Australia: Queensland. Cook District:

Gilbert River, October 1856, F. Mueller s.n.

(lecto: K373279, fide Cabrera & Ragonese

[1978: 254]).

Bean, Pterocaulon section Monenteles

309

Strongly aromatic perennial shrub 35-150 cm

high; stems oneto several from woody rootstock,

with sparse to dense lanate eglandular hairs,

globose sessile glands and sometimes stalked

glandular hairs; stem wings serrate, 2.5-8

mm wide. Leaves elliptical, ovate or broadly-

lanceolate, 20-82 x 8-28 mm, 2.1-4.3 times

longer than wide, apex acute; margins flat or

recurved , serrulate, with 13-45 pairs of teeth.

Indumentum comprising lanate multi-cellular

hairs, glandular hairs present or absent. Upper

surface smooth or slightly bullate; hairs erect

to spreading, sparse to dense, basal cells

short; globose sessile glands present. Lower

surface with lanate hairs erect to spreading,

sparse to very dense, extending 1-1.5 mm

from the surface, basal cells not very short;

globose sessile glands yellow and shining or

pale yellow and opaque, dense, touching or up

to 3 diameters apart. Capitulescence ellipsoid

or cylindrical, sometimes ‘interrupted’, rarely

almost globose, 14-52 mm long and 13-17

mm wide at anthesis, 1.1-3.5 times longer

than wide; side branchlets never continuous,

at 20-45°; peduncles 0-15 mm long. Outer

involucral bracts 2.3-37 mm long, oblong,

with sparse to moderately dense spreading

lanate hairs on margins, glandular-hairy on

both surfaces; apex obtuse. Inner bracts white

to greenish-white on dorsal surface; longest

ones linear, 3.9-6.3 x 0.3-0.5 mm, apex acute,

upper margins with small or large teeth; cilia

6-14, up to 2.4 mm long, attached along the

proximal half and consistently extending to

the distal one third, not exceeding the bract

apex. Filiform (female) florets 29-52, white

to straw coloured, corolla 2.8-4.2 mm long,

styles extending 0.3-0.5 mm beyond corolla.

Hermaphrodite floret solitary, corolla pink,

3.6-47 mm long; corolla lobes 0.4-0.6

mm long, globose sessile glands present,

eglandular trichomes absent. Achenes

narrowly ellipsoidal, 0.7-1 mm long, terete

or almost so, dark brown, with 25-50 twin

hairs each c. 0.1 mm long, evenly distributed;

globose sessile glands at junction between

achene and pappus absent; pappus 2.8-4.5

mm long.

Affinities : Pterocaulon serrulatum is

a distinctive species, but is sometimes

confusable with P. sphacelatum and

P. ciliosum. These are perhaps its closest

relatives, and postulated hybrid specimens

are reported in this paper.

Typification: Vegter (1976) reported that

Montrouzier’s types are at LY, and Guillaumin

& Beauvisage (1914) reported seeing the type

of Monenteles serrulatus there. However, it

appears that Montrouzier’s collection at LY

was destroyed or lost in the 1950’s (P. Morat,

pers. comm. June 2010), and the curator at LY

has confirmed that the type of Monenteles

serrulatus is not present there now. Duplicates

of some of Montrouzier’s collections are held

at MPU, but no type material of Monenteles

serrulatus can be found there (P. Schafer,

pers. comm. June 2010), nor is there an isotype

at P (P. Morat, pers. comm. June 2010). This

is in accord with the information provided

by Guillaumin & Beauvisage (1914), i.e. that

there was only a single type specimen for this

name, the specimen at LY.

There seems little doubt about the

application of the name Pterocaulon

serrulatum. The protologue is fairly detailed

and the description matches the plant currently

called Pterocaulon serrulatum var. serrulatum

in Australia. Guillaumin & Beauvisage (1914)

reported that the type was very similar to

Pterocaulon glandulosum F.Muell., but that it

differed by being totally glabrous.

Notes : Pterocaulon serrulatum is distinctive

in the cylindrical to ovoid capitulescence;

broad serrated stem wings; many female

florets per capitulum; outer involucral bracts

glandular hairy on both surfaces and the

sessile glands on upper leaf surface.

Varietal rank is appropriate for the taxa

included within Pterocaulon serrulatum. The

two varieties differ only in their indumentum,

but these characters do not vary continuously

in Australian material, and a useful

geographical separation is achieved through

their use. All eastern Queensland material can

be readily assigned to var. serrulatum , and all

Northern Territory material is referrable to a

more broadly circumscribed var. velutinum,

as outlined in the key. Some specimens from

the Kimberley region are assigned here to var.

serrulatum. Some specimens from the Jericho

and Hughenden districts of Queensland are

difficultto assign to either variety; for example,

310

Austrobaileya 8(3): 280-334 (2011)

QUEENSLAND HERBARIUM (BRI)

Flora of Queensland Leichhardt

Pterocaulon serrulatum (Montrouz.) Guillaumin

var. serrulatum

QUEENSLAND HERBARIUM (BRI)

Brisbane Australia

aq 649945

(55,512865,7294521 ) (8349-128945) Alt. 400m

118km from Spnngsure, on road to Tambo.

Low woodland or shrubland with Acacia harpophylla, Erythroxylum.

Eremophila mitchellii. Carissa Pink loam near crest of low ridge.

Highly aromatic forto, stems and leaves green, Inflorescence mauve.

Rare at site

Fig. 14. Representative specimen of Pterocaulon serrulatum var. serrulatum ( Bean 22120 [BRI]).

Bean, Pterocaulon section Monenteles

311

Smyrell GS69 has broad based glandular

hairs (a feature of var. serrulatum) and also

has lanate hairs on the upper leaf surface (a

feature of var. velutinum). Similarly, in the

Pilbara region of Western Australia, there are

specimens referred to here as var. velutinum ,

but which closely approach var. serrulatum

e.g. Byrne 2807.

Key to varieties

Lanate hairs absent from upper leaf surface; broad based glandular hairs

common on leaves, stems and stem wings.

.9a. P. serrulatum var. serrulatum

Lanate hairs present on upper leaf surface; broad based glandular trichomes

absent, but globose sessile hairs common on leaves, stems and stem wings

.9b. P. serrulatum var. velutinum

9a. Pterocaulon serrulatum var.

serrulatum

Winged stems and leaf surfaces with many

broad-based glandular hairs 0.1-0.2 mm long;

erect or spreading eglandular multicellular

lanate absent from upper leaf surface, sparse

or absent on lower leaf surface, stem wings

and stems. Figs. IB, 14,16A-D.

Illustration : Stanley & Ross (1986: 532).

Additional selected specimens examined : Australia:

Western Australia. Bell Gorge parking area, Aug

2005, Byrne 1619 (PERTH); western base of Mt

Hamilton, Jul 1905, Fitzgerald 1209 (PERTH); Little

Island, off Long Island, Buccaneer Archipelago, Jun

1982, Hopkins BA0115 (PERTH); Around ‘Beverley

Springs’ Homestead buildings, Jun 1992, Mitchell 2556

(PERTH). Queensland. Cook District: Fossilbrook

Creek, ‘Burlington’, N of Mount Surprise, Aug 1997,

Bean 12229 (BRI); along Kuranda Highway, c. 10

miles [16 km] E of Mareeba, Jun 1962, Hoogland 8486

(BRI, CANB). North Kennedy District: 22.5 km (by

road) SW of Herberton, on the Silver Valley road.

May 1983, Conn & Clarkson 1178 (AD, BRI, CANB,

CNS, DNA, HO, NSW, PERTH); Road from Camel

Creek to Greenvale, Aug 1998, Fox 25 & Bean (BRI);

base of Frederick Peak, 25 km SW of Townsville, May

1991, Bean 3215 (BRI); Lookout, on eastern slopes of

Mt Kelly, c. 14 km south-west of Ayr, May 2009, Bean

28848 (BRI); Bowen, reserve near reservoir, Nov

1978, Ross s.n. (BRI [AQ297444]). Mitchell District:

‘Grant’, c. 70 km ENE of Barcaldine, Jul 1994, Smyrell

GS89 (BRI). South Kennedy District: Redcliff Island,

Seaforth, Aug 1992, Batianoff9208129 (BRI); 19 km E

of‘Doongmabulla’, N of Jericho, Apr 2002, Bean 18984

(BRI, MEL). Leichhardt District: 60 miles [97 km] N

of Comet township, Jul 1962, Story & Yapp 185 (BRI,

CANB); 118 km from Springsure, on road to Tambo,

May 2004, Bean 22120 (BRI); Ka Ka Mundi section,

Carnarvon N.P, on fire trail from Salvator Rosa, Aug

2008, Thomas MBT3780 & Leggett (BRI). Port Curtis

District: ‘Corisande Hills’, Stanage Bay road, N of

Rockhampton, Jul 1997, Bean 12063 (BRI, MEL); 32

km N of Miriam Vale, Jul 1996, Thompson M1R136A

& Turpin (BRI). Burnett District: Gayndah, May

1917, White s.n. (BRI [AQ2703920]); ‘Dykehead’, 70

km W of Mundubbera, Jun 1998, Robinson JR-1 (BRI).

Wide Bay District: Degilbo Creek, near Biggenden,

Aug 1980, Johnson s.n. (BRI [AQ343756]); Rossmore

Road, 10 km SE of Kilkivan, May 2006, Webb s.n. (BRI

[AQ619054]).

Distribution and habitat: Found on Art

Island, New Caledonia, and throughout the

eastern half of Queensland in Australia,

from Cooktown to Kilkivan. It also occurs

in the western Kimberley region of Western

Australia (Map 7). It grows on a wide range

of sites (include coastal headlands) where

drainage is good, usually in eucalypt woodland

or grassland. Soils range from sands to clay-

loams.

Phenology: Flowers and fruits are borne

between April and November.

9b. Pterocaulon serrulatum var. velutinum

(Ewart & O.B.Davies) Guillaumin, Bull.

Soc. Bot. France 84: 57 (1937); Monenteles

glandulosum var. velutinum Ewart &

O. B.Davies, FI. N. Territory 211 (1917). Type:

Australia: Northern Territory. Haast’s Bluff,

17 May 1911, G.F.Hill 184 (holo: MEL).

P. serratum O. Schwarz, Repert. Spec. Nov.

Regni Veg. 24: 108 (1927). Type: Australia:

Northern Territory. Port Darwin, 10 miles

NE, s.dat., F.A.K.Bleeser 712 (holo: B,

destroyed).

Illustrations: Cabrera & Ragonese (1978:

253); Wheaton (1994: 112); Barrs (1999: 118)

as P. serrulatum var. serrulatum.

312

Austrobaileya 8(3): 280-334 (2011)

QUEENSLAND HERBARIUM <BRI)

Flora of Queensland Burke

Pterocaulon serrulatum (Montrouz.) GuiNaumin

Coil. R. Booth LH28-5 7 JUL 2009

Kelman.0.:

(10d 57m 50s 139d 3m 08s) [WGS84J Depth m

54.294910.7902039 (6760-94920) AIL m

Gregory Downs Stn, E of Rrversleigh.

Eucalyptus leucophfoia woodland on Jurrasic sandstone ridgetop.

Hairy upright herb.

Queensland Herbarium (BRI)

Pterocaulon serrulatum var uelutin,

CE-tiaanwajSSSr

QUEENSLAND HERBARIUM (BRI)

Brisbane Australia

850132

Fig. 15. Representative specimen of Pterocaulon serrulatum var. velutinum {Booth LH28-5 & Kelman [BRI]).

Bean, Pterocaulon section Monenteles

313

Fig. 16. Pterocaulon serrulatum var. serrulatum. A. hermaphrodite floret x 32. B. inner involucral bract (cilia not

shown) x 16. C. upper portion of inner involucral bract x 32. D. mature achene and basal portion of pappus x 32.

Pterocaulon sphacelatum. E. hermaphrodite floret x 32. F. inner involucral bract (cilia not shown) x 16. G. upper portion

of inner involucral bract x 32. H. mature achene and basal portion of pappus x 32. Pterocaulon sphaeranthoides. I.

hermaphrodite floret x 32. J. inner involucral bract (cilia not shown) x 16. K. upper portion of inner involucral bract

x 32. L. mature achene and basal portion of pappus x 32. A-C from Bean 28848 (BRI); D from Ross s.n. (BRI

[AQ297444]); E, H from Piercey 51 (DNA); F-G from Halford Q2626 (BRI); I-K from van Leeuwen 0099 (PERTH);

L from van Leeuwen 0092 (PERTH).

314

Winged stems and leaf surfaces without

broad-based glandular hairs; globose sessile

glands c. 0.05 mm long, abundant; erect to

tangled eglandular multicellular lanate hairs

sparse to very dense on leaves, wings and

stems. Figs. 1A, 15.

Additional selected specimens examined: New

Caledonia. Voh, presqu’ile de Gatope, Aug 1970,

McKee 22393 (NOU); Mt Yallein, Jul 1968, Jaffre

57 (NOU); Presqu’ile d’Arama, Aug 1965, Schmid

619 (NOU); West face of Massif de Koniambo, Oct

1963, Green 1288 (A, K, NOU). Australia: Western

Australia. Gibb River Road, 117 km E of Derby, Aug

2001, Courtney 40 (PERTH); Honeymoon Beach, 25

km N of Kalumburu, Jun 1990, Edinger 713 (PERTH);

37.6 km NW of ‘Bonney Downs’ Homestead on road to

Hillside, Oct 1995, Mitchell PRP971 (DNA, PERTH);

McLarty Hills, Great Sandy Desert, Aug 1977, George

14625 (CANB, PERTH); 12 km SSW of Two Sisters,

c. 150 km SE of Shay Gap, Jul 1984, Newbey 10407

(PERTH); E of junction of Kununurra - Wyndham

-Halls Creek roads, Jul 1974, Carr 3203 & Beauglehole

46981 (BRI, CANB, PERTH); Roy Hill, Aug 2007, Byrne

2807 (PERTH); Meentheena Conservation Reserve, 6.5

km SSE of ‘Meentheena’ Homestead, May 2001, van

Leeuwen 4797 (PERTH). Northern Territory. 4.5 km

from Stuart Highway on Edith Falls Road, Jul 2003,

Elick 222 (CANB, CNS); 23 miles [37 km] NW of Rabbit

Flat, Tanami, Apr 1971, Dunlop 2131 (BRI, DNA); base

of Meyers Hill, Alice Springs, Aug 1957, Chippendale

3587 (BRI, CANB, DNA); Mt Bundy, Oct 1968, Harpley

NB982 (BRI, DNA); 269 km N of ‘Barkly’ Homestead

towards Borroloola, Jun 1999, Bean 15047 (BRI, DNA);

Channel Island, Darwin Harbour, Jul 1995, Cowie 5880

(DNA, MEL). Queensland. Burke District: 3 miles [5

km] from Mount Isa towards Lake Moondarra, Jul 1974,

Ollerenshaw 1142 & Kratzing (BRI, CANB, L); Gregory

Downs Station, E of Riversleigh, Jul 2009, Booth LH28-

5 & Kelman (BRI). Cook District: Ortona Station,

102 km by road S of Forsayth, Aug 2010, Bean 29878

6 McDonald (BRI, MEL, NSW). Mitchell District:

Bessie’s Castle, Silsoe Road, c. 100 km W of Longreach,

May 2004, Bean 22246 (BRI); 19.7 km E of Jericho, Jul

1975, Chapman 1281 (AD, BRI, CANB, K, L). Gregory

North District: Winton - Jundah Road, 23.9 km N of

‘Elvo’ Homestead, May 2004, Bean 22561 (BRI).

Distribution and habitat : A widespread

taxon, occurring in New Caledonia, Australia

in central and north-western Queensland,

throughout the Northern Territory, and the

northern half of Western Australia as far

west as the Hamersley Range (Map 7). It is

also recorded for a few locations in northern

South Australia (Anon. 2011). It inhabits a

wide range of habitats; the vegetation can be

Acacia woodland, mallee shrubland, eucalypt

open woodland or Triodia grassland. Soils can

vary greatly, including beach sand, red sandy

Austrobaileya 8(3): 280-334 (2011)

loam, or basaltic clay-loams, and it may grow

on sandstone, limestone or quartzite hills, or

on flats.

Phenology : Flowers and fruits have been

recorded from April to October.

Notes : Cabrera & Ragonese (1978) stated

that Pterocaulon glandulosum var. velutinum

Ewart & O.B.Davies is a nomen nudum , but

that is incorrect. Ewart and Davies provided

an adequate description and the name is

validly published.

The type of Pterocaulon serrulatum var.

velutinum has a very dense cover of lanate

eglandular hairs, and the two varieties have

in the past been distinguished on the density

of these hairs (based on herbarium specimen

determinations). In this paper, I have placed

more emphasis on the presence or absence

of stalked glandular hairs in dividing the

varieties. This provides a useful geographic

separation between the varieties, but it means

that a wide range of lanate hair densities is

included within var. velutinum. The density of

lanate hairs can vary at a single location, e.g.

Latz 7950 comprises pieces collected from

two plants growing at the same place; on one

the lanate hairs are sparse, and on the other

they are very dense.

No type material of Pterocaulon serratum

O. Schwarz has been seen, but from the

description given in the protologue, there

seems little doubt that it is this variety.

10. Pterocaulon sphacelatum (Labill.)

F.Muell., Syst. Census Austral. PI. 79 (1882);

Monenteles sphacelatus Labill., Serf.

Austro-Caledon. 43 (1825). Type: [New

Caledonia] Nova Caledonia. [June 1792],

J.J.H.Labillardiere s.n. (lecto: FI [FI-W

92608], here designated, image!; isolecto:

FI [FI-W 92605, left hand specimen only],

P537804, image!).

Illustrations : Cabrera & Ragonese (1978:

246); Porteners (1992: 204); Milson (2000:

21); Moore (2005: 126).

Strongly aromatic shrub 40-100 cm high,

often almost as wide; stems single or

numerous, densely lanate and sometimes

with a few globose sessile glands; stem wings

entire or sparsely toothed, 0.7-2 mm wide.

Bean, Pterocaulon section Monenteles

315

Leaves oblanceolate or spathulate, 20-66 x

4-15 mm, 3.3-5.8 times longer than wide,

apex obtuse or acute; margins recurved,

denticulate, with 6-14 pairs of blunt teeth.

Indumentum comprising lanate multi-cellular

hairs, glandular hairs absent. Upper surface

moderately to strikingly bullate; hairs erect

to spreading, sparse to moderately dense,

basal cells short; globose sessile glands

absent. Lower surface with lanate hairs erect

to spreading, moderate to dense, extending

0.4-0.7 mm from the surface, basal cells not

very short; globose sessile glands yellow and

shining, dense, touching or up to 3 diameters

apart. Capitulescence globose or ellipsoidal,

10-15 mm long, 10-14 mm wide at anthesis,

1-1.4 times longer than wide; side branchlets

never continuous, at 30-60°; peduncles 0-22

mm long. Outer involucral bracts 2.2-37 mm

long, spathulate, with dense spreading lanate

hairs throughout on the outer surface, sparse

hairs at distal end of the inner surface; glands

absent; apex acute. Inner bracts violet, pink or

white on dorsal surface; longest ones linear,

3.5-4.6 x 0.3-0.5 mm; apex acute, upper

margins conspicuously toothed, or upper

margins lacerate with no single distinct apex;

cilia 6-18, up to 1.7 mm long, mostly attached

in proximal half and few extending to the distal

one third. Filiform (female) florets 17-29, pink,

corolla 2.4-3.2 mm long, styles extending

0.4-0.6 mm beyond corolla. Hermaphrodite

floret solitary, corolla pink, 2.5-37 mm long;

corolla lobes 0.4-0.6 mm long, globose sessile

glands present, eglandular hairs absent.

Achenes narrowly ellipsoid, 0.7-1 mm long,

terete or slightly quadrangular, dark brown,

with 25-50 twin hairs each c. 0.05 mm long,

evenly distributed; globose sessile glands at

junction between achene and pappus rarely

present; pappus 2.3-3.3 mm long. Fruit salad

plant, Apple bush. Figs. 2A, 16E-H, 17.

Additional selected specimens examined : Timor-

Leste. Plateau of Baucau, Dec 1953, Van Steenis 18030a

(L). New Caledonia. Baie Maa, Oct 1982, Suprin

2059 (NOU); s.loc. , Oct 1965, MacKee 13525 (K, L).

Australia: Western Australia. Eagle Bore Study site.

Plot 9, Gibson Desert Nature Reserve, Aug 2002, Bragg

118 (MEL, PERTH); Rocky Pool, Gascoyne River, c.

850 km N of Perth, Oct 1975, Kenneally 4641 (PERTH);

Northern end of Windjana Gorge, Jul 1974, Carr 4281

& Beauglehole 48059 (PERTH); Elder Creek, 2 km

W of Warburton, Aug 1962, George 3828 (PERTH);

Anna Plains Station, Jul 1941, Burbidge 1462 (PERTH);

Lacrosse Island, at head of Cambridge Gulf, Jun 1992,

Kenneally 11328 (DNA, CANB, PERTH). Northern

Territory. Auvergne Station, 35 km NW of Timber

Creek, Jul 1977, Must 1620 (CANB, DNA, SI); c. 4.5 km

by road N of Gilbert Creek crossing by Stuart Highway,

between Wauchope and Tennant Creek, Aug 1978,

Donner 6232 (AD, CANB, NT); Junction Reserve, Jul

1982, Piercey 51 (DNA); Oenpelli, Sep 1948, Specht

1083 (BRI, L); Burt Creek, 4 miles [6 km] S of Yambah

Station, Mar 1953, Perry 3372 (BRI, CANB); 3.2 miles

[5.1 km] N of Alice Springs, Dec 1962, Nelson 559

(CANB, DNA); Andado Station, Mac Clarke Reserve,

Sep 1992, Latz 12706 (DNA, MEL); Horse paddock,

‘Tobermorey’, Oct 1954, Chippendale 418 (DNA); Barkly

Tablelands, N of Telegraph bore, Mittibah Station, Jul

2001, Mangion 1534 & Risler (DNA). Queensland.

Burke District: East bank of Leichhardt River, 72 km

SE of Burketown, Burke Development Road, Jul 1991,

Jobson 1490 (BRI, HO, MEL). Cook District: Topsy

Creek, Kowanyama Aboriginal Reserve, Aug 1980,

Clarkson 3392 (BRI, CNS). Gregory North District:

32 km N of Bedourie along road to Boulia, Jul 1995,

Halford Q2626 (BRI). Mitchell District: Silsoe Road,

4.1 km W of ‘Morelia’ turnoff, W of Longreach, May

2004, Bean 22250 (BRI). Warrego District: 99.3 km

from Blackall towards Adavale, near Listowel Downs

Station, Oct 1983, Canning 6192 & Rimes (CANB, BRI,

DNA, MEL, NSW). Gregory South District: 20.2 km

WSW of Eromanga, on Cooper Developmental Road,

Aug 2010, Bean 30023 (A, BRI, BRIT, MO). New South

Wales. 113 km W of Wanaaring on Milparinka Road,

Nov 1971, Blaxell 613 (BRI, NSW); Lake Pinaroo near

Fort Grey camping area, NW of Tibooburra, Sep 1989,

Coveny 13477 et al. (AD, BRI, CANB, NSW, MEL,

PERTH); Homestead Gorge, Mootwingee N.P, 112

km NE of Broken Hill, Oct 1988, Crawford 1028 (AD,

CANB, NSW). South Australia, around Nent Oura

Research unit. Mount Freeling Station, Oct 1987, Foster

227 (AD, DNA); Stuart Creek, 18 km WSW of ‘Stuart

Creek’ Homestead, Dec 1984, Badman 1597 (AD, DNA);

Bunyeroo Gorge, Flinders Ranges N.P, Nov 1982,

Symon 13040 (AD, CANB, MO).

Distribution and habitat : Pterocaulon

sphacelatum is the most widespread

Australian Pterocaulon species, occurring

in all mainland states except Victoria, and

extending from the west coast of Western

Australia to within 150 km of the Queensland

east coast. It occurs throughout much of

Northern Territory and South Australia, and

in the north-west of New South Wales. It also

occurs in Timor and New Caledonia (Map

8). In grows on a range of sandy to clay-

loam soils, on stony hillsides (higher rainfall

areas) or creek-beds (arid areas), in grassland

(often dominated by Triodia spp.), eucalypt

woodland or low open woodland with Acacia

aneura or other Acacia spp.

316

Austrobaileya 8(3): 280-334 (2011)

oriA

W* m.

northern territory herbarium

DARWIN (DIVA) AUSTRALIA

Pterocsmlon sphacelatum (Labill.) Benlh. & Hook, si F.Muell

Barkly Tablelands. N.of Telegraph bore Millibah Slalion

STATE/DIST.: NT /BT I8"4479"S 137=I710"E

Perennial herb lo 50cm tall on cresl of gravelly clay soil on plain.

Fig. 17. Representative specimen of Pterocaulon sphacelatum (Mangion & Risler 1534 [DNA]).

Bean, Pterocaulon section Monenteles

317

Phenology : Flowers and fruits occur from

June to October in northern parts of its range,

and from August to December in southern

parts.

Affinities : Pterocaulon sphacelatum is

closely related to P. ciliosum, but differs by

the narrower leaves, and the bright pink inner

bracts with few visible cilia on protruding

parts, and strongly toothed to lacerate upper

margins.

Typification: There are two sheets at FI

bearing material that matches the protologue

of Monenteles sphacelatus. On one sheet (FI-

W 92605), specimens of both Pterocaulon

sphacelatum and P. redolens are present, but

neither specimen is named; on the other sheet

(FI-W 92608), there is a single specimen of

P. sphacelatum accompanied by a detailed

description written by Labillardiereandheaded

Monenteles sphacelatus ; this description

agrees perfectly with the protologue. This

latter sheet is selected as the lectotype.

Notes : This species is distinctive in the upper

margins of the inner bracts often lacerate;

capitulescences often ellipsoidal, up to 1.4

times longer than wide; the inner bracts always

partly pink to violet, often predominantly so;

leaves oblanceolate in shape, 3.5-5.8 times

longer than wide; leaf upper surface bullate;

corolla lobes of hermaphrodite floret short

and none or few cilia visible on protruding

part of inner bracts.

11. Pterocaulon sphaeranthoides (DC.)

F.Muell., Syst. Census Austral. PI. 79 (1882);

Monenteles sphaeranthoides DC., Prodr. 5:

456 (1836). Type: Australia: Western Australia.

Enderby Island, Dampier Archipelago, 25

February 1818, A.Cunningham s.n. (holo: G-

DC; iso: PERTH; BM, K n.v).

Illustration : Cabrera & Ragonese (1978:

250).

Strongly aromatic shrub 30-120 cm high;

stems numerous, densely glandular-hairy

and sometimes with sparse lanate eglandular

hairs; stem wings prominently toothed,

0.5-3(-4) mm wide. Leaves narrowly

oblanceolate to linear, 12-48 x 2-10 mm,

3.5-13 times longer than wide, apex acute;

margins recurved to revolute, serrate with

10- 16 pairs of teeth. Indumentum comprising

broad-based glandular hairs, globose sessile

glands and sometimes lanate eglandular hairs.

Upper surface strikingly bullate; glandular

hairs erect, sparse to moderately dense,

0.1-0.2(-0.4) mm long, broad-based; lanate

hairs absent or sparse; globose sessile glands

present. Lower surface glabrous or with

sparse, erect to spreading lanate hairs, basal

cells not very short; globose sessile glands

yellow and shining, dense, touching or up to

3 diameters apart. Capitulescence ± globose,

11- 16 mm long, 11—14 mm wide at anthesis,

1-1.2 times longer than wide; side branchlets

never continuous, at 30-60°; peduncles 1-20

mm long. Outer involucral bracts 2.2-27

mm long, linear-spathulate to oblong, with

dense spreading lanate hairs throughout on

the outer surface, glabrous on inner surface;

glands present; apex acute or obtuse. Inner

bracts violet, pink or white on dorsal surface;

longest ones linear, 3.5-4 x 0.2-0.5 mm,

upper margins with large teeth, or lacerate

with no distinct single apex; cilia 5-12, up

to 1.5 mm long, mostly attached in proximal

half and few extending to the distal one third.

Filiform (female) florets 14-20, pink, corolla

2.3-2.8 mm long, styles extending 0.2-0.9 mm

beyond corolla. Hermaphrodite floret solitary

or very rarely two, corolla pink, 27-3.4 mm

long; corolla lobes 0.4-0.5 mm long, globose

sessile glands present, eglandular trichomes

absent. Achenes narrowly ellipsoid, 0.8-

0.95 mm long, slightly quadrangular, dark

brown, with 25-50 twin hairs each 0.05-0.1

mm long, evenly distributed; globose sessile

glands at junction between achene and pappus

occasionally present; pappus 2.2-2.9 mm

long. Figs. 16I-L, 18.

Additional specimens examined : Australia: Western

Australia. Hermite Island, Montebello Islands, Oct

2000, Kenneally 11558 (PERTH); Depuch Island, NE of

Roebourne, May 1962, Royce 7131 (PERTH); Charles

Knife Road, between picnic area and Cape Range N.P,

Aug 1978, Perry 845 (BRI, PERTH); Legendre Island,

Dampier Archipelago, Jun 1962, Royce 7264 (PERTH);

Barrow Island, Aug 1973, Butler 47 (PERTH); Rosemary

Island, Dampier Archipelago, Jun 1962, Royce 7451

(PERTH); Barrow Island Nature Reserve, 11.3 km

WNW of Town Point, Nov 1991, van Leeuwen 1086

(PERTH); 48.6 km NW of ‘Bonney Downs’ Homestead

on road to Hillside, Nov 1995, Mitchell PRP977 (DNA,

PERTH); Lower end of Upper Bee Gorge, Hamersley

Range N.P, Aug 1973, Weston 8443 (PERTH); Python

318

Austrobaileya 8(3): 280-334 (2011)

WESTERN AUSTRALIAN HERBARIUM, PERTH

Flora of Western Australia

Pterocaulon sp ha era ti th oides (DC.) F.Muell.

Astcraceae

Erect perennial herb.

Rock and proximal colluvium. Steep slope, aspect 330 degrees.

Many coarse fragments to maximum size of 200 mm. Abundant

bedrock outcrop. Brown loam, average depth 12.44 cm.

Scattered Low Trees of Terminalia canescens over Low

Scattered Shrubs over Hummock Grassland of Triodia sp. over

Very Open Herbs of Ptilotus exaltatus.

Coil. S. van Leeuwen et al. 0098 Date: 21/08/2005

Voucher: Pilbara Biological Survey

Loc.: Site: DRW05, West side of Karratha Stn HMS access

track, l.S km S of North West Coastal Highway. 9.2 km SW of

Mt Regal, 19.9 km SSW of Dampicr, Karratha Station, Pilbara

1BRA WA

Lai. 2G°51'14.300 W S Long. 116°40'6.700"E GDA94

Fig. 18. Representative specimen of Pterocaulon sphaeranthoides (van Leeuwen et al. 0098 [PERTH]).

Bean, Pterocaulon section Monenteles

319

Pool, Chichester-MillstreamN.P., Oct 1989, Nordenstam

& Anderberg 302 (PERTH); Yardie Creek, Cape Range

N.P., Dec 1986, Alford 816 (PERTH); Cleaverville Creek,

Roebourne, Jul 2004, Byrne 1075 (PERTH); W of Mt

Leal, on Roebourne to Wittenoon Highway, Sep 1981,

Lander 1132 (PERTH); Karrawingina Pool, Harding

River, Sep 1981, Weston 12772 (PERTH); Site OYE04,

3.4 km E of Pannawonica - Cape Lambert railway

crossing on Pannawonica - Midstream Road, Aug 2006,

van Leeuwen et al. 0102 (PERTH); Site PW17, 13.4 km

N of ‘Mt Flora’, 71.1 km E of Pannawonica, Sep 2006,

van Leeuwen et al. 0092 (BRI, PERTH); Durba Hills,

Heartland district, Jun 1984, Morse 180 (CANB); 3.1 km

SSE of Pulgorah Cone, Warrawagine Station, Pilbara,

Aug 2004, van Leeuwen 0090 (PERTH); West side of

Karratha Station Homestead access track, 1.8 km S of

North West Coastal Highway, Aug 2005, van Leeuwen

0098 (PERTH); North side of Fortescue River Mouth

access track, 6.8 km W of North West Coastal Highway,

Aug 2005, van Leeuwen 0099 (PERTH).

Distribution and habitat : This species is

endemic to Western Australia. It is common

along coastal parts of the Pilbara region,

including offshore islands, from the Cape

Range to Depuch Island, east of Roebourne.

It also occurs away from the coast, to the

south and south-east of Port Hedland, as far

east as the Durba Hills (Map 3). It grows in

Triodia grassland, or in Acacia shrubland

with Triodia understorey, on coastal sand

dunes, rocky hillsides or rocky watercourses.

Soils are often red sandy-loams. The substrate

may be limestone, sandstone or granite.

Phenology : Flowers and fruits have been

recorded from June to December.

Affinities : This species has affinities with

Pterocaulon sphacelatum, however, that

species has stem wings that are entire or

sparsely toothed, lanate hairs frequent and

glandular hairs absent from the upper leaf

surface.

Notes: This species is distinctive in the

blistered bright green narrow leaves; serrated

stem wings; lanate hairs absent or sparse and

the short broad-based glandular hairs present

on upper leaf surfaces.

Pterocaulon sphaeranthoides and

P. sphacelatum are very closely related

and some specimens are hard to identify,

suggesting that they do intergrade in some

areas. For example, Trudgen & Parnell 11608

(PERTH), Backhouse et al. BEM50 (PERTH),

and Lander 1132 (PERTH), all from the

Hamersley Ranges, while assigned here to

Pterocaulon sphaeranthoides , are also close

to P. sphacelatum. In other areas, they are

apparently sympatric; typical specimens of

both species have been seen from Barrow

Island, for example.

I have been unable to confirm the

observation made first by Bentham

(1867) that the involucres of Pterocaulon

sphaeranthoides have “Usually 2, very rarely

1 or 3” disc (hermaphrodite) florets. Cabrera

& Ragonese (1978) stated “male florets 1-

3”, as did Wilson (1992), both presumably

paraphrasing Bentham. Of the hundreds of

capitula examined by the present author,

all except one had a solitary hermaphrodite

floret. The exceptional capitulum had two

hermaphrodite florets.

12. Pterocaulon tricholobum A.R.Bean

species nova affinis P. globulifloro sed alis

caulinis multo latioribus, bracteis interioribus

longioribus, pilis numerosis in lobis

corollarum flosculorum hermaphroditorum,

achenarum pilis paucioribus et pappis

longioribus differens. Typus: Australia:

Northern Territory. East Alligator, 10 June

1971, G.C.Taylor 75 (holo: BRI; iso: CANB,

DNA).

Aromatic shrub 50-120 cm high; stems one to

several from woody rootstock, densely lanate

but without globose sessile glands; stem wings

entire, 2.8-5.5 mm wide. Leaves elliptical to

obovate, 25-92 x 8-30 mm, 2.1-3.7 times

longer than wide, apex obtuse or rarely

acute; margins recurved, denticulate, with

15-30 pairs of very small teeth. Indumentum

comprising lanate multi-cellular hairs,

glandular hairs absent. Upper surface not

or slightly bullate; hairs erect to spreading,

sparse to moderately dense, basal cell short;

globose sessile glands absent. Lower surface

with lanate hairs erect to spreading, dense to

very dense, extending 0.4-0.8 mm from the

surface, basal cells not very short; globose

sessile glands yellow and shining, moderately

dense, 3-6 diameters apart. Capitulescence

globose to broadly ellipsoidal, rarely

truncate-globose, (8—)10—16 mm long and 10-

14 mm wide at anthesis, (0.7-)0.9-1.3 times

longer than wide; side branchlets very often

320

Austrobaileya 8(3): 280-334 (2011)

270504

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HERBARIUM OF NORTHERN TERRITORY

Alice Springs, Australia

N.T.33145.

Pterooaulon sphaeranthoides (DC.) F. Muell,

Loc. Bast Alligator.

Coll. G * c * Taylor 75 Dote 10 June 1971

Notes Erect plant to 36cms. Growing

amongst rooks in sandy grey soil.

Fig. 19. Holotype of Pterocaulon tricholobum ( Taylor 75 [BRI])

Bean, Pterocaulon section Monenteles

321

continuous, at 0-80°; peduncles often absent,

but sometimes up to 11 mm long. Outer

involucral bracts oblong-spathulate, 2.5-3

mm long, with dense spreading lanate hairs

on outer surface, sparse to moderately dense

hairs on the distal end of inner surface; glands

absent; apex obtuse. Inner bracts white, rarely

tinged pink on dorsal surface; longest ones

linear, 4-4.8 x 0.2-0.35 mm, apex acute,

upper margins with some small or large teeth;

cilia 11-22, up to 2.5 mm long, attached along

the proximal half and consistently extending

to the distal one third, not exceeding the bract

apex. Filiform (female) florets 15-21, white

or straw-coloured, corolla 2.3-2.9 mm long,

styles extending 0.6-1 mm beyond corolla.

Hermaphrodite floret solitary, corolla pink,

2.9-3.6 mm long; corolla lobes 0.8-1.2

mm long, globose sessile glands present,

eglandular trichomes 2-8 per lobe. Achenes

narrowly ellipsoidal, 0.8-0.9 mm long, faintly

ribbed, dark brown, with 0-25 twin hairs each

<0.1 mm long, evenly distributed or mostly in

distal half; globose sessile glands at junction

between achene and pappus absent; pappus

2.1-3 mm long. Figs. 19, 20A-D.

Additional selected specimens examined : Australia:

Western Australia. S side of Cockburn Range, c. 13 km

W of King River, Jul 1974, Carr 3338 & Beauglehole

47116 (PERTH); Langi, 4 km S of Prior Point and

33 km SSW of Kuri Bay, Jul 1990, Kenneally 11055

(PERTH); Python Cliffs, Marigui Promontory, Prince

Regent River Reserve, Aug 1974, Kenneally 2138

(CANB, PERTH); The Grotto, Deception Ranges, near

Kimberley Research Station, Jul 1952, Lazarides 3071

(BRI, CANB, DNA); Mt Trafalgar, Prince Regent River

Reserve, Aug 1974, George 12698 (CANB, PERTH);

Inglis Gap, Gibb River Road, west Kimberley, Jun 1988,

Wilson 12959 (PERTH); Mirima [Hidden Valley] N.P,

Jun 1989, Kenneally 10947 (DNA, PERTH); Sunday

Island, Buccaneer Archipelago, Jun 1982, Kenneally

8250 (DNA, MEL, PERTH); Bungle Bungle N.P, Bull

Creek, Jun 1989, Cowie 874 (DNA, MEL, PERTH);

Gibb River Road, Home Valley Station, East Kimberley,

Jun 1992, Menkhorst 1067 (DNA, MEL, PERTH).

Northern Territory. South Bay, Bickerton Island, Gulf

of Carpentaria, Jun 1948, Specht 617 (A, BRI, CANB,

PERTH); Victoria River Gorge, May 2001, Leach &

Cowie 4659 (DNA); Gregory N.P, S of Victoria River

pub. May 2001, Risler & Smith 687 (DNA); Spirit Hills

Conservation Area; Nancy’s Gorge, Aug 1996, Cowie

7136 (DNA); 2 km S of Myra Falls, Jun 1993, Brennan

2355 (DNA); Fitzmaurice River, May 1994, Barritt 1270

(DNA, MEL); Keep River Gorge, Jun 1995, Egan 5004

(DNA); Bartelumba Bay, Groote Eylandt, Aug 1975,

Levitt 437 (CANB, DNA, NE, NT); Mt Brockman near

Koongarra Saddle, 2 km N of Koongarra, May 1980,

Lazarides 8905 (CANB, DNA); Nitmiluk N.P, eastern

boundary. May 2002, Michell 3877 (DNA); Gregory

N.P., c. 29 km NW of Bullita Outstation, Apr 1996, Coles

103 & Duretto (DNA, MEL); Keep River N.P, Apr 1982,

King 65 (DNA); Bauhinia Downs Station, Alligator

Stockyards, May 1985, Wightman 1835 & Leach

(CANB, DNA, NT); Limmen Bight, May 1996, Egan

5396 & Michell (DNA). Queensland. Burke District:

‘Westmoreland’, Big Amphitheatre, May 1997, Forster

BIF21039 & Booth (BRI, DNA).

Distribution and habitat : A widespread

species distributed in the northern Kimberley

of Western Australia, the ‘Top End’ of the

Northern Territory, and extending to the

north-west of Queensland (Map 9). The

occurrences of Pterocaulon tricholobum

are strongly associated with sandstone hills,

escarpments, scree slopes and gorges. The

vegetation type is variously recorded as

Triodia grassland, open savannah or shrubby

woodland. Soils are sandy.

Phenology : Flowers and fruits have been

recorded from May to September.

Affinities : Pterocaulon tricholobum is most

closely allied to P. globidiflorum , but differs

by the stem wings 2.8-5.5 mm wide (0.4-1.5

mm wide for P. globuliflorum ); the longest

inner bracts 4-4.8 mm long (3.2-3.5 mm

long for P. globuliflorum ); trichomes always

present on corolla lobes of hermaphrodite

floret (usually absent for P. globuliflorum );

achene hairs 0-25 (achene hairs 25-50 for

P. globuliflorum)., and pappus 2.1-3 mm long

(1.3-1.7 mm long for P. globuliflorum).

Notes : This species is distinctive in the

broad stem wings, rather small white globose

capitulescences that are often laterally and

the obvious trichomes on corolla lobes of

hermaphrodite floret.

The label of Coles 103 & Duretto records

that the plant “smells like Granny Smith

apples”.

Carr 3338 & Beauglehole 47116 is a mixed

gathering. All pieces on the CANB sheet are

Pterocaulon paradoxum ; both pieces on the

PERTH sheet are P. tricholobum.

Etymology : From the Greek trichos meaning

hair or thread, and lobus meaning lobes. This

is in reference to the conspicuous unicellular

hairs consistently borne on the corolla lobes of

322

Austrobaileya 8(3): 280-334 (2011)

Fig. 20. Pterocaulon tricholobum. A. hermaphrodite floret x 32. B. inner involucral bract (cilia not shown) x 16.

C. upper portion of inner involucral bract x 32, D. mature achene and basal portion of pappus x 32. Pterocaulon

verbascifolium. E. hermaphrodite floret x 32. F. inner involucral bract (cilia not shown) x 16. G. upper portion of inner

involucral bract x 32. H. mature achene and basal portion of pappus x 32. Pterocaulon xenicum. I. hermaphrodite

floret x 32. J. inner involucral bract (cilia not shown) x 16. K. upper portion of inner involucral bract x 32. L. female

floret x 32. M. mature achene and basal portion of pappus x 32. A-C from Egan 5004 (DNA); D from Cowie 7136

(DNA); E-H from von Oertzen 116 (DNA); I-L from Dunlop 1830 (DNA); M from George 14628 (PERTH).

Bean, Pterocaulon section Monenteles

323

the hermaphrodite floret. Other species have

this feature, but in this species the trichomes

are consistently present and obvious.

13. Pterocaulon verbascifolium (Benth.)

F.Muell., Syst. Census Austral PI. 79 (1882);

Monenteles verbascifolius Benth., FI. Austral.

3: 523 (1867). Type: Australia: Northern

Territory. Fitzmaurice River, s.dat., F.Mueller

s.n. (lecto: ¥312>2%6),fide Cabrera&Ragonese

(1978: 245).

Illustration : Cabrera & Ragonese (1978:

243).

Non-aromatic shrub 50-130 cm high; stems

very densely lanate; stem wings entire,

2.5-6(-8) mm wide. Leaves elliptical, ovate

or obovate, 37-70 x 20-43 mm, 1.6-2.3

times longer than wide, apex acute; margins

recurved, denticulate, with 5-24 pairs of

blunt teeth, sometimes obscure. Indumentum

comprising lanate multi-cellular hairs,

glandular hairs absent. Upper surface with

hairs erect to spreading, very dense, basal

cells short; globose sessile glands absent.

Lower surface with lanate hairs adpressed to

spreading, very dense, extending 0.5-0.9 mm

from the surface, basal cells not very short;

globose sessile glands sparsely present, but

hidden by hairs. Capitulescence spiciform

or cylindrical, 35-105 mm long and 17-30

mm wide at anthesis, 1.8-4.3 times longer

than wide; side branchlets never continuous,

at 30-90°; peduncles 3-40 mm long. Outer

involucral bracts 3.3-5.5 mm long, oblong

to oblong-spathulate, with dense spreading

lanate hairs on outer surfaces, inner surface

glabrous or lanate; glands absent; apex acute.

Inner bracts white on dorsal surface; longest

ones linear, 5.3-9.2 x 0.3-0.6 mm, apex acute,

upper margins with numerous small teeth;

cilia 12-30, up to 5.5 mm long, attached along

the proximal half and consistently extending

to the distal one third, not exceeding the

bract apex. Filiform (female) florets 25-58,

straw-coloured, corolla 37-7.5 mm long,

styles extending 0.4-0.9 mm beyond corolla.

Hermaphrodite floret 1(—2), corolla pink, 4-

6.5 mm long; corolla lobes 0.6-1 mm long,

globose sessile glands absent, eglandular

trichomes absent. Achenes narrowly

ellipsoidal, 1.1-1.3 mm long, terete or slightly

angular, dark brown, with 25-50 or more than

50 twin hairs each c. 0.1 mm long; globose

sessile glands at junction between achene

and pappus absent; pappus 3.5-6.9 mm long.

