Editorial Committee

L.W. Jessup (editor)

P.D. Bostock (technical advisor)

B.K. Simon (technical advisor)

Desktop Publishing

Patrick O’Duffy

Yvonne Smith

Austrobaileya

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

Vol. 6, No. 3 was published on 3 December 2003

Vol. 6, No. 4 was published on 6 December 2004*

Austrobaileya is published once per year.

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addressed to: The Editor, Austrobaileya, Queensland Herbarium, Environmental Protection

Agency (EPA), Brisbane Botanic Gardens, Mt Coot-tha, Mt. Coot-tha Road, Toowong Qld

4066, Australia.

ISSN 0155-4131

Austrobaileya is the journal of the Queensland Herbarium and is devoted to publication of

results of sound research and of informed discussion on plant systematics, with special emphasis

on Queensland plants.

Opinions expressed by authors are their own and do not necessarily represent the policies or

views of the Queensland Herbarium.

* this note was added to Acrobat version of Austrobaileya 6(4). It did not appear in the printed version.

Contents

Notes on Tiliaceae in Australia, 4. A revision of the stellate-haired species of

the genus Corchorus L.

D.A. Halford.581

Stictocardia Hallier f. (Convolvulaecae) in Queensland

R.W. Johnson.631

The taxonomy and ecology of Solanum subg. Leptostemonum (Dunal) Bitter

(Solanaceae) in Queensland and far north-eastern New South Wales, Australia

A.R. Bean.639

Vanguerieae A. Rich, ex Dum. (Rubiaceae) in Australia, 3. Psydrax Gaertn.

Sally T. Reynolds & Rodney J.F. Henderson.817

A new species of Gynochtodes Blume (Rubiaceae) from north-east Queensland

D.A. Halford.891

Two new species of Morinda L. (Rubiaceae) from north-east Queensland

D.A. Halford & A.J. Ford.895

The tribe Coffeeae DC. (Rubiaceae: Ixoroideae) in Australia

Paul I. Forster.903

Caelospermum dasylobum (Rubiaceae), a new species from north-east Queensland

D.A. Halford & A.J. Ford.911

New species of Cryptandra Sm. and Stenanthemum Reissek (Rhamnaceae)

from northern Australia

A.R. Bean.917

Rediscovery of Tylophora linearis P.I. Forst. (Apocynaceae: Asclepiadoideae)

from New South Wales, with revision of its conservation status to vulnerable

Paul I. Forster, Doug Binns & Geoff Robertson.941

New Australian species in the lichen genus Siphula Fr.

Gintaras Kantvilas.949

Chromosome records for five trigger plants ( Stylidium\ Stylidiaceae) from

northern Australia

J.A. Wege.957

Gonocarpus hirtus Orchard (Haloragaceae), new from south-eastern

Queensland and north-eastern New South Wales

A.E. Orchard.961

Taxonomic notes on palms (Arecaceae) in catalogues of the Brisbane Botanic

Garden, Australia, of 1875 and 1885

John Leslie Dowe. 967

Notelaea ipsviciensis (Oleaceae), a new species from south-east Queensland

Wayne K. Harris.973

Notes

A new combination in Centratherum Cass. (Asteraceae)

A.R. Bean.977

Nomenclatural changes for two Australian species of Livistona R.Br. (Arecaceae)

John L. Dowe & David L. Jones.979

Reduction of Acacia perangusta to the synonymy of A. fimbriata

Les Pedley.983

0 continued )

Supplement to a synopsis of Racosperma C. Mart. (Leguminosae: Mimosoideae)

Les Pedley.985

Addendum .987

Index .989

Referees consulted for Volume 6 . 988

Notes on Tiliaceae in Australia, 4. A revision of the stellate-haired species of

the genus Corchorus L.

D. A. Halford

Summary

Halford, D. A. (2004). Notes on Tiliaceae in Australia, 4. A revision of the stellate-haired species of the

genus Corchorus L. Austrobaileya 6 (4): 581-629. Twenty-two species are recognised and a key is provided

for their identification. The following species are newly described: C. aulacocarpus Halford,

C. carnarvonensis Halford, C. congener Halford, C. incanus Halford, C. lasiocarpus Halford,

C. mitchellensis Halford, C. obclavatus Halford, C. puberulus Halford, C. subargentus Halford, C. sublatus

Halford and C. tectus Halford. Three new combinations C. sericeus subsp. densiflorus (Benth.) Halford,

based on C. walcottii var. densiflorus Benth.; C. sidoides subsp. rostrisepalus (Domin) Halford, based on

C. rostrisepalus Domin; C. sidoides subsp. vermicularis (F.Muell.) Halford, based on C. vermicularis

F.Muell., are made. Two new subspecies are described: C. incanus subsp. lithophilus Halford and

C. lasiocarpus subsp. parvus Halford. All new taxa are illustrated, while all taxa are described and mapped,

and notes on their distribution, habitat and phenology are given. Lectotypes are chosen for Nettoa

crozophorifolia Baill., C. elderi F.Muell., C.pumilio R.Br. ex Benth., C. rostrisepalus Domin, C. sidoides

F.Muell., C. tomentellus F.Muell., C. walcottii F.Muell. and C. walcottii var. densiflorus Benth. All known

synonyms are listed here including manuscript names that were used to identify taxa prior to their formal

naming in this publication.

Key words: Tiliaceae, Corchorus, Australian flora, taxonomy, nomenclature

D. A. Halford, c/- Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens Mt

Coot-tha, Mt Coot-tha Road, Toowong, Queensland 4066, Australia.

Introduction

This is the fourth paper in a series examining

the family Tiliaceae in Australia and the second

concerned with the genus Corchorus L. This

paper examines the stellate-haired species of

Corchorus. The taxonomy of the other

Australian species (simple-haired species) were

dealt with in Halford (1995). Abrief history of

the taxonomy of the Australian representatives

of this genus was also presented in that paper.

All species treated here are endemic to

Australia. Taxonomic problems certainly

remain in the stellate-haired species of

Corchorus in Australia and there is need for

further collection and study throughout the

range of the group, especially in the Pilbara

and Kimberley regions of Western Australia

and Arnhem Land, Northern Territory. Such

field investigations are beyond the resources

of the present author. A total of 35 species (13

simple-haired and 22 stellate-haired) are

recognised as occurring in Australia.

Accepted for publication 7 March 2004

Materials and methods

This revision is based on an assessment of

morphological characters of approximately 700

dried herbarium collections and collections and

field studies undertaken by the author in 1992

and 1993. Collections from the following

herbaria were studied: AD, BRI, CANB, DNA,

K, MEL, NSW, P and PERTH. These

herbarium acronyms follow Holmgren et al.

(1990). All specimens cited have been seen

unless otherwise indicated (as n.v.).

The species treated in the present paper are

listed alphabetically. Descriptions of colour of

vegetative and flora parts are either from the

herbarium labels or from photographs taken

by the author during field studies. Measurements

listed are based upon the total variation

observed in the herbarium specimens

examined. Plant size, flowering and fruiting

times, and habitat information were obtained

from herbarium labels. All measurements were

made from dried material, material preserved

in 70% ethanol or dried material reconstituted

by placing in boiling water for a few minutes.

The distribution maps are based on herbarium

specimen locality data.

582

Austrobaileya 6 (4): 581-629 (2004)

Taxonomy

Key to stellate-haired Corchorus in Australia

1. Sepals persistent in fruit. 2

Sepals not persistent in fruit. 9

2. Glandular hairs yellow or red-brown, conspicuous, on either stems, petioles,

peduncles, pedicels or abaxial surface of sepals. 3

Glandular hairs white and inconspicuous or absent. 4

3. Sepals < 9 mm long (W.A.). 13. C. parviflorus

Sepals > 9 mm long (W.A.).8. C. laniflorus

4. Annulus densely stellate-pubescent (W.A.). 4. C. crozophorifolius

Annulus glabrous or with a few scattered hairs. 5

5. Fruits < 5 mm long, with indumentum of stellate hairs; ovary 3 or

4-locular; ovules < 10 in each locule. 6

Fruits > 5 mm long, with indumentum of stellate and dendritic-stellate

hairs; ovary 3 to 5-locular; ovules 10-25 in each locule. 7

6. Leaf laminae narrowly to broadly ovate or ovate-elliptic (N.T., Qld). 16. C. sericeus

Leaf laminae narrowly oblong to oblong (W.A.).20. C. tectus

7. Fruits > 12 mm across (including indumentum) (W.A.). 9. C. lasiocarpus

Fruits <12 mm across (including indumentum). 8

8. Leaf laminae narrowly oblong to oblong or narrowly ovate, 0.4-2.5 cm

wide, l:w ratio >2:1 (W.A.). 9. C. lasiocarpus

Leaf laminae ovate to very broadly ovate, 2-7 cm wide, l:w ratio <2:1

(W.A.). 7. C. incanus

9. Fruits with indumentum of stellate and dendritic-stellate hairs. 10

Fruits with indumentum of stellate hairs only. 13

10. Glandular hairs present on stems, petioles, peduncles, pedicels or abaxial

surface of sepals (W.A.) . 22. C. walcottii

Glandular hairs absent. 11

11. Fruits subcylindrical, 10-18 mm long, 1.5-2.5 m across (including

indumentum), 5-12 times longer than wide (W.A.) . 3. C. congener

Fruits narrowly to broadly ellipsoid or narrowly ovoid, 4-10 mm long,

1- 8 mm across (including indumentum), < 5 t im es longer than wide. 12

12. Fruits ellipsoid to broadly ellipsoid, 4-8 mm across (including

indumentum), 4 or 5-valved; inflorescences 2 or 3-flowered; pedicels

2- 7 mm long; adaxial surface of leaf laminae glabrous or with a sparse

indumentum (epidermis clearly visible) (N.T., Qld).6. C. elderi

Fruits narrowly ellipsoid or narrowly ovoid, 1-4 mm across (including

indumentum), 3(rarely 4)-valved; inflorescences 3-7-flowered; pedicels

1-2 mm long; adaxial surface of leaf laminae with a moderately dense

to dense indumentum (epidermis not visible) (W.A.) . 5. C. elachocarpus

Halford, Notes on Tiliaceae 4, Corchorus 583

13. Fruits obtusely angled in transverse section, usually <10 times as long as

wide, not constricted between the seeds. 14

Fruits circular in transverse section, mostly 10 or more times as long as

wide, slightly or markedly constricted between the seeds. 15

14. Leaf laminae ovate, 2-5 cm wide, l:w ratio <.3:1, margin serrate to serrulate;

sepals 9-15 mm long; petals 9-10 mm long, 4-7 mm wide (W.A.) ... 14. C. puberulus

Leaf laminae narrowly ovate, 0.8-2.5 cm wide, l:w ratio 3-7:1, margin

serrulate; sepals 6-9 mm long; petals 6-7 mm long, 2-4 mm wide

(N.T.) .1. C. aulacocarpus

15. Fruits < 15 mm long. 16

Fruits >15 mm long. 18

16. Shrubs to 2 m high; branchlets erect; leaf laminae narrowly ovate, 3-9 cm

long; fruits obclavate (N.T.) .12. C. obclavatus

Shrubs mostly to 1 m high; branchlets spreading or if erect then leaf laminae

oblong or oblong-elliptic, 0.6-3.5 cm long or if ovate and > 3.5 cm long

then fruits cylindrical. 17

17. Indumentum on young shoots and buds ferruginous or if greyish white

then either abaxial surface of sepals obscured by dense indumentum or

leaf laminae narrowly ovate to ovate and > 1.5 cm wide; stamens > 20

(W.A., N.T., Qld) . 17. C. sidoides

Indumentum on young shoots and buds greyish white, or if ferruginous

then abaxial surface of sepals not completely obscured by indumentum;

leaf laminae oblong, oblong-elliptic or narrowly elliptic and <1.5 cm

wide; stamens mostly < 20 (W.A., N.T., Qld). 15. C. pumilio

18. Sepals > 12 mm long (W.A.). 10. C. leptocarpus

Sepals <12 mm long. 19

19. Apex of mature fruits erect or ascending. 20

Apex of mature fruits orientated downward. 22

20. Stellate hairs up to 1.5 mm across; leaves soft to the touch; fruits 3-4 mm

across (including indumentum) (W.A.). 11. C. mitchellensis

Stellate hairs up to 0.5 mm across; leaves felt-like to the touch; fruits

1-2.5 mm across (including indumentum) . 21

21. Peduncles 2-5 mm long (N.T.) . 19. C. sublatus

Peduncles 7-15 mm long (Qld). 18. C. subargentus

22. Sepals 9-11 mm long, 2-3 mm wide; flower buds 3-6 mm across;

indumentum on young shoots ferruginous; fruits 2-3 mm across, not

conspicuously constricted between seeds; stamens > 80 (W.A.) ... 2. C. carnarvonensis

Sepals < 9 mm long, 1-2 mm wide; flower buds 1-3 mm across;

indumentum on young shoots white or if ferruginous then fruits < 2 mm

across and slightly to markedly constricted between seeds; stamens <70. 23

23. Leaf laminae ovate, 1.3-2 times as long as wide; indumentum moderately

dense on adaxial surface of leaf (epidermis clearly visible); fruits

0.7-1.5 mm across (including indumentum) (Qld). 21. C. tomentellus

Leaf laminae oblong, oblong-elliptic, ovate-elliptic or if ovate then more

than 2.5 times as long as wide and with a dense indumentum (epidermis

not visible); fruits 1-2.5 mm across (including indumentum) (W.A.,

N.T., Qld) .

17. C. sidoides

584

Austrobaileya 6 (4): 581-629 (2004)

1. Corchorus aulacocarpus Halford sp. nov.

similis C. leptocarpo autem fructibus

latioribus (3-4 mm latis comparitis 2-3

mm latis) minus quam 10 plo longioribus

quam latis, in sectione transversali

trigonis vel tetragonis non circularibus,

inter semina non constrictis differt.

Corchorus aulacocarpus floribus ex

sepalis 6-9 x 1-2 mm, petalis 6-7 x 2-4

mm, filamentis staminalibus 3-4 mm

longis, stylo 2-4 mm longo compositis,

pedunculis 2-3 mm longis, foliis 0.8-2.5

cm latis praeditus ut videtur maxime arete

affinis C. puberulo qui flores majores ex

sepalis 9-15 x 2-3 mm, petalis 9-10 x

4-7 mm, filamentis staminalibus 4-6 mm

longis, stylo 4-6 mm longo compositos,

pedunculos longiores 2-4 mm longos,

folia interdum latiores 2-5 lata habet.

Typus: Northern Territory. Darwin and

Gulf Region: Mt Basedow Range, 1 June

1973, T.G. Hartley 13886 (holo: DNA;

iso: CANB).

Shrub to 2 m high; stems sparingly branched,

erect; young shoots with ferruginous

indumentum. Indumentum on branchlets,

leaves, stipules, peduncles, pedicels and bracts

greyish-white, dense, comprised of stellate

hairs. Stellate hairs sessile or stipitate, up to

0.2 mm across; stipes red-brown, straight, up

to 0.2 mm long; rays pliable, white or

ferruginous, up to 0.1 mm long. Leaves with

petioles 5-12 mm long; stipules linear, 3-5 mm

long; lamina narrowly ovate, 4-10 cm long,

0.8-2.5 cm wide, l:w ratio 3-7:1, discolorous;

base rounded or rarely cordate; margin

serrulate; apex acute or rarely obtuse.

Inflorescences umbellate, 5-9-flowered, leaf-

opposed or lateral, solitary at upper nodes;

peduncles 2-3 mm long; pedicels 2-4 mm long;

bracts filiform-linear, 3-4 mm long. Flower

buds obovoid-ellipsoid, 4-5 mm across,

longitudinally ridged; apex obtuse with 5

spreading caudae to 1 mm long. Sepals 5, not

persistent, narrowly obovate, 6-9 mm long,

1-2 mm wide; abaxial surface with a

moderately dense to dense indumentum of

stellate hairs up to 0.5 mm long; adaxial surface

glabrous or sometimes sparsely stellate-

pubescent proximally; apex acuminate-caudate,

up to 1 mm long. Petals 5; lamina narrowly

obovate to obovate, 6-7 mm long, 2-4 mm

wide, glabrous; claw c. 1 mm long, stellate-

pubescent on margins. Androgynophore

0.2-0.3 mm long; annulus sinuate or entire,

0.2-0.3 mm long, glabrous. Stamens 45-60;

filaments 3-4 mm long; anthers c. 0.5 mm long.

Ovary subcylindrical, 0.7-1 mm across, densely

stellate-puberulous, 3(rarely 4)-locular, with

26-32 ovules in each locule; style 2-4 mm long.

Fruits on recurved pedicels, subcylindrical,

8-18 mm long, 3-4 mm across, 2-6 times

longer than wide, curved or straight, 3 or 4-

sided, obtusely-angled in transverse section,

longitudinal sulcate, not constricted between

seeds, 3(rarely 4)-valved; apex obtuse or

rounded, orientated upward; indumentum

moderately dense to dense, of stellate hairs up

to 0.3 mm long. Seeds compressed obovoid or

columnar, 1-2 mm long. Fig. 1.

Additional specimens : Northern Territory. Darwin and

Gulf Region: c. 12 miles [c. 19 km] S [of] Mt Brockman, Jul

1972, Byrnes 2713 (CANB, DNA); Mt Basedow, 20 km SE

of Cooinda, Jun 1980, Craven 6310 (CANB, DNA); Kakadu

NP, Upper Koolpin Creek, Jun 1988, Russell-Smith 5509 &

Lucas (BRI, DNA); 1/4 mile [c. 0.4 km] SW [of] El Sharana,

Jan 1973, Martensz & Schodde AE451 (BRI, CANB); 4

mi les [c. 6 km] NW [of] El Sharana, Pine Creek road, Jan

1973, Martensz & Schodde AE503 (BRI, CANB); 2 km SE

of El Sharana Mine, Apr 1992, Halford Q1172 (BRI); 10

miles [c. 16 km] ESE [of] Noranda Mining Camp, Jun 1972,

Schodde AE60 (CANB, DNA).

Distribution and habitat: Corchorus

aulacocarpus is confined to Arnhem Land,

Northern Territory from Mt Brockman

southwards to El Sharana (Map 4). It is

recorded as growing in woodland communities

with spinifex groundcover on shallow sandy

soils, and in Aliosyncarpia forest on talus slopes

and on broken sandstone ridges.

Phenology : Flowers have been recorded from

January and June, fruits from January, April,

June and July.

Affinities: Corchorus aulacocarpus is similar

to C. leptocarpus but differs from that by having

broader fruits (3-4 mm across compared with

2-3 mm across) which are < 10 times as long

as wide, trigonous or tetragonous rather than

circular in transverse section and are not

constricted between the seeds. Corchorus

aulacocarpus seems most closely related to C.

puberulus but differs from that by having

smaller flowers (sepals 6-9 x 1-2 mm, petals

6-7 x 2-4 mm, staminal filaments 3-4 mm

Halford, Notes on Tiliaceae 4, Corchorus

585

Fig. 1. Corchorus aulacocarpus. A. branchlet with fruit, x 1. B. fruit, x 4. C. ventral view of sepal, x 8. A, B from Craven

6310 (DNA); C from Hartley 13886 (DNA). Del. W. Smith.

long, style 2-4 mm long compared with sepals

9-15 x 2-3 mm, petals 9-10 x 4-7 mm,

staminal filaments 4-6 mm long, style 4-6 mm

long), shorter peduncles (2-3 mm long

compared with 5-7 mm long) and generally

narrower leaves (0.8-2.5 cm wide compared

with 2-5 cm wide).

Etymology : The specific epithet refers to the

longitudinal furrows on the fruit of the species;

Greek aulacos furrowed, karpos fruit.

2. Corchorus carnarvonensis Halford sp. nov.

videtur arete affinis C. sidoidi et C.

congenero, ab illo alabastris majoribus

(3-6 mm diam. non 1-3 mm diam.),

sepalis majoribus (9-11 x 2-3 mm non

3- 8 x 1-2 mm), petalis majoribus (6-9 x

4- 5 mm non 2-7 x 0.5-4) ab hoc fructibus

interdum grandioribus (15-35 x 2-3 mm

non 10-18 x 1-2 mm), pilis tantum

stellatis non stellato-dendriticis et stellatis

vesititis, sepalis majoribus (8-11 x 2-3

586

non 1-18 x 1-2 mm) differt. Typus:

Western Australia. Carnarvon District:

near Carnarvon, 22 August 1967, A.M.

Ashby 2321 (holo: PERTH; iso: AD).

Shrub to 0.8 m high; stems much branched,

spreading; young shoots with ferruginous

indumentum. Indumentum on branchlets,

leaves, stipules, peduncles, pedicels and bracts

grey-white, dense, comprised of mostly stellate

hairs but dendritic-stellate hairs also

occasionally present. Stellate hairs sessile or

stipitate, up to 0.5 mm across; stipes red-brown,

straight up to 0.4 mm long; rays firm, white or

ferruginous, up to 0.3 mm long. Leaves with

petioles 7-15 mm long; stipules subulate-linear,

4-5 mm long; lamina narrowly oblong or

narrowly ovate, 2.5-6 cm long, 0.9-2.4 cm

wide, l:w ratio 2.5-3.5:1, concolorous; base

rounded; margin serrate; apex acute to obtuse.

Inflorescences umbellate, 4-6-flowered, leaf-

opposed or lateral, solitary at upper nodes;

peduncles 3-5 mm long; pedicels 5-7 mm long,

spreading to erect in flower, spreading to

recurved in fruit; bracts filiform-linear, 2-3 mm

long. Flower buds obovoid-ellipsoid, 3-6 mm

across; apex obtuse with 5 spreading caudae to

1 mm long. Sepals 5, not persistent, narrowly

obovate, 9-11 mm long, 2-3 mm wide; abaxial

surface with a dense indumentum of stellate

hairs up to 0.5 mm long; adaxial surface

stellate-puberulous proximally, glabrous

distally; apex acute or acuminate-caudate, up

to 1 mm long. Petals 5; lamina obovate, 6-9

mm long, 4-5 mm wide, glabrous; claw c. 1

mm long, sparsely stellate-pubescent.

Androgynophore c. 0.2 mm long; annulus

entire, c. 0.2 mm long, glabrous or with

scattered minute simple hairs. Stamens 100-120;

filaments 4-6 mm long; anthers c. 0.6 mm long.

Ovary cylindrical, c. 0.5 mm across, densely

stellate-puberulous, 3(rarely 4)-locular, with

20-28 ovules in each locule; style c. 4 mm long.

Fruits subcylindrical, 15-35 mm long, 2-3 mm

across, 5-15 times longer than wide, curved,

circular in transverse section, slightly

constricted between seeds, 3 (rarely 4)-valved;

apex rounded to obtuse, orientated downward;

indumentum dense, of stellate hairs up to 0.5

mm long. Seeds compressed obovoid, 1-2 mm

long. Fig. 2.

Austrobaileya 6 (4): 581-629 (2004)

Additional specimens: Western Australia. Carnarvon

District: 170 km N of Carnarvon junction, Oct 1989,

Nordenstam &Anderberg 273 (PERTH); 19.7 km S of Coral

Bay turnoff, Aug 1977, Chinnock 3817 (AD); just N of One

Mile Jetty, Carnarvon, Aug 1986, Chinnock 6772 (AD);

Gascoyne R. flats, Carnarvon, May 1962, Aplin 1556

(PERTH); Browns Range, near Carnarvon, Sep 1967,

Haw son 9 (PERTH); c. 11 km SE of Carnarvon on Geraldton

road, adjacent Motor Cycle Club course, Oct 1983, Forbes

1573 (MEL).

Distribution and habitat: Corchorus

carnarvonensis occurs in the north west of

Western Australia from near Coral Bay

southwards to Carnarvon. A single specimen

of this species is labelled as originating from

the Port Hedland area ( Runich

[PERTH1523708]). This locality is outside the

‘normal’ range of this species. The locality is

considered to be doubtful and has not been

mapped (Map 3). The species is recorded as

growing in shrubland communities, in sandy

soils, on plains and river flats.

Phenology: Flowers and fruits have been

collected from August to October.

Affinities: Corchorus carnarvonensis seems

most closely related to C. sidoides and

C. congener. It can be distinguished from

C. sidoides by its larger floral buds (3-6 mm

across compared with 1-3 mm across), and

larger sepals and petals (sepals 9-11 x 2-3 mm

compared with 3-8 x 1-2 mm; petals 6-9 x 4-5

mm compared with 2-7 x 0.5-4 mm). For

features distinguishing C. carnarvonensis from

C. congener see ‘Affinities’ section under that

species.

Etymology: The specific epithet is derived from

the name Carnarvon, plus the suffix -ensis

indicating place of origin, alluding to the

Carnarvon Botanical Province in Western

Australia where this species occurs.

3. Corchorus congener Halford sp. nov.

videtur maxime arete affinis C. sidoidi et

C. carnarvonensi; ab utroque fructibus

stellato-dendriticis vestitis (stellatis

tantum in fructibus C. carnarvonensis et

C. sidoidis ), et a C. carnarvonensi sepalis

minoribus (5-8 x 1-2 mm comparitis

9-11 x 2-3 mm), interdum fructibus

minoribus (10-18 x 1.5-2.5 mm

comparitis 15-35 x 2-3 mm). Corchorus

congener quoque similis C. elachocarpo

Halford, Notes on Tiliaceae 4, Corchorus

587

Fig. 2. Corchorus carnarvonensis. A. branchlet with flowers and immature fruit, x 2. B. fruit, x 3. C. ventral view of sepal,

x 6. A from Chinnock 3817 (AD); B, C from Ashby 2321 (AD). Del. W. Smith.

autem fructibus cylindricis non anguste

ellipsoideis vel anguste ovoideus pilis

stellato-dendriticis usque 1.5 mm longis

non usque 3 mm longis vestitis differt.

Typus: Western Australia. Fortescue

District: N of Yardie Creek, 27 May

1965, AS. George 6671 (holo: PERTH).

Corchorus interstans Halford ms,

Paczkowska & Chapman (2000).

Shrub to 0.6 m high; stems much branched,

spreading; young shoots with grey-white or

ferruginous indumentum. Indumentum on

branchlets, leaves, stipules, peduncles, pedicels

and bracts grey-white, moderately dense to

dense, comprised of stellate hairs. Stellate hairs

sessile or stipitate, up to 0.5 mm across; stipes

red-brown, straight, up to 0.5 mm long; rays

pliable, white or ferruginous, up to 0.3 mm

long. Leaves with petioles 5-10 mm long;

stipules subulate-linear, 3-5 mm long; lamina

narrowly oblong or narrowly ovate, 1.6-5 cm

long, 0.5-1.2 cm wide, l:w ratio 3-5:1,

discolorous; base rounded to obtuse; margin

serrulate; apex obtuse. Inflorescences

umbellate, 4-6-flowered, leaf-opposed or

lateral, solitary at upper nodes; peduncles 3-7

mm long; pedicels 1-5 mm long, spreading to

erect in flower, spreading to recurved in fruit;

588

bracts filiform-linear, 1-3 mm long. Flower

buds obovoid-ellipsoid, 3-4 mm across; apex

acute with 5 erect caudae to 0.2 mm long.

Sepals 5, not persistent, narrowly obovate-

elliptic, 5-8 mm long, 1-2 mm wide; abaxial

surface with a moderately dense to dense

indumentum of stellate hairs up to 0.5 mm long;

adaxial surface glabrous except for a few

scattered stellate hairs proximally; apex acute

or acuminate-caudate, up to 1 mm long. Petals

5; lamina obovate, 5-7 mm long, 3-5 mm wide,

glabrous; claw 0.6-0.7 mm long, stellate-

pubescent on margins. Androgynophore c. 0.3

mm long; annulus entire, c. 0.2 mm long,

glabrous or with scattered minute simple hairs.

Stamens 45-65; filaments 3-5 mm long;

anthers c. 0.5 mm long. Ovary trigonal-

cylindrical, c. 0.7 mm across, densely stellate

puberulous, 3-locular, with 14-18 ovules in

each locule; style 3-4 mm long. Fruits

subcylindrical, 10-18 mm long, 1.5-2.5 mm

across, 5-12 times longer than wide, curved, ±

circular in transverse section, not conspicuously

constricted between seeds, 3-valved; apex

rounded to obtuse; indumentum dense, of

stellate and dendritic-stellate hairs; dendritic-

stellate up to 1.5 mm long, with stipes pale

yellow, tortuous; rays white, up to 0.2 mm long,

pliable. Seeds compressed obovoid, 1-2 mm

long. Fig. 3.

Selected specimens (from 11 examined): Western

Australia. Fortescue District: Barrow Island, Nov 1965,

Clay & Yardar s.n. [PERTH 1522051] (PERTH). Carnarvon

District: W side of Cape Range, ± 1 mile [c. 1.6 km] S of

lighthouse, Jun 1961, George 2563 (PERTH); Exmouth, Oct

1975, Weber 4992 (AD); + 6 miles [c. 10 km] E of Ningaloo

Station Homestead, Sep 1970, George 10229 (PERTH); 5—

6 miles [c. 8-10 km] S of Exmouth, May 1965, George 6604

(PERTH); Rough Range, c. 7 km by road S of Exmouth

Homestead on main Exmouth-Camarvon road, Aug 1977,

Barker 2134 (AD, MEL, NSW).

Distribution and habitat: Corchorus congener

is confined to north-west of Western Australia,

from Ningaloo Station north-east to Barrow

Island (Map 7). It is recorded as growing in

open shrubland and hummock grassland

communities, mostly on sandy plains and sand

dunes but also on loamy soils derived from

limestone.

Phenology : Flowers have been collected from

April to June and August to November, fruits

in May, August, October and November.

Austrobaileya 6 (4): 581-629 (2004)

Affinities': Corchorus congener seems most

closely related to C. sidoides and

C. carnarvonensis but differs from both by

having dendritic-stellate hairs on the fruit (only

stellate hairs present on the fruits of

C. carnarvonensis and C. sidoides ). In addition,

C. congener differs from C. carnarvonensis by

having smaller sepals (5-8 x 1-2 mm compared

with 9-11 x 2-3 mm), and generally smaller

fruit (10-18 x 1.5-2.5 mm compared with 15-35

x 2-3 mm). Corchorus congener is also similar

to C. elachocarpus but differs from that by

having cylindrical rather than narrowly

ellipsoid or narrowly ovoid fruit with dendritic-

stellate hairs up to 1.5 mm rather than up to 3

mm long.

Notes: The collection Donnell [MEL1599012]

(MEL) from “near the Ord River”, is noted here

because it is similar to C. congener in having

narrowly oblong leaves and 3-valved

subcylindrical fruits with an indumentum of

stellate and dendritic-stellate hairs. However,

it differs from C. congener in having smaller

flowers and a sparser indumentum on its

branchlets and leaves. It is also somewhat

disjunct from the known populations of

C. congener in north-west Western Australia.

The Donnell collection would appear to

represent an undescribed taxon, however,

further collections are required before it is

formally recognised.

Etymology: The specific epithet alludes to the

similarity of this species to C. carnarvonensis ;

Latin congener of the same kind.

4. Corchorus crozophorifolius (Baill.) Burret,

Notizbl. Bot. Gart, Berlin 12: 166 (1934),

(‘ chrozophorifolius , ); Nettoa crozophorifolia

Baill., Adansonia 6: 238 t.7 (1866). Type:

[Western Australia.] Nova Holland, ile sterile,

Leschenault [Baudin Expedition] (lecto, here

chosen: P; isolecto: P).

Corchorus crassifolius Domin, Biblioth. Bot.

89: 384 (1928). Type: [Western

Australia.] upper Murchison River near

Mt Hall, 1884, C. Crossland (holo: K;

iso: MEL).

Shrub to 1 m high; stems much branched,

spreading to erect; young shoots with

ferruginous indumentum. Indumentum on

Halford, Notes on Tiliaceae 4, Corchorus

589

Fig. 3. Corchorus congener. A. branchlet with flower buds and fruit, x 1.5. B. fruit, x 3. C. ventral view of sepal, x 8. D. cross-

section of sepal, x 36. A-D from George 6671 (PERTH). Del. W. Smith.

branchlets, leaves, stipules, peduncles, pedicels

and bracts ferruginous or rarely grey-white,

very dense, floccose, comprised of mostly

dendritic-stellate and stellate hairs but simple

hairs also present. Stellate hairs sessile or

stipitate, up to 0.9 mm across; stipes red-brown

or white, straight, up to 0.2 mm long; rays firm

to pliable, ferruginous or white, up to 0.5 mm

long. Dendritic-stellate hairs up to 4 mm long;

stipes red-brown or pale yellow, tortuous; rays

firm to pliable, white or ferruginous, up to 0.8

mm long. Simple hairs glandular, white,

flexuous, up to 0.3 mm long. Leaves with

petioles 10-25 mm long; stipules subulate-

linear, 4-20 mm long; lamina ovate, (3-)4-9

cm long, 2-6 cm wide, l:w ratio 2-4:1,

concolorous; base rounded; margin dentate-

serrate; apex acute to rounded. Inflorescences

umbellate, 4-15-flowered, leaf-opposed,

solitary at upper nodes; peduncles 5-20 mm

long; pedicels 2-12 mm long, spreading to erect

in flower and fruit; bracts subulate-linear,

5-12 mm long. Flower buds ellipsoid, 6-10 mm

across; apex obtuse occasionally with 5

spreading caudae to 1 mm long. Sepals 5,

persistent, narrowly obovate-elliptic, 10-18

mm long, 2-4 mm wide; abaxial surface with

a very dense indumentum of dendritic-stellate

and stellate hairs, the largest hairs up to 2 mm

long; adaxial surface stellate-villose proximally,

glabrous distally; apex acute or caudate, up to

6 mm long. Petals 5; lamina obovate, 8-11 mm

590

long, 3-6 mm wide, glabrous; claw 1-1.5 mm

long, stellate-villose on abaxial surface and

margins. Androgynophore 0.5-0.7 mm long;

annulus entire, c. 0.5 mm long, densely stellate-

pubescent. Stamens 50-100; filaments 3-8 mm

long; anthers c. 0.8 mm long. Ovary ovoid to

cylindrical, densely stellate-tomentose, 3 to 7-

locular, with 16-30 ovules in each locule; style

c. 5 mm long. Fruits ovoid, 12-17 mm long,

10-12 mm across, 1-1.7 times longer than

wide, circular in transverse section, 4 to 6-

valved; apex rounded; indumentum dense, of

dendritic-stellate and stellate hairs, the largest

hairs c. 4 mm long. Seeds compressed obovoid,

c. 2 mm long.

Selected specimens (from 52 examined): Western

Australia. Fortescue District: 7 km NW of Quarry Hill,

c.130 km W of Tom Price, Jul 1984, Newbey 10586

(PERTH); 33 km SE of Mt Bruce, May 1980, Houston s.n.

[PERTH 1522892] (PERTH). Carnarvon District: 120 km

S of Onslow, May 1962 Aplin 1603 (MEL, PERTH); near

Carnarvon, Jul 1965, Ashby 1563 (AD, CANB); 7 km N of

Gascoyne Junction, Oct 1984, Mitchell 1312 (PERTH); 16

km N of Gascoyne Junction, Sep 1987, Green 5391

(PERTH); 166 km SSE of Carnarvon, on North West Coastal

Highway, Aug 1965, Beauglehole ACB11795 (PERTH);

Woodleigh Station, E of Perth-Camarvon road, Sep 1959,

Burbidge 6366 (PERTH); 30 miles [c. 48 km] E of Hamelin

Pool, Aug 1931, Gardner 2542 (PERTH). Ashburton

District: Barlee Range, Henry R., Aug 1961, Royce 6599

(PERTH); 50 km NW of Cobra Homestead on Gascoyne

Junction to Mt Augustus road near Lyons R., Jul 1986,

Conrick 9851 (MEL); Mount Sandiman Station, Aug 1969,

Wilcox 40 (PERTH); Mt Augustus, Aug 1970, Ashby 3372

(AD, MEL). Austin District: near Mount Gould, flumen

Murchison, Aug 1963, Gardner 14529 (PERTH); 20.7 miles

[c. 33 km] N of Belele, Jul 1958, Speck 945 (CANB); 12

miles [c. 19 km] SE of Berrugarra [Beringarra], Sep 1957,

Speck 675A(CANB); 10 miles [c. 16 km] W of Mileura on

Nookawarraroad, Jul 1958, Speck 1006 (CANB, MEL); 10

miles [c. 16 km] S of Berrugarra [Beringarra], Jul 1958,

Speck 976 (AD, CANB); 2.2 km ENE of Pepper Tree Bore,

Koonanarra [Koonmarra] Station, Aug 1986, Cranfield 5927

(PERTH); Wiluna area, Dec 1970, Morrissey 51 (PERTH).

Distribution and habitat: Corchorus

crozophorifolius is confined to the north-west

of Western Australia, from Exmouth

southwards to Woodleigh Station (south of

Carnarvon) and east to Wiluna (Map 1). It is

recorded as growing in Acacia shrubland and

woodland communities, on sandy soils on

alluvial flats and along watercourses, and on

skeletal soils derived from limestone or granite

on rocky rises and hills.

Phenology: Flowers have been collected from

May to November, fruits from July to

November.

Austrobaileya 6 (4): 581-629 (2004)

Typification: Baillon (1866) cited a collection

from the Baudin expedition in the protologue

of Nettoa crozophorifolius. I have seen two

sheets of original material of

N. crozophorifolius from the Baudin expedition

on loan to BRI from R The sheet with the hand

written label “(Leschenault legit!) Tiliaceae,

Nova Holland ile sterile” is selected as lectotype

because it agrees with the original description

and the fragment at the top left of the sheet

matches the drawing with Baillon’s original

description.

Notes: Corchorus crozophorifolius is a

distinctive species with its floccose

indumentum on the stems, petioles and

inflorescences, and its densely hairy annulus.

There are two indumentum colour forms. The

typical and more widespread form has an

indumentum of ferruginous hairs on the young

shoots and buds. The other form has an

indumentum of white hairs (eg. Newbey 10586

(PERTH), Aplin 1603 (MEL, PERTH) and

Ashby 4143 (AD, PERTH)) and is found mostly

in the northern part of the species range. The

white form of C. crozophorifolius may be

confused with C. incanus but is easily

distinguished by having a densely hairy

annulus.

The collections Weber 492?, (AD), George

1362 (PERTH), McWhae s.n.

[PERTH 1522825] and George 1343 (PERTH)

from Cape Range near Learmonth resemble

C. crozophorifolius in their indumentum and

floral morphology but differ by having

cylindrical fruits (15-20 mm x 4-5 mm) and a

much finer serration on the leaf margin. This

variant needs to be investigated further and may

be worth recognising at least at a subspecific

rank if not as a distinct species.

5. Corchorus elachocarpus F.Muell., Fragm.

8: 6 (1872). Type: [Western Australia.]

Nichol Bay, P. Walcott (lecto: MEL

[MEL223670 l fide B. Rye, Nuytsia 9(3):

418 (1994)).

Shrub to 0.6 m high; stems sparingly to much

branched, spreading; young shoots with grey-

white or rarely ferruginous indumentum.

Indumentum on branchlets, leaves, stipules,

peduncles, pedicels and bracts grey-white,

moderately dense to dense, comprised of stellate

Halford, Notes on Tiliaceae 4, Corchorus

591

hairs. Stellate hairs sessile or sometimes

stipitate, up to 0.6 mm across; stipes red-brown,

straight, up to 0.1 mm long; rays pliable, white,

up to 0.4 mm long. Leaves with petioles 4-13

mm long; stipules subulate-linear, 2-3 mm

long; lamina narrowly oblong, 2-5 cm long,

0.4-1.3 cm wide, l:w ratio 2-5:1, concolorous;

base rounded; margin serrulate; apex obtuse.

Inflorescences umbellate, 3-7-flowered, leaf-

opposed, solitary at upper nodes; peduncles

2- 4 mm long; pedicels 1-2 mm long, spreading

to erect in flower, spreading to recurved in fruit;

bracts subulate-linear, 1-2 mm long. Flower

buds globose, 2-3 mm across; apex obtuse with

5 erect caudae to 0.5 mm long. Sepals 5, not

persistent, narrowly obovate-elliptic, 4-6 mm

long, 1-2 mm wide; abaxial surface with a

dense indumentum of stellate hairs up to 0.5

mm long; adaxial surface glabrous or with a

few scattered stellate hairs proximally; apex

shortly caudate, up to 1 mm long. Petals 5;

lamina obovate to broadly obovate, 3-5 mm long,

3- 4 mm wide, glabrous; claw 0.5-0.7 mm long,

sparsely stellate-pubescent. Androgynophore

0.5-0.8 mm long; annulus entire, c. 0.2 mm long,

glabrous. Stamens 27-42; filaments 3-4 mm

long; anthers c. 0.5 mm long. Ovary ovoid,

1-1.5 mm across, densely stellate-tomentose,

3(rarely 4)-locular, with 6-8 ovules in each

locule; style 3-4 mm long. Fruits narrowly

ellipsoid or narrowly ovoid, 4-10 mm long,

1-4 mm across, 2.5-4 times longer than wide,

not conspicuously constricted between seeds,

circular in transverse section, 3(rarely 4)-

valved; apex rounded to obtuse; indumentum

dense, of dendritic-stellate and stellate hairs,

the largest hairs up to 3 mm long. Seeds

obovoid, c. 2 mm long. Chromosome No. 2n =

14 (Islam & Qaiyum 1961).

Selected specimens (from 13 examined): Western

Australia. Fortescue District: 30-40 km S of Port Hedland,

Jun 1982, Glennon 76 (PERTH); Warralong, Aug 1941,

Burbidge s.n. [PERTH1532278] (PERTH); Muccan Station,

De Grey R., Jun 1941, Burbidge 966 (PERTH); 4.5 km W

of “Warrawagine” Homestead, c. 65 km SE of Shay Gap, Jul

1984, Newbey 10541 (CANB, PERTH); Main Shay Gap-

Marble Bar road, c. 6 km by road SW of turnoff to Kittys

Gap Well, Aug 1977, Barker 2084 (AD); 3 miles [c. 5 km] S

of Stag Arrow Creek, May 1947, Royce 1703 (PERTH); 4

miles [c. 6 km] N of Stag Arrow Creek, May 1947, Royce

1715 (PERTH). Carnarvon District, c. 15 km N of crossing

from North West Coastal Highway towards Yanrey, Oct 1975,

Weber 4890 (PERTH); Marilla Station complex, Oct 1984,

Stretch s.n. [PERTH 1532243] (PERTH); Carnarvon

Geraldton road, Sep 1968, Baird s.n. [PERTH 1532723]

(PERTH).

Distribution and habitat : Corchorus

elachocarpus is confined to the Pilbara region

of Western Australia from near Yanrey Station

eastwards to Warrawagine Station and Stag

Arrow Creek (Map 6). It is recorded as growing

in hummock grassland and open shrubland

communities, on sandy or sandy clay soils on

flats.

Phenology : Flowers have been collected from

May to October, fruits in May, June, August

and October.

Notes: The collections Mitchell PRP1462 (BRI)

and Mitchell PRP1135 (BRI) from Marrillana

Station near Newman represent an undescribed

entity closely related to C. elachocarpus. It

differs from C. elachocarpus in having

narrowly ovate leaves, longer peduncles and

pedicels, and broader fruit. This entity warrants

formal recognition. However, more material is

required before this can be undertaken.

The collection George [PERTH1532251]

(PERTH) from the Great Sandy Desert, is noted

here because it is similar to C. elachocarpus

but differs in having much larger leaves, fruits

and flowers. Even from this single specimen,

it is clear that the entity it represents warrants

formal recognition. However, more material is

required before this can be undertaken.

6. Corchorus elderi F.Muell., Trans. & Proc.

Roy. Soc. South Australia 9: 58 (1887).

Type: [Northern Territory.] N of

Macdonell Range, 1886, Lt Dittrich

(lecto, here chosen: MEL [MEL223672];

isolecto: AD [AD95836005], MEL

[MEL223671]).

Shrub to 0.4 m high; stems much branched,

spreading to erect. Indumentum on young

shoots, branchlets, stipules, peduncles, pedicels

and bracts + white, sparse to moderately dense,

comprised of stellate hairs. Stellate hairs sessile,

up to 0.6 mm across; rays pliable, white, up to

0.4 mm long. Leaves with petioles 2-13 mm

long; stipules subulate-linear, 1-3 mm long;

lamina narrowly oblong-elliptic, 1-4 cm long,

0.5-1.5 cm wide, concolorous; adaxial surface

glabrous or sparsely stellate hairy; abaxial

surface moderately dense to densely stellate

hairy; base obtuse to rounded; margin serrate

or serrulate; apex obtuse. Inflorescences

592

umbellate, 2 or 3-flowered, lateral, solitary at

nodes; peduncles 1-5 mm long; pedicels 2-7

mm long, spreading to erect in flower and fruit;

bracts subulate-linear, 1-2 mm long. Flower

buds broadly obovoid, 2-4 mm across; apex

obtuse or shortly acuminate. Sepals 5, not

persistent, narrowly obovate-elliptic, 6-7 mm

long, 1-2 mm wide; abaxial surface with a

moderately dense indumentum of stellate hairs

up to 0.3 mm long; adaxial surface glabrous;

apex acute or shortly caudate, up to 0.5 mm

long. Petals 5; lamina obovate, 5-7 mm long,

2-4 mm wide, glabrous; claw c. 1 mm long,

stellate-pubescent on margins. Androgynophore

0.2-0.5 mm long; annulus undulate, c. 0.2 mm

long, glabrous. Stamens 40-60; filaments 3-4

mm long; anthers c. 0.5 mm long. Ovary ovoid,

1.5-2 mm across, densely stellate-tomentose,

5 or 6-locular, with c. 20 ovules in each locule;

style 2-3 mm long. Fruits ellipsoid to broadly

ellipsoid, 5-10 mm long, 4-8 mm across,

1.2-2.4 times longer than wide, not

conspicuously constricted between seeds,

circular in transverse section, 4 or 5-valved;

apex rounded; indumentum dense, of stellate

and dendritic-stellate hairs up to 1.5 mm long.

Seeds compressed obovoid, 1-2 mm long.

Selected specimens (from 12 examined): Northern

Territory. Central Northern Region: Trew Bore, Elkedra

Station, Dec 1986, Strong 947 (DNA); 5 mi les [c. 8 km] W

of Tarlton Downs Homestead, Feb 1968, Latz 156 (AD, BRI,

DNA, MEL, NSW); Plenty Highway, 72 km E of Arthur R„

May 1988, Thomson 2433 (DNA). Central Southern

Region: sandy flat on western side of Hay R., c. 14 km SSE

of Mt Winnecke, Jul 1982, Purdie 2323 (CANB, DNA); Lake

Caroline, Oct 1986, Leach 1039 (BRI). Queensland.

Gregory North District: sandplain at mouth of Toko Gorge,

Toko Range, Jul 1982, Purdie 2285 (BRI, CANB); levee

adjacent to Burke R., 2 km S of Boulia, Sep 1978, Purdie

1328 (BRI).

Distribution and habitat: Corchorus elderi is

not a common species over its range from the

Davenport Ranges, Northern Territory

eastwards to Boulia, in western Queensland

(Map 1). It is recorded as growing in low to

tall open Acacia shrubland or open eucalypt

woodland communities, on red sandy soils on

flats or ridges.

Phenology : Flowers have been collected in

February, May, July and from September to

December, fruits in May, July and from

September to December.

Austrobaileya 6 (4): 581-629 (2004)

Typification: In the protologue of Corchorus

elderi , Mueller (1887) cited a collection made

by Lieut. Dittrich from a region north of the

MacDonnell Ranges. Three sheets (two at MEL

[MEL223671; MEL223672] and one sheet at

AD [AD95836005]) of Dittrich’s collection

from this area have been located. The MEL

sheet marked MEL223672 is here chosen as

the lectotype as it is the better preserved of the

three specimens seen.

7. Corchorus incanus Halford sp. nov. similis

C. walcottii et C. lanifloro. Ab utroque

pilis conspicuis simplicibus glandularibus

deficientibus differt. Addite ab illo lobis

calycis persistentibus in fructibus, pilis

longioribus (3-5 non usque 1.5 mm

longis) in fructibus et in pagina abaxiali

sepalium differt. Corchorus incanus

confunderi potest variantia albiflora C.

crozophorifolii autem ab illo annulo

glabro pubescente distinguendus. Typus:

Western Australia. Fortescue District:

Great Northern Highway-Shellborough

track, c. 35 km NNW of Goldsworthy, 6

August 1977, I.R. Telford 6524 & G.

Butler (holo: CANB; iso: BRI, PERTH,

distribuendi).

Shrub to 1 m high, sometimes viscid; stems

much branched. Indumentum on young shoots,

branchlets, leaves, stipules, peduncles, pedicels

and bracts grey-white, moderately dense to

dense, comprised of mostly stellate hairs but

dendritic-stellate and simple hairs also

occasionally present. Stellate hairs sessile or

stipitate, up to 2 mm across; stipes red-brown

to white, straight or tortuous, up to 0.8 mm

long; rays soft, white, spreading, up to 1.5 mm

long. Dendritic-stellate hairs up to 1.5 mm

long; stipes red-brown, tortuous; rays soft,

white, up to 0.8 mm long. Simple hairs

glandular, white, flexuous, up to 0.5 mm long.

Leaves with petioles 15-30 mm long; stipules

subulate-linear, 8-20 mm long; lamina ovate to

very broadly ovate, 3-8.5 cm long, 2-7 cm wide,

l:w ratio 1.2-1.6:1, concolorous or slightly

discolorous; base rounded or slightly cordate;

margin serrate; apex rounded to obtuse.

Inflorescences umbellate, 4-7-flowered, leaf-

opposed, solitary at nodes; peduncles 7-25 mm

long; pedicels 2-10 mm long, spreading to erect

in flower and fruit; bracts filiform-linear; 8-25 mm

Halford, Notes on Tiliaceae 4, Corchorus

593

long. Flower buds broadly ellipsoid, 7-10 mm

across; apex obtuse with 5 spreading caudae to

8 mm long. Sepals 5, persistent, narrowly

elliptic, 8-16 mm long, 1-5 mm wide; abaxial

surface with a dense indumentum of mostly

dendritic-stellate hairs but stellate hairs also

present, the largest hairs c. 3 mm long; adaxial

surface stellate-villose proximally, glabrous

distally; apex caudate, 3-6 mm long. Petals 5;

la min a obovate, 6-9 mm long, 4-5 mm wide,

glabrous; claw c. 1 mm long, with stellate-

pubescent margins. Androgynophore 0.1-0.3

mm long; annulus entire, 0.2-0.4 mm long,

glabrous or rarely with scattered hairs. Stamens

100-120; filaments 4-6 mm long; anthers c.

0.5 mm long. Ovary subglobose, 1-2 mm

across, densely stellate-tomentose; 4 or 5-

locular, with 10-24 ovules in each locule; style

2-6 mm long. Fruits subcylindrical or ovoid,

8-17 mm long, 5-10 mm across, 1.6-2.5 times

longer than wide, not conspicuously constricted

between seeds, + circular in transverse section,

4 or 5-valved; apex obtuse; indumentum dense,

of stellate and dendritic-stellate hairs, the

largest hairs c. 5 mm long. Seeds compressed

obovoid, 2-3 mm long.

Affinities: Corchorus incanus resembles

C. walcottii and C. laniflorus but differs from

both by lacking conspicuous simple glandular

hairs. In addition, C. incanus differs from

C. walcottii by having persistent sepals and

longer hairs on the fruits and the abaxial surface

of the sepals (3-5 mm long compared with 1.5

mm long). Corchorus incanus may be confused

with the white form of C. crozophorifolius but

it can be distinguished from that by its more or

less glabrous rather than densely hairy annulus.

Corchorus incanus can be confused with

C. lasiocarpus but differs from that by its

relative shorter leaves and narrower fruits.

Etymology: The specific epithet alludes to the

overall greyish-white appearance of the whole

plant; Latin incanus quite grey, hoary.

Notes: This species occurs in the north-west of

Western Australia, from Hamersley Range, near

Wittenoom northwards to Port Hedland and

Broome with an isolated record from the Rudall

River on the south eastern edge of the Great

Sandy Desert. Two subspecies are recognised

here.

Indumentum on branchlet dense with stellate hairs mostly < 1 mm across;

plants generally growing on sandy soils on sandhills, plains and along

watercourses. 7a. C. incanus subsp. incanus

Indumentum on branchlets moderately dense with stellate hairs mostly

1-2 mm across; plants growing on stony soils along watercourses and

in gorges. 7b. C. incanus subsp. lithophilus

7a. Corchorus incanus Halford subsp. incanus

Shrub to 1 m high, sometimes viscid. Stellate

hairs sessile or stipitate, up to 1.5 mm across;

stipes straight or tortuous, up to 0.8 mm long;

rays up to 1.5 mm long. Dendritic-stellate hairs

up to 1.5 mm long; rays up to 0.8 mm long.

Leaves with petioles 15-25 mm long; lamina

broadly ovate to very broadly ovate, somet im es

ovate, 3.5-8.5 cm long, 2.5-7 cm wide.

Inflorescences 5-7-flowered; peduncles 7-25

mm long; pedicels 2-10 mm long; bracts 8-12

mm long. Sepals 12-16 mm long, 1-3 mm

wide. Petals obovate, 6-9 mm long, 4-5 mm

wide. Fruits subcylindrical or ovoid, 8-15 mm

long, 5-10 mm across; dendritic-stellate hairs

c. 5 mm long. Fig. 4.

Selected specimens (from 23 examined): Western

Australia. Damper District: 1 mile [c. 1 .6 km] N of Broome,

May 1971, Maconochie 1174 (CANB, DNA, MEL, NSW,

PERTH); 407 km from Port Hedland (PO.) along Great

Northern Highway towards Broome, Apr 1992, Telford 11587

(BRI); 7.5 km SW of the 80 Mile Beach turnoff, Sep 1986,

Chinnock 6941 (AD); Nalgi Station, 80 Mile Beach, June

1941, Burbidge 1261 (PERTH); 2 km E of Nita Downs

Homestead, Oct 1984, Foulkes 37 (PERTH); 86 km NE of

Sandfire roadhouse, Great Northern Highway, Sep 1978,

Beauglehole 59257 & Erroy 2957 (DNA); 124 miles [c.

200 km] SW of Anna Plains, SW of Broome, Aug 1965,

Beauglehole ACB11326 (MEL, PERTH). Fortescue

District: Port Hedland, Feb 1983, Rose 2 (PERTH);

Poondarrah Siding, Port Hedland-Marble Bar railway, May

1941, Burbidge 662 (PERTH); Finucane Island, Mar 1981,

Carr B4 (PERTH); Shellborough track, c. 35 km NNW of

Goldsworthy, Aug 1977, Telford & Butler 6524 (PERTH);

Pier Creek, Warralong Station, May 1941, Burbidge 738

(PERTH); Warralong Station, between Shaw and Coongan

594

Austrobaileya 6 (4): 581-629 (2004)

Fig. 4. Corchorus incanus subsp. incanus. A. branchlet with flowers and fruit, x 0.6. B. fruit, x 2. C. ventral view of sepal,

x 4. A-C from Telford 6524 & Butler (BRI). Del. W. Smith.

Rivers, Jun 1941, Burbidge 1217 (PERTH). Keartland

District: nearRudallR.,May 1971 ,George 10780(CANB,

PERTH); Rudall R., Aug 1971, Wilson 10303 (PERTH).

Distribution and habitat: Corchorus incanus

subsp. incanus occurs from Port Hedland to

Broome with an isolated record from Rudall

River on the south-eastern edge of the Great

Sandy Desert (Map 7). It is recorded as growing

in low open shrubland or low open woodland

communities, on sandy or rarely clayey sandy

soils, on sandhills, plains or along

watercourses.

Phenology: Flowers have been collected in

February and from April to September, fruits

in June and from August and September.

7b. Corchorus incanus subsp. lithophilus

Halford subsp. nov. a subspeciebus ceteris

plantis plerumque viscidis, indumento

pilorum stellatorum grossiorum differt.

Crescit in solis saxosis circum flumina

in tempus et saltus in comparatione in

solis arenosis in clivis sabulosis et

planitiis. Corchorus incanus subsp.

lithophilus confunderi potest C. incano

subsp. lasiocarpo foliis ovatis usque late

ovatis non anguste ovatis usque ovatis

marginibus grossiore serratis, fructibus

minoribus (13-17 x 6-7 mm non 15-20

x 12-16 mm) distinguendus. Typus:

Western Australia. Fortescue District:

Hamersley Range NP, Kalamina Gorge,

below car park, 19 Aug 1977, W.R. Barker

1994 (holo: AD).

Corchorus lithophilus Halford ms,

Paczkowska & Chapman (2000).

Corchorus saxicola Halford ms, Paczkowska

& Chapman (2000).

Halford, Notes on Tiliaceae 4, Corchorus

595

Fig. 5. Corchorus incanus subsp. lithophilus. A. branchlet with flower buds and flowers, x 0.6. B. fruit, x 2. C. ventral view

of sepal, x 4. D. dendritic-stellate hair, x 24. A from Craven 7548 (CANB); B-D from Barker 1994 (AD). Del. W. Smith.

Shrub to 1 m high, usually conspicuously

viscid. Stellate hairs sessile or stipitate, up to 2

mm across; stipes straight, up to 1 mm long;

rays up to 2 mm long. Dendritic-stellate hairs

up to 2 mm long; rays up to 1.5 mm long.

Leaves with petioles 10-35 mm long; lamina

ovate to broadly ovate, 3-8 cm long, 2-5.5 cm

wide. Inflorescences 4-6-flowered; peduncles

8-10 mm long; pedicels 7-10 mm long; bracts

10-25 mm long. Sepals 8-16 mm long, 2-5

mm wide. Petals obovate to broadly obovate,

7-8 mm long, 5-7 mm wide. Fruits

subcylindrical, 13-17 mm long, 6-7 mm

across; dendritic-stellate hairs c. 3 mm long.

Fig. 5.

Additional specimens : Western Australia. Fortescue

District: near racecourse, Wittenoom, Oct 1963, Lullfitz

L2758 (PERTH); Wittenoom, Aug/Sep 1957, Elliott s.n.

[PERTH1522574] (PERTH); near Wittenoom, Aug 1967,

Gittins 1480 (NSW, PERTH); Hamersley Range, near

Wittenoom Gorge, May 1958, Burbidge 6001 (PERTH),

Wittenoom Gorge, Sep 1969, Brooker 2218a (PERTH);

Hamersley Range NP, Fig Tree Soak, c. 10 km by road SW

into Yampire Gorge from Wittenoom-Roy Hill road, Aug

1977, Barker 1966 (AD) c. 17 km E of Wittenoom on Roy

Hill to Wittenoom road, Aug 1996, Mitchell PRP1463 (BRI);

15 km E of Wittenoom on the Roy Hill road, Sep 1982,

Craven 7548 (CANB, MEL, PERTH).

Distribution and habitat : Corchorus incanus

subsp. lithophilus occurs in the Hamersley

Range, near Wittenoom (Map 6). It is recorded

as growing in hummock grassland

communities on stony soils and along

watercourses in low open woodland

communities.

Phenology: Flowers have been collected in May

and from August to October, fruits in August.

Affinities : Corchorus incanus subsp.

lithophilus differs from the other subspecies by

having a sparser indumentum of coarser stellate

hairs and its plants are usually conspicuously

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Austrobaileya 6 (4): 581-629 (2004)

sticky. Its habitat differs also as it grows in stony

soils around creeks and gorges rather than

sandy soils on sandhills, plains or along

watercourses. Corchorus incanus subsp.

lithophilus may be confused with

C. lasiocarpus subsp. lasiocarpus but can be

distinguished from that by its ovate to broadly

ovate rather than narrowly ovate to ovate leaves,

coarser serrated leaf margins and smaller fruit

(13-17 x 6-7 mm compared with 15-20 x

12-16 mm).

Etymology : The subspecific epithet refers to

the rocky habitat; Greek lithos stone, philus

loving, fond.

8. Corchorus laniflorus Rye, Nuytsia 9(3): 416

(1994). Type: Western Australia.

Fortescue District: Red Hill, 20 October

1941, C.A. Gardner 6348 (holo: PERTH

n.v.\ iso: CANB n.v.).

Compact shrub to 1(—1.2) m high; ste m s much

branched, spreading. Indumentum on young

shoots, branchlets, leaves, stipules, peduncles,

pedicels and bracts grey-white, dense, woolly,

comprised of mostly stellate and simple hairs

but dendritic-stellate hairs also occasionally

present. Stellate hairs sessile or stipitate, up to

2 mm across; stipes red-brown, tortuous, up to

0.7 mm long; rays soft, white, up to 2 mm long.

Dendritic-stellate hairs up to 5 mm long; stipes

up to 3 mm long; rays soft, white, 1-2.5 mm

long. Simple hairs glandular, dull yellow to red-

brown, flexuous, up to 1.5 mm long. Leaves

with petioles 8-15(-30) mm long; stipules

subulate-linear, 7-17 mm long; la min a ovate

to very broadly ovate, 1.5-5 cm long, 1-4 cm

wide, l:w ratio 1.1-1.7:1, concolorous or

slightly discolorous; base rounded or slightly

cordate; margin serrate or dentate-serrate; apex

obtuse to rounded. Inflorescences umbellate, 5

or 6-flowered, leaf-opposed, solitary at nodes;

peduncles 10-14 mm long; pedicels 5-9 mm

long, spreading to erect in flower and fruit;

bracts filiform-linear, 6-10 mm long. Flower

buds globose, 5-12 mm across; apex obtuse

with 5 spreading caudae to 4 mm long. Sepals

5, persistent, narrowly obovate-elliptic, 10-15

mm long, 2-3 mm wide; abaxial surface with

a very dense indumentum of dendritic-stellate

and simple hairs, the largest hairs up to 5 mm

long; adaxial surface densely stellate hairy or

± glabrous; apex caudate, 4-6 mm long. Petals

5; lamina obovate to very broadly obovate,

6-10 mm long, 5-9 mm wide, glabrous; claw

c. 1 mm long, stellate-pubescent on abaxial

surface, with ciliate margins. Androgynophore

0.4-0.5 mm long; annulus undulate, up to 0.4

mm long, glabrous. Stamens 80-100; filaments

3-4 mm long; anthers c. 0.5 mm long. Ovary

subglobose, c. 1.5 mm across, densely stellate-

tomentose, 3 or 4(rarely 5)-locular, with 6-8

ovules in each locule; style 3-5 mm long. Fruits

narrowly ellipsoid, 7-9 mm long, 3-4 mm

across, 2.2-2.5 times longer than wide, not

conspicuously constricted between seeds,

circular in transverse section, 3 or 4(rarely 5)-

valved; apex acute or rounded; indumentum

dense, of mostly stellate and glandular hairs

but dendritic-stellate hairs occasionally present,

the largest hairs up to 0.8 mm long. Seeds

obovoid, c. 2 mm long. Fig. 6.

Selected specimens (from 19 examined): Western

Australia. Fortescue District: South Fortescue, Jul 1977,

Pfeiffer 21 (PERTH); Red Hill Station, Aug 1970, Beard

6166 (PERTH); Python Pool, foot of Mt Herbert, Oct 1941,

Gardner 6287 (PERTH); 16 km WNW of Mt York, Mar

1984, Newbey 10002 (PERTH); Nullagine road, S of Mt

Edgar Station, Jun 1941, Burbidge 1183 (PERTH); Cranks

Well on the North West Coastal Highway, Oct 1975, Weber

4873 (AD, PERTH); Nanutarra, Ashburton R., May 1905,

Morrison s.n. [PERTH 1525204] (PERTH); Uaroo Station,

Jul 1964, Beard 3605 (PERTH); 44 km from Duck Creek

along the Mt Stewart-Duck Creek track, Jun 1976, Mitchell

76/118 (PERTH); 3 mile [c. 5 km] N of Roy Hill, Aug 1963,

Beard 2801 (PERTH). Ashburton District: 345 km N of

Carnarvon, Jul 1976, Stacey 453 (PERTH); Towera Station,

Aug 1981, Cranfield 1760 (PERTH); c. 15 km NW of

Lyndon Homestead, Sep 1975, Weber (AD, BRI).

Distribution and habitat: Corchorus laniflorus

is confined to the Pilbara region, Western

Australia, from Lyndon Station eastwards to

Mt Edgar Station (Map 4). It is recorded as

growing on sandy soils on spinifex plains and

stony soils on hills or other rocky localities

sometimes associated with sandstone.

Phenology: Flowers have been collected in

March and from June to October, fruits in June,

September and October.

Notes: Corchorus laniflorus, C. parviflorus and

C. walcottii all have conspicuous simple

glandular hairs present amongst the dense

stellate indumentum on the stems, leaves and

inflorescences. Corchorus laniflorus is most

closely related to C. parviflorus but differs from

that by having generally larger flowers and

Halford, Notes on Tiliaceae 4, Corchorus

597

Fig. 6. Corchorus laniflorus. A. branchlet with flower buds and fruit, x 1. B. fruit with persistent sepals removed, x 3. C.

ventral view of sepal, x 6. D. cross-section of sepal, x 12. E. dendritic-stellate hair, x 24. F. simple glandular hair, x 24. A from

Weber 4873 (AD); B-F from Gardner 6287 (PERTH). Del. W. Smith.

fruits, and a thicker and denser indumentum.

Corchorus laniflorus is distinguishable from

C. walcottii by having a thicker indumentum

on the stems and leaves, persistent sepals that

enclose the fruit and shorter hairs on the fruit.

Corchorus laniflorus resembles C. sericeus in

having persistent sepals that enclose the fruit,

but differs from that in having conspicuous

simple glandular hairs, larger flowers and fruits

and a thicker and softer indumentum on most

parts.

The collections, Forest [MEL227128],

Burbidge 5881 (PERTH), Burbidge 5961

(PERTH) and Mitchell PRP275 (BRI) from

Abydos and Woodstock Stations south of Port

Hedland, resemble C. laniflorus but differ from

the typical form of this species by their smaller

flowers, lack of conspicuous simple glandular

hairs and narrowly ovate to ovate leaves. These

specimens are somewhat similar to C. sericeus

subsp. densiflorus from north-eastern Northern

Territory. However, they differ from C. sericeus

subsp. densiflorus in having a denser

indumentum and larger flowers. These

collections may represent a distinct species but

further collections and study are required.

598

9. Corchorus lasiocarpus Halford sp. nov.

similis C. walcottii et C. lanifloro autem

ab utroque pills conspicuis simplicibus

glandularibus deficientibus differt et ab

illo sepalis angustioribus (1-3 mm latis

non 3-5 mm latis) persistentibus in

fructibus, pilis longioribus (3 mm longis

non 1.5 mm longis) in fructibus et in

pagina abaxiali sepalarum et ab hoc

fructibus majoribus (8-20 x 5-16 mm

non 7-9 x 3-4 mm) differt. Typus:

Western Australia. Fortescue District:

Hamersley Range near Wittenoom Gorge,

7 May 1958, N.T. Burbidge 6005 (holo:

PERTH; iso CANB).

Compact to open shrub to 1 m high, sometimes

viscid; stems much branched, spreading.

Indumentum on young shoots, branchlets,

leaves, stipules, peduncles, pedicels and bracts

grey-white, moderately dense to dense,

comprised of mostly stellate hairs but dendritic-

stellate and simple hairs also occasionally

present. Stellate hairs sessile or stipitate, up to

2 mm across; stipes red-brown or white, straight

or tortuous, up to 0.8 mm long; rays firm to

pliable, white, spreading, up to 2 mm long.

Dendritic-stellate hairs up to 2.4 mm long; rays

firm to pliable, up to 1.5 mm long. Simple hairs

glandular, white, flexuous, up to 0.5 mm long.

Leaves with petioles 7-25 mm long; stipules

subulate-linear; 2-10 mm long; lamina

narrowly ovate to ovate or narrowly oblong to

oblong, 2-6 cm long, 0.4-3 cm wide, l:w ratio

1.8—3.5:1, concolorous; base rounded; margin

serrate to dentate-serrate; apex obtuse to

rounded. Inflorescences umbellate, 3-6-

flowered, leaf-opposed, solitary at nodes;

peduncles 4-25 mm long; pedicels 4-10 mm

long, spreading to erect in flower and fruit;

bracts filiform-linear, 5-15 mm long. Flower

buds ellipsoid, 5-10 mm across; apex obtuse

with 5 spreading caudae to 5 mm long. Sepals

5, persistent, narrowly obovate-elliptic, 9-15

mm long, 1-3 mm wide; abaxial surface with

a dense indumentum of stellate and dendritic-

stellate hairs, the largest hairs up to 3 mm long;

Austrobaileya 6 (4): 581-629 (2004)

adaxial surface stellate-villose proximally,

glabrous distally; apex caudate, up to 6 mm

long. Petals 5; lamina obovate to broadly

obovate, 6-10 mm long, 5-8 mm wide,

glabrous; claw 0.7-1 mm long, with ciliate

margins. Androgynophore 0.1-0.3 mm long;

annulus entire or sometimes sinuate, 0.2-0.4

mm long, glabrous. Stamens 95-140; filaments

4-6 mm long; anthers c. 0.5 mm long. Ovary

subglobose or ellipsoid, 1.6-3 mm across,

densely stellate-tomentose, 4 or 5(rarely 3)-

locular, with 10-24 ovules in each locule; style

3-6 mm long. Fruits narrowly ellipsoid to

broadly ovoid-ellipsoid, 8-20 mm long, 5-16

mm across, 1.2-2.5 t im es longer than wide,

not conspicuously constricted between seeds,

+ circular in transverse section, 4 or 5 (rarely

3)-valved; apex acute to rounded; indumentum

dense, of stellate and dendritic-stellate hairs up

to 5 mm long. Seeds obovoid, c. 2 mm long.

Affinities’. Corchorus lasiocarpus resembles

C. walcottii and C. laniflorus but differs from

both by lacking conspicuous simple glandular

hairs. In addition, C. lasiocarpus differs from

C. walcottii by having narrower sepals (1-3

mm across compared with 3-5 mm across) that

are persistent and longer hairs on the abaxial

surface of the sepals (3 mm long compared with

1.5 mm long). Corchorus lasiocarpus differs

from C. laniflorus by having larger fruit (8-20

x 5-16 mm compared with 7-9 x 3-4 mm).

Corchorus lasiocarpus can be confused

with C. incanus. For differences from

C. incanus refer to ‘Affinities’ section under

that species.

Etymology : The specific epithet refers to the

woolly appearance of this species fruit; Greek

lasios , hairy, woolly and karpos fruit.

Notes: Corchorus lasiocarpus is confined to

the north-west of Western Australia. The

species as circumscribed here exhibits variation

in numerous characters including leaf and fruit

size, and the length of dendritic-stellate hairs.

Two subspecies are formally recognised here.

Fruits 15-20 mm long; dendritic-stellate hairs on fruit up to 5 mm long

. 9a. C. lasiocarpus subsp. lasiocarpus

Fruits 8-12 mm long; dendritic-stellate hairs on fruit up to 2 mm long

. 9b. C. lasiocarpus subsp. parvus

Halford, Notes on Tiliaceae 4, Corchorus

599

Fig. 7. Corchorus lasiocarpus subsp. lasiocarpus. A. branchlet with fruit, x 1. B. fruit, x 2. C. ventral view of sepal, x 4. A-

C from Royce 1707 (PERTH). Del. W. Smith.

9a. Corchorus lasiocarpus Halford subsp.

lasiocarpus

Spreading shrub to 1 m high. Petioles 10-25

mm long. Leaf lamina narrowly ovate to ovate,

3.5-6 cm long, 1.5-3 cm wide, l:w ratio 1.8-2:1.

Inflorescences 3-6-flowered; peduncles 8-25

mm long; pedicels 5-10 mm long. Flower buds

7-10 mm across. Sepals 12-15 mm long, 2-3

mm wide; apex caudate, up to 6 mm long.

Petals obovate to broadly obovate, 8-10 mm

long, 6-8 mm wide. Stamens 130-140;

filaments 5-6 mm long. Ovary subglobose, 2-3

mm across, style 5-6 mm long. Fruits narrowly

to broadly ovoid-ellipsoid, 15-20 mm long,

12-16 mm across; dendritic-stellate hairs up

to 5 mm long. Fig. 7.

Selected specimens (from 20 examined): Western

Australia. Fortescue District: between Woodstock Station

and Hamersley Range (Tablelands), May 1958, Burbidge

5989 (PERTH); 116 miles [c. 187 km] S of Port Hedland, on

Wittenoom road, Aug 1960, George 1091 (PERTH); 4 km

SW of Two Sisters, c. 145 km SE of Shay Gap, Jul 1984,

Newbey 10528 (PERTH); 18 km SSWof Two Sisters, c. 160

km SE of Shay Gap, Jul 1984, Newbey 10477 (PERTH);

142 miles [c. 228 km] S of Port Hedland, on Wittenoom road,

Aug 1960, George 1073 (PERTH); 11 miles [c. 18 km] E of

Wittenoom, Aug 1960, George 1008 (PERTH); Yampire

Gorge, Hamersley Range, Aug 1959, Gardner 12274

(PERTH); Dale Gorge, Hamersley Range, Aug 1960, George

1046 (PERTH); Stag Arrow Creek, Little Sandy Desert, Apr

1979, Mitchell 451 (DNA, PERTH); 1 mile [c. 1.6 km] N of

Stag Arrow Creek, May 1947, Royce 1707 (PERTH); 20

mi les [c. 32 km] N [of] Christmas Creek on R.P.F., May 1947,

Royce 1772 (PERTH); off main road from Paraburdoo to

Tom Price, 1 km along pipeline access track, Paraburdoo,

Sep 1984, Wurm 1496 (PERTH). Keartland District: Little

Sandy Desert, Apr 1979, Mitchell 687 (DNA, PERTH).

600

Austrobaileya 6 (4): 581-629 (2004)

Fig. 8. Corchorus lasiocarpus subsp. parvus. A. branchlet with flower buds and fruit, x 1.5. B. fruit, x 3. C. ventral view of

sepal, x 6. D. cross-section of sepal, x 24. E. dendritic-stellate hair, x 36. A-E from Gardner 6390 (PERTH). Del. W. Smith.

Distribution and habitat’.Corchorus

lasiocarpus subsp. lasiocarpus occurs from

Tom Price eastwards to the Two Sisters on the

western edge of the Great Sandy Desert (Map

8). It is recorded as growing in hummock

grassland and open shrubland communities, on

sandy, stony or gravelly soils along

watercourses.

Phenology : Flowers have been collected in

April, May, December and from July to

September, fruits in April, May, August and

December.

9b. Corchorus lasiocarpus subsp. parvus

Halford subsp. nov. a C. lasiocarpo

subsp. lasiocarpo fructibus minoribus (8-12

Halford, Notes on Tiliaceae 4, Corchorus

601

x 5-7 mm non 15-20 x 12-16 mm), foliis

minoribus (2-5.5 x 0.4-2.5 cm non 3.5-

6 x 1.5-3 cm) pilis stellato-dendriticis

minoribus (usque 2 mm longis non usque

5 mm longis) vestitis distinguendus.

Typus: Western Australia. Fortescue

District: Yathalla Well, near Mt Rica,

Hamersley Range, 22 Oct 1941, C.A.

Gardner 6390 (holo: PERTH).

Open to compact shrub to 0.8 m high. Petioles

7-15 mm long. Leaf lamina narrowly ovate or

narrowly oblong to oblong, 2-5.5 cm long,

0.4-2.5 cm wide, l:w ratio 2.1-3.5:1.

Inflorescences 3 or 4-flowered; peduncles

4-17 mm long; pedicels 4-7 mm long. Flower

buds 5-8 mm across. Sepals 9-12 mm long,

1-3 mm wide; apex caudate, up to 3 mm long.

Petals obovate, 6-8 mm long, 5-6 mm wide.

Stamens c. 95; filaments 4-5 mm long. Ovary

ellipsoid, 1.6-2 mm across; style 3-5 mm long.

Fruits narrowly ellipsoid, 8—12 mm long, 5-7

mm across; dendritic-stellate hairs up to 2 mm

long. Fig. 8.

Selected specimens (from 14 examined): Western

Australia. Fortescue District: near Robe R., Aug 1970,

Beard 6144 (PERTH); Hamersley Range-Bulgeeda to

Pyrton, Aug 1963, Cole WA5040 (PERTH); Hamersley

Range, Aug 1963, Cole WA5094 (PERTH); flats E of East

Prongs, Tom Price, Jul 1980, [Atkins] HI-707 (PERTH); 6

mi les [c. 10km] E of Mt Brockman, Aug 1970, Demarz 2467

(PERTH); Marandoo, Mar 1980, Atkins HI-659 (PERTH);

0.5 km E of Packsaddle, Feb 1987, Mollemans 2279 (AD,

PERTH); 2 km W of the Governor on the Packsaddle-West

Angeles road, 23 km from PS., Feb 1987, Mollemans 2216

(AD, PERTH).

Distribution and habitat: Corchorus

lasiocarpus subsp. parvus is confined to the

Hamersley Range from near Mt Rica south-east

to Mt Bruce (Map 9). It is recorded as growing

in hummock grassland and tree steppe

communities, on stony slopes and plains.

Phenology: Flowers have been collected in

February, March and from July to October,

fruits in March.

Affinities: Corchorus lasiocarpus subsp.

parvus can be distinguished from

C. lasiocarpus subsp. lasiocarpus by its smaller

fruit (8-12 x 5-7 mm compared with 15-20 x

12—16 mm), smaller leaves (2-5.5 x 0.4-2.5

cm compared with 3.5-6 x 1.5-3 cm) and

shorter dendritic-stellate hairs on the fruit (up

to 2 mm long compared with up to 5 mm long).

Etymology: The specific epithet is in reference

to the overall smaller dimensions of this

subspecies; Latin parvus little.

10. Corchorus leptocarpus A.Cunn. ex Benth.,

FI. Austral. 1:278 (1863). Type: [Western

Australia.] Water Island, NW coast, [Sep

1820,] A. Cunningham [No. 247] (holo:

K; iso: MEL [MEL227288], CANB).

Shrub to 2 m high; ste ms sparingly to much

branched, erect; young shoots with ferruginous

indumentum. Indumentum on branchlets,

leaves, stipules, peduncles, pedicels and bracts

grey-white, moderately dense to dense,

comprised of stellate hairs. Stellate hairs sessile

or stipitate, up to 0.4 mm across; stipes red-

brown, straight, up to 0.2 mm long; rays firm,

white or somet im es ferruginous, up to 0.2 mm

long. Leaves with petioles 8-13 mm long;

stipules narrowly triangular to subulate-linear,

3- 5 mm long; lamina narrowly ovate to ovate,

6-10 cm long, 2-4 cm wide, l:w ratio 2.5-3:1,

discolorous; base rounded or slightly cordate;

margin serrulate or crenate; apex obtuse or

rarely acute. Inflorescences umbellate, 3-6-

flowered, leaf-opposed or lateral, solitary at

upper nodes; peduncles 1-4 mm long; pedicels

5-7 mm long, spreading to erect in flower,

recurved to erect in fruit; bracts subulate-linear,

2-3 mm long. Flower buds obovoid-ellipsoid,

4- 5 mm across, longitudinally ridged; apex

obtuse with 5 erect to spreading caudae to 2

mm long. Sepals 5, not persistent, narrowly

obovate, 12-14 mm long, c. 3 mm wide; abaxial

surface with a dense indumentum of stellate

hairs up to 0.2 mm long; adaxial surface

stellate-pubescent proximally, glabrous distally;

apex acuminate-caudate, up to 2 mm long.

Petals 5; lamina obovate to broadly obovate,

8-10 mm long, c. 7 mm wide, glabrous; claw

c. 1 mm long, stellate-pubescent on margins.

Androgynophore c. 0.4 mm long; annulus

entire, c. 0.4 mm long, glabrous. Stamens

80-90; filaments 7-9 mm long; anthers c. 0.5

mm long. Ovary cylindrical, c. 0.9 mm across,

densely stellate-puberulous, 3 (rarely 4)-locular,

with 44-48 ovules in each locule; style 6-7 mm

long. Fruits subcylindrical, 20-60 mm long,

2-3 mm across, 7-20 times longer than wide,

± straight to slightly curved or if on recurved

pedicels then fruit abruptly bent near base so

that the fruit is perpendicular with the apex

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Austrobaileya 6 (4): 581-629 (2004)

pointing upwards, slightly to markedly

constricted between seeds, circular in transverse

section, 3(rarely 4)-valved; apex attenuate,

2- 4 mm long; indumentum moderately dense

to dense, of stellate hairs; stellate hairs up to

0.5 mm long, 0.4 mm across. Seeds compressed

obovoid, c. 2 mm long.

Additional specimens : Western Australia. Gardner

District: 6 km SW of Crystal Head, Port Warrender,

Admiralty Gulf, Jan 1982, Farrell 979 (PERTH); Boomerang

Bay on W side of Bigge Island, Bonaparte Archipelago, May

1987, Kenneally 10018 (BRI, PERTH); E side of Mindjau

Creek, Port Warrender, Admiralty Gulf, Jan 1982, Kenneally

7771 (PERTH); Hunter R„ West Kimberley, May 1987,

Kenneally 9946 (PERTH); Pirn Hill, SE of West Bay, May

1984, Willis s.n. [MEL1599281] (MEL); Osborne Island

(south east island), Bonaparte Archipelago, Jun 1973, Wilson

11095 (PERTH).

Distribution and habitat : Corchorus

leptocarpus occurs on the islands and in coastal

areas of the Kimberley, Western Australia, from

Bigge Island, Bonaparte Archipelago eastwards

to West Bay (Map 6). It is recorded as growing

on soils derived from sandstone, along drainage

lines and in shallow depressions in flat country.

Phenology: Flowers have been collected in

May, fruits in May and June.

Notes: Corchorus leptocarpus is similar to

C. sidoides in that it has narrow cylindrical

fruits that are slightly to markedly constricted

between the seeds. However, C. leptocarpus

differs from C. sidoides by having larger

flowers (sepals 12-14 mm long compared with

3- 9 mm long; petals c. 10 x 7 mm compared

with 2-7 x 0.5-4 mm), larger leaves (6-10 x

2-4 cm compared with 0.6-9 x 0.2-3 cm), and

erect rather than spreading to pendulous fruit.

Corchorus leptocarpus is most closely related

to C. sublatus and C. subargentus. For features

distinguishing C. leptocarpus from C. sublatus

and C. subargentus see ‘Affinities’ under those

species.

11. Corchorus mitchellensis Halford, sp. nov.

affinis C. leptocarpo autem floribus

minoribus (sepalis 6-7 x c. 2 mm et

petalis 6x3 mm non sepalis 12-14 mm

x c. 3 mm et petalis 8-10 x. c. 7 mm),

pilis stellatis majoribus (usque 1.3 mm

diam. non usque 0.4 mm diam.) in

fructibus differt. Typus: Western

Australia. Gardner District: Mitchell

Falls, Mitchell Plateau, 30 May 1992, D.

Halford Q1433 (holo; PERTH; iso: BRI,

DNA, MEL, distribuendi).

Shrub to 1 m high; stems sparingly to much

branched, erect; young shoots with ferruginous

indumentum. Indumentum on branchlets,

leaves, stipules, peduncles, pedicels and bracts

grey-white or ferruginous, moderately dense to

dense, comprised of stellate hairs. Stellate hairs

sessile or stipitate, up to 1.5 mm across; stipes

white, straight, c. 0.1 mm long; rays firm, white

or ferruginous, up to 0.7 mm long. Leaves with

petioles 6-10 mm long; stipules subulate-linear,

1-2 mm long; lamina narrowly ovate to ovate,

3.5-8 cmlong, 1-2.5 cm wide, l:w ratio 3.1—3.5:1,

discolorous; base rounded; margin serrate to

serrulate; apex obtuse to acute. Inflorescences

umbellate 3-6-flowered, leaf-opposed, solitary

at upper nodes; peduncles 1-2 mm long;

pedicels 1-2 mm long, spreading to erect in

flower, recurved in fruit; bracts subulate-linear,

1-2 mm long. Flower buds obovoid-ellipsoid,

3-4 mm across; apex obtuse with 5 erect caudae

to 0.7 mm long. Sepals 5, not persistent,

narrowly obovate, 6-7 mm long, c. 2 mm wide;

abaxial surface with a moderately dense to

dense indumentum of stellate hairs up to 0.7

mm long; adaxial surface stellate-pubescent

proximally, glabrous distally; apex acuminate-

caudate, up to 1 mm long. Petals 5; lamina

obovate, c. 6 mm long, c. 3 mm wide, glabrous;

claw c. 0.8 mm long, stellate-pubescent on

margins. Androgynophore c. 0.4 mm long;

annulus entire, c. 0.4 mm long, glabrous.

Stamens 60-70; filaments 3-4 mm long;

anthers c. 0.5 mm long. Ovary cylindrical, c.

0.8 mm across, densely stellate-tomentose,

3-locular, with 20-24 ovules in each locule;

style c. 4 mm long. Fruits subcylindrical,

20-40 mm long, 3-4 mm across, 7-10 times

longer than wide, curved, circular in transverse

section, slightly constricted between seeds,

3-valved; apex acute to obtuse, orientated

upward; indumentum dense, of stellate hairs;

stellate hairs up to 1 mm long, 1.3 mm across.

Seeds compressed obovoid or columnar, 1-3

mm long. Fig. 9.

Additional specimens : Western Australia. Gardner

District: Mitchell Falls, Feb 1980, Done 120 (DNA);

Mitchell Falls, Mitchell Plateau, May 1992, Halford Q1433a

(BRI).

Halford, Notes on Tiliaceae 4, Corchorus

603

Fig. 9. Corchorus mitchellensis. A. branchlet with flower buds, x 1.5. B. fruit, x 2. C. ventral view of sepal, x 8. A from Done

120 (DNA); B, C from Halford Q1443 (BRI). Del. W. Smith.

Distribution and habitat : Corchorus

mitchellensis is known only from sandstone

country around Mitchell Falls, Mitchell

Plateau, Kimberley, Western Australia (Map

4). It is recorded as growing in shrubland and

low open woodland communities on sandy or

gravelly soils along drainage lines in dissected

sandstone hills.

Phenology: Flowers have been collected in

February, fruits in May.

Affinities: Corchorus mitchellensis is related

to C. leptocarpus but differs from that by having

smaller flowers (sepals 6-7 x c. 2 mm, petals

c. 6 x 3 mm compared with sepals 12-14 x c.

3 mm, petals 8-10 x c. 7 mm) and larger stellate

hairs on the fruit (hairs up to 1.3 mm across

compared with 0.4 mm for C. leptocarpus ).

Etymology: The specific epithet is derived from

the name Mitchell, plus the suffix -ensis

indicating place of origin, alluding to the

Mitchell Plateau from where this species is

known.

12. Corchorus obclavatus Halford, sp. nov.

quoad collocationem et formam et

magnitudinem fructuum C. pumilionis

autem statura altiore et forma

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Austrobaileya 6 (4): 581-629 (2004)

magnitudineque foliis differt (vide

tabulam 2 pro differentiis). Corchorus

obclavatus quoad habitus C. sidoidi

subsp. rostrisepalo et C. sublato similis

autem ab utroque fructibus brevioribus

(4-9 mm longis nec 20-55 mm longis in

C. sidoide subsp. rostrisepalo nec 20-50

mm in C. sublato ), obclavatis non ±

cylindricis differt. Addite C. obclavatus

a C. sublato fructibus pendulis non erectis

differt. Typus: Northern Territory.

Darwin and Gulf Region: Jim Jim Falls,

Kakadu NP, 20 April 1992, D. Halford

Q1150 (holo: DNA; iso: BRI, MEL,

distribuendi).

Shrub to 2 m high; stems sparingly to much

branched, erect; young shoots with ferruginous

indumentum. Indumentum on branchlets,

leaves, stipules, peduncles, pedicels and bracts,

grey-white, dense, comprised of stellate hairs.

Stellate hairs sessile or stipitate, up to 0.7 mm

across; stipes red-brown, straight, up to 0.2 mm

long; rays pliable, white or ferruginous, up to

0.5 mm long. Leaves with petioles 4-15 mm

long; stipules subulate-linear, 2-5 mm long;

lamina narrowly ovate, 3-9 cm long, 0.7-2 cm

wide, l:w ratio 3.6-6:1, discolorous; base

rounded or slightly cordate; margin serrulate

to crenulate; apex acute or obtuse.

Inflorescences umbellate, 4-7-flowered, leaf-

opposed, solitary at upper nodes; peduncles

1-2 mm long; pedicels 1-2 mm long, erect to

spreading in flower, recurved in fruit; bracts

subulate-linear to filiform, 2-3 mm long.

Flower buds obovoid-ellipsoid, 2-3 mm across;

apex obtuse with 4 spreading caudae to 0.7 mm

long. Sepals 4, not persistent, narrowly obovate,

4-6 mm long, 1-2 mm wide; abaxial surface

with a dense indumentum of stellate hairs up

to 0.3 mm long; adaxial surface glabrous; apex

acuminate, up to 0.7 mm long. Petals 4; lamina

narrowly obovate, 4-5 mm long, 2-3 mm wide,

glabrous; claw c. 0.5 mm long, stellate-

pubescent on margins. Androgynophore

0.2-0.3 mm long; annulus entire, c. 0.2 mm

long, glabrous. Stamens 20-35; filaments 2-3

mm long; anthers c. 0.4 mm long. Ovary ovoid,

0.5-0.6 mm across, densely stellate-tomentose,

2-locular, with 2-4 ovules in each locule; style

c. 3 mm long. Fruits obclavate, 4-9 mm long,

1- 2 mm across, 2-5 times longer than wide,

pendulous, straight, circular in transverse

section, slightly constricted between seeds,

2- valved; apex attenuate, 1-4 mm long;

indumentum moderately dense to dense, of

stellate hairs up to 0.5 mm long. Seeds

compressed obovoid or columnar, 1-2 mm long.

Fig. 10.

Aditonalspecimens: Northern Territory. Darwin and Gulf

Region: 12 km E of Mudginberri Homestead, Kakadu NP,

Jan 1991, Russell-Smith 8409 &Brock (BRI); Jim Jim Falls,

Kakadu NP, Apr 1992, Halford Q1151 (BRI).

Distribution and habitat : Corchorus

obclavatus is restricted to the sandstone taluses

and escarpments of Kakadu National Park,

Northern Territory (Map 1). It is recorded as

growing on sandy soils in open woodland

communities near vine thicket margins.

Phenology: Flowers have been collected in

January and April, fruits in April.

Affinities: Corchorus obclavatus is similar in

fruit orientation, shape and size to C. pumilio

but differs from that by its taller stature, and

leaf shape and size. These differences are

summarized in Table 1. Corchorus obclavatus

is similar in habit to C. sidoides subsp.

rostrisepalus and C. sublatus but can be

distinguished from these by having shorter fruit

(4-9 mm long compared with 20-55 mm long

for C. sidoides subsp. rostrisepalus and 20-55

mm long for C. sublatus ) which are obclavate

Table 1. Morphological comparison of Corchorus obclavatus and C. pumilio.

Character

C. obclavatus

C. pumilio

habit

erect shrub to 2 m high

spreading shrub to 0.4 m high

leaf shape

narrowly ovate

oblong, oblong-elliptic or narrowly elliptic

leaf size (cm)

3-9 x 0.7-2

0.6-3.5 x 0.4-1.4

Halford, Notes on Tiliaceae 4, Corchorus

605

Fig. 10. Corchorus obclavatus. A. branchlet with fruit, x 1. B. fruit, x 4. C. ventral view of sepal, x 8. A-C from Halford

Q1150 (BRI). Del. W. Smith.

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Austrobaileya 6 (4): 581-629 (2004)

rather than ± cylindrical. In addition,

C. obclavatus differs from C. sublatus by

having pendulous rather than erect fruit.

Etymology: The specific epithet refers to the

shape of the mature fruit; Latin ob- prefix

reversed-, clavatus club shape, ie the club-

shaped fruit is attached by the thicker end.

13. Corchorus parviflorus (Benth.) Domin,

Biblioth. Bot. 89: 383 (1928); Corchorus

parviflorus (Benth.) Domin var.

parviflorus , Domin, Biblioth. Bot. 89:

383 (1928); Corchorus walcottii var.

parviflorus Benth., FI. Austral. 1: 279

(1863). Type: [Western Australia.] Nichol

Bay, 1862, F. Gregory, (lecto: MEL

[MEL223669]); isolecto: K n.v.,fide B.

Rye, Nuytsia 9(3): 418 (1994); ?isolecto:

MEL [MEL1599091]).

Corchorus parviflorus var. gracilescens

Domin, Biblioth. Bot. 89: 383 (1928).

Type: [Western Australia.] between the

Ashburton and De Gray Rivers, E.

Clement (syn: K); [Western Australia.]

Mons Cupri, Whim Creek, W.A. Michell

(syn: K).

Corchorus parviflorus var. ovatus Domin,

Biblioth. Bot. 89: 383 (1928). type:

[Western Australia.] between the

Ashburton and De Gray Rivers, E.

Clement (holo: K).

Shrub to 1(-1.6) mhigh; stems much branched,

spreading to erect. Indumentum on young

shoots, branchlets, leaves, stipules, peduncles,

pedicels and bracts grey-white, moderately

dense to dense, comprised of mostly stellate

hairs but simple hairs also present. Stellate

hairs sessile or sometimes stipitate, up to 1 mm

across; stipes red-brown, straight, up to 0.2 mm

long; rays soft, white, up to 0.2 mm long.

Simple hairs glandular, dull yellow to red-

brown, flexuous, up to 1.5 mm long. Leaves

with petioles (3-)10-20(-35) mm long; stipules

subulate-linear, 6-8 mm long; lamina ovate to

broadly ovate or elliptic to broadly elliptic,

(1-) 1.5-4 cm long, (0.5-) 1-3 cm wide, 1: w ratio

1.1-2:1, concolorous; base rounded or slightly

cordate; margin serrulate; apex obtuse to

rounded. Inflorescences umbellate, 3-8-

flowered, leaf-opposed, solitary at nodes;

peduncles (2-)6-13 mm long; pedicels 2-5 mm

long, spreading to erect in flower and fruit;

bracts filiform-linear, 3-4 mm long. Flower

buds ellipsoid, 3-4 mm across, apex obtuse with

5 spreading caudae to 1 mm long. Sepals 5,

persistent, narrowly obovate-elliptic, 4-7 mm

long, 1-2 mm wide; abaxial surface with a

dense indumentum of stellate and simple hairs,

the largest hairs up to 1.5 mm long; adaxial

surface stellate-villose proximally, glabrous

distally; apex acuminate-caudate, up to 1 mm

long. Petals 5; lamina obovate, 3-6 mm long,

2-5 mm wide, glabrous; claw 0.5-0.7 mm long,

sparsely stellate-pubescent on margins.

Androgynophore 0.2-0.4 mm long; annulus

entire, c. 0.2 mm long, glabrous. Stamens

43-73; filaments 2-4 mm long; anthers c. 0.4

mm long. Ovary globose, 1.5-2 mm across,

densely stellate-villose, 3 or 4-locular, with

8-10 ovules in each locule; style 3-4 mm long.

Fruits subcylindrical, 4-12 mm long, 2-3 mm

across, 2-4 times longer than wide, spreading,

straight, circular in transverse section, not

conspicuously constricted between seeds, 3 or

4-valved; apex attenuate, 1-5 mm long;

indumentum dense, of mostly stellate hairs but

a few simple hairs also present, largest hairs

up to 0.5 mm long. Seeds compressed

obovoid, c. 2 mm long. Fig. 11.

Selected specimens (from 17 examined): Western

Australia. Fortescue District: North West Coastal Highway,

c.15 kmbyroadWSW ofmain turnoff toDampier, Aug 1977,

Jackson 3033 (AD); Deep Hills runoff gully near Bullgarra

Cell, Karratha, Sep 1985, Glennon 220 (PERTH); Point

Samson, Jul 1981, Craig 214 (PERTH); 65 km N of

Roeboume, Jul 1976, Stacey CIS463 (PERTH); Roeboume,

Oct 1941, Gardner 6329 (PERTH); 15.8 km N of Taiga,

Sep 1986, Chinnock 6985 (AD); Soda Creek, Black Hills,

Eginbah Station, Coongan, Jun 1941, Burbidge 998

(PERTH); 38 km NNW of Abydos, Jul-Aug 1987, Ingleby

HW15 (PERTH); Abydos Station, S of Port Hedland, Sep

1961, Richardson 14 (PERTH); Woodstock Station, May

1958, Burbidge 5972 (AD, PERTH); c. 160 km S of Port

Hedland towards Wittenoom, Apr 1977, Pullen 10.911

(CANB); near Marble Bar, Sep 1968, Blockley s.n. (PERTH);

Mt Edgar, Feb-Mar 1938, Stewart 394 (PERTH); Hamersley

Range, Aug 1932, Gardner s.n. [PERTH1522108]

(PERTH)); head of Nullagine R., 54 km S of Nullagine along

Great Northern Highway, Jun 1977, Telford & Butler 5921

(CANB).

Distribution and habitat : Corchorus

parviflorus is confined to the Pilbara region,

Western Australia, from Karratha eastwards to

Mt Edgar Station (Map 1). It is recorded as

growing in hummock grassland and low open

Halford, Notes on Tiliaceae 4, Corchorus

607

Fig. 11. Corchorusparviflorus. A. branchlet with flowers and fruit, x 2. B. fruit with persistent sepals, x 4. C. ventral view of

sepal, x 8. D. cross-section of sepal, x 24. E. stellate hair, x 48. F. simple glandular hair, x 24. A-F from Jackson 3033 (AD).

Del. W. Smith.

woodland communities, on stony or sandy soils

on hillslopes and plains. It is also recorded

occasionally along watercourses.

Phenology : Flowers have been collected from

March to October, fruits in April and from

August to October.

Notes: Corchorus parviflorus , C. laniflorus and

C. walcottii all have conspicuous simple

glandular hairs present amongst the dense

stellate indumentum on the stems, leaves and

inflorescences. Corchorus parviflorus is most

closely related to C. laniflorus. For a discussion

on the differences between these species see

‘Notes’ under C. laniflorus. Corchorus

parviflorus is distinguishable from C. walcottii

by having persistent sepals and generally

smaller leaves, flowers and fruits.

14. Corchorus puberulus Halford sp. nov.

similis C. leptocarpo autem fructibus

latioribus (3-4 mm latis non 2-3 mm

latis) minus quam 10 plo longioribus

quam latis inter semina non constrictis

differt. Corchorus puberulus floribus ex

sepalis 9-15 x 2-3 mm, petalis 9-10 x

4-7 mm filamentis staminalibus 4-6 mm

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Austrobaileya 6 (4): 581-629 (2004)

longis, stylo 4-6 mm compositis,

pedunculis 5-7 mm longis, foliis 2-5 cm

latis praeditus maxime arete cognatus C.

aulacocarpo qui ex flores minore, sepalis

6-9 x 1-2 mm, petalis 6-7 x 2-4 mm,

stylo 2-4 mm longo compositos,

pedunculos breviores 2-3 mm longos,

interdum folia angustioria 0.8-2.5 cm lata

habet. Typus: Western Australia.

Fitzgerald District: W end of Cockatoo

Island airstrip, W Kimberley, 6 November

1985, P.J. White 27 (holo: PERTH).

Shrub to 1.5 m high; stems sparingly to much

branched, erect. Indumentum on young shoots,

branchlets, stipules, peduncles, pedicels and

bracts grey-white, dense, comprised of mostly

stellate hairs but with a few dendritic-stellate

hairs also present. Stellate hairs sessile or

stipitate, up to 0.2 mm across; stipes red-brown,

straight, up to 0.2 mm long; rays pliable, white,

up to 0.2 mm long. Dendritic-stellate hairs up

to 0.3 mm long; stipes red-brown; rays pliable,

white, up to 0.2 mm long. Leaves with petioles

7-17 mm long; stipules subulate-linear, 4-5

mm long; lamina ovate, 4.5-10 cm long, 2-5

cm wide, l:w ratio 2-2.9:1, discolorous; adaxial

surface sparsely to moderately stellate hairy;

abaxial surface moderately to densely stellate

hairy; base rounded; margin serrate to serrulate;

apex obtuse to acute. Inflorescences umbellate,

3-7-flowered, leaf-opposed or lateral, solitary

at upper nodes; peduncles 5-7 mm long;

pedicels 5-8 mm long, spreading to erect in

flower, recurved to erect in fruit; bracts

subulate-linear, 2-3 mm long. Flower buds

obovoid-ellipsoid, 4-5 mm across,

longitudinally ridged; apex obtuse with 5 erect

caudae to 2 mm long. Sepals 5, not persistent,

narrowly obovate, 9-15 mm long, 2-3 mm

wide; abaxial surface with a dense indumentum

of stellate hairs up to 0.3 mm long; adaxial

surface stellate-pubescent proximally, glabrous

distally; apex caudate, up to 4 mm long. Petals

5; lamina obovate to broadly obovate, 9-10 mm

long, 4-7 mm wide, glabrous; claw c. 1 mm

long, stellate-pubescent on margins.

Androgynophore 0.4-0.7 mm long; annulus

entire, c. 0.5 mm long, glabrous. Stamens 60-75;

filaments 4-6 mm long; anthers c. 0.5 mm long.

Ovary cylindrical, c. 1.5 mm across, densely

stellate-puberulous, 4(rarely 3 or 5)-locular,

with 40-44 ovules in each locule; style 4-6 mm

long. Fruits subcylindrical, 10-30 mm long,

3- 4 mm across, 3-8 times longer than wide,

erect, straight or slightly curved or if on

recurved pedicels then fruit abruptly bent near

base so that the fruit is perpendicular with the

apex pointing upwards, 4(rarely 3 or 5)-sided,

obtusely angled in transverse section, not

constricted between seeds, 4(rarely 3 or 5)-

valved; apex acute to obtuse; indumentum

moderately dense to dense, of stellate hairs;

stellate hairs up to 0.3 mm long, 0.2 mm

across. Seeds compressed obovoid, c. 2 mm

long. Fig. 12.

Additional specimens: Western Australia. Gardner

District: 5.2 km SE of Mount Lochee, Jun 1987, Kenneally

10451 & Hyland (PERTH). Fitzgerald District: Koolan

Island, near Acacia Ore Body in central part of island, Jun

1985, Fryxell 4595 etal. (CANB, MEL); Crocodile Creek,

Yampi Peninsula, W Kimberley Coast, May 1987, Kenneally

10117 (PERTH); Silver Gull Creek at spring, c. 14 km SE of

Cockatoo Island, Apr 1983, Fryxell & Craven 3865 (BRI,

CANB, DNA, MEL); 26 km W of Rankin Island, Collier

Bay, W Kimberley Coast, Jun 1987, Kenneally s.n. & Hyland

(PERTH)

Distribution and habitat : Corchorus puberulus

occurs on the islands and in the coastal areas

of the Kimberley, Western Australia, from

Cockatoo Island, Buccaneer Archipelago

eastwards to Mt Lochee (Map 4). It is recorded

as growing on the edge of vine thickets on soils

derived from sandstone and in eucalypt

woodland communities along dry creeks.

Phenology: Flowers have been collected in

April, fruits in April and June.

Affinities: Corchorus puberulus is similar to

C. leptocarpus but differs from that by having

broader fruits (3-4 mm across compared with

2-3 mm across) which are < 10 t im es as long

as wide, trigonous or tetragonous in transverse

section and are not constricted between the

seeds. Corchorus puberulus is most closely

related to C. aulacocarpus but differs from that

by having larger flowers (sepals 9-15 x 2-3

mm, petals 9-10 x 4-7 mm, staminal filaments

4- 6 mm long mm long, style 4-6 mm long

compared with sepals 6-9 x 1-2 mm, petals

6-7 x 2-4 mm, staminal filaments 3-4 mm

long, style 2-4 mm long), longer peduncles

(5-7 mm long compared with 2-3 long) and

generally broader leaves (2-5 cm wide

compared with 0.8-2.5 cm wide).

Halford, Notes on Tiliaceae 4, Corchorus

609

Fig. 12. Corchoruspuberulus. A. branchlet with flowers, x 1.5. B. fruit, x 3. C. ventral view of sepal, x 4. A from White 27

(PERTH); B, C from Fryxell et ol. 4595 (CANB). Del. W. Smith.

Etymology : The epithet alludes to the

somewhat dense cover of short, fine, soft hairs

on most plant parts; Latin puberulus minutely

pubescent.

15. Corchorus pumilio R.Br. ex Benth., FI.

Austral. 1: 277 (1863). Type: [Northern

Territory.] Carpentaria island r[Burney

Island] No 32 desc., [19 Jan 1803,] R.

Brown (lecto, here chosen: K; ?isolecto:

BRI, CANB, MEL, NSW).

Shrub to 0.4 m high; stems much branched,

spreading; young shoots with greyish white or

rarely ferruginous indumentum. Indumentum

on branchlets, leaves, stipules, peduncles,

pedicels and bracts grey-white, sparse to

moderately dense, comprised of stellate hairs.

Stellate hairs sessile or stipitate, up to 1.2 mm

across; stipes white, straight, up to 0.1 mm

long; rays firm, white, up to 1 mm long. Leaves

with petioles l-6(-25) mm long; stipules

subulate-linear, 2-5 mm long; lamina oblong,

610

oblong-elliptic or rarely narrowly elliptic,

0.6-3.5 cm long, 0.4-1.4 cm wide, l:w ratio

1.5-3:1, concolorous; base cuneate to obtuse;

margin serrulate to serrate; apex rounded or

rarely acute rarely. Inflorescences umbellate,

3-6-flowered, leaf-opposed or lateral, 1 or 2

per node; peduncles c. 1 mm long; pedicels

1- 3 mm long, spreading to erect in flower,

recurved in fruit; bracts filiform-linear, 1-3 mm

long. Flower buds obovoid-ellipsoid, 1-2 mm

across; apex acuminate with 4 or 5 erect caudae

to 0.5 mm long. Sepals 4 or 5, not persistent,

linear to narrowly obovate, 3-6 mm long,

0.5-1 mm wide; abaxial surface with a

moderately dense indumentum of stellate hairs

up to 0.5 mm long; adaxial surface stellate-

pubescent proximally, glabrous distally; apex

acuminate, up to 0.5 mm long. Petals 4 or 5;

lamina narrowly obovate to obovate, 2-5 mm

long, 0.5-2 mm wide, glabrous; claw 0.3-0.5

mm long, stellate-pubescent on adaxial surface

and margins. Androgynophore 0.1-0.3 mm

long; annulus entire, c. 0.1 mm long, glabrous.

Stamens 5-15(-20); filaments 2-4 mm long;

anthers c. 0.5 mm long. Ovary cylindrical or

subglobose, 0.2-0.4 mm across, densely

stellate-tomentose, 2(rarely 3)-locular, with

2- 6 ovules in each locule; style 2-3 mm long.

Fruits subcylindrical, 3-10(-14) mm long,

1-2 mm across, 1.5-7 times longer than wide,

spreading to pendulous, straight, slightly

curved or twisted, circular in transverse section,

slightly or markedly constricted between seeds,

2(rarely 3)-valved; apex attenuate, 1-2 mm

long; indumentum dense, of stellate hairs up

to 0.7 mm long. Seeds compressed obovoid,

1-2 mm long.

Selected specimens (from 58 examined): Western

Australia. Gardner District: c. 10 km SE of shore of King

George R., 5 km W of shore of Timor Sea, Jun 1985, Fryxell

4821 et al. (CANB, MEL, PERTH); headwaters of

Packsaddle Creek, Northern Carr Boyd Ranges, Mar 1978,

Hartley 14359 (CANB, DNA). Fitzgerald District: creek

entering an inlet of Talbot Bay, 23 km SE of Cockatoo Island,

Apr 1983 Fryxell & Carven 3884 (AD, CANB); opposite

Bold Bluff along Milliwindi [Milliewindie] track, Leopold

Range, Apr 1988, Cranfield 6384 (PERTH). Dampier

District: One Arm Point, N Dampier Peninsula, Mar 1989,

Carter 362 (BRI, DNA, PERTH). Canning District:

Godfrey Tank, Southesk Tablelands, Apr 1979, George

15452 (AD, DNA, PERTH). Northern Territory. Darwin

and Gulf Region: Roper Bar road, 80 km E of Stuart

Highway, Apr 1992, Halford Q1090 (BRI); 45 km SSW of

Legune Station, Mar 1989, Russell-Smith 7561 & Brock

(DNA); 500 m N of Larrimah, Stuart Highway, Apr 1992,

Austrobaileya 6 (4): 581-629 (2004)

Halford Q1196 (BRI); 144 miles [c. 232 km] E of Stuart

Highway on Borroloola road, Jun 1971, Dunlop 2180 (AD,

CANB, DNA, MEL). Victoria River Region: Jasper Gorge,

Jul 1974, Carr 2833 8c Beauglehole 46612 (MEL). Central

Southern Region: Simpsons Gap NP, Apr 1974, Latz 4885

(DNA). Queensland. Cook District: Fanneys Creek, 86

km W of Georgetown, E of Gilbert R., Apr 1992, Halford

Q973 (BRI); Turtle Rock area, SE of Laura, Jan 1993, Bean

5503 & Forster (BRI). Burke District: Settlement Creek,

Feb 1923, Brass 257 (BRI, CANB); c. 35 km Wof Cloncurry

on Cloncurry-Mt Isa road, Mar 1977, Schmid AS 179 (BRI);

20 km S of Mt Isa on road to Boulia, Jun 1991, Halford

Q454 (BRI). North Kennedy District: Marble Creek mesa,

SE of Greenvale, Apr 1991, Bean 2940 (BRI). South

Kennedy District: slopes of Mt Hope, Apr 1992, Thompson

& Simon BUC446 (AD, BRI, DNA, NSW).

Distribution and habitat : Corchorus pumilio

is widespread across northern Australia from

the Kimberley, Western Australia to north¬

eastern Queensland (Map 2). It is recorded as

growing in open woodland and hummock

grassland communities, on shallow rocky, stony

or sandy soils, on hills, ridges and rocky

outcrops.

Phenology: Flowers and fruits have been

collected from February to August and in

November.

Typification: In the protologue of Corchorus

pumilio, Bentham (1863) cited two collections

“islands of the Gulf of Carpentaria, R. Brown”

and “Upper Victoria River, F. Mueller”. Seven

sheets of probable type material of Corchorus

pumilio have been located. Six sheets of the R.

Brown collections have been located (two from

K and one each at BRI CANB, MEL, NSW)

and one sheet of the F. Mueller collection

“Tableland between the Victoria River and

Sturts Creek, Feb 1856” located at K with the

name C. pumilio in red pencil in Bentham’s

hand. The sheet at Kew of R. Brown’s collection

labelled “Corchorus No. 32 desc. Carpentaria

island r” is chosen here as lectotype, because it

is part of the original material and has mature

fruit. Whether the other R. Brown sheets (MEL,

BRI, CANB, K, NSW) are all from the same

collection as the lectotype or separate

collections has not been ascertained.

Notes: Corchorus pumilio is confused with

C. sidoides but is distinguished from that by

having shorter fruit, smaller flowers, generally

fewer stamens in each flower and a sparser

indumentum on the leaves and stems.

Halford, Notes on Tiliaceae 4, Corchorus

611

The typical widespread form of C. pumilio

is a slender herbaceous shrub, generally green

in appearance with a sparse to moderately

dense, coarse white indumentum on all parts.

The collections Carr 3784 & Beauglehole

47562 (PERTH)(Geikie Gorge NP) and Hartley

14344 (CANB, DNA, PERTH)(Carr Boyd

Ranges) from the Kimberley, Western Australia

have a more robust habit than the typical form

of C. pumilio and have a ferruginous

indumentum on the young shoots. These

collections may represent a distinct species but

further collections and study are required.

16. Corchorus sericeus Ewart & O.B. Davies,

FI. N. Terr. 178 (1918). Type: Northern

Territory. Darwin and Gulf Region:

Borroloola, 9 Oct 1911, G. F. Hill (holo:

MEL [MEL223676]; iso: DNA, NSW).

Spindly or compact shrub to 1.5 m high; stems

sparingly to much branched, spreading to erect.

Indumentum on young shoots, branchlets,

leaves, stipules, peduncles, pedicels and bracts

grey-white, moderately dense to dense,

comprised of mostly stellate hairs but dendritic-

stellate and simple hairs also present. Stellate

hairs sessile or stipitate, up to 1.2 mm across;

stipes red-brown or white, straight or tortuous,

up to 0.2 mm long; rays soft to pliable, white

or rarely ferruginous, up to 0.6 mm long.

Dendritic-stellate hairs up to 1.5 mm long;

stipes white or red-brown, tortuous; rays

pliable, white, up to 0.6 mm long. Simple hairs

glandular, white, flexuous, up to 0.2 mm long.

Leaves with petioles 4-15(-20) mm long;

stipules subulate-linear, 2-10 mm long; lamina

narrowly to broadly ovate or elliptic-ovate,

1.5-7.5 cm long, 0.6-3 cm wide, l:w ratio

2-3:1, discolorous or concolorous; base obtuse

to rounded or rarely cordate; margin serrulate;

apex acute to rounded. Inflorescences umbellate

or racemose, 3-10-flowered, leaf-opposed,

solitary at upper nodes; peduncles 1-10 mm

long; pedicels 1-7 mm long, spreading to erect

in flower and fruit; bracts subulate-linear,

2-10 mm long. Flower buds ellipsoid, 2-5 mm

across; apex obtuse with 5 spreading caudae to

5 mm long. Sepals 5, persistent, narrowly

obovate-elliptic, 4-14 mm long, 1-3 mm wide;

abaxial surface with a dense indumentum of

stellate and dendritic-stellate hairs, the largest

hairs up to 1 mm long; adaxial surface villose

proximally, glabrous distally; apex caudate, up

to 5 mm long. Petals 5; lamina obovate, 3-7

mm long, 1-5 mm wide, glabrous; claw 0.4-1

mm long, sparsely pubescent. Androgynophore

0.1-0.3 mm long; annulus entire, 0.2-0.4 mm

long, glabrous. Stamens 25-80; filaments 2-5

mm long; anthers c. 0.3. Ovary globose, 0.7-2

mm across, densely stellate-tomentose, 3(rarely

4)-locular, with 2-8 ovules in each locule; style

2-5 mm long. Fruits globose, 2-4 mm across,

circular in transverse section, not constricted

between seeds, 3(rarely 4)-valved; apex rounded

rarely obtuse; indumentum dense of stellate

hairs up to 0.7 mm long. Seeds compressed

obovoid, c. 2 mm long.

Notes: Corchorus sericeus is most closely

related to C. laniflorus but differs from that in

having smaller flowers and fruits, and generally

shorter and coarser indumentum on most parts.

Corchorus sericeus as recognised here,

occurs from the Devils Marbles, Northern

Territory eastwards to Georgetown,

Queensland. Two subspecies are recognised and

can be distinguished using the following key.

Spindly shrubs to 1.5 m high; stems erect; primary stem unbranched for at

least 10 to 20 cm above ground level; inflorescences racemose, 7-10-

flowered; peduncles 4-15 mm long. 16a. C. sericeus subsp. sericeus

Open to compact shrubs to 1 m high; stems spreading; primary stem

branched at ground level; inflorescences umbellate, up to 5-flowered;

peduncles up to 2 mm long . 16b. C. sericeus subsp. densiflorus

16a. Corchorus sericeus Ewart & O.B. Davies

subsp. sericeus

Erect spindly shrub up to 1.5 m high. Primary

stem unbranched for at least 10 to 20 cm above

ground level. Leaves narrowly to broadly ovate,

acute to obtuse at apex. Inflorescences

racemose, 7-10-flowered; peduncles 4-15 mm

long. Indumentum on abaxial surface of sepals

comprised of mostly stellate hairs.

612

Selected specimens (from 23 examined): Northern

Territory. Darwin and Gulf Region: White Islet, May 1977,

McKey 181 (AD, DNA, MEL); 2 km East Lake Eames,

Vanderlin Island, Sir Edward Pellew Group, Jul 1988,

Thomson 2489 (BRI); Favenc Range, c. 160 km from

Borroloola on the road to Daly Waters, May 1974, Pullen

9316 (CANB, DNA); near McArthur R„ May 1947, Blake

17762 (BRI, MEL). Barkly Tablelands Region: Kilgour

Gorge, Mallapunyah Station, May 1984, Thomson 629

(DNA); 30 miles [c. 48 km] S of McArthur River Station, Jul

1948, Perry 1692 (CANB, DNA, MEL); 4 miles [c. 6 km]

N of Wollogorang Station, Jun 1948, Perry 1175 (BRI,

CANB, DNA); Wollogorang Station, Jun 1974, Henshall 423

(CANB, DNA); 15 km SW of Calvert Hills Station, on road

to Barkly Highway, Jun 1991, Halford Q585 (BRI).

Queensland. Burke District: Buchanan Creek W of

“Westmoreland” near the Queensland/Northem Territory

border, May 1974, Pullen 9206 (BRI, CANB, DNA); 3 miles

[c. 5 km] W of Westmoreland Station, Jun 1948, Perry 1350

(CANB, DNA).

Distribution and habitat : Corchorus sericeus

subsp. sericeus occurs in the subcoastal areas

around the Gulf of Carpentaria from Favenc

Range, Northern Territory to Westmoreland

Station in north-west Queensland (Map 8). It

is recorded as growing in open woodland

communities, on shallow sandy or gravelly soils

on sandstone or quartzite ridges. It is also

recorded on alluvial loams along watercourses.

Phenology: Flowers have been collected from

April to July, fruits from April to June.

Notes: The collection Craven 3911 (BRI,

CANB, DNA) from the Mac Arthur River area,

Northern Territory has atypically long racemose

inflorescences and the whole plant is generally

more hairy than in the typical form of

C. sericeus subsp. sericeus.

16b. Corchorus sericeus subsp. densiflorus

(Benth.) Halford comb. nov. & stat. nov.

Corchorus walcottii var. densiflorus Benth,

FI. Austral. 1: 279 (1863), ‘densiflora’.

Type: Gulf of Carpentaria, [without date,]

F. Mueller (lecto, here chosen: K (left

hand element); isolecto: MEL

[MEL227034]).

Open to compact shrub to 1 m high. Primary

stem branched at ground level. Leaves narrowly

ovate to ovate or elliptic-ovate, obtuse to

rounded rarely acute at apex. Inflorescences

umbellate, up to 5-flowered; peduncles up to 2

mm long. Indumentum on abaxial surface of

sepals comprised of stellate and dendritic-

stellate hairs.

Austrobaileya 6 (4): 581-629 (2004)

Selected specimens (from 47 examined): Northern

Territory. Darwin and Gulf Region: Upper Wearyan R., Jan

1989, Russell-Smith 7004 & Lucas (DNA). Barkly

Tablelands Region: 4 km S [of] Spear Waterhole,

Wollogorang, Jan 1989, Russell-Smith 6851 & Lucas

(DNA);4kmWofNo. 16 Bore, Benmara Station, May 1984,

Strong 160 (DNA); 3 km N of No. 19 Bore, Benmara Station,

May 1984, Low 17 (DNA); 10 miles [c. 16 km] NE of

Alexandria Station, Jun 1948, Perry 1487 (AD, BRI, CANB,

DNA, MEL, PERTH); 69 km N of Tennant Creek on Stuart

Highway, Jun 1991, Halford Q537 (BRI); Gibsons Creek,

35 miles [c. 56 km] N of Tennant Creek, Jul 1968, Must 195

(AD, BRI, CANB, MEL). Central Northern Region: Devils

Marbles, Mar 1955, Chippendale 937 (BRI, DNA).

Queensland. Cook District: Blue Hills, “Mount Surprise”,

49 km from Mount Surprise township, Mar 1988, Champion

350 (BRI); on Gulf Development Road, near bridge, 1 km E

of Georgetown, Apr 1990, Batianoff9 00402a & Smith (BRI).

Burke District: Lawn Hill NP, “Island Stack”, Jul 1985,

Williams 85070 (BRI); Bang Bang Jumpup to N of Donors

Hill, Apr 1974, Pullen 8911 (BRI, CANB, DNA); 28.5 km

from Mary Kathleen-Barkly Highway junction on traverse

to Mt MacNamara, May 1975, Can & Cole 9221 (BRI,

CANB); Cloncurry, Aug 1930, Blake 12649 (BRI); 5 kmN

of Barkly Highway on road to Lake Julius, Jun 1991, Halford

Q511 (BRI); 3 miles [c. 5 km] SE of Cloncurry township,

Mar 1954, Lazarides 4411 (BRI, CANB, DNA, MEL,

PERTH).

Distribution and habitat : Corchorus sericeus

subsp. densiflorus occurs from the Devils

Marbles, Northern Territory eastwards to

Georgetown, Queensland and from Sir Edward

Pellew Group, Northern Territory southwards

to Burnham Station, Queensland (Map 9). It

is recorded as growing in shrubland and open

woodland communities, mostly on shallow

sandy or stony soils, on rocky hills or plains

but also rarely on heavy alluvial soils along

drainage lines.

Phenology : Flowers have been collected from

January to November, fruits from March to July.

Notes: The distinguishing characters of this

subspecies are indicated in the key above.

The size of flowers and leaves, and the

thickness of indumentum on the abaxial surface

of the sepals varies greatly in this subspecies

as circumscribed here. The variation appears

to be continuous with no clear gaps that would

allow the subdivision of this taxon based on

any of these characters.

17. Corchorus sidoides F.Muell., Fragm. 3: 9

(1862). Type: [Northern Territory.]

Victoria River, May 1856, F. Mueller

(lecto, here chosen: MEL [MEL220812]).

Halford, Notes on Tiliaceae 4, Corchorus

613

Shrub to 1.5 m high; stems much branched,

procumbent to erect; young shoots with grey-

white or ferruginous indumentum.

Indumentum on branchlets, leaves, stipules,

peduncles, pedicels, and bracts grey-white,

moderately dense to very dense or rarely sparse,

comprised of stellate hairs. Stellate hairs sessile

or stipitate, up to 1.8 mm across; stipes white

or ferruginous, straight, up to 0.3 mm long;

rays firm to pliable, white or ferruginous, up to

1 mm long. Leaves with petioles 1-20 mm long;

stipules subulate-linear, 1-8 mm long; lamina

narrowly oblong, oblong-elliptic, narrowly

elliptic, ovate-elliptic, narrowly ovate to ovate

or narrowly obovate, 0.6-9 cm long, 0.2-3 cm

wide, l:w ratio 2-3.5:1, concolorous or

discolorous; base obtuse, rounded or attenuate;

margin dentate-serrate or serrate to serrulate;

apex acute to rounded. Inflorescences

umbellate, 4-7-flowered, leaf-opposed or

lateral, solitary at upper nodes; peduncles

0.5-5 mm long; pedicels 1-5 mm long,

spreading to erect in flower, recurved in fruit;

bracts subulate-linear to filiform-linear; 0.7-3

mm long. Flower buds obovoid to obovoid-

ellipsoid, 1-3 mm across, sometimes

longitudinally ridged; apex obtuse to

acuminate-caudate with 4 or 5 erect caudae to

1 mm long. Sepals 5(rarely 4), not persistent,

narrowly obovate, 3-9 mm long, 1-2 mm wide;

abaxial surface with a moderately dense to very

dense indumentum of stellate hairs up to 0.5

mm long; adaxial surface glabrous or stellate-

puberulous to stellate-villose proximally; apex

acute or acuminate-caudate, up to 1.5 mm long.

Petals 5, rarely 4; lamina narrowly obovate to

obovate, 2-7 mm long, 0.5-4 mm wide,

glabrous; claw 0.5-0.8 mm long, stellate-

pubescent on margins. Androgynophore

0.1-0.4 mm long; annulus entire or sinuate,

0.1-0.4 mm long, glabrous. Stamens (16-)20-50;

filaments 2-5 mm long; anthers 0.3-0.5 mm

long. Ovary cylindrical, 0.5-1 mm across,

densely stellate-puberulous or densely stellate-

villose, 2 or 3-locular, with 6-24 ovules in each

locule; style 1-4 mm long. Fruits subcylindrical

(5-) 18-60 mm long, 1-3 mm across, mostly

pendulous, straight, curved or very much

twisted, circular in transverse section, slightly

or markedly constricted between seeds, 2 or 3-

valved; apex attenuate up to 4 mm long,

orientated downward; indumentum moderately

dense to dense, of stellate hairs up to 1 mm

long. Seeds compressed obovoid, 1.5-3 mm

long. Chromosome n = 6, 8 and 7; 2n = 14

(Basak 1958).

Typification: In the protologue of Corchorus

sidoides, Mueller (1862) did not cite any

particular collection but stated “In locis

sterilioribus secus flumen Victoriae frequens”

[frequent in barren places along the Victoria

River]. Three sheets [MEL220811,

MEL220812, MEL227306] on loan to BRI

from MEL have been located that are labelled

C. sidoides and have Mueller as the collector

with the locality of collection as Victoria River.

The collection of this material would have

occurred during the Gregory expedition (1855—

1857) to northern Australia and predates the

publication of the name C. sidoides. The MEL

sheet marked MEL220812 is selected as

lectotype of the name Corchorus sidoides

F.Muell. as it agrees with the protologue and is

the more complete collection.

Notes: Corchorus sidoides is the most

widespread and common Corchorus species

across northern Australia. It is characterised

by having generally a spreading habit, relatively

long pendulous subcylindrical fruits, a stellate

indumentum on the fruit, sepals that are not

persistent and leaves that are mostly oblong,

oblong-elliptic or ovate-elliptic in outline. It is

most closely related to C. carnarvonensis,

C .congener, C.pumilio and C. tomentellus. For

differences from these species refer to

‘Affinities’ and ‘Notes’ under each species.

Corchorus sidoides is morphologically

variable, particularly in degree of fruit

distortion, shape and size of leaves and size of

stellate hairs. The types of both C. vermicularis

and C. rostrisepalus fall within the variation

of C. sidoides as circumscribed here. The

extreme forms within this species differ

considerably, but due to the difficulty in

assigning material to one or other forms I have

used the rank of subspecies to recognise these

forms rather than maintaining the previously

recognised species. Three subspecies are

recognised and can be distinguished using the

following key.

614 Austrobaileya 6 (4): 581-629 (2004)

1. Stems erect; indumentum on young shoots and buds ferruginous; hairs

fine, < 0.4 mm across; leaves narrowly ovate or narrowly elliptic, 3.5-9

cm long, 1.5-3 cm wide; fruits straight or slightly curved, prominently

constricted between seeds. 17c. C. sidoides subsp. rostrisepalus

Stems procumbent or spreading horizontally or if erect then other characters

not as above. 2

2. Fruits 2-valved rarely 3-valved, dark purplish-red rarely brown, 1-1.5 mm

across, weakly or strongly twisted; leaf laminae narrowly oblong or

narrowly oblong-elliptic rarely narrowly obovate, 0.6-3.5(-4) cm long,

0.2-1 (-1.5) cm wide; stellate hairs white, up to 0.5 mm across.

. 17b. C. sidoides subsp. vermicularis

Fruits mostly 3-valved occasionally 2-valved, pale-green to brown,

1-2.5 mm across, straight, curved or weakly twisted; leaf laminae ovate-

elliptic or narrowly ovate to ovate, occasionally narrowly oblong-elliptic

or narrowly oblong, 2-6 cm long, 0.8-2.5 cm wide; stellate hairs white

or ferruginous, up to 2 mm across. 17a. C. sidoides subsp. sidoides

17a. Corchorus sidoides F.Muell. subsp.

sidoides

Compact shrub to 0.9 m high; stems much

branched, spreading, rarely erect.

Indumentum on young shoots and buds grey-

white. Stellate hairs up to 2 mm across,

sessile or stalked; stalks white or

ferruginous, up to 0.3 mm long; rays white,

up to 1 mm long, pliable to stiff. Leaves with

petioles (2-)4-20 mm long; lamina narrowly

oblong-elliptic, ovate-elliptic, narrowly

ovate to ovate or narrowly obovate, 2-6 cm

long, 0.8-2.5 cm wide, concolorous or

discolorous; adaxial and abaxial surfaces

moderately dense to very dense stellate hairy;

base rounded; margin serrate; apex acute to

obtuse or rounded. Flower buds obovoid, 2-3

mm across; apex acute with 5 erect caudae

to 0.5 mm long. Sepals 5, narrowly obovate,

(3-)4-8(-9) mm long, 1-2 mm wide; apex

acute or acuminate-caudate, 0.5-1.5 mm

long; abaxial surface with moderately dense

to very dense indumentum of stellate hairs

up to 0.5 mm long; adaxial surface glabrous

or stellate-villose proximally. Stamens

(16-)20-50. Fruits (5-) 18-60 mm long, 1-3

mm across, pale-green to brown, straight,

curved or rarely weakly twisted, weakly to

strongly constricted between the seeds,

3(occasionally 2)-valved; apex attenuate up

to 3 mm long; indumentum moderately dense

to dense, of stellate hairs up to 0.5 mm long.

Chromosome No. 2n= 14 (Dattaeta/. 1966)

Selected specimens (from 167 examined ): Western

Australia. Gardner District: Ivanhoe Station, Ord R., Jun

1944, Gardner 7409 (PERTH). Dampier District: Gogo,

May 1951, Gardner 10254 (PERTH). Fitzgerald District:

Sandy River Gorge, Leopold Gorge, Apr 1988, Cranfield

6570 (PERTH). Fortescue District. 19.9 km S ofWittenoom

turnoff on the Great Northern Highway, Sep 1986, Chinnock

7015 (AD); 9 km SW of Mt Cecelia, c. 90 km SE of Shay

Gap, Jul 1984, Newbey 10553 (CANB, PERTH). Canning

District: Little Sandy Desert, May 1979, Mitchell 921 (AD,

DNA); head of Breaden Valley, Southesk Tablelands, Apr

1979, George 15504 (DNA, PERTH). Keartland District:

Rudall R. area, Aug 1971, Wilson 10585 (MEL, PERTH).

Carnegie District: 39 miles [c. 63 km] W of Jupiter Well,

Jul 1967, George 9087 (PERTH). Northern Territory.

Darwin and Gulf Region: c. 2 km S of Larrimah on Stuart

Highway, May 1985, Fryxell 4429 et al. (DNA, MEL).

Victoria River Region: Bullita Station, Gregory NP, Feb

1986, Wightman 2565 & Clark (DNA); Pinkerton Range,

Mar 1989, Dunlop 8121 & Leach (DNA, MEL); 20.8 miles

[c. 33 km] W [of] Inverway, May 1959, Chippendale 5945

(CANB, DNA, PERTH). Central Northern Region: Native

Gap, 71 miles [c. 114 km] N of Alice Springs, Jan 1969,

Nelson 1828 (AD, BRI, DNA, MEL). Central Southern

Region: 8 miles [c. 13 km] SE of Aileron, Mar 1955,

Winkworth 863 (MEL); Harts Range, 10 km S of Harts Range

Police Station, Oct 1977, Noble 21 (CANB); base of

breakaway - upper talus slope, Ruby Gap, Jul 1982 Purdie

2386 (CANB); Stokes Creek, Oct 1981, Latz 8916 (DNA).

Queensland. Burke District: 10 km SW of Mt Isa on the

road to Dajarra, Jun 1991, Halford Q520 (BRI). Gregory

North District: Warlus VI, Site R13, Mt Datson (ENE of

Boulia), Sep 1977, Purdie 1026 (BRI).

Distribution and habitat : Corchorus sidoides

subsp. sidoides is widespread across northern

Australia from the Pilbara, Western Australia

through the Northern Territory to north-western

Queensland (Map 3). It is recorded as growing

in hummock grassland, open woodland and

open forest communities, on sandy, loamy,

Halford, Notes on Tiliaceae 4, Corchorus

615

gravelly or clay soils, on sand dunes, plains

and hills.

Phenology: Flowers and fruits have been

collected throughout the year.

Notes : A number of collections from around

the Arnhem Land escarpment (eg. Martensz &

Schodde AE701 (BRI, CANB, MEL), Dunlop

4571 (DNA), Telford & Wrigleyl 589 (CANB),

Wightman 1382 & Craven (BRI) and Halford

Q1116 (BRI)) have shorter fruit than typical

(5-10 mm long) and a ferruginous indumentum

on the young shoots and flower buds.

The collections from near Kununurra

(.Pullen 10.879 (CANB) and Mackenzie

710209-6 (CANB) are typical in leaf size and

indumentum but have shorter fruit than

typically found in the species.

The collection Pullen 10.764 (BRI,

CANB) has a fairly erect habit to 1.5 m high

with the lower stem free of branches and has

generally smaller flowers and fruits than

typical, growing on red earths in shrub-

grassland.

Further collection and fieldwork may

show that a number of these entities warrant

formal recognition at specific or subspecific

rank.

17b. Corchorus sidoides subsp. vermicularis

(F.Muell.) Halford comb. nov. & stat.

nov.

Corchorus vermicularis F.Muell., Fragm. 3:

10 (1862). Type: [Western Australia/

Northern Territory.] Head of Sturts Creek,

Feb 1856, F. Mueller (holo: MEF

[MEF220810]).

Scorpia simplicifolia Ewart & A.H.K.Petrie,

Proceedings of the Royal Society of

Victoria ser. 2, 38 (1926). Type: Northern

Territory. Wycliffe, June 1924, A.J. Ewart

(holo: MEF [MEF227302]).

Diffuse shrub to 0.5 m high; stems much

branched, spreading. Indumentum on young

shoots and buds grey-white. Stellate hairs up

to 0.5 mm across, sessile or stalked; stalk

white or reddish-brown, up to 0.1 mm long;

rays white, up to 0.3 mm long, stiff. Feaves

with petioles 1—5(—10) mm long; lamina

narrowly oblong, oblong-elliptic or rarely

narrowly obovate, 0.6-3.5(-4) cm long,

0.2-1 (-1.5) cm wide, concolorous; adaxial

surface sparsely to densely stellate hairy or

rarely glabrous; abaxial surface sparsely to

densely stellate hairy; base obtuse to rounded

or rarely attenuate; margin serrate or

dentate-serrate; apex obtuse to rounded or

rarely acute. Flower buds obovoid-ellipsoid,

1-2 mm across; apex obtuse with 4 or 5 erect

caudae to 0.2 mm long. Sepals 5, rarely 4,

narrowly obovate, 4-7 mm long, 1-2 mm

wide; apex acuminate c. 0.7 mm long;

abaxial surface with moderately dense to

dense indumentum of stellate hairs up to 0.5

mm long; adaxial surface puberulous

proximally, glabrous distally. Stamens

20-30(-40). Fruits 20-35 mm long, 1-2 mm

across, dark purplish-red or rarely brown,

weakly to strongly twisted, strongly

constricted between the seeds, 2(rarely 3)-

valved; apex attenuate up to 4 mm long;

indumentum moderately dense to dense, of

stellate hairs up to 0.3 mm long.

Selected specimens (from 67 examined): Western

Australia. Dampier District: St Mary’s School, Broome, Apr

1987, Foulkes 13 (CANB, PERTH); Munkajarra

[Munkayarra], 20 km S of Derby, Apr 1983, Fryxell 3848

(CANB, MEL, PERTH); 9 km SE of Frazier Downs Station,

Jul 1987, Ingleby JV28 (PERTH), c. 1225 km on Northwest

Coastal Highway, Aug 1971, Ashby 3034 (AD); oldFitzroy

R. crossing, Apr 1988, Cranfield 6414 (CANB, PERTH).

Canning District: N of Dragon Tree Soak, Great Sandy

Desert, Aug 1977, George 14768 (CANB, PERTH); just S

of Tobin Lake, Great Sandy Desert, May 1979, George 15649

(CANB, DNA, PERTH). Northern Territory. Darwin and

Gulf Region: 75 km E of Stuart Highway along Carpentaria

Highway, Apr 1992, Halford Q1073 (BRI). Barkly

Tablelands Region: 8 miles [c. 13 km] S of old Highland

Plains, Jul 1976, Henry 253 (AD, DNA, MEL); 30 km N of

Tennant Creek, Stuart Highway, Apr 1992, Halford Q1201

(BRI). Northern Central Region: Sangsters Bore, Tanami

Desert, Sep 1978, Henshall 2277 (DNA); 2 km W of Lake

Surprise, Tanami Desert, Jun 1985, Latz 10072 (DNA); 77

mil es [c. 124 km] WSW [of] The Granites, Aug 1970, Dunlop

1812 (CANB, DNA); CentralMt Stuart, Jul 1974,Latz 5571

(BRI, CANB). Queensland. Cook District: site EU293,

Barwidgee Homestead, E of Branch Creek, Feb 1992, Godwin

C3718 (AD, BRI, DNA); 22 km E of Croydon, Apr 1992,

Halford Q986 (BRI). Burke District: between Doomadgee

Aboriginal Station and old “Corinda” outstation, May 1974,

Pullen 9077 (CANB, DNA). Gregory North District: Oban

Station, about 62 miles [c. 100 km] SW of Mt Isa, Woodend

Bore, Dec 1947, Everist 3342 (BRI, CANB). Mitchell

District: near Lochnagar, Nov 1935, Blake 10302 (BRI,

CANB). South Kennedy District: 27.5 km W of St Anns

Homestead, Jun 1992, Thompson & Sharpe BUC838 (BRI).

616

Austrobaileya 6 (4): 581-629 (2004)

Distribution and habitat : Corchorus sidoides

subsp. vermicularis is widespread across

northern Australia from the Pilbara, Western

Australia, through the Northern Territory to the

north-east coast of Queensland (Map 5). It is

recorded as growing on sandy, loamy or

gravelly soils, in hummock grassland,

shrubland, open woodland and open forest

communities, on plains, sand dunes and hills.

Phenology: Flowers and fruits have been

collected throughout the year.

Notes: Generally a smaller more diffuse shrub

than the other subspecies with slender stems,

smaller leaves and twisted fruit. In the

Kimberley region this subspecies grades into

small leaf forms of C. sidoides subsp. sidoides.

17c. Corchorus sidoides subsp. rostrisepalus

(Domin) Halford comb. nov. & stat. nov.

Corchorus rostrisepalus Domin, Biblioth.

Bot. 89: 383 (1928). Type: [Northern

Territory.] Carpentaria Island g,

[Vanderlin Island, 15 Dec 1802,] R.

Brown (lecto, here chosen: K; ?isolecto:

BRI, CANB, MEL).

Shrub to 1 m high; stems sparingly to much

branched, erect. Indumentum on young shoots

and buds ferruginous. Stellate hairs up to 0.3

mm across, sessile or stalked; stalks white or

reddish-brown, up to 0.2 mm long; rays white

or ferruginous, up to 0.2 mm long, pliable.

Leaves with petioles 3-13 mm long; lamina

narrowly ovate or narrowly elliptic, 3.5-9 cm

long, (0.5-) 1.5-3 cm wide, discolorous; adaxial

and abaxial surfaces moderately dense to dense

or rarely sparsely stellate hairy; base obtuse or

attenuate; margin serrate to serrulate; apex

obtuse or acute. Flower buds obovoid, 2-3 mm

across, longitudinally ridged distally; apex

acuminate with 5 erect caudae to 1 mm long.

Sepals 5, narrowly obovate, 5-6 mm long, c. 2

mm wide; apex acuminate-caudate up to 1 mm

long; abaxial surface with moderately dense to

dense indumentum of stellate hairs up to 0.2

mm long; adaxial surface stellate-pubescent

proximally, glabrous distally. Stamens 30-35.

Fruits 20-55 mm long, 1-2 mm across, pale-

green to brown, straight or slightly curved,

prominently constricted between seeds, 2(rarely

3)-valved; apex attenuate up to 4 mm long;

indumentum moderately dense, of stellate hairs

up to 0.2 mm long.

Selected specimens (from 15 examined): Northern

Territory. Darwin and Gulf Region: 17 miles [c. 27 km]

NNE [of] Mainoru, Jun 1972, Byrnes 2613 (DNA); South

Bay, Bickerton Island, in the Gulf of Carpentaria, Jun 1948,

Specht 605 (AD, BRI, CANB, MEL, NSW); Groote Eylandt,

4 km W [of] Umbakumba, Jul 1987, Russell-Smith 2738 &

Lucas (DNA); Hemple Bay, Groote Eylandt, in the Gulf of

Carpentaria, Apr 1948, Specht 283 (AD, BRI, CANB, MEL,

NSW); Angurugu, Groote Eylandt, May 1972, Levitt [DNA

4462] (DNA); Nitmiluk Gorge NP, Feb 1990, Evans 2938

(DNA, CANB, MEL); Katherine Gorge [Nitmiluk] NP, Mar

1971, Dunlop & Byrnes 2157 (CANB, DNA, MEL); 12

miles [c. 19 km] NE of Katherine, Jan 1965, Wilson 79

(CANB, DNA); mouth of Rosie Creek, Lorella, Jan 1989,

Russell-Smith 6752 & Lucas (BRI, DNA).

Distribution and habitat: Corchorus sidoides

subsp. rostrisepalus occurs in north-eastern

Northern Territory from near Katherine

eastward to the islands of the Gulf of

Carpentaria (Map 5). It is recorded as growing

in open woodland and open forest communities,

on shallow rocky soils on hills or sandy soils

on alluvial flats.

Phenology: Flowers have been collected June,

July and from December to April, fruits in

December, January and from March to June.

Notes: Corchorus sidoides subsp. rostrisepalus

has a more erect habit, and generally a finer

indumentum on its leaves and stems than the

other subspecies of Corchorus sidoides.

Typification: In the protologue of

C. rostrisepalus, Domin (1928) referred to ‘

Carpentaria Islands, R. Brown als

C. vermicularis'. There are three sheets at K

of original Brown material from the

Carpentaria Islands. The sheet with the small

label with the information ‘Corchorus

vermicularis No. 45 desc. Carpentaria Island

g’ in Browns handwriting is here selected as

lectotype. There are a number of possible

duplicates of this material at a number of

institutions (BRI, MEL, CANB) that have

varying label details. Whether these other R.

Brown sheets are all from the same collection

as the lectotype or separate collections has not

been ascertained.

18. Corchorus subargentus* Halford sp. nov.

quoad staturam et formam foliorum C.

sidoidi subsp. rostrisepalo et C. obclavato

similis autem ab utroque fructibus erectis

* should be subargenteus ' (PDB 2005)

Halford, Notes on Tiliaceae 4, Corchorus

617

non pendulis differt. Corchorus

subargenteus maxime arete affinis C.

sublato et C. leptocarpo\ ab illo floribus

majoribus (sepalis 10-11 mm longis,

petalis 8-10 mm longis, filamentis

staminalibus 5-6 mm longis, stylo 5-6

mm longo comparitis sepalis 7-9 mm

longis, petalis 4-6 mm longis, filamentis

staminalibus 3-4 mm longis, stylo 2-3

mm longo), indumentum caulium

foliorumque (10-11 mm longis non

12-14 mm longis) petalis angustioribus

(2-3 mm latis non c. 7 mm latis),

filamentis staminalibus brevioribus (5-6

mm longis non 7-9 mm longis) differt.

Typus: Queensland. North Kennedy: 13

km along Laroona road, off Paluma to

Ewan road, 19°11\ 145°55\ 15 April

1996, PI. Forster PIF18983 & T. Ryan

(holo: BRI; iso: MEL, distribuendi).

Shrub to 2 m high; stems sparingly to much

branched, erect; young shoots with silvery-grey

or ferruginous indumentum. Indumentum on

branchlets, leaves, stipules, peduncles, pedicels

and bracts silvery-grey, dense, comprised of

stellate hairs. Stellate hairs sessile or stipitate,

up to 0.6 mm across; stipes white or red-brown,

straight, up to 0.2 mm long; rays pliable, white

or ferruginous, up to 0.4 mm long. Leaves with

petioles 5-7 mm long; stipules subulate-linear,

2-3 mm long; lamina narrowly ovate, 3-8 cm

long, 1-1.5 cm wide, l:w ratio 3-5.3:1,

discolorous; base rounded; margin serrulate;

apex rounded. Inflorescences umbellate or

racemose, 5-7-flowered, leaf-opposed or

lateral, solitary at upper nodes; peduncles

7-15 mm long; pedicels 2-7 mm long,

spreading to erect in flower, erect in fruit; bracts

subulate-linear, 2-4 mm long. Flower-buds

obovoid-ellipsoid, 3-4 mm across, slightly

longitudinally ridged; apex acute with 5

spreading caudae up to 1 mm long. Sepals 5,

not persistent, narrowly obovate, 10-11 mm

long, c. 2 mm wide; abaxial surface with a

dense indumentum of stellate hairs up to 0.5

mm long; adaxial surface stellate-pubescent

proximally, glabrous distally; apex caudate, up

to 2 mm long. Petals 5; lamina narrowly

obovate, 8-10 mm long, 2-3 mm wide,

glabrous; claw c. 0.7 mm long, stellate-

pubescent on margins. Androgynophore c. 0.2

mm long; annulus entire, c. 0.1 mm long,

glabrous. Stamens 60-70; filaments 5-6 mm

long; anthers c. 0.5 mm long. Ovary

cylindrical, 0.9-1 mm across, densely

stellate-puberulous, 3-locular, with 34-38

ovules in each locule; style 5-6 mm long.

Fruits subcylindrical, 20-50 mm long, 2-2.5

mm across, 7-20 times longer than wide, +

circular in transverse section, erect , ±

straight, slightly constricted between seeds,

3-valved; apex attenuate, 1-2 mm long,

orientated upward; indumentum moderately

dense to dense, of stellate hairs up to 0.1

mm long. Seeds compressed obovoid or

columnar; 1-3 mm long. Fig. 13.

Additional specimens : Queensland. North Kennedy

District: 2.8 km S of Running River on Ewan-Laroona road,

Feb 1996, Camming 7548 (BRI).

Distribution and habitat: Corchorus

subargentus is confined to north-eastern

Queensland where it is known only from the

Running River area, approximately 90 km

W of Townsville (Map 3). It is recorded as

growing in eucalypt woodland with Triodia

sp. in the understorey, on sandy soils on

granite-quartz ridges.

Phenology: Flowers have been collected in

February and April, fruits in April.

Affinities: Corchorus subargentus is similar

in stature and leaf size to C. sidoides subsp.

rostrisepalus and C. obclavatus. Corchorus

subargentus differs from both of these by

having erect rather than pendulous fruit.

Corchorus subargentus is most closely

related to C. subulatus and C. leptocarpus.

It differs from C. sublatus by having larger

flowers (sepals 10-11 mm long, petals 8-10

mm long, staminal filaments 5-6 mm long,

style 5-6 mm long compared with sepals 7-9

mm long, petals 4-6 mm long, staminal

filaments 3-4 mm long, style 2-3 mm long),

slightly coarser indumentum on the stems

and leaves, and longer and stouter peduncles

(7-15 mm long compared with 2-5 mm

long). Corchorus subargentus differs from

C. leptocarpus by having a more slender

stature and shorter sepals (10-11 mm long

compared with 12-14 mm long), narrower

petals (2-3 mm wide compared with c. 7 mm

wide) and shorter staminal filaments (5-6

mm long compared with 7-9 mm long).

618

Austrobaileya 6 (4): 581-629 (2004)

Fig. 13. Corchorus subargentus. A. branchlet with flower buds and fruit, x 1. B. fruit, x 1.5. C. ventral view of sepal, x 4. A

& B from ForsterPTF 18983 (BRI); C from Cumming 7548 (BRI). Del. W. Smith.

Halford, Notes on Tiliaceae 4, Corchorus

619

Etymology: The specific epithet is from the

Latin sub- somewhat, argentea, silvery, and is

in reference to the appearance of the foliage of

this species.

19. Corchorus sublatus Halford sp. nov. quoad

staturam et amplitudinem foliorum C.

sidoidi subsp. rostrisepalo et C. obclavato

similis autem autem ab utroque fructibus

erectis non pendulis differt. Per

charactereum eius fructuum C. sublatus ,

C. leptocarpo et C. subargenteo similis.

Ab eis C. sublatus floribus minoribus

fructibus angustioribus differt (vide

tabulam 1 pro comparationibus). Addite

C. sublatus a subargenteo indumento

tenuiore pedunculis brevioribus (2-5 mm

longis non 7-15 mm longis) differt.

Typus: Northern Territory. Darwin and

Gulf Region: Baroalba Spring, Kakadu

NP, 16 April 1992, D. Halford Q1114

(holo: DNA; iso: BRI, L, MEL,

distribuendi).

Shrub to 1.5 m high; stems sparingly to much

branched, erect; young shoots with grey-white

or ferruginous indumentum. Indumentum on

branchlets, leaves, stipules, peduncles, pedicels

and bracts grey-white, moderately dense to

dense, comprised of stellate hairs. Stellate hairs

sessile or stipitate, up to 0.2 mm across; stipes

white, straight, up to 0.2 mm long; rays pliable,

white or ferruginous, up to 0.1 mm long. Leaves

petioles 5-15 mm long; stipules subulate-linear,

3-5 mm long; lamina narrowly ovate or rarely

narrowly oblong, 3-11 cm long, 0.7-2.5 cm

wide, l:w ratio 4-5.5:1, discolorous; base

rounded or rarely attenuate; margin serrulate;

apex acute. Inflorescences umbellate, 6-8-

flowered, leaf-opposed or lateral, solitary at

upper nodes; peduncles 2-5 mm long; pedicels

2-5 mm long, spreading to erect in flower,

spreading to recurved in fruit; bracts subulate-

linear, 3-5 mm long. Flower-buds obovoid-

ellipsoid, 4-5 mm across, longitudinally ridged;

apex acute with 5 spreading caudae up to 2 mm

long. Sepals 5, not persistent, narrowly obovate,

7-9 mm long, 1-2 mm wide; abaxial surface

with a moderately dense indumentum of stellate

hairs up to 0.5 mm long; adaxial surface

stellate-pubescent proximally, glabrous distally;

apex caudate, up to 3 mm long. Petals 5; lamina

narrowly obovate to obovate, 4-6 mm long,

2-4 mm wide, glabrous; claw 0.7-1 mm long,

stellate-pubescent on margins. Androgynophore

0.3-0.6 mm long; annulus sinuate or entire,

0.2-0.3 mm long, glabrous. Stamens 40-60;

filaments 3-4 mm long; anthers c. 0.5 mm long.

Ovary cylindrical, 0.7-0.8 mm across, densely

stellate-puberulous, 3-locular, with 20-26

ovules in each locule; style 2-3 mm long. Fruits

subcylindrical, 20-50 mm long, 1-2 mm

across, 10-25 t im es longer than wide, erect,

straight or if on recurved pedicels then abruptly

bent near base so that the fruit is perpendicular

with the apex pointing upwards, circular in

transverse section, slightly constricted between

seeds, 3-valved; apex attenuate, 1-2 mm long;

indumentum moderately dense to dense of

stellate hairs up to 0.2 mm long. Seeds

compressed obovoid or columnar; 1-3 mm

long. Fig. 14.

Selected specimens (from 8 examined): Northern

Territory. Darwin and Gulf Region: Kakadu NP, Baroalba

Springs, May 1983, Fryxell & Craven 4270 (CANB, DNA);

Baroalba Spring, Kakadu NP, Apr 1992, Halford Q1131

(BRI); near mouth of Sawcut Gorge, 28.5 km SSE of Jabiru

East, Jun 1980, Craven 6279 (CANB); c. 7 miles [c. 11 km]

W of Mt Gilruth, Mar 1973, Lazarides 7951 (BRI, CANB,

DNA, NSW); 1 km upstream from Twin Falls, Mar 1988,

Fensham 880 (DNA).

Table 2. Morphological comparison of Corchorus sublatus , C. subargentus and C.

leptocarpus.

Characters

C. sublatus

C. subargentus

C. leptocarpus

sepals (mm)

7-9 x 1-2

10-11 x c. 2

12-14 x c. 3

petals (mm)

4-6 x 2-4

8-10 x 2-3

8-10 x c. 7

fruits width (mm)

1-2

2-3

620

Austrobaileya 6 (4): 581-629 (2004)

Fig. 14 . Corchorus sublatus. A. branchlet with flowers, x 1. B. fruit, x 3. C. ventral view of sepal, x 6. A from Halford Q1114

(BRI); B, C from Halford Q1112 (BRI). Del. W. Smith.

Distribution and habitat : Corchorus sublatus

is confined to Arnhem Land in the Northern

Territory, from Mt Gilruth southwards to Twin

Falls (Map 6). It is recorded as growing in

heathland, woodland and open forest

communities on sandy soils on talus slopes or

on gravelly soils on sandstone plateaus.

Phenology: Flowers have been collected in

March and April, fruits from April to June.

Affinities: Corchorus sublatus is similar in

stature and leaf size to C. sidoides subsp.

rostrisepalus and C. obclavatus. Corchorus

sublatus differs from both of these by having

erect rather than pendulous fruit. In this

character C. sublatus resembles C. leptocarpus

and C. subargentus. Corchorus sublatus differs

Halford, Notes on Tiliaceae 4, Corchorus

621

from both of these species by having smaller

flowers and narrower fruits (see Table 2 for

comparison). In addition, C. sublatus differs

from C. subargentus by having a finer

indumentum and shorter peduncles (2-5 mm

long compared with 7-15 mm long).

Etymology: The specific epithet refers to the

orientation of the fruit; Latin sublatus raised

aloft.

20. Corchorus tectus Halford sp. nov. a C.

sericeio foliis anguste oblongis usque

oblongis distinguenda. Typus: Western

Australia. Fortescue District: 53 miles

[c. 85 km] S of Roebourne on Wittenoom

road, 3 March 1962, A.S. George 3488

(holo: PERTH).

Open shrub to 70 m high; stems much

branched, spreading. Indumentum on young

shoots, branchlets, leaves, stipules, peduncles,

pedicels and bracts grey-white, moderately

dense to dense, comprised of mostly stellate

hairs but dendritic-stellate hairs also present.

Stellate hairs sessile or stipitate, up to 1 mm

across; stipes red-brown or white, straight or

tortuous, up to 0.2 mm long; rays soft to pliable,

white, up to 0.6 mm long. Dendritic-stellate

hairs up to 0.5 mm long; stipes white or red-

brown, tortuous; rays pliable, white, up to 0.6

mm long. Leaves with petioles 7-15 mm long;

stipules subulate-linear, 1-3 mm long; la min a

narrowly oblong to oblong, 2-4.5 cm long,

0.6-1.5 cm wide, l:w ratio 2.8-3.5:1,

discolorous or concolorous; base obtuse to

rounded or rarely cordate; margin crenulate;

apex acute to rounded. Inflorescences umbellate

or racemose, 4-8-flowered, leaf-opposed,

solitary at upper nodes; peduncles 1—10 mm

long; pedicels 1-7 mm long, spreading to erect

in flower and fruit; bracts subulate-linear, 2-5

mm long. Flower buds ellipsoid, 2-4 mm

across; apex obtuse with 5 spreading caudae to

2 mm long. Sepals 5, persistent, narrowly

obovate-elliptic, 5-6.5 mm long, 1-1.5 mm

wide; abaxial surface with a dense indumentum

of stellate and dendritic-stellate hairs, the

largest hairs up to 1 mm long; adaxial surface

villose proximally, glabrous distally; apex

caudate, up to 2 mm long. Petals 5; lamina

obovate, 5-7 mm long, 3-5 mm wide, glabrous;

claw 0.7-0.8 mm long, sparsely pubescent.

Androgynophore 0.1-0.3 mm long; annulus

entire, 0.2-0.4 mm long, glabrous. Stamens 45-95;

filaments 2.5-5.5 mm long; anthers c. 0.5.

Ovary globose, 1.5-2 mm across, densely

stellate-tomentose, 3-locular, with 2-8 ovules

in each locule; style 3-5 mm long. Fruits

globose, 2-4 mm across, circular in transverse

section, not constricted between seeds, 3-

valved; apex rounded rarely obtuse;

indumentum dense of stellate hairs up to 0.7

mm long. Seeds compressed obovoid, 1.2-1.8

mm long. Fig. 15.

Selected specimens (from 11 examined): Western

Australia. Fortescue District: Robe R., between Onslow and

Roebourne, Aug 1966, Butler 20 (PERTH); Fortescue R.,

Jun 1878, Forrest s.n. [MEL227315] (MEL); Fortescue R.,

1895, Cusack s.n. [MEL560600] (MEL); Roebourne, 1897,

Cusack s.n. [MEL1599108] (MEL); 3km NE of Three Peak

Hills, Pannawonica road, Mar 1984, Newbey 9889 (PERTH);

Hamersley Range, Aug 1958, Ride s.n. [PERTH 1524739]

(PERTH); Hamersley Range, Ride s.n. [PERTH 1524690]

(PERTH).

Fig. 15. Corchorus tectus. A. branchlet with flowers, x 1. B.

fruit with persistent sepals removed, x 6. C. ventral view of

sepal, x 6. Afrom van Leeuwen 4376 (BRI); B, C from Bui ter

20 (BRI). Del. W. Smith.

622

Austrobaileya 6 (4): 581-629 (2004)

Distribution and habitat : Corchorus tectus

occurs in north-western Western Australia,

from Robe River eastward to near Millstream

Station (Map 7). It is recorded as growing in

open shrubland communities on gravelly soils

along watercourses.

Phenology : Flowers have been collected in

March, June, August and September, fruits in

March and August.

Affinities'. Corchorus tectus is similar to

C. sericeus but differs from that by having

narrowly oblong to oblong leaves.

Etymology: The specific epithet is from Latin

tectus, meaning ‘covered’, in reference to the

persistent calyx lobes that cover the fruit of this

species.

21. Corchorus tomentellus F.Muell., Fragm.

3: 10 (1862). Type: [Queensland.] Mackenzie

River, [without date,] E Mueller s.n. [lecto, here

chosen: MEL [MEL220813]).

Shrub to 0.3 m high; stems much branched,

spreading. Indumentum on young shoots,

branchlets, leaves, stipules, peduncles, pedicels

and bracts grey-white, moderately dense,

comprised of stellate hairs. Stellate hairs sessile

or stipitate, up to 0.7 mm across; stipes red-

brown, straight, up to 0.3 mm long; rays firm,

white, up to 0.5 mm long. Leaves with petioles

3-5 mm long; stipules subulate-linear, 1-3 mm

long; lamina ovate, 1.5-3.5 cm long, 0.8-2 cm

wide, l:w ratio 1.3-2.2:1, discolorous; base

rounded; margin serrate; apex acute to obtuse.

Inflorescences umbellate, 2 or 3-flowered, leaf-

opposed, solitary at upper nodes; peduncles

1-2 mm long; pedicels 2-4 mm long, spreading

to erect in flower, recurved in fruit; bracts

subulate-linear, 1-3 mm long. Flower buds

ellipsoid, 2-3 mm across, apex acute. Sepals

5, not persistent, narrowly obovate, 6-7 mm

long, 1-2 mm wide; abaxial surface with a

moderately dense indumentum of stellate hairs

up to 0.5 mm long; adaxial surface stellate-

pubescent proximally, glabrous distally; apex

acute to acuminate, up to 0.4 mm long. Petals

5; lamina obovate, 5-7 mm long, 2-5 mm wide,

glabrous; claw c. 0.7 mm long, stellate-

pubescent on abaxial surface and margins.

Androgynophore 0.3-0.4 mm long; annulus

entire, c. 0.2 mm long, glabrous. Stamens

50-60; filaments 2-4 mm long; anthers 2-4

mm long. Ovary cylindrical; c. 0.8 mm

across, densely stellate-puberulous, 2 or 3-

locular, with c. 26 ovules in each locule; style

c. 3 mm long. Fruits subcylindrical, 15-65

mm long, 0.7-1.5 mm across, 7-35 times

longer than wide, pendulous, straight or

slightly curved, circular in transverse

section, markedly constricted between seeds,

2 or 3-valved; apex acute or attenuate, 1-3

mm long, orientated downward;

indumentum moderately dense to dense, of

stellate hairs up to 0.4 mm long. Seeds

compressed obovoid, c. 2 mm long.

Selected specimens (from 18 examined): Queensland.

South Kennedy District: tributary of Hazelwood Creek near

pipeline, Apr 1978, Byrnes & Clarkson 3779 (BRI).

Leichhardt District: near Lake Elphinstone, Jan 1993,

Fensham 441 (BRI) Carborough Range, 1 kmNW of Lake

Elphinstone outlet, Telford 11125 & Rudd (BRI); telecom

road, 14 km E of Comet, Bean 7520 & Forster (BRI); South

Blackwater Mine, Laleham, Jan 1986, Thompson s.n.

[AQ399040] (BRI); South Blackwater Mine, Dec 1990,

Thompson 10 (BRI); Brigalow Research Station, 30 km NW

of Theodore, Apr 1977, Johnson 3517 & Batianoff( BRI);

Brigalow Research Station, 32 km NW of Theodore, Jul 1970,

Johnson 2890 (BRI); ‘Humboldt’, 45 km NE of Rolleston,

Bean 9573 (BRI); near ‘Moorooloo’, E of Springsure, Bean

14172 (BRI). Burnett District: “Narayen”, Mundubbera,

Feb 1967, TothillN325 (BRI); “Narayen” about 30miles [c.

48 km] W of Mundubbera, Feb 1968, Tothill N443 (BRI).

Distribution and habitat : Corchorus

tomentellus occurs in subcoastal areas of

central and southern Queensland from near

Nebo southward to Mundubbera (Map 3). It

is recorded as growing in shrubland,

eucalypt woodland and eucalypt open forest

communities on sandy soils or in brigalow

open forest communities on clay soils.

Phenology: Flowers have been collected in

April, October and from December to March,

fruits in April, July, October and from

December to March.

Typification: In the protologue of Corchorus

tomentellus, Mueller (1862) did not cite any

particular collection but stated “In

graminosis herbidis ad flumina Dawson et

MacKenzie”. A single collection

[MEL220813] (Mackenzie River, trop.

Austr.) which can be considered part of the

original material that Mueller used to draw

up his description of this species has been

located at MEL. The collection has flowers

Halford, Notes on Tiliaceae 4, Corchorus

623

and is chosen as the lectotype to Mueller’s

name C. tomentellus.

Notes: Corchorus tomentellus is closely related

to C. sidoides but can be distinguished from

that by its generally sparser indumentum on

most parts, relatively broader and shorter leaves

and generally narrower fruit.

22. Corchorus walcottii F.Muell., Fragm. 3: 9

(1862); Corchorus walcottii F.Muell. var.

walcottii, Benth., FI. Austral. 1: 279

(1863). Type: [Western Australia.]

Hearson Island, P. Walcott (lecto, here

chosen: MEL [MEL223678]).

Corchorus sp. Burrup (G.Craig 235),

Paczkowska & Chapman (2000).

Shrub to 1.5 m high; stems much branched,

spreading. Indumentum on young shoots,

branchlets, leaves, stipules, peduncles, pedicels

and bracts grey-white, moderately dense to

dense, comprised of mostly stellate hairs but

simple and dendritic-stellate hairs also present.

Stellate hairs sessile or stipitate, up to 1.5 mm

across; stipes, white, straight, up to 0.5 mm

long; rays soft, white, up to 1.5 mm long.

Dendritic-stellate hairs up to 2 mm long; stipes

white or red-brown, tortuous; rays soft, white,

up to 0.5 mm long. Simple hairs glandular, dull

yellow, flexuous, 1-4 mm long. Leaves with

petioles 7-30 mm long; stipules subulate-linear,

4-25 mm long; lamina narrowly ovate to

broadly ovate, 3-9 cm long, 2.5-6 cm wide,

l:w ratio 1.2-2.5:1, concolorous; base rounded

or slightly cordate; margin serrate; apex obtuse

to rounded. Inflorescences umbellate, 3-5-

flowered, leaf-opposed, solitary at nodes;

peduncles 5-25 mm long; pedicels 5-20 mm

long, spreading to erect in flower and fruit;

bracts narrowly ovate, 2-17 mm long. Flower-

buds globose, 7-10 mm across; apex obtuse

usually with 5 spreading caudae up to 2 mm

long. Sepals 5, not persistent, narrowly obovate-

elliptic, 8-18 mm long, 3-5 mm wide; abaxial

surface with a very dense indumentum of

mostly stellate and simple hairs but dendritic-

stellate hairs occasionally present, the largest

hairs up to 1.5 mm long; adaxial surface stellate

hairy over whole surface or restricted to the

proximal third; apex acute, acuminate or

caudate, 1-4 mm long. Petals 5; la min a obovate

to broadly obovate, 7-13 mm long, 6-10 mm

wide, glabrous; claw 0.3-1 mm long, stellate

pubescent on abaxial surface and margins.

Androgynophore c. 0.2 mm long; annulus

entire, c. 0.2 mm long, glabrous. Stamens 80-180;

filaments 3-7 mm long; anthers c. 0.5 mm long.

Ovary subglobose or shortly pentagonal-

cylindrical, c. 1.8 mm across, densely stellate-

tomentose, 5-locular, with 24-26 ovules in each

locule; style 3-7 mm long. Fruits narrowly

ovoid-ellipsoid or subcylindrical, 7-25 mm

long, 3-9 mm across, 2-5 t im es longer than

wide, straight or slightly curved, circular in

transverse section, not constricted between

seeds, 4 or 5-valved; apex rounded;

indumentum dense of dendritic-stellate and a

few simple glandular hairs, the largest hairs

up to 2 mm long. Seeds compressed obovoid,

c. 2 mm long. Fig. 16. Chromosome No. 2n =

14 (Islam & Qaiyum 1961; Datta et al. 1966).

Selected specimens (from 35 examined ): Western

Australia. Fortescue District: 2 km S of landing strip,

Barrow Island, May 1964, Goodall 1520 (PERTH); Sholl

Island, Oct 1949, Seventy s.n. [PERTH1521519] (PERTH);

Dolphin Island, Dampier Archipelago, Jun 1962, Royce 7169

(PERTH); Dampier turnoff (south), Nov 1979, Demarz 2864

(PERTH); North West Coastal Highway, c. 15 km by road

WSW of main turnoff to Dampier, c. 8 km by road ENE of

Karratha Homestead turnoff, Aug 1977, Jackson 3034 (AD);

5 km NE of George R. crossing of North West Coastal

Highway, 42 km from Roebourne, Aug 1977, Telford 6547

(CANB); Karratha, Jul 1981, Craig 235 (PERTH); Pt

Samson, Jul 1981, Craig 212 (PERTH); Woodstock Station,

(S of Port Hedland), May 1958, Burbidge 5974 (AD,

PERTH); Woodstock Station, Apr 1958, Burbidge 5811 (AD,

PERTH); 90 km N of Nullagine, Great Northern Highway,

May 1979, Mitchell 1245 (PERTH). Northern Territory.

Central Northern Region: Mongrel Downs, Apr 1971,

Dunlop 2105 (AD, BRI, MEL); S of Mongrel Downs Station,

Aug 1976, Latz 8211 (DNA, MEL); Tanami Desert, 8 km

NE of Sangsters Bore, Sep 1978, Henshall 2276 (AD, DNA);

42 miles [c. 68 km] NW of Chilla Well, Jul 1970, Dunlop

(DNA). Central Southern Region: 8.4 km E of W. A. Border

on Kintore road, Apr 1988, Leach 1950 (CANB); Mt Liebig,

168 miles [c. 269 km] W of Alice Springs, Jul 1966, Willis

s.n. (DNA, MEL); Mt Liebig, north side, + 210 km WNW of

Alice Springs, Jun 1974, Carr 2368 & Beauglehole 46147

(MEL); + 6.4 km NNW of Mt Zeil, Jul 1968, Beauglehole

27181 (MEL).

Distribution and habitat : Corchorus walcottii

has a disjunct distribution. It occurs in the

Pilbara region, Western Australia, from Barrow

Island to near Marble Bar, and in central

Australia from the Tanami Desert in the

Northern Territory southwards to Mt Agnes

in the north-west of South Australia (Map

2). It is recorded as growing in hummock

grassland, shrubland and low open woodland

communities, on sandy or loamy soils

Fig. 16. Corchorus walcottii. A. branchlet with flower and fruit, x 1. B. fruit, x 02. C. ventral view of sepal, x 6. D. cross-

section of sepal, x 12. E. dendritic-stellate hair, x 24. F. simple glandular hair, x 48. G. simple glandular hair, x 24. A-F from

Telford 6547 (CANB); G from Burbidge 5811 (PERTH). Del. W. Smith.

sometimes associated with limestone, on

plains and hills. It is also recorded on coastal

dunes.

Phenology : Flowers have been collected in

March, April, June, July, September and

November, fruits in March, April, June and

November.

Typification : In the protologue of Corchorus

walcottii, Mueller (1862) cited two

collections “in collibus altioribus rupestribus

prope Nickol Bay et in Hearson island. P.

Walcott” The two collections referred to by

Mueller in his protologue were located

amongst material on loan to BRI from MEL

“elevated rocky hills, Nichol Bay

[MEL223677] and “growing on top of rocky

sandstone hill, Hearson island”

[MEL223678]. The specimen collected from

Hearson Island is selected as lectotype as it

is the better preserved specimen with flowers

and fruits attached.