Figs. 20E-H, 21.

Additional specimens examined: Indonesia. Sumbawa,

s.dat., Zollinger 3421 (GH, L). Australia: Western

Australia. Dillie Gorge, Charnley River Station, Jul

2005, Byrne 1584 (PERTH); Carlton Hill Station, close to

Ningbing Site 3, Jun 2001, Handasyde & Start TH01137

(DNA, PERTH); Sir Graham Moore Island, Jul 1973,

Wilson 11317 (PERTH); SW from Mt Trafalgar, margin

of Saint George Basin, Jul 1990, Willing 212 (PERTH);

Fenelon Island, off Admiralty Gulf, Jul 1977, Kennea/ly

6412 (PERTH); Inglis Gap, King Leopold Range, May

1905, Fitzgerald 752 (PERTH); Bushfire Hill, Prince

RegentRiver Reserve, Aug 1974, George 12261 (PERTH);

around ‘Beverley Hills’ Homestead buildings, Jun 1992,

Mitchell 2557 (PERTH); Euro Gorge, Drysdale River

N.P, Aug 1975, Kenneally 4349 (PERTH). Northern

Territory. Aerodrome paddock, Manbulloo Station,

Apr 1971, Kruizinga s.n. (BRI, CANB); 7 miles NE of

Wilton River - Bulman Crossing, Jun 1972, Maconochie

1437 (BRI); ‘Bupa’, 4 km S Jarrong Yards, May 1994,

Leach 4269 & Walsh (BRI, DNA, MEL); Kakadu N.P,

Coronation Hill, Jul 1988, von Oertzen 116 (DNA);

Cave Creek Station, Snake Track, Jul 2002, Risler 1908

(DNA). Queensland, [unlocalised], s.dat.. Bowman

s.n. (K373287). Burke District: Gorge Creek gorge,

Bowthorn Station, 100 km W of Doomadgee, Aug 2001,

Addicott EPA1034 (BRI, CNS). Cook District: 21.1 km

5 of Georgetown, Aug 2010, Bean 29874 & McDonald

(BRI, CANB); 100 m from Georgetown - Forsayth

Road, 13 km S of Georgetown, Aug 2010, Bean 29861

6 McDonald (AD, BRI, NSW); 5.7 km E of Chillagoe,

Ramparts section, Chillagoe-Mungana Caves N.P, Jun

2011, McDonald 11687 & McDonald (BRI).

Distribution and habitat : Pterocaulon

verbascifolium is found commonly in the

northern Kimberley, including offshore

islands, and occasionally well inland. It is

widespread though sporadic in sub-coastal (but

not coastal) areas of the Northern Territory,

extending into northern Queensland, as far

east as Petford. Outside Australia, it is known

only from the island of Sumbawa in Indonesia

(Map 10). It inhabits lateritic plateaux or

rocky hill-slopes in eucalypt or Terminalia

dominated open woodland. Soil type varies,

and the geological substrate may be basalt or

sandstone.

Phenology : Flowers and fruits are recorded

between May and August.

Affinities : The species is not obviously allied

to any other.

324

Austrobaileya 8(3): 280-334 (2011)

STATE/DIST : NT/DG 14^5156"S 133°1035"E

CC07 Open shrub to 1.2m. Foliage grey green, densely hairy. Uncommon

in midhigh open woodland of Corymbia laiifolia and Erythrophleum

ddorostachys with Pierocaulon serrulalum and P. sphacealamm.

Risler. J.A. 1908 llJul2002

Fig. 21. Representative specimen of Pterocaulon verbascifolium ( Risler 1908 [DNA])

Bean, Pterocaulon section Monenteles

325

Notes : This species is distinctive in the dense

cylindrical capitulescences; corollas, achenes

and pappus longer than in other species; no

globose sessile glands on corolla lobes of

hermaphrodite floret; indumentum of leaves

extremely dense, obscuring surfaces; stem

wings broad and the foliage not aromatic.

Pterocaulon verbascifolium is remarkable for

the large variation between collections in the

sizes of the floral parts. In some collections, the

female florets (for instance) are only 3.7 mm

long, and in other collections they are up to

7.5 mm long. There are correlated differences

in other floral parts; however, there does not

appear to be any geographical or ecological

pattern to this variation.

There is a specimen of Pterocaulon

verbascifolium at K collected by E.M.

Bowman, with the locality given merely

as “Queensland”. It is of interest because

Bowman did not travel any further north or

west than the upper reaches of the Flinders

River, near present-day Hughenden (Blake

1955). Hughenden is more than 200 km beyond

the currently known extent of the species.

14. Pterocaulon xenicum A.R.Bean species

novaaffinisP.ra'veoaliscaulinisangustioribus,

numero majore flosculorum femineorum in

quoque capitulo, lobis brevioribus corollae

flosculorum hermaphroditorum et pappis

longioribus differens. Typus: Australia:

Western Australia. McLarty Hills, Great

Sandy Desert, 5 August 1977, A.S. George

14628 (holo: PERTH; iso: CANB).

Shrub 25-40 cm high; stems numerous,

densely lanate and sometimes with a few

globose sessile glands; stem wings entire,

0.3-0.5 mm wide. Leaves oblanceolate or

obovate, 18-52 x 7—17 mm, 2.6-47 times

longer than wide, apex obtuse or acute;

margins recurved, denticulate, with 10-21

pairs of blunt teeth. Indumentum comprising

lanate multi-cellular hairs, glandular hairs

absent. Upper surface moderately bullate;

hairs erect to spreading, sparse to dense,

basal cells short; globose sessile glands

absent. Lower surface with lanate hairs erect

to spreading, very dense, extending 0.4-0.7

mm from the surface, basal cells not very

short; globose sessile glands yellow and

shining, moderately dense, 3-8 diameters

apart, but usually obscured by indumentum.

Capitulescence globose to ellipsoidal, 13-18

mm long, 13-16 mm wide at anthesis, 1-1.3

times longer than wide; side branchlets never

continuous, at 30-80°; peduncles 0-14 mm

long. Outer involucral bracts 2.5-2.9 mm

long, spathulate, with dense spreading lanate

hairs throughout on the outer surface, glabrous

on the inner surface or with sparse hairs at

distal end; glands absent; apex lacerate to

fimbriate (fimbriae 0.1-0.35 mm long). Inner

bracts pink or white on dorsal surface; longest

ones linear, 4.8-5.6 x 0.25-0.4 mm; apex

acute, upper margins conspicuously toothed

or with small teeth; cilia 6-9, up to 3.8 mm

long, mostly attached in proximal half and

few extending to the distal one third. Filiform

(female) florets 26-44, pink, corolla 3.2-4.1

mm long, styles extending 0.2-0.4 mm beyond

corolla. Hermaphrodite floret solitary, corolla

pink, 3.5-4.3 mm long; corolla lobes 0.3-0.5

mm long, globose sessile glands present,

eglandular hairs absent or present. Achenes

narrowly ellipsoidal, 0.8-0.9 mm long, terete

or slightly angular, dark brown, with 25-50 or

>50 twin hairs each c. 0.05 mm long, evenly

distributed; globose sessile glands at junction

between achene and pappus sparsely present;

pappus 3.3-4.2 mm long. Figs. 20I-M, 22.

Additional specimens examined : Australia: Northern

Territory. 46 miles [74 km] WSW of The Granites, Aug

1970, Dunlop 1830 (DNA). Western Australia, just N

of Dragon Tree Soak, Great Sandy Desert, Aug 1977,

George 14748 (PERTH); 21.1 km ESE of Warrawagine

Station Homestead, Pilbara IBRA (Site PHYE07), Aug

2006, van Leeuwen et al. 0028 (BRI, PERTH); 13.3 km

W of Warrawagine Station Homestead, Pilbara IBRA

(Site PHYE06), Aug 2006, van Leeuwen et al. 0027

(BRI, PERTH); E of Cherrabun Station, Oct 1952, s.coll.

(PERTH536415).

Distribution and habitat: Known from a

few localities in northern Western Australia,

extending from Warrawagine Station to the

Tanami Desert in Northern Territory (Map 4).

At the type locality it occurs on open plains in

association with shrubs including Acacia spp.,

Senna spp., Solanum lasiophyllum Dunal

ex Poir. and Grevillea juncifolia Hook. The

understorey is dominated by spinifex ( Triodia

spp.)

326

Austrobaileya 8(3): 280-334 (2011)

hO/jrfYffsf Queensland Herbarium (BRI)

Pf'To com/ on xen /curt A . Boo/)

Del v&*2/U Zt>{/

PERTH 00541605

\Q-Oob

duplicates

rjicje^MT'D

FLORA OF THE KIMBERLEY REGION - W.A. HERBARIUM

’Pi-esocAvdo^ C£>C-) R NWAJt.

Signed _.. 1=^ 19^

WESTERN AUSTRALIAN HERBARIUM, PERTH

Flora of Western Australia

j£Lr~6 ^5 ~cv*; Arv-cSy .

‘f&A. / .

A t<J -4m*,

x*t /~<M .. Gx~j- S*~A*.

L 00 - A/.X}.

Co ll. <A. S.'^<rr^ - /x-6x§’ J- ^^, 19 ??

Fig. 22. Holotype of Pterocaulon xenicum (i George 14628 [PERTH]).

Bean, Pterocaulon section Monenteles

327

Phenology : Flowers and fruits are recorded

for August and October.

Affinities : The species shows affinity with

Pterocaulon sphacelatum and P. niveum.

Pterocaulon xenicum differs from P. niveum

by the stem wings 0.3-0.5 mm wide (0.5-1.5

mm for P. niveum ); the 26-44 female florets

(11-17 for P. niveum ); the corolla lobes of the

hermaphrodite floret 0.3-0.5 mm long (0.7-

1.1 forP. niveum ); and the pappus 3.3-4.2 mm

long (2-3.1 mm for P. niveum ).

Pterocaulon xenicum differs from

P. sphacelatum by the stem wings 0.3-0.5 mm

wide (0.7-2.0 mm forP. sphacelatum ); the longest

inner bracts 4.8-5.6 mm long (3.5-4.6 mm

for P. sphacelatum ); the female florets 3.2-4.1

mm long (2.4-3.2 mm for P. sphacelatum ); the

hermaphrodite florets 3.5-4.3 mm long (2.5-37

mm long for P. sphacelatum ); the pappus 3.3-

4.2 mm long (2.3-3.3 mm for P. sphacelatum );

and the outer bracts with lacerate or fimbriate

apices (acute apices for P. sphacelatum ).

Notes : This species is distinctive for the

stem wings being very narrow and almost

redundant; leaf indumentum very dense on

lower surface; corolla lobes of hermaphrodite

floret short; none or few cilia visible on

protruding part of inner bracts; apices of outer

bracts lacerate to fimbriate and the relatively

long hermaphrodite florets.

Etymology : Derived from the Greek word

xenos meaning ‘stranger, foreigner’ and the

suffix -icus ‘belonging to’. This alludes to the

specimens of this species found in amongst

collections of Pterocaulon sphacelatum , and

looking like ‘strangers’.

Postulated hybrids

Pterocaulon intermedium x p. paradoxum.

Specimens examined: Australia: Western Australia.

9 km N of Bunda Bunda Mill on Baldwin Creek

track, Dampier Peninsula, Jun 1984, Kenneally 9078

(PERTH).

This specimen has broad stem wings,

somewhat ellipsoidal capitulescences, achenes

c. 0.8 mm long and scarcely acute leaves. These

features do not fit Pterocaulon intermedium ,

nor do they quite fit P. paradoxum , although

the specimen has similarities to both. This

specimen is probably a hybrid between these

two species, both of which occur on the

Dampier Peninsula.

Pterocaulon serrulatum var. velutinum x

P. sphacelatum.

Specimens examined: Australia: Western Australia.

4.8 km E of Donkey Creek, on the Gibb River Road,

Sep 2006, Sweedman 6984 (PERTH); 6.5 km NE of

Mary River Crossing, Great Northern Highway, 92 km

SW of Halls Creek, Jun 1976, Beauglehole ACB53238

(PERTH); Site PHYE09, 10.3 km SW of Warrawagine

Station Homestead, Pilbara, Aug 2006, van Leeuwen et

al. 0035 (PERTH).

These specimens have glandular outer

involucral bracts as in Pterocaulon

serrulatum , but the leaves are smaller and

rather more bullate than is usual for that

species. Furthermore, the stem wings are

almost entire, with just the occasional tooth.

Pterocaulon ciliosum x p. serrulatum var.

serrulatum.

Specimens examined: Australia: Queensland. North

Kennedy District: 22.5 km (by road) SW of Herberton,

on the Silver Valley road. May 1983, Conn & Clarkson

1177 (BRI, CANB, DNA, NSW).

This specimen has narrower stem wings than

Pterocaulon serrulatum , and the toothing

is less than usual for that species. The

capitulescences are almost globose and the

protruding part of inner bracts has many long

cilia, both features of Pterocaulon ciliosum.

Excluded names

Pterocaulon tomentosus Boerl., Handl. FI.

Ned. Ind. (Boerlage) 2(1): 240 (1891).

This is a nomen nudum.

Monenteles tomentosus Sch. Bip., Syst. Verz.

(Zollinger) 122(1854).

This is a nomen nudum.

Gnaphalium redolens G.Forst., FI. Ins. Austr.

91 (1786).

This is a nomen nudum.

Acknowledgements

The Directors of several Herbaria have

kindly sent specimens on loan from their

institutions. I gratefully acknowledge

G.Rouhan (P), M.Pignal (P), P.Morat (P)

328

and P. Schafer (MPU) for their assistance or

advice regarding type specimens collected

from New Caledonia. I am grateful to N. Hind

(K) who dealt with my loan request promptly

and efficiently. V.Fonjallaz (G) and C.Nepi

(FI) kindly sent images of type specimens

held at those institutions. I thank G.von

Poser of Porto Alegre, Brazil, for sending a

reprint of a Pterocaulon chemistry paper.

K.McDonald, R.Booth and D.Kelman have

collected additional Pterocaulon material

at my request. W. Smith (BRI) provided the

illustrations of florets, bracts and achenes,

and PBostock (BRI) expertly translated the

diagnoses into Latin.

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Austrobaileya 8(3): 280-334 (2011)

330

Map 2. Distribution of Pterocaulon ciliosum ■. P. niveum O

Bean, Pterocaulon section Monenteles 331

Map 3. Distribution of Pterocaulon discolor ▲, P. globuliflorum □, P. sphaeranthoides •

Map 4. Distribution of Pterocaulon intermedium •, P. xenicum ▲

332

Austrobaileya 8(3): 280-334 (2011)

Map 5. Distribution of Pterocaulon paradoxum

Map 6. Distribution of Pterocaulon redolens

Bean, Pterocaulon section Monenteles

333

Map 7. Distribution of Pterocaulon serrulatum var. serrulatum •, P. serrulatum var. velutinum ▲

Map 8. Distribution of Pterocaulon sphacelatum

334

Austrobaileya 8(3): 280-334 (2011)

Map 9. Distribution of Pterocaulon tricholobum

Map 10. Distribution of Pterocaulon verbascifolium

Heliotropium microspermum E.J.Thomps.

(Boraginaceae), a new species from Queensland

E. J. Thompson

Summary

Thompson, E.J. (2011). Heliotropium microspermum E.J.Thomps. (Boraginaceae), a new species

from Queensland. Austrobaileya 8(3): 335-339. The new species Heliotropium microspermum

E.J.Thomps., endemic to Cape York Peninsula, Queensland, is described and illustrated.

Key Words: Boraginaceae, Heliotropium , Heliotropium microspermum , Queensland flora, taxonomy,

new species

E.J. Thompson, Queensland Herbarium, Department of Environment & Resource Management,

Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email: john.

thompson@der m. qld. gov. au

Introduction

The genus Heliotropium L. comprises 84

species in Australia (Craven 1996, 2005a,b)

with 43 (38 native, five naturalised) occurring

in Queensland (Thompson 2010). A critical

review of Heliotropium specimens at the

Queensland Herbarium has revealed a new

species for Cape York Peninsula.

Materials and methods

Morphological data were collected from dried

herbarium material at BRI. Description of the

new species largely follows Craven’s (1996)

format.

Indumentum is a useful character in

Australian Heliotropium and is usually

“composed of unicellular, simple, non-

glandular hairs (the ‘standard’ hairs) with

multicellular, simple, glandular hairs also

occurring in some species. The walls of

standard hairs may be smooth or ornamented”

(Craven 1996). Craven (1996) refers to

indumentum as having more than one class of

hair when hairs can be differentiated, such as

long coarse hairs and shorter fine hairs.

Mericarps are essential for identifying

Heliotropium species and the mericarp

characters used were length, commissure (side

faces) pit shape and size; presence or absence

of a food body in the pits; size, distribution

and type of the hairs on the back; and shape

of the apex. Although Craven (1996) provided

Accepted for publication 23 August 2011

images of some mericarp indumentum types

he did not present illustrations of mericarps

for each species. Illustrations of mericarps

are provided here for the new species and its

putative allies. Other significant characters

included leaf width and hair type, inflorescence

characters, and plant growth form.

Taxonomy

Heliotropium microspermum E.J.Thomps.

species nova affinis H rhadinostachyum

Craven sed pilis applanatis fortuito dispositis,

fovea circulari minore in mericarpiis

minoribus et corolla breviore differens. Typus:

Queensland. Cook District: 12.3 km along

Pormpuraaw Road from Gulf Development

Road near Musgrave, 15 November 2010,

K.R.McDonald KRM10095 (holo: BRI).

Prostrate to ascending perennial herbs,

5-15 cm tall and to 15 cm broad. Branches

moderately hairy with ascending standard

hairs of one class. Adult leaves ascending,

linear to narrowly elliptic, 5-13 mm long, 2-3

mm wide, base attenuate, apex acute; petiole

short; abaxial surface moderately densely

hairy with appressed to ascending hairs;

adaxial surface with ornamented hairs, 07-1.2

mm long, similar to the abaxial surface only

hairs tending towards more numerous along

the midrib. Cyme on mature plants at anthesis

straight, simple or branched, bracteate; bracts

conspicuous, linear to narrowly elliptic, 2.5—

4.5 mm long, touching to separated up to one

length along the cymes, with ornamented

hairs on adaxial surface appressed on lamina

.i—----'-'T.

336

Austrobaileya 8(3): 335-339 (2011)

Fig. 1. Heliotropium microspermum. A. corolla *16. B. corolla expanded demonstrating anthers and appressed hairs

in throat x 16. C. pistil x 32. D. plant x 0.6. E. calyx x 16. F. mericarp back view x 40. G. mericarp side face x 40. All

from McDonald KRM10095 (BRI). Del. W.Smith.

Thompson, Heliotropium microspermum

337

and ascending on margin, abaxial surface

glabrous; varying from clearly different to

similar in shape and size to the leaves. Calyx

lobes with ornamented hairs 0.5-0.8 mm long

on the abaxial surface, appressed on the lamina

and ascending on the margin, adaxial surface

glabrous; outermost calyx lobe narrowly

ovate, 2-2.6 mm long, acute. Corolla 2.5-3

mm long, white, tube cylindrical, not swollen

around anthers at anthesis, 1.5-3 mm long, the

outer surface sparsely hairy with appressed

hairs, the inner surface glabrous in the throat;

lobes broadly triangular, 0.7-1 mm long, apex

acute. Anthers ovate, 0.3-0.5 mm long, apex

acuminate, papillate. Gynoecium c. 1.6 mm

long; ovary c. 0.5 mm long; style 0.5-0.7 mm

long; stigma 0.3-0.4 mm long. Mericarps

ovate to broadly ovate, 0.6-0.7 mm long, 0.4-

0.5 mm wide, 1.4 times as long as wide; apex

acute; outer surface with mostly appressed

to slightly ascending microhairs, 0.04-0.1

mm long, flattened, mostly distributed along

minute ridges over most of surface and

somewhat randomly orientated; commissures

with pit to about 14 width of commissure, the

pit elliptic, without a food body. Fig. 1.

Additional specimens examined: Queensland.

Cook District: 36.3 km from Dixie Station on track

to Killarney, Jun 1989, Clarkson 8154B & Neldner

(BRI); 13.9 km along Pormpuraaw Road from Gulf

Development Road, Nov 2010, McDonald KRM10093

(BRI); 6 km N of‘Fairview’ on the Peninsula Road, Apr

1980, Clarkson 3090 (BRI); 12.2 km along Pormpuraaw

Road from Cape York Development Road junction, Dec

2010, McDonald KRM10303 (BRI).

Distribution and habitat: Heliotropium

microspermum is only known from near

Musgrave on Cape York Peninsula (Map 1).

It has been found in woodland of Melaleuca

viridiflora Sol. ex Gaertn. on flat terrain,

and eucalypt woodland dominated by

Eucalyptus tetrodonta F.Muell. or Corymbia

stockeri subsp. peninsularis (K.D.Hill &

L.A.S.Johnson) A.R.Bean or C. nesophila

(Blakely) K.D.Hill & L.A.S.Johnson on

granite hills with sandy soil.

Phenology: Flowers and fruits have been

recorded in June and November.

Affinities: Heliotropium microspermum is

similar to H rhadinostachyum with respect

to growth habit, appearance of the cymose

inflorescences, and leaf and bract shape,

size, and distribution along the branches.

Heliotropium microspermum differs most

notably from H. rhadinostachyum and H.

cunninghamii Benth. by the size and shape

of the corollas and mericarps, as well as

indumentum type and cover (Table 1).

Notes: Juvenile leaves of Heliotropium

microspermum can be larger than adult leaves

and therefore more clearly different from

the bracts. Juvenile plants of this species

may have very short cymes. The corolla of

Heliotropium microspermum can be a little

swollen around the anthers at fruiting.

Conservation status: Heliotropium

microspermum is only known from near

Musgrave. Inappropriate burning regimes

present the most significant potential threat

to this species. Further field observation is

required to determine the species population

size and area of extent.

Etymology: The specific epithet is derived

from the Greek micro (small) and spermum

(seed) alluding to the relatively small

mericarps for this species.

Map 1. Distribution of Heliotropium microspermum in

Queensland

338 Austrobaileya 8(3): 335-339 (2011)

Table 1. Comparison of morphological characters for Heliotropium microspermum,

H. rhadinostachyum and H. cunninghamii

Character State

H. microspermum

H. rhadinostachyum

H. cunninghamii

mericarp length (mm)

0.6-0.7

0.9-1.2

0.6-1.2

mericarp apex shape

acute

rounded

acute

mericarp microhair

length (mm)

0.04-0.1

to c. 0.05 or

occasionally absent

0.05-0.1

mericarp microhair

type

flattened

papillate

circular in cross-

section

mericarp microhair

distribution

most of surface

upper 14

upper 14-14

mericarp pit size

small, up to Vi width

of commissure

large, c. 14 the width of

the commissure

small, up 14 width of

commissure

mericarp pit shape

elliptic

circular

mericarp outer surface

minutely ridged over

most of surface

mostly smooth,

occasionally minutely

ridged in upper 14

smooth

corolla length (mm)

2.5-3

3-4

5-6

corolla tube shape at

anthesis

cylindrical

swollen around anthers

calyx lobe trichome

length (mm)

0.5-0.8

0.3-0.4

bract length (mm)

2.5-4.5

2-4.5

1-2.5

bract

conspicuous; leaves

inconspicuous; leaves

distinctly different to

distinctly different

similar

bract imbrication

touching to separated

overlapping by 14 a

widely separated

by about 1 length

length to separated by

up to 2 lengths

Acknowledgements

I am grateful to David Halford (BRI) and

Tony Bean (BRI) for reading drafts of the

manuscript. I thank Will Smith (BRI) for

the illustrations and map, and Peter Bostock

(BRI) for the Latin diagnosis.

References

Craven, L.R. (1996). A taxonomic revision of

Heliotropium (Boraginaceae) in Australia.

Australian Systematic Botany 9: 521-657.

- (2005a). Malesian and Australian Tournefortia

transferred to Heliotropium and notes on

delimitation of Boraginaceae. Blumea 50: 375-

381.

- (2005b). Seven new species of Heliotropium

(Boraginaceae) from the monsoon and arid

zones of Australia. The Beagle 21: 11-25.

Thompson, E. J. (2010). Boraginaceae. In P.D. Bostock &

A.E. Holland (eds.). Census of the Queensland

Flora 2010 , pp. 29-30. Queensland Herbarium,

Department of Environment & Resource

Management: Brisbane.

Thompson, Heliotropium microspermum

339

Fig. 2. A-D: Heliotropium rhadinostachyum. A. corolla * 16. B. outer calyx lobe x 16. C. mericarp backview x 40.

D. mericarp side view x 40. E-H: Heliotropium cunninghamii. E. corolla x 16. F. outer calyx lobe x 16. G.

mericarp back view x 40. H. mericarp side face x 40. A-D from Clarkson & Neldner 8146 (BRI); E-H from

Thompson & Booth TAN239 (BRI). Del. W.Smith.

Two new species of Pluchea Cass. (Asteraceae:

Plucheinae) from Queensland, Australia

A.R.Bean

Summary

Bean, A.R. (2011). Two new species of Pluchea Cass. (Asteraceae: Plucheinae ) from Queensland,

Australia. Austrobaileya 8(3): 340-346. The new species Pluchea punctata A.R.Bean and P.

xanthina A.R.Bean are described from Queensland, with the latter replacing the misapplied name

P. dioscoridis (L.) DC. Illustrations, a distribution map, and notes on the habitat and conservation

status are provided for each new species. A note is given regarding the apparent sexual dimorphism

of Pluchea baccharoides. A key to the identification of Queensland species of Pluchea is included.

Key Words: Asteraceae, Pluchea, Pluchea baccharoides, Pluchea punctata, Pluchea xanthina,

Australia flora, Queensland flora, taxonomy, new species, identification key

A.R.Bean, Queensland Herbarium, Department of Environment & Resource Management, Brisbane

Botanic Gardens, Mt Coot-tha Road, Toowong 4066, Queensland, Australia. E-mail: tony.bean@

derm.qld.gov.au

Introduction

Pluchea Cass, is a genus of about 50 species,

with representatives throughout the tropics

and subtropics of the world, but especially

in eastern Africa, the Arabian Peninsula and

north-western Australia (King-Jones 2001;

Thulin 2004). The Australian members of

the genus are perennial shrubs or subshrubs,

often woody, with alternate toothed leaves,

and pink tubular florets. The involucral bracts

are acute, and borne in several rows. The

achenes are cylindrical to ellipsoidal, and the

pappus comprises many barbellate capillary

bristles.

Hunger (1996) described the new species

Pluchea dunlopii from Australian material,

and soon after provided a revision of the

genus for Australia wherein six species were

enumerated (Hunger 1997).

Two new species are described here, both

endemic to Queensland. Pluchea xanthina

A.R.Bean is a moderately widespread species

for which the name P. dioscoridis (L.) DC.

has been misapplied. Pluchea punctata

A.R.Bean is a newly discovered species with

a very restricted distribution. Both species

are illustrated and compared with related

taxa. A distribution map is provided, as is an

identification key for the Queensland species

of Pluchea.

Materials and methods

This paper is based on a study of herbarium

specimens at BRI, an image of the type

specimen of Blumea dioscoridis at LINN

and examination of Pluchea populations and

plants in the field. Measurements of florets

were made from material preserved in spirit

or reconstituted in boiling water, while leaves,

bracts and achenes were measured from dried

specimens.

Abbreviations used in the text include N.R

(National Park) and S.F. (State Forest).

Taxonomy

Pluchea xanthina A.R.Bean species nova

affinis P. wallichianae, sed foliis petiolatis

angustioribus, bracteis involucral ibus

exterioribus longioribus, marginibus

superioribus integris in bracteis involucral ibus

omnibus et corollae lobis longioribus

flosculorum discoideorum differens. Typus:

Queensland. South Kennedy District:

Llanarth Back Range Road, 9.4 km S of

junction with Scartwater Road, 15 May 1991,

VJ.Neldner 3479 & E.J.Thompson (holo:

BRI; iso: AD, K).

Pluchea dioscoridis auct. non (L.) DC.;

Mueller (1889); Bailey (1900); Hnatiuk (1990);

Henderson (1997); Holland (2007, 2010).

Accepted for publication 27 June 2011

Beam. Two new Pluchea (Asteraceae)

341

Woody shrub to 150 cm high, rarely higher.

Stems greento brown, terete, erectto spreading,

well-branched, with sessile yellow glands and

multicellular eglandular hairs to 0.3 mm long,

dense on young stems, glabrescent on older

stems. Leaves petiolate, narrowly-elliptic to

oblanceolate, spreading, not decurrent, 24-68

x 4-17 mm, 3.8-6.4 times longer than broad,

margins denticulate or occasionally dentate,

apex acute; both surfaces with many sessile

yellow glands and multicellular eglandular

hairs; petioles 5-15 mm long. Capitula in

small terminal clusters, campanulate, 7-8 x

6-7 mm; peduncles 1—6(—16) mm long, with

1-3 lanceolate bracts near the apex, each l-3(-

9) mm long. Involucral bracts in several rows;

outer bracts ovate, 3.2-4.2 x 1.3—1.7 mm, with

apex acute, outer surface with sessile yellow

glands and dense eglandular hairs, margins

entire distally; median bracts elliptic, 4.7-5 x

1.2-1.6 mm, apex acute, distal half of outer

surface with sessile yellow glands and dense

eglandular hairs, upper margins entire; inner

bracts linear, 6-6.7 x 0.4-0.8 mm, apex acute,

outer surface glabrous, upper margins entire.

Receptacle epaleate, verrucose, flat, 1.6-2.8

mm diameter. Outer florets 75-100, female;

corolla very slender, 4.3-6 mm long, pink;

lobes 3, triangular, 0.2-0.3 mm long, glabrous;

style bifid, exserted, glabrous; pappus 4.5-5.1

mm long, pappus bristles 9-13, all of similar

length, barbellate. Disc florets 4-8, each with

an ovary and divided style but functionally

male; corolla cylindrical, 4.5-6 mm long,

purple; lobes 5, triangular, 0.7-1 mm long,

glandular; anthers c. 2 mm long, including tail

c. 0.2 mm long; style bifid, exserted by 0.5-1

mm, with acute sweeping hairs; pappus 5.2-6

mm long, pappus bristles c. 20, barbellate, all

of similar length. Achenes cylindrical, 0.9-1.2

mm long, with scattered appressed twin hairs;

carpopodium prominent, white. Fig. 1.

Additional selected specimens examined : Queensland.

Burke District: Micky Spring, ‘Clyde Park’, N of

Hughenden, Aug 1984, Godwin C2715 (BRI). North

Kennedy District: Lynd Scrub, c. 64 km SW to SSW of

Mt Garnet, Aug 1948, Smith 3857 (BRI); 15 km S of ‘Mt

McConnell’ Homestead, Jun 1992, Thompson CHA60

& Sharpe (BRI). South Kennedy District: 6 km NE of

‘Moonoomoo’ Homestead in Darkies Range, Apr 1992,

Thompson BUC354 & Simon (BRI); Cudmore N.P.,

Sep 2001, Augustyn JA5 (BRI). Leichhardt District:

Gogango Range, near Edungalba, Sep 1943, Blake 15354

(BRI); Southern end of S.F. 236, SW of Blackwater, Nov

2002, Bean 19528 (BRI); c. 30 miles [48 km] WSW of

Springsure, Sep 1966, Everist 7939 (BRI); Goodliffe

section, Carnarvon N.P, 12 km W of eastern boundary

of park, along road from Springsure, Sep 2009, Halford

Q9690 (BRI); 17 miles [28 km] E of Duaringa, Aug

1963, Speck 1667 (BRI, CANB, NSW); ‘Early Storms’,

Carnarvon Creek, Sep 1974, Gittins 2764 (BRI,

NSW). Port Curtis District: 12.9 km by road SE of

Maryborough towards Rockhampton, Sep 1975, Coveny

6908 & Hind (BRI); ‘Maryvale’, between Byfield and

Yeppoon, Sep 1931, White 8036 (BRI); N of The Caves,

near Yaamba, Rockhampton, Aug 1992, Barton 4 (BRI).

Burnett District: Cole’s Lookout, Coominglah S.F., W

of Monto, Aug 1995, Bean 8856 (BRI, MEL).

Distribution and habitat: Pluchea xanthina is

endemic to Queensland. It occurs from Forty

Mile Scrub N.P. in northern Queensland south

to the Carnarvon Ranges and Monto, with

several collections from the Rockhampton

area (Map 1). The species grows on hilly

terrain in eucalypt woodland, often with

Corymbia citriodora (Hook.) K.D.Hill &

L.A.S. Johnson ox Eucalyptus crebra F.Muell.,

on sandy-loamto clay-loam soils. It sometimes

grows on the margins of brigalow forest, and

near Rockhampton it has also been recorded

growing on serpentinite.

Phenology: Flowers and fruits have been

recorded from May to November.

Notes: Mueller (1889) was the first to apply

the name Pluchea dioscoridis to Australian

material. This name was then employed by

Bailey (1900), and subsequent authors have

followed that usage. A high quality image

of the type of Blumea dioscoridis L. (LINN

992.6) has been viewed on the Linnean

Society website (Anon. 2010). It shows a rather

broad-leaved plant with sessile auriculate-

based leaves, clearly different to the taxon

found in Queensland. According to Boulos

(2002), the type was collected in Egypt by

F. Hasselquist. King-Jones (2001) recorded

Pluchea dioscoridis from central and northern

Africa and the Arabian Peninsula.

Pluchea xanthina differs from

P. dioscoridis in several characters based

on the description of the latter by King-

Jones (2001): leaf base attenuate (auriculate

in P. dioscoridis ); capitula 7-8 mm long

(4.5-(5-7)-9) mm in P. dioscoridis );

involucral bracts with margins entire distally

342

Austrobaileya 8(3): 340-346 (2011)

Fig. 1. Pluchea xanthina. A. flowering branchlet * 0.5. B. portion of leaf surface showing eglandular hairs and sessile

glands x 12. C. capitulum x 6. D. median involucral bract x 12. E. mature achene and pappus x 12. F. receptacle of

spent capitulum and bract-bases x 12. G. disc floret x 8. A, B from Neldner 3479 & Thompson (BRI); C, D from

Halford Q9690 (BRI); E-G from Bean 20718 (BRI).

Beam. Two new Pluchea (Asteraceae)

343

(fimbriate in P. dioscoridis)', inner involucral

bracts 6-6.7 mm long (4.2-5.2 mm long in

P. dioscoridis ); outer florets 4.3-6 mm long

(2.7-4 mm in P. dioscoridis ); and pappus

bristles all the same length (of varying length

in P. dioscoridis ).

Pluchea xanthina is perhaps most closely

allied to P. wallichiana DC., a species from

India and Pakistan. Pluchea xanthina differs

by the leaves 0.4-1.7 cm wide (1.5—2.5(—3) cm

in P. wallichiana ); leaves consistently petiolate

(consistently sessile and sometimes auriculate

in P. wallichiana ); outer involucral bracts 3.2-

4.2 mm long (2-2.5(-2.8) in P. wallichiana );

involucral bract margins entire (fimbriate in

P. wallichiana ); and corolla-lobes of the disc

florets 0.7-1 mm long (0.4-0.6 mm long in

P. wallichiana).

Conservation status : A widespread species,

that it is not currently under threat.

Etymology : The epithet is from the Greek

xanthos meaning yellow and inos, of the

nature of It refers to the shining yellow glands

that are found commonly on many parts of the

plant.

Pluchea punctata A.R.Bean species nova

affinis P. baccharoidi sed capitulo longiore,

marginibus glandularibus bractearum

involucralium, flosculis discoideis corollae

lobis quattuor, pappo longiore et acheniis

ellipsoidalibus brevioribus differens. Typus:

Queensland. Cook District: Gilbert River

Holding, 78 km by road S of Forsayth,

13 August 2010, A.R.Bean 29877 &

K.R.McDonald (holo: BRI [1 sheet + spirit];

iso: AD, B, CNS, MEL, NSW, US).

Woody shrub to 80 cm high. Stems green

to brown, terete, erect to spreading, well-

branched, glabrous, viscid. Leaves sessile,

1 inearto very narrowly oblanc eolate, spreading,

not decurrent, 18-54 x 1-2.5 mm, 13-23 times

longer than broad, viscid or resinous, margins

entire, apex acute; both surfaces punctate

with conspicuous embedded glands. Capitula

in small terminal clusters, narrowly ovoid to

campanulate, 7-10 x 4-5 mm; peduncles 1-10

mm long, with 1-3 lanceolate bracts near the

apex, each 1.5-2.5 mm long. Involucral bracts

in several rows; outer bracts ovate, 2.5-3.5 x

1.5-1.7 mm, apex acuminate, outer surface

and margins with stipitate glandular hairs;

median bracts elliptic, 3.5-47 x 1.2-1.8 mm,

apex obtuse to acuminate, outer surface and

distal margins with stipitate glandular hairs;

inner bracts linear, 5-6.3 x 0.4-0.6 mm, apex

acute, glabrous or with a few stipitate glands

near apex. Receptacle epaleate, verrucose,

flat, c. 2 mm diameter. Outer florets 80-120,

female; corolla very slender, 4.2-5.5 mm

long, pink; lobes 3, triangular, 0.15-0.25 mm

long, glabrous; style bifid, exserted, glabrous;

pappus 3.7-5 mm long, pappus bristles 13-17,

all of similar length, barbellate. Disc florets

8-12, each with an ovary and divided style

and appearing bisexual but functionally male;

corolla cylindrical, 5.2-6.5 mm long, purple;

lobes 4, triangular, each 0.6-0.7 mm long,

glandular; anthers c. 2 mm long, including

a tail c. 0.3 mm long; style bifid, exserted

by 1.5-2 mm, with acute sweeping hairs;

pappus 4.3-5.5 mm long, pappus bristles c.

15, all of similar length, barbellate. Achenes

narrowly ellipsoid, 0.8-1.0 mm long, with

scattered appressed twin hairs; carpopodium

prominent, white. Fig. 2.

Additional specimen examined: Queensland. Cook

District: Gilbert River holding, 40 km by road from

Cobbold Gorge turnoff, Oct 2007, McDonald KRM6961

(BRI, CANB).

Distribution and habitat: Pluchea punctata

is known only from the type locality (Map

1). It occurs on the lower slope of a steep hill

amongst Triodia sp., in an exposed area with

scattered trees of Eucalyptus microneura

Maiden & Blakely and Grevillea striata R.Br.

Soil is a reddish-brown sandy-loam.

Phenology: Flowers and fruits have been

recorded in August and October.

Affinities: Pluchea punctata is superficially

very similar to P. baccharoides (F.Muell.)

Benth. as both are small woody shrubs with

viscid gland-dotted, linear leaves. However,

Pluchea punctata can be distinguished by

the longer leaves 18-54 mm long (6.5-19

mm long for P. baccharoides ); capitula 7-10

mm long (6-7 mm long for P. baccharoides );

involucral bracts with stipitate glandular hairs

on margins (eglandular fimbriate margins

for P. baccharoides ); disc florets 8-12 (1-2

or 13-18 for P. baccharoides ); disc florets

344

Austrobaileya 8(3): 340-346 (2011)

Fig. 2. Pluchea punctata. A. flowering branchlet x 0.5. B. portion of leaf surface showing embedded glands x 16.

C. capitulum x 6. D. median involucral bract x 16. E. disc floret x 8. F. mature achene and pappus x 16. A-E from Bean

29877 & McDonald (BRI); F from McDonald KRM6961 (BRI).

Beam. Two new Pluchea (Asteraceae)

345

with 4 glandular corolla lobes (5 eglandular

lobes for P. baccharoides ); female florets

with pappus 3.7-5 mm long (2.5-3 mm for

P. baccharoides ); and achenes narrowly

ellipsoid, 0.8-1 mm long (linear, 1.4-1.7 mm

long for P. baccharoides ). The hillside habitat

of Pluchea punctata is quite different to the

strictly riparian habitat of P. baccharoides.

Conservation status : There are between 40

and 50 plants at the type locality. Similar

habitats along the roadside north and

south of the type locality were searched

without locating any additional plants. A

conservation status of Critically Endangered

is recommended, based on criterion D of the

IUCN Red List (IUCN 2001).

Etymology : The epithet is from the Latin

punctatus , dotted, which derives from the

Latin punctum ‘a spot, dot, or point’, and

refers to the dot-like glands embedded in the

leaves of this species. Such glands are found

in only a few species of Pluchea.

A note on Pluchea baccharoides

Hunger (1997) noted that Pluchea

baccharoides produces both homogamous

and heterogamous capitula; however, she gave

no indication as to whether homogamous and

heterogamous capitula appear on the one plant

or on different plants. Hunger (1997) recorded

2-30 disc florets and 0-60 female florets

for P. baccharoides ; these observations are

correct but rather misleading, as they imply a

continuum of floret numbers.

There are currently 17 flowering specimens

of Pluchea baccharoides at BRI. Of these,

11 were found to have only heterogamous

capitula with 40-90 female florets and

1 or 2 functionally-male disc florets (not

“hermaphroditic”, as stated by Hunger). The

other six specimens bear only homogamous

capitula in which all 13-18 disc florets are

functionally male, and female florets are

lacking. The absence, in this sample, of any

individual specimen with both homogamous

and heterogamous capitula suggests that the

species is subdioecious.

Key to Queensland species of Pluchea

1 Leaves and stems resinous or viscid.2

1. Leaves and stems neither resinous nor viscid.4

2 Leaves broadly obovate, toothed; hairs (0.1-0.25 mm long) frequent on

stems and leaves.P. dunlopii

2. Leaves linear, entire; hairs absent on stems and leaves.3

3 Margins of involucral bracts fimbriate, eglandular; disc florets with five

corolla lobes; achenes 1.4-1.7 mm long.P. baccharoides

3. Margins of involucral bracts with stalked glandular hairs; disc florets

with four corolla lobes; achenes 0.8-1 mm long.P. punctata

4 Leaves and stems with surfaces obscured by a very dense indumentum

P. ferdinandi-muelleri

4. Leaves and stems glabrous or hairy but surfaces readily visible

5 Leaves and stems with eglandular septate hairs.6

5. Leaves glabrous or with stalked glandular hairs, but eglandular septate hairs absent.... 7

6 Leaves and stems with many yellow sessile glands.P. xanthina

6. Leaves and stems without sessile glands.P. indica

7 Leaf-bases not decurrent; stems and leaves with many stalked glandular

hairs; inner involucral bracts 6-8 mm long.P. dentex

7. Leaf-bases decurrent on stems; stems and leaves glabrous; inner involucral

bracts 3.5-5 mm long.P. rubelliflora

346

Austrobaileya 8(3): 340-346 (2011)

Acknowledgements

I am grateful to Keith McDonald for taking

me to the population of Plucheapunctata that

he discovered. My thanks are also due to Will

Smith for the illustrations and Peter Bostock

for the Latin diagnoses.

References

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- (2010). Asteraceae. In P.D. Bostock & A.E.

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Hnatiuk, R.J. (1990). Census of Australian Vascular

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Willdenowia 26: 273-82.

Map 1. Distribution of Pluchea xanthina • and

P. punctata A

- (1997). A survey of the genus Pluchea

(Compositae, Plucheeae) in Australia.

Willdenowia 27: 207-23.

Iucn (2001). Red List Categories and Criteria: Version

3.1. IUCN Species Survival Commission.

IUCN: Gland, Switzerland/Cambridge, U.K.

King-Jones, S. (2001). Revision of Pluchea Cass.

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Englera 23: 3-136.

Mueller, F. (1889). Second Systematic Census of

Australian Plants. McCarron, Bird & Co.:

Melbourne.

Thulin, M. (2004). Notes on Pluchea (Asteraceae) in

Somalia and Ethiopia. Nordic Journal of Botany

22: 659-666.

Eucalyptus melanophloia subsp. nana D.Nicolle & Kleinig, a new

mallee ironbark (E. series Siderophloiae Blakely; Myrtaceae)

from central Australia and north western Queensland

D.Nicolle 1 * & D.A.Kleinig 2

Summary

Nicolle, D. & Kleinig, D. (2011). Eucalyptus melanophloia subsp. nana D.Nicolle & Kleinig, a new

mallee ironbark (E. series Siderophloiae Blakely; Myrtaceae) from central Australia and north western

Queensland. Austrobaileya 8(3): 347-355. The new subspecies Eucalyptus melanophloia subsp.

nana is described. The subspecies is of widespread but very scattered distribution in arid regions on

rocky sites from the Yaripilangu Range in the Northern Territory in the west (about 250 km WNW of

Alice Springs), eastwards to the Mount Isa area in north western Queensland. It is distinguished from

subsp. melanophloia by its consistently multi-stemmed, bushy mallee habit and this is unique among

the ironbark eucalypts ( Eucalyptus series Siderophloiae and E. series Rhodoxylori). Illustrations of

the new subspecies (including the holotype), a key to the subspecies of Eucalyptus melanophloia and

a distribution map are provided.

Key Words: Myrtaceae, Eucalyptus, Eucalyptus melanophloia , Eucalyptus melanophloia subsp.

nana , Australia flora. Northern Territory flora, Queensland flora, taxonomy, new subspecies,

ironbark, mallee

‘D.Nicolle, Currency Creek Arboretum, RO. Box 808, Melrose Park, South Australia 5039, Australia.