Affinities: Corchorus walcottii, C.

parviflorus and C. laniflorus have

conspicuous simple glandular hairs present

amongst the dense stellate indumentum on

the stems, leaves and inflorescences.

Halford, Notes on Tiliaceae 4, Corchorus

625

For differences with C. parviflorus and C.

laniflorus refer to ‘Notes’ under those

species.

Notes: Corchorus walcottii as treated here

is a variable species. The central Australian

populations are somewhat different in having

a shorter indumentum on the leaves from the

typical form of this species from the Pilbara.

There is another form on the islands off the

Pilbara coast that has smaller leaves and

flowers than the typical form, but it

intergrades with the typical material of this

species.

Excluded names

Corchorus allenii F.Muell., Proc. Linn. Soc.

N.S.W. ser.2 6: 462 (1892). Type: near

Prince Regent River; Bradshaw & Allen

(holo: MEL) = Helicteres sp. (Sterculiaceae)

Corchorus longipes Tate, Transactions of the

Royal Society of S.A. 22: 119 (1898). type:

Mt Lyndhurst Run near Farina, S.A., 1898,

Max Koch s.n. (holo: AD) = Gilesia

biniflora F.Muell. (Sterculiaceae).

Corchorus pachyphyllus Burret, Notizbl.

Bot. Gart. Berlin, 12: 166 (1937). type

citation: [Western Australia.] Gascoyne,

nordlich bei Carnarvon, c. 25 miles ii. d. M.

Sandige Hiigel mit lichtem Gebusch, L. Diels

3718; n.v.

The type of this species has apparently

been destroyed in Berlin during the Second

World War and I have been unable to locate

any material that may be considered an

isotype. The description and discussion in

Burret’s protologue for this species is lengthy

but it has not been possible to say to which

known species it is referable.

Corchorus rothii F.Muell., Second

Systematic Census of Australian Plants. 30

(1889), nom. illeg. Type: based on

Triumfetta pilosa Roth.

Acknowledgements

I would like to thank Gordon Guymer,

Director of BRI, for making working space

and facilities at BRI available to me, the

directors and curators of AD, CANB, DNA,

K, MEL, NSW, P and PERTH for the loan of

their holdings for study at BRI. The

following persons provided assistance and

they are thanked sincerely for their efforts;

Laurie Jessup and Greg Leach for assistance

while they were Australian Botanical Liaison

Officer at K, Will Smith (BRI) for the

illustrations and maps and Les Pedley for

the translation of the diagnoses into Latin.

This work was supported by grants from the

Australian Biological Resource Study,

Environment Australia in 1991, 1992 and

1994, which are gratefully acknowledged.

References

Basak, S.L. (1958). Variations in the haploid chromosome

number of Corchorus sidoides F.Muell. (Family

Tiliaceae). Current Science 27(3): 101.

Baillon, H.E. (1866). Du genre Nettoa et des caracteres

qui separent les Bixacees des Tilacees.

Adansonia 6: 238-242.

Bentham, G. (1863). Tiliaceae. Flora Australiensis 1:

267-282. London: Reeve & Co.

Datta, R. M., Panda, B.S., Roy, K. Bose, M.M. & De,

T.K. (1966). Cytotaxonomic studies of different

Corchorus (Jute) species. 1. Botanical

Magazine (Tokyo) 79(939): 467-473.

Domin, K. (1928). Tiliaceae. In Beitrage zur Flora und

Pflanzengeographie Australiens. Bibliotheca

Botanica 89: 383-384.

Halford, D.A. (1995). Notes on Tiliaceae in Australia,

2. A revision of the simple-haired species of the

genus Corchorus L. Austrobaileya 4: 297-320.

Holmgren, P.K., Holmgren, N.H. & Barnett, L.C.

(1990). Index Herbariorum. Part 1. The

Herbaria of the World. 8 th edn. New York: New

York Botanic Gardens.

Islam, A.S. & Qaiyum, F. (1961). Chromosome numbers

in the genus Corchorus. Current Science

30(11): 433.

Mueller, F. (1862). Fragmenta Phytographiae

Australiae 3: 8-11. Melbourne: Victorian

Government.

Mueller, F. (1887). Description of a new Corchorus from

Central Australia. Transactions and

Proceedings of the Royal Society of South

Australia 9: 58-59.

626

Paczkowska, G. & Chapman, A.R. (2000). The Western

Australia flora: a descriptive catalogue. Perth:

Wildflower Society of Western Australia (Inc.), the

Western Australia Herbarium, CALM and the

Botanic Garden and Parks Authority.

Index to Scientific Names

Names in bold type are accepted names and

those in light are synonyms, etc. The numbers

refer to the number of the species accepted in

the above taxonomic treatment. ‘Excl.’ refers

to a name listed under Excluded names.

Corchorus allenii F.Muell. Excl.

Corchorus aulacocarpus Halford. 1

Corchorus carnarvonensis Halford. 2

Corchorus congener Halford. 3

Corchorus crassifolius Domin. 4

Corchorus crozophorifolius (Baill.) Burret. 4

Corchorus elachocarpus F.Muell. 5

Corchorus elderi F.Muell. 6

Corchorus incanus Halford. 7

Corchorus incanus Halford subsp. incanus . 7a

Corchorus incanus subsp. lithophilus Halford. 7b

Corchorus interstans Halford ms. 3

Corchorus laniflorus Rye . 8

Corchorus lasiocarpus Halford. 9

Corchorus lasiocarpus Halford subsp. lasiocarpus .. 9a

Corchorus lasiocarpus subsp. parvus Halford. 9b

Corchorus leptocarpus A.Cunn. ex Benth. 10

Corchorus lithophilus Halford ms . 7b

Corchorus longipes Tate. Excl.

Corchorus mitchellensis Halford. 11

Austrobaileya 6 (4): 581-629 (2004)

Corchorus obclavatus Halford. 12

Corchorus pachyphyllus Burret. Excl.

Corchorusparviflorus (Benth.) Domin. 13

Corchorus parviflorus (Benth.) Domin var. parviflorus 13

Corchorus parviflorus var. gracilescens Domin .... 13

Corchorus parviflorus var. ovatus Domin. 13

Corchorus puberulus Halford. 14

Corchorus pumilio R.Br. ex Benth. 15

Corchorus rostrisepalus Domin. 17c

Corchorus rothii F.Muell. Excl.

Corchorus saxicola Halford ms. 7b

Corchorus sericeus Ewart & O.B.Davies. 16

Corchorus sericeus Ewart & O.B.Davies subsp. sericeus

. 16a

Corchorus sericeus subsp. densiflorus (Benth.) Halford

. 16b

Corchorus sidoides F.Muell. 17

Corchorus sidoides F.Muell. subsp. sidoides . 17a

Corchorus sidoides subsp. rostrisepalus (Domin) Halford

. 17c

Corchorus sidoides subsp. vermicularis (F.Muell.) Halford

. 17b

Corchorus sp. Burrup (GCraig 235) . 22

Corchorus subargentus Halford. 18

Corchorus sublatus Halford. 19

Corchorus tectus Halford. 20

Corchorus tomentellus F.Muell. 21

Corchorus vermicularis F.Muell. 17b

Corchorus walcottii F.Muell. 22

Corchorus walcottii F.Muell. var. walcottii . 22

Corchorus walcottii var. densiflorus Benth. 16b

Corchorus walcottii var. parviflorus Benth. 13

Nettoa crozophorifolia Baill. 4

Scorpia simplicifolia Ewart & A.H.K.Petrie. 17b

Map I. Distribution of Corchorus spp. C. parviflorus * , C. crozophorifolius * , C. elderi A, ,

C. obclavatus ♦ .

Halford, Notes on Tiliaceae 4, Corchorus

627

Ma p 2, Distribution of Corchorus spp, C. walcottii • , C. /jwmi/io ♦ .

Map 3. Distribution of Corchorus spp, C. sidoides subsp. sidoides ★ , C. carnarvonensis • f

C. fomenfellusA , C. subargentus ♦ .

Map 4, Distribution of Corchorus spp, C, lanijlorus ★ , C, pufrera/us • ,C. mitchellensis ♦ ,

C. aulacocarpus A .

Map 5. Distribution of Corc/rc»r(/,s spp. C. sidoides subsp. vermicularis-k , C. sidoides subsp.

rostrisepalus • .

Map 6. Distribution of Corehorus spp. C elachocarpus ★ . C incanus subsp. liihaphilusA ,

C. leptocarpus • G sublatus ♦

Map 7. Distribution of Corehorus spp. C. incanus subsp. incanus • . C. congener it.

C. tectus A .

Halford, Notes on Tiliaceae 4, Corchorus

629

Map 8. Distribution of Corchorus spp. C. iasiocarpus subsp. iasiocarpus ★ , C. sericeus subsp.

sericeus • .

Map 9, Distribution of Corchorus spp. C. iasiocarpus subsp. parvus A , C. sericeus subsp.

densiflorus • .

Stictocardia Hallier f. (Convolvulaceae) in Queensland

R.W. Johnson

Summary

Johnson, R.W. (2004). Stictocardia Hallier f. (Convolvulaceae) in Queensland. Austrobaileya 6 (4): 631-

637. Two species of Stictocardia are recognised from Queensland. A new combination S. queenslandica

(Domin) R.W.Johnson, based on Argyreia queenslandica Domin, is made. A key to identify both species is

provided, together with descriptions, distribution maps and illustrations of certain diagnostic characters.

Keywords: Convolvulaceae, Stictocardia, Queensland, Australia

R.W. Johnson, Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens Mt

Coot-tha, Mt Coot-tha Road, Toowong, Queensland, 4066, Australia

Introduction

The genus Stictocardia was described by

Hallier (1893) to include three species, one of

which was Stictocardia tiliifolia (Desr.) Hallier

f. A few specimens of S. tiliifolia had been

collected from the Rockhampton and

Rockingham Bay regions of Queensland in the

late 1880’s when they were identified as

Ipomoea grandiflora Lam. (Mueller, 1889;

Bailey, 1901). It was not until Ooststroom

(1943) that the transfer of this taxon to

Stictocardia was recognised in Australia.

In 1928 Domin described Argyreia

queenslandica from a flowering specimen he

collected near Mungana in 1910. He ascribed

the species to the genus Argyreia. At that time

he recognised two other species of Argyreia as

occurring in Australia. A. soutteri (F.M.Bailey)

Domin, which he transferred from the genus

Lettsomia, is now known only from the type

collection and now presumed extinct, while

A. nervosa (Burm.f.) Bojer, an introduced

species, is widely naturalised in north

Queensland.

In recent years, fruiting material of

Argyreia queenslandica has been collected

from the type locality and surrounding areas.

The presence of minute black glands on the

underside of the leaves and the enlarged calyx

which completely encloses the fruit, clearly

requires the transfer of this species to the genus

Stictocardia.

Accepted for publication 15 March 2004

Classification

Stictocardia is regarded as a member of the

tribe Argyreieae (Hallier 1893; Austin &

Demis sew 1997). The only other members of

this tribe in Australia are the introduced

Argyreia nervosa and the enigmatic A. soutteri.

Taxonomy

Stictocardia Hallier f., in Engl. Bot. Jahrb. 18:

159 (1893); Ooststr., Blumea 5: 346, fig.

1, g-h (1943); FI. Males., ser. 1, 4(4):

491 (1953). Type: Stictocardia tiliifolia

(Desr.) Hallier f. (“tiliaefolia”), based on

Convolvulus tiliifolia Desr.

(“tiliaefolia”).

Annual to woody perennial with trailing and

twining stems. Cotyledons bilobed, with the

lobes ovate-oblong, diverging and greatly

exceeding the base. Leaves ovate or orbicular,

mainly cordate at the base, the lower surface

with many minute glands. Inflorescence

axillary, cymose, 1-many flowered; bracts

deciduous. Sepals 5, subequal, elliptic or

orbicular, obtuse to emarginate, subcoriaceous,

often with thinner margin, much accrescent in

fruit. Corolla red or purple, large, funnel-

shaped, the midpetaline bands often somewhat

pilose outside or at the distal end and with

minute glands like the leaves. Stamens and

style included. Filaments inserted near the

corolla base; pollen grains spheroidal and

polypantoporate with a spinulose surface.

Ovary glabrous, 4-celled, each cell with 1

ovule; style 1, stigma biglobular. Fruit much

632

Austrobaileya 6 (4): 631-637 (2004)

enclosed by the much-enlarged calyx, globular,

dissepiments with 2 transverse wings at the

surface of the fruit, dehiscing irregularly

between the wings exposing 4 pubescent seeds.

Distribution : Stictocardia is a pantropical

genus of 10 species (Austin & Eich 2001). The

species occur primarily in the tropics of Africa,

Asia and Australia with only S. tiliifolia

extending into tropical America as an

introduction (Austin & Demissew 1997).

Stictocardia tiliifolia has been recorded from

tropical Australia (Johnson 1983), though

Austin & Eich (2001) in their synopsis did not

cite any records from Australia. A second

species in the genus, described originally as

Argyreia queenslandica, is endemic to north¬

eastern Australia.

Etymology : The generic name is derived from

the Greek, stiktos for dotted, and kardia for

heart-shaped, referring to the broadly ovate,

cordate leaves dotted with glands on the lower

side, a feature characteristic of the genus.

Key to species of Stictocardia in Queensland

1. Outer sepals at flowering 1-1.5 cm long, to 2.5 cm at fruiting; petal

segments 6-7.5 cm long. 1. S. queenslandica

2. Outer sepals at flowering 1.5-2.2 cm long, to 4.5 cm at fruiting; petal

segments 8-10.5 cm long. 2. S. tiliifolia

1. Stictocardia queenslandica (Domin)

R.W.Johnson, comb. nov. Argyreia

queenslandica Domin, Biblioth. Bot. 89:

1087 (1928). Type: Queensland: Nord-

Queensland: am fusse eines Karsthugels

bei Mungana, Domin II. 1910 (holo: PR;

photo BRI).

Annual or perennial with trailing and twining

stems; cotyledons deeply bi-lobed (Fig. 1);

stems terete, herbaceous, sparsely hairy to ±

glabrous, with short, spreading to ± appressed,

tubercle-based hairs, 0.1-0.2 mm long. Leaves

petiolate; petiole 2-15 cm long, slightly

channelled above, hairy as for stem, 0.6-1.4

times as long as the blade; blade simple, broadly

ovate, occasionally ovate- to oblong-elliptic,

often shortly acuminate, 8-18 cm long, 6-15

cm wide with a L:W ratio of 1.1 to 1.4, apex

narrowly obtuse, mucronate, base cordate often

with an oblong sinus, 15-25% of the blade

length, discolorous, darker green above, the

underside covered with dark green to black

glandular pits, sparsely hairy to iglabrous,

occasionally moderately hairy on both sides,

hairs short, weakly erect to ascending, 0.1-0.2

mm long, midrib with 6-8 secondary veins per

side. Inflorescence axillary, cymose, usually

simple, occasionally compound, bearing 1-2,

rarely 3 flowers, occasionally with 2

inflorescences per axil, often with glandular pits

on peduncles, pedicels and sepals; peduncle

terete, stout, 0.4-2.0 cm long, extending to 3.0

cm at fruiting, glabrous to very sparsely hairy;

bracts opposite to sub-opposite, herbaceous,

oblong, 2.5-4 mm long, 1-1.8 mm wide, apex

rounded to barely acute, ciliate, face + glabrous,

with a raised midrib, abscissing when in bud

or early flowering; pedicels terete, thicker than

the peduncle, flattened and dilated and

becoming angular upwards, 0.7-1.2 cm long,

extending to 1.7 cm at fruiting, glabrous or with

an occasional hair. Sepals concave, thick,

ifleshy, with a narrow hyaline margin,

becoming leathery and enclosing the capsule

at fruiting; outer sepals broadly ovate to ovate-

orbicular, 1.0-1.5 cm long, 1.4-2.0 cm wide,

expanding to 2.5 cm x 3.5 cm at fruiting, apex

obtuse, base truncate to slightly cordate,

glabrous except for some short marginal and

ciliate hairs; inner sepals orbicular to ovate or

oblong-orbicular, 1.2-1.6 cm long, 1.0-1.6 cm

wide, expanding to 2.5 cm x 2.7 cm at fruiting

(Fig. 3A), apex obtuse to rounded, emarginate,

base + rounded to truncate, glabrous. Corolla

funnel-shaped, limb purple to reddish-purple

with darker purple inner throat and midpetaline

bands, 5.5-6.5 cm long, tube to base of flare

2.0-3.5 cm long, limb 4.5-6 cm diameter, tube

at base of flare 0.9-1.3 cm diameter, glabrous

except for a few hairs on the midpetaline band

towards the limb and a fringe of slender,

somewhat sinuate, ciliate hairs, 0.2-0.75 mm

long, lining the rim each side of the midpetaline

Johnson, Stictocardia in Queensland

633

Fig. 1 . Cotyledons of Stictocardia queenslandica from glasshouse specimen grown from Clarkson 3789.

band; petal segments 6.0-7.5 cm long, 2.5-3.7

cm wide at the limb, distally rounded-

triangular, sometimes emarginate. Stamens 5,

fused for 9-12 mm from the base of the corolla

tube, fused area not raised; filaments terete

above, flattened and dilated downwards,

unequal in length, 1.3-2.1 cm long, densely

hairy from just below the point of insertion for

2-5 mm along the filament, hairs reddish-

purple, cylindrical, sinuate with small clear

club-shaped to cylindrical terminal cells, up to

1.25 mm long; anthers lanceolate, sagittate,

4.0-4.8 mm long, 1.2-1.6 mm wide, apex

rounded to truncate, basal lobes rounded,

0.6-0.9 mm long, splitting lengthwise at

maturity; pollen globular, spinulose. Ovary

ovoid, constricted in the upper part,

quadrangular in cross-section, glabrous, 4-

celled, with 1 ovule per cell, disk prominent,

donut-shaped, 5-lobed, 0.4-0.7 mm high,

yellowish; style 2.5-3.0 cm long, unbranched,

glabrous, bearing a capitate to almost 2-lobed

stigma, 2.3-2.4 mm x 1.75-2.0 mm in cross-

section, with laterally compressed conical

tubercles bearing short hair-like protuberances.

Capsule ovoid to depressed globular-ovoid,

rounded-quadrangular in cross section, with

thin papery walls, 1.0-1.5 cm high, 1.0-1.7

cm diameter, glabrous, 4-celled, septa persistent

at maturity with distal, transverse wings,

irregularly dehiscing longitudinally between

the wings. Seeds 4, ^-globular-ovoid, olive-

brown to black, 7.5-9 mm long, 6 -8 mm wide,

moderately to densely pubescent, with short

erect to ascending golden brown hairs, 0.1-0.2

mm long somewhat matted, with longer hairs

to 1 mm densely clustered on the inner margin

of the hilum.

Specimens examined : Queensland. Cook District: 16km

E of Rookwood Creek, c. 16km W of Chillagoe, Mar 1980,

Clarkson 3025 (BRI); c. 1km towards Chillagoe from

Mungana railway siding, Jun 1981, Clarkson 3789 (BRI);

Royal Arch Tower, c. 5km SW of Chillagoe, Mar 1987,

Clarkson 6837 & McDonald (BRI, MBA, QRS, K); The

Archways, c. 15km NW of Chillagoe, May 1987, Clarkson

6860 & McDonald (BRI, MBA); 2 km SE of Rookwood

Homestead, c. 6km NW of Mungana, Jun 1983, Conn & De

Campo 1347 (MEL, BRI, MBA); NPR 98, Royal Archway

Cave, Mungana, Apr 2000, Ford 02382 (QRS, BRI);

Wrotham Park - Mungana road, 8km NW of Mungana, Apr

1980, Johnson 4043 (BRI).

Distribution and habitat : It is endemic to

Australia and is restricted to a small area in

the Chillagoe-Mungana area (Map 1). It

appears to be confined to limestone outcrops

and grows in deciduous vine thickets.

634

Austrobaileya 6 (4): 631-637 (2004)

Phenology: Flowers have been recorded from

March to June with fruiting occurring soon after

flowering. Fruit have been recorded from April

to November.

Etymology: The specific epithet refers to

Queensland, the state of Australia in which it

was discovered.

Notes: It resembles Stictocardia discolor

Ooststr., based on the description given by

Ooststroom (1943), which is known only from

one collection from Timor. Both S. queenslandica

and S. discolor have shorter sepals and corollas

than S. tiliifolia. However S. discolor appears

to have shorter sepals and longer and more slender

peduncles and pedicels than S. queenslandica. In

addition, S. discolor is characterised by a red

coloration on the lower leaf surface.

Conservation status: This species is listed as

Rare under Queensland Government

legislation.

2. Stictocardia tiliifolia (Desr.) Hallier f., Bot.

Jahrb. 18: 159 (1893) (“tiliaefolia”);

Ooststr. Blumea 5: 346. (1943); Ooststr.

FI. Males, ser. 1, 4: 491. (1953); Rivea

tiliaefolia (Desr.) Choisy, Mem. Soc.

Phys. Geneve 6: 407 (1834); Argyreia

tiliaefolia (Desr. ) Wight, leones PI. Ind.

Or. 4(2): 121. 1358. (1848); Convolvulus

tiliaefolius Desr., in Lam. Encycl. Meth.,

Bot 3: 544 (1792). Type: Mauritius,

Commerson (P, not seen).

Ipomoea grandiflora Lam., Tabl. Encycl. I:

467 (1791).

Woody perennial liana with twining stems;

stems terete, herbaceous, moderately hairy with

short, weakly ascending, tubercle-based hairs,

0.1-0.4 mm long, becoming woody, reddish-

purple in colour, distinctly lenticular and

glabrous at the base. Leaves petiolate; petiole

2.5-10 cm long, slightly channelled in the

upper half, hairy as for the stem, 0.3-0.9 times

as long as the blade; blade simple, broadly ovate

to ovate-oblong to almost orbicular, often

shortly and abruptly acuminate, (6-) 8-18 cm

long, 6-14 cm wide with a L:W ratio of

0.9-1.4, apex acute, occasionally rounded to

emarginate, mucronate, base cordate with a

broad rounded sinus, 15-27% of the blade

length, discolorous, darker green above, the

underside covered with dark green to black

glandular pits, moderately to sparsely hairy on

both sides, hairs short, erect to ascending,

0.1-0.4 mm long, midrib with 6-9 secondary

veins per side. Inflorescence axillary, cymose,

usually simple, occasionally compound, bearing

1, rarely 2 or 3 flowers, often with glandular

pits on peduncles, pedicels and sepals; peduncle

terete, stout, 1.5-6.5 cm long, moderately hairy

with short sinuate, erect to ascending matted

hairs, 0.1-0.4 mm long,; bracts opposite to

subopposite, herbaceous, oblong, concave,

4.5- 8 mm long, 2.8-4 mm wide, apex rounded,

emarginate, mucronulate, sparsely to

moderately hairy, occasionally ± glabrous,

abscissing when in bud or early flowering;

pedicels terete, stout, slightly dilated and

angular upwards, 1.0-2.0 cm long, moderately

hairy, glabrescent. Sepals concave, thick,

ifleshy, with a narrow hyaline margin,

becoming leathery with distinct longitudinal

and reticulate venation and enclosing the

capsule at fruiting; outer sepals broadly ovate

to orbicular or + reniform, 1.5-2.0 (-2.2) cm

long, 2.0-2.5 cm wide, expanding to 4.5 cm x

5 cm at fruiting (Fig. 3B.), apex rounded, ±

emarginate, base rounded to shallowly cordate,

moderately to sparsely hairy, glabrescent; inner

sepals broadly elliptic to orbicular or obovate-

orbicular, slightly smaller than the outer,

1.3-1.8 (-2.0) cm long and wide at flowering

and expanding at fruiting, apex rounded,

emarginate, base + rounded to truncate,

glabrous. Corolla funnel-shaped, limb purple

with darker violet-purple inner throat and

midpetaline bands, (6.5-) 7.0-8.5 cm long, tube

to base of flare 3-4 cm long, limb 6-9 cm

diameter, tube at base of flare 1.5-2 cm

diameter, ± glabrous or with scattered hairs on

the midpetaline band towards the tip and a

fringe of slender, somewhat sinuate, ciliate

hairs 0.2-0.75 mm long lining the rim each

side of the midpetaline band; petal segments

7.5- 10.5 cm long, 3.5-5 cm wide at the limb,

distally rounded, emarginate. Stamens 5, fused

for 10-13 mm from the base of the corolla tube,

fused area not raised; filaments terete above,

slightly flattened and dilated downwards,

unequal in length, (2.7-) 3.5-4.5 cm long,

densely hairy from about the point of insertion

for 4-8 mm along the filament, hairs reddish-

purple, dense, cylindrical, sinuate with short,

Johnson, Stictocardia in Queensland

635

clear, globular to club-shaped terminal cells,

to 1.2 mm long; anthers lanceolate, sagittate,

5.0-6.5 mm long, 1.7-2.5 mm wide, apex

rounded, bluntly apiculate, basal lobes rounded,

1.0-1.4 mm long, splitting lengthwise at

maturity; pollen globular, spinulose (Fig. 2.).

Ovary ovoid, constricted in the upper part,

rounded quadrangular in cross-section,

glabrous, 4-celled with 1 ovule per cell, disk

prominent, donut-shaped, 5-lobed, 0.6-1.0 mm

high, golden-yellow; style 4.0-5.7 cm long,

unbranched, glabrous, bearing a bi-globular

stigma, 2.5-3.0 mm x 1.5-1.8 mm in cross-

section, covered in distinct tubercles bearing

short hair-like protuberances. Capsule ovoid,

rounded-quadrangular in cross section, 1.5-2

cm high, 1.6-2 cm diameter, glabrous, 4 celled,

septa persistent at maturity with distal,

transverse wings, irregularly dehiscing

longitudinally between the wings. Seeds 4,

^-globular-ovoid, 8-11 mm long and 6-8 mm

wide, brown, densely pubescent with short erect

to ascending brown hairs, 0.1-0.2 mm long,

somewhat matted with longer hairs to 1 mm

densely clustered around the hilum.

Selected specimens examined : Northern Territory.

Darwin and Gulf Region: Goromuru River, c. 7 km SE of

mouth, 27 Apr 1996, Cowie 6659 & Bokarra (DNA, BRI);

Arafura Swamp area near Raminging, Sep 1998, Cowie &

Mangion 7963 (DNA, BRI); Arnhem Land, Bennet Bay, Nov

1987 Dunlop 7350 (DNA, BRI); 9 km SW of West Alhgator

Head, Kakadu, Jun 1988, Russell-Smith 5612 & Lucas.

(DNA, BRI); 11 km E Channel Point, Nov 1988, Russell-

Smith 6406 & (BRI, DNA). Queensland. Cook District:

Low Isles, May 1963, Cribb BRIU2839 (BRI). North

Kennedy District: Mount Bertha, Gloucester Island, 21 Mar

1994, Batianoff 9403318 & Dillewaard (BRI); nr Port

Denison, 1887, Birch s.n. (MEL 95501); Cromarty, Mar

1935, Blake 8318 (BRI); Rockingham Bay, s.d., Dallachy,

(MEL95476); Ayr, s.d., Michael 1526 (BRI); Harvey Range,

Oct 1929, Pollock s.n. (BRI); Long Island, Jul 1935, White

12157 (BRI). Port Curtis District: Howard Point, Middle

Percy Island, Oct 1989, Batianoff 11706, Champion,

Thompson & Dillewaard (BRI); Rockhampton, s.d., Dietrich

(MEL 95502). Moreton District: Burleigh Heads, Mt

Burleigh National Park, 1972, Catherine (BRI).

Distribution and habitat : It is widespread

throughout the old world tropics from central

and southern Africa through south-east Asia to

Australia and islands in the west Pacific Ocean

and is naturalised in tropical America. It has

been recorded along the coast of northern and

north-eastern Australia from east of Darwin to

Burleigh Heads, SE of Brisbane (Map 1). It

grows mainly in coastal vine forests;

occasionally in coastal eucalypt open forests.

Phenology: Flowers have been recorded from

March to July and fruit from July to November.

Etymology: The specific epithet refers to the

shape of the leaves which resemble those of

the Linden or lime tree ( Tilia spp.)

Conservation status: This species is currently

not considered to be rare or endangered.

References

Austin, D.L. & Demissew, S. (1997). Unique fruits and generic

status of Stictocardia (Convolvulaceae). Kew

Bulletin 52: 161-169.

Austin, D.L. & Eich, E. (2001). Synopsis of Stictocardia with

another Madagascan species, S. mojangensis

(Convolvulaceae). Willdenowia 31:79-85.

Bailey, F. M. (1901). The Queensland Flora. Part IV.

Brisbane: H. J. Diddams & Co.

Domin, K. (1928). Argyreia. Beitrage zur Flora und

Pflanzengeographie Australiens. Bibliotheca

Botanica 89(6): 1087.

Hallier, H. (1893). Convolvulaceae Africanae. Botanische

Jahrbuecherfuer Systematik 18: 81-160.

Johnson, R.W. (1983). Convolvulaceae, in R.D Morley &

H.R. Toelken (eds), Flowering Plants of Australia,

pp 259-261. Adelaide: Rigby Publishers.

Mueller, F. (1889). Second Systematic Census of

Australian Plants. Part 1, Vasculares.

Melbourne: Government Printer.

Mabberley, D.J. (1997). The Plant-Book. Second

Edition. Cambridge: Cambridge University

Press.

Oostroom, S.J. van (1943). The Convolvulaceae of

Malaysia IV, XIX. Stictocardia Hall. f.

Blumea 5: 346-352.

-(1953). Convolvulaceae. In FI. Males., Ser. 1, 4:

388-512. Djakarta: Noordhoof-Kolff n.v.

636

Austrobaileya 6 (4): 631-637 (2004)

Map 1. Distribution of □ Stictocardia queenslandica and • S. tilt (folia.

Fig. 2. Pollen grain of Stictocardia tiliifolia from Dunlop 2782 (AD).

Johnson, Stictocardia in Queensland

637

Fig. 3. Fruiting calyx of A. Stictocardia queenslandica - (glasshouse specimen ( Calway BRI-AQ378863), grown from seed

of Clarkson 3789 x 1.5) and B. S. tiliifolia - C Pollock s.n. (BRI-AQ 277029) x 1.5)

The taxonomy and ecology of Solanum subg. Leptostemonum (Dunal)

Bitter (Solanaceae) in Queensland and far north-eastern New South Wales,

Australia

A.R. Bean

Summary

Bean, A.R. (2004), The taxonomy and ecology of Solanum subg. Leptostemonum (Dunal) Bitter (Solanaceae)

in Queensland and far north-eastern New South Wales, Australia. Austrobaileya 6(4): 639-816. 90 species

(82 indigenous and 8 naturalised) of Solanum subg. Leptostemonum are recorded for Queensland and far

north-eastern New South Wales (north of 29°S latitude), with 50 species endemic. 29 species and one

subspecies are described as new and illustrated (S. ammophilum, S. argopetalum, S. cocosoides,

S. crebrispinum, S. ditrichum, S. dryanderense, S. dumicola, S. dysprosium, S. eminens, S. fervens,

S. francisii, S. galbinum, S. graniticum, S. hapalum, S. innoxium, S. intonsum, S. johnsonianum,

S. jucunclum, S. latens, S. longissimum, S. lythrocarpum, S. mentiens, S. parvifolium subsp. tropicum,

S. pusillum, S. rixosum, S. senticosum, S. stenopterum, S. ultimum, S. versicolor and S. vicinum ); six

species are reinstated ( S. angustum Domin, S. erassitomentosum Domin, S. defensum F.Muell.,

S. magnifolium F.Muell., S. mitchellianum Domin and S. shirleyanum Domin) and three are reduced to

synonymy ( S. cleistogamum Symon, S. dallachii Benth. and S. seitheae Symon). S. dianthophorum Dunal

is accepted, but known only from the holotype. S. centrale and S. yirrkalense are newly recorded as native

to Queensland, and S. incanum is newly recorded as naturalised. Lectotypes are chosen for S. defensum and

S. dunalianum, and a replacement lectotype chosen for S. sturtianum. All Australian species are classified

into (mostly pre-existing) informal taxonomic groups. Distribution maps and notes on distribution are given

for all taxa. Other biogeographical data (based on degrees of latitude and longitude) are provided, both at

species and group level. Indumentum characters (particularly those of the stellate hairs) have been used

extensively to elucidate the taxonomy of the group. Ecological requirements of various Solanum species

and the reasons for their infrequent or transient occurrence are discussed. The use of Solanum fruits as food

is reviewed. Identification keys (dichotomous and multi-access) are provided for all Queensland species.

The conservation status of all indigenous taxa is assessed.

Keywords: Solanum, Solanum subg. Leptostemonum, new species, taxonomy, ecology, biogeography,

conservation status, Queensland, New South Wales, Australia, keys, DELTA, identification, indumentum,

stellate hairs.

A.R. Bean, Queensland Herbarium, Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha road, Toowong,

Queensland 4066, Australia

Introduction

Solanum is the fourth largest Angiosperm

genus in the Queensland flora. An

interrogation of the Queensland Herbarium

database in July 2003 returned 73 accepted and

formally named indigenous Solanum species.

Only Acacia (305 spp.), Eucalyptus (188 spp.),

and Cyperus (112 spp.), had more species.

Five subgenera of Solanum (using the

classification of D’Arcy (1972)) are represented

in Queensland, either as natives or

naturalisations:

S. subg. Archaesolanum - the ‘Kangaroo

Apples’, quick growing shrubs without

stellate hairs or prickles, leaves often

deeply lobed {S. aviculare, S. vescum)\

Accepted for publication 2 February 2004

S. subg. Brevantherum - comprising the ‘Wild

Tobaccos’, large shrubs with stellate hairs

and corymbose inflorescences

(S. mauritianum, S. erianthum,

S. abutiloides );

S. subg. Leptostemonum - the subject of this

paper, and comprising the majority of

taxa;

S. subg. Potatoe - the ‘Potato Creepers’, vines

with deeply lobed leaves ( S. laxum,

S. seaforthianum, S. triflorum)\

S. subg. Solanum - comprising the ‘Blackberry

Nightshades’ or ‘Black Nightshades’,

small shrubs with umbel-like

inflorescences (S. americanum,

S. chenopodioides, S. nigrum, S. opacum,

S. physalifolium, S. villosum), and two

species from S. sect. Geminata (Knapp

640

2002) - S. spirale, a tall rainforest shrub

with globose or ellipsoidal orange fruits,

and the small shrub S. pseudocapsicum

(Jerusalem Cherry).

Solanum subg. Leptostemonum consists

of about 500 species distributed throughout the

warmer parts of the world, but with centres of

diversity in the South American tropics and

subtropics, central east Africa, New Guinea and

Australia. Distinguishing morphological

characters for the subgenus include the

attenuate anthers, stellate indumentum, and the

presence of prickles (Whalen 1984). They may

be briefly described as the “prickly” Solanum

species. The few species that lack prickles

invariably possess stellate indumentum, and the

few species that lack stellate indumentum

invariably possess prickles.

Significant contributions to the taxonomy

and knowledge of Queensland members of the

subgenus have been made by Brown (1810),

Mueller (1861), Bentham (1868), Domin

(1929), Symon(1971,1981,1995), Ross (1986)

and Bean (2001, 2002b).

A reappraisal of Queensland species is

necessary for several reasons. Some previously

synonymised species (particularly those

described by Karel Domin) deserve to be re¬

instated; additional taxa have been discovered

or recognised; extra information on

distribution, morphology or ecology has become

available for many species; and some

nomenclatural changes are necessary. The

inclusion of a small portion of New South Wales

(north of 29° S) in this study means that the

biogeographic data are more meaningful.

The ensuing discussion relates entirely to

the members of S. subg. Leptostemonum

occurring in Queensland and far northeastern

New South Wales.

Morphology and terminology

Habit : Habit varies from completely prostrate

shrubs to small trees attaining 9 metres in

height ( S. viridifolium ). Most are shrubs 0.5-

1.5 metres high. Habit is discussed further in

the Ecology section below.

Juvenile plants: These are plants that have not

yet reached reproductive age, or are bearing

Austrobaileya 6 (4): 639-816 (2004)

their first flowers. Their leaves are called

juvenile leaves. The juvenile leaves are

generally larger, pricklier and more deeply

lobed than those occurring on adult (flowering

and fruiting) plants. The stems of juvenile

plants of any given species are usually more

densely prickly than on adult plants. Juvenile

growth may also be found on vigorous shoots

arising from the lower parts of sexually mature

plants, or the shoots arising from a cut stem.

For species exceeding 1 metre in height at

maturity, juvenile characters have been

assessed, where possible, on plants 20-30 cm

high. For smaller species, particularly the

herbaceous resprouters, the juvenile stage is

either absent or of very short duration, and

hence is only rarely described.

Sympodia: Each inflorescence in Solanum is

developmentally terminal, but is quickly

exceeded by subsequent vegetative growth

(Child 1979). Each lateral vegetative shoot is

termed a sympodial unit. The sympodia are

usually difoliate i.e. there are two leaves

between successive inflorescences. These leaves

may arise from the same position on the stem

(geminate) or not (disjunct).

Bark: In most species, the bark is unremarkable

and rather non-descript. It is smooth, green to

dark brown, and often bears raised lenticels.

However, in two species ( S . furfuraceum and

S. sporadotrichum ), the bark is conspicuously

furrowed and corky, especially on the large

stems. In these species, corky outgrowths are

usually visible even on (the thicker stems of)

herbarium specimens. Corky stems (less

obvious than the above) are also found on

S. versicolor, S. francisii and

S. erassitomentosum.

Large stems: Those more than 30 cm from the

growing point, and including the trunks of

small trees.

Branchlets: Stems less than 30 cm from the

growing point {i.e. those usually present on

herbarium specimens). In most species, they

appear to the naked eye smooth and terete, but

in a few species (notably S. quadriloculatum ),

they are conspicuously ridged.

Adult leaves: Leaves that are found adjacent

to, or distal from, the inflorescences. Many

Bean, Taxonomy of Solanum subg. Leptostemonum

641

species have entire adult leaves, but in others,

the adult leaves are variously lobed. A “lobing

index” is used in this paper, which is the length

of the lobe halfway along the lamina divided

by the parallel length of the adjacent sinus (f/e

in Fig. 1). The index is 1 for an entire leaf;

leaves with an index >2 are considered to be

deeply lobed. While the leaf size of a species

can vary considerably depending on prevailing

environmental conditions, the lobing index is

relatively constant. In a few species, leaves can

be pseudo-pinnate (derived from deeply lobed

leaves where the proximal lobes are divided

right to the midrib).

At least some of the adult leaves of all

Queensland Solanum species are bilaterally

asymmetrical, and this is manifested most

conspicuously at the lamina base. In this paper,

such a leaf base is described as oblique.

Obliqueness is given both in absolute terms (the

length “b” on Fig. 1) and as a percentage of

lamina length (100 x b/(a+b) on Fig. 1). Petiole

length is measured in absolute terms (the length

“c” on Fig. 1) and as a percentage of lamina

length (100 x c/(a+b) on Fig. 1). Both are useful

for discriminating some taxa. In a few species,

the petioles are winged, the wings consisting

of a narrow strip of green tissue on either side

of the petiole throughout its length.

Inflorescence: Solanum inflorescences are

ebracteate scorpioid cymes, but often appearing

racemose, with a single axis bearing several

pedicellate flowers. A few species ( e.g.

S. chrysotrichum ) have a much-branched

inflorescence, resembling a panicle. In some

species, particularly those allied to

S. densevestitum, the inflorescence comprises

a single flower, or sometimes 2-3 flowers all

arising from the branchlet without any common

peduncle (pseudo-umbellate). The ‘racemose’

inflorescence may also lack a common

peduncle, with the basal flower emerging very

close to the branchlet, and subsequent flowers

borne on a rachis.

Sex expression : Many species are

andromonoecious, that is, their inflorescences

comprise some bisexual flowers (towards the

base) and some male flowers, where the style

is much reduced in length and is non-functional

(Whalen 1984). A style can be inferred to be

functional when it protrudes beyond the top of

the anthers. Another large group of species bear

only bisexual flowers. One Queensland species

(S. carduiforme ) is dioecious (Symon 1981).

Flowers : Solanum flowers are largely

pentamerous, but tetramery (where there are 4

calyx lobes, four corolla lobes and four stamens)

is found in several groups or species. For most

species, only occasional tetramerous flowers

occur, but in some species (e.g. S. viridifolium)

approximately half of the flowers are

tetramerous, while in a few species (e.g.

S. ammophilum ) all flowers appear to be

tetramerous.

The calyx comprises a fused basal section

or “tube”, and free lobes. The calyx tube is

somewhat invariate in shape, but the shape of

the calyx lobes is often somewhat diagnostic.

The calyx invariably grows between anthesis

and fruiting, but in many species this growth

is minimal and the lobes fail to extend to half

the length of the mature fruit (not accrescent).

In other species, either the calyx lobes grow

until they extend beyond the fruit, or the calyx

tube grows to completely envelop the mature

fruit (accrescent).

Prickliness of the calyx is correlated with

both sex expression and accrescence. In some

Queensland species (mostly naturalised species

such as S. linnaeanum, but also S. stupefactum ),

the calyx of the basal bisexual flowers is much

more prickly than that of the distal male

flowers. In species with a strongly accrescent

calyx, the calyx is always prickly.

The pedicels generally lengthen between

anthesis and fruit maturation, and hence

measurements are provided for both stages.

Corolla shape, size, and sometimes

colour, are useful in identifying Solanum

species. The shape is dependent on the amount

of “interpetalar” tissue present (the tissue

connecting the individual lobes of the corolla).

Species with little or no interpetalar tissue have

deeply lobed corollas; species with abundant

interpetalar tissue have rotate or pentagonal

corollas. In most Queensland species, the

corolla is mauve to purple, but in a few it is

white, while one naturalised species

(S. rostratum ) has a yellow corolla. All

642

Austrobaileya 6 (4): 639-816 (2004)

Fig. 1. Diagrammatic representation of leaves showing the measurements used.

Queensland species (except S. pugiunculiferum )

have stellate hairs on the outer surface of the

corolla, but relatively few have hairs on the

inner surface, and hence this character is useful.

Each stamen comprises a short filament

and a relatively long, attenuate, yellow anther.

The anthers are very similar throughout the

subgenus, although anther length can be

diagnostic. The style is most often erect,

emerging between the stamens, but some

species have a sigmoid or eccentric style that

is strongly bent just above the ovary, so that it

does not touch the anthers at all. The stigma is

for most species unremarkable, being entire or

obscurely lobed. However, in a few species

belonging to the S. dioicum group, the stigma

is conspicuously forked, with each fork up to 5

mm long. The ovary is a rather uniform

structure, but the indumentum pattern on the

surface is very useful.

Fruits and seeds: Mature fruits are very

valuable for species identification and

classification (Nee 1986; Symon 1987). Each

species has a characteristic fruit size, shape,

colour, exocarp thickness and placentation

pattern, and related species usually have si mil ar

fruit characteristics. Unfortunately, dried fruits

preserved on herbarium specimens are totally

unsuitable for determining most of the

characters mentioned, even colour. Collectors

rarely mention fruit colour, and when they do,

there is no guarantee that the fruits they

observed were mature. Hence fresh fruits or

fruits preserved in alcohol and good collector’s

notes are essential for the accurate assessment

of fruiting characters. Such collections are very

infrequent, and much is still to be learned.

Fruit maturity is indicated in the field

either by a distinct colour change (usually from

green to red), or by their becoming relatively

soft and easily detached from the plant.

About 21 of the Queensland species

(belonging to Groups 5, 9A & 13) have bright

red succulent fruits with a thin exocarp, e.g.

S. stelligerum. The three species of Group 13A

similarly have succulent fruits with a thin

exocarp, but are black in colour. All these

species have inflorescences in which usually

all flowers are bisexual. Many other species (of

Groups 22-28) have fruits with a thicker

exocarp that remain pale green (often with some

darker green variegation) at maturity, or

sometimes becoming greenish-white or pale

yellow. This fruit type is correlated with a

weakly or strongly andromonoecious

inflorescence. A small number of species have

fruits which do not fit either of these types e.g.

S. vicinum has a large deep-purple fruit about

the size of a plum; S. stupe)actum has a bright-

orange hairy fruit.

Fruit shape is globular for most species,

but distinctly oblate for S. echinatum, and

distinctly ellipsoidal for S. innoxium and

S. gympiense.

Placentation is a character that varies

widely between species (Nee 1986). The

“standard” placentation pattern for Solanaceae

is a 2-locular fruit with axile placentas, but

many Queensland taxa have 1-locular fruits,

and a few have 3- or 4-locular fruits. The degree

of development of the placenta also varies

greatly.

Seed morphology is rather similar

throughout, and all species have seeds that are

Bean, Taxonomy of Solanum subg. Leptostemonum

643

Fig. 2. Pedicel of S. torvurn , showing mixture of gland-tipped finger hairs and stellate hairs (Wilson GWW 123)

lenticular and orbicular to somewhat reniform

in outline. Differences in colour are useful in

discriminating some species. Most species have

pale (white to pale yellow) seeds, but a few

species have quite dark seeds (brown to black).

Trichomes : Trichome morphology is extremely

useful in Solanum taxonomy (Roe 1971; Seithe

1979; Whalen et al. 1981; Seithe & Sullivan

1990), but has been only superficially utilised

by taxonomists dealing with Australian species.

Comprehensive trichome descriptions are

provided in this paper, as trichomes have

proven to be consistent in form and size for

each plant part for each species, and hence

exceedingly useful for distinguishing taxa.

There is often a bewildering array of

trichome constructions present on a single

species. For example, the pattern observed on

the branchlets is usually different from that

observed on the upper leaf surface, and that

pattern is different from that of the lower leaf

surface etc. Hence, for the comparisons to be

useful, indumentum must be described

separately for each plant part. In this paper,

five plant parts have been chosen for which

indumentum is described in detail: branchlets,

leaf upper surface, leaf lower surface, pedicel/

calyx, and ovary/style.

Some trichome terminology follows Roe

(1971). The trichome types used here are

adapted from Seithe (1979), who recognised

several kinds of trichomes, belonging to two

distinct classes. Type 1 hairs include Finger

hairs and all forms of Stellate hairs, while Type

2 hairs are very short gland-tipped trichomes,

relatively uniform in appearance (see below).

1. Finger hairs (Seithe 1979; Seithe &

Anderson 1982) - uniseriate unbranched hairs

consisting of more than one cell, and usually

0.5-5 mm long. This hair type is found on

several Queensland species (Fig. 2, 3).

Seithe (1979) has shown that they are

ontologically related to stellate hairs, and that

the latter are derived from finger hairs. In a

few Queensland species, the developmental

process can be observed directly e.g. finger hairs

with tiny protuberances near the base; or stellate

hairs with very short lateral rays and very long

644

Austrobaileya 6 (4): 639-816 (2004)

Fig. 3. Branchlet of S. ditrichum, showing abundant finger hairs (Forster 11564 & Leiper)

central ray. In other species, finger hairs may

be present as ‘protostellae’ (see below).

However in most cases, the finger and stellate

hair types are readily distinguishable. The

finger hairs may have a unicellular glandular

tip.

2. Stellate hairs (Roe 1971; Seithe 1979) - made

up of a multiseriate stalk (sometimes obscure

or absent) bearing varying numbers of

unicellular, acicular lateral rays and (usually)

with an erect uniseriate central ray or midpoint

(Whalen 1984) (Fig. 2, 4-10).

Stellate hairs are a feature of nearly all

species in the subgenus (and the subgenus

comprises the majority of Solarium species in

Australia). The morphological characteristics

of stellate hairs (size, form, colour, density,

number of lateral rays, and relative length of

central ray) are all very useful for taxonomic

purposes, in that the indumentum pattern for

each plant part within a species has proven to

be remarkably consistent.

(a) Developmental forms of stellate hairs

In many species, all hairs growing

adjacent to each other are more or less

the same size and shape. In other species,

it appears that the development of the

stellate indumentum of the leaves (and

probably other organs) “switches off’

before all the hairs have reached optimum

development, resulting in stellae of

differing sizes and developments being

found in close proximity. In this paper,

stellate hairs that have not reached their

optimum development are termed

“protostellae”. These protostellae are

smaller in size than ordinary stellae, and

have fewer lateral rays, and they are

interspersed with the ordinary stellae (Fig

5,10). Protostellae are usually mentioned

only for the upper leaf surface, though

they may be present on other parts of the

plant.

(b) Diameter

The average width of the stellate hair

projected at right angles to the subtending

surface.

The absolute distance between

stellae is measured from centre to centre.

Bean, Taxonomy of Solanum subg. Leptostemonum

645

Fig. 4. Calyx of S. longissimum, showing developing prickles with complete stellate hair at the apex (Bean 5617 & Forster)

Leaf indumentum density is measured on

fully expanded adult leaves. In the species

descriptions, the following terminology

is used for stellate hair density:

very dense - ordinary stellae overlapping,

multi-layered, the subtending surface not

visible with hand lens (Fig. 8);

dense - ordinary stellae overlapping, 0.1-

0.5 diameter apart (centre to centre), 1 or

2 layers only, the subtending surface

visible with hand lens;

moderate - ordinary stellae just

overlapping, 0.5-1 diameter apart (centre

to centre) (Fig. 7);

sparse - ordinary stellae 1-2 diameters

apart (centre to centre) (Fig. 5);

very sparse - ordinary stellae > 2 diameters

apart (centre to centre) (Fig. 10).

Hence for “very dense”, “dense” and

“moderate”, the stellae are overlapping;

for “sparse” and “very sparse”, they do

not touch each other.

Stellate hair density is certainly

quite variable for a species. Observations

in the field and herbarium show that

relatively large leaves that have developed

under mesic conditions tend to have more

widely spaced hairs than on small leaves

developed under xeric conditions. It is

hypothesised that the number of stellate

hairs per leaf is fixed for each genotype,

and that density is determined by

environmental conditions.

(d) Orientation of lateral rays

If the lateral rays of the stellate hair are

all in one plane, like the spokes of a

wheel, they are described as “porrect”

(Roe 1971) (Fig. 4,6-10). Sometimes the

lateral rays are all placed at 30-60 degrees

from the central ray. In this case, the

lateral rays (usually 8 or fewer) are

described here as “ascending” (Fig. 5). If

the lateral rays (usually 8-16) are placed

at various angles with respect to the

central ray, the hair is described as

“multiradiate” (Roe 1971).

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Austrobaileya 6 (4): 639-816 (2004)

Fig. 5. Stellate hair pattern on the upper leaf surface of S.furfuraceum, showing ascending lateral rays (Bean 15928)

(e) Central ray

The ratio of the central ray length

compared to the average lateral ray length

is reported. In some species, the central

ray of the stellate hair possesses a

unicellular glandular tip (Seithe 1979)

(Fig. 3).

(f) Stalk length

Often, stellate hairs are sessile, i.e. there

is no stalk below the lateral rays. In all

other cases, the stalk length is reported.

Summary of stellate hair patterns observed

in Queensland species

a) Relative diameter: In general, stellae from

the upper leaf surface have a smaller

diameter than elsewhere on the plant.

Those from the lower leaf surface,

branchlets, calyx and style are usually

about the same size.

b) Absolute diameter : The broadest stellae are

possessed by S. densevestitum (up to 1.8

mm across). Other species with very

broad stellae are S. crebrispinum,

S. quadriloculatum, S. stenopterum,

S. chippendalei and S. stupefactum.

Species with the smallest stellae include

S. corifolium, S. mentiens, S. francisii,

S. macoorai, S. elachophyllum and

S. sturtianum, whose stellae are frequently

less than 0.2 mm across.

c) Lateral rays: Many species have 8, 7 or 8,

or 6-8 lateral rays on each stellate hair.

Some species have many more lateral

rays. S. elaeagnifolium branchlet stellae

have 15-18 lateral rays. Other species

with 13 or more lateral rays are

S. ammophilum, S. cinereum,

S. dianthophorum, S. dryanderense,

S. elachophyllum, S. oligacanthum,

S. sturtianum (Fig. 8) and S. yirrkalense.

Species with very few lateral rays include

S. cookii, S. coracinum, S. densevestitum,

S. dumicola, S. francisii, S. latens,

S. pusillum and S. rostratum.

Bean, Taxonomy of Solanum subg. Leptostemonum

647

Fig. 6. Upper leaf surface of S. pusillum, showing stellate hairs and Type 2 hairs (Bean 2604)

d) Central ray: Five species have branchlet

stellae lacking a central ray: S. corifolium,

S. dimorphispinum, S. lacunarium,

S. mentiens and S. sturtianum.

In most species, the central ray is

about equal in length to the lateral rays

or a little longer.

Several species have branchlet stellae

where the central ray is very short (< 0.5 times

the length of the lateral rays); S. amblymerum,

S. ammophilum, S. chrysotrichum,

S. defensum, S. elachophyllum,

S. elaeagnifolium, S. inaequilaterum,

S. limitare, S. macoorai, S. torvum.

At the other extreme are species

where the central ray is 5-10 times the

length of the laterals, giving the

indumentum a shaggy or velvety

appearance. These include S. cookii,

S. densevestitum, S. fervens, S. hapalum,

S. innoxium, S. johnsonianum,

S. lasiocarpum, S. magnifolium,

S. serpens, S. stelligerum.

The variation in the central ray ratio

shows no consistent pattern within a

species. In other words, the ratio on the

branchlets may differ somewhat from the

ratio of the lower leaf or the calyx, but it

is not consistently longer or shorter on

any of these plant parts. While the ratio

is usually about the same within a species,

a few species have widely different ratios

on different plant parts e.g. S. torvum 0.1-

0.3 vs. 0.8-2; S. latens 0.5-1 vs. 1.5-3;

S. lasiocarpum 0.8-1.5 vs. 4-10;

S. gympiense 1-2 vs. 2-4; S. ditrichum

0.7-1.5 vs. 2-3.

Type 2 hairs are very short trichomes (<0.1

mm long) occurring in Solanaceae, normally

with one stalk cell, and a transparent

multicellular glandular head (Seithe & Sullivan

1990). They have also been called ‘capitate-

glandular trichomes’ (Whalen et al. 1981),

‘multicellular glands’ (Seithe 1979; Seithe &

Anderson 1982) or ‘stipitate glands’ (Merrill

& Perry 1949; Bean 2001). They are present,

though often inconspicuous, on many species,

particularly on the ovary and vegetative

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Austrobaileya 6 (4): 639-816 (2004)

Fig. 7. Stellate hair pattern on the upper leaf surface of S. galbinum (Thomas 1817 &Thompson)

growing points (Fig. 6). Seithe (1979) has

shown that they are not developmentally related

to Type 1 hairs.

A variant of the Type 2 hair has been

observed on the ovary of S. ammophilum. These

hairs are about 0.1 mm long, have a relatively

short stalk and a long ellipsoidal gland,

distinctly opaque and white in colour. The

glandular portion obviously secretes a great deal

of mucilage, as the fruits and the inner surface

of the calyx in S. ammophilum are quite sticky.

Apart from this variant, Type 2 hairs seem quite

uniform in morphology, though this may be

merely a function of the difficulty in observing

such a small structure.

Prickles in Solanum are rigid, shiny, sharply

pointed structures. They are a feature of almost

all species of the subgenus, and may be found

on the large stems, branchlets, petioles,

laminae, peduncles, pedicels, calyx, and

(rarely) the corolla. Their size, shape,

distribution and relative abundance is often

diagnostic for a species. They are straight,

acicular and spreading in the vast majority of

Australian native species, but in many

naturalised species, they are broad-based and

often recurved. Acicular prickles are defined

(for the purposes of this paper) as those that

are more than 7 times longer than wide.

The fact that the prickles are derived from

the lignified stalks of stellate hairs becomes

obvious in a few species ( e.g. S. longissimum ),

where uniseriate lateral rays are mounted near

the apex of the prickles (Fig. 4).

Branchlet prickle density (number of

prickles per decimetre (dm)) has been assessed

on adult branchlets, 1-2 decimetres from the

growing point.

Sometimes prickles bear indumentum i.e.

scattered complete stellate hairs or Type 2 hairs

laterally attached to the lower half of the

prickle, and this can be diagnostic.

In most species, prickles can be found on

the leaf lamina. Usually there are more prickles

present on the upper surface, or roughly equal

numbers on both surfaces. The few exceptional

species (where prickles are more numerous on

the lower surface) are S. capsicoides,

S. chrysotrichum, S. papaverifolium,

S. semiarmatum and S. torvum.

Bean, Taxonomy of Solanum subg. Leptostemonum

649

Fig. 8. Lower leaf surface of S. sturtianum ; stellae sessile, without a central ray, and with 13-16 lateral rays (Anderson 5070)

Materials and methods

Herbarium specimens have been borrowed from

AD, BH, BM, CANB, DNA, K, MEL, NSW,

PERTH, PR, and QRS.

Most measurements have been made from

dried herbarium material, but material

preserved in alcohol has been used for

measurements on the flowers and particularly

the fruits. Where preserved material was not

available, flowers were rehydrated in boiling

water, but dried fruits could not be

reconstituted, and hence measurements of the

exocarp and placenta were not attempted. Leaf

dimensions have been derived from the larger

leaves present on each specimen. Branchlet

indumentum has been assessed from non-

abraded stems 10-20 cm from the growing

point of flowering or fruiting herbarium

specimens.

Whenever possible, the author has

examined species in the field, for the purpose

of learning about the habitat, ecology, flower

and fruit colour, flower sex ratio, and to collect

spirit material for later study.

Comprehensive morphological

descriptions of all native and currently

naturalised species are provided, derived from

a DELTA (DEscriptive Language for

TAxonomy) (Dallwitz, Paine & Zurcher 1993)

dataset of 91 taxa and 153 characters. Data were

entered using the DELTA Editor (Dallwitz,

Paine & Zurcher 1999), and interactive keys

have been produced using Intkey (Dallwitz,

Paine & Zurcher 1995; Dallwitz, Paine &

Zurcher 2000).

Specimen citations have been arranged

according to the Queensland Pastoral districts,

and then chronologically under each district.

Species doubtfully or formerly naturalised in

Queensland are treated briefly at the end of this

paper.

Ecology of Solanum subg. Leptostemonum

There is very little literature relating to the

ecology of Australian species of Solanum subg.

Leptostemonum. Floyd (1966) studied an area

of eucalypt forest near Coffs Harbour in New

South Wales. He recorded the species which

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Fig. 9. Petiole of S. echinatum, showing a long-stalked stellate hair (Alcock 11285)

germinated in a glasshouse from soil samples

taken at an unburnt site, and compared the

species with those recorded in the field (at the

soil sampling site) two months after a fire. He

found that four Solanum spp. were common at

the burnt field site, but that “[seeds] were

virtually absent from the soil before the fire”.

He attributed this difference to bird dispersal

of seeds following the fire. However, the

methodology does not indicate that the soil from

the unburnt site was closely examined to

discover what types of seeds were present, so

the author may have only presumed that the

soil held no Solanum seeds. Bird dispersal of

so many seeds in such a short period, onto a

bare and burnt site seems very unlikely. A more

likely scenario is that the seeds were already

in the soil (gradually dispersed over the years),

and that fire broke the dormancy of the seeds.

In the absence of other published data, I

offer the following observations and comments

on the Queensland members in the hope that

they will stimulate formal study. These

observations stem from a desire to understand

why many species are so reclusive and transitory.

1. Life Form

Various life-forms have been observed for

Queensland species:

(a) herbaceous resprouters - where the above¬

ground portion dies each year, but the root

or rhizome is persistent e.g.

S. adenophorum, S. angustum, S. limitare,

S. multiglochidiatum, S. papaverifolium,

S. stenopterum, S. versicolor. These

species are leafy for only 3-6 months a

year. This term is synonymous with the

often-used ‘herbaceous perennial’. It

would seem that no Queensland species

is truly annual in habit.

(b) perennials - where the above-ground part

of the plant lives for several years,

normally not perishing after the first

flowering and fruiting. Perennials can be

subdivided into two sub-types:

(i) stoloniferous perennials (only three

species; S. acanthodapis, S. serpens and

S. mentiens ), ground-hugging trailing

shrubs that root frequently at the nodes.

Bean, Taxonomy of Solanum subg. Leptostemonum

651

Fig. 10. Upper leaf surface of 5. linnaeanum, showing stellae at various developmental stages (Forster 28894)

(ii) rhizomatous perennials (very

common type), which form clusters of

seemingly separate individuals. These

colonies are clonal, being connected by

an extensive system of horizontal roots

just below the soil surface, and may

extend in size from one to hundreds of

square metres. Species in this category include

S. corifolium, S. densevestitum, S. ellipticum,

S. ferocissimum, S. stelligerum, and

S. stupefactum. In some larger and longer-

lived species occurring in mesic environments

e.g. S.francisii, S. inaequilaterum, S. nobile,

S. semiarmatum, and S. viridifolium,

regeneration may be largely from seed.

2. Major habitats

(a) evergreen notophyll rainforest (with no

eucalypt forest nearby). These species

exploit canopy gaps, where there is

enough light to allow flower and fruit

development. Fire is absent.

S. acanthodapis, S. cookii, S. corifolium,

S. dimorphispinum, S. dryanderense,

S. eminens, S. francisii, S. hamulosum,

S. inaequilaterum, S. lasiocarpum,

S. macoorai, S. mentiens, S. nobile,

S. rixosum, S. semiarmatum, S. serpens,

S. sporadotrichum, S. vicinum, S. viridifolium,

(b) margins of evergreen notophyll rainforest,

or eucalypt forest with rainforest

understorey (often described as “wet

sclerophyll”) on basaltic or other fine¬

grained soils. Fire occurs infrequently.

S. cookii, S. densevestitum,

S. dimorphispinum, S. ditrichum, S. eminens,

S. hapalum, S. intonsum, S. latens,

S. limitare, S. macoorai, S. magnifolium,

S. nobile, S. parvifolium subsp. tropicum,

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Austrobaileya 6 (4): 639-816 (2004)

S. rixosLim, S. semiarmatum, S. shirleyanum,

S. sporadotrichum, S. stelligerum,

S. stupefactum, S. tetrathecum, S. vicinum,

S. viridifolium.

(.Acacia harpophylla ) or Belah

(Casuarina cristata ) dominated

communities on cracking clays. Fire is

normally absent.

S. adenophorum, S. coracinum, S. dissectum,

S. dumicola, S. elachophyllum, S. ellipticum,

S. esuriale, S.jurfuraceum, S. johnsonianum,

S. latens, S. mitchellianum,

S. multiglochidiatum, S. parvifolium

subsp. parvifolium, S. sporadotrichum,

S. stenopterum, S. tetrathecum.

(d) semi-deciduous vine forest on quaternary

sand deposits, laterite or granite hills

(Cape York Peninsula). Fire is normally

absent.

S. defensum, S. discolor, S. dunalianum,

S. dysprosium, S. fervens, S. yirrkalense.

(e) shrubby eucalypt woodland, remote from

rainforest or vine thicket. Fire occurs

frequently.

S. amblymerum, S. angustum, S. carduiforme,

S. chippendalei, S. cinereum, S. coracinum,

S. crassitomentosum, S. crebrispinum,

S. echinatum, S. ellipticum, S. ferocissimum,

S. jurfuraceum, S. galbinum, S. graniticum,

S. gympiense, S. intonsum, S. jucundum,

S. limitare, S. longissimum, S. mitchellianum,

S. multi glochidiatum, S. nemophilum,

S. parvifolium subsp. parvifolium,

S. quadriloculatum, S. senticosum,

S. stelligerum, S. tetrathecum, S. ultimum.

(f) lancewood communities (dominated by

Acacia catenulata, A. shirleyi,

A. burrowii, A. sparsiflora or A. blakei )

on lateritised Cainozoic duricrusts

associated with low mesas or scarps. Soil

may be red earth, brown loam, or skeletal

on plateau margins. Fire occurs

infrequently.

S. argopetalum, S. cocosoides, S. ellipticum,

S. galbinum, S. innoxium, S. jucundum,

S. latens, S. lythrocarpum, S. pusillum,

S. ultimum.

(g) red earths in mulga ( Acacia aneura )

dominated communities.

S. centrale, S. ellipticum, S. ferocissimum,

S. quadriloculatum, S. sturtianum,

S. versicolor.

(h) grassland or open woodland on heavy

cracking clays.

S. esuriale, S. lacunarium, S. papaverifolium,

S. stenopterum.

(i) shrublands in arid zone.

S. ammophilum, S. chenopodinum,

S. oligacanthum, S. sturtianum.

(j) saline littoral zone.

S. pugiunculiferum.

(k) degraded land (where frequent and/or gross

disturbance has allowed exotic species to

partially or completely replace the native

flora).

S. capsicoides, S. chrysotrichum,

S. elaeagnifolium, S. incanum, S. linnaeanum,

S. rostratum, S. sisymbriifolium, S. torvum.

Species from the arid and semi-arid zones

frequently occur underneath trees, which may

reflect the preferential deposition of seeds by

birds, or some requirement for protection from

the sun during the middle of the day, or relief

from grass competition.

No species occurs in heathlands,

mangroves or swamps.

3. Likelihood

The likelihood of encountering species of

Solanum subg. Leptostemonum increases as you

move from:

- low-nutrient soil to high-nutrient soil

- woodland to closed forest/rainforest

- low altitude to high altitude

- poorly drained sites to well drained sites

- frosty areas to frost-free areas

Bean, Taxonomy of Solanum subg. Leptostemonum

In other words, there are few (or no)

species in woodland with sandy soil at low

altitude, and (potentially) numerous species on

basaltic soil in rainforest at high altitude.

4. Response to disturbance

Most (perhaps all?) species can be classified as

pioneer species. They are promoted by

disturbance events, which may be grouped into

three main types:

a) Fire : No species has a lignotuber, hence

above ground parts are killed by moderate

intensity fires, but on the other hand, most

species are rhizomatous. Fire encourages seed

germination, promotes regeneration from

rhizomes and removes competition for light and

nutrients. Absence of fire for a long period may

see the decline or local extinction of Solanum

spp.

b) Cyclone : For those species that grow

in dense rainforest environments, they must

rely on severe storms and cyclones (in the

absence of land-clearing activities) to open the

tree canopy and allow sufficient light for them

to grow and reproduce, and thereby maintain

the soil seedbank.

c) Mechanical : Mechanical disturbance

of the soil can have the same effect as fire i.e.

promoting the seed germination and/or

regeneration from rhizomes. This type of

disturbance is usually man-induced e.g.

roadworks, quarries, logging etc.

In suitable recently disturbed sites,

solanums may occur in large numbers. The

numbers then gradually dwindle until, after a

few years, none is left. Hence, Solanum spp.

(perennials as well as herbaceous resprouters)

often cannot be relocated at (uncleared) sites

where they were recorded only a few years

before.

Native Solanum species do not fare very

well in company with other shrubs, forbs or

graminoids. They are easily out-competed by a

dense sward of grass or by clumps of shrubs.

They are not favoured by frequent disturbance

e.g. soil tilling, as there is insufficient time for

seed production or re-establishment of rhizome

networks.

In summary, indigenous species in

Solanum subg. Leptostemonum are favoured by

653

infrequent major disturbance (cyclonic,

mechanical or fire), provided that such

disturbance does not stimulate the proliferation

of exotic weeds.

Conservation Status

In general, native Solanum species appear to

be in decline. Some species from rainforest-

margins eg. S. ditrichum , S. rixosum, can no

longer be found in many districts where they

were recorded in the early 1900’s. Rainforest

margins are the preferred habitat of the

pernicious weed Lantana camara. It seems clear

that this exotic shrub has caused serious decline

in the populations of many Solanum species

treated in this paper.

The Brigalow communities of central

Queensland have been so reduced by land

clearance, that some species, e.g. S. dissectum,

S. adenophorum, and S. johnsonianum, are now

endangered or critically endangered.

Remaining populations are threatened by

aggressive weeds such as Panicum maximum,

Parthenium hysterophorus, and deliberately

introduced pasture grasses, especially Cenchrus

ciliaris.

All species have been assessed against the

IUCN Red List categories and criteria (IUCN.

2001). With regard to ‘mature individuals’, I

have followed the IUCN recommendation that

“reproducing units within a clone should be

counted as individuals, except where such units

are unable to survive alone (e.g. corals)”, and

that where population size fluctuates, “use a

lower estimate”.

Both of these recommendations have

implications for the Solanum assessments. In

Solanum subg. Leptostemonum , clones are the

rule rather than the exception, and fluctuations

in population size occur in most (if not all)

species. However, I have not interpreted these

fluctuations as “extreme”, as this would have

invoked Criteria Blc(iv) and/or B2c(iv) and

C2b. This would result in many ‘stable’ taxa

(having no apparent threat) being recognised

as threatened. Species assessments have been

done on an Australia-wide basis. Overseas

occurrences have not been considered.

For some species, a herbarium collection

during the last 10-15 years has been accepted

as a current location, even though experience

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Austrobaileya 6 (4): 639-816 (2004)

has shown that solanums can only rarely be

found at old collection sites. Sometimes the

species may still be present (as seed or

rhizomes) and sometimes it has disappeared

altogether.

Of the 82 indigenous taxa treated in this

paper, 2 have been assessed as “Critically

Endangered”, 8 as “Endangered” and 14 as

“Vulnerable”.

Solanum as food

Solarium fruits were an important source of food

for Australian Aborigines, particularly in the

arid areas. Their importance in the arid zone

was due to a number of factors - they are readily

available and collectable and visibly obvious;

they can be stored for later use, and most species

can be eaten without preparation (Peterson

1979).

Several species were used consistently by

aborigines in arid Australia. Solanum centrale

may be eaten when ripe or ground into a paste

and moulded into balls which keep indefinitely.

S. chippendalei is edible, but only after removal

of the bitter seeds and placenta. They may then

be eaten fresh, or dried-out and stored on

skewers. Dried fruits can be reconstituted in

water before consumption. S. ellipticum is

reported to be an important species which can

be eaten fresh. S. esuriale can also be eaten

fresh, or stored as above (Peterson 1979, Latz

1995).

Other arid zone species are strongly

poisonous, e.g. S. quadriloculatum (Everist

1974, Latz 1995), and were avoided by all

aboriginal groups. Other species given by

Everist ( loc. cit.) as definitely poisonous, at

least to livestock, are S. elaeagnifolium,

S. linnaeanum and S. sturtianum. Several other

species were suspected to be poisonous.

Solanum appears to have been of much

less importance as a food in coastal areas,

because of their lesser frequency and the many

other food sources available in these areas.

However, S. stelligerum fruits were reported by

Lampert & Saunders (1973) to be edible.

S. hapalum fruits likewise are edible

(R. Fensham pers. comm.).

Cytology

Randell & Symon (1976) undertook extensive

chromosome counts of Australian Solanum spp.

They found that n = 12 or 24 for all members

of S. subg. Leptostemonum. No other numbers

have been reported.

Hybrids

Naturally occurring hybrids appear to be rare

in Solanum subg. Leptostemonum , at least in

Australia. Symon (1981) made no mention of

hybridisation. I have personally seen only one

instance of a presumed interspecific hybrid in

the field. In an area where Solanum latens and

S. nemophilum were both common, there was

a small colony of plants displaying traits

intermediate between these two species, viz. leaf

size, indumentum density, stem prickliness, and

stellate hair size (voucher material at BRI; AQ

772057).

Classification

With about 500 species world-wide, Solanum

subg. Leptostemonum has a diverse

membership, and classifying them has and will

present difficulties. The conspectus by Whalen

(1984) has provided an excellent framework

for the whole subgenus, and a basis for future

work. He presented 33 informal taxonomic

groups for the subgenus, arranged in systematic

order. By his own admission, his knowledge of

the Old World taxa was limited, and so it is

perhaps not surprising that I have made many

amendments to his classification for taxa

occurring in Queensland. This has entailed the

shifting of many species to a more appropriate

group, and the creation of new groups where

Whalen broadly circumscribed them (especially

the S. ellipticum group). The native Australian

species are here classified into 15 informal

groups, compared to the 8 groups recognised

by Whalen (loc. cit.), but the average Australian

group size is only now comparable to the

average group size for New World taxa in

Whalen (loc. cit.).

I have retained Whalen’s group numbers

except for Group 6 (S. macoorai group), where

the “type” species was obviously misplaced.

New groups have been slotted in with the

addition of letters e.g. 13A. These are not

intended to represent “subgroups”, but rather

are equal in rank to Whalen’s established

groups.

655

Bean, Taxonomy of Solanum subg. Leptostemonum

Informal taxonomic groups for Australian Solanum subg. Leptostemonum , adapted from

Whalen (1984).

Notes: All accepted Australian species are listed. Queensland and north-eastern N.S.W. taxa are

listed in roman script; non-Queensland taxa are given in italics. Naturalised species are denoted

by an asterisk.

Group 5

Group 9A

Group 13

Group 13 A

Group 14

Group 22

Group 23

Group 25

Group 25A

Group 27

Group 27A

Group 27B

Group 27C

Group 27D

Group 27E

Group 28

Group 29

Group 33

Ungrouped

S. dunalianum group; S. dunalianum, S. tetrandrum, S. viridifolium.

S. densevestitum group; S. densevestitum, S. gympiense, S. hapalum, S. innoxium, S.

johnsonianum, S. nemophilum, S. ultimum.

S. ferocissimum group; S. chenopodinum, S. corifolium, S. defensum, S. discolor, S.

dissectum, S. dryanderense, S. dysprosium, S. ferocissimum, S. fervens, S.

inaequilaterum, S. latens, S. lythrocarpum, S. mentiens, S. parvifolium, S. shirleyanum,

S. stelligerum, S. yirrkalense.

S. semiarmatum group; S. coracinum, S. mitchellianum, S. semiarmatum.

S. torvum group; S. chrysotrichum*, S. torvum*.

S. quitoense group; S. lasiocarpum.

S. mammosum group; S. capsicoides*.

S. hystrix group; S. adenophomm, S. campanulatum, S. cookii, S. ditrichum, S.

eardleyae, S. eremophilum, S. graniticum, S. hoplopetalum, S. hystrix, S. lacunarium, S.

multiglochidiatum, S. oligandrum, S. papaverifolium, S. petrophilum, S. prinophyllum,

S. pungetium, S. pusillum, S. stenoptemm, S. vicinum.

S. pugiunculiferum group; S. pugiunculiferum

S. ellipticum group; S. angustum, S. argopetalum, S. crebrispinum, S. crassitomentosum,

S. dianthophorum, S. ellipticum, S. horridum, S. quadriloculatum, S. senticosum, S.

terraneum.

S. hamulosum group; S. dimorphispinum, S. eminens, S. hamulosum.

S. macoorai group; S. acanthodapis, S. amblymemm, S. armourense, S. brownii, S.

celatum, S. centrale, S. cinereum, S. cocosoides, S. curvicuspe, S. dumicola, S. francisii,

S. furfuraceum, S. galbinum, S. intonsum, S. jucundum, S. limitare, S. macoorai, S.

magnifolium, S. neoanglicum, S. nobile, S. rixosum, S. serpens, S. silvestre, S.

sporadotrichum, S. tetrathecum.

S. esuriale group; ammophilum, S. coactiliferum, S. elachophyllum, S. elaeagnifolium*,

S. esuriale, S. hesperium, S. karsense, S. nummularium, S. oldfieldii, S. oligacanthum, S.

orbiculatum, S. plicatile, S. sturtianum, S. versicolor.

S. lasiophyllum group; S. ashbyae, S. gabrielae, S. gilesii, S. lachnophyllum, S.

lasiophyllum.

S. echinatum group; S. echinatum, S. longissimum, S. lucani.

S. dioicum group; S. asymmetriphyllum, S. beaugleholei, S. carduiforme, S.

cataphractum, S. chippendalei, S. clarkiae, S. cunninghamii, S. dioicum, S.

diversiflorum, S. ebumeum, S. heteropodium, S. leopoldense, S. melanospermum, S.

oedipus, S. petraeum, S. phlomoides, S. tudununggae, S. vansittartense.

S. incanum group; S. incanum*, S. linnaeanum*, S. marginatum *, S. stupefactum.

S. rostratum group; S. rostratum*.

S. sisymbriifolium*

Biogeography

With 82 indigenous species, 50 of these

endemic, Queensland and north-eastern New

South Wales is the centre of diversity for

Solanum subg. Leptostemonum in Australia,

and has perhaps a higher concentration of

species than anywhere else in the Old World.

Fig. 11 shows the number of native

species occurring in each lxl degree square.

The highest diversity of 16 spp. is recorded for

a square straddling the Queensland-New South

Wales border, extending from Warwick in the

west to Kyogle in the east, and from

Cunningham’s Gap almost to Tenterfield. This

square contains large areas of rainforest, and

656

Austrobaileya 6 (4): 639-816 (2004)

o o'

0 0

h! o/i o o

10 0

0 1 4

1 2 0 0 0 2 2 8

6 3 1 1 0 5 3 4 4

20 °4 3 4 3 9

-■ - * 1 * 3 2 0 3 ,3

1 0 1 1 0 1 2 3 8 ^

0 2 0 o 0 0 4 2 4 6 0 fi 8 5 X

1 2 2 .3 2.2 0 0 2 2 ® s 8 l 3 * * * * \jP

1 2 2 3 i 6 -g-ffi/.?. 8 4 J 1

2.2 s i 1 4 3 Ifimsh

11 3 4

“"r—/*\ /

140

145

150

Fig. 11 . Number of indigenous species of Solanum subg. Leptostemonum recorded for each 1° x 1° square in Queensland and

far north-eastern New South Wales north of 29° S latitude, based on herbarium records.

includes the highest mountain in southern

Queensland (Mt Superbus, 1381 metres). The

adjoining square to the east boasts 10 native

species, even though half of the square is ocean.

This square includes Lamington N.P., Mt

Warning, Byron Bay and Lismore. Along a

5 km stretch of road near the Lamington N.P.,

I have recorded 8 native species (S. corifolium,

S. ditrichum, S. inaequilaterum, S. limitare,

S. rixosum, S. semiarmatum, S. serpens,

S. stelligerum ), which must surely be the

richest area for Solanum subg. Leptostemonum

in Australia.

The next most diverse square (13 spp.)

surrounds the town of Biloela. It includes the

Kroombit Tops area, Coominglah S.F., the

Callide Range and, to the west and north of

Biloela, some small remnants of Brigalow

forest. All are important havens for Solanum.

The three next most diverse squares (with

12, 12, and 11 spp.) are all in the far southeast

of Queensland, and all contain areas of high

altitude rainforest, of which the most notable

is the Bunya Mountains. There are 6 spp. at

the Bunya Mountains.

Away from the southeast quarter of the

state, the highest diversity (9 spp.) is found in

the square that includes Nebo, Mirani, Dipperu

N.P. and Eungella. The Eungella N.P. is an area

of high altitude rainforest containing an

endemic species (S. francisii ), while Dipperu

N.P. and adjacent uncleared areas are

dominated by Brigalow forest. The critically

endangered S. adenophorum has been recorded

from Dipperu N.P.

Further north, the square that includes

the Atherton Tableland has 9 spp., and the

Bean, Taxonomy of Solanum subg. Leptostemonum

657

. 2 /2,°

t 2 r

j 0 £

j o 'o ' ixo

? 0 0 3 b

"G- T 0

/1 0 0 ,

1 0 0 3 5)1

0 1 3 3 4 ( 2

^ J

3 0 1

3 11

4 5 2

0 0 0225X0

110 4 2 4 3 5> 1

2OO4233 V

’ 21 ° * 1 3 5 k

10 1 2 3 3 0 ^.1

12 2

2 1 3

10 1

0 2 0

1 1 2

xm® 4 4 6 #26/

2 2 1 1 3 2 3 ft f 7 4 /*

j 0

145 150

Fig. 12. Number of informal groups of Solanum subg. Leptostemonum present in each 1° x 1° square in Queensland and far

north-eastern New South Wales north of 29° S latitude, based on herbarium records (includes only those groups that contain

indigenous species).

square immediately to its north has 8 spp. Both

of these squares have large areas of rainforest,

much of it at high altitudes.

In general, as one moves further from the

coast, the species diversity decreases, strongly

correlated with the decreasing rainfall, but even

in the far southwest of Queensland, some

species occur, e.g. S. oligacanthum,

S. sturtianum. In the arid parts of the state, the

highest diversity (6 spp.) is achieved in the

square surrounding Mt Isa.

Two of Queensland’s biogeographic

regions (Thackway & Cres swell 1995) have a

very depauperate Solanum flora. The “Gulf

Plains”, a bioregion characterised by extensive

marine plains, alluvial clay plains and areas of

sandy outwash, apparently has no species at

all, except for S. pugiunculiferum near the

coast. The “Cape York Peninsula”

biogeographic region has solanums only in

rainforest areas; the extensive Eucalyptus and

Melaleuca woodlands are apparently Solanum-

free zones.

Fig. 12 shows the distribution of informal

groups (see Classification section, above) across

Queensland and northeastern New South

Wales. The highest diversity per square is 7

groups, known from four squares (Duaringa

area, Eungella-Nebo area, Kingaroy-Bunya

Mtns area, Dalby-Toowoomba-Milmerran

area). This level of group diversity exceeds all

other parts of the Old World, based on Whalen

(1984), but is still less diverse than some areas

in South America.

658 Austrobaileya 6 (4): 639-816 (2004)

Table 1. The most widespread taxa in Queensland, based on the number of l°x 1° squares

in which they are found.

Species

Number of 1° x 1° squares

S. ellipticum

S. esuriale

S. ferocissimum

S. parvifolium subsp. parvifolium

S. stelligerum

S. mitchellianum

S. jucundum

S. viridifolium

S. nemophilum

S. galbinum

S. furfuraceum

S. corifolium

S. chippendalei

S. quadriloculatum

Taxonomy

Key to informal taxonomic groups of Solanum subg. Leptostemonum (as listed above)

1. Mature fruits with juicy mesocarp, exocarp thin, bright red or black; fruits

relatively small (mostly <12 mm diameter). 2

Mature fruits with mesocarp dry to moist but not juicy, exocarp thick,

often green to yellow, occasionally orange or purple, rarely red; fruits

larger (mostly >12 mm diameter). 5

2. Some or all inflorescences branched. 3

All inflorescences unbranched or pseudo-umbellate. 4

3. Mature fruits black; branchlets very prickly; leaves with stellate hairs;

flowers 5-merous. 13A. S. semiarmatum group

Mature fruits red; branchlets without prickles; fully expanded leaves

glabrous; flowers 4 or 5-merous. 5. S. dunalianum group

4. Plants completely without prickles; calyx lobes elliptic, exceeding mature

fruits; inflorescences pseudo-umbellate, without common peduncle

. 9A. S. densevestitum group

Plants at least sparsely prickly, calyx lobes various, but not elliptic and not

exceeding mature fruits; common peduncle present. 13. S. ferocissimum group

5. Inflorescences with dimorphic flowers (1 or more very prickly bisexual

flowers at base with several less-prickly smaller male flowers beyond,

or plants dioecious). 6

Inflorescences with flowers all of similar size and prickliness, although

functionally male flowers may be frequent

Bean, Taxonomy of Solanum subg. Leptostemonum

659

6. Inflorescence with few distal male flowers; fruiting calyx not markedly

accrescent, not enclosing fruit; style with indumentum. 29. S. incanum group

Inflorescence with numerous male flowers on long rachis; fruiting calyx

accrescent, tube often enclosing fruit; style glabrous. 28. S. dioicum group

7. Calyx prickles absent, or with 1-5 per flower. 8

Calyx moderately to densely prickly (10-200 per flower). 12

8. Stellate hairs absent from stems, leaves and calyx; corolla radius 4-5 mm;

anthers 1.5-2 mm long. 25A. S. pugiunculiferum group

Stellate hairs present; corolla radius 7-20 mm; anthers 3-9 mm long. 9

9. Prickles on branchlets 1-4 times longer than broad, recurved. 10

Prickles on branchlets 5-20 times longer than broad, straight. 11

10. Inflorescence branched; plants shrubby. 14. S. torvum group

Inflorescence unbranched; plants vine-like, scrambling. 27A. S. hamulosum group

11. Stellae with 8 or fewer lateral rays; leaves mostly green on the upper surface

. 27B. S. macoorai group

Many stellae with more than 8 rays; leaves usually with dense indumentum

on both sides. 27C. S. esuriale group

12. Leaves with finger hairs only. 23. S. mammosum group

Leaves with stellate hairs, finger hairs sometimes also present. 13

13. Fruits completely enclosed by accrescent calyx. 14

Fruits readily visible, calyx not accrescent . 15

14. All anthers similar; leaves entire or with shallow lobes. 27E. S. echinatum group

One anther much longer than the other four; leaves deeply lobed

. 33. S. rostratum group

15. Leaves + entire, upper surface moderately to very densely hairy; perennial

shrubs; finger hairs rarely present; petioles not winged. 27. S. ellipticum group

Leaves shallowly to deeply lobed, upper surface glabrous or sparsely hairy;

herbaceous resprouters; finger hairs frequently present; petioles

frequently winged. 25. S. hystrix group

Dichotomous key to the Queensland species of Solanum subg. Leptostemonum

Notes: i.) This key is designed for the

identification of dried herbarium specimens.

Although macroscopic characters have been

used wherever possible, microscopic

examination of the stellate hairs will be

necessary in some instances. Neither open

flowers nor mature fruits are required to

successfully operate this key, but frequently

some fertile material with intact calyces will

be needed.

ii) ‘leaves’ are adult leaves unless otherwise

stated; similarly ‘branchlets’ are adult branchlets

iii. ) leaf lobing index = length of lobe halfway

along lamina divided by parallel length of

adjacent sinus (Fig. 1). Used to quantify degree

of lobing; entire leaves have an index of 1,

deeply lobed leaves have an index >2.

iv. ) the fruit diameters and colours given apply

to fresh mature fruits

v. ) finger hairs - uniseriate unbranched

(simple) hairs consisting of more than one cell,

and usually 0.5-5 mm long.

vi. ) Type 2 hairs - very short trichomes (<0.1

660

mm long), normally with one stalk cell, and a

transparent multicellular glandular head.

vii.) stellate hair density

very dense - stellae overlapping, multi¬

layers, subtending surface not visible with

hand lens

dense - stellae overlapping, 0.1-0.5

Austrobaileya 6 (4): 639-816 (2004)

diameters apart, subtending surface visible

moderate - stellae overlapping, 0.5-1

diameter apart, centre to centre

sparse - stellae not overlapping, 1-2

diameters apart, centre to centre

very sparse - stellae not overlapping, > 2

diameters apart, centre to centre.

1. Leaf margins with 2 or more pairs of deep lobes (lobing index >2) OR leaf base hastate . 2

Leaf margins entire or shallowly lobed throughout (lobing index 1-2). 23

2. Some or all leaves with hastate base. 3

Leaves deeply lobed throughout, never hastate. 6

3. Stellae of lower leaf surface with central ray 1.5-3 times as long as laterals . . 21. S. latens

Stellae of lower leaf surface with central ray 0-1 times as long as laterals. 4

4. Corolla rotate, mature fruits green, 11-16 mm diameter; pedicels 0.8-

0.9 mm thick at anthesis. 73. S. amblymerum

Corolla deeply lobed, mature fruits red, 6-9 mm diameter; pedicels 0.2-

0.7 mm thick at anthesis. 5

5. Lamina 1.1-2.6 cm wide at midpoint; branchlet prickles broad-based (4-7

times longer than wide); fruiting pedicels 8-12 mm long. 24. S. chenopodinum

Lamina 0.2-0.7 cm wide at midpoint; branchlet prickles acicular (9-13

times longer than wide); fruiting pedicels 13—16 mm long. 20. S. ferocissimum

6. Stellate hairs completely lacking from branchlets and leaves. 7

At least some stellate hairs present on branchlets or leaves. 11

7. Finger hairs present on leaves. 8

Finger hairs absent from leaves (Type 2 hairs sometimes present). 9

8. Finger hairs gland-tipped. 39. S. adenophorum

Finger hairs not gland-tipped. 33. *S. capsicoides

9. Calyx with 12-40 prickles; petioles 2.1-3.4 cm long. 38. S. papaverifolium

Calyx unarmed or with 1-4 prickles; petioles 0.3-1.2 cm long. 10

10. Calyx with 1-4 prickles; leaf prickles broad-based; fruiting pedicel 5-8 mm

long; Bk & Co. 44. S. pugiunculiferum

Calyx without prickles; leaf prickles acicular; fruiting pedicels 10-19 mm

long; Pc & Le . 22. S. dissectum

11. Upper leaf surface stellae dense to very dense. 12

Upper leaf surface stellae absent or very sparse to moderate. 13

12. Branchlet stellae 0.25-0.5 mm diameter; prickles 6-70 on lower leaf

surface; fruiting calyx tube accrescent, enclosing fruit; dioecious species

.85. S. carduiforme

Branchlet stellae 0.8-1.3 mm diameter; prickles 0-5 on lower leaf surface;

fruiting calyx tube not accrescent, lobes shorter than mature fruit .. 84. S. chippendalei

Bean, Taxonomy of Solanum subg. Leptostemonum 661

13. Fruits completely enclosed by prickly calyx; corolla yellow; style sigmoid

.89. *S. rostratum

Fruiting calyx tube not accrescent; corolla white to purple; style erect. 14

14. Branchlets with 350-1400 prickles per dm; mature fruits black. 15

Branchlets sparsely to moderately prickly (2-60 prickles per dm); mature

fruits green, yellow or red. 16

15. Lower leaf surface green; calyx without prickles. 27. S. coracinum

Lower leaf surface white to yellowish; calyx with up to 25 short prickles

29. S. semiarmatum

16.

17.

18.

19.

20 .

21 .

22 .

23.

24.

25.

26.

Lower leaf surface very densely stellate-hairy. 17

Lower leaf surface sparsely to moderately stellate-hairy. 19

Calyx prickles absent or 1-5; lower leaf surface with 0-2 prickles. 71. S. nobile

Calyx prickles 15-65; lower leaf surface with 6-45 prickles. 18

Branchlet stellae sparse, central ray absent; branchlet prickles 5-9 times

longer than wide; mature fruits 12-15 mm diameter. 43. S. lacunarium

Branchlet stellae dense to very dense, central ray present; branchlet prickles

11-16 times longer than wide; mature fruits 17-28 mm diameter. 70. S. cinereum

Finger hairs present on lower leaf surface. 90. *S. sisymbriifolium

No finger hairs on lower leaf surface. 20

Branchlet prickles broad-based, 2-4 times longer than wide. 88. *S. linnaeanum

Branchlet prickles acicular, 10-16 times longer than wide . 21

Lamina 1.2-2.6 cm long; petiole 0.15-0.45 cm long. 41. S. graniticum

Lamina 3-7 cm long; petiole 0.5-1.8 cm long. 22

Upper and lower leaf surfaces and calyx with Type 2 hairs; corolla white;

pedicels 3-6 mm long at anthesis. 40. S. pusillum

Type 2 hairs absent; corolla purple; pedicels 21-38 mm long at anthesis

. 42. S. stenopterum

Calyx prickles present on some or all flowers/fruits of inflorescence. 24

Calyx prickles absent from all flowers/fruits. 50

Finger hairs present on branchlets. 25

Finger hairs absent from branchlets. 27

Stellate hairs absent from branchlets and leaves. 33. *S. capsicoides

Stellate hairs present on branchlets and lower leaf surface. 26

Stellate hairs frequent on upper leaf surface, Type 2 hairs absent; calyx

stellae central ray 1-1.5 times as long as laterals; mature fruits 21-26 mm

diameter. 34. S. ditrichum

Stellate hairs absent from upper leaf surface, Type 2 hairs present; calyx

stellae absent or central ray 4-6 times as long as laterals; mature fruits

11-13 mm diameter . 35. S. cookii

27. Branchlet prickles curved, 2-3 times longer than broad . .

Branchlet prickles straight, 5-20 times longer than broad

87. *S. incanum

. 28

662

Austrobaileya 6 (4): 639-816 (2004)

28. Fruiting calyx tube accrescent, covering fruit. 29

Fruiting calyx tube not accrescent, and lobes not exceeding mature fruit. 30

29. Branchlet prickles 200-660 per dm; petioles 25-55% of lamina length;

calyx lobes 3-6 mm long at anthesis; fruiting pedicels 10-18 mm long 82. S. echinatum

Branchlet prickles 3-80 per dm; petioles 65-115% of lamina length; calyx

lobes 1-2.5 mm long at anthesis; fruiting pedicels 20-30 mm long . 83. S. longissimum

30. Branchlet stellae with central ray 2-4 times as long as laterals; Type 2

hairs present on branchlets and upper leaf surface; fruits densely hairy

.86. S. stupefactum

Branchlet stellae with central ray 0-1.8 times as long as laterals; Type 2

hairs absent; fruits glabrous or with a few scattered hairs. 31

31. Stellae of upper leaf surface very sparse to moderate (rarely absent). 32

Stellae of upper leaf surface dense to very dense. 43

32. Leaves 2.8-10 cm wide . 33

Leaves 0.4-2.7 cm wide. 39

33. Calyx prickles 0-11 . 34

Calyx prickles 12-100. 37

34. Leaves 6.5-10 cm wide; Type 2 hairs present on calyx. 64. S. francisii

Leaves 2.8-5.2 cm wide; Type 2 hairs absent from calyx. 35

35. Plants erect; leaf lobing index 1.7-2. 71. S. nobile

Plants prostrate or procumbent; leaf lobing index 1-1.4. 36

36. Upper leaf surface green, stellae very sparse or confined to midrib; calyx

prickles 0-2. 60. S. acanthodapis

Upper leaf surface grey-green, stellae density moderate; calyx prickles

5-11. 45. S. ellipticum

37. Upper leaf surface with 2-20 prickles, stellae 0.1-0.5 mm apart; lower

leaf surface white or yellowish, stellae 0.05-0.3 mm apart. 45. S. ellipticum

Upper leaf surface with 100-400 prickles, stellae 0.6-2.5 mm apart; lower

leaf surface green, stellae 0.3-2.2 mm apart. 38

38. Leaves entire or obscurely lobed, apex obtuse; common peduncle 17-27 mm

long; mature fruits yellowish-green or green; Nk & Co. 36. S. multiglochidiatum

Leaves with distinct acute lobes, apex acute; common peduncle 0-8 mm

long; mature fruits purple; Wb, Mo & Dd. 37. S. vicinum

39. Branchlet stellae sparse; pedicels 21-38 mm long at anthesis; petioles

winged; stellae of lower leaf surface 0.8-3 mm apart. 42. S. stenopterum

Branchlet stellae dense to very dense; pedicels 3-20 mm long at anthesis;

petioles not winged; stellae of lower leaf surface 0.05-0.7 mm apart. 40

40. Leaves 1.6-2.9 times longer than broad . 41

Leaves 3-14 times longer than broad . 42

41. Leaves ± entire, 3.5-14 cm long.

Leaves conspicuously lobed, 1.2-2.6 cm long

. 45. S. ellipticum

41. S. graniticum

Bean, Taxonomy of Solanum subg. Leptostemonum 663

42. Leaves 1.5-4 cm long, lower surface stellae sparse to dense; Co.51. S. angustum

Leaves 7-10.5 cm long, lower surface stellae very dense; Mo, Dd, Bn .... 72. S. limitare

43. Calyx of basal flower(s) within an inflorescence larger and more prickly,

and pedicel(s) thicker, compared to calyces and pedicels of distal flowers

. 84. S. chippendalei

Each inflorescence with all flowers/fruits having calyces all about the same

size and prickliness, and pedicels all about the same thickness. 44

44. Leaves linear to lanceolate (1/b ratio 3.8-14). 45

Leaves ovate to broadly ovate (1/b ratio 1.6-2.9). 46

45. Branchlet stellae 0.4-0.5 mm diameter, lateral rays 15-18; calyx stellae

with 13-20 lateral rays; anthers 7-9 mm long; seeds brown to black

.78. *S. elaeagnifolium

Branchlet stellae 0.7-0.9 mm diameter, lateral rays 8-12; calyx stellae

with 6-9 lateral rays; anthers 3.5-5 mm long; seeds pale yellow. 77. S. esuriale

46. Branchlet prickles absent; calyx prickles 1-4.46. S. dianthophorum

Branchlet prickles present; calyx prickles 5-70. 47

47. Plants erect, 0.3-0.6 m high; calyx prickles 40-70. 48

Plants prostrate or sprawling, 0.1-0.3 m high; calyx prickles 5-40(-50). 49

48. Leaf base cuneate; lower leaf surface rusty; calyx stellae 0.4-0.6 mm

diameter.48. S. senticosum

Leaf base obtuse or cordate; lower leaf surface yellowish; calyx stellae

0.7-0.9 mm diameter.47. S. crebrispinum

49. Calyx prickles stout, rigid; branchlets terete; inflorescence 1-9 flowered,

rachis prickles present; stalks of stellae on upper leaf surface 0-0.3 mm

long.45. S. ellipticum

Calyx prickles flimsy, scarcely rigid; branchlets ridged; inflorescence

9-15 flowered, rachis prickles absent; stalks of stellae on upper leaf

surface 0-1.0 mm long. 49. S. quadriloculatum

50. Leaves linear to lanceolate (1/b ratio 3.3-15). 51

Leaves ovate, broadly ovate, elliptical or orbicular (1/b ratio 0.8-3.2). 76

51. Leaf upper surface stellae dense to very dense. 52

Leaf upper surface stellae absent or density very sparse to moderate. 59

52. Calyx lobes elliptical, exceeding mature fruits; central ray of stellae (upper

leaf surface) 3-6 times as long as laterals. 5. S. innoxium

Calyx lobes deltate to attenuate, not exceeding mature fruits; central ray

of stellae (upper leaf surface) absent or up to 1.7 times as long as laterals. 53

53. Finger hairs present on calyx and upper leaf surface; bark corky.76. S. versicolor

Finger hairs absent from plant; bark not corky. 54

54. Type 2 hairs present on branchlets and lower leaf surface;

flowers 4-merous.79. S. ammophilum

Type 2 hairs absent; flowers 5-merous. 55

55. Upper leaf surface with stellae 0.4-0.7 mm diameter. 56

Upper leaf surface with stellae 0.15-0.4 mm diameter. 57

664

Austrobaileya 6 (4): 639-816 (2004)

56. Branchlet stellae 0.4-0.5 mm diameter, lateral rays 15-18; calyx stellae

with 13-20 lateral rays; anthers 7-9 mm long; seeds brown to black

.78. *S. elaeagnifolium

Branchlet stellae 0.7-0.9 mm diameter, lateral rays 8-12; calyx stellae

with 6-9 lateral rays; anthers 3.5-5 mm long; seeds pale yellow. 77. S. esuriale

57. All stellate hairs lacking a central ray; lower leaf surface stellae with

13-16 lateral rays.80. S. sturtianum

All stellate hairs with an obvious central ray; lower leaf surface stellae

with 7-9 lateral rays. 58

58. Branchlet stellae porrect, central ray 0.3-0.7 times as long as laterals;

branchlet prickles 0.5-6 mm long; calyx stellae sessile or stalks to

0.1 mm long. 61. S. intonsum

Branchlet stellae porrect to multiradiate, central ray 1-1.5 times as long

as laterals; branchlet prickles 6-10 mm long; calyx stellae with stalks

0.1-0.4 mm long.68. S. jucundum

59. Fruiting pedicels 5-13 mm long. 60

Fruiting pedicels 13-33 mm long. 67

60. Calyx lobes elliptic, exceeding mature fruit . 61

Calyx lobes deltate, rostrate or attenuate, never exceeding mature fruit. 62

61. Upper leaf surface stellae moderate to dense; branchlet stellae 0.25-0.4 mm

diameter, central ray 3-4 times as long as laterals; Dd & Ma. 5. S. innoxium

Upper leaf surface stellae very sparse to sparse; branchlet stellae 0.6-0.9 mm

diameter, central ray 0.4-0.8 times as long as laterals; Mi & Bk. 6. S. ultimum

62. Upper leaf surface stellae 0.05-0.4 mm apart (moderate density) . 63

Upper leaf surface stellae 0.4-5 mm apart (very sparse to sparse). 66

63. Leaves linear, 0.3-0.6 cm wide. 74. S. galbinum

Leaves linear to ovate, 0.7-6.5 cm wide. 64

64. Upper leaf surface stellae with lateral rays ascending; inner surface of

corolla sparsely stellate-hairy. 61. S. intonsum

Upper leaf surface stellae with lateral rays porrect; inner surface of corolla

glabrous. 65

65. Stellae of upper leaf surface 0.4-0.7 mm across, lateral rays 6-12, central

ray 0.5-1 times as long as laterals; common peduncle 14-36 mm long .. 77. S. esuriale

Stellae of upper leaf surface 0.15-0.3 mm across, lateral rays 13-15, central

ray absent; common peduncle 3-5 mm long.80. S. sturtianum

66. Upper leaf surface stellae 0.1-0.2 mm diameter and 0.4-0.7 mm apart;

mature fruits 11-16 mm diameter, green.73. S. amblymerum

Upper leaf surface stellae 0.3-0.5 mm diameter and 0.8-5 mm apart; mature

fruits 6-8 mm diameter, red. 19a. S. parvifolium subsp. parvifolium

67. Stellate hairs absent from fully expanded leaves. 23. S. lythrocarpum

Stellate hairs present on fully expanded leaves. 68

68. Lower leaf surface stellae with central ray 1.5-6 times as long as laterals. 69

Lower leaf surface stellae with central ray 0-1 times as long as laterals. 71

665

Bean, Taxonomy of Solanum subg. Leptostemonum

69. Leaves 0.3-0.8 cm wide; lower surface green, with stellae 0.3-2 mm apart .. 21. S. latens

Leaves 0.9-4.5 cm wide; lower surface white, yellowish or rusty, with

stellae 0.05-0.2 mm apart. 70

70. Petioles 3-7% length of lamina; pedicels 4-7 mm long at anthesis; leaves

often with shallow lobes; Cape York Peninsula. 10. S. fervens

Petioles 11-16% length of lamina; pedicels 8-15 mm long at anthesis;

leaves entire; Nk, Pc, Wb, Dd, Bn, Mo .18. S. stelligerum

71. Branchlet stellae 0.7-0.9 mm diameter; common peduncle 14-36 mm long 77. S. esuriale

Branchlet stellae 0.2-0.6 mm diameter; common peduncle 0-7 mm long. 72

72. Upper leaf surface grey-green; herbaceous resprouter; mature fruits green . 72. S. limitare

Upper leaf surface green; perennial shrubs; mature fruits red. 73

73. Leaves lanceolate to ovate, stellae absent from upper surface; mature fruits

10-14 mm diameter; Cape York peninsula only. 13. S. discolor

Leaves linear to narrow-lanceolate; stellae very sparse to moderate on upper

surface; mature fruits 5-9 mm diameter; never on Cape York peninsula. 74

74. Seeds 3.0-3.6 mm long; prickles present on both leaf surfaces.20. S. ferocissimum

Seeds 1.9-2.4 mm long; prickles absent from leaves. 75

75. Leaves 2.3-7 x 0.5-1.5 cm, Type 2 hairs absent; style 3.8-6.5 mm long;

flowers all bisexual. 19a. S. parvifolium subsp. parvifolium

Leaves 7-13.7 x 1.3-3.6 cm, Type 2 hairs present; style 6.5-9.0 mm long;

male flowers present in the inflorescence. 19b. S. parvifolium subsp. tropicum

76. Upper leaf surface with moderate to very dense stellate indumentum. 77

Upper leaf surface glabrous or stellate indumentum very sparse to sparse. 97

77. Calyx lobes elliptic, exceeding mature fruit; branchlet prickles absent. 78

Calyx lobes deltate, rostrate, hemispherical or attenuate, not exceeding

mature fruit; branchlet prickles absent or present. 83

78. Stellae of branchlets, upper and lower leaf surface, and calyx all sessile. 79

At least some stellae on branchlets, leaves and calyx with conspicuous

stalks (i.e. stalks at least 0.2 mm long, and up to 2 mm long). 81

79. Leaves broadly ovate; petioles 30-55% of lamina length; subshrub

0.1-0.25 m high. 4. S. johnsonianum

Leaves lanceolate to ovate; petioles 11-25% of lamina length; shrubs 0.3-1 m high . 80

80. Leaves 2.3-3.5 cm wide; mature fruits globular, 5-6.5 mm diameter; Mo .. 3. S. hapalum

Leaves 0.8-2 cm wide; mature fruits ellipsoidal, 3-4.5 mm diameter; Ma, Dd 5. S. innoxium

81. Some stellae of branchlets, leaves and calyx with gland-tipped central ray;

leaves usually slightly lobed; ovary glabrous. 9. S. gympiense

No stellae with gland-tipped central ray; leaves entire; ovary with stellate

or Type 2 hairs. 82

82. Stellae of upper leaf surface with 4-6 lateral rays, central ray 3-6 times as

long as laterals, stalks absent. 7. S. densevestitum

Stellae of upper leaf surface with 7 or 8 lateral rays, central ray 0.7-1.8

times as long as laterals, stalks 0.1-0.3 mm long.8. S. nemophilum

666

Austrobaileya 6 (4): 639-816 (2004)

83. Leaves 0.8-1.7 times longer than broad

Leaves 1.7-3.2 times longer than broad

84. Lamina 1-2.5 cm long, petioles 0.1-0.3 cm long; stellae of upper surface

with 12-16 lateral rays.81. S. oligacanthum

Lamina 3.5-35 cm long, petioles 0.9-7.2 cm long; stellae of upper surface

with 4-10 lateral rays. 85

85. Stellae of upper leaf surface with central ray 4-10 times as long as laterals;

mature fruits conspicuously tomentose, 25-35 mm diameter.32. S. lasiocarpum

Stellae of upper leaf surface with central ray 0.8-3 times as long as laterals;

mature fruits glabrous, 8-17 mm diameter. 86

86. Branchlet prickles straight, acicular, 10-17 times longer than broad;

inflorescence unbranched or flowers solitary, 1-4 flowered. 52. S. argopetalum

Branchlet prickles curved, broad-based, 1.5-3 times longer than broad;

inflorescence branched, 13-65 flowered . 87

87. Finger hairs absent from calyx and pedicels; new growth rusty .... 30. *S.chrysotrichum

Finger hairs present on calyx and pedicels; new growth white or grey .... 31. *S. torvum

88. Leaves 0.5-1.5 cm long.75. S. elachophyllum

Leaves 2.5-16.5 cm long. 89

89. Branchlet prickles 0-20 per dm. 90

Branchlet prickles 21-1400 per dm. 95

90. Calyx with Type 2 hairs or finger hairs (as well as stellate hairs); bark corky. 91

Calyx with stellate hairs only; bark not corky, non-descript. 92

91. Calyx with stellate hairs and Type 2 hairs; shrub 1-3 m high. 62. S. furfuraceum

Calyx with stellate hairs and finger hairs; subshrub 0.1-0.3 m high.76. S. versicolor

92. Subshrubs 0.3-0.6 m high, herbaceous resprouters. 69. S. centrale

Shrubs 0.6-2 m high, perennials. 93

93. Stellae stalks (branchlets and calyx) up to 2.2 mm long; corolla rotate;

common peduncle 15-33 mm long. 67. S. cocosoides

Stellae stalks (branchlets and calyx) up to 0.4 mm long; corolla shallowly

or deeply lobed; common peduncle 0-12 mm long. 94

94. Type 2 hairs absent from upper leaf surface; branchlet stellae with porrect

lateral rays; stellae of lower leaf surface 0.4-0.6 mm diameter. 61. S. intonsum

Type 2 hairs present on upper leaf surface; branchlet stellae with lateral

rays porrect, ascending, or multiradiate; stellae of lower leaf surface

0.2-0.4 mm diameter.68. S. jucundum

95. Branchlet prickles broad, recurved, 1-2.5 times longer than broad .... 54. S. hamulosum

Branchlet prickles acicular, straight, 10-20 times longer than broad. 96

96. Inflorescence 1 or 2 flowered; branchlet prickles 30-160 per dm; prickles

absent from leaves.50. S. crassitomentosum

Inflorescence 7-25 flowered; branchlet prickles 240-1400 per dm; prickles

present on both leaf surfaces.28. S. mitchellianum

75. S. elachophyllum

. 98

97. Leaves 0.5-1.5 x 0.3-0.7 cm

Leaves 2.5-26 x 0.9-19 cm .

Bean, Taxonomy of Solanum subg. Leptostemonum 667

98. Plants prostrate, stoloniferous, rooting at the nodes. 99

Plants erect or sprawling, but not prostrate or stoloniferous. 101

99. Stellae of lower leaf surface lacking a central ray; mature fruits red. 15. S. mentiens

Stellae of lower leaf surface with clearly visible central ray; fruits green to

yellowish-green. 100

100. Stellae of branchlets with central ray 0.5-1.5 times as long as laterals;

upper leaf surface with 20-60 prickles; prickles present on lower leaf

surface.60. S. acanthodapis

Stellae of branchlets with central ray 4-9 times as long as laterals; upper

leaf surface with 0-8 prickles; lower leaf surface without prickles.59. S. serpens

101. Calyx lobes elliptic and exceeding mature fruit; large stems without

prickles. 6. S. ultimum

Calyx lobes deltate, rostrate or attenuate, not exceeding mature fruit; at

least large stems with some prickles. 102

102. Calyx and pedicel with finger hairs. 103

Finger hairs absent from calyx and pedicel. 105

103. Leaves 1.1-1.5 times longer than broad, lower surface white. 31. *S. torvum

Leaves 1.9-2.7 times longer than broad, lower surface green. 104

104. Calyx lobes 2-3.5 mm long at anthesis; fruiting pedicels c. 15 mm long;

lower leaf surface with 11-17 prickles; Co. 25. S. dysprosium

Calyx lobes 4-9 mm long at anthesis; fruiting pedicels 26-38 mm long;

lower leaf surface with 0-9 prickles; Mo. 26. S. inaequilaterum

105. Stellate hairs absent from petioles and lower surface of fully expanded

leaves; inflorescences mostly branched; branchlet prickles absent. 106

Stellate hairs present on petioles or lower surface of fully expanded leaves;

inflorescences mostly unbranched; branchlet prickles usually present. 107

106. Petioles 7-10% of lamina length; common peduncle 5-8 mm long; calyx

densely stellate-hairy, stellae with central ray deflexed; style 7-8 mm

long . 1. S. dunalianum

Petioles 17-30% of lamina length; common peduncle 12-23 mm long;

calyx sparsely stellate-hairy, stellae with central ray erect; style 4.5-

5.5 mm long. 2. S. viridifolium

107. Branchlet prickles present, curved, broad-based, 1-5 times longer than broad. 108

Branchlet prickles absent or present (and then straight, 6-20 times longer

than broad). 110

108. Leaves 9-19 cm wide; branchlet stellae on thick stalks 0.1-1 mm long;

inflorescence 3 or 4 branched (paniculate).30. *S. chrysotrichum

Leaves 4.5-7.5 cm wide; branchlet stellae sessile or with short thready

stalks; inflorescences pseudo-racemose. 109

109. Leaves 1.9-2.5 times longer than broad, entire or with obtuse lobes;

stellae of branchlets and lower leaf surface lacking a central ray... . 53. S. dimorphispinum

Leaves 1.4-1.8 times longer than broad, with acute lobes; stellae of

branchlets and lower leaf surface with central ray 0.3-0.6 times as

long as laterals. 55. S. eminens

668

Austrobaileya 6 (4): 639-816 (2004)

110. Branchlets with corky outgrowths, large stems very corky.Ill

Branchlets without corky outgrowths, large stems not corky . 112

111. No finger hairs on upper leaf surface; stellae of lower leaf surface with

central ray 0.3-0.7 times as long as laterals; leaves consistently lobed

. 63. S. sporadotrichum

Finger hairs present on upper leaf surface; stellae of lower leaf surface

with central ray 1-2 times as long as laterals; leaves usually entire 62. S. furfuraceum

112. All stellae on lower leaf surface stalked (stalks 0.15-0.6 mm long). 113

Some or all stellae on lower leaf surface sessile. 114

113. Stellae on upper leaf surface absent or confined to midrib; mature fruits

red, 6-7 mm diameter. 17. S. dryanderense

Stellae distributed throughout upper leaf surface; mature fruits yellowish-

green to green, 19-24 mm diameter. 58. S. rixosum

114. Stellae of lower leaf surface with central ray absent or up to 0.6 times as

long as laterals. 115

Stellae of lower leaf surface with central ray 0.7-6 times as long as laterals. 119

115. Lower leaf surface white to yellowish, stellae dense to very dense. 116

Lower leaf surface green, stellae very sparse to moderate. 118

116. Stellae of lower leaf surface without a central ray. 14. S. corifolium

Stellae of lower leaf surface with central ray 0.1-0.5 times as long as laterals. 117

117. Lower leaf surface stellae very dense, c. 0.05 mm apart. 13. S. discolor

Lower leaf surface stellae dense, 0.1-0.3 mm apart. 11. S. yirrkalense

118. Petioles 0.6-1 cm long; stellae absent from upper leaf surface, 0.4-0.6 mm

diameter on lower surface; pedicels 0.15-0.25 mm thick at anthesis;

Cape York. 12. S. defensum

Petioles 1.4-2.6 cm long; stellae present on upper leaf surface, 0.2-0.3 mm

diameter on lower surface; pedicels 0.4-0.7 mm thick at anthesis;

Wet Tropics. 56. S. macoorai

119. Branchlet prickles 30-1400 per dm; petiole prickles present. 120

Branchlet prickles 0-25 per dm; petioles without prickles. 121

120. Branchlet prickles 240-1400 per dm; lamina base obtuse to cordate;

inflorescence 7-25 flowered; mature fruits black, 8-9 mm diameter

.28. S. mitchellianum

Branchlet prickles 30-70 per dm; lamina base cuneate; inflorescence

1-3 flowered; mature fruits green, 16-19 mm diameter. 65. S. dumicola

121. Some leaves shallowly lobed. 122

All leaves entire . 123

122. Petioles 0.4-0.8 cm long; calyx lobes 0.5-1.2 mm long at anthesis; mature

fruits red, 9-12 mm diameter. 10. S. fervens

Petioles 1-3.5 cm long; calyx lobes 3-4 mm long at anthesis; mature

fruits yellowish-green to yellow, 20-27 mm diameter. 57. S. magnifolium

Bean, Taxonomy of Solanum subg. Leptostemonum

669

123. Pedicels 0.8-1.3 mm thick at midpoint; inner surface of corolla stellate-

hairy; fruits 4-locular.66. S. tetrathecum

Pedicels 0.2-0.7 mm thick at midpoint; inner surface of corolla glabrous;

fruits 1-locular. 124

124. Stellae of lower leaf surface all sessile, stellae 0.1-0.3 mm apart; mature

fruits 3.5-6 mm diameter; petioles 5-12% of lamina length.16. S. shirleyanum

Lower leaf surface with some long-stalked stellae (to 0.6 mm long),

stellae 0.05-0.1 mm apart; mature fruits 6.5-9 mm diameter; petioles

11-16% of lamina length .18. S. stelligerum

Synoptic key to Solanum subg. Leptostemonum , using characters applicable to live

material

Notes on the use of this key: Look through the

list of characters and character states below.

Choose applicable character states that are not

commonly occurring. For example, if your

specimen has orange fruits, then fruit colour

will be a good character to use, because only 5

species have orange fruits. Write down all the

numbers adjacent to the first character state you

chose. Choose a second character, decide on

the correct state, and write down the numbers

adjacent to that state. Circle the numbers that

are common to the two lists. Select a third

character, decide on the correct state, and write

down the numbers adjacent to that state. Circle

the numbers that are common to all three lists.

Continue in this way until you reach a single

number, and then match that number with the

species name in the list appearing directly

below the key.

Character

Character State

Taxa possessing that state

Habit

herbaceous resprouter

34-36, 38^16, 49, 51, 52, 66, 69, 72, 76-79, 81, 89

prostrate stoloniferous

15, 59, 60

rhizomatous shrub

1-14, 16-33, 45-50, 53, 56-58, 61-64, 67, 68,

70, 71, 73-75, 80-90

vine or shrubby vine

46, 53-55

Bark

corky

50, 62, 63, 76

non-descript

most species

Prickles, large stems

absent

1, 3-9, 46

curved, broad based

(1-4 times longer than wide)

24, 30, 31, 53-55, 87, 90

straight, broad based

(4-7 times longer than wide)

2, 16, 24, 31, 32, 43, 44, 61, 64, 67, 68,

71, 78-80, 83, 88-90

Straight, acicular

(7-20 times longer than wide)

10-14, 15-23, 25-29, 32^15, 47, 49-52, 56-58,

60, 62-66, 68-78,81-86, 89

Adult leaves

(length-breadth ratio)

0.8-1.7

4, 7, 9, 15, 22, 24, 27-29, 30-39, 40, 41,

44^15, 47, 49, 50, 52, 54, 55, 59, 60,

63, 65,70,71,75,81-83, 86-90

1.8-3

1-18, 22, 24-26, 28, 29, 30, 34, 36-43,

45-51, 53-71, 75, 76, 82-86, 88-90

3.1-26

1, 5, 6, 10, 12, 13, 17-19, 20, 21, 23,

42, 51, 56, 61, 66, 68, 72-74, 76-80, 84

670

Austrobaileya 6 (4): 639-816 (2004)

Lower leaf surface

(density of stellate hairs)

absent

very sparse

sparse

moderate

dense

very dense

1,2, 22, 23, 27, 33,38,39,44

12,21,27, 35,37,42, 56

12, 20, 21, 26, 34-37, 40-42, 51, 56, 60, 75, 88

12, 20, 21, 25, 26, 30, 34-36, 40^12, 51,

56, 57, 59, 60, 63, 64, 75, 88-90

3, 4, 6-11, 16-18, 20, 24, 29-32, 36, 45,

47, 48, 50-52, 54, 55, 57-63, 65, 66, 75-77,

79, 82-84, 86, 87, 89

3-6, 8, 9, 13, 14, 15, 18-20, 24, 28, 29, 32,

43, 45-50, 53, 55, 57, 59, 60, 67-74, 76-82, 84, 85

Corolla colour

white

2, 11, 12, 14, 15, 18, 20, 26, 30-33, 36, 39, 40,

45, 50, 52, 64, 74, 76, 90

yellow

mauve or purple

most species

Mature fruit colour

(or predominant colour)

red

1-24, 26, 56, 90

black

27-29

green to yellowish-green

30, 31, 34, 35, 38—41, 45, 47-52, 54,

57-68, 70, 71-77, 79, 84

yellow

44, 53, 57,80,81,87, 88

orange

32, 33, 55-56, 86

brown

44, 80-83

purple

Mature fruit diameter (fresh)

3-9.0 mm

1-10, 14, 16-25, 27, 28, 44, 46, 80, 81, 89

9.1-13.0 mm

10, 12-15, 23, 26, 29, 31, 35, 42-46, 48, 54, 63,

66, 73-81, 89

13.1-16.0 mm

11-13, 30, 31, 36, 39—41, 43, 45, 47-52, 54, 59-62,

64-68, 72-75, 77, 78, 82, 90

16.1—45 mm

12, 31-34, 36, 37, 45, 49-51, 53, 55-58, 60-62,

64-67, 70-72, 82, 84-88, 90

Calyx prickle number

zero

most species

1-35

29, 33, 35, 38-46, 49, 51, 60, 64, 70-72, 77, 78,

83-88, 90

36-600

34, 36-38, 41, 42, 45, 48, 49, 70, 82-85, 88, 90

Seed colour (fresh)

white to pale yellow

most species

brown to black

27, 28, 32, 50, 66-68, 70, 75, 78, 80-82, 84, 86-89

Distribution (Pastoral district)

Mo or Wb or NE NSW

3, 7-9, 14-16, 18, 26, 29-31, 33, 34, 37, 45,

58-60, 66, 72, 73, 78, 86, 88, 89

5, 8, 14, 18-21, 27-29, 34, 37, 38, 42, 45, 58, 66,

68, 70-73, 77, 78, 86, 89, 90

Bn or Pc

2, 4, 8, 9, 14, 18, 19, 21-23, 28, 30, 31, 33, 42,

45, 46, 58, 62, 66, 68, 72, 77, 78, 86, 87, 89

Le or Sk

2, 4, 7-9, 19-22, 27, 28, 31, 33, 39^11, 45, 47,

62-65, 67, 68, 74, 75, 77, 88

Bean, Taxonomy of Solanum subg. Leptostemonum

671

2, 14, 17-19, 31, 35, 36, 41, 45, 50, 54,

56, 57, 62, 63, 74, 77

1, 2, 10-14, 19, 25, 31-33, 35, 36, 44, 45,

51,53,54-57,61,74, 83, 85

Ma, Wa, Gs, Gn, Mi or Bk

5, 6, 9, 19, 20, 24, 28, 31, 43^15, 48-50,

52, 62, 65, 67-69, 74, 76, 77, 79-85

01. S. dunalianum

02. S. viridifolium

03. S. hapalum

04. S. johnsonianum

05. S. innoxium

06. S. ultimum

07. S. densevestitum

08. S. nemophilum

09. S. gympiense

10 . S.fervens

11 . S. yirrkalense

12 . S. defensum

13 . S. discolor

14 . S. corifolium

15 . S. mentiens

16 . S. shirleyanurn

17 . S. dryanderense

18 . S. stelligerum

19 . S. parvifolium

20 . S. ferocissimum

21 . S. latens

22 . S. dissectum

23 . S. lythrocarpum

24 . S. chenopodinum

25 . S. dysprosium

26 . S. inaequilaterum

27 . S. coracinum

28 . S. mitchellianum

29 . S. semiarmatum

30 . S. chrysotrichum

31 . S. torvum

32 . S. lasiocarpum

33 . S. capsicoides

34 . S. ditrichum

35 . S. cookii

36 . S. multiglochidiatum

37 . S. vicinum

38 . S. papaverifolium

39 . S. adenophorum

40 . S. pusillum

41 . S. graniticum

42 . S. stenopterum

43 . S. lacunarium

44 . S. pugiunculiferum

45 . S. ellipticum

46 . S. dianthophorum

47 . S. crebrispinum

48 . S. senticosum

49 . S. quadriloculatum

50 . S. crassitomentosum

51 . S. angustum

52 . S. argopetalum

53 . S. dimorphispinum

54 . S. hamulosum

55 . S. eminens

56 . S. macoorai

57 . S. magnifolium

58 . S. rixosum

59 . S. serpens

60 . S. acanthodapis

61 . S. intonsum

62 . S. furfuraceum

63 . S. sporadotrichum

64 . S.francisii

65 . S. dumicola

66 . S. tetrathecum

67 . S. cocosoides

68 . S. jucundum

69 . S. centrale

70 . S. cinereum

71 . S. nobile

72 . S. limitare

73 . S. amblymerum

1A . S. galbinum

75 . S. elachophyllum

76 . S. versicolor

77 . S. esuriale

78 . S. elaeagnifolium

79 . S. ammophilum

80 . S. sturtianum

81 . S. oligacanthum

82 . S. echinatum

83 . S. longissimum

84 . S. chippendalei

85 . S. carduiforme

86 . S. stupefactum

87 . S. incanum

88 . S. linnaeanum

89 . S. rostratum

90 . S. sisymbriifolium

672

Solanum subg. Leptostemonum (Dunal) Bitter,

Bot. Jahrb. Syst. 55: 68 (1919); S. sect.

Leptostemonum Dunal in A.DC., Prodr.

13(1): 183 (1852). Type: S. mammosum

L., lecto ,fide D’Arcy, Ann. Missouri Bot.

Gard. 59: 270 (1973).

Herbaceous resprouters, perennial vines,

shrubs, or small trees. Prickles usually present

on stems and petioles, sometimes on leaves and

calyces, rarely on corolla, but in a very few

species absent altogether. Stellate hairs usually

present on many parts of the plant, rarely

absent. Leaves often having two distinct

ontogenetic stages. Juvenile leaves larger, more

deeply lobed and more prickly. Adult leaves

usually two per sympodial unit (in Queensland),

entire or variously lobed, the base oblique on

all or some leaves. Inflorescence cymose, often

appearing racemose or paniculate, or reduced

to a single flower. All flowers bisexual, or distal

flowers functionally male (andromonoecious),

or rarely dioecious. Flowers actinomorphic (in

Queensland), mostly 5-merous, some species

always or predominantly 4-merous. Calyx

fused, often accresent in fruit. Corolla lobes

fused, variously lobed. Stamens equal in size

and shape (in Queensland); anthers attenuate,

yellow, glabrous (in Queensland), dehiscing by

terminal pores. Ovary 1-4-locular, the placenta

variously lobed. Fruit a juicy or mucilaginous

(rarely dry) berry; stone cells absent. Seeds

lenticular, subreniform, the testa with fine

reticulated ornamentation.

About 500 species, throughout the world,

mainly in tropical and subtropical regions.

Group 5 (S. dunalianum group) of Whalen

(1984)

Large shrubs or small trees (100%); adult leaves

entire (100%); branchlet prickles absent

(100%); Stem prickles sparse, broad-based

(100%); stellate hairs present only on young

vegetative growth and inflorescences (100%);

flowers all bisexual (100%); flowers often 4-

merous (100%); mature fruits red, juicy,

succulent, 1-locular, <9 mm diameter, exocarp

<0.5 mm thick (100%); inflorescences 2-3-

branched (83%).

c. 15 species extending from Malesia to

the western Pacific. 3 species indigenous to

Australia, 2 species indigenous to Queensland.

Austrobaileya 6 (4): 639-816 (2004)

The species allied to S. vaccinioides (all

endemic to New Caledonia) were included in

this group by Whalen {loc. cit.). However they

seem to have little in common with S. dunalianum

and related species.

1. Solanum dunalianum Gaudich. in Freyc.,

Voyage Uranie 448 (1829), t. 58 (1828).

Type: Moluccas, Pisang, [December

1818], C. Gaudichaud-Beaupre s.n.

(lecto: P), here designated.

Illustration : Symon (1981: 119)

Erect, rhizomatous perennial shrub, 2-4 m

high. Leaves (in outline) ovate, entire; la min a

c. 35 x 15 cm, without prickles on upper

surface. Adult branchlets brown; prickles

absent; stellae absent or sparse, 0.25-0.35 mm

diameter, sessile; lateral rays 7-9, porrect;

central ray 0.4-0.8 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Adult leaves elliptical or ovate,

entire; la min a 13-23 cm long, 4.3-7.5 cm wide,

2.8-3.1 t im es longer than broad, apex acute,

base cuneate, oblique part 0-15 mm long,

obliqueness index 0-9 percent; petioles 1-1.8

cm long, 7-10% length of lamina, prickles

absent. Upper leaf surface green; prickles

absent; stellate hairs absent, or confined to

midrib; ordinary stellae absent from surface;

finger hairs absent; Type 2 hairs absent. Lower

leaf surface green; prickles absent; stellae

absent; finger hairs absent; Type 2 hairs absent.

Inflorescence supra-axillary, 2-branched,

common peduncle 5-8 mm long, rachis prickles

absent; 15-25-flowered, with all flowers

bisexual and 4 or 5-merous; pedicels 4-7 mm

long at anthesis, markedly thicker distally, 0.4-0.7

mm thick at mid-point, prickles absent. Calyx

tube 1.5-2 mm long, lobes deltate, 0.3-1 mm

long; prickles absent at anthesis; stellae sparse

to very dense, yellow or white, 0.15-0.25 mm

across, sessile, lateral rays 7-9, central ray 0.6-1.2

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Corolla

mauve, 7-9 mm long, deeply lobed, inner

surface sparsely stellate-hairy; anthers 4.5-5.5

mm long; ovary glabrous; functional style 7-8

mm long, erect, with stellate hairs only, stellae

0.25-0.3 mm across, lateral rays 8-11, central

ray 0.5-0.8 times as long as laterals. Fruiting

calyx with lobes less than half length of mature

fruit, prickles absent. Mature fruits 3-13 per

Bean, Taxonomy of Solanum subg. Leptostemonum

inflorescence, globular, 8-9 mm diameter, red,

1-locular (septum absent or incomplete);

mesocarp juicy, succulent; pedicels 8-16 mm

long in fruit, 0.8-1 mm thick at mid-point;

seeds pale yellow, 2-2.7 mm long.

Specimens examined : Queensland. Cook District: Embley

River, 10 mi les [16 km] S of Weipa, Aug 1974, Swan 141

(AD, BRI, CANB); Weipa South, behind old airstrip, Jul

1980, Tucker AM762 (BRI); Possum Scrub, Jun 1994,

Forster PIF15276 & Tucker (AD, BRI).

Distribution and habitat: In Queensland,

Solanum dunalianum is known only from the

vicinity of Weipa on Cape York Peninsula (Map 1).

Also known from Malesia (New Guinea and

some nearby islands). Queensland specimens

are recorded from the edges of semi-deciduous

rainforest, on ‘red lateritic ridges’.

Phenology: Both flowers and mature fruits

have been recorded for June, July and August.

Notes: S. dunalianum is close to S. viridifolium,

but S. dunalianum in Australia has young leaves

and petioles with stellate hairs (vs. glabrous

for S. viridifolium,)', petioles 7-10% of lamina

length (vs. 15-30% for S. viridifolium)',

common peduncle 5-8 mm long (vs. 12-23 mm

long for S. viridifolium ); pedicels and calyx

densely to very densely stellate hairy (vs.

glabrous or sparsely stellate-hairy); central ray

of stellae deflexed (vs. erect) and style 7-8 mm

long (vs. 4.5-5.5 mm for S. viridifolium).

There are two sheets of S. dunalianum at

P that were collected by Charles Gaudichaud-

Beaupre. The sheet with 5 leaves and the prickly

stem, with the label saying “Solanum

dunalianum, Aquartia Dl, lie Pisang, C.G.” is

chosen as lectotype, as that specimen more

closely matches the protologue.

I am uncertain about the correct

application of the name S. dunalianum. The

Australian species is perhaps referable to

S. torricellense Bitter, but further taxonomic

study is required.

Conservation status: Currently listed as

“Vulnerable” under the Queensland Nature

Conservation Act, 1992. S. dunalianum is

known from only 3 locations, none of which is

in a conservation reserve. Applying the IUCN

criteria (IUCN. 2001), the existing category of

“Vulnerable” is endorsed (VU Cl).

673

2. Solanum viridifolium Dunal in A.DC.,

Prodr. 13(1): 73 (1852). Type:

[Queensland. Cook District:] near Cape

Grafton, 9 June 1770, J. Banks & D.

Solander (holo: BM).

S. viride R.Br., Prodr. 445 (1810), nom. illeg.,

non GForst. ex Spreng. (1807).

Illustration : Symon (1981: 123), as S. viride

Erect, rhizomatous perennial shrub, 1.5-9 m

high. Juvenile branchlets with 0-10 prickles

per dm, 1-3 mm long; leaves (in outline) ovate,

entire; lamina 9-14 cm long, 4-6 cm wide,

without prickles on upper surface. Adult

branchlets brown or green; prickles absent (but

large stems prickly); stellae absent; finger hairs

absent; Type 2 hairs absent. Adult leaves ovate,

entire; lamina 5-18.5 cm long, 2-6.5 cm wide,

2.1- 2.9 times longer than broad, apex acute or

acuminate, base cuneate, oblique part 0-8 mm

long, obliqueness index 0-4 percent; petioles

1.2- 3.8 cm long, 15-30% length of lamina,

prickles absent. Upper leaf surface green;

prickles absent; stellate hairs absent, or

confined to midrib; ordinary stellae absent from

surface; finger hairs absent; Type 2 hairs absent.

Lower leaf surface green; prickles absent;

stellae absent; finger hairs absent; Type 2 hairs

absent. Inflorescence supra-axillary, pseudo-

racemose or 2-3-branched, common peduncle

12-23 mm long, rachis prickles absent; 12-60-

flowered, with all flowers bisexual and 4 or 5-

merous; pedicels 5-13 mm long at anthesis,

markedly thicker distally, 0.4-0.7 mm thick at

mid-point, prickles absent. Calyx tube 1-2 mm

long, lobes elliptic or deltate, 0.5-1.5 mm long;

prickles absent at anthesis; stellae sparse, white,

0.2-0.3 mm across, sessile, lateral rays 4-8,

central ray 0.5-1 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Corolla white, mauve or purple, 6-11

mm long, deeply lobed, inner surface sparsely

to densely stellate-hairy; anthers 3.5-5 mm

long; ovary with Type 2 hairs only; functional

style 4.5-5.5 mm long, erect, glabrous or with

stellate and Type 2 hairs, stellae c. 0.3 mm

across, lateral rays 4-6, central ray 1.5-2 times

as long as laterals. Fruiting calyx with lobes

less than half length of mature fruit, prickles

absent. Mature fruits 3-13 per inflorescence,

globular, 5.5-7.5 mm diameter, red, 1-locular

(septum absent or incomplete); placenta not

674

apparent; mesocarp juicy, succulent; exocarp

0.1-0.3 mm thick; pedicels 11-17 mm long in

fruit, 0.6-0.8 mm thick at mid-point; seeds pale

yellow, 2-3 mm long.

Specimens examined : New Guinea. Sabi, lower Wassi

Kussa River, Jun 1973, Henty NGF 49646 (A, BRI, CANB,

L); near Kunini village, Daru district, May 1986, Simaga

768 (CANB). Queensland. Cook District: Boolally, Upper

Barron R., Jun 1899, J.F. Bailey s.n. (BRI); S.F. 185 Danbulla,

Sep 1929, Doggrell A12 (BRI); Mt Spurgeon, Sep 1936,

White 10719 (BRI); Etty Bay, Dec 1941, White 11734 (BRI);

Brown’s Creek, Pascoe River, Jul 1948, Brass 19589 (BRI,

CANB); Ha mm ond Island, Jul 1974, Heatwole 303 (BRI);

Halloran’s Hill, Atherton, Aug 1975, Stocker 1420 (BRI,

QRS); T.R.14, Mcllwraith Range-Leo Ckroad, Sep 1975,

Hyland 8431 (BRI, CANB, QRS); Thursday Island, May

1977, Paton 10 (CANB); S.F. 194, Baldy Mountain S.F., Oct

1987, Foreman 1649 (AD, BRI, CANB, NSW, QRS);

Garraway Hill, 12° 42’S 143° 08’E, Jul 1993, Forster

PIF13540 & Tucker (AD, BRI, MEL, QRS); Mt Misery, S

of Cooktown, Sep 2000, McDonald KRM593 & Hines

(BRI). North Kennedy District: Dunk Island, undated,

Banfield s.n. (BRI); S bank of Tully River, Feb 1965, Everist

7795 (BRI); Edmund Kennedy N.P., near Cardwell, Dec

1991, Bean 3858 (AD, BRI); Kirrama Range, 18.5 km from

Kennedy, Dec 1993, Forster PIF14314 (BRI). South

Kennedy District: Mackay, Oct 1887, Griffith s.n. (BRI);

S.F.. 652 Cauley, Jul 1974, Hyland 7382 (CANB); Mt

Blackwood, Mar 1987, Thompson 80 (BRI); Crediton S.F.,

S ofEungellaN.P., Nov 1990 ,Bean 2545 (BRI). Port Curtis

District: Bobby Range road, S.F.67 Bulburin, E of Builyan,

Mar 1995, Bean 8444 (BRI).

Distribution and habitat : Solanum viridifolium

is distributed along the coast of Queensland,

north from about Monto (Map 1). It also

extends to southern New Guinea. It inhabits

notophyll rainforests in high rainfall areas,

especially at altitudes above 300 metres, but

also on the lowlands. A very common and

widespread species.

Phenology: Flowers and fruits may be found

at any time of the year.

Notes: Young plants of S. viridifolium are

frequently monopodial until about 1.5 metres

in height. Adult plants somet im es have short

broad-based prickles on the large stems,

although the flowering branchlets are

invariably unarmed. One specimen label ( Gray

2156) records a plant 9 metres high and 15 cm

diameter at breast height. This is the tallest

recorded Solanum in Australia.

Robert Brown collected this species on

or near Curtis Island (near Gladstone) in 1802.

It has never been recorded near Gladstone

since then.

Austrobaileya 6 (4): 639-816 (2004)

Conservation status: Widespread. Not

considered at risk.

Group 9A (S. densevestitum group), here

defined; related to Group 9 ( S. jubae

group) of Whalen (1984).

Calyx lobes elliptical, exceeding mature fruits

(100%); large stems without prickles (100%);

branchlet prickles absent (100%); calyx

prickles absent (100%); corolla inner surface

glabrous (100%); mature fruits red, juicy,

succulent, 1-locular, <9 mm diameter, exocarp

<0.5 mm thick (100%); seeds pale yellow, 2-3

mm long (100%); adult leaves entire (93%);

inflorescence with all flowers bisexual (93%);

inflorescence solitary or pseudo-umbellate

(71%); upper leaf surface stellae with central

ray 2-9 times as long as laterals (71%).

7 species endemic to Australia; 7 species

occurring in Queensland.

Solanum densevestitum and its allies form

a very distinctive group. They appear to be

closely related to Whalen’s Group 9, the African

Solanum jubae group, because of “the absence

of prickles, stellae with long central rays,

umbel-like inflorescence, ovate or obovate calyx

lobes, and the succulent red fruits” (Whalen

1984: 227-8).

3. Solanum hapalum A.R.Bean sp. nov. Frutex

usque ad 1 m altus, aculeis carens; folia

integra, ovata, indumento coacto; stellae

sessiles, radio centrali longo, in pagina

inferiore folii 0.4-0.5 mm diametro;

fructus globulares succosi et laete rubri

maturitate; lobi calycis elliptici, fructus

superantes. Typus: New South Wales.

North Coast: 2.3 km along Dingo Range

road, Clouds Creek State Forest, north of

Dorrigo, 9 September 2000, A.R. Bean

16862 (holo: BRI (1 sheet + spirit); iso:

MEL, NE, NSW)

Illustration: Symon (1981: 143), as

S. densevestitum.

Erect, rhizomatous perennial shrub, 0.5-1 m

high. Juvenile branchlets without prickles;

entire or shallowly-lobed. Adult stem prickles

absent. Adult branchlets yellow or brown;

prickles absent; stellae very dense, 0.3-0.4 mm

Bean, Taxonomy of Solanum subg. Leptostemonum

675

Fig. 13. Solanum hapalum. A. flowering branchlet x 1. B. flower x 3. C. style and ovary x 6. D. ovary showing Type 2 hairs

x 20. E. stellate hair from upper leaf surface x 60. F. mature fruit x 2. G. transverse section of fruit x 6. all from Bean 16854.

diameter, stalks 0-0.1 mm long; lateral rays 7

or 8, porrect; central ray 3-7 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Adult leaves ovate, entire;

lamina 4.5-7 cm long, 2.3-3.5 cm wide, 1.9-2.5

times longer than broad, apex obtuse or acute,

base obtuse or cordate, oblique part 1-4 mm

long, obliqueness index 2-6 percent; petioles

0.8-1.7 cm long, 17-25% length of lamina,

prickles absent. Upper leaf surface grey-green;

prickles absent; stellate hairs distributed

throughout; protostellae absent; ordinary stellae

density moderate to dense, 0.15-0.3 mm apart,

0.25-0.4 mm across, sessile; lateral rays 6-8,

porrect; central ray 4-9 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Lower leaf surface white or

yellowish; prickles absent; stellae dense to very

dense, 0.1-0.2 mm apart, 0.4-0.5 mm diameter,

sessile; lateral rays 7 or 8, porrect; central ray

3-5 times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent.

Inflorescence supra-axillary, solitary or pseudo-

umbellate, common peduncle absent; 1-3-

flowered, with all flowers bisexual and 5-

merous; pedicels 2-7 mm long at anthesis, same

thickness throughout, 0.6-0.7 mm thick at mid¬

point, prickles absent. Calyx tube 1-1.5 mm

long, lobes elliptic, 4.5-8 mm long; prickles

676

Austrobaileya 6 (4): 639-816 (2004)

absent at anthesis; stellae very dense,

transparent, 0.3-0.5 mm across, stalks 0-0.1

mm long, lateral rays 7 or 8, central ray 3-5

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Corolla

mauve, 7-12 mm long, shallowly lobed, inner

surface glabrous; anthers 4-5 mm long; ovary

with Type 2 hairs only; functional style 5-8

mm long, erect, glabrous or with Type 2 hairs

only. Fruiting calyx with lobes exceeding

mature fruit, prickles absent. Mature fruits 1

or 2 per inflorescence, globular, 5-6.5 mm

diameter, red, 1-locular (septum absent or

incomplete); placenta not apparent; mesocarp

juicy, succulent; exocarp 0.2-0.4 mm thick;

pedicels 8-11 mm long in fruit, 0.7-1 mm thick

at mid-point; seeds pale yellow, 2.2-2.5 mm

long. Fig. 13.

Specimens examined : Queensland. Moreton District: Mt

Lindesay, Oct 1921, White 1121 (BRI); Border road 4 km S

of the sawmill on Burnett Ck, close to N.S.W. border, Apr

1973, Sharpe 447 (BRI); Mt Barney, south slopes, Oct 1992,

Forster 11857 et al. (AD, BRI). New South Wales. Northern

Tablelands: 30 miles [48 km] NE of Glen Innes, Gibraltar

S.F., Apr 1956, Constable (NSW); Gibraltar Range N.R, 68

km E of Glen Innes, Oct 1969, Coveny 2243a (NSW);

Surface Hill, ESE of Tenterfield, Jan 1975, Moriarty 1640

(BRI, CANB). North Coast: F.R. 121 Raleigh, Bellinger

R., Apr 1910, Swain 157 (NSW); Coffs Harbour, Jun 1911,

Boorman (BRI, NSW); Taree, Apr 1951, Noonan (NSW); 3

miles [5 km] W of Drake, Girard S.F., Apr 1956, Constable

s.n. (NSW); North Obelisk, 2 km SW of Urbenville, Dec

1977, Haegi 1539 (BRI, NSW); 0.4 km W along Dennis

Road, Ingalba S.F., 27 km NW of Kempsey, Dec 1998,

Lepschi 4021 & Connors (CANB, NSW); Forbes Forest

road, NW of Wauchope, Nov 1999, Bean 15702 (BRI,

NSW); Welsh’s Road, Clouds Creek S.F., N of Dorrigo, Sep

2000, Bean 16854 (BRI, NSW); Blacksmiths Shop road,

Dalmorton S.F., SW of Grafton, Jan 2001, Bean 17252 (BRI).

South Coast: Long Beach, Batemans Bay, Apr 1990,

Hancock (AD, CANB, NSW); Blairs Road, Square Head,

Batemans Bay, Jan 1995, Rees 300 (CANB); Eurobodalla

Regional Botanic Gardens, Dec 1999, Booth 2511 (AD, BRI,

NSW). Cultivated: Waite Institute ex Little Smoky, N.S.W.,

Dec 1967, Symon 4701 (AD, BRI, NSW).

Distribution and habitat: Solanum hapalum

extends from the extreme south-east of

Queensland (Mt Barney, Mt Ballow, Mt

Lindesay) to around Port Macquarie in New

South Wales (Map 1), and with a disjunct

occurrence near Batemans Bay. It grows on

rainforest margins or in “wet sclerophyll”

eucalypt forest, preferring disturbed areas such

as roadsides or recently logged sites.

Phenology: Flowers and fruits are recorded for

most months of the year.

Notes: S. hapalum is related to S. densevestitum,

but differing by the smaller stellate hairs on all

plant parts, the stellate hairs sessile on the

lower leaf surface (stalks up to 0.5 mm long

for S. densevestitum), the branchlet stellate

hairs sessile or almost so, and with 7 or 8 lateral

rays (stalks conspicuous, to 2 mm long, and

with 4-6 lateral rays for S. densevestitum) and

the ovary with Type 2 hairs only (Type 2 hairs

and stellate hairs for S. densevestitum ).

Conservation status: Widespread. Not

considered at risk.

Etymology: from the Latin hapalus, meaning

‘soft to the touch’, in reference to the soft, felty

leaves and stems.

4. Solanum johnsonianum A.R.Bean sp. nov.

Subfrutex usque ad 0.25 m altus, aculeis

carens; folia integra late ovata, indumento

coacto; stellae sessiles, radio centrali

longo, in pagina superiore folii 0.25-0.35

mm diametro; fructus globulares, 5.5-8

mm diametro, succosi et laete rubri

maturitate; lobi calycis elliptici, fructus

superantes. Typus: Queensland.

Leichhardt District: “Nirvana”, c. 15 km

WNW of Banana, 18 April 2003, A.R.

Bean 20165 (holo: BRI (1 sheet + spirit);

iso: CANB, MEL, NSW, distribuendi ).

Erect, rhizomatous perennial shrub, 0.15-0.3

m high. Juvenile branchlets without prickles;

leaves (in outline) broadly ovate, shallowly-

lobed, with 2 pairs of lobes; lamina 4.5-5.5

cm long, 3.3-3.8 cm wide, without prickles on

upper surface. Adult stem prickles absent. Adult

branchlets grey to rusty or brown; prickles

absent; stellae very dense, 0.4-0.5 mm

diameter, sessile; lateral rays 7-9, porrect;

central ray 3-5 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Adult leaves broadly ovate, entire;

lamina 2.5-6 cm long, 1.3-4.3 cm wide, 1.4-2

times longer than broad, apex obtuse, base

obtuse or cordate, oblique part 0-1 mm long,

obliqueness index 0-3 percent; petioles 0.9-2.8

cm long, 30-55% length of lamina, prickles

absent. Upper leaf surface green; prickles

absent; stellate hairs distributed throughout;

protostellae present; ordinary stellae dense,

0.15-0.25 mm apart, 0.25-0.35 mm across,

Bean, Taxonomy of Solanum subg. Leptostemonum

677

QUEENSLAND H EH BARIUM (BRI)

Brisbane Australia

QUEENSLAND HERBARIUM (BRI)

Ffcrs □wki'»U«S ^

Selanum

90s 1490 58m S)» (<.S!5W| Oipfirn

HSS.HBSSI.IiSfiHB |W“4S4E3SFS| "■ m

Tir-raKi', S 15ten WNW el BmAS.

OS»n folWi ^ Euc^lJluS BvsWana, AeWs hjFSS^ylU. 'Min

uftSaraiorny *1 Si^cra jjar>iltoT 3 > Mr* ndafthjfl, flat, E 46 W 1 &>£

Small vwixly luMUylr SOcnl Wall, pndUot aSHjnl- Pmite fcnGM Ha.

jplants rtiiMAifliflui with I'wmarsu? rtciaa in cfca* pnKwmv

OesmoiHi aii«. ai*sy* unainiMUi wuffi plants

SplnldiiXilalil BRi

, Queensland Herbarium (BRI)

faLr>7y#f t f

Sofanam joA/sso/t/'aitvm,

0*. A A. £&u* Oiw ) ^ 2<x>3

Fig. 14. Holotype of Solanum johnsonianum.

678

Austrobaileya 6 (4): 639-816 (2004)

sessile; lateral rays 7-9, porrect; central ray 3-

5 times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Lower

leaf surface greenish-white or white; prickles

absent; stellae dense to very dense, 0.1-0.2 mm

apart, 0.3-0.6 mm diameter, sessile; lateral rays

7 or 8, porrect; central ray 2-5 t im es as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Inflorescence supra-

axillary, solitary or pseudo-racemose, common

peduncle 0-2 mm long, rachis prickles absent;

1 or 2-flowered, with all flowers bisexual and

5- merous; pedicels 4-7 mm long at anthesis,

same thickness throughout, c. 0.7 mm thick at

mid-point, prickles absent. Calyx tube 1-3 mm

long, lobes elliptic, 4-6.5 mm long; prickles

absent at anthesis; stellae very dense, yellow,

0.3-0.4 mm across, sessile, lateral rays 7 or 8,

central ray 3-5 times as long as laterals, not

gland-tipped or gland-tipped; finger hairs

absent; Type 2 hairs absent. Corolla mauve,

6- 11 mm long, rotate or shallowly lobed, inner

surface glabrous; anthers 3-4 mm long; ovary

glabrous, or with Type 2 hairs only; functional

style 5-6.5 mm long, erect, glabrous or with

Type 2 hairs only. Fruiting calyx with lobes

exceeding mature fruit, prickles absent. Mature

fruits 1 or 2 per inflorescence, globular, 5.5-8

mm diameter, red, 1-locular (septum absent or

incomplete); placenta not apparent; mesocarp

juicy, succulent; exocarp 0.3-0.5 mm thick;

pedicels 10-16 mm long in fruit, 0.8-0.9 mm

thick at mid-point; seeds pale yellow, 2.2-2.8

mm long. Fig. 14.

Specimens examined : Queensland. Leichhardt District:

‘(Tottenham’, c. 10 miles [16 km] NW of Banana, May 1960,

Johnson 1708, 1709 (BRI); ‘The Rhyddings’, c. 38 km SW

of Moura, Aug 1962, Johnson 2473 (BRI); site of Brigalow

Research Station, 20 miles [32 km] NW of Theodore, Apr

1963, Johnson 2621 (BRI, CANB); c. 4 miles [6 km] E of

Moura, Mar 1967, Henderson 223 (BRI). Port Curtis

District: Orange Creek, c. 20 miles [32 km] NW of Biloela,

Jun 1959, Johnson 854 (BRI); 1.6 km along Hibbs Road, N

of Jambin, Apr 2003, Bean 20225 (BRI).

Distribution and habitat: Solanum

johnsonianum is endemic to Queensland. It

extends from north-west of Theodore to north

of Biloela (Map 3), a distance of around 100

kilometres. All specimens were found in

communities dominated or co-dominated by

Acacia harpophylla (Brigalow), on heavy

cracking clay soils. In some cases, the specimen

labels say “recently burnt” or “recently cleared

Brigalow scrub”.

Phenology : Flowers recorded for March, May,

June and August; mature fruits recorded for

April and May.

Notes: S. johnsonianum is distinguished from

S. nemophilum by the branchlet stellae with a

central ray 3-5 times longer than the lateral

rays (0.7-1.2 times for S. nemophilum ), central

ray similarly much longer on other plant parts,

stellae 0.25-0.35 mm diameter on upper leaf

surface (0.4-0.7 mm for S. nemophilum ), upper

leaf surface lacking Type 2 hairs and stellae

without stalks (Type 2 hairs present, stellae

stalks 0.1-0.3 mm long for S. nemophilum).

S. johnsonianum is also closely related to

S. innoxium. It differs from the latter by the

broader leaves (1.4-2 times longer than wide),

the longer petioles (9-28 mm long), the greater

petiole/lamina ratio (30-55%), the larger stellae

on the lower leaf surface (0.3-0.6 mm diameter)

and the calyx stellae with the central ray often

gland-tipped.

Conservation status: Solanum johnsonianum

can no longer be found at most of the localities

cited above. It is currently known from 3

locations, including the Brigalow Research

Station, where the species is under threat from

abnormally intensive grazing by wallabies. The

other two populations are threatened by weeds

(pasture grasses) and land clearance. No

population is protected within a conservation

reserve. Applying the IUCN guidelines (IUCN.

2001), a category of “Endangered” is

recommended (EN A3c; B lab(iii,v)+2ab(iii,iv,v);

Cl).

Etymology: Named for Robert W. Johnson,

former Director of the Queensland Herbarium,

and an assiduous collector of solanums,

including almost all the specimens of this

species.

5. Solanum innoxium A.R.Bean sp. nov. Frutex

usque ad 0.6 m altus, aculeis carens; folia

integra, lanceolata, indumento coacto;

stellae sessiles, radio centrali radiis

lateralibus 3-5plo longiore, in pagina

superiore folii 0.2-0.25 mm diametro;

fructus ellipsoidales, 3-4.5 mm diametro,

succosi et laete rubri maturitate; calycis

lobi elliptici, fructus superantes. Typus:

Queensland. Maranoa District: slopes of

Bean, Taxonomy of Solanum subg. Leptostemonum

679

Fig. 15. Solanum innoxium. A. flowering branchlet x 1.5. B. style and ovary x 6. D. ovary showing Type 2 hairs x 12. D.

mature fruit still attached x 4. E. mature fruit, detached x 5. F. transverse section of fruit x 8. A, Pedley 792; B-C, Bean

17775; D-F , Bean 18367.

Thomby Range, ‘Glen Fosslyn’, SE of

Surat, 31 August 2001, A.R. Bean 17775

(holo: BRI (1 sheet + spirit); iso: MEL,

NSW).

Erect, rhizomatous perennial shrub, 0.3-0.6 m

high. Leaves (in outline) lanceolate, entire;

without prickles on upper surface. Adult stem

prickles absent. Adult branchlets yellow or

brown; prickles absent; stellae very dense,

0.25-0.4 mm diameter, sessile; lateral rays 7

or 8, porrect; central ray 3-4 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Adult leaves lanceolate or

ovate, entire; la min a 2.5-7 cm long, 0.8-2 cm

wide, 2.6-4.2 times longer than broad, apex

obtuse or acute, base obtuse or cordate, oblique

part 0-2 mm long, obliqueness index 0-5

percent; petioles 0.4-0.8 cm long, 11-19%

length of lamina, prickles absent. Upper leaf

surface grey-green; prickles absent; stellate

hairs distributed throughout; protostellae

absent; ordinary stellae density moderate to

dense, 0.1-0.15 mm apart, 0.2-0.3 mm across,

sessile; lateral rays 7 or 8, porrect; central ray

3-6 times as long as laterals, not gland-tipped;

680

finger hairs absent; Type 2 hairs absent. Lower

leaf surface white or yellowish; prickles absent;

stellae very dense, 0.05-0.1 mm apart,

0.25-0.4 mm diameter, stalks 0-0.1 mm long;

lateral rays 7 or 8, porrect; central ray 3-5 times

as long as laterals, not gland-tipped; finger

hairs absent; Type 2 hairs absent. Inflorescence

supra-axillary, solitary or pseudo-umbellate,

common peduncle absent; 1 or 2-flowered, with

all flowers bisexual and 4 or 5-merous; pedicels

5-11 mm long at anthesis, same thickness

throughout, 0.7-0.8 mm thick at mid-point,

prickles absent. Calyx tube 1-1.5 mm long,

lobes elliptic, 4-5.5 mm long; prickles absent

at anthesis; stellae very dense, transparent,

0.25-0.4 mm across, sessile, lateral rays 7 or

8, central ray 2.5-5 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Corolla mauve or purple, 7-9 mm

long, deeply lobed, inner surface glabrous;

anthers 3-4.5 mm long; ovary with Type 2 hairs

only, or with stellate and Type 2 hairs;

functional style 4-5 mm long, erect, glabrous

or with stellate and Type 2 hairs, stellae

0.2-0.25 mm across, lateral rays 7 or 8, central

ray c. 1 times as long as laterals. Fruiting calyx

with lobes exceeding mature fruit, prickles

absent. Mature fruits 1 per inflorescence,

ellipsoidal, 3-4.5 mm diameter, red, with a few

scattered Type 2 hairs, 1-locular (septum absent

or incomplete); placenta not apparent;

mesocarp juicy, succulent; exocarp 0.2-0.3 mm

thick; pedicels 7-11 mm long in fruit, 0.7-0.8

mm thick at mid-point; seeds pale yellow,

2.2-2.3 mm long. Fig. 15.

Specimens examined : Queensland. Maranoa District: E

of Combididan Farm, ‘Cypress Downs’ [NE of Yuleba], Sep

1961, Jones 166 (BRI); ‘Boxleigh’, S of Surat, Aug 2001,

Bean 17762 (BRI, MEL, NSW); Thomby Range, SW of

Glenmorgan, Jan2002, Bean 18367 (BRI); ‘Silver Springs’,

S of Surat, Jan 2002, Bean 18374 (BRI). Darling Downs

District: ‘Calala’, c. 10 miles [16 km] E of Meandarra, Jun

1960, Johnson 1620A(BRI); ‘Woodlands’, 5 mi les [8 km]

SW of Westmar, Jul 1961, Pedley 792 (BRI, CANB);

Hannaford, 42 km W of Tara, Apr 1963, T.J. McDonald 63

(BRI, CANB).

Distribution and habitat : Solanum innoxium

is endemic to Queensland, where it extends

from near Yuleba to Westmar (Map 3). Habitat

varies from ridges dominated by Lancewood

(Acacia shirleyi ) or Bendee (Acacia

catenulata ), to Eucalyptus populnea woodland,

and one record from an Acacia harpophylla

Austrobaileya 6 (4): 639-816 (2004)

community. Soils also vary from shallow stony

loams, to red-brown clay loams, or heavy grey

clays.

Phenology : Flowers are recorded for January,

and from June to September; mature fruits

recorded for January and April.

Notes: S. innoxium is allied to S. nemophilum

but differs by the lanceolate leaves, the stellae

of the upper leaf surface 0.2-0.3 mm diameter

(0.4-0.7 mm for S. nemophilum), the branchlet

stellae with the central ray 3-4 times longer

than the lateral rays (0.7-1.2 times for

S. nemophilum ), the stellae on the lower leaf

surface sessile or with stalks to 0.1 mm long

(stalks 0.2-0.4 mm for S. nemophilum ), style

4-5 mm long (5.5-8 mm long for

5. nemophilum) and the ellipsoidal fruits

(globular for S. nemophilum ). S. innoxium

differs from S. hapalum by the shorter and

narrower leaves, shorter petioles, smaller stellae

on the lower leaf surface, shorter style, the

relatively long Type 2 hairs on the ovary, and

the ellipsoidal fruits.

Conservation status: Solanum innoxium is

currently known from 3 locations. It does not

occur in a conservation reserve. It is threatened

by land clearance and weeds. Applying the

IUCN guidelines (IUCN. 2001), a category of

“Vulnerable” is recommended (VU A3ce; Cl).

Etymology: From the Latin innoxius, meaning

‘harmless’. This is in reference to the lack of

prickles on plants of this species.

6. Solanum ultimum A.R.Bean sp. nov. Frutex

usque ad 0.6 m altus, aculeis carens; folia

integra, ovata; stellae sessiles, radio

centrali radiis lateralibus 0.4-lplo

longiore, in pagina inferiore folii 0.4-0.6

mm diametro, in pagina superiore folii

indumento absente usque ad moderate

denso; fructus globulares, 6-9 mm

diametro, succosi et laete rubri maturitate;

calycis lobi elliptici, fructus superantes.

Typus: Queensland. Burke District: 3

km by road south then west of ‘Warang’

homestead site, White Mountains

National Park, 11 April 2000, M.B.

Thomas 1572 & E.J. Thompson (holo:

BRI; iso: DNA, NSW).

681

Bean, Taxonomy of Solanum subg. Leptostemonum

Fig. 16. Solanum ultimum. A. flowering branchlet x 1. B. style and ovary x 6. C. ovary showing Type 2 hairs x 20. D. stellate

hair from upper leaf surface x 60. E. mature fruit x 2. all from Cottam AZI 1520.

682

Austrobaileya 6 (4): 639-816 (2004)

Solanum nemophilum var. brachycarpum

Domin, Biblioth. Bot. 89: 585 (1929).

Type: Queensland. North Kennedy

District: near Pentland, March 1910, K.

Domin (holo: PR).

Erect, rhizomatous perennial shrub, 0.3-0.6 m

high. Juvenile stage unknown. Adult stem

prickles absent. Adult branchlets white, grey

or yellow; prickles absent; stellae very dense,

0.6-0.9 mm diameter, stalks 0-0.1 mm long;

lateral rays 7-9, porrect; central ray 0.4-0.8

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Adult

leaves lanceolate or ovate, entire; lamina 4-6

cm long, 1.1-2.3 cm wide, 2.7-3.6 t im es longer

than broad, apex obtuse or acute, base cuneate

or obtuse, oblique part 0-1.5 mm long,

obliqueness index 0-2 percent; petioles

0.7-1.4 cm long, 16-23% length of lamina,

prickles absent. Upper leaf surface green;

prickles absent; stellate hairs confined to

midrib, or distributed throughout; protostellae

present; ordinary stellae very sparse to sparse,

0.4-4 mm apart, 0.25-0.4 mm across, sessile;

lateral rays 7 or 8, porrect; central ray 0.5-1

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Lower

leaf surface white; prickles absent; stellae dense

to very dense, 0.1-0.2 mm apart, 0.4-0.6 mm

diameter, stalks 0-0.1 mm long; lateral rays

8-9, porrect; central ray 0.2-0.8 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Inflorescence

supra-axillary, solitary or pseudo-umbellate,

common peduncle absent; 1-3-flowered, with

all flowers bisexual and 4 or 5-merous; pedicels

5-9 mm long at anthesis, same thickness

throughout or markedly thicker distally, 0.9-1

mm thick at mid-point, prickles absent. Calyx

tube 0.5-1.5 mm long, lobes elliptic, 4-6 mm

long; prickles absent at anthesis; stellae dense

to very dense, white, 0.4-0.5 mm across, stalks

0-0.1 mm long, lateral rays 8-9, central ray

0.4-0.8 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Corolla purple, 7-9 mm long, shallowly lobed,

inner surface glabrous; anthers 3.5-4.5 mm

long; ovary with Type 2 hairs only; functional

style 5.5-7 mm long, erect, glabrous or with

Type 2 hairs only. Fruiting calyx with lobes

exceeding mature fruit, prickles absent. Mature

fruits 1 or 2 per inflorescence, globular, 6-9

mm diameter, red; mesocarp juicy, succulent;

pedicels 7-12 mm long in fruit, 1.5-1.9 mm

thick at mid-point; seeds pale yellow, 2.4-2.7

mm long. Fig. 16.

Specimens examined : Queensland. Burke District: White

Mountains N.P., near ‘Warang’, Mar 2000, Wannan 1624

(BRI, NSW); 0.5 km W of ‘Warang’ HS site, White

Mountains N.R, Apr 2000, Thomas 1803 & Thompson

(BRI). Mitchell District: ‘Narbethong’ SE of Barcaldine,

Jun 1997, Cottam AZI1517 (BRI); ‘Narbethong’, SE of

Barcaldine, Dec 1997, Milson AZI1537 (BRI).

Distribution and habitat: Solanum ultimum is

endemic to Queensland. It is known from the

White Mountains National Park and from near

Barcaldine (Map 3). It grows on red sandy soil

with Corymbia sp., Eucalyptus persistens or

Melaleuca tamariscina.

Phenology: Flowers are recorded for April,

June and December; mature fruits in April and

December.

Notes: S. ultimum differs from S. nemophilum

by the indumentum of the upper leaf surface

absent to sparse (indumentum moderately dense

to dense for S. nemophilum ), upper leaf surface

with stellate hairs sessile and Type 2 hairs

absent (stellate hairs with stalks 0.1-0.3 mm

long and Type 2 hairs present for

S. nemophilum ), calyx stellate hairs with a

central ray 0.4-0.8 times length of lateral rays

(0.8-1.2 times for S. nemophilum ), and the

larger seeds.

S. ultimum differs from all other species

in the S. densevestitum group by the upper leaf

surface with the stellate indumentum absent or

at most moderately dense, and by the stellae

with a comparatively short central ray on all

plant parts.

Conservation status: Data deficient.

Etymology: From the Fatin ultimus, meaning

‘most distant’. This refers to the geographical

remoteness from the eastern Australian coast,

where its relatives are located.

7. Solanum densevestitum F. Muell. ex Benth.,

FI. Austral. 4: 456 (1868). Type:

Queensland. ‘Brisbane River’, December

1856, F. Mueller (lecto: MEF

[MEF12200]), fide Symon (1981).

Bean, Taxonomy of Solanum subg. Leptostemonum

Erect, rhizomatous perennial shrub, 0.4-0.9 m

high. Juvenile branchlets without prickles;

leaves (in outline) broadly ovate, shallowly-

lobed, with 2-3 pairs of lobes; lamina c. 9x6

cm, without prickles on upper surface. Adult

stem prickles absent. Adult branchlets grey;

prickles absent; stellae dense to very dense,

0.7-1.5 mm diameter, stalks 0-2 mm long;

lateral rays 4-6, porrect; central ray 1.5-4 times

as long as laterals, not gland-tipped; finger

hairs absent; Type 2 hairs absent. Adult leaves

ovate or broadly ovate, entire; lamina 5-10 cm

long, 2.6-5.5 cm wide, 1.6-2 times longer than

broad, apex acute, base obtuse or cordate,

oblique part 0-5 mm long, obliqueness index

0-5 percent; petioles 0.9-3.4 cm long, 17-35%

length of lamina, prickles absent. Upper leaf

surface green; prickles absent; stellate hairs

distributed throughout; protostellae present;

ordinary stellae dense, 0.2-0.4 mm apart,

0.5-1.1 mm across, sessile; lateral rays 4-6,

porrect; central ray 3-6 t im es as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Lower leaf surface green; prickles

absent; stellae dense, 0.1-0.3 mm apart,

0.7-1.6 mm diameter, stalks 0-0.5 mm long;

lateral rays 4-8, porrect; central ray 2-5 times

as long as laterals, not gland-tipped; finger

hairs absent; Type 2 hairs absent. Inflorescence

supra-axillary, solitary or pseudo-umbellate,

common peduncle 0-4 mm long; 1-3-flowered,

with all flowers bisexual and 5-merous; pedicels

4-9 mm long at anthesis, same thickness

throughout, 0.5-0.6 mm thick at mid-point,

prickles absent. Calyx tube 1.5-2.5 mm long,

lobes elliptic, 6-10 mm long; prickles absent

at anthesis; stellae dense, transparent, 0.8-1.8

mm across, stalks 0-0.9 mm long, lateral rays

4 or 5, central ray 1.5-3 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Corolla purple, 10-13 mm

long, shallowly lobed, inner surface glabrous;

anthers 4-5.5 mm long; ovary with stellate and

Type 2 hairs; functional style 6-7.5 mm long,

erect, with Type 2 hairs only. Fruiting calyx

with lobes exceeding mature fruit, prickles

absent. Mature fruits 1 or 2 per inflorescence,

globular, c. 7 mm diameter, red, 1-locular

(septum absent or incomplete); placenta not

apparent; mesocarp juicy, succulent; pedicels

9-12 mm long in fruit; seeds pale yellow,

2-2.2 mm long. Felty Nightshade.

683

Selected specimens : Queensland. South Kennedy District:

S of Massey Creek, Eungella N.P., Sep 1991, Bean 3682

(BRI); Eungella Range, Schumanns Road, c. 1.1 km E of

Swampy Ridge, Jun 1995, Pollock 233 (BRI). Burnett

District: 14 km ESE of Elgin Yale, Jan 1991, Pedley 5596

(BRI). Wide Bay District: Lagoon Pocket via Gympie, Aug

1930, Lowe s.n. (BRI); Elanda Point, near Lake Cootharaba,

Jun 1986, Sandercoe C996 & Milne (BRI); S.F.639 Wrattens,

Apr 1996, Forster 19136 & Leiper (AD, BRI); Booloumba

Creek road, SW of Kenilworth, Aug 1999, Bean 15246 (BRI,

NSW); S.F.256 Imbil, Mitchell L.A., Mar 2003, Forster

PIF29294 et al. (BRI, MEL, NE, NSW). Moreton District:

Enoggera, Mar 1912, White s.n. (BRI); Blackall Range, Dec

1918, White s.n. (BRI); near Samford, on House Mtn, Feb

1945, Blake 15506 (BRI); Bruce Hwy, Caboolture, Jun 1960,

Williamson s.n. (BRI); Benarkin S.F. near Blackbutt, Jul

1974, Moriarty 1542 (BRI, CANB); Maiala Reserve, near

Mt Glorious, Jun 1989, Symon 14881 (AD, BRI); Mt

Perserverance S.F.757, Jul 1996, Stephens ESK28 &

Dowling (BRI).

Distribution and habitat : Solanum

densevestitum is endemic to Queensland. From

the north-western outskirts of Brisbane to

Gympie and west to Gallangowan, plus a

remarkable outlier at Eungella, west of Mackay

(Map 2). It inhabits dense eucalypt forest with

a shrubby understorey of rainforest species,

often in areas that have been disturbed e.g. by

roadworks.

Phenology: Flowers are recorded for all months

of the year; mature fruits are recorded between

April and August.

Notes: Solanum densevestitum differs from all

other related species by the stellae on upper

leaf surface being sessile, with only 4 or 5

(rarely 6) lateral rays; the long-stalked stellae

of the branchlets (stalks up to 2.0 mm long);

and the leaf lower surface green.

The collections cited by Bentham in the

protologue belong to three species;

S. densevestitum (as to lectotype), the recently

named S. stupefactum Symon, and S. hapalum

(described in this paper). The description given

in the protologue was clearly derived from a

combination of these taxa.

The isolectotype cited by Symon (1981)

is a mixed specimen. There are three specimens

on the sheet; the lowermost piece is

S. densevestitum , as to lectotype; the upper two

pieces are S. stupefactum Symon.

The locality of “Tweed River district”

given for a collection by E. Betche in April 1896

684

(held at NSW) must be regarded with

considerable suspicion, as there have been no

subsequent collections from there or anywhere

nearby.

Typical specimens of S. densevestitum

and S. nemophilum are readily distinguished

by the stellae of the upper leaf surface (stalk

length and lateral ray number). However,

collections from the Benarkin - Yarraman area

appear to be morphologically intermediate and

difficult to assign to either species.