Email: dn@dn.com.au *author for correspondence

2 D. A.Kleinig, P.O. Box 395, Clare, South Australia 5453, Australia. Email: dendros@bigpond.com

Introduction

Eucalyptus melanophloia F.Muell. belongs

to E. series Siderophloiae Blakely (which

includes all the ironbark taxa in which

all the stamens are fertile - about 22

species) subseries Jugatae Blakely (only

E. melanophloia and E. shirleyi Maiden).

The subseries is distinguished within the

genus by the combination of the following

characteristics (amended from Brooker

2000 ):

Eucalyptus subgenus Symphyomyrtus

(Schauer) Brooker-Cotyledons folded in

seeds; buds bi-operculate; seeds with ventral

or terminal hilum; seed coat formed from

both integuments.

Eucalyptus section Adnataria L.D.Pryor

& L.A.S.Johnson ex Brooker-Cotyledons

reniform; seeds with a ventral hilum; leaf

venation usually densely reticulate; anthers

not versatile.

Accepted for publication 29 June 2011

Eucalyptus subsection Apicales Brooker -

Anthers erect at tip of filament; inflorescences

commonly terminal.

Eucalyptus series Siderophloiae —

Ironbarks; outer operculum shed early;

all stamens fertile, irregularly flexed.

Eucalyptus subseries Jugatae- Juvenile

leaves sessile; mature crown of juvenile

leaves.

Eucalyptus subseries Jugatae (with the

two species E. melanophloia and E. shirleyi )

was recognised by Brooker (2000) on the

basis of the sessile juvenile leaves forming

the mature crown, which distinguish the

subseries from E. subseries Subglaucae

Blakely (the remaining 20 species of E. series

Siderophloiae ), which have petiolate juvenile

leaves and a crown of adult leaves. However,

the distinction between these subseries is

weak and perhaps not representative of the

phylogeny of the group; indeed Eucalyptus

whitei Maiden & Blakely of E. subser.

Subglaucae forms substantial intergrading

populations with E. melanophloia of

E. subseries Jugatae (Holman et al. 2011).

348

Nevertheless, within Eucalyptus subseries

Jugatae , E. melanophloia is distinguished

from E. shirleyi in its less straggly habit,

smaller juvenile leaves, the occasional

development of intermediate or adult leaves

in some individuals (E. shirleyi is not known

to develop intermediate or adult leaves) and

the much smaller flower buds and fruits.

Interestingly, Eucalyptus shirleyi is quite

variable and may be composed of more than

one taxon. The distribution of Eucalyptus

subseries Jugatae can be roughly defined by

the distribution of E. melanophloia , with the

distribution of E. shirleyi (inland of Mackay

north to the Atherton Tableland region)

occurring largely within the northern part of

the distribution of E. melanophloia.

The new taxon described here was first

collected south of Mt Isa in Queensland as early

as 1951 and has subsequently been collected

numerous times south of Mt Isa on the Mt

Isa to Dajarra road, as well as occasionally

from elsewhere on the hills around Mt Isa.

The population on the Mt Isa to Dajarra road

has become well known because it is atypical

compared to Eucalyptus melanophloia

elsewhere in Queensland. Brooker & Kleinig

(1994: 347) mentioned the population in the

text .. occurring as a small mallee south of

Mt Isa”. Brooker & Kleinig (1994) included

an image of both the tree variant (subsp.

melanophloia) and the mallee variant (subsp.

nana) of E. melanophloia. Field and herbarium

studies in conjunction with cultivation trials

in a common garden indicate that the Mt Isa

populations are sufficiently distinct to warrant

taxonomic recognition.

The new subspecies was not collected in

central Australia (Northern Territory) until the

1990s, when Peter Latz collected specimens

from two widely separated localities

(Ennugan Mountains and Yaripilangu

Range). A number of other central Australian

collections have since been made in localities

in between, including at the type locality at

Mt Leichhardt.

Taxonomic recognition of the two variants

of Eucalyptus melanophloia is considered

appropriate due to the conspicuous distinction

in habit between the two geographically

Austrobaileya 8(3): 347-355 (2011)

disjunct variants. This is consistent with

the taxonomic recognition of other tree

versus mallee variants (genetically and not

environmentally determined) within the

eucalypts, such as Eucalyptus loxophleba

Benth. subsp. supralaevis L.A.S.Johnson

& K.D.Hill (tree) and subsp. lissophloia

L.A.S.Johnson & K.D.Hill (mallee).

The new subspecies ( Eucalyptus

melanophloia subsp. nana) has great

horticultural potential in arid regions, because

of its depauperate habit (for a eucalypt), crown

of silvery leaves, and habitat of rocky sites

(i.e. confined soil pockets) in a hot and dry

region.

The relatively recent discovery of this

new subspecies in Central Australia must be

associated with the remoteness and relative

paucity of botanical collections in the area

in which it occurs. The subspecies is not

rare (it can locally dominate some vegetation

communities) and it is conspicuous in the

field.

Material and methods

This paper is based on examination of

herbarium collection at the herbaria AD,

BRI, CANB and NSW, together with visits

to populations in the Northern Territory and

Queensland.

Taxonomy

Eucalyptus melanophloia F.Muell., J. Proc.

Linn. Soc., Bot. 3: 93 (1859). Type citation:

A tractu montano Newcastle Range usque ad

sinum Morton Bay Eucalypto crebra abunde

adsociata, solum sterilibus indicans’. Type:

Queensland: Dawson River, s.dat. [1856],

F.Mueller s.n. (lecto: K, material on upper

half of sheet, fide Bean [2010: 472]).

Bark rough throughout (to branches less

than 25 mm in diameter), moderately to

deeply fissured and hard (ironbark), dark

grey to black. Branchlets terete, waxy on

current season’s growth (wax eroded and

not apparent on old branchlets), lacking pith

glands. Seedling leaves opposite, sessile or

on petiole to 2 mm long, elliptical to ovate,

concolorous to slightly discolorous, dull,

greyish, usually waxy, 35-60 mm long,

Nicolle & Kleinig, Eucalyptus melanophloia

16-35 mm wide. Juvenile leaves opposite,

sessile, (cordate at base) broadly elliptical

to ovate (rounded to slightly pointed at tip),

concolorous, dull, greyish, waxy on current

season’s growth, persisting on reproductively

mature plants. Inflorescences held erect,

terminal or sometimes sub-terminal (growing

tip continuing). Peduncles terete or nearly so.

Pedicels terete. Flower buds with operculum

scar (but outer operculum sometimes still

present at tip at anthesis), diamond-shaped

to broadly fusiform, broadest below or at

hypanthium-operculum join, 6-9 x 2.8-3.2

mm, waxy; operculum bluntly conical to

349

conical, 3-5 mm long. Flower stamens cream,

all anthers fertile. Fruits waxy when young

(wax eroded and not apparent on old fruits).

Seeds compressed ovoid, 1-2 mm long, dark

grey-brown to black, finely reticulate; chaff

smaller, linear, orange brown.

Two geographically disjunct subspecies

are recognised in Eucalyptus melanophloia ,

differing most conspicuously and consistently

in habit but also generally differing (with

some degree of overlap) in leaf spacing and

size, and in fruit size.

Key to subspecies of Eucalyptus melanophloia

Tree, usually single-stemmed, 7 to 18 m tall; most internodes 20 to 50

mm long (central and eastern Queensland and northern New South Wales)

.subsp. melanophloia

Mallee, multi-stemmed, 2 to 6 m tall; most internodes 10 to 30 mm long

(Northern Territory and north western Queensland).subsp. nana

Eucalyptus melanophloia subsp.

melanophloia

Tree, usually single-stemmed at ground level,

7-18 metres tall; forming an inconspicuous

lignotuber. Juvenile leaves well-spaced

(internodes usually 20-50 mm). Juvenile

leaves 35-70 mm long, 22-48 mm wide.

Intermediate and adult leaves sometimes

produced in crown, 75-110 mm long, 13-30

mm wide. Inflorescences 7 or 9-flowered.;

peduncles 5-14 mm long; pedicels 1-5 mm

long. Fruits cupular to truncate-globose to

pyriform, 5-6 mm long, 5-6.5 mm wide, disc

descending then level to slightly ascending,

valves 3, 4 or 5, enclosed to slightly exserted.

Distribution and habitat : Widespread

and relatively common in north-eastern

Australia, from Narromine in New South

Wales northwards to west of Musgrave on

central Cape York Peninsula in Queensland

(Clarkson 2009), and inland in Queensland

to near Adavale and Croydon. A common

component of open woodland vegetation,

where it occurs on level to undulating

topography as well as on hills and ridges, in

skeletal or sandy soils (occasionally including

deep red sands) to well-drained loams.

Numerous woodland eucalypt species have

been recorded as associates, and although

the subspecies sometimes forms pure stands,

it is more commonly co-dominant with one

of a number of Eucalyptus section Adnataria

(ironbark or box) species, especially E. crebra

F.Muell.

Notes: Putative sporadic hybrids are known

with a number of other species, including

intra-serial hybrids with Eucalyptus crebra

(e.g. Forster 16089, 16229, 13104, 16281; see

Brophy et al. 2009), E. cullenii Cambage,

E. decorticans (F.M.Bailey) Maiden and

E. drepanophylla F.Muell. ex Benth. (in the

broad sense when including E. xanthoclada

Brooker & A.R.Bean) and inter-serial

hybrids with E. coolabah Blakely & Jacobs

and E. leptophleba F. Muell. (both E. series

Aquilonares Brooker) and E. populnea

F. Muell. and E. orgadophila Maiden & Blakely

(both E. series Buxeales Blakely). Extensive

intergrading populations between Eucalyptus

melanophloia subsp. melanophloia and

E.whitei are known where their distributions

adjoin in the Pentland to Jericho area of

Queensland (Holman et al. 2011).

350

Eucalyptus melanophloia subsp. nana

D.Nicolle & Kleinig subspecies nova a

subspecie typica habitu minore et pluricauli

“mallee”, foliis juvenilibus minoribus et

internodis brevioribus differt. Typus:

Northern Territory. Western slope of Mt

Leichhardt, 1 July 2008, D.Nicolle 5205 &

M.E.French (holo: AD; iso: CANB, DNA,

PERTH).

Eucalyptus melanophloia subsp. (Dajarra V. J.

Neldner 1523); Bean et al. (2010: 119).

Mallee, several-stemmed from ground

level, 2-6 metres tall; forming a lignotuber.

Juvenile leaves crowded (internodes mostly

10-30 mm). Juvenile leaves 18-55 mm long,

12-40 mm wide. Reproductively mature in

juvenile leaf phase, intermediate and adult

leaves rarely produced. Intermediate leaves

rarely produced. Adult leaves not known.

Inflorescences 7-11-flowered. Peduncles 5-

10 mm long. Pedicels 1-2 mm long. Fruits

cupular, truncate-globose, barrel-shaped

or slightly pyriform, 5-6 mm long, 4.5—5

mm wide, disc descending, valves 3 or 4,

enclosed. Fig. 1-3.

Additional selected specimens examined : Northern

Territory. 4 km NE of Mt Leichhardt, 30 km NNE of

‘Coniston’ Homestead, Sep 2006, Dugnid 22376 & Latz

(CANB, NT); Mt Leichhardt, 29 km NNE of ‘Coniston’

Homestead, Sep 2006, Duguid 22380 & Latz (CANB,

NT); Mt Leichhardt summit, Jul 2008, Nicolle 5207 &

French (AD); Mt Leichhardt, upper S slope, Aug 2005,

Albrecht 11677 & Latz (BRI, NT); Mt Leichhardt, lower

S-slope, Aug 2005, Albrecht 11660 & Latz (BRI, DNA,

NSW, NT); Mt Denison, 23 km NNW of ‘Coniston’

Homestead, Sep 2005, Latz 21028 (BRI, NT); SE end of

Yindjirbi Range, Coniston Station, Jul 2008, Nicolle 5203

& French (AD, CANB, DNA); Ti Tree Station, Ennugan

Mountains, Jul 1992, Latz 12366 (AD, DNA, NT); 13 km

ENE ‘Newhaven’ Homestead, Jul 2002, Harris 4 (NSW,

NT); 13 km ENE ‘Newhaven’ Homestead, Jul 2002, Latz

18773 & Harris (NT); Yaripilangu Range, 12 km ENE

‘Newhaven’ Homestead, Mar 1997, Latz 15097 (CANB,

DNA, NT). Queensland. Burke District: 53 km ENE

of Mt Isa, N of Breakfast Creek, Jul 1988, Harris 246

(BRI); W of Mt Isa on May Downs Station, Sep 2008,

Kleinig 6768 (AD); May Downs Station, W of Mt Isa,

Jul 2004, Booth 3563 & Kelman (BRI, CANB); 11 km

SSW of Mt Isa, Aug 1988, Harris 250 (BRI); S of Mt

Isa on the road to Dajarra, Apr 1995, Nicolle 1330 (AD);

16 km S of Mt Isa, Sybella Creek, Apr 1988, Harris 239

(BRI); South of Mt Isa on the road to Dajarra, Nov 2001,

Nicolle 4183 (AD, BRI, CANB); c. 20 km S of Mt Isa

on Dajarra Road, Sep 2008, Kleinig 6767 (AD); 20.9 km

S of Mt Isa on Dajarra road. Mar 1990, Brooker 10434

Austrobaileya 8(3): 347-355 (2011)

(BRI, CANB, MEL, NSW); 21 km S of Mt Isa on Dajarra

road. May 1994, Brooker 11888 (BRI, CANB, DNA,

NSW); 22.5 km S of Mt Isa, Aug 1973, Williams 138 &

Trapnell (BRI); 29 km S of Mt Isa on stony ridge, Sep

1951, Everist 4455 (BRI).

Distribution and habitat: The new

subspecies is of widespread but has a very

scattered distribution on arid rocky sites

from the Yaripilangu Range in the west

(about 250 km WNW of Alice Springs in

the Northern Territory), eastwards to the

Mount Isa area in north western Queensland

(Map 1). Although the Northern Territory

and Queensland populations are widely

separated geographically, it is likely that

further populations occur in the intervening

area, which is remote and botanically poorly

sampled.

In central Australia (Northern Territory),

the subspecies occurs as very scattered

populations from the Yaripilangu Range to

the Ennugan Mountains but can be locally

common or dominant, such as on the slopes

of Mt Leichhardt. It occurs on granite hills

and ranges, in open mallee vegetation.

Associated eucalypts in Central Australia

include Corymbia aparrerinja K.D.Hill

& L.A.S.Johnson, C. deserticola subsp.

mesogeotica K.D.Hill & L.A.S.Johnson,

C. eremaea (D.J.Carr& S.G.M.Carr) K.D.Hill

& L.A.S.Johnson, E. gillenii Ewart & L.Kerr,

E. normantonensis Maiden & Cambage and

E. pachyphylla F.Muell., with scattered shrubs

including Grevillea wickhamii Meisn. and

Hakea grammatophylla (F.Muell.) F.Muell.

and a groundstorey dominated by Triodia

R.Br.

In north western Queensland, the

subspecies is locally common in a small

area on the Mt Isa to Dajarra road, where

it occurs on rocky outcrops and the

intervening valleys in hilly, quite rugged

topography in gravelly red sand derived from

Sybella granite. The subspecies also occurs

as very scattered populations elsewhere on

the Mt Isa Mines Lease, where Barrs (1999)

describes the vegetation unit as ‘ Eucalyptus

melanophloia dwarf open forest’. Associated

eucalypts in the Mt Isa area include

Corymbia terminalis (F.Muell.) K.D.Hill &

Nicolle & Kleinig, Eucalyptus melanophloia

351

AD219429

D. Nicolle 5205 & M. French

Eucalyptus melanophloia subsp. nana

Western slope ol Ml Leichhardt. Northern Terr

Let 21* 46* 41* Long 132* 33' .

Malleo less than one metre tall to two metros tr

Leaves dull, grey. All parts waxy.

Dominant shrub on western slopes, ridges and

normantononsls and Grevtlba wickhamlla nd T

AD. CANB, PERTH. NT

Photos

Northern Territory. Central Australia North District.

Western slope of Ml Leichhardt.

Coord, ex GPS 2l fl 48'41*S. 132 & 33'5rE; preen I

Dominant shrub. On western slopes, ridges and fool of range in stone with

scattered Corymbut apurrerinja, Eucalyptus normantonensis and Creviliea

wlckhamii and Tnodia groundstorcy.

Malice less than one metre tall to two metres tall. Bark rough throughout, hard,

moderately fissured, dark grey to black. Leaves dull grey. All parts waxy.

D. Nicolle 5205 & M. French

Duplicates to:

Database^ in: ADHERE ghaiucn: gccc.OK)

l JuJ 2008

lllllllil

Fig. 1. Holotype of Eucalyptus melanophloia subsp. nana (Nicolle 5205 & French [AD])

352

Austrobaileya 8(3): 347-355 (2011)

Map 1 . Distribution of Eucalyptus melanophloia subsp. melanophloia (O) and subsp. nana (V).

L.A.S.Johnson, E. leucophloia subsp. euroa

L.A.S.Johnson & K.D.Hill, E. leucophylla

Domin and E. normantonensis. Scattered

Acacia shrubs are present as an under storey

(including A. hilliana Maiden and A. monticola

J.M.Black) and Triodia dominates the

groundstorey. It categorises a distinct regional

ecosystem (RE1.12.2) in Queensland that

has an ‘Of Concern’ status and an estimated

area of <10,000 ha (Accad et al. 2008; http://

www.derm.qld.gov.au/wildlife-ecosystems/

biodiversity/regional_ecosy stems/details.

php?reid=1.12.2.

Notes: Eucalyptus melanophloia subsp. nana

is distinguished from E. melanophloia subsp.

melanophloia by the consistently bushy,

multi-stemmed mallee habit and also in the

generally smaller and more crowded juvenile

leaves. The fruits also tend to be smaller and

the new subspecies more rarely produces

intermediate (and adult) leaves than the

typical subspecies.

This new subspecies is disjunct from the

nearest known locality of subsp. melanophloia

to the north-east (west of Croydon) by about

350 km. No intergrading populations are

known.

Cultivation trials of Mt Isa populations

of the new subspecies and three Queensland

provenances of subsp. melanophloia have

been undertaken since 1994 at Currency

Creek Arboretum in South Australia

(Nicolle 2000). The aim of these trials was

to determine if habit differences apparent

between the Mt Isa populations (subsp.

Nicolle & Kleinig, Eucalyptus melanophloia

353

Fig. 2. Eucalyptus melanophloia subsp. nana at Yindjirbi Range, Northern Territory (population voucher: Nicolle

5203 & French [AD, CANB, DNA]). Photo: D. Nicolle

nana) and E. melanophloia elsewhere in

Queensland (subsp. melanophloia) are

caused by environmental factors (subsp.

nana occurring in a more arid environment)

or due to genetic differentiation. These trials

indicate that subsp. nana retains both its low,

bushy, mallee habit and the generally smaller

adult leaves compared to subsp. melanophloia

(Queensland populations grown from the

Mt Moffatt, Jericho and Miles areas), which

consistently developed into taller, single¬

stemmed trees with a crown composed of

larger juvenile leaves.

Although there are no known intermediate

(‘intergrading’) populations between the two

subspecies, and although the new subspecies

is consistently distinct in habit, subspecific

rather than specific status has been applied to

the new taxon due to the allopatric distribution

of the two taxa and the overlapping distinctions

of leaf and fruit morphology. It is likely

that the two taxa would readily interbreed

if brought into geographical contact, and

the discovery of intermediate populations

remains a possibility.

Sporadic hybrids are known with

Eucalyptus normantonensis Maiden &

Cambage, of E. series Buxeales (see below).

This inter-series hybrid is conspicuous in the

field, having a crown of leaves intermediate in

morphology between the dull greyish, sessile,

broad leaves of Eucalyptus melanophloia

subsp. nana and the glossy green, petiolate,

narrow leaves of E. normantonensis (Fig. 4).

Conservation status : Although populations

are widely scattered, the subspecies is locally

common or dominant where it has been

recorded, and it is not considered to be at

risk. At Mt Leichhardt in Central Australia,

the subspecies is dominant or co-dominant on

the ridges, summit and western slopes, with

the population size likely to exceed 10,000

individuals (Fig. 3).

354

Austrobaileya 8(3): 347-355 (2011)

Fig. 3. Eucalyptus melanophloia subsp. nan a at Mt Leichhardt, Northern Territory (type locality). Photo: D. Nicolle

Etymology : From the Latin nanus (a dwarf)

referring to the consistently low bushy habit

and also in reference to the generally smaller

leaves of the new subspecies in comparison to

E. melanophloia subsp. melanophloia.

Eucalyptus melanophloia subsp. nana x E.

normantonensis hybrids

Specimens examined : Northern Territory. 4 km NE of

Mt Leichhardt, 30 km NNE of ‘Coniston’ Homestead,

Sep 2006, Duguid 22375 & Lat= (CANB, NT); SE end

of Yindjirbi Range, Coniston Station, Jul 2008, Nicolle

5204 & French (AD, CANB, DNA).

Acknowledgments

We wish to thank Ian Brooker for discussions

regarding the new taxon, staff at AD for

assisting with loans of E. melanophloia from

NT (Alice Springs), and staff at BRI and DNA

(Darwin) for allowing access to collections.

The senior author also would like to thank

Malcolm French for field assistance in Central

Australia, Max Lines of Coniston Station for

allowing and assisting with access through

Coniston Station to the Yindjirbi Range and

Mt Leichhardt, and Tony Bean for bringing

to our attention (in 2006) the occurrence of

E. melanophloia in the Northern Territory.

We also thank Paul Forster and an anonymous

referee for providing useful comments on the

manuscript.

Nicolle & Kleinig, Eucalyptus melanophloia

355

Fig. 4. Eucalyptus melanophloia subsp. nan a x E. normantonensis hybrid {Nicolle 5204 & French [AD, CANB,

DNA]). Photo: D. Nicolle

References

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114-126. Queensland Herbarium, Department

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to Eucalypts Vol. 3, Northern Australia.

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D.B. & Punruckvong, A. (2009). Essential oil

variation in Eucalyptus crebra, E. melanophloia

(Myrtaceae) and their hybrids. Australian

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the Cape York Peninsula Bioregion. Department

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Holman, J.E., Hughes, J.M. & Fensham, R.J. (2011).

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252.

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Research. Vol. 1. D.Nicolle: Adelaide.

Cycas terryana P.I.Forst. (Cycadaceae), a

new species from central Queensland

Paul I. Forster

Summary

Forster, P I. (2011). Cycas terryana P.I.Forst. (Cycadaceae), a new species from central Queensland.

Austrobaileya 8(3): 356-363. A new species of Cycas from the Broadsound and Connors Ranges

in central Queensland is described, illustrated and diagnosed as C. terryana P.I.Forst. It is known

from five populations in an area of occurrence of approximately 800 km 2 and does not occur in any

conservation reserves. A conservation status of Vulnerable based on the IUCN criterion of D2 is

recommended for the species.

Key Words: Cycadaceae, Cycas , Cycas terryana , Australia flora, Queensland flora, new species,

taxonomy, identification key, conservation status

P.I.Forster, Queensland Herbarium, Department of Environment & Resource Management, Brisbane

Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email: paul.forster@

derm.qld.gov.au

Introduction

The genus Cycas L. is speciose in Australia

with 28 species recognised (Hill 1998, 2004).

The species generally occur in well defined,

geographically discrete areas in few (in several

instances one) to many populations. Botanical

exploration in Queensland continues to

reveal previously undocumented populations

of Cycas ; such a case pertains to several

populations of plants in the area southwest of

St Lawrence and northwest of Marlborough in

the Broadsound and Connors Ranges.

Hill (1992) was of the opinion, albeit

without supporting evidence, that this

material represented a zone of hybridism

(“intergrading populations”) between Cycas

media R.Br. and C. ophiolitica K.D.Hill.

This hypothesis was developed further

(Hill 1996: 3), viz. “these [populations] are

morphologically intermediate between the

putative parent species and also show the

high degree of variability to be expected from

Mendelian segregation in second and later

generations”. Such introgressed populations

were considered to occur throughout the

range of the genus in Australia (Hill 1996)

and elsewhere (e.g. Hill 1994, 2008; Hill et al.

2004), in some instances between species that

he classified in different series. This ‘multiple

Accepted for publication 18 July 2011

hybridism’ hypothesis remains to be tested

using molecular markers (e.g. haplotype

networks [Schaal & Olsen 2000]) and remains

merely as a plausible supposition in some

cases. Alternative hypotheses are that these

populations are merely variations of more

broadly defined species or in some instances

represent discrete taxa.

Hill’s propensity for designating a large

number of Cycas populations as hybrids

or examples of introgression was probably

influenced by his work on eucalypts (where

this is widespread and by comparison well

documented with genetic studies) and his

mentoring by L.A.S.Johnson who together

with Lindsay Pryor was a keen supporter

of the ‘phantom’ hybrid population concept

(Pryor 1951; Pryor & Johnson 1971; Parsons &

Kirkpatrick 1972; Johnson 1976; Kirkpatrick

& Potts 1987). Eucalypts are bisexual,

often promiscuous outbreeders with non

specialist pollinators and appear to be readily

dispersed; hence, the potential for hybrids

where different taxa adjoin is high, although

recorded instances are relatively low (Griffin

etal. 1988). Populations of hybrid individuals

may lead to extinction of the parental species

via genetic pollution, although in many

cases the progeny are sterile or with reduced

viability (Levin 2000). Alternatively, these

populations may in time lead to separate

speciation events once fertility is restored

Forster, Cycas terry ana

(e.g. via allotetraploidy) and genetic change

via selection or genetic drift occurs (Levin

2000 ).

The situation in Cycas is quite dissimilar,

insomuch as the plants are dioecious, require

pollination by insects (in variations of a

mutualistic relationship) and perhaps wind in

combination (Kono & Tobe 2007), polyploidy

is unknown (Gorelick & Krystle 2011), and

distribution of the populations are limited

by dispersal (Forster 2007). As noted by

Hill (1992) it is perhaps unlikely that strong

genetic barriers to cross pollination between

different taxa exist and so the only real

barrier to hybridism/introgression is isolation

in geographic space. Populations of Cycas

and other Australian cycads are rare in the

landscape for the above reasons, quite unlike

the situation with most eucalypts. Johnson

(1959, 1961) actively promoted the concept of

widespread hybridism in cycads, especially for

Zamiaceae ( Macrozamia ) species, although

some of his examples are merely juveniles of

much larger adults (e.g. M. lucida x M. moorei ;

juveniles of what is now known as M. johnsonii

D.L. Jones & K.D.Hill) or represented discrete

taxa that he was not aware of. Speciation in

Cycas and Macrozamia is perhaps most

likely to occur via genetic drift in isolated

populations, rather than genetic selection per

se (Gorelick 2009). This is reflected in the

occurrence of similar appearing taxa, often in

a geographic replacement series of allopatric

populations. This can be interpreted as an

indication of both continuous and incomplete

speciation in a non-adaptive radiation where

the inter-relationship between environment

and morphology is unclear (Savolainen &

Forest 2005) and in cycads is probably driven

mainly by fragmentation, rather than radical

differences in habitat specialisation ( cf. Linder

2003). Either way, the tempo of speciation is

likely to be slow given the long time period

between germination and individuals reaching

maturity, together with slow turnover of

individuals within populations.

With regard to the so called Cycas media

- C. ophiolitica intermediates south of St

Lawrence, Hill (1992) was probably only aware

of two populations based on his collection

{Hill 3788 & Stanberg ) and that of Hind 2878 ,

357

or others that he examined and determined

as such. There are now five populations of

these plants known and perhaps more in what

is largely a botanically unexplored part of

central Queensland.

Neither of the putative parental species

have populations that are particularly

geographically close to the Broadsound and

Connors Ranges plants. Hill (1996) vaguely

stated that the hybrid populations were

“wherever different species grow in relatively

close proximity”. The closest population of

Cycas media is c. 9 km to the northeast (in

different habitat and geology) and the closest

population of C. ophiolitica some c. 45 km to

the east of the overall area of occupancy for the

five populations. The southern populations of

Cycas media are predominantly coastal and

occur on substrates derived from basalts, finer

grained granites or stabilised sand masses.

The northern populations of Cycas ophiolitica

are restricted to serpentinite substrates. By

comparison the Broadsound and Connors

Ranges plants occur well inland in the ranges

on substrates derived from heavily weathered

porphyritic granites.

These populations of Cycas may well

have originated from mixing of the ancestral

lineages of both C. media and C. ophiolitica

as postulated by Hill, with speciation

following the initial hybridism and subsequent

geographic isolation. However, the situation

could well be much more complex than this

and reflect multiple instances of fragmentation

and integration of populations driven by

climatic changes since the glacial cycles of

the Pleistocene. This hypothetical pattern of

change has been actively promoted as one

of the main drivers of speciation in Mexican

cycads (Vovides et al. 2003) and similar

reproductive and dispersal biology of the

Australian cycads may infer similar patterns

of allopatry (Forster 2004).

Given that the recorded populations of

these Cycas are disjunct over an area of

occurrence of at least 800 km 2 , it is likely

that the hypothesised speciation event is well

advanced with subsequent fragmentation

following an initial introgression. It is

unlikely that these populations are any less fit

358

than the core Cycas media or C. ophiolitica

populations and to all extents they appear

to exist and operate much like any other

healthy cycad populations. If of hybrid origin,

then they probably commenced as a neutral

hybrid zone with genetic compatibility and

superficially similar habitats, although such

zones are usually in close proximity to the

putative parental species (Levin 2000) which

is no longer the case here. Whatever the origin,

these populations possess character states that

enable them to be distinguished from both

Cycas media and C. ophiolitica ; hence, they

are described below as a new species. The

hypothesised species origin is also relevant

for speciation elsewhere in Cycas and other

genera of cycads, particularly Macrozamia

Miq. from Australia (Forster 2004),

Encephalartos Lehm. in Africa (Vorster et al.

2004) and Ceratozamia Brongn. and Zamia

L. in the New World (Vovides et al. 2003,

2004).

Materials and methods

The species description is based on

examination of both live plants in habitat

and herbarium collections at the Queensland

Herbarium.

Taxonomy

Cycas terryana P.I.Forst. species nova a

C. ophiolitica differt praecipue foliis valde

fragilibus (adversum non), multo parcius

carinatis et juvenilibus non glaucis, extra

colore indumenti; quum a C. media subsp.

media differentiae principals sunt folia valde

(non infirme) fragilia, maturitate indumentum

gerentia (adversum indumentum deficiens) et

valde carinata (adversum plus minusve non

vel infirme carinata), cum colore indumenti

in cataphyllis. Typus: Queensland. Port

Curtis District: Broadsound Range, 9 March

2005, P.I.Forster PIF30618 & P.J.Machin

(holo: BRI [4 sheets, $ carpological and spirit

samples]).

Arborescent cycad with stems to 3 m high

(rarely multiheaded), 12-22 cm thick. Leaves

80-190 cm long, straight and becoming wavy

towards the apex, weakly to strongly keeled

in cross-section, pale green to somewhat

glaucous green-grey above; opposing

Austrobaileya 8(3): 356-363(2011)

leaflets inserted at 20-50° to the rhachis, the

rhachis usually terminated by paired leaflets,

tomentose; petiole 17-30 cm long, 7-15 mm

diameter, dull olive grey-green, with 4-40

short teeth (pinnacanths) 1.5-2 mm long and

spaced 8-17 mm apart, sometimes spineless.

Leaflets 184-320 per leaf, 7-10 mm apart,

being evenly spaced in lower half of leaf,

then becoming more interleaved and more

strongly keeled in upper half of leaf, brittle,

margins recurved; median leaflets at 25-45°

to the rhachis, 105-230 mm long, 5-7 mm

wide, green-grey with fawn-tan bloom when

young; weakly convex in cross section,

decurrent for 5-7 mm, up to 13 mm at base

of leaf. New growth densely tomentose with

pale cream-fawn indumentum, glabrescent.

Cataphylls pungent, linear, 8-12 cm long,

densely tomentose for entire length with

pale fawn indumentum. Microsporangiate

cones elongate-ovoid, 21-28 cm long, 6.5-8

cm diameter, with dense ferruginous-brown

indumentum; microsporophylls 28-40 mm

long, fertile zone 17-28 mm long, 4-12

mm wide; apical spine antrorsely recurved,

4-10 mm long. Megasporophylls 20-33

cm long, when young with dense fawn-

tan indumentum, aging grey, eventually

glabrescent and olive-green; ovules 4 to 6;

lamina broadly triangular 50-70 mm long,

30-38 mm wide, strongly dentate with well

developed antrorse teeth 4-7 mm long, apical

spine 17—33 mm long. Seeds ovoid, 37-40

mm long, 30-35 mm diameter, sarcotesta c. 5

mm thick, weakly grey pruinose, olive green

beneath wax covering, becoming orange

when ripe. Figs. 1-5.

Additional specimens examined: Queensland.

Leichhardt District: Broadsound Range, Mar 1964,

Everist 7624 (BRI); ditto loc., Nov 2003, Forster

PIF29714 & Halford (BRI); ditto loc., Nov 2003,

Forster PIF29715 & Halford (BRI); ditto loc.. Mar

2005, Forster PIF30617 & Machin (BRI); ditto loc., Dec

2010, Mathieson MTM965 & 966 (BRI); ditto loc., Jun

2011, Forster PIF38217 & Machin (BRI). Port Curtis

District: St Lawrence towards Croydon, Aug 1990,

Hill 3788 & Stanberg (BRI, NSW); Croydon Station

- St Lawrence Road, Jun 2011, Forster PIF38218 &

Machin (BRI); 5 km W of Prospect Hill, Nov 1992,

Forster PIF12276 & Machin (BRI); near Burwood [as

‘Barwood’], Broadsound, Jun 1981, Maconochie 2757

(BRI; CANB, DNA, NY n.v.).

Forster, Cycas terryana 359

Fig. 1. Cycas terryana. Plants in habitat (population voucher: Forster PIF29714 & Halford [BRI]). Photo: P I.Forster

Distribution and habitat : Cycas terryana

is restricted to the Broadsound and Connors

Ranges northwest of Marlborough and

southwest of St Lawrence. Populations

occur on undulating low hills on stony soil

derived from porphyritic granite in woodland

dominated in various combinations by Corymbia

clarksoniana (D.J.Carr & S.G.M.Carr) K.D.Hill

& L.A.S.Johnson, C. dallachiana (Benth.)

K.D.Hill & L.A.S.Johnson, Eucalyptus crebra

F.Muell., E. drepanophylla F.Muell. ex Benth.

and Petalostigma pubescens Domin.

Notes: Cycas terryana appears to have

first been collected by Selwyn Everist in

1964; however, this herbarium specimen

has been variously identified as C. media

or an intergrade between C. media and

C. ophiolitica (determination of K.D.Hill).

The primary morphological differences

between Cycas terryana , C. media and

C. ophiolitica are summarised in Table 1 . The

leaflets and rhachis of C. terryana are very

brittle and easily snapped, as opposed to the

situation in C. media and C. ophiolitica where

they are less brittle in the former and robustly

flexible in the latter. The megasporophylls in

Cycas terryana are generally broader than in

the other species being compared here and

indumentum on the new growth, cataphylls

and megasporophylls is pale grey-fawn to

fawn-tan whereas in the others it is usually

orange-brown, only becoming greyish if

particularly weather beaten. Cycas terryana

differs from C. ophiolitica mainly in the

leaves being markedly brittle (versus not),

much less strongly keeled and not glaucous

when young, as well as the indumentum

colour; whereas from C. media subsp. media

the main differences are the leaves being

markedly brittle (versus weakly brittle), with

indumentum at maturity (versus none) and

strong keeling (versus ± flat to weakly keeled),

together with the indumentum colour on the

cataphylls.

360

Austrobaileya 8(3): 356-363(2011)

Fig. 2. Cycas terryana. Apex of mature plant showing cataphylls (population voucher: Forster PIF30617 & Machin

[BRI]). Photo: P.I.Forster

Cycas media and C. ophiolitica were

classified in different series by Hill (1996,

1998) on the basis of anatomy and morphology.

Given that species such as C. terryana exist,

there seems little point in the continued

recognition of such artificial supraspecific

taxa.

Etymology : This species is named for

Dr L. Irene Terry of the University of Utah,

U.S.A., in recognition of her ground breaking

work on cycad pollination, both in Australia

and on Guam (e.g. Terry 2001; Terry et al.

2004, 2005, 2007, 2009).

Conservation status : The species is

known from five populations with an area

of occurrence of c. 800 km and an area of

occupancy that is much less. An appropriate

status is Vulnerable based on the D2 criterion

(IUCN 2001). None of the populations occur

in conservation reserves and at least one

has been reduced to non viable numbers of

plants.

Acknowledgements

Thanks to Mike Mathieson for collecting

material and photographing the male

cones; David Halford and Peter Machin for

assistance with fieldwork and Peter Bostock

for translation of the diagnosis into Latin.

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Forster, Cycas terryana 361

Table 1. Comparison of character states for Cycas media, C. ophiolitica and C. terryana

Character State

C. media

C. ophiolitica

C. terryana

Mature leaf

indumentum

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and on rhachis

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Leaflet texture

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Fig. 5. Cycas terry ana. Male plant with microsporangiate

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Photo: M.T.Mathieson.

A taxonomic revision of Trichosanthes L.

(Cucurbitaceae) in Australia, including one

new species from the Northern Territory

W.E. Cooper 1 & H. J. de Boer 2

Summary

Cooper, W.E. & de Boer, H.J. (2011). A taxonomic revision of Trichosanthes L. (Cucurbitaceae) in

Australia, including one new species from the Northern Territory. Austrobaileya 8(3): 364-386.

Trichosanthes is represented by six species in Australia: T. cucumerinaL. var. cucumerina, T. morrisii

W.E.Cooper sp. nov., T, odontosperma W.E.Cooper & A.J.Ford, T. pentaphylla F.Muell. ex Benth.,

T. pilosa Lour, and T. subvelutina F.Muell. ex Cogn. Trichosanthes ovigera Blume has recently been

synonymised with T. pilosa and we now include T. holtzei F.Muell. within this synonymy. All taxa are

illustrated (with the exception of T odontosperma previously illustrated in 2010), and distinguished

from other Australian species. Notes on habitat and distribution are included together with distribution

maps. Three identification keys are presented, two to the sections of Trichosanthes and one to the

species of Trichosanthes in Australia.

Key Words: Cucurbitaceae, Trichosanthes , Trichosanthes cucumerina , Trichosanthes ovigera ,

Trichosanthes pilosa , Trichosanthes odontosperma , Trichosanthes subvelutina , Trichosanthes

pentaphylla , Trichosanthes morrisii , Australia flora. Northern Territory flora, Queensland flora,

taxonomy, identification keys, new species, new combination, probracts

1 W.E. Cooper, Australian Tropical Herbarium, James Cook University, Cairns Campus, McGregor

Road, Smithfield, Queensland, 4878, Australia.

2 H.J. deBoer, Department of Systematic Biology, Uppsala University, Norbyvagen 18 D, 752 36

Uppsala, Sweden. Email: hugo.deboer@ebc.uu.se

Introduction

Trichosanthes L. is a genus of approximately

100 species distributed from India, Sri

Lanka, China, Japan, SE Asia to Malesia

(including New Guinea), Australia, and

the eastern Pacific (Rugayah & de Wilde

1999). Trichosanthes is broadly represented

in Australia with one or two species from

five of the six sections: Trichosanthes L.,

Cucumeroides (Gaertn.) Kitam., Edulis

Rugayah, Aster osperma WJ.de Wilde &

Duyfjes (not in Australia), Foliobracteola

C.Y.Cheng & C.H.Yueh and Involucraria

(Ser.) Wight. Sections are distinguished using

(1) seed shape and structure, (2) colour of fruit

pulp, (3) presence or absence of probracts,

(4) size of male bracts. However, leaf shape

is variable within all species, and shape of

male bracts can vary greatly in Trichosanthes

cucumerina L., T. pentaphylla F.Muell. ex

Benth. and T. subvelutina F.Muell. ex Cogn.

The last comprehensive treatment of

Trichosanthes in Australia (Telford 1982)

Accepted for publication 14 October 2011

described six species in which T. holtzei

F.Muell. was thought to be closely allied

to T. ovigera Blume; however, T. holtzei is

no longer maintained here with the present

authors synonymising it with the widespread

and variable T. pilosa Lour. Examination of

a greater number of herbarium specimens

collected in Australia during recent years, as

well as molecular analysis (de Boer & Cooper

unpublished), have confirmed this finding.

Trichosanthes ovigera has been previously

referred to this synonymy through priority of

T. pilosa (de Wilde & Duyfjes 2008).

Telford(1982) suggestedthat Trichosanthes

pentaphylla (Fig. 1,6) is probably conspecific

with T. trifolia (L.) Merr. and the latter has

now been synonymized with T. wawrae

Cogn. (Rugayah & de Wilde 1997). However,

the two species differ sufficiently to maintain

Trichosanthes pentaphylla as distinct from

T. wawrae.

Taxa in the genus Trichosanthes can

be extremely variable throughout their

distribution with regard to leaf (Fig. 3,4),

Fig. 1 . Trichosanthes pentaphylla from the Mowbray River, showing silvery leaves with dark green veins, typical

juvenile growth of plants in the section Involucraria. Photograph: W.T. Cooper

Fig. 2. Trichosanthes subvelutina. male flowers, Terania Creek NSW. Photograph: H. Nicholson

366

bract and fruit morphology, as well as seed

surface in T. pilosa. The extremes of habitat

preferences within one species can often be

associated with morphological variation.

Trichosanthes sp. Fine Leaf (L.A. Craven

7930) had been recognised as a different taxon

by the Northern Territory Herbarium (DNA);

however, the present authors see it as part of

the widespread and variable T. cucumerina L.

var. cucumerina.

In this revision, six species of

Trichosanthes are recognised for Australia

and of those, T. odontosperma W.E.Cooper

& A.J.Ford, T. pentaphylla , T. subvelutina

and T. morrisii W.E.Cooper are endemic.

Five species are mostly tropical with one

species ( Trichosanthes subvelutina) mainly

subtropical.

Materials and methods

We examined 363 specimens from BR, BRI,

CANB, CNS, DNA, K, L, MEL, P, PERTH

and NSW. All species were observed in the

field by the first author.

Flowers of Trichosanthes are fragile and

difficult to preserve as good quality dried

specimens. They are therefore not easy to

analyze (Duyfjes & Pruespan 2004), and

have not been used as key species characters.

Even with fresh flowers, plants are mostly not

identifiable to species; however, plants having

flowers with fimbriate petal margins belong

to sections Trichosanthes or Cucumeroide s,

and those having petals with fimbriate

apices belong to sections Foliobracteola ,

Involucraria or Edulis. Leaf, probract and seed

features provide good characters (Duyfjes &

Pruespan 2004) and along with male bract

size and sepals, have been predominantly

used here as key features for dichotomous

determination keys.

Measurements of all parts were done on

dried material and specimens preserved in

70% ethanol, as well as on fresh material in

the field. Common abbreviations in the text

include N.P (National Park) and S.F./S.F.R.

(State Forest/State Forest Reserve).

The extensive bibliographic taxonomic

history of Trichosanthes sections and the

species that also occur outside of Australia has

Austrobaileya 8(3): 364—386 (2011)

been repeated several times in recent years

(e.g. Rugayah & de Wilde 1997,1999; Duyfjes

& Pruespan 2004; de Wilde & Duyfjes 2010)

and is largely omitted from this account.

Taxonomy

Trichosanthes L., Sp. PI. 2: 1008 (1753).

Type species: Trichosanthes anguina L.,fide

M.L.Green, Prop. Brit. Bot. 190 (1929) [= T.

cucumerina subsp. anguina (L.) Greb.]

Cucumeroides Gaertn., Fruct. Sem. PI. 2: 485,

t. 4 (1791). Type species: not designated.

Involucraria Ser., Mem. Soc. Phys.

Genev. 3(1): 25, t. 5. (1825). Type species:

Involucraria wallichiana Ser.

Trichosanthes in Australia: Monoecious or

mostly dioecious, trailing or climbing vines or

lianas, annual or mostly perennial, partially

or completely seasonally senescent; bark

corky, warty, flaky, fissured or lenticellate;

stems either weak and slender or robust and

somewhat woody; glabrous, pubescent or

scabrous; roots tuberous. Indumentum of

translucent multicellular trichomes, glandular

or eglandular; cystoliths often present, discoid

or rosette-shaped, drying to white or black

in herbarium specimens. Probracts present

at the nodes in many species, caducous or

persistent, may be pubescent, glands often

present. Tendrils unbranched or 2-, 3- or 4-

branched(upto 9-branched in Asia), positioned

at the node (not truly axillary). Leaves simple

or compound (digitate), alternate, petiolate,

unlobed or 3-7 (rarely 10)-lobed, margin

entire or toothed with a soft mucro or callus

terminating main veins, membranous or

leathery, scabrous, velvety or glabrous, glands

often present on the underside, pedately

veined with 3-5 main veins, intralateral veins

reticulate. Inflorescence beside petiole at the

node (not truly axillary). Flowers unisexual,

mostly nocturnal excepting T. cucumerina,

epigynous, petals 5 (rarely 4); corolla white,

cream or yellowish, actinomorphic; sepals

5 (rarely 4), free, subulate or triangular;

petals hairy, margin fimbriate; receptacle

tube elongate. Male flowers either solitary,

paired, or in racemose or rarely paniculate

inflorescences; bracts present on racemes

at pedicel base and sometimes at peduncle

367

Cooper & de Boer, Trichosanthes in Australia

base, persistent or caducous, glands may be

present; stamens 3, inserted near receptacle

tube orifice, included, free until anthesis

then becoming fused; filaments free; anthers

oblong, s-shaped, two are 2-thecous, one is

1-thecous, basifixed. Female flowers mostly

solitary or paired (one is usually much older),

sometimes in a racemose inflorescence in T.

pilosa , solitary and usually co-axillary with

male raceme in T. cucumerina ; perianth

similar to males; style long, narrow; stigma

3-5-lobed; staminodes absent; ovary inferior,

pubescent or glabrous, elliptic, ovoid, globose

or fusiform; ovules numerous, horizontal.

Fruit apepo or berry, ellipsoid, ovoid, globose

or fusiform, beaked at apex, 30-140 mm

long, mostly solitary; epicarp green with or

without longitudinal white stripes, orange or

red, glabrous or pubescent; epicarp leathery,

chartaceous or woody; mesocarp fleshy or

firm, orange, yellow or whitish, smooth or

fibrous; pulp white, yellow, orange, red or

dark green; seeds few-numerous, horizontal,

1-locular or 3-locular with the two lateral

locules empty, compressed or turgid, smooth

or sculptured, brown, cream, grey or black.

Germination is hypogeal, semi-hypogeal or

epigeal.

Etymology : The generic epithet Trichosanthes

is derived from the Greek tricho- (hairy) and

-anthus (flowered); referring to the petals,

which have fimbriate margins.

Key to sections of Trichosanthes in Australia using mostly seed and fruit features

1 Seeds 3-locular.Section Cucumeroides

1. Seeds 1-locular.2

2 Seeds surrounded by dark green pulp, adult leaves compound or simple;

juvenile leaves silvery with dark green veins.Section Involucraria

2. Seeds surrounded by white, greenish-white, yellow, orange or red pulp, all

leaves simple; juvenile leaves green only.3

3 Seeds surrounded by white or greenish-white pulp.Section Foliobracteola

3. Seeds surrounded by yellow, orange or red pulp.4

4 Seeds toothed; probract present on new growth; plants dioecious.Section Edulis

4. Seeds with an undulate margin; probract absent; plants monoecious

.Section Trichosanthes

Key to sections of Trichosanthes in Australia using leaf, probract and male bract

1 Leaves compound.Section Involucraria

1. Leaves simple.2

2 Probracts present (at least on new growth).3

2. Probracts absent.5

3 Probracts caducous.Section Edulis

3. Probracts persistent.4

4 Leaf upperside with numerous cystoliths.Section Involucraria

4. Leaf upperside without cystoliths.Section Foliobracteola

5 Plants monecious; male inflorescence bracts up to 3 mm long.... Section Trichosanthes

5. Plants dioecious; male inflorescence bracts 4-8 mm long.Section Cucumeroides

368

Austrobaileya 8(3): 364—386 (2011)

Key to species of Trichosanthes in Australia

1 Leaves compound (Qld only).T. pentaphylla

1. Leaves simple.2

2 Leaf upperside glabrous or glabrescent.3

2. Leaf upperside hairy.4

3 Probract linear or narrowly ovate, caducous; seeds toothed, surrounded

in orange or red pulp (Qld only).T. odontosperma

3. Probract obovate, elliptical or oblong, persistent; seeds entire, surrounded

in dark green pulp, (NT only).T. morrisii

4 Plants monoecious; male inflorescence bracts up to 3 mm long; seeds

with an undulate margin (NT, Qld, WA).T. cucumerina var. cucumerina

4. Plants dioecious; male inflorescence bracts 4-33 mm long; seed margin

not undulate.5

5 Male inflorescence bracts 4-8 x 0.5-3 mm; seeds tumid, t-shaped,

surrounded by yellow-orange pulp (NT, Qld, WA).T. pilosa

5. Male inflorescence bracts 15-33 x 7-23 mm; seeds flat, round and entire,

surrounded by greenish-white pulp (NSW, Qld).T. subvelutina

Trichosanthes L. section Trichosanthes

Type species: Trichosanthes anguina L. [= T.

cucumerina var. anguina (L.) Haines]

Monoecious trailing vines. Probracts absent.

Male bracts persistent (caducous in Asia).

Flowers diurnal, petals fimbriate with thread¬

like extensions along the margins. Fruit yellow

or orange; seeds 1-locular, flat, margins broad

and undulate.

Distribution : One species (two varieties)

occurring in China, India, Sri Lanka, SE

Asia, Malesia and Australia (one variety).

1. Trichosanthes cucumerina L., Sp.

PI. 2: 1008 (1753). Type: habitat in India,

“Padavalam ” in Rheede, Hort. Malab. 8: 39,

t. 15 (1688) (lecto: fide Keraudren-Aymonin

[1975: 91]).

The species comprises two varieties

as accepted here with T. cucumerina var.

cucumerina occurring in Australia. The

second variety ( T. cucumerina var. anguina

(L.) Haines) is the common snake gourd of

horticulture. Data from molecular sequencing

support the placement of the cultivated snake

gourd as a cultivated variant of T. cucumerina

(Ali & Al-Hermaid 2010).

Trichosanthes cucumerina var. cucumerina;

Haines, Bot. Bihar Orissa 3: 388 (1922).

Trichosanthes pedatifolia Miq., FI. Ind. Bat.

1: 677 (1856). Type: Indonesia: Java, s.dat.,

T.Horsfield s.n. (holo: BM28187; iso: U).

Trichosanthes reniformis Miq., FI. Ind. Bat.

1: 675 (1856). Type: Indonesia: Java, s.dat.,

T.Horsfield s.n. (holo: BM28189, 28188).

Trichosanthes ambrozii Domin, Biblioth.

Bot. 89: 631 (1929). Type: Australia: Western

Australia, between Ashburton and De Gray

Rivers, s.dat., E.Clement s.n. (syn: B, K, PR

n.v.).

Trichosanthes brevibracteata Kundu, J. Bot.

77: 10 (1939). Type: India. Punjab, Karnal, in

1885-1888, J.R.Drummond 25031 (syn: K);

India. Ahmedabad, July 1920, L.J.Sedgwick

s.n. (syn: K).

Trichosanthes pachyrrhachis Kundu, J. Bot.

77: 9 (1939). Type: Northwest India, in 1844,

M.P.Edgeworth 63 (holo: K).

Trichosanthes sp. Nitmiluk. (C.R.Mitchell

3293); Australian Plant Census (CHAH 2005);

Australian Plant Name Index, http://www.

anbg.gov.au/cgi-bin/apni. Accessed February

2010 .

Cooper & de Boer, Trichosanthes in Australia

369

Fig. 3. Trichosanthes cucumerina var. cucumerina. A. male flower. B. seeds. C. unlobed leaf underside showing

glands, male inflorescence with bracts, fruit and tendril. D. male inflorescence, buds and broadly ovate and orbicular

cupular bracts. E. leaf underside with deeply lobed margin and solitary gland. A-C from Cooper 2119 (CNS); D-E

from Fox 2513 (DNA) (scales as indicated). Del. W.T. Cooper.

Trichosanthes sp. Fine Leaf (L.A.Craven

7930), Australian Plant Census (CHAH 2005);

Kerrigan & Albrecht (2007); Australian Plant

Name Index, http://www.anbg.gov.au/cgi-bin/

apni . Accessed February 2010.

Illustrations : Yueh & Cheng (1974: pi.82);

Telford (1982: 197); Wheeler (1992: 254);

Rugayah & de Wilde (1999: 231); Hyland et

al. (2003); Duyfjes & Pruesapan (2004: 81).

Monoecious slender trailing vine to 2 m.

Stem diameter 1-2 mm, 5-ribbed, pubescent

(trichomes may be sparse); trichomes

short glandular and eglandular and mostly

interspersed with scattered long eglandular

trichomes, denser at nodes or sometimes only

at nodes; cystoliths rarely present. Probracts

absent. Tendrils 2 or 3-branched, sparse

glandular or eglandular trichomes towards

base. Leaves simple, discolorous, membranous;

petioles 8-57 mm long, sparse to dense short

glandular and eglandular trichomes, mostly

with scattered longer trichomes, rarely with a

few discoid cystoliths; lamina cordate, ovate,

370

broad-ovate or reniform, 23-145 mm long, 25-

135 mm wide; unlobed, or shallowly to deeply

3- 7-lobed, rarely 10-lobed, lobe divisions

up to c. 95% of lamina length; central lobes

oblong, elliptical, obovate or spathulate; lateral

lobes ovate, triangular, oblong, asymmetrical,

spathulate; base cordate and shortly cuneate

or truncate, sinus often shallow and wide;

apex acute, acuminate or obtuse, with a

soft mucro; margin undulate, denticulate,

dentate or serrate, 4-40 teeth or soft mucros

along each side; membranous; upperside

pubescent, hirsute or scabrous, trichomes

short eglandular and/or glandular, sometimes

restricted to main veins, and mostly with

scattered long eglandular trichomes often

restricted to near the margin, rarely with a

few scattered discoid cystoliths; underside

pubescent, hirsute or scabrous, trichomes

short glandular and/or eglandular, often

with longer scattered eglandular trichomes

usually on veins or close to margin; glands at

leaf base 1-8 per side, some scattered glands

may be present, gland diameter 0.2-0.3 mm;

rarely with a few scattered discoid cystoliths.

Male inflorescences racemose or solitary

(rarely a panicle), usually arising well after

co-axillary female; raceme 70-240 mm long,

4- 21-flowered, rachis with sparse to dense,

short glandular and eglandular trichomes as

well as long eglandular trichomes, peduncle

25-180 mm long; bracts persistent at base of

pedicels and often at base of peduncle, ovate,

broadly ovate, orbicular, triangular, narrowly

triangular, reniform, obovate, oblong,

elliptical, rhombic or linear, often cupular,

0.25-3 mm long, 0.25-2.5 mm wide, base

truncate or cuneate, apex obtuse, acute or

rarely obcordate, entire, crenate, unlobed or

with 3-5 shallow lobes, mostly orange-brown

or pinkish-brown in herbarium specimens

(linear bracts remain green), glabrous or

trichomes sparse to dense and usually much

sparser than those on the adjoining rachis,

glands absent; venation absent, obscure or

3-veined with a midrib and intramarginal

veins. Male flowers: pedicels 5-45 mm long;

receptacle tube slender-funnelform, 7-35 mm

long; trichomes glandular and eglandular,

sparse to moderately dense; sepals triangular,

entire, 0.5-2.5 mm long, 0.25-1 mm wide,

green, trichomes sparse to dense; corolla

Austrobaileya 8(3): 364-386 (2011)

diameter 8-20 mm (not including fimbrillae);

petals elliptic-ovate, 9-10 mm long (not

including numerous fimbrillae), c. 4 mm wide,

white, pubescent abaxially and adaxially;

anthers c. 2 mm long; filaments c. 0.8 mm

long. Female flowers: solitary or paired, co-

axillary with male racemes (usually emerging

before male inflorescence is fully expanded);

pedicels 3-50 mm long, trichomes glandular;

receptacle tube, slender-funnelform, 12-50

mm long, trichomes glandular and eglandular,

pubescent; sepals triangular or subulate,

entire, 1-3 mm long, 0.5-1.5 mm wide, green,

pubescent abaxially and adaxially; corolla

diameter 9-25 mm (not including fimbrillae),

petals ovate, 5-10 mm long, 2-3 mm wide,

white, trichomes glandular and sparse;

ovary 4-25 mm long, glabrous or trichomes

sparse. Fruit elongated-globular, fusiform

or globose with a beaked apex, 21-50 mm

long, 18-25 mm wide, orange or red (unripe

fruit is longitudinally streaked dark and pale

green), glabrous or glabrescent; pulp meagre,

translucent orange or reddish; mesocarp 2-2.5

mm wide; pedicel 3-9 mm long, trichomes

sparse, bract from base of male raceme usually

persistent at base; exocarp c. 0.3 mm wide;

seeds 3-12, elliptical or obovoid, 1-locular,

margin undulate with 3-5 shallow lobes on

each side, 6-10 mm long, 4-7 wide, 3-5 mm

thick, testa brown or grey. Germination type

unknown. Fig. 3.

Additional selected specimens (from 166 examined ):

Australia: Western Australia. Edge of small bay W

of Cape Ruthiers, Mar 1993, Mitchell 2989 (DNA);

c. 2 km E of Mitchell Falls, Mar 1994, Mitchell 3368

(CANB); Lacrosse Island at head of Cambridge Gulf,

NE Kimberley, Jun 1992, Kenneally 11327 (CANB);

The Dag, Mornington Wildlife Sanctuary, Apr 2005,

Legge 521 et al. (BRI); West Kimberley, Oscar Range,

Brooking Gorge, May 1988, Sands 4732 (CANB);

Dampier Archipelago, Enderby Island, Jul 1980,

Wilson 7268 (CANB); c. 135 km S of Port Hedland on

Great Northern Highway, Apr 1995, Mitchell PRP280

(CANB). Northern Territory, c. 30 km NNE of Jabiru,

Mar 1981, Craven 7930 & Whitebread (CANB); Kakadu

N.P., Mt Brockman, Mar 1995, Egan 4575 (DNA); 6 km

S Mt Gilruth, Arnhem Land, March 1984, Wightman

1345 (DNA); Deaf Adder Gorge, Feb 1977, Fox 2513

(DNA); Nitmiluk gully of Nitmiluk Gorge, Apr 2004,

Crase 1311 & Dixon (DNA); Katherine Gorge, Apr 1972,

Must 973 (DNA); 6 km NE Cape Crawford Hotel, Abner

Range, Jan 1989, Brock 484 (DNA). Queensland. Cook

District: Lankelly Creek, Mcllwraith Range, May 1995,

Hyland 15339 (CNS); Mt White, Coen, Apr 1993, Hyland

14776 (CNS); Jane Table Hill, Lakefield N.P., Mar 1993,

371

Cooper & de Boer, Trichosanthes in Australia

Fell 2903 & Stanton (BRI); Metal Hills, 4 km NNE of

Chillagoe, May 2006, Wannan 4372 (NSW); Royal Arch

N.P., Chillagoe, Feb 1996, Forster PIF18614 & Ryan

(BRI); Royal Arch Tower, Chillagoe N.P., Apr 2010,

Cooper 2119 & FordJ CNS).

Distribution and habitat : Trichosanthes

cucumerina var. cucumerina is widely

distributed from India, Southern China and

Southeast Asia through Malesia to Australia.

Within Australia it is widespread in Western

Australia from the northern Kimberley south

to the Port Hedland area, also in the Northern

Territory from Arnhem Land to the Abner

Range near McArthur, and in Queensland

from Cape York to Chillagoe (Map 1). In

Queensland it is more restricted with a few

isolated collections from Cape York Peninsula

and Chillagoe-Mungana Caves N.P, and

one collection from Black Mountain near

Cooktown within the Wet Tropics bioregion.

It has a strong association with sandstone

habitats but it also occurs in areas of granite

and limestone. It is a climber on rocks, shrubs

and trees in open forest, woodland, shrubland

(often on sandstone escarpments), monsoon

forest, vine thickets, Allosyncarpia forest,

herblands and grasslands.

Phenology : Flowers have been recorded in

January, February, March, April, May, June,

July and October; fruit has been recorded in

January, February, March, April, May, June,

August and October.

Notes : Trichosanthes cucumerina var.

cucumerina is a highly variable species with

leaves varying from unlobed to deeply lobed.

However, all plants have male bracts up to 3 mm

long, flowers that are open during the day, and

seeds with undulate margins. It differs from

all other Australian Trichosanthes by having

flowers that open during daylight, and very

small male bracts. Male inflorescence bracts

are usually persistent through to ripe fruit in

Australian populations, whereas in Malesia

and China the bracts are absent, subpersistent

or caducous (Backer & Bakhuizen van den

Brink 1963; Rugayah & de Wilde 1997; de

Wilde & Duyfjes 2004, 2010; Duyfjes &

Pruesapan 2004; Lu et al. 2011). Leaf glands

are usually present in Australian populations

whereas in Malesia leaf glands are absent or

few (de Wilde & Duyfjes 2010). The leaves

have a pungent smell like those of Momordica

charantia L. mixed with peanut butter.

Etymology : After its similarity to the related

genus Cucumis.

Trichosanthes section Cucumeroides

(Gaertn.) Kitam., J. Jap. Bot. 19: 35 (1943);

Cucumeroides Gaertn., Fruct. Sem. PI. 2: 485,

t. 4, f. 4. (1791). Type species: T. cucumeroides

(Ser.) Maxim. [= T. pilosa Lour.]

Dioecious (sometimes monoecious) vines;

probract absent; male bracts persistent

(caducous or persistent in Thailand, Duyfjes &

Pruesapan 2004), relatively short and mostly

narrow; flowers nocturnal, petals fimbriate

with thread-like margins; fruit pulp yellow

or orange (also whitish or pinkish in Asia,

de Wilde & Duyfjes 2004); seeds 3-locular,

t-shaped, quadrangular and tumid.

Distribution : Approximately 14 species in

China, India, SE Asia, Malesia and Australia

(one species).

2. Trichosanthes pilosa Lour., FI. Cochinch.

1: 588 (1790). Type: Vietnam (holo: lost);

Vietnam. Tu Phap, s.dat.. Bon 4019 (neo: P,

fide de Wilde & Duyfjes [2008: 270]).

Trichosanthes ovigera Blume, Bijdr. FI. Ned.

Ind. 15: 934 (1826). Type: Indonesia: Java. Mt

Salak, s.dat., C. Blume s.n. (holo: LI30442;

iso: LI 30439).

Trichosanthes chinensis Ser. in DC., Prodr.

3: 308 (1828). Type: tab. 13 in Cattley, Ic. PI.

China (1821).

Bryonia cucumeroides Ser. in DC., Prodr. 3:

308 (1828); Trichosanthes cucumeroides (Ser.)

Maxim, in Franch. & Sav., Enum. PI. Jap.

1: 172 (1873). Type: Japan, Cucumeroides

Thunb. ex. Gaertn., Fruct. 2: 485 (1791).

Trichosanthes horsfieldii Miq., FI. Ned. Ind. 1:

677 (1856). Type: Indonesia: Java. Priangan,

s.dat., T.Horsfield s.n. (holo: BM; iso: K, U).

Trichosanthes hearnii F.Muell. ex

Benth., FI. Austral. 3: 315 (1867). Type:

Australia: Queensland. Rockingham Bay,

s.dat., J.Dallachy s.n. (syn: K; isosyn:

MEL100085).

Trichosanthes himalensis C.B.Clarke in

Hook.f., FI. Brit. India 2: 608. (1879). Type:

372

Austrobaileya 8 ( 3 ): 364—386 ( 2011 )

Fig. 4. Trichosanthespilosa. A. seed: lateral and adaxial view. B. seed: adaxial view. C. male inflorescence with bracts

and buds. D. lobed leaf underside showing glands near base. E. unlobed leaf upperside, male inflorescence and tendril.

A & E from Gray 7477 (CNS); B from Cooper 2120 (CNS):

2064 (CNS) (scales as indicated). Del. W.T. Cooper.

India: Sikkim. Yoksun to the plains, ann. coll,

ign., J.D. Hooker s.n. (syn: K); India. Sikkim,

s.dat., Clarke s.n. (syn: K).

Trichosanthes dicoelospermum C.B.Clarke in

Hook.f.,/ 7 /. Brit.Ind. 2:609(1879); T. cucumeroides

C from McDonald 8839 & Dennis (CNS); D from Cooper

var. dicoelosperma (C.B.Clarke) S.K.Chen, Bull.

Bot. Res. 5: 118 (1985). Type: India: Sikkim.

Khasia, s.dat., J.D.Hooker s.n. (syn: K); loc. cit.,

s.dat., J.D.Hooker & T. Thomson s.n. (syn: K,

UPS).

373

Cooper & de Boer, Trichosanthes in Australia

Trichosanthes holtzei F.Muell. Aust. J. Pharm.

1: 447 (1886). Type: Australia: Northern

Territory, near Port Darwin, s.dat., M.Holtze

432 (holo: MEL100084), syn. nov.

Trichosanthes cavaleriei Lev., FI. Kouy-

Tcheou 123 (1914). Type: China: Guizhou

Province: near Guiyang, Bodinier 2306,

Cavalerie 1032,1816 (syn: all E).

Trichosanthes vanoverberghii Merr., Philipp.

J. Sci., C 9: 458 (1915). Type: Philippines.

Luzon Province: Bontoc subprovince, Bauco,

s.dat., M.Vanoverbergh 3662 bis (iso: K).

Trichosanthes baviensis Gagnep., Bull.

Mus. Hist. Nat. Paris 24: 379 (1918). Type:

Vietnam. Tu Phap, s.dat.. Bon [as Balansa ]

4016 (holo: P).

Trichosanthes pierrei Gagnep., Bull. Mus.

Hist. Nat. Paris 28:380 (1918). Type: Vietnam.

Baria, Mt Lu Dinh, s.dat., Pierre 4491 (holo:

P).

Trichosanthes hainanensis Hayata, Icon.

PI. Formosan. 10: 8 (1921); T. cucumeroides

var. hainanensis (Hayata) S.K.Chen, Bull.

Bot. Res. 5: 117 (1985). Type: China. Hainan

Province: s.loc., s.dat., Katsumada s.n. (holo:

TI).

Trichosanthes formosana Hayata, Icon. PI.

Formosan. 10: 7 (1921). Type: Taiwan. Taipei,

Urai, s.dat., Matsuda 267 (holo: TI).

Trichosanthes matsudai Hayata, Icon. PI.

Formosan. 10: 10 (1921). Type: Taiwan.

Qishan, Kosenpo, s.dat., Matsuda s.n. (holo:

TI).

Trichosanthes boninensis Nakai ex Tuyama,

Bot. Mag. Tokyo 50: 133 (1930). Type: Japan.

Bonin Island, Titizima, s.dat., Nakai s.n.

(holo: TI).

Trichosanthes chingiana Hand.-Mazz.,

Sinensia 7: 621 (1936). Type: China: Guangxi

Province: Yema Shan, s.dat., R.C.Ching 7113

(holo: W).

Trichosanthes rostrata Kitam.,Acta Phytotax.

Geobot. 5: 210 (1936). Type: Japan. Ryukyu

Islands. Miyagi Island, s.dat., Koidzumi s.n.

(holo: KYO n.v., iso: US36699).

Trichosanthes cucumeroides var. stenocarpa

Honda, Bot. Mag. Tokyo 54: 223 (1941). Type:

Japan. Musasi, Urawa, s.dat., K.Emori s.n.

(holo: TI).

Trichosanthes okamotoi Kitam., J. Jap. Bot.

19: 40 (1943). Type: Taiwan. Gaoxiong,

‘Benchi, Chushinron’, s.dat., Okamoto s.n.

(holo: KYO).

Trichosanthes mafuluensis Merr. & L. M. Perry,

J. Arnold Arbor. 30: 58 (1949). Type: Papua

New Guinea: Central Province. Mafulu,

1933, L.J.Brass 5257 (holo: A; iso: BO, BRI).

Trichosanthes ascendens C.Y.Cheng &

C.H.Yueh, Act. Phytotax. Sin. 18: 340 (1980).

Type: China: Guangxi Province: He Xian,

s.dat., C.T. Li 604120 (holo: IBX).

Trichosanthes trichocarpa C.Y.Wu ex

C.Y.Cheng & C.H.Yueh, Acta Phytotax.

Sin. 18: 340 (1980). Type: China. Yunnan

Province: Baoshan Xian, s.dat., T.T. Yu 17861

(holo: KUN; iso: A).

Illustrations: Keraudren-Aymonin (1975: 87)

as T. ovigera, Telford (1982:197) as T. ovigera,

Brennan (1986: 26) as T. cucumerina. ; Jones &

Gray (1988: 354) as T. holtzei, Wheeler (1992:

254) as T. ovigera, Rugayah & De Wilde

(1999: 231) as T. ovigera, Hyland et al. (2003)

as T. ovigera, Duyfjes & Pruesapan (2004: 81)

as T. ovigera. Cooper & Cooper (2004: 144)

as T. holtzei.

Dioecious or monoecious trailing vine to 6

m, perennial, partly or completely seasonally

senescent. Stem diameter 1-3 mm, 5-

ribbed, glabrous or glabrescent, trichomes

glandular and/or eglandular, scattered longer

eglandular trichomes may be present; nodes

usually slightly swollen, discoid cystoliths

may be present at nodes. Probracts absent.

Tendrils unbranched or 2- or 3-branched.

Leaves simple, discolorous, membranous

or chartaceous; petioles 17-85 mm long,

eglandular trichomes present, interspersed

with sparse glandular trichomes, rarely with

a few longer trichomes, discoid cystoliths

may be present; lamina cordate, unlobed or

shallowly to deeply 3-lobed, 70-200 mm long,

50-155 mm wide, base cordate, usually shortly

cuneate, sinus deep and narrow (rarely wide),

374

apex acute, acuminate, obtuse or retuse, with

a soft mucro, margin remotely denticulate,

undulate or crenate (mostly almost entire

except for a small mucro terminating each

vein), 16-28 mucros on each side; upperside

pubescent, hirsute or scabrous, sparsely to

densely clothed in short eglandular trichomes,

rarely with some glandular trichomes near

base, sometimes interspersed with longer

trichomes which are often only present near

margin; underside pubescent, hirsute or

scabrous, sparsely to densely covered in short

eglandular or glandular trichomes (usually

denser on underside), often with a few longer

trichomes on main veins especially towards

margin; 1-5 glands at leaf base on each side,

rarely with scattered glands, gland diameter

0.2-0.9 mm; discoid cystoliths may be present

on main veins. Male inflorescence racemose

or sometimes co-axillary with a solitary

flower; raceme up to 200 mm long and up to 11-

flowered, rachis densely covered in glandular

trichomes, peduncle 15-75 mm long; bracts

persistent, lanceolate, ovate, rhomboid, linear,

elliptical or obovate, 4-8 mm long, 0.5-3

mm wide, base truncate or cuneate, apex

acute or rarely obtuse, margin entire or 3-7-

toothed, glabrous or glabrescent, trichomes

glandular and eglandular, glands absent,

discoid cystoliths may be present, venation

pinnate and reticulate. Male flower: pedicels

5-35 mm long; receptacle tube salverform or

slender-funnelform, 23-24 mm long, clothed

in glandular and eglandular trichomes; sepals

triangular, subulate or lanceolate, entire or 3-

toothed, 4-7 mm long, 1-2 mm wide, green,

clothed in glandular and eglandular trichomes;

corolla diameter 12-20 mm; petals ovate, 6-8

mm long (not including fimbrillae), white,

cream, yellow or greenish-yellow, pubescent.

Female flowers in racemose inflorescence or

solitary (rarely intermixed on a male raceme)

on a pedicel 10-30 mm long; racemes 30-

70 mm long, 5-10 flowered; bracts present

on racemes and similar to those on male

inflorescences, trichomes glandular and

eglandular. Female flower: pedicels 12-16

mm long for solitary flowers, c. 5 mm long for

flowers in racemose inflorescence, trichomes

glandular and eglandular; receptacle tube

salverform or slender-funnelform, 18-50 mm

Austrobaileya 8(3): 364-386 (2011)

long (including ovary), trichomes glandular

and eglandular; sepals triangular or subulate,

often recurved, 2-9 mm long, 0.5-3.5

mm wide, green, trichomes glandular and

eglandular; corolla diameter (not including

fimbrillae) 15-17 mm; petals ovate, 8-18 mm

long, 4-9 mm wide (not including fimbrillae),

white or cream, clothed in glandular and

eglandular trichomes; fimbrillae white,

yellow or greenish-yellow; ovary 10-16 mm

long, trichomes glandular and eglandular.

Fruit obovoid or oblong-globose, apex acute

or obtuse with a short beak, 10-ribbed (ribs

sometimes subtle), 31-62 mm long, 31-35

mm wide, orange or red, glabrous or a few

glandular trichomes near base; mesocarp 2-

2.5 mm wide, orange; pedicel 7-10 mm long;

seeds 10-21, t-shaped, subquadrangular,

tumid, 3-locular, central cell warty to smooth

and containing the embryo, lateral cells cone-

shaped and empty, 5-10 mm long, testa brown

or blackish, suspended in yellow-orange pulp.

Germination is hypogeal or semi-hypogeal,

cotyledons straight. Fig. 4.

Additional selected specimens {from 85 examined ):

Western Australia. Mitchell Plateau, N Kimberley, Feb

1979, Beard 8436 (PERTH). Northern Territory. Gurig

N.P, Apr 2006, Brennan 6919 (DNA); Murgenella,

Wunyu Beach, Mar 1987, Russell-Smith 1967 & Lucas

(DNA); 5 km E mouth of Peter John River, NE Arnhem

Land, Feb 1988, Russell-Smith 4768 & Lucas (DNA);

Rocky Bay, Yirrkala, Mar 1988, Russell-Smith 5168 &

Lucas (BRI); 1 km SE of Angurugu, Groote Eylandt,

Mar 1988, Russell-Smith 5135 & Lucas (CANB); 5 km

SW Cutta Cutta Caves, Feb 1989, Russell-Smith 7157 &

Lucas (DNA); Van Dieman Gulf at base of Cape Hotham

Peninsula, Mar 1993, Cowie 3291 (DNA); Gunn Point,

Apr 1984, Dunlop & Wightman 6678 (DNA); Wangi

Falls, Litchfield N.P., Dec 2009, Cooper 2100 & Morris

(CNS). Queensland. Cook District: Old Mapoon Road,

Jun 2000, Hyland 16381 (CNS); Iron Range N.P., Mar

1994, Fell 4118 & Stanton (BRI); Lake Patricia, Weipa,

Mar 1989, O’Reilly 262 (BRI); Turrell Hill, Macrossan

Range, Silver Plains, Jun 1998, Forster PIF23085

et al. (BRI); Brooklyn, May 2010, Cooper 2120 &

Russell (CNS); Barron Gorge, Aug 2001, Cooper 1569

& Cooper (CNS); Eight Mile Mt, Mar 1999, Gray 7477

(CNS); Burke Developmental Road, Royal Arch Tower,

Chillagoe N.P, Feb 2010, Cooper 2102 & Ford (CNS);

Barron Gorge N.P, Apr 2009, Cooper 2064 (CNS);

Kennedy Highway, 300 m N of Herberton turnoff. May

2004, Cooper 1840 & Cooper (BRI); N.P.R. 16, Forty

Mile Scrub, old road. Mar 1999, Ford 2193 (CNS); Wind

Tunnel Lava Tubes, Undara N.P, Mar 2010, McDonald

8839 & Dennis (CNS). South Kennedy District: Carlisle

Island, Sep 1986, Batianoff5038 (BRI).

375

Cooper & de Boer, Trichosanthes in Australia

Distribution and habitat : Trichosanthes

pilosa is widely distributed from India through

Southern China to Japan, Southeast Asia

through Malesia to Australia and the eastern

Pacific. Within Australia it is widespread in

the Northern Territory and in Queensland it

occurs from Cape York south to Carlisle Island

(northeast of Mackay) and west to Chillagoe

(Map 2). There is one specimen from the

Mitchell Plateau in the northern Kimberly area

of Western Australia. It occurs in a variety of

habitats from wet tropical rainforest, monsoon

forest and littoral rainforest to deciduous vine

thickets at an altitudinal range from near sea

level to 1000 m.

Phenology: Flowers have been recorded in

February, March, April and June; fruit has

been recorded in February, March, April,

May, June and August.

Notes : Trichosanthes holtzei F.Muell. was

described as being reminiscent of T. anguina

(synonym of T. cucumerina var. anguina)

by Mueller (1886). His description is of a

specimen with staminate and pistillate flowers.

However, pistillate flowers are not evident on

the type specimen today, and even though

pistillate flowers may have been present,

this feature is not sufficient to distinguish

it, as Trichosanthes pilosa can at times be

monoecious. The inflorescence bracts and

leaves on the type specimen are a good match

for Trichosanthes pilosa , and T. holtzei is here

synonymised with T. pilosa.

A second variety of Trichosanthes pilosa

was described by de Wilde & Duyfjes (2008)

restricted to Thailand. Molecular analyses to

date do not support the recognition of varieties

in this species (de Boer, unpublished). Only

the nominative variety occurs in Australia.

Etymology : The specific epithet pilosa is

derived from the Greek pilos (anything

made of felt), presumably referring to the

indumentum.

Trichosanthes section Edulis Rugayah,

Reinwardtia 11: 232 (1999). Type species:

T. edulis Rugayah

Dioecious trailing vines or lianas; probract

present, caducous; flowers nocturnal, petal

margin fimbriate in upper half; fruit pulp

red; seeds numerous, 1-locular, flat or

quadrangular, toothed or notched.

Distribution : Nine species (New Guinea eight

endemic, Australia one endemic).

3. Trichosanthes odontosperma W.E. Cooper

& A.J.Ford, Austrobaileya 8: 126 (2010).

Type: Australia: Queensland. Cook District:

Topaz, Westcott Road, 15 April 2009,

W.Cooper 2065 (holo: CNS; iso: BRI, CANB,

DNA, L, MO, NSW, UPS).

Illustrations: Cooper & Cooper (1994: 291),

as Trichosanthes sp. (Mt Lewis); Cooper &

Cooper (2004: 145), as Trichosanthes sp. (Mt

Lewis); Hyland et al. (2003), as Trichosanthes

sp. (Mt Lewis BG 167); Jones & Gray (1988:

354 & back cover), as Trichosanthes sp.;

Williams (1987: 307), as Trichosanthes sp.;

Cooper & Ford (2010: 128-129).

Dioecious trailing vine or liana to mid¬

canopy, perennial, partly or completely

seasonally senescent. Stem diameter to 3 cm,

young branchlets 5-angular, glabrescent with

minute trichomes clustered at nodes; bark

fissured and corky on older growth; nodes

often markedly swollen. Probracts caducous,

linear or narrowly ovate, minutely lobed or

with a few teeth, 3-13 mm long, 1-3 mm wide,

glandular, glabrous or glabrescent. Tendrils

unbranched or 2- or 3-branched. Leaves

simple, discolorous, coriaceous; petioles

20-95 mm long; lamina ovate, cordate or

triangular, unlobed or rarely 3-lobed, 50-190

mm long, 41-160 mm wide, base cordate or

rarely hastate, sinus mostly narrow and deep,

apex acuminate to acute, with or without a soft

mucro; margin denticulate with 13-27 teeth

per side; upperside smooth, glabrous or with

sparse minute translucent trichomes on main

veins; numerous small translucent and sunken

multicellular (rosette-shaped) cystoliths,

which in most dried specimens become black;

underside with sparse translucent minute

trichomes on main veins, 1-16 circular and

flat glands on each side of the leaf base. Male

inflorescences with flowers mostly solitary,

rarely in a fascicle of two flowers or a raceme

beside a solitary flower, peduncle 35-110 mm

long; bracts at the base of racemose flowers

narrowly ovate, glabrous, 5-17 mm long,

1-3 mm wide. Male flowers: 45-90 mm

376

diameter, pedicel 43-83 mm long; receptacle

tube salverform, 45-90 mm long, green or

creamy-green with a narrow bright yellow

centre, glabrous or glabrescent abaxially;

sepals usually 5 (rarely 4), triangular, entire

or with 1-3 teeth, 8-20 mm long, 2-4 mm

wide, green, glabrous or glabrescent; petals

5 (rarely 4), obdeltoid, length including

fimbrillae 27-40 mm, 24-40 mm wide, white,

both surfaces villous becoming glabrescent

towards apex. Female inflorescences a

solitary flower on pedicels 31-50 mm long,

bracts absent. Female flowers: 60-73 mm

long, 55-65 mm diameter; receptacle tube

salverform, 39-55 mm long, green or creamy-

green with a narrow bright yellow centre,

adaxial surface of tube yellow, glabrous or

glabrescent abaxially; sepals 5, triangular,

entire, glabrous or glabrescent, 5-12 mm

long, 1-2 mm wide, green; petals 5, obdeltoid,

28-31 mm long, white, both surfaces villous;

ovary 14-24 mm long, glabrous and 10-

ribbed externally. Fruit ovoid or ellipsoid,

apex beaked, 90-140 mm long, 60-90 mm

diameter, glabrous, longitudinally 10-ribbed,

orange to red; mesocarp firm, 10-13 mm

thick, yellow-orange; pedicel 25-40 mm long

(or more, based upon flowering specimens), 4-

10 mm wide; seeds numerous, quadrangular,

1-celled, 2-horned at one end, 4-lobed at the

other end, both sides with 2 rows of 2-10

teeth, 12-18 mm long, 6.3-77 mm wide, 3-4

mm thick, testa brown to blackish, suspended

in orange or red pulp. Germination epigeal.

Distribution and habitat : Trichosanthes

odontosperma is endemic to the rainforests

of the Wet Tropics bioregion in north-east

Queensland. It occurs from the Windsor

Tableland area, west of Cape Tribulation, to

the Cardwell Range, west of Tully (detailed

notes were given previously in Cooper & Ford

[ 2010 ]).

Phenology : Flowers have been recorded in all

months; fruits have been recorded in January,

March, April, June, July, August, October and

November.

Etymology : The specific epithet is derived

from the Greek, odonto (tooth) and -sperma

(seed) and refers to the toothed seeds,

distinguishing it from all other Australian

Trichosanthes.

Austrobaileya 8(3): 364-386 (2011)

Trichosanthes section Foliobracteola

C.H.Yueh & C.Y.Cheng, Act. Phytotax. Sin.

12: 427 (1974). Type species: T. kirilowii

Maxim.

Dioecious vines or lianas; probracts present;

male bracts large; flowers nocturnal, petal

margin fimbriate in upper half; fruit pulp

whitish, cream or greenish-white; seeds 1-

locular, flat, margined.

Distribution : 12 species in China, India, SE

Asia, Malesia and Australia (one species).

4. Trichosanthes subvelutina F.Muell. ex

Cogn., Monogr. Phan. [A.DC. & C.DC.] 3:

366 (1881). Type: Australia: Queensland.

Moreton District: Moreton Bay, Archers

Brush, [November 1843], L.Leichhardt 4

(lecto [here designated]: P731577, photo!);

isolecto: P731576, photo!; MEL100124,

photo!; MEL100125, photo!).

Illustrations : Telford (1982: 197); Harden

(1990: 448); Nicholson & Nicholson (2004:

65); Harden et al. (2007: cover, 95, 96).

Dioecious trailing vines or lianas to 8

m, sometimes extending into the canopy,

perennial, seasonally senescent. Stemdiameter

to 15 mm, ribbed with vertical lenticels, bark

shallowly corky on older stems; young stems

glabrous and 5-angular, diameter c. 3 mm,

clothed in rusty trichomes, discoid cystoliths

may be present, nodes swollen. Probracts

persistent, ovate, quadrate or triangular,

entire or apex 3-toothed, 1-5 mm long, 1-4

mm wide, sparsely to densely pubescent, often

glandular. Tendrils 2-7-branched. Leaves

simple, discolorous, membranous; petioles

22-110 mm long; trichomes translucent

white, fawn or pale rusty eglandular; discoid

cystoliths may be present; lamina cordate or

broadly ovate, 3-5-lobed (rarely 7-lobed),

lobes shallow to deep, 85-260 mm long, 60-

220 mm wide; base cordate, shortly cuneate

or truncate, sinus deep and narrow or wide

and shallow; apex acute, acuminate or obtuse,

usually with a soft mucro; margin denticulate,

undulate or almost entire except for a short soft

mucro terminating each vein, 22-37 mucros

per side; upperside pubescent (sometimes

scabrous) with translucent white or fawn

eglandular trichomes, may be interspersed

Cooper & de Boer, Trichosanthes in Australia

377

Fig. 5.. Trichosanthes subvelutina. A. leaf, probract and tendril. B. rhomboid bract. C. broad-elliptic bract. D. seeds:

left, lateral view; right, adaxial view. A & D from Nicholson 3110 (BRI); B from Boorman s.n. (NSW146961); C from

Telford 9723 (CANB) (scales as indicated). Del. W.T. Cooper.

with longer trichomes; underside pubescent

with translucent white or fawn glandular and

eglandular trichomes, glands at leaf base up

to 4 per side or absent, gland diameter 0.4-0.5

mm; cystoliths absent. Male inflorescences

racemose, paired, solitary or a solitary flower

co-axillary with a raceme, racemes c. 250 mm

long with up to 12 flowers; peduncle 75-165

mm long, pubescent; bracts persistent, broad-

elliptic, rhomboid or obovate, 15-33 mm

long, 7-23 mm wide, apex acute or obtuse,

base truncate or cuneate, densely pubescent,

some trichomes glandular, margin denticulate

or almost entire except for soft mucros,

palmately veined, reticulate, glands sparse

or absent, gland diameter 0.3-0.5 mm. Male

flowers: pedicels 35-155 mm long; receptacle

tube funnelform or slightly urceolate, 31-

45 mm long, green with fawn pubescence

abaxially; sepals triangular, entire, 8-11

mm long, 2-3 mm wide, green, pubescent;

corolla diameter 30-70 mm; petals obovate

or obdeltoid, 18-21 mm long (not including

fimbrillae), white, pubescent on both sides.

Female inflorescences with flowers solitary

or paired (one usually much older) on pedicels

18-50 mm long, pubescent. Female flowers:

receptacle tube tubular, 22-51 mm long,

pubescent, trichomes fawn or rusty; sepals

triangular, entire, 5-11 mm long, 1-3 mm

wide, green, pubescent; corolla diameter

(not including fimbrillae) 40-67 mm; petals

obovate, c. 22 mm long, white, trichomes

mostly on veins adaxially, glabrous abaxially;

ovary 18-20 mm long, pubescent. Fruit

ellipsoid, apex beaked, 75-160 mm long,

60-100 mm wide, green with paler irregular

stripes, sparsely pubescent; mesocarp firm,

c. 5 mm wide, whitish; pedicel 40-92 mm

long, diameter 3.5-6 mm; seeds numerous

378

(about 100), broad-ovate or broad-elliptical,

9-12 mm long, 6-9 mm wide, testa cream-

coloured, marginate, flesh around seeds

white or greenish-white. Germination type

unknown. Silky Cucumber. Figs. 2 & 5.

Additional selected specimens (from 35 examined ):

Queensland. North Kennedy District: Gloucester

Island, Apr 1994, Batianoff 9404129 & Figg (BRI).

Wide Bay District Veteran S.F. [now Gympie N.P.],

c. 9 km NNE of Gympie, May 1999, Bean 14829

(BRI); Booloumba Creek area, Feb 2001, Wright s.n.

(BRI [AQ552068]); Upper Kadanga S.F., 9 km NE

of Gallangowan, Oct 2008, Forster PIF34404 et al.

(BRI); Western Road, Montville, Apr 1959, Gowlett

s.n. (BRI [AQ310073]). Moreton District: Dunumbar

Reserve, London Creek, off McDonalds Road, 2 km

NNE of Peachester, Jul 2006, Forster PIF31757 &

Smyrell (BRI); Geebung, Brisbane, Jan 1977, Gray

s.n. (BRI [AQ310082]); Daisy Hill S.F., Logan City,

Dec 1992, Thompson LOG75 (BRI); O’Possum Creek,

Springfield near Ipswich, Apr 1994, Bird s.n. (BRI

[AQ627235]); Springbrook Plateau, Purlingbrook Falls,

Apr 1984, Telford 9723 (CANB); Currumbin Creek

Road to Tomewin Gap, Jan 1984, Telford 9103 (CANB);

Mt Nathan, Mudgeeraba, May 1963, Morgan s.n. (BRI

[AQ310080]); Lyrebird Ridge Road, Springbrook, Feb

2004, Edginton & Halford s.n. (BRI [AQ763103]); Binna

Burra, Feb 2004, Fechner et al. s.n. (BRI [AQ578156]).

New South Wales. Billynudgel, Feb 1913, Boorman s.n.

(NSW146961); Whian Whian S.F., N of Lismore, Apr

1939, Jouberts.n. (NSW146959); 10.2 km along Terania

Creek Road from The Channon, Apr 2009, Nicholson

3110 (BRI); Victoria Park near Rous, S of Alstonville,

Nov 1964, Williams K22 (NSW).

Distribution and habitat : Trichosanthes

subvelutina is endemic to eastern Australia.

Apart from a highly disjunct population

on Gloucester Island east of Bowen, it is

restricted to southeast Queensland (extant

northern limit at Gympie N.P.) south to Rous

near Alstonville in northern New South Wales

(Map 2). It inhabits open or closed canopy

Eucalyptus forest, disturbed rainforest,

rainforest margins and Araucarian vineforest

at altitudes from near sea level to 600 m.

Phenology : Flowers have been recorded

in January, February, March, April, June,

October and November; fruit has been

recorded in January, February, March, April

and May.

Typification : A diverse range of specimens

were cited in the original description of this

species. These were widely distributed by

Mueller to herbaria in Europe with some

(but not all) having what are presumably

Austrobaileya 8(3): 364-386 (2011)

duplicates retained at MEL. We have selected

as lectotype of this name, a fertile collection

that is labelled as being from “Archers Brush”

and collected by Leichhardt.

Other residual syntypes are as follows

- Australia: Queensland. Moreton District:

Three Mile Scrub [Ashgrove Avenue,

Ashgrove], Burrgan, Moreton Bay, July

1843, F. Mueller s.n. (BR6606213, photo!;

MEL100126, photo!); Moreton Bay, s.dat.,

L.Leichhardt 8 (P731578, photo!); Moreton

Bay, Archers Brush, November [71843],

L.Leichhardt 26 (P731579, photo!); Moreton

Bay, Archers Brush, November-December

[71843], L. Leichhardt s.n. (P731580, photo!);

Moreton Bay, s.dat., F. Mueller s.n. (P731573,

photo!); Brisbane River, Moreton Bay, s.dat.,

F. Mueller s.n. (P731575, photo!; MEL100128,

photo!); New South Wales. Clarence River,

1861, C.Moore s.n., (P731574, photo!).