Juvenile leaves of S. densevestitum are

frequently lobed, in a similar fashion to adult

leaves of S. gympiense.

Conservation status: Moderately widespread.

Not considered at risk.

8. Solanum nemophilum F.Muell., Fragm. 2:

161 (1861); S. nemophilum var.

nemophilum Domin, Biblioth. Bot. 89:

584 (1929); S. nemophilum var. typicum

Domin, Biblioth. Bot. 89: 584 (1929),

nom. inval. Type: [Queensland.

Leichhardt District:] ‘between

Mackenzie and Dawson Rivers’,

November 1856, F. Mueller (lecto: MEL;

isolecto: K, photo seen).

Erect, rhizomatous perennial shrub, 0.3-0.6 m

high. Juvenile branchlets with 0-5 prickles per

dm, 2-3 mm long; leaves (in outline) ovate,

entire; lamina c. 4x2 cm, with 0-4 prickles

on upper surface. Adult stem prickles absent.

Adult branchlets white or grey; prickles absent;

stellae very dense, 0.3-1 mm diameter, stalks

0-0.6 mm long; lateral rays 7 or 8, porrect;

central ray 0.7-1.2 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Adult leaves ovate, entire; lamina

2.5-7 cm long, 1.2-2.8 cm wide, 1.8-3 times

longer than broad, apex obtuse or acute, base

obtuse or cordate, oblique part 0-3 mm long,

obliqueness index 0-5 percent; petioles 0.4-1.4

cm long, 15-35% length of lamina, prickles

absent. Upper leaf surface grey-green; prickles

absent; stellate hairs distributed throughout;

protostellae absent; ordinary stellae dense, 0.1-0.3

mm apart, 0.4-0.7 mm across, stalks 0.1-0.3

mm long; lateral rays 7 or 8, porrect; central

ray 0.7-1.8 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Austrobaileya 6 (4): 639-816 (2004)

Lower leaf surface green or yellowish; prickles

absent; stellae dense to very dense, 0.05-0.15

mm apart, 0.5-0.7 mm diameter, stalks 0.2-0.4

mm long; lateral rays 7 or 8, porrect; central

ray 0.5-1.2 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Inflorescence supra-axillary, pseudo-umbellate,

common peduncle 0-3 mm long; 3-5-flowered,

with all flowers bisexual and 4 or 5-merous;

pedicels 4-7 mm long at anthesis, same

thic kn ess throughout, 0.9-1 mm thick at mid¬

point, prickles absent. Calyx tube 1-2 mm long,

lobes elliptic, 2.5-7 mm long; prickles absent

at anthesis; stellae very dense, white, 0.4-1 mm

across, stalks 0-0.25 mm long, lateral rays 7

or 8, central ray 0.8-1.2 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Corolla purple, 7-10 mm

long, rotate or shallowly lobed, inner surface

glabrous; anthers 3-4 mm long; ovary with

Type 2 hairs only; functional style 5.5-8 mm

long, erect, with Type 2 hairs only. Fruiting

calyx with lobes exceeding mature fruit,

prickles absent. Mature fruits 1 per

inflorescence, globular, 4.5-9 mm diameter,

red, glabrous or with a few scattered Type 2

hairs, 1-locular (septum absent or incomplete);

placenta not apparent; mesocarp juicy,

succulent; exocarp 0.2-0.25 mm thick; pedicels

6-11 mm long in fruit, 0.8-1.2 mm thick at

mid-point; seeds pale yellow, 2-2.3 mm long.

Selected specimens examined : Queensland. Leichhardt

District: Duaringa, Mar 1909, Maiden (NSW); Blackdown

Tableland, c. 35 km SE of Blackwater, Sep 1971, Henderson

H1094 et al. (BRI, CANB); top of Carnarvon Range, N of

Injune, Sep 2003, Bean 20760 (BRI). Port Curtis District:

S.F.69 Dawes Range, SE of Biloela, Dec 1999, Bean 15933

(BRI). Burnett District: c. 14 km NNE of Eidsvold, Oct

1993, Lepschi & Slee 1215 (AD, BRI, CANB); c. 9.2 km

NE of Durong on slopes above Hardy Creek, Jul 1998,

Pollock ABP715 & Dean (BRI); Schellbachs road, Kingaroy,

Mar 1999, Haskard 17 (AD, BRI); Hurdle Gully road,

Coominglah S.F., W of Monto, Jan 2000, Bean 15934 (BRI).

Darling Downs District: 6 miles [10 km] S of Warwick on

Stanthorpe road, Nov 1946, Everist & Webb 1258 (BRI); 25

miles [40 km] WNW of Dalby on road to Kogan, Oct 1969,

Everist s.n. (BRI); Amiens, 10 miles [16 km] NW of

Stanthorpe, Jul 1974, Swan 71 (BRI); Centenary road, W of

Goombungee, May 1997, Bean 12000 (BRI); S.F. 197

Diamondy, 32 km NE of Jandowae, Mar 1999, Forster

PEF24090 & Booth (AD, BRI, MEL); Bony Mountain, 14

km SW of Allora, Aug 1999, Bean 15259 (BRI); Wondul

Range N.P., SWofMilmerran,Nov 1999, Bean 15888 (BRI).

Moreton District: Tarampa, undated, Bailey (BRI); Falls

Creek, 4.5 km NW of West Haldon, May 1987, Forster 2940

& Bird (BRI); Scanlon scrub, Mount Berryman, 15 km SW

of Laidley, Aug 1990, Bird (BRI); near High Camp, 11 km

SE of Cooyar, Feb 2000, Bean 16048 (BRI).

Bean, Taxonomy of Solanum subg. Leptostemonum

Distribution and habitat: Solanum nemophilum

is endemic to Queensland. Widespread in

subcoastal areas of Queensland from Duaringa

to Boonah and Warwick, and west to Kogan

and Miles (Map 4). It grows in shrubby

eucalypt woodland in loamy soils.

Phenology: Flowers and fruits may be borne

at any time of the year.

Notes: S. nemophilum is distinguished from

other species of this group by the relatively

long-stalked stellae on both leaf surfaces, and

the stellae with a central ray more or less equal

in length to the lateral rays.

A Scortechini specimen of S. nemophilum

at BRI, reputedly from Roma, should be

disregarded, as there have been no collections

of this species from the Maranoa district over

the last century.

Conservation status : Widespread. Not

considered at risk.

9. Solanum gympiense Symon, Austrobaileya

4: 433 (1995). Type: Queensland. Wide

Bay District: Gundiah, 21 June 1927,

C.T. White 3527 (holo: BRI).

Solanum sp. 2 in Ross (1986)

Illustration: Symon (1995: 434)

Erect, rhizomatous perennial shrub, 0.2-0.5 m

high. Leaves (in outline) broadly ovate,

shallowly-lobed, with 3 or 4 pairs of lobes;

lamina c. 6x4 cm, without prickles on upper

surface. Adult stem prickles absent. Adult

branchlets yellow or brown or green; prickles

absent; stellae dense to very dense, 0.5-1 mm

diameter, stalks 0-0.8 mm long; lateral rays

6-8, porrect or ascending; central ray 1-2 t im es

as long as laterals, gland-tipped; finger hairs

absent; Type 2 hairs absent. Adult leaves ovate

or broadly ovate, entire or shallowly lobed

throughout; lobes 3-5 on each side, obtuse,

lobing index 1-1.2; lamina 4—10 cm long, 2.2-5.5

cm wide, 1.4-1.9 times longer than broad, apex

obtuse or acute, base obtuse or cordate, oblique

part 0-3 mm long, obliqueness index 0-4

percent; petioles 0.8-2.3 cm long, 17-30%

length of lamina, prickles absent. Upper leaf

surface green or grey-green; prickles absent;

685

stellate hairs distributed throughout;

protostellae present; ordinary stellae dense,

0.2-0.3 mm apart, 0.5-0.9 mm across, stalks

0-0.15 mm long; lateral rays 5-10, porrect or

ascending; central ray 2-4 times as long as

laterals, gland-tipped; finger hairs absent; Type

2 hairs absent. Lower leaf surface white; prickles

absent; stellae dense to very dense, 0.1-0.25 mm

apart, 0.8-1.3 mm diameter, stalks 0.1-0.5 mm

long; lateral rays 5-11, porrect or ascending;

central ray 0.8-2 times as long as laterals,

gland-tipped; finger hairs absent; Type 2 hairs

absent. Inflorescence supra-axillary, pseudo-

racemose, co mm on peduncle 4-21 mm long,

rachis prickles absent; 3-7-flowered, weakly

andromonoecious or with all flowers bisexual

and 4 or 5-merous; pedicels 7-12 mm long at

anthesis, same thickness throughout, 0.7-1.2

mm thick at mid-point, prickles absent. Calyx

tube 1.5-3 mm long, lobes elliptic, 2.5-9 mm

long; prickles absent at anthesis; stellae very

dense, transparent, 0.7-1 mm across, stalks

0-0.7 mm long, lateral rays 7 or 8, central ray

1-3 times as long as laterals, gland-tipped;

finger hairs absent; Type 2 hairs absent. Corolla

mauve or purple, 11-15 mm long, deeply lobed,

inner surface glabrous; anthers 4.5-6 mm long;

ovary glabrous; functional style 6-11 mm long,

erect, glabrous or with Type 2 hairs only or with

stellate and Type 2 hairs, stellae c. 0.5 mm

across, lateral rays c. 7, central ray c. 1.5 times

as long as laterals. Fruiting calyx with lobes

exceeding mature fruit, prickles absent. Mature

fruits 1-4 per inflorescence, ellipsoidal, 4.5-7

mm diameter, red, 1-locular (septum absent or

incomplete); placenta not apparent; mesocarp

juicy, succulent; exocarp 0.2-0.3 mm thick;

pedicels 7-15 mm long in fruit, 1.2-1.6 mm

thick at mid-point; seeds pale yellow, 2.7-2.9

mm long.

Specimens examined : Queensland. Leichhardt District:

crest of Carnarvon Range, Mar 1960, Johnson 1451 (BRI).

Port Curtis District: Rosedale, Mar 1931, Dovey K5 (BRI).

Maranoa District: ‘Claravale’, May 1962, Johnson 2440

(BRI); The Tombs, Maranoa River, c. 110 km NW of Injune,

Jun 1977, Crisp 3113 (AD, BRI, CANB); 1.5 km SW of

‘Kilmorey’ HS, May 1982, Neldner & Thomas 627, 633

(BRI). Wide Bay District: Woondum, Oct 1917, Moore

(BRI); Gundiah, Jun 1927, White 3528 (BRI); c. 6 miles [10

km] NW of Tiaro, Apr 1959, Ridley s.n. (BRI); 8 km SSW

of Howard, Jan 1987, Forster 2855 (AD, BRI); Clifton

Range, S.F. 676,11 kmN of Brooweena, Apr 1992, Forster

PIF9694 (BRI); Veteran S.F., N of Gympie, Oct 1993, Bean

6706 (BRI); S.F.832, Cordalba S.F., c. 19.5 km SE of Gin

Gin, Nov 1995, Sparshott KMS660 (BRI); S.F. 1294 Parish

686

Austrobaileya 6 (4): 639-816 (2004)

of Doongul, c. 27 km S of Childers, May 1996, Sparshott

KMS837 & Shewell (AD, BRI); c. 3 km SE of Fairlies Knob,

Seaview Range, May 2000, Phillips 391 (BRI); S.F.940,10

km SW of Bauple, Mar 2002, Bean 18555 (A, BRI, CANB,

K, MEF, MO, PRE).

Distribution and habitat: Solanum gympiense

is endemic to Queensland where it extends

along the coast from Woondum (near Gympie)

to Rosedale (west of Bundaberg), with disjunct

occurrences in the Carnarvon Range north of

Injune (Map 5). It grows in shrubby eucalypt

forest in sandy to loamy soil.

Phenology: Flowers and fruits have been

recorded for almost every month of the year.

Notes: S. gympiense is distingushed from other

species in the group by the stellae with gland-

tipped central rays on most plant parts, and the

adult leaves usually lobed.

Conservation status: Widespread. Not

considered at risk.

Group 13 (S. ferocissimum group) of Whalen

(1984)

Perennial shrubs (100%); calyx and pedicel

prickles absent, calyx lobes not elliptic (100%);

large stems with prickles (100%); prickles

acicular (100%); upper leaf surface green; fruits

globular, red, juicy, 1-locular (100%); fruiting

calyx less than half length of mature fruit

(97%); inflorescence not branched (97%);

corolla inner surface glabrous (93%); fruit

exocarp <1 mm thick (93%); adult leaves entire

or shallowly lobed (88%); fruits <12 mm

diameter (80%); branchlet prickles present

(77%); Type 2 hairs absent from branchlets

(77%); average petiole length 12% of lamina

length.

17 species endemic to Australia; 17

species occurring in Queensland.

10. Solanum fervens A.R.Bean sp. nov.

Fructus erectus, aculeos aciculares sparse

distributes gerens; folia integra vel non

profunde lobata, lanceolata, supra glabra,

subtus dense stellato-tomentosa; stellae

radio centrali radiis lateralibus 3-6plo

longiore; ovarium stellato-pubescens;

exocarpium fructuum maturorum c. 0.8

mm crassum. Typus: Queensland. Cook

District: eastern bank of Jardine River

mouth, 1 September 1985, J.R. Clarkson

6219 (holo: BRI (1 sheet + spirit); iso:

AD, MBA, QRS).

Solanum sp. (Bamaga V. Scarth-Johnson

1117) in Henderson (2002)

Erect, rhizomatous perennial shrub, 1-2 m

high. Juvenile stage unknown. Adult branchlets

yellow or brown; prickles 3-20 per decimetre,

straight, acicular, 3-5 mm long, 13-16 times

longer than wide; stellae dense to very dense,

0.3-0.4 mm diameter, sessile; lateral rays 7 or

8, porrect; central ray 2-6 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Adult leaves lanceolate or

ovate, entire or shallowly lobed throughout;

lobes 3 or 4 on each side, obtuse, lobing index

1-1.2; lamina 7.5-14 cm long, 2-4.5 cm wide,

2.5-5.2 t im es longer than broad, apex acute,

base cuneate, oblique part 0-4 mm long,

obliqueness index 0-4 percent; petioles 0.4-0.8

cm long, 3-7% length of lamina, prickles

absent. Upper leaf surface green; prickles

absent; stellate hairs absent, or confined to

midrib; ordinary stellae absent from surface;

finger hairs absent; Type 2 hairs absent. Lower

leaf surface yellowish or rusty; prickles absent;

stellae dense, 0.1-0.2 mm apart, 0.4-0.5 mm

diameter, sessile; lateral rays 7 or 8, porrect;

central ray 3-7 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Inflorescence supra-axillary, pseudo-

racemose, common peduncle 0-3 mm long,

rachis prickles absent; 7-20-flowered, with all

flowers bisexual and 5-merous; pedicels 4-7

mm long at anthesis, same thickness

throughout, 0.25-0.4 mm thick at mid-point,

prickles absent. Calyx tube 1.5-2 mm long,

lobes deltate, 0.5-1.2 mm long; prickles absent

at anthesis; stellae dense to very dense, yellow

or white, 0.2-0.3 mm across, sessile, lateral

rays 5-7, central ray 2-3 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Corolla mauve, 5-7 mm

long, deeply lobed, inner surface glabrous;

anthers 3-4.5 mm long; ovary with stellate

hairs only; functional style 5-7 mm long, erect,

with stellate hairs only, stellae 0.2-0.4 mm

across, lateral rays 5-7, central ray 0.5-1 times

as long as laterals. Fruiting calyx with lobes

less than half length of mature fruit, prickles

absent. Mature fruits 1-3 per inflorescence,

globular, 9-12 mm diameter, red, 1-locular

Bean, Taxonomy of Solanum subg. Leptostemonum

687

Fig. 17. Solanum fervens . A. flowering branchlet x 0.6. B. style and ovary x 6. C. ovary showing stellate hairs x 30. D.

stellate hair from lower leaf surface x 60. E. mature fruit x 2. F. transverse section of fruit x 3. A-D, Forster 8968; E-F,

Clarkson 6219.

688

Austrobaileya 6 (4): 639-816 (2004)

(septum absent or incomplete); placenta not

apparent; mesocarp juicy, succulent; exocarp

c. 0.8 mm thick; pedicels 15-23 mm long in

fruit, 0.5-0.8 mm thick at mid-point; seeds pale

yellow, 3-3.5 mm long. Fig. 17.

Specimens examined : Queensland. Cook District: N of

Olive River, Cape York Peninsula, Sep 1974, Webb & Tracey

13480 (BRI, QRS); tributary of Escape River, Jun 1978,

Clarkson 2086 (BRI); Red Island Point, Bamaga, Sep 1980,

Scarth-Johnson 1117A (BRI); 3 km NW of Bolt Head,

Temple Bay, Jul 1991, Forster PIF8968 (AD, BRI, DNA,

QRS); 15 km N of Middle Peak road junction, Mar 1992,

Johnson 5114 (BRI); 0.4 km west of Sach Waterhole, c. 14

km NNW of Ussher Point, Jul 1992, Clarkson 9679 &

Neldner (AD, BRI, MBA); Bolt Head, Jun 1996, Gray 6842,

6875 (BRI, QRS).

Distribution and habitat : Solanum fervens is

endemic to Queensland. Confined to the Cape

York Peninsula, from the Bamaga area to Bolt

Head and Temple Bay (Map 2). It grows on

siliceous sand dunes in semi-deciduous vine-

forest.

Phenology: Flowers have been recorded in

March, June and July; mature fruits in March,

June and September.

Notes: S. fervens is closely related to

S. discolor, but differs by the dense

indumentum on the leaf underside (very dense

for S. discolor ), the stellate hairs of the calyx

and leaf underside having a much longer

central ray, the longer hypanthium, the stellate

pubescent ovary (glabrous for S. discolor ), and

the exocarp c. 0.8 mm thick (0.4-0.5 mm thick

for S. discolor ).

Conservation status: Not considered at risk.

Etymology: From the Latin fervens, meaning

“burning, boiling, hot”. This is in reference to

the high temperatures experienced by the

species in the extreme north of Australia.

11. Solanum yirrkalense Symon, J. Adelaide

Bot. Gard. 4: 137 (1981), as ‘yirrkalensis’.

Type: Northern Territory. Yirrkala

Gardens, 27 February 1976, D. Hinz 7633

(holo: DNA ex NT; iso: BRI, CANB,

DNA).

Illustration: Symon (1981: 139)

Erect, rhizomatous perennial shrub, 0.5-1.5 m

high. Juvenile stage unknown. Adult branchlets

brown; prickles 3-10 per decimetre, straight,

acicular, 1-5 mm long, 10-15 times longer than

wide; stellae sparse to dense, 0.2-0.3 mm

diameter, stalks 0-0.1 mm long; lateral rays

8-13, porrect or multiradiate; central ray absent

or present, 0-1 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Adult leaves ovate, entire or shallowly

lobed throughout; lobes 3-5 on each side, acute

or obtuse, lobing index 1-1.4; lamina 7.5-13

cm long, 3.5-6 cm wide, 2-3 times longer than

broad, apex acute or acuminate, base cuneate,

oblique part 0-3 mm long, obliqueness index

0-2 percent; petioles 0.5-2.2 cm long, 5-17%

length of lamina, prickles absent. Upper leaf

surface green; prickles 0-7, straight, acicular,

3-5 mm long, prickles absent or present on

midvein only or present on midvein and lateral

veins; stellate hairs absent, or confined to

midrib; ordinary stellae absent from surface;

finger hairs absent; Type 2 hairs absent. Lower

leaf surface greenish-white; prickles absent;

stellae dense, 0.1-0.3 mm apart, 0.3-0.5 mm

diameter, sessile; lateral rays 7 or 8, porrect;

central ray 0.1-0.5 t im es as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Inflorescence leaf-opposed,

pseudo-racemose, common peduncle 3-12 mm

long, rachis prickles absent; 5-35-flowered,

weakly andromonoecious, flowers 5-merous;

pedicels 6-10 mm long at anthesis, same

thic kn ess throughout, 0.3-0.4 mm thick at mid¬

point, prickles absent. Calyx tube 1.5-2.5 mm

long, lobes deltate or attenuate, 1-3 mm long;

prickles absent at anthesis; stellae moderate to

dense, transparent, 0.2-0.3 mm across, sessile,

lateral rays 5-8, central ray 1-1.5 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Corolla white or

mauve, 6-8 mm long, deeply lobed, inner

surface glabrous; anthers 3.5-5 mm long; ovary

glabrous; functional style 4.5-5 mm long, erect,

glabrous or with stellate hairs only, stellae

0.2-0.25 mm across, lateral rays 8-9, central

ray 0.7-1.5 times as long as laterals. Fruiting

calyx with lobes less than half length of mature

fruit, prickles absent. Mature fruits 1 or 2 per

inflorescence, globular, 14-16 mm diameter,

red, 1-locular (septum absent or incomplete);

placenta not apparent; mesocarp juicy,

succulent; exocarp c. 0.7 mm thick; pedicels

17-24 mm long in fruit, 0.6-0.8 mm thick at

mid-point; seeds pale yellow, 3.5-3.9 mm long.

Bean, Taxonomy of Solarium subg. Leptostemonum

Specimens examined : New Guinea. Bioto, Jul-Aug 1918,

White 581 (BRI). Northern Territory. Dalywoi Bay, Gove,

Feb 1988, Russell-Smith 4933 & Lucas (BRI, DNA); Rocky

Bay, Yirrkala, Mar 1988, Russell-Smith 5167 & Lucas (BRI,

DNA); 1.5 km NW of Yirrkala, Nov 1989, Forster PIF5976

(BRI, DNA). Queensland. Cook District: Newcastle Bay,

2.5 miles [4 km] S of Somerset, May 1948, Brass 18720

(BRI, CANB); Bamaga district, Galloways Creek area,

undated, Webb & Tracey 6075 (BRI); upstream of Hann Ck,

Jul 1986, Hind 4542 et al. (NSW); Jardine River N.P., 11

km NNW of the Eliot Creek-Jardine River confluence, 38.1

km S of Bamaga, Oct 1993, Fell DGF3647 & Dibella (BRI).

Distribution and habitat: Solanum yirrkalense

is known from the northernmost part of

Queensland, around Bamaga (Map 4). It also

grows near Gove in north-eastern Northern

Territory, and is known from a single New

Guinean specimen. It inhabits coastal vine

thickets on sand dunes.

Phenology: Flowers have been collected in

February and July-August; mature fruits in

March, May and November.

Notes: S. yirrkalense differs from S. discolor

by the mostly broader leaves with longer

petioles, the dense indumentum on the lower

leaf surface (vs. very dense for S. discolor ), the

central ray on the calyx stellae 1-1.5 times as

long as laterals (vs. 0.5-1 times for S. discolor ),

inflorescence common peduncle 3-12 mm long

(vs. 0-2 mm long for S. discolor ).

Conservation status: Not considered at risk.

12. Solanum defensum F.Muell., Fragm. 5:

193 (1866). Type: Queensland. Cook

District: Cape York Peninsula, undated,

E. Daemel (lecto: MEL [MEL12284]),

here chosen.

Erect, rhizomatous perennial shrub, 1-2.5 m

high. Juvenile branchlets with 25-30 prickles

per dm, 6-9 mm long; leaves (in outline)

elliptical, shallowly-lobed, with 4-6 pairs of

lobes; lamina 8-13 cm long, 2.5-4 cm wide,

with 10-20 prickles on upper surface. Adult

branchlets brown; prickles 1-10 per decimetre,

straight, acicular, 4-9 mm long, 13-18 t im es

longer than wide; stellae sparse to dense,

0.25-0.5 mm diameter, sessile; lateral rays 7

or 8, porrect; central ray absent or present, 0-0.4

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Adult

leaves ovate, entire or shallowly lobed

689

throughout; lobes 3-6 on each side, obtuse,

lobing index 1-1.4; lamina 7-14.5 cm long,

2.3-5 cm wide, 2.1-3.2 times longer than

broad, apex acute or acuminate, base cuneate

or obtuse, oblique part 0-2.5 mm long,

obliqueness index 0-2 percent; petioles 0.6-1

cm long, 6-10% length of lamina, prickles

absent or present. Upper leaf surface green;

prickles 0-10, straight, acicular, 5-8 mm long,

prickles absent or present on midvein only or

present on midvein and lateral veins; stellate

hairs absent, or confined to midrib; ordinary

stellae absent from surface; finger hairs absent;

Type 2 hairs absent. Lower leaf surface green;

prickles 0-4, straight, acicular, absent or

present on midvein only or present on midvein

and lateral veins; stellae very sparse to

moderate, 0.5-1.7 mm apart, 0.4-0.6 mm

diameter, sessile; lateral rays 7 or 8, porrect;

central ray 0.2-0.6 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Inflorescence supra-axillary,

pseudo-racemose, common peduncle 0-4 mm

long, rachis prickles absent; 5-13-flowered,

weakly andromonoecious, flowers 5-merous;

pedicels 4-8 mm long at anthesis, markedly

thicker distally, 0.15-0.25 mm thick at mid¬

point, prickles absent. Calyx tube 1.5-3 mm

long, lobes rostrate, 0.2-1 mm long; prickles

absent at anthesis; stellae moderate to dense,

transparent, 0.25-0.3 mm across, sessile, lateral

rays 7 or 8, central ray 0-0.5 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Corolla white, mauve or

purple, 5-8 mm long, deeply lobed, inner

surface sparsely to densely stellate-hairy;

anthers 3-4.5 mm long; ovary with stellate

hairs only; functional style 3.5-4.5 mm long,

erect, with stellate hairs only, stellae c. 0.25

mm across, lateral rays 7-10, central ray 0-1

times as long as laterals. Fruiting calyx with

lobes less than half length of mature fruit,

prickles absent. Mature fruits 1 or 2 per

inflorescence, globular, 13-17 mm diameter,

red, 1-locular (septum absent or incomplete);

placenta in cross-section sessile, elliptical;

mesocarp juicy, succulent; exocarp 1.2-1.7 mm

thick; pedicels 17-28 mm long in fruit,

0.6-0.8 mm thick at mid-point; seeds pale

yellow, 3.8-4.3 mm long.

Specimens examined : Queensland. Cook District: Leo

Creek, Upper Nesbit River, Aug 1948, Brass 19947 (BRI);

Table Range, Dead Horse Creek, Oct 1973, Dockrill 760

690

Austrobaileya 6 (4): 639-816 (2004)

(QRS); Mcllwraith Range, Sep 1974, Hyland 7643, 7658

(BRI, QRS); Lockerbie Scrub, Bamaga, Sep 1974, Webb &

Tracey 13481 (CANB, QRS); ‘Temple Bay’yards, Sep 1976,

Hyland 8964 (QRS); northern slopes Mt Tozer, Nov 1977,

Webb & Tracey 13479 (BRI); near Ginger Mick’s Mine, 2

km S of Punsand Bay, Feb 1990, Forster PIF6403 (BRI);

2.2 km from Captain Billy Landing hut, on the Heathlands

road, Mar 1992, Johnson 5023 (BRI); Claudie Falls, 2.5

km NE of Mt Tozer, Iron Range N.P., May 1992, Fell DF2614

& Jensen (BRI, QRS); Turrel Hill, 10 km WNW of Nesbit

River mouth, 51.6 km N of ‘Silver Plains’ HS., Aug 1993,

Fell DGF3398 etal. (BRI, CANB); 1.5 kmENEof Lamond

Hill, 8.5 km NNW of Lockhart River community, Mar 1994,

FV//DGF4139 & Stanton (BRI); Round Mountain, Embley

Range, ‘Silver Plains’, Jul 1997, Forster PIF21350 et al.

(AD, BRI, QRS).

Distribution and habitat : Solanum defensum

is endemic to Queensland; extending from the

tip of Cape York Peninsula to the Mcllwraith

Range near Coen (Map 5). It grows in

notophyll rainforest, in hilly terrain with

infertile soils derived from metamorphic or

granitic rocks.

Phenology : Flowers and fruits may be found

at any time of the year.

Notes: S. defensum is related to S. discolor ,

S. yirrkalense and S. fervens, but differs from

all these species by the green lower leaf surface

(stellate hairs absent to sparse), the central ray

of the stellae on the calyx only 0-0.5 times as

long as laterals, and the thicker fruiting

exocarp.

Sterile specimens of S. defensum are

readily confused with S. macoorai. These

species are however allopatric.

Conservation status: Not considered at risk.

13. Solanum discolor R.Br., Prodr. 445 (1810).

Type: [Queensland.] ‘Coen River’

[Pennefather River], Carpentaria, 7

November 1802, R. Brown (lecto: BM).

Illustration: Symon (1981: 141)

Erect, rhizomatous perennial shrub, 0.5-2 m

high. Juvenile stage unknown. Adult branchlets

yellow or brown; prickles absent or present,

0-10 per decimetre, straight, acicular, 2-6 mm

long, 10-16 t im es longer than wide; stellae very

dense, 0.25-0.4 mm diameter, stalks 0-0.1 mm

long; lateral rays 7 or 8, porrect; central ray

0.2-0.8 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Adult leaves lanceolate, elliptical or ovate,

entire or shallowly lobed throughout; lobes

3-6 on each side, obtuse, lobing index 1-1.3;

lamina 7-10 cm long, 1.6-4.5 cm wide,

2.3-4.3 times longer than broad, apex acute

or acuminate, base cuneate, oblique part 0-3

mm long, obliqueness index 0-4 percent;

petioles 0.4-0.9 cm long, 4-10% length of

lamina, prickles absent. Upper leaf surface

green; prickles 0-7, straight, acicular, 3-5 mm

long, prickles absent or present on midvein only

or present on mid vein and lateral veins; stellate

hairs absent, or confined to midrib; ordinary

stellae absent from surface; finger hairs absent;

Type 2 hairs absent. Lower leaf surface white

or yellowish; prickles absent; stellae very dense,

c. 0.05 mm apart, 0.3-0.5 mm diameter, stalks

0-0.1 mm long; lateral rays 8-9, porrect;

central ray 0.1-0.5 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Inflorescence leaf-opposed or

supra-axillary, pseudo-racemose, common

peduncle 0-2 mm long, rachis prickles absent;

7-20-flowered, strongly or weakly

andromonoecious, flowers 5-merous; pedicels

2-9 mm long at anthesis, same thickness

throughout, 0.2-0.4 mm thick at mid-point,

prickles absent. Calyx tube 0.5-1.5 mm long,

lobes deltate, 0.5-2 mm long; prickles absent

at anthesis; stellae dense to very dense, white,

0.25-0.4 mm across, sessile, lateral rays 6-8,

central ray 0.5-1 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Corolla mauve, 3-7 mm long, deeply

lobed, inner surface glabrous; anthers 2.5-4

mm long; ovary glabrous. Fruiting calyx with

lobes less than half length of mature fruit,

prickles absent. Mature fruits 1 or 2 per

inflorescence, globular, 10-14 mm diameter,

red, 1-locular (septum absent or incomplete);

placenta not apparent; mesocarp juicy,

succulent; exocarp 0.4-0.5 mm thick; pedicels

13-18 mm long in fruit, 0.5-0.8 mm thick at

mid-point; seeds pale yellow, 2.6-3.6 mm long.

Specimens examined : Queensland. Cook District: between

Rocky River and Massey Creek, Sep 1973, Stocker 1072

(BRI); Mapoon, S of Cullen Point, Port Musgrave, Feb 1981,

Morton AM 1075 (BRI); S bank of Pennefather River, Aug

1983, Clarkson 4914 (BRI); 0.7 km W of Bolt Head, Jul

1990, Clarkson 8806 & Neldner (BRI); 3.5 km NNE of

Massey Ck crossing, ‘Silver Plains’ station, Jul 1993 , Forster

PIF13614 et al. (BRI); 6 km W of the Rocky River mouth,

36.2 km ENE of Coen, ‘Silver Plains’ holding, Aug 1993,

Fell DGF3479 et al. (BRI, MEL); ‘Silver Plains’, S of

Scrubby Creek and W of Colmer Point, Jun 1995, Forster

Bean, Taxonomy of Solanum subg. Leptostemonum

PIF17061 (AD, BRI, MEL, QRS); Bolt Head, Jun 1996,

Gray 6857 (BRI); Temple Bay, Bolt Head, Jun 1996, Forster

PIF19359 (BRI).

Distribution and habitat: Solanum discolor is

endemic to Queensland, occurring on both the

east and west coast of Cape York Peninsula, as

far north as Mapoon and as far south as ‘Silver

Plains’ (Map 7). It occurs in depauperate vine

thicket, sometimes with emergent Araucaria

cunninghamii, on stranded sand dunes.

Phenology: Flowers are recorded for February,

June, July and August; mature fruits from June

to September.

Notes: S. discolor was in the past more broadly

circumscribed to include S. corifolium F.Muell.,

but it differs clearly from that species. See notes

under S. corifolium.

Conservation status: Not considered at risk.

14. Solanum corifolium F.Muell., Fragm. 2:

166 (1861). Type: [Queensland.]

Araucaria Ranges, Burnett River, Bunya

Bunya Ranges, December 1856, F.

Mueller (lecto: MEL [MEL 11617]; isolecto:

K), fide Symon (1981).

Illustration: Symon (1981: 136)

Sprawling or erect, rhizomatous perennial

shrub, 0.6-3 m high. Juvenile branchlets with

15-25 prickles per dm; leaves (in outline) ovate,

shallowly-lobed, with 2-3 pairs of lobes; lamina

9-11 cm long, 3.5-4.5 cm wide, with 0-20

prickles on upper surface. Adult branchlets

yellow; prickles 4-30 per decimetre, straight,

acicular, 3-7 mm long, 12-16 t im es longer than

wide; stellae dense to very dense, 0.15-0.25

mm diameter, sessile; lateral rays 6-8, porrect;

central ray absent; finger hairs absent; Type 2

hairs absent. Adult leaves ovate, entire or

shallowly lobed throughout; lobes 2-3 on each

side, obtuse, lobing index 1-1.2; lamina

4.5-10.5 cm long, 1.9-5 cm wide, 2-2.9 times

longer than broad, apex acute, base cuneate,

oblique part 0-1.5 mm long, obliqueness index

0-2 percent; petioles 0.3-1.1 cm long, 6-14%

length of lamina, prickles absent. Upper leaf

surface green; prickles 0-15, straight, acicular,

3-6 mm long, prickles absent or present on

midvein only or present on midvein and lateral

veins; stellate hairs absent, or confined to

691

midrib; ordinary stellae absent from surface;

finger hairs absent; Type 2 hairs absent. Lower

leaf surface white or yellowish; prickles 0-5,

straight, acicular, absent or present on midvein

only or present on midvein and lateral veins;

stellae very dense, c. 0.05 mm apart, 0.15-0.25

mm diameter, sessile; lateral rays 7 or 8,

porrect, central ray absent; finger hairs absent;

Type 2 hairs absent. Inflorescence supra-

axillary, pseudo-umbellate or pseudo-racemose,

common peduncle 0-3 mm long, rachis prickles

absent; 3-10-flowered, with all flowers bisexual

and 5-merous; pedicels 6-21 mm long at

anthesis, same thic kn ess throughout, 0.3-0.5

mm thick at mid-point, prickles absent. Calyx

tube 1-2.5 mm long, lobes deltate or rostrate,

1.5-3 mm long; prickles absent at anthesis;

stellae moderate to dense, 0.15-0.25 mm

across, sessile, lateral rays 6-8, central ray

absent; finger hairs absent; Type 2 hairs absent.

Corolla white or mauve, 8-10 mm long, deeply

lobed, inner surface glabrous; anthers 4-5.5

mm long; ovary with Type 2 hairs only;

functional style 5.5-7 mm long, erect, with

Type 2 hairs only or with stellate and Type 2

hairs, stellae c. 0.2 mm across, lateral rays

6-7, central ray 0.5-1 times as long as laterals.

Fruiting calyx with lobes less than half length

of mature fruit, prickles absent. Mature fruits

1 or 2 per inflorescence, globular, 7-12 mm

diameter, red, 1-locular (septum absent or

incomplete); placenta not apparent; mesocarp

juicy, succulent; exocarp 0.2-0.7 mm thick;

pedicels 10-22 mm long in fruit, 0.6-0.9 mm

thick at mid-point; seeds pale yellow, 2.8-3.7

mm long.

Selected specimens examined : Queensland. Cook District:

S.F.607, Bridle L.A., Dec 1973, Dansie s.n. (QRS). North

Kennedy District: razorback range NW of Mt Dryander, Jul

1974, Swan 75 (BRI); Paluma Holding, May 1982, Dansie

AFO 05151 (QRS); North Gregory, property of D. & R.

Clarke, adjacent to Dryander S.F., Jul 1997, Champion 1484

& Cali (BRI). Port Curtis District: Callide Valley, Apr 1937,

White 11210 (BRI); ‘Green Horizons’, 8.4 km from Rules

Beach Rd along Fingerfield Rd, c. 60 km NW of Bundaberg,

Oct 1992, Geckeler et al. 49 (CANB); S.F.69, SE of

Thangool, Mar 1996, Bean 10126 (BRI, MEL). Burnett

District: Eidsvold, undated, Bancroft s.n. (BRI); Bunya

Mountains S.F., Mar 1959, Thorne 20043a (BRI); Fontainea

Scrub, Gurgeena Plateau, S.F. 172, Feb 1994, Forster

PIF14805 (AD, BRI). Wide Bay District: base of Guyra

Mount, below Mt Bauple N.P., Feb 1988, Forster PIF3535

(BRI); Mt Glastonbury, S.F.242, Dec 1991, Forster PEF9277

& Sharpe (BRI, MEL). Darling Downs District:

‘Sunnyvale’, Bell, Mar 1927, Langford s.n. (BRI); S.F. 197

Diamondy, 32 km NE of Jandowae, Mar 1999, Forster

PIF24098 & Booth (BRI, MEL, QRS). Moreton District:

692

Mt Glorious, May 1923, White 1953 (BRI); Binna Burra,

Lamington N.P., Apr 1959, Thome 20408 (BRI); S.F.289,

Yarraman, Feb 1972, Moriarty 879 (BRI); P al m Grove N.P,

Tamborine Mountain, Jul 1983, Guymer 1868 & McDonald

(AD, BRI, GUAM); Nineteen L.A., T.R.209, Mt Brisbane,

Jun 1990, Forster PIF6862 et al. (BRI, L, MEL, QRS);

Murray Grey Drive, Dulong, W of Nambour, Jan 2000, Bean

16004 (BRI); Wilkie’s Scrub, Wongawallan, 7 km W of

Coomera, Jul 2001, Bean 17691 (BRI). New South Wales.

North Coast. Richmond River, anno 1876, Fawcett s. n. (NSW).

Distribution and habitat: Solanum corifolium

is common in south-eastern Queensland as far

north as Bundaberg and Biloela, and with

disjunct occurrences near Proserpine, Ingham

and Mareeba. There is an old record from far

north-eastern New South Wales (Map 7). It

grows in Araucarian or other mixed notophyll

rainforest, in hilly to mountainous terrain.

Phenology: Flowers are recorded from

November to March; the exception is a

flowering record from north Queensland in July

{Champion 1484 & Cali); mature fruits are

recorded between January and July.

Notes: S. corifolium is closely related to

S. discolor, but differs by the branchlet stellae

only 0.15-0.25 mm across (0.25-0.4 mm for

S. discolor), central ray of branchlet stellae

lacking (0.2-0.8 times as long as laterals for

S. discolor), all flowers bisexual (some flowers

male for S. discolor), corolla 8-10 mm radius

(3-7 mm for S. discolor), anthers 4-5.5 mm

long (2.5-4 mm long for S. discolor), ovary

with Type 2 hairs (glabrous for S. discolor),

and the mostly smaller fruits.

Conservation status: Widespread. Not

considered at risk.

15. Solanum mentiens A.R.Bean sp. nov.

Frutex prostratus caulibus longis

serpentibus ad nodos radicantibus; folia

ovata 1.5-2.2plo longiora quam latiora,

glabra et atrovirentia supra, densissime

pilosa subtus, apice acuto; stellae omnes

radio centrali carentes; inflorescentia

floribus aliquibus functionale maribus;

fructus maturitate laete rubra, exocarpio

0.8-1.0 mm crasso. Typus: Queensland.

Moreton District: Bahr’s Scrub, 5 km

SSW of Beenleigh, J. Davidson property,

19 December 2001, P.I. Forster 28047 &

G. Leiper (holo: BRI (1 sheet + spirit);

iso: A, K, L, MEL, NSW).

Austrobaileya 6 (4): 639-816 (2004)

Solanum discolor var. procumbens

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

55: 71 (1944). Type: Queensland.

Moreton District: Upper Teviot, undated,

B. Scortechini (holo: MEL).

Prostrate, stoloniferous perennial shrub, c. 0.1

m high. Juvenile stage absent. Adult branchlets

yellow or brown; prickles absent or present,

0-10 per decimetre, straight, acicular, 2-7 mm

long, 8-10 times longer than wide; stellae dense

to very dense, 0.15-0.2 mm diameter, sessile;

lateral rays 6-8, porrect; central ray absent;

finger hairs absent; Type 2 hairs absent. Adult

leaves elliptical or ovate, entire; lamina 5-7.5

cm long, 2.7-4 cm wide, 1.5-2.2 t im es longer

than broad, apex obtuse, base cuneate or obtuse,

oblique part 0-3.5 mm long, obliqueness index

0-6 percent; petioles 0.5-1.1 cm long, 7-20%

length of lamina, prickles absent. Upper leaf

surface green; prickles 0-3, straight, acicular,

2-5 mm long, prickles absent or present on

midvein only; stellate hairs absent, or confined

to midrib; ordinary stellae absent from surface;

finger hairs absent; Type 2 hairs absent. Lower

leaf surface white or yellowish; prickles absent;

stellae very dense, c. 0.05 mm apart, 0.2-0.3

mm diameter, sessile; lateral rays 7 or 8,

porrect; central ray absent; finger hairs absent;

Type 2 hairs absent. Inflorescence supra-

axillary, solitary or pseudo-racemose, common

peduncle 0-3 mm long, rachis prickles absent;

1-6-flowered, weakly andromonoecious,

flowers 5-merous; pedicels 6-18 mm long at

anthesis, same thickness throughout, 0.3-0.5

mm thick at mid-point, prickles absent. Calyx

tube 2-4 mm long, lobes elliptic or deltate, 1-3

mm long; prickles absent at anthesis; stellae

moderate, transparent, 0.15-0.25 mm across,

sessile, lateral rays 6-8, central ray absent;

finger hairs absent; Type 2 hairs absent. Corolla

white, 7-12 mm long, shallowly or deeply

lobed, inner surface glabrous; anthers 4.5-5.5

mm long; ovary with Type 2 hairs only;

functional style 5.5-7 mm long, erect, glabrous

or with Type 2 hairs only. Fruiting calyx with

lobes less than half length of mature fruit,

prickles absent. Mature fruits 1-3 per

inflorescence, globular, 10-12 mm diameter,

red, 1-locular (septum absent or incomplete);

placenta in cross-section sessile, semi-circular

to elliptical; mesocarp moist but not juicy;

exocarp 0.8-1 mm thick; pedicels 22-25 mm

long in fruit, 0.8-1 mm thick at mid-point;

seeds pale yellow, 2.4-3.2 mm long. Fig. 18.

Bean, Taxonomy of Solanum subg. Leptostemonum

693

QUEENSLAND HERBARIUM (BRI>

iFkas Queensland Whwtan

Solanum

QUEENSLAND HERBARIUM (ERI)

Brisbane Australia

ag 553713

fbAoT/Pe 0ljeensliln ' i Herbarium (BRI)

SoLasm. Menftets /t. st

P® 1 ' S?. /f, CWt/dWZcajt

CaftP.LFWHH PIFJKHI laoetMOi

L*rp*r.6.;

SmuanSB* 153a ion W* [ACDK] Deptftni

iM.MBJZiBaHHSj l9$C-1S*»2t Aft lOfra

Barn sen*. 5 tom SSW SC eaanfcdh J OavH™ prapirty.

ft'das^aan vuw&niSl 6 ft *S SS 6 .

E5«xnfc*ffl, prwlrafc buib. leftting |U>) Homo; kiwn wtule; friif

aemt-maio

Uoci^wsy M<wnan

Spits HflinItJil BRI.

0 *L £p latiMMb

Dvp. FJ5l>NSWAItL

■ May t* clml xi Wbtebon Numbet AQ 5»T1 J

tAiabiv* PJR*t>

Fig. 18. Holotype of Solanum mentiens.

694

Specimens examined : Queensland. Moreton District: near

Canungra,May 1917, White s.n. (BRI); end of French’s Creek

road, c. 12 km SW of Boonah, Jul 1984, Bird s.n. (BRI);

Bahr’s Scrub, near Beenleigh, Jan 2000, Leiper s.n. (BRI);

Bahr’s Scrub, 5 km SW of Beenleigh, Feb 2001 ,Bean 17370

(BRI, MEL); Bahr’s Scrub, 6 km SW of Beenleigh, Feb 2002,

Bean 18513 (BRI); slopes of Mt French, W of Boonah, Apr

2003, Bean 20144 (BRI).

Distribution and habitat: Solanum mentiens

is endemic to Queensland. Confined to the

Boonah and Beenleigh areas in far southeastern

Queensland (Map 5). It inhabits Araucarian

notophyll vineforest.

Phenology: Flowers are recorded for December,

January and February; fruits for February and

July.

Notes: This species is very distinct in the field

due to its ground-hugging prostrate habit. It

produces long trailing stems that strike roots

frequently at the nodes. The lack of a central

ray on any stellae immediately distinguishes

S. mentiens from the other two species with

the same habit ( S. serpens and S. acanthodapis).

Solanum mentiens is closely related to

S. corifolium, but differs by the prostrate

stoloniferous habit, leaves 1.5-2.2 times longer

than broad (2.0-2.9 times for S. corifolium );

leaf apex obtuse (acute for S. corifolium );

inflorescence with some flowers functionally

male (all bisexual in S. corifolium ); placenta

sessile, semicircular to elliptical (not apparent

in S. corifolium ); and exocarp 0.8-1.0 mm

thick (0.2-0.7 mm for S. corifolium ).

The locality of “Upper Teviot” given by

Scortechini for his specimen refers to the Teviot

Brook, which flows through the town of

Boonah.

Conservation status: S. mentiens is currently

known from 3 locations. Weeds or land

clearance threaten all populations. One

population may be just within the Mt French

National Park. Applying the IUCN guidelines

(IUCN. 2001), a category of “Endangered” is

recommended (EN Blab(iii,v)+2ab(iii,v); Cl).

Etymology: From the Latin mentiens , meaning

‘imitating’. This refers to its similarity to

S. corifolium.

Austrobaileya 6 (4): 639-816 (2004)

16. Solanum shirleyanum Domin, Biblioth.

Bot. 89: 578 (1929). Type: Queensland.

Moreton District: Tamborine Mountain,

March 1910, K. Domin s.n. (holo: PR (2

sheets)).

Erect, rhizomatous perennial shrub, 0.6-1.8 m

high. Juvenile branchlets with 1-20 prickles

per dm; leaves (in outline) lanceolate or

elliptical or ovate, entire or shallowly-lobed,

with 2-3 pairs of lobes; lamina 9-10 cm long,

3-4.2 cm wide, with 2-8 prickles on upper

surface. Adult branchlets yellow or brown;

prickles absent or present, 0-6 per decimetre,

straight, acicular or broad-based, 2-5 mm long,

6- 12 times longer than wide; stellae dense to

very dense, 0.25-0.4 mm diameter, stalks

0-0.1 mm long; lateral rays 6-8, porrect;

central ray 0.5-3 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Adult leaves elliptical or ovate, entire;

lamina 4-11 cm long, 1.5-4 cm wide, 2-3 times

longer than broad, apex acute or acuminate,

rarely obtuse; base cuneate, oblique part 0-3.5

mm long, obliqueness index 0-3 percent;

petioles 0.25-1.3 cm long, 5-12% length of

lamina, prickles absent. Upper leaf surface

green; prickles 0-5, straight, acicular, 4-6 mm

long, prickles absent or present on mid vein

only; stellate hairs confined to midrib, or

distributed throughout; protostellae absent;

ordinary stellae very sparse, 0.8-4 mm apart,

0.3-0.4 mm across, sessile; lateral rays 6-8,

porrect; central ray 1-2 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Lower leaf surface green or

greenish-white; prickles absent; stellae dense,

0.1-0.3 mm apart, 0.3-0.5 mm diameter,

sessile; lateral rays 7 or 8, porrect; central ray

1-4 times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent.

Inflorescence supra-axillary, pseudo-umbellate,

common peduncle absent; 2-9-flowered, with

all flowers bisexual and 5-merous; pedicels

7- 17 mm long at anthesis, same thickness

throughout, 0.2-0.3 mm thick at mid-point,

prickles absent. Calyx tube 2-3 mm long, lobes

elliptic, 0.3-1 mm long; prickles absent at

anthesis; stellae sparse to moderate,

transparent, 0.15-0.25 mm across, sessile,

lateral rays 5-7, central ray 1-1.5 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent or present. Corolla

mauve, 7-9 mm long, deeply lobed, inner

surface glabrous; anthers 3.5-4.5 mm long;

ovary with Type 2 hairs only; functional style

Bean, Taxonomy of Solanum subg. Leptostemonum

5.5-7 mm long, erect, glabrous. Fruiting calyx

with lobes less than half length of mature fruit,

prickles absent. Mature fruits 1 or 2 per

inflorescence, globular, 3.5-6 mm diameter,

red, glabrous or with a few scattered Type 2

hairs, 1-locular (septum absent or incomplete);

placenta not apparent; mesocarp juicy,

succulent; exocarp 0.1-0.2 mm thick; pedicels

13-21 mm long in fruit, 0.4-0.7 mm thick at

mid-point; seeds pale yellow, 2.6-3 mm long.

Specimens examined : Queensland. Wide Bay District:

Gympie, undated, Kenny (BRI); Peters Ck valley, Conondale

Range S.F., above Funnel Hut site, Oct 1982, McDonald 3627

(BRI); Goods Rd, S.F.274 Conondale, Feb 1991, McDonald

4720 (BRI); Lenthalls Dam scrub, S.F.1294, Oct 1995,

Forster PIF17915 (BRI); S.F.783, 4 km NW of Montville,

May 2001, Forster PIF27114 (BRI). Moreton District:

Maroochie, Oct 1874, Bailey (BRI); Mt Tamborine, 1888,

Simmonds s.n. (BRI); Palmwoods, May 1907, White s.n.

(BRI); Currumbin, 1912, O’Brien (BRI); eastern foothills

of Darlington Range, Upper Ormeau Rd, off Pacific Hwy,

Apr 1984, Williams 84028 (BRI); Bahr’s Scrub, 5 km SW

of Beenleigh, Feb 2001, Bean 17371 (BRI, NSW); Armitage

Creek road, Canungra Army Reserve, 7 km SE of Canungra,

Feb 2001, Bean 17382 (BRI); Rosins Lookout Conservation

Park, Beechmont, Jul 2001, Forster PIF27494 etal. (BRI).

New South Wales. North Coast: Tyalgum Ridge,

Macpherson Range, c. 25 kmWNW of Murwillumbah, Dec

1977, Haegi 1529 (NSW); 1.1 km along South Chowan road,

Nullum S.F., S of Murwillumbah, Apr 2001, Bean 17558

(BRI).

Distribution and habitat: Solanum

shirleyanum occurs in high rainfall areas of

southeastern Queensland, and extends to near

Murwillumbah in N.S.W. (Map 6). It grows

on the margins of notophyll rainforest where

Eucalyptus grandis is often prominent. Soils

are infertile, and associated species may include

Caldcluvia paniculosa and Callicoma

serratifolia. It occurs at relatively low altitudes,

although reaching 560 metres in the Conondale

Ranges.

Phenology: Flowers are recorded for October,

February and April; mature fruits in February,

April, May and July.

Notes: It is related to both S. stelligerum and

S. corifolium, and Domin considered that it was

a hybrid between these two species. However,

my fieldwork has shown that S. shirleyanum

should be regarded as a species in its own right.

It forms populations uniform in morphology,

flowers and fruits freely, and can occur in areas

geographically remote from the postulated parents.

It differs from S. corifolium by the smaller

flowers and fruits, more slender corolla lobes,

695

and leaf stellae with conspicuous central rays.

It differs from S. stelligerum by the shorter

petioles, leaves with stellae all sessile and

having shorter central rays, smaller fruits,

larger seeds and glabrous style.

Conservation status: Moderately widespread.

Not considered at risk.

17. Solanum dryanderense A.R.Bean sp. nov.

Frutex; aculei ramuli praesentes sed

sparse distributi; folia integra ovata,

petiolus longitudine 4-11% laminae

aequans, supra viridia glabra, subtus pilis

stellatis petiolis 0.15-0.25 mm longis,

apice acuto usque acuminato; aculei

calycis absentes; fructus maturitate rubri,

seminibus 3.5-4 mm longis. Typus:

Queensland. North Kennedy District: c.

15 km N of Proserpine, near crest of ridge

leading to east summit of Mt Dryander,

21 July 1974, R.J. Henderson H2214, V.

Moriarty & J. Swan (holo: BRI).

Solanum sp. (Mt Dryander G.P. Guymer

1743) in Henderson (2002).

Erect, rhizomatous perennial shrub, 1-3 m

high. Juvenile stage unknown. Adult branchlets

grey or brown; prickles absent or present, 0-10

per decimetre, straight, acicular, 2-6 mm long,

10-18 times longer than wide; stellae very

dense, 0.4-0.6 mm diameter, stalks 0-0.2 mm

long; lateral rays 6-13, porrect or multiradiate;

central ray 0.7-2 t im es as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Adult leaves ovate, entire; lamina 5-10

cm long, 2-4 cm wide, 2.1-3.2 times longer

than broad, apex acute or acuminate, base

cuneate or obtuse, oblique part 0-1 mm long,

obliqueness index 0-2 percent; petioles 0.2-1.1

cm long, 4-11% length of lamina, prickles

absent. Upper leaf surface green; prickles 0-2,

straight, acicular, 1-6 mm long, prickles absent

or present on midvein only; stellate hairs

absent, or confined to midrib; ordinary stellae

absent from surface; finger hairs absent; Type

2 hairs absent. Lower leaf surface yellowish;

prickles absent; stellae dense, 0.1-0.25 mm

apart, 0.4-0.6 mm diameter, stalks 0.15-0.25

mm long; lateral rays 6-8, porrect; central ray

0.7-1.5 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Inflorescence supra-axillary, solitary or pseudo-

umbellate, common peduncle 0-1 mm long; 1

or 2-flowered, flowers 5-merous; pedicels c. 9

696

Austrobaileya 6 (4): 639-816 (2004)

[aonwimmj mm w* p mM

HO«A Of |Jjpai5tJlia) TOETH KE KPEO * BHTIic t

20 i“> iss 3-' 1 *" 575 ™ : aoaifiraoa. .

"" R«J« Henderson II22X4 21 Ail 1074

V. Itariarty & 7. Siron

So iamb.

discolor R.Er.

I h— | a*- W 1| * | ~-1

I solan, riraiffi - '" Ml “ " ■* l ' w “~‘ T —*"■»*» * I

Ca IS tan H o£ Jraaipinn.

north easterly fating slope near crest of

ridge loading ta easterly peak of Ht* Dtyanrler*

Altitude approsinato* Tropical closod forest,

srcy sandy loss soil in oilght break in over¬

head canopy*

arect shrub with nproadieg canopy*

111794

ffat&jypg Queonsrans Herbarium (Bfli)

So/vjttrun titty 4 -7 duitAtie /4. A, if aiv

Dst /t S. tmit/ticrzoo#

Oueensland Herbarium (Bfll)

.■ToZ+aaf - y (ftt '741 )

/f-S, Stas

Dais

aiFC^K-SL AND

HER BA R ID hi

2:3229

BRISBANE :

Fig. 19. Holotype of Solatium dryanderense.

Bean, Taxonomy of Solanum subg. Leptostemonum

mm long at anthesis, same thickness

throughout, 0.4-0.5 mm thick at mid-point,

prickles absent. Calyx prickles absent at

anthesis; stellae moderate, transparent, c. 0.4

mm across, stalks 0-0.1 mm long, lateral rays

6-8, central ray 0.7-1.2 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. 6-7 mm long, deeply lobed,

inner surface glabrous; anthers c. 5 mm long;

functional style c. 6 mm long, with Type 2 hairs

only. Fruiting calyx with lobes less than half

length of mature fruit, prickles absent. Mature

fruits 1 per inflorescence, globular, 6-7 mm

diameter, red, 1-locular (septum absent or

incomplete); mesocarp juicy, succulent;

pedicels 15-20 mm long in fruit, 0.5-0.6 mm

thick at mid-point; seeds pale yellow, 3.5-4 mm

long. Fig. 19.

Specimens examined : Queensland. North Kennedy

District: upper Dryander Creek, Mt Dryander, NE of

Proserpine, Oct 1969, Webb & Tracey 8354 (BRI); c. 15 km

N of Proserpine, on mountain spur closely parallel to Mt

Dryander Range, Jul 1974 , Henderson H2178Aeta/. (BRI);

c. (5 kmN of Proserpine, on steep SW slopes of Mt Dryander,

Jul 1974, Henderson H2216 et al. (BRI); Mt Dryander, Jul

1974, Swan 77 (BRI); Mt Dryander, Jul 1974, Swan 82

(BRI); headwaters of Box Creek, in vicinity of The

Bloodwoods, Mt Dryander, May 1982, Guymer 1743 (BRI);

ridge above Vine Creek, Mt Dryander, N of Proserpine, May

1992, McDonald 5098 (BRI).

Distribution and habitat : Solanum

dryanderense is endemic to Queensland.

Known only from Mt Dryander, north of

Proserpine (Map 4). Most collections have been

made between 400 and 600 metres altitude,

with one collection given as 200 metres altitude.

It inhabits sunny breaks in notophyll rainforest

on steep ridges.

Phenology : Very little fertile material is

available. A single flower was collected in July,

and mature fruits have been collected in May

and July.

Notes: S. dryanderense is closely related to

S. shirleyanum, but differs by the larger stellae

of the calyx and branchlets, the stellae of the

lower leaf surface with stalks 0.15-0.25 mm

long (sessile for S. shirleyanum ), calyx stellae

c. 0.4 mm diameter (0.15-0.25 mm daimeter

for S. shirleyanum ) and seeds 3.5-4 mm long

(2.6-3 mm long for S. shirleyanum ).

Conservation status: S. dryanderense is known

only from Mt Dryander near Proserpine, where

it is protected within a National Park. Applying

697

the IUCN guidelines (IUCN. 2001), a category

of “Vulnerable” is recommended (VU Dl+2).

Etymology: The epithet refers to Mt Dryander

near Proserpine, the only known locality for

the species.

18. Solanum stelligerum Sm., Exot. Bot. 2:

57, t. 88 (1805). Solanum stelligerum var.

stelligerum Benth., FI. Austral. 451

(1868) Type: New South Wales. Port

Jackson, 1792, J. White (holo: LINN, n.v.,

microfiche 365.13)

Solanum lucorum Domin, Repert. Spec. Nov.

Regni Veg. 12: 130 (1913). Solanum

stelligerum var. lucorum F. Muell. ex

Benth. FI. Austral. 451 (1868). Type:

[Queensland.] Araucaria Ranges, Burnett

River, December 1856, F. Mueller (holo:

K; iso: MEL [MEL 14113]).

Illustration: Symon (1981: 129)

Erect, rhizomatous perennial shrub, 0.8-2.5 m

high. Juvenile branchlets with 20-60 prickles

per dm; leaves (in outline) lanceolate,

shallowly-lobed, with 1 or 2 pairs of lobes;

lamina 4-7 cm long, 1.3-2.4 cm wide, with 1-5

prickles on upper surface. Adult branchlets

yellow or brown; prickles absent or present,

0-10 per decimetre, straight, acicular, 6-10 mm

long, 14-18 times longer than wide; stellae

sparse to very dense, 0.3-0.5 mm diameter,

stalks 0-0.4 mm long; lateral rays 7 or 8,

porrect; central ray 1-4 t im es as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent or sparse. Adult leaves lanceolate

or ovate, entire; lamina 2.5-13 cm long, 0.9—4.5

cm wide, 1.8-3.5 times longer than broad, apex

obtuse or acute, base cuneate or obtuse, oblique

part 0-3 mm long, obliqueness index 0-5

percent; petioles 0.4-1.8 cm long, 11-16%

length of lamina, prickles absent. Upper leaf

surface green; prickles 0-3, straight, acicular,

6-9 mm long, prickles absent or present on

midvein only; stellate hairs confined to midrib,

or distributed throughout; protostellae absent;

ordinary stellae absent from surface to sparse,

0.5-1.5 mm apart, 0.3-0.8 mm across, sessile;

lateral rays 4-8, porrect; central ray 1.5-5 times

as long as laterals, not gland-tipped; finger

hairs absent; Type 2 hairs absent. Lower leaf

surface white or yellowish; prickles absent;

stellae dense to very dense, 0.05-0.1 mm apart,

0.3-0.6 mm diameter, stalks 0-0.6 mm long;

lateral rays 7 or 8, porrect; central ray 1.5-4

698

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent.

Inflorescence supra-axillary, solitary or pseudo-

racemose, common peduncle 1-2 mm long,

rachis prickles absent; 1-5-flowered, weakly

andromonoecious or with all flowers bisexual,

flowers 5-merous; pedicels 8-15 mm long at

anthesis, same thickness throughout or

markedly thicker distally, 0.4-0.6 mm thick at

mid-point, prickles absent. Calyx tube 1.5-3

mm long, lobes deltate or rostrate, 0.5-3 mm

long; prickles absent at anthesis; stellae very

dense, yellow or white, 0.25-0.4 mm across,

stalks 0-0.1 mm long, lateral rays 7 or 8, central

ray 1-4 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent,

or present. Corolla white, mauve or purple, 6-12

mm long, shallowly or deeply lobed, inner

surface glabrous; anthers 4-5.5 mm long; ovary

with Type 2 hairs only, or with stellate and Type

2 hairs; functional style 5.5-8 mm long, erect,

with Type 2 hairs only or with stellate and Type

2 hairs, stellae 0.25-0.4 mm across, lateral rays

6-7, central ray 1-3 t im es as long as laterals.

Fruiting calyx with lobes less than half length

of mature fruit, prickles absent. Mature fruits

1-4 per inflorescence, globular, 6.5-9 mm

diameter, red, glabrous or with a few scattered

stellate hairs, 1-locular (septum absent or

incomplete); placenta not apparent; mesocarp

juicy, succulent; exocarp 0.25-0.5 mm thick;

pedicels 14-21 mm long in fruit, 0.6-0.7 mm

thick at mid-point; seeds pale yellow, 1.9-2.2

mm long. Devil’s Needles.

Selected specimens examined : Queensland. North

Kennedy District: Selheim, 25 km E of Charters Towers,

Mar 1988, Collins MPA44 (BRI); ‘Fanning River’, c. 25

km NW of Mingela, Aug 1989, Fell FAN83 & Cumming

(BRI). Port Curtis District: Marmor near Rockhampton,

Mar 1920, Francis (BRI); Mt Morgan, Jul 1938, Goy 327

(BRI); western slopes of Mt Farcom Range, near Yarwun,

May 1971, Webb & Tracey 10592A(BRI, CANB); MtEtna,

Oct 1976, Hyland 9099 (QRS); Burnett Range, 0.5 km NW

of Mt Fort William, May 1977, Crisp 2701 (AD, BRI,

CANB); Kroombit Tableland, c. 60 km SW of Gladstone,

Jun 1977, Crisp 2850 (AD, BRI, CANB); S.F.69, SE of

Thangool, Jun 1996, Bean 10432 (BRI, MEF). Burnett

District: 2.5 km from Proston on road to weir, May 1996,

Bean 10287 (BRI, MEF); c. 5 km due ENE of Ceratodus

railway junction, E of the Burnett River, Apr 1997, Pollock

ABP575 (BRI); MtBlandy, 4 km W of Mingo Crossing, Mar

1999, Forster PIF24152 & Booth (AD, BRI, QRS); Bunya

Mountains, 1.8 km along Nanango road, Jun 2001, Bean

17676 (BRI); Hoggs Rd, Tingoora, N of Kingaroy, Nov 2001,

Bean 18103 (BRI, MEL). Wide Bay District: Kin Kin, Mar

1916, Francis & White (BRI); near bank of Kolan River,

about N of Gin Gin, Apr 1945, Blake 15519 & Webb (BRI);

Yerra, 15 miles [24 km] W of Maryborough, Jan 1954, Cross

(BRI); south of Anderson road, 10 km W of Cooroy, Nov

1993, Bean 7064 (BRI); Pine Creek scrub, 1 km E of Electra,

Austrobaileya 6 (4): 639-816 (2004)

Oct 1996, Forster PIF 20049 &Leiper (BRI, MEL). Darling

Downs District: Cunningham’s Gap, Main Range, Jul 1930,

White 6867 (BRI); The Head, near border fence, Jun 1980,

Williams 80067 (BRI); behind Greenwood churchyard, N of

Oakey, Oct 1998, Fensham 3505 (BRI); May Road, W of

Clifton, Jan 2002, Bean 18357 (BRI, CANB). Moreton

District: scrub below Enoggera Dam, Aug 1874, Bailey s.n.

(BRI); Benarkin S.F., Blackbutt, Aug 1967, Henderson H293

(BRI); Falls Creek, 4.5 km NW of West Haldon, May 1987,

Forster PIF2933 & Bird (BRI); Mt Davidson, 5 km S of

Withcott, Jul 1990, Forster PIF6903 & Bird (BRI, MEL,

QRS); S.F.258 Mt Binga, 11 km SE of Cooyar, Feb 2000,

Bean 16058 (BRI, MEL); Back Creek road, Canungra Army

Reserve, 5 km SE of Canungra, Feb 2001, Bean 17380 (BRI).

New South Wales. North Coast: Lismore, Jul 1894,

Baeuerlen 1261 (AD, BRI, CANB, MO, NSW); Mt Warning,

Tweed River, Aug 1916, Boorman (AD, BRI, NSW); Branch

Road, Dalmorton S.F., SW of Grafton, Jan 2001, Bean 17257

(BRI); 2.5 km along Carnham Road, Fine Flower, NW of

Grafton, Feb 2001 ,Bean 17335 (BRI, MEL, NSW).

Distribution and habitat : Solanum stelligerum

is a widespread species ranging from Bodalla

in southern New South Wales to Rockhampton

in Queensland, with a disjunct occurrence near

Charters Towers (Map 8). It inhabits margins

of notophyll rainforest or shrubby eucalypt

open-forest, on many soil types and at low to

high altitudes. It is the most commonly

encountered native species in eastern coastal

Australia.

Phenology: Flowers and fruits may be found

at any time of the year.

Notes: Most closely related to S. parvifolium

subsp. parvifolium , but differs by the broader

leaves (1.8-3.5 times longer than wide), stellae

on all plant parts with a long central ray (1-5

t im es as long as laterals), the presence of long-

stalked stellae (to 0.6 mm long) on the lower

leaf surface, the rusty tomentum at least on the

veins of the lower leaf surface, and the

shallowly lobed juvenile leaves (entire for

S. parvifolium subsp. parvifolium).

Conservation status: Widespread. Not

considered at risk.

19. Solanum parvifolium R.Br., Prodr. 446

(1810). Type: [Queensland. Port Curtis

District:] Broadsound, 18 September

1802, R. Brown [Bennett No. 2673] (lecto:

BM; isolecto: K),fide Symon (1981).

Solanum accedens Domin, Repert. Spec.

Nov. Regni Veg. 12: 130-1 (1913). Type:

Queensland. Rockhampton, undated, J.

Dallachy s.n. (holo: K).

Two subspecies are recognised, and are

distinguished by the following key:

Bean, Taxonomy of Solanum subg. Leptostemonum

699

Leaves 2.5-7 x 0.5-1.5 cm, Type 2 hairs absent; ovary with Type 2 hairs

only; style 3.5-6.5 mm long; flowers all bisexual .... 5. parvifolium subsp. parvifolium

Leaves 7.5-13.5 x 1.3-3.5 cm, Type 2 hairs present; ovary either without

indumentum or with both stellate hairs and Type 2 hairs; style 7.0-9.0 mm long; male

flowers consistently present in the inflorescence. S. parvifolium subsp. tropicum

19a. Solanum parvifolium subsp. parvifolium

Illustration : Symon (1981: 131)

Erect, rhizomatous perennial shrub, 0.5-1.2 m

high. Juvenile stage unknown. Adult branchlets

grey or brown; prickles 1-20 per decimetre,

straight, acicular, 0.5-7 mm long, 10-18 t im es

longer than wide; stellae sparse to dense,

0.2-0.4 mm diameter, sessile; lateral rays

6-8, porrect; central ray 0.2-1 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs sparse. Adult leaves linear

or narrow lanceolate, entire; lamina 2.5-7 cm

long, 0.5-1.5 cm wide, 3.7-7.5 times longer

than broad, apex obtuse or acute, base cuneate,

oblique part 0-3 mm long, obliqueness index

0-4 percent; petioles 0.2-0.9 cm long, 7-16%

length of lamina, prickles absent. Upper leaf

surface green; prickles absent; stellate hairs

confined to midrib, or distributed throughout;

protostellae absent; ordinary stellae very sparse,

0.8-5 mm apart, 0.3-0.5 mm across, sessile;

lateral rays 7 or 8, porrect; central ray 0.8-1.2

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Lower

leaf surface white or yellowish; prickles absent;

stellae very dense, c. 0.05 mm apart, 0.2-0.5

mm diameter, stalks 0-0.1 mm long; lateral

rays 7-9, porrect; central ray 0.2-1 times as

long as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Inflorescence

supra-axillary, solitary or pseudo-racemose,

common peduncle 0-4 mm long, rachis prickles

absent; 1-8-flowered, with all flowers bisexual

and 5-merous; pedicels 7-13 mm long at

anthesis, same thic kn ess throughout, 0.4-0.7

mm thick at mid-point, prickles absent. Calyx

tube 1.5-3 mm long, lobes deltate, 1-3 mm

long; prickles absent at anthesis; stellae dense,

yellow or white, 0.25-0.4 mm across, sessile,

lateral rays 7 or 8, central ray 0.2-1 t im es as

long as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Corolla purple, 7-9

mm long, shallowly or deeply lobed, inner

surface glabrous; anthers 3.5-5 mm long; ovary

with Type 2 hairs only; functional style 3.5-6.5

mm long, erect, with Type 2 hairs only or with

stellate and Type 2 hairs, stellae 0.3-0.4 mm

across, lateral rays 7 or 8, central ray c. 0.3

times as long as laterals. Fruiting calyx with

lobes less than half length of mature fruit,

prickles absent. Mature fruits 1-6 per

inflorescence, globular, 6-8 mm diameter, red,

1-locular (septum absent or incomplete);

placenta not apparent; mesocarp juicy,

succulent; pedicels 11-27 mm long in fruit,

0.5-0.7 mm thick at mid-point; seeds pale

yellow, 1.9-2.3 mm long.

Selected specimens examined : Queensland. North

Kennedy District: 14 km E of ‘Mt Cooper’ HS, Jun 1992,

Thompson CHA141 & Sharpe (AD, BISH, BRI). Mitchell

District: base of Great Dividing Range on W side, Sep 1984,

O’Keeffe 600 (BRI). South Kennedy District: Alpha, Sep

1959, Macartney (BRI); ‘Strathmore’, 14 mi les [23 km] W

of Collinsville, May 1960, Johnson 1804 (BRI). Leichhardt

District: Moura-Baralaba road, c. 8 miles [13 km] from

Baralaba, May 1960, Johnson 1701 (BRI); DipperuN.R, c.

24 km S of Nebo, Sep 1971, McDonald 152 (BRI); Yatton

Ck, on Sarina-Marlborough road, c. 93 km from

Marlborough, Jul 1974, Moriarty 1543 (BRI, CANB); Dry

Creek valley, Ka Ka Mundi section, Carnarvon N.R, Aug

1990, McDonald 4615 & Bean (BRI); Melaleuca creek

scrub, ‘Rookwood’, Apr 1991, Forster PIF7937 &

McDonald (BRI, QRS); ‘Moorang’, 30.7 km ENE of

Taroom, Nov 1996, Halford Q3110 & Dowling (BRI);

Cannondale scrub, Expedition N.R, Amphitheatre section,

Nov 1998, Forster PIF23841 & Booth (BRI, MEL, QRS);

20 km S of Injune, Dec 1999, McDonald KRM165 (BRI).

Port Curtis District: Ogmore, Sep 1943, Blake 15312

(BRI); regional experiment Station, Biloela, Feb 1957,

Daniels 30 (BRI); c. 29 km SW of Ridgelands, Fitzroy Shire,

Apr 1990, Anderson 4847 (BRI); 15 km NE of Biloela, 3

km N of Callide Dam, lul 1992, Thompson BIL7 (AD, BRI,

NSW); near Gumigil Mine, Marlborough, Dec 1998,

Batianoff 981227 et al. (BRI). Warrego District: 13 km

NE of ‘Etona’, lul 1977, Purdie 734D (BRI). Maranoa

District: Combabula S.F., 28 km NE of Yuleba, lul 1990,

Warrian CMW518 (BRI); ‘Wombil Downs’ station, c. 40

km NW of Dirranbandi, Dec 1999, Franks AJF9911046

(BRI); Surat-Yuleba road, c. 17 km from Surat, Apr 2001,

Bean 17649 & Pedley (BRI, PRE). Darling Downs District:

Brookvale Park, 8 km from Oakey, lun 1993, Alcock 11265

(AD, BRI, CANB, K, MO); 14 km N of Goondiwindi,

towards Moonie, Feb 1996, Bean 9904 (BRI); 2.0 km S of

‘Wyaga’,NE of Goondiwindi, Nov 1999, Bean 15873 (BRI,

MEL, MO, NSW); 7.8 km N of Miles, Oct 2000, McDonald

KRM249 (BRI). New South Wales. Northwestern Slopes:

Boronga Nature Reserve, 14.4 km E of Boomi, Sep 2001,

Bean 17890 (BRI, MEL, NSW); 13.7 km E of North Star,

700

Austrobaileya 6 (4): 639-816 (2004)

Sep 2001, Bean 17902 (BRI). North West Plains: 8 miles

[13km] from Collarenebri, ontheroadtoWalgett, Nov 1967,

McGillivray 2803 (NSW); Watervale Reserve, 16 miles [26

km] N of Moree, Oct 1968, McBarron 15727 (BRI, NSW);

‘Warivan’, 7.4 km S of North Star on road to Warialda, Sep

1988, Moore 8833 (CANB).

Distribution and habitat : Solanumparvifolium

ssp. parvifolium is widespread from the north¬

western plains of N.S.W. to Charters Towers

in Queensland (Map 6). It grows in Brigalow

scrubs, vine thickets and in shrubby eucalypt

woodland, on a variety of soils.

Phenology: Flowers and fruits may be found

at any time of the year.

Notes: Most closely related to S. stelligerum,

and apparently intergrading with it in the

Rockhampton area, and also in the Bunya

Mtns-Warwick area. S. parvifolium ssp.

parvifolium differs by its narrower leaves,

3.7-7.5 times longer than broad (vs. 1.8-3.5

times for S. stelligerum ), tomentum not rusty

(vs. at least some rusty coloured stellae on leaf

undersides for S. stelligerum ) and with stellae

central ray (leaf undersides) 0.2-1 times as long

as laterals (vs. 1.5-4 times as long as laterals

for S. stelligerum ), and the lack of long-stalked

stellae on midrib of leaf undersides.

Conservation status : Widespread. Not

considered at risk.

19b. Solanum parvifolium subsp. tropicum

A.R.Bean subsp. nov. affinis S. parvifolio

sens. str. sed foliis longioribus

latioribusque, in pagina superiore folii

pilis Type 2 et indumento stellato induta,

stylo longiore et praesentia constante

florum masculorum in inflorescentia

differens. Typus: Queensland. North

Kennedy District: Herberton water

Supply Weir, Wild River head, Moomin

area, 25 April 2002, PI. Forster 28670

(holo: BRI (2 sheets + spirit); iso: AD,

K, L, MEL, MO, NSW).

Erect, rhizomatous perennial shrub, 0.6-1.5 m

high. Juvenile stage unknown. Adult branchlets

grey or brown or green; prickles 1-20 per

decimetre, straight, acicular, 4-7 mm long, 8-18

times longer than wide; stellae sparse to dense,

0.3-0.6 mm diameter, stalks 0-0.2 mm long;

lateral rays 6-8, porrect; central ray 0.1-0.8

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Adult

leaves linear or narrow lanceolate, entire;

lamina 7.5-13.5 cm long, 1.3-3.5 cm wide,

3.3-7 times longer than broad, apex acute, base

cuneate or obtuse, oblique part 0-2 mm long,

obliqueness index 0-3 percent; petioles 0.7-1.3

cm long, 7-13% length of lamina, prickles

absent. Upper leaf surface green; prickles

absent; stellate hairs distributed throughout;

protostellae absent; ordinary stellae very sparse

to moderate density, 0.3-3 mm apart, 0.3-0.5

mm across, sessile; lateral rays 6-8, porrect;

central ray 0.7-1.5 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs present throughout, 0.1-1.5 mm apart.

Lower leaf surface white; prickles absent;

stellae very dense, c. 0.05 mm apart, 0.4-0.6

mm diameter, sessile; lateral rays 7 or 8,

porrect; central ray 0.3-1 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Inflorescence supra-

axillary, pseudo-racemose, common peduncle

0-6 mm long, rachis prickles absent; 2-6-

flowered, strongly or weakly andromonoecious,

flowers 4 or 5-merous; pedicels 14-24 mm long

at anthesis, markedly thicker distally, 0.4-0.8

mm thick at mid-point, prickles absent. Calyx

tube 1.5-3 mm long, lobes attenuate, 2-3.5 mm

long; prickles absent at anthesis; stellae dense

to very dense, yellow or brown or rusty, 0.3-0.5

mm across, sessile, lateral rays 7 or 8, central

ray 0.5-1.5 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Corolla purple, 9-15 mm long, rotate or

shallowly lobed, inner surface glabrous; anthers

4.5-6.5 mm long; ovary glabrous, or with

stellate and Type 2 hairs; functional style 7-9

mm long, erect, glabrous or with Type 2 hairs

only or with stellate and Type 2 hairs, stellae c.

0.4 mm across, central ray c. 0.5 times as long

as laterals. Fruiting calyx with lobes less than

half length of mature fruit, prickles absent.

Mature fruits 1 per inflorescence, globular, 5-7

mm diameter, red, 1-locular (septum absent or

incomplete); placenta not apparent; mesocarp

juicy, succulent; exocarp c. 0.3 mm thick;

pedicels 26-33 mm long in fruit, 0.5-1 mm

thick at mid-point; seeds pale yellow, 2-2.4 mm

long. Fig. 20.

Bean, Taxonomy of Solanum subg. Leptostemonum

701

QUEENSLAND HERBARIUM (BRI)

Ifd Km HMSSffl «l |,*ODM]

(is SKSIMflejWWE f?9SS.3M?«l

HlHwlsn Si*** War, WlH Sms' hart. tteomn <i»l

7*1 open rofHf at Mfoa&usnnii wryteu. Eucalyptus portuwisa &

S>«c*ip* gtomut V* an grante

Sufrs**u& to SOom taJL stef* pWUM *1 along atom to base,

WaofMJipto; Mt amafl, brighUwi safl*ta*hy

2 arm item

QUEENSLAND HERBARIUM i.BFin

Brisbane Australia

556990

, Queensland Herbarium (BREL

/ilMflTW 0 f

Sb/e*um pnrr//t>/iu*i iy>. 7hyHO/*i A.

cm. sf. Qtnfeer Zeo3

Fig. 20. Holotype of Solanum parvifolium subsp. tropicum.

702

Specimens examined: Queensland. Cook District: track

to Wallum Trig, Atherton Tablelands, Sep 1977, Powell 672

& Armstrong (BRI, NSW); Turkey Scrub, ‘Whitewater’, Jan

1993, Fensham 345 (BRI); Mt Baldy S.F., SW of Atherton,

Apr 2002, Bean 18869 & McDonald (BRI). North Kennedy

District: MtFox, Sep 1949, Clemens s.n. (BRI); 41 km from

‘Reedy Brook’ towards ‘Mt Fox’, Aug 1972, Gittins 2508

(BRI); ‘Jervoise’ Holding, May 1979, Hyland 9931 (BRI,

QRS); Forty Mile scrub N.R, 60 km SSW of Mt Garnet, Jan

1990, Batianoff 900133 & Smith (BRI); Mt Abbot, 50 km

W of Bowen, Mar 1992, Bean 4220 (BRI); Mt Moti,

Bluewater Range, WNW of Townsville, May 1996, Cumming

14700 (BRI); Hugh Nelson Range, SE of Herberton, Jan

1998, Jago 4641 & Wannan (BRI).

Distribution and habitat: Solanumparvifolium

ssp. tropicum is endemic to Queensland.

Extends from Mt Abbot (near Bowen) to

Atherton, and west to Forty Mile Scrub N.P.

and Jervoise Holding (Map 6). It inhabits

rainforest margins or “wet sclerophyH” forest

with shrubby understorey.

Phenology : Flowers are recorded from January

to April and for September; mature fruits for

January and August.

Notes : Typical S. parvifolium ssp. tropicum

differs from S. parvifolium s. str. by the longer

and broader leaves, the upper leaf surface with

Type 2 hairs, the longer pedicels, the ovary

either glabrous or with both stellate hairs and

Type 2 hairs, the longer style and the presence

of male flowers in the inflorescence. Some

collections, notably from the Forty Mile Scrub,

are morphologically somewhat intermediate

between the subspecies.

Conservation status : Widespread. Not

considered at risk.

Etymology : From the Fatin tropicus, meaning

tropical. This refers to the distribution of the

subspecies.

20. Solanum ferocissimum Findl. in T.Mitch.,

Three Exped. Australia 2: 58 (1838); S.

ferocissimum var. ferocissimum Domin,

Biblioth. Bot. 89: 580 (1929).iype: New

South Wales, near Burradorgang, 28

April 1836, T.L. Mitchell (holo: CGE; iso:

K, MEF).

Solanum leptophyllum F.Muell., Fragm. 2:

164 (1861). Types: between Mackenzie

and Dawson Rivers, F. Mueller (syn:

?MEF, n.v.); Castlereagh River, E.

Bowman (syn: ?MEF, n.v.); ad oppidulum

Austrobaileya 6 (4): 639-816 (2004)

Warwick, H. Beckler (syn: ?K, n.v.);

Barrier Range, H. Beckler (syn: ?K, n.v.);

Mt Murchison, J. Dallachy (syn: K,

MEF).

Solanum ferocissimum var. rectispinum

Domin, Biblioth. Bot. 89: 580 (1929).

Types: Dividing Range near Jericho,

Queensland, Mar 1910, K. Domin (syn:

PR, n.v.); Peels Range, New South Wales,

undated, Frazer (syn: BM, K, OXF, fide

Symon (1981)); Peels Range, New South

Wales, undated, A. Cunningham (syn:

BM, fide Symon (1981)).

Illustrations : Cunningham et al. (1981:

594); Symon (1981: 133)

Erect, rhizomatous perennial shrub, 0.4-1.5 m

high. Juvenile branchlets with c. 20 prickles

per dm; leaves (in outline) linear or linear-

hastate, with basal lobes only; lamina 7-12 cm

long, 0.8-1.2 cm wide. Adult branchlets white,

grey or brown; prickles 15-60 per decimetre,

straight, acicular, 2-10 mm long, 9-13 times

longer than wide; stellae sparse to very dense,

0.25-0.4 mm diameter, sessile; lateral rays 7

or 8, porrect; central ray absent or present, 0-0.8

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Adult

leaves linear, entire or with obtuse basal lobes

only; lamina 3-7.5 cm long, 0.2-0.7 cm wide,

9-26 times longer than broad, apex obtuse or

acute, base cuneate or hastate, oblique part 0-1

mm long, obliqueness index 0-2 percent;

petioles 0.4-1 cm long, 8-20% length of

lamina, prickles absent. Upper leaf surface

green; prickles 2-6, straight, acicular, 2-9 mm

long, prickles present on midvein only; stellate

hairs confined to midrib, or distributed

throughout; protostellae absent; ordinary stellae

very sparse to sparse, 0.4-0.8 mm apart, 0.25-0.35

mm across, sessile; lateral rays 4-8, porrect;

central ray 0-1 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Lower leaf surface green to white or

yellowish; prickles 0-4, straight, acicular,

absent or present on midvein only; stellae sparse

to very dense, 0.05-0.7 mm apart, 0.2-0.7 mm

diameter, sessile; lateral rays 6-8, porrect;

central ray 0-0.5 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Inflorescence supra-axillary, pseudo-

racemose, common peduncle 0-7 mm long,

Bean, Taxonomy of Solanum subg. Leptostemonum

rachis prickles absent; 2-10-flowered, strongly

or weakly andromonoecious or with all flowers

bisexual, flowers 4 or 5-merous; pedicels 6-13

mm long at anthesis, same thickness

throughout, 0.2-0.4 mm thick at mid-point,

prickles absent. Calyx tube 1-2 mm long, lobes

deltate or rostrate, 0.7-1.5 mm long; prickles

absent at anthesis; stellae moderate to very

dense, yellow, 0.3-0.5 mm across, sessile,

lateral rays 7 or 8, central ray 0.5-1 times as

long as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Corolla white or

mauve, 6-12 mm long, deeply lobed, inner

surface glabrous; anthers 3.5-5.5 mm long;

ovary glabrous, or with stellate and Type 2

hairs; functional style 4.5-6.5 mm long, erect,

glabrous or with stellate and Type 2 hairs,

stellae c. 0.4 mm across, lateral rays 7 or 8,

central ray 0.5-1 times as long as laterals.

Fruiting calyx with lobes less than half length

of mature fruit, prickles absent. Mature fruits

1-4 per inflorescence, globular, 6-9 mm

diameter, red, glabrous or with a few scattered

Type 2 hairs, 1-locular (septum absent or

incomplete); placenta not apparent; mesocarp

juicy, succulent; exocarp c. 0.1 mm thick;

pedicels 13-16 mm long in fruit, 0.5-0.6 mm

thick at mid-point; seeds pale yellow, 3-3.6 mm

long.

Specimens examined : Queensland. Burke District:

Southern Roads, 20 km N of Mt Isa, Dec 1996, Barrs SB93

(BRI). Gregory North District: S of Selwyn, May 1963,

Gittins 700 (BRI). Mitchell District: ‘Oxenhope’ on the

Alroy road, c. 130 km SSW of Longreach, Apr 1989, Emmott

285 (BRI); Blacks Palace, 80 km S of Jericho, May 1989,

Cheffins 377 (BRI). Leichhardt District: c. 9 km SW of

Anakie, Sep 1984, Anderson 3816 (BRI); Telecom road, 14

km E of Comet, Mar 1994, Bean 7524 & Forster (BRI).

WarregoDistrict: ‘GilruthPlains’, Cunnamulla,Apr 1963,

McKee 10347 (BRI); 19 km S of Charleville along road to

Wyandra, Mar 1976, Purdie 215 & Boy land (BRI); 15.1

km S of Corona Creek, on Quilpie-Adavale road, Aug 1990,

Prendergast HDP283 (BRI); Monks Tank road, Idalia N.P,

Feb 2000, Nicholls SN010 (BRI). Maranoa District:

‘Boa t man’ Station, Mar 1947, Everist 2780 (BRI); ‘Glade

Villa’, S of Roma, Aug 1990, Warrian CMW543 (BRI,

NSW); c. 41 km along Shirlo road, NW of Bollon, Mar 2001,

Bean 17540 (BRI, MEL); near ‘Boxleigh’, c. 60 km NE of

St George, Apr 2001, Bean 17641 & Pedley (BRI, MO).

Darling Downs District: Myall Park, 4 m il es [6 km] NW of

Glenmorgan, Apr 1960, Johnson 1605 (BRI); 5 miles [8 km]

W of Westmar, Jul 1961, Pedley 788 (BRI); ‘Coomrith’ area

near Meandarra, Jul 1969, Webb & Tracey 8298 (BRI);

Yelarbon, around cemetery, c. 1 km from P.O., Oct 1983,

Canning 5821 & Rimes (BRI, CANB, NSW); 2.0 km S of

‘Wyaga’, NE of Goondiwindi, Nov 1999, Bean 15868 (AD,

BRI, MEL, NSW).

703

Distribution and habitat: Solanum

ferocissimum is distributed throughout much

of inland Queensland south of about 20° latitude

(Map 9), and extending well into New South

Wales and Northern Territory. Also recorded

by Symon (1981) for Western Australia. It

inhabits stony ridges or flats, on red earths or

loamy soil, in woodlands often dominated by

Eucalyptus populnea, E. melanophloia or

Acacia aneura.

Phenology : Flowers and fruits may be found

at any time of the year, probably in response to

rainfall.

Notes: S. ferocissimum is close to S. parvifolium

ssp. parvifolium, but differs by the often sparse,

sessile indumentum; leaves linear in shape,

frequently with a pair of basal leaf lobes, with

prickles along the midrib on both surfaces; and

by the seeds 3.0-3.6 mm long (1.9-2.3 mm long

for S. parvifolium ssp. parvifolium ).

Ross (1986) suggested that S. ferocissimum

is doubtfully distinct from S. parvifolium. While

it is true that some collections are taxonomically

difficult, the great majority of collections may

be easily determined.

Conservation status : Widespread. Not

considered at risk.

21. Solanum latens A.R.Bean sp. nov. Frutex

parvus; aculei in ramulis frequentes,

aciculares; folia adulta parva, 4.7-7plo

longiora quam latiora, saepe lobata,

aliquando integra; pili stellati in

superficiebus ambabus folii sessiles, radio

centrali radiis lateralibus 1.5-3plo

longiore; corolla profunde lobata; calyx

aculeis carens; fructus maturitate rubri,

seminibus 1.6-2.3 mm longis. Typus:

Queensland. Burnett District: Conservation

Gully, ‘Narayen’, W of Mundubbera, 16

December 2001, A.R. Bean 18292 (holo:

BRI (2 sheets + spirit); iso: AD, MEL, NSW).

Solanum sp. (Kingaroy A.R. Bean 17428) on

BRI database

Erect, rhizomatous perennial shrub, 0.4-1.1 m

high. Juvenile branchlets with 35-50 prickles

per dm, 3-6 mm long; leaves (in outline) linear-

hastate, shallowly to deeply lobed, with 1 or 2

704

Austrobaileya 6 (4): 639-816 (2004)

Fig. 21. Solanum latens. A. fruiting branchlet x 1.2. B. style and ovary x 6. C. ovary showing Type 2 hairs x 30. D. stellate

hair from upper leaf surface x 60. E. mature fruit and pedicel x 3. F. transverse section of fruit x 6. A, D-F, Bean 17432; B-

C ,Bean 18295.

Bean, Taxonomy of Solanum subg. Leptostemonum

pairs of lobes; lamina 2.5-4.5 cm long, 0.5-1.3

cm wide, with 9-16 prickles on upper surface.

Adult branchlets green; prickles 15-30 per

decimetre, straight, acicular, 5-9 mm long, 13-18

times longer than wide; stellae sparse, 0.25-0.5

mm diameter, sessile; lateral rays 4-7, porrect;

central ray 0.5-1 t im es as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

sparse. Adult leaves narrow lanceolate or

lanceolate, entire or shallowly lobed throughout

or with basal lobes only; lobes 1 or 2 on each

side, obtuse, lobing index 1-1.2; lamina 1.5-4.5

cm long, 0.3-0.8 cm wide, 4.5-7.5 times longer

than broad, apex obtuse or acute, base cuneate

or hastate, oblique part 0-2.5 mm long,

obliqueness index 0-6 percent; petioles 0.3-0.6

cm long, 8-16% length of lamina, prickles

absent. Upper leaf surf ace green; prickles 2-6,

straight, acicular, 4-6 mm long, prickles

present on midvein only; stellate hairs confined

to midrib, or distributed throughout;

protostellae absent; ordinary stellae very sparse

to sparse, 0.8-2.5 mm apart, 0.35-0.5 mm

across, sessile; lateral rays 4-7, porrect; central

ray 1.5-3 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Lower leaf surface green; prickles 0-3, straight,

acicular, absent or present on midvein only;

stellae very sparse to moderate, 0.3-2 mm apart,

0.3-0.7 mm diameter, sessile; lateral rays 6-8,

porrect; central ray 1.5-3 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Inflorescence supra-

axillary, solitary or pseudo-umbellate, common

peduncle 0-2 mm long; 1-3-flowered, with all

flowers bisexual and 5-merous; pedicels 8-18

mm long at anthesis, same thickness

throughout, c. 0.3 mm thick at mid-point,

prickles absent or present. Calyx tube 1.5-2

mm long, lobes attenuate, 1.5-2.5 mm long;

prickles absent at anthesis; stellae sparse to

moderate, transparent, 0.2-0.4 mm across,

sessile, lateral rays 6-8, central ray 0.8-1.5

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Corolla

mauve, 6-8 mm long, deeply lobed, inner

surface glabrous; anthers 3-4.5 mm long; ovary

with Type 2 hairs only; functional style 4-6

mm long, erect, with Type 2 hairs only. Fruiting

calyx with lobes less than half length of mature

fruit, prickles absent. Mature fruits 1 per

inflorescence, globular, 4-6.5 mm diameter,

red, glabrous or with a few scattered Type 2

hairs, 1-locular (septum absent or incomplete);

705

placenta not apparent; mesocarp juicy,

succulent; exocarp 0.2-0.3 mm thick; pedicels

14-24 mm long in fruit, 0.5-0.6 mm thick at

mid-point; seeds pale yellow, 1.6-2.3 mm long.

Fig. 21.

Specimens examined : Queensland. Leichhardt District:

‘Kooralbyn’, c. 25 km S of Duaringa, Feb 2003, Bean 19985

(BRI, CANB, MO). Burnett District: Kingaroy, Apr 1947,

Smith 3004 (BRI); c. 14 km NNE of Eidsvold, Oct 1993,

Lepschi & Slee 1216 (BRI, CANB); Semgreen Road, SSE

of Kingaroy, Mar 2001, Bean 17428 (BRI, CANB, MEL,

NSW); Goodger Gully Rd, SSE of Kingaroy, Mar 2001, Bean

17432 (BRI, NSW); Kunioon West, S of Kingaroy, Nov 2001,

Bean 18121 (BRI); Valley Dam paddock, ‘Narayen’, W of

Mundubbera, Dec 2001, Bean 18295 (BRI, MEL, NSW);

S.F.227, c. 30 km E of Cracow, Jul 2002, Bean 19129 (B,

BRI, CANB, L, MEL); S.F.132 Allies Creek, c. 55 km S of

Mundubbera, Nov 2002, Bean 19610 (BRI, MEL). Darling

Downs District: Burraburri Creek, 16 km W of Durong, May

1992, Forster PIF9857 (AD, BRI); 3.8 km NE of Warra, on

Marnhull road, Feb 2003, Bean 19961 (BRI, NSW).

Distribution and habitat: Solanum latens is

endemic to Queensland. It is confined to

subcoastal south-eastern Queensland,

extending from Duaringa to Warra, and near

Kingaroy. (Map 9). It occurs as an understorey

plant in Brigalow-Belah communities, or in

microphyll vine forest, or in Eucalyptus or

Acacia dominated woodland on tertiary-aged

plateaux.

Phenology: Flowers are recorded for February,

May, November and December; mature fruits

in May and December

Notes: Closely related to S. ferocissimum, but

differing by the smaller and broader leaves;

juvenile leaves often with 2 pairs of lobes; upper

and lower leaf stellae with central ray 1.5-3

times as long as laterals (0-1 times for

S. ferocissimum)', the 1-3 flowered inflo rescences

(2-10 flowered for S. ferocissimum ); and the seeds

1.6-2.3 mm long (3.0-3.6 mm for S. ferocissimum).

Conservation status: Moderately widespread.

Not considered at risk.

Etymology: From the Latin latens meaning

‘secret’ or ‘hidden’. This is a reference to the

late recognition of the species, with most

collections being in the last decade.

22. Solanum dissectum Symon, Austrobaileya

4:432-3 (1995). Type: Queensland. Port

Curtis District: west of Thangool, 2 July

1959, R. W. Johnson 858 (holo: BRI; iso: BRI).

706

Illustration : Symon (1995: 432)

Erect, rhizomatous perennial shrub, 0.3-0.8 m

high. Juvenile branchlets with c. 7 prickles per

dm, 7-10 mm long; leaves (in outline) ovate,

deeply lobed, with 2-5 pairs of lobes; lamina

5.5- 7 cm long, 3-4 cm wide, with 2-3 prickles

on upper surface. Adult branchlets grey or

brown; prickles 3-10 per decimetre, straight,

acicular, 4-11 mm long, 9-13 times longer than

wide; stellae absent; finger hairs absent; Type

2 hairs absent. Adult leaves ovate or broadly

ovate, deeply lobed throughout; lobes 2-4 on

each side, acute or obtuse, lobing index 7-22;

lamina 2-5.5 cm long, 0.9-2.5 cm wide,

1.5- 2.9 times longer than broad, apex obtuse

or acute, base cuneate or obtuse, oblique part

0-1.5 mm long, obliqueness index 0-5 percent;

petioles 0.3-1.2 cm long, 12-25% length of

lamina, prickles absent or present. Upper leaf

surface green; prickles 0-4, straight, acicular,

1- 7 mm long, prickles absent or present on

midvein only; stellate hairs absent; finger hairs

absent; Type 2 hairs present only in vein

depressions. Lower leaf surface green; prickles

0-4, straight, acicular, absent or present on

midvein only; stellae absent; finger hairs

absent; Type 2 hairs absent. Inflorescence

supra-axillary, pseudo-racemose, common

peduncle 0-2 mm long, rachis prickles absent;

2- 5-flowered, with all flowers bisexual and 5-

merous; pedicels 6-8 mm long at anthesis,

markedly thicker distally, 0.4-0.5 mm thick at

mid-point, prickles absent. Calyx tube 1-2 mm

long, lobes rostrate, 1-2.5 mm long; prickles

absent at anthesis; stellae absent; finger hairs

absent; Type 2 hairs absent. Corolla mauve,

6-9 mm long, deeply lobed, inner surface

glabrous; anthers 2.5-4 mm long; ovary with

Type 2 hairs only; functional style 4-6.5 mm

long, erect, glabrous. Fruiting calyx with lobes

less than half length of mature fruit, prickles

absent. Mature fruits 1 per inflorescence,

globular, 7-9 mm diameter, red, 1-locular

(septum absent or incomplete); placenta not

apparent; mesocarp juicy, succulent; exocarp

0.2-0.3 mm thick; pedicels 10-19 mm long in

fruit, 0.5-1 mm thick at mid-point; seeds pale

yellow, 3-4.1 mm long.

Specimens examined : Queensland. Leichhardt District:

McCrae property, 50 miles [80 km] S of Duaringa, Jul 1966,

Everist & McDonald 3 (BRI); Portion 2, Parish of Capayan,

Banana Shire, Feb 1989, Philips s.n. (BRI); ‘Nirvana’, c. 15

Austrobaileya 6 (4): 639-816 (2004)

km WNW of Banana, Apr 2003, Bean 20167 (BRI). Port

Curtis District: c. 6 miles [10 km] W of Biloela, Jul 1959,

Johnson 870 (BRI); Biloela near roadside, Sep 1966, Stevens

s.n. (BRI); c. 22kmS of Dululu, Sep 1989, Clamfield4116

(BRI).

Distribution and habitat : Solanum dissectum

is endemic to Queensland, and recorded from

the Biloela-Banana-Baralaba area (Map 11).

It occurs in association with brigalow ( Acacia

harpophylla ), and sometimes Eucalyptus

thozetiana, on heavy cracking-clay soil.

Phenology : Poorly known. Flowers are

recorded for September; mature fruits in

February, April and July.

Notes: S. dissectum is a very distinct species

with no very close relatives. It is probably

closest to S. lythrocarpum (see notes under that

species). It is also close to S.ferocissimum, but

it differs from that species by the total lack of

stellate hairs and the deeply lobed adult leaves.

Conservation status: S. dissectum is currently

known from a total of 17 mature individuals at

one locality, and this locality is not protected

within a conservation reserve. Applying the

IUCN guidelines (IUCN. 2001), a category of

“Critically Endangered” is recommended (CR

A3ce; Blab(iii,v)+2ab(iii,v); Cl+2a(i,ii); D).

Three specimen labels include comments

indicating the probable imminent demise of that

population of S. dissectum viz . “cleared

Brigalow scrub”, “pulled Acacia harpophylla

regrowth”, “recently burnt, pulled brigalow

suckers”. Major threats are continuing land

clearance, and invasion of habitat by exotic

species of grass, introduced as cattle fodder.

This species is undoubtedly the one closest to

extinction in Queensland.

23. Solanum lythrocarpum A.R.Bean sp. nov.

Frutex parvus, aculei in ramulis

praesentes sed sparsi; folia integra,

angusto-lanceolata, utrinque laete viridia;

pili Type-2 in ramulis folii lamina

calyceque; pili stellati in foliis et calyce

sparsissimus; aculei calycis absentes; styli

5.7-6.8 mm longi; fructus maturitate

rubra. Typus: Queensland. Burnett

District: east of Scrubby Dam,

Coominglah State Forest, near Monto, 11

December 1998, A.R. Bean 14430 (holo:

BRI (1 sheet + spirit); iso: AD, MEF, NSW).

Bean, Taxonomy of Solanum subg. Leptostemonum

707

Fig. 22. Solanum lythrocarpum.A. flowering and fruiting branchlet x 1. B. flower showing style and anthers x 3. C. flower

showing hypanthium and calyx lobes x 3. D. ovary and style x 6. E. mature fruit and pedicel x 2. F. transverse section of fruit

x 3. A, Bean 10389; B-D, Bean 14430; E-F, Bean 15936.

708

Solanum sp. (Coominglah A.R. Bean 10389)

in Henderson (2002).

Erect, rhizomatous perennial shrub, 0.3-0.9 m

high. Juvenile stage unknown. Adult branchlets

brown; prickles absent or present, 0-6 per

decimetre, straight, acicular, 0.5-5 mm long,

7-10 times longer than wide; stellae absent or

sparse, 0.2-0.3 mm diameter, stalks 0-0.1 mm

long; lateral rays 4—8, porrect; central ray 0.5-1.5

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs sparse to dense.

Adult leaves narrow lanceolate or lanceolate,

entire; lamina 5-9.5 cm long, 1.1-1.5 cm wide,

4.7-7 times longer than broad, apex acute, base

cuneate, oblique part 0-2.5 mm long,

obliqueness index 0-3 percent; petioles 0.9-2.1

cm long, 11-23% length of lamina, prickles

absent or present. Upper leaf surface green;

prickles 0-4, straight, acicular, 3-7 mm long,

prickles absent or present on midvein only;

stellate hairs absent, or confined to midrib;

ordinary stellae absent from surface; finger

hairs absent; Type 2 hairs present throughout,

0.1-0.5 mm apart. Lower leaf surface green;

prickles 0-3, straight, acicular, absent or

present on midvein only; stellae absent; finger

hairs absent; Type 2 hairs absent. Inflorescence

leaf-opposed, pseudo-racemose, common

peduncle 0-7 mm long, rachis prickles absent;

3-8-flowered, with all flowers bisexual and 4

or 5-merous; pedicels 9-24 mm long at

anthesis, same thickness throughout or

markedly thicker distally, 0.4-0.7 mm thick at

mid-point, prickles absent. Calyx tube 1.5-2.5

mm long, lobes attenuate, 2-6 mm long;

prickles absent at anthesis; stellae absent to

sparse, white, 0.15-0.3 mm across, sessile,

lateral rays 4-8, central ray 0-1 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs present. Corolla purple,

7-11 mm long, shallowly or deeply lobed, inner

surface glabrous; anthers 3-5 mm long; ovary

glabrous, or with Type 2 hairs only; functional

style 5.5-7 mm long, erect, glabrous. Fruiting

calyx with lobes less than half length of mature

fruit, prickles absent. Mature fruits 1-4 per

inflorescence, globular, 8-11 mm diameter, red,

1-locular (septum absent or incomplete); placenta

not apparent; mesocarp juicy, succulent; exocarp

0.4-0.5 mm thick; pedicels 18-24 mm long in

fruit, 0.6-0.8 mm thick at mid-point; seeds pale

yellow, 3-3.7 mm long. Fig. 22.

Austrobaileya 6 (4): 639-816 (2004)

Specimens examined : Queensland. Burnett District:

Sixteen Mile L.A., Coominglah S.F., south-west of Monto,

Mar 1996, Bean 10155 (BRI); W edge of Bogdanoff L.A.,

Coominglah S.F., south-west of Monto, Jun 1996, Bean

10389 (BRI, NSW, MEL); northern boundary of S.F. 132, S

of Mundubbera, Nov 2002, Bean 19627 (BRI).

Distribution and habitat : Solanum

lythrocarpum is endemic to Queensland. It is

known from two small areas near the towns of

Monto and Mundubbera (Map 10). It grows

on lateritised plateaux in ironbark -Acacia

blakei forest with a dense shrubby understorey

including some rainforest species. Associated

species include Croton insularis, Phebalium

nottii, Bertya opponens and Philotheca ciliata.

Phenology: Flowers are recorded for June and

December; mature fruits recorded for March,

November and December.

Notes: S. lythrocarpum is related to

S. dissectum, but differs by the entire adult

leaves (deeply lobed for S. dissectum)', the

presence of Type 2 hairs on the branchlets, leaf

lamina and calyx (absent for S. dissectum ); the

presence of stellate hairs on leaves and calyx

(absent for S. dissectum ) and the fruiting

pedicels 10-19 mm long (18-24 mm long for

S. dissectum ).

Conservation status: S. lythrocarpum known

from two localities, neither of which is within

a conservation reserve. Applying the IUCN

guidelines (IUCN. 2001), a category of

“Vulnerable” is recommended (VU Dl+2). In

the Coominglah State Forest, less than 300

plants are known; 30-40 plants occur at the

Mundubbera site.

Etymology: From the Greek ‘ lythron ’ meaning

blood, and ‘ carpos ’ (fruit), in reference to the

bright-red colour of the mature fruits.

24. Solanum chenopodinum F.Muell., Fragm.

2: 165 (1861). Type: New South Wales.

“Mount Murchison and Darling”,

undated, J. Dallachy s.n. (lecto: MEF

[MEF 11705], fide Symon 1981).

Illustrations: Cunningham et al. (1981:

594); Symon (1981: 149)

Erect, rhizomatous perennial shrub, 0.4-1 m

high. Juvenile branchlets with c. 7 prickles per

dm, c. 6 mm long; leaves (in outline) hastate,

Bean, Taxonomy of Solanum subg. Leptostemonum

shallowly-lobed or with basal lobes only, with

1 or 2 pairs of lobes; lamina 8-11.5 cm long,

4.5-6 cm wide, without prickles on upper

surface. Adult branchlets white or grey or

yellow; prickles absent or present, 0-10 per

decimetre, straight or curved, broad-based,

2-7 mm long, 4-7 times longer than wide;

stellae very dense, 0.2-0.5 mm diameter, stalks

0-0.2 mm long; lateral rays 7-10, porrect or

ascending; central ray present, 0.2-1 times as

long as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Adult leaves ovate

or triangular, with obtuse basal lobes only;

lamina 2.5-6.5 cm long, 1.1-2.6 cm wide, 1.4-3

times longer than broad, apex obtuse or acute,

base cuneate, obtuse, cordate or hastate, oblique

part 0-4 mm long, obliqueness index 0-10

percent; petioles 0.5-2 cm long, 15-35% length

of lamina, prickles absent. Upper leaf surface

green; prickles absent; stellate hairs distributed

throughout; protostellae absent; ordinary stellae

very sparse to sparse, 0.4-0.7 mm apart, 0.2-0.4

mm across, sessile; lateral rays 5-10, porrect;

central ray 0.7-1.5 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Lower leaf surface white or

yellowish; prickles absent; stellae dense to very

dense, 0.05-0.1 mm apart, 0.4-0.5 mm diameter,

stalks 0-0.1 mm long; lateral rays 8-13, porrect;

central ray 0.5-1 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Inflorescence supra-axillary, pseudo-

racemose, common peduncle 0-10 mm long,

rachis prickles absent; 6-20-flowered, with all

flowers bisexual and 5-merous; pedicels 4-6 mm

long at anthesis, same thickness throughout,

0.5-0.7 mm thick at mid-point, prickles absent.

Calyx tube 1.5-2 mm long, lobes deltate, 1-2.5

mm long; prickles absent at anthesis; stellae

dense to very dense, white, 0.25-0.4 mm across,

stalks 0-0.1 mm long, lateral rays 8-10, central

ray 0.7-1.5 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Corolla mauve or purple, 5-8 mm long, deeply

lobed, inner surface glabrous; anthers 3.5-4.5

mm long; ovary glabrous; functional style 5.5-7

mm long, erect, glabrous. Fruiting calyx with

lobes less than or more than half length of mature

fruit, prickles absent. Mature fruits 1-11 per

inflorescence, globular, 7-8 mm diameter, red;

mesocarp juicy, succulent; pedicels 8-12 mm

long in fruit, 0.7-1 mm thick at mid-point; seeds

pale yellow, 3.5-3.8 mm long.

709

Specimens examined : Queensland. Gregory North

District: ‘Currawilla’, Jun 1947, Everist 4025 (BRI); Spring

Creek, 11 km S of ‘Warra’, Jun 1978, Purdie 1243 (BRI);

Simpson Desert, c. 20 km NW of Pulchera Waterhole, Aug

1978, Jahnke 1310 (BRI); Warracoola Waterhole,

DiamantinaN.R,Apr 1995, Mitchell 95\ (BRI); near Mistake

Hut dam, Bladensberg N.R, S of Winton, Mar 1998, Forster

PIF22228 & Booth (BRI). Gregory South District:

Nockatunga, Jun 1936, Blake 11880 (BRI); Murungeri

waterhole, ‘NappaMerrie’ Station, Jun 1988, Conrick 2325

(AD, BRI).

Distribution and habitat: Solanum

chenopodinum is found in arid to semi-arid

areas of south-west Queensland (Map 11). Also

occurs widely in low rainfall parts of New South

Wales, South Australia and Northern Territory.

It occurs in shrubland on flats or watercourses

on sandy or clayey soil.

Phenology: Flowers are recorded in March,

June and August; mature fruits from March to

June.

Notes: This species and S. ferocissimum are

the only red-fruited Solanum species that occur

in arid parts of Australia. Both species may have

a hastate leaf base, but S. chenopodinum differs

by the much broader leaves without prickles

on the upper surface, the broad-based prickles

on the branchlets, and the shorter fruiting

pedicels.

Conservation status: Data deficient.

25. Solanum dysprosium A.R.Bean sp. nov.

Frutex; aculei ramuli leviter vel distincte

recurvi; ramuli classibus duabus pilorum

stellatorum, longitudine radii centralis

differentibus; folia adulta 4 vel 5 paribus

loborum non profundorum, petiolis

aculeos gerentibus et longitudine 19-33%

laminae aequantibus; stellae in pagina

inferiore folii 0.5-0.8 mm diametro; digiti

apicibus glandularibus in calyce

praesentes. Typus: Queensland. Cook

District: Cape Melville National Park,

western slopes, 26 November 2001, K.R.

McDonald 1026 & H. Hines (holo: BRI).

Erect, rhizomatous perennial shrub, c. 0.75 m

high. Juvenile stage unknown. Adult branchlets

mauve or brown; prickles 25-50 per decimetre,

straight, acicular, 3-8 mm long, 9-12 times

longer than wide; stellae sparse, 0.25-0.5 mm

diameter, stalks 0-0.1 mm long; lateral rays

710

Austrobaileya 6 (4): 639-816 (2004)

Fig. 23. Holotype of Solanum dysprosium.

Bean, Taxonomy of Solanum subg. Leptostemonum

4-8, porrect; central ray absent or present, 0-4

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs sparse. Adult

leaves ovate, shallowly lobed throughout; lobes

4 or 5 on each side, acute, lobing index 1.3-

1.6; lamina 7-10.5 cm long, 3.5-5 cm wide,

1.9-2.2 t im es longer than broad, apex acute or

acuminate, base obtuse, oblique part 0-5 mm

long, obliqueness index 0-6 percent; petioles

1.3-3.5 cm long, 20-35% length of lamina,

prickles present. Upper leaf surface green;

prickles 30-50, straight, acicular, 2-8 mm long,

prickles present on midvein and lateral veins;

stellate hairs distributed throughout;

protostellae present; ordinary stellae sparse,

0.5-0.8 mm apart, 0.25-0.5 mm across, sessile;

lateral rays 4 or 5, porrect; central ray 0.5-3

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Lower

leaf surface green; prickles 11-17, straight,

acicular, present on midvein and lateral veins;

stellae moderate, 0.4-0.8 mm apart, 0.5-0.8

mm diameter, sessile; lateral rays 4-6, porrect;

central ray 0-1 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Inflorescence supra-axillary, pseudo-

racemose, common peduncle 17-35 mm long,

rachis prickles present; 12-15-flowered,

flowers 5-merous; pedicels 8-11 mm long at

anthesis, same thickness throughout, 0.3-0.6

mm thick at mid-point, prickles absent. Calyx

tube 2-2.5 mm long, lobes attenuate, 2-3.5 mm

long; prickles absent at anthesis; stellae sparse,

transparent, 0.15-0.2 mm across, sessile, lateral

rays 4 or 5, central ray 1-2 times as long as

laterals, not gland-tipped; finger hairs

abundant; Type 2 hairs absent. Corolla purple,

10-14 mm long, shallowly or deeply lobed;

anthers 5-5.8 mm long; functional style erect,

glabrous. Fruiting calyx with lobes less than

half length of mature fruit, prickles absent.

Mature fruits 1 per inflorescence, globular, c.

7 mm diameter; pedicels c. 15 mm long in fruit,

c. 0.7 mm thick at mid-point; seeds pale yellow,

2.5-3 mm long. Fig. 23.

Specimen examined : Queensland. Cook District: Cape

Melville N.P., western slopes, Nov. 2001, McDonald 1026

& Hines (BRI).

Distribution and habitat : Solanum dysprosium

is endemic to Queensland. Confined to Cape

Melville on Cape York Peninsula (Map 9), it

grows on the edge of a granite boulder-field,

adjacent to vine thicket.

711

Notes: Closely related to S. inaequilaterum, but

differing by the smaller leaves, petioles 20-35%

of lamina length, and bearing prickles (petioles

10-22%, without prickles for S. inaequilaterum ),

stellae on lower leaf surface 0.5-0.8 mm diameter

(0.3-0.5 mm diameter for S. inaequilaterum ),

common peduncle present (absent for

S. inaequilaterum ), calyx lobes 2-3.5 mm

long (4-9 mm for S. inaequilaterum).

Conservation status: Data deficient.

Etymology: from the Greek dysprositos,

meaning hard to get at. This is a reference to

the difficulty in accessing the Cape Melville

area, where the species is endemic.

26. Solanum inaequilaterum Domin, Biblioth.

Bot. 89: 581-582 (1929). Type:

Queensland. Moreton District:

Beechmont, March 1910, K. Domin s.n.

(holo: ?PR), n.v.

Illustration: Symon (1981: 244)

Erect, rhizomatous perennial shrub, 1-3.5 m

high. Juvenile branchlets with 120-400 prickles

per dm, 1.5-6 mm long; leaves (in outline)

elliptical or ovate, deeply lobed, with 2-4 pairs

of lobes; lamina 16-24 cm long, 9-14 cm wide,

with 100-150 prickles on upper surface. Adult

branchlets mauve or brown; prickles 5-55 per

decimetre, straight, acicular, 2-9 mm long,

10-14 times longer than wide; stellae sparse,

0.4-0.5 mm diameter, stalks 0-0.1 mm long;

lateral rays 4-8, porrect; central ray absent or

present, 0-0.5 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Adult leaves ovate, shallowly lobed

throughout; lobes 3-5 on each side, acute,

lobing index 1.1-1.8; lamina 9.5-19 cm long,

3.5-7.5 cm wide, 2.3-2.7 t im es longer than

broad, apex acute or acuminate, base cuneate

or obtuse, oblique part 0-18 mm long,

obliqueness index 0-9 percent; petioles 1.1-2.2

cm long, 10-22% length of lamina, prickles

absent. Upper leaf surface green; prickles 8-40,

straight, acicular, 3-11 mm long, prickles

present on midvein and lateral veins; stellate

hairs distributed throughout; protostellae

present; ordinary stellae very sparse to sparse,

0.8-1.4 mm apart, 0.4-0.5 mm across, sessile;

lateral rays 4-8, porrect; central ray 0.5-1.5

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Lower

712

Austrobaileya 6 (4): 639-816 (2004)

leaf surface green; prickles 0-9, straight,

acicular, absent or present on midvein only or

present on midvein and lateral veins; stellae

sparse to moderate, 0.5-1.3 mm apart, 0.3-0.5

mm diameter, sessile; lateral rays 4—8, porrect;

central ray 0.3-1 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Inflorescence supra-axillary, pseudo-

racemose, common peduncle absent, rachis

prickles absent; 4-13-flowered, weakly

andromonoecious, flowers 5-merous; pedicels

7-24 mm long at anthesis, same thickness

throughout or markedly thicker distally, 0.5-1

mm thick at mid-point, prickles absent. Calyx

tube 1-3 mm long, lobes attenuate, 4-9 mm

long; prickles absent at anthesis; stellae sparse

to moderate, yellow, 0.15-0.25 mm across,

sessile, lateral rays 5-8, central ray 0.5-1.5

times as long as laterals, not gland-tipped;

finger hairs present; Type 2 hairs absent.

Corolla white or purple, 8-15 mm long,

shallowly or deeply lobed, inner surface

glabrous; anthers 4.5-6 mm long; ovary with

Type 2 hairs only; functional style 7.5-8.5 mm

long, erect, glabrous or with Type 2 hairs only.

Fruiting calyx with lobes less than half length

of mature fruit, prickles absent. Mature fruits

I- 4 per inflorescence, globular or ellipsoidal,

II- 13 mm diameter, red, 1-locular (septum

absent or incomplete); placenta not apparent;

mesocarp juicy, succulent; exocarp 0.4-0.5 mm

thick; pedicels 26-38 mm long in fruit, 0.6-2

mm thick at mid-point; seeds pale yellow,

2.6-3.4 mm long. Gin’s Whiskers.

Specimens examined : Queensland. Moreton District:

Roberts Plateau, Lamington N.P., May 1929, White 6074

(BRI); Levers Plateau, Apr 1972, Henderson H1286 (BRI);

beside track at Mt Nothofagus, Sep 1973, Lander 336 (BRI,

NSW); 0.5 km along Duck Creek road, near O’Reilly Guest

House, Dec 1999, Bean 15896 (AD, BRI, MEL, NSW); near

Best of All Lookout, Springbrook, Mar 2000, Forster

PIF25385 & Booth (A, BRI, MEL, QRS). New South Wales.

North Coast: Coopers Creek, viaMullumbimby, Aug 1936,

White 10456 (BRI); Victoria Park, 5 miles [8 km] S of

Alstonville, Feb 1971, O’Hara & Coveny 3491 (BRI, NSW);

summit of Mt Nardi, NE of Nimbin, Feb 2000, Bean 16021

(BRI, NSW); Big Scrub Flora Reserve, c. 20 km N of

Lismore, Dec 2000, Bean 17073 (BRI); off Dingo Flat road,

Clouds Creek S.F., N of Donigo, Jan 2001, Bean 17264 (BRI,

NSW).

Distribution and habitat : Solanum

inaequilaterum extends from Springbrook and

Lamington National Park in Queensland, to

Dorrigo in N.S.W. (Map 11). It grows in

notophyll rainforest in high rainfall areas.

Altitude is often above 750 metres. It is one of

the few species that will flower and fruit under

very low-light conditions.

Phenology: Flowers are recorded from

November to April; fruits may be found

throughout the year.

Notes: Closely related to S. dysprosium (see

Notes under that species). Less closely related

to S. semiarmatum, with which it shares the

deeply lobed juvenile leaves, dense prickles on

stems of juvenile plants, and the succulent 1-

locular fruit.

In 1920, Georg Bitter made the

combination Lycianthes inaequilatera (Rusby)

Bitter for a Bolivian species, based on Bassovia

inaequilatera Rusby in Mem. Torrey Bot. Club

6: 90 (1896). He included the notation

“Solanum inaequilaterum Rusby in sched.”, but

that combination was never actually published.

Since S. inaequilaterum Rusby is a nomen

nudum, it follows that S. inaequilaterum Domin

is a legitimate name.

The type of S. inaequilaterum was sought

from PR, but not received. Symon (1981) stated

that he had not seen the type. It is possible that

the type is missing, but the designation of a

neotype is not warranted until further searches

are made. The application of the name is not

in doubt, as the species is well described in the

protologue.

Conservation status: Not considered at risk.

Group 13A (S. semiarmatum group), here

defined; related to Group 13 ( S.

ferocissimum group) of Whalen (1984).

Mature fruits globular, black, juicy, 1-locular,

<12 mm diameter, exocarp <0.5 mm thick

(100%); Type 2 hairs present on branchlets

(100%); inflorescences with flowers all bisexual

(100%); branchlet prickles abundant

(240-1400 per decimetre), acicular (100%);

mixture of unbranched and branched

inflorescences (100%); corolla inner surface

glabrous (100%); fruiting calyx less than half

length of mature fruit (100%); adult leaves

deeply or shallowly lobed (83%).

Bean, Taxonomy of Solanum subg. Leptostemonum

3 species endemic to Australia; 3 species

in Queensland.

27. Solanum coracinum Symon, Austrobaileya

4: 429-30 (1995). Type: Queensland.

Darling Downs District: c. 22 km east

of Yuleba, on road to Miles, 17 November

1975, R.J. Henderson 2381 (holo: BRI;

iso: AD).

Solanum sp. 3 in Ross (1986)

Illustration : Symon (1995: 430)

Erect, rhizomatous perennial shrub, 0.7-1.5 m

high. Juvenile stage unknown. Adult branchlets

green; prickles 350-800 per decimetre, straight,

acicular, 1-10 mm long, 14-18 t im es longer

than wide; stellae absent or sparse, 0.3-0.5 mm

diameter, sessile; lateral rays 4-6, porrect;

central ray 0.4-0.8 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs sparse. Adult leaves broadly ovate or

orbicular, deeply lobed throughout; lobes 3-5

on each side, acute, lobing index 2-25; lamina

5-11.5 cm long, 3.5-10 cm wide, 1.1-1.5 times

longer than broad, apex acute, base obtuse or

cordate, oblique part 0-7 mm long, obliqueness

index 0-9 percent; petioles 2-3.4 cm long,

25-35% length of lamina, prickles absent.

Upper leaf surface green; prickles 15-100,

straight, acicular, 1-10 mm long, prickles

present on midvein and lateral veins; stellate

hairs confined to midrib, or distributed

throughout; protostellae absent; ordinary stellae

very sparse, 0.9-7.5 mm apart, 0.3-0.45 mm

across, sessile; lateral rays 4-8, porrect; central

ray 0.4-0.8 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Lower leaf surface green; prickles 10-30,

straight, acicular, present on midvein and lateral

veins; stellae absent or very sparse, 0.8-3.2 mm

apart, 0.45-0.6 mm diameter, stalks 0-0.1 mm

long; lateral rays 6-8, porrect; central ray 0.4-0.8

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent.

Inflorescence supra-axillary, pseudo-racemose

or 2-branched, common peduncle 18-39 mm

long, rachis prickles present; 14-30-flowered,

with all flowers bisexual and 5-merous; pedicels

4-8 mm long at anthesis, same thickness

throughout, 0.3-0.4 mm thick at mid-point,

prickles absent. Calyx tube 1.5-2.5 mm long,

lobes deltate or rostrate, 1.5-3 mm long;

prickles absent at anthesis; stellae moderate,

transparent, 0.25-0.4 mm across, sessile, lateral

713

rays 6-8, central ray 0.7-1.3 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs present. Corolla purple, 7-9 mm

long, deeply lobed, inner surface glabrous;

anthers 4-5.5 mm long; ovary with Type 2 hairs

only, or with stellate and Type 2 hairs;

functional style 4.5-6.5 mm long, erect, with

Type 2 hairs only. Fruiting calyx with lobes

less than half length of mature fruit, prickles

absent. Mature fruits 3-9 per inflorescence,

globular, 6-9 mm diameter, black, 1-locular

(septum absent or incomplete); placenta not

apparent; mesocarp juicy, succulent; exocarp

0.15-0.3 mm thick; pedicels 4-10 mm long in

fruit, 0.5-0.8 mm thick at mid-point; seeds pale

yellow, brown or black, 2.1-2.8 mm long.

Specimens examined : Queensland. Leichhardt District:

19 kmW ofWandoan, Aug 1999, Cooks.n. (BRI). Darling

Downs District: ‘Palardo’, May 1934, Blake 5866 (BRI);

10 miles [16 km] E of Texas, Jun 1951, Everist 2539 & Webb

(BRI); ‘Shellboume’, c. 20 miles [32 km] NE of Miles, May

1960, Johnson 1630 (BRI); E of Combididan Farm, ‘Cypress

Downs’, Sep 1961, Jones 164 (BRI); ‘Benandre’, 23 miles

[37 km] SE of Texas, Apr 1962, Pedley 988 (BRI); 20 km W

of Milmerran, Feb 1984, Stower (BRI); Houston-Gillespie

Dam road, N of Grays Gate, Dec 1999, Menkins DDP8

(BRI); 2.6 km E of Arcot, ENE of Texas, May 2000, Bean

16657 (BRI, NSW); Road 105, 7 km S of Milmerran, May

2003, Bean 20305 (BRI); Road 59, SE of Milmerran, May

2003, Bean 20333 (BRI);

Distribution and habitat : Solanum coracinum

is endemic to Queensland, extending from

Wandoan to Texas (Map 10), but not yet

recorded for New South Wales. It inhabits open

forest dominated by brigalow or belah, or

shrubby eucalypt woodland with Callitris. Soils

may be sandy-loams to clays.

Phenology: Flowers are recorded from

February to May and from August to December;

mature fruits recorded from April, May, June

and December.

Notes: S. coracinum is closely related to

S. mitchellianum, differing by the glabrous to

sparsely pubescent branchlets and leaf

undersides, the deeply lobed leaves with more

prickles on the upper surface, and the stellae of

the lower leaf surface with a shorter central ray.

Three specimens (Bean 17776, Thomby Range;

Beasley s.n.. Chinchilla; and Fensham 2877,

NNW of Miles) appear to represent intergrades

between it and S. mitchellianum. S.

mitchellianum occurs to the east, west and north

of the geographical range of S. coracinum.

714

Conservation status: S. coracinum is known

from seven localities, none of which is within

a conservation reserve. Applying the IUCN

guidelines (IUCN. 2001), a category of

“Vulnerable” is recommended (VU A4ce;

Blab(iii)+2ab(iii)).

28. Solanum mitchellianum Domin, Repert.

Spec. Nov. Regni Veg. 12: 131 (1913).

T^pe: Subtropical New Holland, anno

1846, T. Mitchell (lecto: K; isolecto: BM,

L), fide Symon (1981).

Illustration: Symon (1981: 127), as

S. semiarmatum.

Erect, rhizomatous perennial shrub, 0.5-1.8 m

high. Juvenile branchlets with 240-1400

prickles per dm; leaves (in outline) ovate,

deeply lobed, with 3 or 4 pairs of lobes; lamina

c. 11.5 x 8 cm, with c. 40 prickles on upper

surface. Adult branchlets terete or ridged,

green; prickles 240-1400 per decimetre,

straight, acicular, 1-10 mm long, 14-20 times

longer than wide; stellae dense, 0.4-0.7 mm

diameter, stalks 0-0.15 mm long; lateral rays

6-8, porrect; central ray 0.5-1 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs sparse. Adult leaves ovate, entire

or shallowly lobed throughout; lobes 3 or 4 on

each side, acute or obtuse, lobing index 1-1.9;

lamina 5.5-10.5 cm long, 1.8-6.5 cm wide,

1.5-3 times longer than broad, apex acute, base

obtuse or cordate, oblique part 0-3.5 mm long,

obliqueness index 0-5 percent; petioles 1.1-2.8

cm long, 18-30% length of lamina, prickles

present. Upper leaf surface green; prickles 8-

20, straight, acicular, 1-8 mm long, prickles

present on midvein and lateral veins; stellate

hairs distributed throughout; protostellae

absent; ordinary stellae very sparse to moderate

density, 1-3 mm apart, 0.25-0.7 mm across,

stalks 0-0.1 mm long; lateral rays 4-8, porrect;

central ray 0.8-1.5 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs present only in vein depressions. Lower

leaf surface white; prickles 2-10, straight,

acicular, present on midvein only or present

on midvein and lateral veins; stellae very dense,

c. 0.05 mm apart, 0.5-0.8 mm diameter, stalks

0-0.3 mm long; lateral rays 7-9, porrect;

central ray 1-1.5 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Inflorescence supra-axillary, pseudo-

racemose or 2-3-branched, common peduncle

21-40 mm long, rachis prickles present; 7-25-

Austrobaileya 6 (4): 639-816 (2004)

flowered, with all flowers bisexual and 5-

merous; pedicels 3-6 mm long at anthesis, same

thickness throughout or markedly thicker

distally, 0.3-0.6 mm thick at mid-point,

prickles absent or present. Calyx tube 0.5-1.5

mm long, lobes hemispherical, deltate or

attenuate, 2-3.5 mm long; prickles absent at

anthesis; stellae moderate to dense, transparent,

0.25-0.5 mm across, sessile, lateral rays 5-8,

central ray 0.8-1.2 t im es as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Corolla purple, 7-10 mm long,

deeply lobed, inner surface glabrous; anthers

3.5-5.5 mm long; ovary glabrous or with

stellate hairs only; functional style 6.5-8.5 mm

long, erect, glabrous or with stellate hairs only,

stellae c. 0.4 mm across, lateral rays c. 8, central

ray c. 1 times as long as laterals. Fruiting calyx

with lobes less than half length of mature fruit,

prickles absent. Mature fruits 7-17 per

inflorescence, globular, 8-9 mm diameter,

black, 1-locular (septum absent or incomplete);

placenta not apparent; mesocarp juicy,

succulent; exocarp 0.1-0.2 mm thick; pedicels

8-10 mm long in fruit, 0.8-0.9 mm thick at mid¬

point; seeds brown to black, 1.8-2.4 mm long.

Selected specimens examined : Queensland. Leichhardt

District: Moolyamba Gorge, Carnarvon Ranges, Sep 1940,

White 11361 (BRI); ‘Rosedale’ near Baralaba, Sep 1959,

Johnson 917 (BRI); Pegunny North, c. 55 km W of Moura,

May 1962, Johnson 2289 (BRI); Injune-Rolleston road, c.

77 km from Injune, Sep \91A,Moriarty 1558 (BRI, CANB);

Little St Peter, lOmiles [16km] N of Springsure, Sep 1985,

O’Keeffe 783 (BRI); Nathan Gorge, SW of Cracow, Aug

1990, Forster PIF7188 (BRI, MEL); Bunbuncundoo Spring,

Ka Ka Mundi N.P., Sep 1993, Purdie 4339 (BRI, CANB);

Auburn Range S.F., Jun 1996, Bean 10338 (BRI, MEL,

NSW); 14.8 km along Arcturus Road, NE of Springsure, Oct

1998, Bean 14009 (BRI); Expedition N.P., Amphitheatre

section, Nov 1998, Forster PIF23862 & Booth (AD, BRI,

K, MEL, NSW, QRS); Zamia Creek, Palmgrove N.P, Sep

2000, Forster PIF26109 et al. (AD, BRI, MEL); Brigalow

Research Station, SW of Moura, Dec 2001, Bean 18256

(BRI, MEL). Port Curtis District: Callide Valley, Apr 1937,

White 11184 (BRI). Warrego District: ‘Carnarvon’ Station,

Mar 2001, Fensham 4225 (BRI). Maranoa District: Kenniff

Lookout, c. 80 km SW of Rolleston, Jun 1977, Crisp 3084

(AD, BRI, CANB); ‘Stanhope Downs’, c. 44 km by road

NW of Roma on Orallo Road, Dec 1997, Thomas 1267

(BRI). Burnett District: Dingo Trap track, ‘Narayen’, Dec

1973, Leach N1496 (BRI); Monogorilby, Mundubbera shire,

Jan 1982, Forster 1105 (BRI). Darling Downs District:

Chinchilla, Jul 1912, Beasley 3 (BRI); ‘Wyaga’,

Goondiwindi district, Sep 1919, White s.n. (BRI); Cooranga

North, Apr 1925, White 2490 (BRI); ‘Calala’, c. 10 miles

[16 km] E of Meandarra, Apr 1960, Johnson 1620 (BRI);

2.5 km SSW of Gladfield, Jun 1986, Forster PIF2474 et al.

(BRI). New South Wales. North West Slopes: 7 miles [ 11

km] from Crooble on Warialda road, Sep 1950, Roe 258

(CANB); 7.5 km NW of North Star, Sep 1988, Moore 8689

Bean, Taxonomy of Solarium subg. Leptostemonum

(BRI, CANB); ‘Warivan’, 7.4 km from North Star on road

to Warialda, Sep 1988, Moore 8835 (CANB).

Distribution and habitat : Solanum

mitchellianum extends from Springsure and

Blackwater in Queensland to Warialda in New

South Wales (Map 12). It inhabits semi¬

evergreen vine thickets, brigalow-belah

communities or shrubby eucalypt woodlands

often with rock outcrops.

Phenology : Flowers are recorded for all months

of the year; mature fruits are recorded between

September and April.

Notes: Closely related to S. semiarmatum , and

for many years included in synonymy with it.

S. mitchellianum is however amply different

by virtue of the shallowly lobed or entire adult

leaves, stellae on the lower leaf surface sessile

or with stalks <0.3 mm long, ovary glabrous

or with stellae only (with Type 2 hairs only for

S. semiarmatum ), styles glabrous or with stellae

(with Type 2 hairs only for S. semiarmatum ),

and calyx prickles absent (present for

S. semiarmatum). It is also close to S. coracinum

(see notes under that species).

Conservation status: Widespread. Not

considered at risk.

29. Solanum semiarmatum F.Muell., Fragm.

2: 163 (1861). Type: New South Wales.

North Coast: Clarence River, undated,

H. Beckler (holo: MEL [MEL12130]; iso:

K, NSW).

Illustration: Symon (1981: 126)

Erect, rhizomatous perennial shrub, 1.5-3 m

high. Juvenile branchlets with 600-1400

prickles per dm, 0.5-7 mm long; leaves (in

outline) ovate, deeply lobed, with 4 or 5 pairs

of lobes; lamina c. 16 x 10 cm, with c. 70

prickles on upper surface. Adult branchlets

green; prickles 600-1400 per decimetre,

straight, acicular, 1-13 mm long, 15-20 times

longer than wide; stellae sparse to dense, 0.5-0.8

mm diameter, stalks 0-1.5 mm long; lateral

rays 4-8, porrect; central ray 0.5-1.5 times as

long as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs sparse. Adult leaves ovate

or broadly ovate, deeply lobed throughout; lobes

3 or 4 on each side, acute or obtuse, lobing index

2-7; lamina 7-17 cm long, 3.5-10 cm wide,

1.5-1.9 times longer than broad, apex acute,

715

base obtuse or cordate, oblique part 1.5-8 mm

long, obliqueness index 2-7 percent; petioles

1.7-4.5 cm long, 20-40% length of lamina,

prickles present. Upper leaf surface green;

prickles 7-70, straight, acicular, 1-10 mm long,

prickles present on midvein and lateral veins;

stellate hairs distributed throughout;

protostellae present; ordinary stellae sparse or

moderate density, 0.4-0.8 mm apart, 0.15-0.4

mm across, sessile; lateral rays 4-6, ascending;

central ray 0.7-1.2 times as long as laterals,

not gland-tipped; finger hairs present, 0.1-0.8

mm apart, not gland-tipped, 0.1-0.15 mm long;

Type 2 hairs absent. Lower leaf surface white

or yellowish; prickles 35-200, straight,

acicular, present on midvein and lateral veins;

stellae dense to very dense, 0.05-0.1 mm apart,

0.5-0.8 mm diameter, stalks 0-2.5 mm long;

lateral rays 6-8, porrect; central ray 1-2 times

as long as laterals, not gland-tipped; finger

hairs absent; Type 2 hairs absent. Inflorescence

supra-axillary, pseudo-racemose or 2-branched

or 3-branched, common peduncle 7-30 mm

long, rachis prickles present; 10-30-flowered,

with all flowers bisexual and 5-merous; pedicels

3- 10 mm long at anthesis, same thickness

throughout, 0.5-0.7 mm thick at mid-point,

prickles present. Calyx tube 1-2 mm long, lobes

deltate or attenuate, 1-3 mm long; prickles

absent or present at anthesis, 0-25 per flower,

0.5-2 mm long; stellae dense, transparent,

0.6-0.8 mm across, stalks 0-1.2 mm long,

lateral rays 5-7, central ray 1-1.7 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs present. Corolla purple,

10-12 mm long, deeply lobed, inner surface

glabrous; anthers 3.5-5.5 mm long; ovary with

Type 2 hairs only; functional style 7-9 mm

long, erect, with Type 2 hairs only. Fruiting

calyx with lobes less than half length of mature

fruit, prickles 0.5-3 mm long. Mature fruits

4- 9 per inflorescence, globular, 10-12 mm

diameter, black, 1-locular (septum absent or

incomplete); placenta not apparent; mesocarp

juicy, succulent; exocarp 0.1-0.2 mm thick;

pedicels 8-14 mm long in fruit, 1.2-1.4 mm

thick at mid-point; seeds pale yellow, 2-2.8 mm

long. Gin’s Whiskers.

Selected specimens examined: Queensland. Darling

Downs District: Wilsons Peak, May 1933, Michael s.n.

(BRI); 6 km from Mt Colliery towards Gambubal S.F., E of

Warwick, Apr 1999, Bean 14802 (BRI, MEL); Killarney-

The Head road, 7.1 km NE of Queen Mary Falls, Jan 2002,

Bean 18332 (BRI, MEL, NSW). Moreton District: Mt

Mistake, Jun 1887, Simmonds (BRI); Mt Lindesay, Oct 1921,

716

Austrobaileya 6 (4): 639-816 (2004)

White s.n. (BRI); Beech Mountain, Apr 1923, White 1927

(BRI); Roberts Plateau, Lamington N.P., May 1929, White

6072 (BRI); Levers Plateau, c. 90 km SSW of Brisbane, Apr

1972, Henderson H1298 (BRI); 3.3 km along Duck Creek

Road, near O’Reilly’s Guest House, Mar 2001 , Bean 17388

(BRI). New South Wales. North Coast: Toonumbar S.F., c.

26 km NW of Kyogle, Feb 1972, Henderson H1260A &

Parham (BRI); lower slopes ofMtLindesay, 7 miles [11 km]

ENE of Woodenbong, Sep 1972, Coveny 4544 & Rodd (BRI,

NSW); Old Grevillea-Bundgeam road, 25 km NW of Kyogle,

Dec 1977, Haegi 1538 (BRI, NSW).

Distribution and habitat : Solanum

semiarmatum is found along the “scenic rim”

of south-eastern Queensland, from Lamington

N.P. to Mt Mistake, and adjacent areas of New

South Wales south to Kyogle (Map 10). It

inhabits open areas within or on the margins

of tall notophyll rainforest, at altitudes generally

above 800 metres.

Phenology : Flowers are recorded from

September to May; mature fruits are recorded

from January to May.

Notes: Closely related to S. mitchellianum (see

notes under that species).

Conservation status : Not considered at risk.

Group 14 (S. torvum group) of Whalen (1984)

Large shrubs; prickles broad-based, sparse on

large stems and branchlets (100%); adult leaves

entire or shallowly lobed, broadly ovate, 7.5-26

cm long; upper leaf surface without prickles,

protostellae present (100%); inflorescence

2-many branched, with 15-65 flowers (100%);

calyx prickles absent (100%); corolla deeply

lobed, white (100%); ovary with Type 2 hairs

only (100%); mature fruits yellowish-green

(100%).

About 50 species, mainly in montane

areas of the neotropics, but with a few species

in Malesia; 2 species naturalised in

Queensland.

30. *Solanum chrysotrichum Schltdl.,

Linnaea 19: 304 (1847). Type: near Las

Trojes, Mexico, 1825-31, C.J.W. Schiede

81 (holo: HAL, fide Welman (2003)), n.v.

[5. hispidum auct. non Persoon]

Illustrations: Symon(1981:114), as S. hispidum;

Welman (2003: 6).

Erect, rhizomatous perennial shrub, 1.5-4 m

high. Juvenile branchlets with c. 15 prickles

per dm, 3-6 mm long; leaves (in outline)

broadly ovate, shallowly to deeply lobed, with

5 or 6 pairs of lobes; lamina 23-41 cm long,

16-40 cm wide, with 10-30 prickles on upper

surface. Adult branchlets brown; prickles 0-20

per decimetre, curved, broad-based, 1-4 mm

long, 2-3 times longer than wide; stellae sparse

to dense, 0.5-0.6 mm diameter, stalks 0.1-1

mm long; lateral rays 6-8, porrect; central ray

absent or present, 0-0.4 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Adult leaves broadly ovate,

shallowly lobed throughout; lobes 3-5 on each

side, acute or obtuse, lobing index 1.2-2.1;

lamina 12-26 cm long, 9-19 cm wide, 1.3-1.9

times longer than broad, apex acute or

acuminate, base cuneate, obtuse or cordate,

oblique part 0-12 mm long, obliqueness index

0-5 percent; petioles 2-6.3 cm long, 11-25%

length of lamina, prickles absent or present.

Upper leaf surface green; prickles absent;

stellate hairs distributed throughout;

protostellae present; ordinary stellae sparse,

0.3-0.6 mm apart, 0.2-0.6 mm across, sessile;

lateral rays 5-8, porrect; central ray 0.2-2 times

as long as laterals, not gland-tipped; finger

hairs absent; Type 2 hairs absent. Lower leaf

surface green or yellowish; prickles 0-3,

straight or curved, broad-based, prickles absent

or present on midvein only; stellae moderate

to dense, 0.3-0.5 mm apart, 0.4-0.6 mm

diameter, stalks 0-0.3 mm long; lateral rays

6-8, porrect; central ray 0.1-0.7 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Inflorescence leaf-

opposed or supra-axillary, 3 or 4-branched,

common peduncle 6-18 mm long, rachis

prickles absent; 13-65-flowered, weakly

andromonoecious, flowers 5-merous; pedicels

3-13 mm long at anthesis, markedly thicker

distally, 0.7-0.9 mm thick at mid-point,

prickles absent. Calyx tube 2-4 mm long, lobes

rostrate or attenuate, 4.5-10 mm long; prickles

absent at anthesis; stellae dense to very dense,

brown or rusty, 0.3-0.9 mm across, stalks 0-1

mm long, lateral rays 7-9, central ray 0.2-1

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Corolla

white, 14-20 mm long, deeply lobed, inner

surface sparsely stellate-hairy; anthers 6-9 mm

Bean, Taxonomy of Solanum subg. Leptostemonum

long; ovary with Type 2 hairs only; functional

style 10-13.5 mm long, erect, with Type 2 hairs

only. Fruiting calyx with lobes less than or more

than half length of mature fruit, prickles absent.

Mature fruits 7-37 per inflorescence, globular,

14-17 mm diameter, yellowish-green, 2-3-

locular; placenta in cross-section stalked, anvil¬

shaped; mesocarp moist but not juicy; exocarp

0.5-0.8 mm thick; pedicels 11-21 mm long in

fruit, 1.4-1.8 mm thick at mid-point; seeds pale

yellow, 2.5-2.7 mm long. Giant Devil’s Fig.

Specimens examined : Queensland. Burnett District:

Tarong Power station site, 15 km S of Nanango, Nov 1996,

Skillington s.n. (BRI). Wide Bay District: Conondale Range,

S.F.274, Oct 1982, McDonald 3626 & Williams (BRI);

Currymore, NW of Maleny, Apr 1993, Bean 6014 (BRI).

Moreton District: Wooloowin-Windsor etc. (Brisbane), May

1917, White s.n. (BRI): Ferny Grove near Brisbane, Nov

1934, White s.n. (BRI); Joseph Cresent, Deception Bay, Sep

1980, Dillewaard 61 & Olsen (BRI); Samford, Sep 1984,

Bunch JT1114 (BRI); Big Tree L.A., S.F.832, SE of

Bellthorpe, Jun 1993, Bean 6120 (BRI); comer of Alex Rd

& Attunga Lane, Mt Glorious, Oct 1997, Phillips 70 (AD,

BRI); South Stradbroke Island, Jul 1998, Leipers.n. (BRI);

Gold Creek road, Brookfield, W of Brisbane GPO, Feb 2000,

Bean 16034 (BRI, NSW, NY); Brisbane River, Long Pocket,

Indooroopilly, Sep 2000, Batianoff 200905 (BRI, DNA,

NSW); Industry Drive, Caboolture, Apr 2001, Bean 17617

(BRI, MEL, PRE). New South Wales. NORTH COAST:

Tuntable Creekroad, ESE of Nimbin, Sep 1994, Bean 7933

(BRI, NSW). Mexico: 15 km W of Santa Rosa, Yerz Cruz,

Oct 1930, Reddick 616 (BH); San Andres, NW of San

Cristobal, Feb 1931, Souviron & Erlanson 76 (BH); top of

ridge between La Cumbre and Las Joyas, Jan 1979, Nee &

litis 16695 (BH, NY); along highway Mex. 190,10 km SW

of Motozintla, Dec 1985, Nee 32331 (BH, NY).

Distribution and habitat : Solanum

chrysotrichum is indigenous to southern

Mexico, Guatemala, Costa Rica and Panama,

and naturalised in many subtropical parts of

the world. It is naturalised in south-eastern

Queensland, and extreme north-eastern N.S.W.

(Map 13). It inhabits degraded places in

association with a range of other weedy species.

Notes: This species has been erroneously called

S. hispidum Pers. for many years, but true

S. hispidum is from the Peruvian Andes, and

has not been reported as a naturalised plant.

Identification of Queensland material was

achieved by matching with Mexican specimens

identified by neotropical Solanum experts, M.

Whalen and M. Nee.

Phenology: Flowers and fruits may be found

at any time of the year.

717

31. *Solanum torvum Sw., Prodr. 47 (1788).

Type: Jamaica, undated, O. Swartz s.n.

(holo: S), n.v.,fide D’Arcy (1974: 860).

Solanum largiflorum C.T.White, Queensland

Agric. J. 8: 170 (1917). Type:

Queensland. Wide Bay District: Kin Kin,

March 1916, C.T. White & W.D. Francis

(syn: MEL, NSW), photos at BRI.

Illustrations: Symon (1981: 115); Welman

(2003: 4).

Erect, rhizomatous perennial shrub, 0.8-2.5 m

high. Juvenile branchlets with 5-15 prickles

per dm, 2-5 mm long; leaves (in outline)

broadly ovate, deeply lobed, with 2-4 pairs of

lobes; lamina 12-18 cm long, 9-15 cm wide,

with 0-9 prickles on upper surface. Adult

branchlets brown or green; prickles absent or

present, 0-5 per decimetre, straight or curved,

broad-based, 3-7 mm long, 1.5-2 times longer

than wide; stellae dense, 0.6-1 mm diameter,

stalks 0.1-0.4 mm long; lateral rays 6-9,

porrect; central ray 0.1-0.3 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Adult leaves broadly ovate

or orbicular, entire or shallowly lobed

throughout; lobes 1-3 on each side, acute or

obtuse, lobing index 1-1.3; lamina 7.5-16.5

cm long, 4-13 cm wide, 1.1-1.5 t im es longer

than broad, apex acute, base obtuse or cordate,

oblique part 0-4 mm long, obliqueness index

0-5 percent; petioles 0.9-4.3 cm long, 15-26%

length of lamina, prickles absent. Upper leaf

surface green; prickles absent; stellate hairs

distributed throughout; protostellae present;

ordinary stellae sparse to dense, 0.25-0.5 mm

apart, 0.3-0.5 mm across, sessile; lateral rays

5-8, porrect; central ray 0.8-2 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs present throughout, 0.2-0.5 mm

apart. Lower leaf surface white; prickles 0-3,

straight, broad-based, prickles absent or present

on midvein only; stellae dense, 0.1-0.3 mm

apart, 0.6-1 mm diameter, stalks 0-0.4 mm long;

lateral rays 8-9, porrect; central ray 0.3-0.9

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent.

Inflorescence supra-axillary, 2-4-branched,

common peduncle 2-17 mm long, rachis

prickles absent; 15-50-flowered, weakly

andromonoecious, flowers 5-merous; pedicels

718

6-10 mm long at anthesis, same thickness

throughout, 0.6-0.8 mm thick at mid-point,

prickles absent. Calyx tube 1-1.5 mm long,

lobes deltate or rostrate, 1.5-4.5 mm long;

prickles absent at anthesis; stellae sparse to

moderate, white, 0.2-0.5 mm across, sessile,

lateral rays 6-8, central ray 0.8-1.8 times as

long as laterals, not gland-tipped or gland-

tipped; finger hairs present; Type 2 hairs absent.

Corolla white, 9-12 mm long, deeply lobed,

inner surface glabrous; anthers 6-7.5 mm long;

ovary with Type 2 hairs only; functional style

9-10.5 mm long, erect, glabrous. Fruiting calyx

with lobes less than half length of mature fruit,

prickles absent. Mature fruits 2-15 per

inflorescence, globular, 12-17 mm diameter,

yellowish-green, 2-locular; placenta in cross-

section stalked, anvil-shaped; mesocarp moist

but not juicy; exocarp 0.5-0.7 mm thick;

pedicels 14-19 mm long in fruit, 1.8-2.4 mm

thick at mid-point; seeds white or pale yellow,

2.2-2.4 mm long. Devil’s Fig. Fig. 2.

Selected specimens examined’. Northern Territory.

Berrimah Farm, Darwin, Feb 1981, Rankin 2582 (BRI,

CANB, DNA). Queensland. Burke District: Leichhardt

River, Mt Isa, Nov 1999, Stevens & Fox IDF909 (BRI). Cook

District: Daintree, May 1952, Everist 5129 (BRI); Copper

Lode Falls Dam area, Cairns, Dec 1972, Birch 12 (BRI); Mt

Webb near ‘Standee’ HS, N of Hope Yale mission, Aug 1978,

Kanis 1912 (BRI, CANB, L, MO, US); Cooktown rubbish

dump, Jul 1991, Waterhouse 1881 (BRI, DNA, PERTH);

Lakefield N.P., Twelve Mile area, Normanby River, Jan 1993,

Forster PIF12901 & Bean (AD, BRI, QRS). North Kennedy

District: Ayr, undated, Michael 1533 (BRI); Cromarty, Mar

1935, Blake 8317 (BRI); Black River, c. 14 miles [23 km]

N of Townsville, May 1967, Symon 4743 (BRI); c. 15 kmN

of Proserpine, western foot of Mt Dryander, Jul 1974,

Henderson H2228 et al. (BRI); Fletcher Creek, 6 km N of

‘Toomba’ HS, Nov 1986, Bolton MPB740 (BRI). South

Kennedy District: 17 km SE of Kuttabul on Townsville-

Mackay Hwy, Sep 1981, Haegi 2059 (BRI, NSW); Newry

Island, Dec 1986, Dalliston N275 (BRI); Bowen River

crossing, 24 km S of Collinsville, Jun 1989, Jobson 615 (BRI,

CANB, NSW). Leichhardt District: just west of Nebo, Apr

1998, Holland 1192 (BRI). Port Curtis District: Yeppoon,

Jul 1974, Swarbrick 6190 (BRI); 0.9 km from Bruce Hwy,

towards St Lawrence, Apr 2000, Bean 16263 (BRI); T.R.202

Colosseum Creek, Apr 2000, Forster P1F25478 & Booth

(AD, BRI, MEL, QRS). Wide Bay District: Theebine, Nov

1921, White s.n. (BRI); Noosa N.P., north east comer, Aug

1985, Sharpe 3832 et al. (BRI, MEL); Ocean Park estate,

Dundowran, Nov 1991, Forster PIF9177 & Smyrell (AD,

BRI, MEL). Moreton District: Mt Buderim, Jan 1935, Blake

7188 & Middleton (BRI); Naim Road, Morayfield, c. 35 km

N of Brisbane, Mar 2000, Bean 16108 (BRI, CANB, MEL,

MO, NSW); Godfreys road, Bli Bli, c. 1 km N of Maroochy

River, Mar 2000, Bean 16136 (BRI, DNA, NSW).

Austrobaileya 6 (4): 639-816 (2004)

Distribution and habitat : Solanum torvum is

native to the West Indies, but is now naturalised

in many tropical parts of the world. It is

naturalised along the east coast of Queensland,

especially north of the Tropic of Capricorn, and

at Mt Isa (Map 15). It is recorded from the far

north of the Northern Territory, but is not yet

known from New South Wales. It grows on

degraded sites, including roadsides, pasture and

quarries.

Phenology: Flowers and fruits may be found

at any time of the year.

Notes: Boonkerd et al. (1993) list a number of

uses made for S. torvum in south-east Asia. For

example, young immature fruits are eaten raw

or cooked as a vegetable, and the fruits are

commonly available in local markets. S. torvum

is sometimes used as a disease resistant

rootstock for Tomato and Eggplant.

Group 22 (S. quitoense group) of Whalen

(1984)

Mature fruits densely stellate-tomentose, 25-

35 mm diameter; stems, branchlets and leaves

prickly, calyx not prickly; branchlet stellae with

lateral rays ascending or multiradiate; adult

leaves large, broadly-ovate to orbicular,

shallowly lobed; ovary and style with dense

stellate hairs; stellae of upper leaf surface with

central ray many times longer than laterals;

corolla white; seeds brown to black.

12 species in the world; 1 species in

Australia, 1 species indigenous to Queensland.

32. Solanum lasiocarpum Dunal, Hist. Nat.

Solanum 222 (1813); S. ferox var.

lasiocarpum (Dunal) Miq., Flora Indiae

Batavae 2:647 (1856). Type: t. 35, Hortus

Indicus malabaricus Vol. 2 (1680); lecto:

the illustration,//^ Whalen et al. (1981).

Illustration: Symon (1981: 107), as S. ferox.

Erect, rhizomatous perennial shrub, 1-2 m

high. Juvenile stage unknown. Adult branchlets

grey or brown; prickles 20-350 per decimetre,

straight, acicular or broad-based, 1-4 mm long,

5-10 times longer than wide; stellae dense, 0.8-1.2

mm diameter, stalks 0-0.5 mm long; lateral

rays 8-10, ascending or multiradiate; central

ray 0.8-1.5 times as long as laterals, not gland-

Bean, Taxonomy of Solarium subg. Leptostemonum

tipped; finger hairs absent; Type 2 hairs absent.

Adult leaves broadly ovate or orbicular,

shallowly lobed throughout; lobes 3-8 on each

side, acute or obtuse, lobing index 1.1-1.3;

lamina 10-35 cm long, 8.5-26 cm wide, 1.1-1.3

times longer than broad, apex obtuse or acute,

base obtuse or cordate, oblique part 0-15 mm

long, obliqueness index 0-4 percent; petioles

2.7-7.2 cm long, 18-30% length of lamina,

prickles present. Upper leaf surface green;

prickles 0-120, straight, acicular or broad-

based, 2-7 mm long, prickles absent or present

on midvein only or present on midvein and

lateral veins; stellate hairs distributed

throughout; protostellae present; ordinary

stellae density moderate to dense, 0.2-0.4 mm

apart, 0.35-0.5 mm across, sessile; lateral rays

4-8, ascending or multiradiate; central ray

4-10 t im es as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Lower

leaf surface white or yellowish; prickles 0-60,

straight, acicular or broad-based, prickles

absent or present on midvein only or present

on midvein and lateral veins; stellae dense to

very dense, 0.1-0.3 mm apart, 0.5-0.8 mm

diameter, stalks 0-0.4 mm long; lateral rays

8-12, ascending or multiradiate; central ray

1-2 times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent.

Inflorescence supra-axillary, pseudo-racemose,

common peduncle 0-2 mm long, rachis prickles

present; 2-10-flowered, weakly andromonoecious,

flowers 5-merous; pedicels 4-9 mm long at

anthesis, same thic kn ess throughout, 0.7-1 mm

thick at mid-point, prickles absent. Calyx tube

2.5-5 mm long, lobes deltate, 2-8 mm long;

prickles absent at anthesis; stellae very dense,

yellow or white, 0.5-0.8 mm across, stalks

0-1 mm long, lateral rays 6-12, central ray

1-2 times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Corolla

white, 12-18 mm long, deeply lobed, inner

surface glabrous or sparsely stellate-hairy;

anthers 6-8.5 mm long; ovary with stellate

hairs only; functional style erect, glabrous.

Fruiting calyx with lobes less than half length

of mature fruit, prickles absent. Mature fruits

1 or 2 per inflorescence, globular, 25-35 mm

diameter, orange, conspicuously tomentose;

mesocarp moist but not juicy; pedicels 10-15

mm long in fruit, 0.9-1.5 mm thick at mid¬

point; seeds brown to black, 2.2-3.5 mm long.

719

Specimens examined : Queensland. Cook District: near

Lockerbie, Dec 1980, Hyland 10946A (BRI, QRS); 22.4 km

NE of Bamaga, Feb 1994, Fell DGF4064 & Stanton (BRI,

MEL, QRS).

Distribution and habitat : Solanum

lasiocarpum is widespread from India and

southern China, and throughout Indochina and

Malesia. In Australia, it is known only from

the Bamaga area (Map 8). It grows in notophyll

rainforest.

Phenology : Flowers recorded in February;

fruits (maturity unknown) recorded in

December.

Notes: A photograph of the iconotype is shown

in Whalen et al. (1981: 102). Heiser (1996)

treated S. lasiocarpum as a variety of S. ferox

L., but the latter differs significantly by the

aculeate and accresent calyx.

Conservation status: S. lasiocarpum is known

only from near Bamaga, and is not protected

in a conservation reserve. Its occurrence equates

to a single location, using the definition

provided by the IUCN (2001). Applying the

IUCN guidelines (IUCN. 2001), a category of

“Vulnerable” is recommended (VU Dl+2).

Group 23 (S. mammosum group) of Whalen

(1984)

Small prickly shrubs; leaves broad, shallowly

lobed; seeds 4.8-5.5 mm long; branchlets and

leaves without stellate hairs but with many

finger hairs; mature fruits globose, 25-45 mm

diameter, orange, with dry mesocarp; corolla

deeply lobed; calyx prickly; Type 2 hairs present

on branchlets.

About 20 species in the neotropics; 1

species naturalised in Queensland.

33. *Solanum capsicoides All., Melanges

Philos. Math. Soc. Roy. Turin 5: 12

(1773). Type: cultivated at Turin,

undated, C. Allioni s.n. (holo: TO), n.v.,

fide Welman (2003).

Solanum ciliatum Lam., Tab. Encyc. Meth.

Bot. 2: 21 (1794). Type: locality

unknown, undated, coll, unknown (syn:

P-LA, microfiche 467.18).

Illustrations: Symon (1981: 102); Welman

(2003: 6).

720

Erect, rhizomatous perennial shrub, 0.3-1 m

high. Juvenile branchlets with c. 100 prickles

per dm; leaves (in outline) broadly ovate, deeply

lobed, with 3 pairs of lobes; lamina c. 11 x 13

cm, with c. 10 prickles on upper surface. Adult

branchlets brown or green; prickles 30-80 per

decimetre, straight, acicular, 1.5-8 mm long,

8-16 times longer than wide; stellae absent;

finger hairs present, not gland-tipped, 3-6 mm

long; Type 2 hairs sparse to dense. Adult leaves

broadly ovate or orbicular, entire or shallowly

to deeply lobed throughout; lobes 2-4 on each

side, acute or obtuse, lobing index 1-3; lamina

7.5-16.5 cm long, 6-13 cm wide, 1-1.3 times

longer than broad, apex acute, base obtuse or

cordate, oblique part 0-6 mm long, obliqueness

index 0-4 percent; petioles 3.8-6.2 cm long,

35-55% length of lamina, prickles present.

Upper leaf surface green; prickles 4-30,

straight, acicular or broad-based, 3-10 mm

long, prickles present on midvein and lateral

veins; stellate hairs absent; finger hairs present,

0.5-2 mm apart, not gland-tipped, 1-4 mm

long; Type 2 hairs present only in vein

depressions. Lower leaf surface green; prickles

30-50, straight, acicular or broad-based,

prickles present on midvein and lateral veins;

stellae absent; finger hairs present, 0.6-6 mm

apart, not gland-tipped, 1.5-3 mm long or

absent; Type 2 hairs present only on veins.

Inflorescence supra-axillary, solitary or pseudo-

racemose, common peduncle absent, rachis

prickles present; 1-5-flowered, weakly

andromonoecious, flowers 5-merous; pedicels

8-21 mm long at anthesis, same thickness

throughout, 0.3-0.4 mm thick at mid-point,

prickles present. Calyx tube 1-3 mm long, lobes

deltate, 1.5-3 mm long; prickles present at

anthesis, 20-30 per flower, 1.5-6 mm long;

stellae absent; finger hairs present; Type 2 hairs

present. Corolla white, 7-10 mm long, deeply

lobed, inner surface glabrous; anthers 4.5-6

mm long; ovary with Type 2 hairs only;

functional style 5.5-7 mm long, erect, glabrous.

Fruiting calyx with lobes less than half length

of mature fruit, prickles 3-7 mm long. Mature

fruits 1-3 per inflorescence, globular, 25-45

mm diameter, orange, 1-locular (septum absent

or incomplete); placenta not apparent;

mesocarp dry; exocarp c. 2 mm thick; pedicels

17-28 mm long in fruit, 1-1.4 mm thick at

mid-point; seeds pale yellow, 4.8-5.5 mm long.

Devil’s Apple.

Austrobaileya 6 (4): 639-816 (2004)

Selected specimens examined : Queensland. Cook District:

Malanda Falls N.P., Jun 1965, Cunningham s.n. (BRI);

S.F.310, Gadgarra, c. 4 km ESE of Lake Barrine, Mar 1976,

Moriarty 1995 (BRI, QRS); edge of Lake Eacham, c. 45 km

SSW of Cairns, Aug 1976, Henderson H2402 (BRI); Dismal

Ck, 6 km W of Kuranda, Sep 1999, Wannan 1401 et al.

(BRI). South Kennedy District: 9 miles [14 km] SW of

Pinnacle, Jan 1964, Mayne s.n. (BRI); Eungella, Apr 1978,

Byrnes 3696 & Clarkson (BRI); Slade Point dunal system,

Mar 1993, Champion 782 (BRI). Burnett District:

Cherbourg, c. 6 miles [10 km] SE of Murgon, Oct 1969, coll,

unknown (BRI). Wide Bay District: Sandy Ck, Biggenden-

Childers road, Dec 1969, Colbran s.n. (BRI); Lake

Cootharaba, side of Mill Point track, Apr 1986, Sandercoe

Cl 160 & Milne (BRI); Currymore, NW of Maleny, Apr 1993,

Bean 6016 (BRI, NSW). Moreton District: Virginia near

Brisbane, Nov 1915, White s.n. (BRI); Myer’s Ferry,

Southport, Apr 1920, Francis s.n. (BRI); 2 km S of

Alstonville, Aug 1985, Reynolds & Calway s.n. (BRI);

Natural Bridge N.P., Numinbah Valley, Feb 2000, Bean

16006 (BRI); North Tamborine Environmental Park, Mt

Tamborine, Mar 2001, Boyle TPB185 & Phillips (BRI). New

South Wales. North Coast: Kings Beach, Broken Head, Mar

1982, Hind 3039 (BRI, NSW); Tuckean Island road, W of

Warded, Apr 2001 ,Bean 17577 (BRI, NSW).

Distribution and habitat: Solanum capsicoides

is indigenous to coastal Brazil (Whalen 1984).

It is widely naturalised in coastal and subcoastal

areas of Queensland and northern New South

Wales (Map 14). It inhabits degraded sites, or

rainforest margins, prefering moist shady areas.

Phenology: Flowers and fruits may be found

at any time of the year.

Notes: S. capsicoides was first recorded as

naturalised for Queensland by Bailey (1881),

under the name S. aculeatissimum Jacq., to

which it is closely related. The differences were

outlined by Welman (2003).

S. capsicoides completely lacks stellate

hairs; the indumentum comprises finger hairs

and Type 2 hairs.

Group 25 (S. hystrix group) of Whalen (1984)

Small plants <1 m high (100%); the leaves

green on upper surface (100%); corolla inner

surface glabrous (100%); seeds pale yellow

(100%); adult leaves lobed (100%); calyx

prickles >5 per flower (100%); inflorescence

weakly to strongly andromonoecious (100%);

calyx lobes not exceeding mature fruits (100%);

prickles present on both upper leaf surface

(100%); branchlet prickles acicular (95%);

prickles present on lower leaf surface (95%);

the herbaceous resprouter habit (90%); mature

fruits yellow-green to green (90%); adult leaves

Bean, Taxonomy of Solanum subg. Leptostemonum

with winged petioles (50%); finger hairs on

upper leaf surface (50%); style sigmoid (30%).

19 species endemic to Australia; 10

species indigenous to Queensland.

34. Solanum ditrichum A.R.Bean sp. nov.

Frutex fusus humilis; ramuli pilis digitatis

abundis sed pilis stellatis raris vel

absentibus; folia adulta 3-6 paribus

loborum acutorum non profundorum, et

usque ad 100 aculeos in superficiebus

ambabus, stellis superficiei superioris

radio centrali radiis lateralibus 2-3plo

longiore; calyx aculeatus sed non

accrescens; stylus functionalis

sigmoideus; semina flaveola. Typus: New

South Wales. North Coast: Branch road,

Dalmorton State Forest, SW of Grafton,

7 January 2001, A.R. Bean 17255 (holo:

BRI (2 sheets + spirit); iso: CANB, K,

MEL, NSW).

Solanum sp. (Mt Maroon P.I. Forster+

PIF11564) in Henderson (2002).

Prostrate or sprawling, herbaceous resprouter,

0.1-0.6 m high. Juvenile stage absent. Adult

branchlets green; prickles 80-180 per decimetre,

straight, acicular, 1-7 mm long, 10-17 times

longer than wide; stellae sparse, 0.5-0.6 mm

diameter, stalks 0-0.1 mm long; lateral rays

5-8, porrect; central ray present, 0.7-1.5 times

as long as laterals, not gland-tipped; finger

hairs abundant, gland-tipped, 0.1-0.7 mm long;

Type 2 hairs absent. Adult leaves ovate or

broadly ovate, shallowly lobed throughout;

lobes 3-6 on each side, acute, lobing index

1.2-1.4; lamina 6-12.5 cm long, 4.5-9.5 cm

wide, 1.3-1.8 times longer than broad, apex

acute, base obtuse or cordate, oblique part 1-10

mm long, obliqueness index 2-11 percent;

petioles 2-5.6 cm long, 20-50% length of

lamina, prickles present. Upper leaf surface

green; prickles 15-100, straight, acicular, 2-8

mm long, prickles present on midvein and

lateral veins; stellate hairs distributed

throughout; protostellae absent; ordinary stellae

sparse, 0.8-1.7 mm apart, 0.3-0.4 mm across,

sessile; lateral rays 4-7, porrect; central ray 2-3

times as long as laterals, not gland-tipped;

finger hairs present, 0.4-1.2 mm apart, not

gland-tipped or gland-tipped, 0.1-0.6 mm long;

Type 2 hairs absent. Lower leaf surface green;

721

prickles 30-100, straight, acicular, present on

midvein and lateral veins; stellae sparse to

moderate, 0.4-1.5 mm apart, 0.5-0.7 mm

diameter, sessile; lateral rays 4-8, porrect;

central ray 1-1.7 times as long as laterals, not

gland-tipped; finger hairs present, 0.3-1 mm

apart, gland-tipped, 0.1-0.6 mm long; Type 2

hairs absent. Inflorescence supra-axillary,

pseudo-racemose, common peduncle 2-16 mm

long, rachis prickles present; 2-5-flowered,

weakly andromonoecious, flowers 5-merous;

pedicels 5-18 mm long at anthesis, markedly

thicker distally, 0.4-1 mm thick at mid-point,

prickles present. Calyx tube 3-5 mm long, lobes

deltate or attenuate, 4-8 mm long; prickles

present at anthesis, 36-70 per flower, 2-7 mm

long; stellae sparse, transparent, 0.4-0.5 mm

across, sessile, lateral rays 4-8, central ray 1-1.5

times as long as laterals, not gland-tipped;

finger hairs present; Type 2 hairs absent.