Etymology : The specific epithet is derived

from the Latin sub- (almost or somewhat) and

velutinus (velvety), probably referring to the

indumentum.

Trichosanthes section Involucraria (Ser.)

Wight, Madras J. Lit. Sci. 12: 52 (1840);

Involucraria Ser., Mem. Soc. Phys. Geneve

3: 27, t. 5 (1827). Type species: Involucraria

wallichiana Ser. [= Trichosanthes wallichiana

(Ser.) Wight]

Dioecious vines or lianas; probracts persistent;

leaves simple or compound; juvenile leaves

silvery with dark green veins; male bracts

medium or large, glandular; petal margin

fimbriate in upper half; fruit red, pulp dark

green; seeds 1-locular, flat or slightly swollen,

with a narrow margin or not margined.

Distribution : Approximately 50 species in

China, India, SE Asia, Malesia (including

New Guinea) and Australia (two species).

379

Cooper & de Boer, Trichosanthes in Australia

5. Trichosanthes pentaphylla F.Muell. ex

Benth., FI. Austral. 3: 314 (1867). Type:

Australia: Queensland. North Kennedy

District: “Mt Grame” [Mt Graham, west

of Rockingham Bay], June 1864, [probably

J.Dallachy s.n. for F.Mueller] (lecto [here

designated]: MEL100120, photo!).

Illustrations : McCubbin(1971: cover, 58-59);

Jones & Gray (1988: 354); Cooper & Cooper

(1994: cover, 290); Hyland et al. (2003);

Cooper & Cooper (2004: 145).

Dioecious vine or liana to canopy, perennial,

seasonally senescent. Stem diameter to 40 mm,

bark on older stems slightly flaky, fissured,

lenticellate, not ribbed, adventitious roots

at nodes when trailing; young branchlets 7-

ribbed, diameter 4-6 mm, glabrescent or some

trichomes clustered at nodes; nodes slightly

swollen; discoid cystoliths present, usually

denser at nodes. Probracts persistent, ovate,

rotund, oblong, rhomboid or broadly elliptic,

often cupular, base truncate, apex acute or

acuminate; margin entire, crenate or serrate

with 1-6 teeth, 2-14 mm long, 1.5-7 mm

wide, glands 2-8, glabrous or sparse minute

trichomes on both sides; discoid cystoliths

may be present. Tendrils unbranched or 2- or

3-branched. Leaves compound, digitate with

3-5 leaflets (juveniles simple or 3-foliolate,

cordate, ovate or elliptical, lobed or unlobed),

coriaceous; petioles 15-60 mm long, trichomes

sparse and minute, discoid cystoliths may be

present; central leaflets sessile or petiolules

up to 13 mm long; lamina elliptic, obovate,

elliptic-obovate or ovate-elliptic, 45-160 mm

long, 18-95 mm wide, base cuneate, rounded

or decurrent, apex acute or acuminate with a

soft mucro; margin serrate, remotely serrate

or undulate; 3-7 teeth, mucros or calluses

per side; lateral leaflets sessile or petiolules

up to 12 mm long, ovate or oblique-cordate,

50-170 mm long, 22-110 mm wide, lobed

or unlobed, base oblique, rounded, cuneate,

attenuate or cordate, apex acute or acuminate

with a soft mucro; margin serrate, remotely

serrate or undulate with 3-13 teeth, mucros

or callouses per side; upperside coarsely

scabrous, glabrous or with a few minute

trichomes along main veins especially near

base, numerous small translucent rosette¬

shaped cystoliths; underside scabrous or

smooth, glabrous or with a few translucent

trichomes near base, discoid cystoliths on

main veins; glands at base absent or up to

3, a few scattered usually present, diameter

0.6-1 mm. Male inflorescences with flowers

racemose or coaxillary with a solitary flower,

racemes 92-180 mm long with 3-14 flowers,

rachis thickened and 6-ribbed and with sparse

minute trichomes; peduncle 10-85 mm long,

3-5.25 mm wide, 6-ribbed; bracts obovate or

rhomboid, 25-40 mm long, 23-33 mm wide,

base truncate, apex obtuse or broadly obtuse,

margin fimbriate in upper half, venation

parallel, reticulation sparse, both sides

pubescent, glands scattered, gland diameter

0.5-1 mm. Male flowers: pedicels 3-5 mm

long; receptacle tube slender-funnelform,

10-angled or 10-ribbed, 38-46 mm long,

sparse minute trichomes; sepals subulate

or triangular, entire or 5-toothed, 22-38

mm long, 6-8 mm wide, green, 5-veined,

trichomes sparse and minute; corolla diameter

c. 50 mm (not including fimbrillae); petals

obdeltoid, 14-25 mm long, 15-20 mm wide,

white, pubescent. Female inflorescences

with flowers solitary or paired (one much

older). Female flowers: pedicel 10-19 mm

long; receptacle tube, salverform, 10-angled

or 10-ribbed, 40-47 mm long, glabrous

or glabrescent; sepals narrow-triangular,

entire, 5-veined, 11-15 mm long, green or

yellowish-green, clothed in minute trichomes;

corolla diameter 34-40 mm (not including

fimbrillae); petals obdeltoid, 14-18 mm long,

c. 9 mm wide, white, trichomes dense and

minute on both sides; stigma green; ovary

13-16 mm long, trichomes sparse and minute;

ovules several, in vertical rows, c. 1 mm long.

Fruit globose, ellipsoid, apex beaked, 10-

ribbed, 40-75 mm long, 50-65 mm wide,

red, glabrous or with a few trichomes at base;

mesocarp 11-14 mm wide, orange; pedicel 6-

36 mm long, 3-7 mm thick, trichomes sparse,

cystoliths absent or few to numerous; seeds

several, oblong or elliptical, flat or slightly

swollen, base rounded and may be flattened,

apex obtuse or acute, seeds 7-15 mm long,

5-8 mm wide, 2-4 mm thick, testa brown,

pulp dark green. Germination is hypogeal,

cotyledons straight. Figs. 1 & 6.

380

Austrobaileya 8(3): 364—386 (2011)

Fig. 6. . Trichosanthes pentaphylla. A. glandular probract at node. B. fruit, probract and tendril. C. adaxial view of

seeds. D. lateral view of seed. E. underside of leaf showing veins and glands. F. male inflorescence with glandular

bracts and bud. A-D from Cooper 2124 & Ford (CNS); E & F from Cooper 2134 & Jensen (CNS) (scales as indicated).

Del. W.T. Cooper.

Additional selected specimens (from 59 examined ):

Queensland. Cook District: 22 km NE of Bamaga, Feb

1994, Fell 4040 & Stanton (BRI); Lockerbie Scrub, Feb

1994, Cooper 814 (CNS); Bamaga, May 1981, Hyland

21136V (CNS); Merluna, Oct 1999, Hyland 16278

(CNS); Mowbray River Road, 3.5 km from highway, Jan

2000, Gray 7762 (CNS); Oak Beach, May 1974, Foley

431 (CNS); Kuranda, May 1993, Cooper 535 (CNS);

Kuranda Range Road, May 1976, Gray 163 (CNS); Bank

of Freshwater Creek, Freshwater, Nov 1936, Hunter s.n.

(CNS [NQNC2541); Aloomba, edge of Mulgrave River,

May 1998, Kitchener 73 (CNS); SFR310, Mar 1979, Gray

20088V (CNS); Royal Arch Tower, Chillagoe NP, Jun

2010, Cooper 2124 & Ford (CNS); Royal Arch Tower,

Chillagoe NP, Feb 2011, Cooper 2134 & Jensen (CNS);

Bank of North Johnstone River, Goondi near Innisfail,

Mar 1973, Colman s.n. (BRI [AQ9203]). North Kennedy:

Ingham, Jun 1997, Waterhouse 4431 (BRI); Townsville,

Jun 1936, Rowse s.n. (BRI [AQ310052]); Herbert River,

Euramo Station, Dansie s.n. (CNS [QRS119011]); Ayr,

Jan 1980, McGuire s.n. (BRI [AQ319650]); Home Hill

district, 1964, Wyatt s.n. (BRI [AQ310049]).

381

Cooper & de Boer, Trichosanthes in Australia

Distribution and habitat: Trichosanthes

pentaphylla is known from northeast

Queensland between Cape York and Home

Hill south of Townsville (Map 3). It mostly

occurs on riverbanks where it climbs into the

canopy and hangs along the forest edge in

wet lowland rainforest. It also occurs in the

Chillagoe-Mungana Caves N.P. at Chillagoe

where the habitat is deciduous vine thicket on

limestone.

Phenology: Flowers have been recorded in

January, February, March, April and May;

fruit has been recorded in January, February,

March, April, May, June and December.

Typification: A diverse range of specimens

were cited in the original description of this

species. These were widely distributed by

Mueller to herbaria in Europe with some

(but not all) having what are presumably

duplicates retained at MEL. We have selected

as lectotype of this name, a fertile collection

that is labelled as being from “Mt Grame”

and collected in June 1864, presumably by

Dallachy.

Other residual syntypes are as follows

- Australia: Queensland. North Kennedy

District: Rockingham Bay, s.dat., [probably

J.Dallachy s.n. for F.Mueller] (BR6605889,

photo!; K, photo!); Burdekin River, s.dat.,

[probably J.Dallachy s.n. for F.Mueller]

(MEL100121; K742661, photo!); Burdekin,

s.dat., [probably J.Dallachy s.n. forF. Mueller]

(K, photo!).

Notes: Trichosanthes pentaphylla specimens

from the Chillagoe area are rather variable

and could represent a new subspecies.

However, DNA analysis has not confirmed

this (de Boer & Cooper, unpublished). One

collection with male inflorescences is typical

of specimens from the lowland rainforests,

with a thickened rachis; bracts obovate or

rhomboid, margin fimbriate in upper half

and sparse reticulation. A second specimen

(Cooper 2101 & Ford ) from the same vicinity

has a slender rachis, bracts broadly ovate or

reniform, with an acute or acuminate apex,

margin denticulate not fimbriate, reticulate

venation and female receptacle tube deeply

ribbed. The morphology of this particular

specimen has not been included in the general

description.

Affinities: Telford (1982) suggested that

Trichosanthes pentaphylla may be conspecific

with T. trifolia (L.) Merr. (syn. T. wawrae).

However, Trichosanthes wawrae has narrowly

ovate probracts, 3-5 x 1.5-3 mm; suborbicular

juvenile leaves; adult leaves finely scabrous

above, margin entire or minutely sparsely

dentate, central leaflet base long-cuneate;

male raceme not thickened; bracts obovate or

oblong, 7-25 x 4-12 mm; sepals 4-6 mm long;

fruit pedicel 2-3 mm thick. T. pentaphylla has

probracts ovate, rotund, oblong, rhomboid

or broadly elliptic, 2-14 mm long, 1.5-7

mm wide; juvenile leaves cordate, ovate or

elliptical; adult leaves coarsely scabrous

above, margin serrate, remotely serrate or

undulate; central leaflet base cuneate, rounded

or decurrent; male raceme thickened; bracts

obovate or rhomboid, 25-40 mm long, 23-33

mm wide; sepals 22-38 mm long; fruit pedicel

3-7 mm thick.

Etymology: The specific epithet is derived

from the Greek penta- (five) and -phyllus (-

leaved) referring to the digitate compound

leaves, which often have 5 leaflets.

6. Trichosanthes morrisii W.E.Cooper

species nova a Trichosanthi pentaphyllae

similis, foliis simplicibus et sepalis margine

laciniato differt. Type: Australia: Northern

Territory. Kakadu National Park, near Gubara

Saddle, 28 December 2010, W.E.Cooper 2128,

I.Morris & R.Dempster (holo: CNS [2 sheets

+ spirit], iso: 8 sheets to be distributed to BR,

BRI, CANB, DNA, L, MO, P, UPS).

Trichosanthes sp. (D55403); Liddle et al.

(1994).

Trichosanthes sp. Kakadu (C.R. Dunlop

6639), Australian Plant Census (CHAH 2005);

Australian Plant Names Index, http://www.

anbg.gov.au/cgi-bin/apni Accessed February

2010 .

Dioecious trailing vine or liana to about c.

6 m, perennial, seasonally senescent. Stem

diameter to 20 mm, cream with numerous

black warts; young branchlets 7-ribbed with

cystoliths near nodes. Probracts persistent,

broadly elliptical, obovate or oblong; apex

acute or acuminate; margin denticulate,

crenate or serrate with 5-7 teeth, base

382

truncate or cuneate, 5-10 mm long, 2-4 mm

wide, adaxial surface with 4-7 glands, sparse

trichomes on both sides, cystoliths may be

present abaxially. Tendrils 2-4-branched.

Leaves simple, discolorous, membranous;

petioles 22-60 mm long, sparsely hairy,

discoid cystoliths present, sparse minute

trichomes may be present; lamina cordate,

3-5-lobed (juveniles cordate-deltoid, unlobed

or remotely 3-lobed), 80-170 mm long,

68-160 mm wide, base truncate, cuneate

or cordate, sinus deep and narrow or wide

(juveniles with shallow and wide sinuses),

apex acute or acuminate, usually with a soft

mucro; margin remotely denticulate with

17-24 mucros or teeth per side; upperside

scabrous, trichomes may be present on midrib

and main lateral veins; cystoliths numerous,

multicellular, rosette-shaped, diameter 0.3-1

mm; underside glabrous or a few trichomes

may be present near base; glands sparse near

base and towards apex, gland diameter 0.5-

1.25 mm; cystoliths discoid, sparse, scattered

Austrobaileya 8(3): 364-386 (2011)

along main veins. Male inflorescences with

flowers (from mature buds) racemose, usually

beside a solitary flower, racemes 140-240 mm

long, up to 11-flowered, peduncle 95-105 mm

long, rachis slender, trichomes sparse, discoid

cystoliths present; bracts persistent, rhomboid

or ovate, unlobed or shallowly lobed, 27-34

mm long, 17-24 mm wide, base attenuate,

cuneate or decurrent, apex acuminate; margin

denticulate, laciniate or fimbriate in upper

half only (entire in lower half); venation

palmate-pinnate, reticulation dense, both

sides with sparse minute trichomes denser

near margin towards apex, rarely with glands,

gland diameter c. 0.5 mm; aberrant leaf¬

like bracts may be present towards base of

inflorescence, ovate-cordate, 3-lobed, 26-57

mm long, 18-63 mm wide, base short cuneate

or decurrent, apex acute or acuminate, margin

denticulate, venation palmate-pinnate, 3-5-

veined (venation similar to leaves), reticulate,

trichomes as in main bracts. Male flowers:

pedicel on solitary flowers c. 40 mm long,

Fig. 7. Trichosanthes morrisii. A. male bract. B. seeds: left, lateral view; right, adaxial view. C. male sepal. D. leaf

underside with glands, tendril & male inflorescence. A, C, D from Cooper 2128, Morris & Dempster (CNS); B from

Dunlop 6626 (DNA) (scales as indicated). Del. W.T. Cooper.

383

Cooper & de Boer, Trichosanthes in Australia

sepals rhomboid, 3-veined, margin laciniate,

hairy, reticulate; petals not seen. Female

flowers: not seen. Fruit globose, apex beaked,

36- 55 mm long, 35-60 mm wide, orange to

red, probract persistent at base; mesocarp firm,

7-9 mm wide; pedicel up to 18 mm long and

5 mm wide, cystoliths present; seeds several,

flat, oblong, both ends obtuse, base slightly

bulbous, 11.2-12 mm long, 6.8-7.4 mm wide,

37- 4.2 mm deep, testa brown, suspended

in greenish-black pulp. Germination type

unknown. Fig. 7.

Additional selected specimens (from 8 examined ):

Australia: Northern Territory. Arnhem Land, SE Mt

Howship, Feb 1984, Dunlop 6626 (DNA); Arnhem Land,

SE Mt Howship, Feb 1984, Dunlop 6639 (DNA); Groote

Eylandt, Umbakumba, Mar 1988, Russell-Smith 5099 &

Lucas (DNA); Groote Eylandt, 8 km SW Umbakumba,

Jul 1987, Russell-Smith 2750 & Lucas (DNA); Groote

Eylandt, Jul 1987, Russell-Smith 2798 & Lucas (DNA);

Mt Brockman Outlier, Apr 1989, Russell-Smith 8056

(DNA); 1 km east of the Gubara Saddle, Kakadu N.P.,

Dec 2009, Cooper 2098 & Morris (CNS).

Distribution and habitat : Trichosanthes

morrisii is endemic to the Northern Territory

where it is currently known to occur in Kakadu

N.P., Arnhem Land and on Groote Eylandt

(Map 3). It inhabits Allosyncarpia-dominated

forest in gorges on sandstone escarpments

and in riparian evergreen vineforest on

coastal dunes. It appears to favour areas with

perennial water, which retain some moisture

through the drier seasons. Other species it co¬

occurs with are Remusatia vivipara (Roxb.)

Schott, Drynaria quercifolia (L.) J.Sm.,

Calophyllum sil Lauterb., Myristica insipida

R.Br., Tylophora benthamii Tsiang and

Desmos wardianus (F.M.Bailey) Jessup.

Phenology : Male inflorescences without

fully expanded flowers were collected in

December, and flowering probably occurs

between December and February; fruit has

been collected in February and April.

Affinities: Trichosanthes morrisii is closely

related to T. pentaphylla with similar juvenile

leaves and fruit. The former has simple

leaves, male bracts which are reticulate with

an acuminate apex, attenuate or cuneate base,

and lacinate sepals, whereas, T. pentaphylla

has compound leaves, male bracts are not

reticulate and have an obtuse apex, a truncate

base and entire sepals (excepting one aberrant

collection [Cooper 2101 & Ford ] in which the

bracts are ovate, reticulate and acute).

Etymology: Named for Northern Territory

naturalist, Ian Morris (1951—), in honour of

his extensive contribution to natural history

knowledge.

Acknowledgements

The authors thank the curators of the following

herbaria for allowing the examination of their

collections, either at the herbarium or through

loans or high resolution scans: A, BR, BRI,

CANB, CNS, DNA, E, K, KUN, L, MEL,

NSW, P, PERTH, S and UPS. Some specimen

data were sourced from Australia’s Virtual

Herbarium with permission of the Council of

Heads of Australasian Herbaria Inc.

The first author thanks Frank Zich &

Darren Crayn for support and access to CNS

herbarium; Andrew Ford, Rigel Jensen & Ian

Morris for much appreciated expertise and

companionship in the field; Bill Cooper for

the illustrations, Lyn Craven for the Latin

diagnosis, Steve Murphy for maps, Yumiko

Baba for translation; Sarah Legge, Keith

McDonald, Stephen McKenna, Hugh & Nan

Nicholson, Caroline Puente-Lelievre, Rupert

Russell and Jeremy Russell-Smith for various

contributions.

The first author is grateful for permits to

collect, which were issued by the Queensland

Department of Environment and Resource

Management as well as the Australian

Government (Kakadu National Park).

The second author was supported by

SIDA-SAREC grant SWE-2005-338.

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Map 1. Distribution of Trichosanthes cucumerina var. cucumerina (•) in Australia.

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Austrobaileya 8(3): 364—386 (2011)

Map 3. Distribution of Trichosanthespentaphylla (A) and T. morrisii (•).

Five new species of Plectranthus L.Her. (Lamiaceae)

from New South Wales and Queensland

Paul I. Forster

Summary

Forster, P I. (2011). Five new species of Plectranthus L.Her. (Lamiaceae) from New South Wales

and Queensland. Austrobaileya 8(3): 387-404. Plectranthus caldericola P.I.Forst. from New South

Wales and P. bellus P.I.Forst., P. fragrantissimus P.I.Forst., P. insularis P.I.Forst. and P. venustus

P.I.Forst. from Queensland are described as new species. All species are illustrated together with

notes on their habitat, distribution, affinities and conservation status.

Key Words: Lamiaceae, Plectranthus, Plectranthus bellus, Plectranthus caldericola, Plectranthus

fragrantissimus, Plectranthus insularis, Plectranthus venustus, Australia flora. New South Wales

flora, Queensland flora, taxonomy, new species, conservation status

PI. Forster, Queensland Herbarium, Department of Environment & Resource Management, Brisbane

Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email:paul.forster@

derm.qld.gov.au

Introduction

The genus Plectranthus L.Her. is highly

diverse in Australia with at least 35 described

species (Blake 1971; Forster 1992,1994,1996,

1997b, 1999, 2008). As noted previously,

speciation in the genus has occurred where

populations occur in isolated areas such as

mountain peaks or ranges and where suitable

habitat exists on areas of exposed rock

pavements and outcrops. This suite of habitat

factors has resulted in the ‘islands on islands’

effect (Porembski et al. 2000) and is widely

recognized as being a driving force in genetic

diversity and speciation (Carlquist 1974;

Grant 1981; Hopper 2000; Seine et al. 2000;

Kruckeberg 2002). The swift maturation

of Plectranthus individuals from seed (one

growing season) with the potential for rapid

generation turnover enables speciation within

a given locality if a suitable niche is present and

once new genetic changes arise in a population.

These changes are perhaps most likely to

occur via genetic drift, rather than selection

for markedly different growth forms. This has

resulted in subtle differences in morphology

with a ‘non-adaptive radiation’ and a ‘high

lineage diversification rate’ (Savolainen &

Forest 2005), i.e. there are a lot of similar

appearing species in microhabitats that share

edaphic and topographic similarities and that

are often highly disjunct (Linder 2003).

Accumulation of a number of field

collections, together with a critical

examination of previously described taxa and

herbarium collections, now enable a further

five new species to be described, including

one long recognized, unnamed species in the

group (Forster 2002, 2007a; Bean & Forster

2007,2010). Two of these species are currently

known from single populations, whereas the

others are more widely distributed. All of

the newly described species occur in rocky

areas of pavements and slabs, on a range of

geological substrates.

Materials and methods

This work is based on herbarium specimen

collections in BRI, CANB (including CBG),

CNS (incorporating QRS) and MEL and

a large collection of live material that was

maintained in Brisbane in the 1990s. All

taxa were examined in habitat. Descriptive

terminology is similar to that previously used

(Forster 1994). Conservation assessments are

made using the IUCN (2001) criteria.

Accepted for publication 16 July 2011

388

Taxonomy

1. Plectranthus bellus RI.Forst., species nova

affinis P. spectabili S.T.Blake sed indumento

in caulibus et lamina foliorum e trichomatibus

glandulosis constanti (in illo absentibus) et

eglandulosis divaricatis (adversum antrorsis),

dentibus foliorum 10-19 in quoque latere (in

illo 7-12) et lobo superiore calycis oblongo-

ovato (non lanceolato-ovato) differens. Typus:

Queensland. Cook District: Daintree National

Park, Adeline Creek headwaters, 18 May 1999,

P.I.Forster PIF24610 & R.Booth (holo: BRI [2

sheets]; iso: MEL).

Erect to decumbent herb or subshrub to 100

cm high; foliage with very faint ‘lemon’

scent when crushed, clammy; non-glandular

and glandular trichomes uncoloured, sessile

glands 8-celled and yellow. Roots thickened

and tuberous to fibrous. Stems square, erect

to straggling, succulent, the lower parts up

to 12 mm diameter, pale green, upper parts

with persistent indumentum, non-glandular

trichomes dense and shaggy, divaricate,

6-16-celled up to 2 mm long, glandular

trichomes sparse, sessile glands absent.

Leaves discolorous, petiolate; petioles 4-18 x

2.5-3 mm, channelled on top, non-glandular

trichomes dense and shaggy, divaricate, 6-16-

celled up to 2 mm long, glandular trichomes

sparse, sessile glands absent; laminae ovate

to broadly ovate, ± succulent, 10-80 x 10-65

mm, crenate with 10-19 teeth up to 2 mm

long on each margin, widest above middle,

secondary teeth usually present; tip acute to

obtuse; base rounded to truncate; upper surface

silver-green, velutinous, veins impressed,

non-glandular trichomes dense, divaricate,

4-12-celled up to 1 mm long, glandular

trichomes absent or scattered to sparse,

sessile glands absent; lower surface silver-

white (due to the indumentum), velutinous,

veins raised, non-glandular trichomes dense

and shaggy, divaricate, 6-16-celled up to

2 mm long, glandular trichomes sparse,

sessile glands absent. Inflorescence up to 210

mm long, single or with 1-4 side branches,

pedunculate for 15-38 mm; axis square in

cross-section, non-glandular trichomes dense

and shaggy, divaricate, 4-16-celled up to 2

mm long, glandular trichomes sparse, sessile

glands absent; bracts broadly ovate, 3-4.2

Austrobaileya 8(3): 387-404 (2011)

x 2.8-3 mm, usually forming a tight coma,

non-glandular trichomes dense and shaggy,

divaricate, 6-10-celled up to 1 mm long,

glandular trichomes sparse, sessile glands

occasional; cymes sessile; verticillasters 5 or

6-flowered, up to 7 mm apart; pedicels 2-2.5

x c. 0.5 mm, non-glandular trichomes dense

and shaggy, antrorse to divaricate, 6-12-

celled up to 1mm long, glandular trichomes

sparse, sessile glands absent. Flower calyces

3-3.5 mm long, non-glandular trichomes

dense and shaggy, divaricate, 6-12-celled up

to 1 mm long, glandular trichomes sparse,

sessile glands absent. Corolla 8-10 mm

long, lilac to purple; tube 5-6 mm long,

curved at 90-100° 3-4 mm from the base,

slightly curved upwards, ± glabrous; upper

lobes suborbicular, reflexed, 2-2.2 x 1.2-1.5

mm, non-glandular trichomes occasional

to sparse, divaricate, 4-6-celled up to 0.8

mm long, glandular trichomes and sessile

glands absent; lateral lobes oblong, 1.8-2

x 1-1.2 mm, glabrous; lower lobe broadly

ovate, 4.5-5 x 3.8-4 mm, non-glandular

trichomes dense, divaricate, 4-6-celled up

to 1 mm long, glandular trichomes sparse,

sessile glands absent; filaments filiform, 9-10

x c. 0.1 mm, lilac, fused for c. 4 mm from

the base; anthers 0.6-0.7 x 0.4-0.5 mm; style

filiform, 10-11 x c. 0.2 mm, lilac, bifid for c.

0.3 mm. Fruit calyces 3-4 mm long; upper

lobe oblong-ovate, 1.8-2 x 0.8-1 mm; lateral

lobes lanceolate-falcate, 2.6-3 x 0.7-0.8 mm;

lower lobes lanceolate-falcate, 2.8-3 x 0.3-0.4

mm. Nutlets ± circular in outline, flattened to

somewhat convex, 0.7-0.8 mm long, 0.7-0.8

mm wide, 0.4-0.5 mm thick, glossy brown,

surface smooth. Fig. 1-3.

Additional specimens examined : Cook District:

Hann Tableland, track to Radar tower station. May

2006, Forster PIF31735 & McDonald (BRI); Hann

Tableland N.P., Jul 2004, McDonald KRM2982 & Bean

(BRI). Cultivated: Tolga (ex plant collected by G. &

N. Sankowsky from base of Hann Tableland), Jul 1995,

Forster PIF17166 (BRI); Mt Coot-tha Botanic Gardens

(ex plant [collector unknown] from Hann Tableland),

Mar 2007, Forster PIF32451 (BRI).

Distribution and habitat : Plectranthus

bellus is endemic to north-east Queensland

and has been recorded once from the western

edge of the Daintree N.R and at two localities

on the Hann Tableland. Sterile plants of

what appears to be this species were also

Forster, five new species of Plectranthus

389

Fig. 1. Plectranthus bellus. A. flowering stem x0.6. B. abaxial view of leaf xl. C. adaxial view of leaf xl. D. portion of

inflorescence with two verticillasters, buds and flowers x2. E. lateral view of flower x4. F. face view of flower x4. All

from Forster PIF17166 (BRI). Del. W. Smith.

390

observed at several additional localities in

the foothills of the Hann Tableland in May

2010. Plectranthus bellus is invariably found

on rock surfaces and crevices on small (30-

200 m 2 ) areas of granite pavements that form

irregular mosaics (‘islands’) within a greater

area of eucalypt woodlands (Fig. 2 & 3). Only

scattered perennial shrubs and small annual

grasses and forbs are found in association.

Notes : There are some morphological

similarities to the much taller growing

Plectranthus spectabilis (Forster 1996), that

has been found close by with at least two

populations within 10 km on upland parts of

the Hann Tableland. Observed morphological

differences between the two species are

Austrobaileya 8(3): 387-404 (2011)

summarized in Table 1; however, the most

important differences relate to the overall

colour of the plants (due to a combination

of epidermis colour and indumentum cover)

(Fig. 3), together with the form of the flowers.

Other species of Plectranthus that also occur

on the Hann Tableland, but were not observed

to be locally sympatric with P. bellus , include

P. apreptus S.T.Blake and P. mirus S.T.Blake.

Plectranthus bellus is a quite spectacular

appearing plant and has considerable potential

in amenity horticulture and landscaping on

account of its attractive foliage, upright form

and the ability to grow in a range of soil types

in cultivation. Clones are self sterile, further

adding to its attraction as an amenity plant.

Fig. 2. Plectranthus bellus. Habitat at Hann Tableland (population voucher: Forster PIF31735 & McDonald [BRI])

Grasses in background are predominantly exotic species. Photo: P.I. Forster

Forster, five new species of Plectranthus 391

Table 1. A comparison of selected morphological character states for Plectranthus bellus

and P. spectabilis

Character state

P. bellus

P. spectabilis

life form

perennial, herb or subshrub to 1 m tall

annual or biennial, herb or

subshrub to 1.6 m tall

indumentum on

stems and leaf

laminae

non-glandular trichomes dense

and shaggy, divaricate; glandular

trichomes generally present

non-glandular trichomes

sparse to dense, antrorse;

glandular trichomes absent

number of teeth

on leaf laminae

margins

10-19

7-12

inflorescence

up to 210 mm long with 1-4 branches

from near the base; non-glandular

trichomes divaricate

up to 320 mm long with 1-11

branches from near the base

or further up the axis; non

glandular trichomes antrorse

calyx indumentum

divaricate

antrorse

upper lobe of calyx

oblong-ovate, 1.8-2 x 0.8-1 mm

lanceolate-ovate, 2.3-4 x

0.8-1.3

Fig. 3. Plectranthus bellus. Habit of individual plant (population voucher: Forster PIF31735 & McDonald [BRI]). Photo:

P I. Forster

392

Conservation status: This species is currently

known from at least three populations with a

known area of occupancy of less than 5 km 2 .

The population in the Daintree N.P. occurs in

an area that is more or less pristine with little

evidence of weed infestation. The populations

at the Hann Tableland are besieged by invasive

alien weeds that have been spread widely

over the tableland by grazing cattle and

uncontrolled wildfires prior to gazettal of the

area as National Park. These small islands of

granite pavement are under direct and current

threat from weeds such as Andropogon

gay anus Kunth, Bidens pilosa L., Lantana

camara L., Melinis minutiflora PBeauv., M.

repens (Willd.) Zizka and Praxelis clematidea

R.M.King & H.Rob. that compete for the

scarce resources in these habitats and greatly

increase fuel loads both on the pavements and

in adjacent vegetation. These Plectranthus

species are unable to survive hot fires with the

plants being killed outright or being forced to

resprout from subterranean rootstocks. Due

to the shallow soils on these pavements, the

heat and scorch from any fires often tends to

also kill the rootstocks of Plectranthus. The

infrequent occurrence of this species on small

pavements (<20 m 2 ) (Fig. 2) may be regulated

by fires that occur not necessarily only on the

pavement, but in adjacent grassy vegetation.

Map 1 . Distribution of Plectranthus bellus in north¬

east Queensland. Shaded area on map indicates

conservation reserves (National Parks, Forest Reserves

and Conservation Parks).

Austrobaileya 8(3): 387-404 (2011)

The recommended conservation status

for Plectranthus bellus is Vulnerable (criteria

B2,a,b(i,ii,iii,iv,v), D2) based on the IUCN

(2001) categories and criteria.

Etymology: The specific epithet is formed

from the Latin word bellus (beautiful) and

alludes to the pleasing form of this plant.

2. Plectranthus caldericola P.I.Forst., species

nova a P. graveolenti R.Br. foliis caulibusque

siccis non viscidis leviter citriodoris (in illo

humidis viscidis et praeditis odore anti-

septico gravi), foliis dentibus paucioribus

praeditis (14-18 non 20-42) et apicibus

rotundatis (in illo acutis usque ad obtusis),

verticillastris 2-14-floris (adversum 12-15) et

floribus pedicellis brevioribus praeditis (0.8-

1.8 mm non 2-4.5 mm) differt. Typus: New

South Wales. The Pinnacle, 13 km W of Mt

Warning, 6 April 1994, P.I. Forster PIF15093

& G.Leiper (holo: BRI [3 sheets ± spirit]; iso:

MEL, NSW).

Subshrub to 50 cm high; foliage with faint

Temon’ scent when crushed, not clammy;

non-glandular and glandular trichomes

uncoloured to silver, sessile glands 8-celled

and orange. Roots thickened and tuberous.

Stems square, erect to straggling, fleshy,

the lower parts up to 15 mm diameter, dark

purple, non-glandular trichomes dense and

shaggy, divaricate, 6-10-celled up to 3 mm

long, glandular trichomes sparse, sessile

glands scattered. Leaves discolorous, petiolate;

petioles 5-33 x 2-2.5 mm, channelled on top,

non-glandular trichomes sparse, divaricate,

6-10-celled up to 3 mm long, glandular

trichomes sparse, sessile glands scattered;

laminae broadly ovate to almost lobate, fleshy,

10-50 x 10-50 mm, crenate with 7-9 teeth up

to 3 mm long on each margin, widest above

middle, secondary teeth absent; tip rounded;

base truncate, cordate or lobate; upper surface

dark-green, veins impressed, non-glandular

trichomes sparse, divaricate, 2-8-celled up

to 1 mm long, glandular trichomes scattered,

sessile glands sparse; lower surface pale-green,

veins raised, non-glandular trichomes dense,

divaricate, 2-10-celled up to 2 mm long,

glandular trichomes sparse, sessile glands

dense. Inflorescence up to 320 mm long,

comprising 1-5 pedunculate branches from

near the base; axis square in cross-section,

Forster, five new species of Plectranthus

393

Fig. 4. Plectranthus caldericola. A. flowering stem x0.4. B. abaxial view of leaf xl. C. adaxial view of leaf xl. D. portion

of inflorescence with two verticillasters, buds and flower x3. E. lateral view of flower x6. F. face view of flower x6. G.

lateral view of fruiting calyx x6. All from Forster PIF15093 & Leiper (BRI). Del. W. Smith.

394

non-glandular trichomes sparse, divaricate,

6-10-celled up to 5 mm long, glandular

trichomes dense, sessile glands scattered;

bracts broadly-ovate to obovate, 3-5 x 2.5-4

mm, not forming a coma but approaching one,

non-glandular trichomes dense, divaricate, 6-

10-celled up to 3 mm long, glandular trichomes

absent, sessile glands scattered; cymes sessile;

verticillasters 2-14-flowered, 5-15 mm

apart; pedicels 0.8-1.8 x c. 0.2 mm diameter,

non-glandular trichomes dense, divaricate,

6-10-celled up to 2 mm long, glandular

trichomes absent, sessile glands scattered.

Flower calyces 1.8-2 mm long, non-glandular

trichomes dense and shaggy, divaricate,

6-10-celled up to 3 mm long, glandular

trichomes absent, sessile glands dense. Corolla

6-9 mm long, dark-purple; tube 3-4 mm long,

± straight or weakly curved at 160-170° 2-2.5

mm from base, slightly inflated upwards,

glabrous; upper lobes orbicular, erect, 1-1.8

x 1-2 mm, non-glandular trichomes dense,

divaricate, 2-4-celled up to 0.6 mm long,

glandular trichomes absent, sessile glands

scattered; lateral lobes oblong, 1-1.5 x 0.5-0.6

mm, glabrous; lower lobe ± orbicular, 3-4.5 x

3.5- 4.5 mm, non-glandular trichomes sparse,

divaricate, 2-4-celled up to 0.3 mm long,

glandular trichomes absent, sessile glands

scattered; filaments filiform, 7-11 x c. 0.2 mm,

lilac, fused for 3-4 mm from base; anthers c.

0.5 x 0.3 mm; style filiform, 8-9 mm long,

lilac, bifid for c. 0.8 mm. Fruit calyces 3-3.5

mm long; upper lobe orbicular-ovate, 1.2-1.6 x

1.5- 1.7 mm; lateral lobes lanceolate, 1.2-1.5 x

O. 9-1 mm; lower lobes lanceolate-falcate, 1.2-

1.5 x 0.8-1 mm. Nutlets ± circular in outline, ±

flattened, 0.6-0.7 mm long, 0.6-0.7 mm wide

and c. 0.5 mm thick, glossy brown, surface

smooth. Fig. 4.

Additional specimens examined: New South Wales.

The Pinnacle, Jan 1993, McDonald s.n. (BRI, NSW).

Distribution and habitat : Plectranthus

caldericola is known only from The Pinnacle

in the Border Ranges of northern New South

Wales. Plants grow on rock outcrops and

pavements formed of porphyritic basalts

between 920 and 1000 m with a shrubland

dominated by Leptospermum polygalifolium

Salisb. Other species of Plectranthus that

occur sympatrically are P. graveolens and

P. parviflorus Willd.

Austrobaileya 8(3): 387-404 (2011)

Notes : This species was first brought to my

attention by Graham McDonald in April

1994. Plectranthus caldericola is distinctive

in the very broad leaves that are sometimes

almost lobate, the shaggy indumentum on the

foliage and the very hairy calyces and very

short pedicels. It differs from Plectranthus

graveolens in the non-clammy foliage with

a faint Temon’ scent (versus clammy with a

strong antiseptic or foetid scent), leaves with

fewer teeth to the lamina (14-18 versus 20-42)

and rounded tips (versus acute to obtuse), the

2-14 flowered verticillasters (versus 12-15)

and the flowers with short pedicels (0.8-1.8

mm versus 2-4.5 mm).

The possibility that this species is an

example of an allotetraploid derived from an

initial hybridization between Plectranthus

graveolens and P. parviflorus needs

investigation. Plectranthus caldericola is not

typical of what one would expect from an FI

hybrid in terms of its morphology and ‘wild’

hybrids of Plectranthus have proven to be

rarely encountered (Blake 1971: 29; Forster

1991). Further localities should be sought for

this species in adjacent areas and the status of

somewhat similar plants (perhaps also from

putative hybridization events) at the North

Obelisk in New South Wales and Buchanan’s

Fort in Queensland requires further work.

Conservation status : Plectranthus caldericola

is presently known only from the type locality

where it is a very common plant. Although

this locality is within National Park, the

plants may be subjected to trampling from

bushwalkers and from the occasional wildfire.

The recommended conservation coding is

Vulnerable based on the criterion D2 (IUCN

2001 ).

Etymology : The specific epithet is derived

from calderus (a caldera) and cola (loving)

and refers to the occurrence of this species on

part of the Mt Warning caldera.

3. Plectranthus fragrantissimus P.I.Forst.,

species nova affinis P. suaveolenti S.T.Blake,

sed lamina foliorum minore angustiore

(lanceolato-ovata usque ovata, 10-40 x

6-25 mm non ovata usque late ovata, 30-

75 x 20-60 mm), florum pedicellis multo

longioribus (4-5.5. mm adversum 1.5-4 mm),

Forster, five new species of Plectranthus

395

Map 2. Distribution of Plectranthus caldericola ★,

P. fragrantissimus • and P. insularis A in southern

Queensland and northern New South Wales. Shaded area

on map indicates conservation reserves (National Parks,

Forest Reserves and Conservation Parks).

calyce lobis inferioribus lobos laterales plus

minusve aequantibus et lobis superioribus

ovatis (non subcircularibus) differens. Typus:

Queensland. Port Curtis District: State

Forest 316, Kroombit Tops, 16 February 1995,

P.I.Forster PIF16246 (holo: BRI [2 sheets +

spirit]; iso: MEL, NE).

Low growing herb to 30 cm high; foliage

with sweetly aromatic scent when crushed,

clammy; non-glandular and glandular

trichomes uncoloured, sessile glands absent.

Roots thickened, fibrous. Stems square, erect

to straggling, fleshy, the lower parts up to 4

mm diameter, pink-purple, non-glandular

trichomes sparse to dense, retrorse, 2-6-celled

up to 0.3 mm long, glandular trichomes absent

or scattered to sparse. Leaves discolorous;

petioles 2-19 x 1-1.7 mm, channelled on

top, non-glandular trichomes dense, retrorse,

2-6-celled up to 0.8 mm long, glandular

trichomes sparse; laminae lanceolate-ovate to

ovate, fleshy to succulent, 10-40 x 6-25 mm,

crenate with 7-14 teeth up to 2 mm long on

each margin, widest above middle, secondary

teeth sometimes present; tip acute; base

cuneate to truncate; upper surface dark-green

to silver-green, veins impressed, non-glandular

trichomes sparseto dense, antrorseto divaricate,

6-8-celled up to 1 mm long, glandular

trichomes sparse to dense; lower surface silver-

green, veins raised, non-glandular trichomes

dense (particularly on veins), retrorse, 6-

8-celled up to 1 mm long, glandular trichomes

sparse to dense. Inflorescence up to 230 mm

long, usually unbranched (rarely with two

side branches); axis square in cross-section,

non-glandular trichomes sparse to dense,

retrorse, 2-6-celled up to 0.2 mm long,

glandular trichomes scattered to sparse; bracts

obovate, 1.8-2 x 1.4-2 mm, not forming a

coma, non-glandular trichomes sparse to

dense, antrorse to divaricate, 2-8-celled up

to 0.6 mm long, glandular trichomes sparse;

cymes sessile; verticillasters 6-14-flowered,

4-20 mm apart; pedicels 4-5.5 x c. 0.2 mm

diameter, non-glandular trichomes sparse to

dense, antrorse to divaricate, 2-4-celled up

to 0.2 mm long, glandular trichomes scattered

to sparse. Flower calyces 2.5-3.5 mm long,

non-glandular trichomes scattered to sparse,

antrorse to divaricate, 2-6-celled up to 0.5

mm long, glandular trichomes sparse to dense.

Corolla 9-10 mm long, lilac; tube 5-5.5 mm

long, weakly curved at 10-20° c. 2 mm from

base, slightly inflated upwards, non-glandular

trichomes scattered, divaricate to retrorse, 2-4-

celled up to 0.2 mm long, glandular trichomes

absent; upper lobes suborbicular, erect, 1.8-2 x

1.8-2 mm, non-glandular trichomes scattered

to sparse, divaricate, 2-4-celled up to 0.2

mm long, glandular trichomes scattered to

sparse; lateral lobes oblong, 1.3-1.8 x 0.6-0.8

mm, glabrous or as in upper lobes; lower lobe

oblong-ovate, 4.8-5 x 3.8-4 mm, indumentum

as in upper lobes; filaments filiform, 6-8 x c.

0.2 mm, lilac, fused for 3-4 mm from base;

anthers c. 0.4 x 0.3 mm; style filiform, 8-9 mm

long, lilac, bifid for c. 0.3 mm. Fruit calyces

3.5- 4.8 mm long; upper lobe ovate, 1.5-2 x

1.7-2 mm; lateral lobes lanceolate, 1.5-2 x

0.8-0.9 mm; lower lobes lanceolate-falcate,

1.5- 2.2 x c. 0.5 mm. Nutlets ± circular in

outline, ± flattened, 0.7-0.9 mm long, 0.7-0.9

mm wide, 0.4-0.5 mm thick, glossy dark

brown, surface smooth. Fig. 5.

Additional specimens examined : Queensland. Port

Curtis District: S.F. 316 Kroombit Tops, Feb 1995,

396

Austrobaileya 8(3): 387—404 (2011)

Fig. 5. Plectrantlius fragrantissimus. A. flowering stem *1. B. abaxial view of leaf x2. C. adaxial view of leaf x2. D.

portion of inflorescence with two verticillasters, buds and flowers x4. E. lateral view of flower x6. F. face view of flower

x9. G. pedicel and calyx x9. All from Forster PIF29770 & Halford { BRI). Del. W.Smith.

Forster, five new species of Plectranthus

Forster PIF16252 (BRI, MEL, NSW); loc. cit., Feb 1995,

Forster PIF16254 (BRI); Kroombit Tops F.R., near radio

tower, Nov 2003, Forster PIF29770 & Halford (BRI,

MEL); Kroombit Tops, S.F. 316, Mar 1995, Thompson

BIL156 et al. (BRI); Kroombit Creek, Kroombit Tops

N.P., Feb 2009, Mathieson MTM307 (BRI); top of

range. Tableland Road, Kroombit Tops N.P., Dec 2010,

Mathieson MTM980 (BRI).

Distribution and habitat: Plectranthus

fragrantissimus is endemic to Kroombit Tops

National Park (formerly State Forest or Forest

Reserve) where it occurs in a series of locally

disjunct populations that may represent

subpopulations of one large metapopulation.

The plants occur at altitudes between 800

and 900 m on large outcrops or pavements

of metamorphosed quartzose sandstone that

are scattered within woodland dominated by

Eucalyptus eugenioides Sieber ex Spreng.,

E. longirostrata (Blakely) L.A.S.Johnson &

K.D.Hill and E. montivaga A.R.Bean. Typical

Plectranthusparviflorus is also widespread at

Kroombit Tops.

Notes: This species has obvious affinities

to Plectranthus suaveolens, from which it

is disjunct by about 450 km with the most

northerly recorded population of that species

west of Mt Huntley in the Main Range

near the Queensland/New South Wales

border. Plectranthus suaveolens occurs

on granophyre (Mt Barney), trachyte (Mt

Ballow), rhyolite (Mt Ernest) or granite

(Granite Belt and northern New South Wales),

but has not yet been recorded from sandstone.

As with Plectranthus suaveolens there is an

absence of sessile glands on the foliage of

P. fragrantissimus\ however, the two species

differ by the latter with smaller, narrower leaf

laminae (lanceolate-ovate to ovate, 10-40

x 6-25 mm versus ovate to broadly ovate,

30-75 x 20-60 mm), much longer flower

pedicels (4-5.5 mm versus 1.5-4 mm), the

calyx with lower lobes much the same length

as the lateral lobes (versus much longer) and

the ovate upper lobes (versus subcircular).

Plectranthus fragrantissimus is the

third angiosperm taxon endemic to the

Kroombit Tops tableland, the others being

Bulbophyllum weinthalii subsp. striatum

D.L. Jones and Parsonsia kroombitensis

J.B Williams. The tableland is also botanically

notable for a number of significant species

397

disjunctions (northern limits, indicated *) or

for geographically restricted species with a

significant proportion of their populations

being present there, including *Amperea

xiphoclada (Sieber ex Spreng.) Druce var.

xiphoclada, Astrotricha brachyandra

A.R.Bean, Boronia palasepala Duretto,

*Bossiaea rupicola A.Cunn. ex Benth.,

*Bulbine vagans E.M.Watson, Bulbophyllum

globuliforme Nicholls, *Callicomaserratifolia

Andrews, *Ceratopetalum apetalum

D.Don, Chiloglottis sylvestris D.L. Jones

& M.A.Clem., *Lasiopetalum ferrugineum

Sm. var. ferrugineum, *Maytenus silvestris

Lander & L.A.S.Johnson, Myrsine ireneae

subsp. curvata Jackes, *Persoonia volcanica

P.H.Weston & L.A.S.Johnson, *Thismia

rodwayi F.Muell., Triplarina volcanica subsp.

borealis A.R.Bean and Zieria distans Duretto

& RI.Forst. Consequently Kroombit Tops is

often regarded as an important Pleistocene

refugial area for both plant communities and

species (e.g. Rozefelds & Barnes 2001) and

can be considered a cooler upland ‘island’

analogous with the ‘sky island’ concept

applied elsewhere (Heald 1951; Watling &

Donnelly 2006). The small number of endemic

plants does not justify it being considered

as even a minor centre of endemism ( sensu

Crisp et al. 2001); however, when combined

with an increasing number of vertebrate and

invertebrate animals that are being recognized

as endemic to the area (e.g. Ponniah &

Hughes 2004), it easily qualifies as such. A

logical hypothesis is to regard Plectranthus

fragrantissimus and P. suaveolens as sister

taxa originating from an allopatric speciation

event following ‘stranding’ of the lineage on

‘sky islands’ at Kroombit Tops and the New

England Tableland and Border Ranges/Scenic

Rim of southern Queensland and adjacent

New South Wales.

Conservation status: Plectranthus

fragrantissimus is abundant at most of its

recorded localities; however, the overall area

of occurrence and occupancy is very small.

The recommended conservation status is

Vulnerable based on the criterion D2 (IUCN

2001 ).

Etymology: The specific epithet is derived

from the Latin fragrans (fragrant) and -issima

398

(most) and alludes to the sweetly smelling

foliage of this plant.

4. Plectranthus insularis RI.Forst. species

nova a P. intraterranea S.T.Blake lamina

foliorum lanceolata, lanceolato-ovata vel

oblanceolata (in illo ovata usque late ovata),

dentibus foliorum longitudine plus quam 1/3

partes latitudinis laminae ad medianum ejus,

et calyce fructificanti majore (5.8-6.5 mm

non 3.8-5.5 mm) differt. Typus: Queensland.

Darling Downs District: Mingimarny State

Forest 131, Pine Hill, 20 km S of Millmerran,

28 April 1995, P.I.ForsterPIF16468 & S.J.Figg

(holo: BRI [2 sheets + spirit]; iso: MEL).

Low multistemmed herb to 50 cm high; foliage

with faint aromatic scent, not clammy; non-

glandular and glandular trichomes clear, sessile

glands 8-celled and red. Roots thickened-

tuberous, but not ‘carrot’-like. Stems square,

erect to decumbent, fleshy with the lower

parts slightly woody and up to 5 mm diameter,

green to pink-purple, non-glandular trichomes

dense, retrorse, 2-4-celled up to 0.1 mm long,

glandular trichomes absent, sessile glands

dense. Leaves discolorous, petiolate; petioles

3-8 x 1-1.8 mm, grooved on top, non-glandular

trichomes dense, retrorse, 2-4-celled up to

0.2 mm long, glandular trichomes scattered,

sessile glands dense; laminae lanceolate,

lanceolate-ovate or oblanceolate, fleshy, 15-60

x 5-28 mm, markedly crenate with 5-8 teeth

1.5-5 mm long on each margin, 2-6 mm wide

at base, secondary teeth absent; tip acute;

base attenuate; upper surface pale green,

veins impressed, non-glandular trichomes

sparse, antrorse, 4-8-celled up to 2 mm long,

glandular trichomes sparse, sessile glands

sparse; lower surface pale green, veins raised,

non-glandular trichomes sparse, retrorse, 2-8-

celled up to 1 mm long (usually much shorter),

glandular trichomes scattered, sessile glands

dense. Inflorescence up to 180 mm long,

single-stemmed; axis square in cross-section,

non-glandular trichomes dense, retrorse, 2-4-

celled up to 0.1 mm long, glandular trichomes

scattered, sessile glands sparse; cymes sessile;

verticillasters 10-12 flowered, up to 22 mm

apart; bracts lanceolate-ovate, 2.5-3 x c.

1.5 mm, not forming a coma, non-glandular

trichomes sparse, antrorse, 2-4-celled to 0.2

mm long, glandular trichomes absent, sessile

Austrobaileya 8(3): 387-404 (2011)

glands sparse; pedicels 4.5-5 x c. 0.3 mm,

non-glandular trichomes dense, divaricate,

2-celled up to 0.1 mm long, glandular

trichomes dense, sessile glands scattered.

Flower calyces 3.5-4 mm long, non-glandular

trichomes sparse, divaricate, 2-4-celled up

to 0.1 mm long, glandular trichomes absent,

sessile glands dense. Corolla 10-11 mm long,

purple; tube 2-2.5 mm long, weakly curved at

30-45° 2-2.5 mm from base, barely inflated

upwards, glabrous; upper lobes subcircular,

reflexed, 2.2-2.5 x 2.2-2.5 mm, non-glandular

trichomes sparse, divaricate to retrorse, 2-4-

celled up to 0.1 mm long, glandular trichomes

absent, sessile glands dense; lateral lobes

oblong, 2-2.2 x 0.9-1 mm, glabrous; lower

lobe orbicular-ovate, 5-5.5 x 4.5-5.5 mm, non-

glandular trichomes scattered, divaricate, 2-

celled up to 0.1 mm long, glandular trichomes

absent, sessile glands sparse; filaments

filiform, 12-13 x c. 0.3 mm, lilac, fused for c.

4 mm from base; anthers 0.4-0.5 x c. 0.3 mm;

style filiform, 13-14 x 0.2-0.3 mm, lilac, bifid

for c. 0.5 mm. Fruit calyces 5.8-6.5 mm long;

upper lobe broadly ovate, 2.3-3.1 x 2.2-2.8

mm; lateral lobes lanceolate-falcate, 2.5-3 x

0.5-0.7 mm; lower lobes lanceolate-falcate, 3-

3.5 x 0.3-0.5 mm. Nutlets ± circular in outline,

convex, 0.8-0.9 mm long, 0.8-0.9 mm wide, c.

O. 6 mm thick, glossy brown, surface smooth.

Fig. 6.

Additional specimens examined: Queensland. Darling

Downs District: Pine Hill S.F., c. 22 km S ofMillmerran,

Nov 1997, Grimshaw PG2925 & van Baalen (BRI); Pine

Hill, Mingimarny S.F. 131,20 km S ofMillmerran, May

2000, Menkins ILM96 (BRI).

Distribution and habitat : Plectranthus

insularis is currently known only from the

type locality in an isolated State Forest south of

Millmerran in south-east Queensland where it

is the only species of this genus present. This

locality is unusual in being an outlier of granite

(with pavements and outcrops) supporting low

eucalypt woodland or shrubland surrounded

by dissimilar geology (mainly heavy clay

soils) that used to support brigalow open

forest prior to conversion to agricultural land.

The granite appears similar to that found

near Texas and Stanthorpe; however, the

only species of Plectranthus found in those

areas are P. graveolens, P. parviflorus and

P. suaveolens.

Forster, five new species of Plectranthus

Notes: Plectranthus insularis is perhaps one

of the most distinctive species in the genus

in Australia on account of the extreme lobing

(in comparison with the overall leaf laminae

size) with the marginal teeth commonly up to

5 mm long. Suggested affinities are somewhat

difficult to deduce based purely on morphology,

but include Plectranthusparviflorus (which is

closest geographically) and P. intraterraneus

399

(which is markedly disjunct with the nearest

locality in western Queensland, but perhaps

closest on morphological similarity). A

comparison of morphological character states

for these three species is given in Table 2 with

notable distinctions for Plectranthus insularis

being the leaf laminae shape, floral bract

shape and the length of the fruiting calyx.

Table 2. Comparison of some morphological character states for Plectranthus insularis ,

P. intraterraneus and P. parviflorus

Character state

P. insularis

P. intraterraneus

P. parviflorus

Habit

Erect to decumbent herb

to 50 cm high

Erect subshrub to 1

m high

Erect to decumbent

herb to 70 cm high

Root system

thickened tuberous

‘carrot’ tuberous

distinct tuber at

stem base with

fibrous roots

Foliage

faintly scented, not

clammy

sweetly scented,

clammy

faintly scented, not

clammy

Lamina shape

lanceolate, lanceolate-

ovate or oblanceolate

ovate to broadly ovate

ovate to oblong-

ovate; more rarely

± orbicular or

truncate-ovate

Leaf teeth

markedly crenate;

5-8 pairs with the teeth

length being over 1/3 of

the lamina width at its

midpoint

crenate to dentate -

crenate; 4-9 pairs

with the teeth length

being less than 1/3 of

the lamina width at its

midpoint

crenate to dentate-

crenate; 4-8

(-12) with the teeth

length being less

than 1/3 of the

lamina width at its

midpoint

Marginal teeth

length (mm)

1.5-5

1-4

0.8-2 (-4)

Number of

flowers per

verticillaster

10-12

6-10

Floral bracts

lanceolate-ovate, 2.5-3

mm long

broadly ovate to

circular ovate, 1.2-2.7

mm long

broadly ovate to

obovate, 1.1-2.5

mm long

Fruiting calyx

length (mm)

5.8-6.5

3.8-5.5

37-5.5

Conservation status: State Forest 131 is an

isolated pocket of intact vegetation within a

much greater area of cleared land. The granite

geology of this State Forest is also unique

within the context of the local Millmerran

area. Plectranthus insularis is locally very

common on pavement areas within the State

Forest, particularly at the western end. The

plants have been observed to either die, or

shrivel back to the rootstocks during periods

of drought.

400

Austrobaileya 8 ( 3 ): 387—404 ( 2011 )

Fig. 6. Plectranthus insularis. A. flowering stem x0.4. B. abaxial view of leaf xl.5. C. adaxial view of leaf xl.5. D.

portion of inflorescence with two verticillasters, buds and flowers x2. E. lateral view of flower x4. F. face view of flower

x4. All from Forster PIF16468 & Figg (BRI). Del. W. Smith.

The recommended conservation status for

Plectranthus insularis is Vulnerable (criterion

D2) based on the IUCN (2001) categories

and criteria. This species is known from a

single population (or single metapopulation

with subpopulations) with a very restricted

area of occupancy (<20 km 2 ) that is prone to

stochastic events in the short to long term.

Etymology : The specific epithet is derived

from the Latin adjective insularis (pertaining

to islands) and alludes to the occurrence of

this species on an island of granite pavement,

further surrounded by a ‘sea’ of cleared land.

Forster, five new species of Plectranthus

5. Plectranthus venustus P.I.Forst., species

nova affinis P. gratae S.T. Blake sed

caulibus praeditis trichomatibus antrorsis

non-glandularibus (adversum divaricatis),

foliis numero dentium marginalium majore

(22-33 in latere quoque non 13-15), bracteis

inflorescentiae obovatis quae comam format

(adversum ovatis et comam non formans) et

floris pedicellis multo brevioribus (0.5-1 mm

non 2.5-3 mm) differens. Typus: Queensland.

Cook District: Kennedy Hill Gorge, 21

June 1989, P.I. Forster PIF5388 & M.C. Tucker

(holo: BRI [1 sheet + spirit]; iso: CNS, MEL).

Plectranthus sp. (Restoration Island J. Le

Cussan 285); Forster (1997: 98); Forster

(2002: 96); Bean & Forster (2007: 95); Bean

& Forster (2010: 90).

Prostrate to semi-erect herb or subshrub to 40

cm high; foliage with very faint ‘lemon’ scent

when crushed, not clammy; non-glandular and

glandular trichomes uncoloured to purplish,

sessile glands 8-celled and orange-red. Roots

fibrous. Stems square, erect to straggling,

succulent, the lower parts up to 10 mm

diameter, green to pink, upper parts with

indumentum, non-glandular trichomes sparse,

retrorse, 6-10-celled up to 1.2 mm long,

glandular trichomes absent, sessile glands

absent. Leaves discolorous, petiolate; petioles

2-16 x 2.5-4 mm, slightly channelled on top,

non-glandular trichomes dense, divaricate,

6-12-celled up to 1.5 mm long, glandular

trichomes absent, sessile glands scattered to

dense; laminae lanceolate to narrow-ovate, ±

succulent, 26-90 x 14-50 mm, crenate with

22-33 teeth up to 1 mm long on each margin,

widest above middle, secondary teeth usually

present; tip acute; base cuneate to rounded;

upper surface silver-green, veins impressed,

velutinous, non-glandular trichomes sparse,

antrorse to divaricate, 6-10-celled up to 0.8

mm long, glandular trichomes absent, sessile

glands absent; lower surface silver, veins

raised, velutinous, non-glandular trichomes

dense, divaricate, 6-12-celled up to 1.5 mm

long, glandular trichomes absent, sessile glands

scattered to dense. Inflorescence up to 150 mm

long, with 3-5 pedunculate branches from

near the base; axis square in cross-section,

non-glandular trichomes scattered, retrorse,

6-10-celled up to 2 mm long, glandular

401

trichomes absent, sessile glands absent; bracts

obovate to broadly-ovate, 2-3 x 2.4-3.2

mm, usually forming a coma, non-glandular

trichomes dense, antrorse, 6-8-celled up to 1.8

mm long, glandular trichomes absent, sessile

glands scattered; cymes sessile; verticillasters

18-22-flowered, up to 10 mm apart; pedicels

0.5-1 x 0.2-0.3 mm, non-glandular trichomes

dense, divaricate, 6-10-celled up to 1.8 mm

long, glandular trichomes absent, sessile

glands scattered. Flower calyces 2.2-2.8

mm long, non-glandular trichomes dense,

divaricate, 4-10-celled up to 2 mm long,

glandular trichomes absent, sessile glands

sparse. Corolla 10-12.5 mm long, pale-lilac;

tube 4.8-5.5 mm long, curved at 100-120°

1.7-2 mm from base, slightly inflated upwards,

± glabrous; upper lobes suborbicular, reflexed,

2-2.2 x 1.8-2.2 mm, non-glandular trichomes

sparse, divaricate, 4-6-celled up to 0.7 mm

long, glandular trichomes absent, sessile

glands sparse; lateral lobes oblong, 1.5-2 x

0.9-1 mm, non-glandular trichomes scattered,

divaricate, 4-6-celled up to 0.7 mm long,

glandular trichomes absent, sessile glands

scattered; lower lobe broadly ovate, 4-4.5 x

4.2- 4.8 mm, non-glandular trichomes sparse,

divaricate, 6-celled up to 1 mm long, glandular

trichomes absent, sessile glands scattered;

filaments filiform, 7-8 x c. 0.2 mm, lilac,

fused for 4-4.5 mm from base; anthers c. 0.5

x 0.4 mm; style filiform, 7-8 mm long, lilac,

bifid for c. 0.5 mm. Fruit calyces 3.5-4 mm

long; upper lobe oblong-ovate, 1.2-2 x 1.3-1.5

mm; lateral lobes lanceolate-falcate, 1.2-2 x

0.7-0.8 mm; lower lobes lanceolate-falcate,

1.2- 1.6 x 0.5-0.6 mm. Nutlets ± circular in

outline, ± flattened, 0.8-0.9 mm long, 0.9-1

mm wide, 0.5-0.6 mm thick, glossy brown,

surface smooth. Fig. 7.

Additional specimens examined : Queensland. Cook

District: 1.8 km W of Mosquito Point Just E of Hunter

Inlet, Temple Bay, Lockhart River Aboriginal Reserve,

May 1992, Fell DF2575 (BRI); Restoration Island, Apr

1995, Le Cussan 285 (BRI); South Pap, Tozers Gap, Jul

1991, Forster PIF9072 (BRI); Puff de Looney Ridge

[Mt Tozer, Iron Range N.P], Jul 1972, Irvine 264 (BRI,

QRS); Brown’s Creek, Pascoe River, Jul 1948, Brass

19590 (BRI); 19 km from the Peninsula Development

road on a track to Wolverton via the Cook tin mine, Jun

1993, Clarkson 10114 & Neldner (BRI).

402

Distribution and habitat : Plectranthus

venustus occurs on Cape York Peninsula

in far north-eastern Queensland in an area

bounded by Mosquito Point in the north to

‘Wolverton’ in the south. Plants occur on

granite pavements and outcrops, but never on

granite rock piles where it is replaced by the

much taller growing Plectranthus excelsus

PI.Forst.

Notes : At Iron Range both Plectranthus

excelsus and P. venustus may occur in close

geographical proximity but have not been

observed growing sympatrically.

Plectranthus venustus appears to have

morphological similarities to P. gratus (Forster

1996) from the Wet Tropics, Queensland.

Observed morphological differences between

these three species are summarized in Table

Austrobaileya 8 ( 3 ): 387-404 ( 2011 )

Map 3. Distribution of Plectranthus venustus in far

north Queensland. Shaded area on map indicates

conservation reserves (National Parks, Forest Reserves

and Conservation Parks).

Table 3. Comparison of some morphological character states between Plectranthus gratus

and P. venustus

Character state

P. gratus

P. venustus

stem indumentum

non-glandular trichomes

antrorse

non-glandular trichomes

divaricate

petiole indumentum

non-glandular trichomes

antrorse, glandular trichomes

sparse, sessile glands absent

non-glandular trichomes

divaricate, glandular

trichomes absent, sessile

glands scattered to dense

laminae form and

colour

ovate; bluish-green above,

silver-green below

lanceolate to narrow-ovate;

silver-green above, silver

below

number of marginal

teeth per side of leaf

lamina

13-15

22-33

pedicels

2.5-3 x c. 0.2 mm; antrorse non-

glandular trichomes

0.5-1 x 0.2-0.3 mm;

divaricate non-glandular

trichomes

inflorescence bracts

not forming a coma; ovate,

1.3-1.8 x 1.3-1.8 mm

usually forming a coma;

obovate to broadly ovate, 2-3

x 2.4-3.2 mm

Conservation status : This species is known

from six localities (two in Iron Range N.P

and one in Restoration Island N.P.) and is not

considered to be threatened at this time. It is

likely that further populations exist within the

area of occurrence which is poorly explored

from a botanical view.

Forster, five new species of Plectranthus

403

Fig. 7. Plectranthus venustus. A. flowering stem x0.6. B. abaxial view of leaf xl. C. adaxial view of leaf xl. D. portion

of inflorescence with two verticillasters, buds and flowers x3. E. lateral view of flower x6. F. face view of flower x6. All

from Forster PIF5388 & Tucker (BR1). Del. W.Smith.

404

Etymology : The specific epithet is based

on the Latin word venustus (beautiful) and

alludes to the form of this plant.

Acknowledgements

The assistance of R. Booth, G. Leiper,

M.T. Mathieson, G. McDonald, K. R. McDonald,

I.Menkins, G. &N. Sankowsky, E. J.Thompson

and M.C.Tucker with new discoveries and

collections of both herbarium and live material

of Plectranthus is gratefully acknowledged.

W. Smith (BRI) provided the illustrations.

Translation of the diagnoses into Latin were

undertaken by P. Bostock (BRI).

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in Australasia. Contributions from the

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Queensland Plants: names and distribution , pp.

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a new species from north-eastern Queensland,

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Names and Distribution of Queensland Plants,

Algae and Lichens , pp. 94-98. Queensland

Herbarium, Environmental Protection Agency:

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(Lamiaceae), a new species from north-east

Queensland. Austrobaileya 7: 707-710.

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History 60: 56-63.

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inselberg vegetation. In S. Porembski & W.

Barthlott (eds.), Inselbergs. Ecological Studies

146: 391-407.

Iucn (2001). Red List Categories and Criteria: Version

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Wilkiea kaarruana Zich & A.J.Ford (Monimiaceae),

a new species from north-east Queensland

F.Zich 1 & A.J.Ford 2

Summary

Zich, F. & Ford, A.J. (2011). Wilkiea kaarruana Zich & A.J.Ford (Monimiaceae), a new species from

north-east Queensland. Austrobaileya 8(3): 405-411. Wilkiea kaarruana Zich & A.J.Ford endemic

to the Wet Tropics of north-east Queensland is described, illustrated and distinguished from related

Australian species. Notes on habitat, distribution, and conservation status are provided.

Key Words: Monimiaceae, Wilkiea , Wilkiea kaarruana , Australia flora, Queensland flora, new

species, taxonomy, identification key, hyperstigma

'F.Zich, Australian Tropical Herbarium, Sir Robert Norman Building, James Cook University Cairns

Campus, PO Box 6811, Cairns, Queensland 4870, Australia

2 A.Ford, CSIRO, Ecosystem Sciences, Tropical Forest Research Centre, P.O. Box 780, Atherton,

Queensland 4883, Australia

Introduction

Wilkiea F.Muell. (Monimiaceae) is a genus

of approximately 12 species distributed from

Papua New Guinea to Australia (Philipson

1980; Whiffin & Foreman 2007). In their

treatment of the genus for the Flora of

Australia , Whiffin & Foreman (2007) elected

not to describe two poorly known taxa from

north Queensland. Recent collections of

one of these taxa, Wilkiea sp. A (Whiffin &

Foreman 2007) by the authors have provided

for the first time female flowers, and abundant

new male flowering specimens, which have

enabled confident placement of the taxon in

the genus Wilkiea.

The flowers of Wilkiea have been recorded

as possessing a hyperstigma (Endress 1979,

1980); that is, a sticky surface receiving pollen

that is not the stigma itself Mucilaginous

secretion from the stigmas as a reward for

pollinators has been reported for at least one

other genus of Monimiaceae, namely the

Madagascan Tambourissa Sonn. (Lorence

1985), which is pollinated by flies and beetles,

while the pollination of Australian Wilkiea

huegeliana (Tul.) A.DC. appears to be mainly

by thrips (Williams et al. 2001).

Staedler & Endress (2009) have revealed

that all taxa in Monimiaceae that possess a

hyperstigma also have a complex, whorled

floral phyllotaxis, in which merism may

increase or decrease once or several times

between whorls. They call organs tepals when

they are anywhere on the floral cup (regardless

of whether on top of the cup or toward the base

of the cup), and bracts when they are below

the floral cup on the pedicel.

Materials and methods

The study is based upon the examination

of herbarium material from BRI and

CNS (formerly QRS) combined with field

observations. All specimens cited have been

seen by one or both authors.

Measurements of the floral parts of Wilkiea

kaarruana are based on fresh material and

material preserved in 70% ethanol. Common

abbreviations in the specimen citations are:

L.A. (Logging Area), N.R/R. (National Park/

Reserve), S.F./R. (State Forest/Reserve) and

T.R. (Timber Reserve).

NCA is an acronym for the Queensland

Nature Conservation Act (1992) and its

associated schedules. Discussions of

conservation status are made in reference to

the criteria of the IUCN (2001).

Accepted for publication 29 June 2011

406

The abbreviation RE in the distribution and

habitat notes refers to Regional Ecosystem,

descriptions of which can be viewed at (in

this case):

http://www.derm.qld.gov.au/wildlife-ecosystems/

biodiversity/regional_ecosy stems/search.

php?&page=31

Extent of occurrence estimates were

derived from the validation of original

collection localities. These data points were

loaded into ESRI ArcView 3.2 and the draw

polygon feature was used to calculate the area

between the points. The area of occupation

estimates were principally derived from a

digital Regional Ecosystem map supplemented

by expert opinion of vegetation types and

habitats within the Wet Tropics bioregion

(hereafter referred to as the Wet Tropics)

(Environment Australia 2005).

Taxonomy

Wilkiea kaarruana Zich & A. J.Ford, species

nova affinis Wilkiea smithii Whiffin a qua

cortice suberoso, ramulis glabris, pedicellis

glabris, floribus femineis longioribus (22-24

mm longis), et stigmate conico differt. Typus:

Queensland. Cook District: Wooroonooran

National Park, 9.5 km along Maple Creek

Road from South Johnstone River bridge, 20

November 2007, A.Ford 5205 (holo: BRI; iso:

A, BO, CANB, CNS, H, K, L, LAE, MO, NSW,

SYD, MEL, SING, US, Z distribuendi).

Wilkiea sp. Palmerston (B.P.Hyland 80);

Jessup (2007, 2010).

Wilkiea sp. Palmerston (BH 80RFK); Hyland

etal. (2003).

Wilkiea sp. Palmerston; Cooper & Cooper

(2004: 320).

Wilkiea sp. A; Whiffin & Foreman (2007:

81).

Illustration : Cooper & Cooper (2004: 320),

as Wilkiea sp. (Palmerston).

Shrub or small tree 5-12 m tall, androdioecious

or monoecious; bark corky, thick and

longitudinally fissured. Leaves glabrous,

slaty green above when fresh; petiole 5-12

mm long, pink-red; lamina elliptic to oblong-

Austrobaileya 8(3): 405-411 (2011)

elliptic, 6-15 cm long, 2.5-6 cm wide, cuneate

to attenuate at base, entire, acuminate at apex,

mucronate; midrib raised above and below.

Male inflorescence axillary, terminal and

ramiflorous, often fasciculate, 1-4 cm long;

occasionally solitary, mostly triads (botryoids

sensu Endress [2010]), sometimes fascicles

of triads (botryoids), in total 1-15 flowers;

pedicels 5-18 mm long, glabrous; bracts

lanceolate, c. 1.1 mm long. Male flowers

globose to depressed globose, c. 2-3.5 mm

wide, yellow to yellow-orange; tepals rounded

to truncate 4-6, c. 1.8 mm long, glabrous on

both surfaces, two pairs around ostiole, third

pair if present near base of floral cup; stamens

4, c. 1.1 mm long, in equal pairs, outer 2

positioned on floral cup wall and higher than

inner two which are positioned on the base

of receptacle, anthers opening by apical slit

curving over apex, filaments very short, hairy.

Female inflorescence terminal, 3-5 cm long,

fascicles of triads (botyroids), in total 6-14

flowers, pedicels 11-24 mm long, glabrous.

Female flowers depressed globose, c. 7 mm

diameter, yellow-orange fading to green at

base; tepals 4-6, 2 pairs around ostiole, outer

pair thin, inner pair greatly thickened, forming

a floral cup, third pair when present often

consisting of one tepal near the base of floral

cup and the opposing tepal on the side of the

floral cup; carpels 21-26, angular, tapering,

densely pubescent with golden, more or less

appressed hairs; stigma short, conical and

black; staminodes absent. Fruiting receptacle,

6—10(—15) mm wide, pubescent; mature fruit

not known. Fig. 1.

Additional selected specimens examined: Queensland.

Cook District: Big Tableland via Cooktown, s.dat, Jack

inEldridge s.n. (BRI [AQ63833]); Gap Creek, c. 22 miles

[36 km] S by E of Cooktown, Oct 1969, Smith s.n. (BRI

[AQ339983]); loc. cit., Smith s.n. (BRI [AQ339984]); loc.

cit., Smith s.n. (BRI [AQ339988]); Daintree N.R, Cedar

Bay Section, Gap Creek, Nov 2000, Forster PIF26462

et al. (BRI); Daintree N.R, Daintree River headwaters.

May 1998, Forster PIF22947 et al. (BRI); Mitchell

Bomber crash site. East Bartle Frere, Wooroonooran

N.R, Nov 1994, Hunter 2147 (BRI); Jordan L.A., 16.5

km SE of Millaa Millaa, site 2, Oct 1988, Jessup GJM

2095 et al. (BRI); Wooroonooran N.R, 1.1 km along K-

Tree road from Kaarru Creek crossing, Nov 2007, Ford

5208 1 (BRI, CNS); Wooroonooran N.R, 1.1 km along K-

Tree road from Kaarru Creek crossing, Nov 2008, Zich

626 x (BRI, CNS); Wooroonooran N.R, Maple Creek

Road, CSIRO Climate Change Plot, Charappa, Nov

Zich & Ford, Wilkiea kaarruana (Monimiaceae)

2008, Zich 62? (BRI, CANB, CNS); Palmerston F.R.,

old logging road towards Karang-Garee Falls, off South

Johnstone Forestry road, Oct 2003, Ford 4174 & Holmes

(BRI, CNS); ( - flowering specimen).

Distribution and habitat : Wilkiea kaarruana

is endemic to the Wet Tropics bioregion

in north-eastern Queensland, where it is

currently known to occur from the Cooktown

area (Big Tableland and Mt Finnigan environs)

to the South Johnstone River (south-east of

Millaa Millaa) (Map 1). However, within this

large range Wilkiea kaarruana is confined to

four distinct and disjunct populations (from

north to south):

1. Mt Finnigan area (from Big Tableland to

Gap Creek)

2. Daintree River headwaters

3. Mt Bartle Frere

4. South Johnstone River catchment.

It inhabits predominantly the wetter and

more mountainous notophyll vine-forests/

rainforests on soils derived from granite, fine

grained metasediments (including mudstone)

and basalt. However, it is more commonly

encountered on granitic substrates, with

occurrences on basalt and metasediments

being less common. Common canopy

species that grow in association with Wilkiea

kaarruana include: Cardwellia sublimis

F.Muell., Carnarvonia araliifolia F.Muell.,

Cryptocarya mackinnoniana F.Muell.,

Doryphora aromatica (F.M.Bailey) L.S.Sm.,

Elaeocarpus ruminatus F.Muell., Flindersia

bourjotiana F.Muell., Flindersia pimenteliana

F.Muell., Franciscodendron laurifolium

(F.Muell.) B.Hyland & Steenis, Sloanea

macbrydei F.Muell., Syzygium johnsonii

(F.Muell.) B.Hyland, Syzygium papyraceum

B. Hyland and Xanthophyllum octandrum

(F.Muell.) Domin. Common small trees and

shrubs throughout most of its range include:

Apodytes brachystylis F.Muell., Ardisia

brevipedata F.Muell., Bobea myrtoides

(F.Muell.) Valeton, Bubbia semecarpoides

(F.Muell.) B.L.Burtt, Chionanthus axillaris

R.Br., Citronella smythii (F.Muell.)

R.A. Howard, Gossia spp., Pilidiostigma

tropicum L.S.Sm., Pittosporum rubiginosum

A.Cunn., Polyscias australiana (F.Muell.)

407

Philipson and Psychotria spp. Altitudinal

range, from existing specimens, is 60-1090

Wilkiea kaarruana has been collected

or reliably reported in the following REs:

7.8.2a (rarely); 7.11.1a (occasionally); 7.12.1a

(occasionally), 7.12.16a (commonly) and

7.12.19a (rarely).

Phenology : Flowers have been recorded in

October and November; immature fruits have

been recorded in March.

Affinities: Wilkiea kaarruana is

morphologically most similar to W. smithii

and W. hylandii Whiffin. However these

species are easily distinguished by several

key features and geographic distribution.

All three species have pubescent carpels and

glabrous to glabrescent adaxial leaf surfaces.

A comparison of diagnostic differences

between these three species (and Wilkiea

austroqueenslandica ) is provided in Table 1.

Whiffin (2007) suggests that Wilkiea

austroqueenslandica, W. hylandii and

W. smithii form a “distinctive group”, as

they all have pubescent inflorescences and

carpels. Although W. kaarruana has glabrous

inflorescences, the carpels are distinctly

pubescent. Molecular studies by Renner et

al. (2010, Fig. 2) and Renner pers. comm.

(2009) place Wilkiea kaarruana close to

W. austroqueenslandica Domin., W. smithii

and W. hylandii.

Notes : In habitat Wilkiea kaarruana is easily

recognized by the thick and longitudinally

fissured pale corky bark. A specimen from the

Big Tableland ( Jack in Eldridge s.n .) consists

of a piece of bark 20mm thick mounted on a

sheet and the name “Corkwood Tree” on the

original label. No other species of Wilkiea , or

Monimiaceae, in Australia have this striking

feature though it has been reported for at least

two species of Monimiaceae on the island of

New Guinea, currently assigned to the island

endemic genus Kairoa Philipson (Renner &

Takeuchi 2009). Recent molecular research

on Kairoa indicates it should be included with

Wilkiea , along with Kibara Endl., which is

likewise embedded among species of Wilkiea

(Renner et al. 2010).

408

Austrobaileya 8(3): 405-411 (2011)

Table 1. Morphological comparison between Wilkiea austroqueenslandica , W. hylandii ,

W. kaarruana and W. smithii

Character

state

W. austroqueenslandica

W. hylandii

W. kaarruana

W. smithii

Bark

finely wrinkled

not recorded (but

not corky)

corky

tessellated

flaky

Leaf

indumentum

glabrous

pubescent when

young, glabrescent

or persisting on

midrib

glabrous

densely

pubescent

when young,

esp. midrib,

glabrescent

Leaf margin

prominently, irregularly

toothed

prominently

toothed in upper

half

entire

Male

inflorescence

position

axillary

axillary,

sometimes

fasciculate

axillary,

terminal and

ramiflorous,

often fasciculate

axillary,

sometimes

fasciculate

Male

inflorescence

triad or fascicle of triads

triad or fascicle of

triads

occasionally

solitary, mostly

2-flowered or

sometimes triads

or fascicle of

triads

triad or

fascicle of

triads

Male flower

c. 8 mm

4-5 mm

5-18 mm

6-8 mm

pedicel

length &

indumentum

densely pubescent

glabrous

densely

pubescent

Male flowers

tepal number

8 as 4 pairs

6, with 2 pairs

around ostiole,

and 1 lower on

floral cup

4 to 6. 2 as

pairs around an

ostiole. 3rd pair,

if present at base

of floral cup

8 in 2 whorls

Male flowers

stamen

number

(6)8(10)

usually 4

Female

inflorescence

position

axillary

axillary or

terminal

axillary or

terminal

Female

10-15 mm

7-15 mm

22-24 mm

5-8 mm

flower pedicel

length &

indumentum

densely silky pubescent

densely pubescent

glabrous

densely

pubescent

Female flower

tepal number

4 or 6

Carpel

number

21-26

20-40

Stigma shape

hemispherical

conical

hemispherical

Zich & Ford, Wilkiea kaarruana (Monimiaceae)

Although Wilkiea kaarruana is a small

tree, it is remarkable that there are so few

fertile specimens in herbaria. The few

flowering collections have sometimes made

it difficult to determine levels of dioecy and

monoecy. As noted by Whiffin & Foreman

(2007) the genus Wilkiea is monoecious or

perhaps rarely dioecious, and “It is likely

that many (perhaps all) species described

as dioecious have male and female flowers

on the same plant but usually not at the

same time. For example, many plants with

mature fruit have young flowers that are

male”. Observations made during this study

corroborate these statements, with Ford 5208

having mature female flowers and younger

male flowers. These male flowers would

likely come into anthesis after the female

flowers were fertilized. Also, despite regular

monitoring of several plants before and after

flowering, we found no evidence of male plants

(Ford 5205 and Zich 628 ) ever producing

female flowers. Accordingly, we suggest that

Wilkiea kaarruana is androdioecious (male

dioecious) and monoecious (having pure male

plants growing together with monoecious

plants). Further field observations are

required to assess the reproductive strategy

within Wilkiea kaarruana , and indeed for all

Australian Monimiaceae.

Floral observations of female Wilkiea

kaarruana conducted as part of this study

have confirmed the presence of a hyperstigma.

The outline of the flower is globular, the inner

pair of tepals is greatly thickened and appears

to produce mucilage at anthesis. The amount

of mucilage in dissected flowers varied but

was observed to fill the floral cup creating a

continuous cover over the carpels to the apical

pore (see Fig. 1). Despite examination of many

flowers no insects or their larvae were found,

though possible evidence of insect damage

was observed in the partial destruction of the

apical pore as described and illustrated by

Endress (1980: 85, fig.35).

The presence of a third whorl of tepals in

both male and female flowers in W. kaarruana

is not unusual within Wilkiea ; however, it is

more usual for male flowers to possess more

whorls than female flowers (Whiffin 2007).

409

Leaf oils in Australian species of

Wilkiea have been studied by Brophy et

al. (2009). Wilkiea kaarruana produced a

sesquiterpenoid oil similar to the related

Wilkiea smithii Whiffin; although this

latter species also contained very small

amounts of monoterpenes. Wilkiea

austroqueenslandica Domin, another

closely related species, yielded an oil also

dominated by sesquiterpenes, but had a

much higher proportion of monoterpenes.

Wilkiea hylandii Whiffin was not sampled

in their study.

Conservation status : Most existing collections

of this species have been made within the

World Heritage Area of the Wet Tropics.

Wilkiea kaarruana has been collected in

Daintree and Wooroonooran National Parks.

The species has a wide, albeit restricted and

disjunct geographical range with an area of

occupation estimated to be 110 km 2 . It is not

considered at risk.

Etymology : The specific epithet honours

Kaarru (pronounced Kaa-roo) Creek and

Kaarru Logging Area, where several

collections of the species, including the first

female flowering collection were made. Now

known as Wooroonooran National Park, the

area was previously referred to as State Forest

Reserve 756.

Acknowledgements

The authors wish to thank Will Smith

for the illustrations. Peter Bostock provided

the distribution map and Lyn Craven the

translation of the diagnosis into Latin. Peter

Endress and Susanne Renner improved an

earlier draft. Permits to collect in the Wet

Tropics were issued by the Department of

Environment & Resource Management. The

curators and staff at BRI are thanked for

allowing access to specimens and the use of

their facilities.

410

Austrobaileya 8(3): 405-411 (2011)

Fig. 1. Wilkiea kaarruana. A. large branch showing longitudinal corky fissures x 1. B. transverse section of large

branch x 2. C. branchlet with male inflorescences x 0.5. D. leaf adaxial surface with extra venation detail x l.E. lateral

view of male flower showing off-set tepal on floral tube and bract on pedicel x 10. F. face view of male flower showing

tepals around ostiole and anthers x 12. G. lateral view of female flower showing off-set tepals on floral cup x 10. H.

face view of female flower showing tepals around ostiole and off-set tepal on floral cup x 12.1. section of female flower

showing hairy carpels and mucilaginous cover from the hyperstigma x 12. A-B from Forster PIF26462 et al. (BRI);

C-F from FordAF5205 (CNS); G-I from FordAF5208 (CNS). Del. W.Smith

Zich & Ford, Wilkiea kaarruana (Monimiaceae)

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(1965). Flora of Java, Vol. 2. N.V.P. Noordhoff:

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Australian Tropical Rainforest. Nokomis

Editions: Melbourne.

Endress, P.K. (1979). Noncarpellary pollination and

‘hyperstigma’ in an angiosperm ( Tambourissa

religiosa, Monimiaceae). Experientia 35: 45.

- (1980). Ontogeny, function and evolution of

extreme floral construction in Monimiaceae.

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- (2010). Disentangling confusions in

inflorescence morphology: Patterns and

diversity of reproductive shoot ramification

in angiosperms. Journal of Systematics and

Evolution 48(4): 225-239.

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Biogeographic Regionalisation for Australia

(IBRA) and Development of Version 5.1

- Summary report (2000). Updated, IBRA

Version 6.1 (Digital Data, metadata), [cited 24

June 2009], http://www.environment.gov.au/

parks/nrs/science/bioregion-framework/ibra

Hyland, B.P.M., Whiffin, T., Christophel, D.C., Gray,

B. & Elick, R.W. (2003). Australian Tropical

Rain Forest Plants. Trees, Shrubs and Vines.

CD-ROM. CSIRO Publishing: Melbourne.

Jessup, L.W. (2007). Monimiaceae. In P.D.Bostock &

A.E.Holland (eds.). Census of the Queensland

Flora 2007 ’ p.l 16. Environmental Protection

Agency: Brisbane.

- (2010). Monimiaceae. In P.D.Bostock &

A.E.Holland (eds.). Census of the Queensland

Flora 2010 , p. 111. Queensland Herbarium,

Department of Environment & Resource

Management: Brisbane.

Lorence, D.H. (1985). A monograph of the Monimiaceae

(Laurales) in the Malagasy Region (Southwest

Indian Ocean). Annals of the Missouri Botanical

Garden 72: 1-165.

Philipson, W.R. (1980). The genus Wilkiea (Monimiaceae)

in Papua New Guinea. Blumea 26: 365-366.

Renner, S.S. & Feil, J.P (1993). Pollinators of tropical

dioecious angiosperms. American Journal of

Botany 80: 1100-1107.

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Renner, S.S., Strijk, J.S., Strasberg, D. & Thebaud,

C. (2010). Biogeography of the Monimiaceae

(Laurales): a role for East Gondwana and long¬

distance dispersal, but not West Gondwana.

Journal of Biogeography 37: 1227-1238.

Renner, S.S. & Takeuchi, W.N. (2009). A phylogeny

and revised circumscription for Kairoa

(Monimiaceae), with the description of a new

species from Papua New Guinea. Harvard

Papers in Botany 14: 71-81.

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I. Floral morphology and gametophyte

development of Hedycarya arborea J.R. et G.

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Atherospermataceae, Siparunaceae,

Monimiaceae). International Journal of Plant

Sciences 170: 522-550.

Whiffin, T. & Foreman, D.B. (2007). Monimiaceae.

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Publishing: Melbourne.

Williams, G.A., Adam, P. & Mound, L.A. (2001). Thrips

(Thysanoptera) pollination in Australian

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1 - 21 .

Map 1. Distribution of Wilkiea kaarruana in Queensland.

Shaded areas indicate conservation reserves.

New and reinstated species of the Solanum

ellipticum R.Br. (Solanaceae) species group

A.R.Bean

Summary

Bean, A.R. (2011). New and reinstated species of the Solanum ellipticum R.Br. (Solanaceae) species

group. Austrobaileya 8(3): 412-430. The Solanum ellipticum species group is endemic to Australia,

and is here considered to comprise 13 species. Solanum lithophilum F.Muell. and S. cleistogamum

Symon are reinstated. Solanum aridicola A.R.Bean, S. callosum A.R.Bean, S. emmottii A.R.Bean

and S. unispinum A.R.Bean are described as new. The four new species and Solanum lithophilum are

illustrated and distribution maps are provided for them. A key to the identification of species in the

Solanum ellipticum group is presented.

Key Words: Solanaceae, Solanum, Solanum aridicola, Solanum callosum, Solanum ellipticum,

Solanum emmottii, Solanum lithophilum, Solanum unispinum, Australia flora. New South Wales

flora. Northern Territory flora, Queensland flora. Western Australia flora, taxonomy, new species,

identification key

A.R. Bean, Queensland Herbarium, Department of Environment & Resource Management, Brisbane

Botanic Gardens, Mt Coot-tha Road, Toowong 4066, Queensland, Australia. E-mail: tony.bean@

derm.qld.gov.au

Introduction

Solanum ellipticum was named by Robert

Brown in 1810, based on type material

collected near St Lawrence on the central

Queensland coast. For many years, the name

has been widely applied to Solanum species

with the following characteristics: prostrate or

low growing, stems moderately prickly, calyx

prickly; leaves ovate to elliptical with entire

(or repand) margins, densely stellate-hairy

at least on the lower surface; inflorescences

racemose; fruits green, yellow-green or

almost white at maturity; calyx not accrescent.

Solanum specimens matching these criteria

occur in all mainland states except Victoria.

The current author was able to distinguish

some taxa (, Solanum crebrispinum A.R.Bean,

S. senticosum A.R.Bean) within S. ellipticum

sens. lat. based on herbarium material (Bean

2004). Subsequent field studies have provided

new insights and revealed characters that

are not readily apparent from the standard

herbarium specimen. For instance, there

are consistent differences between taxa in

growth habit (Figs. 1 & 2), corolla shape

and indumentum, style orientation (erect or

eccentric), stigma size and colour, fruit colour

and placentation, and pericarp thickness.

Accepted for publication 27 June 2011

Bean (2004) included 10 species within

the informal Solanum ellipticum group

(Group 27), which is endemic to Australia.