Corolla purple, 9-17 mm long, rotate, inner

surface glabrous; anthers 4-5 mm long; ovary

glabrous, or with Type 2 hairs only; functional

style 9-10.5 mm long, sigmoid, glabrous.

Fruiting calyx with lobes less than half length

of mature fruit, prickles 2-7 mm long. Mature

fruits 1 or 2 per inflorescence, globular, 21-26

mm diameter, yellowish-green or pale green

with dark green streaks, 2-locular; placenta in

cross-section stalked, anvil-shaped; mesocarp

moist but not juicy; exocarp c. 2.5 mm thick;

pedicels 15-28 mm long in fruit, 1.3-2 mm

thick at mid-point; seeds pale yellow, 2.3-2.6

mm long. Fig. 3, 24.

Specimens examined : Queensland. Wide Bay District:

Gympie, undated, Kenny (BRI); Kin Kin, Mar 1916, Francis

& White s.n. (BRI). Darling Downs District: top of Mt

Mitchell, Main Range, Jul 1930, White 6881 (BRI); S.F.401,

2 km W of Mt Huntley, Oct 1992, Forster PIF11852 et at.

(AD, BRI, K, L, MEL, NSW); Portion 90, Wyberba, near

Girraween N.P., Sep 1993, Bean 6409 & Forster (BRI); 6

km from Mt Colliery towards Gambubal S.F., E ofWarwick,

Apr 1999, Bean 14806 (BRI). Moreton District: Yandina,

Mar 1891, Simmonds s.n. (BRI); Springbrook, Sep 1929,

White 6244 (BRI); foot of Mt Ernest, Oct 1932, Blake 4287

(BRI); Campbell’s Folly, 4 km SW of Tylerville, Sep 1992,

Forster PIF 11523 8cLeiper (AD, BRI); Mt Maroon, summit

area, Sep 1992, Forster PIF 11564 & Leiper (BRI, MEL);

Mt Gillies, Oct 1992, Forster PIF12076 & Reilly (AD, BRI);

Duck Creek road, near O’Reilly’s Guest House, Jan 2000,

Bean 15972 (BRI, NSW); Wilkies Scrub, Wongawallan, 7

km W of Coomera, Jul 2001, Bean 17689 (BRI, CANB, L,

MO). New South Wales. Northern Tablelands: Gibraltar

Range N.P, c. 67 km E of Glen Innes, Oct 1969, Coveny

2236 (AD, BRI, NSW); c. 12 km from Tenterfield on road to

Bluff Rock, Sep 1976, Pearce 87 (NSW); c. 3 km S of

722

Austrobaileya 6 (4): 639-816 (2004)

QUEENSLAND HERBARIUM (BR1|

North C 6 Ht

Solarium *fr. |JYll Mansart P.l.Fonrt®r+ PIF11S64)

QUEENSLAND HERBARIUM m)

Brisbane Australia

AO 493863

C4ll.AK.lfel* 17255 TiWiffH

SSd 49m 44 i 15 SH S*» (AGQ 66 J

(S 5 . 4 jam 67 t)MTSi 'S«w«»wb

BniwURoMt. Oturaton £li» f«SSL S«flhWM! CtSfonoo

Open fctisl 4f EucaJs-jiluS ssnsnotJM, E- eflSMtofs. £ vdSiWhlM

f camsa. Syscsipa fftmiUtrn. LoflUoUsman ciutfertut.

M»chSii<JiUafS«n«fl* SJiiutOy R**««a»l (twin# |u*i

4»™n*losw L4K4rt'* rats at W»

Sjmrartwj thrjf Is Wan t>«H. hrovos dull g FBMI* pun*S.

FruW mssura. nua-gwi will! d**«« fliwn t1r*si*

OXUB4I a tit

Sara rttlirsi it CSI

dk let) stf s*«m*

Cup. H 5 WMEL CAMB K

■ Ulsy l» sfiod Si «mpu1(rmiil osiSfliSh MjnlSsr AQ485SJ*

[Antfirtral PflpS^

HhLQTypg Queensland Heitatium (BRl)

Sofo/tm^ e&trithtm, A A. 6**^

Af. a*„io<r Zlkj 2

Fig. 24. Holotype of Solanum ditrichum.

Bean, Taxonomy of Solanum subg. Leptostemonum

723

Wallangarra on main road to Tenterfield, Dec 1977, Haegi

1547 (BRI, NSW); 29 km N of Tenterfield, off the Mt

Lindesay Hwy, Bungoona track to summit in Bald Rock N.R,

Oct 1993, Coveny 16560 & Whalen (AD, BRI, MO, NSW).

North Coast: Toonumbar S.F., c. 26 km NW of Kyogle, Feb

1972, Henderson H1265 (BRI); Mountaineer track,

Chichester S.F., Dungog, Jun 1974, Swan 24 (AD, BRI);

Whian Whian S.F., N of Lismore, Aug 1975, Moriarty 1707

(BRI); summit of Bald Knob, 10 km W of Woodenbong, Sep

1998, Bean 13799 (BRI); Mt Warning, Oct 1963, Johnson

2730 (BRI); Ewingar S.F., S of Tabulam, Feb 2001, Bean

17331 (BRI, NSW).

Distribution and habitat : Solanum ditrichum

is a widespread species extending from Dungog

in New South Wales to Mt Mee in Queensland

(plus historical records from Yandina, Kin Kin

and Gympie) (Map 16). It grows in wet

sclerophyll eucalypt forest or on rainforest

margins.

Phenology : Flowers and fruits are recorded for

all months except May and June.

Notes: S. ditrichum is closely related to

S. campanulatum R.Br. It differs from

S. campanulatum by having pale seeds, acute

leaf lobes, calyx scarcely accresent, finger hairs

very short on upper leaf surface, and the stellate

hairs rare or absent on branchlets. S. campanulatum

is restricted to the greater Sydney area, as far

north as Denman and Scone.

The mature fruits of S. ditrichum remain

green to yellowish-green at maturity, although

they may have some light streaking of purple

at the pedicel end (R. Fensham pers. comm.)

Conservation status: Widespread. Not

considered at risk.

Etymology: From the Greek di- meaning two

or double, and -trichos meaning hair. This

refers to presence of both stellate and finger

hairs on the leaves of this species.

35. Solanum cookii Symon, J. Adelaide Bot.

Gard. 4: 233 (1981). Type: cultivated at

Waite Institute (ex Upper Lankelly Creek,

Cape York Peninsula, Queensland), 25

February 1972, D.E. Symon s.n. (holo:

AD; iso: BRI, CANB, K).

Solanum adenophorum var. indivisum

Domin, Biblioth. Bot. 89: 586 (1929).

Type: Queensland. North Kennedy District:

Rockingham Bay, undated, J. Dallachy (holo:

K, n.v.),fide Symon (1981).

Illustration: Symon (1981: 234)

Prostrate or sprawling, herbaceous resprouter,

0.1-0.5 m high. Juvenile stage absent. Adult

branchlets green; prickles 30-300 per decimetre,

straight, acicular, 1-8 mm long, 12-17 times

longer than wide; stellae sparse to dense,

0.6-1 mm diameter, stalks 0-0.2 mm long;

lateral rays 4 or 5, porrect; central ray 2-5 times

as long as laterals, gland-tipped; finger hairs

present, gland-tipped, 0.2-1.7 mm long; Type

2 hairs absent. Adult leaves ovate or broadly

ovate, shallowly lobed throughout; lobes 4 or 5

on each side, acute, lobing index 1.1-1.2;

lamina 5-10.5 cm long, 4-7.5 cm wide, 1.3-1.7

times longer than broad, apex acute, base obtuse

or cordate, oblique part 0-8 mm long,

obliqueness index 0-8 percent; petioles 3.4-5.4

cm long, 40-55% length of lamina, prickles

present. Upper leaf surface green; prickles 30-145,

straight, acicular, 1-7 mm long, prickles

present on midvein and lateral veins; stellate

hairs absent; finger hairs present, 0.1-0.6 mm

apart, not gland-tipped or gland-tipped, 0.2-2

mm long; Type 2 hairs present throughout, 0.1-1

mm apart. Lower leaf surface green; prickles

20-65, straight, acicular, present on midvein

only or present on midvein and lateral veins;

stellae very sparse to moderate, 0.5-3 mm apart,

0.5-0.9 mm diameter, stalks 0-0.2 mm long;

lateral rays 4-6, porrect; central ray 2-5 times

as long as laterals, not gland-tipped; finger

hairs present, 0.1-0.3 mm apart, not gland-

tipped or gland-tipped, 0.1-1.4 mm long; Type

2 hairs absent. Inflorescence supra-axillary,

pseudo-racemose, common peduncle 18-27

mm long, rachis prickles present; 2-7-flowered,

with all flowers bisexual and 5-merous; pedicels

4-7 mm long at anthesis, same thickness

throughout, 0.2-0.4 mm thick at mid-point,

prickles present. Calyx tube 1.5-3 mm long,

lobes attenuate, 7-11 mm long; prickles present

at anthesis, 25-35 per flower, 2.5-6 mm long;

stellae absent or sparse or moderate,

transparent, 0.5-0.7 mm across, stalks 0-0.1

mm long, lateral rays 2-4, central ray 4-6 times

as long as laterals, not gland-tipped; finger

hairs present; Type 2 hairs absent. Corolla

mauve, 8-12 mm long, shallowly lobed, inner

surface glabrous; anthers 3.5-5 mm long; ovary

with Type 2 hairs only; functional style 5-6

mm long, erect, glabrous. Fruiting calyx with

lobes more than or exceeding mature fruit,

724

prickles 2.5-6 mm long. Mature fruits 1-6 per

inflorescence, globular, 11-13 mm diameter,

green, 2-locular; placenta in cross-section

stalked, circular to elliptical; mesocarp moist

but not juicy; exocarp 0.6-0.7 mm thick;

pedicels 5-9 mm long in fruit, 0.6-1 mm thick

at mid-point; seeds pale yellow, 1.8-2 mm long.

Specimens examined: Queensland. Cook District: c. 6

miles [10 km] SW ofYungaburra, Curtain Fig site, Dec 1968,

Tracey (BRI); Upper Lankelly Creek, western slopes of

Mcllwraith Range, NE of Coen, Oct 1969, Webb & Tracey

8355 (BRI); S.F.144, Agapetes L.A., Dec 1979, Hyland

10174 (BRI, QRS); S.F.191, Parish of Barron, Dec 1991,

Hyland 14405 (BRI, QRS); S.F.194, Parish of Western, Mar

1994, Hyland 15052 (QRS); Klondike Mine road, May 1995,

Hyland 15329 (QRS); Shiptons Flat, Jun 1996, Jago 4025

& Roberts (BRI); Wongabel S.F.191, Nov 1997, Forster

PEF21929 etal. (BRI, MEF, QRS). North Kennedy District:

Tully Falls, Oct 1948, Fielding (QRS); Elphinstone Ck,

Coldwater, 24 km W of Ingham, Jun 1972, Set on (BRI);

‘Bellview’, Evelyn, Jan 1974, Collins (BRI); Elliots Toe, Mt

Elliot, Jun 1990, Camming 9965 (BRI); Keoghs Scrub,

Portion 205, Parish of Herberton, Oct 1993, Gray 5717 (QRS).

Distribution and habitat : Solanum cookii is

endemic to Queensland. Sporadically

distributed along the east coast from Coen to

Townsville (Map 12). It grows on disturbed

sites in microphyll or tall notophyll rainforest,

or on the margin with woodland communities.

Altitude is usually between 500-1100 metres,

but it has occasionally been found near sea level.

Phenology : Flowers are recorded from

November to February, and also June; mature

fruits are recorded from October to February,

and also in June.

Notes: S. cookii has very long finger hairs on

upper leaf surface, while on the lower surface,

finger hairs are mixed with stellate hairs (lateral

rays often poorly developed).

Conservation status: S. cookii has been

collected from about 7 locations in north

Queensland over the last 10-15 years. It is not

currently known from a conservation reserve.

Applying the IUCN guidelines (IUCN. 2001),

a category of “Near Threatened” is

recommended.

36. Solanum multiglochidiatum Domin,

Biblioth. Bot. 89: 586 (1929). Type:

Queensland. Cook District: near

Mungana, February 1910, K. Domin

(holo: ?PR), n.v.

Austrobaileya 6 (4): 639-816 (2004)

Illustration: Symon (1981: 232)

Prostrate or sprawling, herbaceous resprouter,

0.2-0.4 m high. Juvenile stage absent. Adult

branchlets brown; prickles 140-320 per

decimetre, straight, acicular, 1-6 mm long,

13-20 times longer than wide; stellae sparse

to dense, 0.3-0.45 mm diameter, stalks 0-0.1

mm long; lateral rays 5-8, porrect; central ray

0.3-0.7 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Adult leaves broadly ovate or orbicular, entire

or shallowly lobed throughout; lobes 2 or 3 on

each side, obtuse, lobing index 1-1.1; lamina

6.5-9.5 cm long, 3.4-6 cm wide, 1.1-2.1 times

longer than broad, apex obtuse, base cuneate

or obtuse, oblique part 0-6 mm long, obliqueness

index 0-6 percent; petioles 1.2-1.7 cm long,

13-25% length of lamina, prickles present.

Upper leaf surface green; prickles 100-400,

straight, acicular, 1-6 mm long, prickles

present on midvein and lateral veins; stellate

hairs distributed throughout; protostellae

absent; ordinary stellae very sparse to sparse,

0.8-2.5 mm apart, 0.25-0.45 mm across,

sessile; lateral rays 3-7, porrect; central ray

0.8-1.6 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Lower leaf surface green; prickles 40-250,

straight, acicular, present on midvein and

lateral veins; stellae sparse to dense, 0.3-1.4

mm apart, 0.5-0.7 mm diameter, stalks 0-0.1

mm long; lateral rays 6-8, porrect; central ray

0.5-1 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Inflorescence supra-axillary, pseudo-racemose,

common peduncle 17-27 mm long, rachis

prickles present; 6-10-flowered, weakly

andromonoecious, flowers 5-merous; pedicels

8-14 mm long at anthesis, same thickness

throughout, 0.5-0.8 mm thick at mid-point,

prickles present. Calyx tube 2.5-4 mm long,

lobes attenuate, 3-6 mm long; prickles present

at anthesis, 36-50 per flower, 1.5-4 mm long;

stellae moderate to dense, transparent, 0.4-0.5

mm across, stalks 0-0.1 mm long, lateral rays

4-8, central ray 0.8-1.2 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Corolla white or mauve,

13—18 mm long, rotate, inner surface glabrous;

anthers 6.5-8.5 mm long; ovary with Type 2

hairs only, or with stellate and Type 2 hairs;

functional style 11-14 mm long, sigmoid, with

Type 2 hairs only or with stellate and Type 2

Bean, Taxonomy of Solanum subg. Leptostemonum

hairs, stellae 0.2-0.4 mm across, lateral rays

6-8, central ray 1-2 times as long as laterals.

Fruiting calyx with lobes less than or more than

half length of mature fruit, prickles 1.5-4 mm

long. Mature fruits 1-6 per inflorescence,

globular, 16-21 mm diameter, yellowish-green

or green, 1-locular (septum absent or

incomplete); placenta in cross-section sessile,

elliptical; mesocarp moist but not juicy; exocarp

0.4-0.5 mm thick; pedicels 18-23 mm long in

fruit, 1.4-2 mm thick at mid-point; seeds pale

yellow, 3.3-3.5 mm long.

Specimens examined : Queensland. Cook District: 24 km

W of Petford and c. 48 km SE of Chillagoe, May 1967, Symon

4873 (AD, BRI, CANB, K, L, NSW); Palmer River, Feb

1978, Hinton 61 (BRI); Holmes Creek, 3.5 km WNW of Mt

Carbine, Jan 1984, Clarkson 5105 (AD, BRI, CANB, K,

MEL, MO, QRS); 12.6 km SE of Mt Janet, 11.5 km SW of

Lakeland Downs township, Jan 1986, Clarkson 6293 (AD,

BRI, DNA, PERTH, QRS); 9 km SW of Lakeland Downs

township, Jan 1986, Clarkson 6298 (AD, BRI); 4 km SE of

Chillagoe, Jan 1996, Gray 6506 (QRS); 4.7 km from Walsh

River crossing, N of Chillagoe, Mar 2000, McDonald

KRM335 (BRI); 14 km W of Chillagoe, Apr 2002, Bean

18734 & McDonald (BRI). North Kennedy District: 40 Mile

Scrub, [SW of Mt Garnet], Mar 1989, Sankowsky 979 &

Sankowsky (BRI).

Distribution and habitat : Solanum

multiglochidiatum is endemic to Queensland,

extending from Lakeland Downs to Forty Mile

Scrub (Map 8). It usually grows in shrubby

eucalypt woodland dominated by Eucalyptus

cullenii, E. leptophleba or E. dallachiana, on

gently undulating terrain, in clay-loam soils.

There is one record from deciduous microphyll

vine thicket.

Phenology: Flowers are recorded from January

to March; mature fruits from January to May.

Notes: The type of S. multi glochidiatum was

sought from PR, but not received. Symon

(1981) stated that he had not seen the type. It

is possible that the type is missing, but the

designation of a neotype is not warranted until

further searches are made. The application of

the name is not in doubt, as the species is well

described in the protologue.

Conservation status: S. multiglochidiatum has

been collected from about 7 locations in north

Queensland over the last 15 years. It is not

known from a conservation reserve. Applying

the IUCN guidelines (IUCN. 2001), a category

of “Near Threatened” is recommended. It is

725

currently listed as “Rare” under the Queensland

Nature Conservation Act, 1992.

37. Solanum vicinum A.R.Bean sp. nov. Frutex

parvus, aculeis abundis in ramulis foliis

calycibusque; stellae ramulorum 0.4-0.5

mm diametro; folia non profunde lobata,

ovata usque late ovata, utrinque viridia;

pedunculus communis in inflorescentia

praesens; corolla 11-18 mmlonga; stylus

functionalis sigmoideus; fructus

maturitate purpureus, globularis, 24-30

mm diametro. Typus: Queensland.

Darling Downs District: adjacent to

Gambubal State Forest, NE of Killarney,

7 October 2000, A.R. Bean 16891 (holo:

BRI (1 sheet + spirit); iso: MEL, MO,

NSW).

[S. prinophyllum auct., non Dunal]

Erect, rhizomatous perennial shrub, 0.2-0.9 m

high. Juvenile branchlets with 200-500 prickles

per dm, 4-9 mm long; leaves (in outline)

broadly ovate, shallowly-lobed, with 3-5 pairs

of lobes; lamina 6-7 cm long, 5-6 cm wide,

with c. 100 prickles on upper surface. Adult

branchlets brown or green; prickles 90-180 per

decimetre, straight, acicular, 1-11 mm long,

12-18 times longer than wide; stellae sparse,

0.4-0.5 mm diameter, stalks 0-0.15 mm long;

lateral rays 5-8, porrect; central ray 0.5-1.2

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Adult

leaves ovate, shallowly lobed throughout; lobes

4-6 on each side, acute, lobing index 1.4-1.9;

lamina 7.5-14 cm long, 4-9.5 cm wide,

1.5- 2.1 times longer than broad, apex

acute, base cuneate or obtuse, oblique part

1.5- 7 mm long, obliqueness index 1-8

percent; petioles 1.7-3.7 cm long, 18-30%

length of lamina, prickles present. Upper

leaf surface green; prickles 100-200,

straight, acicular, 3-11 mm long, prickles

present on midvein and lateral veins;

stellate hairs distributed throughout;

protostellae absent; ordinary stellae density

sparse to moderate, 0.6-1.3 mm apart, 0.4-0.5

mm across, sessile; lateral rays 4-8, porrect;

central ray 0.8-1.5 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Lower leaf surface green; prickles

100-200, straight, acicular, present on midvein

and lateral veins; stellae very sparse to sparse,

726

Austrobaileya 6 (4): 639-816 (2004)

QUEENSLAND HERBARIUM (BRI)

Flora ct Que«ii*l*rtd

Oirfcng Downs

QUEENSLAND HERBAHIUM (BRI)

Brisbane AusifiSsi

SolliiUfn prfnoptiyNum Dunal

Col. A ft Bean 16W*

2801*11164 4Ss

t&_44V&2JW7«29B}

Adj«*nt to GanUjuMl SF. HE pr KjPamey

Hotels* r*ntor*»i a&Pul 2-3 )W‘ a)?®

Rod b**aHc sod

SpreadS*vn* 80cm high, flowers py*»t*

Hunditf* or plain SMrk. f**«y pn intg Ira-^s.

5p.nl material *1 BRI

D4C

Ogp MOKSWMfL

■ May bo cited as cwr.pute.-rM0 CtiflKW Humber

(Artfuval Paper)

(&$9/

AQ 491705

tt&L&TyfBt,/ QueansJand Haiti atium <8RI>

So/enia* t'/onot#* st.A.£e*w

oct Di!t tecrZoos

Fig. 25. Holotype of Solanum vicinum.

Bean, Taxonomy of Solanum subg. Leptostemonum

0.7-2.2 mm apart, 0.5-0.7 mm diameter,

sessile; lateral rays 4-8, porrect; central ray

0.5-1.5 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Inflorescence supra-axillary, pseudo-racemose,

common peduncle 0-8 mm long, rachis prickles

present; 3-8-flowered, flowers 5-merous;

pedicels 6-15 mm long at anthesis, same

thickness throughout, 0.3-0.7 mm thick at mid¬

point, prickles present. Calyx tube 3-4.5 mm

long, lobes attenuate, 3.5-10 mm long; prickles

present at anthesis, 50-100 per flower, 1-8 mm

long; stellae moderate, yellow or white, 0.3-0.5

mm across, sessile, lateral rays 4-7, central ray

0.5-1 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Corolla purple, 11-18 mm long, rotate or

shallowly lobed, inner surface glabrous; anthers

3.5-5 mm long; ovary with Type 2 hairs only;

functional style 8-12 mm long, sigmoid,

glabrous or with Type 2 hairs only. Fruiting

calyx with lobes less than or more than half

length of mature fruit, prickles 1-8 mm long.

Mature fruits 1 or 2 per inflorescence, globular,

24-30 mm diameter, purple, 2-locular; placenta

in cross-section stalked, anvil-shaped;

mesocarp moist but not juicy; exocarp 3-3.5

mm thick; pedicels 20-34 mm long in fruit,

1.3-2 mm thick at mid-point; seeds pale yellow,

1.9-2.4 mm long. Fig. 25.

Specimens examined : Queensland. Wide Bay District:

Peters L.A., Conondale Ranges, Dec 1990, Bean 2699 (BRI).

Darling Downs District: c. 1 km ENE of Gambubal Forest

Station, E of Warwick, Dec 1996, Bean 11426 (BRI);

Goomburra S.F., c. E5 km W of Mt Castle Lookout carpark,

NE of Warwick, Jan 2003, Bean 19929 (BRI). Moreton

District: Mt Glorious, Jan 1965, Henderson H104 (BRI);

D’Aguilar Range, on top of Tenison Woods Mt, Jul 1974,

Swan 55 (BRI); Lewington Road, Mt Mee, Sep 1992, Symon

s.n. (BRI). New South Wales. Northern Tar t. e l ands:

HillgrovenearArmidale, Sep l91l,McBarron, 20291 (BRI,

NSW). North Coast: Dorrigo S.F., Oct 1930, White 7809

(BRI); c. 6 miles [10 km] along Doyles River road, c. 48

miles [77 km] W of Wauchope, Oct 1951, Ford s.n. (AD,

BRI, NSW); Toonumbar S.F., 21 miles [34 km] NW of

Kyogle, Sep 1972, Coveny 4571 & Rodd (BRI, NSW); near

Mt Boss, NW of Wauchope, Nov 1999, Bean 15712 (BRI,

NSW); Welsh’s Road, Clouds Creek S.F., N of Dorrigo, Sep

2000, Bean 16855 (BRI, CANB, MEL, NSW); Bellangry

S.F. lookout, NW of Wauchope, Dec 2000, Bean 17230 (BRI,

NSW); Armidale - Kempsey road, 10.7 km S of Styx River,

Dec 2000, Bean 17233 (BRI, NSW).

Distribution and habitat : Solanum vicinum is

uncommon in Queensland, known from just a

few locations as far north as Conondale Ranges

(Map 17). Reasonably widespread in north-

727

eastern N.S.W., as far south as Wauchope. It

grows in notophyll rainforest, or on the margins

of same, usually at relatively high altitude.

Phenology: Flowers are recorded from

September to February; mature fruits are

recorded in May, September and December.

Notes: Closely related to Solanum

prinophyllum Dunal, and regarded by Symon

(1981) as the northern form of that species.

S. vicinum differs from S. prinophyllum by:

branchlets with 90-180 prickles per decimetre

(30-65 for S. prinophyllum ), branchlet stellae

0.4-0.5 mm diameter (0.25-0.3 mm diameter

for S. prinophyllum ), shallowly-lobed leaves

(lobing index of 1.4-1.9 vs. index of 2-4 for

S. prinophyllum), inflorescences 3-8 flowered

with common peduncle usually present and up

to 8 mm long (inflorescences 1-3 flowered,

common peduncle absent for S. prinophyllum ),

and corolla 11-18 mm long (7-9 mm long for

S. prinophyllum ).

Solanum prinophyllum sens. str. is

perhaps more closely related to S. pungetium

R. Br. than it is to S. vicinum. S. prinophyllum

and S. pungetium are not easily separable, but

I have noted the following differences, based

on collections from central and southern coast

of N.S.W.: The leaves and branchlets of

S. prinophyllum are glabrous to sparsely hairy

(always moderately to densely hairy for

S. pungetium)-, the stellae of the upper leaf

surface in S. prinophyllum (when present) are

0.2-0.35 mm diameter, with 6-8 lateral rays

and the central ray 0-0.7 times as long as

laterals. In S. pungetium, the corresponding

stellae are 0.35-0.5 mm diameter, with 4 or 5

lateral rays and a central ray 1-1.5 times as

long as laterals; the leaves, pedicels and calyces

of S. prinophyllum tend to have a greater

number of prickles; S. prinophyllum leaves have

4-6 pairs of lobes, and the lobes may themselves

have small lobes or angles, whereas S. pungetium

leaves have 3 or 4 pairs of lobes, and the lobes

are always “simple”.

The few specimens I have seen from

Victoria suggest that the patterns of variation

are different again, and if both species are

present there, they are even less distinct.

While I feel sure that S. prinophyllum and

S. pungetium differ sufficiently to warrant

728

Austrobaileya 6 (4): 639-816 (2004)

species status for both, it will require a detailed

field study to determine the exact relationships

involved. The character most often used to

separate S. pungetium in identification keys,

namely the reduced 1 or 2 flowered inflorescence

without a common peduncle, is not diagnostic.

Both species can have this feature.

Conservation status : Widespread. Not

considered at risk.

Etymology: From the Latin vicinus - near,

neighbouring. This is a reference to its close

affinity to S. prinophyllum.

38. Solanum papaverifolium Symon, Trans.

& Proc. Roy. Soc. South Australia 95:

233-4 (1971). Type: New South Wales.

‘Maneroo’, Graman, c. 56 km north-west

oflnverell, 11 June 1969, V.N. Gidley s.n.

(holo: NSW; iso: AD, BRI, CANB, K,

NSW).

Illustration: Symon (1981: 180)

Prostrate or sprawling, herbaceous resprouter,

0.2-0.4 m high. Juvenile stage absent. Adult

branchlets brown or green; prickles 15-40 per

decimetre, straight, acicular, 1-6 mm long,

10-13 times longer than wide; stellae absent;

finger hairs absent; Type 2 hairs absent, or

sparse. Adult leaves broadly ovate, deeply lobed

throughout; lobes 4-6 on each side, acute,

lobing index 4-27; lamina 4-9 cm long, 2.6-6.5

cm wide, 1.4-1.8 times longer than broad, apex

acute, base cuneate, oblique part 0-4 mm long,

obliqueness index 0-5 percent; petioles 2.1-3.4

cm long, 25-60% length of lamina, winged,

prickles present. Upper leaf surface green;

prickles 20-50, straight, acicular, 0.5-7 mm

long, prickles present on midvein and lateral

veins; stellate hairs absent; finger hairs absent;

Type 2 hairs absent. Lower leaf surface green;

prickles 30-150, straight, acicular, present on

mid vein and lateral veins; stellae absent; finger

hairs absent; Type 2 hairs absent. Inflorescence

leaf-opposed, pseudo-racemose, common

peduncle 24-33 mm long, rachis prickles

present; 3-5-flowered, strongly or weakly

andromonoecious, flowers 5-merous; pedicels

4-23 mm long at anthesis, same thickness

throughout, 0.5-0.6 mm thick at mid-point,

prickles present. Calyx tube 2-4 mm long, lobes

deltate or attenuate, 2.5-6 mm long; prickles

present at anthesis, 12-40 per flower, 1-5 mm

long; stellae absent; finger hairs absent; Type

2 hairs present. Corolla purple, 7-11 mm long,

rotate, inner surface glabrous; anthers 3.5-5

mm long; ovary glabrous, or with Type 2 hairs

only; functional style 4.5-6 mm long, erect,

glabrous or with Type 2 hairs only. Fruiting

calyx with lobes more than half length of

mature fruit, prickles 1-6 mm long. Mature

fruits 1 or 2 per inflorescence, globular,

yellowish-green or green; mesocarp moist but

not juicy; pedicels 28-37 mm long in fruit,

0.9-1.3 mm thick at mid-point.

Specimens examined : Queensland. Darling Downs

District: Jimbour, Dec 1875, Bailey (BRI); ‘Kuyura’, Dec

1931, Cadell (BRI); Dalby, Nov 1941, McMahon (BRI);

‘Yandilla’, Nov 1951, Everist s.n. (BRI); Macalister, Jan

1942, McCoy (BRI); ‘Paxton’, Pirrinuan, via Dalby, Feb

1953, Guard (BRI); Brookstead, Apr 1959, Keeley (BRI);

Hermitage Regional Experimental station, Warwick, Apr

1959, Seton 8 (BRI); Branch Creek road, Dalby, Dec 1970,

Mohen (BRI); ‘Kuyura’, near Jimbour, Jan 1985, Jensen

(BRI); 19 km SSW of Dalby, Feb 1995, Fensham 1912

(BRI); 5 km SE of Oakey, Feb 1995, Fensham 2085 (BRI);

Clifton-Leybum road, near Millbrook road turnoff, Jan 2002,

Bean 18353 (BRI, MET., NSW). New South Wales. North

West Slopes : Bingara, Mar 1901, McColl s. n. (NSW). North

West Plains. ‘Karina’, Moree, May 1971, Strange s.n.

(NSW).

Distribution and habitat: Solanum

papaverifolium has been recorded from between

Jimbour and Warwick in Queensland (Map 19).

In New South Wales, it has been found from

Inverell to Quirindi and Singleton, and west to

Narrabri and Moree. It grows on heavy clay

soil, in grassland or open eucalypt woodland.

Phenology: Flowers are recorded from October

to March; mature fruits from November to

April.

Notes: F.M. Bailey, in 1875, was the first to

collect the species in Queensland. Charles

Moore had earlier collected it from “Liverpool

Plains” in New South Wales. The next

Queensland collection was in 1931, when a

specimen was forwarded to C.T. White. He

identified it as S. hystrix R.Br., a species from

southern Australia, though he probably did not

have access to authentic S. hystrix specimens.

In 1939, while at Kew, he examined it again

and determined that it was not S. hystrix, but

“evidently a native”.

In S. papaverifolium, the outer surface of

the corolla often bears prickles. Only a few other

Bean, Taxonomy of Solanum subg. Leptostemonum

species (including S. hystrix ) display this

characteristic.

Conservation status : S. papaverifolium is

currently known from 3 locations. It grows on

soils that are utilized for agriculture. All

populations are threatened by weeds, roadworks

and agriculture, and none is protected in a

conservation reserve. Currently listed as

“Endangered” under the Queensland Nature

Conservation Act, 1992.

Applying the IUCN guidelines (IUCN.

2001), the existing category of “Endangered”

is endorsed (EN A3ce; B2ab(iii,v); Cl).

The most recent N.S.W. collection was

made in 1982, from Moree.

39. Solanum adenophorum F.Muell., Fragm.

2: 162 (1861); S. adenophorum var.

adenophorum Domin, Biblioth. Bot. 89:

586 (1929); S. adenophorum var. typicum

Domin, Biblioth. Bot. 89: 586 (1929),

nom. inval. Type: [Queensland.] between

the Dawson and Mackenzie Rivers, 17-

20 November 1856, F. Mueller (holo:

MEL).

Prostrate or sprawling, herbaceous resprouter,

0.15-0.3 m high. Juvenile stage absent. Adult

branchlets grey or brown; prickles absent or

present, 0-15 per decimetre, straight, acicular,

6-10 mm long, 12-17 times longer than wide;

stellae absent; finger hairs present, gland-

tipped, 0.4-0.8 mm long; Type 2 hairs dense.

Adult leaves ovate or broadly ovate, deeply

lobed throughout; lobes 3 or 4 on each side,

obtuse, lobing index 2.3-5; lamina 3.5-5.5 cm

long, 2-4 cm wide, 1.3-1.8 t im es longer than

broad, apex obtuse, base cuneate to cordate,

oblique part 0-6 mm long, obliqueness index

0-10 percent; petioles 1.2-4 cm long, 30-80%

length of lamina, winged, prickles present.

Upper leaf surface green; prickles 6-20,

straight, acicular, 3-13 mm long, prickles

present on midvein and lateral veins; stellate

hairs absent; finger hairs present, 0.1-0.3 mm

apart, gland-tipped, 0.5-0.9 mm long; Type 2

hairs absent. Lower leaf surface green; prickles

11-25, straight, acicular, present on midvein

and lateral veins; stellae absent; finger hairs

present, 0.3-0.5 mm apart, gland-tipped, 0.3-0.6

mm long; Type 2 hairs absent. Inflorescence

supra-axillary, pseudo-racemose, common

729

peduncle 9-26 mm long, rachis prickles present;

4- 8-flowered, strongly andromonoecious,

flowers 5-merous; pedicels 5-10 mm long at

anthesis, same thickness throughout, 0.4-0.5

mm thick at mid-point, prickles absent or

present. Calyx tube 2-3 mm long, lobes deltate,

2-4.5 mm long; prickles present at anthesis,

5- 20 per flower, 3-6 mm long; stellae absent;

finger hairs present; Type 2 hairs absent.

Corolla white, 8-10 mm long, deeply lobed,

inner surface glabrous; anthers 4.5-5.5 mm

long; ovary with Type 2 hairs only; functional

style 6-7 mm long, erect, with Type 2 hairs

only. Fruiting calyx with lobes less than or more

than half length of mature fruit, prickles 1-6

mm long. Mature fruits 1 per inflorescence,

globular, c. 15 mm diameter, yellowish-green

or green; pedicels 9-18 mm long in fruit,

0.9-1.2 mm thick at mid-point; seeds pale

yellow, 2.8-3.5 mm long.

Specimens examined: Queensland. South Kennedy

District: Logan Downs Pty Ltd, ‘Wentworth’, 68 miles [109

km] N of Clermont, Oct 1957, Saclier 3 (BRI); 115 kmNW

of Clermont, Jul 1977, Dale 148 (BRI). Leichhardt District:

Blackwater Creek, anno 1871, Bowman s.n. (MEL);

Marlborough-Sarina road, c. 45 miles [72 km] from

Marlborough, May 1960, Johnson 1779 (BRI); Dipperu N.R,

c. 24kmS ofNebo, Sep 1971, McDonald 143 (BRI); along

boundary fence, Taunton N.R, Oct 1995, Brnshe JB408 et

al. (BRI); Taunton N.R, Oct 1996, Porter JB836 (BRI); Red

Hill section of Taunton N.P., just N of Dingo, Aug 1998,

Melzer 994 & Clarke (BRI).

Distribution and habitat : Solanum

adenophorum is endemic to Queensland, in the

Dingo-Nebo-Clermont area, west and north¬

west of Rockhampton (Map 13). It is recorded

mainly from brigalow ( Acacia harpophylla )

communities, but also from gidgee ( Acacia

cambagei) woodland. Soils are deep cracking clays.

Phenology: flowers recorded in October;

mature fruits recorded for May, September and

October.

Notes: S. adenophorum has very long petioles

compared to most other Australian species. The

longest petioles are on the lowermost leaves,

hence it seems that the petioles continue to

elongate after the leaf has fully expanded. The

taxon known as S. adenophorum in New South

Wales (Conn 1992) and Victoria (Jeanes 1999) is

referable to S. eremophilum F.Muell. (Bean 2002a).

Conservation status: Currently listed as

‘Endangered’ under the Queensland Nature

730

Conservation Act, 1992. It is threatened by

habitat clearance, and by introduced weeds such

as Parthenium hysterophorus, Cenchrus

ciliaris, Opuntia spp. and Acanthocereus

tetragonus. It is currently known only from a

single location, viz. Taunton N.P.

Applying the IUCN guidelines (IUCN.

2001), a category of “Critically Endangered”

is recommended (CR B2ab(iii,v)).

40. Solanum pusillum A.R.Bean sp. nov.

Frutex prostratus vel fusus; aculei in

ramulis foliis calycibusque praesentes;

folia viridia, profunde lobata, in paginis

superioribus stellis et digitis commixtis;

petioli alati, longitudine 8-18% laminae

aequantes; pili Type 2 in foliis calyce

ovario styloque; inflorescentia pseudo-

umbellata; corolla alba, 7-10 mm longa;

fructus maturitate virides. Typus:

Queensland. Leichhardt district:

western road, Junee State Forest, Junee

Tableland, north of Dingo, 5 October

2002, A.R. Bean 19411 (holo: BRI).

Prostrate or sprawling, herbaceous resprouter,

0.1-0.3 m high. Juvenile stage absent. Adult

branchlets brown or green; prickles 10-50 per

decimetre, straight, acicular, 1-9 mm long,

10-16 times longer than wide; stellae sparse

to dense, 0.8-1.2 mm diameter, stalks 0-0.1

mm long; lateral rays 4-7, porrect; central ray

0.8-1.5 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent,

or sparse. Adult leaves elliptical or ovate, deeply

lobed throughout; lobes 2-4 on each side, acute

or obtuse, lobing index 2.3-4; lamina 3-5.5

cm long, 1.4-2.5 cm wide, 1.6-2.2 times longer

than broad, apex obtuse or acute, base cuneate,

oblique part 0-3 mm long, obliqueness index

0-7 percent; petioles 0.8-1.8 cm long, 8-18%

length of lamina, winged, prickles absent or

present. Upper leaf surface green; prickles

10-50, straight, acicular, 2-8 mm long, prickles

present on midvein and lateral veins; stellate

hairs distributed throughout; protostellae

present; ordinary stellae very sparse to sparse,

0.8-1.7 mm apart, 0.3-1.1 mm across, sessile;

lateral rays 1-4, porrect or ascending; central

ray 1-4 times as long as laterals, not gland-

tipped; finger hairs present, 0.5-3 mm apart,

not gland-tipped, 0.8-1.3 mm long; Type 2

hairs present throughout, 0.1-0.4 mm apart.

Austrobaileya 6 (4): 639-816 (2004)

Lower leaf surface green, or yellowish; prickles

4-17, straight, acicular, present on midvein and

lateral veins; stellae sparse to moderate, 0.2-1

mm apart, 0.7-1 mm diameter, stalks 0-0.1 mm

long; lateral rays 4-7, porrect; central ray

0.8-1.5 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs present

throughout, 0.05-0.2 mm apart. Inflorescence

supra-axillary, solitary or pseudo-umbellate,

common peduncle absent; 1-3-flowered,

strongly andromonoecious, flowers 5-merous;

pedicels 3-6 mm long at anthesis, same

thickness throughout or markedly thicker

distally, 0.4-0.6 mm thick at mid-point,

prickles present. Calyx tube 1.5-2 mm long,

lobes deltate, 1.5-3 mm long; prickles present

at anthesis, 20-35 per flower, 2.5-6 mm long;

stellae sparse to moderate, white, 0.7-1.1 mm

across, sessile, lateral rays 4-8, central ray 1-1.5

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs present.

Corolla white, 7-10 mm long, deeply lobed,

inner surface glabrous; anthers 3-4 mm long;

ovary with Type 2 hairs only; functional style

4-4.5 mm long, erect, with Type 2 hairs only.

Fruiting calyx with lobes less than or more than

half length of mature fruit. Mature fruits 1 per

inflorescence, globular, c. 15 mm diameter,

green, 2-locular; placenta in cross-section sessile,

semi-circular; mesocarp moist but not juicy;

exocarp 0.8-1.3 mm thick; pedicels 12-18 mm

long in fruit, 0.9-1.5 mm thick at mid-point;

seeds pale yellow, 3.8-4 mm long. Fig. 6,26.

Specimens examined : Queensland. Leichhardt District:

Junee Tableland, N of Dingo, Nov 1990, Bean 2604 (BRI);

near eastern road, Junee Tableland, N of Dingo, Oct 2002,

Bean 19399 (BRI); eastern edge of S.F.236, SW of

Blackwater, Nov 2002, Bean 19519 (BRI, MEL); ditto, Bean

19525 (BRI); S of Triumph Ck, c. 30 km SW of Blackwater,

Nov 2002, Bean 19538 (BRI).

Distribution and habitat : Solanum pusillum is

endemic to Queensland. Known from two areas

in the central east of the state (Map 14). It

usually grows in shallow yellowish soil on the

edges of low plateaux, where the dominant

species may be Acacia catenulata, Eucalyptus

trachyphloia, E. tenuipes or E. exserta.

Phenology : Poorly known. Flowers and fruits

are recorded for October and November.

Notes: Closely related to S. adenophorum, but

differing by having only stellate hairs on the

Bean, Taxonomy of Solanum subg. Leptostemonum

731

Fig. 26. Solanumpusillum. A. flowering branchlet x 1. B. ovary and style x 9. C. stellate hair with 4 lateral rays, upper leaf

surface x 30. D. finger hair, upper leaf surface x 30. E. stellate hair with 2 lateral rays, upper leaf surface x 30. F. mature fruit,

calyx and pedicel x 2. G. transverse section of fruit x 3. all from Bean 19411.

732

branchlets and lower leaf surface, petioles

8-18% of lamina length (30-80% for

S. adenophorum ), presence of Type 2 hairs on

the lower leaf surface and calyx, 1-3 flowered

pseudo-umbellate inflorescence (4-8 flowered

and pseudo-racemose for S. adenophorum) and

style 4-4.5 mm long (6-7 mm for S. adenophorum).

Conservation status: S. pusillum is known to

exist in 7 subpopulations, of which four are on

the Junee Tableland. It does not occur in any

conservation reserve. Applying the IUCN

guidelines (IUCN. 2001), a category of “Near

Threatened” is recommended.

Etymology : From the Latin pusillus meaning

tiny or puny, a reference to the small size of

the plant.

41. Solanum graniticum A.R.Bean sp. nov.

Repullulator herbaceus prostratus vel

fusus; aculei in ramulis praesentes; folia

adulta 1.2-2.6 cm longa, profunde vel

non profunde lobata, stellis in pagina

inferiore sparse praeditis; inflorescentia

1 vel 2-flora, pedunculo communi 0-1

mm longo; pedicelli sub anthesi 9-15 mm

longi; calyx aculeis 18-50 praeditus;

corolla malvina, non profunde lobata;

fructus maturitate virides. Typus:

Queensland. North Kennedy District:

Cape Gloucester, SW of Montes Resort,

17 March 1997, I.G. Champion 1419

(holo: BRI; iso: MEL).

Solanum sp. (Gloucester Island G.N.

Batianoff+ 9403312) in Henderson

( 2002 ).

Prostrate or sprawling, herbaceous resprouter,

0.15-0.3 m high. Juvenile stage absent. Adult

branchlets yellow or brown; prickles 10-45 per

decimetre, straight, acicular, 2.5-9 mm long,

10-16 times longer than wide; stellae dense,

0.25-0.5 mm diameter, sessile; lateral rays 7

or 8, porrect; central ray 0.3-0.7 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Adult leaves

elliptical or ovate, shallowly to deeply lobed

throughout; lobes 2 or 3 on each side, obtuse,

lobing index 1.5-3; lamina 1.2-2.6 cm long,

0.6-1.3 cm wide, 1.6-2.4 times longer than

broad, apex obtuse, base cuneate, oblique part

0-1 mm long, obliqueness index 0-4 percent;

petioles 0.15-0.45 cm long, 13-22% length of

Austrobaileya 6 (4): 639-816 (2004)

lamina, prickles absent. Upper leaf surface

green; prickles 1-3, straight, acicular, 1-5 mm

long, prickles present on midvein only; stellate

hairs confined to midrib, or distributed

throughout; protostellae absent; ordinary stellae

very sparse to sparse, 0.3-1 mm apart, 0.1-0.3

mm across, sessile; lateral rays 5-8, porrect;

central ray 0.5-1 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Lower leaf surface green; prickles

0-2, straight, acicular, absent or present on

midvein only; stellae sparse to moderate, 0.4-0.7

mm apart, 0.25-0.4 mm diameter, sessile;

lateral rays 7 or 8, porrect; central ray 0.5-1.2

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent.

Inflorescence leaf-opposed or supra-axillary,

solitary or pseudo-umbellate, common peduncle

0-1 mm long; 1 or 2-flowered, strongly

andromonoecious, flowers 5-merous; pedicels

9-15 mm long at anthesis, same thickness

throughout, 0.4-0.6 mm thick at mid-point,

prickles present. Calyx tube 2.5-3 mm long,

lobes deltate or rostrate, 2-5.5 mm long;

prickles present at anthesis, 18-50 per flower,

1.5-5 mm long; stellae moderate to dense,

transparent, 0.2-0.3 mm across, sessile, lateral

rays 6-8, central ray 0.5-1 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Corolla mauve, 9-14 mm

long, shallowly lobed, inner surface glabrous;

anthers 3.5-4.5 mm long; ovary with Type 2

hairs only; functional style 5.5-8.5 mm long,

erect, with Type 2 hairs only. Fruiting calyx

with lobes less than or more than half length

of mature fruit, prickles 2-5 mm long. Mature

fruits 1 per inflorescence, globular, c. 15 mm

diameter, green; mesocarp moist but not juicy;

pedicels 15-27 mm long in fruit, 0.7-0.9 mm

thick at mid-point; seeds pale yellow, 2.7-3.2

mm long. Fig. 27.

Specimens examined : Queensland. North Kennedy

District: ridge 1 km S of Mt Bertha, Gloucester Island, Mar

1994, Batianoff 9403312 et al. (BRI); SE Point, above

Chinaman Rock, Gloucester Island, May 1994, Batianoff

940592 & Dillewaard (BRI); c. 1.5 km due E of Nelly Bay,

Mar 1999, Kemp TH347 & Allison (BRI). South Kennedy

District: lookout at Eungella Dam, W of Eungella, Feb 2003,

Bean 20061 & Champion (BRI).

Distribution and habitat: Solanum graniticum

is endemic to Queensland. Found on Gloucester

Island (near Bowen), and adjacent parts of the

mainland, and with a disjunct occurrence at

Eungella Dam (Map 16). It grows in open

Bean, Taxonomy of Solanum subg. Leptostemonum

733

QUEENSLAND HERBARIUM (BRI|

Flora trf Queensland

Norm Kennedy

Solanum elnoreum R-Br.

Co®. l e-CHamjMOfl 1419

If MAR 1997

QUEENSLAND HERBARIUM [BR!)

Brisbana AusUa'ta

AQ 656070

i o-T J

20’0-'S 14,B’2-'£

AH rrt. Queensland Herbarium (BRl)

oopih m. fiUeryPFaf

Queensland Herbarium |BRI>

xj, m 4«gs^&’ ft, /■ f$0}}!2,)

cape Gloucester SW d Montes Resort _

Open woodland with scattered targe shrubs on lop of rocky

terrace. soii-orarUe derived-hard packed.

Prickly sub shrub, stellate hales preseni. Rswe * 5 .

wdhsinguVar lobes Frurt globose, pala green ™« h <***

geeen streaks, widest at baso

Scarce to occasional

DM. M.A.MC.Gcwan APR 1997 Sofaweae

cried as compare™* afcdon nuodwr AO 656(179

[Art)** riapeh

Do).

Cht u

Fig. 27. Holotype of Solanum graniticum.

734

Austrobaileya 6 (4): 639-816 (2004)

eucalypt woodland on hillsides with shallow

soil derived from granite or granodiorite.

Phenology: Flowers are recorded for March;

mature fruits for March and May.

Notes : S. graniticum is closely related to

S. pusillum, but S. graniticum differs by the

relatively small leaves, petioles without wings,

the lack of finger hairs and Type 2 hairs, the

smaller stellae on all plant parts, the mauve,

shallowly-lobed corolla, and the longer

pedicels.

Conservation status: S. graniticum is known

from 4 locations, 3 of them in close proximity.

It occurs in the Gloucester Island N.P.. Most

populations are threatened by road and housing

construction, weeds, and grazing. Applying the

IUCN guidelines (IUCN. 2001), a category of

“Endangered” is recommended (EN

B1 ab(ii,iii,v)+2ab(ii,iii,v); C1 +2a(i)).

Etymology: The epithet refers to the granite

substrate where this species is found.

42. Solanum stenopterum A.R.Bean sp. nov.

Repullulator herbaceus; folia adulta

lobata vel integra utrinque viridia; petioli

alati; stellae in pagina superiore folii

radiis lateralibus 3-5 et radio centrali

comparate longo praeditae; inflorescentia

1 vel 2-flora pedunculo communi carens;

pedicelli 21-28 mm longi; aculei calycis

praesentes; corolla purpurea. Typus:

Queensland. Darling Downs District:

Warrego Highway, 10 km NW of Oakey,

13 December 2001, A.R. Bean 18190

(holo: BRI (1 sheet + spirit); iso: AD,

CANB, MEL, NSW).

Solanum sp. (Dalby R.F. Kelsey 56) in

Henderson (2002).

Sprawling or erect, herbaceous resprouter,

0.2-0.4 m high. Juvenile stage absent. Adult

branchlets green; prickles 7-15 per decimetre,

straight, acicular, 2-8 mm long, 10-15 times

longer than wide; stellae sparse, 0.4-1.2 mm

diameter, sessile; lateral rays 4-8, porrect;

central ray 0.7-1.4 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Adult leaves linear to ovate, entire

or shallowly to deeply lobed throughout; lobes

2 or 3 on each side, acute or obtuse, lobing index

1- 5; lamina 4-7 cm long, 0.5-2.6 cm wide,

2.2-12 times longer than broad, apex acute,

base cuneate, oblique part 0-3 mm long,

obliqueness index 0-5 percent; petioles 0.5-1.3

cm long, 10-18% length of lamina, winged,

prickles absent. Upper leaf surface green;

prickles 3-15, straight, acicular, 1-4 mm long,

prickles present on midvein only or present on

midvein and lateral veins; stellate hairs

confined to midrib, or distributed throughout;

protostellae absent; ordinary stellae very sparse

to moderate density, 0.8-2.5 mm apart, 0.3-1.1

mm across, sessile; lateral rays 3-5, porrect;

central ray 1-3 times as long as laterals, not

gland-tipped; finger hairs absent or present, 1-3

mm apart, not gland-tipped, 0.6-1.2 mm long;

Type 2 hairs absent. Lower leaf surface green;

prickles 1-15, straight, acicular, present on

midvein only or present on midvein and lateral

veins; stellae very sparse to moderate, 0.8-3

mm apart, 0.5-1 mm diameter, sessile; lateral

rays 4-7, porrect; central ray 1-2 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Inflorescence leaf-

opposed, solitary or pseudo-umbellate, common

peduncle absent; 1 or 2-flowered, strongly or

weakly andromonoecious, flowers 5-merous;

pedicels 21-38 mm long at anthesis, same

thickness throughout, 0.5-0.7 mm thick at mid¬

point, prickles absent or present. Calyx tube

2- 3 mm long, lobes deltate, 1.5-2.5 mm long;

prickles present at anthesis, 5-40 per flower,

1-4 mm long; stellae sparse to dense,

transparent, 0.3-0.6 mm across, sessile, lateral

rays 2-6, central ray 1-3 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Corolla purple, 14-18 mm

long, shallowly lobed, inn er surface glabrous;

anthers 3.5-5 mm long; ovary with Type 2 hairs

only; functional style 7.5-8.5 mm long, erect,

glabrous or with Type 2 hairs only. Fruiting

calyx with lobes more than half length of

mature fruit, prickles 2-4 mm long. Mature

fruits 1 per inflorescence, globular, c. 10 mm

diameter; pedicels 21-38 mm long in fruit, c.

1.5 mm thick at mid-point. Fig. 28.

Specimens examined: Queensland. Burnett District: c.

64 miles [103 km] WSW of Gayndah, Oct 1969, McPherson

(BRI). Darling Downs District: c. 5 km SW of Dalby, Mar

1958, Kelsey 56 (BRI); Condamine River, 4-5 miles [6-8

km] S of Dalby, Sep 1958, Kelsey 64 (BRI); ‘Burradoo’, 50

km NE of Goondiwindi, Mar 1979, Lahey (BRI); Myall Park,

Bean, Taxonomy of Solanum subg. Leptostemonum

735

Fig. 28. Solanum stenopterum. A. flowering branchlet x 0.8. B. base of leaf showing winged petiole x 3. C. ovary and style

x 6. D. ovary with Type 2 hairs x 12. E. stellate hair with 4 lateral rays, upper leaf surface x 60. F. finger hair, upper leaf surface

x 60. all from Bean 18190.

Glenmorgan, Dec 1984, Gordon 8887 (AD, BRI, CANB); 6

km SE of Cecil Plains, Feb 1995, Fensham 2012 (BRI);

Warrego Highway, c. 7 km W of Jackson, Jan 2000,

McDonald KRM262 (BRI); SE corner of Oakey Army

Airfield, Dec 2001, Menkins ILM76 (BRI); 4 km E of Cecil

Plains, Jan 2002, Franzmann 66 (BRI). New South Wales.

Northwest Slopes: Ashford, Mar 1908, Hayes s.n. (NSW).

Distribution and habitat: In Queensland,

Solanum stenopterum extends from Gayndah

to Moonie, and west to Glenmorgan and Yuleba

(Map 15), and there is an old collection from

Ashford in New South Wales. It inhabits

grassland, Belah forest or Eucalyptus populnea

woodland, on clayey soil.

Phenology: Flowers are recorded from October

to March; mature fruits recorded for March.

Notes: S. stenopterum differs from S. lacunarium

by the more sparsely distributed, longer and

more acicular prickles; smaller leaf lobing

index (sometimes entire); stellae very sparse

to moderate on lower leaf surface (vs. very dense

for S. lacunarium); stellae consistently with

3-5 lateral rays on upper leaf surface (vs. 7-10

lateral rays for S. lacunarium ); central ray

relatively long; inflorescences comprising 1 or

2 flowers and common peduncle absent (vs.

5-7 flowers, common peduncle 26-46 mm long

for S. lacunarium).

In S. stenopterum , the style extends

further beyond the anthers than in most other

species. It very rarely sets fruit (pers. obs.; I.

Menkins pers. comm.). Only 2 fruits are present

736

in the collections at BRI, and both of these are

insect damaged.

Conservation status: Currently listed as

“Vulnerable” under the Queensland Nature

Conservation Act 1992.

Solanum stenopterum is known from 5

locations, none of which is protected within a

conservation reserve. It is threatened by land

clearance, agricultural practices, weed

encroachment, and mowing of road reserves,

where most of the existing stands are located.

Applying the IUCN guidelines (IUCN. 2001),

a category of “Endangered” is recommended

(EN A3ce; B2ab(ii,iii,v)).

Etymology : From the Greek stenos meaning

‘narrow’ and pteron meaning ‘wing’. This is

in reference to the proximal extension of the

leaf lamina resulting in a winged petiole.

43. Solanum lacunarium F.Muell., Trans.

Philos. Soc. Victoria 1: 18 (1854). Type:

near the junction of the rivers Darling and

Murray, December 1853, F. Mueller

(lecto: MEL [MEL11745]), fide Symon

(1981).

Illustration : Cunningham et al. (1981: 593);

Symon (1981: 185).

Prostrate or sprawling, herbaceous resprouter,

0.2-0.4 m high. Juvenile stage absent. Adult

branchlets grey or brown; prickles 5-35 per

decimetre, straight, acicular or broad-based,

0.5-5 mm long, 5-9 times longer than wide;

stellae sparse, 0.25-0.3 mm diameter, sessile;

lateral rays 8-9, porrect; central ray absent;

finger hairs absent; Type 2 hairs absent. Adult

leaves elliptical or ovate, deeply lobed

throughout; lobes 3-5 on each side, obtuse,

lobing index 5-12; lamina 2.5-6.5 cm long,

1.3-3.5 cm wide, 1.8-2.2 times longer than

broad, apex obtuse, base cuneate or obtuse,

oblique part 0-2 mm long, obliqueness index

0-4 percent; petioles 0.9-3.2 cm long, 35-55%

length of lamina, winged, prickles present.

Upper leaf surface green; prickles 35-50,

straight, broad-based, 1-4 mm long, prickles

present on midvein and lateral veins; stellate

hairs distributed throughout; protostellae

absent; ordinary stellae very sparse, 0.4-1.3

mm apart, 0.2-0.3 mm across, sessile; lateral

rays 7-10, porrect; central ray 0-0.5 times as

Austrobaileya 6 (4): 639-816 (2004)

long as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Lower leaf surface

white; prickles 6-20, straight, broad-based,

prickles present on midvein only or present on

midvein and lateral veins; stellae very dense,

0.05-0.1 mm apart, 0.3-0.4 mm diameter,

sessile; lateral rays 8-9, porrect; central ray

0-0.4 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Inflorescence supra-axillary, pseudo-racemose,

common peduncle 26-46 mm long, rachis

prickles present; 5-7-flowered, weakly

andromonoecious, flowers 5-merous; pedicels

7-18 mm long at anthesis, same thickness

throughout, 0.4-0.8 mm thick at mid-point,

prickles absent or present. Calyx tube 1.5-2.5

mm long, lobes deltate, 1-2 mm long; prickles

present at anthesis, 15-25 per flower, 0.5-3 mm

long; stellae moderate to dense, white, 0.2-0.3

mm across, sessile, lateral rays 7 or 8, central

ray 0-0.4 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Corolla mauve or purple, 8-10 mm long, rotate,

inner surface glabrous; anthers 3.5-4.5 mm

long; ovary with stellate hairs only; functional

style 5-7.5 mm long, erect, with stellate and

Type 2 hairs, stellae 0.25-0.35 mm across,

lateral rays 4-8, central ray 0-0.4 times as long

as laterals. Fruiting calyx with lobes less than

half length of mature fruit, prickles 1-3 mm

long. Mature fruits 1-3 per inflorescence,

globular or ellipsoidal, 12-15 mm diameter;

pedicels 14-21 mm long in fruit, 0.9-1.1 mm

thick at mid-point. Lagoon Nightshade.

Specimens examined : Queensland. Maranoa District:

Dirranbandi, anno 2000, Christodoulou (BRI). New South

Wales. North Far Western Plains: Arrara-Lake Eliza, Oct

1912, Boorman s.n. (NSW); Gidgee Tank, Mount Wood,

Tibooburra, Oct 1949, Constable 10489 (NSW); floodplain

of Cuttaburra Channels, 35 km NE of Yantabulla, Nov 1979,

Paijmans 3282 (CANB).

Distribution and habitat : In Queensland,

Solanum lacunarium is known from a single

collection near Dirranbandi (Map 15). It occurs

widely in semi-arid parts of New South Wales,

Victoria and South Australia. The Queensland

record was from a grassy plain with Mitchell

Grass ( Astrebla spp.) and scattered Eucalyptus

coolabah.

Phenology: Flowers and fruits are recorded for

January and October.

Bean, Taxonomy of Solanum subg. Leptostemonum

Notes: The bright orange prickles borne on the

branchlets and both leaf surfaces of

S. lacunarium are distinctive.

Its habitat is similar to that of S. stenopterum,

but S. lacunarium occurs in areas of much lower

rainfall.

Conservation status : Not considered at risk in

an Australian context, although its occurrence

in Queensland is probably very limited.

Group 25B (S. pugiunculiferum group), here

defined; related to Group 25 ( S . hystrix

group) of Whalen (1984)

Large stems hollow; stellate and finger hairs

absent; prickles straight, broad-based; adult

leaves deeply lobed, obliqueness index 15-24%;

prickles present on both leaf surfaces; corolla

4-5 mm long; anthers 1.5-2.5 mm long; calyx

with 0-4 prickles; mesocarp dry in mature

fruits.

1 species endemic to Australia, and

indigenous to Queensland.

44. Solanum pugiunculiferum C.T.White,

Proc. Roy. Soc. Queensland 53: 225

(1942). Type: Queensland. Burke

District: Settlement Creek, November

1922, L.J. Brass 244 (holo: BRI).

Illustration : Symon (1981: 100)

Prostrate or sprawling, herbaceous resprouter,

0.2-0.6 m high. Juvenile stage absent. Adult

branchlets grey, yellow or brown; prickles

10-200 per decimetre, straight, acicular or

broad-based, 2-16 mm long, 6-8 times longer

than wide; stellae absent; finger hairs absent;

Type 2 hairs absent, or sparse. Adult leaves

elliptical or ovate, deeply lobed throughout;

lobes 2 or 3 on each side, acute, lobing index

2.3-10; lamina 1.6-4.5 cm long, 1.1-2.8 cm

wide, 1.4-1.7 t im es longer than broad, apex

acute, base obtuse or cordate, oblique part

0.5-11 mm long, obliqueness index 2-24

percent; petioles 0.5-1 cm long, 20-30% length

of lamina, prickles absent or present. Upper

leaf surface green or grey-green; prickles

3-10, straight, broad-based, 1-17 mm long,

prickles present on midvein only or present on

midvein and lateral veins; stellate hairs absent;

finger hairs absent; Type 2 hairs present only

737

in vein depressions. Lower leaf surface green;

prickles 2-8, straight, broad-based, prickles

present on midvein only or present on midvein

and lateral veins; stellae absent; finger hairs

absent; Type 2 hairs absent. Inflorescence

supra-axillary, pseudo-racemose, common

peduncle 2-4 mm long, rachis prickles present;

2-5-flowered, with all flowers bisexual and

5-merous; pedicels 3-5 mm long at anthesis,

same thickness throughout, 0.4-0.6 mm thick

at mid-point, prickles absent or present. Calyx

tube 1-1.5 mm long, lobes deltate, 0.5-1 mm

long; prickles present at anthesis, 1-4 per

flower, 2-6 mm long; stellae absent; finger

hairs absent; Type 2 hairs absent. Corolla

mauve, 4-5 mm long, rotate, inner surface

glabrous; anthers 1.5-2.5 mm long; ovary

glabrous; functional style 2.5-3.5 mm long,

erect, glabrous. Fruiting calyx with lobes less

than half length of mature fruit, prickles

present, 2.5-9 mm long. Mature fruits 1-4 per

inflorescence, globular, 8-11 mm diameter,

brown or yellow; mesocarp dry; pedicels 5-8

mm long in fruit, 0.7-1 mm thick at mid-point;

seeds pale yellow, 3-3.8 mm long.

Specimens examined : Northern Territory. Red Lily

Lagoon, 8 miles [13 km] E of ‘Elsey’ HS, Oct 1958,

Chippendale 5064 (BRI); 3 km E of ‘Bing Bong’ HS, Sep

1985, Latz 10243 (BRI). Queensland. Burke District:

Burketown, May 1919, Higgins (BRI); Normanton, May

1935, Blake 8989 (BRI); Karumba, Jun 1966, Pedley 2103

(BRI); 6 miles [10 km] SE of Burketown, Jun 1967, Symon

4998 (BRI); 1.6 km SW of Burketown, Jul 1974,

Ollerenshaw & Kratzing 1373 (BRI, CANB); Denham

Island, north end, just W of Momington Island, Sep 1981,

Fosberg 62034 (CANB); 32 km NNE of ‘Inverleigh’ HS,

duckhole, Jul 1987, Dalliston HC137 (BRI); Karumba, Sep

1998, Gunther MG107 (BRI). Cook District: Mapoon, N

of Weipa, Sep 1980, Godwin A55 (BRI); c. 7 km N of

‘Inkerman’ HS, Jun 1990, NeIdner 2917 & Clarkson (AD,

BRI, NSW, QRS); road from Kowanyama to ‘Topsy’, c. 16

km from ‘Topsy’, Oct 1999, Thomas & Lansdown (BRI).

Distribution and habitat : In Queensland,

Solanum pugiunculiferum is found around the

Gulf of Carpentaria (Map 13), but it also occurs

in Northern Territory and the Kimberley of

Western Australia. It grows near the coast on

heavy clay soils, sometimes in areas that are

periodically flooded by very high tides. In the

latter situation, it is associated with Sporobolus

virginicus grassland.

Phenology: Flowers recorded from May to

September; mature fruits from May to October,

plus a single record in January.

738

Notes: Solarium pugiunculiferum is a highly

distinctive species, worthy of the monotypic

sectional status accorded it by Symon (1981).

It is totally without stellate hairs or finger hairs,

it has hollow stems, very oblique leaf bases,

tiny flowers (corolla 4-5 mm long) with

disproportionately long filaments, and fruits

that are quite dry at maturity. The often saline

habitat is also very unusual for a Solarium.

Solarium oligandrum Symon, recently

described from tropical Western Australia

(Symon 2001) does not appear to be particularly

closely related to S. pugiunculiferum.

White (1942) cited two collections in the

protologue, without any indication of a type.

However, on the Brass 244 sheet, he has written

“Solanum pugiunculiferum C.T.White, Type ”.

Hence the choice of a lectotype is unnecessary.

Conservation status: Moderately widespread.

Not considered at risk.

Group 27 (S. ellipticum group) of Whalen

(1984)

Calyx prickly, not accrescent in fruit (100%);

corolla shallowly to deeply lobed (100%);

inflorescences andromonoecious, male flowers

and bisexual flowers same size and prickliness

(100%); branchlet stellae dense to very dense

(100%); fruits green to yellowish-green,

mesocarp moist but not succulent (100%);

branchlets prickly (87%); adult leaves entire

(87%); upper leaf suface with dense to very

dense stellate hairs (71%).

10 species endemic to Australia; 8 species

indigenous to Queensland.

45. 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 (lecto: BM; isolecto: MPU).

Solanum ellipticum var. chillagoense Domin,

Biblioth. Bot. 89: 588 (1929). T>pe:

Queensland. Cook District: near

Austrobaileya 6 (4): 639-816 (2004)

Chillagoe, February 1910, K. Domin (syn:

PR [2 collections], photos at BRI).

Solanum ellipticum f. albiflora Domin,

Biblioth. Bot. 89: 588 (1929). Type:

Queensland. Cook District: near

Chillagoe, February 1910, K. Domin

(holo: PR?).

Solanum cleistogamum Symon, Trans. &

Proc. Roy. Soc. South Australia 95: 227

(1971), syn. nov. Type: Western

Australia. 20 miles [32 km] north of

Onslow, 1 July 1967, D.E. Symon 5418

(holo: PERTH; iso: AD, CANB, K, L).

Solanum sp. (Newcastle Range D.E. Symon

4907) in Henderson (2002).

[S. dianthophorum auct., non Dunal]

Illustration: Cunningham et al. (1981: 592);

Symon (1981: 192), as S. dianthophorum.

Prostrate, herbaceous resprouter or rhizomatous

perennial shrub, 0.1-0.3 m high. Juvenile stage

absent. Adult branchlets white to mauve or

brown; prickles 10-250 per decimetre, straight,

acicular, 1-8 mm long, 7-15 t im es longer than

wide; stellae dense to very dense, 0.5-0.8 mm

diameter, stalks 0-0.3 mm long; lateral rays

7-12, porrect or ascending; central ray 0.5-1

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Adult

leaves elliptical or ovate, entire; lamina 3.5-14

cm long, 1.5-5.2 cm wide, 1.7-2.9 t im es longer

than broad, apex obtuse or acute, base cuneate

or obtuse or cordate, oblique part 0-9 mm long,

obliqueness index 0-6 percent; petioles 0.8^1.6

cm long, 20-65% length of lamina, prickles

present. Upper leaf surface grey-green or grey;

prickles 2-20, straight, acicular, 1-7 mm long,

prickles present on midvein only or present on

midvein and lateral veins; stellate hairs

distributed throughout; protostellae absent;

ordinary stellae density moderate to very dense,

0.1-0.5 mm apart, 0.4-0.7 mm across, stalks

0-0.3 mm long; lateral rays 7 or 8, porrect;

central ray 0.5-1 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Lower leaf surface white or yellowish;

prickles 0-40, straight, acicular, absent or

present on midvein only or present on midvein

and lateral veins; stellae dense to very dense,

0.05-0.3 mm apart, 0.5-0.8 mm diameter,

Bean, Taxonomy of Solanum subg. Leptostemonum

stalks 0-0.4 mm long; lateral rays 7 or 8,

porrect; central ray 0.5-1 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Inflorescence supra-

axillary, solitary or pseudo-racemose, common

peduncle 0—40 mm long, rachis prickles present;

1- 9-flowered, weakly andromonoecious or with

all flowers bisexual, flowers 5-merous; pedicels

4-16 mm long at anthesis, same thickness

throughout, 0.7-0.9 mm thick at mid-point,

prickles absent or present. Calyx tube 1.5-3

mm long, lobes deltate or attenuate, 2-10 mm

long; prickles present at anthesis, 5-50 per

flower, 1-5 mm long; stellae very dense, white

to brown, 0.4-1 mm across, stalks CM).4 mm long,

lateral rays 7 or 8, central ray 0.8-1 times as

long as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Corolla white,

mauve or purple, 5-12 mm long, shallowly or

deeply lobed, inner surface glabrous or sparsely

stellate-hairy; anthers 3-6 mm long; ovary

glabrous, or with Type 2 hairs only; functional

style 5.5-10 mm long, erect or sigmoid,

glabrous or with Type 2 hairs only. Fruiting

calyx with lobes less than or more than half

length of mature fruit, prickles 1-5 mm long.

Mature fruits 1-5 per inflorescence, globular,

13-17 mm diameter, yellowish-green or green,

2- or 4-locular; placenta in cross-section

stalked, anvil-shaped; mesocarp moist but not

juicy; exocarp 0.6-2.1 mm thick; pedicels 7-19

mm long in fruit, 0.8-2.1 mm thick at mid¬

point; seeds pale yellow, 2.2-2.7 mm long.

Selected specimens : Queensland. Cook District:

Guguyalangi Gallery, 13 km S of Laura on the Peninsula

Development road, Jun 1981, Clarkson 3669 (AD, BRI,

PERTH, QRS); Royal Arch cave, Chillagoe N.P., Mar 2000,

McDonald KRM337 (BRI). North Kennedy District: Mt

Remarkable, near Pentland, Jun 1934, Blake 6124 (BRI); S

side of Red Falls road, 42 km NW of Charters Towers, Jul

1989, Jobson 672 (BRI, MEL). Gregory North District:

‘Kamaran Downs’, 10 km NNW of Bedourie, Jun 1995,

Edmunds 23 (BRI). Mitchell District: 61 km E of

Barcaldine on road to Jericho, Oct 1993, Lepschi & Slee 1173

(AD, BRI, CANB); 4 km SE of ‘Lake Dunn’ HS, NE of

Aramac, Feb 1994, Bean 7504 & Forster (AD, BRI, DNA).

South Kennedy District: 13 miles [21 km] SW of ‘Alpha’

station, Oct 1964, Adams 1339 (BRI, CANB); 11 km NW

of Belyando Crossing on edge of Gregory Development road,

Jun 1992, Thompson BUC818 & Sharpe (AD, BRI).

Leichhardt District: ‘Pamaroo’, 25 miles [40 km] SE of

Injune, Apr 1961, Johnson 2129 (BRI); ‘Wandobah’, c. 1

km NE of Dingo, Jul 1988, Anderson 4492 (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). Gregory

739

South District: 10 miles [16 km] W of Betoota, Jul 1936,

Blake 12184 (BRI). Warrego District: c. 11 km SW of

Thargomindah, on road to Hungerford, Sep 1973, Henderson

H2061 & Boyland (AD, BRI); 0.4 km W of Gee Gee Gap,

Mt Moffatt N.P., Dec 1997, Bean 12905 (BRI). Maranoa

District: c. 41 km along Shirlo road, NW of Bollon, Mar

2001, Bean 17543 (BRI). Burnett District: ‘Neaavie’, NE

of homestead towards Barambah Creek, Aug 1996,

Grimshaw PG2536 & Turpin (BRI); c. 6 km NE of Allies

Creek and 7.5 km SW of ‘Weir Weir’, Jul 1998, Pollock

ABP617 & Dean (BRI). Wide Bay District: Bingera, Dec

1896, J.F. Bailey (BRI). 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); 2 km ESE of Laidley, Jun 2002, Bean 19062

(BRI). New South Wales. North West Plains: 4 km SSW

of ‘The Glen’ HS, c. 70 km W of Moree, Jan 1999, Wannan

1036 et al. (AD, BRI, NSW). North Far Western Plains:

Urisino-Yamba station road, 30 miles [48 km] W of

Wanaaring, Oct 1963, Constable 4583 (BRI, NSW); Golden

Gully mining site, adjacent to Deadhorse Gully camping area,

1 kmN of Tibooburra, Sturt N.P., Sep 1989, Coveny 13598

et al. (AD, BRI, MO, NSW).

Distribution and habitat : Solanum ellipticum

occurs throughout Queensland, except for the

north-west, Cape York Peninsula and much of

the east coast (Map 18). It inhabits eucalypt

woodlands, Lancewood communities, semi¬

evergreen vine thicket and Mulga woodlands.

Soils vary greatly from sands to heavy clays.

Phenology : Flowers and fruits are recorded for

every month of the year.

Notes: On the date of collection of the type of

Solanum ellipticum , Robert Brown was

botanising near the mouth of the Styx River,

ESE of St Lawrence (Vallance et al. 2001).

S. ellipticum is probably the most

taxonomically difficult species in Australia. It

is common and widespread, but also highly

variable. There are some (ill-defined)

geographical variants, while plants growing

side by side in natural habitat can be quite

different in morphology e.g. prickliness, leaf

colour, flower colour, indumentum density.

Symon (1981) highlighted its taxonomic

difficulties, and my own efforts to classify its

variation have largely failed.

Specimens of S. cleistogamum (including

the holotype) borrowed from PERTH are a very

good match for material found in eastern

Queensland, where the type of S. ellipticum was

collected, and I do not think it is possible to

740

Austrobaileya 6 (4): 639-816 (2004)

maintain the former as a distinct taxon. The

Western Australian specimens do generally

have longer, consistently glabrous prickles, long

petioles and leaves with an obtuse apex, but

there is no qualitative difference. The perceived

cleistogamous habit was apparently a major factor

in Symon’s decision to name S. cleistogamum.

However, the occurrence of cleistogamy does

not seem sufficient to warrant the recognition

of a separate species. It is very difficult to assess

the degree of cleistogamy in any specimen in

the herbarium, nor is it easy in the field. Perhaps

cleistogamy occurs throughout the range of

S. ellipticum, and indeed I have observed it on

occasion for S. ellipticum in Queensland. The

other key morphological characters given by

Symon (1981: 34) for S. cleistogamum (stems

sprawling; corolla 1-1.5 cm diameter; mature

berry pale yellow-green or slightly purplish)

apply equally well to S. ellipticum from around

its type locality. The alternative statement

(which eventually leads to S. ellipticum in the

key) starts with “stems erect”, but S. ellipticum

is always prostrate or procumbent.

Conservation status: Widespread. Not

considered at risk.

46. Solanum dianthophorum Dunal, Hist. Nat.

Solanum 183 (1813), nom. nov.; S.

biflorum R.Br., Prodr. 445 (1810), nom.

illeg., non Lour. (1790). Type:

[Queensland. Port Curtis District:] “Port

II”, undated [Port Clinton, 21-23 August

1802], R. Brown (holo: BM [Bennett No.

2668]).

Prostrate? shrub. Juvenile stage unknown.

Adult branchlets rusty or brown; prickles

absent; stellae very dense, 0.4-0.5 mm

diameter, stalks 0-0.1 mm long; lateral rays

8-13, porrect; central ray 0.5-1 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Adult leaves ovate,

entire; lamina 1.5-3.5 cm long, 0.9-1.8 cm

wide, 1.9-2.4 times longer than broad, apex

obtuse or acute, base obtuse or cordate, oblique

part 0-1.5 mm long, obliqueness index 0-4

percent; petioles 0.4-0.85 cm long, 20-25%

length of lamina, prickles absent. Upper leaf

surface grey-green; prickles 0-2, straight,

acicular, 0.5-1.5 mm long, prickles absent or

present on midvein only; stellate hairs

distributed throughout; protostellae absent;

ordinary stellae dense, 0.1-0.2 mm apart, 0.3-0.4

mm across, sessile; lateral rays 8, porrect;

central ray 0.5-1 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Lower leaf surface yellowish; prickles

absent; stellae very dense, 0.05-0.15 mm apart,

0.3-0.4 mm diameter, stalks 0-0.1 mm long;

lateral rays 8, porrect; central ray 0.5-1 times

as long as laterals, not gland-tipped; finger

hairs absent; Type 2 hairs absent. Inflorescence

supra-axillary, pseudo-racemose, common

peduncle absent, rachis prickles absent; 2-4-

flowered, flowers 5-merous; pedicels 4-5 mm

long at anthesis, same thickness throughout,

c. 0.5 mm thick at mid-point, prickles absent.

Calyx tube c. 1.5 mm long, lobes attenuate, c.

3 mm long; prickles present at anthesis, 1-4

per flower, 0.5 mm long; stellae very dense,

brown or rusty, 0.3-0.45 mm across, stalks

0-0.1 mm long, lateral rays 8, central ray 0.6-1

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Corolla

c. 7 mm long, deeply lobed, inner surface

glabrous; anthers c. 3.9 mm long. Fruiting

calyx with lobes less than half length of mature

fruit, prickles present, 0.5-1 mm long. Mature

fruits 1 per inflorescence, globular, 9-10 mm

diameter (maturity unknown); pedicels 7-12

mm long in fruit, 0.9-1.2 mm thick at mid¬

point. Fig. 29.

Specimens examined: known only from the type.

Distribution and habitat: The type collection

of Solanum dianthophorum was from Port

Clinton, now part of the Shoalwater Bay

Military Reserve, N of Rockhampton (Map 16).

Brown, on the specimen label, recorded the

habitat as “ arenosus prope littus ” (sandy area

near beach).

Phenology: Brown’s collection, made in the

month of August, bears flowers and fruits, but

the state of maturity of the fruits is unknown.

Notes: The type of S. biflorum R.Br. (on which

S. dianthophorum is based) differs from the

hundreds of available S. ellipticum specimens

in several respects. These differences are: the

smaller stellae on the branchlets, upper leaf

surface and lower leaf surface; the leaves only

1.5-3.5 cm long (3.5-14 cm for S. ellipticum );

the lack of prickles on the branchlets and

petioles; the shorter petioles; and the calyx less

Bean, Taxonomy of Solanum subg. Leptostemonum

741

Fig. 29. Holotype of Solanum dianthophorum.

742

Austrobaileya 6 (4): 639-816 (2004)

than half the fruit length and with 1-4 tiny

prickles (calyx > half fruit length, with 5-50

stout prickles for S. ellipticum).

No other collections are known which

resemble this specimen. Bentham (1868) cited

two other specimens for S. dianthophorum, i.e.

“Bay of Inlets” Banks & Solander, and “Percy

Islands” A. Cunningham, but I have not seen

these.

Either S. dianthophorum is a very

restricted endemic, or the type represents an

extreme form of S. ellipticum. A field survey

around Port Clinton should clarify the matter.

Conservation status : Data deficient.

47. Solanum crebrispinum A.R.Bean sp. nov.

Frutex perennis erectus, ramulis teretibus

aculeis 180-420 per decimetrum; folia

ovata, integra; pagina superiore aculeis

praedita et dense stellato-pilosa; stellae

0.8-1.2 mm diametro, pedicellis 0.1-0.9

mm longis; pagina inferior folii

densissime stellato-pilosa, floccosiuscula;

calyx aculeis 50-70 validissimis

praeditus, stellis proximale affixis; lobi

calycis attenuati; stylus functionalis

tantum pilis Type-2 praeditus. Typus:

Queensland. South Kennedy District: 23

km SE of ‘Teamass’ Homestead, 10 June

1992, E.J. Thompson BUC457 & P.R.

Sharpe (holo: BRI; iso: AD).

Erect, rhizomatous perennial shrub, 0.3-0.5 m

high. Juvenile stage unknown. Adult branchlets

yellow or rusty; prickles 180-420 per

decimetre, straight, acicular, 2-11 mm long,

11-15 times longer than wide; stellae very

dense, 0.9-1.2 mm diameter, stalks 0-1 mm

long; lateral rays 6-8, porrect or ascending;

central ray 1-1.7 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Adult leaves ovate or broadly ovate,

entire; lamina 5-9.5 cm long, 2.8-5.5 cm wide,

1.6-1.8 times longer than broad, apex obtuse,

acute or acuminate, base obtuse or cordate,

oblique part 0-3 mm long, obliqueness index

0-6 percent; petioles 1.1-4.4 cm long, 20-45%

length of lamina, prickles present. Upper leaf

surface grey-green; prickles 3-35, straight,

acicular, 1-5 mm long, prickles present on

midvein only or present on midvein and lateral

veins; stellate hairs distributed throughout;

protostellae present; ordinary stellae dense,

0.2-0.4 mm apart, 0.8-1.2 mm across, stalks

0.1-0.9 mm long; lateral rays 7 or 8, porrect or

ascending; central ray 1-1.7 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Lower leaf surface

yellowish; prickles 5-20, straight, acicular,

present on midvein only or present on midvein

and lateral veins; stellae dense to very dense,

0.05-0.25 mm apart, 0.7-1.2 mm diameter,

stalks 0-1 mm long; lateral rays 6-8, porrect

or ascending; central ray 1-1.7 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Inflorescence

supra-axillary, pseudo-racemose, common

peduncle 0-25 mm long, rachis prickles

present; 3-5-flowered, flowers 5-merous;

pedicels 4-6 mm long at anthesis, same

thickness throughout, 0.9-1.2 mm thick at mid¬

point, prickles present. Calyx tube 1.5-2 mm

long, lobes attenuate, 3-5.5 mm long; prickles

present at anthesis, 50-70 per flower, 2-6 mm

long; stellae very dense, yellow or white, 0.7-0.9

mm across, stalks 0-0.7 mm long, lateral rays

7 or 8, central ray 1-1.7 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Corolla purple, 8-10 mm

long, shallowly lobed, inner surface glabrous;

anthers c. 3.8 mm long; ovary with Type 2 hairs

only; functional style c. 5.8 mm long, erect,

with Type 2 hairs only. Fruiting calyx with

lobes more than half length of mature fruit,

prickles 3-6 mm long. Mature fruits 1 or 2 per

inflorescence, globular, 14-16 mm diameter,

yellowish-green or green; mesocarp moist but

not juicy; pedicels 8-9 mm long in fruit,

1.5-1.8 mm thick at mid-point. Fig. 30.

Specimens examined : Queensland. South Kennedy

District: on crest of Great Dividing Range, c. 25 km NNW

of ‘Yarrowmere’ HS, c. 92 km SSE of Pentland, Oct 1983,

Henderson H2821 et al. (BRI); on Llanarth Back Range

road, 9.4 km S of junction with Scartwater road, May 1991,

Neldner 3525A & Thompson (BRI); 4 km N of

‘Moonoomoo’ in headwaters of Carmichael Creek, Apr 1992,

Thompson BUC247 & Simon (BRI); 10 km W of ‘St Anns’

HS, Jun 1992, Thompson BUC454 & Sharpe (BRI).

Distribution and habitat : Solanum

crebrispinum is endemic to Queensland.

Apparently restricted to the Lake Buchanan

area south of Charters Towers (Map 17). It

grows on sandstone ridges or “jump-ups” with

Eucalyptus similis, E. leichhardtii or Acacia

Bean, Taxonomy of Solanum subg. Leptostemonum

743

hehmiih <hi>

FlWa i,i Qut*r 4 l*id SfajUl KAfiMdj-

SoJanm a) i fp<t i-cut 1 R. Hr.

Coll, e.J.Thc^tcn BOWS? 10 AJW s m

*P'R, 5Kin»

2 rws i«c* 34 'e ah, zso ■.

Ow*h ■-

U fc=- SE Twroass NmtHd (i«U W6-12H J&S

21*13*25-, WJt'H”}

Woodland of Eucalyptus brawftll. Occasional s*all *«ct

shA* with c*.rpl# Flewor* l,5c=a new*-,

Cteli*iert»l Mil OPOCt Shrub with prvpl*

tow,

QUEENSLAND herbar<‘j?,i ^-1)

Bfisbaruo AijiL d

>Q 544033

Queensland Herbarium (BRI)

SoWvw f. IAmiH.

Dm. fcum. D«,d«V4 De

Hoi-oTfPg Queensland Herbarium {Bftij

e of

Da?p ?&CT* ajTffc g

Fig. 30. Holotype of Solanum crebrispinum.

744

shirleyi. One specimen records Eucalyptus

brownii as the dominant tree.

Phenology: Poorly known. Flowers are

recorded for April, June and October; fruits are

recorded for April and October.

Notes: S. crebrispinum is related to S. ellipticum,

but differs by the erect habit, the branchlet

stellae 0.9-1.2 mm across (0.5-0.8 mm for S.

ellipticum ) on stalks up to 1.0 mm long (up to

0.3 mm long for S. ellipticum), the presence of

protostellae on the upper leaf surface, normal

stellae 0.8-1.2 mm across (0.4-0.7 mm for

S. ellipticum ), on very thick stalks 0.1-0.9 mm

long (thin stalks 0-0.3 mm long for S. ellipticum),

and the 50-70 calyx prickles (5-50 for

S. ellipticum). On the branchlets, upper and

lower leaf surface, and on the calyx, the central

ray of the stellae is 1-1.7 times as long as the laterals;

this compares with 0.5-1 times for S. ellipticum.

Conservation status: Collection density in this

area is low. Not considered at risk.

Etymology: From the Latin crebra - abundant,

and spina - spines or prickles. This refers to

the abundant prickles on the branchlets and

calyx.

48. Solarium senticosum A.R.Bean sp. nov.

Frutex perennis erectus; ramuli teretes

brunnei vel ferruginei, aculei 180-400 per

decimetrum; folia ovata, integra, ad basim

cuneata; pagina inferior folii ferruginea,

stellis radio centrali 0.6-1.2plo lateralibus

longiore; inflorescentia 1-3-flora; stylus

functionalis 7.5-8.5 mm longus, glaber;

calyx aculeis 40-65 validissimis et lobi

attenuati praeditus. Typus: Queensland.

Burke District: 4.5 miles [7 km]

southwest of Mount Isa, 5 July 1974, P.

Ollerenshaw 1220 & D. Kratzing (holo:

BRI; iso: AD, n.v., CANB).

Solanum ellipticum var. horridum Domin,

Biblioth. Bot. 89: 588 (1929). type:

Queensland. Burke District: around

Cloncurry, February 1910, K. Domin

(holo: PR, photo at BRI).

Erect, rhizomatous perennial shrub, 0.3-0.6 m

high. Juvenile stage unknown. Adult branchlets

rusty or brown; prickles 180-400 per decimetre,

straight, acicular, 1.5-6 mm long, 11-18 times

longer than wide; stellae very dense, 0.6-0.9

Austrobaileya 6 (4): 639-816 (2004)

mm diameter, stalks 0-0.4 mm long; lateral

rays 7 or 8, porrect or ascending; central ray

0.6-1.2 t im es as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Adult leaves ovate, entire; lamina 3.5-9 cm

long, 1.2-3.4 cm wide, 1.9-2.7 t im es longer

than broad, apex acute, base cuneate, oblique

part 0-5 mm long, obliqueness index 0-9

percent; petioles 0.7-1.7 cm long, 19-30%

length of lamina, prickles present. Upper leaf

surface grey; prickles 0-6, straight, acicular,

1-4 mm long, prickles absent or present on

midvein only; stellate hairs distributed

throughout; protostellae absent; ordinary stellae

dense, 0.1-0.25 mm apart, 0.6-1 mm across,

stalks 0-0.25 mm long; lateral rays 7 or 8,

porrect; central ray 0.8-1.2 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Lower leaf surface rusty;

prickles 0-10, straight, acicular, absent or

present on midvein only or present on midvein

and lateral veins; stellae dense to very dense,

0.1-0.2 mm apart, 0.7-1 mm diameter, stalks

0-0.6 mm long; lateral rays 7 or 8, porrect;

central ray 0.6-1.2 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Inflorescence supra-axillary,

solitary or pseudo-racemose, common peduncle

0-2 mm long, rachis prickles present; 1-3-

flowered, flowers 5-merous; pedicels 5-11 mm

long at anthesis, same thickness throughout,

0.9-1.2 mm thick at mid-point, prickles absent

or present. Calyx tube 2-3.5 mm long, lobes

attenuate, 4-9 mm long; prickles present at

anthesis, 40-65 per flower, 1-4 mm long;

stellae very dense, brown or rusty, 0.4-0.6 mm

across, stalks 0-0.2 mm long, lateral rays 6-8,

central ray 0.6-1.2 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Corolla purple, 7-11 mm long,

shallowly or deeply lobed, inner surface

glabrous; anthers 4-4.5 mm long; ovary

glabrous; functional style 7.5-8.5 mm long,

erect, glabrous. Fruiting calyx with lobes more

than half length of mature fruit, prickles 2-5

mm long. Mature fruits 1 or 2 per inflorescence,

globular, 12-14 mm diameter, green; mesocarp

moist but not juicy; pedicels 6-14 mm long in

fruit, 1.2-1.5 mm thick at mid-point. Fig. 31.

Specimens examined : Queensland. Burke District: Mtlsa,

Apr 1935, Blake 8780 (BRI); 3 km from kiosk road, near

Lake Moondarra, NW of Mt Isa, Aug 1973, Swan 110 (BRI);

60 km E of Mount Isa on the Mount Isa-Cloncurry road,

Aug 1973, Swan 116 (BRI); hill with radio repeater 4141 on

Bean, Taxonomy of Solanum subg. Leptostemonum

745

ChMssfusiand Hwtaoum iBRIf

109124

t- herbarium, cjwherea botanic qabdews

n>. oSV/rp-

JOlaBila qua.lrilQ'CrLil t i*ij?: 'f . k Kiwll,

July 1974

D*t ftA. oa^t"] 00 Lh. 4i*Il«a (7ka.J S.W. of Haunt Ian, ^d*

Queensland HeeOarium (SRH

h&U'Ty?? ,f

iiuaty aj]»iinar;ce.

r iil.TfcHS'. AfcC

□»««

f Brisbane -

Fig. 31. Holotype of Solanum senticosum.

746

Barkly Highway 24 km NNW of Mount Isa, May 1974, Kanis

1701 (BRI, CANB, K); Mount Isa, May 1952, Morris 143

(BRI); Mount Isa, Oct 1974, Specht 75 & Rogers (BRI);

north end of dam, 4 km SW of Mt Isa, Aug 1986, Harris 66

(BRI); Millican Creek, track to Telstra tower, Barkly Hwy,

Apr 1997, Forster PIF20821 & Holland (BRI); S0 2 study

site MR6,8.27 km to Mt Isa copper stack at 339°, Apr 1998,

Fell DGF5235 & Barrs (BRI); ‘Riversleigh’, Campbells

Camp on the Gregory River, Jul 1998, Symon 15794 (AD,

BRI). Gregory North District: ‘Elderslie’, Winton, Sep

1934, Kennedy 21 (BRI); Duchess, May 1936, Blake 11528

(BRI); track to Mistake Hut, Bladensberg N.R, S of Winton,

Mar 1998, Forster PIF22230 & Booth (AD, BRI, MEL).

Mitchell District: ‘Springdale’, N of Aramac, Jul 2000,

Fensham 3946 (BRI).

Distribution and habitat : Solanum senticosum

is endemic to Queensland. Most collections are

from the Mount Isa area, but it extends to

Winton and Aramac (Map 17). It is most often

recorded from rocky ridges dominated by

Eucalyptus leucophloia subsp. euroa with

Triodia understorey.

Phenology: Flowers are recorded from March

to October; fruits from March to August.

Notes: S. senticosum is close to S. ellipticum,

but differs by the erect shrubby habit; the rusty

indumentum on the stems, leaves and calyces;

the more densely prickly branchlets; the

consistently cuneate leaf bases; the larger stellae

on the upper leaf surface; and the calyx with

many stout prickles in longitudinal rows.

The Symon collection from Riversleigh

is atypical as it lacks the rusty colouration of

the stems and leaves.

Conservation status: Widespread. Not

considered at risk.

Etymology: from the Latin senticosus - full of

thorns or prickles.

49. Solanum quadriloculatum F.Muell.,

Fragm. 2: 161 (1861). Type:

[Queensland. Burke District:] Gulf of

Carpentaria, Nicholson River, 21-24

August 1856, F. Mueller (lecto: MEL

[MEL 11735]),//Jc Symon (1981).

Illustration: Symon (1981: 211).

Sprawling, herbaceous resprouter or

rhizomatous perennial shrub, 0.1-0.3 m high.

Juvenile stage unknown. Adult branchlets

ridged, yellow or rusty; prickles 240-700 per

Austrobaileya 6 (4): 639-816 (2004)

decimetre, straight, acicular, 2-8 mm long,

11-20 times longer than wide; stellae very

dense, 0.5-1 mm diameter, stalks 0-0.8 mm

long; lateral rays 7 or 8, ascending; central ray

0.8-1.2 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Adult leaves ovate, entire; lamina 5.5-14.5 cm

long, 2.5-7 cm wide, 1.7-2.3 times longer than

broad, apex obtuse or acute, base cuneate or

obtuse, oblique part 0-6 mm long, obliqueness

index 0-5 percent; petioles 0.8-4.4 cm long,

15-60% length of lamina, prickles present.

Upper leaf surface grey; prickles 1-10, straight,

acicular, 2-5 mm long, prickles present on

midvein only or present on midvein and lateral

veins; stellate hairs distributed throughout;

protostellae absent; ordinary stellae dense to

very dense, 0.05-0.2 mm apart, 0.6-1.2 mm

across, sta lks 0-1 mm long; lateral rays 7 or 8,

porrect or ascending; central ray 0.8-1.2 times

as long as laterals, not gland-tipped; finger

hairs absent; Type 2 hairs absent. Lower leaf

surface white or yellowish; prickles 3-20,

straight, acicular, present on midvein only or

present on midvein and lateral veins; stellae

very dense, 0.05-0.1 mm apart, 0.7-1.2 mm

diameter, stalks 0-1.6 mm long; lateral rays 7

or 8, porrect or ascending; central ray 0.8-1.2

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent.

Inflorescence supra-axillary, pseudo-racemose,

common peduncle 23-60 mm long, rachis

prickles absent; 9-15-flowered, with all flowers

bisexual and 5-merous; pedicels 4-11 mm long

at anthesis, same thickness throughout, 1-1.2

mm thick at mid-point, prickles absent. Calyx

tube 2-4 mm long, lobes deltate or attenuate,

1.5-4.5 mm long; prickles present at anthesis,

5-40 per flower, 1-3 mm long; stellae very

dense, yellow or white, 0.5-0.9 mm across,

stalks 0-1.2 mm long, lateral rays 7 or 8, central

ray 0.8-1.2 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Corolla mauve or purple, 9-15 mm long,

shallowly or deeply lobed, inner surface

glabrous; anthers 5.5-6.5 mm long; ovary

glabrous; functional style 9-10.5 mm long,

erect, glabrous. Fruiting calyx with lobes less

than or more than half length of mature fruit,

prickles 1-3 mm long. Mature fruits 1-6 per

inflorescence, globular, 14-17 mm diameter,

yellowish-green, 4-locular; pedicels 11-17 mm

long in fruit, 1.1-1.9 mm thick at mid-point;

seeds pale yellow, 2.3-2.9 mm long.

Bean, Taxonomy of Solarium subg. Leptostemonum

Specimens examined: Queensland. Burke District:

Cloncurry,Nov 1935, Blake 10109 (BRI); ‘Barkly Downs’,

c. 50 miles [80 km] SE of Camooweal, Dec 1947, Everist

3381 (BRI); Dugald River, Apr 1962, Cole 128 & Provan

(BRI); c. 24kmW of Cloncurry, on road to Mt Isa, Apr 1973,

Henderson H1854 (BRI); 5 km SW of Mt Isa, Apr 1976,

Farrell TF369 (BRI); 105 km N of Cloncurry, Apr 1993,

Milson JM357 (BRI); 85 km from Cloncurry on the

Normanton highway near ‘Coolulla’ station, Jul 1993, Alcock

11283 (AD, BRI); Riversleigh road, 22 km ESE of

‘Riversleigh’ HS towards Burketown-Camooweal road, Aug

1993, Lolly 99 (BRI, CANB); 34 km NW of McKinlay, Apr

1996, Milson JM1011 (BRI); 15 km from Mt Isa on Duchess

road, May 1997, Forster PEF21180 & Booth (BRI, DNA);

Camooweal Caves N.R, Aug 1999, Fox IDF707 & Middleton

(BRI); between Cloncurry and Mt Isa on Barkly Highway,

Apr 2001, McDonald KRM826 (AD, BRI). Gregory North

District: Georgina River, May 1933, Whitehouse (BRI);

between Glenormiston and Toko Range, Feb 1935 , Boyle s.n.

(BRI); Duchess-Mt Isa road, May 1963, Gittins 735 (BRI);

QT Bore, 29 km SE of ‘Kollala’, May 1985, Neldner 1963

& Stanley (BRI); 3 km N of ‘Bushy Park’ HS, 21 km NW of

Duchess, May 1985, Neldner 2113 & Stanley (BRI); Trough

Tank, Placer Pacific Osborne exploration lease, 30 km N of

Pathungra, May 1993, Gunness AG2188 (BRI).

Distribution and habitat: In Queensland,

Solanum quadriloculatum is confined to the

north-west, from Lawn Hill N.P. to Duchess,

and east to Cloncurry (Map 19). It also occurs

in Northern Territory and the Kimberley of

Western Australia. It inhabits low open eucalypt

woodland on gravelly hills and flats.

Phenology : Flowers and fruits are recorded

from April to December.

Notes: S. quadriloculatum is closely related to

S. ellipticum, but differs by the conspicuously

ridged branchlets, the presence of long-stalked

stellae (stalks > 0.4 mm long and up to 1.6

mm long) on all plant parts, the mostly larger

stellae on the upper and lower leaf surfaces,

the 9-15 flowered inflorescences (1-9 flowered

for S. ellipticum ), the flimsy (non-rigid)

prickles on the calyx, and the absence of

prickles on the rachis of the inflorescence.

Solanum quadriloculatum is consistently

4-locular, but some forms of S. ellipticum may

also be 4-locular.

The type of S. quadriloculatum was

collected from the Nicholson River at about 18°

latitude. All subsequent collections of

S. quadriloculatum have been made further

south (19-22.5° latitude). The type (which is

rather scrappy) does not match the subsequent

747

collections terribly well. It has broad based

prickles, < 10 times longer than wide, the stellae

of the upper leaf surface are relatively widely

spaced (moderate to dense) and the stellae stalks

are very short. Collections from the type locality

are sorely needed to determine if the name has

been correctly applied. The full description

given above is based on the modem collections.

Conservation status: Widespread. Not

considered at risk.

50. Solanum crassitomentosum Domin,

Biblioth. Bot. 89: 584 (1929). Type:

Queensland. North Kennedy District:

Dividing range west of Pentland,

February 1910, K. Domin (holo: PR).

Erect, rhizomatous perennial shrub, 0.3-1 m

high. Juvenile stage unknown. Adult stems bark

furrowed, corky. Adult branchlets terete or

ridged, yellow or rusty or brown; prickles

30-160 per decimetre, straight, acicular, 1-5

mm long, 10-16 t im es longer than wide; stellae

very dense, 0.25-0.7 mm diameter, stalks

0-0.7 mm long; lateral rays 6-9, ascending;

central ray 1-2 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Adult leaves ovate or broadly ovate,

entire; lamina 3.5-7.5 cm long, 1.5-3.5 cm

wide, 1.7-2.3 times longer than broad, apex

acute, base obtuse or cordate, oblique part 0-3

mm long, obliqueness index 0-6 percent;

petioles 0.8-2.6 cm long, 16-35% length of

lamina, prickles absent or present. Upper leaf

surface grey; prickles absent; stellate hairs

distributed throughout; protostellae absent;

ordinary stellae very dense, c. 0.05 mm apart,

0.5-0.7 mm across, stalks 0-0.7 mm long;

lateral rays 6-8, porrect; central ray 1.2-2.5

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Lower

leaf surface white or yellowish; prickles absent;

stellae dense to very dense, 0.05-0.1 mm apart,

0.4-0.8 mm diameter, stalks 0-1 mm long;

lateral rays 7 or 8, porrect; central ray 1.5-2

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent.

Inflorescence leaf-opposed, solitary or pseudo-

racemose, co mm on peduncle 5-14 mm long,

rachis prickles absent; 1 or 2-flowered, strongly

or weakly andromonoecious, flowers 5-merous;

pedicels 4-11 mm long at anthesis, same

748

thickness throughout or markedly thicker

distally, 1.1-1.5 mm thick at mid-point,

prickles absent. Calyx tube 3.5-5 mm long,

lobes deltate, 4-7 mm long; prickles absent at

anthesis; stellae very dense, transparent or

brown or rusty, 0.5-0.7 mm across, stalks

0.3-0.9 mm long, lateral rays 7 or 8, central

ray 1-2 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Corolla white, mauve or purple, 12-17 mm

long, shallowly or deeply lobed, inner surface

densely stellate-hairy; anthers 5.5-7.5 mm

long; ovary glabrous, or with Type 2 hairs only;

functional style 10.5-12.5 mm long, erect,

glabrous. Fruiting calyx with lobes less than

or more than half length of mature fruit,

prickles absent. Mature fruits 1 per

inflorescence, globular, 14-18 mm diameter,

yellowish-green or green, 4-locular; placenta

in cross-section stalked, circular to elliptical;

mesocarp moist but not juicy; exocarp 2.4-3

mm thick; pedicels 7-13 mm long in fruit,

1.3-1.5 mm thick at mid-point; seeds brown

to black, 2.5-2.8 mm long.

Specimens examined : Queensland. Burke District:

‘Warang’ area, White Mountains N.P., Apr 2000, McDonald

KRM385 (BRI); NW of ‘Warang’, White Mountains N.P.,

Apr 2000, McDonald KRM467 (BRI); 20 km Wof ‘Warang’

HS site, White Mountains N.P, 57 km by road N of Torrens

Creek, Apr 2000, Thomas 1835 & Thompson (BRI, NSW);

3 km NW of ‘Warang’ HS site, White Mountains N.P., Apr

2000, Thomas 1840 & Thompson (BRI, NSW). North

Kennedy District: W of Pentland, between Warrigal and

Burra, Oct 1935, Blake 9921 (BRI); Burra Range, W of

Pentland, Jul 1985, Williams 85048 (BRI); White Mountains

N.P, east of ‘Warang’, Mar 2000, Wannan 1629 &

Martindale (BRI, MEL). Mitchell District: Poison Valley

road, White Mountains N.P., Apr 2000, McDonald KRM426

(BRI); 7 km E of ‘Warang’ HS on track to Sandstone Wall,

Apr 2000, Thomas 1452 & Thompson (BRI, NSW).

Distribution and habitat: Solanum

erassitomentosum is endemic to Queensland.

Confined to the White Mountains-Burra Range

area, W of Townsville (Map 19). It grows in

eucalypt woodland on sandstone. The dominant

tree species include Eucalyptus exilipes,

E. lamprophylla, E. leichhardtii, Acacia

shirleyi or Lysicarpus angustifolius. Triodia is

often present in the ground layer.

Phenology: Flowers are recorded for March,

April, July and October; mature fruits recorded

for April.

Austrobaileya 6 (4): 639-816 (2004)

Notes: S. eras sitomento sum is related to

S. quadriloculatum, but differs by the 1 or 2

flowered inflorescence (9-15 flowered for

S. quadriloculatum ), calyx and leaf laminae

without prickles (leaves and calyx prickly for

S. quadriloculatum ), inner surface of corolla

densely stellate hairy (vs. glabrous for

S. quadriloculatum), central ray of stellae on

lower leaf surface 1.5-2 times lateral rays (vs.

0.8-1.2 times for S. quadriloculatum) and the

brown to black seeds (vs. pale yellow for

S. quadriloculatum).

Conservation status: S. eras sitomento sum is

currently known from 4 locations, all within

the White Mountains National Park. There are

no substantial threats to the species, and it is

not considered to be at risk.

51. Solanum angustum Domin, Biblioth. Bot.

80: 588-589 (1929). Type: Queensland.

Cook District: Walsh River, north of

Chillagoe, February 1910, K. Domin

(holo: PR).

Sprawling, herbaceous resprouter, c. 0.3 m

high. Juvenile stage absent. Adult branchlets

grey or yellow; prickles 5-30 per decimetre,

straight, acicular, 3-6 mm long, 10-15 times

longer than wide; stellae very dense, 0.3-0.5

mm diameter, stalks 0-0.1 mm long; lateral

rays 6-8, porrect; central ray present, 0.5-1

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Adult

leaves narrow lanceolate to ovate or elliptical,

entire or shallowly lobed throughout; lobes 2

or 3 on each side, obtuse, lobing index 1-1.3;

la min a 1.5-4 cm long, 0.4-1.4 cm wide, 3-7.5

times longer than broad, apex obtuse, base

cuneate, oblique part 0-1 mm long, obliqueness

index 0-3 percent; petioles 0.3-2.2 cm long,

17-55% length of lamina, prickles absent.

Upper leaf surface green; prickles 0-3, straight,

acicular, 3-6 mm long, prickles absent or

present on midvein only; stellate hairs

distributed throughout; protostellae absent;

ordinary stellae sparse to moderate density,

0.3-0.9 mm apart, 0.25-0.4 mm across, sessile;

lateral rays 6-8, porrect; central ray 0.5-1 times

as long as laterals, not gland-tipped; finger

hairs absent; Type 2 hairs absent. Lower leaf

surface green or yellowish; prickles absent;

stellae sparse to dense, 0.1-0.7 mm apart,

0.3-0.6 mm diameter, stalks 0-0.1 mm long;

Bean, Taxonomy of Solanum subg. Leptostemonum

lateral rays 7 or 8, porrect; central ray 0.5-1

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent.

Inflorescence leaf-opposed, solitary or pseudo-

racemose, common peduncle 0-15 mm long,

rachis prickles present; 1-4-flowered, flowers

5-merous; pedicels c. 7 mm long at anthesis,

same thickness throughout, c. 0.8 mm thick at

mid-point, prickles present. Calyx tube 2.5-3

mm long, lobes deltate, 2-2.5 mm long;

prickles present at anthesis, 2-10 per flower,

1-4 mm long; stellae dense, transparent,

0.3-0.4 mm across, sessile, lateral rays 7 or 8,

central ray 0.5-1 t im es as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Corolla c. 10 mm long. Fruiting calyx

with lobes more than half length of mature fruit,

prickles 1-4 mm long. Mature fruits 1 per

inflorescence, globular, 16-20 mm diameter,

green; pedicels 15-20 mm long in fruit,

0.9-1.2 mm thick at mid-point; seeds pale

yellow, 2.5-3 mm long.

Specimens examined : Queensland. Cook District:

Stannary Hills, 1908, Bancroft 173 (BRI); Alma-den, Nov

1939, Thurston 575 (QRS); 26 km W of Einasleigh on road

to Forsayth, on top of Newcastle Range, Apr 1992, Halford

Q958 (AD, BRI, MEL).

Distribution and habitat : Solanum angustum

is endemic to Queensland and known only from

the type and two other collections (cited above)

(Map 20). The label of the only recent

collection indicated that the species was

growing in “low eucalypt woodland with

shallow clay soil”.

Phenology: Very poorly known. Fruits have

been collected in April.

Notes: The affinities of this species are not

known with any certainty, and it is only

tentatively placed in the S. ellipticum group.

Conservation status: S. angustum has been

collected just four times, in 1908, 1910, 1939

and 1992. A recent visit to the 1992 collection

site failed to find the species. It has not been

seen since 1992, despite deliberate searches

during the summer months, especially in the

Chillagoe-Walsh River area. Although there

are no known extant populations, a

presumption of extinction is premature. There

would appear to be much available habitat

within the extent of occurrence, although the

749

required habitat for the species is still not

known.

Applying the IUCN guidelines (IUCN.

2001), and using the 1992 location as an

existing location, a category of “Endangered”

is recommended (EN C2a(ii); D).

52. Solanum argopetalum A.R.Bean sp. nov.

Frutex prostratus vel fusus; folia late

ovata, non profunde lobata, basi cordata

et petiolis 12-35 mm longis; pili stellati

aliqui apicibus glandularibus in ramulis

foliisque praesentes; aculei in ramulis

sparse distributi, 1-5 mm longi, calyci

absentes; corolla albida, 7-9 mm longa;

fructus maturi virides usque flavi. Typus:

Queensland. Maranoa District: near

‘Boxleigh’, c. 60 km NE of St George,

24 April 2001, A.R. Bean 17644 & L.

Pedley (holo: BRI; iso: AD, CANB, K,

MEL, MO, NSW, PRE, distribuendi ).

Prostrate or sprawling, herbaceous resprouter,

0.1-0.4 m high. Juvenile branchlets with

50-100 prickles per dm, 2-5 mm long; leaves

(in outline) broadly ovate, shallowly-lobed, with

2-4 pairs of lobes; lamina c. 5 x 4.5 cm, with

20-40 prickles on upper surface. Adult

branchlets yellow, rusty or brown; prickles

5-25 per decimetre, straight, acicular, 1-5 mm

long, 10-17 times longer than wide; stellae

dense, 0.4-0.7 mm diameter, stalks 0-0.25 mm

long; lateral rays 6-11, porrect or ascending;

central ray present, 1.5-3 times as long as

laterals, not gland-tipped or gland-tipped;

finger hairs absent; Type 2 hairs sparse. Adult

leaves broadly ovate or orbicular, entire or

shallowly lobed throughout; lobes 2 or 3 on each

side, obtuse, lobing index 1-1.2; lamina

3.5-8.5 cm long, 2-5.5 cm wide, 1.2-1.7 times

longer than broad, apex obtuse, base cordate,

oblique part 0-3.5 mm long, obliqueness index

0-5 percent; petioles 1.2-3.5 cm long, 30-65%

length of lamina, prickles absent. Upper leaf

surface grey-green; prickles 0-2, straight,

acicular, 1-3 mm long, prickles absent or

present on midvein only; stellate hairs

distributed throughout; protostellae present;

ordinary stellae density moderate, 0.4-0.6 mm

apart, 0.6-0.9 mm across, stalks 0-0.3 mm

long; lateral rays 7-10, porrect or ascending;

central ray 1-3 times as long as laterals, not

gland-tipped or gland-tipped; finger hairs

750

Austrobaileya 6 (4): 639-816 (2004)

Fig. 32. Solatium argopetalum. A. flowering branchlet x 1. B. ovary and style x 6. C. ovary with Type 2 hairs x 12. D. stellate

hair, upper leaf surface x 50. E. mature fruit and calyx x 2. F. transverse section of fruit x 3. A-D, Bean 17644 & Pedley; E-

F, Bean 17768.

Bean, Taxonomy of Solanum subg. Leptostemonum

absent or present, 0.3-0.6 mm apart, gland-

tipped, 0.1-0.3 mm long; Type 2 hairs absent.

Lower leaf surface green, or greenish-white;

prickles absent; stellae dense, 0.1-0.2 mm

apart, 0.5-0.6 mm diameter, stalks 0-0.1 mm

long; lateral rays 8-11, porrect; central ray

1-1.5 t im es as long as laterals, not gland-tipped

or gland-tipped; finger hairs absent; Type 2

hairs absent. Inflorescence leaf-opposed or

supra-axillary, solitary or pseudo-racemose,

common peduncle 0-2 mm long, rachis prickles

absent or present; 1-4-flowered, weakly

andromonoecious or with all flowers bisexual,

flowers 4 or 5-merous; pedicels 3-10 mm long

at anthesis, same thickness throughout,

0.5-0.6 mm thick at mid-point, prickles absent.

Calyx tube 1.5-3 mm long, lobes deltate,

rostrate or attenuate, 2-7 mm long; prickles

absent at anthesis; stellae dense, transparent,

0.4-0.7 mm across, stalks 0-0.2 mm long,

lateral rays 6-10, central ray 1-2 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Corolla white,

7- 9 mm long, deeply lobed, inner surface

glabrous; anthers 3-4 mm long; ovary with

Type 2 hairs only; functional style 4.5-6 mm

long, erect, with Type 2 hairs only. Fruiting

calyx with lobes less than or more than half

length of mature fruit, prickles absent. Mature

fruits 1-3 per inflorescence, globular, c. 15 mm

diameter, yellowish-green or green, 1-locular

(septum absent or incomplete); placenta in

cross-section sessile, elliptical; mesocarp moist

but not juicy; exocarp 1-1.5 mm thick; pedicels

8- 20 mm long in fruit, 1-1.2 mm thick at mid¬

point; seeds pale yellow, 2.7-3. 1 mm long. Fig. 32.

Specimens examined : Queensland. Maranoa District: 8

km E of Mungallala, Mar 1960, Johnson 1483 (BRI); near

‘Boxleigh’, c. 60 km NE of St George, Apr 2001, Bean 17643

& Pedley (BRI); 1.1 km W of ‘Eulorel’, W of Surat, Aug

2001, Bean 17747 (BRI, MEL); ‘Boxleigh’, S of Surat, Aug

2001, Bean 17761 (BRI); SWof ‘Boxleigh’, S of Surat, Aug

2001, Bean 17768 (BRI); ‘Glen Fosslyn’, SW of

Glenmorgan, Jan 2002, Bean 18368 (BRI); 54 km from

Mitchell towards Bollon, Jan 2002, Bean 18406 (BRI).

Distribution and habitat : Solanum

argopetalum is endemic to Queensland.

Confined to a limited area of southern

Queensland, recorded from Mungallala to the

Thomby Range south-east of Surat (Map 20).

It grows on disturbed sites on low hills and

ridges in shallow red sandy to loamy soil, often

in association with Acacia catenulata (Bendee),

751

Eucalyptus melanophloia and Phebalium

glandulosum.

Phenology: Flowers have been recorded in

April and August; mature fruits recorded in

August.

Notes: This species is quite distinctive. It does

seem fairly close to S. crassitomentosum

because of its short prickles, unarmed calyx and

relatively small flowers, but S. argopetalum is

easily distinguished by its often gland-tipped

stellate hairs with 8-11 lateral rays, the lobed

leaves, the long petioles (30-65% of lamina

length) and the white corolla.

The populations of S. argopetalum so far

recorded have all been on disturbed sites. At

the type locality, plants were quite common on

a strip of roadside land that had been cleared

in preparation for the erection of a fence. The

species could not be found on the opposite

(relatively undisturbed) side of the road, even

though the vegetation type was identical.

Conservation status: S. argopetalum is

currently known from 4 locations, none of

which is within a conservation reserve. It is

threatened by land clearance and weeds

(especially Cenchrus ciliaris). Applying the

IUCN guidelines (IUCN. 2001), a category of

“Vulnerable” is recommended (VU B2ab(iii,v);

Cl+2a(i)).

Etymology: From the Greek argos- white and

petalos- petals, in reference to the white corolla.

Consistently white flowers are found in only a

few native Queensland species.

Group 27A (S. hamulosum group), here

defined; related to Group 27B (S. macoorai

group).

Style and ovary bearing abundant stellate

hairs (100%); scrambling or vine-like habit

(100%); leaves consistently geminate

(100%); branchlets with broad recurved

prickles (100%); calyx prickles absent

(100%); lower leaf surface white or yellowish

(100%); inflorescences supra-axillary

(100%); inner surface of corolla sparsely to

densely stellate-hairy (100%); juvenile leaves

(and sometimes adult leaves) with shallow

lobes and very numerous prickles (100%);

flower buds ovoid (100%).

752

Austrobaileya 6 (4): 639-816 (2004)

c. 6 species from Malesia to Australia; 3

species endemic to Australia, all in Queensland.

Symon (1981) included S. dimorphispinum

and S. hamulosum with S. sect. Micracantha

Dunal, a group of species from tropical

America. Whalen (1984) pointed out several

morphological differences from S. sect.

Micracantha, that precludes any close

affinity. Symon (1981) further hypothesised

that S. dimorphispinum and S. hamulosum

were both naturalised from an unknown

American source. This is untenable, given the

close relationship between these species and

others from the same part of Queensland,

particularly S. macoorai, and the presence of

related species in Malesia. The Malesian

species probably belonging to the S. hamulosum

group are S. heteracanthum Merr. & L.M.Perry

and S. scheferi F.Muell. from New Guinea, and

S. lianoides Elmer from Philippines.

53. Solanum dimorphispinum C.T.White,

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

Type: Queensland. Cook District: Mt

Spurgeon, September 1936, C.T. White

10619 (holo: BRI).

Illustration : Symon (1981: 253)

Erect, rhizomatous perennial shrub or vine,

2-5 m high. Juvenile branchlets with c. 20

prickles per dm, 2-3 mm long; leaves (in

outline) ovate, shallowly-lobed, with 6-7 pairs

of lobes; lamina c. 22 x 15 cm, with c. 90

prickles on upper surface. Adult branchlets

yellow or brown; prickles 15-30 per decimetre,

curved, broad-based, 2-6 mm long, 2-5 times

longer than wide; stellae sparse to dense,

0.15-0.25 mm diameter, sessile; lateral rays

6-8, porrect; central ray absent; finger hairs

absent; Type 2 hairs absent. Adult leaves ovate,

entire or shallowly lobed throughout; lobes

2-6 on each side, obtuse, lobing index 1-1.3;

lamina 10-14.5 cm long, 4.5-7 cm wide,

1.9-2.5 times longer than broad, apex acute or

acuminate, base cuneate, oblique part 3-7 mm

long, obliqueness index 2-6 percent; petioles

1.3-2.4 cm long, 13-18% length of lamina,

prickles absent or present. Upper leaf surface

green; prickles 0-15, straight, broad-based,

5-7 mm long, prickles absent or present on

midvein only or present on midvein and lateral

veins; stellate hairs confined to midrib, or

distributed throughout; protostellae absent;

ordinary stellae very sparse, 0.7-4.5 mm apart,

0.25-0.35 mm across, sessile; lateral rays 7-9,

porrect; central ray 0-0.5 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Lower leaf surface white

or yellowish; prickles 0-4, straight, acicular or

broad-based, prickles absent or present on

midvein only or present on midvein and lateral

veins; stellae very dense, c. 0.05 mm apart,

0.25-0.35 mm diameter, sessile; lateral rays

7-10, porrect, central ray absent; finger hairs

absent; Type 2 hairs absent. Inflorescence

supra-axillary, pseudo-racemose, common

peduncle 0-10 mm long, rachis prickles absent;

5-17-flowered, weakly andromonoecious,

flowers 5-merous; pedicels 12-23 mm long at

anthesis, same thickness throughout, 0.6-0.9

mm thick at mid-point, prickles absent. Calyx

tube 1-3 mm long, lobes deltate, 3-5 mm long;

prickles absent at anthesis; stellae very dense,

transparent or purple, 0.2-0.3 mm across,

sessile, lateral rays 8-10, central ray 0-0.5

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Corolla

mauve, 8-14 mm long, deeply lobed, inner

surface sparsely to densely stellate-hairy;

anthers 5-6.5 mm long; ovary with stellate

hairs only; functional style 7-8.5 mm long,

erect, with stellate hairs only, stellae 0.2-0.4

mm across, lateral rays 11-15, central ray

0.5-1 times as long as laterals. Fruiting calyx

with lobes less than half length of mature fruit,

prickles absent. Mature fruits 1-3 per

inflorescence, oblate or globular, 30-35 mm

diameter, yellow, 2-locular; placenta in cross-

section stalked, anvil-shaped; mesocarp moist but

not juicy; exocarp 3.8-4.2 mm thick; pedicels

29-39 mm long in fruit, 0.9-1.8 mm thick at

mid-point; seeds pale yellow, 2.3-2.8 mm long.

Specimens examined : Queensland. Cook District:

McDowall Range between Mossman and Bloomfield River,

May 1969, Webb & Tracey 8302 (BRI, CANB); Mt Lewis,

Oct 1969, Webb & Tracey 8352 (BRI, CANB); T.R.140,

Cow L.A., Sep 1973, Hyland 6874 (QRS); S.F.143, South

Mary L.A., Feb 1974, Hyland 7205 (BRI, QRS); 23 miles

[37 km] N of Mt Molloy on road to Mossman, Aug 1974,

Swan 138 (BRI); T.R.66, Oct 1977, Moriarty 2278 (BRI);

S.F. 143, Kanawarra, Carbine L.A., Oct 1991, Gray 5336

(BRI, QRS); Black Mountain, 16°38’S 145°29’E, Jul 1999,

Jago 5314 et al. (BRI); 29.5 km along Mt Lewis road, Mt

Lewis Forest Reserve, Nov 2001, McDonald KRM1013

(BRI, HO).

Bean, Taxonomy of Solanum subg. Leptostemonum

Distribution and habitat : Solanum

dimorphispinum is endemic to Queensland. It

is known only from the Mt Lewis and Mt

Spurgeon areas (Map 2). It inhabits notophyll

rainforest at altitudes above 900 metres.

Phenology: Flowers are recorded between May

and November; mature fruits in November and

December.

Notes : S. dimorphispinum is one of the few

species for which all stellae lack a central ray.

The mature fruits are pale yellow and globose,

30-35 mm diameter. Smaller immature fruits

(seen in spirit collection) are quite ellipsoidal

in shape.

Conservation status: Currently listed as “Rare”

under the Queensland Nature Conservation Act,

1992. The extent of occurrence is small, but

the species is not considered at risk.

54. Solanum hamulosum C.T.White, Contr.

Arnold Arbor. 4: 95 (1933). Type:

Queensland. Cook District: Boonjie,

Atherton Tableland, 23 September 1929,

S.F. Kajewski 1222 (holo: BRI).

Illustration: Symon (1981: 254)

Erect, perennial vine, 1.5-7 m high. Juvenile

branchlets with 30-40 prickles per dm, 2-3 mm

long; leaves (in outline) broadly ovate,

shallowly-lobed, with 4-6 pairs of lobes; lamina

18-22 cm long, 9-13 cm wide, with 50-100

prickles on upper surface. Adult branchlets grey

to rusty or brown; prickles 50-140 per

decimetre, curved, broad-based, 1-3.5 mm

long, 1-2.5 t im es longer than wide; stellae very

dense, 0.3-0.4 mm diameter, stalks 0-0.4 mm

long; lateral rays 7-10, porrect or ascending;

central ray 0.5-1 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Adult leaves ovate, entire; lamina

11-15 cm long, 5-6.5 cm wide, 1.7-2.4 times

longer than broad, apex acute or acuminate,

base cuneate, obtuse or cordate, oblique part

0-6 mm long, obliqueness index 0-6 percent;

petioles 1.9-3.8 cm long, 17-25% length of

lamina, prickles present. Upper leaf surface

green or grey-green; prickles 0-10, straight,

broad-based, 2.5-7 mm long, prickles absent

or present on midvein only or present on

midvein and lateral veins; stellate hairs

distributed throughout; protostellae present;

753

ordinary stellae sparse to dense, 0.2-0.3 mm

apart, 0.25-0.4 mm across, sessile; lateral rays

6-8, porrect; central ray 0.7-1.2 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Lower leaf surface

white or yellowish; prickles 1-6, curved, broad-

based, prickles present on midvein only; stellae

dense, 0.1-0.15 mm apart, 0.4-0.5 mm

diameter, stalks 0-0.1 mm long; lateral rays

8-9, porrect; central ray 0.7-1.5 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Inflorescence

supra-axillary, pseudo-racemose, common

peduncle 0-5 mm long, rachis prickles absent

or present; 5-11-flowered, weakly

andromonoecious, flowers 5-merous; pedicels

8-18 mm long at anthesis, markedly thicker

distally, 0.5-0.6 mm thick at mid-point,

prickles absent. Calyx tube 2-3.5 mm long,

lobes deltate, 2-3.5 mm long; prickles absent

at anthesis; stellae dense to very dense, white,

brown or rusty, 0.25-0.5 mm across, stalks

0-0.2 mm long, lateral rays 7 or 8, central ray

0.7-1.2 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Corolla mauve, 9-15 mm long, shallowly or

deeply lobed, inner surface sparsely stellate-

hairy; anthers 4.5-6 mm long; ovary with

stellate hairs only; functional style 8-8.5 mm

long, erect, with stellate hairs only, stellae

0.3-0.5 mm across, lateral rays 6-8, central

ray 0.7-1.2 t im es as long as laterals. Fruiting

calyx with lobes less than half length of mature

fruit, prickles absent. Mature fruits 1 or 2 per

inflorescence, globular, 13-15 mm diameter,

yellowish-green or green, 2-locular; placenta

in cross-section sessile, semi-circular, or

stalked, anvil-shaped; mesocarp moist but not

juicy; exocarp 0.8-1 mm thick; pedicels 18-25

mm long in fruit, 0.8-1.2 mm thick at mid¬

point; seeds pale yellow, 2-2.5 mm long. Dirran

Curse.

Specimens examined : Queensland. Cook District: Tarzali,

the Dirran and Russell River road, Feb 1918, White s.n. (BRI);

Millaa Millaa, Sep 1937, Henry s.n. (QRS); R310, Boonjie,

Aug 1961, Hyland AFO 2041 (QRS); near Boonjie, May

1967, Symon 4750 (BRI); Davies Creek S.F, Aug 1973,

Moriarty 1371 (BRI, CANB); western foo thi lls of Mt Bartle

Frere, Aug 1973, Moriarty 1434 (BRI, CANB); S.F.755,

T.R. 1230, Boonjie L.A., Dec 1974, Irvine 1094 (BRI, QRS);

Herberton Range, McKell Road turnoff, Aug 2003,

McDonald KRM1165 (BRI). North Kennedy District:

Ravenshoe, Sep 1937, Brass & White 132 (BRI); Keogh’s

block, ‘Bellview’, Evelyn, Jan 1974, Collins C74-12 (BRI);

754

S.F.194, Parish of Herberton, Dec 1991, Hyland 14421 (BRI,

QRS).

Distribution and habitat: Solanum hamulosum

is endemic to Queensland, and known only

from the Atherton Tableland (Map 21). It grows

in notophyll rainforest on basaltic soils, between

600-1100 metres altitude.

Phenology: Flowers are recorded between

August and December; mature fruits in

November and December.

Notes: S. hamulosum and S. dimorphispinum

are very similar in habit, habitat, leaf size and

prickle shape. They can readily be distinguished

on characters of the stellate hairs, but it was

not known whether these differences correlated

to differences on other organs of the plant.

During this study I was able to procure mature

fruits of both species; those of S. dimorphispinum

are 30-35 mm in diameter, while those of

S. hamulosum are only 13-15 mm diameter.

This correlation is seen as a vindication for

using stellate hair morphology in diagnosing

and separating Solanum taxa where there is

incomplete data about the flowering or fruiting

characters. The hairs are always available for

study, and are unaffected by drying or

preserving. Many other useful characters are

rarely available or visible only on fresh

material, or difficult to determine from dried

specimens.

Conservation status: Currently listed as “Rare”

under the Queensland Nature Conservation Act,

1992.

Solanum hamulosum was considered a

weed of secondary regrowth during the 1930’s

and 1940’s, when the rainforests of the Atherton

Tableland were being cleared, and was called

‘Dirran Curse’. Now, little of the original

habitat remains, and the species is exceedingly

difficult to locate. The rainforest edges (the most

likely habitat) are usually swathed in Lantana

camara. Applying the IUCN guidelines (IUCN.

2001), acategory of “Endangered” is recommended

(EN Blab(iii,iv,v)+2ab(iii,iv,v); Cl).

55. Solanum eminens A.R.Bean sp. nov. Vitis

perennis vel frutex scandens; folia adulta

viridia, late ovata, non profunde lobata,

aculeis utrinque rectis numerosis; ramuli

moderate aculeati (70-90 per dm.), aculei

Austrobaileya 6 (4): 639-816 (2004)

curvi, ad basim lati; inflorescentia

pseudo-racemosa, 7-10-flora; ovarium

tantum pilis stellatis praeditum; calycis

aculei absentes; fructus maturi aurantiaci,

pedicellis 23-28 mm longis. Typus:

Queensland. Cook District: Bellenden

Ker Range, 11 October 1974, B. Hyland

7772 (holo: BRI; iso: QRS).

Sprawling perennial vine, 1-2 m high. Juvenile

stage unknown. Adult branchlets brown;

prickles 70-90 per decimetre, curved, broad-

based, 2-4 mm long, 2.5-4 times longer than

wide; stellae sparse to dense, 0.2-0.3 mm

diameter, stalks 0-0.1 mm long; lateral rays 7

or 8, porrect; central ray 0.4-0.8 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Adult leaves

broadly ovate, shallowly lobed throughout;

lobes 5-7 on each side, acute, lobing index

1.1-1.3; lamina 8-12 cm long, 5-7.5 cm wide,

1.4-1.8 t im es longer than broad, apex acute or

acuminate, base cuneate, oblique part 0-7 mm

long, obliqueness index 0-7 percent; petioles

1.6-3 cm long, 20-30% length of lamina,

prickles present. Upper leaf surface green;

prickles 40-120, straight, acicular, 3-11 mm

long, prickles present on midvein and lateral

veins; stellate hairs distributed throughout;

protostellae absent; ordinary stellae very sparse

to sparse, 0.6-1.4 mm apart, 0.2-0.35 mm

across, sessile; lateral rays 7 or 8, porrect;

central ray 0.5-1 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Lower leaf surface white or yellowish;

prickles 15-30, straight, acicular or broad-

based, prickles present on midvein and lateral

veins; stellae dense to very dense, 0.05-0.2 mm

apart, 0.25-0.35 mm diameter, sessile; lateral

rays 7 or 8, porrect; central ray 0.3-0.6 times

as long as laterals, not gland-tipped; finger

hairs absent; Type 2 hairs absent. Inflorescence

supra-axillary, pseudo-racemose, common

peduncle 6-12 mm long, rachis prickles absent

or present; 7-10-flowered, weakly

andromonoecious, flowers 5-merous; pedicels

12-18 mm long at anthesis, same thickness

throughout, 0.5-0.6 mm thick at mid-point,

prickles absent or present. Calyx tube 2.5-3.5

mm long, lobes deltate or rostrate, 0.5-3 mm

long; prickles absent at anthesis; stellae dense,

yellow or white or purple, 0.2-0.3 mm across,

sessile, lateral rays 7 or 8, central ray 0.4-1

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Corolla

Bean, Taxonomy of Solanum subg. Leptostemonum

755

Fig. 33. Solanum eminens. A. flowering branchlet x 0.8. B. ovary and style x 6. C. ovary with stellate hairs x 20. D. stellate

hair, upper leaf surface x 60. A, D, Powell 796 et al. ; B-C, Clarkson 6575.

mauve, 9-14 mm long, deeply lobed, inner

surface sparsely stellate-hairy; anthers 5.5-6.5

mm long; ovary with stellate hairs only;

functional style c. 8.5 mm long, erect, with

stellate hairs only, stellae c. 0.2 mm across,

lateral rays 6-8, central ray c. 1 times as long

as laterals. Fruiting calyx with lobes less than

half length of mature fruit, prickles absent.

Mature fruits 1 per inflorescence, c. 17 mm

diameter, orange; pedicels 23-28 mm long in

fruit, 1-1.3 mm thick at mid-point; seeds pale

yellow, 1.9-2.4 mm long. Fig. 33.

Specimens examined : Queensland. Cook District:

Bellenden Ker, Nov 1972, Hyland 6584 (BRI, QRS);

Bellenden Ker Range, Oct 1974, Hyland 7757 (BRI, CANB,

QRS); summit of Bellenden Ker, S of Cairns, Sep 1977,

Powell 796 et al. (BRI, NSW); Mount Bellenden Ker, near

the cableway terminus, Sep 1986, Clarkson 6575 (AD, BRI,

MBA, QRS); central peak of Bellenden Ker, Wooroonooran

N.P., Jun 1995, Hunter JH5302 (BRI).

Distribution and habitat: Solanum eminens is

endemic to Queensland. Apparently confined

to the highest altitudes (above 1500 metres) on

Mt Bellenden Ker (Map 23), in wet microphyll

756

to notophyll fern forest.

Phenology : Flowers are recorded in October;

mature fruits in June and November

Notes: Two collectors have independently

stated that the mature fruit colour is orange.

Solanum eminens differs significantly

from S. hamulosum by the adult leaves that have

5-7 pairs of conspicuous acute lobes, and with

40-120 prickles on the upper leaf surface

(leaves entire, 0-10 prickles for S. hamulosum );

the very sparse to sparse indumentum of the

upper leaf surface (moderate to dense

indumentum for S. hamulosum ); the stellae of

the lower leaf surface 0.25-0.35 mm diameter

with central ray 0.3-0.6 times as long as laterals

(0.4-0.5 mm diameter and central ray 0.7-1.5

times as long as laterals for S. hamulosum ) and

the orange mature fruit (green for S. hamulosum ).

Conservation status: S. eminens is known only

from the highest parts of Mt Bellenden Ker in

Wooroonooran N.P. It is threatened by low

population size and restricted area of

occupancy. Applying the IUCN guidelines

(IUCN. 2001), a category of “Vulnerable” is

recommended (VU Dl).

Etymology: From the Latin, eminens meaning

“projecting, high”. This is in reference to the

altitude at which this species grows (1500-1600

metres), the highest for any Australian

Solanum.

Group 27B (S. macoorai group) = Group 6 of

Whalen (1984)

Calyx not accrescent in fruit (100%);

inflorescences andromonoecious, male flowers

and bisexual flowers same size and prickliness

(100%); branchlet stellae not gland-tipped

(100%); branchlet finger hairs absent (100%);

mature fruits green to yellowish-green (94%);

branchlet prickles acicular (89%); adult leaves

entire or shallowly lobed (87%); seeds white

to pale yellow (83%); adult leaves entire (78%);

calyx without prickles (77%); rhizomatous

perennial shrubs (72%); inner surface of corolla

stellate-hairy (71%).

24 species endemic to Australia, 18

species indigenous to Queensland and north¬

eastern N.S.W.

Austrobaileya 6 (4): 639-816 (2004)

56. Solanum macoorai F.M.Bailey,

Queensland Dept. Agric. Bull. 8: 80

(1893). Type: Queensland. Cook District:

summit of south peak, Mt Bellenden-Ker,

June 1889, F.M. Bailey s.n. (holo: BRI).

Illustration: Symon (1981: 241)

Erect, rhizomatous perennial shrub, 1-4 m

high. Juvenile branchlets with 90-200 prickles

per dm, 2-12 mm long; leaves (in outline)

ovate, shallowly-lobed, with 7-9 pairs of lobes;

lamina c. 12x5 cm, with 100-150 prickles on

upper surface. Adult branchlets yellow, rusty

or brown; prickles absent or present, 0-55 per

decimetre, straight, acicular, 3-7 mm long,

7-12 times longer than wide; stellae sparse to

dense, 0.15-0.25 mm diameter, sessile; lateral

rays 6-8, porrect; central ray absent or present,

0-0.2 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Adult leaves elliptical or ovate, entire or

shallowly lobed throughout; lobes 3-5 on each

side, acute or obtuse, lobing index 1-1.2;

lamina 9-16 cm long, 3-7.5 cm wide, 2.1-3.2

times longer than broad, apex acute or

acuminate, base cuneate or obtuse, oblique part

0-9 mm long, obliqueness index 0-6 percent;

petioles 1.4-2.6 cm long, 10-18% length of

lamina, prickles absent. Upper leaf surface

green; prickles 0-80, straight, acicular, 3-8 mm

long, prickles absent or present on midvein only

or present on midvein and lateral veins; stellate

hairs confined to midrib, or distributed

throughout; protostellae absent; ordinary stellae

very sparse, 0.8-5 mm apart, 0.2-0.3 mm

across, sessile; lateral rays 6-8, porrect; central

ray 0-0.5 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Lower leaf surface green; prickles 0-16,

straight, acicular, absent or present on midvein

only or present on midvein and lateral veins;

stellae very sparse to moderate, 0.2-0.5 mm

apart, 0.2-0.3 mm diameter, sessile; lateral rays

7-10, porrect; central ray 0-0.3 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Inflorescence

supra-axillary, pseudo-racemose, common

peduncle 1-7 mm long, rachis prickles absent;

7-30-flowered, weakly andromonoecious,

flowers 5-merous; pedicels 6-12 mm long at

anthesis, markedly thicker distally, 0.4-0.7 mm

thick at mid-point, prickles absent. Calyx tube

Bean, Taxonomy of Solanum subg. Leptostemonum

2-4 mm long, lobes deltate or rostrate, 1.8-3.5

mm long; prickles absent at anthesis; stellae

dense to very dense, yellow, transparent or

purple, 0.2-0.25 mm across, sessile, lateral rays

6- 9, central ray 0-1 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Corolla mauve or purple, 8-10

mm long, deeply lobed, inner surface sparsely

stellate-hairy; anthers 4-5.5 mm long; ovary

with stellate hairs only, or with stellate and Type

2 hairs; functional style 5-7 mm long, erect,

with stellate hairs only or with stellate and Type

2 hairs, stellae 0.25-0.4 mm across, lateral rays

7- 9, central ray 0.5-2 t im es as long as laterals.

Fruiting calyx with lobes less than half length

of mature fruit, prickles absent. Mature fruits

1 or 2 per inflorescence, globular, 23-25 mm

diameter, red or orange, 2-locular; placenta in

cross-section stalked, anvil-shaped; mesocarp

moist but not juicy; exocarp 4.7-5 mm thick;

pedicels 22-31 mm long in fruit, 1.1-1.8 mm

thick at mid-point; seeds pale yellow, 3-3.6 mm

long.

Specimens examined : Queensland. Cook District:

Johnstone River, Jul 1917, Michael (BRI); Boonjie, Atherton

Tableland, anno 1929, Kajewski 1255 (BRI); LakeBarrine,

Jul 1938, Goy 432 (BRI); near Mt Haig, c. 12 miles [19 km]

SEof Mareeba, May 1969, Tracey s.n. (BRI); MtBellenden

Ker, c. 1 mile [1.6 km] SE of Centre Peak, Jun 1969, Smith

14682 (BRI); Pine Creek Forestry road, Malbon Thompson

Range, Jul 1973, Webb & Tracey 13383 (BRI, QRS); SW

slopes of May Peak, ESE of Cairns, Sep 1974, Moriarty 1580

(BRI, CANB); S.F.607, West L.A., Dec 1974, Hyland 7903

(BRI);MtBartleFrere, 1.8kmWSW ofBobbin Bobbin Falls,

Nov 1988, Jessup GJM1077 etal. (BRI); S.F. 194 Herberton

Range, c. 9.5 km S of Rifle Range road turnoff, Aug 1989,

Bostock 1023 & Guymer (BRI); S.F. 185 Danbulla, Breach

F.A., Dec 1991, Gray 5361 (BRI, QRS); Smiths Track,

Barron Gorge N.P., Apr 1997, Jago 4327 (BRI); S.F. 144,

Chowchilla F.A., Sep 2000, Ford 2428 (BRI); Wooroonooran

N.P., East Mulgrave River, Nov 2000, Forster PIF26433 et

al. (A, BRI, MEF). North Kennedy District: Koolmoon

Creek, Sep 1959, Smith 10809 (BRI); ‘Bellview’, Evelyn,

Jan 1974, Collins C74-5 (BRI); Kirrama Range, S.F.344, c.

38 km NW of Kennedy, Nov 1989, Fell DF2006 (AD, BRI,

MEF); Koombooloomba S.F., near Tully Falls road, S of

Ravenshoe, Apr 2002, Bean 18717 & McDonald (BRI,

MEF).

Distribution and habitat: Solanum macoorai

is endemic to Queensland, extending from

Windsor Tableland (near Mount Carbine) to the

Kirrama Range near Cardwell (Map 22). It

occurs in notophyll rainforest or the ecotone

between eucalypt forest and rainforest, in high

rainfall areas. Altitude is commonly between

500 and 1500 metres, but it extends almost to

sea level in some places.

757

Phenology : Flowers and fruits are recorded for

almost every month of the year.

Notes: S. macoorai is closely related to

S. magnifolium, but differs by the juvenile

plants with longer and more abundant prickles;

adult leaves usually lobed and bearing prickles,

and with sparser indumentum on the lower

surface; the stellae (on all organs) with a much

shorter central ray, and stellae of the lower leaf

surface lacking a central ray; and fruits red or

orange at maturity (vs. yellow).

In the protologue, Bailey described the

fruit of S. macoorai, but no trace of this (these)

fruit(s) can now be found at BRI. The holotype

comprises 4 detached leaves mounted on a

single sheet. Fortunately the very distinctive

indumentum pattern on the leaves of S. macoorai

means that there is no doubt about the correct

application of the name.

Conservation status: Widespread. Not

considered at risk.

57. Solanum magnifolium F.Muell., Fragm.

6: 27 (1867); S. stelligerum var.

magnifolium (F.Muell.) Benth., FI.

Austral. 4:451 (1868). Type: Queensland.

North Kennedy District: Murray River

[N of Cardwell], August 1867, J.

Dallachy s.n. (holo: MEL [MEL11658]).

Solanum dallachii Benth., FI. Austral. 4: 456

(1868), syn. nov. Type: Queensland.

North Kennedy District: Rockingham

Bay, anno 1864, J. Dallachy (lecto: K;

isolecto: MEL [MEL11655, MEL11659]),

fide Symon (1981), but see discussion

below.

[Solanum repandum auct. non G. Forster: F.

Muell., Fragm. 6: 145 (1868)].

Illustration: Symon (1981: 246), as S. dallachii.

Erect, rhizomatous perennial shrub, 1.5-4 m

high. Juvenile branchlets with 30-50 prickles

per dm, 4-5 mm long; leaves (in outline) ovate,

shallowly-lobed, with 4 or 5 pairs of lobes;

lamina 9.5-11.5 cm long, 4.3-5.5 cm wide,

with 20-40 prickles on upper surface. Adult

branchlets yellow, rusty or brown; prickles

absent or present, 0-25 per decimetre, straight,

758

acicular or broad-based, 1.5-5.5 mm long,

7-10 times longer than wide; stellae dense,

0.2-0.3 mm diameter, sessile; lateral rays 7 or

8, porrect; central ray 3-6 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Adult leaves elliptical or

ovate, entire or shallowly lobed throughout;

lobes 4-6 on each side, obtuse, lobing index

1-1.1; lamina 9-18 cm long, 4-8 cm wide,

2.2-2.7 t im es longer than broad, apex acute or

acuminate, base cuneate, oblique part 0-4.5 mm

long, obliqueness index 0-3 percent; petioles

1- 3.5 cm long, 8-19% length of lamina,

prickles absent. Upper leaf surface green;

prickles 0-10, straight, acicular, 1-4 mm long,

prickles absent or present on midvein only or

present on midvein and lateral veins; stellate

hairs confined to midrib, or distributed

throughout; protostellae absent; ordinary stellae

very sparse to sparse, 0.25-1 mm apart,

0.2-0.3 mm across, sessile; lateral rays 6-8,

porrect; central ray 4-8 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Lower leaf surface yellowish;

prickles absent; stellae moderate to very dense,

0.05-0.3 mm apart, 0.25-0.4 mm diameter,

sessile; lateral rays 7 or 8, porrect; central ray

1.5- 3 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Inflorescence supra-axillary, pseudo-racemose,

common peduncle 2-8 mm long, rachis prickles

absent; 6-18-flowered, weakly andromonoecious,

flowers 5-merous; pedicels 4-10 mm long at

anthesis, same thickness throughout, 0.4-0.7

mm thick at mid-point, prickles absent. Calyx

tube 1.5-2.5 mm long, lobes deltate, 3-4 mm

long; prickles absent at anthesis; stellae very

dense, yellow or brown or rusty, 0.25-0.35 mm

across, sessile, lateral rays 7 or 8, central ray

3-6 times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Corolla

mauve or purple, 6-11 mm long, deeply lobed,

inner surface sparsely stellate-hairy; anthers

3.5- 4.5 mm long; ovary with stellate hairs only;

functional style c. 6 mm long, erect, with

stellate hairs only, stellae 0.25-0.3 mm across,

lateral rays c. 8, central ray 1.5-3 times as long

as laterals. Fruiting calyx with lobes less than

half length of mature fruit, prickles absent.

Mature fruits 1 per inflorescence, globular,

20-27 mm diameter, yellow or yellowish-green,

2- locular; placenta in cross-section stalked,

anvil-shaped; mesocarp moist but not juicy;

Austrobaileya 6 (4): 639-816 (2004)

exocarp 4.5-6 mm thick; pedicels 19-23 mm

long in fruit, 1.2-2.5 mm thick at mid-point;

seeds pale yellow, 2.7-3.1 mm long.

Specimens examined : Queensland. Cook District:

Endeavour River, Sep 1872, Hann (BRI); Upper Parrot

Creek, Annan River, Sep 1948, Brass 20283 (BRI); c. 6 miles

[10 km] SW of Yungaburra, Curtain Fig site, Dec 1968,

Tracey s.n. (BRI); between Macnamee Ck and Downey Ck,

W of Innisfail, May 1972, Webb & Tracey 10755 (BRI);

Macdowall Range between Daintree River and China Camp,

Aug 1972, Webb & Tracey 10752 (BRI); Mt Sampson Wof

The Forks on Cooktown-Bloomfield road, Jun 1973, Webb

& Tracey 10793 (BRI, CANB); T.R.55, lul 1974, Hyland

7342 (BRI); S.F.144, Fantail F.A., Mar 1981, Unwin 823

(QRS); S.F.756 Jordan, Fower Downey F.A., Nov 1991,

Hyland 14332 (BRI, QRS); Daintree N.P., Adeline Creek

headwaters, top of hill 929, May 1999, Forster PIF24543 &

Booth (BRI); T.R.176, Shipton F.A., Oct 1999, Forster

PIF25007 & Booth (BRI, QRS); Shipton’s Flat, S of

Cooktown, Apr 2002, Bean 18775 & McDonald (BRI).

North Kennedy District: Herbert River, anno 1873, Stone

(BRI); Koombooloomba on road S towards Tully River

crossing, May 1972, Webb & Tracey 10907 (BRI); Echo

Creek walking track, off North Davidson road, west of Tully,

Aug 2002, Ford AF3564 & Holmes (BRI, NSW).

Distribution and habitat : Solanum

magnifolium is endemic to Queenland. It

extends from Cooktown to S of Ravenshoe.

There are historical records (including the type)

from the Ingham and Cardwell areas (Map 24).

It inhabits rainforest margins or where

rainforest is invading adjacent eucalypt forest.

Altitude ranges from 100-900 metres.

Phenology: Flowers are recorded for May and

from August-November; mature fruits for

March and May and September-December.

Notes: Mueller’s description of S. ‘repandum’

in Fragmenta 6: 145 is based on 3 collections

made by Dallachy in 1864. One of these

specimens has a label that states the plant is

“12 feet high” and has “fruits yellow”. Mueller

repeats these statements in his description.

Another specimen has a strip of bark taken from

an older stem that bears numerous short

prickles. The leafy specimen is without prickles.

Mueller in his description, states that the

branchlets of S. repandum are on one part

conspicuously aculeate, in another part

unarmed. Bentham named S. dallachii in

December 1868, in what amounts to a nom.

nov. for S. repandum F.Muell. non GForst., and

the same specimens serve as types for S. dallachii.

Symon (1981) chose as lectotype of

S. dallachii one of the sheets at K. Furthermore,

Bean, Taxonomy of Solanum subg. Leptostemonum

he cited three MEL sheets as isolectotypes.

While the sheet MEL 11655 and its duplicate

(MEL11659) may well be isolectotypes, the

sheet MEL 11656 certainly is not as its label

clearly shows that the date and locality of

collection are different.

Mueller (1867) named Solanum

magnifolium, based on a Dallachy specimen or

specimens. The exact identity of the type

specimen(s) has until now been unclear. A

specimen collected by Dallachy from “Murray

River” in August 1867 (MEL11658), precisely

matches the details given in the protologue. In

particular, this specimen has unusually small

prickles on the branchlets (2-3 mm long), the

leaves of the specimen are very large (up to 20

x 11 cm), and there are no fruits. These

characteristics are confirmed in the protologue

{aculei 1-1.5”’ longi; foliis ad 9” longa, ad 5”

lata; and “ baccae ignotae”). The repand-

dentate leaves, calyx length and anther length

offer further confirmation that Mueller’s

description was taken from this specimen, and

this specimen alone.

Significantly, Bentham (1868) cited

Dallachy’s “Murray River” specimen when

reducing S. magnifolium to a variety of

S. stelligerum Sm., thereby confirming that this

was the specimen used by Mueller to describe

his species.

Conservation status : Although not seen in the

Ingham-Cardwell area (the type locality) for

130 years, S. magnifolium appears to be

moderately common further north, particularly

around Bloomfield. No conservation code is

recommended.

58. Solanum rixosum A.R.Bean sp. nov. Frutex

perennis, erectus; folia ovata, integra vel

non profunde lobata; aculei in ramulis

sparsi; cortex tenuis nondescriptus; pili

digitati pagina superiore folii absentes;

calycis aculei absentes; pagina interna

corollae sparse stellato-pilosa; ovarium

pilis stellatis et Type-2 praeditum; fructus

maturi virides, 19-24 mm diametro;

pedicellis fructiferis 17-26 mm longis.

Typus: Queensland. Moreton District:

0.5 km along Duck Creek road, near

O’Reillys Guest House, 11 December

1999, A.R. Bean 15899 (holo: BRI (1

sheet + spirit); iso: AD, CANB, NSW).

759

Solanum sp. 1 in Ross (1986); Solanum sp.

(Benarkin R.J. Henderson H297) in

Henderson (2002).

Illustration : Symon (1981: 248), as

S. furfuraceum.

Erect, rhizomatous perennial shrub, 1-2 m

high. Juvenile branchlets with 10-30 prickles

per dm; leaves (in outline) broadly ovate,

shallowly-lobed, with 1-3 pairs of lobes; lamina

9-15 cm long, 4-8 cm wide, with 10-40

prickles on upper surface. Adult branchlets

yellow or rusty; prickles absent or present,

0-15 per decimetre, straight, acicular, 3-10 mm

long, 10-14 t im es longer than wide; stellae very

dense, 0.3-0.4 mm diameter, stalks 0-0.2 mm

long; lateral rays 4-8, porrect; central ray 1-2

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Adult

leaves ovate, entire or shallowly lobed

throughout; lobes 1-3 on each side, obtuse,

lobing index 1-1.3; lamina 6-12 cm long,

2.9-4.8 cm wide, 1.8-2.6 times longer than

broad, apex acute, base cuneate or obtuse,

oblique part 0-4 mm long, obliqueness index

0-4 percent; petioles 0.7-1.4 cm long, 8-14%

length of lamina, prickles absent. Upper leaf

surface green; prickles 0-10, straight, acicular,

4- 10 mm long, prickles absent or present on

midvein only or present on midvein and lateral

veins; stellate hairs distributed throughout;

protostellae absent; ordinary stellae very sparse

to sparse, 0.7-1.6 mm apart, 0.4-0.8 mm

across, stalks 0-0.2 mm long; lateral rays 6-9,

porrect or ascending; central ray 1—1.5 times

as long as laterals, not gland-tipped; finger

hairs absent; Type 2 hairs absent. Lower leaf

surface yellowish; prickles absent; stellae

dense, 0.1-0.4 mm apart, 0.5-0.8 mm diameter,

stalks 0.2-0.6 mm long; lateral rays 7-9,

porrect; central ray 1-1.5 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Inflorescence supra-

axillary, pseudo-racemose, common peduncle

1-6 mm long, rachis prickles absent; 2-15-

flowered, weakly andromonoecious, flowers

5- merous; pedicels 6-11 mm long at anthesis,

same thickness throughout, 0.5-0.8 mm thick

at mid-point, prickles absent. Calyx tube

1.5-3 mm long, lobes attenuate, 7-9 mm long;

prickles absent at anthesis; stellae very dense,

yellow or brown or rusty, 0.5-0.6 mm across,

stalks 0-0.2 mm long, lateral rays 8-9, central

760

Austrobaileya 6 (4): 639-816 (2004)

QUEENSLAND HERBARIUM (BRI)

Fiona of Quaensland

Meiaton

QUEENSLAND HERBARIUM (BRI)

&»&•»• Australia

SoJSnuiTj

Cot A.H.B&in 15839

2fl*1ZS 153WE

R7D1 22

O.SJtm along Duck Creek Road, near O'fiddtiis Goesl Housa.

(GPS 2ft 12 50 153 Iff 17),

Ndophyll ra'mtoml with ArgyrOdentden. Bwchychifon

aeenlalius. Pefflacaras.

Shrub to im high, prickles sparse; baft with lenlwels but not

corky, flowers latge. puipta: Injit graan, net quite rnalure.

Occasional at site.

Spirit material at BRI.

□<".:r,

280 SolBnaoeaa

/fa-oTy/g- ° UMl1s * an <l Haibarium

rikosttm. 4. 4.

Om s'** '? OiHjt0O-2ae3

Cat.

Oups. NSW CANS AD

+ Ktpy ibfl cited as eornputcrised coilGCtipn number AO 679122

{tonne! Piped

Fig. 34. Holotype of Solanum rixosum.

Bean, Taxonomy of Solanum subg. Leptostemonum

ray 1-1.5 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Corolla purple, 9-14 mm long, shallowly lobed,

inner surface sparsely stellate-hairy; anthers

5-6 mm long; ovary with stellate and Type 2

hairs; functional style 7.5-10 mm long, erect,

glabrous or with Type 2 hairs only or with

stellate and Type 2 hairs, stellae 0.4-0.5 mm

across, lateral rays 6-9, central ray 1-1.5 times

as long as laterals. Fruiting calyx with lobes

less than half length of mature fruit, prickles

absent. Mature fruits 1-3 per inflorescence,

globular, 19-24 mm diameter, yellowish-green

or green, 2 or 3-locular; placenta in cross-

section stalked, anvil-shaped; mesocarp moist

but not juicy; exocarp 0.9-1.2 mm thick;

pedicels 17-26 mm long in fruit, 1-1.4 mm

thick at mid-point; seeds pale yellow, 2.2-2.7

mm long. Fig. 34.

Specimens examined : Queensland. Burnett District:

BunyaMtns, Oct 1919, White s.n. (BRI); Bunya Mountains,

Oct 1958, Mitchener (BRI); 2.5 km E of Mt Mowbullan,

Bunya Mountains, Sep 1988, Forster PIF5795 & Bird (AD,

BRI, MEL). Wide Bay District: Blackall Range, May 1910,

Keys (BRI); Blackall Range, Dec 1916, White s.n. (BRI);

Conondale Range (S.F.274 Conondale), BooloumbaL.A., Oct

1982, McDonald 3575 & Williams (BRI, MEL, NSW); Mt

Glastonbury, Sep 1986, Sharpe 4515 & Bean (BRI); Peters

L.A., Conondale Ranges, Nov 1990, Bean 2694 (BRI).

Darling Downs District: Cunninghams Gap, undated, Bailey

(BRI); Killamey, Oct 1891, coll, unknown (BRI); Westcott,

Bunya Mountains, Feb 1995, Fairfax 61 (BRI); 800 m S of

Cherry Plain picnic area [Bunya Mountains], Nov 1996,

Holland 1147 (BRI). Moreton District: Peecheys

[Ashgrove, Brisbane], Aug 1888, Simmonds (BRI);

Rosewood, May 1918, White s.n. (BRI);TamborineMtn, Dec

1921, White 1789 (BRI); Blackbutt Range near Moore, May

1967, Griffin (BRI); Benarkin S.F., c. 15 miles [24km] E of

Yarraman, May 1967, Henderson H253 (BRI); Levers

Plateau, on Qld/N.S.W. border, c. 90 km SSW of Brisbane,

Apr 1972, Henderson H1284 (BRI); D’Aguilar Range, NW

of Brisbane, Apr 1972, Moriarty 906 (BRI); Ravensboume,

N of Toowoomba, Nov 1985, Swarbrick8331 (BRI); S.F.809

Laceys Creek, D’Aguilar Range, Oct 1993, Forster

PIF13992 (AD, BRI, K, MEL, QRS); 3.2 km along Duck

Creek road, near O’Reillys Guest House, Jan 2000, Bean

15970 (BRI, NSW). New South Wales. North Coast:

Toonumbar S.F., c. 26 km NW of Kyogle, Feb 1972,

Henderson HI266, HI267 & Parham (BRI); Eden Creek

Falls, Toonumbar S.F., Sep 1972, Coveny 4410 (BRI, NSW);

lower slope of Mt Lindesay, 7 miles [11 km] ENE of

Woodenbong, Sep 1972, Coveny 4542 & Rodd (BRI, NSW).

Distribution and habitat: Solanum rixosum has

a rather limited extent, from the Bunya

Mountains and Mt Glastonbury in Queensland

to the Kyogle district of far north-eastern New

South Wales (Map 21). It grows in disturbed

761

areas of notophyll rainforest, or in shrubby

eucalypt forest close to rainforest.

Phenology : Flowers are recorded for April and

May and from August-December; mature fruits

from November-May.

Notes: Solanum rixosum is related to S.jurfuraceum,

but differs by the thin, non-descript bark; the

stellae of the upper leaf surface very sparse to

sparse; the fruiting pedicels 17-26 mm long

(vs. 4-8 mm long for S. furfuraceum)\ the ovary

with both stellate and Type 2 hairs; the calyx without

Type 2 hairs, and the fruiting exocarp 0.9-1.2 mm

thick (2-2.6 mm thick for S. jurfuraceum).

S. rixosum differs from S. magnifolium

by the larger stellae with a relatively shorter

central ray, stellae stalked on the lower leaf

surface, adult leaves (when lobed) with only

1-3 pairs of lobes, the thinner exocarp of the

fruit and the broadly ovate juvenile leaves.

Conservation status: Still moderately common

in some areas, and not currently considered at

risk.

Etymology: From the Latin rixosus meaning

quarrelsome. This refers to the differences of

opinion of various botanists regarding the

taxonomic status of this species.

59. Solanum serpens A.R.Bean, Austrobaileya

6: 248 (2002). Type: Queensland.

Moreton District: Cainbable Creek

track, Lamington National Park, 11

December 1999, A.R. Bean 15903 (holo:

BRI; iso: CANB, MEL, NSW).

Solanum stelligerum var. procumbens

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

55: 72 (1944). Type: Queensland.

Moreton District: Lamington National

Park, 27 November 1942, C.T. White

11889 (holo: BRI, 2 sheets).

Illustration: Bean (2002b: 249)

Prostrate, stoloniferous perennial shrub, 0-0.3

m high. Juvenile stage absent. Adult branchlets

grey or rusty; prickles 15-35 per decimetre,

straight, acicular, 3-6 mm long, 10-14 times

longer than wide; stellae dense, 0.25-0.35 mm

diameter, sessile; lateral rays 7 or 8, porrect;

central ray 4-9 times as long as laterals, not

762

gland-tipped; finger hairs absent; Type 2 hairs

absent. Adult leaves elliptical, ovate or broadly

ovate, entire; lamina 5.5-10 cm long, 2.5-4

cm wide, 1.4-2.9 times longer than broad, apex

obtuse or acute, base cuneate or obtuse, oblique

part 0-3 mm long, obliqueness index 0-4

percent; petioles 0.6-1.5 cm long, 10-19%

length of lamina, prickles present. Upper leaf

surface green; prickles 0-8, straight, acicular,

2-4 mm long, prickles absent or present on

midvein only or present on midvein and lateral

veins; stellate hairs confined to midrib, or

distributed throughout; protostellae absent;

ordinary stellae very sparse to sparse, 0.6-3 mm

apart, 0.2-0.5 mm across, sessile; lateral rays

7 or 8, porrect; central ray 3-6 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Lower leaf surface white,

yellowish or rusty; prickles absent; stellae

moderate to very dense, 0.05-0.3 mm apart,

0.3-0.4 mm diameter, sessile; lateral rays 7 or

8, porrect; central ray 3-5 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Inflorescence supra-

axillary, solitary or pseudo-racemose, common

peduncle 0-3 mm long, rachis prickles absent;

1- 4-flowered, weakly andromonoecious,

flowers 5-merous; pedicels 11-28 mm long at

anthesis, same thickness throughout, 0.6-0.8

mm thick at mid-point, prickles absent or

present. Calyx tube 1.5-2.5 mm long, lobes

attenuate, 4-7 mm long; prickles absent at

anthesis; stellae dense, transparent, 0.3-0.4 mm

across, sessile, lateral rays 7 or 8, central ray

2- 5 times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Corolla

purple, 10-13 mm long, shallowly or deeply

lobed, inner surface glabrous; anthers 3.5-5.5

mm long; ovary with stellate and Type 2 hairs;

functional style 7.5-8 mm long, erect, with

stellate hairs only, stellae 0.5-0.7 mm across,

lateral rays 8-9, central ray 1-2 times as long

as laterals. Fruiting calyx with lobes less than

half length of mature fruit, prickles absent.

Mature fruits 1 or 2 per inflorescence, globular,

14-16 mm diameter, yellowish-green or green,

1-locular (septum absent or incomplete);

mesocarp moist but not juicy; exocarp 0.6-0.8

mm thick; pedicels 20-32 mm long in fruit,

0.6-0.8 mm thick at mid-point; seeds pale

yellow, 3.5-4 mm long.

Specimens examined : Queensland. Moreton District:

Tamborine Mtn, anno 1888, Simmonds 346 (BRI); Little

Austrobaileya 6 (4): 639-816 (2004)

Nerang River, Jan 1916, White s.n. (BRI); Lamington N.R,

Nov 1942, White 11889 (BRI); Cainbable Ck track,

Lamington N.R, c. 1 km SSE of Cainbable Falls, Nov 1995,

McDonald 6176 (BRI); Nicholl’s scrub, c. 6 km SW of

Currumbin, Mar 2001, Bean 17398 (BRI); Darlington

Range, Upper Ormeau, Feb 2002, Bean 18521 & Leiper

(BRI); Rosins Lookout, Beechmont, Dec 2002, Bean 19681

(BRI). New South Wales. North Coast: Three Mile scrub,

near Byron Bay, Nov 1898, Forsyth (NSW); Byron Bay, Nov

1903, Maiden & Boorman s.n. (NSW); Levers Plateau on

Qld-N.S.W. border, c. 90 km SSW of Brisbane, Apr 1972,

Henderson H1299 (BRI); Brunswick Heads Nature Reserve,

Oct 2000, Bean 16929 (BRI).

Distribution and habitat : Solanum serpens is

found from Mt Tamborine in Queensland to

Byron Bay In N.S.W. (Map 23), although the

Byron Bay population is probably extinct. It

inhabits notophyll rainforest, especially where

there are canopy gaps.

Phenology : Flowers are recorded between

October to April; fruits in March and April.

Notes: S. serpens is rather variable in morphology,

and intermediate forms with S. acanthodapis exist

(Bean 2002b).

Conservation status : It is known from several

locations, some of which are on protected land.

No conservation status is recommended at this

time.

60. Solanum acanthodapis A.R.Bean,

Austrobaileya 6: 250 (2002). Type: New

South Wales. North Coast: Big Scrub

Flora Reserve, c. 20 km N of Lismore, 2

December 2000, A.R. Bean 17075 (holo:

BRI; iso: AD, K, L, MEL, NSW).

Illustration: Bean (2002b: 251)

Prostrate, stoloniferous perennial shrub, 0-0.3

m high. Juvenile stage absent. Adult branchlets

terete or ridged, brown; prickles 25-65 per

decimetre, straight, acicular, 3-7 mm long,

9-20 times longer than wide; stellae sparse to

dense, 0.2-0.35 mm diameter, sessile; lateral

rays 7 or 8, porrect; central ray 0.5-1.5 times

as long as laterals, not gland-tipped; finger

hairs absent; Type 2 hairs absent. Adult leaves

elliptical or ovate, entire or shallowly lobed

throughout; lobes 2-4 on each side, acute,

lobing index 1-1.4; lamina 5.5-9.5 cm long,

2.8-4.5 cm wide, 1.6-2.3 times longer than

broad, apex acute, base cuneate or obtuse,

oblique part 0-5 mm long, obliqueness index

Bean, Taxonomy of Solanum subg. Leptostemonum

0-5 percent; petioles 1-1.7 cm long, 14-29%

length of lamina, prickles present. Upper leaf

surface green; prickles 20-60, straight,

acicular, 2-8 mm long, prickles present on

midvein and lateral veins; stellate hairs

confined to midrib; ordinary stellae absent from

surface; finger hairs absent; Type 2 hairs absent.

Lower leaf surface green to white or yellowish;

prickles 10-30, straight, acicular, present on

midvein and lateral veins; stellae sparse to very

dense, 0.05-0.25 mm apart, 0.25-0.35 mm

diameter, sessile; lateral rays 7 or 8, porrect;

central ray 0.2-0.8 t im es as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Inflorescence supra-axillary,

pseudo-umbellate or pseudo-racemose,

common peduncle 0-7 mm long, rachis prickles

absent or present; 2-8-flowered, strongly or

weakly andromonoecious, flowers 5-merous;

pedicels 6-20 mm long at anthesis, same

thickness throughout, 0.25-0.4 mm thick at

mid-point, prickles absent or present. Calyx

tube 2-3 mm long, lobes attenuate, 1.5-3.5 mm

long; prickles absent or present at anthesis,

0-2 per flower, 0.5-1 mm long; stellae

moderate, transparent or purple, 0.2-0.3 mm

across, sessile, lateral rays 7 or 8, central ray

0.5-1 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Corolla mauve or purple, 8-12 mm long,

shallowly lobed, inner surface glabrous; anthers

4-5 mm long; ovary with Type 2 hairs only;

functional style 6-7.5 mm long, erect, glabrous

or with Type 2 hairs only. Fruiting calyx with

lobes less than half length of mature fruit,

prickles absent. Mature fruits 1 per

inflorescence, globular, 15-18 mm diameter,

yellowish-green, 1-locular (septum absent or

incomplete); placenta in cross-section sessile,

elliptical; mesocarp moist but not juicy; exocarp

c. 0.8 mm thick; pedicels 21-30 mm long in

fruit, c. 0.8 mm thick at mid-point; seeds pale

yellow, 3.5-4 mm long.

Specimens examined : New South Wales. North Coast:

Victoria Park, 4 mi les [6 km] SSW of Alston ville, Aug 1969,

Clark 1260 et al. (BRI, NSW); Rotary Park, Lismore, Dec

2000, Bean 17081 (BRI); Booyong Flora Reserve, ENE of

Lismore, Dec 2000, Bean 17083 (BRI); Victoria Park Nature

Reserve, Apr 2001, Bean 17564 (AD, BRI, MEL, NSW).

Distribution and habitat : Solanum

acanthodapis is endemic to the extreme north¬

east of New South Wales, around Lismore

763

(Map 22). It grows in canopy-gaps in

notophyll rainforest.

Phenology: Flowers recorded between August

and February; fruits in March and April.

Notes : Well-developed plants of this species

form a dense prickly carpet, no more than 20

centimetres high. The fruits are borne virtually

at ground level, and hidden from view by the

foliage. I have not observed any clues that

would indicate the method of seed dispersal.

Conservation status: Applying the IUCN

guidelines (IUCN. 2001), a category of

“Vulnerable” is recommended (VU C2a(i)).

61. Solanum intonsum A.R.Bean sp. nov.

Frutex perennis erectus; aculei

ramulorum recti, ad basim lati, sparse

distributi vel saepe absentes; folia adulta

integra, lanceolata usque ovata; pagina

superior folii proto Stellas gerens; stellae

vulgatae sessiles, latitudine 0.15-0.25

mm, parcae usque densae; aculei calycis

absentes; ovarium pilos type-2 tantum

gerens; fructus 15-17 mm diametro,

virides maturitate; pedicelli fructiferi 7-

10 mm longi. Typus: Queensland. Cook

District: Smiths track, Barron Gorge

National Park, 6 May 2000, A.R. Bean

16542 (holo: BRI (1 sheet + spirit); iso:

MEL, NSW).

Solanum sp. (Font Hill s J.R. ClarksonE 7901)

in Henderson (2002).

Erect, rhizomatous perennial shrub, 0.8-2 m

high. Juvenile branchlets with 30-50 prickles

per dm, 1-7 mm long; leaves (in outline) ovate,

entire or shallowly-lobed, with 2 or 3 pairs of

lobes; lamina 9.5-14 cm long, 5.5-6 cm wide,

with 0-7 prickles on upper surface. Adult

branchlets yellow; prickles absent or present,

0-5 per decimetre, straight, broad-based,

0.5-6 mm long, 4-7 times longer than wide;

stellae dense to very dense, 0.3-0.5 mm

diameter, stalks 0-0.2 mm long; lateral rays

6-8, porrect; central ray 0.3-0.7 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Adult leaves

lanceolate or ovate, entire; lamina 4.5-16.5 cm

long, 1.3-6.5 cm wide, 2.1-4.5 times longer

than broad, apex acute, base cuneate or obtuse,

oblique part 0-5 mm long, obliqueness index

0-6 percent; petioles 0.3-2.5 cm long, 7-30%

764

length of lamina, prickles absent. Upper leaf

surface green or grey-green; prickles absent;

stellate hairs distributed throughout;

protostellae present; ordinary stellae density

moderate to dense, 0.1-0.2 mm apart,

0.15-0.25 mm across, sessile; lateral rays

4-10, ascending; central ray 0.5-1 times as

long as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Lower leaf surface

white; prickles absent; stellae dense to very

dense, 0.05-0.1 mm apart, 0.4-0.6 mm

diameter, stalks 0-0.2 mm long; lateral rays 7

or 8, porrect; central ray 0.2-0.6 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Inflorescence

supra-axillary, pseudo-umbellate or pseudo-

racemose, common peduncle 0-5 mm long,

rachis prickles absent; 2-7-flowered, weakly

andromonoecious, flowers 5-merous; pedicels

4-9 mm long at anthesis, same thickness

throughout, 0.5-0.8 mm thick at mid-point,

prickles absent. Calyx tube 3-4.5 mm long,

lobes attenuate, 1-5 mm long; prickles absent

at anthesis; stellae dense to very dense, yellow,

0.25-0.4 mm across, stalks 0-0.1 mm long,

lateral rays 7 or 8, central ray 0.5-1 times as

long as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Corolla purple,

9-13 mm long, shallowly lobed, inner surface

sparsely stellate-hairy; anthers 4.5-5.5 mm

long; ovary with Type 2 hairs only; functional

style 8-9 mm long, erect, with stellate hairs

only or with Type 2 hairs only, stellae c. 0.3

mm across, lateral rays 5-6, central ray 0.5-1

times as long as laterals. Fruiting calyx with

lobes less than half length of mature fruit,

prickles absent. Mature fruits 1 or 2 per

inflorescence, globular, 15-17 mm diameter,

green, 2-locular; placenta in cross-section

sessile, linear; mesocarp moist but not juicy;

exocarp c. 2 mm thick; pedicels 7-10 mm long

in fruit, 0.8-1 mm thick at mid-point; seeds

pale yellow, 2.3-2.8 mm long. Fig. 35.

Specimens examined : Queensland. Cook District: sources

of Stuart’s River [Stewart River near Coen], 1891, Johnson

s.n. (MEL); Hartley’s Creek, Cook Highway, Jul 1936,

Flecker 1972,1986 (BRI); Mt Carbine, Nov 1967, Cassels

(BRI, QRS); S.F.700, Mar 1968, Hyland 4068 (BRI, QRS);

Flinders Island, c. 7 km N of Bathurst Head, Jun 1978,

Clarkson 2215 (BRI); Earl Hill near Cairns, Oct 1979,

Batianoff 1292 & McDonald (AD, BRI); Richards Creek,

Mount Mulligan, Apr 1984, Clarkson 5268 (AD, BRI, QRS);

‘Font Hills’, c. 15 km W of Mount Molloy, Apr 1989,

Austrobaileya 6 (4): 639-816 (2004)

Clarkson 7901 & Henderson (AD, BRI, DNA); Mt Alto, c.

4 km SSW of Mt Carbine, Apr 1989, Clarkson 7939 &

Henderson (BRI); Cape Melville N.R, 2 km SSE of Temple

Hill, May 1993, Fell DGF3172 & Stanton (BRI, CANB);

S.F.144, Chowchilla L.A., Mar 1994 , Hyland 15058 (QRS);

Bakers Blue Mtn, Apr 1995, Hyland 15294 (QRS); Smiths

Track, Barron Gorge N.R, Mar 1997, Jago 4284 (BRI).

Distribution and habitat : Solanum intonsum

is endemic to Queensland. Currently known to

extend from Cape Melville to just south of

Cairns, and west to Mt Mulligan (Map 25).

There is also an historical record from the

Stewart River near Coen. It grows on hilly to

mountainous terrain, in shrubby eucalypt

woodland or on the margins of rainforest or

vine thicket.

Phenology : Flowers are recorded from March

to November; mature fruits in October and

November.

Notes: S. intonsum is related to S. magnifolium,

but differs by the upper leaf surface with

protostellae and moderate to dense ordinary

stellae; the central ray of the stellate hairs much

shorter on all plant parts; the longer

hypanthium and attenuate calyx lobes; the

smaller fruits, green at maturity and on shorter

pedicels and the ovary with Type 2 hairs only.

Conservation status : Moderately widespread.

Not considered at risk.

Etymology: From the Latin in- meaning not,

and tonsus meaning shaven, referring to the

dense but uniform indumentum of the leaves.

62. Solanum furfuraceum R.Br., Prodr. 446

(1810). Type: [Queensland. Port Curtis

District:] “Broadsound”, September 1802,

R. Brown (lecto: BM (Bennett No. 2672);

isolecto: MEL [MEL11620]), fide Symon

(1981).

Erect, rhizomatous perennial shrub, 1-3 m

high. Juvenile branchlets with 60-90 prickles

per dm; leaves (in outline) ovate, entire or

shallowly-lobed, with 2 or 3 pairs of lobes;

lamina 6-8 cm long, 2.5-4 cm wide, with

2-10 prickles on upper surface. Adult stems

bark furrowed, corky. Adult branchlets brown;

prickles absent or present, 0-15 per decimetre,

straight, acicular, 4-10 mm long, 9-12 times

longer than wide; stellae dense, 0.4-0.9 mm

Bean, Taxonomy of Solanum subg. Leptostemonum

765

Fig. 35. Solanum intonsum. A. flowering branchlet x 0.8. B. ovary and style x 6. C. ovary, bearing a few Type 2 hairs x 12.

D. stellate hair, upper leaf surface x 100. E. mature fruit and calyx x 2. F. transverse section of fruit x 3. all from Bean 16542.

766

Austrobaileya 6 (4): 639-816 (2004)

diameter, stalks 0.1-0.6 mm long; lateral rays

4-8, porrect; central ray 1-1.5 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Adult leaves ovate, entire;

lamina 3-11.5 cm long, 1.1-4.5 cm wide,

2.1-2.7 times longer than broad, apex acute,

base cuneate or obtuse, oblique part 0-7 mm

long, obliqueness index 0-7 percent; petioles

0.5-1.3 cm long, 8-18% length of lamina,

prickles absent. Upper leaf surface green;

prickles 0-2, straight, acicular, 5-7 mm long,

prickles absent or present on midvein only;

stellate hairs distributed throughout;

protostellae present; ordinary stellae density

sparse to moderate, 0.4-0.7 mm apart, 0.4-0.8

mm across, stalks 0-0.3 mm long; lateral rays

4- 7, porrect to ascending; central ray 1-2 times

as long as laterals, not gland-tipped; finger

hairs present, 0.1-0.3 mm apart, not gland-

tipped, 0.1-0.3 mm long; Type 2 hairs absent.

Lower leaf surface yellowish or rusty; prickles

absent; stellae dense, 0.1-0.3 mm apart, 0.6-1

mm diameter, stalks 0.1-0.7 mm long; lateral

rays 4-8, porrect; central ray 1-2 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Inflorescence leaf-

opposed, pseudo-racemose, common peduncle

9-12 mm long, rachis prickles absent; 5-8-

flowered, weakly andromonoecious, flowers

5- merous; pedicels 2-8 mm long at anthesis,

same thickness throughout, 0.5-0.7 mm thick

at mid-point, prickles absent. Calyx tube

1.5- 3.5 mm long, lobes deltate or attenuate,

2-5 mm long; prickles absent at anthesis;

stellae dense, transparent, 0.5-0.8 mm across,

stalks 0-0.2 mm long, lateral rays 4-8, central

ray 1-1.5 times as long as laterals, not gland-

tipped or gland-tipped; finger hairs absent;

Type 2 hairs present. Corolla mauve or purple,

9-17 mm long, shallowly lobed, inner surface

glabrous or sparsely stellate-hairy; anthers

4.5- 7 mm long; ovary glabrous; functional style

8.5- 10 mm long, erect, glabrous or with stellate

hairs only, stellae c. 0.4 mm across, lateral rays

c. 4, central ray c. 2 times as long as laterals.

Fruiting calyx with lobes less than half length

of mature fruit, prickles absent. Mature fruits

1- 3 per inflorescence, oblate or globular,

15-21 mm diameter, yellowish-green or green,

2- locular; placenta in cross-section stalked,

anvil-shaped; mesocarp moist but not juicy;

exocarp 2-2.6 mm thick; pedicels 4-8 mm long

in fruit, 0.6-0.8 mm thick at mid-point; seeds

pale yellow, 2.5-2.6 mm long. Corky

Nightshade. Fig. 5.

Specimens examined : Queensland. North Kennedy

District: Wild Horse Mtn, W of Townsville, May 1996,

Cumming 14644 (BRI). South Kennedy District: ‘Havilah’,

Dec 1992, Fensham 965 (BRI). Leichhardt District:

Rosedale, near Baralaba, Sep 1959, Johnson 919 (BRI);

Dipperu N.R, Sep 1971, McDonald 116 (BRI); Isla Gorge,

c. 28 km SW of Theodore, Aug 1973, Sharpe 645 &

Hockings (BRI); near Yatton Ck, c. 93 km from Marlborough,

Sep l913,Moriarty 1467 (BRI); ‘Clifton’, northern Boomer

Range, Feb 1993, FenshamlYl (BRI); Auburn Range S.F.,

c. 50 km S of Thangool, Jun 1996, Bean 10339 (BRL MEL,

NSW). Port Curtis District: Marmor, near Rockhampton,

Mar 1920, Francis s.n. (BRI); Callide valley, Apr 1937, White

10773 (BRI); Ogmore, Sep 1943, Blake 15311 (BRI); Jim

Crow Mtn, Rockhampton district, Apr 1963, McKee 10272

(BRI); Dan Dan Scrub, S.F.53,20 km SW of Calliope, Nov

1987, Gibson 1116 (AD, BRI); c. 37 km WSW of Ridgelands,

Fitzroy Shire, Apr 1990, Anderson 4846 (BRI); 15 km NE

of Biloela, 3 km N of Callide Dam, Jul 1992, Thompson

BIL24 (BRI); E boundary of Triangle Creek, Taunton N.P.,

Oct 1995, Melzer RM632 (BRI); S.F.114, 14 km from top

ridge near Fairview HS, Dec 1998, Batianoff 98125 et al.

(BRI); western edge of S.F.69, SE of Biloela, Dec 1999, Bean

15928 (BRI, MEL, MO). MARANOA DISTRICT: c. 20

mi les [32 km] W of Mitchell, Mar 1950, Everist s.n. (BRI,

CANB). Burnett District: Coalstoun crater, Nov 1970, Bell

263 (BRI); Mt Blandy, Burnett River, Dec 1981, Forster

1051 (BRI); c. 15 km NW of Abercom, near S edge of

Coominglah S.F., Jan 2003, Bean 19727 (BRI).

Distribution and habitat : Solanum

furfuraceum is endemic to Queensland. In

subcoastal areas extending from west of

Townsville to the Gayndah area, and with a

disjunct occurrence west of Mitchell (Map 21).

Most records come from within 200 km of

Rockhampton. It grows in vine thicket,

communities dominated by brigalow (Acacia

harpophylla ) or bottle trees ( Brachychiton

rupestris ), or in nearby shrubby eucalypt

woodland. Soils are moderately to very fertile.

Phenology: Flowers are recorded for most

months of the year; mature fruits from March-

June and September-December.

Notes: Solanumfurfuraceum and S', sporadotrichum

are notable for their corky bark, very

conspicuous on adult plants, but usually visible

even on herbarium specimens. S. furfuraceum

forms a handsome shrub with its yellowish bark

and often abundant purple flowers.

Conservation status: Widespread. Not

considered at risk.

Bean, Taxonomy of Solanum subg. Leptostemonum

63. Solanum sporadotrichum F.Muell., Chem.

& Druggist (Melb. Chemist) Oct. 1882

p. 48 (1882). Type: Queensland. North

Kennedy District: Mount Dryander,

undated, Kilner & E. Fitzalan (lecto:

MEL [MEL12282]), fide Symon (1981).

Erect, rhizomatous perennial shrub, 1.5-4 m

high. Juvenile branchlets with c. 130 prickles

per dm, 1-5 mm long; leaves (in outline)

broadly ovate, shallowly-lobed, with 3-5 pairs

of lobes; lamina 6.5-8.5 cm long, 5-6 cm wide,

with 80-120 prickles on upper surface. Bark

on adult stems furrowed, corky. Adult

branchlets brown; prickles absent or present,

0-3 per decimetre, straight, acicular, 4-7 mm

long, 10-12 times longer than wide; stellae

sparse, 0.3-0.6 mm diameter, sessile; lateral

rays 5-8, porrect; central ray 0.3-0.7 times as

long as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Adult leaves ovate,

shallowly lobed throughout; lobes 3 or 4 on each

side, obtuse, lobing index 1.1-1.4; lamina

6.5-11 cm long, 3.5-5.5 cm wide, 1.6-2.1 t im es

longer than broad, apex obtuse or acute, base

obtuse, oblique part 0-6 mm long, obliqueness

index 0-8 percent; petioles 1-2.2 cm long,

13-25% length of lamina, prickles absent.

Upper leaf surface green; prickles 0-30,

straight, acicular, 1-6 mm long, prickles absent

or present on midvein only or present on

midvein and lateral veins; stellate hairs

distributed throughout; protostellae present;

ordinary stellae sparse, 0.9-1.6 mm apart,

0.4-0.5 mm across, sessile; lateral rays 4-8,

porrect; central ray 0.5-1 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Lower leaf surface green;

prickles 0-15, straight, acicular, absent or

present on midvein only or present on midvein

and lateral veins; stellae moderate to dense,

0.4-0.8 mm apart, 0.5-0.9 mm diameter, stalks

0-0.1 mm long; lateral rays 6-8, porrect;

central ray 0.3-0.7 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Inflorescence supra-axillary,

pseudo-racemose, common peduncle 11-16

mm long, rachis prickles absent; 3-7-flowered,

weakly andromonoecious, flowers 5-merous;

pedicels 5-10 mm long at anthesis, same

thickness throughout, 0.5-0.8 mm thick at mid¬

point, prickles absent. Calyx tube 2-3 mm long,

lobes deltate or attenuate, 2.5-7.5 mm long;

767

prickles absent at anthesis; stellae moderate to

dense, transparent, 0.3-0.5 mm across, sessile,

lateral rays 5-7, central ray 0.7-1.5 times as

long as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Corolla purple,

7-12 mm long, shallowly or deeply lobed, inner

surface glabrous; anthers 4.5-7.5 mm long;

ovary glabrous; functional style 10-11.5 mm

long, erect, glabrous. Fruiting calyx with lobes

less than or more than half length of mature

fruit, prickles absent. Mature fruits 1 or 2 per

inflorescence, globular, c. 13 mm diameter,

green, 1-locular (septum absent or incomplete);

placenta in cross-section sessile, semi-circular;

mesocarp moist but not juicy; exocarp 2.3-2.5

mm thick; pedicels 6-13 mm long in fruit,

0.5-0.8 mm thick at mid-point; seeds pale

yellow, c. 2.9 mm long.

Specimens examined : Queensland. North Kennedy

District: near crest of ridge leading to easterly peak of Mt

Dryander, Jul 1974, Henderson H2213 et al. (BRI);

Magnetic Island, near top of Mt Cook, Aug 1982, Sandercoe

904 (BRI); Mingela Bluff, Aug 1989, Camming 9300 (BRI);

‘Fanning River’, 25 km NW of Mingela, Sep 1989, Fell 64

& Cumming (BRI); Mount Wickham, Jul 1993, Fensham

991 (BRI); Leichhardt Range, Jul 1993, Fensham 992 (BRI);

Squally Bay, Gloucester Island, Apr 1994, Batianoff 94941%

& Figg (BRI); East Coast bay, S of Mt Sunter, Conway N.R,

May 1994, Batianoff 940579 & Dillewaard (BRI); Flame

Tree Hill, c. 3 kmESE of Airlie Beach, Jun 1994, McDonald

5874 & Champion (BRI); headland W of Horseshoe Bay,

Magnetic Island, Jan 1998, Cumming 16758 (BRI). South

Kennedy District: Calder Island, May 1992, Halford Q1321

& Crombie (AD, BRI); Calder Island, c. 50 km NE of

Mackay, Jun 2000, Bean 16693 & Champion (BRI, DNA,

MEL, NSW).

Distribution and habitat : Solanum

sporadotrichum is endemic to Queensland

ranging from west of Townsville to Airlie

Beach, and on Calder Island (Map 20). It grows

in semi-evergreen vine thicket, margins of

littoral notophyll rainforest and (at Airlie Beach

and Mt Dryander) in well-developed

Argyrodendron- dominated rainforest.

Phenology : Flowers are recorded for January,

May, June, July and September; mature fruits

in May, June and August.

Notes: Closely related to S. furfuraceum (both

have conspicuously corky bark), but differing

by the lobed or angled leaves, the very sparse

to sparse stellate indumentum and absence of

finger hairs on the upper leaf surface, and the

(usually) prickly adult leaves.

768

Conservation status: S. sporadotrichum is

listed as “Rare” under the Queensland Nature

Conservation Act, 1992. The species is more

widespread than was known in 1992, and I do

not consider it to be currently at risk.

64. Solanum francisii A.R.Bean sp. nov.

Frutex perennis altus foliis atrovirentibus

non profunde lobatis; ramuli porcati,

aculeis numerosis et stellis 0.15-0.25 mm

diametro; pagina superior folii aculeis

50-90 praedita; pedicelli floriferi 7-13

mm longi sub anthesi; aculei calycis

4-11 vel interdum absentes; pagina

interna corollae sparse stellato-pilosa;

fructus maturi virides, globulares, 14-22

mm diametro. Typus: Queensland. South

Kennedy District: Eungella, 2 September

1938, C.T. White 12972 (holo: BRI; iso:

CANB, MEL, distribuendi ).

Erect, rhizomatous perennial shrub, 2.5-4 m

high. Juvenile stage unknown. Adult branchlets

ridged, brown; prickles 10-140 per decimetre,

straight, acicular or broad-based, 4-10 mm

long, 5-11 times longer than wide; stellae

sparse, 0.15-0.25 mm diameter, sessile; lateral

rays 5-7, porrect; central ray absent or present,

0-0.5 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Adult leaves ovate, shallowly lobed throughout;

lobes 3-5 on each side, acute, lobing index

1.1-1.4; lamina 13-21 cm long, 6.5-10 cm

wide, 1.9-2.4 times longer than broad, apex

acute, base obtuse or cordate, oblique part 2-6

mm long, obliqueness index 1-3 percent;

petioles 1.6-3.5 cm long, 11-18% length of

lamina, prickles absent or present. Upper leaf

surface green; prickles 50-90, straight,

acicular, 4-8 mm long, prickles present on

midvein and lateral veins; stellate hairs

distributed throughout; protostellae absent;

ordinary stellae very sparse to sparse, 0.4-3 mm

apart, 0.2-0.3 mm across, sessile; lateral rays

7-9, porrect; central ray 0.2-0.7 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Lower leaf surface

green; prickles 5-20, straight, acicular, present

on midvein and lateral veins; stellae moderate,

0.3-0.8 mm apart, 0.2-0.3 mm diameter,

sessile; lateral rays 7-9, porrect; central ray

0-0.5 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Austrobaileya 6 (4): 639-816 (2004)

Inflorescence supra-axillary, pseudo-racemose,

common peduncle 1-22 mm long, rachis

prickles absent; 9-14-flowered, with all flowers

bisexual and 5(6)-merous; pedicels 7-13 mm

long at anthesis, same thickness throughout,

0.5-0.7 mm thick at mid-point, prickles absent

or present. Calyx tube 3.5-4.5 mm long, lobes

attenuate, 1.5-7 mm long; prickles absent or

present at anthesis, 0-11 per flower, 3-6 mm

long; stellae sparse to moderate, transparent,

0.2-0.3 mm across, sessile, lateral rays 7 or 8,

central ray 0.3-1 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

present. Corolla white or mauve, 12-17 mm

long, shallowly or deeply lobed, inner surface

sparsely stellate-hairy; anthers c. 5.8 mm long;

ovary with Type 2 hairs only; functional style

c. 10 mm long, erect, with Type 2 hairs only.

Fruiting calyx with lobes less than half length

of mature fruit, prickles 4-8 mm long. Mature

fruits 1-8 per inflorescence, globular, 14-22

mm diameter, yellowish-green or green, 2-

locular; placenta in cross-section stalked, anvil¬

shaped; mesocarp moist but not juicy; exocarp

1-1.4 mm thick; pedicels 16-23 mm long in

fruit, 0.9-1.2 mm thick at mid-point; seeds

white or pale yellow, 2-2.3 mm long. Fig. 36.

Specimens examined : Queensland. South Kennedy

District: Eungella Range, via Mackay, Oct 1922, Francis

s.n. (BRI); catchment of Mt William Creek, Eungella N.R,

Dec 2002, Meyer & Bloor s.n. (BRI); Snake Road, S.F.62,

NE of Eungella township, Feb 2003, Bean 20031 (BRI,

MEL); Chelman’s Road, S.F.62, NE of Eungella, Feb 2003,

Bean 20057 (BRI).

Distribution and habitat : Solanum francisii is

endemic to Queensland. Recorded only from

the Eungella area, west of Mackay (Map 12).

It grows in high-rainfall notophyll rainforest

at altitudes of 1000-1100 metres.

Phenology: Flowers recorded for September

and October; mature fruits in February.

Notes: S. francisii is related to S. sporadotrichum,

but differs by: the bark non-corky, except at the

base of large plants; the longer and wider leaves

with acute lobes; smaller stellae on all plant

parts; inner surface of corolla sparsely stellate-

hairy; calyx prickles usually present; and calyx

with Type 2 hairs.

Conservation status: S. francisii is known only

from a small section of the Eungella N.R and

an adjacent State Forest. It is threatened by low

Bean, Taxonomy of Solanum subg. Leptostemonum

769

QuE£ft3LAM0 HERBAJtMfeN rfiftll

Souf* KPW(

StMkmum

C 4 fl CTIM 4 . mn

E*pm

C-t AMD HERBARIUM (BRI)

L^uba/W AfcfK-hiiJ4,

" 620250

Mter/K ^<*«™** ***** ««'t

-fr^cLsii

p*1fl iQCjrZtol

Dft

Dufii AD MEL NSW

■ OM^ rww AO flJOiSS

Fig. 36. Holotype of Solanum francisii.

770

population size and weeds (especially Lantana

camara). Applying the IUCN guidelines

(IUCN. 2001), a category of “Vulnerable” is

recommended (VU Blab(iii,v)+2ab(iii,v);

Cl+2a(i); Dl).

Etymology : Named for W.D. Francis (1889-

1959), the first collector of the species. Mr

Francis was the author of the well known book

“Rainforest Trees of Australia”, and was for a

time the Director of the Queensland Herbarium.

65. Solanum dumicola A.R.Bean sp. nov.

Fruticulus; cortice tenui, nondescripto;

ramulorum aculei 30-70 per decimetrum;

folia adulta integra, ovata usque late

ovata; pagina superior folii viridis, stellis

sparsis praedita, pilis digitatis carens;

inflorescentiae 1-3-florae; pagina interna

corollae sparse stellato-pilosa; calyx

stellis 0.25-0.4 mm latitudine ornatus,

pilis type-2 gerens sed aculeis carens;

fructus maturi virides, pedicellis 10-24

mm longis. Typus: Queensland.

Leichhardt District: Cannondale scrub,

Amphitheatre section, Expedition

National Park, 6 November 1998, P.I.

Forster PIF23858 & R. Booth (holo: BRI

(1 sheet + spirit); iso: MEL, QRS).

Sprawling or erect, herbaceous resprouter,

0.3-0.5 m high. Juvenile stage unknown. Adult

branchlets brown or green; prickles 30-70 per

decimetre, straight, acicular, 6-10 mm long,

13-20 times longer than wide; stellae dense,

0.25-0.6 mm diameter, stalks 0-0.2 mm long;

lateral rays 4-7, porrect; central ray 1-1.5 times

as long as laterals, not gland-tipped; finger

hairs absent; Type 2 hairs absent. Adult leaves

ovate or broadly ovate, entire; lamina 4.5-12.5

cm long, 3-7.5 cm wide, 1.6-1.9 t im es longer

than broad, apex obtuse or acute, base cuneate,

oblique part 0-7 mm long, obliqueness index

0-7 percent; petioles 0.6-2.8 cm long, 13-25%

length of lamina, prickles present. Upper leaf

surface green; prickles 0-20, straight, acicular,

3-7 mm long, prickles absent or present on

midvein only or present on midvein and lateral

veins; stellate hairs distributed throughout;

protostellae absent; ordinary stellae sparse,

0.4-1.7 mm apart, 0.3-0.7 mm across, stalks

0-0.15 mm long; lateral rays 5-8, porrect;

central ray 1-2 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

Austrobaileya 6 (4): 639-816 (2004)

absent. Lower leaf surface greenish-white;

prickles 0 or 1, straight, acicular, absent or

present on midvein only; stellae dense,

0.15-0.4 mm apart, 0.6-0.8 mm diameter,

stalks 0-0.2 mm long; lateral rays 5-8, porrect;

central ray 1-1.5 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Inflorescence supra-axillary, solitary or

pseudo-racemose, common peduncle 1.5^1 mm

long, rachis prickles present; 1-3-flowered,

strongly andromonoecious, flowers 5-merous;

pedicels 8-13 mm long at anthesis, same

thickness throughout, 0.9-1.1 mm thick at mid¬

point, prickles absent or present. Calyx tube

3-5 mm long, lobes attenuate, 4-6 mm long;

prickles absent at anthesis; stellae dense, white,

0.25-0.4 mm across, sessile, lateral rays 5-8,

central ray 1-1.5 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

present. Corolla mauve, 7-11 mm long,

shallowly lobed, inner surface sparsely stellate-

hairy; anthers 4-5 mm long; ovary with stellate

and Type 2 hairs; functional style 5-6 mm long,

erect, with stellate and Type 2 hairs, stellae

c. 0.25 mm across, lateral rays 6-8, central ray

1-1.5 times as long as laterals. Fruiting calyx

with lobes less than half length of mature fruit,

prickles absent. Mature fruits 1 or 2 per

inflorescence, globular, 16-19 mm diameter,

green; mesocarp moist but not juicy; exocarp

0.8-1 mm thick; pedicels 10-24 mm long in

fruit, 1.1-2 mm thick at mid-point; seeds pale

yellow, 2-2.3 mm long. Fig. 37.

Specimens examined : Queensland. Leichhardt District:

Injune-Rolleston road, c. 77 km from Injune, Sep 1974,

Moriarty 1556 (BRI, CANB). Warrego District: above

Cattle Creek, ‘Carnarvon’ station, NW of Injune, Jun 1999,

McDonald 6768 (BRI); ‘Carnarvon’ station, Mar 2001,

Fensham 4249 (BRI). Maranoa District: The Tombs,

Maranoa River W branch, c. 110 km NW of Injune, Jun 1977,

Crisp 3113A (BRI).

Distribution and habitat : Solanum dumicola

is endemic to Queensland. Known from a few

scattered locations in south central Queensland

(Map 23). It usually grows in semi-evergreen

vine thicket or Belah ( Casuarina cristata) forest

on clayey soil, but there is one record from

ironbark woodland on sandy soil.

Phenology: Poorly known. Flowers are

recorded for June and November; mature fruits

for March and November.

Notes: S. dumicola is related to S.furfuraceum,

but differs by its small size; the thin non-

Bean, Taxonomy of Solanum subg. Leptostemonum

771

Fig. 37. Solanum dumicola. A. flowering branchlet x 1. B. ovary and style x 6. C. ovary with stellate and Type 2 hairs x 12.

D. stellate hair, upper leaf surface x 60. A, D, Moriarty 1556; B-C, Forster 23858.

descript bark, the more frequent prickles on the

branchlets; the broader adult leaves; the sparse

stellate indumentum and absence of finger hairs

on the upper leaf surface; the reduced 1-3

flowered inflorescence; the smaller stellae on

the calyx; and the longer fruiting pedicels.

Conservation status: S. dumicola has been

recorded from 4 locations, and is currently

known from 2 locations. It is threatened by

clearing of regrowth forest. Applying the IUCN

guidelines (IUCN. 2001), a category of

“Vulnerable” is recommended (VU B2ab(iii)).

One population is within the Expedition

National Park.

Etymology : The epithet is from the Latin, and

means “dweller in thickets”, a reference to the

usual habitat.

66. Solanum tetrathecum F.Muell., Fragm. 2:

165 (1861). Type: [Queensland.]

“Eutassa Ranges, Upper Brisbane”

[River], December 1856, F. Mueller

(lecto: MEL [MEL12231]), fide Symon

(1981).

772

Austrobaileya 6 (4): 639-816 (2004)

Erect, herbaceous resprouter, 0.2-0.5 m high.

Juvenile stage absent. Adult branchlets grey or

yellow; prickles 1-20 per decimetre, straight,

acicular, 3-10 mm long, 6-15 times longer than

wide; stellae very dense, 0.4-0.6 mm diameter,

stalks 0-0.1 mm long; lateral rays 6-9, porrect;

central ray absent or present, 0-1 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Adult leaves

elliptical or ovate, entire; lamina 3-7 cm long,

1.1-3.2 cm wide, 2.2-3.2 times longer than

broad, apex obtuse, base cuneate or obtuse,

oblique part 0-3 mm long, obliqueness index

0-4 percent; petioles 0.4-2.2 cm long, 11-30%

length of lamina, prickles absent. Upper leaf

surface green; prickles 0-3, straight, acicular,

5-11 mm long, prickles absent or present on

midvein only; stellate hairs confined to midrib,

or distributed throughout; protostellae absent;

ordinary stellae density very sparse to moderate,

0.4-1.6 mm apart, 0.25-0.5 mm across, sessile;

lateral rays 7 or 8, porrect; central ray 0.6-1.2

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent or

present throughout, 0.1-0.5 mm apart. Lower

leaf surface white or yellowish; prickles absent;

stellae dense, c. 0.1 mm apart, 0.4-0.7 mm

diameter, stalks 0-0.1 mm long; lateral rays

7-9, porrect; central ray 0.7-1.2 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Inflorescence leaf-

opposed or supra-axillary, solitary or pseudo-

racemose, common peduncle 3-11 mm long,

rachis prickles absent; 1-3-flowered, weakly

andromonoecious, flowers 5-merous; pedicels

5-20 mm long at anthesis, same thickness

throughout, 0.8-1.3 mm thick at mid-point,

prickles absent. Calyx tube 3-4.5 mm long,

lobes deltate, 1-3.5 mm long; prickles absent

at anthesis; stellae very dense, yellow,

0.25-0.4 mm across, stalks 0-0.1 mm long,

lateral rays 7-10, central ray 0.7-1.3 times as

long as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Corolla mauve or

purple, 8-15 mm long, deeply lobed, inner

surface sparsely to densely stellate-hairy;

anthers 5-7 mm long; ovary with Type 2 hairs

only; functional style 8.5-12.5 mm long, erect,

with stellate and Type 2 hairs, stellae 0.1-0.2

mm across, lateral rays c. 6, central ray c. 1

times as long as laterals. Fruiting calyx with

lobes less than or more than half length of

mature fruit, prickles absent. Mature fruits 1

per inflorescence, globular, 12-17 mm

diameter, yellowish-green or pale green with

dark green streaks, 4-locular; placenta in cross-

section sessile, semi-circular, or stalked, anvil¬

shaped; mesocarp moist but not juicy; exocarp

0.6-1.1 mm thick; pedicels 8-20 mm long in

fruit, 0.8-1.3 mm thick at mid-point; seeds pale

yellow, brown or black, 2.7-2.8 mm long.

Specimens examined : Queensland. Burnett District:

Kingaroy, Oct 1945, Michael 2955 (BRI); ‘Kragra’, 60 km

N of Chinchilla, Aug 1986, Hando (BRI); Meandu Mine,

Tarong Coal, adjacent to Dobby L.A., S.F.289, Feb 1994,

Forster PIF14837 & Smyrell (AD, BRI); S.F.132 Allies

Creek, c. 55 km S of Mundubbera, Nov 2002, Bean 19609

(BRI, NSW). Darling Downs District: Tara, Feb 1938,

White 11181 (BRI); ‘Parkhurst’ via St George, Mar 1955,

Tay lor (BRI); ‘Woodlands’, 38 mil es [61 km] N of Bungunya,

Jun 1960, Johnson 1602 (BRI); 10 miles [16 km] N of

Chinchilla, May 1966, Redgen 39 (AD, CANB); ‘Dilbong’,

35 miles [56 km] WSW of Tara, May 1968, Clague (BRI);

Bullock Head road, 21 km S of Hannaford, Nov 1974,

Johnson 2999 (BRI); 20.5 km SE of Moonie Hwy, SE of

Westmar, Nov 1999, Bean 15861 (BRI); ‘Karriba’, SWof

Milmerran, Nov 1999, Bean 15889 (BRI); cnr Schwennsen’s

Road & Riverglen road, N of Glenmorgan, Apr 2001 , Bean

17665 & Pedley (BRI); off Lee’s Road, 17 km W of

Chinchilla, Feb 2003, Bean 19964 (BRI). Moreton District:

Cooyar Range, S.F.289, c. 6 miles [10 km] W of Yarraman,

Jul 1969, Smith 14736 (BRI, CANB); S.F.289, c. 5 kmNW

of Yarraman, Jan 2000, Bean 15991 (BRI, CANB, MEL,

NSW).

Distribution and habitat : Solanum tetrathecum

is endemic to Queensland. Known from two

separate areas: the main area is on the western

Darling Downs from Allies Creek to Westmar;

the other is around Kingaroy and Yarraman

(Map 22). It usually inhabits heavy clays soils

in Brigalow-Belah forest, but also grows on the

margins of notophyll rainforest, in Eucalyptus

populnea woodland or (rarely) on stony

ironbark ridges.

Phenology: Flowers are recorded for all months

of the year; mature fruits from November to

June.

Notes: Closely related to S.jucundum. See notes

under that species.

Conservation status: Widespread. Not

considered at risk.

67. Solanum cocosoides A.R.Bean sp. nov.

Frutex sparse aculeatus usque ad 1.5 m

alto; folia adulta integra ovata sine

aculeis; indumento stellato in paginis

ambabus densissime praedita; stellae

Bean, Taxonomy of Solarium subg. Leptostemonum

ramulorum calycumque pedicellis usque

ad 2 mm longae; florae pedicellis 1-4 mm

longae sub anthesi; corollae facies interna

dense stellato-pilosa; aculei calycis

absentes; fructus maturi 4-loculares,

virides usque flaveoli; semina brunnea

usque atra. Typus: Queensland. Leichhardt

District: 21.1 km W of Blackwater, 16

September 1999, A.R. Bean 15403 (holo:

BRI (1 sheet + spirit); iso: CANB, MEL,

NSW, NY).

Erect, rhizomatous perennial shrub, 1-1.5 m

high. Juvenile stage unknown. Adult branchlets

terete or ridged, rusty or brown; prickles absent

or present, 0-5 per decimetre, straight, broad-

based, 4-10 mm long, 4-7 times longer than

wide; stellae very dense, 0.5-1 mm diameter,

stalks 0.2-2.2 mm long; lateral rays 7-11,

ascending; central ray 1-2 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Adult leaves ovate, entire;

lamina 4.5-8.5 cm long, 1.7-3.5 cm wide, 2.3-

2.7 times longer than broad, apex acute, base

obtuse or cordate, oblique part 0-3 mm long,

obliqueness index 0-6 percent; petioles 0.9-

2.5 cm long, 17-35% length of lamina, prickles

absent. Upper leaf surface grey; prickles absent;

stellate hairs distributed throughout;

protostellae absent; ordinary stellae very dense,

c. 0.05 mm apart, 0.4-0.8 mm across, stalks

0-0.1 mm long; lateral rays 7 or 8, porrect or

ascending; central ray 1-1.5 t im es as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Lower leaf surface white

or yellowish; prickles absent; stellae very dense,

c. 0.05 mm apart, 0.5-0.8 mm diameter, stalks

0-0.3 mm long; lateral rays 7-9, porrect;

central ray 1-1.5 t im es as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Inflorescence supra-axillary, pseudo-

racemose, co mm on peduncle 15-33 mm long,

rachis prickles absent; 2-5-flowered, weakly

andromonoecious, flowers 5-merous; pedicels

1-4 mm long at anthesis, same thickness

throughout or markedly thicker distally, c. 0.7

mm thick at mid-point, prickles absent. Calyx

tube 3-4 mm long, lobes attenuate, 3-9 mm

long; prickles absent at anthesis; stellae very

dense, brown or rusty, 0.4-1 mm across, stalks

0-2 mm long, lateral rays 7 or 8, central ray

1-2 times as long as laterals, not gland-tipped;

773

finger hairs absent; Type 2 hairs absent. Corolla

purple, 12-20 mm long, rotate, inner surface

densely stellate-hairy; anthers 5.5-6.5 mm

long; ovary with Type 2 hairs only; functional

style 9.5-11.5 mm long, erect, with Type 2 hairs

only. Fruiting calyx with lobes less than half

length of mature fruit, prickles absent. Mature

fruits 1-3 per inflorescence, globular, 14-20

mm diameter, yellowish-green or pale green

with dark green streaks, 4-locular; placenta in

cross-section stalked, circular to elliptical, or

stalked, anvil-shaped; mesocarp moist but not

juicy; exocarp 0.7-2.5 mm thick; pedicels 4-8

mm long in fruit, 1-1.2 mm thick at mid-point;

seeds brown to black, 2.6-3 mm long. Fig. 38.

Specimens examined : Queensland. Leichhardt District:

‘Berrigurra’, c. 17 km NW of Blackwater, Aug 1984,

Anderson 3796 (BRI); 21.1 km W of Blackwater, Sep 1999,

Bean 15401 (BRI, MEL); eastern edge of S.F.236, SW of

Blackwater, Nov 2002, Bean 19522 (A, BRI, MEL, MO);

20.8 km W of Blackwater, Aug 1973, Trapnell 61 & Williams

(BRI). Maranoa District: Great Dividing Range, c. 80 km

SW of Rolleston, 7 km SE of Mt Sugarloaf, Jun 1977, Crisp

3102 (AD, BRI, CANB).

Distribution and habitat : Solanum cocosoides

is endemic to Queensland. Known from a

limited area of central Queensland, around

Blackwater and near Rolleston (Map 24). It

inhabits low ridges dominated by Acacia

catenulata (Bendee) and Eucalyptus exserta,

with grey loamy soil.

Phenology: Flowers are recorded from June to

November; fruits are recorded for August and

November.

Notes: S. cocosoides is morphologically close

to S. jucundum, but differs by the branchlet

stellae 0.5-1 mm across, with stalks 0.2-2.2

mm long (0.15-0.4 mm across and stalks 0-

0.4 mm long for S. jucundum ); petioles 17-

35% length of lamina (10-20% for S. jucundum)’,

corolla rotate (deeply lobed for S. jucundum ); the

larger stellae on both leaf surfaces; and the often

larger fruits.

Conservation status: S. cocosoides has been

collected from 4 locations in central

Queensland. It is not known from a conservation

reserve. Land clearance is a minor threat.

Applying the IUCN guidelines (IUCN. 2001),

a category of “Near Threatened” is

recommended.

774

Austrobaileya 6 (4): 639-816 (2004)

QUEENSLAND HERBARIUM (BR1)

Psora el Queensland

Laieiilwrall

Eetinwrt

Coll. A.R.Bean 15403

16 SEP 1993

QUEENSLAND HERBARIUM (BRl)

Brisbane Aust'Slia

g7<> t on

Alt

Depih

23-36'S 148'40'E

21.1 fen Weal 61 BJechwetor.

iSPS 23 36 04 14B 40 291

Low ridges dommaled lw AMCiS cawnglata, w»

hetarophytu* war. lurtus. Alpdilonia emcalsa. Eremophiia-

Swillaw atrKf/ soil.

SJin* 16 1-Sm tiigh. pricStlO* sparse.

Flowers penlagCftfil, deep purple.

Occasional al sile-

Spinl material St BRl.

0flL 200 ScilanarsaM

■maj us ciled ewrpuWrUed cclkictksi number AQ E7&135

rArcim-aJ PsptP

fiOl^Tffe ^! eErts1ar>d Hertarium (BRl)

So/tm*v CoCejQ/'&s ,t 6 /* *<. ,

D * L ckit* / ocr

Fig. 38. Holotype of Solanum cocosoides.

Bean, Taxonomy of Solanum subg. Leptostemonum

Etymology : resembling Cocos, a genus of

palms; an allusion to the very long-stalked

stellae of the branchlets, which resemble

miniature palm-trees.

68. Solanum jucundum A.R.Bean sp. nov.

Frutex perennis usque ad 1.8 m altus;

aculei ramuli sparsi vel nulli; cortex

tenuis non-descriptus; folia adulta

integra, lanceolata usque ovata, aculeis

carentia; pagina superior folii stellis

densis usque densissimis et pilis type-2

praesentes; corolla profunde lobata,

pagina interna dense stellato-pilosa; calyx

aculeis carens, lobis rostratis usque

attenuatis; fructus maturi globulares, 14-

lb mm diametro, flavo usque viridi.

Typus: Queensland. Darling Downs

District: Spinifex Road, 16 km N of Tara,

28 May 2000, A.R. Bean 16662 (holo:

BRI (2 sheets + spirit); iso: MEL, NSW).

Solanum sp. (Monto A.R. Bean 8817) in

Henderson (2002).

Illustration : Symon (1981: 177), as

S. tetrathecum.

Erect, rhizomatous perennial shrub, 0.7-1.8 m

high. Juvenile stage unknown. Adult branchlets

grey to yellow or rusty; prickles absent or

present, 0-10 per decimetre, straight, acicular

or broad-based, 6-10 mm long, 6-14 times

longer than wide; stellae very dense, 0.15-0.4

mm diameter, stalks 0-0.4 mm long; lateral

rays 8-11, porrect, ascending or multiradiate;

central ray 1-1.5 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Adult leaves lanceolate or ovate, entire;

lamina 2.5-8.5 cm long, 1.1-2.3 cm wide,

2.3-3.6 t im es longer than broad, apex acute,

base obtuse or cordate, oblique part 0-3.5 mm

long, obliqueness index 0-4 percent; petioles

0.4-1.3 cm long, 10-20% length of lamina,

prickles absent. Upper leaf surface grey-green

or grey; prickles absent; stellate hairs

distributed throughout; protostellae absent;

ordinary stellae dense to very dense, 0.05-0.1

mm apart, 0.2-0.4 mm across, sessile; lateral

rays 6-8, porrect or ascending; central ray

0.7-1.3 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs present

throughout, 0.5-1 mm apart. Lower leaf surface

yellowish, or rusty; prickles absent; stellae very

775

dense, 0.05-0.1 mm apart, 0.2-0.4 mm

diameter, stalks 0-0.3 mm long; lateral rays

7-9, porrect; central ray 0.7-1.4 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Inflorescence

supra-axillary, pseudo-racemose, common

peduncle 6-12 mm long, rachis prickles absent;

2-5-flowered, weakly andromonoecious,

flowers 5-merous; pedicels 3-10 mm long at

anthesis, markedly thicker distally, 1-2 mm

thick at mid-point, prickles absent. Calyx tube

2.5- 4.5 mm long, lobes attenuate or rostrate,

1-10 mm long; prickles absent at anthesis;

stellae very dense, white to brown or rusty,

0.3-0.5 mm across, stalks 0.1-0.4 mm long,

lateral rays 7-10, central ray 0.8-1.5 times as

long as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Corolla mauve or

purple, 10-14 mm long, deeply lobed, inner

surface densely stellate-hairy; anthers 4.5-6.5

mm long; ovary with Type 2 hairs only, or with

stellate and Type 2 hairs; functional style 8-10

mm long, erect, with Type 2 hairs only. Fruiting

calyx with lobes less than half length of mature

fruit, prickles absent. Mature fruits 1 per

inflorescence, globular, 14-16 mm diameter,

yellowish-green or pale green with dark green

streaks, 4-locular; placenta in cross-section

stalked, anvil-shaped; mesocarp moist but not

juicy; exocarp 0.8-1.1 mm thick; pedicels

5—15 mm long in fruit, 0.8-2.5 mm thick at

mid-point; seeds pale yellow or brown to black,

2.5- 3.5 mm long. Fig. 39.

Specimens examined : Queensland. Mitchell District: E

of Jericho, Jul 1934, Blake 6814 (BRI). Leichhardt District:

Injune-Rolleston road, c. 10 km from Injune on Rolleston

road, Sep 1974, Moriarty 1554 (BRI, CANB); ‘Anchor’, 28

km SW of Duaringa, Mar 1982, Anderson 2969 (BRI);

‘Morabinda’, Taroom Shire, Jun 1993, Schefe CS7534

(BRI); Isla Gorge N.P., Jan 2000, McDonald KRM237

(BRI). Port Curtis District: S.F.69 Dawes Range, SE of

Thangool, Mar 1996, Bean 10113 (BRI); 16 km NNE of

Biloela, T.R.170, Mar 1996, Thompson BIL236 & Price

(AD, BRI). Warrego District: 0.4 km W of Gee Gee Gap,

Mt Moffatt N.P., Dec 1997, Bean 12904 (BRI). Maranoa

District: c. 10 mi les [16 km] W of Mitchell, May 1949,

Everist 3745 (BRI); Moonie Highway, 61.2 km E of St

George, Aug 1961, Phillips 442 (BRI, CANB); ‘Deemfem’,

53 miles [85 km] SW of Roma, Feb 1962, Nancarrow (BRI);

‘Windamore’, c. 40 miles [64 km] ESE of St George, Feb

1967, Pedley 2187 (BRI); Chesterton Range, c. 54 km N

from Mungallala, Sep 1993, Purdie 4414 (BRI, CANB); near

Coomrith Road, c. 25 km SSW of Glenmorgan, Apr 2001,

Bean 17664 & Pedley (BRI, MEL, NSW). Burnett District:

Fontainea Scrub, S.F.172, Gurgeena Plateau, Mar 1994,

Forster PIF15067 (AD, BRI, MEL); S.F.215, c. 20 km ESE

of Monto, Aug 1995, Bean 8817 (BRI). Darling Downs

776

Austrobaileya 6 (4): 639-816 (2004)

0UEEHSLANO HERBARIUM (BR!)

Flora of Queensland failing Down*

Coll- A.R.B**n 16*162

QUEENSLAND HERBARIUM (BRI)

Brisbane Auilroku

37'00'S 150^0'E

Spinire* Road,. 16 km: hi pi Tara.

(GPS 27 08 20 150 28 50).

Flal area

Woodland or Eucalyptus c ns tun, E- exsarta. Acao* btirrown.

Sandy scki.

Branched shrub, l.4m high, Prickles vary spars* on imnk amt

large branches. Fruits ivd, green wflh dam brawn marks.

Common at site.

Spirit inatenal al BRI

Specimen at juvenile pUml Included.

Dei.

Dups. NSW MEL

■May bo tiled si HHnpiilCJiHid colfeelien number AQ JM59Q

tAKhn-Jl P*X*i)

AQ 490 53<

L’.’ll

280 SoiansceaE

ffotgjyfg Queensland Har&arium (BRI)

■Sob* u*i JUC-Uiltium,

DW- A. Din /{5<pr-4=oJ

Fig. 39. Holotype of Solanum jucundum.

Bean, Taxonomy of Solanum subg. Leptostemonum

District: NNWof Bungunya, Jul 1945, Blake 15860 (BRI);

‘Cypress Downs’, c. 15 miles [24 km] NW of Jackson, Apr

1951, Everists.n. (BRI); ‘Lapunyah’, 42 mi les [68 km] NW

of Goondiwindi, Jul 1958, Johnson 511 (BRI, CANB);

Coomrith area near Meandarra, Jul 1969, Webb & Tracey

8300 (AD, BRI, CANB, K, L); Chinchilla-Miles road, c. 10

km from Miles, Sep 1974, Moriarty 1553 (AD, BRI, CANB);

Gurulmundi-Woleebee road, 11 km W of Gurulmundi, Oct

1975, Williams 75053 (BRI). New South Wales. North West

Plains: Narrabri West, Jun 1907, Boorman s.n. (NSW).

Central Western Slopes: Newell Highway, 35 mi les [56 km]

SW of Dubbo towards Peak Hill, Nov 1969, Coveny 2514

(BRI, NSW).

Distribution and habitat : Solanum jucundum

is widely distributed in Queensland south of

the Tropic of Capricorn, especially in the

western Darling Downs and Maranoa districts

(Map 25). It also extends to central-western

New South Wales, as far south as Peak Hill. It

inhabits stony or sandy ridges in shrubby

eucalypt woodland; the dominant species often

include Eucalyptus crebra, E. citriodora,

E. melanophloia or Acacia catenulata.

Phenology : Flowers are recorded for every

month of the year; mature fruits from

September to June.

Notes: S. jucundum is closely related to

S. tetrathecum, but differs by the moderate to

very dense indumentum of the upper leaf

surface, perennial habit and greater height (up

to 1.8 m high), the acute leaf apex, the rostrate

to attenuate calyx lobes, the calyx stellae with stalks

0.1-0.4 mm long (0-0.1 mm for S. tetrathecum)

and the style bearing Type 2 hairs only (Type 2

hairs and stellate hairs for S. tetrathecum). The

distributions of S. jucundum and S. tetrathecum

overlap considerably, especially around

Glenmorgan, Hannaford and Westmar, but they

never grow together and there is no

intergradation between them.

Conservation status : Widespread. Not

considered at risk.

Etymology : From the Latin jucundus, pleasing

or agreeable. Well developed plants have a

dense canopy of leaves and provide a showy

display of flowers.

69. Solanum centrale J.M.Black, Trans. &

Proc. Roy. Soc. South Australia 58: 180

(1934). Type: Northern Territory.

‘Macdonald Downs’ Station, anno 1932,

J. Chalmers s.n. (lecto: K), fide Symon

(1981), photo seen.

ill

Erect, herbaceous resprouter, 0.2-0.4 m high.

Juvenile stage unknown. Adult stem prickles

present. Adult branchlets yellow, rusty or

brown; prickles absent or present, 0-2 per

decimetre, straight, acicular, 4-7 mm long,

7-10 times longer than wide; stellae very dense,

0.5-0.6 mm diameter, stalks 0-0.2 mm long;

lateral rays 7 or 8, porrect; central ray 0.8-1.2

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Adult

leaves ovate, entire; lamina 2.8-8.5 cm long,

1.1-4.2 cm wide, 2-2.5 times longer than

broad, apex obtuse or acute, base cuneate to

cordate, oblique part 0-1.5 mm long,

obliqueness index 0-4 percent; petioles

0.8-2.6 cm long, 17-40% length of lamina,

prickles absent. Upper leaf surface grey-green

or grey; prickles absent; stellate hairs

distributed throughout; protostellae absent;

ordinary stellae dense to very dense, 0.05-0.2

mm apart, 0.5-0.7 mm across, stalks 0-0.2 mm

long; lateral rays 7 or 8, porrect; central ray

0.7-1.1 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Lower leaf surface white, yellowish or rusty;

prickles absent; stellae very dense, 0.05-0.1

mm apart, 0.5-0.8 mm diameter, stalks

0.1-0.3 mm long; lateral rays 6-8, porrect;

central ray 0.7-1.1 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Inflorescence supra-axillary,

pseudo-racemose, common peduncle 4-23 mm

long, rachis prickles absent; 1-4-flowered,

flowers 5-merous; pedicels 5-7 mm long at

anthesis, same thickness throughout or

markedly thicker distally, 0.9-1.3 mm thick at

mid-point, prickles absent. Calyx tube c. 4 mm

long, lobes rostrate, 3-4 mm long; prickles

absent at anthesis; stellae very dense, brown or

rusty, 0.4-0.5 mm across, stalks 0-0.2 mm

long, lateral rays 7 or 8, central ray 0.7-1.2

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Corolla

mauve or purple, 10-13 mm long, shallowly

or deeply lobed, inner surface sparsely to

densely stellate-hairy; anthers 4.3-6.2 mm

long; ovary glabrous; functional style c. 9 mm

long, erect, glabrous. Fruiting calyx with lobes

less than or more than half length of mature

fruit, prickles absent. Mature fruits 1 per

inflorescence, globular; pedicels c. 15 mm long

in fruit, c. 1 mm thick at mid-point.

778

Austrobaileya 6 (4): 639-816 (2004)

Specimens examined: Queensland. Mitchell District:

Idalia N.P., WSW of Blackall, along Emmet Pocket road,

Sep 1992, Bennie s.n. (BRI). Warrego District: IdaliaN.P.,

Round Hole, Mar 1996, Forster PIF18891 & Ryan (BRI);

Chesterton Range N.P., south-west corner, Jul 2001, Dollery

266 (BRI).

Distribution and habitat: Solanum centrale

is of restricted occurrence in Queensland,

known only from Idalia and Chesterton Range

National Parks (Map 24). It is widespread in

the southern part of Northern Territory and

adjacent areas of South Australia and Western

Australia. The habitat for Queensland

populations is recorded as “ Acacia aneura

woodland or tall shrubland on red sand”.

Phenology : Flowers are recorded for March,

July and September; mature fruits not seen.

Notes: Solanum centrale has not been recorded

for Queensland before. The specimens cited

above are a good match for specimens from the

Northern Territory, including the type.

However, the N.T. material tends to have leaves

more rusty in colour, shorter prickles, and

stellae with a somewhat shorter central ray.

Further Queensland material, particularly of the

fruits, is needed to confirm the identity. S. centrale

is closely related to S. jucundum, but differs by

its lower stature; the much larger stellae of the

branchlets, upper and lower leaf surfaces;

absence of Type 2 hairs from the upper leaf

surface; and the glabrous ovary.

S. centrale produces an edible fruit, which

was highly prized by the Aboriginal tribes of

central Australia (Latz 1995).

Conservation status: Although very restricted

in Queensland, this species is common and

widespread in central Australia.

70. Solanum cinereum R.Br., Prodr. 446

(1810). Type: New South Wales: “banks

of the Grose” [River], “1804” [October-

November 1803], R. Brown (holo: BM

(Bennett No. 2678)).

Illustration: Cunningham etal. (1981: 595);

Symon (1981: 259).

Sprawling or erect, rhizomatous perennial

shrub, 0.3-0.7 m high. Leaves (in outline)

broadly ovate; lamina 5-9.5 cm long, 4-9 cm

wide. Adult branchlets grey or mauve; prickles

15-60 per decimetre, straight, acicular, 3-10

mm long, 11-16 times longer than wide; stellae

dense to very dense, 0.35-0.5 mm diameter,

stalks 0-0.1 mm long; lateral rays 8-14,

porrect; central ray 0.5-1 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Adult leaves ovate or

broadly ovate, deeply lobed throughout; lobes

3- 5 on each side, acute or obtuse, lobing index

2.7-5; lamina 5.5-12.5 cm long, 3.5-5.5 cm

wide, 1.4-2.5 times longer than broad, apex

obtuse or acute, base cuneate or obtuse, oblique

part 0-12 mm long, obliqueness index 0-12

percent; petioles 1.7-3.3 cm long, 20-50%

length of lamina, prickles present. Upper leaf

surface green; prickles 13-65, straight,

acicular, 2-10 mm long, prickles present on

midvein and lateral veins; stellate hairs

confined to midrib, or distributed throughout;

protostellae absent; ordinary stellae very sparse

to sparse, 0.9-2.5 mm apart, 0.2-0.4 mm

across, sessile; lateral rays 6-8, porrect; central

ray 1-2 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Lower leaf surface white or yellowish; prickles

20-45, straight, acicular, present on midvein

and lateral veins; stellae very dense, c. 0.05

mm apart, 0.4-0.7 mm diameter, stalks 0-0.2

mm long; lateral rays 7 or 8, porrect; central

ray 0.5-1.5 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Inflorescence supra-axillary, pseudo-racemose,

common peduncle 0-18 mm long, rachis

prickles present; 3-6-flowered, weakly

andromonoecious, flowers 5-merous; pedicels

7-16 mm long at anthesis, same thickness

throughout, 0.8-1 mm thick at mid-point,

prickles present. Calyx tube 2.5-3.8 mm long,

lobes attenuate, 2.5-4 mm long; prickles

present at anthesis, 20-65 per flower, 1-7 mm

long; stellae dense, transparent or purple,

0.4-0.5 mm across, sessile, lateral rays 7-11,

central ray 0-0.5 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Corolla purple, 11-15 mm long, rotate,

inner surface sparsely stellate-hairy; anthers

4- 5.5 mm long; ovary glabrous; functional style

6-10 mm long, erect, with stellate hairs only.

Fruiting calyx with lobes less than half length

of mature fruit, prickles 2-7 mm long. Mature

fruits 1-4 per inflorescence, globular, 17-28

mm diameter, yellowish-green or pale green

with dark green streaks, glabrous or with a few

Bean, Taxonomy of Solanum subg. Leptostemonum

scattered stellate hairs, 2-locular; mesocarp

moist but not juicy; exocarp 2-4.5 mm thick;

pedicels 15-27 mm long in fruit, 1.2-1.4 mm

thick at mid-point; seeds brown to black, 2.9-

3.7 mm long. Narrawa Burr.

Specimens examined : Queensland. Darling Downs

District: The Summit, Apr 1954, Hall (BRI); Glenlyon-

Mingoola area, Apr 1957, Taylor (BRI); Severnlea,

Stanthorpe district, May 1961, Morwood (BRI); Bendee near

Stanthorpe, Mar 1973, Newton (BRI); Inglewood, Sep 1973,

O ’Donohue (BRI); 5.4 km E of ‘Arcot’, ENE of Texas, May

2000, Bean 16651 (BRI, MEL, NSW); S.F.444, Durikai,

Herries Range, Nov 2001, Forster PIF27745 (BRI, MEL).

New South Wales. Northwest Slopes: 60 km E of Bonshaw

towards Tenterfield, Aug 1975, Coveny 6665 (BRI, NSW)

Distribution and habitat : In Queensland,

Solanum cinereum is confined to the Warwick-

Texas-Stanthorpe area (Map 25), but it is

widespread in New South Wales. It grows in

shrubby eucalypt or Eucalyptus-Callitris

woodland.

Notes: for more information, see Bean (2001).

Conservation status: Widespread. Not

considered at risk.

71. Solanum nobile A.R.Bean, Telopea 9: 645

(2001). Type: New South Wales:

Northern Tablelands: Gwydir Highway,

Gibraltar Range National Park, 0.7 km

east of watershed, 8 September 2000, A.R.

Bean 16850 (holo: BRI; iso: AD, K, MEL,

NSW).

Solanum sp. 5 in Ross (1986).

Illustration: Bean (2001: 647); N. & H.

Nicholson, Australian Rainforest Plants

IV, p. 64 (1994), as S. sp. aff. cinereum.

Erect, rhizomatous perennial shrub, 1.5-4 m

high. Leaves (in outline) ovate or broadly ovate,

deeply lobed, with 2-4 pairs of lobes; lamina

9-24 cm long, 5.5-17 cm wide, with c. 60

prickles on upper surface. Adult branchlets

white or grey; prickles 2-30 per decimetre,

straight, acicular or broad-based, 3-9 mm long,

6-10 t im es longer than wide; stellae dense,

0.2-0.3 mm diameter, sessile; lateral rays 6-7,

porrect; central ray absent or present, 0-1 times

as long as laterals, not gland-tipped; finger

hairs absent; Type 2 hairs absent. Adult leaves

ovate or broadly ovate, shallowly to deeply

779

lobed throughout; lobes 2-4 on each side, acute,

lobing index 1.7-4; lamina 7-14 cm long, 3-5

cm wide, 1.4-2.7 times longer than broad, apex

acute, base obtuse or cordate, oblique part 1-8

mm long, obliqueness index 1-6 percent;

petioles 1.3-3.5 cm long, 17-26% length of

lamina, prickles absent or present. Upper leaf

surface green; prickles 0-10, straight, acicular,

4-10 mm long, prickles absent or present on

midvein only or present on midvein and lateral

veins; stellate hairs distributed throughout;

protostellae present; ordinary stellae sparse,

0.4-0.5 mm apart, 0.15-0.25 mm across,

sessile; lateral rays 6-8, porrect; central ray

0-0.5 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Lower leaf surface white or yellowish; prickles

0-2, straight, acicular, absent or present on

midvein only; stellae very dense, c. 0.05 mm

apart, 0.35-0.45 mm diameter, sessile; lateral

rays 6 or 7, porrect; central ray 0-0.5 times as

long as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Inflorescence

supra-axillary, pseudo-racemose, common

peduncle 0-6 mm long, rachis prickles absent

or present; 4-9-flowered, weakly

andromonoecious, flowers 5-merous; pedicels

10-17 mm long at anthesis, markedly thicker

distally, 0.7-1.2 mm thick at mid-point,

prickles absent or present. Calyx tube 3-6 mm

long, lobes attenuate, 5-8 mm long; prickles

absent or present at anthesis, 0-5 per flower,

2-5 mm long; stellae dense, transparent or

purple, 0.4-0.7 mm across, stalks 0-0.3 mm

long, lateral rays 6-8, central ray 1-1.5 times

as long as laterals, not gland-tipped; finger

hairs absent; Type 2 hairs absent. Corolla

purple, 10-17 mm long, rotate, inner surface

glabrous; anthers 4-5.5 mm long; ovary with

Type 2 hairs only; functional style 9-10 mm

long, erect, with Type 2 hairs only or with

stellate and Type 2 hairs, lateral rays 7-9.

Fruiting calyx with lobes less than half length

of mature fruit, prickles absent or present, 4-7

mm long. Mature fruits 1-3 per inflorescence,

globular, 18-24 mm diameter, pale green with

dark green streaks, 2-locular; placenta in cross-

section stalked, anvil-shaped; mesocarp moist

but not juicy; exocarp 0.8-1 mm thick; pedicels

15-24 mm long in fruit, 1.2-1.6 mm thick at

mid-point; seeds pale yellow, 2.5-2.8 mm long.

Specimens examined : Queensland. Darling Downs

District: Main Range, 24 km ENE of Killarney, Oct 1968,

780

Austrobaileya 6 (4): 639-816 (2004)

Everist 8124 (BRI); The Heads, 36 km SSW of Boonah on

road to Killamey, Aug 1972, Henderson 1335 & Sharpe

(AD, BRI); The Head, E of Killamey, Sep 1982, Bird (BRI);

0.7 Ion W of Moss Gardens, E of Killamey, Sep 2002, Bean

19367 (AD, BRI, NSW). New South Wales. North Coast:

on top of Tooloom Range, near Acacia Creek, Sep 1908, Dunn

(BRI); at Rory’s roadT/O, Ewingar S.F., south of Tabulam,

Feb 2001, Bean 17329 (BRI).

Distribution and habitat : In Queensland,

Solanum mobile is known only from the

Killamey area, but several populations are

known in New South Wales as far south as

Bellingen River (Map 27). It grows in

notophyll rainforest, or in tall wet sclerophyll

forest dominated by Eucalyptus saligna.

Notes: for more information, see Bean (2001).

Conservation status : Applying the IUCN

guidelines (IUCN. 2001), a category of

“Vulnerable” is recommended (VU

Blab(ii,iii)+2ab(ii,iii); C2a(i)).

72. Solanum limitare A.R.Bean, Telopea 9:

653 (2001). Type: Queensland. Moreton

district: adjacent to Mt Binga S.F., 11 km

SE of Cooyar, 21 February 2000, A.R.

Bean 16080 (holo: BRI; iso: MEL, NSW).

Illustration: Symon (1981:147), as S. elegans;

Bean (2001: 647).

Erect, herbaceous resprouter, 0.3-0.8 m high.

Juvenile branchlets with 10-30 prickles per dm,

2-7 mm long; leaves (in outline) lanceolate or

ovate, shallowly to deeply lobed, with 1-3 pairs

of lobes; lamina 6.5-13 cm long, 3-8 cm wide,

with 5-15 prickles on upper surface. Adult

branchlets grey or rusty; prickles 1-6 per

decimetre, straight, acicular, 3-7 mm long,

7-9 times longer than wide; stellae very dense,

0.25-0.4 mm diameter, stalks 0-0.1 mm long;

lateral rays 6-8, porrect; central ray absent or

present, 0-0.5 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Adult leaves narrow lanceolate or

lanceolate, entire; lamina 7-10.5 cm long,

0.9-1.7 cm wide, 4.5-8.5 times longer than

broad, apex acute, base obtuse, oblique part

1-3.5 mm long, obliqueness index 1-4 percent;

petioles 0.7-2.2 cm long, 8-30% length of

lamina, prickles absent or present. Upper leaf

surface grey-green; prickles 0-7, straight,

acicular, 1-6 mm long, prickles absent or

present on midvein only; stellate hairs

distributed throughout; protostellae absent or

present; ordinary stellae sparse to moderate

density, 0.2-0.6 mm apart, 0.3-0.5 mm across,

sessile; lateral rays 6-8, porrect; central ray

0-0.5 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Lower leaf surface white or yellowish; prickles

0-6, straight, acicular, absent or present on

midvein only; stellae very dense, c. 0.05 mm

apart, 0.35-0.45 mm diameter, sessile; lateral

rays 7 or 8, porrect; central ray 0.5-1 t im es as

long as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Inflorescence

supra-axillary, pseudo-racemose, common

peduncle 0-5 mm long, rachis prickles absent;

3-8-flowered, weakly andromonoecious,

flowers 5-merous; pedicels 11-20 mm long at

anthesis, same thickness throughout, 0.7-0.9

mm thick at mid-point, prickles absent. Calyx

tube 2.5-4 mm long, lobes deltate or attenuate,

3-5 mm long; prickles absent or present at

anthesis, 0-2 per flower, 1-4 mm long; stellae

dense, transparent, 0.3-0.45 mm across, stalks

0-0.1 mm long, lateral rays 7 or 8, central ray

0.5-1 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Corolla purple, 9-15 mm long, rotate, inner

surface sparsely stellate-hairy; anthers 3.5-5.5

mm long; ovary with stellate and Type 2 hairs;

functional style 8-12 mm long, erect, with

stellate hairs only, stellae 0.3-0.4 mm across,

lateral rays 8-20, central ray 1-1.5 times as

long as laterals. Fruiting calyx with lobes less

than half length of mature fruit, prickles absent.

Mature fruits 1-3 per inflorescence, globular,

14-17 mm diameter, pale green with dark green

streaks, 2-locular; placenta in cross-section

stalked, anvil-shaped; mesocarp moist but not

juicy; exocarp c. 0.5 mm thick; pedicels 16-20

mm long in fruit, 1.3-1.6 mm thick at mid¬

point; seeds white or pale yellow, 2.7-2.9 mm

long.

Specimens examined : Queensland. Burnett District:

Bunya Mtns, Oct 1919, White s.n. (BRI). Darling Downs

District: Highfield, Main Range, Dec 1875, Bancroft (BRI);

Bunya Mtns, between Ghinghion Lookout and Summerhill

View, Oct 1989, Bird (BRI); Condamine Gorge, Paddys Knob,

Lot 13, Parish of Emuvale, Mar 1993, Sparshott KM40

(BRI); Mt Mitchell walking track, Jan 2003, Hines (BRI).

Moreton District: Mt Mistake, Jun 1887, Simmonds (BRI);

Main Range, between Spring Bluff and Murphys Creek, Aug

1930, White 7017 (BRI); Ben Lomond Peak, Jan 1988, Bird

(BRI); near Boonah border gate, Croftby road, Jan 1990,

Forster 6210 (BRI); 3.6 km along Duck Creek road, near

Bean, Taxonomy of Solanum subg. Leptostemonum

O’Reillys Guest House, Mar 2001, Bean 17393 (BRI). New

South Wales. North Coast: Tooloom, Nov 1949, Webb 2188

(BRI); E of Mt Boorabee, near Kyogle, Jan 1963, Salasoo

2583 (NSW); 20 metres S of border fence, 2 km W of Mt

Lindesay, Jan 2001, Bean 17238 (BRI, NSW).

Distribution and habitat: Solanum limitare

extends from the Bunya Mountains in

Queensland to Kyogle in New South Wales

(Map 26). It grows in grassy to somewhat

shrubby eucalypt woodland, usually not far

from a rainforest edge.

Notes: for more information, see Bean (2001).

Conservation status: Applying the IUCN

guidelines (IUCN. 2001), a category of

“Vulnerable” is recommended (VU

B lab(ii,iii)+2ab(ii,iii); C2a(i)).

73. Solanum amblymerum Dunal in A.DC.,

Prodr. 13(1): 294 (1852); S. violaceum

var. amblymerum (Dunal) Maiden &

Betche, Census N.S.W. PL 181 (1916).

Type: New South Wales: Macquarie

River, October 1822, A. Cunningham 90

(holo: G-DC n.v., microfiche 13/1:

294.692; iso: K, NSW).

Erect, rhizomatous perennial shrub, 0.8-1.8 m

high. Juvenile branchlets with 15-25 prickles

per dm, 5-9 mm long; leaves (in outline)

lanceolate, shallowly to deeply lobed, with 1

or 2 pairs of lobes; lamina 7-10 cm long,

1.5-2.5 cm wide, with 3-5 prickles on upper

surface. Adult branchlets grey; prickles 1-10

per decimetre, straight, acicular, 5-9 mm long,

7-10 times longer than wide; stellae very dense,

0.3-0.4 mm diameter, sessile; lateral rays

7-10, porrect; central ray absent or present,

0-0.5 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Adult leaves linear or narrow lanceolate, entire

or with obtuse basal lobes only; lamina 5-11

cm long, 0.5-1.3 cm wide, 6—13 times longer

than broad, apex obtuse or acute, base cuneate,

obtuse or hastate, oblique part 0-1.5 mm long,

obliqueness index 0-2 percent; petioles

0.3-0.9 cm long, 5-15% length of lamina,

prickles absent or present. Upper leaf surface

green; prickles 1-7, straight, acicular, 4-10 mm

long, prickles present on midvein only; stellate

hairs distributed throughout; protostellae

present; ordinary stellae very sparse to sparse,

0.4-0.7 mm apart, 0.1-0.2 mm across, sessile;

781

lateral rays 7 or 8, porrect; central ray 0-0.5

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Lower

leaf surface white or yellowish; prickles 0-5,

straight, acicular, absent or present on midvein

only; stellae very dense, c. 0.05 mm apart,

0.2-0.45 mm diameter, sessile; lateral rays

7 or 8, porrect; central ray 0.5-1 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Inflorescence

supra-axillary, pseudo-racemose, common

peduncle 1-11 mm long, rachis prickles absent;

3-6-flowered, weakly andromonoecious,

flowers 5-merous; pedicels 5-10 mm long at

anthesis, same thic kn ess throughout, 0.8-0.9

mm thick at mid-point, prickles absent. Calyx

tube 1.5-2.5 mm long, lobes deltate or

attenuate, 2.5-4.5 mm long; prickles absent at

anthesis; stellae dense, transparent or purple,

0.25-0.4 mm across, stalks 0-0.1 mm long,

lateral rays 7 or 8, central ray 0.5-1 times as

long as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Corolla purple,

8-14 mm long, rotate, inn er surface glabrous

or sparsely stellate-hairy; anthers 4.5-6.5 mm

long; ovary with Type 2 hairs only; functional

style 8-11 mm long, erect, with stellate and

Type 2 hairs, stellae 0.4-0.5 mm across, lateral

rays 8-11, central ray c. 1 times as long as

laterals. Fruiting calyx with lobes less than half

length of mature fruit, prickles absent. Mature

fruits 1-3 per inflorescence, oblate or globular,

11-16 mm diameter, pale green with dark green

streaks, 2-locular; placenta in cross-section

stalked, anvil-shaped; mesocarp moist but not

juicy; exocarp 0.7-0.8 mm thick; pedicels 9-13

mm long in fruit, 0.9-1.4 mm thick at mid-point;

seeds white or pale yellow, 2.3-3 mm long.

Specimens examined : Queensland. Darling Downs

District: Stanthorpe, Jul 1904 , Boorman (BRI); Silverwood,

Sep 1922, White 1765 (BRI); granite hill W of Glen Aplin,

Nov 1946, Everist 1362 (BRI); ‘Benandre’, 23 miles [34

km] SE of Texas, Apr 1962, Pedley 985 (BRI); Thulimbah,

4 miles [6 km] N of Stanthorpe, Oct 1967, Greeness (BRI);

1.6 km W of Cottonvale, Oct 1975, Williams 75096 (BRI);

S.F.595, near Mt Gammie North, Sep 1992, Forster 11723

(BRI); Sundown N.R near Red Rock Gorge road to Severn

R., Jan 1993, Forster PIF12677 (AD, BRI); Mt Janet road,

Passchendaele S.F., NW of Stanthorpe, Oct 1997, Bean 12471

(BRI); 1 km S of Crystal Mt turnoff, N of Dalveen, Nov 1999,

McDonald KRM94 (BRI). Moreton District: ridge SW of

Mt Hennessy, Black Duck Ck, ‘Glenrock’, Mar 1997,

Grimshaw PG2694 (AD, BRI). New South Wales. Northern

Tablelands: Roberts Range, W of ‘Donnybrook’, via

782

Austrobaileya 6 (4): 639-816 (2004)

Wallangarra, Feb 1990, Bean 1358 (BRI); 9.9 km S of

Boonoo Boonoo River, Mt Lindesay Hwy, Jan 1992, Wilson

1321 (BRI, NSW); lookout 1 km S of Emmaville, Jan 2001,

Bean 17293 (BRI, NSW).

Distribution and habitat : In Queensland,

Solanum amblymerum is known from the

Granite Belt and adjacent areas (Map 28). It is

also common on the north-western slopes of

N. S.W. It grows in shubby eucalypt or

Eucalyptus-Callitris woodland on sandy to

loamy soils.

Notes: for more information, see Bean (2001).

Conservation status : Widespread. Not

considered at risk.

74. Solanum galbinum A.R.Bean sp. nov.

Frutex erectus sparse foliatus, in ramulis

sparse aculeatus; folia adulta linearia,

3-6 mm lata, petiolis 2-3 mm longis

(3-7% longitudinis laminae); protostellae

a pagina superiore folii absentes; calycis

aculei absentes; corolla profunde lobata,

pagina interna sparse stellato-pilosa;

fructus maturi 10-15 mm diametro,

virides cum striis obscurioribus viridibus;

placenta sessilis; semina 3.5-4 mm longa.

Typus: Queensland. South Kennedy

District: Mt Coolon-Collinsville road,

0.8 km W of Caves Creek, 20 January

1996, 1.G. Champion 1310 &A.B. Pollock

(holo: BRI).

Solanum sp. (Mt Coolon I.G. Champion+

1310) in Henderson (2002).

Erect, rhizomatous perennial shrub, 0.5-1.5 m

high. Juvenile stage unknown. Adult branchlets

rusty or brown; prickles 10-40 per decimetre,

straight, acicular, 3-8 mm long, 8-13 times

longer than wide; stellae very dense, 0.35-0.5

mm diameter, sessile; lateral rays 7 or 8,

porrect; central ray 0.4-0.8 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Adult leaves linear, entire;

lamina 4-7 cm long, 0.3-0.6 cm wide, 8.5-17

times longer than broad, apex obtuse or acute,

base cuneate, oblique part 0-2 mm long,

obliqueness index 0-3 percent; petioles

O. 2-0.3 cm long, 3-7% length of lamina,

prickles absent. Upper leaf surface green;

prickles 0-6, straight, acicular, 1.5-7 mm long,

prickles absent or present on midvein only;

stellate hairs distributed throughout;

protostellae absent; ordinary stellae sparse to

moderate density, 0.15-0.4 mm apart, 0.2-0.3

mm across, sessile; lateral rays 7 or 8, porrect;

central ray 0.7-1.3 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Lower leaf surface white; prickles

absent; stellae very dense, c. 0.05 mm apart,

0.4-0.5 mm diameter, stalks 0-0.15 mm long;

lateral rays 7 or 8, porrect; central ray 0.4-0.8

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent.

Inflorescence supra-axillary, pseudo-umbellate

or pseudo-racemose, common peduncle 0-4

mm long, rachis prickles absent; 4-9-flowered,

weakly andromonoecious, flowers 5-merous;

pedicels 4-6 mm long at anthesis, same

thickness throughout, 0.4-0.6 mm thick at mid¬

point, prickles absent. Calyx tube 1-2 mm long,

lobes deltate, 1.5-2.5 mm long; prickles absent

at anthesis; stellae very dense, purple, brown

or rusty, 0.25-0.35 mm across, stalks 0-0.1 mm

long, lateral rays 8 or 9, central ray 0.6-1.4

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Corolla

white or mauve, 7-11 mm long, deeply lobed,

inner surface sparsely stellate-hairy; anthers

4-5.5 mm long; ovary with stellate hairs only,

or with Type 2 hairs only; functional style

8-9.5 mm long, erect, with Type 2 hairs only

or with stellate and Type 2 hairs, stellae

0.3-0.4 mm across, lateral rays 5-7, central

ray 0.5-1 times as long as laterals. Fruiting

calyx with lobes less than half length of mature

fruit, prickles absent. Mature fruits 1-4 per

inflorescence, globular, 10-15 mm diameter,

pale green with dark green streaks, 1-locular

(septum absent or incomplete); placenta in

cross-section sessile, semi-circular; mesocarp

juicy or moist; exocarp 0.3-0.4 mm thick;

pedicels 5-9 mm long in fruit, 0.5-0.9 mm

thick at mid-point; seeds pale yellow, 3.5-4 mm

long. Fig. 7, 40.

Specimens examined : Queensland. Burke District: White

Mountains N.R, NWof ‘Warang’, Apr 2000, Wannan 1713

& Martindell (BRI, CANB, NSW); 13 km NW of ‘Warang’

HS site, White Mountains N.R, Apr 2000, Thomas 1817 &

Thompson (BRI). Cook District: Newcastle Range, Feb

1928, Brass 1781 (BRI, CANB). North Kennedy District:

5 miles [8 km] S of ‘Wairuna’, Jun 1967, Morain 12 (BRI);

near Reeve’s Lake, c. 25 km W of junction of Lolworth Ck

and Burdekin River, Jul 1981, Henderson H2658 (AD, BRI);

‘Valley of Lagoons’, c. 10.5 km E of HS, Apr 1989, Clarkson

7929 & Henderson (BRI); 2.5 km W of Clermont turn-off

Bean, Taxonomy of Solanum subg. Leptostemonum

783

Fig. 40. Solanum galbinum. A. flowering branchlet x 1.5. B. ovary and style x 6. C. ovary with Type 2 hairs x 16. D. stellate

hair, upper leaf surface x 100. E. mature fruit and pedicel x 2. F. transverse section of fruit x 4. A, D, Thompson BUC455 &

Sharpe ; B-C, E-F, Thomas 1437 & Thompson.

on Flinders Hwy, Charters Towers, Sep 1990, Wilson 607 &

Rowe (BRI, NSW); 46 km SWof Greenvale, towards ‘Wando

Vale’, Jun 2000, Cumming 19647 (BRI). Mitchell District:

2.6 km S of road to ‘Mundoo Bluff’ HS, on road to

‘Thirlestone’, May 1991 ,Neldner 3349 & Thompson (BRI):

‘Fortuna’, Aramac, Aug 1998, House KZX 1576 (BRI); Poison

Valley road, White Mountains N.P., Apr 2000, McDonald

KRM439 (BRI). South Kennedy District: on Great Dividing

Range, c. 21 km NNW of ‘Yarrowmere’ HS, c. 97 km SSE

of Pentland, Oct 1983, Henderson H2852 et al. (BRI,

CANB); Peak Downs Highway, 17 km W of Moranbah

turnoff, Mar 1989, Champion 437 (BRI); 20 km SE of

‘Teamass’ HS, Jun 1992, Thompson BUC455 & Sharpe

(AD, BRI); 7 8 km from Collinsville on Mt Coolon road, Feb

1994, Bean 7360 & Forster (BRI); 14 km SW of ‘Bowie’,

near Fake Buchanan, Feb 1994, Bean 7494 & Forster (BRI).

Feichhardt District: 12.5 km W of Middlemount on Dysart

road, Jul 1998, Thompson 1094 & Fox (BRI).

Distribution and habitat : Solanum galbinum

is endemic to Queensland. Distributed from

Newcastle Range near Einasleigh to

Middlemount and Moranbah (Map 27). It

grows on sandstone ridges and lateritised

plateaux in open eucalypt woodland or Acacia

shirleyi forest in shallow sandy soil.

Phenology: Flowers are recorded from January

to October; mature fruits from January to June

and in October.

Notes: S. galbinum seems closest to S. amblymerum,

but differs from that species by the linear leaves

3-6 mm wide and petioles only 2-3 mm long;

shorter calyx lobes at anthesis and shorter

fruiting pedicels; the deeply-lobed corolla; the

larger seeds and the stellae of the upper leaf

surface having a longer central ray.

784

Austrobaileya 6 (4): 639-816 (2004)

It is of very similar appearance to

S. ferocissimum, but that species has smaller

red fruits with longer pedicels, and the upper

leaf surface is glabrous or almost so.

Conservation status : Widespread. Not

considered at risk.

Etymology: From the Latin galbinus meaning

yellowish. A reference to the yellow-green fruit.

Both S. parvifolium and S. ferocissimum , with

which this species has been confused, have

bright red fruits.

Group 27D (S. esuriale group), here defined;

related to Group 27 ( S . ellipticum group)

of Whalen (1984)

Upper leaf surface stellae with 8-18 lateral rays

(100%); inner surface of corolla glabrous

(100%); calyx not accrescent in fruit (100%);

mature fruits green, yellowish-green or brown

(100%); inflorescences andromonoecious, male

flowers and bisexual flowers same size and

prickliness (100%); inflorescence solitary or

pseudo-racemose (93%); lower leaf surface

indumentum dense to very dense (90%); adult

leaves entire (86%); calyx without prickles

(86%); Type 2 hairs absent from leaves (86%);

upper leaf surface indumentum dense to very

dense (70%); seeds brown to black (50%).

14 species endemic to Australia, 8 species

indigenous to Queensland.

75. Solanum elachophyllum F.Muell., Fragm.

2: 164 (1861). Type: [Queensland.

Leichhardt District:] between the

Mackenzie and Dawson Rivers,

November 1856, F. Mueller (lecto: K;

isolecto: MEL [MEL 12234]), fide Symon

(1981).

Illustration : Symon (1981: 179).

Sprawling or erect, rhizomatous perennial

shrub, 0.1-0.4 m high. Juvenile branchlets with

c. 30 prickles per dm, 6-12 mm long; leaves

(in outline) elliptical, entire; lamina 1.2-3.4

cm long, 0.6-1.5 cm wide, with 0-2 prickles

on upper surface. Adult branchlets white or grey

or brown; prickles 15-50 per decimetre,

straight, acicular, 7-10 mm long, 11-14 times

longer than wide; stellae very dense, 0.15-0.3

mm diameter, sessile; lateral rays 10-15,

porrect; central ray absent or present, 0-0.3

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Adult

leaves elliptical, entire; lamina 0.5-1.5 cm

long, 0.3-0.7 cm wide, 1.7-2.5 times longer

than broad, apex obtuse, base cuneate, oblique

part 0-0.3 mm long, obliqueness index 0-3

percent; petioles 0.1-0.3 cm long, 18-35%

length of lamina, winged, prickles absent.

Upper leaf surface green or grey-green; prickles

0-2, straight, acicular, 2-7 mm long, prickles

absent or present on midvein only; stellate hairs

confined to midrib, or distributed throughout;

protostellae absent; ordinary stellae very sparse

to dense, 0.3-0.6 mm apart, 0.15-0.25 mm

across, sessile; lateral rays 8-10, porrect;

central ray 0-0.2 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Lower leaf surface greenish-white,

white, or yellowish; prickles absent; stellae

sparse to dense, 0.1-0.4 mm apart, 0.15-0.3

mm diameter, sessile; lateral rays 8-16, porrect;

central ray 0-0.1 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Inflorescence leaf-opposed, solitary or

pseudo-racemose, common peduncle 0-1 mm

long, rachis prickles absent; 1 or 2-flowered,

with all flowers bisexual and 5-merous; pedicels

3-15 mm long at anthesis, same thickness

throughout, 0.5-0.8 mm thick at mid-point,

prickles absent. Calyx tube 2-2.5 mm long,

lobes deltate, 1-2.5 mm long; prickles absent

at anthesis; stellae sparse to dense, white,

0.2-0.3 mm across, sessile, lateral rays 8-12,

central ray 0-0.3 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Corolla mauve or purple, 7-10 mm

long, deeply lobed, inner surface glabrous;

anthers 4-5 mm long; ovary with Type 2 hairs

only; functional style 6-6.5 mm long, erect,

glabrous or with Type 2 hairs only. Fruiting

calyx with lobes less than half length of mature

fruit, prickles absent. Mature fruits 1 per

inflorescence, globular, 12-14 mm diameter,

pale green with dark green streaks, 2-locular;

placenta in cross-section sessile, semi-circular;

mesocarp moist but not juicy; exocarp 1.2-1.8

mm thick; pedicels 6-15 mm long in fruit,

0.6-0.9 mm thick at mid-point; seeds pale

yellow or brown to black, 2.9-3.7 mm long.

Bean, Taxonomy of Solanum subg. Leptostemonum

Specimens examined'. Queensland. Leichhardt District:

Blackwater, E of Emerald, Mar 1920, Francis (BRI);

Emerald, undated, Carey (BRI); near ‘Warwick’ HS, Jul

1962, Story & Yapp 174 (BRI, CANB); 39 miles [63 km] S

of Blackwater, Jul 1964, Gittins 862 (BRI); c. 4 m il es [6

km] E of Moura, March 1967, Henderson H220 (BRI);

‘Thomby’, 20 miles [32 km] NW of Theodore, Sep 1969,

Johnson 2874 (BRI); 0.5 km N of ‘Bogandilla’ HS,

Broadsound Shire, Feb 1979, Anderson 769 (BRI); Brigalow

Research Station, May 1981, Dillewaard 621 & Johnson

(BRI); Taunton N.R, Oct 1995, Melzer RM688 & Hendry

(BRI); 3 km E of Dingo, Feb 1998, Fairfax 267 & Holman

(BRI); Brigalow Research Station, SW of Moura, Dec 2001,

Bean 18254 (BRI); Crooked Creek Dam, Junee Tableland,

Oct2002, Bean 19378 (BRI); ‘Nirvana’, c. 15kmWNWof

Banana, Apr 2003, Bean 20166 (BRI).

Distribution and habitat : Solanum

elachophyllum is endemic to Queensland.

Confined to the central east of the state, from

Middlemount to Theodore (Map 26). It grows

on fertile cracking-clay soils in association with

Brigalow ( Acacia harpophylla ), Belah (i Casuarina

cristata ), Bonewood ( Macropteranthes

leichhardtii ) or Dawson River Blackbutt

(.Eucalyptus cambageana ).

Phenology : Flowers recorded for February,

March, July and September; mature fruits in

March, May and July.

Notes: S. elachophyllum can be distinguished

from all other Queensland Solanum species by

the very small leaves (5-11 x 3-7 mm). It is

apparently related to S. esuriale and S. sturtianum

by virtue of its stellate hair morphology.

It would probably make an interesting

horticultural subject as a pot or tub plant.

Conservation status: S. elachophyllum was

first collected by Ferdinand Mueller in 1856

and has been collected in many locations since

then. However, its habitat has been greatly

reduced in recent decades and it is also

threatened by weeds, particularly introduced

pasture grasses. It is currently known from 4

locations, and it does not occur within a

conservation reserve. Applying the IUCN

guidelines (IUCN. 2001), a category of

“Endangered” is recommended (EN A2ce;

B2ab(iii,v); Cl).

76. Solanum versicolor A. R.Bean sp. nov.

Repullulator erectus herbaceus usque ad

0.3 m altus; cortex suberosus; ramuli

sparse aculeati, aculeis ll-20plo

785

longioribus quam latioribus; folia sine

aculeis, pagina superior digitis ornata,

pagina inferior albida; inflorescentia

1-3-flora, pedunculus communis absens;

pedicelli floriferi 11-26 mm longi sub

anthesi; calyx digitis praeditus sed aculeis

carens; alibastra purpurascentia-albida et

corolla albida; fructus maturi virides.

Typus: Queensland. Warrego District:

27 km SE of Charleville, on road to

Bollon, 28 January 2002, A.R. Bean

18456 (holo: BRI (1 sheet + spirit); iso:

AD, MEL, NSW).

Erect, herbaceous resprouter, 0.15-0.3 m high.

Juvenile stage absent. Bark on adult stems

furrowed, corky. Adult branchlets yellow or

brown; prickles 1-10 per decimetre, straight,

acicular, 1-5 mm long, 13-20 times longer than

wide; stellae dense, 0.5-0.8 mm diameter,

stalks 0-0.1 mm long; lateral rays 7-9, porrect;

central ray present, 0.2-0.5 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs sparse. Adult leaves lanceolate or

elliptical, entire; lamina 3.5-6 cm long,

1.1-2.2 cm wide, 2.5-4 times longer than

broad, apex obtuse, base cuneate, oblique part

0-1.5 mm long, obliqueness index 0-3 percent;

petioles 0.6-1.4 cm long, 17-25% length of

lamina, prickles absent. Upper leaf surface

grey-green; prickles absent; stellate hairs

distributed throughout; protostellae absent;

ordinary stellae dense, 0.15-0.3 mm apart,

0.5-0.7 mm across, stalks 0-0.15 mm long;

lateral rays 8-9, porrect; central ray 0.5-1 times

as long as laterals, not gland-tipped; finger

hairs present, 0.1-0.3 mm apart, gland-tipped,

0.05-0.1 mm long; Type 2 hairs absent. Lower

leaf surface white; prickles absent; stellae dense

to very dense, 0.05-0.1 mm apart, 0.6-0.9 mm

diameter, stalks 0.1-0.2 mm long; lateral rays

7 or 8, porrect; central ray 0.4-1 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Inflorescence

supra-axillary, solitary or pseudo-racemose,

common peduncle absent, rachis prickles

absent; 1-3-flowered, with all flowers bisexual

and 5-merous; pedicels 11-26 mm long at

anthesis, same thic kn ess throughout, 0.5-0.6

mm thick at mid-point, prickles absent. Calyx

tube 1.5-3 mm long, lobes attenuate, 3-4.5 mm

long; prickles absent at anthesis; stellae

moderate to dense, transparent, 0.5-0.7 mm

786

Austrobaileya 6 (4): 639-816 (2004)

QUEENSLAND HERBAHRJM [BRI]

Brisbane Austrafia

QUEENSLAND HERBARIUM (BRI)

F*St df ChtcdmLidd Wjn*ps

AQ 695293

C*n A R Elean 1

26d 33 m 21s K5d26m 47* |AC3D66]

15 *.+t*p&i. 7 P 626 r 9 | (S 1 +M 456 J 61

ZJVn SE ef C*irt*v**,on rood Id 0 $hw

Woodland or Eucafypguf popuinaa, A coca ancura. SI bOHom of

noar ipOOn dhk^n. Loamy sc*!

Slvub 20cm high; Apuwr* while, it-mfirous Groan Ihjfis jlso

PcbswtI.

OecMionf I aj

$pirri material a£ Bftl

ffa£0 7 ypg> ^ueenaand HeifcariLim (brij

S&fx/utn ters/c^/or

tW. 0*'»fy/f/(>3

Fig. 41. Holotype of Solanum versicolor.

Bean, Taxonomy of Solarium subg. Leptostemonum

across, stalks 0-0.1 mm long, lateral rays 6-8,

central ray 0.5-1 times as long as laterals, not

gland-tipped; finger hairs present; Type 2 hairs

absent. Corolla white, 7-11 mm long, shallowly

lobed; anthers 4-5 mm long; ovary with Type

2 hairs only; functional style 5-6.5 mm long,

erect, with Type 2 hairs only. Fruiting calyx

with lobes less than half length of mature fruit,

prickles absent. Mature fruits 1 or 2 per

inflorescence, globular, 11-13 mm diameter,

green, 2-locular; placenta in cross-section

sessile, semi-circular; mesocarp moist but not

juicy; exocarp 0.4-0.5 mm thick; pedicels

12-26 mm long in fruit, 0.6-0.9 mm thick at mid¬

point; seeds pale yellow, 2.7-3 mm long. Fig. 41.

Additional specimen examined : Queensland. Warrego

district: 44 km E of Charleville, towards Morven, Jan 2002,

Bean 18507 (AD, BRI, NSW).

Distribution and habitat: Solanum versicolor

is endemic to Queensland. Known only from

around Charleville in south-western

Queensland (Map 7). It grows on lower

hillslopes with Acacia aneura and Eucalyptus

populnea, on deep red earths.

Phenology: Poorly known. Flowers and fruits

recorded in January.

Notes: S. versicolor is related to S. esuriale,

but differs by: the corky bark; the presence of

finger hairs on both the upper leaf surface and

the calyx; the absence of a common peduncle;

pedicels 11-26 mm long at anthesis (5-9 mm

long for S. esuriale ); the white corolla; and the

moderate to dense stellae on the calyx (very

dense for S. esuriale ).

Conservation status: Data deficient.

Etymology: From the Latin versicolor meaning

different or varying colour, a reference to the

green, sparsely hairy pedicels, which contrast

strongly with the white, densely hairy rachises

and branchlets.

77. Solanum esuriale Lindl. in Mitchell, Three

Exped. Australia 2: 43 (1838); S. esuriale

var. esuriale Domin, Biblioth. Bot. 89:

583 (1929). Type: New South Wales.

[South Western Plains:] interior of New

Holland [near Hillston], 19 April 1836,

T. Mitchell (holo: CGE; iso: K).

787

Solanum esuriale var. sublobatum Domin,

Biblioth. Bot. 89: 583 (1929). Type:

Queensland. Mitchell District: near

Longreach, March 1910, K. Domin s.n.

(holo: PR, photo at BRI).

Illustration: Cunningham et al. (1981: 590);

Symon (1981: 173).

Sprawling or erect, herbaceous resprouter,

0.1-0.4 m high. Juvenile stage absent. Adult

branchlets terete or ridged, grey or brown;

prickles absent or present, 0-100 per decimetre,

straight, acicular or broad-based, 1-4 mm long,

8-12 times longer than wide; stellae very dense,

0.7-0.9 mm diameter, stalks 0-0.2 mm long;

lateral rays 8-12, porrect; central ray 0.3-1

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Adult

leaves linear to lanceolate, entire or shallowly

lobed throughout; lobes 3-5 on each side,

obtuse, lobing index 1-1.1; lamina 3-10 cm

long, 0.7-1.4 cm wide, 3.8-14 times longer

than broad, apex obtuse or acute, base cuneate,

oblique part 0-2 mm long, obliqueness index

0-4 percent; petioles 0.4-1.4 cm long, 13-20%

length of lamina, prickles absent or present.

Upper leaf surface grey-green or grey; prickles

absent; stellate hairs distributed throughout;

protostellae absent or present; ordinary stellae

density moderate to very dense, 0.05-0.4 mm

apart, 0.4-0.7 mm across, sessile; lateral rays

6-12, porrect; central ray 0.5-1 t im es as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Lower leaf surface

white or yellowish; prickles absent; stellae

dense to very dense, 0.05-0.4 mm apart,

0.5-0.8 mm diameter, stalks 0-0.1 mm long;

lateral rays 8-14, porrect; central ray 0.3-1

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent.

Inflorescence supra-axillary, pseudo-racemose,

common peduncle 14-36 mm long, rachis

prickles absent; 3-6-flowered, with all flowers

bisexual and 5-merous; pedicels 5-9 mm long

at anthesis, same thickness throughout,

0.6-0.7 mm thick at mid-point, prickles absent.

Calyx tube 2-3 mm long, lobes rostrate or

attenuate, 2-3.5 mm long; prickles absent or

present at anthesis, 0-3 per flower, 0.5-1 mm

long; stellae very dense, white to rusty-brown,

0.5-0.7 mm across, stalks 0-0.1 mm long,

lateral rays 6-9, central ray 0.3-1 times as long

788

Austrobaileya 6 (4): 639-816 (2004)

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Corolla mauve or

purple, 8-13 mm long, shallowly lobed, inner

surface glabrous; anthers 3.5-5 mm long; ovary

glabrous, or with stellate hairs only; functional

style 6-7 mm long, erect, glabrous or with

stellate hairs only, stellae 0.5-0.7 mm across,

lateral rays 6-8, central ray 0.7-1 times as long

as laterals. Fruiting calyx with lobes less than

half length of mature fruit, prickles absent or

present, 1-1.5 mm long. Mature fruits 1-4 per

inflorescence, globular or ellipsoidal, 12-15

mm diameter, yellowish-green or green,

glabrous or with a few scattered stellate hairs,

2-locular; placenta in cross-section stalked,

anvil-shaped; mesocarp moist but not juicy;

exocarp 1—1.3 mm thick; pedicels 7-20 mm

long in fruit, 0.9-1.1 mm thick at mid-point;

seeds pale yellow, 3-4.2 mm long. Quena.

Selected specimens examined : Queensland. Burke

District: Lake Mary, near Camooweal, May 1935, Blake

8882 (BRI); Julia Ck-Normanton road, 45 km N of Julia

Creek, Jul 1985, Williams 85057 (BRI); 14 km NE of

‘Eulolo’ HS, 41 km NE of McKinlay, Nov 1986, Neldner

2695 & Nicolson (BRI). North Kennedy District: Suttors

River, undated, Bowman (BRI, MEL); 20 kmN of Charters

Towers, Mar 1992, Dorney 295 (BRI). Gregory North

District: Eyre Creek, 8 km SE of Bedourie, Sep 1978, Purdie

1338 (BRI); Chartwage Bore, ‘Headingley’, May 1985,

Neldner 1779 & Stanley (BRI). Mitchell District: 12 miles

[19km] S of Tambo on Ward road, Feb 1968, Beasley (BRI);

c. 30 km S of Hughenden on Muttaburra road, Jun 1977,

McDonald 2509 (BRI). South Kennedy District: ‘Laglan’

Station, c. 105 km WNW of Clermont, Mar 1958, Smith

10321 (BRI); 26.5 km NW of Belyando Crossing on Gregory

Development road, Jun 1992, Thompson BUC619 & Sharpe

(BRI). Leichhardt District: 9 miles [14 km] WNW of

Springsure, Oct 1964, Adams 1394 (BRI, CANB);

‘Berrigurra’, c. 19 km NW of Blackwater, Duaringa shire,

Dec 1983, Anderson 3621 (BRI); ‘Homevale’ Station, Mar

1994, Champion 1029 etal. (BRI). Gregory South District:

Old Canterbury Hotel, on Betoota-Windorah road, Mar 1990,

Sandercoe 4096 (BRI); 21.8 km NW of Eromanga turnoff,

on Quilpie-Windorah road, Aug 1990, Prendergast HDP293

(BRI). Warrego District: Oakwood Station, N of Charleville,

Apr 1932, Francis s.n. (BRI); Eulo, Apr 1941, White 12016

(BRI). Maranoa District: ‘Tamanick’, 80 km SE of Mitchell,

Nov 1989, Warrian CMW396 (BRI); 44.7 km along road to

‘Combo’, S of Bollon, Jan 1998, Bean 13050 (BRI). Darling

Downs District: ‘Calala’, 10 m il es [16 km] EofMeandarra,

Apr 1960, Johnson 1622 (BRI); 2 km NW of Dalby, Apr

1994, Fensham 1259 (BRI); Mamhull, SE of Jandowae, Dec

2001, Bean 18201 (BRI). New South Wales. Northwest

Plains: 10 km NNE of Ashley on Rosedale road, Mar 1987,

Dunn 61 et al. (BRI, NSW); 0.4 km S of Gurley Ck, on

Moree-Narrabri road, Jan 1996, Bean 9503 (BRI, NSW).

Distribution and habitat : Solanum esuriale is

widely distributed in Queensland, being absent

only in coastal areas and the far north of the

state (Map 29). It occurs in all mainland states.

It inhabits heavy clay soils and occurs in several

plant communities including grassland, open

woodland of Eucalyptus coolabah or E. populnea,

open forest of Acacia harpophylla or A. cambagei.

Phenology : Flowers and fruits may be found

at any time of the year.

Notes : S. esuriale is often completely unarmed,

but sometimes bears numerous short prickles

on the stems, rarely extending to the calyx.

These prickly forms appear to be randomly

distributed and hence have no taxonomic

significance. It is closely related to S. elaeagnifolium

(see Notes under that species).

Conservation status : Widespread. Not

considered at risk.

78. *Solanum elaeagnifolium Cav., Icon. 3:

22, t. 243 (1795). Type: cultivated in

Madrid (originating from America),

undated, A.J. Cavanilles s.n. (iso: C, MA,

P-JU), n.v.Jide D’Arcy (1974).

Illustrations : Cunningham et al. (1981:

593); Symon (1981: 153).

Erect, herbaceous resprouter, 0.3-0.6 m high.

Juvenile stage unknown. Adult branchlets terete

or ridged, white or grey; prickles absent or

present, 0-30 per decimetre, straight, acicular

or broad-based, 0.5-1.5 mm long, 6-10 times

longer than wide; stellae very dense, 0.4-0.5

mm diameter, stalks 0-0.1 mm long; lateral

rays 15-18, porrect; central ray absent or

present, 0-0.5 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Adult leaves lanceolate, entire or

shallowly lobed throughout; lobes 3-6 on each

side, obtuse, lobing index 1-1.1; lamina 3-6.5

cm long, 0.8-1.4 cm wide, 3.9-5.2 t im es longer

than broad, apex acute, base cuneate, oblique

part 0-3 mm long, obliqueness index 0-6

percent; petioles 0.4-1.3 cm long, 13-25%

length of lamina, prickles absent or present.

Upper leaf surface grey-green or grey; prickles

absent; stellate hairs distributed throughout;

protostellae absent; ordinary stellae dense to

very dense, 0.05-0.2 mm apart, 0.4-0.5 mm

across, sessile; lateral rays 10-15, porrect;

central ray 0-0.5 times as long as laterals, not

Bean, Taxonomy of Solanum subg. Leptostemonum

gland-tipped; finger hairs absent; Type 2 hairs

absent. Lower leaf surface white; prickles

absent; stellae very dense, c. 0.05 mm apart,

0.3-0.6 mm diameter, stalks 0-0.2 mm long;

lateral rays 13-16, porrect; central ray 0.2-0.6

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent.

Inflorescence supra-axillary, pseudo-racemose,

common peduncle 6-21 mm long, rachis

prickles absent; 4-8-flowered, with all flowers

bisexual and 5-merous; pedicels 7-14 mm long

at anthesis, same thickness throughout,

0.7-0.9 mm thick at mid-point, prickles absent

or present. Calyx tube 3-4.5 mm long, lobes

attenuate, 3-6 mm long; prickles absent or

present at anthesis, 0-35 per flower, 1-2.5 mm

long; stellae very dense, white, 0.4-0.5 mm

across, stalks 0-0.1 mm long, lateral rays

13-20, central ray 0.1-1 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Corolla mauve or purple,

12-14 mm long, shallowly or deeply lobed,

inner surface glabrous; anthers 7-9 mm long;

ovary with stellate hairs only; functional style

11-13 mm long, erect, glabrous or with stellate

hairs only, stellae 0.6-0.7 mm across, lateral

rays 11—14, central ray 0.4-1 times as long as

laterals. Fruiting calyx with lobes more than

half length of mature fruit, prickles absent or

present, 1-2.5 mm long. Mature fruits 2-5 per

inflorescence, globular, 13-15 mm diameter,

yellowish-green, with a few scattered stellate

hairs; mesocarp moist but not juicy; pedicels

16-27 mm long in fruit, 1.2-1.5 mm thick at

mid-point; seeds brown to black, 2.9-3.5 mm

long. Silver-leaf Nightshade.

Specimens examined : Queensland. Burnett District: 1

mile [1.6 km] N of Dappil railway siding, 7 miles [11 km] N

of Gayndah, Oct 1966, Marlowe (BRI); County of Bowen,

near Gayndah, Oct 1973, James (BRI). Darling Downs

District: Elbow Valley, Dec 1959, Seton (BRI); 6 mi les [10

km] N of Oakey, May 1963, Inglis (BRI); 2 miles [3 km]

NW of Jandowae, Jan 1966, Colborn (BRI); 10.5 miles [17

km] ENE of Dalby, Nov 1971, Cook (BRI); Parish of Daadine

near Dalby, Oct 1973, James (BRI); near Jandowae, Oct

1973, James (BRI); between Oakey and Biddeston, Jan 1979,

Kay (BRI); Oakey, 50-100 m along Bacon’s Road, Jan 2000,

Menkins DDP17 (BRI); Lime vale, on Texas-Inglewood road,

May 2002, Halford Q7531 & Batianoff (BRI). Moreton

District: 3 miles [5 km] SW of Laidley, Feb 1965, Noffke

(BRI); 3 km E of Laidley, Nov 1973, James 5 (BRI); Gatton,

Dec 1977, Olsen s.n. (BRI); 5 miles [8 km] NNE of Laidley,

Feb 1979, Armstrong (BRI); Pearsons Road, Kalbar, Mar

2002, Pocknee s.n. (BRI). New South Wales. Bonshaw, Dec

1976, Sharp (BRI).

789

Distribution and habitat: Solanum

elaeagnifolium is native to southern U.S.A.,

Mexico, Argentina and Paraguay. It is

sporadically naturalised in south-eastern parts

of Queensland (Map 31). It occurs commonly

in New South Wales, Victoria and South

Australia, and sporadically in south-western

Western Australia. It is found on degraded sites,

such as disused cultivation paddocks, improved

pasture and roadsides. Soil type is apparently

unimportant, and varies from sandy to clayey types.

Phenology: Flowers have been recorded from

October to February; mature fruits in February.

Notes: S. elaeagnifolium is close to S. esuriale,

but differs by the branchlet stellae 0.4-0.5 mm

diameter (vs. 0.7-0.9 mm for S. esuriale ) with

15-18 lateral rays (8-12 lateral rays for S. esuriale),

prickles usually present on pedicels and calyx

(absent for S. esuriale ), calyx stellae with

13-20 lateral rays (vs. 6-9 for S. esuriale ) and

anthers 7-9 mm long (vs. 3.5-5 mm for S. esuriale ).

It is remarkable that the American species

S. elaeagnifolium is so similar to the Australian

species S. esuriale, to the extent that

microscopic examination is often necessary to

determine them.

S. elaeagnifolium has been declared a

noxious weed for the whole of New South Wales

(Conn 1992), but does not appear to be a serious

weed in Queensland.

79. Solanum ammophilum A.R.Bean sp. nov.

Frutex erectus parvus; ramuli aculei

sparsi, recti, ad basim lati; ramuli stellae

radiis lateralibus 8-13; folia linearia

usque anguste lanceolata, pilis type-2 in

paginis ambabus; inflorescentia 1 vel 2-

flora, floris constanter 4-meris; calycis

aculei absentes; ovarium papillis albidis

glandularibus praeditum; fructus maturi

flavovirentes, biloculares. Typus:

Queensland. Maranoa District: 73 km

from Charleville towards Bollon, 28

January 2002, A.R. Bean 18423 (holo:

BRI (1 sheet + spirit); iso: AD, CANB,

DNA, MEL, MO, NSW, distribuendi).

Erect, herbaceous resprouter, 0.2-0.4 m high.

Juvenile stage absent. Adult branchlets white,

grey or yellow; prickles absent or present,

790

Austrobaileya 6 (4): 639-816 (2004)

0-10 per decimetre, straight, broad-based,

1.5-5 mm long, 5-7 times longer than wide;

stellae dense to very dense, 0.3-0.5 mm

diameter, sessile; lateral rays 8-13, porrect;

central ray absent or present, 0-0.3 times as

long as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs dense. Adult leaves linear

or narrow lanceolate, entire; lamina 2-7 cm

long, 0.3-0.7 cm wide, 6-10 times longer than

broad, apex obtuse, base cuneate, oblique part

0-1 mm long, obliqueness index 0-3 percent;

petioles 0.3-0.8 cm long, 10-17% length of

lamina, prickles absent. Upper leaf surface

grey-green; prickles absent; stellate hairs

distributed throughout; protostellae absent;

ordinary stellae dense, 0.1-0.2 mm apart,

0.2-0.5 mm across, sessile; lateral rays 8-10,

porrect; central ray 0-0.3 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs present throughout, 0.1-0.2 mm

apart. Lower leaf surface white; prickles absent;

stellae dense to very dense, 0.05-0.1 mm apart,

0.4-0.8 mm diameter, stalks 0-0.1 mm long;

lateral rays 8-10, porrect; central ray 0-0.3

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs present

throughout, 0.05-0.1 mm apart. Inflorescence

leaf-opposed or supra-axillary, solitary or

pseudo-racemose, common peduncle absent,

rachis prickles absent; 1 or 2-flowered, with

all flowers bisexual and 4-merous; pedicels

5-17 mm long at anthesis, same thickness

throughout, 0.8-1.1 mm thick at mid-point,

prickles absent. Calyx tube 2.5-4 mm long,

lobes deltate, 2.5-6 mm long; prickles absent

at anthesis; stellae very dense, white, 0.4-0.6

mm across, stalks 0-0.1 mm long, lateral rays

8-11, central ray 0.2-0.8 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Corolla purple, 7-15 mm

long, deeply lobed, inner surface glabrous;

anthers 4.5-7.5 mm long; ovary with white

glandular papillae; functional style 8-11.5 mm

long, erect, with white glandular papillae.

Fruiting calyx with lobes less than or more than

half length of mature fruit, prickles absent.

Mature fruits 1 or 2 per inflorescence, oblate

or globular, c. 13 mm diameter, yellowish-

green, 2-locular; placenta in cross-section

sessile, elliptical; mesocarp moist but not juicy;

exocarp 0.7-0.9 mm thick; pedicels 9-18 mm

long in fruit, 0.8-1.1 mm thick at mid-point;

seeds pale yellow, 2.8-3 mm long. Fig. 42.

Specimens examined : Queensland. Warrego District:

‘Gilruth Plains’, E of Cunnamulla, May 1939, Blake 14062

(BRI); ‘Glenbar’, c. 42 [miles?] SEof Charleville,Apr 1948,

Everist 3395 (BRI, CANB); ‘Gilruth Plains’, Apr 1963,

McKee 10366 (BRI); 22 miles [35 km] E of Cunnamulla on

road to Bollon, Sep 1963, Phillips 534 (BRI); Top Cane

paddock, ‘Gilruth Plains’, Cunnamulla, Apr 1967, Barker

784 (CANB). Maranoa District: ‘Dingwall’, c. 90 miles

[145 km] SSE of Charleville, Jul 1948, Everist 3483 (BRI);

‘Calabah’, 75 miles [121 km] SE of Charleville, Mar 1962,

Ebersohn (BRI); c. 21 km W of Boatman road, ENE of

Cunnamulla, Dec 1998, Bean 14485 (BRI, MEL); 73 km

from Charleville towards Bollon, Jan 2002, Bean 18426 (AD,

BRI, CANB); 70 km from Charleville towards Bollon, Jan

2002, Bean 18434 (BRI. MEL); 64 km from Charleville

towards Bollon, Jan 2002, Bean 18440 (BRI). New South

Wales. North West Plains: Bourke district, Jan 1951 ,Darley

15 (NSW); ‘Widgee Downs’, Weilmoringle, Jun 1967,

Jensen B6 (NSW); 32 mil es [52 km] N of Bourke on Mitchell

Highway, Apr 1977, Moore 7502 (CANB); ‘Lila Springs’,

c. 25 km SSE of Enngonia, Feb 1978, Barker A2A (NSW).

Distribution and habitat : Solanum

ammophilum is found in the Charleville and

Cunnamulla districts of Queensland (Map 27),

and near Bourke in New South Wales. It

inhabits low open woodland with Eucalyptus

melanophloia, and often with Acacia aneura,

Angophora melanoxylon, E. populnea, Triodia

sp. on yellow to red sandy soils, often adjacent

to shrubland areas featuring Grevillea

juncifolia.

Phenology: Flowers are recorded from

September to May; mature fruits from January

to July.

Notes: S. ammophilum is close to S. coactiliferum,

but differs by the leaves 6-10 times longer than

wide (4-6 times for S. coactiliferum)', branchlet

stellae with 8-13 lateral rays; leaf stellae with 8-10

lateralrays (vs. 7 or 8 lateralrays for S', coactiliferum)',

stellae stalks <0.1 mm long (vs. 0.1-0.2 mm

long for S. coactiliferum ); the corolla glabrous

on the inner surface (stellate-hairy for

S. coactiliferum), and the presence of Type 2

hairs on the lower leaf surface.

The glandular papillae on the ovary and

lower half of style are unique among

Queensland species. These papillae persist on

the fruits, and give them a distinctly sticky

texture.

Conservation status: Moderately widespread.

Not considered at risk.

Etymology: from the Greek ammo- (of the

Bean, Taxonomy of Solanum subg. Leptostemonum

791

Fig. 42. Solanum ammophilum. A. flowering branchlet x 1.5. B. flower x 3. C. ovary and style x 6. D. ovary, bearing

glandular papillae x 12. E. stellate hair, upper leaf surface x 100. F. mature fruit, calyx and pedicel x 2. G. transverse section

of fruit x 3. all from Bean 18423.

792

sand) and -philus (loving). This species occurs

on soils that are much sandier than the

prevalent soil type in the area.

80. Solanum sturtianum F.Muell., Trans.

Philos. Soc. Victoria 1: 18-19 (1854);

Hook. J. Bot. & Kew Gard. Misc. 8: 166

(1856). Type: interior of South Australia,

1844-46, C. Sturt No. 87 (lecto: MEL

[MEL 11651]), here chosen.

Illustrations : Cunningham et al. (1981:

589); Symon (1981: 205); P.S. Green,

Curtis’s Botanical Magazine 177: t. 543

(1969).

Erect, rhizomatous perennial shrub, 0.3-1.5 m

high. Juvenile stage unknown. Adult branchlets

white or grey; prickles absent or present, 0-10

per decimetre, straight, broad-based, 2-4 mm

long, 5-7 times longer than wide; stellae very

dense, 0.3-0.5 mm diameter, sessile; lateral

rays 12-16, porrect; central ray absent; finger

hairs absent; Type 2 hairs absent. Adult leaves

lanceolate, entire; lamina 4.5-7 cm long,

1-1.8 cm wide, 3.8-5.7 times longer than

broad, apex obtuse or acute, base cuneate or

obtuse, oblique part 0-3 mm long, obliqueness

index 0-6 percent; petioles 0.6-1.3 cm long,

15-25% length of lamina, prickles absent or

present. Upper leaf surface grey-green or grey;

prickles absent; stellate hairs distributed

throughout; protostellae absent; ordinary stellae

density moderate to dense, 0.1-0.2 mm apart,

0.15-0.3 mm across, sessile; lateral rays

13-15, porrect; central ray absent; finger hairs

absent; Type 2 hairs absent. Lower leaf surface

white or yellowish; prickles absent; stellae very

dense, 0.05-0.1 mm apart, 0.2-0.3 mm

diameter, sessile; lateral rays 13-16, porrect;

central ray absent; finger hairs absent; Type 2

hairs absent. Inflorescence supra-axillary,

pseudo-racemose, common peduncle 3-5 mm

long, rachis prickles absent; 5-7-flowered,

strongly or weakly andromonoecious, flowers

5-merous; pedicels 4-8 mm long at anthesis,

same thickness throughout, 0.5-0.8 mm thick

at mid-point, prickles absent. Calyx tube

1.5-3 mm long, lobes deltate, 1.5-2.5 mm long;

prickles absent at anthesis; stellae very dense,

yellow or white, 0.15-0.25 mm across, sessile,

lateral rays 12-15, central ray absent; finger

hairs absent; Type 2 hairs absent. Corolla

mauve or purple, 8-10 mm long, shallowly

lobed, inner surface glabrous; anthers 4.5-5.5

Austrobaileya 6 (4): 639-816 (2004)

mm long; ovary with stellate and Type 2 hairs;

functional style 7-8 mm long, erect, with

stellate and Type 2 hairs, stellae 0.15-0.2 mm

across, lateral rays 6-13, central ray 1-1.5 times

as long as laterals. Fruiting calyx with lobes

less than half length of mature fruit, prickles

absent. Mature fruits 1-3 per inflorescence,

globular, 9-13 mm diameter, brown or yellow;

mesocarp dry; pedicels 7-12 mm long in fruit,

0.8-1 mm thick at mid-point; seeds brown to

black, 4-5.1 mm long. Thargomindah

nightshade. Fig. 8.

Specimens examined : Queensland. Gregory South

District: South Galway, Jun 1949, Everist 4042 (BRI); c.

61 km W of Eromanga, Sep 1989, Wilson 437 & Pickering

(AD, BRI, MO, NSW); 76.5 km from Thargomindah, Sep

1990, Zalucki (BRI); W of Eromanga, Aug 1997, Anderson

5070 (BRI). Warrego District: Thargomindah, Sep 1896,

Maiden (BRI, NSW); 5 miles [8 km] S of Thargomindah,

May 1910, Little (BRI); Norley Station [c. 25 km NNE of

Thargomindah], May 1919, Watts (BRI); ‘Thargomindah’

Station, Oct 1950, Grummit (BRI); 19 km S of Thagomindah,

Nov 1954, Smith 6055 (BRI); Thargomindah-Hungerford

Stock Route, Feb 1965, Carr (BRI); c. 11 km SW of

Thargomindah, on road to Hungerford, Sep 1973, Henderson

H2060 & Boyland (AD, BRI, SP, US); Grey Range, c. 74

km from Thargomindah on road to Noccundra, Jul 1977,

Purdie 735 (BRI); 12 km W of Thargomindah, May 1978,

Olsen 760 & Boyland (BRI); 27.1 km W of Pinidary turnoff,

on Thargomindah-Nockatunga road, Sep 1990, Prendergast

HDP341 (BRI). New South Wales. North Far Western

Plains: 78.1 km from ‘Olive Downs’ HS via Jump-up Loop

road, ENE of Tibooburra, Sep 1989, Coveny 13583 et al.

(AD, BRI, MO, NSW).

Distribution and habitat: In Queensland,

Solanum sturtianum is confined to the far south

western parts, especially around Thargomindah

(Map 26). It also occurs widely in arid parts of

New South Wales, South Australia, Western

Australia and Northern Territory. It grows on

plains or stony ridges, in chenopod shrubland

or Acacia open woodland. Associated species

include Acacia aneura, A. cambagei and Senna

spp.

Phenology: Flowers are recorded from June to

October; mature fruits from May to November,

plus a single record from February.

Notes: Brown (1849) stated that Captain Sturt

“placed at my disposal the collection of plants

formed in his recent expedition ....”, and this

explains the presence of a syntype at BM.

Brown {loc. cit.) named numerous new species

from this collection, but decided not to name

several species that he considered were probably

Bean, Taxonomy of Solanum subg. Leptostemonum

new due to “the incomplete state of the

specimens”. Presumably Solanum sturtianum

was one of these. Mueller named S. sturtianum

in 1854.

Symon (1981) chose the specimen

residing in BM as the lectotype of S. sturtianum,

“in view of the quality of the specimen”.

Mueller never visited England, nor did he

borrow specimens from BM. Since Mueller

could never have seen the BM specimen, it

cannot be considered in the choice of the

lectotype. I therefore overturn Symon’s

lectotypification, which is contrary to Article

9.10 (Greuter et al. 2000), and choose instead

MEL 11651 as lectotype.

Conservation status: Widespread. Not

considered at risk.

81. Solanum oligacanthum F.Muell., Trans.

Philos. Soc. Victoria 1: 18-19 (1854).

Type: Central Australia, 1844-46, C.

Sturt (holo: MEL [MEL11991]).

Illustrations: Cunningham et al. (1981:

591); Symon (1981: 207).

Erect, herbaceous resprouter or rhizomatous

perennial shrub, 0.2-0.6 m high. Juvenile stage

unknown. Adult branchlets white or grey;

prickles absent or present, 0-30 per decimetre,

straight, acicular, 0.5-13 mm long, 8-11 times

longer than wide; stellae very dense, 0.2-0.4

mm diameter, stalks 0-0.1 mm long; lateral

rays 13-15, porrect or multiradiate; central ray

absent or present, 0-0.7 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Adult leaves broadly ovate

or orbicular, entire; lamina 1-2.5 cm long,

0.9-2.8 cm wide, 0.8-1.4 times longer than

broad, apex obtuse, base obtuse or cordate,

oblique part 0-1 mm long, obliqueness index

0-6 percent; petioles 0.1-0.3 cm long, 6-18%

length of lamina, prickles absent. Upper leaf

surface green to grey; prickles 0-2, straight,

acicular, 1-7 mm long, prickles absent or

present on midvein only; stellate hairs

distributed throughout; protostellae absent;

ordinary stellae density moderate to dense,

0.1-0.25 mm apart, 0.15-0.3 mm across,

sessile; lateral rays 12-16, porrect or

multiradiate; central ray 0.1-0.5 times as long

as laterals, not gland-tipped; finger hairs

793

absent; Type 2 hairs absent. Lower leaf surface

white; prickles absent; stellae very dense, c.

0.05 mm apart, 0.2-0.35 mm diameter,

sessile; lateral rays 9-18, porrect or

multiradiate; central ray 0.3-0.8 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Inflorescence

supra-axillary, pseudo-racemose, common

peduncle 0-4 mm long, rachis prickles absent

or present; 4-6-flowered, flowers 5-merous;

pedicels 2.5-7 mm long at anthesis, same

thickness throughout, 0.5-0.8 mm thick at mid¬

point, prickles absent. Calyx tube 2-3 mm long,

lobes deltate, 2.5-3.5 mm long; prickles absent

at anthesis; stellae very dense, yellow or white,

0.3-0.4 mm across, stalks 0-0.2 mm long,

lateral rays 10-16, central ray 0.7-1.3 times as

long as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Corolla purple,

8-11 mm long, rotate, inner surface glabrous;

anthers 3.5-5.5 mm long; ovary with Type 2

hairs only; functional style 7-8 mm long, erect,

with Type 2 hairs only. Fruiting calyx with

lobes less than half length of mature fruit,

prickles absent. Mature fruits 1-6 per

inflorescence, globular, 8-11 mm diameter,

brown or yellow; mesocarp moist but not juicy;

pedicels 3-8 mm long in fruit, 0.6-0.9 mm

thick at mid-point; seeds brown to black,

3.6-5.3 mm long.

Specimens examined : Queensland. Gregory South

District: Cuppa Creek, 12 miles [19 km] W of Durrie, Jan

1937, Everist & Smith 82 (BRI); N of Betoota, Aug 1953,

Davidson 433 (BRI); ‘Tanbar’ Station, W of Cooper’s Creek,

Feb 1968, Lewis (CANB); east of Simpson Desert, Jul-Aug

1969, Cockburn BPS3 (BRI); 8 km from Birdsville on new

Betoota road, Sep 1977, Purdie 1139 (BRI); NW edge of

Lake Yamma Yamma, Mar 1990, Sandercoe 4065 (BRI);

50 km W of Birdsville, on track to Poeppel Comer, Sep 1998,

Halford Q3651 (BRI). WarregoDistrict: 38 miles [61 km]

S of Thargomindah, Feb 1968, Lewis (CANB). New South

Wales. North Far Western Plains: Ford Grey Basin, c. 100

km NW of Tibooburra, May 1980, Paijmans 3456 (CANB);

S edge of Lake Pinaroo near Fort Grey camping area, Sturt

N.P., Sep 1989, Coveny 13475 et al. (AD, BRI, MO, NSW).

Distribution and habitat : In Queensland,

Solanum oligacanthum is confined to the

extreme south-west of the state, near Birdsville

(Map 28). It is also found in adjacent areas of

New South Wales and South Australia. It grows

in shrubland, on creek channels, claypans, lake

margins and interdunal flats. Soils may be

sandy or clayey.

794

Phenology: Flowers are recorded for January,

March, July, August and September; mature

fruits in January and September.

Notes: S. oligacanthum is immediately

recognisable by its adult leaves. They are entire,

orbicular or broadly ovate, and less than 25 mm

long. S. orbiculatum from the Northern

Territory and Western Australia also has

orbicular or broadly ovate leaves, but they are

larger. It also has longer petioles and larger

fruits.

Conservation status: Widespread. Not

considered at risk.

Group 27F (S. echinatum group), here

defined; related to Group 27 ( S . ellipticum

group) of Whalen (1984).

Adult leaves entire to shallowly-lobed (100%);

fruiting calyx prickly, accrescent, covering the

fruit completely (100%); habit prostrate or

procumbent (100%); ovary glabrous (100%);

mature fruits brown, oblate, 4-locular (100%);

common peduncle >25 mm long (100%);

corolla rotate, inner surface glabrous (100%);

seeds brown to black (100%).

3 species endemic to Australia; 2 species

indigenous to Queensland.

82. Solanum echinatum R.Br., Prodr. 447

(1810). Type: [Northern Territory.] North

Island, Sir Edward Pellew Group, 16

December 1802, R. Brown (holo: BM; iso:

MEL [MEL 12416]).

Solanum seitheae Symon, J. Adelaide Bot.

Gard. 4: 201 (1981), syn. nov. Type:

Queensland. Burke District: 116 km SW

of Normanton, top of Donors Hills, 29

May 1967, D.E. Symon (holo: AD; iso:

BRI, CANB, K, NSW).

Illustrations: Symon (1981: 197); Symon

(1981: 203), as S. seitheae.

Prostrate, rhizomatous perennial shrub,

0.1-0.3 m high. Juvenile stage unknown. Adult

branchlets yellow, rusty or brown; prickles

200-660 per decimetre, straight, acicular, 1-8

mm long, 10-20 times longer than wide; stellae

very dense, 0.6-0.8 mm diameter, stalks 0-0.8

mm long; lateral rays 7 or 8, porrect; central

ray 0.8-1.8 times as long as laterals, not gland-

Austrobaileya 6 (4): 639-816 (2004)

tipped; finger hairs absent; Type 2 hairs absent.

Adult leaves ovate, entire or shallowly lobed

throughout; lobes 3 or 4 on each side, obtuse,

lobing index 1-1.1; lamina 3.5-9.5 cm long,

2- 5.5 cm wide, 1.7-2 times longer than broad,

apex acute, base obtuse or cordate, oblique part

0-3.5 mm long, obliqueness index 0-9 percent;

petioles 1.1-5 cm long, 25-55% length of

lamina, prickles present. Upper leaf surface

grey-green; prickles 0-12, straight, acicular,

3- 5 mm long, prickles absent or present on

midvein only or present on midvein and lateral

veins; stellate hairs distributed throughout;

protostellae absent; ordinary stellae density

moderate to dense, 0.1-0.4 mm apart, 0.4-0.8

mm across, stalks 0-0.2 mm long; lateral rays

7-9, porrect; central ray 1-1.5 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Lower leaf surface white

or yellowish; prickles 0-9, straight, acicular,

absent or present on midvein only or present

on midvein and lateral veins; stellae dense to

very dense, 0.05-0.1 mm apart, 0.6-0.9 mm

diameter, stalks 0-0.2 mm long; lateral rays 7

or 8, porrect; central ray 1-1.5 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Inflorescence supra-

axillary, pseudo-racemose, common peduncle

25-38 mm long, rachis prickles present;

2-5-flowered, with all flowers bisexual and

5-merous; pedicels 3-9 mm long at anthesis,

same thickness throughout or markedly thicker

distally, 1-1.2 mm thick at mid-point, prickles

present. Calyx tube 3-6 mm long, lobes rostrate

or attenuate, 3-6 mm long; prickles present at

anthesis, 60-300 per flower, 1-3 mm long;

stellae very dense, yellow, brown or rusty,

0.7-0.9 mm across, stalks 0-1.1 mm long,

lateral rays 7 or 8, central ray 1-1.8 t im es as

long as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Corolla mauve or

purple, 9-14 mm long, rotate, inner surface

glabrous; anthers 4.5-6 mm long; ovary

glabrous; functional style 7-7.5 mm long, erect,

glabrous. Fruiting calyx tube accrescent,

enclosing most or all of mature fruit, prickles

1-8 mm long. Mature fruits 1-4 per

inflorescence, oblate, 15-20 mm diameter,

brown, 4-locular; placenta in cross-section

stalked, anvil-shaped; mesocarp moist but not

juicy; exocarp 2-2.3 mm thick; pedicels

10—18 mm long in fruit, 1-1.8 mm thick at

mid-point; seeds brown to black, 2-2.3 mm

long. Fig. 9.

Bean, Taxonomy of Solarium, subg. Leptostemonum

Specimens examined : Queensland. Burke District:

Leichhardt River, Jun 1935, Blake 9292 (BRI); ‘Adels

Grove’, via Camooweal, Mar 1947, DeLestang 347 (BRI);

9 miles [14 km] W of ‘Riversleigh’ Station, Jun 1948, Perry

1440 (BRI, CANB); Calton Hills, 45 miles [72 km] N of Mt

Isa, May 1963, Gittins 762 (BRI); 1 mile [1.6 km] E of

‘Wernadinga’, May 1967, Symon 4995 (AD, BRI, CANB,

K); 17 miles [27 km] W of Nicholson River crossing, Jun

1967, Symon 5017 (AD, BRI); 14 km from Gunpowder on

Quamby road, Oct 1972, Althofer 288 (BRI); 60 km E of Mt

Isa on Mt Isa-Cloncurry road, Aug 1973, Swan 117 (BRI);

c. 8 miles [13 km] NW of ‘Riversleigh’ station on road to

‘Lawn Hill’, Jul 1974, Ollerenshaw 1317 & Kratzing (BRI,

CANB); Burketown, c. 24 km on Doomadgee Road, Jul

1974, Ollerenshaw 1354 & Kratzing (BRI, CANB); 120

km W of Mt Isa, Jul 1974, Swan 94 (BRI); 60 miles [97 km]

NE of Mt Isa on road to Julius Dam, Aug 1974, Swan 126

(BRI, CANB); Dugald River Lode in vicinity of One Tree

Hill, NW of Quamby, May 1976, Simon 3010 & Farrell

(BRI); 24 km N of Mt Isa, Dec 1986, Harris 107 (BRI); W

side of Lake Moondarra, 16 km NE of Mt Isa, Dec 1989,

Harris 432 (BRI); 0.7 km NW of Magazine Hill, 10 km N

of Silver Star Mine, Apr 1991, Jones 403A (BRI);

Musselbrook Gorge, Lawn Hill N.R, Jul 1992, McDonald

KM 1117 & Johnson (BRI); main highway 109 km from

Normanton, 88 km from Burke and Wills Roadhouse, Jul

1993, Alcock 11285 (AD, BRI); bank of Nicholson River,

near Kingfisher Camp, Apr 1996, Milson JM1108 (BRI);

Hilton shaft, Mt Isa shire, Jun 1996, Barrs SB28 (BRI); Lawn

Hill, plateau above canoe portage area, Jul 1998, Symon

15791 (AD, BRI, L, NY).

Distribution and habitat : Solanum echinatum

is found in the northwestern corner of

Queensland, as far south as Mt Isa (Map 28).

It is also common in the Northern Territory,

including the islands of the Gulf of Carpentaria.

It grows on stony ridges or gullies in low

eucalypt woodland, on a variety of substrates,

including laterite.

Phenology: Flowers are recorded from April

to August, with a single record in December.

Fruits are recorded from March to August.

Notes: Closely related to S. longissimum, see

Notes under that species. Both S. echinatum

and S. longissimum have numerous conspicuously

stalked stellae, especially on the branchlets and

rachis, but also to a lesser degree on the upper

and lower leaf surfaces (Fig. 9). The calyx

prickles are extremely numerous, and there are

always some smaller prickles with stellae

mounted at the top (or, from the other

perspective, long-stipitate stellae which are

becoming prickles) (Fig. 4). By contrast, the

stellae of S. senticosum (which grows in the

same area) have stalks all about the same length

with none conspicuously long. Also, none of

the calyx prickles of S. senticosum have stellae

795

mounted at the top. Hence S. echinatum and

S. longissimum can be readily separated from

S. senticosum in the absence of fruiting

material.

The type of S. seitheae is indistinguishable

from the type of S. echinatum. They both have

fruits of about the same size, a dense rusty

tomentum, some leaves with shallow lobes, and

petioles less than half as long as the lamina.

There are additional unnamed taxa

related to S. echinatum in the Northern

Territory, and S. wilkinsii S.Moore probably

deserves to be reinstated.

Conservation status: Widespread. Not

considered at risk.

83. Solanum longissimum A.R.Bean sp. nov.

Frutex prostratus vel procumbens; ramuli

aculeis 3-80 per decimetrum; folia adulta

ovata usque late ovata, integra, petiolis

longitudine 67-116% laminae; inflorescentia

pseudo-racemosa, pedunculo communi 35-

67 mm longo; calycis aculei saepe non

manifesti sub anthesi; corolla rotata;

ovarium glabrum; calyx fructifer

densissime aculeatus, accrescens, fructum

includens; fructus maturi oblati, brunnei,

pedicellis 20-30 mm longis. Typus:

Queensland. Cook District: Pannikan

Springs area, 29 km W of Mungana, 26

January 1993, A.R. Bean 5617 & P.I.

Forster (holo: BRI; iso: DNA).

Prostrate, rhizomatous perennial shrub,

0.15-0.3 m high. Juvenile stage unknown.

Adult branchlets rusty or brown; prickles

3-80 per decimetre, straight, acicular or broad-

based, 1-5 mm long, 6-10 times longer than

wide; stellae dense, 0.6-1.1 mm diameter,

stalks 0-0.6 mm long; lateral rays 7 or 8,

porrect; central ray 1-1.8 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Adult leaves ovate or

broadly ovate, entire; lamina 3-8 cm long,

1.7-4 cm wide, 1.6-2.1 times longer than

broad, apex acute, base cuneate or obtuse,

oblique part 0-5 mm long, obliqueness index

0-12 percent; petioles 2.4-5.5 cm long,

65-115% length of lamina, prickles absent.

Upper leaf surface green or grey-green; prickles

absent; stellate hairs distributed throughout;

protostellae absent; ordinary stellae sparse to

796

dense, 0.25-0.5 mm apart, 0.6-0.7 mm across,

stalks 0-0.1 mm long; lateral rays 6-8, porrect;

central ray 1-1.8 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Lower leaf surface yellowish; prickles

absent; stellae dense, 0.15-0.3 mm apart,

0.6-0.8 mm diameter, stalks 0-0.2 mm long;

lateral rays 7 or 8, porrect; central ray 0.8-1.5

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent.

Inflorescence supra-axillary, pseudo-racemose,

common peduncle 35-67 mm long, rachis

prickles present; 3 or 4-flowered, flowers

5-merous; pedicels 4-6 mm long at anthesis,

same thickness throughout or markedly thicker

distally, 0.5-0.9 mm thick at mid-point,

prickles present. Calyx tube 2.5-3.5 mm long,

lobes deltate, 1-2.5 mm long; prickles absent

or present at anthesis, zero or 20-100 per

flower, 1-1.5 mm long; stellae very dense,

yellow, 0.5-0.9 mm across, stalks 0-1.5 mm

long, lateral rays 5-8, central ray 1-1.8 times

as long as laterals, not gland-tipped; finger

hairs absent; Type 2 hairs absent. Corolla

purple, 7-10 mm long, rotate, inner surface

glabrous; anthers 3.5-4.5 mm long; ovary

glabrous; functional style 6-6.5 mm long, erect,

glabrous. Fruiting calyx tube accrescent,

enclosing most or all of mature fruit, prickles

1-5 mm long. Mature fruits 1 or 2 per

inflorescence, oblate, brown; mesocarp moist

but not juicy; pedicels 20-30 mm long in fruit,

0.9-1.2 mm thick at mid-point. Fig. 4, 43.

Specimens examined : Northern Territory. Nicholson

River area, Tin Hole Creek, Jun 1974, Maconochie 2025

(BRI, CANB, DNA); 4 km W of ‘Newcastle Waters’, Feb

1980, Latz 8335 (BRI, DNA). Queensland. Burke District:

16 km (by road) W of Musselbrook Mining Camp on road to

Border Waterhole, Apr 1995, Thomas MRS 179 & Johnson

(AD, BRI). Cook District: Gugu Yalungi Main Camp, Laura,

Mar 1975, Hyland 8089 (BRI, CANB, QRS).

Distribution and habitat : Solanum

longissimum has a scattered distribution across

northern Queensland (Map 31) and adjacent

areas of the Northern Territory. It is recorded

from sandstone ridges or lateritic rises

supporting low eucalypt woodland with spinifex

(Triodia spp.) in the understorey.

Phenology : Flowers are recorded for January,

February, March and June. Fruits are recorded

for April.

Austrobaileya 6 (4): 639-816 (2004)

Notes: Related to S. echinatum, but differing

by the sparsely prickly branchlets with 3-80

prickles per dm (vs. 200-660 for S. echinatum );

branchlet prickles 6-10 times longer than wide

(10-20 times for S. echinatum ); the dense

stellate hairs on the branchlets (vs. very dense

for S. echinatum ); the very long petioles, 65-115%

of lamina length (vs. 25-55% for S. echinatum)’,

and the deltate calyx lobes 1-2.5 mm long (vs.

rostrate to attenuate, 3-6 mm long for S. echinatum).

Conservation status : Data deficient.

Etymology: From the Latin longissimus

meaning ‘longest’, in reference to the

exceptionally long petioles of this species,

sometimes exceeding the lamina in length.

Group 28 (S. dioicum group) of Whalen

(1984)

One bisexual flower per inflorescence (100%);

male flowers smaller and less prickly than

bisexual flowers (100%); mature fruits large,

23-30 mm diameter, green to yellowish-green

(100%); fruiting calyx prickly, accrescent

(100%); corolla rotate (100%); upper and lower

leaf surfaces with dense to very dense

indumentum (100%); seeds brown to black

(100%); stigma of bisexual flower forked (50%).

18 species endemic to Australia; 2 species

indigenous to Queensland.

84. Solanum chippendalei Symon, J. Adelaide

Bot. Gard. 4: 272 (1981). type: Western

Australia, base of the Sir Frederick

Range, 1 August 1962, D.E. Symon 2272

(holo: AD; iso: AD, CANB, PERTH).

Illustration: Symon (1981: 273)

Erect, rhizomatous perennial shrub, 0.4-1 m

high. Juvenile stage unknown. Adult branchlets

terete or ridged, white; prickles 35-400 per

decimetre, straight, acicular, 1-9 mm long,

7-11 times longer than wide; stellae very dense,

0.8-1.3 mm diameter, stalks 0-1.2 mm long;

lateral rays 7 or 8, porrect or ascending; central

ray present, 1-1.5 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Adult leaves lanceolate, elliptical or

ovate, shallowly to deeply lobed throughout,

rarely entire; lobes 2-4 on each side, obtuse,

lobing index 1-5; lamina 4-9.5 cm long,

1.4-3.7 cm wide, 1.6-3.5 times longer than

Bean, Taxonomy of Solanum subg. Leptostemonum

797

QUEENSLAND HERBARIUM (BRfV

Australia

ao 564031

a of QjMfnlitnd

m t-Br-

qurusuw icjauxiktf tail)

irws wore *it. sen •,

ceeih

Parent*-! Strict In, SSi» wt of Mmi,

fti undttona ilopa wlHi Euc. ^UbortaniH. TrlWaiU

SO" ska IotaL

houMHit iht Jus. Mof,, flown bToo,

Haro at 4 luv

*-W

Gueenslaftd Hartrarium |SR!|i

■So/tmt/mr /oy /£sif!&*v /f. A,

tw. A. 4. Cat* /S<T 2ao_2

Fig. 43. Holotype of Solanum longissimum.

798

broad, apex obtuse or acute, base cuneate or

obtuse, oblique part 0-5 mm long, obliqueness

index 0-8 percent; petioles 0.8-2.1 cm long,

20-30% length of lamina, prickles present.

Upper leaf surface grey-green or grey; prickles

0-8, straight, acicular or broad-based, 3-10 mm

long, prickles absent or present on midvein only

or present on midvein and lateral veins; stellate

hairs distributed throughout; protostellae

absent; ordinary stellae dense, 0.15-0.3 mm

apart, 0.7-1 mm across, stalks 0-0.2 mm long;

lateral rays 6-9, porrect or ascending; central

ray 1-2.5 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Lower leaf surface white or yellowish; prickles

0-5, straight, acicular or broad-based, prickles

absent or present on midvein only or present

on midvein and lateral veins; stellae dense to

very dense, 0.1-0.2 mm apart, 0.8-1.2 mm

diameter, stalks 0-0.5 mm long; lateral rays

6-8, porrect; central ray 1-1.8 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Inflorescence supra-

axillary, pseudo-racemose, common peduncle

0-3 mm long, rachis prickles present; 7—11-

flowered, strongly andromonoecious, flowers

5-merous. Flowers markedly dimorphic, with

larger pricklier basal flower(s); pedicels 5-23

mm long at anthesis, same thickness

throughout, 0.7-1.3 mm thick at mid-point,

prickles absent or present. Calyx prickles on

basal bisexual flowers 40-125, 2-6 mm long;

prickles on distal male flowers 0-15, 1-3 mm

long. Calyx tube 3-5 mm long, lobes attenuate,

6.5-11 mm long; stellae very dense, white,

0.7-0.9 mm across, stalks 0-1.5 mm long,

lateral rays 7 or 8, central ray 1—1.5 times as

long as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Corolla purple,

10-20 mm long, rotate, inner surface glabrous;

anthers 4-6.5 mm long; ovary glabrous;

functional style 10-12.5 mm long, erect,

glabrous. Fruiting calyx with lobes more than

half length of mature fruit, prickles 2-6 mm

long. Mature fruits 1 per inflorescence,

globular, 23-30 mm diameter, yellowish-green

or green; mesocarp moist but not juicy; pedicels

23-48 mm long in fruit, 1.3-2.2 mm thick at

mid-point; seeds brown to black, 3.3-3.6 mm

long.

Selected specimens examined : Queensland. Burke

District: Leichhardt River, Apr 1935, Blake 8722 (BRI); 2

Austrobaileya 6 (4): 639-816 (2004)

miles [3 km] N of Selwyn, May 1963, Gittins 716 (BRI);

1.6 km E of Mary Kathleen on road to Cloncurry, Apr 1974,

Hockings 49 (BRI); 1.6 km S of Mt Isa on Urandangi road,

Jul 1974, Kratzing & Ollerenshaw 1148 (BRI, CANB); 21.2

km N of Wills River on the Duchess-Mt Isa road, Sep 1990,

Wilson 639 & Rowe (AD, BRI, NSW); c. 8 km along road to

Lady Loretta Mine, turnoff c. 65 km from Mt Isa, Aug 1993,

Lolly 101 (BRI, CANB, DNA); 16 km (by road) W of

Musselbrook Mining camp on road to Border Waterhole, 175

km N of Camooweal, Apr 1995, Thomas MRS 174 &

Johnson (BRI); 106 km N of Dajarra on Mt Isa road, May

1997, Forster PIF21159 & Booth (BRI, DNA); Barkly

Highway, 52 km NW of Mt Isa, Jul 1998, Symon 15787

(AD, BRI, L); Spring Creek, NW of Mt Isa, Apr 2001,

McDonald KRM827 (AD, BRI). Gregory North District:

between Dajarra and Duchess, Jun 1978, Althofer 8350

(BRI); 15 km NE of ‘Burnham’, Jun 1979, Purdie 1551

(BRI); Boulia-Winton road, 112 km E of Middleton, Jul

1981, Williams 81188 (BRI).

Distribution and habitat : In Queensland,

Solanum chippendalei is confined to the north¬

west between Winton and Lawn Hill (Map 30).

It is also widely distributed in semi-arid parts

of Northern Territory and Western Australia.

It grows commonly in low open eucalypt

woodland with Spinifex ( Triodia spp.) in the

understorey, on a variety of substrates. It has

also been recorded from alluvium in association

with Eucalyptus camaldulensis.

Phenology: Flowers are recorded between

January and October; fruits between February

and October.

Notes: In this species, the basal flower of each

inflorescence is larger and pricklier than all

other flowers, and only the basal flower

develops into a fruit. It is closely allied to

S. melanospermum F.Muell., a species of

restricted occurrence in the Northern Territory.

Conservation status: Widespread. Not

considered at risk.

85. Solanum carduiforme F.Muell., Fragm. 2:

163 (1861). Type: [Queensland. Burke

District:] Gulf of Carpentaria, Nicholson

River, 21 August 1856, F. Mueller (lecto:

K; isolecto: MEL), fide Symon (1981).

Illustration: Symon (1981: 301)

Erect, rhizomatous perennial shrub, 0.5-1 m

high. Juvenile stage unknown. Adult branchlets

terete or ridged, white, grey or rusty; prickles

140-600 per decimetre, straight, acicular,

Bean, Taxonomy of Solarium subg. Leptostemonum

1-11 mm long, 10-14 times longer than wide;

stellae very dense, 0.25-0.5 mm diameter,

stalks 0-0.1 mm long; lateral rays 7-11, porrect

or ascending; central ray present, 0.7-2 times

as long as laterals, not gland-tipped; finger

hairs absent; Type 2 hairs absent. Adult leaves

elliptical or ovate, deeply lobed throughout;

lobes 3-5 on each side, obtuse, lobing index

2.5-12; lamina 5-7.5 cm long, 2-3 cm wide,

1.9-3 times longer than broad, apex obtuse or

acute, base cuneate or obtuse, oblique part 4-7

mm long, obliqueness index 6-10 percent;

petioles 0.7-1.4 cm long, 15-25% length of

lamina, prickles present. Upper leaf surface

grey-green or grey; prickles 4-55, straight,

acicular, 1-10 mm long, prickles present on

midvein only or present on midvein and lateral

veins; stellate hairs distributed throughout;

protostellae absent; ordinary stellae dense to

very dense, 0.05-0.15 mm apart, 0.2-0.35 mm

across, sessile; lateral rays 7 or 8, porrect;

central ray 0.2-1.2 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Lower leaf surface white or

yellowish; prickles 6-70, straight, acicular,

present on midvein only or present on midvein

and lateral veins; stellae very dense, c. 0.05

mm apart, 0.25-0.4 mm diameter, stalks 0-0.1

mm long; lateral rays 7 or 8, porrect; central

ray 0.2-1.2 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Inflorescence supra-axillary, pseudo-racemose

(male) or solitary (female), common peduncle

14-18 mm long, rachis prickles present;

12-28-flowered (male), with all flowers

unisexual (dioecious species), flowers 5-merous.

Flowers markedly dimorphic, with larger

pricklier basal flower(s); pedicels 2-6 mm long

at anthesis, same thickness throughout, 0.4-0.8

mm thick at mid-point, prickles absent or present.

Calyx prickles on basal female flower 300-600,

1-5 mm long; prickles on distal male flowers

18-30,1-4 mm long. Calyx tube 2-4 mm long,

lobes deltate, 2-3.5 mm long; stellae very

dense, white, brown or rusty, 0.3-0.4 mm

across, stalks 0-0.1 mm long, lateral rays 7 or

8, central ray 0.7-1.4 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Corolla purple, 11-14 mm long

(male), rotate, inner surface densely stellate-

hairy; anthers 4-4.5 mm long; ovary glabrous;

functional style 9.5-12 mm long, erect,

799

glabrous; stigma forked. Fruiting calyx tube

accrescent, enclosing most or all of mature fruit,

prickles 3-10 mm long. Mature fruits 1 per

inflorescence, globular, c. 25 mm diameter;

pedicels 3-5 mm long in fruit, 1.5-2 mm thick

at mid-point.

Specimens examined : Queensland. Burke District: Lawn

Hill, Oct 1887, Hann (BRI); Lawn Hill Gorge, Aug 1979,

Farrell 922 (AD, BRI, CANB, K, MO); Lawn Hill N.P., Jul

1985, Williams 85082 (BRI); Musselbrook Creek Gorge,

27.6 km by road NE of Musselbrook Mining Camp, Apr 1995,

Thomas MRS614 & Johnson (AD, BRI); Amphitheatre, 40

km N of Musselbrook Mining Camp, May 1995, Johnson

MRS787 & Thomas (BRI); ‘Bowthom’, Jun 2001, Fens ham

4591 (BRI). Cook District: c. 50 km SW of Forsayth, anno

1998, Hughes (BRI); Cobbold Gorge, Apr 1999, Wannan

1187 (BRI, CANB, NSW).

Distribution and habitat : In Queensland,

Solanum carduiforme is known from the Lawn

Hill National Park and surrounding area, and

the Forsayth area (Map 14). Also known from

the eastern Kimberley of Western Australia, but

has not been recorded for the Northern

Territory. It inhabits depressions or crevices on

skeletal sandstone ridges and plateaux, in or

adjacent to low eucalypt woodland.

Phenology : Flowers are recorded from April

to July and in October; mature fruits recorded

in April and May.

Notes: S. carduiforme is the only dioecious

species indigenous to Queensland. The female

plants have reduced inflorescences bearing only

a single flower.

S. carduiforme is close to S. chippendalei

but differs by the dioecious habit, the more-

dissected leaves with a much “closer”

tomentum, smaller fruits, and accrescent calyx.

Conservation status: Currently listed as

“Vulnerable” under the Queensland Nature

Conservation Act, 1992.

Solanum carduiforme is restricted in its

area of occupancy, but has been collected

several times in recent years. It is conserved in

Lawn Hill N.P. The only perceived threat to its

continued existance is the small population

size. Applying the IUCN guidelines (IUCN.

2001), a category of “Near Threatened” is

recommended.

800

Austrobaileya 6 (4): 639-816 (2004)

Group 29 (S. incanum group) of Whalen

(1984).

Inflorescences with one (rarely two) basal

bisexual flower(s) with very prickly calyx, and

several male flowers with unarmed or sparsely

prickly calyx (100%); calyx prickles present in

fruit (100%); mature fruits 2-locular, 15-30

mm diameter, yellow or orange (100%); finger

hairs absent (100%); branchlet prickles broad-

based (67%); adult leaves lobed (67%).

4 species in Australia, 3 species occurring

in Queensland (2 naturalised and 1 native).

This is not a very cohesive group, as

S. stupefactum is rather different in several respects.

86. Solanum stupefactum Symon,

Austrobaileya 4: 435 (1995). Type:

Queensland. Moreton District: northern

outskirts of Yarraman township, 3 April

1975, R.J. Henderson 2286 (holo: BRI;

iso: CANB).

Solanum sp. 4 in Ross (1986)

Illustration: Symon (1995: 436)

Erect, rhizomatous perennial shrub, 1-2 m

high. Juvenile stage unknown. Adult branchlets

white or brown; prickles 25-45 per decimetre,

straight, acicular, 4-10 mm long, 9-15 times

longer than wide; stellae dense, 0.5-1.2 mm

diameter, stalks 0-2 mm long; lateral rays

6-8, porrect; central ray 2-4 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs dense. Adult leaves ovate, entire;

lamina 4.5-9 cm long, 2.2-4.3 cm wide,

1.7-2.4 times longer than broad, apex obtuse

or acute, base cuneate to cordate, oblique part

0-3 mm long, obliqueness index 0-4 percent;

petioles 0.8-2 cm long, 17-25% length of

lamina, prickles absent or present. Upper leaf

surface green; prickles 0-2, straight, acicular,

5-9 mm long, prickles absent or present on

midvein only; stellate hairs distributed

throughout; protostellae present; ordinary

stellae density moderate to dense, 0.3-0.5 mm

apart, 0.5-0.9 mm across, stalks 0-0.3 mm

long; lateral rays 6-9, porrect; central ray 2-4

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs present

throughout, 0.3-0.8 mm apart. Lower leaf

surface green; prickles absent; stellae dense,

0.1-0.2 mm apart, 0.5-0.9 mm diameter, stalks

0-1.1 mm long; lateral rays 7 or 8, porrect;

central ray 2-3 times as long as laterals, not

gland-tipped; finger hairs absent; Type 2 hairs

absent. Inflorescence supra-axillary, pseudo-

racemose, common peduncle 7-21 mm long,

rachis prickles present; 5-9-flowered, strongly

andromonoecious, flowers 5-merous. Flowers

markedly dimorphic, with larger pricklier basal

flower(s); pedicels 10-25 mm long at anthesis,

same thickness throughout, 0.5-0.7 mm thick

at mid-point, prickles present. Calyx prickles

on basal bisexual flowers 10-18,1-7 mm long;

prickles absent from distal male flowers. Calyx

tube 3-5 mm long, lobes deltate, 6-8 mm long;

stellae very dense, white to brown or rusty,

0.6-1.5 mm across, stalks 0.3-1.7 mm long,

lateral rays 5-7, central ray 1-3 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Corolla mauve or

purple, 9-20 mm long, rotate or shallowly

lobed, inner surface glabrous; anthers 4.5-7

mm long; ovary with stellate hairs only;

functional style 7-11 mm long, erect, with

stellate hairs only, stellae 0.5-0.9 mm across,

central ray 2-4 times as long as laterals; stigma

forked, or lobed. Fruiting calyx with lobes less

than half length of mature fruit, prickles 1-7

mm long. Mature fruits 1 per inflorescence,

globular or ellipsoidal, 18-26 mm diameter,

orange, conspicuously tomentose, 2-locular;

placenta in cross-section stalked, anvil-shaped;

mesocarp moist but not juicy; exocarp 2.2-3

mm thick; pedicels 15-25 mm long in fruit,

1.1-1.6 mm thick at mid-point; seeds pale

yellow, brown or black, 2.8-3.5 mm long.

Specimens examined : Queensland. Darling Downs

District: Gowrie Mountain, undated, Bailey (BRI). Moreton

District: Araucaria Ranges, Dec 1856, Mueller s.n. (MEL

[MEL12199]); Rockmount, 25 km S of Helidon, Aug 1986,

Bird (BRI); Paradise Range, Mt Sylvia via Gatton, Sep 1986,

Williams 86016 (BRI); S.F.289, c. 5 km NW of Yarraman,

Jan 2000, Bean 15992 (BRI, MEL, NSW); adjacent to Mt

Binga S.F., 11 km SE of Cooyar, Feb 2000, Bean 16060

(AD, BRI, CANB, MEL, NSW).

Distribution and habitat : Solanum stupefactum

is endemic to Queensland, and only in the

south-east, from Yarraman to Mt Sylvia (Map

30). It grows on the edge of notophyll rainforest

or in sclerophyll forest close to rainforest.

Phenology : Flowers recorded from January to

September; mature fruits recorded for February

and March.

Bean, Taxonomy of Solanum subg. Leptostemonum

Notes: A distinctive species without close

relatives in Australia, but with some affinity to

the African species S. incanum. It differs from

S. incanum by the entire adult leaves and bright

orange fruits with persistent dense stellate hairs.

Sterile specimens of S. stupefactum are

very similar to S. densevestitum, particularly in

leaf size, shape and indumentum, but S. stupefactum

is a more robust plant with prickly stems and

petioles.

Conservation status: S. stupefactum is

currently known from a few locations. It is not

known from a conservation reserve. It is

threatened by land clearing and competition

from weeds, particularly Lantana camara.

Applying the IUCN guidelines (IUCN. 2001),

a category of “Vulnerable” is recommended

(VU B2ab(ii,iii); Cl).

87. *Solanum incanum L., Sp. PI. 188 (1753).

Type: Herb. J. Burser Vol. IX No. 20 (neo:

UPS), fide Hepper & Jaeger, Kew Bull.

40: 388 (1985).

Erect, rhizomatous perennial shrub, 0.5-1.5 m

high. Juvenile stage unknown. Adult branchlets

brown or green; prickles 25-50 per decimetre,

curved, broad-based, 3-7 mm long, 2-3 times

longer than wide; stellae dense, 0.7-1 mm

diameter, stalks 0-0.2 mm long; lateral rays

6-8, porrect; central ray 0.7-1.5 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Adult leaves ovate

or broadly ovate, shallowly lobed throughout;

lobes 2 or 3 on each side, obtuse, lobing index

1.1-1.6; lamina 3.5-8.5 cm long, 2.3-5 cm

wide, 1.5-1.7 t im es longer than broad, apex

obtuse or acute, base cuneate or obtuse, oblique

part 0-2 mm long, obliqueness index 0-2

percent; petioles 0.8-1.7 cm long, 20-30%

length of lamina, prickles present. Upper leaf

surface grey-green; prickles 3-11, straight,

broad-based, 5-10 mm long, prickles present

on midvein only or present on midvein and

lateral veins; stellate hairs distributed

throughout; protostellae absent; ordinary stellae

density moderate to dense, 0.3-0.5 mm apart,

0.4-0.7 mm across, sessile; lateral rays 4-8,

porrect; central ray 0.8-1.2 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Lower leaf surface green

to white; prickles 5-15, straight, broad-based,

801

prickles present on midvein only or present on

midvein and lateral veins; stellae dense,

0.1-0.2 mm apart, 0.5-0.7 mm diameter, stalks

0-0.2 mm long; lateral rays 7 or 8, porrect;

central ray 0.8-1.2 times as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs absent. Inflorescence supra-axillary,

solitary or pseudo-umbellate, common

peduncle absent; 1-3-flowered, strongly

andromonoecious, flowers 5-merous. Flowers

markedly dimorphic, with larger pricklier basal

flower(s); pedicels 5-10 mm long at anthesis,

same thickness throughout, 0.4-1 mm thick at

mid-point, prickles absent or present. Calyx

prickles on basal bisexual flowers 10-30, 2-7

mm long; prickles absent from distal male

flowers. Calyx tube 3-4 mm long, lobes deltate

or rostrate, 1.5-2.5 mm long; stellae very dense,

yellow or white, 0.6-0.8 mm across, stalks

0-0.2 mm long, lateral rays 7 or 8, central ray

0.8-1.2 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs absent.

Corolla purple, 9-12 mm long, rotate, inner

surface sparsely stellate-hairy; anthers 4.5-5

mm long; ovary with stellate and Type 2 hairs;

functional style 6.5-7 mm long, erect, with

stellate and Type 2 hairs, stellae 0.6-0.7 mm

across, lateral rays c. 9, central ray 0.5-1 times

as long as laterals. Fruiting calyx with lobes

less than half length of mature fruit, prickles

4-7 mm long. Mature fruits 1 per inflorescence,

globular, 18-22 mm diameter, yellow, 2-

locular; placenta in cross-section stalked, anvil¬

shaped; mesocarp moist but not juicy; exocarp

c. 1 mm thick; pedicels 15-20 mm long in fruit,

0.8-1.2 mm thick at mid-point; seeds pale

yellow or brown to black, 2-2.6 mm long.

Specimens examined : Queensland. Port Curtis District:

Castletower Creek, Awoonga Dam, Calliope Shire, Aug 1987,

Gibson 852 (BRI); 2 Tabala Rd, Calliope, ex Kroombit road,

Nov 1998, Worthington 1930 (BRI); Awoonga Dam, 1 km

east of Charters Crossing, Aug 2000, Bean 16804 (BRI, K,

MEL, NSW).

Distribution and habitat: Solanum incanum is

native to Africa. In Australia, it is naturalised

only in the Calliope area south-west of

Gladstone (Map 30), where it inhabits

degraded sites, including rodsides and cleared

alluvial flats.

Phenology: Flowers recorded for August only;

mature fruits recorded for August and

November. The species probably produces

802

Austrobaileya 6 (4): 639-816 (2004)

flowers and fruits for several months of the year.

Notes: S. incanum was first collected in

Queensland in 1987, but not identified until

recently.

88. *Solanum linnaeanum Hepper & P. Jaeger,

Kew Bull. 41: 435 (1986). type: South

Africa. Somerset Div., Bosch Berg, June

1813, W.J. Burchell 3238 (holo: K, n.v.).

Solarium sodomeum, nom. rej., fide Hepper,

Taxon 27: 555 (1979)

Solarium hermannii Dunal, nom. illeg.

(= S. sodomeum )

Illustrations : H. Heine, Solanaceae in Flore

de la Nouvelle Caledonie, p. 147 (1976),

as S. sodomeum; Cunningham et al.

(1981: 594), as S. sodomaeum ; Symon

(1981: 265), as ‘S. hermanni’ (sic).

Erect, rhizomatous perennial shrub, 0.7-1.5 m

high. Juvenile stage unknown. Adult branchlets

grey, brown or green; prickles 10-25 per

decimetre, straight, broad-based, 3-7 mm long,

2-3 times longer than wide; stellae sparse,

0.25-0.4 mm diameter, stalks 0-0.2 mm long;

lateral rays 7 or 8, porrect; central ray 0.1-1.5

times as long as laterals, not gland-tipped;

finger hairs absent; Type 2 hairs absent. Adult

leaves ovate or broadly ovate, deeply lobed

throughout; lobes 3 or 4 on each side, obtuse,

lobing index 5-13; lamina 5-12.5 cm long,

2.8-8 cm wide, 1.4-1.9 times longer than

broad, apex obtuse, base cuneate or obtuse,

oblique part 0-10 mm long, obliqueness index

0-8 percent; petioles 0.6-2.7 cm long, 10-25%

length of lamina, prickles present. Upper leaf

surface green; prickles 9-25, straight, broad-

based, 3-13 mm long, prickles present on

midvein and lateral veins; stellate hairs

distributed throughout; protostellae absent;

ordinary stellae very sparse, 0.9-2 mm apart,

0.25-0.4 mm across, sessile; lateral rays 6-9,

porrect; central ray 0.7-2 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Lower leaf surface green;

prickles 8-20, straight, broad-based, prickles

present on midvein and lateral veins; stellae

sparse to moderate, 0.5-1 mm apart, 0.4-1 mm

diameter, stalks 0-0.1 mm long; lateral rays

6-8, porrect; central ray 0.8-2 times as long as

laterals, not gland-tipped; finger hairs absent;

Type 2 hairs absent. Inflorescence supra-

axillary, pseudo-umbellate, common peduncle

absent; 3-5-flowered, strongly or weakly

andromonoecious, flowers 5-merous. Flowers

markedly dimorphic, with larger pricklier basal

flower(s); pedicels 4-12 mm long at anthesis,

same thickness throughout, 0.6-0.9 mm thick

at mid-point, prickles present. Calyx prickles

on basal bisexual flowers 60-100, 0.5-5 mm

long; prickles on distal male flowers 12-25,

0.5-3 mm long. Calyx tube 3-5 mm long, lobes

elliptic or deltate, 2-5.5 mm long; stellae

moderate to dense, white or transparent,

0.2-0.5 mm across, sessile, lateral rays 4-7,

central ray 0.5-1.5 t im es as long as laterals,

not gland-tipped; finger hairs absent; Type 2

hairs present. Corolla purple, 8-15 mm long,

shallowly or deeply lobed, inner surface

sparsely stellate-hairy; anthers 5-6 mm long;

ovary glabrous or with stellate hairs only;

functional style 7.5-9 mm long, erect, with

stellate hairs only, stellae 0.2-0.25 mm across,

lateral rays 6-8, central ray 1.5-2 times as long

as laterals. Fruiting calyx with lobes less than

half length of mature fruit, prickles 1-5 mm

long. Mature fruits 1 or 2 per inflorescence,

globular, 23-30 mm diameter, yellow, 2-locular;

placenta in cross-section stalked, anvil¬

shaped; mesocarp moist but not juicy;

exocarp 1-2.5 mm thick; pedicels 17-21 mm

long in fruit, 1.5-3 mm thick at mid-point;

seeds brown to black, 2.9-3.5 mm long.

Apple of Sodom. Fig. 10.

Selected specimens: Queensland. South Kennedy District:

Plateau road, Crediton, SE of Eungella, Apr 2002, Bean

18656 (BRI, MEL). Wide Bay District: 9.6 km S of Gunalda,

Mar 1973, Price s.n. (BRI); Mary River, Conondale on the

Maleny-Kenilworth road, Aug 1975, Coveny 6730 & Hind

(BRI, NSW); Currymore, NW of Maleny, Apr 1993, Bean

6017 (BRI); 0.8 km S of Kenilworth P.O., Aug 1999, Bean

15243 (BRI). Moreton District: Victoria Park, Oct 1888,

Simmonds (BRI); Caloundra, North Coast Line, Aug 1933,

Everist 402 (BRI); Flagstone via Tamborine, Nov 1963,

Wood (BRI); Mt Glorious-Samford road, Jan 1965,

Henderson H105 (BRI); Dunwich, North Stradbroke Is, Sep

1969, Coveny 2070 (BRI, NSW); Zillmere, Brisbane, Sep

1980, Dillewaard 16 & Olsen (BRI); Long Pocket road,

Indooroopilly, Brisbane, May 1981, Dillewaard 603 &

Stanley (AD, BRI); 3 km W of Ripley, S of Swanbank

powerhouse, Jan 1986, Dowling s.n. (BRI); 1 km N of

Bellthorpe Forest station, Conondale Range, Oct 1993,

Halford Q1909 (BRI); Banks Creek road, 6 km NE of

Fernvale, Jul 2000, Bean 16711 (BRI, MEL); 1.2 km along

Tabletop road, S of Canungra, Apr 2001 ,Bean 17598 (BRI).

Distribution and habitat : Solanum linnaeanum

is native to South Africa. In Queensland, it is

Bean, Taxonomy of Solanum subg. Leptostemonum

naturalised in areas south from Gunalda, within

about 50 km of the coast, and then on the

Eungella plateau west of Mackay (Map 32). It

is also naturalised in New South Wales. It grows

on degraded sites, including pasture and

roadsides.

Phenology : Flowers are recorded from July to

December; fruits may be found throughout the year.

Notes: S. linnaeanum is one of Australia’s first

recorded introduced weeds. It was found by

Robert Brown around Port Jackson in 1802.

The first Queensland naturalisation (around

Brisbane) was recorded by Bailey (1881).

Group 33 (S. rostratum group) of Whalen

(1984)

Corolla yellow, rotate; one anther much longer

than the remainder; calyx prickles abundant;

lower leaf surface and calyx with Type 2 hairs;

style sigmoid; seeds brown to black;

hypanthium accrescent, enclosing the fruit;

adult leaves deeply lobed.

12 species in North and South America;

1 species naturalised in Australia, and present

in Queensland.

89. *Solanum rostratum Dunal, Hist. Nat.

Solanum 234, t. 24 (1813); Nycterium

rostratum (Dunal) Link, Enum. Hort.

Berol. 1: 189 (1821); Androcera rostrata

(Dunal) Rydb., Bull. Torrey Bot. Club 33:

150 (1906). Type: cultivated at

Montpellier, undated, coll, unknown

(holo: MPU; iso: G-DC, P \fide Whalen

(1979).

Illustration: Cunningham et al. (1981: 595);

Symon (1981: 109)

Sprawling or erect, herbaceous resprouter or

rhizomatous perennial shrub, 0.3-0.6 m high.