The membership of that group needs

alteration. On morphological grounds, the

author now recognises that the affinities

of S. crassitomentosum Domin lie with

S. furfuraceum R.Br. (Group 27B), that

S. angustum Domin belongs near A graniticum

A.R.Bean in Group 25, and that A argopetalum

A.R.Bean belongs in the S. esuriale Lindl.

group (Group 27C). Additions to the group

comprise the reinstated S. cleistogamum

Symon and S. lithophilum F.Muell., and the

four species named in this paper (S. unispinum,

S. emmottii, S. callosum and S. aridicola).

With these changes, Group 27 now comprises

13 species, namely S. aridicola, S. callosum,

S. cleistogamum, S. crebrispinum A.R.Bean,

S. dianthophorum Dunal, S. ellipticum,

S. emmottii, S. horridum Dunal ex Poir.,

S. lithophilum, S. quadriloculatum F.Muell.,

S. senticosum A.R.Bean, S. terraneum Symon

and S. unispinum.

Solanum terraneum is tentatively retained

in the S. ellipticum group following Symon

(1981), but no specimens have been seen by

the present author, and so it is not included

in the key to species and is not considered

further in this paper.

Fig. 1. Plant of Solanum ellipticum at Homevale National Park ( Bean 30650 et ah). Photograph: A.R.Bean

Full descriptions are given for the new

species, the reinstated species {Solanum

cleistogamum and S. lithophilum ), and for the

recircumscribed Solanum ellipticum.

Materials and methods

Relevant specimens have been examined in

the herbaria BRI, MEL, NSW and NT; as have

specimens of Solanum ellipticum sens. lat.

from CANB, and material of S. cleistogamum

from PERTH.

Measurement techniques and terminology

are as per Bean (2004). All species except

Solanum dianthophorum and S. terraneum

have been examined in the field. Species

treatments are in alphabetical order.

414

Austrobaileya 8(3): 412^130 (2011)

Taxonomy

Key to the species in the Solanum ellipticum group

1 Stems and branchlets without prickles.S. dianthophorum

1. Stems and branchlets sparsely to densely prickly.2

2 Style protruding between anthers or slightly bent, displacing one anther,

erect for most of its length but recurved near apex.3

2. Style bent strongly near base, displaced from the centre of the flower

(eccentric).10

3 Stems ultimately ascending or erect; bushy plants.4

3. Stems ultimately prostrate or procumbent; ground-covering plants.8

4 Calyx prickles 40-70, very stout and pungent.5

4. Calyx prickles rather flimsy, 1-40, or sometimes absent.6

5 Leaves 1.6-1.8 times longer than broad, base obtuse or cordate; ovary

and style with very shortly stalked glandular hairs; calyx stellate hairs

0.7-0.9 mm across.S. crebrispinum

5. Leaves 1.9-2.7 times longer than broad, base cuneate; ovary and style

glabrous; calyx stellate hairs 0.4-0.6 mm across.S. senticosum

6 Leaves 2.2-3.5 cm long; calyx prickles 0-5; inflorescences 1 or 2

flowered.S. unispinum

6. Leaves 4.9-14.5 cm long; calyx prickles 5-40; inflorescences 7-15 flowered

7 Stellate hairs consistently present on inner surface of corolla; branchlets

terete; prickles glabrous.S. callosum

7. Stellate hairs consistently absent from inner surface of corolla; branchlets

usually ridged; prickles with stellate hairs attached at their bases

.S. quadriloculatum

8 Calyx prickles >50; fruits 18-22 mm diameter; inflorescences 1(—2)-

flowered.S. horridum

8. Calyx prickles 5-50; fruits 9-17 mm diameter; inflorescences 1-6-

flowered.9

9 Fruits 9-12 mm diameter; anthers 2-2.8 mm long; central ray of stellate

hairs 1-2 times as long as laterals.S. cleistogamum

9. Fruits 13-17 mm diameter; anthers 4-5.5 mm long; central ray of stellate

hairs 0.5-1 times as long as laterals.S. ellipticum

10 Plant prostrate to procumbent; calyx prickles pungent, 3.5-8 mm long

.S. aridicola

10. Plant bushy, with ascending stems; calyx prickles flimsy, 0.5-4 mm

long.11

11 Branchlet hairs not floccose (stalks 0-0.2 mm); calyx prickles 0.5-2 mm

long; corolla 9-13 mm long.S. emmottii

11. Branchlet hairs floccose (stalks 0-2 mm); calyx prickles 2-4 mm long;

corolla 14-20 mm long.S. lithophilum

Bean, Solarium ellipticum group

415

Fig. 2. Population of Solarium lithophilum , north of Alice Springs (site of collection for Bean 25457 & Albrecht).

Photograph: A.R.Bean

Solanum aridicola A.R.Bean species nova

S. horrido affinis sed ab eo inflorescentiis

3-8-floris, praesentia pilorum stellatorum

longe stipitatorum in pagina superiore

foliorum et marginibus foliorum aequabiliter

integris differens. Typus: Northern Territory.

Larapinta trail, Standley Chasm area. c.

40 km W of Alice Springs, 1 August 2006,

A.R.Bean 25449 & D.E.Albrecht (holo: BRI;

iso: CANB, MEL, NSW, NT).

S. ellipticum form “b”; Symon (1981).

S. ellipticum var. Ranges (D.E.Albrecht

5807); Kerrigan & Albrecht (2007), Albrecht

et al. (2007: 176).

Herbaceous resprouter or rhizomatous

perennial shrub 0.1—0.3 m high. Stems initially

ascending, then prostrate or sprawling. Adult

branchlets yellow or rusty; prickles 35-56

per cm, straight, acicular, 4-12 mm long,

14-20 times longer than wide, glabrous;

stellate hairs very dense, lateral rays 8-9,

porrect or ascending, central ray 1-2 times

as long as laterals; Type 2 hairs absent. Adult

leaves elliptical or ovate, entire or shallowly

lobed throughout with 2-3 on each side;

lobes obtuse, lobing index 1-1.1; lamina

2.8-9.8 cm long, 1.3-4.4 cm wide, 2.1-2.8

times longer than broad, apex acute, base

cuneate or obtuse, oblique part 0-4.5 mm

long, obliqueness index 0-6%; petioles 1.3-4

cm long, 26-56% length of lamina, prickles

present. Upper leaf surface green; prickles

present on midvein only or present on midvein

and lateral veins, 1-20, straight, acicular,

3.5-13 mm long; stellate hairs distributed

throughout, very dense; lateral rays 8, porrect

or ascending, central ray 1-2 times as long

as laterals; simple hairs absent; Type 2 hairs

absent. Lower leaf surface yellowish; prickles

present on midvein only or present on midvein

416

and lateral veins, 3-35, straight, acicular;

stellate hairs very dense, lateral rays 8-12,

porrect, ascending or multiradiate, central ray

0.9-2 times as long as laterals; simple hairs

absent; Type 2 hairs absent. Inflorescence

supra-axillary, cymose (pseudo-racemose),

common peduncle 11-27 mm long, rachis

prickles present, 2-8-flowered. Flowers 5-

merous. Pedicels at anthesis 3-5 mm long,

same thickness throughout, prickles absent

or present; calyx tube 2-3 mm long; lobes

attenuate, 3.5-8 mm long; prickles 22-50

per flower, 3.5-8 mm long; stellate hairs very

dense, lateral rays 7-12, central ray 1-2 times

as long as laterals; simple hairs absent; Type

2 hairs absent. Corolla rotate, 9-15 mm long,

purple, inner surface glabrous; anthers 3.6-

4.3 mm long; filaments c. 1 mm long; ovary

glabrous. Functional style 8-8.5 mm long,

strongly bent near base, not touching anthers,

eccentric, glabrous; stigma expanded, darker

than style, obscurely lobed. Fruiting calyx

lobes less than half length of mature fruit.

Mature fruits 1-3 per inflorescence, oblate

or globular, 18-21 mm diameter, brown or

yellow, 2-locular; placenta stalked, anvil¬

shaped, pericarp 0.5-0.7 mm thick; pedicels

7-12 mm long. Seeds 1.8-2.5 mm long, white

or pale yellow. Fig. 3.

Additional selected specimens examined: Northern

Territory. Un-named gorge between Hugh and Spencer

Gorge, Mar 1994, Albrecht 5807 (AD, NT); Valley of

Eagles, 35 miles [58.3 km] ENE of Alice Springs, Oct

1966, Beauglehole 20671 (NT); James Range, 3 miles

[5 km] N of Hugh River, Aug 1967, Beauglehole 24592

(AD, NT); Standley Chasm, MacDonnell Ranges, Jul

1968, Orchard 818 (AD, NT); Palm Valley, Sep 1970,

Late 790 (NT); Kings Canyon, Dec 1968, Latz 321 (NSW,

NT); ‘Derry Downs’, Dulcie Range, Oct 1974, Latz 5828

(NT); Redbank Gorge, 3 Jul 1985, Leach 666 (MEL,

NT); Emily Gap, 8 miles [13 km] E of Alice Springs, Feb

1968, Nelson 1618 (MEL, NSW, NT); Stuart Highway,

10.7 km N of Aileron, Aug 2006, Bean 25455 & Albrecht

(BRI, NT).

Distribution and habitat : Solanum aridicola

is known only from the southern parts of

the Northern Territory (Map 1), although it

probably occurs in adjacent parts of Western

Australia and South Australia. It inhabits

granite or quartzite hills and ranges with

shallow sandy loam soils. Associated species

include Eucalyptus intertexta R.T.Baker,

E. terminalis F.Muell. and Grevillea

wickhamii Meisn.

Austrobaileya 8(3): 412-430 (2011)

Phenology : Flowers have been recorded

for July and August; fruits in August and

February.

Notes : Solanum aridicola is most closely

related to S. horridum as both species are more

or less prostrate, with very prickly stems and

leaves and have flowers and fruits of about

the same size and colour. Solanum aridicola

is most readily distinguished by the 3-8-

flowered inflorescences (1 or 2-flowered in S.

horridum ), and by the eccentric style in the

flower. It also differs from Solanum horridum

by the entire leaves (often shallowly lobed in

S. horridum) and the upper leaf surface with

many long-stalked stellate hairs (all ± sessile

in S. horridum).

Etymology : From the Latin aridus meaning

‘arid’ and -cola meaning ‘to inhabit’. This

refers to the arid landscape that the species

inhabits.

Solanum callosum A.R.Bean species nova

S. quadriloculato affinis sed ab eo fasciculis

pilorum stellatorum in corollam, lobis

calycum apicibus longis linearibus praeditis,

caulibus teretibus et ovario interdum 2-

loculari differens. Typus: Queensland.

Gregory South District: 20.2 km WSW of

Eromanga, on Cooper Development Road, 27

August 2010, A.R. Bean 30027 (holo: BRI [1

sheet + spirit]; iso: NSW).

S. ellipticum R.Br., and. non Bean (2004),

pro parte.

Illustrations : Cunningham et al. (1981: 592),

as S. quadriloculatum ; Brooke & McGarva

(1998: 112), as S. ellipticum.

Rhizomatous perennial shrub 0.15-0.6

m high. Stems initially sprawling, then

spreading to erect. Adult branchlets yellow,

rusty or brown; prickles 8-70 per cm, straight,

acicular, 2-7 mm long, 14—18 times longer

than wide, glabrous; stellate hairs very dense,

0.6-1.1 mm diameter, stalks 0-0.2 mm long,

lateral rays 7-8, porrect or ascending, central

ray 0.7-1.4 times as long as laterals; Type 2

hairs absent. Adult leaves ovate or broadly

ovate, entire, lamina 4.9-9 cm long, 3-4.8

cm wide, 1.5-2.4 times longer than broad,

apex obtuse or acute, base cuneate or obtuse,

oblique part 1-3.5 mm long, obliqueness

Bean, Solanum ellipticum group

417

Fig. 3. Solanum aridicola. A. flowering branchlet * 0.8. B. flower with corolla and calyx lobes removed x 4. C. pedicel,

calyx and style x 4. D. corolla x 2. E. ovary and style x 4. F. cross-section of mature fruit x 2. All from Bean 25449 &

Albrecht (BRI).

index 1-4%; petioles 1.2-3 cm long, 17-33%

length of lamina, prickles present. Upper leaf

surface grey; prickles absent or with 1-4 on

midvein only, straight, acicular, 2-4.5 mm

long; stellate hairs distributed throughout,

dense or very dense; 0.05-0.2 mm apart,

0.5-0.7 mm across, stalks 0-0.15 mm long,

lateral rays 6-8, porrect or ascending, central

ray 0.7-1.3 times as long as laterals; simple

hairs absent; Type 2 hairs absent. Lower leaf

surface white or grey; prickles 3-10, present

on midvein only or present on midvein and

lateral veins, straight, acicular; stellate hairs

very dense, 0.05-0.1 mm apart, 0.6-0.9 mm

diameter, stalks 0-0.3 mm long, lateral rays

7-8, porrect, central ray 0.7-1.3 times as long

as laterals; simple hairs absent; Type 2 hairs

absent. Inflorescence supra-axillary, cymose

(pseudo-racemose), common peduncle 15-

41 mm long, rachis prickles present, 7-11-

418

flowered, with all flowers bisexual. Flowers

5-merous. Pedicels at anthesis 5-11 mm long,

same thickness throughout, prickles absent

or present. Calyx tube at anthesis 2.5-4 mm

long; lobes deflate or attenuate, 1.5-7.5 mm

long; prickles 14-40 per flower, 1-3 mm long;

stellate hairs very dense, white or purplish,

0.4-0.8 mm across, stalks 0.1-0.5 mm long,

lateral rays 7-8, central ray 0.8-1.4 times as

long as laterals; simple hairs absent; Type 2

hairs absent. Corolla shallowly lobed, 9-14

mm long, purple, inner surface densely

stellate-hairy at distal end; anthers 4.4-6.1

mm long; ovary glabrous; style 7.5-8.5 mm

long, protruding between anthers, glabrous;

stigma unexpanded, same colour as style,

obscurely lobed. Fruiting calyx lobes less

than or more than half length of mature fruit,

prickles 2-4 mm long. Mature fruits 2-4 per

inflorescence, globular, 18-20 mm diameter,

green, 2-locular or 4-locular, placenta stalked,

anvil-shaped, interior moist but not juicy,

pericarp 3-3.3 mm thick. Pedicels at fruiting

stage 8-21 mm long. Seeds pale yellow, 2.5-

2.7 mm long. Fig. 4.

Additional specimens examined: Queensland. Mitchell

District: 42.7 km along Silsoe Road, W of Longreach,

May 2004, Bean 22213 (BRI); Longreach, Thompson

River, Apr 1963, Gittins 670 (BRI); Milo Station, Old

Gooyea Outstation ruins, near Powell Creek, NNW of

Adavale, Aug 2009, Forster PIF35347 & Thomas (BRI,

NSW); ‘Springdale’, N of Aramac, Jul 2000, Fensham

3947 (BRf); road from Adavale to Windorah, 64 miles

[102 km] WNW of Adavale, Jun 1967, Gittins 1232

(BRI); Idalia N.P., SW of Blackall, Jun 1990, Rogers s.n.

(BRI [AQ624125]). Gregory North District: Kangaroo

Mountains, between Bedourie and Windorah, Jun 1978,

Althofer 8363 (BRI). Gregory South District: 3.7 km

From Poison Tank Bore, GiGi Paddock, Mt Margaret

Station, 30 km SW of Eromanga, Oct 1995, Stol &

Landsberg QSL382 (CANB); near Jackson, south-west

Qld, Jul 2003, Hines s.n. (BRI [AQ779478]); c. 40 miles

[64 km] W of Windorah, Jun 1949, Everist 3894 (BRI);

19.1 km W of central Tank turnoff, Thargomindah

- Nockatunga Road, Sep 1990, Prendergast HDP346

(BRI). Warrego District: Eulo sports ground, 0.8 km

S of Eulo, Aug 2006, Bean 25634 (BRI); 42.3km from

Charleville, towards Quilpie, Aug 2010, Bean 29925

(BRI, MEL); ‘Gilruth Plains’, E of Cunnamulla, May

1939, Blake 14034 (BRI); Currawinya N.P, 4 km WNW

of Gidgee Bore, Mar 1997, Forster PIF20444 & Watson

(BRI, DNA, MEL); c. 27 km NE of Thargomindah, on

road to Quilpie, Sep 1973, Henderson H2083 & Boyland

(BRI); ‘Yarronvale’, 66 miles [106 km] from Charleville

on Quilpie road, Sep 1963, Everist 7520 (BRI). Maranoa

District: ‘Murra Murra’, River paddock, at Brumby

Lake, SW of Bollon, Feb 2007, Eddie CPE1091 (BRI).

Austrobaileya 8(3): 412-430 (2011)

New South Wales. North Far Western Plains: Ursino -

Yamba Station road, 30 miles [48 km] W of Wanaaring,

Oct 1963, Constable 4583 (BRI, NSW); 14 miles [23 km]

W of Wanaaring, Dec 1969, Martensz 260 (AD, CANB,

NSW); Peery Station, Apr 1963, Bailey 2862 (CANB).

North Western Plains: 65.7 km from Bourke - Cobar

Road, towards Coolabah, Mar 2008, Bean 27637 (BRI);

0.1 km W of Condobolin - Cobar Road, c. 40 km N of

Condobolin, Mar 2008, Bean 27694 (BRI); Coolabah,

Apr 1900, Helms s.n. (NSW).

Distribution and habitat : Solanum callosum

is widespread from north of Muttaburra in

central Queensland to Condobolin in central

New South Wales (Map 2). It grows on plains

and low stony hills on sandy loam to clay

loam soils. Associated tree species include

mulga {Acacia aneura F.Muell. ex Benth.),

A. excelsa Benth., Eucalyptus terminalis

F.Muell. and E. populnea F.Muell.

Phenology : Flowers are recorded for all

months of the year. Fruits are recorded

between February and September.

Notes: This species is most closely allied to

Solanum quadriloculatum ; however, there

a number of differences; S. callosum has a

cluster of stellate hairs on the distal portion

of the inner surface of the corolla, with

scattered stellate hairs extending down the

midvein for about half the corolla length (in

S. quadriloculatum the corolla is glabrous,

except for two or three collections which bear

simple hairs at the distal end of the lobes);

calyx lobes narrowly deflate with long linear

apices (broadly deflate, with or without short

linear apices for S. quadriloculatum ); stems

terete (usually ridged for S. quadriloculatum );

no stellate hairs on prickles (stellate hairs on

at least some prickles for S. quadriloculatum );

ovary 2- or 4-locular (consistently 4-locular

for S. quadriloculatum) ; fruits 18-20 mm

diameter (14-17 mm for S. quadriloculatum).

Etymology: The species epithet is from the

Latin callosus meaning tough, hard, or thick-

skinned. This is given in reference to the

fruits that are hard when dry, and for the very

thick pericarp.

Bean, Solanum ellipticum group

419

Fig. 4. Solanum callosum. A. flowering branchlet x 0.6. B. oblique view of an open flower x 2. C. base of flower

showing calyx and corolla x 2. D. mature fruits attached to rachis x 1.5. E. cross-section of a 4-locular fruit x 2. F.

cross-section of a 2-locular fruit x 2. A from Gittins 670 (BRI); B,C from Bean 29925 (BRI); D,F from Bean 30027

(BRI); E from Hines s.n. (BRI [AQ779478]).

Solanum cleistogamum Symon, Trans.

& Proc. Roy. Soc. South Australia 95: 227

(1971). Type: Western Australia, c. 32 km

N of Onslow, 1 July 1967, D.E. Symon 5418

(holo: PERTH; iso: AD, CANB, K, L).

S. petrophilum var. pedicellatum Ewart &

Davies, FI. Northern Territory 243 (1917).

Type: Northern Territory. Lat. 19° long.

132°, 4 July 1911, G.F. Hill 435 (holo: MEL

[2 sheets]).

420

Illustrations : Cunningham et al. (1981: 592);

Symon(1981: 194)

Herbaceous resprouter, 0.15-0.25 m high.

Stems initially ascending, then prostrate

and trailing. Adult branchlets white to

rusty-brown; prickles 7-18 per cm, straight,

acicular, 1-6 mm long, 15-20 times longer

than wide, glabrous; stellate hairs dense

or very dense, lateral rays 8-15, porrect,

ascending or multiradiate, central ray 1-

2.5 times as long as laterals. Adult leaves

lanceolate or ovate, entire or with 2 or 3

shallow obtuse lobes on each side, lobing

index 1-1.1; lamina 2.8-47 cm long, 1.2-3.6

cm wide, 1.2-2.4 times longer than broad,

apex obtuse, base obtuse or cordate, oblique

part 0-4.5 mm long, obliqueness index

0-10%; petioles 1.4-3.1 cm long, 40-70%

length of lamina, prickles present. Upper leaf

surface grey-green, prickles absent; stellate

hairs moderately dense or dense, stalks 0-0.1

mm long, lateral rays 6-8, porrect, central

ray 1.1-2 times as long as laterals. Lower

leaf surface green or greenish-white, prickles

absent or with 1-10 on midvein only, straight,

acicular. Lower leaf surface stellate hairs

moderately dense or dense, stalks 0-0.1 mm

long, lateral rays 7-8, porrect; central ray 1.2-

2 times as long as laterals, sometimes gland-

tipped. Inflorescence supra-axillary, cymose

(pseudo-racemose), common peduncle

8- 21 mm long, rachis prickles present, 1-5-

flowered, with all flowers bisexual. Flowers

5-merous. Pedicels at anthesis 7-14 mm long,

prickles present. Calyx tube at anthesis 2-3.5

mm long; lobes deflate or rostrate, 2-4 mm

long; prickles present, 10-25 per flower,

1.5-3 mm long; stellate hairs dense, lateral

rays 6-8; central ray 1.2-2.2 times as long

as laterals, sometimes gland-tipped. Corolla

rotate, 6-9 mm long, white or mauve, inner

surface glabrous; anthers 2-2.8 mm long;

filaments 1.2-1.3 mm long; ovary glabrous;

style 5-6 mm long, erect near base, then

bent, displacing one anther, glabrous; stigma

slightly expanded, obscurely lobed. Fruiting

calyx lobes less than or more than half length

of mature fruit, prickles 1.5-5 mm long.

Mature fruits 1-3 per inflorescence, globular,

9- 12 mm diameter, yellow, 2-locular; placenta

sessile, semi-circular; interior juicy, pericarp

Austrobaileya 8(3): 412-430 (2011)

0.4-0.8 mm thick; pedicels 15-23 mm long.

Seeds 2-2.5 mm long, white or pale yellow.

Additional selected specimens examined: Western

Australia. Horridum Hill, near highway, c. 50 km E

of Port Hedland, Apr 2006, Bean 25014 & Lepschi

(BRI, CANB, PERTH). Northern Territory. Track

to Lake Lewis, S of Tilmouth Bore, c. 160 km NW

of Alice Springs, Aug 2006, Bean 25472 & Albrecht

(BM, BRI, CANB, MEL, NSW, NT, NY); North-west

Simpson Desert, Sep 1973, Latz 4658 (AD, BRI, NT);

Curlew Waterhole, Lander River, 70 miles [113 km] NW

of Willowra Homestead, Central Australia, Jul 1958,

Chippendale 4799 (BRI, NT); W margin of Lake Ruth, c.

5 km SE of Mongrel Downs Homestead, Tanami Desert,

Aug 1978, Donner 6283 (AD, BRI). Queensland. Burke

District: Roxmere Station, S of Cloncurry, Apr 2003,

Booth 3168 & Kelman (AD, BRI, NSW). South Kennedy

District: Lake Buchanan, May 1980, Hyland 10525

(BRI, CNS). Warrego District: Thargomindah, Jun

1936, Blake 11781 (BRI); 10.9 km from Adavale towards

Blackall at tank, Oct 1983, Canning 6206 (BRI, CANB).

Maranoa District: Karoola Road, S of Dirranbandi,

10 km N of state border, Sep 2004, Bean 22847 (BRI).

New South Wales. 3.6 km NE of Cumborah, towards

Lightning Ridge, Sep 2004, Bean 22893 (BRI); Fowlers

Gap, N of Broken Hill, Oct 1975, Jacobs 2337 (BRI,

NSW); ‘Tundulya’, c. 25 miles [40 km] SE of Louth, Apr

1967, Moore 4949 (CANB). South Australia. E side of

Lincoln Highway, 17 km N of Whyalla, Oct 1998, Jobson

5736 & Downs (BRI, NSW).

Distribution and habitat: Solarium

cleistogamum is widespread in mainland

Australia south of about latitude 19°, from the

west coast of Western Australia to the slopes

of the Great Divide in New South Wales and

Queensland. In South Australia, it occurs as

far south as Spencer Gulf.

It grows on a range of habitats but prefers

sandy or sandy-loam soils, often on rather

low-lying terrain, but also sometimes on steep

hillsides.

Phenology: Flowers and fruits may be found

throughout the year.

Notes: This species was previously included

in the synonymy of Solarium ellipticum (Bean

2004). I now recognise that this was incorrect,

and that the two are distinct. While the two

species are closely related and rather difficult

to distinguish in the herbarium, the relatively

small tasty yellow fruits, the small corolla, and

the ± cordate leaf bases of S. cleistogamum

readily distinguish it.

Bean, Solarium ellipticum group

Solanum ellipticum R.Br., Prodr. 446

(1810); S. ellipticum var. ellipticum , Benth.,

FI. Austral. 4: 464 (1868); S. ellipticum f.

ellipticum , Wawra, Itin. Princ. S. Coburgi

100 (1883); S. ellipticum var. typicum Domin,

Biblioth. Bot. 89: 588 (1929), nom. inval.

Type: [Queensland. Port Curtis District:]

‘Broadsound’, 25 September 1802, R.

Brown [Bennett Number 2674] (lecto: BM

[000596888]; isolecto: MPU, fide Symon

[1981: 188]).

Herbaceous resprouter, or rhizomatous

perennial shrub 0.1-0.25 m high. Stems

initially ascending, then prostrate and

trailing. Adult branchlets grey, purple, yellow

or brown; prickles 1-10 per cm, straight,

acicular, 1-8 mm long, 8-14 times longer

than wide, glabrous or with dense stellate

hairs on lower part; stellate hairs dense to

very dense, 0.5-0.8 mm diameter, stalks

0-0.1 mm long, lateral rays 7-10, lateral rays

porrect or ascending, central ray 0.5-1.2 times

as long as laterals. Adult leaves elliptical or

ovate, entire; lamina 3.5-14 cm long, 1.5-5.2

cm wide, 1.7-2.9 times longer than broad,

apex obtuse or acute, base cuneate or obtuse,

oblique part 0-9 mm long, obliqueness index

0-6%; petioles 0.8-4 cm long, 20-45% length

of lamina, prickles absent or present. Upper

leaf surface green or grey-green, prickles

2-20, present on midvein only or on midvein

and lateral veins, straight, acicular, 1-7 mm

long; stellate hairs distributed throughout,

sparse to dense, 0.1-0.5 mm apart, 0.4-0.7

mm across, stalks 0-0.1 mm long, lateral rays

7-8, porrect, central ray 0.5-1 times as long

as laterals; simple hairs absent; Type 2 hairs

absent. Lower leaf surface greenish-white to

grey or yellowish; prickles 0-20, absent or

present on midvein only or on midvein and

lateral veins, straight, acicular; stellate hairs

moderately dense to very dense, 0.05-0.3

mm apart, 0.5-0.8 mm diameter, stalks 0-0.2

mm long, lateral rays 7-8, porrect, central ray

0.5-1.2 times as long as laterals; simple hairs

absent; Type 2 hairs absent. Inflorescence

supra-axillary, solitary or cymose (pseudo-

racemose), common peduncle 0-6 mm long,

rachis prickles present, 1-5-flowered, with

some bisexual and some male flowers. Flowers

5-merous. Pedicels at anthesis 4-11 mm long,

421

0.7-0.9 mm thick at mid-point, prickles

absent or present. Calyx tube at anthesis 1.5-

3 mm long; lobes deflate or attenuate, 2-6

mm long; prickles 5-50 per flower, 1-5 mm

long; stellate hairs moderately dense to very

dense, white to yellow or rusty, 0.4-0.8 mm

across, stalks 0-0.3 mm long, lateral rays 7-

8, central ray 0.8-1.3 times as long as laterals;

simple hairs absent; Type 2 hairs absent.

Corolla shallowly or deeply lobed, 7-12 mm

long, mauve or purple, inner surface with a

few stellate hairs near lobe apices. Anthers

4-5.5 mm long. Ovary glabrous or with Type

2 hairs only; functional style 5.5-10 mm long,

protruding between anthers, glabrous or with

Type 2 hairs only; stigma unexpanded, same

colour as style. Fruiting calyx lobes less than

or more than half length of mature fruit,

prickles 1-5 mm long. Mature fruits 1-2 per

inflorescence, globular, 13-17 mm diameter,

yellowish-green, 2-locular; placenta stalked,

anvil-shaped; interior moist but not juicy,

pericarp 0.6-1 mm thick; pedicels 7—19 mm

long, 0.8-2.1 mm thick at mid-point. Seeds

2.2-27 mm long, pale yellow. Fig. 1.

Additional selected specimens examined : Queensland.

Leichhardt District: Slopes of Marling Spike, Homevale

N.P., NNW of Nebo, Dec 2010, Bean 30650 et al. (BRI).

Port Curtis District: Orange Creek, c. 20 miles [32

km] NW of Biloela, Jun 1959, Johnson 857 (BRI); 4

km WSW of St Lawrence, Apr 2000, Bean 16250 (BRI,

MEL). Burnett District: ‘Neaavie’, NE of homestead

towards Barambah Creek, Aug 1996, Grimshaw PG2536

& Turpin (BRI); c. 6 km NE of Allies Creek and 75 km

SW of ‘Weir Weir’, Jul 1998, Pollock ABP617 & Dean

(BRI). Wide Bay District: Bruce Highway, between

Gin Gin and Miriam Vale, Jan 2006, Bohs 3540 & Bean

(BRI); Bingera, Dec 1896, Bailey s.n. (BRI [AQ39062]).

Darling Downs District: 8 km WSW of Oakey, Apr

1994, Fensham 1444 (BRI); 14 km N of Goondiwindi,

towards Moonie, Feb 1996, Bean 9907 (BRI, MEL,

NSW). Moreton District: Gatton, Nov 1916, Bick s.n.

(BRI [AQ39059]); 2 km ESE of Laidley, Jun 2002, Bean

19062 (BRI). New South Wales. North West Plains:

4 km SSW of ‘The Glen’ Homestead, c. 70 km W of

Moree, Jan 1999, Wannan 1036 et al (AD, BRI, NSW).

Distribution and habitat : The limits of

distribution of Solanum ellipticum sens. str.

are not yet known with certainty, but it is

found in the Queensland Pastoral districts of

Moreton, Darling Downs, Wide Bay, Burnett,

Leichhardt and Port Curtis. It also occurs on

the north-western plains of New South Wales

near Moree.

422

Phenology : Flowers and fruits can be found

throughout the year.

Notes : Solanum ellipticum is a prostrate to

semi-erect plant with trailing stems (Fig. 1).

It has a small inflorescence with 1-5 flowers,

and some of these are functionally male

(styles very short and non-functional). It is

perhaps the only species of the group in which

the flowers are not all bisexual. The common

peduncle is usually absent, although it can be

up to 6 mm long. The number of fruits is one

or two.

Solanum ellipticum var. chillagoense

Domin, S. ellipticum f. albiflora Domin, and

S. sp. (Newcastle Range D.E. Symon 4907)

were included by Bean (2004) in the synonymy

of S. ellipticum. The author now recognises

that this is incorrect; however, further study

is required to accurately place these names

beyond synonymy with S. ellipticum.

Solanum emmottii A.R.Bean species nova

S. lithophilo affinis sed ab eo corolla minore,

calycis aculeis paucioribus brevioribusque

et pilis stellatis omnibus stipitibus quam 0.2

millimetris longitudine brevioribus praeditis

differens. Typus: Queensland. Gregory

North District: Near homestead, Ethabuka

Reserve, 8 November 2010, A. J. Emmott B690

(holo: BRI [1 sheet + spirit]; iso: AD, NT, NY,

W, distribuendi ).

Rhizomatous perennial shrub 0.2-0.5 m

high. Stems initially sprawling, then more

or less erect. Adult branchlets white, grey

or yellowish; prickles 1-28 per cm, straight,

acicular, 1-5 mm long, 10-14 times longer

than wide, glabrous; stellate hairs very dense,

0.4-0.8 mm diameter, stalks 0-0.2 mm

long, lateral rays 7-13, porrect, ascending

or multiradiate, central ray 0.8-1.3 times as

long as laterals; Type 2 hairs absent. Adult

leaves ovate, entire; lamina 3.5-8.8 cm long,

1.4-3.8 cm wide, 1.7-2.9 times longer than

broad, apex obtuse or acute, base cuneate,

obtuse or cordate, oblique part 0-4 mm long,

obliqueness index 0-7%; petioles 0.6-4.2

cm long, 18-48% length of lamina, prickles

absent or present. Upper leaf surface grey-

green or grey, prickles absent; stellate hairs

dense to very dense, 0.05-0.2 mm apart, 0.4-

0.7 mm across, stalks 0-0.1 mm long, lateral

Austrobaileya 8(3): 412-430 (2011)

rays 6-10, porrect, central ray 0.7-1.2 times

as long as laterals; simple hairs absent; Type 2

hairs absent. Lower leaf surface white or grey;

prickles absent or present on midvein only,

0-2, straight, acicular; stellate hairs dense to

very dense, 0.05-0.1 mm apart, 0.4-0.6 mm

diameter, stalks 0-0.3 mm long, lateral rays

7-11, porrect or ascending, central ray 0.7-1.2

times as long as laterals; simple hairs absent;

Type 2 hairs absent. Inflorescence supra-

axillary, cymose (pseudo-racemose), common

peduncle 15-50 mm long, rachis prickles

absent or present, 2-6-flowered, with all

flowers bisexual. Flowers 5-merous; pedicels

at anthesis 4-15 mm long, same thickness

throughout, prickles absent or present; calyx

tube at anthesis 2.5-4 mm long; lobes deflate

or attenuate, 2-7 mm long; prickles 1-25 per

flower, 0.5-2 mm long; stellate hairs very

dense, yellow or white, 0.4-0.8 mm across,

stalks 0-0.3 mm long, lateral rays 8-10,

central ray 0.7-1.3 times as long as laterals;

simple hairs absent; Type 2 hairs absent.

Corolla rotate or shallowly lobed, 9-13 mm

long, purple, inner surface sparsely stellate-

hairy; anthers 4.2-5.5 mm long; filaments

1.2-1.6 mm long. Ovary glabrous; functional

style 8-9.5 mm long, strongly bent near base,

not touching anthers, eccentric, glabrous or

rarely with stellate hairs; stigma expanded,

darker than style, obscurely lobed. Fruiting

calyx lobes less than or more than half

length of mature fruit. Mature fruits 1-3 per

inflorescence, globular, c. 13 mm diameter,

colour unknown, 2-locular; placenta sessile,

semicircular; interior moist but not juicy,

pericarp 0.3-0.4 mm thick; pedicels 8-13 mm

long, 1.2-1.5 mm thick at mid-point. Seeds

2.5-3.3 mm long, pale yellow. Fig. 5.

Additional specimens examined : Northern Territory.

c. 5 km east of Lake Caroline, northern Simpson

Desert, Jul 2007, Albrecht 12300 & Duguid (BRI,

NT); c. 1 km west of Lake Caroline, northern Simpson

Desert, Jul 2007, Albrecht 12340 & Duguid (BRI, NT);

Near Hale River, Simpson Desert, Sep 1955, Perry

5421 (BRI, CANB); 12 miles [20 km] NW of Lucy

Creek Homestead, Jul 1957, Chippendale 3523 (BRI).

Queensland. Mitchell District: Valetta Bore paddock,

‘Valetta’, near ‘Noonbah’, c. 170 km SW of Longreach,

May 2010, Bean 29753 & Emmott (AD, BM, BRI, NSW,

NT); south boundary of Valetta Bore paddock, ‘Valetta’,

c. 180km SW of Longreach, May 2010, Bean 29757 &

Emmott (BRI, MEL). Gregory North District: 10 km

NNW of Bedourie, (1.6 km W of Duck Hole), Kamaran

Downs Station, Jun 1995, Edmunds 23 (BRI). Gregory

Bean, Solanum ellipticum group

423

Fig. 5. Solanum emmottii. A. flowering branchlet x 1.5. B. lateral view of flower x 4. c. stamens, style and ovary x 4.

D. immature fruit x 2. All from Bean 29753 & Emmott (BRI).

South District: Mt Howitt Station, 80 miles [133 km] W

of Eromanga, Jul 1936, Blake 11926 (BRI); 10 miles [16

km] W of Betoota, Jul 1936, Blake 12184 (BRI); ‘Kyra’,

40 miles [67 km] W of Quilpie, Jul 1965, Emmerson s.n.

(BRI [AQ39068]); 35 km W of Betoota on Birdsville

road, Aug 1979, Williams 78187 (BRI). South Australia.

54 km by road S of Birdsville on the Birdsville Track to

Marree, Mar 2001, Thomas 2049 & Fechner (BRI).

Distribution andhabitat: Solanum emmottii is

known from the far south-west of Queensland,

and adjacent areas of Northern Territory and

South Australia (Map 3). It grows almost

exclusively on or adjacent to sand dunes,

in red to orange sandy soil, associated with

shrubs such as Crotalaria eremaea F.Muell.

However, at the most north-easterly location,

it occurs on the slopes of gravelly hills in

brown or yellow loam, and in association with

Atalaya hemiglauca (F.Muell.) F.Muell. ex

Benth., Acacia ensifolia Pedley, Eremophila

mitchellii Benth. and Triodia longiceps

J.M.Black.

Phenology : Flowers have been recorded from

March to November; a potentially mature

fruit was collected in July.

Notes : Solanum emmottii differs from

S. lithophilum by the non-floccose branchlet

indumentum and the stellate hairs stalks less

than 0.2 mm (floccose branchlet indumentum

with stalks 0-2 mm long for S. lithophilum );

424

stellate hairs with shorter central rays; shorter

calyx lobes; calyx prickles 1-25, each 0.5-2

mm long (25-50 calyx prickles, each 2-4 mm

long for S. lithophilum ); corolla 9-13 mm long

(16-20 mm for S. lithophilum ); the presence

of stellate hairs on the inner corolla surface

(glabrous for S. lithophilum ); and the smaller

fruits.

Edmunds 23 (cited above) differs from

the other cited specimens by the presence of

numerous stellate hairs on the style, but is in

other regards quite typical.

Conservation status : This species appears

to be fairly common and is not currently

considered to be under threat. A conservation

status of ‘Least Concern’ is recommended.

Etymology : The species is named for Angus

James Emmott (1962-), grazier, accomplished

naturalist, and dedicated advocate for the

preservation of natural ecosystems in arid

Australia.

Solanum lithophilum F.Muell., Linnaea

25: 434 (1853). Type: South Australia. Near

Cudnaka [Kanyaka Homestead], Flinders

Ranges, October 1851, F. Mueller s.n. (lecto:

MEF11873 ,fide Symon [1981: 188]).

S. ellipticum var. mollibaccalis J.M.Black,

Trans. & Proc. Roy. Soc. South Australia 52:

227 (1928). Type: South Australia. Finniss

Springs near Fake Eyre, s.dat., F.D.Warren

s.n. (syn: AD, fide Symon [1981: 188], n.v).

S. ellipticum var. Foot hills (G.J.Leach 1145)-,

Kerrigan & Albrecht (2007), Albrecht et al.

(2007: 176).

S. ellipticum R.Br. auct. non Symon (1981),

pro parte.

Herbaceous resprouter, 0.3-0.45 m high.

Stems initially sprawling, then more or less

erect. Adult branchlets grey, yellow or rusty;

prickles 9-40 per cm, straight, acicular,

2-7 mm long, 13-18 times longer than wide,

glabrous; stellate hairs very dense, stalks

0-2 mm long, lateral rays 8—13, porrect or

multiradiate, central ray 1.1-2.2 times as long

as laterals. Adult leaves elliptical or ovate,

entire; lamina 3.6-10.1 cm long, 1.5-5.3 cm

wide, 1.9-3.8 times longer than broad, apex

acute, base cuneate or obtuse, oblique part 0-

Austrobaileya 8(3): 412-430 (2011)

6 mm long, obliqueness index 0-7%; petioles

1.3-6 cm long, 33-60% length of lamina,

prickles present. Upper leaf surface grey-

green or grey; prickles absent or present on

midvein only, 0-2, straight, acicular, 3-7 mm

long; stellate hairs dense, stalks 0-2 mm long,

lateral rays 7-10, porrect or ascending, central

ray 1.2-2 times as long as laterals. Fower leaf

surface white, grey or silvery; prickles absent

or present on midvein only, 0-4, straight,

acicular; stellate hairs very dense, lateral

rays 8-15, porrect or multiradiate, central ray

1.2-2 times as long as laterals. Inflorescence

supra-axillary, cymose (pseudo-racemose),

common peduncle 20-52 mm long, rachis

prickles present, 3-7-flowered, with all

flowers bisexual. Flowers 5-merous. Pedicels

at anthesis 7-16 mm long, same thickness

throughout, prickles absent or present. Calyx

tube at anthesis 3-4 mm long; lobes attenuate,

5-11 mm long; prickles present, 25-50 per

flower, 2-4 mm long; stellate hairs very

dense, yellowish-white or purple, lateral rays

8-13, central ray 1-2 times as long as laterals.

Corolla rotate, 14-20 mm long, purple, inner

surface glabrous. Anthers 4.7-6 mm long.

Filaments 1-1.2 mm long. Ovary glabrous;

style 9-13 mm long, strongly bent near base,

not touching anthers, eccentric, glabrous;

stigma expanded, darker than style, obscurely

lobed. Fruiting calyx lobes less than or more

than half length of mature fruit, prickles 2.5-4

mm long. Mature fruits 1-2 per inflorescence,

globular or ellipsoidal, 18-21 mm diameter,

purplish-green or almost white, 2-locular;

placenta stalked, anvil-shaped; interior

juicy, succulent; pericarp 0.3-0.8 mm thick;

pedicels 12-25 mm long. Seeds 2.8-3.2 mm

long, pale yellow or white. Figs. 2 & 6.

Additional selected specimens examined: Western

Australia. 100 miles [161 km] N ofEucla, s.dat., Batts.n.

(MEL). Northern Territory. Stuart Highway, 10.7 km N

of Aileron, Aug 2006, Bean 25457 & Albrecht (AD, BRI,

NT); track to Lake Lewis, S of Tilmouth Bore, c. 160 km

NW of Alice Springs, Aug 2006, Bean 25473 & Albrecht

(BRI, NT); Western side of Kings Canyon Range, 1 km

south of Carmichael Crag, Aug 1986, Menkhorst s.n.

(MEL); Orange Creek, 1 km S of Homestead on Stuart

Highway, 24 km S of Alice Springs on Adelaide Road,

Jun 1968, Must 17 (MEL); 4 miles [6 km] W of Central Mt

Wedge Homestead, Jun 1955, Chippendale 1283 (BRI);

Uluru N.P.: Kata Tjuta, on the Docker River Road, 35.7

km WNW of Ranger Station, May 1988, Lazarides &

Palmer 108 (CANB); Beside Todd River, N of Schwarz

Bean, Solatium ellipticum group

Crescent, Alice Springs, Aug 2006, Bean 25494 (BRI,

CANB, NT); N’Dahla Gorge, Ross River area, E

MacDonnell Ranges, Jun 1974, Williss.n. (MEL); S bank

of upper Finke River opposite Mt Sonder, Jul 1966, Willis

& Morrison s.n. (MEL). Queensland. Gregory South

District: on track 13.4 km due NW of ‘Old Karmona’

cottage, Aug 2008, Turpin GPT1280 & May (BRI, CNS);

on track 26.9 km due NW of Orientos Station, Aug 2008,

Turpin GPT1279 & May (BRI, CNS). New South Wales.

Tarella, Aug 1887, Baeuerlen 118 (MEL); Golden Gully

mining area adjacent to the Deadhorse Gully camping

area, 1 km N of Tibooburra in Sturt N.R, Sep 1989,

Coveny 13598 et al. (AD, BRI, MO, NSW); ridge at end

of Mootwingee Gorge Road, Mootwingee N.R, Aug

1989, Porteners 33 & Plaza (NSW, NT); 2 km W of Big

Wallaby tank, 5 km NNW of Koonenberry Mountain,

Sep 1971, De Nardi 847 (NSW); Silverton near Barrier

Range, Nov 1884, Harris s.n. (MEL); Mt Koonenberry,

Dec 1860 , Beckler s.n. (MEL). South Australia. nearMt

Everard, 1882, Giles s.n. (MEL); Fogarty’s claypan. Mar

1984, Jackson 5255 (MEL); Strangways Springs, 1895,

Gratwick 16 (MEL); NW of Ooldea, s.dat., Tietkins s.n.

(MEL); Brachina Creek, Jun 1960, Filson 2115 (MEL); 2

miles [3.3 km] S of Emu, Aug 1956, Forde 389 (MEL);

Innamincka Homestead, Jun 1980, Lay 1493 (MEL);

Lake Torrens Basin, east Mernmerna, c. 150 km NE of

Port Augusta, on road to Marree, Oct 1968, Weber 729

(MEL).

Distribution and habitat : Solanum

lithophilum is widespread in South

Australia (except in the south-east) and

the southern one-third of the Northern

Territory. It also extends to far eastern

parts of Western Australia and north¬

western New South Wales and it has

recently been recorded from the extreme

south-west of Queensland (Map 3). In

South Australia, it is frequently recorded

from rocky hill slopes, but in the Northern

Territory it often grows on rather sandy

colluvial and alluvial soils, with only

limited occurrences on steep hillsides.