Juvenile stage unknown. Adult branchlets

brown or green; prickles 180-240 per

decimetre, straight, acicular or broad-based,

1-7 mm long, 6-9 times longer than wide;

stellae sparse to dense, 0.3-1.3 mm diameter,

stalks 0-0.1 mm long; lateral rays 4-7, porrect

or ascending; central ray 0.8-1.5 times as long

as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs absent. Adult leaves

elliptical, ovate or broadly ovate, deeply lobed

throughout; lobes 3 or 4 on each side, obtuse,

803

lobing index 9-50; lamina 4-8 cm long,

2.1-4.2 cm wide, 1.5-1.9 times longer than

broad, apex obtuse, base cuneate to cordate,

oblique part 0-3 mm long, obliqueness index

0-4 percent; petioles 1.3-2.9 cm long, 35-50%

length of lamina, not winged or winged,

prickles present. Upper leaf surface green;

prickles 9-35, straight, broad-based, 2-7 mm

long, prickles present on midvein and lateral

veins; stellate hairs distributed throughout;

protostellae present; ordinary stellae density

moderate, 0.3-0.8 mm apart, 0.6-0.9 mm

across, sessile; lateral rays 4-6, porrect; central

ray 1-1.7 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs present

throughout, 0.1-0.2 mm apart. Lower leaf

surface green; prickles 10-20, straight, broad-

based, prickles present on midvein and lateral

veins; stellae moderate to dense, 0.2-0.5 mm

apart, 0.8-1.1 mm diameter, stalks 0-0.1 mm

long; lateral rays 4-8, porrect; central ray

1-1.7 times as long as laterals, not gland-

tipped; finger hairs absent; Type 2 hairs present

throughout, 0.2-0.4 mm apart. Inflorescence

supra-axillary, pseudo-racemose, common

peduncle 5-20 mm long, rachis prickles

present; 5-10-flowered, weakly

andromonoecious, flowers 5-merous; pedicels

3-6 mm long at anthesis, markedly thicker

distally, 0.5-0.7 mm thick at mid-point,

prickles present. Calyx tube 2.5-3.5 mm long,

lobes deltate or attenuate, 4-5.5 mm long;

prickles present at anthesis, 100-250 per

flower, 1-9 mm long; stellae dense, transparent,

0.6-1.3 mm across, stalks 0-0.3 mm long,

lateral rays 4-8, central ray 0.8-1.5 times as

long as laterals, not gland-tipped; finger hairs

absent; Type 2 hairs present. Corolla yellow,

9-15 mm long, rotate, inner surface glabrous;

anthers 3.5-7.5 mm long; ovary glabrous;

functional style c. 13 mm long, sigmoid,

glabrous. Fruiting calyx tube accrescent,

enclosing most or all of mature fruit, prickles

1-11 mm long. Mature fruits 4-8 per

inflorescence, globular, 9-12 mm diameter, 2-

locular; mesocarp dry; pedicels 6-10 mm long

in fruit, 1-1.2 mm thick at mid-point; seeds

brown to black, 2-2.7 mm long. Buffalo Burr.

Specimens examined : Queensland. Burnett District: 5

miles SW of Gayndah, Nov 1971, Sauer (BRI). Darling

Downs District: Jandowae, Portion 69, Parish Canaga, Nov

1961, Sperling (BRI); 28 km W of Miles, Dec 1973, Pollock

(BRI); ‘Rosewood Downs’, Limevale, Mar 1981, Major

(BRI); Portion 28V, Shire of Clifton, Dec 1983, Achilles

804

Austrobaileya 6 (4): 639-816 (2004)

(BRI); Clifton, Dec 1983, Beitz (BRI). Moreton District:

Gatton, Feb 1906, Bailey (BRI). New South Wales. North

West Slopes: Warialda district, Dec 1961, Vet. Inspector

(NSW).

Distribution and habitat : Solanum rostratum

is indigenous to the U.S.A. and Mexico. There

have been sporadic naturalisation records from

several parts of south-eastern Queensland (Map

24), with the most recent record being in 1983.

It is naturalised in all southern Australian states

except Tasmania. It grows on degraded sites

(pastures and crops).

Phenology: Flowers are recorded in November

and December; fruits are recorded in March,

November and December.

Notes: This is the only species of the American

S. sect. Androceras that is naturalised in

Australia (Whalen 1979), and the only

Australian species of Solanum (native or

naturalised) with yellow flowers.

I have been unable to locate populations

of S. rostratum in Queensland, and it is rather

doubtful whether it still occurs here.

Ungrouped. Whalen (1984) did not assign this

species to any group

90. *Solanum sisymbriifolium Lam., Tabl.

Encycl. 2: 25 (1794). Type: Buenos Aires,

Argentina, undated, P. Commerson (syn:

P-LAM), microfiche 469.14.

Illustration: Symon (1981: 111)

Erect, rhizomatous perennial shrub, 0.8-1.5 m

high. Juvenile stage unknown. Adult branchlets

brown or green; prickles 30-60 per decimetre,

straight or curved, broad-based, 1—12 mm long,

5- 7 times longer than wide; stellae dense,

0.3-0.5 mm diameter, sessile; lateral rays

6- 12, ascending or multiradiate; central ray

1.5- 3 times as long as laterals, not gland-tipped

or gland-tipped; finger hairs present, gland-

tipped, 0.2-0.4 mm long; Type 2 hairs absent.

Adult leaves broadly ovate, deeply lobed

throughout; lobes 4-6 on each side, acute,

lobing index 2.5-11; lamina 8-12 cm long,

5.5- 8.5 cm wide, 1.4-1.8 times longer than

broad, apex acute, base obtuse or cordate,

oblique part 0-5 mm long, obliqueness index

0-5 percent; petioles 2.1-4.6 cm long, 21-40%

length of lamina, winged, prickles present.

Upper leaf surface green; prickles 9-30,

straight, broad-based, 1-10 mm long, prickles

present on midvein and lateral veins; stellate

hairs distributed throughout; protostellae

absent; ordinary stellae sparse to moderate

density, 0.3-0.5 mm apart, 0.3-0.5 mm across,

sessile; lateral rays 4-6, ascending; central ray

1-3 times as long as laterals, not gland-tipped;

finger hairs present, 0.2-0.9 mm apart, gland-

tipped, 0.2-0.3 mm long; Type 2 hairs absent.

Lower leaf surface green; prickles 7-20,

straight, broad-based, prickles present on

midvein and lateral veins; stellae moderate,

0.3-0.6 mm apart, 0.4-0.6 mm diameter,

sessile; lateral rays 4-7, ascending; central ray

1-2 times as long as laterals, not gland-tipped;

finger hairs present, 0.2-0.4 mm apart, gland-

tipped, 0.2-0.3 mm long; Type 2 hairs absent.

Inflorescence supra-axillary, pseudo-racemose,

common peduncle 15-52 mm long, rachis

prickles present; 4-11-flowered, weakly

andromonoecious, flowers 5-merous. Flowers

all similar; pedicels 5-12 mm long at anthesis,

same thickness throughout, 0.5-0.7 mm thick

at mid-point, prickles present. Calyx tube

1.5- 3 mm long, lobes deltate, 5-6.5 mm long;

prickles present at anthesis, 25-90 per flower,

I- 7 mm long; stellae dense, transparent,

0.4-0.6 mm across, sessile, lateral rays 6-10,

central ray 2-3 times as long as laterals, not

gland-tipped or gland-tipped; finger hairs

present; Type 2 hairs absent. Corolla white,

II- 16 mm long, rotate or shallowly lobed, inner

surface glabrous; anthers 8-10 mm long; ovary

with Type 2 hairs only; functional style

15.5- 17 mm long, erect, with Type 2 hairs only

or with stellate and Type 2 hairs, stellae c. 0.25

mm across, lateral rays c. 6, central ray 2-3

times as long as laterals. Fruiting calyx with

lobes more than half length of mature fruit,

prickles 2-10 mm long. Mature fruits 1-3 per

inflorescence, globular, 15-20 mm diameter,

red; pedicels 17-25 mm long in fruit, 1.1-1.9

mm thick at mid-point; seeds pale yellow,

2.9-3.2 mm long.

Specimens examined : Queensland. Darling Downs

District: near Toowoomba, Bellview St, Jan 1921, Leslie

(BRI); near Toowoomba, Mar 1953, Clydesdale (BRI);

Toowoomba, S of Picnic Point, Oct 1964, Everistllll (BRI);

17 Trafalgar St Toowoomba, Mar 1996, Story (BRI); Nelson

St Reservoir, Toowoomba, Jul 1996, Bean 10448 (BRI);

corner Nelson St and Ramsay St, Toowoomba, Dec 2001,

Bean 18166 (BRI, MEL, NSW).

Bean, Taxonomy of Solarium subg. Leptostemonum

Distribution and habitat: Solanum

sisymbriifolium is indigenous to Bolivia, Brazil,

Paraguay and Argentina in South America. In

Queensland it is known only from Toowoomba

(Map 32), where it has been naturalised for

over 80 years. It grows on degraded sites in

urban bushland, although once recorded as a

weed of cultivation paddocks. Also naturalised

in New South Wales and Western Australia.

Phenology: Flowers are recorded between

October and March; mature fruits recorded in

July and December.

Notes: Not assigned to a group by Whalen

(1984), but appears to have affinity with the S.

incanum group (strongly andromonoecious

inflorescences, large fruits, somewhat

dimorphic flowers).

Species doubtfully naturalised or formerly

naturalised in Queensland

Solanum aculeastrum Dunal

Shrub to 2 m high, with large broad-based

prickles, leaves that are white on the underside,

and fruits about 3 cm across.

Specimens examined : Queensland. Moreton District:

Victoria Park, Brisbane, Oct 1887, Simmonds s.n. (BRI);

Cleveland, Aug 1906, Rowney s.n. (BRI); Sandgate, Moreton

Bay, Sep 1915, White s.n. (BRI); Currumbin Heights, Nov

1962, Wallace s.n, (BRI).

Notes: S. aculeastrum is native to South Africa,

and formerly naturalised in Queensland. Bailey

(1881) stated that it “is met with in places

covering acres of land, with shrubs of from 6

to 9 feet in height”. It has not been recorded

since 1962.

Solanum dimidiatum Raf.

Shrub to 50 cm high, with deeply lobed leaves,

acicular prickles on branchlets but none on the

calyx.

Specimens examined: Queensland. Wide Bay District:

Bundaberg, Dec 1963, Arnold s.n. (BRI); Greatheads Rd,

Bundaberg, Mar 1966, Draper (BRI); Bundaberg, May 1972,

Henderson 1322 (BRI).

Notes: S. dimidiatum was actively targeted for

eradication in the Bundaberg area, and has not

been seen since 1972.

805

Solanum lasiophyllum Dunal

A shrub with broad felty entire leaves and

prickly stems. Calyx prickly, accrescent in fruit.

Specimens examined : Queensland. Wide Bay District:

Bingera, Gibbsons and Howes Sugar Mill, Oct 1981,

Sarnadsky s.n. (BRI)

Notes: S. lasiophyllum is an Australian species

occurring commonly in Western Australia. A

small population was documented from near

Bundaberg in 1981, but it did not persist.

Solanum pseudolulo Heiser

A small shrub with broad shallowly-lobed

leaves, prickles abundant on leaves and stems,

calyx without prickles, fruits large and

tomentose.

Specimens examined: Queensland. Cook District: Cairns

Airport sewage treatment plant, Jun 2000, King s.n. (BRI).

Moreton District: Cross St, Sinnamon Farm, Goodna,

Ipswich, Jun 1988 , Perrotts.n. (BRI).

Notes: The fruits of S. pseudolulo are edible,

and it is often grown for that reason. These

records are non-cultivated, but neither

population has persisted. It is related to S.

lasiocarpum Dunal.

Solanum pyracanthos Lam.

A shrub with deeply lobed, sparsely pubescent

leaves and abundant orange prickles.

Specimen examined: Queensland. Moreton District:

Bowen Park, Feb 1907, White s.n. (BRI).

Notes: S. pyracanthos was recorded by Bailey

(1881) as being naturalised in Brisbane, with

the comment that “this species was introduced

some years ago as an ornamental plant by the

Queensland Acclimitisation Society, from

whose grounds it has spread into the pasture”.

It is a native of Madagascar.

Excluded Species

Solanum tumulicola Symon

All Queensland specimens previously assigned

to this species have proven to be S. esuriale

Lindl.

806

Austrobaileya 6 (4): 639-816 (2004)

Solanum coactiliferum J.M.Black

All Queensland specimens previously assigned

to this species are referrable to S. ammophilum

A.R.Bean.

Acknowledgements

Numerous people have assisted me in locating

various species of Solanum in the field, or have

collected material for me. I thank Andrew Ford,

Keith McDonald, Bill McDonald, Paul Forster,

Harry Hines, Bob Philips, Lana Little, Rod

Fensham, Ed Meyer, Matt Bloor, Irene

Champion, Steve Pearson, Megan Thomas and

John Thompson. I am grateful to the Directors

of AD, BH, BM, DNA, K, MEL, NSW, PERTH,

PR and QRS for loan of specimens; Laurie

Jessup for guiding me through the intricacies

of DELTA; Will Smith for the art work and

distribution maps; Peter Bostock for

photographing some types while Australian

Botanical Liaison Officer at Kew, and also for

the Latin descriptions; Donovan Sharp for

assistance with the stellate-hair images; and

Wayne Harris for assistance with the

photographing of type specimens.

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808

Austrobaileya 6 (4): 639-816 (2004)

1-10 145 150 155

Map 1 , Distribution of Solanum dmatianum A .

S.viridifoliumA and S.hapalum'A;.

Map 2. Distribution of Solatium densevestitum •,

5. fervensA and S. dimorphispimtm'A .

Map 3, Distribution of Solanum/ohnsoniartum • Map 4, Distribution of Solatium nemophilumA

S. itmoxium A r and S. ultimum w S' yirrkalense • and S. dtyanderense.it -

Bean, Taxonomy of Solanum subg. Leptostemonum

809

Map 5. Distribution of Solanum gympieme • Map 6. Distribution of Solanum shirleyanum • ,

S. defensunfk and S. men tie ns A . S. parvifoUum ssp. parvi/oliumA and

S. parvifolium ssp. tropicum'k ,

810

Austrobaileya 6 (4): 639-816 (2004)

Map 9. Distribution of Solatium ferocissimum • . Map 10. Distribution of Solarium lythrocarpum 9

S. dysprosium A and S. la,emir S - coracinumA and S. semiarmatom*.

Map 11. Distribution of Solarium dissectum A.

S. inaequilaterwn • ,and S. chenopodinum ^

Map 12. Distribution of Solatium mitchelliamtm 9 .

S. cookii A and S. frauds! i'Jt .

Bean, Taxonomy of Solanum subg. Leptostemonum

811

Map 13. Distribution of Solatium chrysotrichum •.

S.pugnmcit life rum ★ and S. adenaphontm A.

Map 14. Distribution of Solatium cap$icoides% .

S, pusiUum^C and S. carduiforme A.

140 145 150 155

Map 15, Distribution of Solanum lacunarium •

S. torvum A and S. stenopterum ★.

140 145 150 155

Map 16, Distribution of Solanum ditrichum A .

S. graniticum% and S. dianthophorum'k .

812

Austrobaileya 6 (4): 639-816 (2004)

Ma p 17. Distribution of Solatium vicinum • Map 18. Distribution of Solanum ellipticum • .

S.senticosum^ and 5 crebrispirmm A.

Map 19. Distribution of Solatium papaver(folium •. Map 20. Distribution of Solatium angustum A ,

S. quadriloculatum A .. and S crassitomentosum'fc. S, sporadotrichum # and S. argopetalum'^.

Bean, Taxonomy of Solanum subg. Leptostemonum

813

Map 21, Distribution of Solanumjurfuraceum% . Map22 , Distribution of Solatium macoorai A ,

S.rixosumA and S. hamulosum ★ S<acartihodapis% and S. tetrathecumif.

Map 23, Distribution of Solatium dumicoh'k

S. emimm •, and $ serpens A

Map 24. Distribution of Solatium magnifolium •.

S.cocosoidesW. S. centrale A and S. rostra turn O

814

Austrobaileya 6 (4): 639-816 (2004)

Ma p 25, Distribution of Solarium cinereumw , Map 26, Distribution of Solarium elachophylhtm A ,

S intonsum •and S. jucundum A. Slim itare • an d S. stun iauum^k.

Map 27, Distribution of Solarium ammophilum A . Map 28. Distribution of Solarium ambfymerum A .

S. galbimm • , and S. mb tie ★ S. echinatomit and S. oligacanthum •.

Bean, Taxonomy of Solanum subg. Leptostemonum

815

Map 29, Distribution of Solanum esuriale • Map 30. Distribution of Solanum chippendaki • .

S.incanum^ and S stupefaction A.

Map3l. Distribution of Solanum elaeagnifolium A ,

and S bngissitmm'A

Map 32, Distribution oi'Solanum linnaeanum • , and

S. sisymbriifolhtm ★.

816

Index

Austrobaileya 6 (4): 639-816 (2004)

New names are printed in boldface; synonyms

and misapplied names are printed in italics

Name. Taxon number

Solanum acanthodapis. 60

S. accedens . 19

S. adenophorum. 39

S. adenophorum var. indivisum . 35

S.amblymerum. 73

S.ammophilum. 79

S.angustum. 51

S. argopetalum . 52

S. capsicoides . 33

S. carduiforme. 85

S. centrale. 69

S. chenopodinum. 24

S. chippendalei . 84

S. chrysotrichum. 30

S. ciliatum . 33

S.cinereum. 70

S. cleistogamum . 45

S. cocosoides. 67

S. cookii. 35

S.coracinum. 27

S.corifolium. 14

S. crassitomentosum. 50

S. crebrispinum. 47

S. dallachii . 57

S. defensum. 12

S. densevestitum. 7

S. dianthophorum. 46

S. dimorphispinum. 53

S. discolor. 13

S. discolor var. procumbens . 15

S. dissectum. 22

S. ditrichum. 34

S. dryanderense. 17

S.dumicola . 65

S. dunalianum. 1

S. dysprosium . 25

S.echinatum. 82

S. elachophyllum. 75

S. elaeagnifolium. 78

S. ellipticum. 45

S. ellipticum var. chillagoense . 45

S. ellipticum var. horridum . 48

S. eminens. 55

S.esuriale. 77

S. ferocissimum. 20

S. ferocissimum var. rectispinum . 20

S. fervens. 10

S. francisii. 64

S. furfuraceum. 62

S. galbinum . 74

S.graniticum. 41

S. gympiense. 9

S.hamulosum. 54

S. hapalum. 3

S. hispidum . 30

S. inaequilaterum . 26

S. incanum . 87

S. innoxium. 5

S. intonsum. 61

S. johnsonianum. 4

S.jucundum. 68

S. lacunarium. 43

S. largiflorum . 31

S. lasiocarpum. 32

S.latens. 21

S. leptophyllum . 20

S. limitare. 72

S. linnaeanum. 88

S. longissimum. 83

S. lythrocarpum. 23

S. lucorum . 18

S. macoorai. 56

S. magnifolium. 57

S.mentiens. 15

S. mitchellianum. 28

S. multiglochidiatum. 36

S. nemophilum . 8

S. nemophilum var. brachycarpum . 6

S. nobile . 71

S. oligacanthum. 81

S. papaverifolium. 38

S. parvifolium ssp. parvifolium. 19a

S. parvifolium ssp. tropicum . 19b

S. prinophyllum . 37

S. pugiunculiferum. 44

S.pusillum. 40

S. quadriloculatum. 49

S. rixosum . 58

S. rostratum. 89

S. seitheae . 82

S. semiarmatum. 29

S. senticosum. 48

S. serpens. 59

S. shirley anum. 16

S. sisymbriifolium. 90

S. sporadotrichum. 63

S. stelligerum. 18

S. stelligerum var. magnifolium . 57

S. stelligerum var. procumbens . 59

S. stenopterum. 42

S. stupefactum. 86

S. sturtianum. 80

S. tetrathecum. 66

S.torvum. 31

S.ultimum. 6

S. versicolor. 76

S. vicinum . 37

S. viride . 2

S. viridifolium. 2

S. yirrkalense. 11

Vanguerieae A.Rich. ex Dum. (Rubiaceae) in Australia, 3.

Psydrax Gaertn.

Sally T. Reynolds and Rodney J.F. Henderson

Summary

Reynolds, S.T. & Henderson, R.J.F. (2004). Vanguerieae A.Rich. ex Dum. (Rubiaceae) in Australia, 3.

Psydrax Gaertn. Austrobaileya 6(4): 817-889. The genus Psydrax Gaertn. (Rubiaceae, Vanguerieae), as

it occurs in Australia, is revised. Several species of this genus have previously been included in Canthium

Lam. but are now considered distinct from that genus. Of the twenty two Australian species currently

recognised as belonging to Psydrax , thirteen are described here as new, namely Psydrax ammophila

S.T.Reynolds & R.J.F.Hend., P. banksii S.T.Reynolds & R.J.F.Hend., P.forsteri S.T.Reynolds & RJ.F.Hend.,

P. johnsonii S.T.Reynolds & R.LF.Hend., P. laxiflorens S.TReynolds & RJ.F.Hend., P. lepida S.TReynolds

& RJ.F.Hend., P. longipes S.T.Reynolds & RJ.F.Hend., P. pallida S.T.Reynolds & R.LF.Hend., P. paludosa

S.T.Reynolds & R.J.F.Hend., P. pendulina S.T.Reynolds & R.J.F.Hend., P. rigidula S.T.Reynolds &

R.J.F.Hend., P. saligna S.T.Reynolds & R.J.F.Hend. and P. tropica S.T.Reynolds & R.J.F.Hend. New

combinations are made for six of the other nine species, namely Psydrax attenuata (Benth.) S.T.Reynolds

& R.J.F.Hend., P. graciliflora (Merr. & L.M.Perry) S.T.Reynolds & R.J.F.Hend., P. latifolia (F.Muell. ex

Benth.) S.T.Reynolds & R.J.F.Hend., P. oleifolia (Hook.f.) S.T.Reynolds & R.J.F.Hend., P. reticulata

(C.T.White) S.T.Reynolds & R.J.F.Hend. and P. suaveolens (S.Moore) S.T.Reynolds & R.J.F.Hend., while

P. montigena S.T.Reynolds & R.J.F.Hend. is provided as a new name for the plant previously known as

Ixora orophila C.T.White, which belongs in Psydrax as well. New infraspecific taxa recognised here are

Psydrax attenuata var. myrmecophila S.T.Reynolds & R.J.F.Hend. with Psydrax attenuata forma

myrmecophila S.T.Reynolds & R.J.F.Hend. and P. attenuata forma megalantha S.T.Reynolds &

R. J.F.Hend., P. attenuata var. tenella S.T.Reynolds & R.J.F.Hend., P. lamprophylla forma latissima

S. T.Reynolds & R.J.F.Hend., P. odorata subsp. amhemica S.T.Reynolds & R.J.F.Hend., P. odorata subsp.

australiana S.T.Reynolds & R.J.F.Hend., with P. odorata forma australiana S.T.Reynolds & R.J.F.Hend.,

P. odorata forma foveolata S.T.Reynolds & R.J.F.Hend. and P. odorata forma subnitida S.T.Reynolds &

R.J.F.Hend., and P. saligna forma filiformis S.T.Reynolds & R.J.F.Hend. Canthium buxifolium Benth. is

reduced to a subspecies of Psydrax odorata (Forst.f.) A.C.Sm. & S.P.Darwin containing P. odorata forma

buxifolia (Benth.) S.T.Reynolds & R.J.F.Hend. and P. odorata forma parviflorifra S.T.Reynolds &

R. J.F.Hend., and Canthium lineare E.Pritz. is accepted as conspecific with Psydrax suaveolens (S.Moore)

S. T.Reynolds & R.J.F.Hend. Lectotypes for Canthium attenuatum Benth. and Canthium lamprophyllum

F.Muell., basionyms for Psydrax attenuata (Benth.) S.T.Reynolds & R.J.F.Hend. and P. lamprophylla

(F.Muell.) Bridson respectively, are designated. Keys to the species and infraspecific taxa, distributional

maps and illustrations of some of the species are provided. A list of currently accepted names of Australian

taxa previously considered to belong in Canthium Lam. or Plectronia L., is given in Supplement A. New

combinations are made in Appendix 1. for two related species occurring in New Guinea (but not in

Australia) considered to belong in Psydrax, namely Psydrax cymigera (Valeton) S.T.Reynolds &

R.J.F.Hend. and P. suborbicularis (C.T.White) S.T.Reynolds & R.J.F.Hend.

Key words: Rubiaceae, Vanguerieae, Psydrax, Australian flora, taxonomy, nomenclature

Sally T. Reynolds and Rodney J.F. Henderson, Queensland Herbarium, Environmental Protection Agency,

Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha Road, Toowong, Queensland 4066

Introduction

As stated previously (Reynolds & Henderson

1999, 2001), taxa of Psydrax Gaertn. and

Cyclophyllum Hook.f., together with those of

Everistia S.T.Reynolds & R.J.F.Hend., have,

until comparatively recent times, been included

in Canthium Lam. Following critical studies

from the 1960s evaluating the genus Canthium

in Africa, recognition of some of the

subordinate taxa previously included in it as

Accepted for publication 13 July 2004

distinct genera has resulted. Cuparon (1969)

proposed that many species of Canthium from

Africa, Madagascar and Asia be transferred to

Psydrax and this led Bridson (1985) to reinstate

Psydrax and include in it African species

previously included in Canthium and one from

Australian she named Psydrax lamprophylla

(F.Muell.) Bridson.

Most of the Australian species previously

included in Canthium are referrable to Psydrax.

In this country, species of this latter genus are

818

Austrobaileya 6 (4): 817-889 (2004)

very complex and most of them are difficult to

delimit because they have similar flowers and

fruits, and the leaves, which are used to

distinguish most species, are very variable and

sometimes vary on the one branchlet. Moreover,

most species are poorly known, poorly

represented by herbarium material or

represented in herbaria by incomplete material,

and specimens exist which appear to represent

intergrades between them.

Three of the taxa Bentham (1867)

accepted as species of Canthium , and dealt with

under the names C. attenuatum, C. lucidum and

C. oleifolium, were each found to contain more

than one species. Some of these species were

undescribed while duplicates of the one

collection of others were found filed under

different species names in various herbaria.

Moreover, it was discovered that the application

of C. attenuatum and C. lamprophyllum had

been inconsistent from the time the species were

described. Examination of the syntypes of

C. attenuatum and C. lamprophyllum showed

that in both cases they represent more than one

taxon at species rank. This is discussed more

fully under the relevant Psydrax species below.

Canthium attenuatum had also been confused

with C. oleifolium, probably because the species

concerned are morphologically very similar and

their protologue descriptions are scanty and not

clearly diagnostic. Bentham (1867) added to

the confusion when he cited a Ferdinand

Mueller collection from the ‘Burdekin river’

(in Queensland) for both these species (once as

a syntype of Canthium attenuatum Benth.).

Consequently, matching specimens had been

filed under either of these names in various

herbaria. This had the effect of making the

species concerned seem very variable and

difficult to delimit. Moreover, although

Bentham considered certain Australian

specimens he included under the name

Canthium lucidum (= Psydrax odorata in this

paper) to be conspecific with those from the

Pacific Islands with this name, Merrill & Perry

(1945) and Bridson (1985) considered the

Australian ones specifically distinct from those

from the Pacific Islands and referable to

Canthium lamprophyllum F.Muell. (former) or

Psydrax lamprophylla (F.Muell.) Bridson

(latter). We accept, in part at least, Merrill’s,

Perry’s and Bridson’s point of view that two

species, not one as Bentham thought, are

involved. However, we have found that both

these species actually do occur in Australia.

This confusion in the nomenclature and

taxonomy of these species, especially regarding

the variability allowed in Australian specimens

previously filed under the name Canthium

odoratum in herbaria, began when Bentham

(1867) combined Canthium lamprophyllum

F.Muell. under C. lucidum. This was because

the former, as Bentham conceived it, actually

comprised two distinct taxa one of which is

Psydrax odorata and the other is P. lamprophylla

(F.Muell.)Bridson as delimited in this revision

(see under P. odorata below).

Although the previous confusion in

relation to P. odorata and most other Psydrax

species in Australia has been clarified, and once-

undescribed taxa allied to P. odorata, P attenuata

and P. oleifolia have been described (in this

paper), many species still remain poorly known

and specimens which cannot satisfactorily be

placed in the taxa recognised here also exist.

More collections of these species, especially of

specimens with flowers, are needed to be made

and more field studies undertaken before their

affinities or dissimilarities can be fully

understood. The variation accepted for some

of the species recognised here is sometimes

limited, mainly because they are insufficiently

known, so their status may change as more

material and more field observations become

available.

Because most species are similar and

often difficult to delimit, study of a combination

of characters is often necessary before

specimens can be identified. For this reason,

lists of diagnostic attributes and the taxa that

possess them are provided below to assist in

identification of mainly sterile material. In

addition, species which are closely related are

grouped together and secondary keys to separate

them from other group members are provided.

As before (Reynolds & Henderson 2001),

this study was based mostly on herbarium

material, but measurements given for leaves,

inflorescences, flowers and fruits are based on

dried, fresh or spirit material. Some measurements,

such as heights for shrubs/trees, rely on notes

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

provided by the collector. In the list of

specimens cited, only the herbaria from which

specimens have been seen are recorded. These

herbaria are indicated by acronyms following

Holmgren et al., 1990. State subdivisions

(pastoral districts) are provided for Queensland

collections only. The acronym ‘dbh’, used in

the forestry industry sense for ‘diameter at

breast height’ and provided by the collector with

some specimens, is used in relation to the tree/

shrub trunk in some descriptions. The

taxonomic concepts accepted here are those of

the first author and result from her many years

of detailed herbarium studies.

Taxonomy

Psydrax Gaertn., Fruct. 1: 125, t.26, f.2 (1788);

from Greek psudros, wrong, and

pseudein, to lie, an allusion to the

‘knobbly’ seeds found in this genus and

the belief that pimples grow on a liar’s

tongue. Type: Psydrax diococcos Gaertn.

Trees or shrubs; branches usually borne at

right angles to the stem; branchlets occasionally

spinescent. Leaves opposite or whorled,

stipulate, petiolate; stipules interpetiolar, free

or shortly confluent and sheathing at base,

mostly triangular or lanceolate, keeled and

attenuated into a folded lobe at apex (except in

P. lamprophylla), persistent or sometimes

caducous; blades entire, glabrous or hairy.

Inflorescences axillary, pedunculate, of

dichotomously branched cymes with bifurcate

cymose branches, the flowers mostly secund on

the rachis and usually with a comparatively

long-stalked solitary (central) flower near forks

of the dichotomous branches; bracts small,

usually near the distal end of the main peduncle.

Flowers usually bisexual, 4- or 5-merous,

fragrant. Calyx tube short, obconical or broad

ellipsoid, with limb short cupular, subtruncate

or shallowly lobed with 4 or 5, usually minute

lobes. Corolla white, cream or lemon yellow,

with tube usually campanulate, glabrous

819

abaxially and usually with a ring of retrorse

hairs at throat adaxially; lobes as long as the

tube, valvate in bud, erect, spreading or

recurved in flower, elliptic-oblong or lanceolate,

obtuse or acute, slightly incurved and ± hooded

at apex. Stamens 4 or 5, inserted in throat of

corolla tube; filaments comparatively short,

slender, sometimes nearly obsolete; anthers

exserted or + included in corolla tube,

oblongoid or ellipsoid, obtuse and mucronate

at apex and often tailed at base, dorsifixed,

extrorse. Disc annular, inconspicuous. Ovary

2-locular, with a solitary, + pendulous ovule in

each locule; style filiform, longer than corolla

tube, usually much exserted; stigma attached

to the style by its concave hollow base; stigmatic

knob ovoid or oblongoid, usually bilobed at

apex. Fruit a drupe, usually blackish coloured

when dry, subglobose, ellipsoid or obovoid but

laterally compressed, slightly fleshy; pyrenes

1 or 2 in each fruit, these usually thick and

woody, slightly ellipsoid, broad ovoid or

hemispherical, usually depressed distally,

rugose or smooth externally, 1-seeded; seeds

oblongoid, with testa membranous.

Distribution: 100 species in the Old World

Tropics according to Mabberley (1997); 22

species in Australia.

Affinities: Psydrax is closely related to

Canthium from which it was segregated by

Cuparon (1969) primarily on the orientation

of the cotyledons. In Psydrax, he accepted the

cotyledons are aligned parallel to the ventral

surface of the seed whereas in Canthium they

are aligned perpendicular to the ventral face of

the seed.

Cuparon provided additional diagnostic

characters for each genus which Bridson, when

reinstating Psydrax as a distinct genus, noted

before providing more characteristics to

distinguish this genus from Canthium (Bridson

1985, p.691). A synopsis of these is as follows.

820

Austrobaileya 6 (4): 817-889 (2004)

Plants in habit erect small trees or shrubs, rarely scandent but never lianas;

leaves usually restricted to new growth at the apex of stems; stipules

with or without villous hairs adaxially; style equalling the corolla tube

in length, shortly exceeding it or occasionally twice as long as it; stigmatic

knob ± as long as wide, rarely longer; anthers partly or completely

exserted from corolla tube, straight, never reflexed. Canthium

Plants in habit erect tall trees or shrubs or lianas; leaves dispersed along

stems or restricted to apices in a very few species; stipules never with

fine hairs adaxially; style exceeding the corolla tube in length by about

twice or often much more; stigmatic knob distinctly longer than wide;

anthers fully exserted from corolla tube, reflexed or straight

.Psydrax (and Keetia, which is restricted to Africa)

Within the African species of Psydrax,

Bridson recognised two subgenera namely P. subgen.

Psydrax and P. subgen. Phallaria (Schum. &

Thonn.)Bridson. Using her key, the Australian

species are referrable to the former subgenus.

The Australian species of Psydrax agree

morphologically with those from outside

Australia, differing from them only in their

erect or subpatent rarely reflexed corolla lobes

and anthers, which are always reflexed in the

African species (Bridson 1985, p.688, f.l;

p.704, f.3; p.718, f.6; p.720, f.7), and by their

spinescent branchlets or thorns on the branches

of young plants in P. oleifolia.

Notes: In the keys and descriptions that follow,

the main peduncle is taken to be the main stalk

of the whole inflorescence, and the central or

solitary flower is the usually long-stalked

solitary flower which is situated in/near the

forks of the dichotomously branched cymes.

The measurement of the style includes that of

the stigma.

Key to species of Psydrax in Australia

1. Leaves whorled with 3-5 per node or opposite on the same branchlet,

canescent or subglabrous; branchlets canescent; N NT & N QLD. 1. P. paludosa

Leaves opposite or clustered on brachyblasts; branchlets and leaves never

canescent; hairs if present usually very fine and occurring on young

parts and inflorescence axes only. 2

2. Leaf blades linear or narrow elliptic, 2.5-4.0 (-5.5) mm wide, with margins

recurved or re volute; central Australia (WA, NT & SA). 3. P. suaveolens

Leaf blades variously shaped, usually more than 5.0 mm wide, with margins

flat or rarely recurved. 3

3. Leaves usually more than 10 cm long. 4

Leaves usually less than 10 cm long. 9

4. Leaf blades broad elliptic or elliptic ovate, about twice as long as wide, up

to 18 cm long and (3.1-) 4.0-8.7 cm wide; peduncles hairy. 5

Leaf blades mostly narrow elliptic or lanceolate, 2-3 times as long as

wide, rarely up to 18 cm long and 0.6-2.7 (-3.0) cm wide; peduncles glabrous. 6

5. Stipules with a prominent keel and a folded lobe at apex; leaf blades with

prominent, finely reticulate veins forming a closely arranged network;

peduncles slender, sparsely hairy; NE QLD. 9. P. tropica

Stipules without a keel and an apical lobe; leaf blades with prominent or

obscure, loosely arranged reticulate veins; peduncles robust, densely

hairy; E QLD & NE NSW . 7. P. lamprophylla

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

821

6. Leaf blades 3-10 times as long as wide, 0.6-1.5 (-2.0) cm wide, obscurely

veined; domatia present; inflorescences 5-15 (-22)-flowered; pedicel of

solitary flower (5-) 10-18 mm long; branchlets dark green coloured;

N NT & NE QLD. 17. P. saligna

Leaf blades 3-4 times as long as wide, 1.4-3.0 cm wide, prominently

veined; domatia present or absent; inflorescences more than 15-flowered;

pedicel of solitary flower (3.5-) 8-10 mm long; branchlets reddish brown,

brown or grey . 7

7. Leaf blades narrow elliptic, subfalcate, 11.5-17.0 (-18.5) cm long,

conspicuously nerved with lateral nerves ascending; domatia absent;

young branchlets 4-angular, red coloured; N WA & N NT.18. P. pendulina

Leaf blades narrow elliptic or lanceolate, usually less than 11 cm long;

lateral nerves oblique; domatia present or absent; young branchlets terete

or slightly angular, brown, red brown or gray coloured. 8

8. Leaf blades + shiny on adaxial surface; domatia usually present on most

leaves of a branchlet; flowers 5-merous; pedicels comparatively long

(those of solitary flowers 8-17 mm long); bark of branchlets tessellated;

NE QLD . 16. P. lepida

Leaf blades dull or with a slight sheen on adaxial surface; domatia usually

present only on some leaves of a branchlet, sometimes totally absent;

flowers 4- or 5-merous; pedicels comparatively short (those of solitary

flowers 8-11 mm long); bark of branchlets smooth; tropical WA, NT & QLD. . 15. P. attenuata

9. Leaf blades up to twice as long as broad, with conspicuous raised nerves

and close secondary reticulate venation (especially in dried specimens);

main lateral nerves in (3 or) 4-8 pairs. 10

Leaf blades not with the above set of characters, broad or narrow, with

lateral nerves and reticulate venation prominent or obscure, or veins

absent; main lateral nerves in 1-4 (-6) pairs. 12

10. Leaves with petiole 0.2-0.3 cm long; base of blade usually narrow and

attenuate into the petiole; domatia usually present on leaves; N QLD .... 14. P. pallida

Leaves with petiole 0.2-1.3 cm long; base of blade usually subcordate,

truncate or obtuse; domatia present or absent on leaves. 11

11. Domatia absent on leaves; leaf blades broad elliptic or ovate or

± suborbicular, dull, glabrous, smooth or scrabridulous; petioles

0.2-1.2 cm long; arid WA, NT, SA, SW QLD & NW NSW.11. P. latifolia

Domatia usually present on leaves; leaf blades obovate or broad elliptic,

shiny on adaxial surface, glabrous, smooth; petioles 0.2-0.6 cm long;

Cape York, QLD. 13. P. reticulata

12. Emits obovoid, 1.3-2.0 cm long x 0.8-1.1 cm wide. 13

Emits obovoid or ellipsoid, 0.4-1.0 cm long x 0.3-0.9 cm wide. 14

13. Inflorescences on slender peduncles 0.3-1.6 cm long; branching lax;

branches ending in 3-7-flowered cymules; flowers 5-merous; leaf blades

6.0-9.0 cm long x 3.0-6.0 cm wide, shiny on adaxial surface; domatia

present on leaves, conspicuous; fruits 1.3-1.5 cm long x 0.8-1.0 cm

wide; NE QLD. 8. P. laxiflorens

Inflorescence on long, stout, + erect peduncles 2.0-6.0 cm long; branches

and pedicels comparatively robust when dry; terminal cymes 8-12-

flowered; flowers 4-merous; leaf blades usually 3.6-6.6 cm long x 1.5-3.3 cm

wide, dull on adaxial surface; domatia on leaves small and inconspicuous

or absent; fmits 1.5-2.0 cm long x 0.9-1.1 cm wide; NE QLD. 5. P. montigena

822 Austrobaileya 6 (4): 817-889 (2004)

14. Branchlets and peduncles finely hairy. 15

Branchlets and peduncles glabrous. 20

15. Leaves usually more than 5.0 cm long. 16

Leaves usually less than 5.0 cm long. 17

16. Leaf blades narrow elliptic or sublanceolate, 0.8-1.8 cm wide, dull on

adaxial surface; reticulate v eins inconspicuous; domatia absent; central WA. 4. P. rigidula

Leaf blades elliptic, elliptic-ovate or subobovate, (2.2-) 3.0-4.4 (-5.2) cm

wide, usually very glossy on adaxial surface; reticulate veins apparent;

domatia usually present; N WA, N NT, E QLD & NE NSW. 6. P. odorata

17. Domatia present on leaves, prominent; leaf blades narrow elliptic or elliptic,

2.6-4.5 cm long x 0.7-2.0 cm wide, dull on adaxial surface; lateral

nerves indistinct; E QLD. 21. P. johnsonii

Domatia absent from leaves or if present obscure; leaf blades elliptic, narrow

elliptic, suborbicular or obovate, 0.8-5.0 cm long x 0.4-3.0 cm wide,

usually shiny on adaxial surface; lateral nerves distinct. 18

18. Inflorescences 5-21-flowered, comparatively small, slender; main

peduncles slender, 0.15-1.0 cm long; corolla 4.5-5.5 mm long,

± chartaceous; leaf blades elliptic, (0.8-) 2.2-4.6 cm long x (0.4—) 1.0-1.8 cm

wide, rarely more, slightly shiny on adaxial surface; NE QLD & PNG . . . 2. P. graciliflora

Inflorescences usually more than 21-flowered; main peduncles slender or

stout, (0.3-) 0.8-4.6 cm long; corolla 4.0-7.5 mm long, chartaceous or

fleshy; leaf blades elliptic or obovate, 1.5-5.0 cm long x 0.9-3.0 cm

wide , usually very glossy on adaxial surface. 19

19. Leaf blades obovate, 3.2-5.0 cm long x 1.6-3.0 cm wide, rarely more;

primary veins and secondary, reticulate veins prominent; inflorescence

with main peduncles 1.5-4.6 cm long, slender; corolla 6.0-7.5 mm long,

chartaceous, drying straw coloured; corolla tube cylindrical; Cape York,

QLD. 10. P. banksii

Leaf blades elliptic or suborbicular, 1.5-5.0 cm long x 0.6-2.5 cm wide;

primary veins prominent; secondary veins obscure; inflorescence with

main peduncles (0.3-) 0.8-2.0 cm long, stout or slender; corolla 4.0-5.0 mm

long, fleshy, drying brown; corolla tube broad campanulate; N WA,

N NT, E QLD & NE NSW. 6. P. odorata

20. Leaf blades 2.0-2.8 cm long x 1.1—1.3 cm wide; inflorescences 5-17-

flowered; petioles and peduncles 0.2-0.4 cm long; central & NW QLD . . 22. P. forsteri

Leaf blades more than 3.0 cm long and 1.3 cm wide; inflorescences more

than 17-flowered; petioles and peduncles usually more than 0.4 cm long. 21

21. Domatia present on leaf blades, prominent. 22

Domatia absent from leaf blades or if present inconspicuous. 24

22. Leaf blades 1.4-4.4 cm wide, usually up to twice as long as wide, mostly

very glossy on adaxial surface; flowers usually crowded on the branches

of the inflorescence; pedicels stout, comparatively short, those on

branches of the cyme almost obsolete; flowers mostly 4-merous, corolla

fleshy; N WA, N NT, E QLD & NE NSW. 6. P. odorata

Leaf blades 0.6-2.7 (-3.0) cm wide, usually more than 3 times as long as

wide, dull or slightly shiny on adaxial surface; flowers loosely arranged

along branches of the inflorescence; pedicels comparatively long and

slender; flowers mostly 5-merous, corolla chartaceous. 23

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax 823

23. Leaf blades 0.6-1.5 (-2.0) cm wide, (3-) 4-10 times as long as wide;

nerves and veins usually indistinct; inflorescences 5-15 (-22)-flowered,

branched once only; branchlets dark green; N NT & NE & central QLD. 17. P. saligna

Leaf blades 1.0-2.7 (-3.0) cm wide, up to 4 t im es as long as wide; nerves

and veins prominent; inflorescences (8-) 13-41-flowered, usually

branched more than once; branchlets brown, reddish brown or grey;

tropical WA, NT & QLD. 15. P. attenuata

24. Leaf blades (0.7-) 1.7-2.4 (-3.3) cm wide, concolorous, with obscure or

distinct nerves; reticulate veins and domatia very rarely present;

inflorescences usually compact, the flowers closely arranged on branches;

S QLD & N NSW. 19. P. oleifolia

Leaf blades (1.6-) 2.4-4.8 cm wide, somewhat discolourous; nerves and

reticulate veins prominent; domatia usually present on some leaves of a

branchlet; inflorescences usually open, the flowers loosely arranged on

branches. 25

25. Leaf blades to 4.8 cm wide, with 3-6 pairs of conspicuous lateral nerves;

flowers mostly 4-merous; corolla 5.0-6.5 mm long; pedicels

comparatively short, that of solitary flowers 0.2-0.35 cm long; young

branchlets usually stout, reddish brown coloured; central Australia (WA,

NT & SA). 12. P. ammophila

Leaf blades 1.0-3.0 (-4.0) cm wide, with 2 or 3 pairs of prominent lateral

nerves; flowers 4- or 5-merous; corolla 6.0-11.0 mm long; pedicels

comparatively long, that of solitary flowers (0.3-) 0.5-1.0 cm long; young

branchlets slender, reddish brown, brown or grayish coloured. 26

26. Petioles (1.2-) 2.0-3.5 cm long; leaf blades 2.6-4.0 cm wide, rarely less,

up to twice as long as wide; corolla 7.5-11.0 mm long; pedicel of solitary

flowers (0.3-) 0.5-0.7 cm long; branchlets whitish coloured, or pale

grey mottled with white; central & SE QLD. 20. P. longipes

Petioles 0.5-2.2 cm long; leaf blades 1.6-2.7 (-3.0) cm wide, 3-4 times as

long as wide; corolla 5.5-8.0 mm long; pedicel of solitary flowers (0.5-)

0.8-1.0 cm long; branchlets brown, yellowish brown, reddish brown or

grayish coloured; tropical WA, NT & QLD. 15. P. attenuata

A conspectus of some diagnostic attributes

of Psydrax in Australia

Leaves usually in whorls and plants usually

canescent: P. paludosa

Leaf blades linear with revolute or recurved

margins: P. suaveolens

Leaf blades comparatively large, (8.5-)

10.5- 20 cm long; stipules without a

keel or an apical lobe: P. lamprophylla

Leaf blades comparatively small, 0.9-8.3 (-9.3) cm

long: P. ammophila, P banksii, P. forsteri,

P. graciliflora, P odorata, P. oleifolia

Both leaf blades and inflorescences

comparatively small (1.0-6.0 cm long):

P ammophila, P. forsteri, P. graciliflora,

P. odorata

Leaf blades shiny or exceedingly glossy on adaxial

surface: P. banksii, P. lamprophylla,

P laxiflorens, P lepida, P. odorata, P tropica

Leaf blades with a slight sheen on adaxial surface:

P. attenuata, P. johnsonii, P. lepida,

P montigena, P. odorata, P. saligna

Leaf blades dull on adaxial surface: P. ammophila,

P. attenuata, P. johnsonii, P. oleifolia,

P rigidula, P. saligna

824

Austrobaileya 6 (4): 817-889 (2004)

Leaf blades concolorous: P. oleifolia

Leaf blades comparatively broad, less than 4

times as long as wide, with reticulate

venation conspicuous: P. latifolia,

P. pallida, P. reticulata, P. tropica

Leaf blades comparatively broad, less than 4

times as long as wide, with reticulate

venation inconspicuous: P ammophila

Leaf blades comparatively narrow, 4-10 times

as long as wide: P. attenuata, P. lepida,

P. pendulina, P. saligna

Leaves, branchlets and peduncles exceedingly

stout in dried specimens: P. montigena

Domatia on leaves mostly prominent: P. attenuata,

P. johnsonii, P lamprophylla, P. laxiflorens,

P lepida, P. montigena, P. odorata, P pallida,

P reticulata, P saligna

Domatia on leaves mostly small and obscure:

P. ammophila, P. attenuata, P. banksii,

P longipes, P montigena, P. tropica

Domatia absent from leaves: P. ammophila,

P. attenuata, P. odorata, P. forsteri,

P. latifolia, P. longipes, P. oleifolia,

P. pendulina, P. rigidula

Species Groups

Although species such as Psydrax paludosa,

P suaveolens, P. lamprophylla, P. montigena

and P. graciliflora are very distinctive and

readily recognisable as indicated in the key and

conspectus above, others belong to groups of

related somewhat similar species and are not

so easy to distinguish. These groups are as

follows.

(a) The Psydrax odorata group:

Psydrax odorata and its related species

P banksii, P. lamprophylla, P. laxiflorens and

P. tropica are characterised by their glossy,

coriaceous leaf blades with distinct nerves and

veins and usually prominent domatia, their

usually large, many-flowered inflorescences

with short and stout or slender pedicels, and

their flowers with fleshy or coriaceous corollas.

These species may be distinguished as follows.

1. Stipules neither keeled nor attenuated into a folded lobe at apex; leaf blades

(8.5-) 10.5-18.0 (-20.0) cm long x (3.1-) 4.0-8.7 (-9.7) cm wide; main

peduncles exceedingly robust, hairy. 7. P. lamprophylla

Stipules keeled and attenuated into a folded lobe at apex; leaves and

peduncles not with above set of characters. 2

2. Fruits obovoid, 1.3-1.5 cm long x 0.8-1.0 cm wide; inflorescences

exceedingly loosely branched, with long slender branches and scattered,

mostly 3-7-flowered cymules; domatia present on the leaves, conspicuous

. 8. P. laxiflorens

Fruits obovoid or ellipsoid, 0.5-0.8 cm long x 0.3-0.9 cm wide;

inflorescences more or less compact, usually with many-flowered short

branches, the ultimate cymules mostly more than 7-flowered; domatia

present or absent on leaves, prominent or obscure. 3

3. Leaf blades 10.0-14.5 (-18.0) cm long x 5.2-7.0 (-8.0) cm wide, usually

shortly acuminate or subcaudate at apex, with prominent reticulate veins

fine and forming a closely arranged network; stipules usually broad ovate,

acuminate, scarious along margins. 9. P. tropica

Leaf blades 1.5-8.3 (-9.2) cm long x 0.6-4.2 (-5.2) cm wide, obtuse,

rounded or subacute at apex, with veins if visible, usually loosely

reticulate; stipules ovate, with a long or short apical lobe, coriaceous. 4

4. Leaf blades elliptic, elliptic-ovate, subrhombic or narrow elliptic, 5.0-9.3 cm

long x 2.5-5.2 cm wide, rarely smaller; domatia usually prominent. 6. P. odorata

Leaf blades obovate, elliptic, suborbicular or narrow elliptic, 1.5-5.0 cm

long x 0.6-2.5 (-3.0) cm wide, rarely larger; domatia prominent or

obscure, or absent. 5

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax 825

5. Leaf blades obovate or subelliptic, rounded at apex, narrowed into petiole

proximally; inflorescence on long slender stalks 1.5-4.6 cm long; pedicels

slender; corolla scarious, tube narrow campanulate. 10. P. banksii

Leaf blades elliptic, narrow elliptic or suborbicular, obtuse or subacute at

apex and base; inflorescence on stout or slender peduncles 0.3-2.0 cm

long; pedicels usually stout and short, sometimes almost obsolete; corolla

fleshy, tube broad campanulate. 6. P. odorata

(b) The Psydrax latifolia group:

Psydrax latifolia and its related species

P. ammophila, P. pallida and P. reticulata are

characterised by broad, usually thick, rigid,

conspicuously nerved and reticulate veined leaf

blades especially in dried leaves. These species

may be distinguished as follows.

1. Petioles 2.0-3.0 cm long; leaf blade base subacute or acute and attenuate

into the petiole; lateral nerves in 3-5 pairs; domatia present. 14. P. pallida

Petioles 0.2-1.3 cm long; leaf blade base broad obtuse, truncate, subcordate

or rounded though sometimes subacute and attenuate into the petiole;

lateral nerves in 4-9 pairs; domatia present or absent. 2

2. Petioles 0.2-0.6 cm long; leaf blades obovate or broad elliptic, slightly

shiny on adaxial surface; domatia usually present. 13. P. reticulata

Petioles (0.2-) 0.5-1.3 cm long; leaf blades elliptic, broad ovate or

suborbicular, dull on adaxial surface; domatia absent or occasionally present. 3

3. Domatia absent on leaves; leaf blades (3.0-) 4.2-6.0 (-8.7) cm wide, with

apex slightly rounded or truncate and base subcordate or obtuse; lateral

nerves in (4-) 5-9 pairs; reticulate veins prominent, usually closely

arranged, raised in dried specimens, usually visible on abaxial surface. 11. P. latifolia

Domatia present on some leaves of a branchlet or absent; leaf blades 1.2-3.5

(-4.8) cm wide, with apex and base obtuse or subacute; lateral nerves in

(3-) 4-6 pairs; reticulate veins obscure, openly arranged, usually

not visible on abaxial surface. 12. P. ammophila

Psydrax attenuata and its related species

P. lepida, P. pendulina and P. saligna are

characterised by narrow elliptic, lanceolate or

elliptic leaf blades which are usually provided

with domatia (though domatia are absent in

P. pendulina and sometimes in P. attenuata) and

prominent oblique ascending nerves (which are

sometimes obscure in P. saligna ), open, laxly

branched inflorescences and chartaceous 4- or

5-merous flowers on long slender pedicels.

These species may be distinguished as follows.

1. Domatia present on all leaves or only on one or two leaves on the one branchlet. 2

Domatia absent on leaves. 4

2. Leaf blades 0.6-1.8 cm wide, 4-10 times as long as wide; nerves distinct

or indistinct; veins indistinct; inflorescences comparatively small, 5-15

(-22)-flowered; branchlets dark reddish brown or blackish grey coloured ... 17. P. saligna

Leaf blades 1.0-3.0 (-4.2) cm wide, usually 3 to 4 times as long as wide;

nerves and veins distinct; inflorescences comparatively large, to

50-flowered; branchlets reddish-brown, brown or greyish coloured

826

Austrobaileya 6 (4): 817-889 (2004)

3. Domatia usually present on all the leaves of a branchlet; leaf blades shiny

on adaxial surface; peduncle of inflorescence 1.0-1.5 cm long; flowers

5-merous, densely hairy at mouth of corolla tube; pedicel of solitary

flowers 0.8-1.7 cm long. 16. P. lepida

Domatia rarely present on all the leaves of a branchlet; leaf blades dull on

adaxial surface; peduncle of inflorescence 0.3-0.6 cm long; flowers 4-

or 5-merous, sparsely hairy at mouth of corolla tube; pedicel of solitary

flowers (0.35-) 0.8-1.1 cm long. 15. P. attenuata

4. Leaf blades usually 11.0 to 18.5 cm long, more than 5 times as long as

wide, usually thin coriaceous; lateral nerves conspicuous, exceedingly

oblique; young branchlets quadrangular distally, reddish coloured . 18. P. pendulina

Leaf blades usually less than 11.0 cm long, about 3-4 times as long as

wide, thick coriaceous; lateral nerves prominent, oblique or exceedingly

oblique; young branchlets terete or slightly angular distally, greyish or

brownish coloured. 15. P. attenuata

(d) The Psydrax oleifolia group:

Psydrax oleifolia and its related species

P. forsteri, P. johnsonii and P. longipes are

characterised by dull or slightly shiny thick leaf

blades with obscure or conspicuous nerves, with

or without reticulate veins and usually without

domatia (though domatia are present in

P. johnsonii and sometimes in P. longipes ), and

usually 4-merous flowers (though 5-merous

flowers occur in P. forsteri and sometimes in

P. longipes ). These species may be

distinguished as follows.

1. Axes of young branchlets and inflorescences hairy; domatia present on

leaves, prominent. 21. P. johnsonii

Axes of young branchlets and inflorescences glabrous; domatia absent on

leaves or, if present, inconspicuous . 2

2. Leaves and inflorescences comparatively small; leaf blades 2.0-2.8 cm

long x 1.1-1.3 cm wide; inflorescences 5-17-flowered; peduncles

2-4 mm long. 22. P. forsteri

Leaves and inflorescences usually comparatively large; leaf blades (3.5-)

5.5-10.0 cm long x (0.7-) 1.7-4.0 cm wide; inflorescences 15-39-

flowered; peduncles usually longer than 4 mm. 3

3. Petioles 0.6-1.2 cm long; leaf blades (0.7-) 1.7-2.4 (-3.3) cm wide,

concolorous;domatia and reticulate veins usually absent; nerves obscure

or distinct; inflorescences usually much smaller than leaves and compact;

flowers with corolla 4.0-6.5 mm long; fruits 4.5-6.0 mm across. 19. P. oleifolia

Petioles 1.2-2.5 (-3.5) cm long; leaf blades (1.7-) 2.6-4.0 cm wide,

discolorous; domatia present on some leaves of a branchlet; nerves and

reticulate veins distinct; inflorescences open; flowers with corolla

7.5-11.0 mm long; fruits (0.4-) 0.6-0.7 cm across. 20. P. longipes

1. Psydrax paludosa S.T.Reynolds &

R.J.F.Hend., sp. nov. a speciebus

Australiae omnibus ceteris generis foliis

plerumque ternatis et oppositis et

canescentibus differt. Typus: Queensland.

Cook District: Weipa, track between

Telegraph line and 20 Mile, S of Marmoss

Creek, 12°41’S, 142°05’E, 4 January

1983, A. Morton AM 1773 (holo: BRI; iso:

MEL).

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

Canthium sp. (Weipa, A.Morton AM 1773),

S.T. Reynolds (1997, p. 181), P.I. Forster

& D.A. Halford (2002, p.174).

Canthium sp. (Wenlock River, J.R.Clarkson

8418+)* 1 , S.T. Reynolds (1997, p.181).

Shrubs 2-8 m high, with branches horizontal;

bark mottled; branchlets finely hairy or

glabrous, dark reddish-brown, reddish coloured

distally. Leaves whorled, with 3-5 leaves at

each node, or opposite; stipules comparatively

small, broad ovate with a short lobe at apex,

hairy or glabrous abaxially, hairy at base

adaxially; petioles 0.1-0.8 cm long; blades

elliptic, narrow elliptic-subobovate or narrow

obovate, (1.0-) 2.2-5.2 cm long x (0.4-) 1.2-1.7

cm wide, with apex obtuse and base acute or

subacute, thin coriaceous; adaxial and abaxial

surfaces finely white hairy or glabrous; lateral

nerves consisting of 1 or 2 pairs, obscure,

oblique to midrib and ascending distally, or

absent; reticulate veins absent; domatia one on

each side of midrib and obscure, or absent.

Inflorescences 1.8-2.5 cm long x 1.8-2.5 cm

wide, 8-30-flowered; peduncles hairy or

subglabrous, the basal one 6.0-8.0 mm long

with a pair of small ovate bracts at its distal

end; axis branches 3.0-5.0 mm long. Flowers

4-merous; pedicel of solitary flowers 3.0-5.0

mm long, that of others 0.2-2.0 mm long; calyx

2.0-2.5 mm long, with tube campanulate and

limb 4-denticulate; lobes ovate, 0.5-0.8 mm

long x c. 0.75 mm wide; corolla 3.5-7.0 mm

long, cream or lemon yellow; tube inflated, ±

campanulate, 1.5-3.5 mm long x 2.0-2.5 mm

wide at top, densely hairy at throat; lobes

elliptic, 2.0-4.5 mm long x c. 1.5 mm wide,

subacute or obtuse, erect or recurved; stamens

exserted; filaments 1.5-2.0 mm long; anthers

1.5-2.0 mm long, erect; disc hairy; style (with

stigma) 6.0-8.0 mm long; stigma ellipsoid,

c.0.75 mm long, bifid at apex. Fruits (only

immature ones seen) ellipsoid or obovoid,

1.0-1.2 cm long x 0.8-1.0 cm wide, smooth,

exceedingly fleshy; pyrenes ellipsoid, smooth,

thin, crustaceous; endosperm thick. (Fig. 1)

Additional representative specimens : Northern Territory.

BingBong Station (15°44’S, 136°20’E), Nov 1977, Craven

4694 (DNA). Queensland. Cook District: Archer River,

13°25’S, 142°10’E, Sep 1974, Hyland 7712 (BRI, QRS);

ditto, Archer Bend, Sep 1974, Tracey 14355 (BRI, QRS);

along road to Inkerman, from Burke Development Road,

16°24’S, 142°34’E, Jan 1987, Dalliston 52 (BRI); 4 km S

827

of Dunbar Homestead, 16°04’S, 142°20’E, Jul 1987,

Dalliston 18 (BRI); track to James Creek, 4 km N of Escott

Homestead, 17°29’S, 139°14’E, Jul 1987, Dalliston 216

(BRI); 56 kmW of Rokeby Homestead, 13°27’S, 142°16’E,

Jul 1988, Dalliston 531 (BRI); 8.8 km S of Wenlock River

on Peninsula Development Road, 12°31’S, 142°39’E,Apr

1990, Clarkson 8418 8cNeldner{ BRI)* 1 ; 2.4km Wof Lydia

Creek on Mission River Road, 12°33’S, 142°34’E, Apr 1990,

Clarkson 8600 & Neldner (BRI)* 1 ; Cape York, Edward River

near Mitchell Downs, 14°46’S, 141°40’E, Apr 1990, Taplin

CY26 (BRI); about 45 km due E of Kowanyama Aboriginal

Community, 15°28’S, 142°13’E, Jun 1992, Blackman G33

(BRI).

Distribution and habitat: Northern Australia

from Arnhem Land, Northern Territory, to Cape

York Peninsula, Queensland; usually in

standing water on the edge of waterways and

flooded drainage lines in Eucalyptus tetrodonta

woodland and in swampy areas with Melaleuca

species. (Map 1)

Diagnostic attributes: Psydrax paludosa is

readily distinguishable from other Australian

species of this genus by its whorled (with

usually 3-5 leaves at each node) and opposite

leaves on the same branchlet, canescent

branchlets and leaf blades, comparatively small

inflorescences, and conspicuously fleshy fruits

with thin crustaceous, smooth-surfaced pyrenes.

Notes: Although most of the specimens seen

have the typical canescent branchlets, a form

with glabrous or subglabrous branchlets (as

represented by specimens marked *' above) also

exists.

Etymology: The specific epithet paludosus ,

Latin for ‘marshy, swampy or boggy’, refers to

the usual habitat of this species.

2. Psydrax graciliflora (Merr. & L.M.Perry)

S.T.Reynolds & R.J.F.Hend., comb, nov.;

Canthium graciliflorum Merr. &

L.M.Perry, J. Arnold Arb. 26: 230-231

(1945). Type: Papua New Guinea.

Tarara, Wassi Kussa River, December

1936, L.J. Brass 8596 (holo: n.v.; iso:

BRI, K).

Shrubs or small trees (1-) 2-8 (-10) m high;

trunk 10-15 cm dbh, coppicing at base, fluted;

bark whitish coloured; branchlets quadrangular

and ridged distally, minutely hairy, dark

greyish-brown, young ones often slightly

zigzagged. Leaves opposite; stipules broad

triangular, keeled, attenuated into a long, broad,

828

Austrobaileya 6 (4): 817-889 (2004)

Fig. 1 . Psydraxpaludosa (typical form). A. flowering branch x 1. B. inflorescence x 3. C. flower x 6. D. LS of flower x 6. E.

Fruit x 3. F. FS of fruit x 3. G. pyrene x 6. A-G, Clarkson 8600 & Neldner (BRI). Psydrax paludosa (hairy form). H. part

of inflorescence x 3. H, Morton AM1773 (BRI).

lanceolate lobe or sometimes into a small and

short lobe at apex, somewhat scarious, shiny;

petioles (0.1-) 0.2-0.4 cm long, glabrous or

sparsely hairy; blades elliptic or sometimes

narrow elliptic, (0.9-) 2.2-4.6 (-5.2) cm long

x (0.4-) 1.0-1.8 (-2.0) cm wide, with apex and

base usually subacute, glabrous on both surfaces

with adaxial surface bright green and slightly

shiny, the abaxial one slightly darker and dull;

lateral nerves in 3-5 pairs, slightly patent or

suboblique and looping at margins, fine,

obscure or absent on abaxial surface; domatia

absent or rarely when present 1 or 2 on each

leaf and inconspicuous. Inflorescences 1.2-1.4

cm long x 1.3-2.2 cm wide, 5-17 (-21)-

flowered; peduncles and pedicels minutely

hairy, terminating in 2 or 3 branched cymes,

the main peduncle 1.5-10.0 mm long, with

minute bracts at its distal end; axis branches

4.0-5.0 mm long, ultimate cymules usually 5-

flowered. Flowers 4(or 5)-merous; pedicel of

solitary flowers 4.5-7.0 mm long, that of others

2.0-5.0 mm long; calyx 1.75-2.0 mm long,

with tube subturbinate and limb 4-denticulate;

lobes ovate, 0.25-0.5 mm long x 0.75-1.0 mm

wide; corolla 4.5-5.5 mm long, white or cream;

tube campanulate, 1.0-1.75 mm long x c.1.5

mm wide at top, hairy at throat; lobes narrow

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

elliptic, 3.5-4.5 mm long x 1.0-1.5 mm wide,

obtuse, erect or ± spreading; stamens as long

as the corolla lobes; filaments 1.5-2.0 mm long;

anthers 2-3 mm long, erect, ± recurved at

anthesis; style (with stigma) 4-6 mm long;

stigma oblongoid, c.1.5 mm long, bilobed at

apex. Fruits subglobose or broad ellipsoid,

3.0-4.5 mm long x 3.0-6.5 mm wide, black

when ripe; pyrenes ellipsoid, slightly rugose.

Representative specimens : Papua New Guinea: Tarara,

Wassi Kussa River, Dec 1936, Brass 8596 (BRI, K).

Australia: Queensland. Cook District: Timber Reserve 14,

Parish of Kesteven, 13°43’S, 143°19’E,Mar 1962, Hyland

11757 (BRI); Lankelly Creek, Mcllwraith Range, Oct 1969,

Webb & Tracey 9559 (BRI, QRS); Mt Tozer near Iron Range,

12°45’S, 143°12’E, Nov 1977, Tracey 14871 (BRI); near

Lockerbie, 10°47’S, 142°28’E, Feb 1980, Hyland 10281

(BRI); Summit of Mt Tozer, 12°44’S, 143°12’E, Jun 1980,

Morton AM847 (BRI); National Park Reserve 8, Parish of

Weymouth, 12°37’S, 143°21’E, Mar 1982, Hyland 11739

(BRI); Endeavour River, 15°25’S, 145°15’E, Dec 1983,

Langford s.n. (BRI, QRS); Crusher Creek, Moa Island,

10°15’S, 142°15’E, Apr 1987, Budworth 975 (BRI).

Distribution and habitat : South coast of Papua

New Guinea to Cape York Peninsula, Australia,

including Torres Strait Islands; on ranges,

ridges, creek and riverbanks, in evergreen or

semi-deciduous notophyll vine forests. (Map

2 - Queensland only)

Diagnostic attributes: Psydrax graciliflora is

characterised by small, thick, finely nerved,

elliptic leaf blades and small, fragile

inflorescences with small flowers on slender

pedicels, hairy branchlets and inflorescences,

and ± scarious stipules which are keeled and

attenuated into a long broad (though sometimes

short) lobe at the apex.

Notes: A specimen recorded under the name

Canthium sp. (Altanmoui Range D.G.Fell+

DGF2702) by Reynolds (1997, p.180) and

others previously filed under that name in BRI

have small leaves but they probably represent

distinct forms of this species. However, as they

are sterile it is not possible to be certain of this.

These specimens are as follows.

Queensland. Cook District: Altanmoui Range, Cape

Melville National Park, 7 km E of Wakooka Outstation,

14°34’S, 144°26’E, Oct 1992, Fell DGF2702 & Stanton

(BRI). North Kennedy District: northern end of Little

Ramsay Bay, Hinchinbrook Island, 18°20’S, 146°19’E,Aug

1975, Sharpe 1678 (BRI); Mulligan Bay, Hinchinbrook

Island, 18°27’S, 146°20’E, Jan 1987, Godwin & Stanton

C3013 (BRI); tributary of Mulligan Creek, Hinchinbrook

829

Island, 18°26’S, 146°19’E, Sep 1994, Cumming 13353

(BRI).

If correctly placed, the Cape Melville

collection by Fell & Stanton has the smallest

leaves seen for this species whereas the

Hinchinbrook Island collections have smaller

and narrower leaves than are usual for it.

Moreover, P. graciliflora has sofar not

definitely been collected from as far south as

these latter specimens were.

3. Psydrax suaveolens (S.Moore) S.T.Reynolds

& R.J.F.Hend., comb, nov.; Canthium

suaveolens S.Moore, J. Linn. Soc. Bot.

34: 194 (July 1899). Type: Western

Australia. Repperi juxta Mt Margaret,

1894-95, S. Moore s.n. (holo: BM).

Canthium lineare E.Pritz., Feddes Repert.

15(3): 359 (Dec 1918), Plectronia

linearis (E.Pritz.) J.M.Black, Trans. &

Proc. Roy Soc. S. Aust. 59: 261 (1935),

syn. nov. Type: Northern Territory. Central

Australia, Hermannsburg ad flumen

Fincke, 1906-1908, Strehlow 88 (n.v.).

Slender shrubs 2-3.5 m high; bark smooth,

light or bluish grey coloured; young parts and

inflorescences hairy; branchlets greyish or

reddish coloured, occasionally with a whitish

bloom distally. Leaves opposite or sometimes

clustered on brachyblasts; stipules ovate,

slightly keeled, attenuated into a folded lobe at

apex, with colleters present at the base

adaxially; petioles 0.1-0.2 cm long; blades

linear, very narrow elliptic or narrow oblong

obcuneate, (2.1-) 4.0-6.2 cm long x 0.25-0.4

(-0.55) cm wide, with apex obtuse, base

subacute and narrowing into the very short

petiole, and margins revolute or recurved, rigid

on drying; adaxial and abaxial surfaces

glabrous; midrib deeply channelled above,

raised below; nerves and reticulate venation not

apparent. Inflorescences with main peduncles

2-5 mm long with minute bracts at its distal

end, terminating in 2 branched cymes; axis

branches c.2.0 mm long, cymules 2-5-flowered.

Flowers 5-merous; pedicel of solitary flowers

3.5-5.0 mm long, of others 2.0-4.5 mm long;

calyx 2.0-2.5 mm long, with tube cupular,

flared to a 5-dentate limb; lobes ovate-

acuminate or ovate; corolla 4.0-7.5 mm long,

white or cream; tube ± cylindrical, 1.5-2.5 mm

830

Austrobaileya 6 (4): 817-889 (2004)

long x 2.0-2.5 mm wide, sparsely hairy at

throat; lobes elliptic, 3.0-6.5 mm long x 2.0-2.5

mm wide, subacute, erect; stamens slightly

exserted; filaments 1.5-2.5 mm long; anthers

2.0-3.0 mm long, erect or patent; disc shorter

than the calyx limb, glabrous; style (with

stigma) 7.0-10.0 mm long; stigma oblongoid,

bifid at apex. Fruits transversely ellipsoid or

subglobose, 5.0-6.0 mm long x 6.5-8.5 mm

wide, 2-celled, black and shiny when ripe,

fleshy and juicy; pyrenes exceedingly rugose

on abaxial surface. (Fig. 2A-2E)

Additional representative specimens : Western Australia.

Rawlinson Range, S of Sladen Water (24°5-’S, 128°1-’E),

Feb 1935, Finlayson s.n. (AD); 24 mi les (c.38.4 km) East

Blackstone Ranges (25°56’S, 128°08' E), Jun 1958,

Chippendale 4524 (DNA, PERTH); 60 miles (c.96 km) N

of Paynes Find, Apr 1960, George 710a (PERTH); Yerilla

Station (29°28’S, 121°41’E), Aug 1985, Burnside s.n.

(PERTH); 1.4 km W of No 17 Bore, Berwidgie Station, Jun

1988, Cranfield 6845 (PERTH); West Angelas, Dec 1995,

Janicke Ub6 (PERTH). Northern Territory. Napperby,

about 100 mi les (c. 160 km) NW of Alice Springs, 22°35’S,

132°45’E, Jan 1930, Everist 4194 (AD, BRI); 11 miles

(c. 17.6 km) W of Ayers Rock along Ayers Rock and Docker

Creek road (25°20’S, 130°49’E), Jan 1969, Maconochie 650

(AD, CANB, DNA); CSIRO Mulga enclosure (23°28’S,

133°48’E), Dec 1971, Latz 1867 (DNA); MacDonnell

Ranges, Nov 1885, Schwarz s.n. (MEL); Mt Windsor

(23°50’S, 130°51’E), May 1988, Leach 1993 & Latz (DNA).

South Australia, about 19 km W of Tallaringa Well, 28 °51 ’ S,

133°05’S, May 1964, Lothian 2746 (AD, BRI, DNA);

Everard Ranges between Boundary Bore and Gap Well

(27°05’S, 132°3-’E), Dec 1981 ,Kalotas 1003 (AD, DNA).

Distribution and habitat: Central Australia

from Barker and Rawlinson Ranges to south

of Leonora, Western Australia, ranges near

Alice Springs, Northern Territory (Haast’s Bluff

to Palm Valley Chalet), and northwestern plains

of South Australia (Everard Ranges to west of

Tallaringa Well); on top of scree slopes of

foothills and on sandy plains, usually with

Mulga (Acacia aneura F.Muell. ex Benth.:

Mimosaceae) in grassland, on red to red-brown,

sandy to sandy-clayey soils. (Map 2)

Diagnostic attributes : Psydrax suaveolens is

readily recognisable by its narrow, usually

linear leaves with re volute or recurved margins.

It is related to P. rigidula with which it shares

the hairy young branchlets and hairy, few-

flowered inflorescences with 5-merous flowers,

but that species differs from P. suaveolens in

its broader, distinctly nerved leaf blades with

flat or slightly recurved margins. There are,

however, occasional specimens which appear

to be intergrades between the two.

Uses: The fruits of this species are reported by

certain collectors to be edible.

4. Psydrax rigidula S.T.Reynolds &

R.J.F.Hend., sp. nov. aemulans P.

suaveolens (S.Moore) S.T.Reynolds &

R.J.F.Hend. cujus flores habet sed folii

lamina latiore et plusminusve rigida,

marginibus recurvis constructis differt.

Typus. Western Australia. 6 miles (c.9.6

km) N of Meekatharra, 6 November 1968,

H. Demarz D591 (holo: PERTH).

Small trees or shrubs 2-3 m high; bark

smooth, purplish grey coloured; branchlets

slightly quadrangular distally; young parts and

inflorescence axes finely hairy, reddish

coloured. Leaves opposite; stipules ovate,

keeled, attenuated into a long folded lobe at

apex; petioles 0.25-0.6 cm long; blades narrow

sublanceolate or elliptic, 52-1.5 cm long x 0.8-1.8

cm wide, with apex subacute or obtuse, base

subacute and attenuate into the petiole, and

margins slightly recurved, coriaceous, drying

coriaceous, yellowish coloured adaxially, dark

green abaxially; midrib sunken adaxially, raised

abaxially; lateral nerves in 4-7 pairs, oblique,

ascending; reticulate venation not apparent;

domatia absent. Inflorescences 1.5-2.3 cm

long x 2.8-3.5 cm wide, 17-45-flowered; main

peduncle 4.5-6.0 mm long, provided with

minute bracts medially; branches 2.5-4.5 mm

long; cymes 8-22-flowered. Flowers 5-merous;

pedicel of solitary flowers 3.5-4.5 mm long, of

others 2.5-3.0 mm long; calyx 2.5-3.0 mm

long x 1.5-2.5 mm wide, with tube cupular,

puberulous and limb 5-dentate, flared,

puberulous, sometimes ciliolate; lobes ovate,

c.0.5 mm long x 0.5 mm wide; corolla 6-7 mm

long, white or cream; tube cylindrical, 2.0-3.5

mm long x 2.0-2.5 mm wide, sparsely hairy at

throat; lobes elliptic to lanceolate, 4.0-5.5 mm

long x 1 . 5-2.0 mm wide, obtuse or subacute,

slightly hooded and papillose at apex; stamens

exserted; filaments 1.5-2.5 mm long; anthers

2.0-3.0 mm long, erect or ± patent; disc shorter

than calyx limb, glabrous; style (with stigma)

7-10 mm long. Fruits broad ellipsoid, 5.0-7.0

mm long x 7.0-9.0 mm wide; pyrenes

hemispherical, exceedingly rugose (especially

on the abaxial side). (Fig. 2F-2J)

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

831

Fig. 2. Psydrax suaveolens. A. flowering branch x 1. B. flower x 5. C. LS of flower x 5. D. fruit x 3. E. pyrene x 5. A-C,

Janicke U6b (PERTH); D&E, Symon 9961 (AD). Psydrax rigidula. F. flowering branch x 0.6. G. flower x 5. H. LS of flower

x 5.1. fruit x 4. J. pyrene x 5. F-H, Demarz D591 (PERTH); I&J, Gardner & Blackall 122 (PERTH).

832

Austrobaileya 6 (4): 817-889 (2004)

Additional representative specimens : Western Australia.

Kimberley, exact locality unknown, May 1874, Forrest’s

Expedition [MEL1538055] (MET,); N of Meekatharra, Jul

1931, Gardner s.n. (PERTH); Cue, between Mt Magnet and

Meekatharra (27°27’S, 117°53’E), Jul 1931, Gardner &

Blackall 122 (PERTH); Boolardy Station, Murchison River,

May 1936, Melville 37 (K, PERTH); 483 miles (c.773 km)

N of Meekatharra, Oct 1963, Lullfitz 2585 (PERTH); 10

miles (c.16 km) N of Callytharra, Oct 1972, Demarz 3968

(PERTH); 14 km SE of Leonora, Jul 1977, Parker 92

(PERTH); 2 km W of MidshinnerWell, Milly Milly Station

(25°48’S, 116 0 51’E), Apr 1986, Cranfield 5385 (PERTH).

Distribution and habitat : Western Australia,

from Mt Wendell to Warbuton and Robinson

Ranges (between 21° to 30°S and 115° to

126°E); along water courses, dry creek beds,

on lateritic soil. (Map 3)

Diagnostic attributes: Psydrax rigidula is

characterised by shortly stalked, thick, narrow,

sublanceolate or elliptic leaf blades which dry

a greenish yellow colour, and shortly stalked,

comparatively small, hairy inflorescences with

5-merous flowers. This species is related to

P suaveolens, with which it has similar bark,

leaf texture, inflorescence and flower attributes,

but that species differs from P. rigidula by its

narrow, linear leaf blades which usually have

re volute margins.

Uses: The leaves and fruits of Psydrax rigidula

are reported by collectors to be edible to stock

but the leaves are usually inaccessible to sheep.

Etymology: The specific epithet rigidulus,

Latin for ‘somewhat rigid’, refers to the more

or less rigid leaves in this species.

5. Psydrax montigena S.T.Reynolds &

R.J.F.Hend., nom. nov.; Ixora orophila

C.T.White, Proc. Roy. Soc. Qld 53: 220

(1942), nom. illeg. non Bremek., J. Bot.

London 75: 321 (1937). Type:

Queensland. Cook District: Thornton

Peak, 16°15’S, 145°25’E, alt. 4,500 feet,

14 December 1940, H. Flecker

NQNC7110 (holo: BRI).

Canthium sp. (Thornton Peak H.Flecker

NQNC7110), S.T. Reynolds (1997,

p.181), PI. Forster & D.A. Halford (2002,

P-174).

Shrubs or small trees 2.5-12 m tall; trunk

10.0-18.3 cm dbh, usually buttressed;

branchlets usually robust, ± angular distally,

glabrous, lenticellate. Leaves opposite; stipules

ovate, 4.0-6.0 mm long, keeled, attenuated into

a long folded lobe at apex; petioles (0.2-) 0.4-0.7

cm long; blades elliptic, (3.6-) 4.5-6.6 (-7.2)

cm long x (1.5-) 2.1-3.3 (-4.0) cm wide, with

apex and base obtuse or apex shortly

subacuminate, coriaceous, drying exceedingly

thick; both adaxial and abaxial surfaces

glabrous, the adaxial ones slightly glossy, the

abaxial ones dull; midrib conspicuous on

abaxial surfaces, drying paler than other parts

of the blade; lateral nerves in 4-6 pairs, slender,

obliquely arched from midrib; reticulate

venation not apparent; domatia small, usually

one on each side of midrib, sometimes absent.

Inflorescences 6.5-8.0 cm long (including

peduncle), 4.0-6.0 cm wide, 17-25-flowered;

peduncles glabrous, the main one robust, more

or less erect, 2.0-3.8 (-6.0) cm long; axis

branches 0.8-1.2 cm long; cymes 8-12-

flowered. Flowers 4-merous; pedicel of solitary

flowers 6.0-9.0 mm long, of others 1.5-2.0 mm

long; calyx 2.0-3.5 mm long, with tube ±

urceolate and limb 4-denticulate; lobes ovate,

c.0.5 mm long x 1.5 mm wide; corolla 5.0-9.0

mm long, white or cream; tube broad

campanulate, 1.0-3.0 mm long, 2.0-3.5 mm

wide at top, sparsely hairy at mouth; lobes

elliptic to sublanceolate, 4.5-6.0 mm long x

1.5-2.5 mm wide, obtuse or subacute with apex

incurved, fleshy, usually recurved; disc

glabrous; stamens shorter than corolla lobes;

filaments 1.5-2.0 mm long; anthers 3.0-3.5

mm long x 0.7-1.0 mm wide, exserted, erect;

style (with stigma) 6.5-9.0 mm long, exserted;

stigma 2.0-3.0 mm long. Fruits obovoid,

15.0-20.0 mm long x 9.0-11.0 mm wide, black

when ripe; pyrenes exceedingly rugose.

Additional representative specimens : Queensland. Cook

District: Russell River, in 1892, Johnson [MEL1538182,

MEL1538183] (MEL); N Mary Logging Area, Mt Lewis,

Oct 1973, Sanderson 458 (QRS); Thornton Peak, Nov 1973,

Hyland 7047 (BRI, K); Mt Hermet, Oct 1975, Hyland 3343

(BRI, K, QRS); Broken Nose, Mt Bartle Frere, Dec 1978,

Jago 251 (QRS); Thornton Peak, Jan 1986, Godwin 2912

(BRI); Daintree National Park, Mt Sorrow track before

razorback, 2.5 km W of Cape Tribulation, 16°04’S,

145°27’E, Dec 1997, Forster PIF21979, Booth , Jago &

Jensen (BRI).

Distribution and habitat: North Queensland,

from Thornton Peak to Mt Bartle Frere; in

montane rainforests bordering shrublands at

altitudes of 1000-1500 m. (Map 6)

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

Diagnostic attributes: Psydrax montigena is

readily distinguishable by its compact habit,

robust branchlets, exceedingly thick leaves,

long robust suberect peduncles, fleshy flowers

and comparatively large obovoid fruits with

very rugose pyrenes.

Notes: Psydrax montigena is a new name

published here for the plant previously known

as Ixora orophila C.T.White. White’s name is

illegitimate being a later homonym of

Bremekamp’s Ixora orophila , dating from

1937, which is applicable to a different species.

Etymology: The specific epithet montigena ,

Latin for ‘mountain-born’, refers to the habitat

of this species.

6. Psydrax odorata (Forst.f.) A.C.Sm. &

S.P.Darwin, FI. Vitiensis Nova 4: 230

(1988); Cojfea odorata Forst.f., FI. Ins.

Austr. Prod. 16 (1786); Canthium

odoratum (Forst.f.) Seem., FI. Vitiensis

132 (1866). Type: Vanuatu. Tanna Island,

date unknown, J.R. & G.Forster s.n.

(lecto: BM, fide A.C. Smith & S.P.

Darwin, op. cit. p.232).

Canthium lucidum Hook. & Am., Bot. Beechey

Voy. 65 (1832), nom. illeg. non R.Br.;

Canthium beecheyi Steud., Nomen. Bot.

ed.2, 1: 275 (1841); Plectronia

hookeriana Domin, Biblioth. Bot. 89: 622

(1929). Type: Gambier Islands, collector

unknown (holo: K n.v., fide A.C. Smith

& S.P. Darwin, op. cit. p.232).

Shrubs or small trees 2-12 m high, with

branches usually at right angles to the stem;

bark smooth or rough, dark or pale creamy grey,

mottled with white; branchlets, petioles, young

leaf blades and inflorescence axes with fine

minute hairs or glabrous; branchlets usually

slightly quadrangular and resinous distally, pale

brown or greyish white, drying brownish

coloured, lenticellate. Leaves opposite, very

variable; stipules triangular, 3.0-8.0 mm long,

keeled, abruptly contracted and attenuated into

a long folded lobe at apex, with the lobe either

conspicuous, + foliaceous, to 9.0 mm long or

narrow and acuminate, hairy or glabrous, with

colleters occasionally present at base adaxially;

petioles 0.1-1.1 cm long; blades elliptic,

elliptic-oblong, ovate-elliptic, suborbicular,

833

narrow elliptic, slightly rhombic or subobovate,

1.5-8.3 (-9.3) cm long x 0.6-4.4 (-5.6) cm

wide, with apex obtuse, subacute or shortly,

usually bluntly, acuminate, and base obtuse or

subacute and usually decurrent into petiole, thin

or thick coriaceous; adaxial and abaxial

surfaces glabrous, or hairy proximally along

the midrib on the abaxial surfaces on young

leaves; adaxial surfaces exceedingly glossy,

green or ± bluish green with nerves pale green,

or slightly matt then yellowish green with

whitish coloured nerves, and abaxial surfaces

pale green and dull, both drying brown with

nerves and margins paler coloured adaxially,

and pale brown abaxially, with both surfaces

sometimes blotched; lateral nerves in 1-4 (-6)

pairs, oblique with the angle to midrib usually

between 30°-45°, or subpatent, looping near

the margins, prominent or obscure on adaxial

surfaces, usually obscure or not apparent

abaxially; reticulate venation distinct or obscure

on adaxial surface; domatia 1-4 on each side

of the midrib, usually present in the axils of

the middle pairs of nerves, or absent.

Inflorescences 1.2-6.0 cm long x 2-6 cm wide,

(2-) 12-93-flowered; peduncles stout or

slender, finely hairy or glabrous, the main one

3.0-30.0 mm long provided with minute bracts

proximally or near the middle and terminated

by 2(or 3)-branched cymes; axis branches 1.0-11.0

mm long, usually branched at apex with

ultimate cymes to 35-flowered. Flowers 4- or

5-merous, sweetly perfumed; pedicels slender

or stout, hairy or glabrous, those of solitary

flowers 3.0-9.0 mm long, of others 0.5-4.0 mm

long; calyx 1.0-2.0 mm long with tube cupular

and limb 4- or 5-denticulate; lobes broad ovate,

c.0.5 mm long and wide, glabrous; corolla 3.0-7.0

mm long, fleshy, white or cream; tube 0.5-2.0

mm long, usually sparsely deflexed hairy at

throat; lobes lanceolate or + oblanceolate,

2.0-5.0 mm long x 0.75-1.5 mm wide, obtuse,

thick coriaceous, papillose at apex and along

margins, erect or slightly recurved; disc

glabrous or puberulous; stamens exserted;

filaments 1.0-3.0 mm long; anthers 1.5-3.5

mm long, + sagittate at base, erect, rarely

recurved; style with stigma 4.0-10.0 mm long,

exserted; stigma 1.5-2.0 mm long. Fruits

subglobose, broad ellipsoid or obovoid, slightly

compressed, 5.0-6.0 mm long x 3.0-8.0 mm

wide, black and glossy when ripe; pyrenes

ellipsoid, subovoid or hemispherical, usually

slightly rugose.

834

Austrobaileya 6 (4): 817-889 (2004)

Distribution and habitat : Pacific Islands,

including Hawaii and Mariana Islands, Fiji,

Vanuatu and New Caledonia, Papua New

Guinea and Australia; in a wide variety of

habitats including along beaches, along creeks,

on hillsides, ridges and sandstone, mostly in

dry rainforests.

Diagnostic attributes: Psydrax odorata is

readily distinguishable by its usually

exceedingly glossy, sometimes vernicose,

usually thick leaf blades which are mostly

provided with domatia, its usually ovate-

cuspidate, prominently keeled and lobed

stipules, its pedunculate, usually many-

flowered inflorescences with 4- or 5-merous

flowers, short and stout or long and slender

pedicels and flowers of branches of the cymes

secund and sometimes subsessile, and by its

fleshy corollas.

Notes: Psydrax odorata was found to be one of

the most variable and complex species of the

genus Psydrax in Australia. Within the material

broadly belonging to this species available for

study, quite distinct taxa are distinguishable.

One of these has keel-less, usually obtuse

stipules, whereas the others have stipules with

a distinct keel which is attenuated into a folded

lobe at apex.

Some of the past confusion within this

species started when Bentham (1867) combined

Canthium lamprophyllum F.Muell. under

Canthium odoratum (for which he used the

illegitimate name Canthium lucidum ) and cited

many collections including syntypes of

C. lamprophyllum F.Muell. (see under

C. lamprophyllum below).

Although Bentham considered all the

Australian specimens he called Canthium

lucidum to be ‘precisely’ those of the Pacific

Islands now know as Psydrax odorata , Merrill

& Perry (1945) and Bridson (1985) considered

the Australian specimens to be specifically

distinct from the Pacific species and referrable

to Psydrax lamprophylla. However, as indicated

previously, we agree these species are distinct

but consider that both occur in Australia.

The name Plectronia hookeriana was

provided in 1929 by Domin for the Australian

plants previously known as Canthium lucidum

Hook. & Arn. (of which C. lamprophyllum

F.Muell. was then considered to be a synonym).

This was because when Canthium Lam. was

transferred to the genus Plectronia L., the

combinations Plectronia lucida K.Schum. &

Krause (= Psydrax horizontalis (K.Schum. &

Thonn.) Bridson) and Plectronia lamprophylla

K.Schum. (= Psydrax micans (Bullock)

Bridson) already existed for taxonomically

different species from Africa thus inhibiting

transfer of Hooker and Arnott’s, and Mueller’s

epithets. The taxon Domin called Plectronia

hookeriana var. dietrichiae Domin, however,

belongs to Tarenna Gaertn. sensu lato, not

Psydrax.

Variability: Psydrax odorata varies greatly,

both in Australia and in the Pacific Islands, in

the shape, size, glossiness and texture of its

leaf blades, the hairiness of branchlets and

inflorescences, the size of its flowers, and in

its habitat of occurrence. Specimens from Fiji

and that from New Guinea examined for this

study resemble the type specimen from Vanuatu

and others from the latter country in their thin

coriaceous, shiny, elliptic leaf blades, their

regularly parallel lateral nerves, fine reticulate

venation, very open fragile inflorescences with

slender peduncles and pedicels, and their small,

4-merous flowers. Most of the collections of

this taxon from other Pacific Islands and some

from Vanuatu, particularly Flower s.n.(BM,

BRI, K, PVNH), are usually provided with thick

glossy leaf blades with either loose-flowered

slender inflorescences or dense-flowered

compacted inflorescences. Some of these

approach the Australian collections either in

attributes of their leaf blades or inflorescences,

whereas some, especially collections from

Hawaii, appear to be quite different from the

Australian ones and the type. However, the

variation observed in specimens from both

Australia and the Pacific region suggests we

are dealing with only local variants of one very

variable species. However, till now, with the

exception of C. odoratum var. tiniantense

(Kanehira) Fosberg from Micronesia, none of

this infraspecific variation has been formally

recognised.

Since none of the Australian collections

of this taxon match exactly the type of Psydrax

odorata , they are treated here as belonging to

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

segregate subspecies of this species. Within the

Australian material, three such subspecies are

distinguishable. They are the newly recognised

P. odorata subsp. arnhemica, a taxon from

northern Australia which is most like the

specimens from the Pacific regions in its leaf

blades, loose inflorescences and slender

pedicels, and P. odorata subsp. australiana, a

835

taxon from eastern Australia which differs from

the above by its compact glabrous

inflorescences, short thick pedicels and small

or medium-sized very glossy leaf blades, as well

as P. odorata subsp. buxifolia, based on

Canthium buxifolium Benth. These three

subspecies may be distinguished from the

typical subspecies as follows.

Key to Australian subspecies of Psydrax odorata and the extra-Australian

P. odorata subsp. odorata

1. Leaf blades 1.5-4.2 cm long x 0.6-2.0 cm wide; inflorescences 2-22

(-49)-flowered. 2

Leaf blades usually more than 4 cm long x 2 cm wide; inflorescences 12-93-flowered. 3

2. Young branchlets and inflorescence axes glabrous or subglabrous; domatia

present on the leaves; leaf blades thick coriaceous.... 6c. P. odorata subsp. australiana

Young branchlets and inflorescence axes hairy; domatia usually absent on

the leaves; leaf blades thick or thin coriaceous . 6d. P. odorata subsp. buxifolia

3. Young branchlets and inflorescence axes usually glabrous; inflorescences

usually compact, much-branched and densely flowered; flowers on

branches of the cyme subsessile or on short stout pedicels; stems usually

swollen (myrmecophilous). 6c. P. odorata subsp. australiana

Young branchlets and inflorescence axes usually hairy; inflorescences

usually open, laxly branched and flowered; flowers on branches of the

cyme on slender pedicels; stems not myrmecophilous. 4

4. Inflorescences (28-) 40-70-flowered, compact, much branched with flowers

densely arranged on branches; flowers mostly 4-merous; corolla

3.0-5.0 mm long; disc glabrous; leaf blades thin or thick coriaceous;

petioles 0.5-0.8 cm long. 6a. P. odorata subsp. odorata

Inflorescences 12-48-flowered, loose-branched and loose-flowered; flowers

mostly 5-merous; corolla 5.0-7.0 mm long; disc hairy; leaf blades usually

thick coriaceous; petioles 0.6-1.1 cm long. 6b. P. odorata subsp. arnhemica

6a. P. odorata subsp. odorata

Branchlets finely hairy or glabrous. Leaf blades

4.5-8.3 cm long x 2.5-4.0 cm wide, with lateral

nerves usually regularly parallel, subpatent or

oblique, and looping at margins; domatia

present or absent. Flowers 4.0-6.0 mm long,

usually chartaceous, 4-merous; pedicel of

solitary flower 2.5-10.0 mm long.

Representative specimens (typical form only):

New Guinea. Western Division: Mabaduan,

Apr 1936, Brass 6539 (BRI). Vanuatu.

Erromanga: Dillons Bay, Feb 1982, Cabalin

1455 (PVNH); ditto, Dec 1984, Sam 298

(PVNH). Efate: Ouen Tora Park, near the

PVNH herbarium, Sep-Oct 1995, Flower s.n.

(BRI). New Caledonia. Dothio River valley

near Thio, Mar 1983, McPherson 5577 (BRI).

Fiji. Macuata Coast, Jan 1924, Greenwood 555

(BRI).

Distribution and habitat : Papua New Guinea,

Vanuatu, New Caledonia and Fiji; chiefly

coastal, on beaches, ridges and flats, in vine

thickets.

Diagnostic attributes : Psydrax odorata subsp.

odorata is characterised by its finely hairy

branchlets and inflorescence axes, usually thin

coriaceous leaf blades, fragile inflorescences

with usually very slender peduncles and

836

Austrobaileya 6 (4): 817-889 (2004)

Fig. 3. Psydrax odorata subsp. arnhemica. A. fruiting habit x 0.8. B. flower x 6. C. LS of flower x 6. D. fruit x 3. E. LS of

fruit showing embryo x 3. F. pyrene x 3. A, Dunlop 7629 (CANB); B&C, Byrnes 2830 (DNA); D-F, Dunlop 3258 (BRI).

Psydrax odorata subsp. buxifolia. G. flowering branch x 0.8. H. flower x 6.1. fruit x 3. J. FS of fruit showing embryo x 3.

K. pyrene x 3. G from Febler 1978, p.530; H. McDonald 4131 & Williams (BRI); I-K, Forster PIF2492 et al. (BRI).

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

pedicels, and its comparatively small, 4-merous

flowers.

Notes: The Flower s.n. specimen cited above

has slightly thicker leaf blades and peduncles

and also more flowers in each inflorescence

than do the other collections cited above.

6b. P. odorata subsp. arnhemica S.T.Reynolds

& R.J.F.Hend., subsp. nov. a P. odorata

subsp. australiana inflorescentiis laxis,

paucifloris, ramis puberulentis, petiolis

pedicellisque longioribus et floribus

plerumque 5-meris differt. Typus:

Northern Territory. Gunn Point, 12°10’S,

131°05’E, 20 December 1989, J. Russell-

Smith 8171 Sc Lucas (holo: BRI; iso: DNA).

Young branchlets, stipules, petioles, young leaf

blades and inflorescence axes finely hairy or

sometimes glabrous. Petioles 0.6-1.1 cm long;

leaf blades (4.5-) 6.3-8.7 (-9.0) cm long x

(2.1-) 3.0-4.4 (-5.6) cm wide, with apex

subacute or shortly acuminate and base

subacute or abruptly obtuse and shortly

attenuate and decurrent into the petiole, drying

dark brown or red brown adaxially; lateral

nerves in 4-6 pairs, exceedingly slender, at an

angle of more than 45° to the midrib; domatia

small, usually present along the midrib

Inflorescences loosely branched, 12-48-

flowered; main peduncle (0.8-) 2.0-3.0 cm

long, the ultimate cymules 2-5-flowered.

Flowers usually 5-merous; pedicel of solitary

flowers 4.0-9.0 (-12.0) mm long, of the others

(1.0—) 2.5-4.5 mm long; corolla lobes elliptic

or lanceolate, 3.5-5.0 mm long, papillose at

apex and margins, finely reticulate veined.

Pyrenes depressed ovoid, usually rugose;

fruiting pedicels to 15.0 mm long. (Fig. 3A-3F)

Representative specimens: Western Australia. 4.5 km NW

of Cliffy Point on Middle Island, Bonaparte Archipelago

(14°19’S, 126°00’E), Jun 1987 ,Kenneally 10233 & Hyland

(PERTH); 6.7 km SSE of Walsh Point (14°37’S, 125°51’E),

Jun 1987, Kenneally 10340 & Hyland (PERTH); 9 km NNE

of Meeloyoo Hill (14°46 ’ S, 126°27’E), Jun 1987, Kenneally

10380 & Hyland (PERTH). Northern Territory, between

Groote Eylandt & mainland, Jan 1803, Brown s.n. (CANB);

Cannon Hill (H°58’S, 132°56’E), Dec 1972, Byrnes 2830

(DNA, K); ditto, Nov 1976, Dunlop 4300 (BRI, K); NE coast

of Cape Van Diemen, Melville Island, 11°10’S, 130°15’E,

May 1978, Webb & Tracey 12281 (BRI); 2 kmN of Nabarlek

airstrip, Apr 1979, Rankin 2106 (BRI, DNA, K); Waterfall

Creek, 13°28’S, 132°26’E, Nov 1980, Dunlop 5605 (BRI,

DNA); Nourlangie Rock, 12°52’S. 132°49’E, Dec 1983,

Russell-Smith 920 (BRI, DNA); Yirrkala, Shady Beach

837

(12°08’S, 136°55’E), Sep 1985, Wightman 2251 (DNA);

Groote Eylandt, Angyowmanja Creek (13°59’S, 136°41 ’E),

Jul 1987, Russell-Smith 2820 & Lucas (DNA); East Alligator

River (12°50’S, 133°22’E), Dec 1987, Dunlop 7629

(CANB); Gunn Point (12°10’S, 131°05’E), Dec 1989,

Russell-Smith 8171 & Lucas (DNA); ditto, Feb 1990, Leach

2697 & Dunlop (CANB, DNA); Wigram Island, English

Company Islands (ll 0 45’S, 136°37’E), Jul 1992, Leach

3065 (DNA); Craw Claw Island (12°42’S, 130°38’E), Dec

1993, Egan 2879 (DNA).

Distribution and habitat: Northern Australia,

including offshore islands, common in Arnhem

Land; usually on sandstone, sandstone

outcrops, lateritic cliff faces and gorges, in

coastal vine thickets, on sandy soil. (Map 4)

Diagnostic attributes: Psydrax odorata subsp.

arnhemica is characterised by laxly branched,

few-flowered inflorescences with long slender

peduncles and pedicels, and by its usually finely

hairy young branchlets, young leaves and

inflorescence axes. It may be distinguished from

P odorata subsp. australiana, which has more

or less similar leaves, by its open inflorescences,

larger flowers and comparatively long slender

pedicels.

Note: The inclusion of the collections from

Western Australia above is tentative because

the specimens concerned are incomplete.

Etymology: The subspecific epithet arnhemica,

Latin for ‘from Arnhem Land’ (in the Northern

Territory), refers to where this taxon is

common.

6c. P. odorata subsp. australiana S.T.Reynolds

& R.J.F.Hend., subsp. nov. a P. odorata

subsp. odorata inflorescentiis plerumque

glabris, pedunculis pedicellisque

brevioribus crassioribus, foliis lamina

crassiore adaxialiter pemitenti et floribus

corollis camosis non plerumque chartaceis

differt. Typus: Queensland. Wide Bay

District: Stony Creek, 4 km E of Didcot,

25°28’S, 151°51’E, 25 October 1993, PI.

Forster PIF14127 (holo: BRI; iso: A,

CANB, DNA, K, L, MEL, NSW, QRS).

Canthium odoratum subsp. (Didcot

RI.Forster PIF14127), PI. Forster & D.A.

Halford (2002, p.174).

Young branchlets, peduncles and pedicels

glabrous, rarely subglabrous. Petioles 0.25 - 0.6

cm long; leaf blades 3.5-8.3 (-9.3) cm long x

838

Austrobaileya 6 (4): 817-889 (2004)

Fig. 4. Psydrax odorata forma australiana. A. flowering habit x 0.6. B. flower x 6. A&B from Lebler 1978, p.527. Psydrax

odorata forma subnitida. C. part of inflorescence showing calyx and pedicel x 2. D. flower x 6. E. LS of flower x 6. F. fruit x 3.

G. pyrene x 3. C-E, Forster 14257 (BRI); F&G, Batianojf 900127 (BRI).

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

1.4-4.4 (-5.2) cm wide, with apex obtuse,

acute, subacute or shortly acuminate and base

subacute and attenuate into petiole, usually

glossy on adaxial surface, dull and opaque

below; lateral nerves in (1-) 2-4 pairs, slender,

suboblique or oblique; domatia 1-3 on each side

of midrib. Inflorescences with densely flowered

branches, (15-) 53-93-flowered; main peduncle

0.5-2.5 cm long. Flowers 4(or rarely 5)-merous;

pedicel of solitary flowers 3.0-7.0 mm long, of

the others 0.5-3.5 mm long; corolla lobes

sublanceolate, 3.5-5.0 mm long. Pyrenes broad

depressed ovoid, slightly rugose.

Diagnostic attributes: Psydrax odorata subsp.

australiana is readily recognisable by its usually

exceedingly adaxially glossy thick leaf blades,

glabrous branchlets, glabrous many-flowered

usually compact inflorescences, short thick

pedicels with flowers on the lateral branches

of the cyme often subsessile, and usually 4-

merous flowers with fleshy corollas. It differs

839

from the typical subspecies by its thicker leaves,

compact, many-flowered inflorescences,

subsessile flowers with short stout pedicels, and

by its distended myrmecophilous stems and

branches. The plants in Vanuatu belonging to

the typical subspecies are reported to have a

different growth form from the Australian ones

and are without insect galls (P. Flower, pers.

comm, to STR).

Etymology: The subspecific epithet australiana,

Latin for ‘pertaining to Australia’, refers to the

distribution of this subspecies.

Variability: Psydrax odorata subsp. australiana

varies considerably in attributes of its leaves.

On this basis, several forms or subforms were

distinguishable in the specimens of it available

for study. Though only three forms are formally

recognised here, they are sometimes connected

by intergrading specimens.

Key to forms of Psydrax odorata subsp. australiana

1. Leaves with very conspicuous domatia; leaf blades broad elliptic-ovate,

usually between 4.0-5. 2 cm wide. 6c(ii). P. odorata forma foveolata

Leaves with usually small, inconspicuous domatia; leaf blades elliptic,

elliptic-obovate, elliptic-ovate or subrhombic, 1.4-4.4 cm wide. 2

2. Leaf blades pale or dark green, glossy on the adaxial surface; reticulate

veins usually distinct on adaxial surface; domatia 1-3 on each side of

the midrib, prominent or obscure. 6c(i). P. odorata forma australiana

Leaf blades pale or yellowish green, slightly shiny or dull on the adaxial

surface; reticulate veins usually not apparent on the adaxial surface;

domatia 1 or 2 on each side of the midrib, usually small

.6c(iii). P. odorata forma subnitida

6c(i). P. odorata forma australiana

S.T.Reynolds & R.J.F.Hend., forma nov.

ab formis aliis P. odoratae subsp.

australianae foliis adaxialiter nitidis et

domatiis inconspicuis differt Type: As for

P. odorata subsp. australiana.

Young branchets and inflorescences glabrous,

rarely subglabrous; leaf blades elliptic, narrow

elliptic, elliptic-ovate, elliptic-oblong or slightly

rhombic; lateral nerves oblique or suboblique,

in (1-) 2-4 (-6) pairs with usually 2 or 3 pairs

more obvious than others and looping at

margins. Inflorescences (15-) 53-92-flowered; main

peduncles (5-) 15-25 mm long. (Fig. 4A, 4B)

Additional representative specimens : Queensland. Cook

District: Timber Reserve 14, Massey, 13°52’S, 143°23’E,

Nov 1980, Hyland 10874 (BRI); Bakers Blue Mountain,

Font Hills Station, 19 km S of Mt Carbine, 16°43’S,

143°10’E, Jan 1989, Fell 1591 (BRI); West Claudie River,

10.3 kmWNW of Lockhart River, 12°44’S, 143°15’E, Mar

1994, Fell 4145 (BRI). North Kennedy District: Barrabas

Scrub (20°10’S, 146°45’E), May 1972, Hyland 6076

(QRS)* * 1 2 ; Townsville Road, 14 km N of Bowen, 19°30’S,

147°57’E, Nov 1976, Sharpe 68 (BRI)* 2 ; Bucks Gully Scrub

(18°37’S, 142°08’E), May 1979, Hyland 9841 (QRS)* 2 ;

Jervoise Holding (18°34’S, 144°43’E), May 1979, Hyland

9937 (QRS)* 2 ; 40 Mile Scrub National Park, 1.6 km north

of Mt Surprise road junction on Kennedy Highway, 18°07’S,

144°49’E, Nov 1983, Stocker 1806 (BRI); ditto, Mar 1987,

Clarkson 6886 & McDonald (BRI)* 2 ; ditto, Feb 1990,

Hyland 13989 (BRI); Whitsunday Island, 20°16’S,

148°56’E, Nov 1985, Batianoff 3020 & Dalliston (BRI);

840

Austrobaileya 6 (4): 817-889 (2004)

Mt Stuart, 9 km S of Townsville, 19°21’S, 146°47’E, Dec

1991, Bean 3866 (BRI); Port Denison, date unknown,

Fitzalan s.n. [MEL 1538534 & MEL1538536](MEL). South

Kennedy District: southern end of Lake Elphinstone,

21°33’S, 148°13’E, Nov 1987, Champion 322 (BRI);

Hazelwood Gorge, 21°15’S, 148°27’E, 13 km SSW of

Eungella, Dec 1992, Bean 5269 (BRI). Leichhardt District:

Carnarvon Development Road, 80kmNof Injune, 25°05’S,

148°15’E, Nov 1988, Schefe CMW1147 (BRI); near

Theodore, 24°55’S, 150°05’E, Dec 1989, Phillips s.n. (BRI).

Port Curtis District: Mt Etna, 23°10’S, 150°25’E,Oct 1976,

Hyland 9104 (BRI); ditto, Nov 1987, Vavryn 26 (BRI).

Burnett District: about 20 miles (32 km) NNW of Monto,

in 1983, Barlett s.n. (BRI); 1.5 km along Archookoora Road

from Kingaroy-Cooyar road, about 18 km SSE of Kingaroy,

Dec 1986, Beesley 975 & Ollerenshaw (CANB);

Coominglah State Forest 28,24°53’S, 151°00’E,Apr 1990,

Forster PIF6702 (BRI). Wide Bay District: CoongaraRock,

18 kmS of Biggenden, 25°35’S, 152°05’E,Dec 1977, Young

8 (BRI); Coast Range, 9 km E of N of Goomeri, 26°07’S,

152°06’E, Oct 1990, Pedley 5579 (BRI, NSW). Moreton

District: Samford Range, Jan 1930, Meebold & White s.n.

(BRI); Whiteside, North Pine River near Brisbane, 27° 13’ S,

152°57’E, Jan 1932, Blake 3196 (BRI, K); Bangaree

Enviromental Park, 2 km ENE of Crows Nest, 27°16’S,

152°04’E, Apr 1993, Halford 1664, Thomas & Thompson

(BRI); 10.5 km NW of Ballandean, 28°45’S, 151°45’E,Dec

1994, Halford 2361 (BRI); 10kmSEofPittsworth,27°46’S,

151°42 ‘E, Feb 1995, Halford 2394a (BRI), 2394b (BRI).

New South Wales. Richmond River, in 1876, Fawcett

[NSW193681](NSW); Lismore, Nov 1894, Bauerlen s.n.

(NSW); Kerrabia via Rylstone, Nov 1895, Dawson s.n.

(NSW); 9 miles (c. 14.4 km) E of Aberdeen, Hunter Valley

(32°09’S, 151°03’E),Aug 1959, Storey 6605 (CANB). (For

specimens above marked * 2 , see 40-Mile Scrub form below.)

Distribution and habitat: Eastern Australia,

from Cape York Peninsula, Queensland, to

Hunter River, New South Wales; along creeks

and rivers, steep hillsides, stony ridges, rocky

outcrops, rocky water courses, usually in dry

rainforests, also on beach dunes, on sandy

loamy, clay or stony soils. (Map 4)

Note : The use of the epithet ‘ australiana ’ in

the name of this form follows Recommendation

26A.3. in the current International Code of

Botanical Nomenclature (Greuter et al., 2000).

Variability : The leaf blades of Psydrax odorata

forma australiana are variable and several ill-

defined subforms exist. Specimens from along

creek and river banks and usually from dry

rainforests are typical of this form. They have

very shiny, mostly medium-sized, elliptic,

ovate-elliptic, subrhombic to subobovate leaf

blades with prominent oblique nerves and

reticulate venation, and large many-flowered

inflorescences. Specimens from coastal areas,

especially those from off-shore islands, usually

have smaller or thicker leaves which are usually

only slightly shiny on the adaxial surface. These

are not unlike those of P. odorata forma

subnitida except that they are larger and broader

(broad elliptic or obovate) than what is allowed

for in that form. Specimens from steep ridges

and hillsides tend to have narrower, thicker and

often more shiny leaf blades than those on other

specimens. Moreover, some of the collections

from north Queensland approach P. odorata

subsp. arnhemica in their leaves, subglabrous

or sparsely hairy branchlets and inflorescence

axes, slender peduncles and pedicels but differ

from that in their general aspect, compact

inflorescences with greater numbers of flowers,

shorter pedicels and smaller flowers. On the

other hand, collections from south-eastern

Queensland, which possess small, elliptic or

narrow elliptic, thick coriaceous leaf blades and

small inflorescences resemble some of the

collections filed under P. odorata subsp.

buxifolia, but that taxon differs from them by

its pubescent branchlets and peduncles, and

usually efoveolate leaf blades. However, these

subspecies ( P. odorata subsp. australiana and

P. odorata subsp. buxifolia ) are sometimes

connected by specimens with hairy branchlets

and inflorescences, and foveolate leaf blades.

40-Mile Scrub form: Collections from deciduous

vine thickets in the 40 Mile Scrub (as indicated

by * 2 in the list above) probably represent a

distinct form or subform but they contain only

fruiting material. Flowering material is

necessary to be certain of this. However, these

specimens differ from the other collections in

their narrow infructescences and narrow, thick

shiny leaves. These leaves usually have narrow

elliptic blades with a subacute apex and base

(or an acute base which is decurrent into the

short petiole), which are dark green on the

adaxial surface and pale green below. The

lateral nerves are slender and obliqueand in 2

or 3 pairs. The reticulate veins are not apparent.

One or two small domatia occur on each side

of the midrib, or somet im es domatia are absent.

The infructescences are 2.5-6.5 cm x c.3.0 cm,

branched only towards the apex with the main

peduncles usually to 4.5 cm long. Fruits are

black when ripe. (Map 1)

6c(ii). P. odorata forma foveolata S.T.Reynolds

& R.J.F.Hend., forma nov. P. odoratae

formae australianae primo adspectu

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

maxime similis, sed foliis latioribus et

domatiis valde conspicuis differt. Typus:

Queensland. Cook District: Aeroglen,

16°55’S, 145°45’E, November 1978, B.

Jago 221 (holo: QRS).

Leaf blades broad elliptic-ovate or elliptic, thick

coriaceous, drying dark brown and glossy

adaxially, pale brown abaxially; lateral nerves

in 4 or 5 pairs, obliquely arched and looping at

margins of blade; domatia usually conspicuous,

few on each side of the midrib. Peduncles and

pedicels glabrous or subglabrous. Flowers

mostly 5-merous; pedicels slender; corolla lobes

slender.

Additional representative specimens : Queensland. Cook

District: Endeavour River, in 1882, Persieh [MEL

1533874](MEL); Hartleys Creek (16°39’S, 145°34’E), Apr

1946, Flecker s.n. (QRS); ditto, Dec 1987, Sankowsky 791

& Sankowsky (BRI); 2 km from Cooktown along Mclvor

River Road (15°29’S, 145°14’E), Feb 1983, Telford 9398

& Butler (CANB); State Forest Reserve 607, Parish of Cairns,

Shoteel Logging Area, 16°55’S, 145°36’E,Feb 1985, Gray

3909 (BRI); Lizard Island, Cook’s Lookout, 14°47’S,

145°28’E, Oct 1988, Batianoff 10274 (BRI); Round

Mountain, Embly Range, 13°33’S, 143°30’E, Jun 1992,

Forster PIF10495 & Tucker (BRI); Jane Table Hill, Lakefield

National Park, 46.6 km N of Lakefield homestead, 14°30’S,

144°07’E, Mar 1993, Fell 2925 & Stanton (BRI). North

Kennedy District: Round Mountain, SSW of Townsville,

19°27’S, 146°42’E, Jan 1996, Cumming 14040 (BRI).

Distribution and habitat : North Queensland,

common around Cooktown; on open slopes,

rocky creek banks and frontal dunes, in semi-

deciduous vine thickets, woodlands and

grasslands on sandy loamy soil. (Map 3)

Diagnostic attributes: This form resembles

P. odotata forma australiana in its leaves and

inflorescences, but differs by its very prominent

domatia and usually broader leaves.

Etymology: The forma epithet foveolata, Latin

for ‘minutely pitted’, alludes to the very

conspicuous domatia on the leaves of this form.

6c(iii). P. odorata forma subnitida

S.T.Reynolds & R.J.F.Hend., forma nov.

a P. odorata forma australiana foliis

brevioribus crassioribusque, folii laminae

paginis leviter nitentibus, nervis

lateralibus indistinctis venatione

reticulata obscurapraeditis differt. Typus:

Queensland. Darling Downs District:

near Texas on the road to Stanthorpe,

841

28°51’S, 151°14’E, 7 November 1993,

PI. Forster PIF14257 (holo: BRI).

Canthium oleifolium var. pedunculatum

Maiden & Baker, Proc. Linn. Soc. N.S.W.

9: 460 (1894). Type: New South Wales.

Palistan, 30 miles (c.48 km) NW of

Condobolin, Feb. 1892, Clements s.n.

(syn: NSW [NSW155950]); ditto, Dec

1893, Clements s.n. (syn: NSW

[NSW155949]).

Canthium odoratum forma (Texas P.I.Forster

PIF14257), S.T. Reynolds (1997, p.180),

PI. Forster & D.A. Halford (2002, p. 174).

Leaves with petioles 0.2-0.5 cm long; blades

elliptic or subobovate, 3.5-6.5 cm long x

1.4-2.6 cm wide, with apex obtuse and base

narrowed and attenuate into the petiole, matt

green or pale green or slightly yellowish-green,

thick and stiff especially when dry; lateral

nerves in 1 or 2(or 3) pairs; domatia small, 1

or 2 on each side of midrib. Inflorescences 20-36-

flowered; main peduncles 0.4-1.2 cm long.

(Fig. 4C-4G)

Additional representative specimens : Queensland.

L eichhardt District: 24 miles (c.38.4 km) E of Taroom, Jul

1963, Lazarides 6943 (BRI, K, NSW); Mt Britton Mine,

Homevale Station, Nebo, Dec 1973, Webb & Tracey 13725

(CANB); Bull Creek Gorge, Castlevale Station, about 13 km

SW of homestead, 24°33’S, 146°45’E, Sep 1990 , McDonald

4664 & Bean (BRI); 7 km NE of Taroom, 25°33’S,

149°55’E, Nov 1996, Halford Q3008 & Dowling (BRI).

Warrego District: W boundary of Chesterton Range

National Park, N of Morven, 26°10’S, 147°16’E, Sep 1995,

Bean 8987 & Grimshaw (BRI). Maranoa District: 4 km W

of Euloral, 27°08’S, 148°44’E, May 1982, Neldner &

Thomas 783 (BRI). Darling Downs District: Kindon

Station, 54 miles (c.86 km) NNE of Goodiwindi, 28°05’S,

150°45’E,Dec 1938, Smith 593 (BRI); ‘TheRanch’, 7 miles

(c. 11.2 km) N of Tara, Aug 1946, Everist 2671 (BRI). New

South Wales. Gunnedah, Oct 1886, Betche [NSW193693]

(NSW); Lachlan River near Lake Cudgellico, Jun 1891,

Maiden s.n. (NSW); Murrumbo Creek, Goulbum River, Sep

1895, Baker [NSW193707] (NSW); Condobolin, Euabolong

Road, Aug 1897, Maiden [NSW 193705] (NSW)* 3 ;

Gunnedah, Oct 1899, Forsyth [NSW193691] (NSW); Terry

Hie Hie District, May 1914, Julius s.n. (NSW); Gungal, Nov

1914, Boorman s.n. (K, NSW [NSW 193700]; Ticketty

Wells, W of Wallangara, Jul 1917, Julius s.n. (NSW);

Warialda, May 1932, Vickery s.n. (NSW); about 2 km S of

Warialda, Dec 1963, Williams 28 (NSW); Waa Gorge, Mt

Kaputar National Park, 68 km NE of Narrabri, Nov 1976,

Coveny 8989 & Roy (BRI, K); 14 km from Narrabri on road

to Mt Kaputar, 30°19’S, 149°54’E, Dec 1994, Forster

PIF 15934 (BRI).

842

Austrobaileya 6 (4): 817-889 (2004)

Distribution and habitat : Central and south

western Queensland and western New South

Wales; usually in dry scrubs, on sandy soil

amongst rocks, on slopes, ridges and hill sides.

(Map 5)

Diagnostic attributes: This form is characterised

by its thick, pale green or yellowish green,

subobovate or elliptic usually small leaf blades

which are slightly shiny on the adaxial surface

and dull and opaque on the abaxial surface,

which have 1 or 2 pairs of obscure lateral nerves

and 1 or 2 domatia on each side of the midrib.

This form can be confused with P. oleifolia

which also has dull leaf blades with indistinct

nerves and short small inflorescences, and the

plants also often grow in the same general area.

However, P. oleifolia differs from this by its

much duller, concolorous, usually efoveolate

leaf blades, usually smaller flowers and fruits

and has very slender pedicels.

Notes: Canthium oleifolium var. pedunculatum

Maiden & Baker is included as a synonym here

because its type, collected from the Condobolin

area of New South Wales, belong to the same

taxon as that of P odorata forma subnitida.

Maiden also collected specimens of this taxon

from the Condobolin area (see specimen

marked * 3 in the above list).

Etymology: The forma epithet subnitida, from

Latin sub-, ‘somewhat’, and nitidus, ‘shiny’,

refers to the slightly shiny adaxial surface of

the leaf blades of this form.

6d. P. odorata subsp. buxifolia (Benth.)

S.T.Reynolds, comb. & stat. nov.;

Canthium buxifolium Benth., FI. Austral.

3: 422 (1867). Type: Queensland. Ad

fluvium Burnett [et] Dawson, Mueller

[MEL153843] (syn: MEL); Burnett

River, Mueller s.n. (syn: K). ? New South

Wales, exact locality unknown, Leichhardt

s.n. (syn: MEL [MEL1538144]; ? isosyn:

NSW [NSW193767]).

Branchlets, young leaves, petioles, peduncles,

pedicels and calyx lobes minutely hairy.

Stipules usually very conspicuous with a long

(to 9.0 mm long), broad, more or less foliaceous,

folded lobe at apex. Petioles 0.1-0.3 cm long;

leaf blades elliptic, ovate or suborbicular,

1.5-4.2 (-5.0) cm long x (0.6-) 1.4-2.0 (-2.5)

cm wide, with apex and base obtuse or subacute

or with apex slightly rounded and both adaxial

and abaxial surfaces glabrous or finely hairy

towards the base, thin or thick coriaceous, the

adaxial surface dark or pale green, drying dark

brown but usually paler brown at the margin;

lateral nerves in 1-3 pairs, exceedingly slender,

indistinct; reticulate veins not apparent;

domatia very rarely present Inflorescences

(2* 4 -) 14-22 (-49)-flowered; peduncles

pubescent, the main one (3-)8-20 mm long;

branches 1.0-3.5 mm long; pedicels pubescent,

slender in solitary flowers, 3.5-7.0 mm long,

stout in others and nearly obsolete, 0.5-2.0 mm

long; calyx lobes sparsely hairy distally;

corolla 3.0-5.0 mm long; tube 0.5-1.25 mm

long; lobes elliptic, 2.5-3.5 mm long x 0.75-1.0

mm wide; filaments of stamens 1.0-1.5 mm

long; anthers erect, 1.5-2.0 mm long; style

(with stigma) 4.0-5.2 mm long. Fruits broad

ellipsoid or obovoid but retuse at apex, 5.0-7.0

mm long x 3.0-8.0 mm wide; pyrenes

depressed ovoid, slightly rugose. (Fig. 3G-3K)

Diagnostic attributes: Psydrax odorata subsp.

buxifolia is readily recognisable by its

comparatively small, very glossy leaves, hairy

branchlets and small inflorescences. It

resembles P. odorata subsp. australiana,

especially the small-leaved form, in its leaves,

inflorescences, 4-merous flowers on short stout

pedicels and fleshy corollas. However, that

subspecies differs from P. odorata subsp.

buxifolia by its foveolate leaves, usually

glabrous branchlets and inflorescences.

Moreover, the leaves of that variety are usually

borne at right angles to the main axis of the

branchlet whereas in P. odorata subsp. buxifolia

the leaves are borne in the same plane as the

main axis of the branchlet. These subspecies,

however, are connected by intergrading

specimens, ones with thicker leaves with or

without domatia and with hairy branchlets and

inflorescences.

Variability: The leaves and inflorescences are

variable in this subspecies; based on these

attributes, two forms are recognised here.

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

843

Key to forms of P. odorata subsp. buxifolia

1. Leaves and inflorescences comparatively small; blades narrow elliptic,

1.5-2.0 mm long x 0.6-0.9 cm wide, thick; inflorescences 2-5-

flowered* 4 ; main peduncles 0.3-0.5 cm long . .. 6d(ii). P. odorata forma parviflorifra

Leaves and inflorescence comparatively larger; blades narrow elliptic or

suborbicular, 2.5-5.0 cm long x 0.7-2.5 cm wide, elliptic, thick or thin;

inflorescences 14-22 (-49)-flowered; main peduncles 0.8-2.0 cm long

. 6d(i). P. odorata forma buxifolia

Note: The character marked * 4 in the

description and key above is uncertain because

only the type material has been seen and the

specimens concerned have only very old

inflorescences with fruit on them.

6d(i). P. odorata forma buxifolia (Benth.)

S.T.Reynolds & R.J.F.Hend., comb. &

stat. nov.; Canthium buxifolium Benth,

FI Austral. 3: 422 (1867). Type: as for

P. odorata subsp. buxifolia above.

Most of the description of P. odorata subsp.

buxifolia provided above applies to this form.

Its leaves are variable though its broad elliptic,

broad ovate or suborbicular, usually thin,

efoveolate leaf blades, which are usually borne

in a one plane along the branchlets, are typical

of this form. However, specimens of it with

thicker, often broader or larger leaf blades

approach those of P. odorata subsp.

australiana, but differ from the latter by their

usually efoveolate leaves, and hairy branchlets

and inflorescences.

Additional representative specimens: Queensland. Burnett

District: Mt Wooroolin near Kingaroy, 26°35’S, 151°45’E,

Apr 1947, Michael 3002 (BRI); Munro’s Camp, 26°—’S,

151°—’E, Dec 1954, Smith 6283 (BRI, K); Bunya

Mountains State Forest (State Forest 151 Tureen), Maidenwell

road, about 3.5 km NNE of Munro’s Camp, 26°52’S,

151°38’E, Dec 1987, McDonald 4131 & Williams (BRI); N

side of Bunya Mountains, 26°55’S, 151°35’E, Mar 1988,

Sterling 1 (BRI). Darling Downs District: 2.5 km SSW of

Gladfield, 28°06’S, 152°10’E, Jun 1986, Forster PIF2492,

Bird & Grimshaw (BRI); MacKinnemy’s Irvingdale-Moola

road, 27°10’S, 151 0 31’E, Jan 1993, Smith 34 (BRI); 5.6 km

along Linthorpe road, N of Pittsworth, 27°39’S, 151°39’E,

Feb 1996, Bean 9947 (BRI). Moreton District: Rosewood,

May 1913, White s.n. (BRI); Hoya, Dec 1935, Michael 2267

(BRI); ditto, Mar 1936, Michael 2268 (BRI); Mt French,

Jan 1936, Smith s.n. (BRI); Brisbane, Upper Brookfield,

27°25’S, 152°55’E, Feb 1978, Jessup 57 (BRI)* 5 ; N of

Boonah, Dec 1981, Bird s.n. (BRI); Indooroopilly, 27°31’S,

153°00’E, Nov 1982, Wood [AQ339614] (BRI); on Boonah-

Ipswich road, 7 km N of Boonah, 28°00’S, 152°41’E, Feb

1989, Williams s.n. (BRI). New South Wales: Acacia Creek

via Killamey, Dec 1905, Dunn [NSW193789] (NSW)* 5 ;

River Tree area, c.39 miles (62.4 km) E of Fiston, c.25 m il es

(40 km) NW of Tabulam, Jul 1949, Clark, Pickard & Coveny

1780 (NSW); Sandy Hollow, Aug 1963, Burgess s.n.

(CANB).

Distribution and habitat : South-eastern

Queensland and northern New South Wales;

usually in semi-evergreen vine thickets on

ridges and river banks. (Map 6)

Notes: The collections marked * 5 in the above

list have thick foveolate leaf blades and hairy

inflorescences, and appear to be intergrades

between P. odorata subsp. buxifolia and P. odorata

subsp. australiana.

The use of the epithet buxifolia in the

name of this form follows Recommendation

26A.3. in the current International Code of

Botanical Nomenclature (Greuter et al., 2000).

6d(ii). P. odorata forma parviflorifra*

S.T.Reynolds & R.J.F.Hend., forma nov.

a P. odorata forma buxifolia foliorum

laminis minoribus, inflorescentiis

minoribus paucifloris differt. Typus.

Queensland. Moreton District: 4.5 km

WSW of Mt Alford, 28°05’S, 152°23’E,

27 April 1986, L.H. Bird [AQ408941]

(holo: BRI; iso: BRI).

Canthium sp. (Mt Alford L.H.Bird

AQ408941), S.T. Reynolds (1997, p.180).

Canthium buxifolium var. (Mt Alford

L.H.Bird AQ408941), P.I. Forster & D.A.

Halford (2002, p.174).

Stipules ovate; leaf blades elliptic or narrow

elliptic, 1.5-2.0 cm long x 0.6-0.9 cm wide,

with apex obtuse and base subacute and

narrowed into petiole, glossy on the adaxial

surface; lateral nerves slender, obscure; domatia

absent. Inflorescences (remnant ones only seen)

* should be parviflorifera ' (PDB 2005)

844

Austrobaileya 6 (4): 817-889 (2004)

2-5-flowered; pedicels slender, those of solitary

flowers c. 3.5 mm long, of others c.2.0 mm long.

Fruits obovoid, 5.0-5.5 mm long x 3.0-5.0 mm

wide, 2-lobed, bluish black; pyrenes rugose.

Distribution and habitat : Known only from the

type collection; in Araucarian microphyll vine

forest. (Map 8)

Etymology: The forma epithet parviflorifra,

from Latin parvus, ‘small’, and florifer,

‘flowerbearing’, refers to the small

inflorescences in this form.

6d(iii). Mundubbera form Canthium

buxifolium forma (Brigooda P.I.Forster

PIF5657), P.I. Forster & D.A. Halford

(2002, p.174).

The collections of P. odorata from around

Mundubbera, southeastern Queensland, listed

below probably represent a distinct form of P. odorata

subsp. buxifolia but as the specimens are

without well-developed flowers, it is not

possible to be certain of this.

Queensland. Burnett District: Monogorilby, Mundubbera

Shire, 26°01’S, 15F00’E, Dec 1981, Forster 213B (BRI);

2.5 km E of Brigooda, 26° 15 ’S, 151°26’E,Aug 1989 , Forster

PIF5657 (BRI); Fontainea Scrub, State Forest 172, Gurgeena

Plateau, 25°26’S, 151°23’E, Mar 1994, Forster PIF15068

(BRI).

The taxon represented by these specimens may

be described as having: Branchlets greyish

coloured mottled with white; young branchlets,

petioles, peduncles and pedicels finely

pubescent. Stipules comparatively small, keeled

and attenuated into an acuminate lobe at apex;

petioles 0.15-0.3 cm long; leaf blades narrow

elliptic or subobovate elliptic, 2.5-4.5 (-5.0)

cm long x 0.9-1.5 (-2.5) cm wide, with apex

subacute or obtuse and base cuneate,

exceedingly glossy on adaxial surface, drying

dark brown with usually paler coloured margins

adaxially, pale brown or pale olive-green

abaxially, thick coriaceous; lateral nerves in 3

pairs, obliquely arching, looping at margins;

reticulate veins not apparent; domatia (1-) 2-4

on each side of midrib, prominent.

Inflorescences 1.3-1.8 cm long x 1.0-2.0 cm

wide, 11-19-flowered; main peduncle 4.0-10.0

mm long; flowers 4-merous; pedicel of solitary

flowers 2.0-2.5 mm long, of others 0.5-1.0 mm

long; calyx 1.0-1.5 mm long; corolla 4.5-6.0

mm long, corolla tube 2.0-2.5 mm long,

sparsely hairy at throat; corolla lobes subacute

at apex, 2.5-3.5 mm long; filaments c.1.0 mm

long; anthers c.2.5 mm long. Fruits broad

ellipsoid, c.6.0 mm long x 4.0 mm wide, black

when ripe; pyrenes rugose.

Distribution and habitat: Occurring around

Mundubbera, southeastern Queensland; in

semi-evergreen vine thickets in hilly rocky

country. (Map 1)

Diagnostic attributes: The above specimens

resemble P odorata forma buxifolia in their

small very shiny leaves, dark brown adaxial

surface of dried leaves (with paler margins),

and small inflorescences, but differ from that

form as follows:

Domatia present on the leaves, usually 2-4 on each side of the midrib

. P. odorata ‘Mundubbera form’

Domatia rarely present on the leaves but when so, only one on each side of

the midrib. P. odorata forma buxifolia

The above specimens also resemble the small¬

leaved form of P. odorata forma australiana,

but that form differs from them by its glabrous

branchlets and inflorescences, less shiny and

pale brown adaxial surface of dried leaves, and

fewer domatia (only 1 or 2 domatia on each

side of the midrib).

7. Psydrax lamprophylla (F.Muell.) Bridson,

Kew Bull. 40: 724 (1985); Canthium

lamprophyllum F.Muell, Fragm. 2: 133

(1861). Type: Queensland. Moreton

District: near Brisbane River, in 1855,

F. Mueller s.n. (lecto, here designated:

MEL [MEL1538219]; isolecto: K, MEL

[MEL1538225]).

Canthium sp. L, Ross, E.M. in T.D. Stanley

& E.M. Ross, FI. south-eastern

Queensland 2: 343 (1986).

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

[Canthium lucidum auct. non Hook. & Arn.:

G. Bentham, FI. Austral. 3: 421 (1867

‘1866’) quoad speciminibus Brisbane

River, Moreton Bay, [leg.] F. Mueller et

Tweed River, [leg.] C. Moore].

Trees 5-20 m high; trunk to c.30 cm dbh; bark

dark brown, smooth or closely longitudinally

fissured; young parts and inflorescence axes

with short spreading hairs; branchlets

quadrangular towards their apices, lenticellate,

glabrous. Leaves opposite; stipules lacking a

keel and an apical lobe, c.3.0 mm long x 2.0

mm wide, ± membranous, usually connate to

above the middle, with broad ovate, obtuse

lobes, or stipules slightly truncate and very

rarely provided with a minute, obscurely keeled

lobe, hairy outside, with colletors present at

base adaxially; petioles (0.4-) 1.0-3.0 cm long,

usually flexuose, deeply channelled above;

blades elliptic, broad elliptic, elliptic-oblong,

ovate-elliptic or ovate-oblong, (8.5-) 10.5-18.0

(-20.0) cm long x (3.1-) 4.0-8.7 (-9.7) cm

wide, with apex and base acuminate or

subacute, or apex abruptly shortly acuminate

or obtuse and base obtuse, slightly rounded,

subcordate or truncate and shortly decurrent

into the petiole, coriaceous; both surfaces

glabrous, the adaxial surface dark green, glossy,

drying brown or with a yellowish tinge,

sometimes dotted with insect galls, the abaxial

surface pale or dull green; lateral nerves in 4-7

pairs, oblique or subpatent, the lower pairs ±

acutely angled to the midrib and ascending

distally; reticulate venation loosely arranged,

fine, distinct or indistinct; midrib and nerves

usually raised in dried specimens; domatia

present or absent. Inflorescences robust,

2.5-8.0 cm long x 2.5-8.5 cm wide, 130-145-

flowered; peduncles pubescent or puberulent,

basal one 1.5-3.5 cm long, thickened towards

the middle or base, provided with minute

denticulate bracts usually about a third of its

length from the base, usually terminated by 3

branches; branches usually thickened, bifurcate,

0.5-2.5 cm long, terminated by 23-45-flowered

cymes. Flowers 4(or 5)-merous; pedicel of

solitary flowers (1.5-) 4.0-6.0 (-9.0) mm long,

of others 0.5-3.0 mm long; calyx c.1.0 mm

long, minutely patent hairy, with tube cupular

and limb 4(or 5)-denticulate; lobes ovate,

minute; corolla 3.5-6.5 mm long with tube

narrow or broad campanulate, 1.5-3.5 mm long

845

x 1.5-3.0 mm wide, with sparse or dense

reflexed hairs at the throat and puberulent

abaxially; lobes elliptic or lanceolate, 2.0-3.5

mm long x c.1.0 mm wide, thick, papillose,

obtuse and reflexed at apex, sparsely puberulous

abaxially especially distally; disc usually

sparsely minutely hairy; stamens included;

filaments erect, about 1 mm long; anthers erect

or ± recurved, 1.5-2.0 mm long, slightly

papillose; style (with stigma) 5.0-10.0 mm

long; stigmatic knob oblongoid, 2-lobed at

apex. Fruits broad ellipsoid, divaricately

2-lobed, 0 5-1.0 cm long x 0.7-1.2 cm wide,

usually with a red glandular layer under

pericarp; pyrenes ellipsoid, usually exceedingly

rugose.

Diagnostic attributes : Psydrax lamprophylla

is readily distinguishable from other species of

Psydrax in Australia by its slightly membranous

stipules which are neither keeled nor lobed at

the apex. It resembles Psydrax cymigera

(Valeton) S.T.Reynolds & R.J.F.Hend. from

New Guinea in its large leaves and

inflorescences, but the stipules in that species

are keeled and attenuated into a folded lobe at

the apex.

Notes: The taxonomy and nomenclature of

Psydrax lamprophylla and P. odorata have been

confused since Bentham (1867) treated the

species concerned as conspecific and used the

name Canthium lucidum Hook. & Arn. for his

taxon. He did not appreciate that that name is

illegitimate, being a later homonym of

Canthium lucidum R.Br., and was possibly

unaware that Canthium odoratum (Forst.f.)

Seem, should have been used for his species

when treating it as belonging to Canthium.

Mueller cited several syntypes in his

protologue for Canthium lamprophyllum

F.Muell. Some of these, i.e. those from

Rockhampton collected by Thozet (MEL), from

between the Dawson and Burnett Rivers

collected by Mueller (MEL) and from Port

Molle and from Port Denison collected by

Fitzalan (BM, MEL), have keeled stipules

which are attenuated into a folded lobe at the

apex, slender glabrous inflorescences and small

to medium-sized glossy leaf blades. These

resemble specimens of Psydrax odorata from

the Pacific Islands and are therefore referred

here to that species. The remainder of the

846

syntypes, i.e. Brisbane River and Pine River

collected by Hill and Mueller, Brisbane River

collected by Mueller (MEL, K), and Wide Bay

collected by C. Moore (MEL), have stipules

which are neither keeled nor lobed at the apex,

robust hairy inflorescences and comparatively

large leaf blades.

Thus, we believe two discrete taxa are

represented by the syntypes of Mueller’s

Canthium lamprophyllum, namely (1) a taxon

with stipules which are keeled and lobed at the

apex, slender glabrous inflorescences and small

or medium-sized shiny leaf blades, which is

referrable to Psydrax odorata, and (2), a taxon

with stipules which are without a keel and an

apical lobe, robust hairy inflorescences and

large leaf blades.

Austrobaileya 6 (4): 817-889 (2004)

To fix the applicability of P. lamprophylla

(F.Muell.) Bridson, Mueller’s name is

lectotypified here by the Mueller collection from

near the Brisbane River, i.e., specimen

MEL1538219 in MEL. An isolectotype is

present in K and also in MEL (MEL1538225).

The lectotype, which is in flower, is the best of the

syntypes of Mueller’s Canthium lamprophyllum

seen in this study. It also has the hairy

inflorescences described in Mueller’s

protologue (which is not the case in the syntypes

referrable to Psydrax odorata which have

glabrous inflorescences). Thus lectotypified,

Mueller’s name applies to the second taxon

represented by the above syntypes which is, we

believe, specifically distinct from P. odorata.

Variability : Psydrax lamprophylla varies

consistently in its leaves and fruits. On this

basis, two forms are formally recognised here.

Key to forms of Psydrax lamprophylla

1. Petioles 1.0-3.0 cm long; leaf blades usually narrow at base and attenuate

into the petiole; lateral nerves usually oblique; reticulate venation

prominent; corolla 3.5-4.5 mm long; fruits 5.0-5.5 mm long x (4.0-)

7.0-7.5 mm wide; pyrenes usually slightly rugose

. 7a. P. lamprophylla forma lamprophylla

Petioles 0.4-1.8 mm long; leaf blades usually broad at base, with the base

obtuse or subcordate; lateral nerves widely spaced, patent or suboblique;

reticulate veins usually obscure; corolla 4.5-6.5 mm long; fruits 7.0-

8.0 mm long x (0.8-) 1.0-1.2 cm wide; pyrenes usually exceedingly

rugose . 7b. P. lamprophylla forma latissima

7a. P. lamprophylla forma lamprophylla

Stipules truncate; leaf blades (8.5-) 10.5-13.5

cm long x (3.1-) 4.0-5.2 cm wide, usually

narrowed at both ends, attenuate at base into a

petiole up to 3.0 cm long, or the base obtuse;

surfaces usually dotted with insect galls; lateral

nerves usually exceedingly oblique, with

prominent cross veins and reticulate veins

between them; main peduncles 1.3-3.5 cm

long; corolla tube 1.5-2.0 mm long x c.1.5 mm

wide; fruits didymous, 5.0-5.5 mm long x

7.0-7.5 mm wide, or rarely l-lobedthenc.5.0mm

long x 4.0 mm wide, smooth; pyrenes mgose.

Additional representative specimens’. Queensland. Port

Curtis District: Bulburin State Forest, 24°3-’S, 151°2-’E,

Apr 1980, McDonald 3162, Fisher & Ryan (BRI); Mount

Atherton, 22°46’S, 150°45’E, Dec 1992, Melzer 63 (BRI);

Brolga Mine, GlenGeddes, 23°01’S, 150°18’E, May 1993,

Batianoff 930539, Specht & Reeves (BRI); State Forest 69,

24°38’S, 150°53’E, 33.9 km NW of Monto, Mar 1996,

Halford Q2808 (BRI). Burnett District: Mill North Logging

Area, State Forest 95,3 km N of Kalpowar, Aug 1984,24°4-

’S, 151°1-’E, Grant s.n. (BRI). Wide Bay District: Kin Kin,

26°16’S, 152°53’E, Jan 1916, White s.n. (BRI); Dundowran,

Jul 1928, Tryon s.n. (BRI); Fraser Island, 25°15’S, 153°20’E,

May 1964, Webb & Tracey 6345 (BRI); Pine Mountain, S of

Biggenden, 25°—’S, 152°—’E, Mar 1980, Young & Randall

342 (BRI); Vicinity of Fairlies Knob, 25 km E of Biggenden,

25°30’S, 152°18’E, Jul 1981, Young & Randall 392 (BRI);

Kenilworth Bluff about 8 km N of Kenilworth, 26°32’S,

152°43’E, May 1987, Sharpe 4683 & Bean (BRI); 1 km N

of Mt Mittarula, SW of Gympie, 26°18’S, 152°32’E, Oct

1993, Bean 6720 (BRI). Moreton District: Moreton Bay,

in 1872, Eaves s.n. (MEL); Benarkin, Apr 1924,

Cameron s.n. (BRI); Indooroopilly, Brisbane, Apr 1936,

Cribb s.n. (BRI); Yarraman, Aug 1957, McGillivray

[NSW193674] (NSW); Palen Creek State Forest near Mt

Lindesay, 28°20’S, 152°45’E, Nov 1993, Grimshaw 102

(BRI). New South Wales. Ballina, Jul 1893, Bauerlen s.n.

(NSW); ditto, Feb 1894, Bauerlen s.n. (NSW, K); Brunswick

River, Pacific Highway, May 1964, Schodde 3552 & Hayes

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

(CANB, NSW); 3 km S of southern edge of Broken Head,

Feb 1981, Williamss.n. (NSW); Tweed River, date unknown,

Moore s.n. (MEL [MEL1538240], K, NSW).

Distribution and habitat: Central and south¬

eastern Queensland to far northern New South

Wales; usually in dry rain forests on hill crests,

slopes, gullies and along creeks, on red brown

soil. (Map 7)

Notes: The specimens from Bulburin State

Forest and Dundowran have broad leaves with

a broad base, but short petioles as in P.

lamprophylla forma latissima. However, they

differ from that in their aspect and venation.

Specimens which are sterile appear to represent

young growth of this form.

7b. P. lamprophylla forma latissima

S.T.Reynolds & R.J.F.Hend., forma nov.

a P. lamprophylla forma lamprophylla

foliorum laminis latis basi lata praeditis,

petiolis brevioribus (12.0-18.0 mm

longis) inflorescentiis robustioribus et

pyrenis valde rugosis differt. Typus:

Queensland. North Kennedy District:

about 7 km W of Paluma, 19°0-’S, 146°0-

’E, 20 July 1986, G. Duff 18 (holo: BRI).

[Plectronia diococca auct. non (Gaertn.)

F.Muell.: F. Mueller, Fragm. 9: 185-186

(1875) quoad specimini Rockingham

Bay, [leg.] Dallachy].

[Canthium cymosum auct. non Pers.: F.

Mueller, Fragm. 9: 185 (1875), Syst.

census Austral. PI. 75 (1882) & Second

syst. census Austral. PI. 126 (1889)]

[Canthium didymum auct. non C.F.Gaertn.:

F. Mueller, Fragm. 9: 185 (1875) et F.M.

Bailey, Qld FI 3: 764 (1900) quoad

specimini Rockingham Bay, [leg.]

Dallachy .]

Stipules truncate or very rarely with a vestigial

apical lobe; petioles (0.4-) 1.2-1.8 cm long;

leaf blades (9.5-) 12.0-18.0 (-20.0) cm long x

(5.1-) 6.0-8.7 (-9.7) cm wide, with apex

abruptly short acuminate, subobtuse or obtuse,

and usually truncate, subcordate, slightly

rounded or obtuse and shortly attenuate into

petiole proximally; lateral veins widely spaced

with reticulate veins obscure; domatia 1 or 2

on each side of midrib, sometimes absent; main

847

peduncles 1.5-3.0 cm long, hairy, robust, with

minute denticulate bracts towards the middle

or towards the base, thickened from the bracts

to its base; inflorescences branches bifurcate,

8.0-12.0 mm long; cymes about 23-flowered;

pedicels finely hairy, those of solitary flowers

3.0-6.0 mm long, of others 1.5-3.0 mm long;

calyx finely puberulent; corolla tube broad

campanulate, 2.5-3.5 mm long x 2.5-3.0 mm

wide, puberulent abaxially, densely reflexed

hairy at throat; lobes somewhat elliptic,

2.0-3.0 mm long, obtuse; fruits 2-lobed,

transversely ellipsoid, slightly compressed,

7.0-8.0 mm long x 8.0-12.0 mm wide;

endocarp slightly resinous, usually with a red

glandular layer below the pericarp; pyrenes

strongly rugose.

Additional representative specimens'. Queensland. Cook

District: Leo Creek, upper Nesbitt River, 13°45’S, 143°25’E,

Aug 1948, Brass 19899 (BRI, K); Upper Parrot Creek, Annan

River, 15°45’S, 145°15’E, Sep 1948, Brass 20299 (BRI, K);

Big Tableland Logging Area, about 27 km S by E of

Cooktown, 15°45’S, 145°17’E, Sep 1960, Smith 11206 (BRI,

K); Lankelly Creek(13°55’S, 143°20’E), Sep 1911, Hyland

5412 (BRI, K, QRS); State Lorest Reserve 144, Windsor

Tableland, I6°30’S, 145°05’E, Oct 1971, Hyland 5558 (BRI,

K, QRS); State Lorest Reserve 607, Bridle Logging Area,

17°00’S, 145°35’E, near Mareeba, Apr, 1972, Irvine 184

(BRI, K, QRS); Leo Creek Road, 13°40’S, 143°20’E, Jul

1972, Hyland 6365 (BRI, QRS); Timber Reserve 146,

Tableland Logging Area, 15°45’S, 145°15’E, nearRossville,

N of MtLinnegan, Jul 1975, Hyland 8330 (BRI, QRS); State

Lorest Reserve 144,16°17’S, 145°,05’E, WofDaintree,Apr

1976, Hyland 8732 (BRI, CANB, K, QRS); Moa Island,

Torres Strait, 10°U’S, 142°15’E, May 1987, Budworth 1264

(BRI). North Kennedy District; Gregory Creek, near

Proserpine, Dec 1993, Perry s.n. (BRI); Portello’s Crossing,

Dryander Creek, base of Mt Dryander, 20°16’S, 148°35’E,

Leb 1994, Forster PIP 14858 (BRI); Rockingham Bay, date

unknown, Dallachy s.n. (MEL)* 6 . South Kennedy District:

Pinch Hatton Gorge, Eungella National Park, 21°09’S,

148°38’E, Jan 1991, Pearson 432 (BRI).

Distribution and habitat: Northern

Queensland, from Torres Strait, Cape York

Peninsula to near Eungella; on steep slopes and

gullies, near permanent water at altitudes of

400-1100 m, in wet or dry rainforests, usually

with Agathis robusta, on soils derived from

granite. (Map 7)

Diagnostic attributes: Psydrax lamprophylla

forma latissima may be distinguished from the

typical form by the broad base of its leaf laminas

which are very broad obtuse or truncate, widely

spaced lateral nerves with obscure reticulate

veins, robust peduncles, larger flowers and

fruits with very rugose pyrenes.

848

Austrobaileya 6 (4): 817-889 (2004)

Notes: Dallachy’s collection from Rockingham

Bay cited above (* 6 ) was included under

Plectronia diococca (Gaertn.) F.Muell. by

Mueller (1875), when making the new

combination, and under Canthium didymum

C.F.Gaertn. (as ‘Roxb.’) by Bailey (i900).

These names are now considered synonyms and

the species they relate to does not occur in this

region (Bridson 1985, p.687).

Etymology: The forma epithet latissima, Latin

for ‘very broad’, refers to the broad leaf blades

with a broad base present in this form of the

species.

8. Psydrax laxiflorens S.T.Reynolds &

R. J.F.Hend., sp. nov. quoad folia P.

lamprophyllae (F.Muell.) Bridson simile

sed stipulis carinatis inflorescentiis laxis,

ramis paucioribus constructis, floribus

fructibusque majoribus, pedicellis

longioribus differt. Typus: Queensland.

Cook District: Herberton Range,

17°30’S, 145°30’E, 19 November 1929,

S. F. Kajewski 1377 (holo: BRI; iso: K,

MEL, NSW).

Canthium sp. (Herberton Range S.F.Kajewski

1377), S.T. Reynolds (1997, p.180), P.I.

Forster & D.A. Halford (2002, p.174).

Trees 10-20 m high; trunk to c.25 cm dbh,

fluted and buttressed; bark flakey; branchlets

glabrous, usually with ± globose or irregular

shaped lenticels. Leaves opposite; stipules

broad ovate, 2.5-4.0 mm long, abruptly

acuminate with a long folded lobe at apex;

petioles 0.6-1.5 cm long, flexuose, slightly

channelled adaxially; blades elliptic or elliptic-

ovate, (6.0-) 8.0-9.0 (-10.5) cm long x (3.0-)

4.0-6.0 (-6.3) cm wide, with apex and base

obtuse or with apex abruptly shortly acuminate

and base subacute or subtruncate, glabrous, thin

coriaceous, shiny adaxially; lateral nerves in

3-5 pairs, very slender, arcuate, looping

distally; reticulate venation very fine, open;

domatia conspicuous, 1 or 2 on each side of

the midrib, very rarely absent. Inflorescences

open, very laxly branched and flowered,

2.5-4.0 cm long x 3.5-4.5 cm across, 6-20-

flowered; peduncles glabrous, the basal one

3.0-16.0 mm long with small bracts near its

distal end; axis branches 8.0-14.0 mm long,

bifurcate, each terminated by a 3-7-flowered

cymule, the solitary flower at forks of the

dichotomous branches often absent. Flowers

5-merous, strongly scented; pedicel of solitary

flowers 7.0-10.0 mm long, of others 5.0-7.0

mm long; calyx 2.0-2.5 mm long, glabrous,

with tube ± turbinate and limb 5-denticulate;

lobes ovate; corolla 6.0-8.0 mm long, cream-

yellow or yellow; tube broad campanulate,

2.0-3.0 mm long, densely hairy at the throat;

lobes elliptic, 4.0-5.0 mm long x c.1.5 mm

across, obtuse and ± incurved at apex, thick,

papillose at margins, longer than tube,

recurved; stamens shorter than tube plus lobes;

filaments erect, c.1.5 mm long; anthers erect

or reflexed, c.3.0 mm long, apiculate; style

(with stigma) 8.0-9.0 mm long; stigma oblong,

2-lobed at apex. Fruits obovoid, 13.0-15.0 mm

long x 8.0-10.0 mm across; pyrenes ellipsoid,

exceedingly rugose.

Additional representative specimens : Queensland. Cook

District: Davies Creek, Lamb Range, between Mareeba and

Kuranda, 16°55’S, 145°35’E, in 1962, Webb & Tracey 7377

(BRI); State Forest Reserve 194, about 6 km WSW of

Atherton, 17°17’S, 145°26’E,Nov 1974, Hyland7S6S (BRI,

K, QRS); ditto, Mar 1976, Moriarty 2022 (BRI, QRS); ditto,

Dec 1978, Sanderson 1601 (BRI, QRS); State Forest Reserve

607, Emerald Logging Area, SE of Mareeba, 17°05’S,

145°35’E, Apr 1979, Gray 1370 (BRI, NSW, QRS); Bartle

Frere, Boonjie Logging Area, 17°23’S, 145°45’E,Nov 1991,

Hyland 14312 (QRS). North Kennedy District: Paluma

village, near police station, 19°00’S, 146°12’E, Dec 1984,

Jackes 5 (BRI).

Distribution and habitat: Northern Queensland

from the Atherton Tableland to Paluma Range;

mostly on ranges in mountainous country, in

vine forests at altitudes of 720-1220 m.

(Map 5)

Diagnostic attributes: Psydrax laxiflorens is

readily distinguishable by its comparatively

large obovoid fruits, very loosely branched,

sparsely-flowered inflorescences with slender

branches and 2-7-flowered cymules with 5-

merous flowers that are densely hairy at the

mouth of the corolla tube, and prominent

domatia on its leaf blades. In respect to leaves,

it is similar to P. lamprophylla (F.Muell.)

Bridson but differs from that with its keeled

stipules, lax inflorescences with fewer branches

and bigger flowers and fruits on longer pedicels.

It is closely related to P. odorata, under which

t hi s taxon was previously included, but P. odorata ,

especially P odorata subsp. australiana which

occurs in the same area, differs from P. laxiflorens

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

in its compact inflorescences, ellipsoid fruits

5.0-6.0 mm long x 3.0-7.0 mm across, and

usually 4-merous flowers which are only

sparsely hairy at the mouth of the corolla tube.

Etymology: The specific epithet laxiflorens,

from Latin laxus, ‘loose’, and florens,

‘flowering’, refers to the laxly branched, open

inflorescences in this species.

9. Psydrax tropica S.T.Reynolds &

R.J.F.Hend., sp. nov. P. lamprophyllae

(F.Muell.) Bridson simile sed stipulis

acuminatis carinatis, petiolis brevioribus,

foliorum lamina + membranaceiore

nervis et venis reticulatis pertenuibus

praedita differt. Typus: Queensland.

Cook District: State Forest Reserve 310,

Upper Goldsborough Logging Area, 17

km SE of Little Mulgrave Township,

17°16’S, 145°47’E, November 1988, L.W.

Jessup 1819, G.P. Guymer & W.J.

McDonald (holo: BRI; iso: BRI).

Canthium sp. (Whitfield Range B.RHyland

1020); S.T. Reynolds (1997, p.181), P.I.

Forster & D.A. Halford (2002, p.174).

Small trees to 10 m high; branchlets brown or

reddish brown, quadrangular distally and

usually resinous, glabrous. Leaves opposite;

stipules conspicuous, broad ovate, 6.5-8.0 mm

long, keeled, attenuated into a long folded lobe

at apex, paler coloured and ± membranous at

margins, those distal on branchlets resinous;

petioles 0.3-0.7 (-0.9) cm long, slightly

flattened, flexuose; blades elliptic, ovate-elliptic

or elliptic-oblong, 10.0-14.5 (-18.0) cm long

x 5.2-7.0 (-8.0) cm wide, abruptly narrowed

at both ends, or the apex abruptly shortly

acuminate (more or less caudate in young

leaves) or subacute and the base obtuse or

subacute and attenuate into short petiole;

margins thick and slightly recurved; both

adaxial and abaxial surfaces glabrous with the

adaxial surface glossy, with prominent nerves

and reticulate secondary venationed; lateral

nerves in 4-8 (-9) pairs, obliquely arched and

ascending and with a fine, delicate network of

secondary veins between them, which is

prominently raised in dried specimens; domatia

present or very rarely absent. Inflorescences

open with bifurcate branches, 4.0-5.0 cm long

x 5.0-6.5 cm wide, (13-) 30-75-flowered;

849

peduncles and pedicels sparsely short hairy,

with main peduncles 1.5-2.3 cm long and

secondary peduncles 4.0-11.0 mm long,

bifurcate. Flowers 5-merous; pedicels slender,

that of solitary flowers 2.5-5.0 mm long, of

others 0.5-2.0 mm long; calyx 1.0-2.0 mm

long, glabrous, with tube ± campanulate and

5-denticulate limb wider and paler coloured

than tube, the lobes broad ovate; corolla 6.5-8.5

mm long; tube broad cylindrical, 1.0-3.5 mm

long, to 1.5 mm wide at mouth, densely long

hairy at throat; lobes erect or slightly spreading,

elliptic, 2.5-5.0 mm long x 1.5-2.0 mm wide,

obtuse but cucullate at apex, usually delicately

reticulate veined; stamens usually as long as

the corolla lobes; filaments erect, about 2.0 mm

long, filiform; anthers 3.0-3.5 mm long, erect

or sometimes ± recurved; style (with stigma)

about 10.0 mm long; stigma oblong, 2-lobed

at apex. Fruits transversely ellipsoid, about 8.0

mm long x 9.0 mm wide; pericarp verrucose

when dry; pyrenes about 8.0 mm long x 5.0

mm wide, exceedingly hard and rugose.

Representative specimens : Queensland. Cook District:

Jo hns tone River, Mar 1916, Michael s.n. (BRI); The Boulders

near Babinda, Jan 1967, Rijkers [NQNC14772] (BRI, QRS);

Whitfield Range, Oct 1967, Hyland 1020 (QRS); Alexandra

Creek, N of Thornton Peak, Jul 1973, Webb & Tracey 13220

(BRI, QRS); foot of MacAlister Range, 2.5 km ENE of Saddle

Mountain, 16°49’S, 145°41’E, Dec 1987, Lyons 54 (BRI);

State Forest Reserve 310, Upper Goldsborough Logging Area,

17 km SEofLittle Mulgrave Township, 17°16’S, 145°47’E,

Nov 1988, Jessup 1819, Guymer & McDonald (BRI).

Distribution and habitat: North Queensland,

from Thornton Peak to the Atherton Tableland;

in vineforests, on gentle slopes, creek and river

banks, usually on alluvium. (Map 9)

Diagnostic attributes: Psydrax tropica is

characterised by its usually large elliptic

acuminate or subacute leaf blades with

prominent, very fine, delicately arranged

network of secondary veins, and by its ovate,

keeled stipules with usually paler coloured

thinner margins. It is related to P. odorata

subsp. australiana, which has more or less

si mil ar attributes of the leaves and inflorescences,

but that taxon differs from P. tropica by its usually

smaller leaves which are up to 8.3 cm long and

4.4 cm wide, rarely more, and which have an

obtuse or subacute apex and loosely arranged

reticulate venation.

850

Austrobaileya 6 (4): 817-889 (2004)

Etymology: The specific epithet tropicus , Latin

for ‘tropical’, refers to the usual habitat of this

species, namely the Wet Tropics of northern

Queensland.

10. Psydrax banksii S.T.Reynolds &

R.J.F.Hend., sp. nov. P. suborbiculari

(C.T.White) S.T.Reynolds & R.J.F.Hend.

arte simile sed foliorum lamina obovata

ellipticave, basi anguste attenuata,

venatione reticulata prominente, et

inflorescentiarum pedunculis longioribus

differt. Typus: Queensland. Cook

District: Mouth of Jardine River,

10°56’S, 142°14’E, 2 February 1983, A.

Morton AM 1782 & L. Hiddins (holo:

BRI; iso: MEL).

Canthium sp. (Friday Island L.J.Brass

18158), S.T. Reynolds (1997, p.180), P.I.

Forster & D.A. Halford (2002, p.174).

Shrubs or small trees to 5 m high; branchlets

+ angular and usually puberulous distally,

lenticellate. Leaves opposite; stipules ovate or

± triangular, 4.0-6.0 mm long, keeled and

attenuated into a long folded lobe distally;

petioles (0.1-) 0.2.-0.4 cm long; blades obovate

or subelliptic, 3.2-5.5 (-6.2) cm long x

1.6-3.0 (-3.6) cm wide, with apex rounded and

base subacute or attenuate and decurrent into

petiole, the margins flat or slightly recurved in

dried specimens; adaxial and abaxial surfaces

glabrous, the adaxial ones shiny, the abaxial

ones dull, usually resin-dotted; lateral nerves

in (2-) 4 or 5 pairs, oblique, ascending distally

and looping at margins; reticulate veins loosely

arranged, distinct or indistinct on adaxial

surface; domatia small, usually few on each side

of the midrib, sometimes absent.

Inflorescences 6.0-9.0 cm long x 4.0-5.0 cm

across, 27-59-flowered; peduncles slender,

puberulous, the basal one (1.5-) 3.0-4.6 cm

long; axis branches 7.0-14.0 mm long; cymes

12-15-flowered. Flowers 4(or 5)-merous;

pedicels slender, puberulent, that of solitary

flowers 4.0-8.0 mm long, that of others

1.0-4.0 mm long; calyx about 2.0 mm long;

tube slightly turbinate, limb 4- or 5-denticulate;

lobes broad ovate, minute; corolla 6.0-7.5 mm

long; tube cylindrical or narrow campanulate,

2.0-2.5 mm long xc.1.0 across, sparsely hairy

at throat; lobes recurved, sublanceolate,

subacute or obtuse, 4.0-5.5 mm long x

1.0-1.25 mm across; stamens with filaments

subulate, 1.0-2.5 mm long, erect; anthers

2.5-3.0 mm long; style (with stigma) 6.0-7.0

mm long, exserted; stigma oblong, shortly

bilobed at apex. Fruits obovoid to ellipsoid,

0.9-1.0 cm long x 0.7-0.9 cm across, usually

resin-dotted; endocarp thick, resinous;

pyrenes ellipsoid or depressed ovoid, rugose.

(Fig. 5F-5K)

Additional representative specimens : Queensland. Cook

District: Lizard Island, Aug 1770, Banks & Solander s.n.

(BM); Torres Strait, Friday Island, 10°36’S, 142°10’E, Feb

1948, Brass 18158 (BRI); between Cape York and

Galloway’s Hill, Oct 1965, Smith 12540 (BRI); Olive River,

12°10’S, 143°05’E, Sep 1974, Hyland 1441A (BRI); Starke-

Cape Flattery Road, Jul 1976, Tracey 14579 (BRI); Morgan

River, 15°06’S, 145°14’E, Jan 1979, Duke 1209 (BRI);

Torres Strait, Badu Island, Jan 1980, Garnett 344 & 356

(BRI); coast S of Thorpe Point, Shelburne Holdings, 11°55’S,

143°08’E, Nov 1985, Gunness 1961 (BRI); Pom 2V, Parish

of Annan, 15°36’S, 145°19’E, Dec 1988, Hyland 13773

(BRI); 0.8 km N of Captain Billy Landing, 11°37’S,

142°51’E, Mar 1992, Clarkson 9269 & Neldner (BRI);

Lakefield National Park, 32.5 km from Laura on road to New

Laura Homestead, 15°18’S, 144°25’E, Jan 1993, Forster

PIF12 819 & Bean (BRI); Evans B ay, 26 km NE of B amaga,

10°42’S, 142°32’E, Feb 1994, Fell 3917, Stanton & Roberts

(BRI); Cape York, date unknown, Hill s.n. (K). North

Kennedy District: Hinchinbrook Island, 18°1-’S, 146°2-’E,

Dec 1978, Thorsboume & Travers 492 (BRI, K).

Distribution and habitat : North Queensland,

from Torres Strait Islands and Cape York

Peninsula to Hinchinbrook Island; in coastal

woodlands, heaths and vine thickets, mostly on

sand dunes or sand hills. (Map 8)

Diagnostic attributes: Psydrax banksii is

characterised by its usually small, shiny,

obovate, finely reticulate-nerved leaf blades,

hairy, long-stalked inflorescences, and delicate

flowers with a narrow, ± cylindrical corolla

tube.

Affinities: Psydrax banksii is closely related

to P. suborbicularis (C.T.White) S.T.Reynolds

& R.J.F.Hend. from Papua New Guinea with

which it shares the slender inflorescences and

corollas, but that species differs from P. banksii

in its broad suborbicular leaf blades as indicated

in the key to distingish the two below.

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax 851

Leaf blades suborbicular with base truncate or subcordate and reticulate

veins indistinct; inflorescences with main peduncle 2.0-2.5 cm long;

corolla with tube 3.0-3.5 mm long, densely hairy at throat, and lobes

2.0-2.5 mm long. P. suborbicularis

Leaf blades obovate with base narrowed and reticulate veins prominent;

inflorescences with main peduncles (1.5-) 3.0-4.6 cm long; corolla with

tube 2.0-2.5 mm long, sparsely hairy at throat, and lobes 4.0-5.5 mm

long. P. banksii

Notes: Though the majority of specimens of

this species seen had the typical small obovate

leaf blades, a few had somewhat larger leaves

which approached those of P odorata, but that

species, especially in P. odorata subsp.

australiana, which occurs in the same area as

P. banksii , differs from P. banksii by its glabrous

branchlets and inflorescence axes, short and

stout pedicels, and fleshy corollas with a

comparatively broad tube.

Etymology: The specific epithet banksii

honours Sir Joseph Banks for his valuable

contribution to Australian botany. He, with

Daniel Solander, was the first (according to our

records) to collect this species when visiting

Lizard Island, off the north Queensland coast,

in August 1770 during Captain Cook’s voyage

in the Endeavour.

11. Psydrax latifolia (F.Muell. ex Benth.)

S.T.Reynolds & R.J.F.Hend., comb, nov.;

Canthium latifolium F.Muell. ex Benth.,

FI. Austral. 3: 421 (1867). Type: New

South Wales. In the interior towards

Barrier Range, date unknown, Nielsen

s.n. (syn : MEF [MEF1538208]); NW

interior, date unknown, J. McDouall

Stuart s.n. (syn: n.v.).

Shrubs or small trees 2-5 m high with

branches usually borne at right angles to the

main stem; bark whitish to light or dark grey,

usually finely fissured; branchlets reddish-

brown and usually covered with a white bloom

distally, sometimes swollen and galled

(myrmecophilous); young branchlets and young

leaves usually with short spreading hairs.

Leaves opposite; stipules ± triangular, 5.0-7.0

mm long, keeled and attenuated into a long,

apiculate, keeled lobe distally; petioles (0.2-)

0.5-1.2 cm long, stout, flattened, sometimes

slightly winged by decurrent leaf blade; blades

broad ovate or suborbicular, elliptic-ovate or

elliptic, (3.6-) 5.0-7.0 (-8.5) cm long x (3.0-)

4.2-6.0 (-8.7) cm wide, with apex rounded,

obtuse or truncate, base subcordate, obtuse or

abruptly obtuse and attenuate into the petiole,

and margins entire or slightly wavy (in young

leaves), glabrous, or slightly scabrid on adaxial

surface, coriaceous, usually exceedingly so and

rigid, with the raised nerves and dense

reticulate venation rendering the blade very stiff

when dry, pale or dull green, drying yellowish-

green, yellowish-brown or brown; lateral nerves

in (4-) 5-9 pairs, conspicuous, obliquely arched

and ascending, usually looping at margins;

secondary veins closely reticulate, usually

conspicuous on both surfaces. Inflorescences

shorter than leaves, 2.2-4.5 cm long x 2.5-4.5

cm wide, usually shortly stalked and

trichotomously branched; peduncles glabrous

or sparsely hairy, the basal one 3.0-12.0 mm

long, with minute bracts towards its apex and

terminated by 3 branched cymes; branches

(3.0-) 9.0-25.0 mm long; flowering branches

3.0-6.0 mm long; cymules 14-41-flowered.

Flowers 4(or 5)-merous; pedicel of solitary

flowers 2.5-6.0 mm long, of others 1.0-3.0 mm

long; calyx 0.5-2.0 mm long, with tube ±

turbinate, 0.25-0.75 mm long, and limb thin,

± scarious, 4(or 5)-toothed; lobes ovate, to 0.5

mm long, glabrous or with tufts of hairs at the

apex; corolla 5.5-8.0 mm long, with tube

shorter than the lobes, ± campanulate, 1.5-3.0

mm long, sparsely hairy at throat; lobes

subelliptic, 3.5-4.5 mm long, obtuse,

occasionally papillose outside, erect or

recurved; disc shorter than the calyx limb,

glabrous; stamens nearly as long as the corolla

lobes; filaments 0.5-2.0 mm long; anthers

2.0-3.0 mm long, usually erect; style (with

stigma) 7.0-10.0 mm long, slightly exserted

from corolla tube; stigmatic knob oblongoid,

2-lobed at apex. Fruits obovoid or broad

ellipsoid, slightly laterally compressed, 0.5-0.9

cm long x 0.6-1.2 cm wide, black and shiny

when ripe; pyrenes ellipsoid, very deeply rugose

on the dorsal side. (Fig. 5A-5E)

852

Austrobaileya 6 (4): 817-889 (2004)

Fig. 5. Psydrax latifolia. A. flowering branch x 0.6. B. flower x 6. C. LS of flower x 6. D. fruit x 3. E. pyrene x 4. A-C,

Nelson 1818 (DNA); D&E, Willis [MEL 1538609] (MEL). Psydrax banksii. F. fruiting branch x 0.6. G. flower x 6. H. LS of

flower x 6.1. fruit x 3. J. LS of fruit showing embryo x 4. K. pyrene x 4. F,I&K, Clarkson 9649 & Neldner (BRI); G&H,

Morton AM1782 & Hiddins (BRI); J, Brass 18158 (BRI).

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

Additional representative specimens: Western Australia.

6 mi les (c.9.6 km) E of Willenom Gorge, Apr 1952, Broadway

s.n. (PERTH); 65 miles (c.104 km) E of Leonora (28°5-’S,

120°2-’E), May 1959, Vollprecht 16 (PERTH); 3 miles (c.4.8

km) S of Meekatharra, Oct 1965, Blockey 12 (PERTH);

Dale’s Gorge between Millstream and Willenom (21°—’S,

117°—’E), Nov 1968, Young 159 (CANB); Stony Creek, 1

mile (c.1.6 km) S of Meekatharra (26°36’S, 118°29’E), Aug

1976, Demarz 6088 (PERTH); Doolgunna Homestead

(25°01’S, 119°13’E), Dec 1983, Mitchell 1179 (PERTH);

16 km N of Two Sisters about 130 km SE of Shay Gap (21 0 —

’S, 121°—’E), Jul 1984, Newbery 10378 (PERTH).

Northern Territory. 8 miles (c.12.8 km) E of Haast’s Bluff,

MacDonnell Ranges, May 1911, Hill 176 (MEL); Idracowra

(25°00’S, 133°47’E), May-Sep 1920, Basedow s.n. (AD)* 7 ;

MacDonald Downs (22°25’S, 135°07’E), Aug 1933, Ising

s.n. (AD)* 7 ; 5 miles (c.8 km) NNE of Barrow Creek

Township, Aug 1956, Lazarides 5820 (AD, BRI, DNA, K,

PERTH); Reedy Creek, George Gill Range, May 1962,

Madden 10372 (DNA); Bagot Springs, George Gill Range,

Jul 1966, Willis s.n. (MEL); 60 miles (c.96 km) NW of Alice

Springs, just outside Desert Block (23°00’S, 133° 13’E), Jan

1967, Latz 34 (DNA); Stuart Highway, 5 miles (c.8 km) S of

Tea Tree (22°12’S, 133°26’E), Feb 1968, Maconochie 573

(DNA); No 1 Desert Bore, Amburla Station (23°20’S,

133°10’E), Jan 1969, Nelson 1818 (DNA, K); Docker River,

Nov 1970, Woenne 108 (PERTH); Marque Station (22°58’S,

137°40’E), May 1972, Dunlop 2597 (DNA); Ormiston

Gorge (23°37’S, 132°43’E), Feb 1973, Griffin 346 (DNA);

New Crown Station, Beddome Range (25°52’S, 134°28’E),

Apr 1977, Henshall 1525 (CANB); Yuendumu, 5 km N of

Ngana Outstaion (22°25’S, 131°30’E),Feb 1981 , Henshall

3332 (DNA); The Granites Tenements, Tanami Desert

(20°32’S, 130°18’E), Dec 1984, Kalotas 1771 (DNA).

South Australia: Lake Eyre Basin, Alberga River, 20 miles

(c.32 km) W of Todmorden, Apr 1950, Black s.n. (AD);

Tom kin son Range, Mt Davies area(26°13’S,129 0 16’E),Jan

1956, South Australia Pastoral Board (AD). Queensland.

Gregory North District: Ardmore, 25 miles (c.40 km) W of

Dajarra, 21°39’S, 139°ll’E,Nov 1947, Everist 3218 (BRI);

about 1 km W of Lark Quarry, 23°03’S, 142°20’E, Oct 1989,

White s.n. (BRI); about 29 km N of Pathungra, 22°07’S,

140°34’E, May 1993, Gunness 2155 (BRI); 40 km E of

Bedourie, date unknown, Pulsford 598 (BRI). Mitchell

District: 17 km NW of Withywene Homestead, NW of

Stonehenge, 23°44’S, 143°19’E, Apr 1986, Neldner 2345

& Stanley (BRI). Gregory South District: between

Nockatunga and New South Wales border, Mar 1924, Allison

s.n. (AD, BRI); Momey Station, 25°22’S, 141°28’E, Aug

1973, Trapnell & Williams 223 (BRI); Nockatunga, 27°43’S,

142°43’E, Apr 1977, Hughes s.n.(AD)* 7 ; 50 km N of

Orientos homestead, 24°4-’S, 141°1-’E, Oct 1980, Bolton

116 (BRI). Warrego District: Wittenburra Station, 36 miles

(c.57.6 km) S of Eulo, 28°35’S, 144°45’E, Jan 1937, Everist

& Smith 50 (BRI); Dynevor Downs, 28°05’S, 144°21 ’E, Apr

1941, White 11821 (BRI); about30kmNofThargomindah,

Jun 1955, Smith 6348 (BRI); Grey Range, 79 km W of

Thargomindah, 27°35’S, 143°15’E, Oct 1980, Bolton

MPB64 &Ashwell (BRI). New South Wales. Main Barrier

Range, Mootwingee, about 110 km NE of Broken Hill

(31°17’S, 142° 18’E), Aug 1962 Jackson 390 (AD)* 7 ; Peery

Hills, Wilcannia (31°34’S, 143°2-’E), Sep 1976,

Cunningham 4824 & Milthorpe (NSW); Brindingabbe via

Bourke (29°00’S, 144°54’E), Oct 1983, O’Neill s.n. (NSW).

853

Distribution and habitat : Central Australia; in

low woodlands with Eucalyptus spp. and/or

Mulga (Acacia aneura F.Muell. ex Benth.:

Mimosaceae) on a variety of soils but mostly

in red sandy soil, on sandy plains, ridges and

slopes. (Map 9)

Diagnostic attributes: Psydrax latifolia is

readily recognisable by its very stiff, broad

ovate, suborbicular or elliptic leaf blades with

very conspicuous nerves and reticulate veins,

its comparatively small inflorescences, its

usually 4-merous flowers, and by its hairy or

glabrous, reddish brown usually robust young

branchlets (which mostly turn pale to dark grey

with age).

Affinities: Psydrax latifolia is closely related

to P. ammophila, with which it shares the

branchlet colour and attributes of the

inflorescences and flowers, but that species

differs from P. latifolia in its usually narrow

bases of its leaf blades which have a finer

reticulate venation, and the presence of a small

domatium on some leaves of the branchlet.

These species, however, are connected by

integrades so that more field collections and

notes are necessary before the total variability

of each of these species can be fully understood.

P. latifolia also intergrades into P. attenuata

and P. oleifolia so that it is sometimes difficult

to delimit those species as well.

Notes: Psydrax latifolia varies greatly in the

shape and size of its leaf blades and in the

prominence of its reticulate venation. Two

forms are distinguishable in the specimens

available for study, but they are not formally

recognised here because of the presence of so

many intermediate specimens. However, these

forms are generally as follows.

The ‘large-leaved’form. The majority of

specimens of this species from central Western

Australia, Northern Territory and South

Australia examined usually have comparatively

large leaf blades which are broad elliptic, broad

ovate or suborbicular in shape, are broad and

truncate apically, and are subcordate or obtuse

at base. These leaf blades have very conspicuous

lateral nerves and secondary reticulate veins,

are glabrous or slightly scabrid on both surfaces,

and are exceedingly pale brown or yellowish

brown when dry.

854

The ‘Barrier Ranges ’ form. Specimens

of this species from New South Wales and

south-western Queensland examined usually

have comparatively smaller, mostly elliptic leaf

blades which are usually abruptly obtuse

apically, and are narrowed proximally into the

petiole. They have comparatively much finer

lateral nerves and reticulate venation, and they

dry a brownish color. These specimens,

especially those indicated by * 7 in the list above,

closely resemble the syntype of P. latifolia from

the Barrier Ranges cited above.

Although both forms are recorded from a

variety of habitats, the ‘Barrier Ranges’ form

is found mostly in rocky situations, whereas

the ‘broad leaf’ form usually occurs on sandy

flats, plains or ridges usually in red loamy soil.

However, both forms sometimes grow in the

same area where they are connected by

intermediate specimens, for example, ones with

large, wide leaf blades but with much finer

lateral nerves and secondary reticulate

venation, or by specimens with comparatively

smaller leaf blades with coarse conspicuous

lateral nerves and secondary venation. The

existence of such specimens indicates that these

forms need futher study before they can be

satisfactorily delimited.

The branchlets of this species are

sometimes swollen due to insects living in their

tissues.

Uses: Fruits of Psydrax latifolia are reported

by collectors to be edible.

12. Psydrax ammophila S.T.Reynolds &

R.J.F.Hend., sp. nov. foliis latis

nervatione prominenti omata P. latifoliae

(F.Muell. ex Benth.) S.T.Reynolds &

R.J.F.Hend. simulans autem foliorum

base angusta, venatione reticulata obscura

differt. Typus: Queensland. Gregory

North District: 17 miles (c.27.2 km)

ENE of Headingly Station, 21°—’S,

138°—’E, 18 May 1948, R.A. Perry 848

(holo: CANB; iso: BRI, DNA, K,

PERTH).

Canthium sp. (Headingly Station R.A.Perry

848), PI. Forster & D.A. Halford (2002,

P-174).

Austrobaileya 6 (4): 817-889 (2004)

Shrubs or small trees 1-5 m high, sometimes

multistemmed; bark light grey; branchlets

glabrous, stout, terete, the young ones reddish

brown, the older ones dark grey. Leaves

opposite; stipules ovate-triangular, keeled and

attenuated into a folded lobe distally; petioles

(0.4-) 0.8-1.3 cm long, usually slightly winged

by decurrent leaf blade; blades elliptic or

elliptic-ovate, 5.2-7.0 (-8.5) cm long x 2.6-3.5

(-4.8) cm wide, with apex and base obtuse or

subacute, thick coriaceous, rigid; both surfaces

glabrous, dark green, drying yellowish green

or pale brown with age; lateral nerves in (3-)

4-6 pairs, obliquely arched, usually looping at

margins, prominent on adaxial surfaces;

reticulate venation fine, loosely arranged,

prominent on adaxial surfaces but not visible

abaxially; domatia absent or if present, then

only on one or two leaves of a branchlet where

1 or 2 occur on each side of the midrib, obscure.

Inflorescences shorter than the leaves, 2.2-3.5

cm long x 2.2-3.5 cm wide, 26-32-flowered;

main peduncle 7.0-12.0 mm long, with minute

bracts towards the middle or at the distal end,

terminated by 2 or 3 branched cymes. Flowers

comparatively small, 4-merous; pedicels of

solitary flowers 2.0-3.5 mm long, of others

0.5-2.0 mm long; calyx 1.5-2.0 mm long; limb

denticulate; corolla 5.0-6.5 mm longwith tube

campanulate, 1.5-3.0 mm long, sparsely or

densely hairy at the throat; lobes subelliptic,

2.5-4.5 mm long, obtuse, more or less cucullate

at apex, papillose abaxially near apex; stamens

with filaments 0.5-2.0 mm long and anthers

2.0-3.0 mm long; disc usually puberulous; style

(with stigma) 6.0-7.5 mm long. Fruits broad

ellipsoid, 5-8 mm long x 6-8 mm wide;

pyrenes rugose.

Additional representative specimens : Western Australia.

c.5 km W of Gahnda Rockhole, Gibson Desert (26°36’S,

125°49’E), Apr 1982, Kalotas 1127 (DNA, PERTH).

Northern Territory, between Mt Olga and Ayers Rock, Jun

1953, Symon 9434 (AD); No.2 Desert Bore, Hamilton Downs

(23°31’S, 133°16’E), Sep 1955, Chippendale 1695 (BRI,

NSW); Palm Valley (24°04’S, 132°43’E), Sep 1958,

Chippendale 4895 (DNA); Wallenby Gorge area, George

Gill Range, 1 mile (c.1.6 km) NE of Reedy Rock Hole

(24°20’S, 131°41’E), Jul 1968, Beauglehole 26483 (MEL);

90 km W of Stewart Highway, Victory Downs (25°58’S,

132°21’E), Jul 1982, Conrickll9 (AD, DNA); Edmirringee

Rockhole (20°36’S, 134°51’E), Sep 1983, Halford 83934

(DNA); 5 km NW of Mt Olga (25°18’S, 130°44’E), May

1985 , Bates 5606 (AD); 4 km S of Finke (Apatula) (25°35’S,

134°35’E), Jun 1988, Smith 1180 (DNA); 14 km W of

Waterloo Bore, Ti-Tree Station (22°09’S, 133°04’E), Jul

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

1992, Latz 12400 (DNA); 8 km NW of No. 14 Bore, Georgina

Station (20°56’S, 137°04’E), Sep 1992, Latz 12738 (DNA);

26 km NW of Barkly Homestead, Alyawarr Desert Survey

Site 15 (19°37’S, 135°37’E), Apr 1993, Parsons 392 (DNA);

100 km N of Annitowa Homestead, Wakaya Desert (20°24’ S,

136°13’E), May 1993, Latz 13085 (DNA); 50 km NNW of

Kantju Gorge, Uluru (25°21’S, 131°02’E), Jul 1995 , Nelson

2905 (DNA). South Australia. Upper Arkaringa Valley, Wa

WaWaterhole (Elder Expedition), May 1891, Helms s.n. (K,

MEL; NSW); Mulgrave Ranges, Mulgrave Park about 55

kmWNW of Mt Woodroffe, Jan 1964, Lange s.n. (AD); about

105 km NNW of Coober Pedy, Jun 1984, Badman 1096

(AD); Mt Crombie approx. 60 km SSW of Am ata (26°38’S,

130°48’E), 26 Sep 1985, Copely 1409 (AD). Queensland.

Gregory North District: Oban Station, 60 miles (c.96 km)

SW of Mt Isa, Woodend Bore, 21 0 —’ S, 139°—’E, Nov 1938,

Everist 1709 (BRI).

Distribution and habitat : Central Australia,

from Gibson Desert, Western Australia to

western Queensland; on red sand plains or in

rocky gullies or gorges, often near bores, on

sandy soil. (Map 14)

Diagnostic attributes : Psydrax ammophila is

characterised by its usually broad, prominently

nerved and finely reticulate-veined leaf blades

which are sometimes provided with domatia,

stout reddish coloured branchlets and

comparatively small inflorescences with 4-

merous flowers on short pedicels. However, this

species is imperfectly known, it being

represented in herbaria by mainly sterile

collections with very variable leaves. Collection

of more material is necessary to be certain of

this species’ characteristics.

Affinities: Psydrax ammophila is related to

P. latifolia and P. attenuata and some of the

specimens cited above have previously been

identified with those species or as an intergrade

between them. It resembles the former species

superficially in its leaves, branchlets,

inflorescence and flowers, but P. latifolia differs

from it by the usually broad base of its leaf

blades, absence of domatia, much more

conspicuous lateral nerves and reticulate

venation and shorter petioles (see also the key

to the Psydrax latifolia group above). These

species however are sometimes connected by

intergrades.

Psydrax attenuata may be readily

distinguished from P. ammophila by its usually

narrower leaf blades with fewer (2 or 3(or 4))

pairs of lateral nerves which are very oblique,

longer petioles which are 7.0-22.0 mm long,

slender reddish brown or grey branchlets,

855

longer pedicels which are usually 8.0-11.0 mm

long for solitary flowers, and 4- or 5-merous

flowers.

Notes: The following collections from south

of Alice Springs were previously identified as

P. latifolia, P. attenuata or as an intergrade

between these species but are now considered

to belong in P. ammophila. They differ from

the former two species by their comparatively

very small leaves, small inflorescences and

small flowers. In addition, they are not typical

of P. ammophila either and probably represent

a distinct form of it or an intergrade between

this species and P latifolia. However, as the

material available at present is too incomplete,

it is not possible to be certain of the actual

standing of these specimens. This taxon may

be described as follows.

Branchlets stunted; leaf blades elliptic,

c.3.2 cm long x 1.2 cm wide, subacute at both

ends, efoveolate, finely reticulate-veined;

inflorescences about 26-flowered with the main

peduncle 3.0-7.0 mm long and with small

bracts at its distal end; flowers 4-merous;

pedicels 2.0-2.5 mm long (solitary flowers) or

0.5-1.5 mm long (other flowers); calyx c.1.5

mm long; corolla 5.0-6.0 mm long with tube

2.5-3.0 mm long, densely hairy at throat, and

lobes 2.5-3.0 mm long, obtuse and slightly

cucullate at apex, papillose above the middle;

anthers 2.0-2.5 mm long on filaments 0.5-1.5

mm long; disc puberulous; and style with

stigma 6.0-7.0 mm long.

Northern Territory: Deep Well Road, 23.5 miles (c.38 km)

S of Alice Springs (24°05’S, 133°55’E), Oct 1964, Nelson

1366 (AD, BRI, NSW); c.37 km S of Alice Springs, Jul 1965,

Kuchel 2285 (AD).

Etymology : The specific epithet ammophila,

from Greek ammo-, ‘sand’, and -philus,

‘loving’, refers to the sandy soil on which this

species is usually found growing.

13. Psydrax reticulata (C.T.White)

S.T.Reynolds & R.J.F.Hend., comb, nov.;

Plectronia odorata var. reticulata

C.T.White, Proc. Roy. Soc. Qd. 50: 78

(1939). Type: Queensland. Cook

District: Thursday Island, date unknown,

E. Cowley 10 (holo: BRI).

856

Canthium sp. (Thursday Island E.Cowley

10), S.T. Reynolds (1997, p.181), P.I.

Forster & D.A. Halford (2002, p.174).

Shrubs or small trees to 6 m high, usually

stunted; bark grey, tessellated; branchlets brown

or reddish brown, slightly 4-angular distally,

glabrous. Leaves opposite; stipules subovate,

5.5-9.0 mm long, keeled and attenuated into a

folded lobe distally; petioles 0.2-0.6 cm long;

blades obovate or broad elliptic, (4.2-) 6.0-8.5

(-9.2) cm long x (3.0-) 4.4-6.2 (-7.3) cm wide,

with apex and base slightly rounded, or apex

retuse and base obtuse and shortly attenuate

into petiole, coriaceous, slightly stiff when dry

(due to prominent raised nerves and reticulate

veins); both surfaces glabrous, the adaxial

surface glossy, the abaxialy surface dull, usually

slightly bullate between the nerves; lateral

nerves in 4-8 pairs, obliquely arched and

ascending, looping at margins, with secondary

veins loosely reticulate, both lateral nerves and

reticulate venation usually prominent on both

surfaces (especially on dried specimens);

domatia usually small, sometimes absent.

Inflorescences 2.5-4.0 cm long x 4.5-6.0 cm

wide, 24-44-flowered; peduncles stout,

glabrous, the basal peduncle 1.0-2.0 cm long

with minute bracts near its middle and

terminated by 2-branched cymes; axis branches

0.8-2.5 cm long. Flowers 4-merous; pedicel

of solitary flowers 2.5-6.0 mm long, that of

others 1.0-3.0 (-4.0) mm long; calyx 1.5-1.7

mm long, with tube turbinate and limb 4-

denticulate; corolla 4.0-5.0 mm long; tube

broad campanulate, 2.0-2.5 mm long, with a

ring of dense hairs at throat; lobes elliptic or

ovate, 2.5-3.0 mm long x c.1.5 mm wide,

obtuse and more or less cucullate distally,

mostly recurved; stamens nearly as long as the

corolla, exserted; filaments + erect, c. 1.0 mm

long; anthers 2.0-2.5 mm long, erect or patent,

slightly exserted; disc shorter than calyx limb,

glabrous; style (with stigma) 6.5-8.0 mm long,

exserted, + sigmoid in younger flowers; stigma

oblong with 2, small, slightly recurved lobes

distally. Fruits (when didymous) transversely

ellipsoid or (when 1-lobed) subglobose, 4.0-4.5

mm long x 4.5-7.5 mm wide, black when ripe;

pyrenes hemispherical, rugose.

Additional representative specimens : Queensland. Cook

District: Prince of Wales Island, Feb 1975, Cameron 20270

(QRS); Moa Island, Aug 1985, Budworth s.n. (BRI); Mt

Austrobaileya 6 (4): 817-889 (2004)

Bremer, 26.0 km NE of Bamaga, 10°42’S, 142°31’E, Feb

1994, Fell 3920, Stanton & Roberts (BRI); 26.7 km NE of

Bamaga, 0.7 km NofMt Bremer summit, 10°41’S, 142°32'

E, Feb 1994, Fell 4048 & Stanton (BRI); Hammond Island,

date unknown, Heatwole & Cameron 254 (QRS).

Distribution and habitat : North Queensland

from Bamaga, Cape York Peninsula and Torres

Strait Islands; chiefly on steep hill and gully

slopes and creek banks close to the coast, on

sandy soil at the fringe of semi-deciduous vine

thickets (e.g. Pajinka Scrub near Bamaga).

(Map 6)

Diagnostic attributes: Psydrax reticulata is

readily recognisable by its shortly stalked,

comparatively broad, shiny, usually foveolate

leaves with prominent nerves and reticulate

venation. It resembles Psydrax latifolia in its

leaves, but that species differs from it by having

stiffer, dull leaves which lack domatia. It also

resembles P. suborbicularis from Papua New

Guinea because of its leaves, but that species

usually has comparatively small leaves with a

finer, less distinct reticulate venation, and hairy

inflorescences.

Notes: Psydrax reticulata is accepted here at

specific rank because of its distinctive,

comparatively large, broad, conspicuously

reticulate-veined leaf blades which clearly

distinguishes it from P. odorata under which

(as Plectronia odorata ) C.T. White described

it at varietal level in 1939. In Psydrax odorata ,

the leaf blades are usually thicker, more glossy

and have fewer primary nerves, and their

reticulate venation is more open and less

obvious.

14. Psydrax pallida S.T.Reynolds &

R. J.F.Hend., sp. nov. foliis latis

nervatione prominent reticulatoque

ornatis P. reticulatae (C.T.White)

S. T.Reynolds & R.J.F.Hend. simulans

autem foliorum base angusta, petiolis

longis differt. Typus: Queensland. Cook

District: Dixie Station, 15°03’S,

143°27’E, 1 December 1994, S.T Garnett

1545 (holo: BRI).

Canthium sp. (Dixie Station S.T.Garnett

1545), S.T. Reynolds (1997, p.180), P.I.

Forster & D.A. Halford (2002, p.174).

Shrubs or small trees 3-5 m high; bark

smooth, exceedingly pale grey or almost white

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

and blotched; branchlets greyish-white or very

pale brown, usually with scattered small

lenticels, the older ones with pale grey, finely

tessellated bark. Leaves opposite; stipules

deltoid, keeled and attenuate distally into a

usually long, folded decurrent lobe; petioles

2.0-3.0 cm long, flexuose; blades broad elliptic,

elliptic-obovate or elliptic, 5.0-9.0 cm long x

2.5- 5.0 cm broad, with apex obtuse or slightly

rounded and base subacute or acute and

attenuate into the long petiole, coriaceous with

both surfaces glabrous and often slightly rough,

rigid, drying yellowish green; lateral nerves and

reticulate veins very conspicuous (raised on

adaxial surface in dried specimens); lateral

nerves in 3-5 pairs, obliquely arched and

ascending towards the apex and looping near

the margins, or nerves sometimes very oblique

and ascending; tertiary veins ± closely

reticulate, obscure below; domatia small,

usually a few occurring on each leaf.

Inflorescences with spreading branches,

12-40-flowered; peduncles glabrous,

sometimes slightly resinous, the basal one

3.0-15.0 mm long with small bracts and usually

three branches at its distal end; branches

0.6-1.8 cm long, terminated by 5-18-flowered

cymes. Flowers 4(rarely 5)-merous; pedicel of

solitary flowers (4.0-) 7.0-10.0 mm long, that

of others 2.0-4.0 mm long; calyx about 1.5 mm

long; corolla 6.5-8.0 mm long, with tube

campanulate, 2.5-3.0 mm long x c.2.0 mm

wide, densely hairy at throat; lobes 4.0-5.0 mm

long, obtuse; stamens exserted; filaments erect,

2.5- 3.0 mm long; anthers 2.5-3.0 mm long;

disc shorter than the calyx limb; style (with

stigma) 7.0-8.0 mm long. Fruits subglobose

or broad ellipsoid, 5.0-6.0 mm long x 5.0-7.0

mm wide; pyrenes exceedingly rugose.

Additional representative specimens: Queensland. Cook

District: Princess Charlotte Bay, Marrett River, 14°25’S,

144°15’E, May 1979, Elsol & Stanley 667 (BRI)* 8 ; Dixie

Station, Sugarloaf, 15°02’S, 143°27’E, Dec 1994, Garnett

1555 (BRI); Battle Camp Range, 15°23’S, 144°40’E, Mar

1995, Garnett 1578 (BRI); Dixie Station, Lower Emu Creek,

15°04’S, 143°32’E, Apr 1995, Garnett 1579 (BRI).

Distribution and habitat: North Queensland;

usually in sandy soil on sandridges and

drainage flats, in open woodlands. According

to collector Garnett, this species occurs at the

base of mounds of the termite Amitermes

laurensis, in low open woodlands dominated

by Eucalyptus papuana F.Muell. and E.

857

clarksoniana K.D.Hill & L.A.S.Johnson

(Myrtaceae), and also with Melaleuca

viridiflora Sol. ex Gaertn. (Myrtaceae). This

species is known only from the above

specimens. (Map 14)

Diagnostic attributes: Psydrax pallida is

characterised by its long petiolate leaves with

broad, prominently nerved and reticulate,

usually foveolate blades, and open

inflorescences with flowers on slender pedicels.

Affinities: In its broad leaf blades with

prominent nerves and reticulate venation,

Psydrax pallida most resembles P. reticulata

but differs from that by having the base of the

leaf blade acute and attenuate into a much

longer petiole rather than rounded or obtuse

and shortly attenuate into a comparatively

shorter petiole. It also appears to be related to

P. latifolia in its broad, prominently nerved leaf

blades, e.g. in the collections from Dixie Station and

Battle Camp Range cited above, and to P. attenuata

in its inflorescences, flowers and colour of

branchlets. It differs from P. latifolia in its long

petioles, usually foveolate leaf blades with less

dense reticulate venation, exceedingly pale

brown branchlets and flowers on longer

pedicels. It differs from P attenuata in its

comparatively long petioles, its broad leaf

blades with prominent nerves and reticulate

venation, and its very pale brown branchlets.

However, P pallida is imperfectly known and

more specimens are necessary to be certain that

it is constantly distinct from these species.

Note: The Elsol & Stanley collection cited

above (* 8 ), which is in bud only, has a slightly

different aspect due to its leaves and

inflorescence. Its inclusion here as Psydrax

pallida is therefore tentative.

Etymology: The specific epithet pallida , Latin

for ‘pale’, refers to the very pale grey or pale

brown, almost whitish stems and very pale grey

twigs in this species.

15. Psydrax attenuata (Benth.) S.T.Reynolds

& R.J.F.Hend., comb, nov.; Canthium

attenuatum Benth., FI. Austral. 3: 421

(1867). TyP e: [Queensland.] North Coast

(Carpentaria islands ‘a’ (Sweers Island)

and ‘b’ (Bentinck Island), Nov 25-28

1802), R. Brown [3443] (lecto, here

designated: BM; isolecto: BM; K; MEL

[MEL1538097]).

858

Small trees 3-7 m high; bark pale or dark grey,

usually rough towards base, smooth distally;

branchlets pale brown, reddish brown or grey

mottled with white, but reddish brown, pale

grey or brownish coloured distally, the older

ones with somewhat rough bark, the younger

ones smooth and angular or subterete. Leaves

opposite; stipules ovate, prominently keeled and

attenuate into an apical lobe; petioles 0.5-1.7

(-2.2) cm long, subterete; blades elliptic,

narrow elliptic or lanceolate, 4.2-11.2 (-13.0)

cm long x 1.0-3.0 (-4.2) cm wide, with apex

acute, subacute or obtuse, and base acute or

subacute, attenuate and decurrent into the

petiole (usually to below middle of petiole),

coriaceous; both surfaces glabrous, yellowish

green or pale green, drying yellowish brown

or brown; midrib slender, mostly ridged

adaxially, raised abaxially; lateral nerves

usually distinct on both surfaces, in 2 or 3

(or 4) pairs, the lower 2 pairs usually decurrent

along midrib, opposite or alternate, slender,

oblique and looping at margins, or acutely

angled to the midrib; reticulate venation loosely

arranged, usually prominent; domatia absent

or present on all leaf blades or only on one or

two of the one branchlet, when present 1-3 on

each side of the midrib, usually small and

inconspicuous and situated in the axils of the

median pairs of nerves. Inflorescences 2.4-4.5

cm long x 3.5-5.5 cm wide, exceedingly open,

(8-) 13-90-flowered; basal peduncles 0.2-1.0

cm long, with minute bracts distally and

terminated by 2 or 3 branched cymes; branches

slender, 0.6-1.5 cm long, terminated by 3—11-

flowered cymes. Flowers 4- or 5-merous;

pedicels slender, those of solitary flowers (3.5-)

8.0-10.0 mm long, that of others 1.5-9.0 mm

long; calyx 1.0 -2.0 mm long, with tube

somewhat turbinate and limb shallowly 4- or

5-denticulate, the lobes broad ovate; corolla

5.5-8.0 mm long; tube obconic, dilated to

mouth, 1.5-3.5 mm long x 2.5-3.0 mm wide

at mouth, usually sparsely hairy at its throat;

lobes narrow ovate or elliptic, 3.0-5.5 mm long

x 1.0-1.5 mm wide, acute or subacute, erect or

subpatent; stamens erect, exerted; filaments

1.0-3.0 mm long; anthers 2.0-3.0 mm long;

style (with stigma) 6.0-10.0 mm long; stigma

broad and shallowly 2-lobed at apex. Fruits

obovoid or ellipsoid, 0.4-0.7 cm long x 0.5-0.8

cm across, 2-lobed or sometimes 1-lobed,

verrucose when dry; pyrenes broad ellipsoid,

usually exceedingly rugose.

Austrobaileya 6 (4): 817-889 (2004)

Distribution and habitat : From Dampier

Peninsula, Western Australia, to northern and

central Queensland, and on offshore islands;

in coastal vine thicket remnants or open

woodlands, on creek banks, rocky slopes and

ridges, on sandy, rocky or clay soils.

Diagnostic attributes: Psydrax attenuata is

characterised by its narrow elliptic or

lanceolate, usually foveolate leaf blades which

have domatia present on at least on some leaves

of a branchlet, and which are narrowed

proximally and decurrent into the petiole and

have prominent nerves and reticulate venation,

its open laxly branched inflorescences with

fragile 4- or 5-merous flowers on long slender

pedicels, and by its brownish, reddish brown

or greyish coloured branchlets.

Notes: Psydrax attenuata was found to be one

of the most complex and variable of Psydrax

species based on the many different-looking

specimens filed under this name in various

herbaria. However, on closer examination,

some of these specimens were found to be

identical with others filed as P. oleifolia (or

Canthium oleifolium ), while yet others

represented undescribed taxa associated with

the species concerned.

Bentham (1867) in describing what he

took to be a single species Robert Brown in

manuscript had called ‘ Pilidium attenuatum 1 ,

in effect cited ten syntypes for the name he gave

it with his brief diagnosis. Although his

description (especially with respect to

measurement of leaves) covered the specimens

he cited, three (possibly four) distinct species,

two (possibly three) of which are undescribed,

are distinguishable in these syntypes in addition

to Canthium attenuatum. Moreover, one of the

syntypes, namely Burdekin River, leg. F. Mueller

(identified as P. saligna in this paper), was cited

again by him under Canthium oleifolium

(Bentham 1867, p.422). Consequently specimens

which matched any of the specimens or came

from any of the localities cited by Bentham were

filed under the name C. attenuatum in herbaria,

making this species seem extremely variable

and very difficult to delimit.

To fix the application of Bentham’s name,

Canthium attenuatum is lectotypified here.

Bentham (1867), as indicated above, cited ten

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

syntypes under his brief diagnosis of

C. attenuatum namely “Brunswick Bay, N.W.

coast, A. Cunningham ; Victoria River and

Arnhem’s Land, F. Mueller (= 2 separate

collections); N. coast, R. Brown * 1 2 * * * * * * 9 ; Sweers

Island, Henne\ Burdekin and Burnett rivers

F. Mueller (= 2 separate collections); Port

Denison, W. Hill, Bowman (= 2 separate

collections); St George’s Bridge on the Balonne,

Mitchell”

Considering that Bentham primarily

intended to recognise the species that Brown

named ‘ Pilidium attenuatum ’ in manuscript,

Brown’s collection from the north coast of

Australia Brown annotated with this manuscript

name is here chosen as lectotype of Canthium

attenuatum. This lectotype, housed in

herbarium BM, agrees with Brown’s

manuscript description of this species (as

“No.29 spec”; microfilm of manuscript held in

BRI) and with Bentham’s protologue. An

illustration prepared contemporaneously with

Brown by Ferdinand Bauer from relevant

material has been published by Mabberley &

Moore (1999).

The isolectotype at K and that at MEL

are labelled “North Coast, leg. Brown ”, whereas

the isolectotype at BM is labelled “Carpentaria

island ‘a’ and ‘b’, Nov 25-28 1802, leg. Brown

3443”, and is annotated ‘ Pilidium ’ by Brown.

859

The other collections cited by Bentham

(now lectoparatypes) are referred to the

following species in this paper.

(a) The specimens from Brunswick Bay,

Victoria River and Arnhem Land are Psydrax

pendulina.

(b) The specimen from Sweers Island (K)

is referrable to Psydrax attenuata as delimited

in this revision (see under P. attenuata var.

attenuata).

and Port Denison are referrable to P. saligna.

(d) The specimen from St George’s bridge

on the Balonne [River] is referrable to P. oleifolia.

(e) The specimen from the Burnett River

was not seen in this study but probably belongs

in P. longipes, which occurs in that area.

Since Brown did not suggest the name

Canthium attenuatum in his manuscript,

Bentham only should be cited as the author of

that name in spite of what he said when

publishing it.

Affinities: Psydrax attenuata, in the sense

accepted in this paper, may be distinguished

from its close relatives P. lepida, P. longipes,

P. pendulina and P. saligna, and from P. oleifolia

as follows.

1. P. attenuata from P. oleifolia

Leaf blades discolorous, prominently nerved and reticulate veined; domatia

usually present; inflorescences open, loosely branched; pedicel of solitary

flowers usually 8.0-17.0 mm long; flowers 4- or 5-merous; corolla tube

usually densely hairy at its mouth . P. attenuata

Leaf blades concolorous, usually without domatia and conspicuous reticulate

venation; nerves obscure or distinct; inflorescences mostly small and

compact with flowers congested on branches; pedicel of solitary flowers

3.5-5.0 (-7.0) mm long; flowers 4-merous; corolla tube sparsely hairy

at its mouth. P. oleifolia

2. P. attenuata from P. saligna

Leaf blades 0.6-1.8 cm wide, 4-10 times as long as wide; reticulate veins

indistinct; domatia always present on leaf blades; inflorescences branched

once only, to 22-flowered; branchlets dark coloured. P. saligna

Leaf blades 1.0-3.0 (-4.2) cm wide, 3 or 4 times as long as wide; reticulate

veins prominent; domatia not always present on leaf blades;

inflorescences many-branched, to 50-flowered; branchlets reddish brown,

brown or grey . P. attenuata

860

Austrobaileya 6 (4): 817-889 (2004)

3. P. attenuata from P. longipes

Leaves with petioles 0.5-1.7 (-2.2) cm long and blades usually 1.4-3.0 cm

wide, 3-4 times as long as wide; domatia present on most or only some

leaves of a branchlet; pedicel of solitary flowers usually 8.0-11.0 mm

long; corolla 5.5-8.0 mm long. P. attenuata* 10

Leaves with petioles 1.2-3.5 cm long and blades usually 2.6-4.0 cm wide,

about twice as long as wide; domatia present only on some leaves of a

branchlet; pedicel of solitary flowers usually 5.0-7.0 mm long; corolla

7.5-11.0 mm long. P. longipes

(For * 10 above, see in particular P.

attenuata forma megalantha of this species.)

4. P. attenuata and P. lepida resemble

each other in their leaves, but the latter differs

from the former in its slightly shiny leaves

which always have domatia, and its 5-merous

flowers (see the key to the Psydrax attenuata

group above).

5. P. attenuata and P. pendulina may

readily be distinguished by the comparatively

long, strongly nerved, efoveolate leaves and

reddish coloured, 4-angular young branchlets

of the latter species (see the key to the Psydrax

attenuata group above).

Variability'. Psydrax attenuata, as circumscribed

here, varies greatly in its general appearance,

leaves (especially in the presence or absence of

domatia) and branchlet colour, the presence or

absence of galls on branchlets, the length of its

petioles and pedicels and the type of cymule

terminating each inflorescence branch. Though

three varieties are recognised here, they are

poorly known so that collection of more

specimens with associated field information in

the future may well result in a change of status

of all or some of these taxa.

Key to varieties of Psydrax attenuata

1. Domatia usually present on most leaves of a branchlet, 1-3 on each side of

the midrib; petioles usually 0.5-1.0 cm long. 15a. P. attenuata var. attenuata

Domatia absent or if present then only on one or two leaves of a branchlet

and usually one on each side of the midrib; petioles usually 0.7-2.2 cm long. 2

2. Pedicel of solitary flowers much longer than those of other flowers of the

cyme; domatia usually present (at least on one or two leaves a branchlet);

insect galls usually present on branchlets and stems . . . 15b. P. attenuata var. myrmecophila

Pedicels of all flowers of the cyme equal or subequal in length; domatia

rarely present on the leaves; insect galls usually absent on branchlets

and stems. 15c. P. attenuata var. tenella

15a. P. attenuata (Benth.) S.T.Reynolds &

R.J.F.Hend. var. attenuata

Leaf blades elliptic, 4.2-6.0 cm long, decurrent

into a comparatively short petiole; domatia 1-

3 on each side of midrib, usually prominent.

Additional representative specimens: Northern Territory.

Bessie Springs, 8 km ESE of Cape Crawford (16°40’S,

135°52’E), Oct 1975, Cardale s.n. (CANB)* 11 ; Calvert River

mouth (16°16’S, 137°46’E), Jun 1987, Thomson 1862

(DNA); Vanderlin Island, 7 km SE of Lake Eames (15°43 ’ S,

137°05’E), Aug 1988, Latz 11105 (DNA). Queensland.

Burke District: Sweers Island, SE of resort near the currently

used well, 17°07’S, 139°36’E, Nov 2002, Pedley & Thomas

SWI153 (BRI); Sweers Island, date unknown, Henne s.n.

(K); Gulf of Carpentaria (probably same as previous), date

unknown, Henne s.n. (K).

Distribution and habitat: Around theGulf of

Carpentaria and on close-by offshore islands;

usually in coastal scrubs. (Map 11)

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

Note : Psydrax attenuata var. attenuata is

poorly known and represented in this study by

the lectotype of the species name and a few

collections as indicated above. It is very si mil ar

to P. attenuata var. myrmecophila except for

its leaves with comparatively shorter petioles

and shorter blades. Collection of more

specimens in the future may indicate that the

latter variety is but a form of the former.

Of the specimens cited above, only the

Cardale specimen (* u ) is in flower (apart from

the lectotype specimen) but its leaves are too

young to be certain of their mature characteristics

so its inclusion here is tentative.

15b. P. attenuata var. myrmecophila

S.T.Reynolds & R.J.F.Hend., var. nov. a

P. attenuata var. attenuata petiolis

longioribus, foliis interdum foveolatis,

ramulis plerumque myrmecophlis differt.

Typus: Queensland. Burke District:

Lawn Hill National Park, 18°34’S,

138°38’E, November 1986, B. O’Keefe

2 (holo: BRI).

Bark dark grey or grey, usually rough at base

of stem and smooth distally; branchlets terete

or angular distally (sometimes dilated at nodes),

brown, reddish brown, yellowish brown or grey;

stems and branchlets often with prominent

insect galls. Leaf blades 4.7-13.0 cm long x

1.0-3.0(-4.2) cm wide, bright green, yellowish

green, pale or dull green, drying yellowish

brown or brown, dull or with a slight sheen on

the adaxial surface; lateral nerves in (2), 3 or 4

pairs, the middle one more obvious and acutely

angled to midrib and usually provided with a

domatium in each axil (domatia present only

861

on some leaves of a branchlet). Inflorescences

10-90-flowered; main peduncle 0.2-1.0 cm

long; branches 0.3-0.7 cm long. Flowers 4- or

5-merous; pedicel of solitary flowers (3.5-)

8.0-11.0 mm long, of others (1.0-) 2.5-6.0mm

long; calyx 1.0-2.0 mm long; corolla 6.0-8.0

mm long, usually sparsely hairy at the throat;

lobes narrow ovate, 4.0-5.5 mm long, obtuse.

Fruits broad ellipsoid, 4.0-7.0 mm long x

5.5-7.0 mm wide, exceedingly pale brown or

off-white and ribbed when dry; pericarp thin;

pyrenes exceedingly rugose.

Diagnostic attributes : Psydrax attenuata var.

myrmecophilia is characterised by its long

petiolate, narrow elliptic foveolate or efoveolate

leaves and galled stems and branchlets. It

appears to be scarcely distinct from P. attenuata

var. tenella, with which it shares the same leaf-

blade, domatia and long-petiole attributes, but

that variety differs by its subumbelliform

cymules and usually efoveolate leaves. More

collections especially of flowering plants are

necessary to be certain that these varieties are

constantly different.

Etymology : The varietal epithet myrmecophila ,

from Greek myrmeco-, ‘pertaining to ants’, and

-philus , ‘loving’, refers to the insect galls, home

to species of ants, found on stems and

branchlets of this variety.

Variability : The nature of the leaves, the colour

of branchlets and the size of inflorescence and

flowers are variable in specimens of this variety

available for study and a number of forms are

distinguishable within them. Two are

considered distinctive enough to be formally

recognised here.

Key to forms of Psydrax attenuata var. myrmecophila

1. Inflorescences (16-) 25-90-flowered; pedicel of solitary flowers 0.8.-1.0 cm

long, of others usually 0.5.-0.75 cm long; corolla 6.0-7.5 mm long;

branchlets usually reddish brown or brown

. 15b(i). Psydrax attenuata forma myrmecophila

Inflorescences (8-) 12-21-flowered; pedicel of solitary flowers (0.5-) 0.8-1.1 cm

long, of others 0.2-0.35 cm long; corolla 6.5-8.0 mm long; branchlets

brown, yellowish brown or grey. 15b(ii). Psydrax attenuata forma megalantha

862

15b(i). P. attenuata forma myrmecophila

S.T.Reynolds & R.J.F.Hend., forma nov.

a P. attenuata forma megalantha

inflorescentiis plerumque floribus 25-90,

floris solitarii pedicello bis pedicellos

ceteros in longitudine et floribus corolla

6.0-7.5 mm longa differt. Typus: As for

P. attenuata var. myrmecophila

Branchlets usually reddish brown and dilated

at nodes; leaf blades yellow brown when dry,

with prominent nerves and reticulate venation;

inflorescences (16-) 25-90-flowered.

Additional representative specimens: Northern Territory.

Larrimah to Darwin, Aug 1942, Huang s.n. (AD); 18.8 miles

(c.30.08 km) W of Soudan (20°04’S, 136°30’E), Oct 1957,

Chippendale 3830 (BRI, DNA, NSW); 10-12 miles (c.16-

19 km) NW of Cleanskin Yards, Sep 1967, Nicholls 719

(DNA)* 12 ; Cox River Station, Tanumbirini Creek (16°01’S,

134°47’E), Jul 1977, Latz 7311 (DNA); Gregory National

Park (15°37’S, 131°08’E), Feb 1986, Thomson 1281

(DNA)* 12 ; Barkly Stock Route (19°08’S, 136°46’E), Mar

1988, Brock 322 (DNA); Dunmarra (16°37’S, 133°19’E),

Mar 1991, Wilson 107 (DNA). Queensland. Burke District:

Carpentaria, in 1875, Gulliver 39 (MEL); Mt Isa, Oct 1930,

MacCallum 95 (BRI); Adel’s Grove, Jan 1946, deLastang

110 (QRS); 107 miles (c.172 km) NW of Mt Isa, on Barkly

Highway, 19°45’S, 138°05’E, Jul 1974, Ollerenshaw

P01276 & Kratzing (BRI); 25 km N of Mt Isa, Mt Isa to

Camooweal Road (20°33’S, 139°29’E), Jan 1987, Russell-

Smith 1882 & Lucas (DNA); Lawn Hill National Park, Jan

1989, O’Keefe [AQ454831] (BRI); 34 km N of Barkly

Highway on Thorntonia-Yelvertoft Road, 19°46’S,

138°48’E, Apr 1990, Latz 11624 (BRI); Mussellbrook

section of Lawn Hill National Park, 42.5 km N of

Mussellbrook Mining Camp, 18°21’S, 138°09’E,May 1995,

Thomas 818 & Johnson (BRI); 8.9 km SE of Mussellbrook

Mining Camp, 18°38’S, 138°ll’E,May 1995, Thomas 991

& Johnson (BRI).

Distribution and habitat : North-eastern

Northern Territory to north-west Queensland,

common on the Barkly Tableland; on ridges,

slopes and along creeks, on sandy and rocky

soil. (Map 11)

Diagnostic attributes : Psydrax attenuata forma

myrmecophila is characterised by its usually

foveolate, green or greenish yellow, narrow

elliptic leaf blades which attenuate proximally

into the petiole and which have exceedingly

oblique lateral nerves, and its laxly flowered,

open inflorescences with long-stalked, usually

5-merous flowers.

This form resembles P attenuata var.

tenella and P. attenuata forma megalantha in

its long, narrow leaf blades with very oblique

Austrobaileya 6 (4): 817-889 (2004)

nerves and its long petioles, but the former

taxon differs by its subumbelliform cymules,

whereas the latter one differs by its larger

flowers.

Variability : The leaves are quite variable in this

form. Specimens from shallow soil on rocky

slopes and ridges usually have narrow elliptic

or elliptic, pale green or greenish yellow leaf

blades with very oblique, fine lateral nerves and

obscure domatia, whereas specimens from

deeper soil on creek and river banks and

lateritic plains have elliptic usually efoveolate

leaf blades with often more conspicuous nerves.

Plants from lateritic country are reported by

collectors to be very green in colour with bright

green leaves whereas the leaves of plants from

other areas are reported as pale green or yellow

green.

Notes: The collections from Cleanskin Yards

and Gregory National Park cited above (* 12 )

are tentatively included here because in most

respects they appear to be of this variety.

However, as they are sterile, fertile specimens

would be necessary to be certain of their

placement.

The use of the epithet ‘ myrmecophila ’ in

the name of this form follows Recommendation

26A.3. in the current International Code of

Botanical Nomenclature (Greuter etal., 2000).

Uses: Plants of Psydrax attenuata forma

myrmecophila are reported by collectors to be

readily eaten by stock; sheep especially are

apparently very partial to it and graze foliage

to as far up as they can reach.

15b(ii). Psydrax attenuata forma megalantha

S.T.Reynolds & R.J.F.Hend., forma nov.

aemulans P. attenuata forma

myrmecophila S.T.Reynolds & R.J.F.Hend.

a quo inflorescentiis plerumque floribus

12-21 floris solitarii pedicello plerumque

plus quam ter pedicellos ceteros in

longitudine et floribus corolla 6.5-8.0

mm longa differt. Typus: Queensland.

North Kennedy District: Fletcher Creek,

HannHighway, 19°4-’S, 146°0-’E, November

1985, E.M. Jackes 19 (holo: BRI).

Bark dark or pale grey, usually finely tessellated

proximally, smooth distally; branchlets

brownish or greyish coloured mottled with

white; stems and branchlets often galled;

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

branchlets, petioles and nerves drying brown

or with a yellowish tinge. Leaves with blades

narrow elliptic, 8.0-13.0 cm long x 1.4-2.5

(-2.8) cm wide, with apex obtuse, and base

acuminate and attenuate into the comparatively

long petiole; adaxial surface yellow green;

abaxial surface bright green; lateral nerves

prominent, in 3 pairs, acutely angled to the

midrib, obscure or not apparent abaxially; veins

finely reticulate, prominent on young leaves,

indistinct on adult ones, not apparent on abaxial

surface; domatia present, inconspicuous, one

on each side of midrib, usually present only on

one or two leaves of the one branchlet.

Inflorescences (8-) 12-21-flowered; peduncles

and pedicels often drying reddish brown.

Flowers 5(rarely 4)-merous; pedicel of solitary

flowers (5.5-) 8.0-11.0 mm long, of others 2.0-3.5

mm long; calyx 1.5-2.0 mm long; corolla

6.5-8.0 mm long. Fruits broad ellipsoid, 1- or

2-celled, 4.0-7.0 mm long x 5.0-7.0 mm wide;

pericarp thin, shiny when dry; pyrenes rugose.

Additional representative specimens: Queensland. Cook

District: on Einasleigh-Lynd Roads nearEinasleigh, 18°3-

’S, 144°0-’E, Dec 1967, Sankowsky 736 & Sankowsky

(BRI). North Kennedy District: approx. 45 miles (72 km)

SE of Mt Garnett, 18°15’S, 145°25’E, Jan 1968, Morain

278 (BRI, K); Felspar, 10 km WNW of Charters Towers,

19°50’S, 145°15’E, Feb 1993, Rolfe 510 (BRI); Black

Mountain, NW of Pentland, 20°23’S, 145°04’E, Jul 1993,

Fensham 1040 (BRI). South Kennedy District: 3.5 miles

(c.5.6 km) SE of Yarrowmere Station, 21°30’S, 145°55’E,

May 1964, Adams 983 (BRI, CANB, K, PERTH); 8 kmNNE

of Carmichael Homestead, 21°54’S, 146°08’E, Apr 1992,

Thompson 206 & Simon (BRI). Mitchell District: 2.9 km

from turn-off to Cherhill, on road to Lake Dunn, 22°41’S,

145°31’E, Sep 1990, Wilson 526 & Rowe (BRI); about 42

kmEof Aramac, Mailmans Gorge, 22°—’S, 146°—’E, Oct

1994, Smyrell 85 (BRI). Leichhardt District: 1 mile (c.1.6

km) E of Langley Homestead, 80 miles (c.128 km) NE of

Emerald, 22°40’S, 148°59’E, Nov 1969, Auldist 24 (BRI,

K); Junee Tableland, N of Dingo, 22°47’S, 149°03’E, Nov

1990, Bean 2589 (BRI).

Distribution and habitat: Known only from the

above collections from northern and central

eastern Queensland; usually on sandy loam in

open woodlands. (Map 11)

Diagnostic attributes and affinities: Psydrax

attenuata forma megalantha is imperfectly

known as specimens of it available for study

are mostly poor and variable in morphological

attributes. More specimens, especially

flowering ones, are needed to be certain of its

characteristics and affinities. It is tentatively

863

included under P. attenuata because it

resembles P attenuata forma myrmecophila in

its long leaves, long petioles, domatia, lax

inflorescences, usually 5-merous flowers and

myrmecophilous branchlets. It has, however,

a different aspect from that form in its

branchlets and leaves, and also differs from that

by its yellowish tinged dried branchlets, petioles

and nerves, and larger flowers.

Variability: The leaves of this form are quite

variable. Specimens with shorter leaves

approach some of the collections included here

under P. longipes and are often difficult to

distinguish from that species because they have

the same general aspect, colour of dried

inflorescences and large 5-merous flowers.

However, P longipes generally has leaves with

wider blades and longer petioles, larger flowers

and usually whitish coloured branchlets which

are usually without insect galls. Specimens with

small, narrow, indistinctly nerved leaf blades

resemble those of P saligna but in that species

the leaf blades are usually foveolate and its

branchlets are dark coloured.

Etymology: The forma epithet megalantha ,

from Greek mega , Targe’, and anthos, ‘flower’,

refers to the comparatively large flowers found

in this form.

15(c). Psydrax attenuata var. tenella

S.T.Reynolds & R.J.F.Hend., var. nov.

quoad folios et adspectum P. attenuatam

var. myrmecophilam similem sed cymulis

subumbelliformis (florum pedicellis sub-

aequalibus), foliis plerumque efoveolatis

et ramulis non myrmecophilis differt.

Typus: Western Australia. Bobby Creek,

11 km ENE of Beagle Bay, Dampier

Peninsula, 16°58’S, 122° 47’E, 23 April

1988, B.J. Carter 267 (holo: BRI, iso:

PERTH).

Small, multistemmed trees; trunks with bark

somewhat dark grey, deeply furrowed

proximally, smooth distally; branchlets

exceedingly pale grey or dark grey, terete, very

rarely with insect galls. Leaves with blades

5.5-11.2 cm long x 1.7-2.5 cm wide, drying

greyish green or brown; lateral nerves in 3 or 4

pairs, with reticulate veins fine; domatia absent

or rarely present when 1 (rarely 2) on each side

of midrib. Inflorescences 22-50-flowered;

864

cymules subumbelliform. Flowers 4-merous;

pedicels slender, 0.5-1.0 cm long, those of

solitary flowers more or less of the same length

as those of other flowers; calyx 4-toothed;

corolla 6.0-7.5 mm long, with tube 2.5-3.5 mm

long and sparsely hairy at its mouth, and lobes

narrow ovate, 3.0-4.0 mm long. Fruits broad

ellipsoid, 6.5-7.0 mm long x 6.0-7.0 mm wide;

pericarp thin, pale brown when dry; pyrenes

rugose.

Additional representative specimens : Western Australia.

Emu Creek, Kununurra (15°45’S, 128°45’E), Dec 1982,

Done 639 (DNA); Broome (17°58’S, 122°14’E), Jun 1984,

Kenneally 9020 (PERTH); 3.2 km E of Bungle Bungle

Outcamp on track to Osmond Valley Palms Yard (17°20’S,

128°22’E), Jul 1984, Forbes 2547 (CANB, MEL); ditto,

Scarlett 84-296 (CANB, PERTH); 14.7 km S of campsite

adjacent to Red Rock Creek, 10 km ENE of Bungle Bungle

Outcamp, (17°26’S, 128°26’E), Jul 1984, Kenneally 9227

(K, PERTH); about 2 km SW of Broome (17°58’S,

122°13’E), Nov 1986, Wilson 12561 (PERTH).

Distribution and habitat : Western Australia,

from Dampier Peninsula to east Kimberley;

usually along creeks on sandy and clay soils.

(Map 11)

Diagnostic attributes: Psydrax attenuata var.

tenella differs from P. attenuata var.

myrmecophila in its subumbelliform cymules

terminating inflorescence branches and usually

efoveolate leaves. However, as only two of the

collections of it seen for this study are in flower,

it is difficult to be certain of all its

characteristics.

Variability: The leaves on specimens available

for study are variable. Their blades are

lanceolate or elliptic, prominently nerved and

finely reticulate-veined on the type specimen,

but are shorter on those from Bungle Bungle

Outcamp (which are probably only young

leaves), and longer and narrower on the

collection seen from Broome. The leaves of the

latter approach those of P. pendulina which

occurs in that area, but in that species the leaves

are usually more elongate, thin coriaceous, and

the young branchlets are quadrangular and

reddish in colour.

Etymology : The varietal epithet tenella, Latin

for ‘delicate, slender’, refers to the delicate,

subumbelliform cymules with flowers on long

slender pedicels in the inflorescences of this

variety.

Austrobaileya 6 (4): 817-889 (2004)

16. Psydrax lepida S.T.Reynolds &

R.J.F.Hend., sp. nov. P. attenuatae

(Benth.) S.T.Reynolds & R.J.F.Hend.

valde similis autem foliis foveolatis

subnitentibus, inflorescentiis lepidis,

pedicellis comparate longis et gracilibus,

floribus 5-meris corolla tubo in orificio

dense pubescenti differt. Typus:

Queensland. Cook District: Cooktown,

creek off Me Ivor River, January 1981, VT

Scarth-Johnson 1151A [AQ347294]

(holo: BRI).

Canthium sp. (Cooktown V.Scarth-Johnson

AQ314008), P.I. Forster & D.A. Halford

(2002, p.174).

Small trees with bark greyish coloured and

smooth; branchlets somewhat brown or usually

somewhat red coloured and slightly 4-angular

distally and dilated at nodes, the older ones with

brown, rough, usually tessellated bark. Leaves

opposite; stipules ovate, acuminate with a

folded lobed at apex and keeled; petioles

0.5-0.8 (-1.0) cm long; blades elliptic or

lanceolate, 4.2-8.0 (-9.5) cm long x 1.0-2.8

(-4.2) cm wide, attenuate at apex and base, or

with apex subacute or obtuse, and base acute

and attenuate into petiole, coriaceous, stiff when

dry; adaxial surfaces slightly shiny somewhat

dull abaxially, with both surfaces minutely

resin-dotted; midrib prominent on both

surfaces, raised when dry; lateral nerves in 1-3

pairs, prominent, the lower 2 pairs decurrent

with midrib, alternate, oblique, erect and

ascending, looping distally, (raised on adaxial

surface, indistinct on abaxial one in dried

specimens); reticulate veins distinct, fine,

loosely arranged; domatia present on most

leaves of a branchlet, small, with 1-3 on each

side of the mi drib. Inflorescences 2.5-4.5 cm

long x 3.5-5.5 cm wide, eceedingly slender,

laxly branched, (8-) 16-29-flowered; basal

peduncle 1.0-1.5 cm long, terminated by 2 or

3 branched cymes, axis branches 1.3-1.5 cm

long; cymes 6-10-flowered. Flowers 5-merous;

pedicel of solitary flower 0.8-1.7 cm long, that

of others (0.15-) 0.3-0.4 cm long; calyx 1.5-2.0

mm long, 5-toothed; corolla 5.5-7.0 mm long,

fragile; tube 2.0-3.0 mm long, inflated, dilated

to orifice where 2.0-2.5 mm wide, usually with

dense long white hairs projecting from the

orifice; lobes 3.5-5.0 mm long x c.1.5 mm

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

wide, subacute at apex; stamens with filaments

2.5-2.75 mm long, erect; and anthers 2.0-2.5

mm long; style (with stigma) 6.0-9.0 mm long.

Fruits obovoid or ellipsoid, c. 7.0 mm long x

8.0 mm wide; pyrenes broad ovoid, depressed

distally, exceedingly deeply rugose and

verrucose all over. (Fig. 6A-6E).

Representative specimens : Queensland. Cook District:

Finch Bay, near Cooktown, 15°25’S, 145°15’E, May 1966,

Smith 13100 (BRI); on road between Peach Creek and Leo

Creek, approximately 30-40 miles (48-64 km) NE of Coen,

approx. 13°44’S, 143°15’E, Oct 1969, Webb & Tracey 9896

(BRI); 10 km NE of Silver Plains Homestead (13°54’S,

143°36’E), Aug 1978, Paijmans 2959 (CANB); Quarantine

Bay, near Cooktown, Apr 1979, Scarth-Johnson s.n. (BRI);

19 km W of Peninsula Development Road, 7 km W of Emu

Creek, on the track to New Dixie Station, 15°03’S, 143°27’E,

Apr 1980, Clarkson 3132 (BRI, K)*' 3 ; Cooktown, creek off

Me Ivor River, 15°2-’S, 145°1-’E, Jan 1981, Scarth-Johnson

1151A (BRI); 1 km S of the Peninsula Development Road

on the track to Honey Dam about 4 km WSW of Lakeland

Downs Township, 15°52’S, 144°49’E, Jan 1986, Clarkson

6300 (BRI); Artemis Station, 15°00’S, 143°30’E,Dec 1994,

Garnett 1553 (BRI); Flinders Island, 14°11’S, 144°15’E,

May 1995, Le Cussan 506 (BRI). North Kennedy District:

approx. 45 miles (72 km) SE of Mt Garnett, 18°15’S,

145°25’E, Jan 1968, Morion 278 (BRI); The Bluff E of

Mingela, about 70 km S of Townsville, 19°53’S, 146°44’E,

Jan 1990, Cummings 10302 (BRI)* 13 ; Magnetic Island, date

unknown, Sandercoe 311 p.p. (BRI).

Distribution and habitat: Northern

Queensland, from Cape York Peninsula to near

Townsville; in remnant coastal vine thickets

inland of mangroves, on sand ridges, along

cliffs and creek banks, on sandy and rocky

sandstone soils. (Map 10)

Diagnostic attributes : Psydrax lepida is

characterised by its shiny foveolate leaves,

usually delicate inflorescences, 5-merous

flowers usually with dense hairs projecting from

the orifice of the corolla tube and red- brown

or brown branchlets with tessellated bark.

Affinities: Psydrax lepida is closely related to

P. attenuata and may be found with further

study to be conspecific with that species. It

resembles P. attenuata var. attenuata in its

leaves with oblique nerves and short petioles,

but in that taxon domatia are usually present

only on some leaves of a branchlet, the leaf

blade surfaces are dull, and its flowers are 4-

or 5-merous with only sparse hairs at the orifice

of the corolla tube (see the key to the Psydrax

attenuata group above).

865

Notes: The majority of the specimens seen

typically have narrow elliptic or lanceolate leaf

blades with domatia on all leaves. A few

specimens however have narrower leaf blades

(for example, see specimens indicated by * 13 in

the list above) which approach those of P. saligna

but that species differs from P. lepida in its

usually dull leaf-blade surfaces, small, few-

flowered inflorescences and dark coloured

branchlets.

The following collections from

northern Queensland, which have been

identified as P. attenuata or P. odorata in

the past, though mostly incomplete are

tentatively included here because in their

branchlet and leaf characters they appear to

be of this species. They differ, however, from

the other collections of P. lepida by their

wider, obscurely nerved leaf blades with

usually a solitary obscure domatium, shorter

petioles and few-flowered inflorescences.

Queensland. Cook District: 9 miles (c. 14.4 km) S of Laura

on main Mareeba-Coen Road, approx. 15 0 41’S, 144°34’E,

Oct 1969, Webb & Tracey 9925 (BRI); Split Rock to Gugu

Yelandji, S of Lake, Lakefield Downs-Laura road, 15°4-’S,

144°2-’E, May 1975, Byrnes 3374 (BRI); 31 km from the

Peninsula Development Road on the Fairview to Fairlight

Road, 15°36’S, 144°02’E, Nov 1983, Clarkson 5038 (BRI,

K, QRS).

Etymology: The specific epithet lepida , Latin

for ‘neat, fine or elegant’, refers to the graceful

inflorescences and dainty flowers found in this

species.

17. Psydrax saligna S.T.Reynolds &

R.J.F.Hend., sp. nov. ad P. attenuatae

(Benth.) S.T.Reynolds & R.J.F.Hend.

affinis sed foliis comparate angustis vel

angustissimis, nervis lateralis obscuris

tenuibus ornata, inflorescentiis

brevioribus paucifloris differt. Typus.

Queensland. South Kennedy District: off

Range Road, approximately 4 km from

Lake Elphinstone Turnoff, 21°32’S,

148° 16’E, 14 November 1987, I.G.

Champion 308 (holo: BRI).

[Canthium attenuatum auct. non Benth.: G.

Bentham, FI. Austral. 3: 421 (1867)

quoad specimini Burdekin River, [leg.]

F. Mueller ] et Port Denison, [leg.] W. Hill.

866

Austrobaileya 6 (4): 817-889 (2004)

Fig. 6. Psydrax lepida. A. flowering branch x 1. B. flower x 4. C. LS of flower x 4. D. fruit x 4. E. pyrenes x 4. A-C, Scarth-

Johnson 1151A(BRI); D, Volck 4418 (BRI); E, Smith 13100 (BRI ). Psydrax saligna. F. flowering branch x 0.6. G. flower

x 4. H. LS of flower x 4. F-H, Champion 308 (BRI).

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

[Canthium oleifolium auct. non Hook.f.: G.

Bentham, FI. Austral. 3: 422 (1867)

quoad specimini Burdekin River, [leg.]

F. Mueller].

Small trees 2-6 m high; trunk with bark dark

brown or dark grey, smooth or rough;

branchlets dark grey or dark reddish brown,

with finely tessellated bark, the young ones

reddish brown especially distally. Leaves

opposite; stipules triangular, keeled and

attenuated into an apical acuminate lobe, to 5.5

mm long; petioles (0.2-) 0.5-1.4 cm long,

subterete; blades narrow elliptic or lanceolate,

sometimes somewhat linear, (6.0-) 7.5-11.5

(-14.5) cm long x 0.6-1.5 (-2.0) cm wide, with

apex acuminate or obtuse and base narrowed

and decurrent into petiole, coriaceous, drying

dull and opaque; both adaxial and abaxial

surfaces glabrous; lateral nerves in 1 or 2 (or 3)

pairs, the lower pair usually acutely angled to

the midrib, slender, usually indistinct on

adaxial surface, not apparent on abaxial

surface; reticulate veins obscure; domatia

present, small, 1-3 on each side of midrib.

Inflorescences 1.2-3.0 cm long x 1.2-3.5 cm

across, 5-15 (-22)-flowered; basal peduncle

1.0-8.0 mm long, usually with minute bracts

medially and terminated by 2 branched few-

flowered cymes; axis branches 5.0-10.0 mm

long. Flowers 5(rarely 4)-merous; pedicels very

slender, that of solitary flowers (5.0-) 10.0-18.0

mm long, of others 3.0-5.5 mm long; calyx

about 1.5 mm long, with tube turbinate and

limb 4- or 5-toothed; corollas 6.0-8.0 mm long;

tube campanulate, occasionally slightly inflated

and obscurely striped, 2.0-3.5 mm long x 1.5-2.0

mm wide, densely hairy at the orifice; lobes

elliptic or ± lanceolate, 3.0-5.0 mm long x 1.5-2.0

mm wide, obtuse, erect or slightly recurved

distally; stamens with filaments 2.0-3.0 mm

long and anthers about 2.5 mm long, erect or

slightly patent; style (with stigma) 6.0-7.0 mm

long. Fruits broad ellipsoid or obovoid, 4.0-5.0

mm long x 4.0-6.0 mm wide, black and shiny

when ripe; pyrenes rugose. (Fig. 6F-6H)

Distribution and habitat: Arnhem Land,

Northern Territory, and northern and central

867

Queensland; usually in sandy soil among

sandstone rocks and granite boulders, crests of

sandy ridges, sandstone escarpments, on steep

slopes and granitic outcrops, in deciduous vine

thickets.

Diagnostic attributes: Psydrax saligna is

readily distinguishable by its comparatively

narrow, finely-nerved, foveolate leaves, its

comparatively small, 5-15 (-22)-flowered

inflorescences which have a short peduncle and

axis branched only once, its 5-merous flowers

on long slender pedicels, and its dark grey or

dark reddish brown branchlets.

Affinities: Psydrax saligna is closely related

to P. attenuata and P. oleifolia. In the past, some

of the specimens cited below have been

included in those species but P. saligna may be

distinguished from them as follows.

P. saligna and P. attenuata both have

foveolate leaves and usually 5-merous flowers

on comparatively long slender pedicels but

P. attenuata differs from the former in its leaves

with broader, prominently nerved and

reticulate-veined blades and longer petioles,

and its larger inflorescences with a greater

number of flowers (see the key to the Psydrax

attenuata group above).

P. saligna and P. oleifolia resemble each

other in their leaves (especially in the typical

form of P. oleifolia ) and their comparatively

small inflorescences, but P oleifolia differs

from the former by its usually efoveolate leaves,

its many-flowered (to 39-flowered) compact

inflorescences, its comparatively short pedicels

(those of solitary flowers being 3.5-5.0 (-7.0)

mm long) and its 4-merous flowers.

Etymology: The specific epithet saligna, Latin

for ‘willow-like’, refers to the leaves of this

species which resemble those of a Salix species

(Salicaceae).

Variability: The leaves and inflorescences of

Psydrax saligna are quite variable. Based on

these attributes, two forms are recognised here.

868

Austrobaileya 6 (4): 817-889 (2004)

Key to forms of Psydrax saligna

1. Pedicel of solitary flowers, filiform, 1.0-1.8 cm long; leaves with petioles

0.8-1.4 cm long and blades 6.0-10.0 cm long x 0.6-1.6 (-2.0) cm wide

. 17(b). P. saligna forma filiformis

Pedicel of solitary flowers comparatively stout, 0.5-1.0 cm long; leaves

with petioles 0.2-0.7 cm long and blades 7.5-12.5 (-14.5) cm long

x 0.9-1.5 (-1.8) cm wide. 17(a). P. saligna forma saligna

17(a). Psydrax saligna S.T.Reynolds &

R.J.F.Hend. forma saligna

Canthium sp. (Charters Towers T.Stuart

TWR116), S.T. Reynolds (1997, p.180),

PI. Forster & D.A. Halford (2002, p.174).

Small trees with usually smooth grey bark on

the trunk. Leaves with blades narrow elliptic

to almost linear, the lateral nerves and reticulate

venation exceedingly fine, sometimes obscure,

and domatia 1 or 2 on each side of the midrib.

Inflorescences 7-15 (-22)-flowered; pedicel of

solitary flowers 5.0-10.0 mm long, that of

others 3.0-5.5 mm long; corolla 7.0-8.0 mm

long with tube campanulate, not striped. (Fig. 6B)

Additional representative specimens : Queensland. Cook

District: Giant Horse Gallery, 15°40’S, 144°30’E, Mar

1975, Hyland 8102 (BRI, QRS); Mt Mulligan, about 40 km

NW of Dimbulah, 16°51’S, 144°50’E, Apr 1985, Clarkson

5899 (BRI). Burke District: Georgetown (18°48’S,

143°48’E), Nov 1942, Blake 14722 (DNA); Pairie Gorge,

45 km NNE of Hughenden, 20°30’S, 144°30’E, Jun 1986,

Murray 64 & Morgan (BRI)* 14 ; Warang Holding, White

Mountains, about 37 km NNW of Torrens Creek Township,

20°29’S, 144°44’E, Jul 1988, Fell 1310 & Swain (BRI)* 14 ;

Westmoreland Valley behind Little Amphitheatre, 17°24’S,

138°16’E, May 1997, Forster PIF21138 & Booth (BRI).

North Kennedy District: Burdekin River, Oct 1856, Mueller

s.n. (BM, K, MEL [MEL 1537684])* 15 ; Port Denison, in 1874,

Fitzalan [MEL1537674] (MEL); Ewan, Nov 1930, Miller

s.n. (BRI); 14kmN ofBowen, on Townsville Road, 19°27’S,

147°57’E, Nov 1971, Sharpe 57 (BRI); between Burdekin

Dam andRavenswood, 19°5-’S, 145°2-’E, Dec 1988 , Perry

2 (BRI); Lucky Downs, Greenvale Area, 18°59’S, 144°58’E,

Apr 1991, Batianoff 9104042 & Franks (BRI)* 14 ; Round

Mountain, 3 km Wof Ross River Dam, Townsville, 19°28’S,

146°42’E, Jun 1991, Bean 3298 (BRI); Gloucester Island,

southern end, 20°02’S, 148°27’E, Sep 1992, Batianoff

9209542 (BRI); Sally’s Mesa, 34 km from Greenvale towards

Charters Towers, 19°05’S, 145°14’E,Feb 1994, Bean 7461

& Forster (BRI)* 14 ; Yahoo Waterhole, 500 m N of Warrang

homestead, White Mountain National Park, 20°26’S,

144°49’E, Jan 1995, Anchen 176 (BRI); NW of Townsville,

19°15’S, 146°36’E, Sep \991,Fensham 3327 (BRI). South

Kennedy District: Moonoomoo Station, about 4 km N of

homestead, 21°46’S, 146°05’E, Oct 1983, Henderson

H2783, Guymer & Dilleward (BRI)* 14 . Leichhardt District:

7 km from Nebo on Nebo-Clermont Road, 22°15’S,

148° 15’E, May 1962, Johnson 2375 (BRI); Taunton

National Park, 23°30’S, 149°12’E, Dec 1996, Stansk T914

(BRI)* 14 . Port Curtis District: The Springs Grazing Trail,

Livingstone Shire, 22°51’S, 150°17’E, Nov 1983, Anderson

3602 (BRI); Table Mt, Rockhampton, O’ShanesyAl (MEL).

Distribution and habitat : Northern and central

Queensland; mostly on sandstone on

escarpments, steep slopes and rocky outcrops,

on sandy soil amongst boulders or in sandy

rocky soil, in open forests. (Map 12)

Variability: The leaves of Psydrax saligna

forma saligna are very variable. Specimens

with leaf blades usually narrow elliptic, finely

nerved and reticulately veined, and with one

or two domatia on each side of the midrib are

typical of it. However, specimens with shorter

or longer or smaller leaves are also present.

Those with shorter leaves, especially ones from

central Queensland, are not unlike some of the

collections included in typical P. oleifolia and

consequently have been identified as that

species in the past. Specimens with usually

smaller, narrow elliptic leaf blades, which have

a much duller adaxial surface and fainter

nerves, for example, the collections from Mt

Mulligan and Giant Horse Gallery above, and

specimens with leaf blades 10.5-14.5 cm long

and 0.6-0.8 cm wide that are acute or

acuminate at the apex and base and occasionally

have slightly recurved margins and stipules

with a prominent long, folded lobe at the apex

(indicated by * 14 in list above) probably

represent distinct subforms of this form.

However, since these specimens are mostly

sterile or have young flower buds only, this

cannot be ascertained at the present time.

Notes: Ferdinand Mueller’s collection from

Burdekin River in October 1856, marked * 15

in the above list, was cited by Bentham (1867)

under both Canthium oleifolia and Canthium

attenuatum Benth. (and hence is one of the

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

syntypes of the latter name). Of the three sheets

of this Mueller collection seen in this study,

two of them (those at BM and MEL) are

referrable to Psydrax saligna, whereas the sheet

at K represents a mixed collection of P. saligna

(the part with long, narrow, foveolate leaves)

and P. oleifolia (the part with broader,

efoveolate leaves).

17(b). Psydrax saligna forma filiformis

S.T.Reynolds & R.J.F.Hend., forma nov.

a P. saligna forma saligna florum

pedicellis filiformis duplo saltern

longioribus differt. Typus: Northern

Territory. Echo Gorge, Wollogorang

Station, 17°11’S, 137°43’E, November

1984, B. Thomson 795 (holo: BRI; iso:

DNA, PERTH).

Small trees with smooth or rough bark on the

trunk. Leaves with blades narrow elliptic,

obscurely nerved and with domatia 1-3 on each

side of the midrib. Inflorescences (5-) 7-9-

flowered; flowers with pedicels filiform, that

of solitary flowers (6.0-) 14.0-18.0 mm long,

of others 2.0-4.0 mm long; corolla 6.0-7.0 mm

long with tube broad campanulate, inflated,

obscurely striped.

Additional representative specimens : Northern Territory.

Jabiru, Bukbukluk Lookout jumpup on Kakadu Highway,

13°35’S, 132°15’E, Nov 1991, Brennan 1614 (BRI)* 16 ;

Jabiru, Ranger U Mine lease, 12°55’S, 132°55’E,Nov 1994,

Brennan 2953 (BRI)* 16 ; Mt Gilruth area (12°58’S,

133°10’E), Jun 1978, Dunlop 4900 (DNA); Centre Island,

Sir Edward Pellew Group (15°41’S, 136°46’E), Feb 1976,

Craven 3787 (CANB); Caranbirini Waterhole, 33 km SW

of Borroloola (16° 16’S, 136°05 , E),Nov. 1975, Cardale s.n.

(CANB); Wollogorang Station, Echo Gorge, 17°H’S,

137°43’E, Nov 1984, Halford 841161 (BRI).

Distribution and habitat : Known only from the

above collections; usually on sandstone. (Map 12)

Affinities: P. saligna forma filiformis differs

from the typical form in its leaves with longer

petioles, which are (0.4-) 0.8-1.4 cm long, its

smaller, 5-9 (-15)-flowered inflorescences, and

its flowers with a comparatively long filiform

pedicel and usually broader corolla tube which

is sometimes slightly inflated and obscurely

striped.

Notes: Collections from near Jabiru, marked

* 16 in the above list, are only tentatively

included here because they differ from the

869

others in having very pale coloured smooth bark

on the stems, wider leaf blades, which are

1.6-2.0 cm wide, and shorter pedicels, those

of solitary flowers being (5.0-) 7.0-9.0 mm

long, of others 2.0-3.0 mm long. However, in

their aspect, leaves with short petioles which

are 0.4-0.6 cm long, small inflorescences with

12-15 flowers on a very short basal peduncle

3.0-5.0 mm long, they belong with P salicina

forma filiformis.

Etymology: The forma epithet filiform is, Latin

for ‘thread-like’, refers to the comparatively

long, very slender pedicels of the flowers of

this form.

18. Psydrax pendulina S.T.Reynolds &

R.J.F.Hend. sp. nov. aemulans P.

attenuatam (Benth.) S.T.Reynolds &

R.J.F.Hend. a qua foliis longioribus,

lamina subfalcata, nervis lateralibus

divergentibus (a costa ad angulum

40°-50°), efoveolata, ramulis 4-angulis,

floribus 4-meris differt. Typus: Western

Australia. Brunswick Bay, 3rd Voyage of

the Mermaid, October 1820, A.

Cunningham 178 (holo: BM; iso: K).

[Canthium attenuatum Benth., FI. Austral.

3: 421 (1867) quoad specimenibus

Brunswick Bay, [leg.] A. Cunningham ;

Victoria River and Arnhem Land, [leg.]

F. Mueller, et Victoria River, [leg.] F.

Mueller ].

[Canthium sp. A, B.L. Koch in J.R. Wheeler,

ed., FI. Kimberley Region 906, 927, Fig.

285, Aj & A, (1992), pro parte.

Shrubs or small trees 2-8 m high, usually with

slender, drooping stems and leaves; bark dark

grey, rough; young branchlets quadrangular

distally, usually drying dark reddish brown and

often covered with white thin flaky bark, the

older ones usually blackish coloured and

verrucose. Leaves opposite; stipules towards

apex of branchlets subulate with a fine apical

point, those proximally on branchlets smaller

and ovate; petioles 1.0-1.7 (-2.0) cm long,

flexuose, erect or subpatent, usually drying pale

yellow or whitish (as pale as midrib); blades

narrow elliptic though mostly wider below

middle, usually subfalcate, (8.5-) 11.5-17.0

(-18.5) cm long x 1.5-2.2 (-2.7) cm wide, with

870

Austrobaileya 6 (4): 817-889 (2004)

apex acuminate, subacute or rarely obtuse, base

cuneate and attenuate into petiole, and both

surfaces glabrous, usually thin coriaceous,

drying brownish coloured with a pale yellow

midrib and nerves; midrib ridged above; lateral

nerves conspicuous, in 2 or 3 (-5) pairs,

divergent, angled 40°-50° to the midrib, erect

and ascending to near apex of leaf blade,

looping at margins; reticulate venation very

open, obscure; domatia absent. Inflorescences

in axil of new leaves or above scar of fallen

leaves, 2.4-3.5 cm long x 2.4-4.7 cm across,

12-23-flowered, glabrous; basal peduncle (2.0-)

4.0-8.0(-14.0) mm long, with 2 or 3 branched

cymes and minute bracts at its distal end; axis

branches 7.0-16.0 mm long; ultimate cymules

6-10-flowered. Flowers 4-merous; pedicels of

solitary flowers (5.0-) 8.0-11.0 mm long, that

of others 3.0-6.0 mm long, or occasionally the

pedicels of all the flowers more or less equal

when cymules are subumbelliform; calyx 2.0-3.0

mm long, with limb 4-toothed, the lobes broad

ovate; corolla 6.5-7.5 mm long, with tube

2.5- 3.0 mm long, dilated to the orifice, 1.5-2.0

mm wide at base, 2.0-4.0 mm wide at orifice,

usually with dense hairs projecting from orifice;

lobes ovate-oblong, 4.0-5.0 mm long x

1.5- 2.0 mm wide, obtuse, erect to spreading;

disc shorter than calyx limb, glabrous or

subglabrous; stamens with filaments 1.0-2.0

mm long and anthers 2.0-3.0 mm long, erect

or slightly recurved; style (with stigma) 7.0-10.0

mm long. Fruits obovoid or transversely

ellipsoid, obscurely 2-lobed, 6.0-10.0 mm long

x 7.0-10.0 mm across; pericarp drying soft and

very thin; pyrenes hard, usually slightly rugose.

Additional representative specimens : Western Australia.

Drysdale River Mission, Kimberley, Aug 1921, Gardner

1049 (PERTH); Napier Broome Bay (14°02’S, 126°36’E),

Aug 1921, Gardner 1549 (NSW); Bushfire Hill, Prince

Regent River Reserve (15°28’S, 125°39’E), Aug 1974,

George 12303 (PERTH); Phython Cliffs, Prince Regent River

Reserve, W Kimberley (15°20’S, 124°56’E), Aug 1974,

Kenneally 2184/B (CANB, PERTH); Ashton Range,

Drysdale River National Park (15°16’S, 126°43’E), Aug

1975, George 13302 (PERTH); Hidden Valley, 3.2 km E of

Kununurra (15°43’S, 128°48’E), Jul 1976, Beauglehole

54207 (PERTH); S of Ernest River (15°25’S, 127°27’E),

Mar 1978, Hartley 14688 (PERTH); Dampier Peninsula,

Djulanan, western-most inl et of Curlew Bay (16°24’S, 123°1-

’E), Aug 1986, Smith 867 (PERTH); Dampier Peninsula, 8

kmNW of One Arm Point (16°25’S, 123°01’E), Jan 1988,

Carter 190 (PERTH); Dampier Peninsula, S of Hunter Creek,

7 km ESE of Cape Leveque (16°25 ’ S, 122°59’E),Feb 1993,

Carter 616 (PERTH); Koolan Island (16°07’S, 123°43’E),

Feb 1993, Keighery & Gibson 16 (PERTH). Northern

Territory. Victoria River (15° 13’S, 129°48’E), Oct 1855,

Mueller s.n. (K, MEL [MEL1537680 & MEL1537688]);

Eva Valley Station, 14°15’S, 132°19’E, Oct 1973, Dunlop

3097 (BRI, DNA, K); 1 km N of Pine Creek-El Sharana Rd

on Jim Jim road, 13°32’S, 132°19’E, Oct 1978 .Rankin 1487

(BRI, DNA); above Moline Rock Pool (13°34’S, 132°16’E),

Nov 1978, Rankin 1638 (DNA); El Sharana Road, E Pine

Creek, 13°33’S, 132°17’E, Dec 1979, Dunlop 5215 (BRI,

DNA, K, NSW); Katherine Gorge National Park (14°19’S,

132°27’E), Oct mi,Silvertsen 624 (DNA); ditto, Apr 1987,

Purdie 3389 (BRI, CANB); Kakadu National Park, near

Kurundie Creek (13°34’S, 132°29’E), Apr 1990, Cowie 1166

& Leach (DNA); Nitmiluk, 14°19’S, 132°25’E, Dec 1990,

Evans 3494 (BRI, K).

Distribution and habitat : Northern Australia,

from Dampier Peninsula, Western Australia, to

Arnhem Land, Northern Territory; amongst

sandstone rocks, on sandstone plateau, base of

sandstone outcrops and hills, on sandy soil, in

open forests. (Map 13)

Diagnostic attributes : Psydrax pendulina is

readily distinguishable by its 4-angular, reddish

brown young branchlets, comparatively long,

narrow elliptic, efoveolate leaf blades with

conspicuous, suberect-divergent nerves (nerves

acutely angled to midrib) and 4-merous flowers

on comparatively long slender pedicels.

Affinities: Psydrax pendulina is closely related

to P attenuata and specimens of the former

have been included in that species in the past

by Bentham (1867), namely the syntypes of

P. attenuata from Victoria River and Arnhem

Land collected by Mueller , and from Brunswick

Bay collected by Cunningham. P. attenuata ,

however, differs from the former by its terete

or slightly angular, brown, reddish brown or

greyish coloured juvenile branchlets, and its

comparatively smaller, thicker, usually

foveolate leaf blades which are 4.7-11.0 (-13.0)

cm long x 1.5-3.0 cm wide (see the key to the

Psydrax attenuata group above).

Variability: The majority of relevant specimens

seen for this study are what are considered

typical of this species. That is, they have thin,

coriaceous, narrow elliptic, subfalcate leaf

blades and usually expanded spreading

inflorescences. However, one or two collections

have either typical narrow long thin leaves but

short inflorescences while others have thick or

shorter leaf blades with expanded spreading

inflorescences.

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

Note: The following sterile collection from low

open eucalypt woodland, on red sandy soil, is

tentatively included here. It differs from the

collections cited above by having broad elliptic

leaf blades which are 12.5-13.5 cm long x 3.5-5.0

cm wide, but in its leaf texture, nervature and

branchlet characters, it is of P. pendulina. This

collection probably represents an aberrant form

of this species because, according to the label,

the plants sucker in groups and the specimens

may have been taken from one of these suckers

with, perhaps, juvenile foliage.

Western Australia, approximately 1 km N of Derby airport,

17°19’S, 123°39’E, Tracey 13977 (BRI, QRS).

Etymology: The specific epithet pendulina,

Latin for ‘hanging down’, refers to the usually

more or less pendulous branchlets and leaves

of this species.

19. Psydrax oleifolia (Hook.f.) S.T.Reynolds

& R.J.F.Hend. comb, nov.; Canthium

oleifolium Hook.f. in Mitchell, Trop.

Australia 397 (1848). Type: New South

Wales. Plains of the Gwydir (between the

Gwydir and Barwon Rivers), December 1846,

T.L. Mitchell 491 (holo: K; iso: K, NSW).

[Canthium attenuatum Benth., FI. Austral.

3: 421 (1867) pro parte quoad specimini

St. George’s Bridge on the Balonne, [leg.]

Mitchell].

Erect shrubs or small trees 4-7 m high,

usually with straight, erect trunk and stiff,

spreading, divaricate branches borne more or

less at right angles to the main trunk, hispid

and often provided with thorns in the juvenile

or regrowth state; bark grey, smooth, or rough

proximally and smooth distally; branchlets

terete, usually resinous distally, glabrous,

smooth or usually spinulose in the juvenile or

regrowth state, pale brown or grey, with

brownish or greyish coloured bark mottled with

white distally. Leaves opposite or occasionally

clustered on brachyblasts on lower part of

branchlets but opposite distally, more or less

erect; stipules comparatively small, ovate with

a long, subulate lobe at apex, keeled, viscid in

young growth; petioles 0.6-1.2 (rarely to 1.7)

cm long, flexuose; blades elliptic, narrow

elliptic, elliptic-ovate, elliptic-oblong or

subobovate, (3.5-) 4.5-6.5 (-8.5) cm long x

871

(0.7-) 1.7-2.4 (-3.3) cm wide, with apex

obtuse, ± rounded, subtruncate or retuse and

slightly recurved, base subacute, acute or

subobtuse and abruptly narrowed and decurrent

into petiole, and both surfaces dull, glabrous,

smooth or hispid in juvenile or regrowth plants,

usually concolorous, pale green and slightly

glaucous, or yellowish green, thick coriaceous,

drying thick and rigid, pale yellowish green to

pale brown; midrib raised below, slightly

flattened above; lateral nerves either obscure

and subpatent, then in 2-4 (or 5) pairs, or

distinct suboblique or oblique and sometimes

looping near margins, then in 2 or 3 pairs with

2 pairs usually more obvious; nerves not visible

on abaxial surface; reticulate venation not

apparent; domatia absent or very rarely present

when usually solitary and inconspicuous.

Inflorescences usually exceedingly small,

1.0-2.6 cm long x 1.5-4.0 cm across, densely

flowered with 23-39 flowers, with basal

peduncle usually between 2.5-4.0 mm long, the

flowers usually congested on the short branches

which are 3.0-7.0 mm long and slender, or the

inflorescences occasionally loosely flowered

with basal peduncles to 10 mm long; peduncles,

pedicels and calyces glabrous, usually drying

reddish brown or brown. Flowers 4(rarely 5)-

merous; pedicel of solitary flowers 3.5-5.0

(-7.0) mm long, that of others 0.5-2.5 (-4.0)

mm long; calyx 1.0-2.0 mm long, with tube

campanulate and limb 4(or 5)-lobed, the lobes

ovate, minute; corolla 4.0-6.5(rarely 9.0) mm

long; tube broad cylindrical or broad turbinate,

1.5-2.0 mm long, 1.5-2.0 mm wide at orifice,

sparsely hairy at throat; lobes subelliptic or

lanceolate, 3.0-5.0 (-6.5) mm long x 1.0-1.75

mm across, obtuse, erect or spreading; disc

shorter than calyx limb, glabrous; stamens with

filaments 1.5-2.5 mm long, erect, and anthers

2.0-2.5 mm long, erect or rarely patent; style

(with stigma) 7.0-8.0 mm long, exserted;

stigmatic knob oblongoid. Fruits obovoid or

broad ellipsoid, 4.5-6.0 (-7.0) mm long x 4.5-7.0

mm across; pyrenes broad ovoid, depressed

distally or slightly ellipoid, usually rugose

distally and laterally. (Fig. 7A-7F)

Additional representative specimens: Queensland.

Mitchell District: Noondoo, 28°35’S, 148°25’E, Dec 1934,

Everist 751 (BRI, K); Blackall, 24°25’S, 145°45’E, Sep

1937, Brass & White 5 (BRI); Warren Point Station, 26°32’S,

148°01’E, May 1968, Martensz s.n. (BRI)* 17 . Leichhardt

District: Blackwater, E of Emerald, 23°35’S,148°55’E, Mar

872

Austrobaileya 6 (4): 817-889 (2004)

Fig. 7. Psydrax oleifolia. A. fruiting branch (of typical form) x 0.6. B. branchlet (of juvenile growth form) x 1. C. flower x 6.

D. LS. of flower x 6. E. fruit x 3. F. pyrene x 6. A,C-F, Auldist 10 (BRI); B. Everist & Smith 6 (BRI). Psydrax longipes. G.

flowering branch x 1. H. portion of leaf showing venation x 1.1. flower x 4. J. FS of flower x 4. K. fruit x 2. F. pyrene x 4.

G&H, Forster PIF2765 (BRI); I&J, Forster PIF6142 (BRI); K&F, Sankowsky 460 (BRI).

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

1920, Francis s.n. (BRI)* 17 ; 4 miles (c.6.4 km) S of Brigalow

Research Station, 24°55’S, 149°47’E, Jun 1969, Auldist 10

(BRI, K); Banana Shire, about 1 km S of Brigalow Research

Station, 24°50’S, 149°48’E, Nov 1984, Anderson 3886

(BRI); Springsure, 24°00’S, 148°00’E, Nov 1987, O’Keeffe

884 (BRI); Autumnvale, Bingle Shire, 25°51’S, 148°41’E,

Dec 1991, Schefe 1157 (BRI); 4 km N of Springsure,

24°04’S,148°05’E, Mar 1995, Fensham 2650 (BRI); 3 km

NE of Taroom, 25°37’S, 149° 49’E, Nov 1996, Halford

Q3065 & Dowling (BRI). Warrego District: Cunnamulla,

28°04’S, 145°41’E, Jan 1937, Everist & Smith 6 (BRI)* 17 ;

Gilruth Plains, Dec 1940, Allen 20 (CANB)* 17 ; 25 miles

(c.40km) N of Augathella, 25°4-’S, 145°3-’E, Dec 1961, Cull

s.n. (BRI). Maranoa District: Maranoa River, on range

bearing SW of camp, Sep 1846, Mitchell s.n. (BM); Boorara

near Yalleroi, 24°05’S, 145°35’E, Dec 1935, Everist 1461

(BRI); 20 miles (c.32 km) NE of Surat on Yuleba Rd,

26°55’S, 149°15’E, Aug 1956, Everist 5819 (BRI); 70 miles

(c.112 km) SE of Charleville, ‘Wheatleigh’, Boatman,

27°10’S, 146°50’E, Jul 1962, Ebersohn E73 (BRI); 9 miles

(c.14.4 km) W of Mitchell, in 1970, Hanger s.n. (BRI).

Darling Downs District: about 40 km N of Goondiwindi

along Leichhardt Highway (28°20’S, 150°17’E),Feb 1983,

Telford 9515 & Butler (CANB); Broadwater Lagoon, Dalby,

27°21’E, 151°06’E, Nov 1985, collector unknown

[AQ398749] (BRI); 15 km SE of Inglewood, 28°30’S,

151°11’E, Oct 1993, Halford Q1988 (BRI). New South

Wales. Narran Swamps (29°—’S, 147°—’E), Mar 1846,

Mitchell 50 (CGE, K); Lachlan River (33°13’S, 147°20’E),

in 1881, Ramsay [MEL1537973] (MEL); Bourke to

Barrington Road, Nov 1893, collector unknown

[NSW193735] (NSW)* 17 ; Bourke (30°05’S, 145°56’E),in

1896, McDougall [NSW193740] (NSW)* 17 ; ditto, Dec

1939, Beuzeville s.n. (NSW)* 17 ; Coolabah (31°05’S,

146°38’E), Dec 1898, Peacock s.n. (NSW)* 17 ; Narrabri, Nov

1899, Maiden s.n. (NSW); Boppy Mountain, 28 miles (c.45

km) E of Cobar, towards Nyngan (31°32’S, 146° 17’E), Nov

1903, Boorman [NSW193727] (NSW); ditto, Oct 1972,

Sikkes 169 & Telford (CANB)* 17 ; Nyngan (31°39’S,

147°12’E), Jan 1905, Rogers s.n. (NSW)* 17 ; Warialda

(29°32’S, 150°35’E), Jul 1905, Bowman s.n. (NSW); Cobar

(31°30’S, 145°50’E), in 1910, Abrahams [NSW193728]

(NSW)* 17 ; Barwon River, Narrabri (30°20’S, 149°47’E),

Nov 1912, Julius s.n. (NSW); Moree (29°28’S, 149°51’E),

Apr 1919, Gilmour s.n. (NSW); Narrabri-Gunnedah, Sep

1926, Welch s.n. (NSW); Pera Bore, about 10 miles (16 km)

W of Bourke, Aug 1928, Benton s.n. (NSW)* 17 ; 25 miles

(c.40 km) W of Nyngan (31°33’S, 146° 46’E), Nov 1949,

Reik & Common s.n. (CANB, K)* 17 ; Yantabulla District

(28°20’S, 145°—’E), Apr 1965, McReadie (NSW)* 17 ; 30

km NNW of Cobar, 0.5 km N of Little Tank (31°15’S,

145°15’E), Sep 1978, Crisp 4232 (CBG); Castlereagh River

(30°—’S, 147°—’E), date unknown, Blake [MEL1537974]

(MEL).

Distribution and habitat: In drier parts of

western Queensland and New South Wales,

from near Emerald, central Queensland to

Lachlan River, New South Wales; on sandy

plains, sandy ridges usually on red or yellow

sand, or on clay or on hard ridges, in open

woodlands, usually occurring in small groups.

(Map 13)

873

Diagnostic attributes: Psydrax oleifolia as

circumscribed here is a very variable species

which is characterised by thick, dull, obscurely

or distinctly nerved, concolorous, elliptic or

narrow elliptic leaves which are without

reticulate veins and usually without domatia,

and by its usually small, compact inflorescences

with short peduncles and pedicels, and usually

small flowers and fruits.

Variability: The shape and size of leaf blades,

inflorescences and flowers, and the distinctness

of nerves on the leaf surface are all very variable

in the specimens available for study. Two

distinct forms and a number of subforms are

distinguishable but, as they are all connected

by intermediate forms throughout their range,

they are not formally recognised here. The

distinct forms are as follows.

(1) Typical form: Specimens with narrow,

elliptic oblong or subobovate leaf blades,

usually obscure, suboblique or subpatent nerves,

shortly stalked, comparatively small

inflorescences, usually comparatively small

flowers and obovoid fruits, as represented by

those indicated * 17 in the above list (which

resemble the type specimen), are typical of this

species. The majority of specimens from New

South Wales seen, especially those from the

north-west plains of that state, are of this form.

It is usually found on sandy plains, undulating

country or sandy ridges and hillsides, in red or

yellow sandy soil.

(2) Broad-leaved form: The majority of

specimens seen are of this form. They differ

from the typical form by their usually

comparatively broader, larger or longer, elliptic

or elliptic-oblong leaf blades with distinct

oblique nerves, usually larger inflorescences

and larger flowers on longer stalks. This form

usually occurs on heavy (clay) soils and hard

ridges or sometimes on sandy ridges. It is

reported to occur in association with Brigalow

{Acacia harpophylla F.Muell. ex Benth.:

Mimosaceae), whereas the typical form has not

been reported as occurring in such a

community.

However, as indicated above, these forms

are connected by intermediates, namely

specimens with larger or broader leaf blades

with indistinct nerves and large open or small

874

congested inflorescences, or specimens with the

small or narrow leaf blades of the type specimen

but with distinct nerves and often larger

inflorescences. In addition, these forms also

sometimes grow sympatrically making it

extremely difficult to delimit them.

In addition, the broad-leaved form often

intergrades into P. attenuata and P. longipes,

and is, therefore, often difficult to delimit from

those species as well.

Notes: Psydrax oleifolia is related to Pforsteri,

P johnsonii, P. longipes and P saligna and

some of the specimens cited under those species

here were previously filed under the names

P. oleifolia, P. attenuata or P. buxifolia (now

applicable to a subspecies of P. odorata) in

various herbaria. P. oleifolia may be distinguished

from all these species as indicated below.

(1) P oleifolia and P attenuata may be

distinguished using the key under the latter

species above.

(2) P. oleifolia and P. forsteri may be

distinguished by the comparatively small leaves

and the 5-17-flowered inflorescences of the

latter species (see the key under P. forsteri ).

Austrobaileya 6 (4): 817-889 (2004)

(3) P. oleifolia and P. johnsonii have

similar leaves, inflorescences and flowers, but

the latter differs from P oleifolia in its foveolate

leaves and hairy branchlets and inflorescence

axes (see the key under P. johnsonii ).

(4) P. oleifolia and P. longipes may be

distinguished by the long petioles, the usually

broader, often foveolate leaf blades with

prominent lateral and reticulate nervation, and

the larger flowers of the latter species, as in the

key to the Psydrax oleifolia group above.

However, specimens of P. oleifolia with wider

elliptic leaf blades or longer or larger leaves

which are distinctly nerved with oblique nerves

are often difficult to separate from P. longipes,

except that in P. oleifolia the petioles are

relatively shorter, the lateral nerves do not

extend to the apex of the leaf as in P. longipes,

and both reticulate veins and domatia are

usually absent on the leaf blades. The species

are also connected by intergrades, for example

specimens with narrow leaf blades, obscure

nerves and prominent domatia, or specimens

with wide leaf blades, prominent nerves and

domatia but without obvious reticulate

venation.

(5) P. oleifolia and P. odorata subsp.

buxifolia may be distinguished using the

following key.

Leaf blades shiny, discolorous; branchlets and inflorescence axes hairy;

pedicels stout; corolla fleshy. P. odorata subsp. buxifolia

Leaf blades dull, concolorous; branchlets and inflorescence axes glabrous;

pedicels slender; corolla thin. P. oleifolia

(6) P. oleifolia and P. saligna may be

segregated using the key under the latter species

above.

Common names: wild lemon, native orange

or myrtle tree.

Uses: Psydrax oleifolia is reported by collectors

as being a good fodder for stock and that sheep

are partial to it.

20. Psydrax longipes S.T.Reynolds &

R.J.F.Hend., sp. nov. P. oleifoliae

(Hook.f.) S.T.Reynolds & R.J.F.Hend.

affinis a qua imprimis differt foliorum

petiolo longiore lamina majore latiore

nervis prominentibus, venatione reticulata

ornatis, floribus majoribus pedicellis

longioribus. Typus: Queensland. Port

Curtis District: Larcom Gully, Calliope

Shire, 23°44’S, 150°53’E, 24 November

1987, N. Gibson 1057 (holo: BRI).

Canthium sp. (Larcom Gully N.Gibson

1057), S.T. Reynolds (1997, p.180), P.I.

Forster & D.A. Halford (2002, p.174).

Shrubs or small trees 3-7 m high; trunk with

bark greyish, slightly blotched, rough and

tessellated proximally, smooth distally;

branches usually divaricate; branchlets terete

or subterete, whitish or greyish coloured

mottled with white. Leaves opposite; stipules

ovate, keeled, attenuated into a short or long

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

folded lobe distally; petioles 1.2-2.5 (-3.5) cm

long; blades elliptic, obovate or subobovate,

(6.0-) 8.0-10.0 cm long x (1.7-) 2.6-4.0 cm

wide, with apex obtuse, rounded or subacute

and base acute or subacute and attenuate and

decurrent into the petiole, thick coriaceous, with

adaxial and abaxial surfaces glabrous, pale

green or yellowish green when fresh, drying

yellowish green or brownish coloured; lateral

nerves in 2 or 3 pairs, arched and ascending,

or acutely angled to the midrib, ascending to

near apex of the lamina, the adaxialmost pairs

usually forming a distinct loop, prominent on

adaxial surface, usually indistinct abaxially;

reticulate venation fine, open, usually distinct;

domatia present at least on one or two leaves

of a branchlet with one or few on each side of

the midrib, inconspicuous, or domatia absent.

Inflorescences open, 2.0-2.6 cm long x

2.4- 3.8 cm across, 12-28 (-39)-flowered with

peduncles and pedicels drying pale yellow or

reddish-brown; basal peduncle 4.0-9.0 mm

long with minute bracts at its distal end; axis

branches 5.0-7.0 mm long. Flowers 4- or 5-

merous; pedicel of solitary flowers (3.0-)

5.0-7.0 mm long, that of others 1.0-3.0 (-4.0)

mm long; calyx 1.0-2.0 mm long, with tube

turbinate and limb 4- or 5-denticulate; corolla

7.5- 9.0 (-11.0) mm long; tube campanulate,

2.0-3.5 mm long, c.2.5 mm wide at orifice,

densely or sparsely hairy in its throat; lobes

ovate-oblong or elliptic, 5.0-6.5 (-8.0) mm

long x 1.0-2.0 mm wide, subacute or obtuse,

erect or slightly recurved; stamens with

filaments 2.0-2.5 mm long, erect, and anthers

2.0-5.0 mm long, erect; disc shorter than calyx

limb, glabrous; style (with stigma) 7.0-12.0

mm long. Fruits ellipsoid, obscurely 2-lobed,

(4.0-) 6.0-7.0 mm long x 5.0-7.0 mm across;

pericarp usually exceedingly wrinkled when

dry; pyrenes hemispherical or depressed ovoid,

usually exceedingly rugose distally and

laterally. (Fig. 7G-7L)

Additional representative specimens : Queensland. Port

Curtis District: Rockhampton, Nov 1865, Dietrich 2074

(MEL); Dawson Highway, Calliope Shire, 24°—’S, 151°1-

’E, Jan 1983, Gibson 494 (BRI); The Springs, Livingstone

Shire, 22°51’S, 150°17’E,Nov 1983 , Anderson 3602 (BRI).

Burnett District: 12 km W of Mundubbera near Burnett

River, 25°35’S, 151°15’E, Jun 1984, Kent s.n. (BRI). Wide

Bay District: Nora Creina, Didcot, Biggenden Shire,

25°58’S, 151°53’E, Dec 1981, Forster 138B (BRI); Mt

Perry, 25°27’S, 151°52’E, Dec 1986, Forster PIF2765

(BRI); 1 km NE of Didcot, 25°28’S, 151°52’E, Feb 1994,

875

Forster PIF14809 (BRI). Maranoa District: Roma, 26°3-

’S, 148°4-’E, Feb 1938, Blake 13284 (BRI). Darling Downs

District: 7.5 km W of Chinchilla on Warrego Highway,

26°44’S, 150°34’E, Dec 1994, McKenzie s.n. (BRI).

Distribution and habitat: Central and

southeastern Queensland; usually on rocky hill

slopes, gravelly ridges and rocky creek banks,

in eucalypt woodlands on rocky or sandy-clay

loamy soils. (Map 10)

Diagnostic attributes: Psydrax longipes is

characterised by its comparatively long

petiolate leaves with elliptic or subobovate

blades which are sometimes provided with a

small domatium, its large, usually 5-merous

flowers in loosely branched inflorescences, and

by its very pale brown or whitish coloured

branchlets.

Affinities: Psydrax longipes is related to P oleifolia

and P attenuata and some of the specimens

cited above have been included in herbaria

under either of those species names. It

resembles the former species in its leaves,

general aspect, whitish coloured terete

branchlets, and in the colour of dried

inflorescences, but P oleifolia differs form it

by its efoveolate, usually narrower leaves,

obscure or distinct nerves without any reticulate

veins between them, and usually compact,

comparatively small inflorescences, small

flowers and fruits (see the key to the Psydrax

oleifolia group above). However, specimens of

P. oleifolia with larger or wider leaf blades with

distinct oblique nerves are sometimes difficult

to delimit from some of the collections included

here under P. longipes, but the latter species

differs from the former by its leaves with blades

with distinct reticulate veins and the usual

presence of domatia on at least some leaves of

a branchlet, and longer petioles. These species,

however, are connected by intergrades and

further study of more specimens in the field

may indicate that P. longipes represents the

extreme of the range of variation of the very

variable P. oleifolia. Until more collections of

P longipes are seen, however, these species are

kept distinct here because their extremes are

very diverse.

Psydrax attenuata may be distinguished

from P. longipes by its leaves with usually

narrower blades, which are usually provided

with domatia, and shorter petioles, its smaller

876

Austrobaileya 6 (4): 817-889 (2004)

flowers with longer pedicels, its brown or

reddish brown usually angular young

branchlets, and by its usually galled branchlets

and stems (see under P. attenuata above).

However, P. longipes can sometimes have

narrow leaf blades approaching those of

P. attenuata forma megalantha , which also has

large flowers and occasional domatia on its

leaves, but differs from that form by its larger

flowers with shorter pedicels, shorter leaves and

pale grey branchlets.

Etymology : The specific epithet longipes, from

Latin longus, ‘long’, and pes, ‘foot’, refers to

the long petioles of the leaves in this species.

21. Psydrax johnsonii S.T.Reynolds &

R.J.F.Hend., sp. nov. quoad faciem

foliorum inflorescentiarumque P.

oleifoliae (Hook.f.) S.T.Reynolds &

R.J.F.Hend. maxime similis sed foliis

foveolatis, inflorescentiis pubescentibus

in pedunculis brevioribus portatis,

floribus plerumque 5-meris differt.

Typus: Queensland. Leichhardt District:

Duaringa Shire, Berrigurra Station, about

6 km N of Tolmies, 23°32’S, 148°44’E,

9 Dec 1981, E.R. Anderson 2829 (holo:

BRI).

Canthium sp. (Berrigurra Station

E.R.Anderson 2829), S.T. Reynolds

(1997, p. 180), P.I. Forster & D.A. Halford

(2002, p.174).

Small trees 3-8 m high, usually much

branched with numerous divaricate branches;

trunk with bark grey brown mottled with white,

dark grey or blackish coloured, rough

proximally, smooth distally; branchlets terete,

dark or pale grey, drying + yellowish coloured

distally, the young branchlets minutely

puberulous. Leaves opposite; stipules ovate,

keeled, attenuate into a extended lobe distally;

petioles 0.4-1.1 cm long, minutely puberulous

when young; blades elliptic, narrow elliptic or

subobovate, 2.6-4.5 cm long x 0.7-1.5 (-2.0)

cm wide, with apex obtuse or more or less

rounded, and base usually cuneate and attenuate

into the petiole, thick coriaceous, glaucous,

drying thick and rigid; adaxial and abaxial

surfaces glabrous, or sparsely hairy in young

leaves, slightly shiny to somewhat dull on

adaxial surfaces, pale green to olive-brown or

yellow green; lateral nerves in 2 or 3 (or 4)

pairs, oblique, arched and looping at margins,

obscure; reticulate venation not apparent;

domatia 1-3 on each side of the midrib, usually

prominent, or very rarely absent.

Inflorescences open, 2.0-2.8 cm long x

2.0-2.8 cm wide, 22-25-flowered; peduncles

slender, the basal one 12.0-15.0 mm long,

usually with minute bracts towards the middle,

minutely puberulous when young; axis

branches divaricate, 1.0-3.0 mm long, minutely

puberulous when young. Flowers (4 or) 5-

merous; pedicel of solitary flowers 3.0-5.0

(-6.5) mm long, of others 1.0-1.5 (-2.5) mm

long, minutely puberulous when young; calyx

1.0-1.5 mm long, minutely puberulous when

young, with tube somewhat urceolate and limb

denticulate; lobes 4, minute; corolla 5.0-6.0

mm long; tube somewhat cylindrical, 1.5-1.7

mm long, with dense long hairs at orifice; lobes

elliptic or sublanceolate, 3.5-4.0 mm long x

1.0-1.5 mm wide, obtuse or subacute, erect or

recurved; stamens with filaments 1.5-2.0 mm

long, erect, and anthers c.2.0 mm long, erect

or slightly recurved; disc shorter than calyx

limb, glabrous; style (with stigma) 7.0-8.0 mm

long, exserted; stigma oblongoid, 2-lobed

distally. Fruit broad ellipsoid, depressed

proximally and distally, 4.0-6.0 mm long x

4.5-6.5 mm across, slightly rugose when dry;

pyrenes usually slightly rugose on the dorsal

side. (Fig. 8A-8E)

Additional representative specimens : Queensland. North

Kennedy District: Foot Print Scrub, Fanning River Station,

19°42’S, 146°26’E, Jan 1993, Fensham 632 (BRI). South

Kennedy District: Exmoor, 21°00’S, 148°04’E,Dec 1992,

Fensham 640 (BRI); Mount Blackjack, 21°01’S, 147°56’E,

Jan 1993, Fensham 639 (BRI); Bluegrass Creek, Ceitah,

21°14’S, 147°38’E, Jan 1993, Fensham 643 (BRI).

Leichhardt District: Nogoa River, in 1890, Foot [MEL

1538093] (MEL); Scott’s Peak, 22°51’S, 149°13’E, Jun

1951, Everist 4425 (BRI); 20 miles (c.32 km) S of Banana,

24°45’S, 150°09’E, May 1964, Speck 1996 (BRI, CANB,

K, NSW); Cabbage Creek Crossing, 35.6 km from Cracow,

25°29’S, 150°13’E, Jul 1990, Forster PIF7046 (BRI); Lotus

Creek, 22°24’S, 149°07’E, Feb 1993, Fensham 665 (BRI);

38 kmNE of Taroom, 25°27’S, 150°07’E, Oct 1996, Halford

Q3066 & Dowling (BRI). Port Curtis District: Callide

Valley, Apr 1917, White s.n. (BRI); Biloela, 24°24’S,

150°30’E, Oct 1947, Smith 3544 (BRI, K). Burnett District:

Narayan, 25°3-’S, 150°5-’E, Nov 1969, collector unknown

(BRI); Murgon-Kingaroy area, 26°—S, 151°—E, Dec 1995,

Haager s.n. (BRI).

Distribution and habitat : Eastern Queensland,

from Fanning River to Burnett River; in dry

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

877

Fig. 8. Psydrax johnsonii. A. flowering branch x 1. B. flower x 6. C. LS of flower x 6. D. fruit x 3. E. pyrene x 6. A, Fensham

643 (BRI); B-E, Evens! 4425 (BRI). Psydrax forsteri. F. flowering branch x 1. G. flower x 6. H. LS of flower x 6.1. fruit

x 3. J. pyrene x 6. F, Speck 1819 (BRI); G&H , Anderson 3604 (BRI); I&J, Kerswell [AQ424459] (BRI).

878

Austrobaileya 6 (4): 817-889 (2004)

rainforests usually on rocky hills and rocky

creek beds, usually in association with Brigalow

(Acacia harpophylla F.Muell. ex Benth.:

Mimosaceae). (Map 14)

Diagnostic attributes: Psydrax johnsonii is

distinguishable by its foveolate, elliptic,

obscurely nerved, thick, more or less dull leaf

blades with prominent domatia, hairy

peduncles and branchlets, and usually 5-merous

flowers. It is closely related to P. oleifolia of

which it has more or less the aspect, leaves and

flowers, but differs from that species by the

prominent domatia on its leaves, and hairy

branchlets and inflorescence axes. Moreover,

P. johnsonii is reported mostly as growing in a

Brigalow community, usually on rocky hills or

creek beds, whereas P oleifolia is usually found

on sandy plains and ridges, not in association

with Brigalow.

Variability : The leaves of Psydrax johnsonii

are variable in the specimens available for

study. Specimens with narrow, elliptic-obovate

leaf blades are typical of this species whereas

specimens with broader or shorter leaves

(mostly from rocky situations) probably

represent a distinct form of it. As many

specimens are incomplete, more material,

especially specimens with flowers and/or fruit,

is necessary to ascertain this.

Etymology'. This species is named P. johnsonii

in honour of Dr Robert William (Bob) Johnson,

a former Director of the Queensland

Herbarium. He was particularly interested in

the ‘Canthiuiri’ species growing in association

with brigalow ( Acacia harpophylla ) while

studying the ecology of brigalow scrubs in

central Queensland over many years.

22. Psydrax forsteri S.T.Reynolds &

R.J.F.Hend., sp. nov. P. oleifoliae

(Hook.f.) S.T.Reynolds & R.J.F.Hend.

valde similis sed foliis minoribus,

inflorescentiis paucifloris minoribus in

pedunculis brevioribus portatis, floribus

5-meris differt. Typus: Queensland.

North Kennedy District: Yahoo

Waterhole, 500 m N of Warang

Homestead, White Mountains National

Park, 20°26’S, 144°49’E, 16 January 1995,

G. Anchen GA174 (holo: BRI; iso: BRI).

Canthium sp. (Duaringa N.H.Speck 1819),

S.T. Reynolds (1997, p.180), P.I. Forster

& D.A. Halford (2002, p.174).

Shrubs or small trees 2-4 m high; trunk with

bark greyish white or brown, smooth or rough;

branches usually at an angle to the main axis,

or more or less intricately branched; branchlets

terete, whitish coloured, somewhat reddish

distally when dry, glabrous. Leaves opposite;

stipules usually ovate, keeled, attenuated into

an apical lobe; petioles 0.2-0.4 cm long; blades

narrow obovate or obovate-elliptic, 2.0-2.8 cm

long x 1.1-1.3 cm wide, with apex obtuse,

slightly rounded or ± truncate and base acute

or obtuse, thick coriaceous; adaxial and abaxial

surfaces glabrous, the adaxial one glossy dull

green, the abaxial one pale green and dull;

lateral nerves in 1 or 2 pairs, obscure adaxially,

not apparent abaxially, the proximal pair arched

and ascending; reticulate venation not

apparent; domatia absent, or very rarely present

when 1 or 2 on each leaf and inconspicuous.

Inflorescences much shorter than the leaves,

1.0-1.7 cm long x 1.8-2.0 cm across, 5-17-

flowered, with basal peduncle 0.2-0.4 cm long,

terminated by 2 branched cymes and ciliated

ovate bracts; cymes 2-5-flowered. Flowers

5-merous, strongly perfumed; pedicel of solitary

flower 2.0-3.0 mm long, that of others 0.5-2.0

mm long; calyx 1.5-2.0 mm long, with tube ±

cupular and limb 5-toothed; corolla 6.0-7.0

(-8.0) mm long; tube ± campanulate, 2.5-3.0

mm long, densely hairy at orifice; lobes

sublanceolate, 3.0-3.5 mm long, subacute or

obtuse, erect or recurved; stamens with filaments

1.5-2.5 mm long, erect; anthers 2.0-2.5 mm

long, erect or ± patent; disc shorter than calyx

limb, glabrous; style (with stigma) 7.0-7.5 mm

long; stigma oblongoid, 2-lobed distally. Fruits

obovoid or broad ellipsoid, 0.5-0.7 cm long x

0.55-0.6 cm across; pyrenes depressed ovoid or

subellipsoid, ± deeply mgose on lateral sides, slightly

mgose distally. (Fig. 8F-8J)

Additional representative specimens: Queensland. Cook

District: Gilbert River, date unknown, Daintree

[MEL1538145] (MEL). Burke District: Warang Holding,

White Mountains, about 37 km NNW of Torrens Creek

Townships, 20°29’S, 144°49’E, Aug 1988, Fell 1378 &

Swain (BRI); 55 km NE of Hughenden, at Porcupine Gorge

National Park Lookout, 20°24’S, 144°26’E, Nov 1992,

Thompson 68 & Turpin (BRI). North Kennedy District:

Black Rock, 16 miles (c.25.6 km) S of Lynd Junction on

Hann Highway between Hughenden and Mt Garnet,

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

c. 19°05’S, c.144°21’E, May 1970, Webb & Tracey 10260

(BRI); Doongara, S of Homestead, SSE of Charters Towers,

20°33’S, 146°29’E, Nov 1986, Stuart 117 (BRI); ditto, Mar

1989, Forster PIF3725 & Bolton (BRI); Sally’s Mesa, 34

km from Greenvale on Charters Towers Road, 19°05’S,

145°14’E, Feb 1994, Forster PIF14972 & Bean (BRI).

South Kennedy District: 61 km N of Clermount, 22°14’S,

147°08’E, Dec 1986, Jackes 8622 (BRI). Mitchell District:

Enniskillen, 24°35’S, 146°05’E, Nov 1943, White 12358

(BRI, US). Leichhardt District: 7 miles (c.ll km) E of

Mantuan Downs, Tambo-Springsure Road, 24°25’S,

147°25’E, Apr 1946, Everist 2548 (BRI); 9.1 miles (c.14.6

km) W of Duaringa, Oct 1963, Speck 1819 (BRI, CANB, K,

NSW); Duaringa Shire, Berrigurra Station, about 20 km NW

of Blackwater, 23°32’S, 149°42’E, Nov 1983, Anderson

3604 (BRI, CANB, NSW); St Peter, NNW of Springsure,

Sep 1984, O’Keefe 750 (BRI); Novrindilla homestead, 25

km NE of Dingo, 23°30’S, 149°32’E, Oct 1989, Anderson

4803 (BRI); Humboldt, S of Blackwater, 24°13’S, 148°57’E,

Jan 1997, Fensham 3001 (BRI).

Distribution and habitat: North and central

Queensland; on sandy ridges, low hillsides,

sandstone slopes, rocky outcrops, in open

Eucalypt forests on red sandy loam. (Map 8)

879

Diagnostic attributes: Psydrax forsteri is

characterised by its comparatively small leaves

with usually obovate-elliptic, thick, more or less

dull, obscurely nerved leaf blades, and short

petioles, and its 5-17-flowered inflorescences

on exceedingly short peduncles. It differs from

other small-leaved species of Psydrax in

Australia by the above set of characters.

Notes: Psydrax for steri is related to P. oleifolia

and P. odorata subsp. buxifolia, and some of

the specimens cited above have previously been

included under the names Canthium oleifolium

or C. buxifolium in various herbaria. This

species may be distinguished from the latter

ones as follows:

1. Leaf blades exceedingly glossy on the adaxial surface; branchlets

and inflorescence axes hairy; corolla coriaceous; petioles stout

. P. odorata subsp. buxifolia

Leaves somewhat shiny or dull on the adaxial surface; branchlets and

inflorescence axes glabrous; corolla chartaceous; petioles slender. 2

2. Leaves with blades 2.0-2.8 cm long x 1.1-1.3 cm wide; inflorescence

with axis not branched, 5-17-flowered; flowers 5-merous; petioles

0.2-0.4 cm long. P. forsteri

Leaves with blades 3.5-6.5 (-8.5) cm long x 0.7-2.4 (-3.3) cm wide;

inflorescence with axis branched, 23-39-flowered, the flowers congested

on short branches; flowers mostly 4-merous; petioles 0.6-1.2 (-1.7) cm long .. P. oleifolia

The leaves of Psydrax forsteri are

variable. Specimens with usually comparatively

small, obovate-elliptic or narrow obovate leaf

blades are typical of this species. However,

specimens with narrower leaves, namely the

collections from Porcupine Gorge, White

Mountains and Sally’s Mesa cited above,

possibly represent a distinct form of it but more

material and field observations/notes would be

necessary to be certain of this.

Etymology: This species is named P. forsteri

in honour of Mr Paul Lorster, a botanist on the

staff of the Queensland Herbarium. His

collections of various ‘ Canthium ’ species from

throughout Queensland have been most

valuable in gaining an understanding of some

of the Psydrax species complexes. Most of the

illustrations provided in this paper are drawn

from his collections housed in BRI.

Acknowledgements

We thank Les Pedley and Peter Bostock for

assistance with the Latin diagnoses, colleagues

at BRI for collecting material of many of the

‘Canthium ’ species and for their comments

about the habitat of these species, Paul Blower

for his collections and photographs of P.

odorata from Vanuatu, Clyde Dunlop for his

comments on the manuscript, and Laurie

Jessup, Gordon Guymer and Clyde Dunlop for

their assistance in locating specimens and

photographing types and literature during their

term as Australian Botanical Liaison Officer

at Kew Gardens, England. Diane Bridson is

880

thanked for her comments on Psydrax and

Canthium , and for assistance with specimens

during visits to K (S.T.R. and, later, R.J.F.H.),

Will Smith for the illustrations and maps, David

Halford and Kym Sparshot for assistance with

measurements, maps and illustrations, and the

Directors/Keepers of herbaria AD, BM, CANB,

CGE, DNA, K, L, MEL, NSW, P, PERTH,

PVNH, QRS and UNSW for allowing me

(S.T.R.) full access to specimens in their

institutions and for the loan of herbarium

material. The Australian Biological Resource

Study, ABRS, Environment Australia, is

thanked for a grant (to S.T.R. in the 1990s) to

undertake research in the genus ‘Canthium’ in

Australia.

References

Bailey, F.M. (1900). Canthium Lam. Queensland Flora 3:

762-764. Brisbane: Queensland Government.

Bentham, G. (1867). Canthium, in Flora Australiensis 3:

420-423. Lovell Reeve & Co., London.

Bridson, D.M. (1985). The reinstatement of Psydrax

(Rubiaceae, subfamily Cinchonoideae, tribe

Vanguierieae ) and a revision of the African species.

Kew Bulletin 40(4): 687-725.

Cuparon, R. (1969). Apropos des Rubiacees Vangueriees

de Madagascar. Adansonia ser. 2, 9(1): 47-55.

Forster, RI. & Halford, D.A. (2002). Canthium, in R.J.F.

Henderson (ed), Names and Distribution of

Queensland Plants, Algae and Lichens, p.174.

Brisbane: Queensland Herbarium, Queensland

Government Environmental Protection Agency.

Austrobaileya 6 (4): 817-889 (2004)

Greuter, W., McNeil, J., Barrie, F.R., Burdet, H.M.,

Demoulin, V., Filgueiras, T.S., Nicolson, D.H.,

Silva, P.C., Skog, J.E., Trehane, R, Turland N.J. &

Hawksworth, D.L. (2000). International Code of

Botanical Nomenclature (St Louis Code).

Konigstein, Germany: Koeltz Scientific Books.

Holmgren, K., Holmgren, N.H. & Barnett, L.C. eds. (1990).

Index Herbariorum Ptl: The Herbaria of the World,

ed.8. RegnumVegetabile 120. New York, U.S.A.,

The New York Botanical Garden.

Lebler, B.A. (1978). The Canthiums of South-eastern

Queensland. Queensland Agricultural Journal 104

(6): 527-532. Brisbane: Government Printer.

Mabberley, D.J. (1997). The Plant-Book, Ed.2. Cambridge:

Cambridge University Press.

Mabberley, D.J. & Moore, D.T. (1999). Catalogue of the

holdings in The Natural History Museum (London)

of the Australian botanical drawings of Ferdinand

Bauer (1760-1826) and cognate materials relating

to the Investigator voyage of 1801-1805 Bull. nat.

Hist. Mus. London (Bot.) 29(2): 123,124.

Merrill, E.D. & Perry, L.M. (1945). Canthium. Plantae

PapuanaeArchboldianae 16. Journal of the Arnold

Arboretum 26(3): 230-233.

Mueller, F. (1875). Fragmenta 9: 185-186.

Reynolds, S.T. (1997). Canthium, in R.J.F. Henderson (ed),

Queensland Plants, Names and Distribution,

pp. 180-181. Brisbane: Queensland Herbarium,

Department of Environment.

Reynolds, S.T. & Henderson, R.J.F. (1999). Yanguerieae

A.Rich. ex Dum. (Rubiaceae) in Australia, 1.

Everistia S.T.Reynolds & R.J.F.Hend.

Austrobaileya 5(2): 353-361.

Reynolds, S.T. & Henderson, R.J.F. (2001). Vanguerieae

A.Rich. ex Dum. (Rubiaceae) in Australia, 2.

Cyclophyllum Hook.f. Austrobaileya 6(1): 41-66.

Supplement A

Currently accepted names for Australian taxa previously considered to belong in Canthium Lam.

or Plectronia L. are as follows. Numbers (and letters) where given with the currently accepted

name refer to the relevant position of the taxon concerned in the above revision. Accounts of

Everistia or Cyclophyllum taxa may be found in Reynolds & Henderson (1999 or 2001).

Previous identification Currently accepted name

Canthium attenuatum Benth.

Canthium attenuatum sensu G. Bentham (1867) pro parte

Canthium brevipes Merr. & L.M.Perry.

Canthium buxifolium Benth.

Canthium buxifolium forma (Brigooda P.LForster PIF5657)

. 15. Psydrax attenuata

.18. Psydrax pendulina

. 17. Psydrax saligna

.19. Psydrax oleifolia

.Cyclophyllum brevipes

6d. Psydrax odorata subsp. buxifolia

6d(i). Psydrax odorata forma buxifolia

. 6d(iii). Psydrax odorata subsp. buxifolia

‘Mundubbera form’

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

881

Canthium buxifolium var. (Mt Alford L.H.Bird AQ408941) . . 6d(ii). Psydrax odorata forma parviflorifra

Canthium coprosmoides F.Muell.Cyclophyllum coprosmoides

Canthium costatum C.T.White. Cyclophyllum costatum

Canthium cymosum sensu F. Mueller (1875,1882 &1889). 7b. Psydrax lamprophylla forma latissima

Canthium didymum sensu F. Mueller (1875) & F.M. Bailey (1900) pro parte. 7b. Psydrax lamprophylla

forma latissima

Canthium graciliflorum Merr. & L.M.Perry.2. Psydrax graciliflora

Canthium lamprophyllum F.Muell .7. Psydrax lamprophylla

Canthium latifolium F.Muell. ex Benth. 11. Psydrax latifolia

Canthium lineare E.Pritz. 3. Psydrax suaveolens

Canthium lucidum Hook. & Am., nom. illeg. non R.Br.6. Psydrax odorata

Canthium lucidum sensu G. Bentham (1867) pro parte. 7. Psydrax lamprophylla

Canthium microphyllum F.Muell.Everistia vacciniifolia var. nervosa

Canthium odoratum subsp. (DidcotP.I.ForsterPIF14127). 6c. Psydrax odorata subsp. australiana

Canthium odoratum forma (Texas P.I.Forster PIF14257). 6c(iii). Psydrax odorata forma subnitida

Canthium oleifolium Hook.f.19. Psydrax oleifolia

Canthium oleifolium sensu G. Bentham (1867) pro parte. 17. Psydrax saligna

Canthium oleifolium var. pedunculatumMaiden & Baker. 6c(iii). Psydrax odorata forma subnitida

Canthium schultzii (O. Schwarz) Chippendale. Cyclophyllum schultzii

Canthium sp. (Altanmoui Range D.G.Fell+DGF2702). 2. Psydrax graciliflora

Canthium sp. (Berrigurra Station E.R.Anderson 2829). 21. Psydrax johnsonii

Canthium sp. (CooktownV.Scarth-JohnsonAQ314008) . 16. Psydrax lepida

Canthium sp. (Cooroy S.T.Blake+ 15507) .Cyclophyllum longipetalum

Canthium sp. (Copper-Lode Falls C.H.Gittins 2211). Cyclophyllum protractum

Canthium sp. (Dixie Station S.T.Garnett 1545). 14. Psydrax pallida

Canthium sp. (DuaringaN.H.Speck 1819). 22. Psydrax forsteri

Canthium sp. (Friday Island L.J.Brass 18158). 10. Psydrax banksii

Canthium sp. (Headingly Station R.A.Perry 848) . 12. Psydrax ammophila

Canthium sp. (Herberton Range S.F.Kajewski 1377) . 8. Psydrax laxiflorens

Canthium sp. (Kuranda G.Sankowsky+ 680).Cyclophyllum multiflorum

Canthium sp. (Larcom Gully N.Gibson 1057). 20 Psydrax longipes

Canthium sp. (Lizard Island R.L.Specht+LI181). Cyclophyllum maritimum

Canthium sp. (Massy Creek P.I.Forster+PIF10568). Everistia vacciniifolia

Canthium sp. (Mt Alford L.H.Bird AQ408941). 6d(ii). Psydrax odorata forma parviflorifra

Canthium sp. (Mt Rose A.R.Bean 1978). Cyclophyllum rostellatum

Canthium sp. (ThorntonPeak H.FleckerNQNC7110). 5. Psydrax montigena

Canthium sp. (Thursday IslandE.Cowley 10). 13. Psydrax reticulata

Canthium sp. (Weipa, A.Morton AM 1773). 1. Psydrax paludosa

Canthium sp. (Wenlock River, J.R.Clarkson 8418+).1. Psydrax paludosa

Canthium sp. (Whitfield Range B.P.Hyland 1020) . 9. Psydrax tropica

Canthium sp.A, Koch, B.L. in J.R. Wheeler (1992) pro parte.18. Psydrax pendulina

Canthium sp. L, Ross, E.M. in T.D. Stanley & E.M. Ross (1986). 7. Psydrax lamprophylla

Canthium suaveolens S.Moore. 3. Psydrax suaveolens

Canthium vacciniifolium F.Muell.Everistia vacciniifolia

Plectronia attenuata (Benth.) C .A. Gardner. 15. Psydrax attenuata

Plectronia barbata sensu F. Mueller (1875).Cyclophyllum coprosmoides

Plectronia coprosmoides var. spathulata O.Schwarz. Cyclophyllum coprosmoides var. spathulatum

Plectronia diococca sensu F. Mueller (1875) pro parte.7b. Psydrax lamprophylla forma latissima

Plectronia hookeriana Domin. 6. Psydrax odorata

Plectronia hookeriana var. dietrichiae Domin. Tarenna sp.

Plectronia linearis (E.Pritz.) J.M.Black. 3. Psydrax suaveolens

Plectronia odorata var. reticulata C.T.White. 13. Psydrax reticulata

Plectronia schultzii O.Schwarz. Cyclophyllum schultzii

882

Austrobaileya 6 (4): 817-889 (2004)

Map L Distribution of Psydrax odorata subsp. australiana (40 Mile Scrub form) A , Psydrax

odor at a subsp, buxifolia (Mundubbcraform)* and Psydrax pa ludosa • ,

US 120 ]2S 130 135 NO 145 150 155

Map 2. Distribution of Psydrax graciliflora • and Psydrax suaveolensA ,

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

883

Map 3. Distribution of Psydrax odorata subsp australiana {P. odorata forma foveolata A ) and

Psydrax rigidula • ,

Map 4, Distribution of Psydrax odorata subsp. arnhemicaA and Psydrax odorata subsp.

australiana ( P. odorata forma australiana • ).

884

Austrobaileya 6 (4): 817-889 (2004)

.Map 5. Distribution of Psydrax laxijlorem ★and Psydrax odorata subsp. australiana (P

odorata forma subnitida A ).

.Map 6. Distribution of Psydrax montigena • , Psydrax odorata subsp, buxifolia {P odorata

forma buxifolia A ) and Psydrax reticulata *

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

885

Map 1. Distribution of Psydrax lamprophylla (P lamprophylla forma lamprophylla A and P.

lamprophylla forma latissima* ).

215 120 125 130 155 m 145 150 155

Map 8. Distribution of Psydrax banksii *, Psydrax forsteri • and Psydrax odorata subsp.

buxifolia (P odorata forma parvijlorijra A ).

886

Austrobaileya 6 (4): 817-889 (2004)

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

887

Map 11. Distribution of Psydrax attenuata var. attenuata A. , Psydrax attenuata var.

myrmecophila (P, attenuata forma megalantha ♦ and P attenuata forma myrmecophila + ) and

Psydrax attenuata var. tenella • .

Map 12. Distribution of Psydrax saligna (P. saligna fonna Jiliformis A and R saligna forma

saligna % ).

Austrobaileya 6 (4): 817-889 (2004)

Map 13, Distribution of Psydrax oleifoliaA and Psydrax pend id met + ,

Map 14, Distribution of Psydrax ammophila • , Psydrax johmonii ★ and Psydrax pallida A .

Reynolds & Henderson, Rubiaceae in Australia 3, Psydrax

Appendix 1

There are two species related to those dealt with

in the above account of Psydrax in Australia

which do not occur on this continent but which

are referred to in our account for comparison

purposes. As we consider these species also

belong in this genus but they lack a name under

Psydrax , we provide the following new

combinations for them.

Psydrax cymigera (Valeton) S.T.Reynolds &

R.J.F.Hend., comb, nov.; Plectronia

cymigera Valeton, Engl. Bot. Jahrb. 61:

54 (1927); Canthium cymigerum

(Valeton) B.L.Burtt, ( Kew) Bull. Misc.

Information 1936: 463 (1936). Type:

Northeast New Guinea. In den Walden

des Maboro, May 1909, Schlechter

n.17513 (719513) (holo: n.v.; iso: ?A.,

fide Merrill & Perry (1945), ?K, fide

B.L.Burtt, loc. cit.).

889

The situation regarding the numbering (or

possible mis-numbering) of parts of the type

collection has been commented upon by Merrill

and Perry (1945). Psydrax cymigera is related

to P lamprophylla.

Psydrax suborbicularis (C.T.White)

S.T.Reynolds & R.J.F.Hend., comb, nov.;

Plectronia suborbicularis C.T.White,

Proc. Linn. Soc. N.S. Wales 51: 296

(1926); Canthium suborbiculare

(C.T.White) Merrill & Perry, J. Arnold

Arb. 26: 230 (1945). Type: Papua New

Guinea. Rigo District, R. Lister Turner

(holo: BRI).

Psydrax suborbicularis is related to P. banksii

and P. reticulata.

A new species of Gynochtodes Blume (Rubiaceae) from north east

Queensland

D.A. Halford

Summary

Halford, D.A. (2004). Anew species of Gynochtodes Blume (Rubiaceae) from north east Queensland.

Austrobaileya 6 (4): 891-894. Gynochtodes sessilis Halford is described, illustrated and compared with

Gynochtodes australiensis J.T. Johanss. Notes on habitat and distribution are provided.

Key words: Gynochtodes , taxonomy, Australian flora, Gynochtodes sessilis, Rubiaceae

D.A. Halford, c/- Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens Mt

Coot-tha, Mt Coot-tha Road, Toowong, Queensland 4066, Australia

Introduction

Johansson (1988) reported the genus

Gynochtodes Blume from Australia for the first

time with the description of G. australiensis

J.T.Johanss. based on material from the

headwaters of East Alligator River in the

Northern Territory. In his circumscription of

this species he included two collections from

north eastern Queensland, namely A.S. & M.G.

Thorsborne 337 (BRI) collected from

Hinchinbrook Island and V. Scarth-Johnson

1109A (BRI) collected from near Bamaga. In

regards to the collection from Hinchinbrook

Island, he noted that it differed from the

Northern Territory collections by having longer

acuminate leaf-tips and more compressed

inflorescences. However, Johansson considered

that there was insufficient material to draw any

taxonomic conclusion.

As a result of the examination of a number

of fertile collections that are now available at

the Queensland Herbarium (BRI) from north

east Queensland I believe that the taxon

represented by the collection (A.S. & M.G.

Thorsborne 337) is specifically distinct from

G. australiensis and is here named Gynochtodes

sessilis.

The collection from Bamaga, V. Scarth-

Johnson 1109A, is here maintained under the

name G. australiensis as it has inflorescences

that are shortly pedunculate. However, it differs

from Johansson’s circumscription of that

species by its generally shorter and broader

leaves, its sparse to moderately dense

Accepted for publication 14 May 2004

indumentum of erect unicellular hairs (< 0.1 mm

long) on young branchlets and is reportedly a

tree (collectors notes). This population warrants

further study to establish the significance of

these differences.

Measurements of flowers and fruit in this

account are taken from material preserved in

spirit at BRI. This work is part of current studies

being undertaken by the author into the tribe

Morindeae (Rubiaceace) in Queensland.

Gynochtodes sessilis Halford sp. nov. arete

affinis G. australiensi autem floribus

fructibusque sessilibus, foliis plerumque

grandioribus (9.5-19 x 3-6 (-7.5) cm

comparitis 5-13 x 1.6-5 cm) apicibus

longioribus acuminatis instructis,

calycibus pubescentibus non glabris

differt. A G. coriacea Bl. floribus

fructibusque sessilibus, floribus

comparate grandioribus, calycibus

pubescentibus domatiis carentibus

distinguenda. Typus: Queensland. Cook

District: State Forest 607 Dinden, Bridle

Creek track, 15 Jan 2002, P.I. Forster

PIF28124 R. Booth & R. Jensen (holo:

BRI; iso: A, BISH, DNA, F, MEF, MO,

P, Z, distribuendi).

Gynochtodes sp. (Hinchinbrook Island

A.S.Thorsborne+ 337), (Forster &

Halford 2002).

Woody liana , becoming somewhat shrubby in

open situations. Young branchlets terete,

glabrous, smooth. Stems blackish in colour with

wart-like lenticels. Bark and sap wood becoming

purple to dark violet when cut. Raphides

present. Stipules interpetiolar, sheathing, with

892

Austrobaileya 6(4): 891-894 (2004)

short triangular lobe, 1-2 mm long,

chartaceous, glabrous except for minute hairs

(< 0.2 mm long) on margin, fragmenting as

node thickens. Leaves decussate, glabrous,

usually drying black; petioles plano-convex, 1-2

cm long; laminae narrowly obovate to obovate

or narrowly elliptic to elliptic, 9.5-19 cm long,

3-6 (-7.5) cm wide, length/width ratio 2-3:1,

coriaceous, glossy dark green adaxially and

paler abaxially when fresh, turning + black

when dried; base obtuse to broadly cuneate;

margins entire; apex obtuse or usually

acuminate with acumen up to 1 cm long;

venation brochidodromous, with midrib raised

slightly adaxially and strongly abaxially when

dried; lateral, intramarginal and interlateral

veins slightly raised on both surfaces when

dried; lateral veins 7-10 per side of midrib, at

c. 60-70° to the midrib; intramarginal veins

2-4 mm from margin; interlateral veins

reticulate; domatia absent. Flowers unisexual

(?), axillary, fasciculate, with up to 12 in each

cluster, sessile; calyx cup-shaped, c. 2 x 2 mm,

green, sparsely hairy with minute erect hairs

(< 0.05 mm long); limb c. 0.5 mm long,

glabrous adaxially, truncate, with minute erect

hairs on margin; corolla cream-yellow or white

when fresh, turning black when dried, sparsely

hairy abaxially with minute erect hairs; corolla

tube ± cylindrical, 1-2.5 mm long, glabrous

abaxially except for scattered minute erect hairs

proximally; corolla lobes (3) 4 (5), narrowly

obovate, 6-7 mm long, 1.5-3.3 mm wide,

densely villose for the proximal two thirds

adaxially, ± cucullate at apex; stamens 4,

exserted, inserted at corolla lobe sinuses;

filaments terete, 2-3 mm long, glabrous;

anthers oblong c. 2.7 mm long; ovary bilocular,

ovules 2 in each locule, with false septum in

the upper part appearing to divide each into 2,

funicule inserted at base of dissepiment; style

absent; stigma bifid, c. 0.2 mm long. Fruit a

drupe, globose to ellipsoid, 12-14 mm long,

9-14 mm diameter, 1-4-seeded, glabrous;

epicarp, black, brittle; mesocarp black, pulpy

with pungent aroma. Fig. 1.

Selected specimens: Queensland: Cook District. Mt Cook

NP, 0.5 km NE of summit, Feb 1993, D.G. Fell DGF2858 &

J.P. Stanton (BRI); State Forest 607 Dinden, Bridle Creek

track, Jan 2002, P.I. Forster PIF28127, R. Booth & R. Jensen

(BRI); NPR 164, Parish of Noah, Oliver Creek, Jun 1987, B.

Gray 04483 (BRI); SFR 607, Shoteel logging area, May

1982, B. Gray 20203v (BRI); Cairns, Hills Creek, Jan 1994,

C. Lyons 152 (BRI). North Kennedy District. SFR 299,

Conway, Cedar Creek, May 1975, B. Hyland 08238 (BRI);

Hinchinbrook Island, Dec 1976, A.S. & M.G. Thorsborne

337 (BRI); upper reach of Deluge Inlet, Hinchinbrook Island,

May 1972, L.J. Webb & J.G. Tracey 11186 (BRI).

Distribution and habitat : Occurs in north east

Queensland, where it has been collected in

coastal and subcoastal areas from Mt Cook

National Park near Cooktown southwards to

Hinchinbrook Island, with a disjunct population

in the Conway Range near Proserpine (Map 1).

It is commonly recorded growing from near sea-

level to 520 m altitude in simple or complex

notophyll rainforest mostly on alluvial soils

derived from various substrates.

Phenology: Flowers have been recorded from

June to November, fruits from April to June

and August.

Affinities: Gynochtodes sessilis is closely

related to G. australiensis but differs from that

by its sessile flowers and fruits, its generally

larger leaves (9.5-19 x 3-6 (-7.5) cm compared

with 5-13 x 1.6-5 cm), its longer acuminate

leaf apices and its hairy rather that glabrous

calyces. Gynochtodes sessilis can be

distinguished from G coriacea Blume by its

sessile flowers and fruits, comparatively larger

flowers, hairy calyces and lack of domatia.

Notes: All flowering collections (C. Lyon 152;

A.S. & M.G. Thorsborne 337; P.I. Forster

PIF28124, R. Booth & R. Jensen; P.I. Forster

PIF28127, R. Booth & R. Jensen ) of G. sessilis

that have been examined during this study are

tentatively interpreted as having unisexual

flowers as they lacked a well developed style

and stigma. However, the ovaries are well

developed with what appears to be functional

ovules. Further flowering material is required

to assess what reproductive strategies are

present in this species.

The flowers of G. sessilis are reported to be

either sweetly scented (P.I. Forster PIF28124,

R. Booth & R. Jensen) or to have a pungent

aroma (C. Lyons 152) at anthesis.

Etymology: The specific epithet is Latin and

refers to the sessile flowers and fruits of the

species.

Halford, Anew species of Gynochtodes

893

QUEENSLAND HERBARIUM iBRl)

Brisbane Australia.

QUEENSLAND HERBARtUM (BRI)

Ffepa ctOuMMlind Ox*

Gynochtodes sp. (Hlttchlnbroolr Inland

A.S.Tliorsborno* 1J7|

HOtOTYFE

&TK)dilwtciK5silis Halford

l)il. LI X tblmjiEWIl

Figure 1. Photograph of holotype of Gynochtodes sessilis Halford.

894

Austrobaileya 6 (4): 891-894 (2004)

Acknowledgements

I would like to thank Les Pedley for the

translation of the diagnosis into Latin and

Gordon Guymer, Director of BRI, for making

available working space and facilities at BRI.

References

Forster, P.I. & Halford, D.A. (2002). Rubiaceae. In R.J.F.

Henderson (ed.), ‘Names and Distribution of

Queensland Plants, Algae and Lichens.’ pp. 173—

177. (Environmental Protection Agency: Brisbane).

Johansson, J.T. (1988). A new species of Gynochtodes

(Rubiaceae, Rubioideae) from Australia. Australian

Systematic Botany 1(4): 369-372.

Two new species of Morinda L. (Rubiaceae) from north east Queensland

D.A. Halford and A.J. Ford

Summary

Halford, D. A. & Ford, A. J. (2004). Two new species of Morinda L. (Rubiaceae) from north east Queensland.

Austrobaileya 6(4): 895-902. Morinda podistra and M. ammitia are described, illustrated and diagnosed

against related species. Notes on habitat and distribution are provided for each species.

Key words: Morinda, taxonomy, Australian flora, Morinda podistra, Morinda ammitia, Rubiaceae

D.A. Halford, Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens Mt

Coot-tha, Mt Coot-tha Road, Toowong, Queensland 4066, Australia

A.J. Ford, CSIRO, Sustainable Ecosystems and R ai nforest. CRC, Tropical Forest Research Centre, PO Box

780, Atherton, Queensland 4883, Australia

Introduction

The genus Morinda L. comprises approximately

80 species mostly in the Old World tropics

(Mabberley 1997). Currently six named species

of Morinda are recorded in Australia, namely

M. bracteata var. celebica (Miq.) Valeton,

M. canthoides (F.Muell.) Halford & RJ.F.Hend.,

M. citrifolia L., M. jasminoides A.Cunn. ex Benth.,

M. reticulata Benth. and M. umbellata L. In

this paper we describe two new species endemic

to north east Queensland.

Methods

The study is based upon the examination of

herbarium material from BRI and QRS with

field observations by the second author of both

new species. The Herbarium acronyms follow

Holmgren et al. (1990). All specimens cited

have been seen by one or both authors.

Measurement of floral parts are based on

material preserved in 70% ethanol or dried

material reconstituted by placing in boiling

water for a few minutes. The compound fruit

formed by the fusion of a number of ovaries in

species of Morinda as well as in some other

genera of the tribe Morindeae has been

described by the term syncarp by many authors

(e.g. Johansson 1987, Robbrecht 1988,

Johansson 1988). Igersheim and Robbrecht

(1993) have pointed out that this is an

inappropriate term and should be avoided. In

this paper the term “compound syncarpous

drupe” is used to describe the compound fruit.

Accepted for publication 29 March 2004

The term “drupe” is used here as defined by

Clifford and Dettmann (2001).

Taxonomy

Morinda podistra Halford & A. J.Ford, sp. nov.

M. umbellatae similis autem foliis

maturis apice acuminato plus extenso

praeditis et indumento pilis antrorsis in

ramulosis, in pedunculis et in pagina

inferiore folii vestita differt. Typus:

Queensland. Cook District: Daintree

River National Park, Black Mountain

area, Daintree River headwaters, 25 May

1998, P.I. Forster PIF22959, R. Jago, R.

Jensen & R. Booth (holo: BRI).

Morinda sp. (Mt Lewis L.W.Jessup+

GJM228), Forster and Halford (2002).

Twining vine, to 4m high, or rarely a scandent

shrub. Stem and main branches slender, ±

smooth to longitudinally striated, 3-6 mm

diameter. Branchlets terete, scabrid, moderately

dense to densely hairy, glabrescent; hairs

simple, antrorse, ascending to spreading, up to

0.6 mm long. Leaves petiolate, opposite;

stipules interpetiolar, sheathing, 1-5 mm long,

truncate at apex, moderately dense to densely

hairy with hairs as for branchlets; petioles 1-5

mm long; laminae + chartaceous, narrowly

obovate-elliptic or narrowly elliptic, 2.5-10 cm

long, 0.7-3 cm wide; base attenuate; margins

entire, slightly recurved (when dry); apex

acuminate; adaxial surface usually glabrous and

smooth or rarely with scattered, short, antrorse

hairs and then scabrid; abaxial surfaces sparsely

to moderately hairy especially along midvein

896

Austrobaileya 6 (4): 895-902 (2004)

and secondary veins, scabrous; hairs simple,

antrorse, appressed to ascending, up to 0.9 mm

long; venation brochidodromus with 4-8 lateral

veins per side of midvein, slightly raised on

adaxial surface, prominent on abaxial surface;

midvein conspicuously raised on adaxial

surface, prominent on abaxial surface; tuft-

domatia present in lateral vein axils on abaxial

surface. Flowers usually 4-merous (rarely 3 or

5-merous), bisexual or unisexual (?), sessile,

in congested capitula, the flowers joined at least

by the base of the gynoecium. Capitula

pedunculate in terminal umbels with 2-4 (often

2) capitula per umbel, ± globose, 3—4 mm across

(excluding corollas), 6-12 flowered, with

colleters present between flowers; peduncles

2- 10 (-15) mm long, moderately dense to

densely hairy with hairs as for branchlets. Calyx

tube green, 0.6-0.9 mm long, c. 1.5 mm across,

glabrous to minutely and sparsely hairy

abaxially and glabrous adaxially, truncate.

Corolla valvate, deciduous, cream, turning

creamy yellow with age, sparsely to densely

hispidulous on abaxial surface with hairs up to

0.1 mm long; tube 1.3-2 mm long, the middle

of the tube is fenestrated by short longitudinal

slits, slightly widened at the apex, glabrous

adaxially but densely hairy at mouth; hairs

simple, up to 0.8 mm long; lobes spreading,

ovate, 1.8-2.2mmlong, 1.1-1.7 mm wide, with

a small subapical appendage on abaxial surface,

sparsely to densely hairy adaxially, acute and

+ cucullate at apex. Stamens exserted; filament

0.5-1 mm long, inserted c. 0.3 mm below the

sinuses of the corolla lobes; anthers dorsifixed,

linear-oblong, 1.0-1.3 mm long, dehiscing

laterally. Disc entire, convex, c. 0.3 mm high,

glabrous. Ovary 2-celled, biovulate, with false

septum in the upper part appearing to divide

each cell into 2; style c. 1 mm long, glabrous;

stigma bifid, with erect lobes 0.3-0.8 mm long,

adaxial surface papillate, abaxial surface with

minute, scattered, antrorse hairs. Fruit a

compound syncarpous drupe, orange when ripe,

subglobose, 8-10 mm across, persistent calyx

tubes prominent on surface, minutely and

sparsely hairy; colleters enlarged and fleshy;

mesocarp fleshy, containing several pyrenes;

pyrenes + ellipsoid, dorsiventrally compressed,

3- 3.5 mm long, 2.5-3mm wide, c. 1.7 mm

thick, 1-seeded; endocarp cartilagineous,

brown, ± rugose, with basal marginal groove.

Seeds irregularly shaped but usually 3-faced,

c. 2.8 mm long, c. 1.7 mm wide, c. 1.2 mm

thick; testa membraneous, dark brown;

endosperm starchy and hard; embryo c. 1 mm

long, straight; cotyledons very thin, about the

same length as, but slightly wider than, the

radicle. Fig. 1.

Additional specimens examined : Queensland. Cook

District, slopes of Mt Lewis, Feb 1932, Brass 2083 (BRI);

Mt Lewis, a few km down from hut, Mt Lewis Forest Reserve,

Jan 2002, Cooper WWC1659 & Ford (BRI); Mt Lewis

Forest Reserve, 26 km along Mt Lewis road, near creek down

from hut, Dec 2001, Cooper WWC1632 et al. (BRI, QRS);

edge of Zarda Camp Clearing Upper Mossman River, Sep

1936, Flecker NQNC2331 (QRS); Mt Lewis FR, 1 km E of

peak “1243”, Dec 2002, Ford 3782 & Holmes (BRI); ditto.

Ford 3784 & Holmes (BRI); summit of Devil’s Thumb,

Mossman Gorge NP, Apr 1986, Godwin C3091 (BRI); SFR

143, North Mary logging area, Dec 1977, Gray 828 (QRS);

Mt Lewis, Sep 2002, Halford Q7400 & Ford (BRI); SFR

143, North Mary logging area, Dec 1974, Hyland 7909 (BRI,

QRS); SFR 143, North Mary logging area, Nov 1974, Hyland

7887 (QRS); SFR 143, Nov 1974, Irvine 1066 (BRI, QRS);

Mt Lewis Rd, S Mary logging area, 16 km NNW [of] Mt

Molloy, Nov 1988, Jessup GJM1472 etal. (BRI); Mt Lewis

Rd near Half Ton Creek crossing, about 30 km from Rex

Highway, Nov 1988, Jessup GJM29 etal. (BRI); Mt Misery

E of Mt Spurgeon, 15.4 km NNE [of] Mt Carbine, Nov 1988,

Jessup GJM971 et al. (BRI); Mt Lewis Rd, 28 km from Mt

Molloy - Mossman Rd, Jan 1981, Jessup 282 & Clarkson

(BRI); Mt Lewis Rd boundary of Round logging area and

Carbine logging area, 20 km NNW [of] Mt Molloy, Nov

1988, Jessup GJM288 etal. (BRI); Mossman Coast Range,

Apr 1992, Russell 28 (BRI); Near Schillers Hut, Mt

Spurgeon, Sep 1972, Webb & Tracey 11799 (BRI); MtLewis,

Oct 1971, Webb & Tracey 10510 (BRI).

Distribution and habitat : Morinda podistra is

endemic to north east Queensland, where it is

currently known from the Mt Lewis-Mt

Spurgeon area, west of Mossman, on the Mount

Carbine Tableland and the Main Coast Range

(Map 1). It is recorded as growing in simple

and complex notophyll vine forest, and simple

microphyll vine-fern thicket communities on

soils derived from granite substrates at altitudes

of 1000 to 1330 m. Morinda podistra appears

to be more common in the microphyll vine-

fern thicket communities amongst granite

boulders, which have a low and broken canopy,

than in the notophyll vine forests. Common

canopy trees include: Sphalmium racemosum

(C.T. White) B.G.Briggs, B.Hyland &

L. A. S.Johnson, Syzygium endophloium

B.Hyland, Garcinia brassii C.T.White,

Flindersia pimenteliana F.Muell. f.

pimenteliana, Niemeyera sp. (Mt Lewis

A. K.Irvine 1402), Balanops australiana

Halford & Ford, two new species of Morinda

897

Fig. 1. Morindapodistra. A. branchlet with flowers x 1. B. branchlet node with stipules and leaf bases x 6. C. inflorescence x 8.

D. compound fruit x 4. E. adaxial view of pyrene x 12. F. lateral view of pyrene x 12. A from Ford 3782 & Holmes (BRI); B

from Halford Q7400 & Ford (BRI); C from Ford 3784 & Holmes (BRI); D-F from Ford 3871 & Holmes (BRI). Del. W. Smith.

898

F.Muell., Halfordia kendack (Montrouz.)

Guillaumin and Uromyrtus metrosideros

(F.M.Bailey) A.J.Scott. Common shrubs and

understorey plants include: Chionanthus

axillaris R.Br., Mischocarpuspyriformis subsp.

retusus (Radik.) R.W.Ham, Alyxia orophila

Domin, Agapetes meiniana F.Muell.

Oraniopsis appendiculata (F.M.Bailey)

J.Dransf., A.K.Irvine & N.W.Uhl, Rapanea sp.

(Zarda LA B.RHyland 6898) and Lomandra

hystrix (R.Br.) L.R.Fraser & Vickery.

Phenology: Flowers have been recorded from

November to January whilst fruits have been

recorded in April and May.

Affinities: Morinda podistra resembles

M. umbellata, but differs from that species by

its mature leaves that have a longer drawn out

acuminate apex and the antrorse hairs that are

present on the branchlets, peduncles and

abaxial leaf surface. Morinda podistra and M.

umbellata are known to co-exist in the Mt

Lewis area.

Notes: Morinda podistra is commonly observed

as growing on soil substrate. However, it has

also been observed as a lithophyte, where it

produces adventitious roots along the stem.

The new leaf growth of M. podistra is

pale green. Expanding stipules, recently

produced nodes, petioles, twigs and peduncles

are usually purplish in colour. Flower bud

colour has been recorded as being pinkish, or

purplish at the base of the corolla.

The flowers from the collections Ford

AF3784 (BRI), Jessup GJM228 et al. (BRI)

and Jessup GJM971 etal. (BRI) are tentatively

interpreted as being unisexual as they have

pollen producing anthers but lack a style of any

description, although the ovary is well

developed with what appear to be functional

ovules. The flowers from collections Gray 828

(QRS), Cooper WWC1632 etal (BRI), Jessup

GJM771 etal (BRI),Anon. [AQ456135] (BRI)

and Irvine 1066 (BRI) appear to be bisexual

with a well developed style, stigma, ovary and

pollen producing anthers. Further flowering

material and field investigations are required

to assess what reproductive strategies are

present in this species.

Austrobaileya 6 (4): 895-902 (2004)

Conservation status: Morinda podistra has an

apparently restricted and narrow distribution.

However it is locally common within its habitat

in this range. All of the cited specimens have

been collected from within the World Heritage

Area of the Wet Tropics bioregion. Also, a

number of collections have been made in the

Daintree National Park. It is not considered at

risk at this time.

Etymology: The specific epithet podistra is

Greek meaning ‘foot trap’. As intrepid

bushwalkers will testify, the wiry stems of this

species often form dense stands in which

backpacks and feet get caught.

Morinda ammitia Halford & A.J.Ford, sp.

nov. M. bracteatae var. celebicae

nonnihil similis autem caulibus

pubescentibus, foliis pubescentibus,

fructibus pubescentibus (glabris in M.

bracetata var. celebica ), corollae tubis

brevioribus (1.5-3.5 mm vice c. 10 mm

in M. bracteata var. celebica ), habitu

scandenti (habitu fruticis usque arboris

parvae in M. bracteata var. celebica) et

lobis calycis foliaceis carentibus differt.

Typus: Queensland. Cook District:

Cooktown - Isabella Road, c. 200 m E of

Isabella Falls, 13 March 2001, A. Ford

AF2697 & J. Holmes (holo: BRI; iso:

DNA, NSW distribuendi).

Morinda sp. (Altanmoui Range B.P.Hyland

6323), Forster and Halford (2002).

Vine or scandent shrub with long arching, non¬

twining, stems. Stem and main branches not

seen. Branchlets + quadrangular when young

becoming + terete and striated with age,

moderately dense to densely hairy; hairs simple,

ascending to spreading, 0.1-0.7 mm long.

Feaves petiolate, opposite; stipules

interpetiolar, sheathing, 1-2 mm long,

produced into a narrow triangular lobe up to 2

mm long, hairs as for branchlets; petioles 2-5

mm long; laminae + coriaceous, narrowly

obovate-elliptic, narrowly elliptic to elliptic,

4.5-9 cm long, 2.5-5.6 cm wide; base cuneate;

margins entire, slightly recurved; apex obtuse

to rounded, acute or sometimes shortly

acuminate; adaxial and abaxial surfaces

glabrous, scabrid or sparsely to moderately

hirtellous, indumentum usually slightly denser

Halford & Ford, two new species of Morinda

899

on abaxial surface; hairs simple, spreading to

ascending, up to 0.8 mm long; venation

brochidodromus with 5-10 lateral veins per side

of midvein, slightly raised on adaxial surface,

prominent on abaxial surface; midvein slightly

raised on adaxial surface, prominent on abaxial

surface; tuft-domatia present in lateral vein

axils on abaxial surface. Flowers 4 or 5-merous,

bisexual or unisexual (?), sessile, in congested

capitula, the flowers joined at least by the base

of the gynoecium. Capitula pedunculate in

terminal umbels with 2-4 capitula per umbel

or solitary in upper leaf axils, rarely a solitary

sessile flower or a rudimentary capitulum

present at base of peduncles, + globose, 8-12

mm across (excluding corollas), 12-30-

flowered with colleters absent between flowers;

peduncles 10-23 mm long, (elongating to 30

mm in fruit), terete, often striated, moderately

dense to densely hairy with hairs as for

branchlets. Calyx tube yellow-green, c. 2 mm

long, c. 2.5 mm across, moderately to densely

hispidulous adaxially with simple hairs up to

0.2 mm long, densely hairy abaxially with

appressed to spreading simple hairs up to 0.3

mm long, truncate, undulate or irregularly and

shallowly toothed. Corolla valvate, deciduous,

greenish cream to white, hispidulous on abaxial

surface; tube 1.5-3.5 mm long, the middle of

the tube is fenestrated by short longitudinal

slits, slightly widened at the apex, glabrous

adaxially but densely hairy towards mouth;

hairs simple, up to 1 mm long; lobes spreading

and recurved distally, narrowly triangular to

triangular, 3-4 mm long, 1.5-2 mm wide, acute

and ± cucullate at apex, sparsely to densely

hairy adaxially, glabrous towards apex.

Stamens exserted; filament 0.5-1.7 mm long,

inserted c. 0.5 mm below the sinuses of the

corolla lobes; anthers, dorsifixed, linear-oblong,

1.3-2 mm long, minutely hairy, dehiscing

laterally. Disc entire, flat, c. 0.5 mm high,

sparsely covered with minute simple hairs.

Ovary 2-celled, biovulate, with false septum in

the upper part appearing to divide each cell into

2; style 0.8-1.5 mm long, included in corolla

tube, glabrous; stigma bifid, with erect lobes

c.1.5 mm long, adaxial surface papillate,

abaxial surface glabrous. Fruit a compound

syncarpous drupe, black when ripe, subglobose,

20-30 mm across, hispidulous, persistent calyx

tubes prominent on surface, mesocarp fleshy,

containing several pyrenes; pyrenes ± ellipsoid

or obovoid, dorsiventrally compressed, 5-6 mm

long, 2.5-4mm wide, 1.8-2.2 mm thick,

1-seeded; endocarp cartilagineous, pale brown,

± smooth, with basal marginal groove. Seeds

c. 4 mm long, c. 2 mm wide, c. 1.6 mm thick;

testa membraneous, dark brown; endosperm

corneous, + white; embryo (most seeds lacking

an embryo) c. 2mm long, straight, surrounded

by a sticky and gelatinous membrane;

cotyledons thin, shorter than but slightly wider

than the radicle. Fig. 2.

Additional specimens examined : Queensland. Cook

District. Turtle Rock, 12 km SSE of Laura, Nov 1991, Bean

3803 (BRI); Isabella Creek, about 27 miles [c. 43 km] NW

of Cooktown, Jun 1968, Brass 33836 (QRS); Laura

sandstone area N of Laura River near Early Man site, May

1975, Byrnes 3312 (BRI); Split Rock Gallery, 14 km S of

Laura on the Peninsula Development Rd, Oct 1984, Clarkson

5635A (BRI, QRS); Cape Melville NP, Altanmoui Range

section, 7 km SE of Wakooka Outstation, Oct 1992, Fell

DGL2703 & Stanton (BRI, QRS); Split Rock Gallery, 14

km S of Laura on the Peninsula Development Rd, Jul 1994,

Gray 5764 (QRS); Isabella - Mclvor road, 200 m from

Isabella Palls, NW of Cooktown, Nov 2002, Ford 3719 &

Holmes (BRI); Gugu Yalangi (Laura), Sep 1976, Hyland

9041 (BRI, QRS); Altanmoui, Jul 1972, Hyland 6323 (BRI,

QRS); Cape Melville, Sep 1970, Hyland 4660 (BRI); 16

km SE of Laura, Jun 1976, Jackes [AQ168531] (BRI);

between Battle Camp and Isabella Ck on Laura Station -

Cooktown Rd, Nov 1979, Webb & Tracey 13411 (BRI,

QRS); 9 miles [c. 14 km] S of Laura on main Mareeba -

Coen Rd, Oct 1969, Webb & Tracey 9917A (BRI); Laura

Galleries, a few km S of Laura on Mareeba - Coen Rd, Nov

1979, Webb & Tracey 13395 (BRI); Kennedy River, Aug

1966, VolckAFO3303 (QRS).

Distribution and habitat : Morinda ammitia is

endemic to north east Queensland, where it is

known from Cape Melville to the Cooktown -

Laura area (Map 1). It is recorded as growing

in open forest and low woodland communities,

at the base of sandstone cliffs, in deciduous vine

thicket, in low open eucalypt woodland and

heath-like communities on sandstone

escarpments. Associated dominant trees

include: Corymbia hylandii (D.J.Carr &

S.G.M.Carr) K.D.Hill & L.A.S.Johnson,

Eucalyptus phoenicea F.Muell., Eucalyptus

crebra F.Muell. and Blepharocarya

involucrigera F.Muell. Large shrubs include:

Acacia flavescens A.Cunn. ex Benth., Parinari

nonda F.Muell. ex Benth. and Erythrophleum

chlorostachys (F.Muell.) Baill. Small shrubs

include: Alyxia spicata R.Br., Hibbertia banksii

(R.Br. ex DC.) Benth. and Phyllanthus/

Sauropus spp. Conspicuous understorey genera

include: Xanthorrhoea, Pandanus and

Heteropogon.

900

Austrobaileya 6 (4): 895-902 (2004)

Fig. 2. Morinda ammitia . A. branchlet with flowers x 0.8. B. branchlet node with stipules and leaf bases x 4. C. inflorescence x 6.

D. flower viewed from above x 8. E. compound fruit x 2. F. adaxial view of pyrene x 6. G. lateral view of pyrene x 6. A-D from

Ford 3719 & Holmes (BRI); E-G from Ford AF2697 & Holmes (BRI). Del. W. Smith.

Halford & Ford, two new species of Morinda

901

Phenology : Flowers have been recorded from

June to December, whilst fruits have been

recorded in March, May and July.

Affinities: Morinda ammitia is somewhat

similar to M. bracteata var. celebica but differs

from that species in its hairy stems, leaves and

fruits; (glabrous forM. bracteata var. celebica)',

shorter corolla tubes (1.5-3.5 mm long

compared with c. 10 mm long forM. bracteata

var. celebica ); scandent habit (shrub to small

tree for M. bracteata var. celebica)', and the

lack of foliaceous calyx lobes.

Notes: Flower buds have been recorded as

having yellowish corolla lobes. The flowers

from collections Hyland 6323 (QRS), Clarkson

5635A (BRI), Bean 3803 (BRI) and Fell &

Stanton DGF2703 are tentatively interpreted

as being unisexual as they have pollen

producing anthers but lack a style of any

description, although the ovary is well

developed with what appear to be functional

ovules. The summit of the ovary has an orifice,

as does the bisexual flower after the style has

shed. The flowers from collections Ford 3719

(BRI, QRS) and Ford 3720 (BRI, QRS) appear

to be bisexual with a well developed style,

stigma, ovary and pollen producing anthers.

Further flowering material and field

investigations are required to assess what

reproductive strategies are present in this

species.

Conservation status: Morinda ammitia is

assessed as data deficient. Three collections

have been made from Cape Melville National

Park while other collections are from

populations outside of conservation reserves.

Etymology: The specific epithet is from the

Greek ammites, sandstone; in reference to the

species occurrence on sandstone substrates.

Acknowledgements

The authors wish to thank Will Smith for the

illustrations, Peter Bostock for the maps, Les

Pedley for the translation of the diagnoses into

Latin and suggestions regarding the specific

epithet “ ammitia ”. The first author would like

to thank Gordon Guymer, Director of BRI, for

making available working space and facilities

at BRI. Permits to traverse and collect in the

Mount Lewis Forest Reserve were issued by the

Queensland Parks and Wildlife Service. Jenny

Holmes (Atherton) provided invaluable field

assistance.

References

Clifford, H.T. & Dettmann, M.E. (2001). Drupe - a term in

search of a defi ni tion. Austrobaileya 6(1): 127-131.

Forster, P.I. & Halford, D. (2002). Rubiaceae. In R.J.F.

Henderson (ed.), Names and Distribution of

Queensland Plants, Algae and Lichens, 68-74.

Brisbane: Environmental Protection Agency.

Holmgren, P.K., Holmgren, N.H. & Barnett, L.C. (1990).

Index Herbariorum. Part 1. The Herbaria of the

World. 8 th edn. New York: New York Botanic

Gardens.

Igersheim, A. & Robbrecht, E. (1993). The character state

and relationships of the Prismatomerideae

(Rubiaceae - Rubioideae): comparisons with

Morinda and comments on the circumscription of

the Morindeae s.str. Opera Botanica Belgica 6:61-

79.

Johansson, J.T. (1987). Motleyia, a new genus of the

Rubiaceae from Borneo. Blumea 32: 149-155.

-(1988). Revision of the Genus Caelospermum Blume

(Rubiaceae, Rubioideae, Morindeae). Blumea

33(2): 265-297.

Mabberley, D.J. (1997). The Plant-Book: A portable

Dictionary of Higher Plants. 2 nd edn. Cambridge:

Cambridge University Press.

Robbrecht, E. (1988). Tropical woody Rubiaceae. Opera

Botanica Belgica 1: 1-27 E

902

Austrobaileya 6 (4): 895-902 (2004)

Map 1. Distribution of Morinda podistra% and Morinda ammitia A .

The tribe Coffeeae DC. (Rubiaceae: Ixoroideae) in Australia

Paul I. Forster

Summary

Forster, P.I. (2004). The tribe Coffeeae DC. (Rubiaceae: Ixoroideae) in Australia. Austrobaileya 6 (4):

903-909. Two genera in the tribe Coffeeae occur in Australia, namely Coffea L. (with the naturalised

species C. arabica L. and the doubtfully naturalised C. liberica Hiern.) and Psilanthus Hook.f. (with the

single non-endemic species P. brassii (J.-F.Leroy) A.P.Davis). A key to genera is provided, together with an

amplified description and illustration for Psilanthus brassii.

Keywords: Coffea, Psilanthus - Australia

Paul I.Forster, Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens

Mt Coot-tha, Mt Coot-tha Road, Toowong Queensland 4066 Australia

Introduction

This paper provides an overview of the Tribe

Coffeeae (Rubiaceae: Ixoroideae) as it occurs

in Australia. The tribal classification within the

Rubiaceae was reviewed by Robbrecht & Puff

(1986) and summarised by Robbrecht (1988)

wherein a narrow circumscription for the tribe

Coffeeae DC. was proposed with only two

genera included, namely Coffea L. and

Psilanthus Hook.f. These two genera are closely

allied and are distinguished primarily on

growth habit (evergreen or deciduous) and

floral features (e.g. length of corolla tube, style

and stamen exsertion), all characters that might

be logically accommodated in a single genus.

Limited (as to the number of taxa sampled)

molecular work (chloroplast DNA) on

Psilanthus does tend to indicate that this genus

is discrete from Coffea, as the two species of

Psilanthus sampled clustered as an outgroup

to twenty-three taxa of Coffea (Cros et al.