Phenology : Flowers are recorded from all

months of the year; fruits are recorded from

March to September.

Affinities: Solanum lithophilum may be readily

distinguished from S. ellipticum. Solanum

lithophilum is a multi-stemmed plant to 45

cm high; the stems are at first sprawling, but

finally erect and have a floccose tomentum;

the lower leaves have very long petioles, the

inflorescences are well displayed and the very

showy flowers (corolla 14-20 mm long) have

a prominently eccentric style. The fresh fruits

are large, 18-21 mm in diameter, and often

425

bone-white when mature, at least at the distal

end.

By contrast, Solanum ellipticum rarely

exceeds 20 cm in height, the stems are

ascending near the base, but then prostrate

or procumbent, the tomentum is not floccose,

all leaves have petioles of a similar length,

the inflorescences are not well displayed and

the flowers could not be described as showy

(corolla 7-12 mm long). The style is erect

rather than eccentric, protruding between

the anthers. The fruits are smaller, 13-17

mm across, and green to yellow-green at

maturity.

There are further differences at the

microscopic level. Some stellate hairs on

the branchlets of Solanum lithophilum are

multiradiate with 10-13 lateral rays, and

the central ray on all plant parts is 1.1-2.2

times longer than the laterals. The stigma

of Solanum lithophilum is conspicuously

swollen, and green in colour (cf. S. ellipticum

where the stigma is scarcely broader than the

style, and is the same colour as it).

Notes: Well-developed plants of Solanum

lithophilum form a small multi-stemmed

bush with numerous, large, well-displayed

purple flowers (Fig. 2). Such plants are very

attractive and have considerable horticultural

potential.

Solanum unispinum A.R.Bean species

nova S. cleistogamo affinis sed ab eo corolla

profunde lobata, antheribus longioribus, radio

centrali pilorum stellatorum breviore, calyce

aculeis pauculis vestito et habitu fruticoso

differens. Typus: Queensland. Gregory

North District: Winton - Jundah Road, 2.7

km N of Mayne River crossing, 25 May 2004,

A.R. Bean 22512 (holo: BRI; iso: CANB,

NSW, distribuendi).

Herbaceous resprouter or rhizomatous

perennial shrub, to 0.3 m high. Stems initially

sprawling, then more or less erect. Adult

branchlets purplish, yellow or rusty. Adult

branchlet prickles 1-4 per cm, straight,

acicular, 3-8 mm long, 12-16 times longer

than wide, glabrous; stellate hairs very dense,

0.4-0.7 mm diameter, stalks 0-0.1 mm

long; lateral rays 8-10, porrect; central ray

426

Austrobaileya 8(3): 412-430 (2011)

Fig. 6. Solatium lithophilum. A. flowering branchlet x 0.8. B. leaf from lower part of plant * 0.8. C. oblique view of an

open flower x 2. D. lateral view of a flower bud x 2. E. mature fruit with attached calyx x 1.5. F. cross-section of mature

fruit x 2. A, E, F from Bean 25457 & Albrecht (BR1); B-D from Bean 25494 (BRI).

427

Bean, Solanum ellipticum group

0.4-0.8 times as long as laterals, not gland-

tipped. Adult leaves ovate or broadly ovate,

entire; lamina 2.2-3.5 cm long, 1.4-2.1 cm

wide, 1.6-1.7 times longer than broad, apex

obtuse or acute, base obtuse, oblique part

0-2 mm long, obliqueness index 0-9%.

Petioles 1.4-2.7 cm long, 58-77% length of

lamina, prickles present. Upper leaf surface

grey-green, prickles absent; stellate hairs

distributed throughout, dense, 0.15-0.2 mm

apart, 0.4-0.5 mm across, stalks 0-0.1 mm

long, lateral rays 8, central ray 0.6-0.9 times

as long as laterals, not gland-tipped; simple

hairs absent; Type 2 hairs absent. Lower

leaf surface white or grey; prickles absent

or present on midvein only, 0-3, straight,

acicular; stellate hairs dense or very dense,

0.05-0.1 mm apart, 0.4-0.55 mm diameter,

stalks 0-0.1 mm long, lateral rays 8-9, porrect;

central ray 0.4-1 times as long as laterals, not

gland-tipped; simple hairs absent; Type 2 hairs

absent. Inflorescence supra-axillary, cymose

(pseudo-racemose), common peduncle 3-10

mm long, rachis prickles absent or present, 1-

2-flowered, with all flowers bisexual. Flowers

5-merous. Pedicels at anthesis 6-13 mm long,

same thickness throughout, 0.5-0.6 mm thick

at mid-point, prickles absent. Calyx tube at

anthesis 2.5-3 mm long; lobes attenuate, 1.5-

2 mm long. Calyx prickles absent or present,

0-4 per flower, 1-4 mm long. Calyx stellate

hairs very dense, yellow or white or purple,

0.3-0.5 mm across, stalks absent, lateral rays

7-8, central ray 0.5-1 times as long as laterals,

not gland-tipped; simple hairs absent; Type 2

hairs absent. Corolla deeply lobed, 7-10 mm

long, purple, inner surface glabrous. Anthers

37-4.6 mm long. Ovary glabrous; style 7.5-

9 mm long, straight, protruding between

anthers or slightly bent, displacing one anther,

glabrous; stigma expanded, yellow, obscurely

lobed. Fruiting calyx lobes less than or more

than half length of mature fruit; prickles

absent or present, 2-4 mm long. Mature

fruits 1-2 per inflorescence, globular, 11-13

mm diameter, yellowish-green, 2-locular;

placenta sessile elliptical to semi-circular;

interior juicy, succulent; pericarp 0.2-0.5 mm

thick; pedicels 12-20 mm long. Seeds 2.5-27

mm long, pale yellow. Fig. 7.

Additional specimens examined: Queensland.

Mitchell District: c. 160 km S of Longreach, Apr 1989,

Emmott 286 (BRI); Noonbah Station, in Back Paddock,

SW corner of Back Paddock, Jan 2008, Emmott B516

(BRI, CANB); SW corner of Back Paddock, ‘Noonbah’,

c. 170 km SW of Longreach, May 2010, Bean 29771 &

Emmott (BRI); SW corner of Back Paddock, ‘Noonbah’,

c. 170 km SW of Longreach, May 2010, Bean 29772 &

Emmott (BRI, MEL). Gregory North District: 86.6 km

W of Winton on the Winton - Boulia Road, Nov 2010,

Emmott B682 & Denny { BRI, NY); 98.9 km W of Winton

on the Winton - Boulia Road, Nov 2010, Emmott B683 &

Denny (BM, BRI); Winton - Jundah Road, 4.4 km S of

‘Elvo’ Homestead, May 2004, Bean 22553 (BRI).

Distribution and habitat : Endemic to south¬

western Queensland between Stonehenge

and Winton (Map 1). It grows in shrubland

or low open woodland, on stony hills and

ranges, with shallow sandy soil. Associated

species include Eucalyptus normantonensis

Maiden & Cambage, Acacia shirleyi Maiden,

A. aneura and Triodia sp.

Phenology: Flowers have been recorded in

April, May and November; fruits in January,

May and November.

Notes: Solanum unispinum is similar in

appearance to S. cleistogamum , but differs

by the bushy habit and ascending branches,

branchlet prickles 1-4/cm (7-18/cm for S.

cleistogamum ), the central ray of the stellate

hairs 0.4-1 times as long as laterals (1-2.5

times for S. cleistogamum ), the leaf bases not

cordate, the calyx prickles 0-5 (10-25 for S.

cleistogamum ), the deeply lobed corolla (rotate

for S. cleistogamum ), and the anthers 37-4.6

mm long (2-2.8 mm for S. cleistogamum).

Conservation status: At the two sites on the

Winton - Boulia road, four plants and one

plant were found respectively. At ‘Noonbah’,

it is known from fewer than 20 plants. Only

two plants were seen near ‘Elvo’ Homestead,

while at the type locality there were fewer

than 10 plants. Using the criteria of the IUCN

Red List (IUCN 2000), a conservation status

of Critically Endangered (Criterion D) is

recommended.

Etymology: From the Latin uni meaning one

and spinus meaning spine. The calyces of this

species have no more than a single prickle on

each lobe, and sometimes only one calyx lobe

bears a prickle.

428

Austrobaileya 8(3): 412-430 (2011)

Fig. 7. Solanum unispinum. A. flowering branchlet x 1. B. oblique view of an open flower x 3. C. lateral view of flower

x 3. C. stamens, style and ovary x 4. D. cross-section of mature fruit x 3. All from Bean 22512 (BRI).

Acknowledgements

I am grateful to David Albrecht (NT) for taking

me into the field around Alice Springs to study

various Solanum taxa, and for convincing me

that S. cleistogamum is a distinct species.

Angus Emmott of Noonbah Station kindly

escorted me around his property to see

noteworthy solanums that he had found, and

supplied many images of them; he has also

made excellent collections from other parts

of western Queensland at my request. Will

Smith (BRI) provided the illustrations and

the distribution maps. Peter Bostock expertly

translated the diagnoses into Latin.

Bean, Solarium ellipticum group

References

429

Albrecht, D.E., Duguid, A.W., Coulson, H., Harris,

M.G. & Latz, P.K. (2007). Vascular Plant

Checklist for the Southern Bioregions of the

Northern Territory , 2nd edition. Northern

Territory Government: Alice Springs.

Bean, A.R. (2004). The taxonomy and ecology of

Solanum subg. Leptostemonum (Dunal) Bitter

(Solanaceae) in Queensland and far north¬

eastern New South Wales. Austrobaileya 6:

639-816.

Brooke, G. & Mcgarva, L. (1998). The Glove Box

Guide to plants of the NSW rangelands. NSW

Agriculture: Sydney.

Cunningham, G.M., Mulham, W.E., Milthorpe, P.L. &

Leigh, J.H. (1981). Plants of Western New South

Wales. Soil Conservation Service and N.S.W.

Government Printing Office: Sydney.

Henderson, R.J.F. (2002). Names and Distribution

of Queensland Plants, Algae and Lichens.

Queensland Herbarium, Environmental

Protection Agency: Toowong.

Iucn (2001). Red List Categories and Criteria: Version

3.1. IUCN Species Survival commission. IUCN:

Gland, Switzerland/Cambridge, U.K.

Kerrigan, R.A. & Albrecht, D.E. (2007). Checklist

of NT Vascular species, http://www.nt.gov.au/

nreta/wildlife/plants/pdf/200701nt_checklist.

pdf Accessed 17 th October 2010.

Symon, D.E. (1981). A revision of the genus Solanum

in Australia. Journal of the Adelaide Botanic

Gardens 4: 1-367.

Map 1 . Distribution of Solanum aridicola • and S. unispinum ▲

430

Austrobaileya 8(3): 412-430 (2011)

Map 2. Distribution of Solarium callosum

Map 3. Distribution of Solarium emmottii ▲ and S. lithophilum •

Austrobaileya 8(3): 431-434 (2011)

431

SHORT COMMUNICATION

Luvunga monophylla (DC.) Mabb. (Rutaceae):

a new species for Queensland

David G. Fell 13 & David J. Stanton 2

'3D Environmental, P.O. Box 337, Alstonville NSW 2477, Australia. Email:

davidfell@3denvironmental.com.au; Corresponding author

2 3D Environmental, PO Box 959, Kenmore, Queensland 4069, Australia.

The genus Luvunga Buch.-Ham. ex Wight

& Arn. contains twelve species occurring

in Asia, Malesia and Australia (Hyland et

al. 2003). Luvunga monophylla is the sole

Australian representative of Rutaceae subtribe

Triphasiineae in the tribe Aurantieae Rchb.

(Mabberley 1998); however, infrafamilial

relationships in this part of the family remain

to be fully resolved utilising molecular data

and this group may not be recognized in the

future (Kubitzki et al. 2010).

Luvunga monophylla (DC.) Mabb. is

a facultatively deciduous, spiny shrub or

climber, 0.5-3 m high with white, cream,

yellow flowers, obovoid fleshy fruit to 10 mm

diameter, and one or two seeds per fruit (Pedley

1987; Russell-Smith & Dunlop 1992; Hyland

et al. 2003; Department of Environment and

Conservation 2010). It was first collected from

Australia in the Sir Edward Pellew Group of

islands, Gulf of Carpentaria, in December

1802 by Robert Brown during Flinders’

Investigator circumnavigation of Australia

and was collected in Timor by J.B.L.C.T.

Leschenault de la Tour during Baudin’s

1800-1803 expedition in Le Geographe and

Le Naturaliste (Mabberley 1998).

It has now been recorded on lama (Yam

Island), Torres Strait which is the first record

of the species for Queensland. Prior to this

collection it had been recorded from Australia

in the North Kimberley and Dampierland

bioregions of Western Australia (Department

of Environment & Conservation 2010) and in

the Arnhem Coast, Central Arnhem, Tiwi,

Gulf Coastal, Gulf Fall and Uplands and

Darwin Coastal Bioregions of the Northern

Territory. The habitat in these areas includes

monsoon forest, deciduous vine thicket and

semi-deciduous vine thicket on coastal dunes

and basalt scree slopes (Pedley 1987; Russell-

Smith & Dunlop 1992; Liddle et al. 1994;

Department of Planning & Infrastructure

2010). It is also found in the Philippines,

Indonesia (Java) and Timor-Leste (Hyland et

al. 2003; Cowie 2006).

A survey of the vegetation of the Torres

Strait Islands, north Queensland, Australia,

was carried out in 2007 (Stanton et al.

2008). The survey’s primary objective was

to map vegetation communities at 1: 25,000

and Regional Ecosystems at 1: 50,000,

supplemented by floristic inventory and

collections of voucher specimens for

Australian herbaria. Collections of Luvunga

monophylla from lama (also known as Yam

Island) were made in October 2007 (Fell 8851

& Stanton [BRI]). This occurrence extends

its geographical range eastwards from the

eastern Arnhem Land coast of the Northern

Territory.

lama (Yam Island)

lama is a continental island of 186 ha located

at 9°54'S 142°46'E. It is situated 91 km north-

north-east of Cape York Peninsula and belongs

to the Central Group of Torres Strait Islands.

The island is the homeland of the Iamalgal

people. As at the 2006 Census, the population

was 309, of whom over 90% were Indigenous

(Australian Bureau of Statistics 2007). Land

tenure is Deed of Grant in Trust (DOGIT) and

Native Title.

Accepted for publication 8 August 2011

432

The primary geology of the island is granite

with a range of Holocene aged landforms

fringing its margins. Coarse-grained granite,

which forms the dominant landform feature,

represents the most easterly extension of the

Badu Granite batholith. Soil development is

typically skeletal, with large granite boulders

on lower slopes and littoral margins. Small

areas of acid volcanic rock form a headland

in the island’s east. Holocene features include

ridges of coralline sand and fine-grained

estuarine deposits associated with a broad

embayment on the island’s northern coast

(Willmott & Powell 1977; Stanton et al.

2008).

Remnant vegetation occurs over 75%

of the island with the balance supporting

regrowth, exotics, bamboo groves and cleared

land. Broad vegetation types are deciduous

and semi-deciduous vine forest and thicket,

Acacia dominated open forests and woodlands,

shrublands and shrubland complexes, coastal

dune complexes, grasslands and mangrove

forest (Stanton et al. 2008).

Luvunga monophylla on lama

Our vegetation survey recorded Luvunga

monophylla at eight sites on the island,

within open forest dominated by Acacia

auriculiformis A.Cunn. ex Benth. and in

deciduous vine forest and semi-deciduous

vine thicket. These vegetation types occur

on rocky granitic hillslopes and the Acacia

dominant vegetation is considered to have

developed in response to past disturbance.

The Acacia open forest habitat is endemic

to lama and the Torres Strait Islands and is

mapped as Regional Ecosystem 3.12.35f

(Stanton et al. 2008). Vine forest species

such as Canarium australianum F.Muell.,

Terminalia subacroptera Domin, Bombax

ceiba var. leiocarpum A.Robyns and

Diospyros hebecarpa A.Cunn. ex Benth.

are typical canopy associates. Groves of

the naturalised Bamboo ( Bambusa vulgaris

Schrad.) and scattered Mango trees ( Mangifera

indica L.) occur throughout this habitat and

are indicative of a long history of traditional

use by local people.

Austrobaileya 8(3): 431-434 (2011)

Deciduous vine forests and semi-deciduous

vine thickets (RE3.12.35a and RE3.12.21a)

are extensive on the lower exposed rocky

slopes and feature canopy species such as

Erythrina insularis F.M.Bailey, Antiaris

toxicaria var. macrophylla (R.Br.) Corner,

Terminalia subacroptera , Canarium

australianum , Bombax ceiba var. leiocarpum ,

Acacia auriculiformis , Manilkara kauki (L.)

Dubard, Gyrocarpus americanus Jacq. subsp.

americanus, Diospyros sp. (Mt White PI.

Forster PIF14415), Sterculia quadrifida R.Br.

and Premna dallachyana Benth.

The habitats in which Luvunga monophylla

occurs are in good condition and at the time of

survey were relatively undisturbed and free of

weeds. A conservative estimate of population

size for Luvunga monophylla is between 50

and 100 individuals. The area of extent is 50

ha.

Discussion

The nature of the disturbance history in the

Acacia open forest is unknown, although the

community would be expected to burn in hot

conditions. Old charred stumps indicate that

it is occasionally burnt, largely in a wildfire

regime. Resilience to fire is documented

from Northern Territory populations of

Luvunga which have an ability to resprout

from epicormic and basal lignotubers and

display less than 30% mortality rate when

subject to 100% leaf scorch. Life span is over

20 years with first seeding occurring between

6-20 years (Tropical Savannas Cooperative

Research Centre 2010).

The fact that the Acacia dominant

community is found only on lama presents

another puzzle. The nearby continental islands

of Mabuyag (Mabuiag Island) and Gebar

are dominated by open and closed forests of

Welchiodendron longivalve (F.Muell.) Peter

G. Wilson & J.T.Waterh., although not a single

individual of this species was found on lama.

This may suggest that forests on lama have

developed independently from forest types

on these neighbouring islands (Stanton et

al. 2008). Extensive field surveys by Stanton

et al. (2008) in similar habitats across the

majority of the Torres Strait islands did not

locate Luvunga elsewhere.

Fell & Stanton, Luvunga monophylla , new for Queensland

Prior to European contact the people

of lama practised traditional horticulture

growing crops such as banana, yam and sweet

potato (Haddon 1935; Neal 1989; Fuary 1991)

with archaeological evidence suggesting that

some gardening took place on rocky hillslopes

(Neal 1989). It is likely that a combination of

human disturbance, wind and fire have played

a role in the development of Luvunga habitat.

No uses are documented for Australian

populations of Luvunga monophylla.

Specimens were shown to numerous members

of the community during our visit to lama in

November 2010. Although they recognized the

plant, they stated that they had no name or use

for it. In Java, it is occasionally cultivated as a

hedge plant (Kruse 2001: 1013). In India, the

dried fruits of Luvunga scandens Roxb. are

used in the production of a medicinal oil said

to be effective in treating dermal infections

and baldness (Garg & Jain 1999) and the roots

and fruits are used for treatment of scorpion

stings (Lien et al. 2002).

It is interesting to note that the occurrence

of Luvunga monophylla across northern

Australia appears to be coterminous with

those areas visited in the past by Macassan

beche-de-mer fishermen from southern

Sulawesi in what is now Indonesia, Timor-

Leste and between Cape Leveque in Western

Australia and the Sir Edward Pel lew Group

in the Northern Territory (Macknight 1969: 2,

27; Stacey 2007: 58). There is no firm evidence

that this trade included Torres Strait, although

there is an 1881 record of an Ambonese prau

(boat) being driven into the region during

the northwest monsoon season (Anonymous

1881). This raises the possibility that Luvunga

monophylla was introduced to Australia by

human vectors.

Conservation status of Luvunga

monophylla

In the Northern Territory this species is

assigned ‘least concern’ status under the

Territory Parks and Wildlife Conservation

Act 2000, and in Western Australia it is

‘Not Threatened’ according to the Wildlife

Conservation Act 1950.

433

Cowie (2006) records the taxon from

dry deciduous forest in Timor-Leste,

noting that the forest type is under ongoing

pressure from conversion to agriculture.

The heavy exploitation of dry deciduous

forests elsewhere in the Timor region

for swidden agriculture has resulted in

large-scale conversion to anthropogenic

grassland -Chromolaena shrubland (Cowie

2006). The extent of its occurrence and

condition in habitats in the Philippines and

Java is not known.

Potential threats to the lama population

include habitat disturbance from infrastructure

and residential development, weed incursion

and perhaps wildfire. Pressures on the limited

availability of land suitable for housing and

infrastructure development is intensified

by population growth and coastal erosion

associated with regular tidal surges. This

presents a considerable challenge for the

island community and planning agencies.

In combination, these threats represent a

risk of stochastic extinction of the species in

Queensland due to its small population size

and highly localised occurrence. Accordingly,

the species should be assigned regional

significance in the Cape York Peninsula

Bioregion. Integration of ecological and

genetic studies will be required to determine

the management requirements of Luvunga

monophylla on lama. Further assessments of

population size, phenology and recruitment is

achievable at the local level with the assistance

of the local ranger program.

Acknowledgements

The field survey on lama was funded by the

Land and Sea Management Unit of the Torres

Strait Regional Authority. Access to lama was

gratefully provided by the Magani Lagaugal

(Torres Strait Islanders) Corporation. Advice

on identification and species distribution

was provided by the staff of the Queensland

Herbarium. Comments on an earlier version

of this paper from Garrick Hitchcock and Rob

Neal are greatly appreciated.

434

References

Anonymous (1881). The Brisbane Courier 21 February:

Australian Bureau Of Statistics (2007). Population

Distribution, Aboriginal and Torres Strait

Islander Australians, Australia - 2006. Cat.

No. 4705.0. Australian Bureau of Statistics:

Canberra.

Cowie, I. (2006). A Survey of Flora and Vegetation of

the Proposed Jaco-Tutuala-Lore National

Park, Timor-Leste (East Timor). Report to

Birdlife International from NT Herbarium

(DNA) Department of Natural Resources,

Environment and the Arts: Palmerston, N.T.

Version 1.0. Available from: http://www.dpif.

nt.gov.au/nreta/publications/wildlife/science/

pdf/2006CowieI.pdf

Department Of Environment And Conservation,

Western Australian Herbarium (2010).

Florabase. Online at http://florabase.calm,

wa.gov.au/ Accessed 30 October 2010.

Department Of Planning And Infrastructure (2010).

Australian Virtual Herbarium. NT Herbarium

map tool. Northern Territory DPI, Darwin.

Available from: http:/ipe.nt.gov.au/cgi-bin/avh.

cgi.

Garg, S.C. & Jain, R. (1999). Antifungal activity of

Luvunga scandens against some keratinophilic

fungi. Indian Journal of Pharmaceutical

Sciences 61: 248-249.

Fuary, M.M. (1991). In so many words: an ethnography

of life and identity on Yam Island, Torres Strait.

PhD thesis, James Cook University of North

Queensland: Townsville.

Haddon, A.C. (1935). Reports of the Cambridge

anthropological expedition to Torres Straits.

Vol. 1. general ethnography. Cambridge

University Press: Cambridge.

Hyland, B.P.M., Whiffin, T., Christophel, D.C., Gray,

B. & Elick, R.W. (2003). Australian tropical

rain forest plants: trees and shr ubs and vines.

CSIRO Publishing: Melbourne.

Kruse, J. (2001). Rutaceae. In P. Hanelt (ed.), Mansfeld’s

encyclopedia of agricultural and horticultural

crops, pp. 996-1038. Springer Verlag: Berlin.

Kubitzki, K., Kallunki, J.A., Duretto, M. & Wilson,

PG. (2010). Rutaceae. In K. Kubitzki (ed.).

The Families and Genera of Vascular Plants.

X. Flowering Plants Eudicots Sapindales ,

Cucurbitales , Myrtaceae , pp. 276-356. Springer

Verlag: Berlin/Heidelberg.

Austrobaileya 8(3): 431-434 (2011)

Liddle, D.T., Russell-Smith, J., Brock, J., Leach, G.J.

& Connors, G.T. (1994). Atlas of the vascular

rainforest plants of the Northern Territory.

Flora of Australia Supplementary Series No.

3. Australian Biological Resources Study:

Canberra.

Lien, T.P, Kamperdick, C., Schmidt, J., Adam, G. & Van

Sung, T. (2002). Apotirucallane triterpenoids

from Luvunga sarmentosa (Rutaceae)

Phytochemistry 60: 747-754.

Mabberley, D.J. (1998). Australian Citreae with notes

on other Aurantioideae (Rutaceae). Telopea 7:

333-344.

Macknight, C.C. (1976). The voyage to Marege’:

Macassan Trepangers in Northern Australia.

Melbourne University Press: Carlton.

Neal, R.A. (1989). An Archeological Inspection of

Alternative Telecom Locations on Mabuiag

and Yam Islands, Torres Strait. Unpublished

report to Department of Community Services

and Ethnic Affairs. Pirriport Pty Ltd: Helidon,

Queensland.

Pedley, L. (1987). Paramignya Wight (Rutaceae:

Citreae) in Australia. Austrobaileya 2: 416.

Russell-Smith, J. & Dunlop, C.R. (1992). The status of

monsoon vine forests in the Northern Territory:

a perspective. In G.L. Werren & A.P Kershaw

(eds.) The rainforest legacy. Australian

National Rainforests Study. Vol. 1, The Nature,

Distribution and Status of Rainforest types, pp.

227-288. Australian Government Publishing

Service: Canberra.

Stacey, N. (2007). Boats to burn: Bajo fishing activity

in the Australian fishing zone. Asia-Pacific

Environment Monograph 2. ANU E Press:

Canberra.

Stanton D.J., Fell, D.G. & Gooding, D.O. (2008).

Vegetation Communities and Regional

Ecosystems of the Torres Strait Islands,

Queensland, Australia. Unpublished report

to the Torres Strait regional Authority, Land

and Sea Management Unit. 3D Environmental

Trust: Brisbane.

Tropical Savannas Cooperative Research Centre

(2010). North Australian Land Manager

Website. Available from: http://landmanager,

org.au/view/311424/fire - responses-of--

luvunga-monophvlla.html

Willmott, W.F. & Powell, B.S. (1977). Torres Strait-

Boigu-Daru, Queensland 1:250 000 Geological

Series - Explanatory Notes, Sheets SC/ 54-12,

SC/ 54-7 and SC/ 54-8. Bureau of Mineral

Resources, Geology and Geophysics. Australian

Government Publishing Service: Canberra.

Austrobaileya 8(3): 435-437 (2011)

435

SHORT COMMUNICATION

Hydrocharis dubia (Blume) Backer (Hydrocharitaceae)

is an alien species in Australia

A.R. Bean

Queensland Herbarium, Department of Environment & Resource Management, Brisbane Botanic

Gardens, Mt Coot-tha Road, Toowong 4066, Queensland, Australia. Email: tony.bean@derm.qld.

gov.au

Hydrocharis dubia (Blume) Backer, is an

aquatic plant with a white 3-petaled flower

that occurs widely in eastern Asia. It has

been accepted as an indigenous species in

Australia but recently this has been repeatedly

questioned by weed control authorities. This

note examines the evidence relating to its

origin status and concludes that it is not native

in Australia.

The genus Hydrocharis L. was revised by

Cook & Luond (1982), with the recognition of

three species,//. morsus-ranaeL.,H. chevalieri

(De Wild.) Dandy and H. dubia. Hydrocharis

morsus-ranae is widely distributed in Europe,

with scattered occurrences in western Asia. It

has become a serious weed in south-eastern

Canada (Cook & Luond 1982) and in the

north-eastern U.S.A. (Catling et al. 2003;

O’Neill 2007). Hydrocharis chevalieri is

restricted to central Africa, and is not recorded

as a weed. Hydrocharis dubia is widely

distributed in south-eastern Asia, from India

to Japan and south to Java and New Guinea,

and in Australia (Cook & Luond 1982). It has

been referred to as a “noxious weed” in Japan

(Oki 1994). Shaffer-Fehre (1991) transferred

Hydrocharis dubia to Limnobium, a genus

which until then had comprised two species

confined to the Americas; however, this

transfer has not been followed in the Flora

of Australia (Jacobs & McColl 2011) and is

not supported by molecular data (Les et al.

2006).

Hydrocharis dubia was first recorded

for Australia by Bentham (1873) under the

misapplied name// morsus-ranae. The record

Accepted for publication 8 September 2011

was based on a specimen (now at K) collected

by J. Bidwill, from “Wide Bay” between 1848

and 1853. This specimen was cited by Cook

& Luond (1982) under Hydrocharis dubia.

Bentham (1873) entertained the idea that it

could be an introduced species in Australia.

Bailey (1902) recorded Hydrocharis

morsus-ranae from “the still waters about the

Brisbane River”. There is a specimen record

of Hydrocharis dubia on the BRI database

with the locality “Brisbane River”, but the

specimen cannot currently be located.

Hydrocharis dubia has been found at 13

Australian locations in Queensland (Qld) and

New South Wales (NSW) based on herbarium

records (Table 1).

Hydrocharis dubia (as Limnobium dubium

by some authors) has been regarded as an

indigenous species in Australia (Aston 1973;

Stanley & Ross 1989; Jacobs 1993; Holland

& Hansen 2009; Jacobs & McColl 2011). The

reasons for this have not been expounded, but

they are probably two-fold: that the species

is indigenous to south-east Asia, and natural

dispersal from that region to Australia is not an

unreasonable hypothesis; and that the species

is included in Bentham’s Flora Australiensis.

Thomas & McDonald (1989) gave

Hydrocharis dubia a status of ‘2V’, meaning

a Vulnerable species with a geographic range

in Australia of less than 100 km. Presumably

the authors, at that time, considered it native

to south-eastern Queensland. Subsequently

it was classified as “Vulnerable” under the

Queensland Nature Conservation Act 1992,

and “Vulnerable” under the Environment

Protection and Biodiversity Conservation Act

1999.

436

Austrobaileya 8(3): 435^37 (2011)

Table 1. Locations for Hydrocharis dubia in Australia over time

Localities

Time period of collections

Wide Bay, Qld

1848-1853

Clarence River, NSW

1882

Brisbane River, Qld

pre-1902

Harrisville, Qld

1960, 1985, 1986

Bountiful Creek, Richmond River, NSW

1967

Frederickton near Kempsey, NSW

1968

Beaudesert, Qld

1971, 1977

Toomba, Qld

1977, 1996, 1997, 2001, 2008

16 km W of Casino, NSW

1978

Ayr, Qld

1994

Rosewood, Qld

1999, 2001, 2009

Fletcher Ck, Qld

2002, 2004

Giru, Qld

2004

The following observations are made on

Hydrocharis dubia in Australia drawing on

criteria used by Bean (2007) to assess its

origin status:

1. it grows in disturbed or modified sites, and

has not been found in remote “intact” areas.

This is consistent with a species introduced

by man, and transported to other sites via

machinery or with soil, or dumping of plants

2. it is persistently invasive (one label reports

it covering two dams), and is not in ecological

balance with surrounding biota

3. it does not appear to have any pests or

diseases, suggesting it has arrived recently in

evolutionary terms

4. it has substantially expanded its

geographical range over the last 40 years (see

list of locations above)

5. its distribution is highly discontinuous, and

the discontinuities cannot be explained by

climatic or other natural criteria

6. the initial locations in Australia are highly

disjunct geographically from nearest known

occurrences in Asia

7. the first record for Australia (Wide Bay,

between 1848 and 1853, by JohnBidwill) is not

early enough to preclude human introduction

Hydrocharis dubia fails all of the

ecological criteria for an indigenous species,

and the historical criteria are equivocal. In the

key of Bean (2007) it readily keys to “alien”

(lx, 2).

In conclusion, there is a strong weight

of evidence to support an alien status

for Hydrocharis dubia with regard to its

occurrences in Australia. I recommend that

this species be recorded as naturalised for

Australia, and de-listed from the Queensland

Nature Conservation Act 1992 , and the

Environment Protection and Biodiversity

Conservation Act 1999.

References

Aston, H.I. (1973). Aquatic Plants of Australia.

Melbourne University Press: Carlton.

Bailey, F.M. (1902). Hydrocharis. InFlora of Queensland

5: 1510. H. J. Diddams & Co.: Brisbane.

Bean, A.R. (2007). A new system for determining

which plant species are indigenous in Australia.

Australian Systematic Botany 20: 1-43.

Bentham, G. (1873). Hydrocharis. In Flora Australiensis

6: 256. L. Reeve & Co.: London.

Catling, P.M., Mitrow, G., Haber, E., Posluszny, U.

& Charlton, W.A. (2003). The biology of

Canadian weeds. 124. Hydrocharis morus-

ranae L. Canadian Journal of Plant Science 83:

1001-1016.

437

Bean, Hydrocharis dubia alien in Australia

Cook, C.D.K. & Luond, R. (1982). A revision of the

genus Hydrocharis (Hydrocharitaceae). Aquatic

Botany 14: 177-204.

Holland, A.E. & Hansen, D. (2009). Species

Information - Hydrocharis dubia. Wetland

Info, Queensland Government. http://www.

epa. qld .gov. au/wetlandinfo/site/factsfigures/

FloraAndFauna/Flora/IndicatorSpeciesList/121

14.html Accessed 29 March 2011.

Jacobs, S.W. (1993). Hydrocharitaceae. In G. Harden

(ed.). Flora of New South Wales 4: 11-16. New

South Wales University Press: Sydney.

Jacobs, S.W. & McColl, K.A. (2011). Hydrocharis. In

Flora of Australia 39: 21-22. ABRS/CSIRO:

Melbourne.

Les, D.H., Moody, M.L. & Soros, C.L. (2006). A

reappraisal of phylogenetic relationships in the

monocotyledon family Hydrocharitaceae. Aliso

22: 211-230.

Oki, Y. (1994). Integrated Management of aquatic

weeds in Japan. Food & Fertilizer Technology

C enter, http: II www. agnet. org/1 ibrary/bc/45008/

Accessed 29 March 2011.

O’Neill, C.R. (2007). European Frog-bit ( Hydrocharis

morsus-ranae ) - Floating invader of Great

Lakes Basin Waters. NYSG Invasive Species

Factsheet Series: 07-1. http://www.seagrant.

sunysb.edu/ais/pdfs/Frog-bitFactsheet.pdf

Accessed 29 March 2011.

Shaffer-Fehre, M. (1991). The endotegmen tuberculae:

an account of little-known structures from

the seed coat of the Hydrocharitoideae

(Hydrocharitaceae) and Najas (Najadaceae).

Botanical Journal of the Linnean Society 107:

169-188.

Stanley, T.D. & Ross, E.M. (1989). Flora of south¬

eastern Queensland , Volume 3. Department of

Primary Industries: Brisbane.

Thomas, M.B. & McDonald, W.J.F. (1989). Rare and

threatened plants of Queensland , 2 nd edition.

Department of Primary Industries: Brisbane.

Austrobaileya 8(3): 438-440 (2011)

438

SHORT COMMUNICATION

Reinstatement of Vigna suberecta Benth. (Fabaceae: Phaseoleae)

Ailsa E. Holland

Queensland Herbarium, Department of Environment & Resource Management, Brisbane Botanic

Gardens, Mt Coot-tha, Toowong, Queensland 4066, Australia. Email: Ailsa.Holland@derm.qld.gov.au.

The genus Vigna Savi is a pantropical,

herbaceous legume genus of about 200

species distributed throughout the tropics

and subtropics of both hemispheres. In

Australia, there are a number of endemic

Vigna taxa, including those in the wide

ranging V lanceolata Benth. complex (Lawn

& Holland 2003).

Vigna lanceolata and Vigna suberecta

Benth. were both originally described by

Bentham in Mitchell (1848). On the 10 th of

October 1846 Mitchell arrived at the summit

of Mt Farraday (Mitchell 1848: 347) where

he first encountered “the yellow Vigna

lanceolata ” and collected specimen number

376, the left hand specimen on Kew sheet

H/1215/89 4, labelled “ Vigna lanceolata

Benth. in Mitch Austr. Trop. p. 350. 1846. 376.

Oct 10 402, small twining herb, fl yellow, top

of Mount Farraday” and annotated as the

type by Marechal, although he later refers

to specimen 370 (Marechal et al:. 1978: 168)

which is probably an error.

On 13 th of October 1846, Mitchell had

advanced to Mt Owen (Mitchell 1848: 350),

where he found “a new species of Vigna

with yellow flowers” and collected specimen

number 401. This is the right hand specimen

labeled as “Oct 13 420 on Mt Owen... probably

label changed with Swainsona phasoides ”,

on another Kew sheet H/1215/89 3 and is the

same taxon as the previous. This species was

described by Bentham in the footnote on this

page (translated): “V. LANCEOLATA (Benth.

MS.). Glabrous, twining. Leaflets lanceolate,

reticulate, entire, hastate or lobed at the base.

Peduncles much longer than leaves, with few

flowers clustered at apex. Calyx glabrous,

campanulate, the lobes shorter than the tube;

keel beak acute”

Accepted for publication 9 September 2011

Mitchell arrived on the Moonie River

on the 18 th November 1846 (Mitchell 1848:

387) and collected another specimen “in the

thick forest, a new species of Vigna very

near V. lanceolata , though very different in

habit”. This is specimen number 466, Kew

sheet H/1215/89 5, label “1846. 466. Vigna

suberecta Benth. in Mitch Austral, trop.”

This species was described by Bentham in

a footnote (Mitchell 1848: 388) translated

as: “V. SUBERECTA (Benth. MS.); slightly

pubescent, sub-erect, branched. Leaflets

broadly lanceolate leaflets, entire, hastate or

3-lobed at base. Peduncles somewhat shorter

than the leaves, few-flowered at the apex.

Calyx pubescent, campanulate; calyx lobes

subequal to the tube; keel beaked, acute. Pod

sparsely pubescent .”

Bentham later (1864) placed Vigna

suberecta in synonymy under V. lanceolata ,

but did not give a reason. Examination

of the type material (the sheets indicated

above) shows that the specimens labelled “V

lanceolata ” (376 and 401) do indeed belong

to the same taxon, but the specimen labelled

“E suberecta ” (466) is quite distinct and so

this name is here reinstated for a relatively

widespread species.

Vigna suberecta Benth., in T.L.Mitchell,

J. Exped. Trop. Australia[Mitchell] 388 (1848).

Type: subtropical New Holland [Queensland.

Darling Downs District: Moonie River], [18

November] 1846, T.L. Mitchell 466 (holo: K).

Vigna sp. (Jimbour A.R.Bean 12534); Holland

& Pedley (2007: 84).

Herb with perennial tuberous rootstock; tubers

cylindrical, up to 15 mm diameter. Aerial

parts annual or short-lived perennial. Stems

to 80 cm long, initially semi-erect, becoming

twining in upper parts at maturity, with a few

appressed, retrorse white hairs to 0.8 mm

Holland, Reinstatement of Vigna suberecta (Fabaceae)

long. First leaves unifoliolate, petiolate; blade

narrowly triangular, 2-2.5 cm long, 4-7 mm

wide, truncate to slightly sagitate at base.

Remaining leaves trifoliolate; petiole 2.2-

5.7 cm long; rachis 5-15 mm long; stipels

lanceolate, 1-2.3 mm long. Terminal leaflets

lanceolate, becoming linear towards the plant

apex, 2.5-7 cm long, 3-17 mm wide; 3 to 13

times as long as wide, apex acute, base cuneate

with rounded or acute lobes, 1-4 mm long

and wide; upper surface glabrous or nearly

so, lower surface very sparsely appressed

hairy; midrib uniformly coloured (not silver

striped), secondary nerves forming an angle

of 45-80° to the midrib, looped at margin.

Lateral leaflets similar, slightly shorter

than terminal leaflet. Stipules triangular

to lanceolate, 1.8-3.5 mm long, 3 or 5-

veined, acute, base ± cordate. Subterranean

inflorescences absent. Aerial inflorescences

with 2-10 flowers crowded towards the apex;

flowers developing sequentially in pairs with

a swollen extrafloral nectary between each

pair. Peduncle terete, striate, 2-16 cm long;

pedicel 1-2.2 mm long; bracts and bracteoles

ovate, acute, 0.8-1.2 mm long; bracteoles

inserted at base of calyx. Calyx campanulate,

2.2-4.5 mm long, glabrous or minutely ciliate;

tube 1.5-2.5 mm long; lobes triangular, equal

to or slightly shorter than tube, upper two

joined almost to apex, lower one longest.

Corolla pale yellow, glabrous. Standard

transversely elliptic, emarginate, 6-9 mm

long, 10—14 mm wide; the inner surface with

two U-shaped pockets with vertical ridges, c.

2 mm long; apex emarginate; base somewhat

auriculate; claw 1-1.5 mm long. Wings 6-10

mm long, 4-7 mm wide, expanding from

a narrow base into a broadly rounded apex

covering the keel; upper margin with a small

spur near base; lower margin without a spur;

claw 1.5-2 mm long. Keel 5-9 mm long, 4-6

mm wide; upper margin with a shallow pocket

near base; lower margin strongly curved;

beak strongly upturned and twisted through

c. 180°; claw 2-3 mm long. Filament tube

c. 10 mm long, strongly upturned; filaments

free for 1/3 - 1/2 their length; anthers 2-3

mm long. Ovary pubescent, 4-6 mm long;

style terete, twisted, anterior face hairy for

1/3—1/2 the length; stigma situated near apex,

the style slightly prolonged past stigma into a

439

beak 2-3 mm long, straight or slightly curved

by less than 90° Pods slightly curved, 2.5-4

cm long, 3-6 mm wide, sparsely puberulent

when young, glabrescent, brown or black at

maturity. Seeds 6-10, oblong, dark brown or

reddish, 3-4.2 mm long, 3-3.7 mm wide, 2-

2.6 mm thick; hilum central, 1-1.3 mm long.

Additional selected specimens examined : Queensland.

Leichhardt District: 169 km from Clermont toward

Mackay on Peak Downs Highway, Dec 1987, Lawn &

Cottrell ACC906 (BRI); 9 km N of Capella, Mar 1995,

Fensham 2769 (BRI); 9 miles [14.4 km] SSW of Comet,

Feb 1960, Johnson 1382 (BRI). Burnett District:

Wooroolin, Portion 31, Nov 1961, Bailey s.n. (BRI

[AQ239116]). Maranoa District: Roma, Mar 1936, Blake

10854A (BRI). Darling Downs District: Marnhull, SE of

Jandowae, Dec 2001, Bean 18199 (BRI); Jimbour Plain,

Mar 1951, BissetS758 (BRI); CSIRO experimental area,

Nandi, Dalby, Apr 1983, Watkinson ACC207 (BRI);

19 km SSW of Dalby, Feb 1995, Fensham 1909 (BRI);

Jondaryan, Feb 1935, Blake 7753 (BRI); 8 km S of Cecil

Plains turnoff (E of Dalby) at Duncan’s road turnoff,

Nov 2001, Holland 1388 et al. (BRI, US); Cecil Plains to

Milmerran Road at River Road turnoff. Mar 1987, Lawn

et al. ACC261 (BRI). New South Wales. Yalleroi, s.dat.,

Wilson ACC210 (BRI); Moree, Mar 1962, Cutting s.n.

(NSW); Delungra, Feb 1968, Gilroy s.n. (NSW); Wee

Waa, c. 32 km NW of Narrabri, Feb 1981, Nehrkorn s.n.

(NSW); Edgeroi, 16 miles [25.6 km] N of Narrabri, Feb

1966, Quick s.n. (NSW); Gunnedah, May 1938, O’Reilly

s.n. (NSW).

Distribution and habitat: Vigna suberecta

is a widespread species occurring mainly in

the Brigalow Belt bioregion of Queensland

from Clermont in the north to as far west as

Roma, and as far south as Gunnedah in New

South Wales. It is confined to heavy clay soils

in grassland or open eucalypt woodland with

grassy under storey.

Phenology: Flowers in summer, mostly

between November and March and fruits

through to June, depending on rainfall.

Notes: Vigna suberecta is distinguished

from V. lanceolata by the leaflets lacking a

silver midrib, wing petals rounded, keel beak

twisted c. 180°, and seeds with a longer (1-1.3

mm) hilum (Table 1).

While Vigna suberecta is a semi-erect

herb in the early stages, it becomes vine-like

and twining at maturity. Vigna suberecta

often resprouts from tubers in spring, or after

fire.

440 Austrobaileya 8(3): 438-440 (2011)

Table 1. Morphological character differences between Vigna lanceolata Benth. and

V. suberecta Benth.

Character

V lanceolata

V suberecta

Underground inflorescences

yes

leaflets midrib silver striped

yes

standard with raised vertical ridges

yes

wing shape

triangular

rounded

keel beak twist

c. 90°

c. 180°

seed hilum length

0.7-1.1 mm

1-1.3 mm

Acknowledgements

I would like to thank the director, Royal

Botanic Gardens Kew for the loan of type

material, and the director of the National

Herbarium of New South Wales for the

specimen loan. Thanks to Prof R. Lawn and

Dr L. Bielig (both JCU) for their support

and encouragement, and to Peter Bostock

for the excellent Latin translation program

TRANSLAT. This work was done in partial

fulfillment of a PhD degree at James Cook

University (JCU).

References

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Lawn, R.J. & Holland, A.E. (2003). Variation in

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