1998). Extremely limited hybridization work

between single species from the two genera

(Coffea arabica L. and Psilanthus ebracteolatus

Hiern) (Couturon et al. 1998) found few barriers

to intergeneric gene transfer and these authors

considered recognition of two genera was not

warranted. Until further evidence is presented

on the monophyly or otherwise of the genera

in the Coffeeae, the status quo is maintained

here.

The genus Coffea has about 90 species

that are native to Africa, Madagascar and the

Mascarenes (Bridson 1988), with a couple of

Accepted for publication 10 May 2004

species widely cultivated in the tropics for

coffee. A single species of Coffea (C. arabica

L.) is naturalised in Queensland (Halford &

Reynolds 1994; Reynolds & Halford 1997;

Forster & Halford 2002) and on Lord Howe

and Norfolk Islands (Green 1994), although no

detail has been provided to date on the

Queensland naturalisations. A second species

of Coffea, C. liberica Hiern. has been recorded

from a single locality near Cape Tribulation;

but is not considered fully naturalised,

particularly as efforts have been made to

eradicate it from the site.

The genus Psilanthus Hook.f. was newly

recorded for Queensland, Australia by Bridson

(1987) with a note that Paracoffea brassii J.-

F.Leroy occurred in the Torres Strait, however,

no combination was made in Psilanthus at that

time. Davis (2003) has recently made the new

combination P. brassii (J.-F.Leroy) A.P.Davis

and recorded the species for Australia on the

basis of several collections from Queensland

(Cunningham 7 (K) and Wannan [1818]

NSW443387 (NSW)). Davis (2003) recorded

Psilanthus brassii from Central Province in

Papua New Guinea and from Australia (“Torres

Strait Island and Queensland”). He treated the

Torres Strait Islands as being a separate political

entity to the state of Queensland, which they

are not, and the few specimens that he cited

would indicate that the species only occurs in

the Torres Strait and tip of Cape York in

Queensland. Davis did not consult collections

at the Queensland Herbarium (BRI), the major

repository of material from Queensland,

otherwise the known distribution of this species

may have been better documented by him.

904

Apart from the original publication of

Paracoffea brassii J.-F.Leroy (basionym for

Psilanthus brassii ) where there is a description

in French (Leroy 1968), there is no description

of this plant in English that includes coverage

of the flowers, nor have the flowers been

illustrated, hence these deficiencies are

Taxonomy

Coffeeae DC., Ann. Mus. Hist. Nat. 9: 217

(1807), emend Robbrecht & Puff (1986).

Type: Cojfea L.

Austrobaileya 6 (4): 903-909 (2004)

remedied in the current paper. The genus

Psilanthus has about twenty-one species that

are found in tropical Africa, India and Malesia

(Bridson 1987; Sivarajan et al. 1992) as well

as the single species in Australia and New

Guinea.

Refer to Robbrecht & Puff (1986), Robbrecht

(1988) and Bridson (1988) for a tribal

description.

Key to genera in Australia

Evergreen shrubs; corolla-tube + same length as lobes; style and anther

fully exserted. Coffea

Deciduous shrubs; corolla tube longer than lobes; style and anthers included

(anthers may be part exserted). Psilanthus

Coffea L., Sp. PI. 172 (1753). Type: C. arabica L.

Refer to Bridson (1988) for a generic description.

Key to the Naturalised (or Doubtfully Naturalised) species of Coffea in Australia (N.B.

these characters may not work for material of these species outside of this region)

Leaf lamina elliptic; flowers 5-merous

Leaf lamina obovate; flowers 6-merous

1. Coffea arabica L., Sp. PL 172 (1753).

Type: cultivated in Holland, Hort. Clifford

(holo: BM).

Refer to Bridson (1988) or Green (1994) for a

species description.

Specimens examined : Queensland. Cook District: Curtain

Fig site, SWof Yungaburra, 17° 17’S, 145° 34’E, May 2000,

Bean 16559 (BRI); Daintree, Sep 1937, Brass 176 (BRI;

CANB, L, MO n.v.)\ Peeramon Scrub, 4 km NE of Malanda,

17° 18’S, 145° 37’E, Oct 2000, Forster PIF26330 etal. (A,

AD, BRI, K, MEL); Topaz, Towalla road, 17° 28’S, 145°

43’E, Oct2001, Forster PIF27610 etal. (BRI, MEL,NSW);

Lake Eacham N.P., S end of Picnic area, 17° 15,145° 35’E,

May 1985, Melville S15 (BRI); Rocky Creek, Atherton -

Mareebaroad, 17° 12’S, 145°27’E, Sep 1959, Smith 10795

(BRI); Prior’s Creek, Atherton, 17° 15’S, 145° 25’E, Oct

1992, Swarbrick 10551 (BRI); Kuranda, 16° 49’S, 145°

38’E, Nov 1992, Swarbrick 10621 (BRI). Moreton District:

Brisbane River, Long Pocket, Indooroopilly, 27° 3l’S, 152°

59’E, Aug 2000, Batianoff 200828 (BRI, NSW); Property

of D.Cunnington, Gwynore Crescent, Buderim, 26° 41’S,

153°04’E, Aug 1995, Bean 8871 (BRI, MEL); Off Paynter

Creek road, c. 1.5mileEofNambour, 26°38’S, 153°00’E,

May 1977, Elsol 111 (BRI); Buderim Mt., Fountain road,

26°45’S, 153° 05’E, Jul 1989 ,Nielsen [AQ456714] (BRI).

New South Wales. Lord Howe Island. Stevens Reserve,

.C. arabica

.C. liberica

31° 31’S, 159° 03’E, Jan 2001, Le Cussan 1102 (BRI; ex

NSW). Norfolk Island. Louis Evan’s Market Garden,

Duncombe Bay, 29° 00’S, 167° 55’E, Oct 1999, Waterhouse

BMW5528 (BRI). Map 1.

Distribution and habitat: Coffea arabica

(Arabian Coffee) is slowly naturalising in a

number of localities in the ‘Wet Tropics’ of

north-eastern Queensland, and around

habitation in the south-eastern corner of the

state (Map 1). The origins of these multiple

naturalisations in Queensland are undoubtedly

from cultivated plants, particularly in the ‘Wet

Tropics’ around the Atherton Tableland, and

near Buderim in the south-east. There are also

localised naturalisations on both Lord Howe

and Norfolk Islands (Green 1994). Arabian

Coffee has been grown occasionally as a

commercial crop or garden plant in these areas

and is commonly encountered in gardens.

Naturalised colonies of coffee have been

recorded from the margins of rainforest,

adjacent to old gardens or in disturbed

situations where there is considerable shrubby

undergrowth. It is presumed that bird mediated

Forster, tribe Cojfeeae in Australia

dispersal of Coffea arabica fruit is occurring,

but no direct observations are known. At this

stage, Cojfea arabica is lowly ranked in terms

of its invasiveness potential (ranked 189/200

by Batianoff & Butler 2002) and is not

considered a high priority invasive weed

(Batianoff & Butler 2003).

Notes: Material of Coffea arabica from

elsewhere, particularly within its natural

distribution range, has a broader range of leaf

lamina shape and floral part diversity than

indicated in the above key (Bridson 1988).

2. Coffea liberica Hiern., Trans. Linn. Soc.

Bot, ser. 2, 1: 171, t. 24 (1876).

Type: Sierra Leone, cultivated on Mr

Effenhausen’s farm, Daniell s.n. (lecto:

BM, fide Bridson (1985: 806).

Refer to Bridson (1988) for a species

description.

Specimens examined : Queensland. Cook District: Cooper

Creek, Cape Tribulation road, 16° ll’S, 145°25’E,Oct 1998,

Gray 7392 (BRI). Map 2.

Distribution and habitat : Cojfea liberica is

doubtfully naturalised at Cooper Creek in the

‘Wet Tropics’ of Queensland (Map 2) where it

has been recorded in lowland complex

mesophyll vineforest on substrates derived from

metamorphics.

Notes: The material from Cooper Creek appears

to fall within the variation ascribed to Coffea

liberica var. liberica, viz. corolla 6-9-merous,

tube distinctly widened at the throat, lobes

usually 5-10 mm wide (Bridson 1988).

It is likely that this species may continue

to pose a problem as a naturalised plant in the

Cape Tribulation area unless follow-up

eradication procedures are employed.

Psilanthus Hook.f., Gen. PI. 2: 115 (1873).

Refer to Bridson (1988) for a generic

description.

A genus of c. 22 species. One non¬

endemic species in Australia (Queensland).

Psilanthus brassii (J.-F.Leroy) A.P.Davis,

Novon 13: 183 (2003).

905

Paracoffea brassii J.-F.Leroy, J. Agric. Trop.

Bot. Appl. 14: 598 (1967).

Type: Papua New Guinea. Central Province:

Baroka, Nakeo district, 8 April 1933, L.J.

Brass 3743 (holo: BO n.v.\ iso: A n.v.;

BRI).

[Randia sp. (Coen B.P.Hyland 13278)

(Reynolds & Halford 1997)]

Illustration: Leroy (1968: 599).

Shrub to 3 m high, deciduous. Indumentum

absent or of simple, uniseriate trichomes. Leaf

bearing stems slender, covered with thin

reddish-brown bark, glabrous or with scattered

to sparse trichomes. Stipules truncate to

irregularly triangular, basal tubular portion

0.8-1.5 mm long, free apiculate portion 1.8-4

mm long. Leaves petiolate; petioles 4-5 mm

long; lamina elliptic to obovate, 25-95 mm

long, 12-60 mm wide, thin, papery, discolorous

and drying grey-green, glabrous or with

scattered to sparse indumentum abaxially on

midrib and some lateral veins; domatia absent

or present abaxially as small tufts of

indumentum in angle between midrib and

lateral veins; midrib and some lateral veins

whitish on abaxial surface; tip acute to short

acuminate; base cuneate. Flowers solitary and

terminal on stems, sometimes on a short shoot

to 5 mm long; flowers pedicellate for 1.5-2 mm,

immediately subtended by short bracteoles, that

are + truncate, 0.4-1 mm long, then subtended

(or not) by a leaf pair (shed in fruit), finally

subtended by 4 bracts (shed in fruit) that are

lanceolate, 3-5.8 mm long and 0.8-1.2 mm

wide. Calyx tube 1.8-3 mm long, c. 1 mm

diameter, bearing 4 or 5 unequal lobes that are

shortly acute, 0.3-0.5 mm long. Corolla

glabrous; tube 16-18 mm long, 2.8-3 mm wide

at top, 1.2-1.6 mm wide at base; lobes acute,

10-13 mm long, 4.5-6 mm wide, attached 14-15

mm from base. Stamens 5; anthers attached

near apex of tube with only the tips exserted,

6-6.5 mm long, c. 0.6 mm wide. Style c. 5 mm

long; stigma lanceolate, c. 2.5 mm long,

positioned 5-6 mm below the anthers. Fruit

bilobed (rarely trilobed), soft-fleshy, 7-8 mm

long, 7.5-8 mm wide, black to plum coloured;

seeds 2 (3) per fruit, oblong-ellipsoid, c. 6 mm

long and 5 mm wide, fawn, distinctly grooved

on adaxial face. Fig. 1.

906

Austrobaileya 6 (4): 903-909 (2004)

Fig. 1. Psilanthus brassii. A. branchlet with fruit, x 0.8. B. undersurface of leaf showing venation, x 1. C. indumentum on

undersurface of leaf (area indicated in circle in B) with domatium in angle between vein and midrib, x 16. D. node with stipule,

x 8. E. node with inflorescence bearing four bracts, x 4. F. inflorescence with leaf pair' and flower, x 4. G dissected corolla with

stamens and pistil, x 4. H. node with fruiting inflorescence. X 3.1. seed showing groove on adaxial face, x 4. A & D from Fell

DGF2919 (BRI); B, C, H, I from Forster PIF19452 (BRI); F & G from Wannan 1418 (BRI); E from Puttock & King

UNSW16901 (BRI). Del. W.Smith.

907

Forster, tribe Cojfeeae in Australia

Specimens examined : Papua New Guinea. Central

Province: Tovobada Hills, 12 miles [20 km] N of Port

Moresby, May 1965, Heyligers 1199 (BRI; CANB, L, LAE

n.v.); Near SE side of Little Mt Lawes, c. 16 miles [26.7 km]

N of Port Moresby, Apr 1967, Pullen 6814 (BRI; A, CANB,

K, L, LAE ruv.y, Tovobada Hills, c. 9 miles [15 km] NNW of

Port Moresby, May 1965, Pullen 6980 (BRI; A, BO, CANB,

K, L, LAE, US n.v.). Australia. Cook District: 10.8 km S of

Batavia Downs on the Peninsula Development road, 12°45’S,

142° 43’E, Apr 1990, Clarkson 8463 & Neldner (BRI, L,

MBA); Jane Table Hill, Lakefield N.P., 46.6 km N of

Lakefield Homestead, 14° 30’S, 144° 07’E, Mar 1993, Fell

DGF2919 & Stanton (BRI, MEL); Cape Melville N.P.,

Altanmoui Range section, 1.6 km E of Flat Hill, 62.6 km NE

of Lakefield Homestead, 14° 30’S, 144° 35’E, May 1993,

Fell DGF3215 & Stanton (BRI); Altanmoui Range, Cape

Melville N.P., 1.6 km E of Flat Hill, 62.6 km NE of Lakefield

RangerBase, 14° 30’S, 144° 35’E, May 1994, Fell DGF4350

& McDonald (BRI); Equina Scrub, 42.4 km W of Coen,

Holroyd Pastoral Holding, 13° 59’S, 142° 48’E, Jun 1994,

Fell DGF4397 & Buck (BRI); Round Mountain, Embley

Range, 13° 33’S, 143° 30’E, Jun 1992, Forster PIF10467 &

Tucker (BRI); Klondyke, Station Creek track, Mcllwraith

Range, 14° 00’S, 143° 19’E, Jun 1996, Forster PIF19452

(BRI); Gore Island, Great Barrier Reef, 11°59’S, 143° 14’E,

Apr 1995, Gray 6125 (BRI; ex QRS); T.R. 14, Parish of

Kesteven, 13° 42’S, 143° 18’E, Nov 1991, Hyland 14378

(BRI; ex QRS); Restoration Island, 12° 36’ S, 143° 26’E, Apr

1995, Le Cussan 286 (BRI); Haggerstone Island, 12° 03’S,

143° 18’E, May 1995, Le Cussan 339 (BRI); Portland Roads,

c. 0.5 km S of headland, 12° 36’S, 143° 24’E, Puttock &

King UNSW16901 (BRI); 1 km S of Seisia, 10° 51’S, 142°

21’E, Nov 1999, Wannan 1418 (BRI, NSW).

Distribution and habitat : Psilanthus brassii

has been recorded from Central Province of

Papua New Guinea (in the vicinity of Port

Moresby) and from the Cook District of

Queensland with a northern occurrence on

islands in Torres Strait and a southern limit at

Altanmoui Range, Cape Melville (Map 2).

Plants grow in “monsoon” forest (semi-

deciduous microphyll to notophyll vineforest)

on substrates derived from stabilised sand-

dunes, laterite, sandstone or metamorphics at

altitudes from near sea level to 300 m.

Notes: The identity of this species was first

debated by Merrill & Perry (1945) who referred

collections from Papua New Guinea ( Brass

3743; Carr 11220 & 12389) and the Torres

Strait (Sunday Island) ( Cunningham 7) to

Cojfea (Lachnostoma ) with a ? Nearly 60 years

on and with the addition of the collection by

Wannan , Davis (2003) finally classified this

species in Psilanthus. This delay in

identification has undoubtedly been influenced

by the lack of flowering material (only Hyland

14378 and Wannan 1418 have flowers). The

deciduous habit of Psilanthus brassii means

that the plant is probably leafless between the

months of September and November. Flowering

commences with the onset of storm rain in

November and December just prior to, or as

the plants produce new leaves after the end of

the ‘dry’ season. Fruits have been collected

between March and June and are borne on

plants where the leaves are fully expanded and

hardened. This pattern of floral and fruiting

behaviour is not unusual and is repeated in a

wide range of plant species from diverse

families that grow in the so-called ‘wet - dry’

tropics. Good examples of species with this

phenological behaviour are Croton simulans

P.I.Forst. (Euphorbiaceae) (Forster 2003),

Drypetes deplanchei (Brongn. & Gris) Merr.

(Euphorbiaceae) (Forster 1997), Gyrocarpus

americanus Jacq. (Hernandiaceae), Mallotus

surculosus P.I.Forst. (Euphorbiaceae) (Forster

1999) and Turraea pubescens Hellen.

(Meliaceae). Nevertheless this phenological

behaviour can make identification of this

Rubiaceous plant difficult when it is leafless or

with only immature leaves.

Davis (2003) noted that Psilanthus brassii

was similar to P. mabesae (Elmer) J.-F.Leroy

from the Philippines but differed in the “leaves,

flowers (corolla), and fruits... glabrous (not

puberulous to pubescent as in P. mabesaey\

With the greater amount of material of

Psilanthus brassii now available, these

distinctions are not strictly correct. Although

the flowers (corolla) and fruits of P brassii are

glabrous, not all collections have glabrous

foliage. The collections Fell DGF4397, Forster

PIF19452, Le Cussan 266 & 399, have from

between a few scattered hairs on the leaf midrib

below, to being sparsely hairy on the leaf

undersurface and young shoot tips. While most

collections lack leaf domatia, those that are

pubescent to some degree also have occasional

small domatia formed in the angle between the

midrib and a lateral vein.

There is some slight variation in the

arrangement of bracts and bracteoles in the

inflorescence of this species. In its simplest

form, the terminal inflorescence is subtended

by two small bracteoles (retained in the fruit)

that are in turn subtended by four linear-

lanceolate bracts that are caducous. In its most

complex form, the terminal inflorescence is

908

once again subtended by the two small

bracteoles (retained in the fruit), that are

subtended by the four linear-lanceolate bracts,

then further subtended by a pair of true leaves,

the stipular pair, then a short shoot, followed

by the four linear-lanceolate bracts and a pair

of stipules again. Whether the most complex

form has been derived from an inflorescence

that has flowered and then produced a new

shoot terminally, followed by yet another

inflorescence is unknown without actual

observation of live plants.

Conservation status : Psilanthus brassii is

widespread on Cape York Peninsula and some

islands in Torres Strait. It has been recorded

from Cape Melville and Lakefield National

Parks. Davis (2003) listed it as of Least

Concern; however, this was based on limited

information about its distribution or perceived

threats. It is not considered rare or threatened

at present.

Acknowledgements

Field collections of the two species concerned

were facilitated with the assistance of R.Booth

(BRI), R. Jensen, G. Sankowsky and M.C. Tucker.

Artwork was executed by W. Smith (BRI).

References

Batianoff, G..N. & Butler, D.W. (2002). Assessment of

invasive naturalized plants in south-east Queensland.

Plant Protection Quarterly 17: 27-34.

-(2003). Impact assessment and analysis of sixty-six

priority invasive weeds in south-east Queensland.

Plant Protection Quarterly 18: 11-17.

Bridson, D.M. (1985). The lectotypification of Coffea

liberica (Rubiaceae). Kew Bulletin 40: 805-807.

-(1987). Nomenclatural notes on Psilanthus, including

Coffea sect. Paracoffea (Rubiaceae tribe Coffeeae).

Kew Bulletin 42: 453-460.

-(1988). Psilanthus. In R.M.Polhill (ed.), D.M.Bridson

& B.Verdcourt, Flora of Tropical Africa, Rubiaceae,

Part 2, pp. 723-727. Rotterdam/Brookfield:

Balkema.

Couturon, E., Lasermes, P. & Charrier, A. (1998). First

intergeneric hybrids ( Psilanthus ebracteolatus

Hiern. x Coffea arabica L.) in coffee trees.

Canadian Journal of Botany 76: 542-546.

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Cros, J., Combes, M.C., Trouslot, R, Anthony, E, Hamon,

S., Charrier, A. & Lashermes, P. (1998).

Phylogenetic analysis of chloroplast DNA variation

in Coffea L. Molecular Phylogenetics & Evolution

9: 109-117.

Davis, A.P. (2003). A new combination in Psilanthus

(Rubiaceae) for Australasia, and nomenclatural

notes on Paracoffea. Novon 13: 182-184.

Forster, PI. (1997). A taxonomic revision of Drypetes Vahl

(Euphorbiaceae) in Australia. Austrobaileya 4:

477-494.

-(1999). A taxonomic revision of Mallotus Four.

(Euphorbiaceae) in Australia. Austrobaileya 5:

457-497.

-(2003). A taxonomic revision of Croton F.

(Euphorbiaceae) in Australia. Austrobaileya 6:

349-436.

Forster, PI. & Halford, D.A. (2002). Rubiaceae. In

R.J.F.Henderson (ed.), Names and Distribution of

Queensland Plants, Algae and Lichens, pp. 173—

177. Brisbane: Environmental Protection Agency.

Green, PS. (1994). Rubiaceae. Flora of Australia 49: 350-

359. Canberra: Australian Government Publishing

Service.

Halford, D.A. & Reynolds, S.T. (1994). Rubiaceae. In

R.J.F.Henderson (ed.), Queensland Vascular Plants

Names and Distribution, pp. 294-301. Brisbane:

Queensland Department of Environment & Heritage.

Eeroy, J.-F. (1967, published 1968). Un Cafeier du genre

Paracoffea en Nouvelle-Guinee. Journal

d Agriculture Tropicale et de Botanique Appliquee

14: 598-600.

-(1981). Les cafeiers du genre Psilanthus (Rubiacees) en

Afrique orientale et en Asie etiles du Pacifique. Bull.

Mus. Natl. Hist., Nat., B, Adansonia 3: 251-258.

Merrill, E.D. & Perry, L.M. (1945). Plantae Papuanae

Archboldianae, XVI. Journal of the Arnold

Arboretum 26: 229-266.

Reynolds, S.T. & Halford, D.A. (1997). Rubiaceae. In

R.J.F.Henderson (ed.), Queensland Plants Names

and Distribution, pp. 180-184. Brisbane:

Department of Environment.

Robbrecht, E. (1988). Tropical Woody Rubiaceae. Opera

Botanica Belgica 1: 1-271.

Robbrecht, E. & Puff, C. (1986). Asurvey of the Gardenieae

and related tribes (Rubiaceae). Botanische

Jahrbticher fiir Systematik, Pflanzengeschichte

und Pflanzengeographie 108: 63-137.

Sivarajan, V.V., Bhu, S.D. & Mathew, P. (1992). Revision of

the genus Psilanthus Hook.f. (Rubiaceae tribe

Coffeeae) in India. Botanical Bulletin of Academia

Sinica (Taipei) 33: 209-224.

Forster, tribe Cojfeeae in Australia

909

Map I. Distribution in 1° grids in Australia for Cojfea liberica T and Psilanthus brassii #

Map 2, Distribution in 1 6 grids in Australia (excluding Lord Howe and Norfolk Islands) for

Cojfea arabica A .

Caelospermum dasylobum (Rubiaceae), a new species from north-eastern

Queensland

D.A. Halford and A.J. Ford

Summary

Halford, D.A. & Ford, A.J. (2004). Caelospermum dasylobum (Rubiaceae), a new species from north¬

eastern Queensland. Austrobaileya 6 (4) 911-915. Caelospermum dasylobum Halford & A.J.Ford is

described, illustrated and diagnosed against related species. Notes on its habitat and distribution are provided.

Key words: Caelospermum , taxonomy, Australian flora, Caelospermum dasylobum, Rubiaceae

D.A. Halford, Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens Mt

Coot-tha, Mt Coot-tha Road, Toowong, Queensland 4066, Australia

A. J. Ford, CSIRO, Sustainable Ecosystems and Rainforest-CRC, Tropical Forest Research Centre, PO Box

780, Atherton, Queensland 4883, Australia

Introduction

Caelospermum Bl. is a genus of mostly lianas

distributed throughout South-east Asia,

Malesia, Melanesia, New Guinea and Australia.

In the most recent revision of the genus,

Johannson (1988) recognised a total of seven

species, with a single species C. paniculatum

F.Muell. present in Australia. Johannson

recognised two varieties of C. paniculatum:

C. paniculatum var. paniculatum distributed

from the Claudie River, north-eastern

Queensland southwards to the Clarence River,

north-eastern New South Wales and C. paniculatum

var. syncarpum J.T. Johansson occurring on the

Atherton Tableland, north-eastern Queensland.

A second Australian species of Caelospermum,

from the Mossman-Julatten area, is described

here.

Methods

The study is based upon the examination of

herbarium material from BRI and QRS and

augmented by field observations made by the

second author. The Herbarium acronyms follow

Holmgren et al. (1990). All specimens cited

have been seen by one or both authors.

Measurements of floral parts are based

on material preserved in 70% ethanol. In this

paper the term “compound syncarpous drupe”

is used to describe the compound fruit of the

species. The term “drupe” is used here as

defined by Clifford and Dettmann (2001).

Accepted for publication 18 June 2004

Taxonomy

Caelospermum dasylobum Halford &

A.J.Ford, sp. nov. in clavi de Johannson

(1988) advenit ad C. salomonense a quo

corollae lobis dense pubescentibus et

brevioribus (6-7 mm longis vice 9-11

mm longis), antheris brevioribus (3-3.5

mm longis vice 5.6-7.4 mm longis),

ramulis inflorescentiae glabris (hispidulosis

in C. salomonensi ) et fructibis ubi maturis

plus minusve flavo-aurantiacis usque

flavo-brunneis (rubris in C. salomonensi)

differt. Caelospermum dasylobum in

fructibus drupam compositum syncarpum

formentibus C. paniculato var. syncarpo

similis autem ramulis inflorescentiae

glabris (hispidulosis in C. paniculato var.

syncarpo ) fructibus ubi maturis plus

minusve flavo-arantiacis usque flavo-

brunneis (rubris usque purpureis in

C. paniculato var. syncarpo), lobis

corollae dense pubescentibus (glabris in

C. paniculato var. syncarpo) et tubo

corollae interdum longiore (5-7 mm

longis vice 3-6 mm longis in C. paniculato

var. syncarpo) differt. Typus: Queensland.

Cook District: Chapman road, Kingfisher

Park, Julatten, Dec 2002, A. Ford

AF3785, J. Holmes & D. McJannet (holo:

BRI; iso: K, L, MEL, MO, NSW, QRS,

distribuendi).

Twining vine or scandent shrub with long

arching stems. Stems terete, bark corky and

fissured to tessellated, to 4cm diameter. Wood

912

Austrobaileya 6 (4): 911-915 (2004)

and roots yellow-orange. Branchlets ± terete,

rarely angled, glabrous, longitudinally striated

and becoming fissured with age. Leaves

petiolate, opposite; stipules interpetiolar,

sheathing, c. 1.5 mm long, produced into a

narrow triangular lobe, glabrous, fragmenting

as node thickens; petioles 5-12 mm long;

laminae chartaceous when young becoming

hard and stiff with age, narrowly elliptic or

narrowly ovate, 6.5-13 cm long, 3.5-6.5 cm

wide; adaxial and abaxial surfaces glabrous;

venation brochidodromus with 5-12 lateral

veins per side of midvein, slightly raised on

adaxial surface, prominent on abaxial surface;

base rounded to cuneate-obtuse; margins entire;

apex acute or acuminate; pit domatia with a

ring of hairs around the orifice present on most

leaves in lateral vein axils on abaxial surface.

Flowers fragrant, 4(5)-merous, bisexual, sessile,

in congested capitula, joined at least by the base

of the gynoecium. Capitula pedunculate, 4-6

mm across (excluding corollas), 3 or 4 (rarely

6)-flowered with colleters absent between

flowers, in terminal paniculate umbel-like

dichasial cymes or rarely axillary dichasial

cymes, with peduncles 0.5-6 mm long,

glabrous, terete. Primary axis of inflorescence

10-30 mm long, glabrous, terete; bracts c. 1mm

long, glabrous; lower bracts broadly triangular

or rarely foliaceous; upper bracts narrowly

triangular. Calyx tube (including hypanthium)

green, 1.5-2 mm long, 1.7-2 mm across,

abaxial surface glabrous, with a single bristle¬

like bracteole (?) c. 0.8mm long occasionally

present towards the base of the gynoecium;

adaxial surface glabrous, with a ring of minute

colleters at base; apex truncate, undulate or

irregularly and shallowly toothed. Corolla

valvate, deciduous, white and pale green

turning yellow with age, glabrous on abaxial

surface; tube 5-7 mm long, + cylindrical,

slightly widened at the mouth, fenestrated by

short longitudinal splits in lower third of tube,

glabrous and papillose on adaxial surface in

proximal half but hairy and smooth distally;

hairs simple, up to 0.5 mm long, white,

spreading; lobes spreading and strongly

recurved distally at anthesis, narrowly ovate,

6-7 mm long, 2-2.5 mm wide, acute and ±

cucullate at apex, densely hairy adaxially; hairs

as for corolla tube. Stamens exserted; filaments

1.6-4.5 mm long, inserted at the sinuses of the

corolla lobes; anthers dorsifixed, linear-oblong,

3-3.5 mm long, glabrous, dehiscing laterally

through longitudinal slits. Disc entire, flat, c.

0.5 mm high, glabrous. Ovary 2-celled,

biovulate; style 5.5-8 mm long, exserted,

glabrous; stigma bifid, with spreading lobes

2.5-4 mm long, adaxial surface and margin

papillose, abaxial surface glabrous. Fruit a

compound syncarpous drupe, subglobose or

irregularly lobate, 20-25mm long and 15-30

mm across, + yellow-orange to yellow-brown

when ripe, glabrous, persistent calyx tubes not

prominent on surface; pericarp firm, shell-like;

mesocarp fleshy, containing several pyrenes.

Pyrenes oblong in outline, 8-11 mm long, 5-6

mm wide, 2-3 mm thick, 1-seeded; endocarp

cartilaginous, pale brown, rugose, with basal

marginal groove. Seed 7-9 mm long, 3-4 mm

wide, c. 1 mm thick; testa membraneous, dark

brown; endosperm corneous, white; embryo

3.7-4.3 mm long, straight; cotyledons 1.6-2

mm long, 0.9-1.1 mm wide, thin, c. twice as

broad as the radicle; radicle 1.9-2.3 mm long,

0.5-0.6 mm wide. Fig. 1.

Additional specimens examined : Queensland. Cook

District. Kingfisher Park, Julatten, Oct 2002, Cooper &

Cooper WWC1791 (BRI); SFR 72, Pinnacle Mountain, Aug

1983, Dansie AF05283 (QRS); SFR 1229, Black Mountain

road, near Rifle Creek bridge, Aug 2003, Ford 4148 (BRI,

QRS); Mt Perseverence road, near Nissen Creek via Julatten,

Feb 2003, Ford AF3840 & Holmes (BRI); SFR 143, Little

Mossman logging area, Jul 1980, Gray 20141V (QRS); ditto,

Aug 1981, Gray 20193V (QRS); c. 1.5 km N of Julatten,

adjacent to Kingfisher Park Bird Lodge, Sep 2002, Halford

Q7385 & Ford (BRI); Kingfisher Park, Julatten, Oct 1998,

Holmes 69 (QRS); SFR 1229, Danbulan, Feb 1993, Hyland

14638 (QRS); ditto, Aug 1992, Hyland 14506 (QRS); c.

lkm S of “The Pinnacle” & c. 12km SSE of Mossman, Sep

1978, Moriarty 2461 (QRS); bank of Mossman River, near

Mair [Marr] Creek, Apr 1998, Sankowsky 1625 (QRS);

Rumula Creek, Oct 1939, Twort per Sparvell NQNC6570/

71 (QRS).

Distribution and habitat : Caelospermum

dasylobum is endemic to north-eastern

Queensland, where it is currently known from

the Julatten-Mossman district (Map 1). It is

recorded as growing in mesophyll or notophyll

rainforest on clay soils derived from mudstone.

Common canopy trees include: Cardwellia

sublimis F.Muell., Darlingia darlingiana

(F.Muell.) L.A.S.Johnson, Elaeocarpus grandis

F.Muell., Castanospora alphandii (F.Muell.)

F.Muell., Castanospermum australe A.Cunn.

& Fraser ex Hook., Carnarvonia araliifolia

F.Muell. var. araliifolia, Myristica globosa

subsp. muelleri (Warb.) W.J.deWilde and

Halford & Ford, Caelospermum dasylobum

913

Fig. 1. Caelospermum dasylobum. A. branchlet with flowers x 0.4. B. part of inflorescence x 2. C. flower x 4. D. style x 8.

E. fused ovaries with bracteole x 8.F. compound fruit x 1.5. G. adaxial view of pyrene x 6. H. lateral view of pyrene x 6.

I. embryo x8. A-E from Ford el al. AF3785 (BRI); F from Cooper & Cooper WWC1791 (BRI); G-I from Halford Q7385

& Ford (BRI). Del. W. Smith.

914

Austrobaileya 6 (4): 911-915 (2004)

Alstonia scholaris (L.) R.Br. Polyscias

australiana (F.Muell.)Philipson is a conspicuous

sub-canopy species. At the type locality the

rainforest is very fragmented from clearing and

agriculture. Very small fragments exist with

little or no canopy structure. In this case

C. dasylobum occurs with Guioa acutifolia

Radik., Cryptocarya triplinervis R.Br. and

Rhamnella vitiensis (Benth.) A.C.Sm.

Altitudinal range is 20-600m, with most

collections recorded at 400-450m.

Phenology: Flowers have been recorded in

December, January and February whilst mature

fruits have been recorded in September and

October.

Affinities'. Caelospermum dasylobum will key

to C. salomonense in Johansson’s (1988) key.

It can be distinguished from C. salomonense

by its densely hairy and shorter corolla lobes

(6-7 mm long compared to 9-11 mm long for

C. salomonense), shorter anthers (3-3.5 mm long

compared to 5.6-7.4 mm long for C. salomonense),

glabrous inflorescence branches

(hispidulous for C. salomonense) and ±

yellow-orange to yellow-brown fruit when

ripe (red for C. salomonense).

Caelospermum dasylobum is similar to

C. paniculatum var. syncarpum in its fruit that

form a compound syncarpous drupe. However,

it differs in having glabrous inflorescence

branches (hispidulous for C. paniculatum var.

syncarpum), ± yellow-orange to yellow-brown

fruit when ripe (red to purple for C. paniculatum

var. syncarpum), densely hairy corolla lobes

(glabrous for C. paniculatum var. syncarpum)

and generally longer corolla tube (5-7 mm long

compared to 3-6 mm long for C. paniculatum

var. syncarpum).

Conservation status: Caelospermum

dasylobum has been collected within the World

Heritage Area of the Wet Tropics bioregion in

the vicinity of Black Mountain (Harris Peak),

via Kuranda and also from the Julatten area

(ex-SFR 143 and ex-SFR 72). However, it has

not yet been recorded in a National Park area.

No conservation status is recommended at this

time, although a thorough search of adjacent

lands might elucidate more records of this

seldom seen species.

Notes: Leaves on strictly juvenile plants or

seedlings in the understory have very narrow

leaves compared to adult leaves. This vegetative

feature is common within the genus of Morinda.

Mature fruit of C. dasylobum lack the distinctive

and pungent smell of rotting cheese, which is

a well known attribute of Morinda fruit.

Etymology: The specific epithet is derived from

the Greek dasys - very hairy, and lobos - lobe,

and refers to the hairy corolla lobes of this

species.

Acknowledgements

The authors wish to thank Will Smith for the

illustration, Peter Bostock for the map and Les

Pedley for the translation of the diagnosis into

Latin. Jenny Holmes (Atherton) provided field

assistance and showed the authors additional

locations for distributional information. Staff

at QRS (CSIRO, Plant Industry, Atherton)

provided access to specimens. Permits to collect

and traverse, in State Forest Reserve 1229, were

issued by the Queensland Parks and Wildlife

Service. The first author would like to thank

Gordon Guymer, Director of BRI, for making

available working space and facilities at BRI.

References

Clifford, H.T. & Dettmann, M.E. (2001). Drupe - a term in

search of a definition. Austrobaileya 6(1): 127-131.

Holmgren, P.K., Holmgren, N.H. & Barnett, L.C. (1990).

Index Herbariorum. Part 1. The Herbaria of the

World. 8 th edn. New York: New York Botanic

Gardens.

Johansson, J.T. (1988). Revision of the genus Caelospermum

Blume (Rubiaceae, Rubioideae, Morindeae).

Blumea 33(2): 265-297.

Halford & Ford, Caelospermum dasylobum

915

Map 1. Distribution of Caelospermum dasylobum m

New species of Cryptandra Sm. and Stenanthemum Reissek (Rhamnaceae)

from northern Australia

A.R. Bean

Summary

Bean, A.R. (2004). New species of Cryptandra Sm. and Stenanthemum Reissek (Rhamnaceae) from northern

Australia. Austrobaileya 6 (4): 917-940. Seven new species of Cryptandra, and one new species of

Stenanthemum are described and illustrated. One (C. gemmata ) is from the Northern Territory, four

(C. debilis, C. filiformis, C. pogonoloba, S. argenteum ) are from northern Queensland, and three

(C. ciliata, C. orbicularis, C. rigida) are from southern Queensland. Full diagnostic descriptions are given

for all Queensland taxa and for C. gemmata. Distribution maps and an identification key are provided

covering all Queensland species. Photographic images are provided for all newly described taxa. C. armata

is newly recorded for New South Wales.

Keywords: Queensland, Northern Territory, Cryptandra, Stenanthemum, taxonomy, key, Australian flora,

new species, Rh a mnaceae.

A.R. Bean, Queensland Herbarium, Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha road, Toowong,

Queensland 4066, Australia.

Introduction

Cryptandra Sm. and Stenanthemum Reissek are

endemic Australian genera found mostly in the

southern non-arid parts of the continent, but

with some representation in the subtropics and

tropics. Cryptandra comprises about 40 species,

and Stenanthemum about 30 species (Rye 1995;

Walsh & Udovicic 1999).

Taxonomic problems associated with the

Tribe Pomaderreae (e.g. Cryptandra,

Stenanthemum, Spyridium Fenzl and

Trymalium Fenzl) usually relate to the rather

ill-defined distinctions between the genera.

Rye (1995) published numerous new

species of both Cryptandra and Stenanthemum

from Western Australia. She did not explicitly

state the differences she perceived between

these genera, but comparisons reveal that she

interpreted Stenanthemum to have several to

many flowers aggregated into dense head-like

clusters surrounded by bracts and leaves, leaf

margins incurved (conduplicate in bud),

stipules on the upper side of the petiole (often

connate), and a glabrous disc; with Cryptandra

having a 1-flowered inflorescence usually

subtended by a leaf, leaf margins recurved to

revolute, stipules connate on lower side of

petiole, and a densely stellate-hairy disc.

Accepted for publication 15 April 2004

Walsh & Udovicic (1999) stated that (in

Victoria) Stenanthemum has a several-flowered

head-like inflorescence, sometimes surrounded

by ‘floral leaves’, the individual flowers with

0-2 bracts, and the disc glabrous.

This paper presents new species from

Queensland and the Northern Territory, and an

identification key for all Queensland taxa. One

of the new species clearly belongs in

Stenanthemum, while the remainder have been

placed in Cryptandra, although some of the

northern species have some features not usually

associated with Cryptandra.

While I have not had the opportunity to

view many of the types of existing species, the

respective protologues have been sufficiently

detailed to satisfy me that none of the new

species are synonymous with them. Indeed,

some of the new taxa occur thousands of

kilometres from related species and are very

distinctive.

All Cryptandra and Stenanthemum

species grow on infertile sandy soils, or

sometimes on almost bare rock. Weed species

are few on these substrates, and the threat from

clearing or grazing is minimal. Only one of

the species described in this paper is considered

threatened under the criteria developed by the

International Union for the Conservation of

Nature (IUCN 2001).

918

Austrobaileya 6 (4): 917-940 (2004)

Taxonomy

Key to the Queensland species of Cryptandra and Stenanthemum

1. Lateral branchlets terminating in a spine. 2

Lateral branchlets not terminated by a spine. 3

2. Leaf margins strongly recurved to re volute; calyx with simple adpressed

hairs throughout. 4. C. armata

Leaves flat or margins weakly recurved; calyx with stellate hairs throughout

. 2a. C. amara var. amara

3. Upper leaf surface with dense stellate hairs. 4

Leaf upper surface glabrous, muricate, or with short simple hairs. 5

4. Stipules 2.8-4.4 mm long, thread-like, stellate-hairy. 13. C. filiformis

Stipules 1.7-2 mm long, broad-based, glabrous. 12. C. pogonoloba

5. Leaf margins re volute, lower surface rarely if ever visible (dried material). 6

Leaves flat or margins recurved or incurved, lower surface always easily visible. 11

6. Calyx tube and calyx lobes thickly covered by tangled woolly hairs. 7. C. lanosiflora

Calyx tube and lobes glabrous or hairy, but without tangled woolly hairs. 7

7. Bract margins conspicuously ciliate (cilia 0.4-0.6 mm long). 10. C. ciliata

Bract margins entire or with cilia 0.05-0.2 mm long. 8

8. Bracts confined to very base of calyx tube, longest bracts 1.0—1.3 mm long. 9

Bracts covering most or all of calyx tube, longest bracts 2.1-3.8 mm long. 10

9. Leaf apex acute to mucronate; style densely hairy; bracts glabrous on outer

surface. 1. C. debilis

Leaf apex obtuse; style glabrous; bracts stellate-hairy on outer surface

.3. C. amara var. floribunda

10. Calyx tube glabrous (except adjacent to lobes); bracts (at anthesis) covering

50-90% of tube. 9. C. rigida

Calyx tube stellate-hairy throughout; bracts (at anthesis) covering all of

calyx tube and part of calyx lobes. 8. C. propinqua

11. Lower leaf surface white, densely hairy. 12

Lower leaf surface green, glabrous or very sparsely hairy. 14

12. Leaves 1.8-4 mm long; stipules 0.6-1.1 mm long. 6. C. longistaminea

Leaves 5-16 mm long; stipules 2-3.8 mm long. 13

13. Leaves recurved; inflorescence with conspicuous masses of woolly hair;

south Qld.15. Stenanthemum scortechinii

Leaves flat or incurved; inflorescence without woolly hairs; north Qld

.14. Stenanthemum argenteum

14. Leaves oblanceolate to obovate, 2-4 times longer than broad; petals

protruding 0.4-0.6 mm beyond calyx tube; longest bracts 0.7-0.8 mm

long, covering 0-20% of calyx tube. 2a. C. amara var. amara

Leaves orbicular to reniform, 0.8-1.3 times longer than broad; petals

extending 0.8-1.1 mm beyond calyx tube; longest bracts 1.2-1.5 mm

long, covering 50-100% of calyx tube. 5. C. orbicularis

Bean, new species of Cryptandra & Stenanthemum

1. Cryptandra debilis A.R.Bean sp. nov.

affinis C. amarae var. floribundae Maiden

& Betche, sed stylo longiore pills densis

in dimidio inferiore, calyce longiore,

apicibus foliorum acutis et ramulis saepe

glabris differens. Typus: Queensland.

Cook District: Baal Gammon, E of

Watsonville, 18 May 1997, R.L. Jago

4379 & B. Wannan (holo: BRI; iso:

NSW).

Cryptandra amara var. (Mt Mulligan J.R.

Clarkson 5865) in Bean (2002)

Shrub to 0.3 m high, lateral branchlets 1-8 cm

long, not terminating in a spine. Young

branchlets often glabrous, sometimes with

simple adpressed hairs overlying small stellate

hairs. Older branches glabrous. Stipules

connate on lower side of petiole, persistent,

narrowly triangular, 1.3-1.6 mm long. Leaves

in fascicles (2-8 per node), petioles 0.3-0.5 mm

long; lamina terete, linear, 2.5-6.5(-12) mm

long, 0.5-0.7(-1.0) mm wide, 5-10 times

longer than broad, apex acute to mucronate,

margins strongly revolute; upper surface green,

glabrous; lower surface (rarely visible) white,

indumentum dense throughout, with simple

hairs overlying small stellate hairs.

Inflorescences 1-4 flowered, anthopodia

0.1-0.5 mm long; floral bracts 3-6, broadly

ovate to orbicular, 1.1-1.3 mm long, brown,

outer surface glabrous, apex obtuse or acute,

margin entire or with cilia 0.05-0.1 mm long.

Calyx white; calyx tube cylindrical, 1.2-1.5 mm

long, indumentum dense throughout, with

simple adpressed hairs overlying small stellate

hairs, 0-20% of tube obscured by bracts at

anthesis; calyx lobes erect, 1.0-1.2 mm long,

indumentum dense throughout, with simple

adpressed hairs overlying small stellate hairs.

Petals hooded, white, protruding 0.4-0.5 mm

beyond calyx tube. Disc densely stellate hairy,

obscurely 10-lobed. Style 0.6-0.7 mm long,

densely hairy on lower half, stigma 3-lobed.

Schizocarps ellipsoidal, c. 2.6 mm long,

3-locular, c. 20% inferior, ovary roof with

sparse stellate and simple indumentum, disc

forming a narrow densely stellate-hairy annulus

at the base of the persistent calyx, mericarps

dehiscent by a ventral slit. Seeds flattened,

elliptical in outline, c. 1.8 mm long excluding

aril. Aril white, terminal. Fig. 1.

919

Selected specimens : Queensland. Cook District: near Mt

Emerald, SW of Walkamin, Jul 1998, Bean 13751 (BRI);

Mt Mulligan, 0.5 km SE of dam on summit of Mt, c. 40 km

NW of Dimbulah, Apr 1985, Clarkson 5865 (BRI); Toys

Creek, W of Herberton, Powerline access road, Feb 1996,

Forster PIF18442 (BRI). North Kennedy District: TR245,

Snubby Ck, near Ravenshoe, Apr 1999, McDonald 2 (BRI);

11 km NE of Innot Hot Springs, on the Silver Valley road,

Jan 2001, Wannan 2034 & Younghusband (BRI).

Distribution and habitat: Found on the western

parts of the Atherton Tableland, and on Mt

Mulligan further north (Map 2). It grows on

granite or sandstone ridges in shrubland.

Phenology: Flowers mostly from April to July,

but also in January and February.

Affinities: Cryptandra debilis is related to C. amara

var. floribunda, but differs by the often glabrous

branchlets (vs. always hairy), bract outer surface

glabrous (vs. stellate hairy), acute to mucronate

leaf apices, the longer calyx (both tube and

lobes), the style 1.1-1.3 mm long with dense

hairs on lower half (vs. 0.6-0.7 mm long,

glabrous throughout), and the schizocarp only

about 20% inferior (vs. 60-70%).

Etymology: From the Latin debilis, meaning

weak or feeble. This is a reference to the small

stature of the plant, compared with other related

species.

2. Cryptandra amara Sm. in Rees, Cycl. 10,

sect. 2 (1808). Type: New South Wales,

in the neighbourhood of Port Jackson,

undated, J. White s.n. (holo: LINN, n.v.,

microfiche 294.1).

There are two varieties treated separately in the

key to species.

2a. Cryptandra amara var. amara

Cryptandra amara var. longiflora F.Muell.

ex Maiden & Betche, Proc. Linn. Soc.

New South Wales 28: 905 (1904). Type:

not cited.

Cryptandra sp. (Thulimbah C. Schindler 6)

in Bean (2002)

Cryptandra sp. 3, in Briggs & Leigh (1996)

Shrub 0.3-0.9 m high, lateral branchlets 0.5-2

cm long, sometimes terminating in a spine.

Young branchlets with small stellate hairs only.

Older branches glabrescent. Stipules connate

920

Austrobaileya 6 (4): 917-940 (2004)

Fig. 1 . Cryptandra debilis. A. flowering branchlet (McDonald 2); B. side view of flower, showing predominantly simple hairs

on calyx and very short bracts ( McDonald 2).

Bean, new species of Cryptandra & Stenanthemum

on lower side of petiole, persistent, triangular,

0.9-1.3 mm long. Leaves only rarely in

fascicles (1(—3) per node), petioles 0.2-0.4 mm

long; lamina oblanceolate or obovate, 2.2-6.0

mm long, 0.8-2.3 mm wide, 2-4 t im es longer

than broad, apex obtuse, lamina flat or margins

recurved; upper surface green, glabrous; lower

surface green, glabrous or with a few simple

hairs on midrib. Inflorescences 1- flowered,

anthopodia 0.4-1.2 mm long; floral bracts 5-7,

orbicular, inner ones 0.7-0.8 mm long, brown,

outer surface glabrous, apex obtuse, margin

with cilia c. 0.05 mm long. Calyx white to

creamy; calyx tube cylindrical to urceolate,

0.7-1.5 mm long, with dense small stellate

hairs throughout, 0-20% of tube obscured by

bracts; calyx lobes erect to spreading, 0.9-1.3

mm long, with the same indumentum as the

tube. Petals hooded, white, protruding 0.4-0.6

mm beyond calyx tube. Disc densely stellate

hairy, obscurely 10-lobed. Style 0.7-0.9 mm

long, glabrous, stigma entire or 3-lobed.

Schizocarps ellipsoidal, 3-locular, c. 2.4 mm

long, 30-40% inferior, with ovary roof sparsely

stellate-hairy, disc forming a narrow densely

stellate-hairy annulus at the base of the

persistent calyx, mericarps dehiscent by a

ventral slit. Seeds not seen.

Selected specimens: Queensland. Darling Downs District:

c. 1 km ENE of Gambubal Forest Station, E of Warwick, Oct

1996, Bean 10987 (BRI, MEL, NSW); Bald Rock Creek,

Mt Norman fire trail, Girraween N.R, Aug 1995, Forster

PIF17609 & Figg (BRI, CANB, MEL); Rhumbalara Railway

crossing, Fletcher Lane, Fletcher, Aug 1996, Grimshaw

PG2544 & Bean (BRI). Moreton District: Double Peak,

Mt Ballow area, McPherson Range, Sep 1990, Forster

PIF7421 & Leiper (BRI, CANB, K, MEL, NSW). Leichhardt

District: Boyd Creek, SF46, c. 70 km W of Taroom, Sep

2002, Bean 19263 (BRI).

Distribution and habitat : In Queensland, C.

amara var. amara is known mainly from the

‘Granite Belt’ surrounding the town of

Stanthorpe, but also occurs disjunctly at Mt

Ballow, and near Taroom (Map 3). It inhabits

rocky ridges and mountaintops on sandstone,

granite or other acidic substrates.

Phenology: Flowers from August to October;

fruits in October.

Notes: This taxon sometimes has spinose lateral

branchlets, and hence it can be confused with

C. armata.

921

3. Cryptandra amara var. floribunda Maiden

& Betche, Proc. Linn. Soc. New South

Wales 29: 736 (1905). Types: Howell,

July 1904, J.L. Boorman (syn: BRI!,

NSW?); Stanthorpe, Queensland, July

1904, J.L. Boorman (syn: NSW?).

Shrub to 0.6 m high, lateral branchlets 1-4 cm

long, not terminating in a spine. Young

branchlets with simple adpressed hairs

overlying small stellate hairs. Older branches

glabrescent. Stipules connate on lower side of

petiole, persistent, narrowly triangular, 1.0-1.3

mm long. Leaves often in fascicles (1-8 per

node), petioles 0.2-0.3 mm long; lamina terete,

linear, 1.8-3.5 mm long, 0.4-0.6 mm wide, 3-8

times longer than broad, apex obtuse, margins

strongly revolute; upper surface green,

glabrous, sometimes muricate; lower surface

(rarely visible) white, indumentum dense

throughout, with simple hairs overlying small

stellate hairs. Inflorescences 1-3 flowered,

anthopodia 0.3-0.5 mm long; floral bracts 4-6,

broadly ovate to orbicular, 1.0-1.2 mm long,

brown, outer surface stellate hairy, apex obtuse,

margin entire or with cilia 0.05-0.1 mm long.

Calyx white; calyx tube campanulate, 0.8-1.0

mm long, indumentum dense throughout, with

simple adpressed hairs overlying small stellate

hairs, 0-20% of tube obscured by bracts at

anthesis; calyx lobes erect, 0.7-0.9 mm long,

indumentum dense throughout, with simple

adpressed hairs overlying small stellate hairs.

Petals hooded, white, protruding 0.3-0.5 mm

beyond calyx tube. Disc densely stellate hairy,

obscurely 10-lobed. Style 0.6-0.7 mm long,

glabrous throughout, stigma entire. Schizocarps

ellipsoidal, 2.0-2.6 mm long, 3-locular, 60-70%

inferior, ovary roof with sparse stellate

indumentum, disc forming a narrow densely

stellate-hairy annulus at the base of the

persistent calyx, mericarps dehiscent by a

ventral slit. Seeds flattened, elliptical in outline,

c. 1.2 mm long excluding aril. Aril white,

terminal.

Selected specimens: Queensland. Darling Downs District:

Portion 90, Wyberba, Aug 1995, Forster 17587 & Figg

(BRI); Ballandean, Oct 1933, White 9397 (BRI).

Distribution and habitat: Widespread in New

South Wales, but in Queensland it is confined

to the ‘Granite Belt’, around Stanthorpe (Map 2).

It grows on granitic hills and ridges with

shallow soil.

922

Phenology: Flowers are recorded from August

to October.

Notes: This taxon is amply distinct from C.

amara var. amara , and should be accorded

species rank. Walsh & Udovicic (1999)

foreshadowed this, and suggested that C. tomentosa

Lindl. may be the appropriate name.

4. Cryptandra armata C.T.White &

W.D.Francis, Proc. Roy. Soc. Queensland

33: 153 (1922). Type: Queensland.

Darling Downs District: Barakula, a few

mil es north of Chinchilla, July 1919, J.E.

Young s.n. (holo: BRI).

Shrub to 1.5 m high, lateral branchlets 0.5-1.5

mm long, terminating in a spine. Young

branchlets with simple adpressed hairs

overlying small stellate hairs. Older branches

glabrescent. Stipules connate on lower side of

petiole, persistent, narrowly triangular, 0.9-1.4

mm long. Leaves often in fascicles (1-5 per

node), petioles 0.3-0.5 mm long; lamina

oblanceolate, obovate or spathulate, 1.4-5.2

mm long, 0.4-1.4 mm wide, 2-5 times longer

than broad, apex obtuse or acute, margins

recurved; upper surface green, glabrous; lower

surface green, glabrous or with a few simple

hairs on midrib. Inflorescences l-(2-) flowered,

anthopodia 0.5-1 mm long; floral bracts

several, orbicular, 1.3-1.6 mm long, brown,

outer surface glabrous, apex obtuse, margin

with cilia 0.05-0.15 mm long. Calyx white to

creamy; calyx tube cylindrical to campanulate,

2.5-3 mm long, with dense adpressed simple

hairs overlying small stellate hairs, 15-25% of

tube obscured by bracts at anthesis; calyx lobes

erect to spreading, 1-1.6 mm long, with the

same indumentum as the tube. Petals hooded,

white, protruding 1-1.2 mm beyond calyx tube.

Disc densely stellate hairy. Style 1.5-2.8 mm

long, with dense stellate hairs on lower third;

stigma entire or 3-lobed. Schizocarps

ellipsoidal, 3-locular, 3-3.5 mm long, ovary

superior or 10% inferior, ovary roof with sparse

stellate indumentum, disc forming an annulus

of dense stellate hairs at base of the persistent

calyx, mericarps dehiscent by a ventral slit.

Seeds flattened, elliptical in outline, c. 2.3 mm

long excluding aril. Aril white, terminal. Fig. 2.

Selected specimens : Queensland. Port Curtis District:

Castletower N.P., east slopes Many Peaks Range, Feb 1995,

Austrobaileya 6 (4): 917-940 (2004)

Forster PIF16331 (BRI); Castletower N.P, site 297, Oct

1995, Thompson MIR87 & Price (BRI). Maranoa District:

“Boxleigh”, S of Surat, Aug 2001, Bean 17765 (BRI);

Chesterton Range N.P., c. 15 km E of Wineba house, Apr

1998, Dollery s.n. (BRI). Burnett District: on SF130, 2

km NW of Nantglyn, Sep 1989, Forster 5738 & Bean (BRI);

northern boundary of SF132, 35 km SSW of Mundubbera,

Jul 1997, Halford Q3305 & Holland (BRI); “Narayan”, Bull

Paddock, Jul 1968, McHarg H529 (BRI). Darling Downs

District: 3.7 km along Booroondoo Rd, S of Moonie, Sep

1997, Bean 12368 (BRI); 11.6 km along Boondandilla Rd,

W of Milmerran, Oct 1998, Bean 13903 (BRI); Spinifex

Heath, Wondul Range, Aug 1995, Forster 17508 & Figg

(BRI); SF132, c. 8.5 km WNW of Limevale, Oct 1994,

Sparshott KMS404 & Bean (BRI). New South Wales. North

Western Slopes: Severn River, 16.1 km from Bukkulla, c.

21 km SE of Ashford, Oct 1990, Coveny 14582 (BRI, NSW).

Distribution and habitat: C. armata is found

as far north as Gladstone, west to Morven and

south to Ashford in New South Wales (Map

2). It grows in sandy soil (sometimes skeletal),

on granite or sandstone substrate.

Phenology: It flowers from July to September,

and fruits from September to February.

Notes: C. armata is the only consistently

spinose species in northern Australia. The

specimen from near Ashford (Coveny 14582)

is the first record of the species from New South

Wales.

5. Cryptandra orbicularis A.R. Bean sp. nov.

affinis C. longistamineae F.Muell., sed

folia orbicularia usque reniformibus

pagina inferiore glabra vel glabriuscula,

calycis tubo pilis sparsis usque densis

praedito et petalis brevibus differens.

Typus: Queensland. Leichhardt District:

Expedition National Park, NW of

Taroom, Robinson Creek headwaters, 14

September 2000, PI. Forster PIF26173,

R. Booth & F. Carter (holo: BRI; iso:

MEL, NSW).

Cryptandra sp. (Isla Gorge P. Sharpe 627)

in Bean (2002)

Shrub 0.3-0.9 m high, lateral branchlets 1-5

cm long, not terminating in a spine. Young

branchlets with small white stellate hairs only,

or somet im es also with a few simple patent to

adpressed hairs. Older branches glabrescent.

Stipules connate on lower side of petiole,

persistent, narrowly triangular, 0.9-1.2 mm

long. Leaves usually solitary, but occasionally

in fascicles of up to 4, orbicular, reniform, or

Bean, new species of Cryptandra & Stenanthemum

923

Fig. 2. Cryptandra armata. A. flowering branchlets with spinose tips (Bean 17765); B. mature schizocarp (Forster 5738 &

Bean).

924

Austrobaileya 6 (4): 917-940 (2004)

broadly obovate, 1.3-2.5 mm long, 1.3-2.2 mm

wide, 0.8-1.3 times longer than broad, apex

obtuse, lamina flat or margins recurved; upper

surface green, glabrous; lower surface green,

glabrous or with a few simple hairs on midrib.

Inflorescences 1- flowered, anthopodiaO.3-0.6

mm long; floral bracts several, elliptical, inner

ones 1.2-1.5 mm long, brown, outer surface

stellate-hairy, apex obtuse, margin with cilia

0.05-0.15 mm long. Calyx white to creamy;

calyx tube cylindrical to campanulate, 0.9-1.1

mm long, with sparse to dense small stellate

hairs throughout, or sometimes almost

glabrous, 50-100% of tube obscured by bracts;

calyx lobes spreading, 1.3-1.5 mm long, with

a mixture of adpressed simple hairs and small

stellate hairs. Petals hooded, white, protruding

0.8-1.1 mm beyond calyx tube. Disc densely

stellate hairy. Style 1.3-2.0 mm long, glabrous

throughout, distinctly 3-lobed. Mature

schizocarps not seen. Immature schizocarps

ellipsoidal, 50% inferior, ovary roof sparsely

stellate hairy, disc forming a narrow densely

stellate-hairy annulus at the base of the

persistent calyx. Seeds not seen. Fig. 3.

Selected specimens : Queensland. Leichhardt District:

Precipice N.P., catchment of Precipice Ck, Sep 1996, Forster

PIF19741 (BRI, MEL); 38 km NE of Taroom, ‘Spring Creek’

station, Oct 1996, Halford Q3181 (BRI); Isla Gorge, c. 28

km SW of Theodore, Aug 1973, Sharpe 627 & Hockings

(BRI).

Distribution and habitat: C. orbicularis is

confined to south-eastern Queensland, between

Cracow and Rolleston (Map 2). It grows on

sandstone ridges in shrubby eucalypt woodland.

Phenology : Flowers recorded from August to

October.

Affinities: Cryptandra orbicularis is clearly

related to C. longistaminea, but differs by the

leaves orbicular to reniform (vs. elliptical to

obovate), the lower leaf surface green, glabrous

or with a few simple hairs on midrib (vs. white,

very densely hairy throughout); calyx tube with

sparse to dense stellate hairs (vs. glabrous);

calyx lobes with a mixture of adpressed simple

hairs and small stellate hairs (vs. stellate hairy

throughout, sometimes with a few simple

adpressed hairs); style 1.3-2.0 mm long (vs.

2.0-2.3 mm long); and petals protruding

0.8-1.1 mm beyond calyx tube (vs. 1.4-1.8 mm

beyond calyx tube).

Etymology: the epithet refers to the frequently

orbicular leaves in this species.

6. Cryptandra longistaminea F.Muell.,

Fragm. 3: 64 (1862). Type: Severn River,

New England, undated, C. Stuart s.n.

(holo: ?MEL, n.v.).

Shrub to 1 m high, lateral branchlets 1-5 cm

long, not terminating in a spine. Young

branchlets with small white stellate hairs only,

each hair c. 0.1 mm diameter. Older branches

glabrescent. Stipules connate on lower side of

petiole, persistent, narrowly triangular, 0.6-1.1

mm long. Leaves usually solitary, but

occasionally in fascicles of up to 4, petioles

0.2-0.6 mm long; lamina elliptical to obovate,

1.8-4.0 mm long, 0.9-1.7 mm wide, 1.5-2.5

times longer than broad, apex acute, margins

recurved; upper surface green, glabrous; lower

surface white, densely stellate-hairy, with a few

erect simple hairs on midrib. Inflorescences

1- flowered, anthopodia 0.3-0.8 mm long;

floral bracts several, elliptical, to 1.0-1.2 mm

long, brown, outer surface glabrous, apex

obtuse, margin with cilia 0.05-0.1 mm long.

Calyx white to creamy; calyx tube cylindrical

to campanulate, 1.0-1.3 mm long, glabrous,

10-70% of tube obscured by bracts; calyx lobes

erect to spreading, 1.7-2.0 mm long, with a

thin layer of small stellate hairs throughout,

and somet im es with a few adpressed simple

hairs at the apex. Petals hooded, white,

protruding 1.4-1.8 mm beyond calyx tube. Disc

densely stellate hairy. Style 2.0-2.3 mm long,

glabrous throughout, stigma distinctly trifid.

Mature schizocarps not seen. Immature

schizocarps ellipsoidal, c. 50% inferior, ovary

roof with sparse stellate indumentum, disc

forming an annulus of dense stellate hairs at

the base of the persistent calyx. Seeds not seen.

Selected specimens: Queensland. Port Curtis District: near

tributary of East Boyne River, Many Peaks Range, Jun 1995,

Thompson CAL325 & Turpin (BRI). Darling Downs

District: 9 km W of Bringalily Forestry lookout tower, Sep

1992, Forster PIF11639 & Machin (BRI, MEL). Burnett

District: 7.8 km W of Cynthia, S of Monto, Sep 1999, Bean

15313 & McDonald (BRI); about 13.5 km SSE of Allies

Creek, Jul 1998, Pollock ABP663 & Dean (BRI). Moreton

District: 1 km E of Swanbank Power station, Nov 1988,

Bostock & Forster s.n. (BRI); Buchanans Fort, Christmas

Creek area, Sep 1995, Forster PIF17675 & Leiper (BRI,

CANB).

Bean, new species of Cryptandra & Stenanthemum

925

Fig. 3. Cryptandra orbicularis. A. flowering branchlet (Pollock 1261); B. flowers, showing the bracts, recurved calyx lobes

and relatively long petals (Pollock 1261).

926

Distribution and habitat : Found in south¬

eastern Queensland, south from about

Gladstone (Map 3). It is widespread in New

South Wales.

Phenology : Flowers recorded from June to

September; fruits in November.

Notes: Some specimens from the Burnett

District are atypical, because of the dense

simple hairs on the upper leaf surface. It is not

yet known whether these specimens represent

an unnamed taxon.

7. Cryptandra lanosiflora F.Muell., Fragm.

3: 65 (1862). Type: Severn River, New

England, undated, C. Stuart s.n. (syn:

?MEL, n.v.)\ Mt Mitchell towards the

Clarence river, undated, H. Beckler (syn:

?MEL, n.v.).

Shrub to 0.6 m high, lateral branchlets 1-3 cm

long, not terminating in a spine. Young

branchlets with simple adpressed hairs

overlying small stellate hairs. Older branches

glabrescent. Stipules connate on lower side of

petiole, persistent, narrowly triangular, 0.9-1.8

mm long. Leaves in fascicles (2-7 per node),

petioles 0.2-0.5 mm long; lamina terete,

ellipsoidal, 1.3-3.2 mm long, 0.6-0.8 mm

wide, 2-5 times longer than broad, apex obtuse

or acute, margins strongly re volute; upper

surface green, glabrous, muricate, or with short

simple hairs; lower surface (rarely visible)

white, indumentum dense throughout, with

simple hairs overlying small stellate hairs.

Inflorescences 1- flowered, anthopodiaO.3-0.5

mm long; floral bracts several, orbicular, 1.8-2.1

mm long, brown, outer surface glabrous or with

a few simple hairs, apex obtuse, margin with

cilia 0.1-0.15 mm long. Calyx white; calyx tube

ovoid, 1.5-1.8 mm long, with dense tangled

simple hairs, 50-75% of tube obscured by bracts

at anthesis; calyx lobes erect, 1.2-1.3 mm long,

with the same indumentum as the tube. Petals

hooded, white, protruding 0-0.3 mm beyond

calyx tube. Disc densely stellate hairy, obscurely

10-lobed. Style 0.6-0.8 mm long, glabrous

throughout, stigma entire or 3-lobed.

Schizocarps obovoid, 3-locular, 2.3-2.6 mm

long, 60-70% inferior, ovary roof with sparse

stellate indumentum, disc forming a narrow

densely stellate-hairy annulus at the base of the

persistent calyx, mericarps dehiscent by a

ventral slit. Seeds flattened, elliptical in outline,

Austrobaileya 6 (4): 917-940 (2004)

1.2-1.4 mm long excluding aril. Aril white,

terminal.

Selected specimens: Queensland. Darling Downs District:

upper reaches Bald Rock Creek, Girraween N.R, Sep 1993,

Forster 13833 & Bean (BRI); Mt Jibbenbar, Sundown N.R,

34 km WSW of Stanthorpe, Sep 1996, Halford Q2949 (BRI);

Wyberba, Bald Rock Creek, Oct 1963, Pedley 1549 (BRI).

Distribution and habitat: In Queensland,

confined to the Stanthorpe area. It also occurs

on the northern tablelands of N.S.W. It grows

on granite hills in eucalypt woodland.

Phenology: Flowers and fruits recorded for

September and October.

Notes: C. lanosiflora is immediately

distinguishable by the dense white woolly hairs

on the calyx.

8. Cryptandra propinqua Fenzl in Endl.,

Enum. PL 23 (1837). Type: “New South

Wales”, A. Cunningham (?W, n.v.).

Shrub 0.6-1 m high, lateral branchlets 1-3 cm

long, not terminating in a spine. Young

branchlets with simple adpressed hairs

overlying small stellate hairs. Older branches

glabrescent. Stipules connate on lower side of

petiole, persistent, narrowly triangular, 1.0-1.5

mm long. Leaves often in fascicles (1-5 per

node), petioles 0.2-0.3 mm long; lamina terete,

ellipsoidal, 1.2-3.5 mm long, 0.4-0.7 mm

wide, 2.5-7 times longer than broad, apex

obtuse, margins strongly revolute; upper surface

green, glabrous, sometimes muricate; lower

surface (rarely visible) white, indumentum

dense throughout, with simple hairs overlying

small stellate hairs. Inflorescences 1- flowered,

anthopodia 0.3-0.6 mm long; floral bracts

c. 15, ovate to elliptical, inner ones 2.9-3.8 mm

long, brown, outer surface glabrous, apex acute,

margin with cilia 0.1-0.2 mm long. Calyx

white; calyx tube campanulate, 2.0-2.3 mm

long, indumentum sparse throughout, with

small stellate hairs only, 100% of tube obscured

by bracts at anthesis; calyx lobes erect, 1.8-2.7

mm long, indumentum dense throughout, with

simple adpressed hairs overlying small stellate

hairs, lobes partially obscured by bracts at

anthesis. Petals hooded, white, protruding 0.2-0.5

mm beyond calyx tube. Disc densely stellate

hairy, obscurely 10-lobed. Style 1.0-3.3 mm

long, glabrous throughout or stellate-hairy on

lower half, stigma 3-lobed. Schizocarps broadly

Bean, new species of Cryptandra & Stenanthemum

ellipsoidal, 2.8-4 mm long, 3-locular, ovary

roof with sparse stellate indumentum, disc

forming a narrow densely stellate-hairy annulus

at the base of the persistent calyx, mericarps

dehiscent by a ventral slit. Seeds not seen.

Selected specimens: Queensland. Warrego District: near

Bouden’s Dam, Chesterton Range N.R, Aug 2001 ,Dollery

280 (BRI).Maranoa District: Mt Moffatt N.R, fire

monitoring site 5, May 1997, Addicott MM45 (BRI).

Leichhardt District: Planet Downs Pastoral holding, adjacent

to Planet Creek, Mar 1998, Brushe JB 1518 (BRI). Burnett

District: S.R 132, 9 km ESE of Brovinia, Jun 1997, Bean

12037 (BRI). Da rti ng Downs District: Cecil Plains, Jun

1962, Hockings s.n. (BRI).

Distribution and habitat: Widespread from

Springsure to Inglewood, and west to Morven

(Map 1). It is also widespread in New South

Wales. It grows on sandy soils derived from

sandstone.

Phenology: Flowers recorded from June to

September, with one record in December.

9. Cryptandra rigida A.R.Bean sp. nov. affinis

C. propinquae Fenzl sed ramulis pilis

stellatis tantum praeditis, habitu rigido

invenusto, bracteis floralibus obtusis

interioribus sub anthesi totum calycis tubi

haud tegentibus et calycis tubo maximam

partem glabro differens. Typus:

Queensland. Burnett District: “Cooya”,

west of junction of Barambah and

Boonara Creeks, 17 July 1996, P.

Grimshaw 2486 & R. Price (holo: BRI;

iso: MEL).

Cryptandra sp. (NgungunL.S. Smith 13973)

in Bean (2002).

Shrub 0.4-1 m high, lateral branchlets 1-2.5

cm long, not terminating in a spine. Young

branchlets with small stellate hairs only. Older

branches glabrescent. Stipules connate on lower

side of petiole, persistent, narrowly triangular,

0.9-1.6 mm long. Leaves in fascicles (2-6 per

node), petioles 0.2-0.5 mm long; lamina terete,

ellipsoidal to lanceolate, 1.7-3.0 mm long, 0.4-0.9

mm wide, 2.5-6 times longer than broad, apex

obtuse, margins strongly revolute; upper surface

green, glabrous; lower surface (rarely visible)

white, indumentum dense throughout, with

simple hairs overlying small stellate hairs.

Inflorescences 1- flowered, anthopodia0.3-0.5

mm long; floral bracts c. 10, elliptical, inner

ones 2.1-2.3 mm long, brown, outer surface

glabrous, apex obtuse, margin with cilia 0.1-0.15

927

Fig. 4. Cryptandra rigida. A. flowering branchlet ( Grimshaw

2486 & Price).

mm long. Calyx white; calyx tube cylindrical,

1.8-2.3 mm long, glabrous except for a few

stellate hairs near the lobes, 50-90% of tube

obscured by bracts at anthesis; calyx lobes erect,

1.6-2.0 mm long, indumentum rather sparse

throughout, predominantly small stellate hairs

with a few simple adpressed hairs. Petals

hooded, white, protruding 0.5-0.8 mm beyond

calyx tube. Disc densely stellate hairy, obscurely

lobed. Style 2.5-2.7 mm long, glabrous

throughout, stigma entire. Schizocarps broadly

ellipsoid, 2.3-2.6 mm long, 3-locular, 50%

inferior, ovary roof with sparse stellate

indumentum, disc forming a narrow densely

stellate-hairy annulus at the base of the

persistent calyx, mericarps dehiscent by a

ventral slit. Seeds flattened, elliptical in outline,

1.4-1.5 mm long excluding aril. Aril white,

terminal. Fig. 4.

Selected specimens: Queensland. Burnett District: Mt

Lorna, 3 km W of Toondahra HS, Aug 1988, Forster 4637

(BRI, CANB); Campbell Creek, W of Mt Brian, Nov 1996,

Grimshaw 2621 & Turpin (BRI). Wide Bay District: 1.5

km SSWof Biggenden Bluff, Mt Walsh N.P., Sep 2002, Bean

19229 (BRI) Moreton District: Ngungun, NE comer below

sum mi t basin, Jul 1968, Smith 13973 (BRI).

Distribution and habitat : Known from several

rhyolitic or granitic mountains in the far south

east of Queensland (Map 1).

Phenology: Llowers recorded from July to

September; fruits in November.

928

Austrobaileya 6 (4): 917-940 (2004)

Affinities : Cryptandra rigida is clearly related

to C. propinqua, from which it differs by the

branchlets having stellate hairs only (vs. simple

+ stellate); the rigid unattractive habit; the

obtuse inner floral bracts 2.1-2.3 mm long, not

covering whole of calyx tube at anthesis (vs.

acute inner floral bracts 2.9-3.8 mm long,

covering all of calyx tube and partly covering

calyx lobes); the mostly glabrous calyx tube;

and the predominantly stellate indumentum of

the calyx lobes.

Etymology : the epithet refers to the rigid

branchlets, that make field encounters with the

species somewhat unpleasant.

10. Cryptandra ciliata A.R. Bean sp. nov.

affinis C. propinquae Fenzl sed folia

conspicue fasciculatis 4-11 per nodum,

bracteis pallidis brunneis obtusis,

marginibus bracteae ciliis 0.4-0.6 mm

praeditis, et calycis tubo glabro differens.

Typus: Queensland. Leichhardt District:

28 km from Cracow on Nathan Gorge

road, 15 July 1990, P.I. Forster PIF7037

(holo: BRI; iso: AD, CANB, K, MEL,

NSW, distribuendi ).

Cryptandra sp. (Gurulmundi G.W. Althofer

'8418) in Bean (2002)

Cryptandra sp. 1, in Briggs & Leigh (1996)

Shrub to 0.5 m high, lateral branchlets 10-50

mm long, not terminating in a spine. Young

branchlets with simple adpressed hairs

overlying small stellate hairs. Older branches

glabrescent. Stipules connate on lower side of

petiole, persistent, triangular with slender apex,

1.3-1.8 mm long. Leaves in fascicles (usually

4-11 per fascicle), petioles 0.1-0.2 mm long;

lamina terete, linear, 1.7-2.6 mm long, 0.4-0.5

mm wide, 4-5.5 times longer than broad, apex

obtuse, margins strongly revolute; upper surface

green, glabrous, sometimes muricate; lower

surface (rarely visible) white, with dense stellate

hairs and simple hairs. Inflorescences

1-flowered, anthopodia 0.5-0.8 mm long; floral

bracts c. 13, obovate, inner ones 1.7-2.1 mm

long, pale brown, apex obtuse, margin with cilia

0.4-0.6 mm long. Calyx white to brown; calyx

tube campanulate, 1.8-2.0 mm long, glabrous

except adjacent to calyx lobes, 100% of tube

obscured by bracts at anthesis; calyx lobes erect

to spreading, 1.4-1.7 mm long, with dense

white simple hairs. Petals hooded, white,

protruding 0.5-0.6 mm beyond calyx tube. Disc

densely stellate hairy, obscurely 10-lobed. Style

0.7-1.2 mm long, glabrous throughout, stigma

entire. Schizocarps ellipsoidal, 3-locular,

2.4-2.9 mm long, 50-65% inferior, ovary roof

with sparse stellate indumentum, disc forming

an annulus of dense stellate hairs at the base of

the persistent calyx, mericarps dehiscent by a

ventral slit. Seeds flattened, oblong in outline,

I. 8-1.9 mm long excluding aril. Aril white,

terminal. Fig. 5.

Selected specimens : Queensland. Leichhardt District:

Cracow-Taroom road, S of “Fairyland”, Sep 1990, Bean

2302 (BRI); Gwambegwine, Ruined Castle Creek catchment,

Sep 1996, Forster 19653 (BRI); 16 km SSW of Cracow

township, Jul 1963, Lazarides 6948 (BRI).

Distribution and habitat : C. ciliata is

sporadically distributed from Barakula State

Forest to west of Theodore (Map 1). It grows

on sandstone ridges and slopes in eucalypt

woodland.

Phenology : flowers and fruits recorded from

July to September.

Affinities : Cryptandra ciliata is allied to

C. propinqua but differs the conspicuously

fasciculate leaves with 4-11 per node, the pale

brown obtuse bracts up to 2.1 mm long (vs.

dark brown, acute, 2.9-3.8 mm long for

C. propinqua ), bract cilia 0.4-0.6 mm long (vs.

0.1-0.2 mm long), calyx tube 1.4-1.7 mm long,

glabrous (vs. 1.8-2.7 mm long, stellate hairy)

and calyx lobes with simple adpressed hairs

(vs. simple and stellate).

Etymology : the epithet refers to the conspicuously

ciliate bracts, which serve to distinguish it from

other Queensland species.

II. Cryptandra gemmata A.R. Bean sp. nov.

Folia 0.8-1.1 mm longa, inflorescentiae

terminales cymosaeque, 3-5-florae,

alabastra in axillis bractearum

plurimarum, bracteae apex aristatus,

discus floralis annularis dense pilosus,

mericarpia dehiscentia. Typus: Northern

Territory. Arnhem Land, 19 km E of

Jabiru, 18 April 1989, R.W. Johnson 4552

(holo: BRI; iso: AD, BISH, CONN, DNA,

MEL, NSW, distribuendi).

Bean, new species of Cryptandra & Stenanthemum

929

Fig. 5. Cryptandra ciliata.A. flowering branchlet, showing revolute leaves and ciliate bracts (Forster 7037); B. schizocarp,

showing persistent bracts and calyx, and dehiscent mericarps (Forster 19668).

930

Austrobaileya 6 (4): 917-940 (2004)

Shrub 0.2-0.3 m high, lateral branchlets 1-3

cm long, not terminating in a spine. Young

branchlets with small stellate hairs only. Older

branches glabrescent. Stipules connate on lower

side of petiole, persistent, narrowly triangular

with slender apex, 1.0-1.3 mm long. Leaves

in fascicles (3-9 per node), petioles absent;

lamina terete, ellipsoidal, 0.8-1.1 mm long,

0.3-0.4 mm wide, 2.5-3.5 times longer than

broad, apex obtuse, margins revolute; upper

surface green, glabrous, muricate; lower surface

(rarely visible) white, densely hairy.

Inflorescences cymose, 3-5 flowered, clustered

at very end of branchlets; anthopodia 0-0.2 mm

long, floral bracts several (3 or 4 fertile bracts

and 2 sterile), elliptical but with an aristate

apex, 1.9-2.2 mm long, brown, outer surface

glabrous, margin entire or with tiny cilia <0.1

mm long. Calyx white to creamy; calyx tube

cylindrical to urceolate, 1.6-1.7 mm long, with

dense small stellate hairs and scattered

adpressed simple hairs, 50-100% of tube

obscured by bracts at anthesis; calyx lobes erect,

0.8-1.0 mm long, with the same indumentum

as the tube. Petals hooded, white, protruding

0.5-0.6 mm beyond calyx tube. Disc densely

stellate hairy, obscurely lobed. Style 1.2-1.6

mm long, glabrous throughout, stigma entire.

Schizocarps obovoid, 3-locular, 2.5-2.9 mm

long, 60-70% inferior, ovary roof sparsely

stellate hairy, disc forming a narrow densely

stellate-hairy annulus at the base of the

persistent calyx, mericarps dehiscent by a

ventral slit. Seeds flattened, elliptical in outline,

1.8-2.0 mm long excluding aril. Aril white,

terminal. Fig. 6.

Additional specimen: Northern Territory. Arnhem Land,

19 km E of Jabiru, Apr 1989, Johnson 4618 (AD, BRI, DNA,

MEL, NSW).

Distribution and habitat : Known only from the

far north of the Northern Territory, near Jabiru.

It inhabits shrubland on sandstone pavement,

with little or no soil.

Phenology: flowers and fruits recorded for

April.

Affinities: Cryptandra gemmata is a distinctive

species with no obvious allies. The leaves are

only 0.8-1.1 mm long, and the bracts have an

aristate apex. Most significantly, the

inflorescences appear to be truly terminal, and

cymose. Each flower is surrounded by several

bracts, of which only two are sterile; the

remainder subtend flower buds and have the

potential to form new flowers. Hence it is

common to see various aged buds, flowers and

fruits on the same inflorescence.

Etymology: The epithet is from the Latin

gemmatus, meaning ‘provided with buds,

budded’. This refers to the development of buds

around the existing flowers and fruits on the

inflorescences.

12. Cryptandra pogonoloba A.R.Bean sp. nov.

ab omnibus aliis speciebus queenslandicis

stipulis in pagina superiore petiolorum

positis, bracteis floralibus apice aristato

ornatis et pilis longis simplicibus erectis

ad apicem calycis lobi differens. Typus:

Queensland. Cook District: Bulleringa

National Park, 80 km north-west of Mt

Surprise, Red River area, 23 April 1998,

P.I. Forster PIF22542 & R. Booth (holo:

BRI; iso: DNA, K, MEL, NSW, QRS,

distribuendi ).

Shrub 0.5-1 m high, lateral branchlets 0.3-1

cm long, not terminating in a spine. Young

branchlets with small stellate hairs only. Older

branches glabrescent. Stipules overlapping but

not connate on upper side of petiole, persistent,

triangular to ovate, with slender apex, 1.7-2.0

mm long. Leaves not in fascicles (1 per node),

petioles 0.3-0.9 mm long; lamina linear to

oblanceolate, 4.0-8.5 mm long, 0.8-1.2 mm

wide, 4.5-9 times longer than broad, apex

obtuse or acute, margins revolute; upper surface

grey, densely hairy, with stellate hairs or short

simple hairs; lower surface white, with dense

stellate hairs throughout and some simple hairs

along midrib. Inflorescences 1- flowered,

anthopodia absent; floral bracts several,

identical to stipules but larger, inner ones

2.0-2.3 mm long, brown, glabrous except for

a few simple hairs along midrib, apex aristate,

margin with cilia 0.3-0.6 mm long. Calyx white

to creamy; calyx tube ovoid to ellipsoid, 0.7-0.8

mm long, with dense small stellate hairs

throughout, bract apices 70-100% length of

tube, only base of tube obscured; calyx lobes

erect, 1.0-1.2 mm long, with dense stellate

hairs throughout and long erect simple hairs,

especially towards apex. Petals hooded, white,

protruding 0.2-0.4 mm beyond calyx tube. Disc

Bean, new species of Cryptandra & Stenanthemum

931

Fig. 6. Cryptandra gemmata. A. branchlet, showing fasciculate leaves and prominent stipules (Johnson 4552); B. schizocarp,

showing persistent calyx and dehiscent mericarps (Johnson 4552).

932

Austrobaileya 6 (4): 917-940 (2004)

Fig. 7. Cryptandra pogonoloba. A. flowering branchlet ( Forster 22542); B. dehisced mericarp, showing long erect hairs on

calyx, and seed (Forster 22608).

Bean, new species of Cryptandra & Stenanthemum

densely stellate hairy, lobing obscure. Style

0.6-0.8 mm long, glabrous, stigma entire.

Schizocarps ellipsoidal, 3-locular, 2.1-2.4 mm

long, 60-70% inferior, ovary roof densely

stellate-hairy, disc forming a narrow densely

stellate-hairy annulus at the base of the

persistent calyx, mericarps dehiscent by a

ventral slit. Seeds flattened, elliptical in outline,

1.6-1.9 mm long excluding aril. Aril white,

terminal. Fig. 7.

Selected specimens: Queensland. Cook District: 54 km

along Bulimba Station road, off Chillagoe-Wrotham Park

road, Jun 1991, Forster PIF8438 (AD, BRI, MEL);

Bulleringa N.P, 80 km NW of Mt Surprise, Apr 1998, Forster

PIF22608 & Booth (BRI); ditto, Forster PIF22642 & Booth

(BRI, DNA, MEL); ditto, Forster PIF22689 & Booth (BRI,

DNA, MEL, NSW, QRS).

Distribution and habitat : Cryptandra

pogonoloba is confined to north Queensland,

in the area north of Georgetown and west of

Chillagoe (Map 1). It grows on sandstone

ridges and slopes in eucalypt woodland.

Phenology : Flowers and fruits are recorded for

April and June.

Affinities : C. pogonoloba differs from all other

Queensland species by the stipules placed on

the upper side of the petiole, the floral bracts

with an aristate apex and the long erect simple

hairs at the apex of the calyx lobes. The upper

leaf surface is densely stellate-hairy, a feature

shared only by C. filiformis.

Etymology: the epithet refers to the

conspicuous erect simple hairs on the apex of

the calyx lobes (Gk. pogon - beard, and lobus -

lobe).

13. Cryptandra filiformis A.R.Bean sp. nov.

ab omnibus aliis speciebus a

Queenslandia cognitis inflorescentia

cymosa, absentia bractearum floralium et

stipulis filiformibus differens. Typus:

Queensland. Cook District: Mt Carbine

to Maytown, 46 km along Whites Creek

road, 20 April 2002, A.R. Bean 18766 &

K.R. McDonald (holo: BRI; iso: CANB,

MEL, NSW).

Shrub 0.5-0.7 m high, without distinct lateral

branchlets, not spinose. Young branchlets with

small stellate hairs only. Older branches stellate

hairy. Stipules connate on lower side of petiole,

933

persistent, thread-like, 2.8-4.4 mm long,

stellate-hairy throughout. Leaves not in

fascicles (1 per node), petioles 0.5-0.8 mm

long; lamina narrowly elliptic to narrowly

lanceolate, 7.5-30 mm long, 0.9-3.5 mm wide,

5-9 times longer than broad, apex acute,

margins recurved to re volute; upper surface

grey, with dense stellate hairs only; lower

surface white, with dense stellate hairs only.

Inflorescences 2-11- flowered, cymose;

anthopodia 0.7-1.2 mm long; floral bracts

absent. Calyx white to creamy; calyx tube

cylindrical to urceolate, 1.0-1.3 mm long, with

dense small stellate hairs throughout; calyx

lobes erect, 1.1-1.3 mm long, with dense

stellate hairs throughout. Petals hooded, white,

protruding 0.4-0.5 mm beyond calyx tube. Disc

densely stellate hairy, 5-lobed. Style 1.6-1.8

mm long, stellate hairy on proximal one-third,

otherwise glabrous, stigma entire. Schizocarps

obovoid, 3-locular, 3.0-3.8 mm long, 60-70%

inferior, ovary roof stellate-hairy, disc forming

a narrow densely stellate-hairy annulus at the

base of the persistent calyx, mericarps dehiscent

by a ventral slit. Seeds flattened, ellipsoidal,

2.2-2.4 mm long excluding aril. Aril white,

terminal. Fig. 8.

Selected specimens : Queensland. Cook District: 23 km E

of Forsay th on road to Einasleigh, Oct 2000, Addicott EPA851

& Newton (BRI); near Groganville, N of Chillagoe, Apr 2002,

Bean 18754 & McDonald (BRI); 26 km W of Einasleigh on

road to Forsayth, on top of Newcastle Range, Apr 1992,

Halford Q957 (BRI).

Distribution and habitat : C. filiformis is known

from two areas of north Queensland about 250

kilometres apart - on the Newcastle Range (W

of Einasleigh), and in the Maytown-

Groganville area (Map 3). These populations

are apparently disjunct. It grows on ridges and

plateaux on metamorphics or lateritised

sandstone.

Phenology : Llowers are recorded for April;

fruits in October.

Affinities: Cryptandra filiformis differs from

all other Queensland species by the thread-like

stipules, the complete absence of simple hairs,

the cymose inflorescence, and the absence of

floral bracts. It seems most closely related to

C. intratropica W.Litzg., a species from the

Kimberley region of Western Australia, which

also has strictly stellate indumentum,

934

Austrobaileya 6 (4): 917-940 (2004)

Fig. 8. Cryptandrafiliformis. A. flowering branchlet, showing revolute leaves and filiform stipules (Bean 18766); B. half-

lower, showing stellate-hairy disc and glabrous style (Bean 18766).

Bean, new species of Cryptandra & Stenanthemum

rudimentary bracts and a contracted cymose

inflorescence. While these characteristics are

unusual in Cryptandra, both C. filiformis and

C. intratropica display enough typical

Cryptandra character states (e.g. the annular

stellate-hairy disc, the conspicuous calyx tube

and the dehiscent mericarps) for them to be

regarded as members of Cryptandra.

Etymology : the epithet refers to the slender

thread-like stipules.

14. Stenanthemum argenteum A.R.Bean sp.

nov. affinitate ad S. leucophractum

(Schltdl.) Reissek, sed statura majore,

partibus hornotinis argenteis, foliis

longioribus, bracteis floralibus aristatis et

calycis tubo pilos et simplices et stellatos

ferenti differens. Typus: Queensland.

Cook District: The Pepperpot, Mount

Mulligan, c. 40 km NW of Dimbulah, 31

May 1985, J.R. Clarkson 5949 (holo:

BRI; iso: L, MEL, MO, NSW,

distribuendi).

Cryptandra sp. (Mt Mulligan J.R. Clarkson

5949) in Bean (2002).

Cryptandra sp. 2, in Briggs & Leigh (1996).

Shrub c. 2 m high, without distinct lateral

branchlets, not spinose. Young branchlets with

simple adpressed hairs only. Older branches

remaining hairy. Stipules on upper side of

petiole, not connate but sometimes overlapping,

persistent, very narrowly triangular with

slender apex, 2.0-3.1 mm long. Leaves solitary,

not in fascicles, petioles 2-3 mm long; lamina

oblanceolate to obovate, 7.5-16 mm long, 2.0-4.9

mm wide, 2.5-4.5 times longer than broad,

apex acute or obtuse, lamina flat or margins

incurved; upper surface green, glabrous; lower

surface white, with dense adpressed simple

hairs throughout. Inflorescences c. 7- flowered,

in a terminal head, surrounded by c. 3 bi-

aristate stipule-like bracts, and 1 or 2 spathulate

floral leaves that are densely hairy on both

surfaces. Llowers without anthopodia; floral

bracts 2 or 3, elliptical, 2.7-3.0 mm long,

brown, outer surface glabrous except along

midrib, apex obtuse, margin with cilia 0.2-0.25

mm long. Calyx white to creamy; calyx tube

cylindrical to urceolate, 2.5-3.0 mm long, with

mixture of long simple hairs and minute stellate

935

hairs, 80-100% of tube obscured by bracts;

calyx lobes erect, 1.9-2.0 mm long, with long

simple adpressed hairs throughout. Petals

hooded, white, protruding 1.1-1.4 mm beyond

calyx tube. Disc not apparent. Style c. 3.3 mm

long, glabrous throughout, stigma entire.

Schizocarps and seeds not seen. Fig. 9.

Additional specimen: Queensland. Cook District: Mt

Mulligan, c. 40 km NW of Dimbulah, Apr 1985, Clarkson

5765 (BRI).

Distribution and habitat : Known only from Mt

Mulligan, in north-eastern Queensland (Map 1).

It grows on sandstone pavement in heathland.

Phenology : Llowers are recorded for May.

Notes: S. argenteum is similar to S. leucophractum.

Both species have petiolate, obovate leaves,

stipules united above the petiole, a head-like

inflorescence with 1 or 2 specialised floral

leaves, and the disc obscure and glabrous.

However, S. argenteum differs by the greater

stature, the silvery new growth, the longer

leaves, the aristate floral bracts, and the calyx

tube with a mixture of simple and stellate hairs.

Conservation status: S. argenteum is currently

known only from Mt Mulligan. It is threatened

by its small population size and restricted area

of occupancy. Applying the IUCN guidelines

(IUCN. 2001), a category of “Vulnerable” is

recommended (VU Dl+2).

Etymology: The epithet refers to the silvery

branchlets and leaves (L. argenteus - silvery).

15. Stenanthemum scortechinii (L.Muell.)

Maiden & Betche, Proc. Linn. Soc. New

South Wales 27: 57 (1902); Cryptandra

scortechinii L.Muell., Australas. Chem.

Druggist 6 (69): 72 (1884). Type: on the

Severn [River], undated, B. Scortechini

(holo: ?MEL, «.v.).

Shrub 0.3-1 m high, lateral branchlets 1-8 cm

long, not spinose. Young branchlets with dense

stellate hairs only. Older branches remaining

hairy. Stipules on upper side of petiole, not

connate but overlapping, persistent, triangular,

2.2-3.8 mm long. Leaves solitary, not in

fascicles, petioles 1-1.9 mm long; lamina

lanceolate, 5-14 mm long, 1.7-3.9 mm wide,

3-6 times longer than broad, apex acute or

mucronate, margins recurved to re volute; upper

936

Austrobaileya 6 (4): 917-940 (2004)

Bean, new species of Cryptandra & Stenanthemum

Fig. 9. Stenanthemum argenteum. A. flowering branchlet

showing inflorescence with a single floral leaf ( Clarkson

5949); B. flowering branchlet, showing stipules and silvery

lower leaf surface ( Clarkson 5949); C. half-flower, showing

glabrous style and lack of obvious disc ( Clarkson 5949).

surface green, glabrous; lower surface white,

with dense stellate hairs throughout and

scattered simple hairs along midrib.

Inflorescences compound, each terminal ‘head’

comprising 5-6 shortly pedunculate clusters of

c. 6 flowers, each surrounded by very dense

mass of simple long woolly hairs. Clusters

subtended by 2 outer + 3 or 4 inner, orbicular

bracts, up to 5 mm long, brown, outer surface

glabrous except for simple and stellate hairs

along midrib, apex acute, margin entire. Calyx

white; calyx tube cylindrical, 0.9-1.0 mm long,

glabrous; individual flowers not subtended by

bracts; calyx lobes erect, 0.9-1.0 mm long, with

long simple hairs and small stellate hairs. Petals

hooded, white, protruding c. 0.6 mm beyond

calyx tube. Disc not apparent. Style 0.8-1.2 mm

long, glabrous throughout, stigma entire.

Schizocarps ellipsoidal, 3-locular, 2.7-3.0 mm

long, 90% inferior, ovary roof stellate-hairy,

mericarps indehiscent, chartaceous. Seeds

flattened, ellipsoidal, 1.9-2.0 mm long. Aril

absent.

Selected specimens’. Queensland. Darling Downs District:

upper reaches of Bald Rock Creek, Girraween N.R, Sep 1993,

Bean 6357 & Forster (BRI, CANB, MEL); Lyra, Nov 1959,

Blake 21092 (BRI, CANB); Stanthorpe, Jul 1904, Boorman

s.n. (BRI, NSW); 9 km NW of Ballandean, Murphys Ck,

Portion 91 Tartini Pty Ltd, Dec 1994, Halford Q2363 (BRI);

slopes near summit of Mt Norman, Sep 1977, Powell &

Armstrong s.n. (BRI, NSW).

937

Distribution and habitat : In Queensland,

S. scortechinii is confined to the Stanthorpe

district (Map 1), but is moderately widespread

in New South Wales. It grows on lower slopes

of hills in coarse sandy soils derived from

granite, in eucalypt forest with a heathy

understorey.

Phenology : Flowers and fruits recorded

between September and December.

Notes: S. scortechinii has chartaceous,

indehiscent mericarps. All other Stenanthemum

species have dehiscent mericarps. Indehiscent

mericarps are a regular feature of both

Trymalium and Spyridium, but in those genera,

the mericarps are very hard, thick and rugose.

Acknowledgements

I am grateful to Peter Bo stock for the Latin

diagnoses, Donovan Sharp for assistance with

the images, and Paul Forster for the numerous

helpful plant collections.

References

Bean, A.R. (2002). Rhamnaceae. In: R.J.F. Henderson (ed.).

Names and Distribution of Queensland Plants, Algae

and Lichens. Environmental Protection Agency:

Brisbane.

Briggs, J.D. & Leigh, J.H. (1996). Rare or Threatened

Australian Plants, 1995 Revised edition. CSIRO:

Canberra.

IUCN. (2001). IUCN Red List Categories and Criteria:

Version 3.1. IUCN Species survival Commission.

IUCN, Gland, Switzerland and Cambridge, UK.

Rye, B. (1995). New and Priority taxa in the genera

Cryptandra and Stenanthemum (Rhamnaceae) of

Western Australia. Nuytsia 10: 255-305.

Walsh, N.G. & Udovicic, F. (1999). Cryptandra. In: Flora

of Victoria, Volume 4, Dicotyledons, Cornaceae to

Asteraceae. Inkata Press: Melbourne.

938

Austrobaileya 6 (4): 917-940 (2004)

Map 1. Distribution of Cryptandra propinqua □ „ C. rigida* . C. cilia (a A ,

C. pogonolobaO , Sienanthemum argemeum ★ S. seortechinii + .

Bean, new species of Cryptandra & Stenanthemum

939

Map 2. Distribution of Cryptandra debilis k , C amara var .floribunda □ ,

C. armata % , C. oM/cu/ans* „

940

Austrobaileya 6 (4): 917-940 (2004)

MapJ, Distribution of Cryptandra amara var, amara • , C. hngistaminea A .

C, filiformis ★.

Rediscovery of Tylophora linearis P.I.Forst. (Apocynaceae:

Asclepiadoideae) from New South Wales, with revision of its conservation

status to vulnerable

Paul I. Forster, Doug Binns and Geoff Robertson

Summary

Forster, P.I., Binns, D. & Robertson, G. (2004). Rediscovery of Tylophora linearis RI.Forst. (Apocynaceae:

Asclepiadoideae) from New South Wales, with revision of its conservation status to vulnerable. Austrobaileya

6(4): 941-947. Newly discovered populations of Tylophora linearis (last collected in 1969) are documented

from central New South Wales. An amplified description and diagnostic illustrations are provided. The

conservation status of Tylophora linearis is reviewed with the recommendation that it be downgraded to

Vulnerable. Tylophora linearis is newly recorded as a host-plant for the butterfly Danaus chrysippus petilia.

Keywords: Tylophora linearis; Queensland flora; New South Wales flora; Apocynaceae; Asclepiadaceae;

Danaus chrysippus petilia; Vulnerable flora

Paul I. Forster, Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens, Mt

Coot-tha Road, Toowong, Queensland 4066, Australia, email: paul.forster@ epa.qld.gov.au

Doug Binns, State Forests of New South Wales, PO. Box J19, Coffs Harbour Jetty, New South Wales, 2450.

Australia, email: DougB@sf.nsw.gov.au

Geoff Robertson, New South Wales National Parks & Wildlife Service, PO. Box 2111, Dubbo, New South

Wales, 2830, Australia, email: Geoff.Robertson@npws.nsw.gov.au

Introduction

Tylophora linearis P.I.Forst. was named in the

most recent revision of this genus for Australia

(Forster 1992). A further concise account of

the species was provided in the ‘Flora of

Australia’ (Forster 1996). Only four gatherings

(including the type collection) have previously

been made of Tylophora linearis, in 1913,1915,

1960 and 1969. These four gatherings were all

made from widely scattered localities in

southern Queensland (near Glenmorgan)

through to southern New South Wales (near

Temora). The vegetation in all of these four

localities has been greatly altered since these

collection dates, and the species has been listed

as Endangered under the Australian

Commonwealth Environment Protection and

Biodiversity Conservation Act, 1999 ( EPBC ),

the Queensland Natu re Conservation Act, 1994

(NCA ) and the New South Wales Threatened

Species Conservation Act, 1995 ( TSCA ). The

paucity of adequate herbarium material for this

species has meant that no detailed illustrations

have been provided to date; an essential aid if

rapid identification is to be achieved. This

identification problem is compounded by the

Accepted for publication 10 May 2004

superficial similiarity of vegetative plants of

Rhyncharrhena linearis (Decne.) K.L.Wilson

and Marsdenia viridiflora R.Br.

A small number of populations of

Tylophora linearis have been recently located

in central New South Wales. This has enabled

information to be gathered about extant

distribution, abundance, habitat and phenological

patterns. It has also enabled provision of an

expanded morphological description, together

with illustrations. A new assessment of the

conservation status of Tylophora linearis is

undertaken using the IUCN criteria (Anon.

2001 ).

Materials and Methods

This paper is based on herbarium collections

at BRI and NSW, together with new collections

by D. Binns and G. Robertson (deposited in

BRI). The first of the new collections was made

as a result of pre-harvest survey in Eura State

Forest conducted as part of the standard survey

requirement for logging in State forests in

NSW. Although Tylophora linearis was one of

the target species for survey, its discovery was

partly serendipitous. Subsequently, on the

premise that the species may exhibit

942

Austrobaileya 6 (4): 941-947 (2004)

synchronous flowering over a broader area,

specific searches for flowering material were

conducted elsewhere. Searches were conducted

opportunistically in habitats in the Dubbo area

similar to that at the Eura S.F. site, and at a

selection of systematic flora survey sites in the

Pilliga forests where vegetatively similar plants

had been recorded previously. Most of these

previous records were sight records, but some

were based on vegetative vouchers held at State

Forests of New South Wales herbarium at Coffs

Harbour, NSW. Sites were selected for ease of

access and to represent a relatively wide

geographical spread. Opportunities were also

taken to search for the species during other flora

surveys being conducted at the time.

Taxonomy

Tylophora linearis P.I.Forst., Austral. Syst.

Bot. 5: 31 (1992). type: New South

Wales. Temora, November 1915, Dwyer

802 (holo: NSW).

Herbaceous twiner to c. 2 m long, latex clear.

Stems cylindrical, up to 3 mm diameter,

glabrous or with scattered indumentum;

internodes up to 100 mm long. Feaves

petiolate; petioles 1-7 mm long, c. 0.5 mm

diameter, channelled on top, with scattered to

sparse indumentum, extrafloral nectaries absent

from base. Feaf lamina linear-lanceolate, up to

100 mm long and 4 mm wide, glabrous or with

scattered indumentum, + concolorous, venation

+ obscure, tip acute, base attenuate to cuneate,

adaxial surface dark green, glabrous; abaxial

surface dark to medium green, glabrous or with

scattered indumentum. Cyme umbelliform, up

to 20 mm long and with up to 6 flowers;

peduncle cylindrical, up to 23 mm long and c.

0.5 mm diameter, glabrous or with scattered to

sparse indumentum; bracts linear to linear-

lanceolate, 1-3.5 mm long, 0.3-0.5 mm wide,

glabrous or with scattered indumentum.

Flowers 2-2.5 mm long, (6-) 18-22 mm

diameter; pedicels (3-) 5-8 mm long, glabrous

or with scattered indumentum. Sepals

lanceolate to lanceolate-ovate, weakly apiculate,

1.5-2 mm long, 1-1.6 mm wide, glabrous, base

of each sinus with 1 extrafloral nectary. Corolla

rotate, externally olive-green, internally dark

purple; tube 1-1.2 mm long, 1.5-2.8 mm

diameter, glabrous; lobes lanceolate to

lanceolate-ovate, 3.5-5.5 mm long, 1.7-2.5

mm wide, with short indumentum on internal

surface that is more concentrated towards tip.

Staminal corona 0.5-0.8 mm long, 1.5-2 mm

diameter, purple, each lobe rounded to flat-

topped, 0.5-0.8 mm long, 0.6-1.1 mm wide.

Staminal column c. 1 mm long, 1-1.3 mm

diameter; anther appendages truncate to

elliptic, 0.4-0.5 mm long, 0.3-0.5 mm wide;

alar fissure c. 0.2 mm long. Style-head

pentagonal globose, 0.5-0.7 mm long, c. 1 mm

diameter, green; ovaries c. 1 mm long and 1

mm diameter, glabrous. Pollinaria 0.2-0.3 mm

long, 0.2-0.5 mm wide; pollinia globose, held

horizontally to erect, 0.16-0.22 mm long,

0.12-0.18 mm wide; corpusculum oblong,

0.1-0.13 mm long, 0.04-0.05 mm wide;

caudicles 0.05-0.1 mm long, c. 0.02 mm wide.

Follicles fusiform, 95-100 mm long, c. 5 mm

diameter, glabrous, seeds not seen. Fig. 1.

Specimens examined : Queensland. Darling Downs

District: “Myall Park”, Glenmorgan, May 1960, Blake

21234 (BRI). New South Wales. Eura S.F., c. 18 km SSE of

Gilgandra, 31 0 51 ’ S, 148° 42’E, Mar 2003, Binns 9873 (BRI,

NSW); 2 mi les [3 km] SW of Mendooran, towards Dubbo,

Nov 1969, Coveny 2492 (NSW); Goobang N.P., 32° 41’S,

148° 17’E, May 2003, Robertson s.n. (BRI); Eura S.F., c.

18 km SSE of Gilgandra, 31° 51’S, 148° 42’E, May 2003,

Robertson s.n. (BRI); northern boundary of Goonoo S.F. &

Coolbaggie Nature Reserve, 31° 58’S, 148° 44’E, May 2003,

Robertson s.n. (BRI); Crow Mt, Barraba, Nov 1913, Rupp

(NSW); Pilliga West S.F., c. 30 km NW of Baradine, 30°

45’S, 149° 50’E, May 2003, Binns 9879 (BRI).

Habitat and distribution : Tylophora linearis

occurs most commonly in dense shrubland that

may be overtopped by eucalypts. At the

Mendooran site, Coveny noted that this species

grew in association with Acacia hakeoides,

A. lineata, Myoporum sp. and Casuarina sp.

At the newly documented localities in the

Dubbo area (Eura S.F., Goobang N.P. and

Goonoo S.F./Coolbaggie N.R.), the plants were

encountered among dense shrubland of Melaleuca

uncinata with scattered M. erubescens and other

shrub species, and in adjacent areas of

woodland to open woodland with M. uncinata

shrub understorey, on flat to gently undulating

landscapes at 300-400 m altitude. In Eura S.F.,

the woodland is variously of Eucalyptus crebra,

E. sideroxylon and Callitris glaucophylla, in

Goobang N.P. it is of E. microcarpa and in

Goonoo S.F./Coolbaggie N.R., the dominant

trees are E. crebra and E. dumosa. Tylophora

linearis individuals have been observed twining

Forster, Binns & Robinson, rediscovery of Tylophora linearis

943

Fig. 1. Tylophora linearis. A. habit of flowering stem, x 0.5. B. face view of flower, x 0.6. C. side view of flower, x 6. D. side

view of gynostegium. x 12. E. pollinarium. x 40. All from Binns 9873 (BRI). Del. W. Smith.

around the stems of Melaleuca uncinata, with

the occasional plant twining around Calytrix

tetragona, Acacia tindaleae, Lissanthe strigosa

and Leptospermum divaricatum. The critical

habitat defining feature in these areas appears

to be the presence of the dense understorey of

Melaleuca uncinata, however survey has been

biased towards these habitats. Recently another

location was found in Goobang National Park

in Eucalyptus fibrosa woodland where

Melaleuca uncinata is a minor component of

the understorey. At this locality Tylophora

linearis was found on Dodonaea viscosa and

Acacia doratoxylon.

In contrast, the collection from Pilliga

West S.F. was from woodland of E. pilligaensis

and Callitris glaucophylla, with a moderately

dense subcanopy and understorey of

Allocasuarina luehmannii and scattered Acacia

hakeoides. Plants were twining on A. luehmannii

stems. There is also a sight record of a flowering

specimen in Cumbil S.F. (30° 46’E, 149° 04’S,

Binns obs. 8 May 2003) from regrowth Callitris

glaucophylla forest with occasional

E. melanophloia and E. crebra. The

understorey comprised scattered shrubs and a

sparse grassy ground layer dominated by

Aristida ramosa, Digitaria diffusa, Aristida

944

Austrobaileya 6 (4): 941-947 (2004)

leichhardtiana, Whalleya subxerophylla and

Eulalia fulva. This site included more than 100

separate shoots over an area of about 0.5 ha,

and several shoots with senescent umbels, but

only a single flowering specimen. Mature

shoots were climbing on fallen dead branches,

shrubs of Geijera parviflora. Acacia deanei and

Hakea decurrens, and small trees of Callitris

glaucophylla.

Sight records of vegetatively similar

plants have been made over the past ten years

from 43 systematic survey sites over the

geographic range (28° 55’ -31° 57’S and 148°

40' - 150° 55’E). These sites represent a wide

range of habitats, but Callitris glaucophylla is

common in almost half and Allocasuarina

luehmannii is common in about 25% of sites.

Other dominant overstorey species include E. crebra,

E. populnea subsp. bimbil, Casuarina cristata,

E. albens and E. viridis. Melaleuca uncinata

is common in only two of these sites. If these

records actually do represent Tylophora.

linearis, as seems likely, then the habitat range

is much greater than indicated by the known

sites described above.

The recorded, historical latitudinal

distribution is from Glenmorgan in southern

Queensland in the north (last seen 1960) to

Goobang National Park in central New South

Wales in the south (Map 1).

Notes: The new collections of Tylophora

linearis have enabled a number of additions

and changes to the morphological description

of this species (cf. Forster 1992,1996). Perhaps

most significant of these is that plants of this

species do not form subshrubs, nor are they

particularly woody. Instead Tylophora linearis

appears to be an herbaceous twiner, a life form

that is widespread in Australian asclepiads,

particularly those that occur in non-rainforest

communities. Plants appear to be perennial, and

regenerate from a thickened rootstock. A

number of Australian asclepiads are geophytic

in lifeform, with the annually produced aerial

parts being shed in times of drought (e.g.

various Marsdenia spp., Rhyncharrhena

linearis, Nichols et al. 1991; Forster 1995).

Such a lifeform also tends to reduce the annual

timeframe when plants may be noticeable.

A few plants were partially excavated.

These were seen to have thickened vertical

rootstocks from ground level to a depth of

between 5 and 20 cm, where the vertical portion

joined a thinner horizontal rhizome. In the one

case where the latter was further excavated, it

was traced to another shoot. This suggests that

the species is clonal to at least some extent,

and may be extensively clonal. Large numbers

of shoots may represent few genets. Each

population included varying proportions of

flowering shoots with long internodes that

appear to rapidly elongate, and other, shorter

shoots with smaller leaves, where apical

elongation appears to have slowed or ceased.

It appeared that the latter stem type occurred

predominantly in open patches where no

supporting shrub stems were in close proximity.

This was especially evident at the Cumbil S.F.

site where numerous short shoots occurred on

the cleared road verge and longer flowering

shoots were restricted to adjacent forest. It is

possible that shoot elongation and subsequent

flowering is stimulated to some extent by a

shoot encountering a support for climbing.

When compared to the earlier collections,

recent collections have significantly larger and

longer leaves and larger flowers with longer

pedicels and corolla lobes. Whether these

differences are population based or more an

artefact of the process of specimen preparation

is unknown, but the latter seems likely. Several

of the early collections are of poor standard.

Floral anthesis is quite variable in this

plant and may be correlated with light. Flowers

close during the late afternoon with the corolla

lobes reflexing forward so that the staminal

column is obscured. Under bright light, the

corolla lobes reflex so that the corolla lobes lie

flat and the staminal column is fully exposed.

In one case, flowers which were closed at about

3 pm when collected, opened after brief (less

than 30 minutes) exposure to fluorescent light

later that evening. Flowers open non-

synchronously on different umbels with only

one or two flowers open at any one time on any

single umbel. These traits of floral behaviour

are not unusual in Tylophora and have been

observed in other species such as T. benthamii

Tsiang, T. colorata C.T.White, T. erecta

F.Muell. ex Benth., T.flexuosa R.Br., T. glabriflora

(Warb.) Schltr., T. grandiflora R.Br., T. rupicola

RI.Forst. and T. williamsii RI.Forst. (Forster

1992, 1994).

Forster, Binns & Robinson, rediscovery of Tylophora linearis

Flowering cues are unknown, but may be

partly related to rainfall. The first collection

from Eura S.F. was several weeks after brief

but relatively intense rain (estimated about 20

mm), following a long period of drought. Buds

and developing inflorescences were numerous

and no umbels had bare pedicels from which

flowers had fallen, suggesting that anthesis had

begun only a few days earlier. Three weeks later,

all flowers had fallen from that cohort of umbels

and another cohort had developed to the point

of anthesis. Further rain fell during that period.

Individuals of Tylophora linearis are

difficult to distinguish from those of

Rhyncharrhena linearis or Marsdenia

viridiflora R.Br. when only juvenile or sterile

plants are encountered. Marsdenia viridiflora

has copious white latex, but T. linearis and

R. linearis both have clear latex. Tylophora

linearis has a corolla that is valvate and purple,

and staminal coronal lobes that are rounded to

somewhat flat-topped and do not extend above

the style-head, whereas Rhyncharrhena linearis

has a corolla that is imbricate and purple, and

staminal coronal lobes that are saccate and

elongated, with tips extending above the style-

head (see illustration in Forster 1996) and

Marsdenia viridiflora has a campanulate

corolla that is green-yellow in colour (see

illustration in Forster 1995). Rhyncharhena

linearis usually occurs in drier vegetation

communities than Tylophora linearis and has

a more inland distribution, whereas Marsdenia

viridiflora is at its southern limit in southern

Queensland (apart from a couple of populations

in northern New South Wales) and tends to

grow in vegetation communities on heavier clay

soils.

Ecology : We have little or no information on

biotic interactions with Tylophora linearis. The

pollinator(s) for Tylophora linearis, or any

other species of Tylophora are unknown

(Ollerton & Liede 1997). As with most

asclepiads, insect-mediated transfer of

pollinaria from flower to flower is necessary

for pollination.

Larvae of the Lesser Wanderer butterfly

(.Danaus chrysippus petilia (Stoll, 1790)) have

been observed at Eura S.F., Goobang N.P. and

Cumbil S.F. feeding on foliage of Tylophora

linearis (G. Robertson and D. Binns, pers. obs.,

945

April 2003 at Eura S.F.) and this represents a

new host-plant record for this species. Other

native host plant records (all Apocynaceae:

Asclepiadoideae) for the Lesser Wanderer

include Brachy stelma glabriflorum

F.Muell., Cynanchum carnosum (R.Br.)

Schltr., C. floribundum R.Br., Marsdenia

australis (R.Br.) Druce, Oxystelma esculenta

(L.f.) R.Br. ex Schult., Rhyncharrhena linearis,

Vincetoxicum christineae (RI.Forst.) S.Liede

and V. liebianum (F.Muell.) S.Liede (Braby

2000). Naturalised asclepiad hosts include

Asclepias curassavica L., Calotropis gigantea

(L.) W.T.Aiton, C. procera (Aiton) W.T.Aiton,

Gomphocarpus cancellatus (Burm.f.) Bruyns

and G. fruticosus (L.) W.T.Aiton (Braby 2000).

All of the native host plants are herbs or twiners,

usually with reduced foliage (lamina linear to

lanceolate). Several of the native host plants have

annual stems ( Brachystelma glabriflorum,

Oxystelma esculenta, Vincetoxicum christineae,

V. liebianum ) and as noted previously for the

first listed species (Forster 1991), the larvae

are only able to feed on the plants in the

relatively short period of time when the shoots

are actively being produced. One plant in

Goobang N.R supported two larvae of different

instars and had been substantially depleted by

feeding. Each larva is likely to require more

than one shoot to mature and the effects of larval

predation on shoot, flower and fruit production

may be locally substantial.

Many of the plants in Goobang N.R and

Eura S.F. were twining on regrowth stems of

M. uncinata following commercial harvest for

broombush, which completely removes stems

at about 20 cm above ground level. The size of

the regrowth stems indicated the harvest event

occurred about 10 years previously. It is

probable that the Tylophora survived the

harvest, but it may also have colonised the area

since harvest. Part of the population in Goobang

N. P. was burnt by a moderate intensity fire about

12 months prior to observation. The fire killed

the aerial parts of the Melaleuca uncinata and

burnt the foliage and small twigs, but most

shrubs were regrowing from basal shoots. The

Tylophora linearis plants were clearly

resprouting from stems which existed prior to

the fire, indicating that the plant has an ability

to survive at least moderate intensity fire.

946

Conservation status : Tylophora linearis is

currently listed as Endangered under Australian

Federal ( EPBC ), Queensland ( NCA ) and New

South Wales (TSCA) legislation. It is an

inconspicuous plant that is likely to be

overlooked by less than ardent botanical

collectors. The relative ease with which several

additional localities were found during a short

period of specific searching following the initial

rediscovery suggests that the apparent rarity

may be an artefact of undercollection, at least

in the Dubbo-Pilliga area. However, the total

known population size and area of occupancy

remain small. The broad historical distribution

for the species (range of c. 900 km), together

with broad scale land clearing during the 20 th

century, may indicate that the paucity of

collections elsewhere is real as well as being

an artefact of undercollection. Searches after

rain are required in other parts of its historical

range.

At Eura State Forest, at least 270 separate

shoots, in over 150 clumps, were seen over 0.5

ha, twining around plants of Melaleuca

uncinata and occasionally other shrubs. At

Pilliga West S.F., about 20 shoots were seen over

0.2 ha, mostly twining around Allocasuarina

luehmannii. At Cumbil S.F., over 100 shoots

were seen over 0.2 ha, but at least 80% were

small shoots not of flowering size.

Whilst most plants of Tylophora linearis

have been found closely associated with

Melaleuca uncinata, this has not meant that

the Tylophora is correspondingly common

where large areas of the Melaleuca are present.

Nevertheless, the occurrence of this plant,

apparently preferentially, in closed shrubland

of Melaleuca uncinata, may lend clues to

extending the known occurrence of Tylophora

linearis. Melaleuca uncinata is widespread in

inland Australia from Lake Lucy in Queensland

(18° 33’S) to near Bendigo in Victoria (36°

30’S) and westwards to south-west Western

Australia. Although the largest and most

floriferous populations of Tylophora linearis are

associated with Melaleuca uncinata

shrublands, its distribution is not limited to an

association with this species.

Despite recent records, the conservation

status of Tylophora linearis using the criteria

defined by the IUCN (Anon. 2001) remains

uncertain, particularly in relation to criteria

based on estimated population changes.

However, the IUCN criteria are based on a time

Austrobaileya 6 (4): 941-947 (2004)

scale of, at most, several decades. The absence

of records, other than those reported here, over

the last several decades, renders any assessment

of recent population changes rather speculative.

Known populations occur in habitats which are

not subject to any evident significant threat and

there is no reason to expect recent or future

decline. However, it is unknown whether

populations may be subject to extreme

fluctuations through unidentified causes. Using

a precautionary assessment which assumes such

fluctuations are possible, and using known

populations only, Tylophora linearis meets

Endangered criterion B2ac. Alternatively,

assuming that extreme fluctuations are unlikely,

it meets only the Vulnerable criterion D (fewer

than 1000 mature individuals, in five or fewer

populations). This is suggested as the most

realistic assessment with current information.

Acknowledgements

Thanks to W. Smith (BRI) for the illustrations

and map.

References

Anonymous (2001). IUCN Red List Categories and criteria:

Version 3.1. IUCN Species Survival Commission.

IUCN: Gland, Switzerland, Cambridge, UK. ii + 30

pp.

Braby, M.F. (2000). Butterflies of Australia. Melbourne:

CSIRO Publishing.

Forster, PI. (1991). Host records (Family Asclepiadaceae)

and distribution of Danaus chrysippus petilia (Stoll)

(Lepidoptera: Nymphalidae) in Australia.

Australian Entomological Magazine 18: 97-99.

-(1992). A taxonomic revision of Tylophora R.Br.

(Asclepiadaceae: Marsdenieae) in Australia.

Australian Systematic Botany 5: 29-51.

-(1994). A taxonomic revision of Tylophora R.Br.

(Asclepiadaceae: Marsdenieae in Papuasia.

Australian Systematic Botany 7: 485-505.

-(1995). Circumscription of Marsdenia (Asclepiadaceae:

Marsdenieae) with a revision of the genus in

Australia and Papuasia. Australian Systematic

Botany 8: 703-933.

-(1996). Asclepiadaceae. Flora of Australia 28: 197—

283. Melbourne: CSIRO Publishing.

Nichols, K.M., Browne, J.H. & Parsons, R.F. (1991).

Ecology of two asclepiadlianes in semi-arid Victoria.

Proceedings of the Royal Society of Victoria 103:

93-112.

Ollerton, J. & E tf. de, S. (1997). Pollination systems in the

Asclepiadaceae: a survey and preliminary analysis.

Biological Journal oftheLinnean Society 62: 593-

610.

Forster, Binns & Robinson, rediscovery of Tylophora linearis

947

Map 1. Distribution of Tylophora linearis in Australia. Records pre 1970 • ,

records post 1970 ▼

New Australian species in the lichen genus Siphula Fr.

Gintaras Kantvilas

Summary

Kantvilas, G. (2004). New Australian species in the lichen genus Siphula Fr. Austrobaileya 6 (4) 949-955.

Two species of Siphula are described and illustrated: S. australiensis Kantvilas sp. nov. from the Central

and Southern Tablelands of New South Wales, and S. parhamii Kantvilas sp. nov. from the humid wet

tropics of Queensland. Their relationships with other taxa in the genus are discussed. An unusual, peltate

form of Siphula coriacea Nyl. from Central Queensland is briefly noted and illustrated.

Keywords: lichens, lichenized fungi, Siphula, Australia

Gintaras Kantvilas, Tasmanian Herbarium, Private Bag 4, Hobart, Tasmania 7001, Australia.

Introduction

The lichen genus Siphula Fr. is characterised

by a foliose to fruticose thallus with well-

developed root-like rhizines and a green,

unicellular photobiont. Ascomata are

completely unknown for any species of the

genus and consequently, without the benefit of

characters of apothecial ontogeny, anatomy and

morphology, ascus structure and ascospore

morphology, the taxonomy of Siphula is

notoriously difficult. The situation is

complicated further by the broad range of

morphological variation displayed by most

species, often in response to subtle variations

in ecological factors. The current species-level

classification is based extensively on chemical

composition, and correlations between this and

morphology, ecology and geographical

distribution; this approach has been applied to

the delimitation of species from Tasmania

(Kantvilas 1996, 1998) and South America

(Kantvilas & Elix 2002).

The taxonomic position of the genus is

still unclear although several molecular studies

have indicated that the closest relatives of at

least some Siphula species are to be found in

the family Icmadophilaceae (Stenroos & De

Priest 1998, Platt & Spatafora 1999), an

hypothesis based on molecular data but not

inconsistent with morphological, chemical and

ecological information (Kantvilas 2002).

A recent word-wide review of the genus

(Kantvilas 2002) recognised 23 described

species, with main centres of diversity in

temperate Australasia, tropical America,

southern South America, and southern Africa

and the Macarene Islands. Several undescribed

species are also known and two of these, both

from mainland Australia, are described here.

An unusual form of the Australasian endemic,

S. coriacea Nyl., is also discussed briefly.

Materials and Methods

This study is based on herbarium specimens

held in CANB and HO, although comparative

material and types from most major herbaria

have also been examined over the last decade.

Thallus anatomy was studied by high-power

microscopy of hand-cut sections mounted in

water, 10% KOH, lactophenol Trypan Blue and

ammoniacal Congo Red. Chemical composition

was determined routinely via thin-layer

chromatography using standard methods

(Orange et al. 2001), with selected, critical,

confirmatory analyses undertaken by Prof. J.A.

Elix, Canberra, using high performance liquid

chromatography (Feige et al. 1993).

Taxonomy

1. Siphula australiensis Kantvilas, sp. nov.

Aliquot similis Siphulae decumbenti Nyl.

et item acidum thamnolicum continens

sed habitu folioso, lobis late rotundatis

concavisque et rhizinis sparsis differens.

Typus: Australia: New South Wales.

Pigeon House Mountain, 35°21’S

150°16’E, c. 650 m altitude, on vertical

rock face in a west-facing, rather moist

cleft, 21 October 1999, G Kantvilas 344/

Accepted for publication 11 March 2004

950

Austrobaileya 6 (4): 949-955 (2004)

Fig. 1. Siphula australiensis Kantvilas (holotype). Scale = 10 mm.

99 (holo: HO; iso: GZU, NSW).

Thallus foliose, loosely attached to soil or soft

sandstone. Lobes markedly flattened,

decumbent, generally discrete, spreading and

broadly rounded at the thallus margins, much

divided, contiguous to imbricate and at times

± erect and subfruticose in the thallus centre,

undulate to concave, somet im es ± cochleate,

1-3 (-6) mm wide, with dorsal and ventral

surfaces ± identical; surface chalky white,

sometimes developing a faint beige tinge in the

herbarium, generally smooth in younger parts

of the thallus but becoming verrucose in older

parts, very intensely and coarsely scabrid-mealy

throughout; margins ± entire or minutely and

irregularly crenulate, undulate or ascending,

not thickened, very brittle and frequently

fractured. Thallus in section 120-230 pm thick;

well-defined cortex absent, but outermost layers

of the thallus with a 10-50 pm thick layer of

tiny crystals not dissolving in KOH, visible at

high-power magnification in polarised light;

photobiont cells spherical, 7-13 pm diam.,

irregularly clumped, especially beneath the

dorsal surface of the the thallus; medullary

hyphae rather loosely and irregularly

interwoven, 5-8 pm thick, with walls to 3 pm

thick. Rhizines mostly uncommon and typically

widely scattered, pale brownish, 0.25-0.5 mm

thick at point of attachment. Fig.l.

Chemistry: thamnolic and decarboxythamnolic

acids; K+ intense yellow, sometimes slowly

becoming brownish red, KC-, C-, P+ orange,

UV-.

Additional specimens: New South Wales. Katoomba, Mar.

1965, G.C. Bratt & J.A. Cashin 2040 (HO); Evans Lookout,

c. 4 km E of Blackheath, Apr. 2002, G Kantvilas 178/02

(HO); Pigeon House Mountain, on sandstone rock, Sept.

1977, D. Verdon 3126 (CANB); same locality and date, J.A.

Elix 3930 (CANB); same locality, Nov. 2000, G. Kantvilas

488/00 (HO).

Remarks: Siphula australiensis is most closely

related to the widespread S. decumbens Nyl.,

and both species share a chalky white thallus

and thamnolic acid as the major chemical

constituent. The main difference between the

two taxa lies in their growth habit: foliose with

spreading, generally rounded, cochleate,

marginal lobes in the former, but distinctly

fruticose with crowded, elongate lobes in the

latter. Although S. decumbens is extremely

variable (see Kantvilas 1998), none of the

Kantvilas, new Australian species in Siphula Fr.

951

Fig. 2. Siphulaparhamii Kantvilas (isotype). Scale = 5 mm.

material studied, encompassing a very wide

range of localities and habitats, even remotely

approaches the distinctive growth habit of

S. australiensis . The differences in morphology

are manifest in subtle anatomical differences.

Thus in S. decumbens , the relatively thick

medullary hyphae are periclinal whereas in

S. australiensis they are irregular or anticlinal.

Similarly the rhizines of the two species differ,

with those of S. decumbens occurring as a thick,

basal tuft, whereas those of S. australiensis are

scattered on the ventral surface.

Perhaps the lichen that is superficially

most similar to S. australiensis is Icmadophila

splachnirima (Hook.f. & Taylor) D.J. Galloway,

a widespread species that grows on wet peaty

soil in cool to cold temperate areas of

Australasia. Like S. australiensis, this species

also has a foliose habit, rounded, often cochleate

lobes and contains thamnolic acid, but the lobes

are generally thinner and not markedly scabrid

and mealy. The presence of pink apothecia,

characteristic of the Icmadophilaceae in general,

and the absence of rhizines make 7. splachnirima

easily identifiable, but the resemblance between

the two species is striking and perhaps offers

some insight into their relationships at a higher

taxonomic rank.

Distribution and habitat : Siphula australiensis

is locally common at the type locality and at

other scattered locations on the sandstone

escarpment of the New South Wales Tablelands.

It grows in moist, sheltered clefts and on ledges,

usually on large cliffs and bluffs. Vertical rock

faces are especially favoured. That the new

species is restricted to such habitats probably

indicates a relict distribution in fire-protected

refugia, rather than necessarily a predisposition

to grow on moist, vertical rocks. In this habitat,

it is typically intermixed with an algal film,

small ferns and bryophytes, but other lichens

with which it was associated at the type locality

include Cystocoleus ebeneus (Dillwyn)

Thwaites, Leioderma duplicatum (Mull. Arg.)

D. J. Galloway & P.M. Jprg. and depauperate

tufts of Cladia aggregata (Sw.) Nyl. and

Cladonia pertricosa Kremp.

2. Siphula parhamii Kantvilas, sp. nov. Arete

affinis speciei austroafricanae endemicae

Siphulae torulosae (Thunb. ex Ach.) Nyl.

a qua lobis decumbentibus robustioribusque,

caespes laxiores formantibus imprimis differt.

Typus: Australia: Queensland. Mt

Finnigan, Mt Finnigan Range, Cedar Bay

National Park, 15°49’S 145°16’E, 1090

952

Austrobaileya 6 (4): 949-955 (2004)

Fig. 3. Variation in baeomycesic acid and squamatic acid-containing species of Siphula. A: S. parhamii Kantvilas (part of

holotype); B: S. torulosa (Thunb. ex Ach.) Nyl. ( Brusse 2527, HO); C: S.fastigiata (Nyl.) Nyl. ( Moscal 11956, HO). Scale

= 10 mm. (A-B drawn by Lauren Black; C drawn by Georgina Davis)

m altitude, on exposed rocky ground in

exposed heathy-grassy area with large

rock outcrops, 20 October 1995, H.

Streimann 57192 (holo: CANB; iso: HO).

Thallus fruticose, forming scattered or

contiguous clumps mostly 10-30 mm wide over

stones and gravelly soil. Lobes generally

markedly flattened, decumbent, + discrete or,

more commonly, loosely overlapping, 30-50 (-

100) mm long, 0.4-1 (-1.5) mm wide, with

dorsal and ventral surfaces ± identical; surface

dull whitish grey, matt, generally unevenly

puckered and dimpled, neither mealy nor

scabrid, patchily discoloured by a sparse to

moderately dense ‘veil’ of black, unidentified

fungal hyphae composed of chains of rhomboid

cells 5-6 pm wide; margins entire or irregularly

crenulate, unevenly thickened in places,

sometimes with knob-like projections; apices

erect or ascending, sometimes with short,

subterete extensions, or rather thickened and

knob-like. Thallus in section 250-600 pm

thick, lacking a well-defined cortex but with a

brownish grey outer layer c. 10 pm thick;

photobiont cells irregularly roundish, 8-15 pm

wide, mostly single but sometimes in pairs or

tetrads to 20 pm wide, typically concentrated

in a continuous or interrupted layer beneath

the dorsal surface, sometimes also occurring

in bundles through the entire thickness of the

thallus or at the ventral surface; medullary

hyphae periclinal, densely entwined and ±

conglutinated, 1-2 pm thick. Rhizines pale

grey-brown, c. 0.25 mm at point of attachment.

Figs 2, 3A.

Chemistry: baeomycesic and squamatic acids;

K+ pale yellowish, C-, KC-, P+ yellow-orange,

UY+ pale yellow-orange with whitish patches.

These two compounds appear to be irregularly

distributed in the thallus, with squamatic acid

mainly in the internal, medullary area and

baeomycesic acid near the thallus surface.

Hence, in UV light, the thallus surface appears

pale orange-yellow, but abraded or fractured

areas where squamatic acid containing tissues

are exposed react UV+ whitish.

Additional specimen: Queensland. MtFinnigan, on exposed

boulder, Oct 1995, H. Streimann 57206 (CANB).

Remarks: Siphula parhamii is a distinctive

species in the Australian flora, characterised

by the relatively robust, decumbent lobes that

form loose tufts, and by the presence of

baeomycesic and squamatic acids. Its closest

relative is S. torulosa (Thunb. ex Ach.) Nyl.,

Kantvilas, new Australian species in Siphula Fr.

953

Fig. 4. Variation in Siphula coriacea Nyl. A: ‘typical’ erect form (Kantvilas & Jarman 375/92, HO); B: compact, decumbent

form on rock (Bratt 67/515, HO); C: compact, flattened form on arid soil (Bratt 67/121, HO). Scales: A= 10 mm; B,C = 5mm.

endemic to the Cape region of South Africa

(Fig. 3B). Siphula torulosa is chemically

identical but has more slender, erect or

ascending lobes, 4-12 mm tall and 0.3-0.8 mm

wide, that form relatively dense tufts or swards

(see also Mathey 1974). Although many species

of Siphula are very widespread geographically

and very variable morphologically (see

Kantvilas 2002), S. torulosa is a very well-

defined, localised taxon. It seems untenable to

broaden its concept to include S. parhamii,

which is localised in north Queensland and is

morphologically rather extreme.

Also rather similar and chemically

identical is S.fastigiata (Nyl.) Nyl. This species

has generally erect, strap-like, chalky white

lobes with a usually scabrid, mealy, sometimes

intensely puckered and verruculose surface

(Kantvilas (1998) (Fig. 3C). It ranges from

Tasmania to New Zealand, southern South

America and northward into tropical America.

It displays extreme morphological variation but

does not encompass the distinct morphologies

of S. parhamii and S. torulosa (Kantvilas 2002).

Distribution and habitat : Siphula parhamii is

known only from the type locality, a peak in

tropical north Queensland. On the basis of

specimen label data, it grows in open, grassy-

954

Austrobaileya 6 (4): 949-955 (2004)

Fig. 5. An unsual peltate form of S. coriacea Nyl. (Purdie 4214, HO). Scale = 5 mm.

heathy vegetation, directly on rocks and on

gravelly soil, inter mi xed with moss. The late

Heinar Streimann, who collected the material,

described this species as being locally abundant.

It should be sought for on other peaks in the

region.

Etymology: The new species is named in

honour of Mr John Willoughby Parham, friend

and mentor of the author, in recognition of his

contributions to the Botany of Fiji, Queensland

and Tasmania (see Kantvilas 2003).

An unusual form of Siphula coriacea Nyl.

Siphula coriacea is a very distinctive species,

characterised by typically erect or ascending,

flattened, distinctly bluish grey lobes containing

barbatic acid. This chemical composition is

unique in the genus. It is known from the

mainland States of Australia and New Zealand

and typically occurs on soil in heathland and

grassland. As with most other species of the

genus, this lichen is extremely variable, with

elongate, loosely entangled, strap-shaped lobes

when occurring in moister, sheltered habitats

(Fig. 4A), but with very compact, short,

convoluted lobes in drier, exposed locations

such as on soil in the rangelands of the

Australian interior (Figs 4B-C).

Two specimens, both from the same

locality in Central Queensland, represent a very

unusual form of this pecies. They have a thallus

composed of discrete, flat, peltate lobes, 2.5-

8.5 mm wide, 0.5-0.85 mm thick, with a plane

to undulate to rather bullate and puckered,

bluish grey upper surface, recurved, ± thickened

margins, and a pale brownish underside with

scattered, very prominent, white, dichotomously

branched rhizines (Fig. 5). Although

morphologically disjunct from S. coriacea as

generally understood, these unusual specimens

contain barbatic acid and are anatomically

identical with S. coriacea : they have an

epinecral layer to c. 15 pm thick, a

‘pseudocortex’ to c. 60 pm thick, composed of

anticlinal hyphae c. 5 pm thick that extend from

the medulla, a photobiont layer composed of a

unicellular, green alga, 6-10 pm diam, and a

medulla of irregularly interwoven hyphae,

inspersed with crystals that dissolve in KOH.

According to notes (by R.W. Rogers)

accompanying one specimen, the collection is

from microhabitats that are possibly

waterlogged after rain but otherwise dry. This

unusual form may be an environmental

modification. Certainly given the degree to

which Siphula species can vary in response to

habitat, any taxonomic recognition of this odd

form would need to be preceded by careful study

of the populations in the field. ‘Typical’ S. coriacea

also occurs at the same general locality.

Specimens : Queensland. Idalia National Park, c. 12 km S

of Idalia homestead, on walk to Mountain Rock Hole, 24°57 ’ S

144°47’E, 22 Sept 1992, R.W. Purdie 4214 (CANB, HO);

Kantvilas, new Australian species in Siphula Fr.

Idalia National Park, old Idalia homestead, 24°55’S

144°43’E, Feb 1996, R.W. Rogers 10478 (BRI).

Acknowledgements

I thank Jean Jarman for preparation of the

figures, and Jack Elix and Pat McCarthy for

their companionship and assistance during field

work in New South Wales. Jack Elix also

undertook the critical h.p.l.c. analyses. The late

Heinar Streimann and Rod Rogers are

acknowledged for placing at my disposal their

collections.

References

Feige, G.B., Lumbsch, H.T., Huneck, S. & Elix, J.A. (1993).

The identification of lichen substances by a

standardized high-performance liquid

chromatographic method. Journal of

Chromatography 646: 417-427.

Kantvilas, G. (1996). Studies on the lichen genus Siphula in

Tasmania I. S. complanata and its allies. Herzogia

12: 7-22.

955

-(2002). Studies on the lichen genus Siphula Fr.

Bibliotheca Lichenologica 82: 37-53.

-(2003) Obituary, John W. Parham 1929-2002.

Austrobaileya 6: 575-579

Kantvilas, G & Elix, J.A. (2002). The taxonomy, chemistry

and morphology of some South American species

of Siphula. Herzogia 15: 1-12

Mathey, A (1974). Contribution a F etude du genre Siphula

(Lichens) en Afrique. Nova Hedwigia 22:795-878

Orange, A., James, PW. & White, FJ. (2001). Microchemical

Methods for the Identification of Lichens. British

Lichen Society

Platt, J.L. & Spatafora, J.W. (1998). Are-examination of

generic concepts of baeomycetoid lichens based on

phylogenetic analyses of nuclear SSU and LSU

ribosomal DNA. Lichenologist 31: 409-418

Stenroos, S.K. & De Priest, PT. (1998). SSU rDNA

phylogeny of Cladoniiform lichens. American

Journal of Botany 85: 1548-1559

-(1998). Studies on the lichen genus Siphula in Tasmania

II. The S. decumbens group. Herzogia 13: 119—

138

Chromosome records for five trigger plants ( Stylidium ; Stylidiaceae) from

northern Australia

J.A. Wege

Summary

Wege, J.A. (2004). Chromosome records for five trigger plants ( Stylidium ; Stylidiaceae) from northern

Australia. Austrobaileya 6 (4): 957-959. Chromosome counts are reported for five species of Stylidium

from northern Australia. In contrast to previous hypotheses, variation in number is evident: S. lobuliflorum

and S. ensatum possess n = 11, S. schizanthum n = 14, whilst S. semipartitum and S. turbinatum possess n = 15. As

for their south-western counterparts, cytogenetic factors are postulated to be involved in the speciation of

trigger plants of northern Australia.

Keywords: Stylidium, trigger plants, chromosomes, speciation, breeding systems.

J.A. Wege, Western Australian Herbarium, Department of Conservation and Land Management, Locked

Bag 104, Bentley Delivery Centre, Western Australia 6983. Email: julietw@calm.wa.gov.au

Introduction

The majority of cytogenetic research performed

on the trigger plant genus Stylidium has

focussed on taxa endemic to the south-west of

Western Australia. This region is a primary

centre of species diversification for Stylidium,

containing approximately 70% of the currently

described taxa (Wagstaff & Wege 2002).

Chromosome counts have been made for a high

proportion of these taxa with numbers ranging

from n = 5 to n = 16, with polyploidy occurring

on 13, 14 and 15 (James 1979; Coates 1982;

Burbidge & James 1991). Studies on

morphologically allied species have shown that

chromosome number change is often a feature

of species differentiation (James 1979; Banyard

& James 1979; Farrell & James 1979; Coates

1982). The hypothesized base number for the

genus is n = 15 and several independent

dysploid reduction series are thought to have

evolved (James 1979).

Since James’ (1979) landmark study,

taxonomic research has seen a two-fold increase

in the number of species of Stylidium known

from northern Australia. With over 60 taxa

currently described (see Bean 1999, 2000) this

region must now be regarded as a second area

of trigger plant diversity; however, there is only

one published chromosome record. James

(1979; pg 22) reported a count of n = 15 made

by B. & G. Keighery, although the species in

question was not identified and no voucher

details were given. He considered all northern

Accepted for publication 13 November 2003

species to possess the basal number of n = 15.

Chromosome numbers for additional species

from northern Australian were sought in order

to assess whether variation in chromosome

number exists in this region.

Methods

Buds were fixed in 3:1 absolute ethanol:glacial

acetic acid for 24 hours, rinsed in 70% ethanol

and subsequently stained with alcoholic

hydrochloric acid carmine (Snow 1963). At

least three separate counts were obtained from

pollen mother cell meiotic material using the

squash technique. Photographs were taken

using a Zeiss Axiophot microscope and images

captured using 6ASA imagelink film.

Herbarium voucher specimens are lodged at

PERTH.

Results

Variation in chromosome number between

northern Australian species of Stylidium is

demonstrated for the first time in the present

study (Table 1). A haploid chromosome number

of 11 was recorded for both S. ensatum and

S. lobuliflorum (Fig. 1). Stylidium turbinatum

was found to have a haploid chromosome

number of 15.

An unidentified herbarium voucher

specimen from south of Darwin with an

annotation “chromosome count of n = 15” was

recently uncovered at The University of Western

Australia. The collection and count, performed

by Bronwyn Keighery, is likely to correspond

958

>V»y i J-

T>* L ■.» '

V gfc.

Fig. 1. Chromosomal preparations (n=ll) for

A) Stylidium ensatum and B) 5. lobuliflorum

to the n = 15 quoted by James (1979) as

occurring in species from northern Australia.

This specimen has since been identified as

S. semipartitum and has been lodged at PERTH,

with a duplicate sent to BRI.

A second voucher specimen with an

unpublished chromosome count was found in

the collection at PERTH. The count for

S. schizanthum of n = 14 is likely to have been

performed by Sid James (G. Keighery pers.

comm.).

Discussion

Whilst the data presented here do not constitute

a comprehensive survey, it is likely that

chromosome change is associated with

speciation of Stylidium across northern

Australia. This notion raises a number of

interesting questions. Do the tropical species

exhibit similar variation in chromosome

number as their south-west congeners? Is

Austrobaileya 6 (4): 957-959 (2004)

chromosome number differentiation a feature

of morphologically allied species? What is the

nature of the breeding systems in tropical

trigger plants?

The last question is raised in view of the

findings of Burbidge & James (1991) that

dysploidy in Stylidium is typically associated

with post-zygotic seed-aborting systems.

Although the trigger plant flower appears

designed to promote cross-pollination, high

levels of inbreeding may be generated by

geitonogamous self-pollination. The seed¬

aborting systems function to significantly

reduce the amount of seed set after self-

pollination as compared to cross-pollination.

They have been shown to occur in many trigger

plants from both south-western and south¬

eastern Australia (Banyard & James 1979;

Coates & James 1979; James 1979; Burbidge

& James 1991; Willis & Ash 1990). Burbidge

& James (1991) also demonstrated that seed¬

aborting systems operate to varying degrees in

Stylidium depending on the species in question.

They can be absent, weak or highly efficient in

species with the hypothesized base number of

n = 15; however, those species with reduced

chromosome numbers almost always exhibit

efficient systems.

If one were to apply the results of

Burbidge & James (1991) to the present study,

one would expect efficient seed-aborting

systems to be present in those tropical trigger

plants with reduced chromosome numbers;

Table 1. Chromosome numbers (n) of species of Stylidium from Northern Australia

Classification (Milbraed 1908) Species

Voucher

Approximate Locality

Subgenus Andersonia

(R.Br) Mildbr.

S. ensatum A.R. Bean

JAW 470

(PERTH 05596513)

McMinns Lagoon,

E of Darwin

S. lobuliflorum F.Muell

JAW 478

(PERTH 05596521)

SW of Jabiru,

Kakadu National Park

S. schizanthum F.Muell

G.J. Keighery 4738

(PERTH 03163296)

Mitchell Plateau,

Kimberleys

Subgenus Tolypangium

(Endl.) Mildbr.

section Debiles Mildbr.

S. semipartitum F.Muell

G.J. Keighery s.n.

(PERTH 05596548)

Burrell’s Creek

S of Darwin

section Floodia Mildbr.

S turbinatum Fowrie & Kenneally JAW All

(PERTH 05596483)

Arhnem Highway,

Kakadu National Park

Wege, Stylidium in northern Australia

959

however, this phenomenon would be an

evolutionary disadvantage given the majority

of these species are annuals and rely on seed

set for regeneration. Burbidge & James (1991)

found seed-aborting systems to be absent in the

south-west annuals from S. section Despectae

(n = 15). They noted that self-pollination is less

likely to occur in these species because

individuals are few-flowered and populations

typically consist of very large numbers of plants

over a small area. In contrast, geitonogamous

self-pollination is a more likely occurrence in

tropical trigger plants given the branched,

many-flowered inflorescences and multiple

scapes of many species. Further, Carlquist

(1979) suggested that selfing is a favorable

adaptation in tropical trigger plants and

outlined a number of morphological features

that may facilitate it, such as the widespread

occurrence of a column pouch into which pollen

can be shed, and from which the stigma can

later receive. If such features have indeed

evolved to promote self-pollination, then it is

doubtful whether efficient seed-aborting

systems have been associated with chromosome

change in these species.

An alternative scenario is that tropical

trigger plants with reduced chromosome

numbers may possess weak seed-aborting

systems, as is the case in S. calcaratum (n =

11) and S. ecorne (n = 13) from S. subgenus

Centridium (for which there are northern

Australian representatives) (Farrell & James

1979). Prior to this study, these two species were

the only annual trigger plants recorded as

having reduced chromosome numbers. Clearly

further research is warranted on chromosome

number variation in northern Australian trigger

plants in conjunction with studies on the nature

of their breeding systems.

Acknowledgments

Thanks are extended to David Coates for

comments and discussion; Bronwyn Keighery

for granting permission to publish her

chromosome count; Greg Keighery for

assistance with vouchering queries; the

Northern Territory Herbarium (DNA) for

providing access to collection information; and

The Northern Territory Government and

Australian National Parks and Wildlife Services

for permission to collect in the Northern

Territory and Kakadu National Park. This

project formed part of a PhD dissertation

completed at the Department of Botany, The

University of Western Australia and was

supported by an Australian Postgraduate Award.

References

Banyard, B J. & James, S.H. (1979). Biosystematic studies

in the Stylidium crassifolium species complex

(Stylidiaceae). Australian Journal of Botany

27:27-37.

Bean, A.R. (1999). A revision of Stylidium sect. Debilia

Mildbr., S. sect. Floodia Mildbr. and S. sect. Lanata

A.R. Bean (Stylidiaceae). Austrobaileya 5:427-

455.

-(2000). A revision of Stylidium subg. Andersonia (R.Br.

ex G.Don.) Mildbr. (Stylidiaceae). Austrobaileya

5:589-649.

Burbidge, A.H. & James, S.H. (1991). Postzygotic seed

abortion in the genetic system of Stylidium

(Angiospermae: Stylidiaceae). Journal of Heredity

82:219-28.

Carlquist, S.J. (1979). Stylidium in Arnhem land: new

species, modes of speciation on the sandstone

plateau, and comments on floral mimicry. Aliso

9:411-461.

Coates, D.J. (1982). Chromosome variation and species

relationships in the scale-leaved triggerplants

(,Stylidium Section Squamosae ). Australian Journal

of Botany. 30:121-130.

Farrell, RG. & James, S.H. (1979). Stylidium ecorne

(F.Muell. ex Erickson & Willis) comb, et stat. nov.

(Stylidiaceae). Australian Journal of Botany

27:39-45.

James, S.H. (1979). Chromosome numbers and genetic

systems in the triggerplants of Western Australia

(Stylidium ; Stylidiaceae). Australian Journal of

Botany 27:17-25.

Mildbraed, J. (1908). Stylidiaceae. In Engler, A. (ed.) Das

Pflanzenreich. IV: 278 (Wilhelm Engelmann:

Leipzig.)

Snow, R.R. (1963). Alcoholic hydrochloric acid carmine as a

stain for chromosomes in squash preparations. Stain

Technology 38:9-13.

Wagstaff, S.J. & Wege, J. (2002). Patterns of Diversification

in New Zealand Stylidiaceae. American Journal of

Botany 89(5):865-874.

Willis, A.J. & Ash, J.E. (1990). The breeding systems of

Stylidium graminifolium and S. productum

(Stylidiaceae). Australian Journal of Botany

38:217-227.

Gonocarpus hirtus Orchard (Haloragaceae), new from southeastern

Queensland and northeastern New South Wales

A.E.Orchard

Summary

Orchard, A.E. (2004). Gonocarpus hirtus Orchard (Haloragaceae), new from southeastern Queensland

and northeastern New South Wales. Austrobaileya 6 (4): 961-965. Anew species from the ranges southwest

of Brisbane, and the northeastern highlands of New South Wales, is described and illustrated. G. hirtus,

found in dry sclerophyll forest, belongs to a complex of weak scrambling subshrubs found throughout the

Great Dividing Range, from Victoria to Queensland, and is most closely allied to G. longifolius and G.

effusus.

A.E.Orchard, c/o Centre for Plant Biodiversity Research, CSIRO Division of Plant Industry, GPO Box

1600, Canberra ACT 2601, Australia.

Introduction

The genus Gonocarpus comprises 42 species,

widespread in Australia and New Zealand, with

a few species also extending to New Guinea,

Indonesia, the Philippines, Japan and SE Asia.

There are 37 species in Australia, the majority

in dry to damp sclerophyll forest, but with some

species found in almost all habitats. The genus

was revised by Orchard (1975), with additional

species described in Orchard (1977, 1986). A

summary of the then-known Australian species

was published by Orchard (1990).

With their often sparse, scrambling habit

and inconspicuous flowers, Gonocarpus species

are frequently overlooked by all but the most

thorough collectors, and are probably under¬

represented in most collections. Despite this,

sporadic collecting has led to a steady increase

in numbers of new taxa described since the last

revision, particularly from the ranges of the

eastern seaboard. In the dry to damp sclerophyll

forests of the eastern ranges of Victoria, New

South Wales and Queensland the understorey

often contains one or more species of

Gonocarpus. In the driest areas, particularly

on rocky outcrops, the most common species is

the alternate-leaved G. elatus. In slightly

damper areas, subject to enhanced runoff or

higher rainfall G. elatus is replaced by the most

common Gonocarpus of all, G tetragynus. In

even damper situations the scrambling G humilis

or the more erect shrubby G. teucriodes replaces

G tetragynus, to be replaced in turn by members

of the G. longifolius complex. G. tetragynus,

Accepted for publication 2 February 2004

G humilis, G. teucrioides, and members of the

G. longifolius complex (G. longifolius,

oreophilus, G. effusus) all have opposite leaves,

which usually become alternate above as they

merge into the bracts of the inflorescence.

Gonocarpus elatus differs from all others

below in having uniformly alternate leaves. Its

fruit is irregularly rugose with about 4 rows of

rounded tubercles, while that of the opposite¬

leaved species is uniformly 8-ribbed with 3-4

oblique calluses between the pairs of ribs, often

sloping in alternate directions in each vertical

row, forming a chevron pattern.

Gonocarpus tetragynus is a small

subshrub, rarely exceeding 30 cm tall, and is

easily recognisable by its lanceolate leaves with

a distinctive indumentum of evenly distributed,

closely appressed stiff hyaline hairs. The

remaining species are usually much larger

plants.

G. longifolius is a subshrub 35 to 100 cm

tall, found mainly in shrub communities on

sandstone soils from sealevel to about 600 m,

mainly in New South Wales, from the Blue

Mountains to New England, and extending just

over the border into SE Queensland. See Bell

(2001) for a discussion of recent range

extensions. It is characterised by its long, soft,

spreading hairs on stems and sometimes on the

leaves, and by its linear-oblong leaves, 1.3-3.2

mm long, with 20-30 small cuspidate teeth.

G. humilis is usually a much weaker,

scrambling plant, with relatively long soft hairs

on the stems and leaves, more or less ovate

962

Austrobaileya 6 (4): 961-965 (2004)

leaves, and differs from other species of the

complex in having very short (to 1.5 mm) petals

and anthers, the latter reduced to 4 instead of

the usual 8. The missing whorl of stamens may

be replaced by small staminodes. G. humilis

seems to prefer damp or swampy areas within

sclerophyll forest, from sealevel to about 1200 m.

G.. teucrioides is a more robust species

than G. humilis, with more hispid hairs, usually

seated on swollen multicellular bases. It has

the usual 8 stamens, and can be distinguished

from all other members of the complex by its

large, green, fleshy, lanceolate to narrowly ovate

bracteoles which often more or less conceal the

ovary of the flower. The bracteoles of other

members of the complex are usually brown, narrow,

and much smaller than the ovary. G teucrioides is

usually found in the understorey of dry

sclerophyll forest, from sealevel to about 1200

m, but has also been recorded on the margins

of rainforest and in subalpine communities.

G. oreophilus has a similar distribution

to G. longifolius, but is found in rainforest or

wet sclerophyll forest at 300-1500 m. It is

distinguished from G longifolius by its larger

(1.0-3.5 cm) ovate to oblong leaves with 15-

30 rounded teeth, and by an indumentum on

the stems and leaves which is fine, very short,

and almost velvety.

G. effusus is a further member of the

complex, confined to the Glasshouse Mountains

of SE Queensland at around 180 m, where it

occurs in open, rocky situations. It has very

small (4-5 mm long) ovate leaves with

thickened margins and only 4-6 minute teeth.

At the time of preparation of my revision

of the genus (Orchard, 1975) an anomalous

collection from Mt Maroon, SW ofBeaudesert,

Queensland, was known, but its status was

unclear. It was tentatively identified as a possible

hybrid between G longifolius and G teucrioides,

or a possible new species. Additional material

has now been collected from nearby Mt Moon,

Mt Gillies and Mt Greville, and from near

Moonbi Gap in New South Wales, and it now

seems clear that these collections represent

another species of restricted distribution within

the G. longifolius complex. The new species,

described below, has the long silky stem

indumentum of G. longifolius, but differs in its

narrowly ovate leaves with only about 10, much

coarser, teeth. The hairs on the upper surface

of the leaves of G. hirtus are silky, spreading

and dense. In G. longifolius in the southern part

of its range the leaf hairs are relatively sparse

and short, and usually more or less appressed.

In the northern part of its range they are longer

and more erect, but never quite as exuberant as

in G hirtus. The dentition of the leaves is also

clinal, with southern G longifolius having up

to 30 small teeth per leaf, but the numbers of

teeth decrease and their size increases as one

moves north. However, despite these

intergradations, G. hirtus is distinguishable by

its narrowly ovate leaf shape, teeth rarely

exceeding 10 per leaf, teeth about 1 mm long

and coarse, and dense softly hirsute upper leaf

indumentum. G. longifolius has narrowly

oblong leaves, with (12-) 20-30 small teeth to

0.5 mm long, and subglabrous to appressed

hairs on the upper leaf surface. The only other

species which might be confused with G hirtus

is G. humilis, which has very short petals,

and stamens reduced to 4 (usually plus

4 staminodes).

Taxonomy

Gonocarpus hirtus Orchard, sp. nov. Species

G longifolium simulans, sed foliis ovatis,

(8-) 10-14 mm longis, 5-8 mm latis,

dentibus usque 10 grossis et 1 mm longis,

et pilis foliorum patentibus hirsutis usque

1 mm longis, differt. Typus (here

designated): Queensland. Moreton

District. I. R. Telford 11987 &

J.Nightingale, 15 Nov. 1993, Mount

Greville, southeast ridge. Holotypus:

CBG9313900. Isotypi (n.v.): BRI, NSW,

MEL.

Weak scrambling subshrub 25-40 cm or more

tall. Stems mainly annual from a persistent

woody rootstock, c. 1.0-1.5 mm diam., round

in section (not ribbed), reddish brown, covered

with dense soft spreading (slightly antrorsely

curved) hairs. Hairs c. 1 mm long, hyaline, of

about 5 cells, uniseriate, seated on a very

slightly swollen basal cell. Leaves opposite,

becoming alternate near inflorescence, on

petioles 1.5-3 mm long. Lamina dark green,

narrowly ovate, (8-) 10-14 mm long, 5-8 mm

wide, gradually becoming smaller towards tips

Orchard, new species Gonocarpus hirtus

of branches, soft, coarsely serrate with about 5

cuspidate teeth c. 1 mm long on each margin;

margin with distinct hyaline thickening;

moderately semiappressed hirsute on both

surfaces with hairs as for stem. Inflorescence

of flowers borne singly in axils of upper leaves

(=bracts). Bracteoles reddish brown, linear-

lanceolate, 1 mm long, 0.25-0.3 mm wide,

tapering gradually to acute tip, densely hirsute

with hairs as for stems. Flowers 4-merous, on

pedicels 0.7 mm long. Sepals purplish green,

narrowly oblong, 0.7 mm long, 0.3 mm wide,

blunt at tip, with prominent white callus at base;

margins white-hyaline, slightly thickened.

Petals reddish purple, paler at margins, 7 mm

long, 1.5 mm keel to margin, strongly hooded,

non-unguiculate (sessile), with moderately

dense hairs (as for stem) on dorsal surface.

Stamens 8; filaments 0.25 mm long; anthers

reddish, linear, 2 mm long, non-apiculate.

Styles 4, clavate, reddish-yellow fimbriate.

Ovary silvery grey, ovoid, 1.0-1.3 mm long,

0.9 mm diam., 8-ribbed with c. 3 oblique

calluses between each pair of ribs, with

occasional hyaline scabrous hairs on calluses

and/or ribs. Fruit as for ovary, silvery grey to

pale reddish purple, 1.2-1.3 mm long, 1.0 mm

diam.; sepals green, persistent. Fig. 1.

Specimens examined : Queensland: Moreton District.

A.R.Bean 6647,2.X.1993, Mt Gillies, south of Rathdowney

(BRI, CANB); P.I.Forster, A.R.Bean & L.H.Bird PIF6633,

14.iv.1990, Mount Moon, 5 km SW of Mount Alford township

(BRI, CANB); I.R.Telford 3525, 4.x. 1973, Mt Maroon, 36

km SW of Beaudesert (CANB). New South Wales: Northern

Tablelands: L.M.Copeland 3277,22.xi.2001, Stony Batter

Creek Nature Reserve, c. 60 km WNW of Armidale (CANB,

NE, NSW). Northwest Slopes: J.R.Hosking 1382 & 1385,

1. Stamens 4, opposite sepals; staminodes 4

Stamens 8, all functional.

963

10.xii.1996, ~1.5 km Wof Moonbi Gap (CANB, MEL, NE,

NSW, TARCH).

Distribution, habitat and ecology : Present

collections suggest that in Queensland G. hirtus

is confined to the Moreton District, in a small

area of mountains ca 50 km southwest of

Brisbane, where it is recorded from "eucalypt

forest with E. dura, E. tindaliae, E.

acmenoides, shallow soil, alt. 600m."

(A.R.Bean 6647), "rhyolitic rock outcrops, with

Eucalyptus dura and Eucalyptus acmenoides ;

alt. 600 m" (P.I.Forster et al., PIF6633), "dry

sclerophyll forest on rocky slopes, alt. c. 700

m altitude" (I.R.Telford 3525), and "rocky

ridge, SE aspect, skeletal soil on trachyte, shrub

community, Eucalyptus notabilis (mallee

habit), Westringia sericea, Leptospermum spp.,

alt. 350 m." (I.R.Telford 11987 &

J.Nightingale). In New South Wales it is known

from the North Western Slopes and Northern

Tablelands regions in "shrubland on brown silt

and sand over granite at 790 m." (J.R.Hosking

1382 & 1385) and "shallow sandy loam

amongst boulders; layered woodland dominated

by Eucalyptus andrewsii and E. prava; 920 m."

(L.M.Copeland 3277). It flowers October-

November, and fruit is present until April.

Key to related species

The following key is based on leads 10 to 15 in

Orchard (1990), with the addition of G. teucriodes

and G hirtus. G. chinensis is included because it

is a species of northern Queensland and the

Northern Territory, but is not part of the

complex under discussion.

. G. humilis

2. Subshrub usually 50-100 cm tall; main stems distinctly woody. 3

Perennial herb usually less than 35 cm tall; main stems herbaceous or

only slightly woody. 7

3. Bracts of inflorescence opposite, at least at base; bracteoles green, fleshy,

concealing ovary; indumentum of young ste m s and leaves of stiff scabrous

hairs 0.2-0.5 mm long, seated on swollen multicellular bases. G. teucrioides

Bracts of inflorescence all alternate; bracteoles red-brown, membranous,

much smaller than ovary; indumentum of young stems and leaves long

(to 1 mm) and soft, or if shorter and stiff, not seated on swollen

multicellular bases. 4

964

Austrobaileya 6 (4): 961-965 (2004)

4. Leaves and young stems with dense indumentum of soft, spreading 2-5-

celled hyaline hairs 0.3-1 mm long. 5

Leaves and young stems with sparse to dense indumentum of stiff, spreading

1-2-celled hyaline hairs 0.1-0.3 mm long. 6

5. Leaves linear-oblong, (1.3-) 1.5-2.5 (-3.2) cm long, 0.3-0.6 cm wide,

with (12-) 20-30 teeth to 0.5 mm long. G. longifolius

Leaves narrowly ovate, 0.8-1.4 cm long, 0.5-0.8 cm wide, with ca 10

teeth to 1 mm long. G. hirtus

6. Indumentum dense, velvety; leaves ovate to oblong, 1.0-3.5 cm long.G. oreophilus

Indumentum sparse, scabrous; leaves ovate, 0.4-0.5 cm long.G. effusus

7. Indumentum of young stems and leaves soft, spreading; leaves narrowly ovate . . G. hirtus

Indumentum of young stems and leaves stiff, scabrous, appressed; leaves

lanceolate to linear-lanceolate . 8

8. Leaves lanceolate, widest near centre; leaves, bracts, bracteoles and sepals

lacking prominent white thickened margins; indumentum of young stems

and leaves moderately dense, even, appressed, of stiff, straight unicellular

hairs 0.2-0. 3 mm long. G. tetragynus

Leaves linear-lanceolate, widest towards base; leaves, bracts, bracteoles

and sepals with prominent white thickened margins; indumentum of

young stems and leaves sparse, appressed, scabrous, of curved 1-2 celled

hairs 0.2-0. 3 mm long. G. chinensis

Etymology: The epithet 'hirtus' refers to the

long soft spreading indumentum of the young

stems and leaves, which this species shares with

G. longifolius.

Acknowledgements

I thank Tony Bean (BRI) for advice on

collections held in BRI, and for undertaking

fieldwork on my behalf in the Mt Maroon area.

Jim Croft (CANB) kindly provided working

space in that herbarium and access to the

collections.

References

Bell, S.A J. (2001). Notes on the distribution and conservation

status of some restricted plant species from sandstone

environments of the upper Hunter Valley, New South

Wales. Cunninghamia 7: 77-88.

Orchard, A.E. (1975). Taxonomic revisions in the family

Haloragaceae. I The genera Haloragis,

Haloragodendron, Glischrocaryon, Meziella and

Gonocarpus. Bull. Auckland Inst. Mus. 10: 1-299.

-(1977). Taxonomic revisions in the family Haloragaceae.

II Further notes on Haloragis, Haloragodendron

and Gonocarpus. Nuytsia 2: 126-144.

-(1986). New taxa in Gonocarpus and Haloragis

(Haloragaceae). Nuytsia 5: 327-339.

-(1990). Haloragaceae, pp.5-85, in A.S.George (ed.).

Flora of Australia vol. 18, Podostemaceae to

Combretaceae. AGPS, Canberra.

Orchard, new species Gonocarpus hirtus

965

Fig. 1. Gonocarpus hirtus. A. Habit. B. Leaf. C. Flower in axil of bract. D. Fruit. E. Distribution. (A-C. Telford 3525; D.

Forster, Bean & Bird PIF6633). Scales represent 1 cm (A), 5 mm (B), 1 mm (C, D).

Taxonomic notes on palms (Arecaceae) in catalogues of the Brisbane

Botanic Garden, Australia, of 1875 and 1885

John Leslie Dowe

Summary

Dowe, John Leslie (2004). Taxonomic notes on palms (Arecaceae) in catalogues of the Brisbane Botanic

Garden, Australia, of 1875 and 1885 , Austrobaileya 6 (4): 967-972. Two catalogues of plants cultivated in

Brisbane Botanic Gardens, Queensland, Australia, namely W. Hill’s Catalogue of the plants in the

Queensland Botanic Gardens , published in 1875, and F. M. Bailey’s Catalogue of plants in the two

metropolitan gardens, the Brisbane Botanic Garden and Bowen Park (The Garden of the Queensland

Acclimatisation Society), published in 1885, were studied in regards to palm nomenclature. Citations and

notes are provided for the entries Desmoncus minor, Jubaea speciosa, Pinanga smithii and Sagus blackalli.

Keywords:Arecaceae, Brisbane Botanic Garden, W. Hill, F.M. Bailey

John Leslie Dowe, Australian Centre for Tropical Freshwater Research, James Cook University, Townsville,

Queensland 4811, Australia

Introduction

While investigating the taxonomy of Australian

palms for the Flora of Australia treatment, I

examined two catalogues of the plants that were

being grown in Brisbane Botanic Garden,

Queensland, Australia. These catalogues,

published in 1875 and 1885 respectively, listed

both economically important and ornamental

species. Each catalogue was prepared in a

systematic format, with that by Hill (1875)

arranged according to Lindley (1836), the

Natural System of the Vegetable Kingdom,

while the other by Bailey (1885) was arranged

in the system of Bentham and Hooker (1862-

1883) as used in their Genera Plantarum. Both

catalogues were produced in hardbound

editions and widely distributed. In this paper I

will discuss the nomenclatural and taxonomic

implications of names in publications such as

botanic garden plant catalogues (see Mabberley,

1983; Ewan, 1993), with reference to the

International Code of Botanical Nomenclature

(ICBN) (Greuter etal., 2000). The introduction

and perpetuation of misapplied names, both

legitimate and illegitimate, to some palms in a

horticultural context, have been recognised

(Moore, 1971; Zona, 1990). Four names of

palms that appeared in Hill’s catalogue are

discussed, and their nomenclatural status

reconciled.

Accepted for publication 18 March 2004

Walter Hill’s Catalogue of 1875

Walter Hill (b.1820, d.1904) was appointed as

the first Superintendent of Brisbane Botanic

Gardens in 1855, first Queensland Colonial

Botanist in 1859, and retained both positions

until 1881 (Orchard, 1999). In Hill’s (1875)

catalogue, the palm family is termed

‘ Palmaceae\ the name used by Lindley (1836),

but now considered an obsolete name. The total

number of plant names included in the

catalogue was about 10 000. This number

comprised many hundreds of cultivar names.

Pyrus and Malus cultivars alone numbered

almost 300 entries. The catalogue presented

information in a tabulated form with the

column headings ‘Systematic name, and

authority’; ‘English or local name’; ‘Habit’;

and ‘Locality’.

The catalogue listed 155 names of palms

in 44 genera. Of these 46 (30%) are valid names

in current use. All but four of the remaining

119 names are recognised as validly published

synonyms of otherwise valid names in current

use. Therefore, 151 of Hill’s names can be

nomenclaturally reconciled. Three of the four

names - ‘ Desmoncus minor R. et R ’, ‘ Jubaea

speciosa H. K.’ and ‘ Sagus blackalli W. H.’ -

do not appear on available taxonomic databases

(Chapman, 1991; Index Kewensis, 1993; IPNI,

1999; APNI, 2001; TROPICOS, 2001); the

fourth name, ‘ Pinanga smithi W. H.’, did not

appear in either Chapman (1991) or APNI

(2001), but was present in Index Kewensis

968

Austrobaileya 6 (4): 967-972 (2004)

(1993) and IPNI (1999).

Names published in systematically

arranged lists, such as Hill’s catalogue, become

part of the taxonomic literature and may be

accounted for in subsequent accounts and

revisions (Barker and Barker, 1990). However,

the validity of a name is dependent upon there

being a description that, ideally, allows

recognition of the species, or if there is a

reference to a specimen that provides identity

for the name. Illegitimate names are rejected

according to the rules in the ICBN (Greuter et

al., 2000). However, such names may be

accounted for in subsequent taxonomic

treatments of Australian palms, albeit within

the nomina dubia et excludenda section.

A search was instigated of the two

herbaria, BRI and MEL, where Hill was known

to deposit most of his specimens, but no

specimens related to any of the obscure names

were located. A search was also made of the

records of the Brisbane Botanic Garden, but

apart from the original citation in the catalogue,

no further evidence of the names was revealed.

In regards to those names in genera of American

palms, i.e. Desmoncus minor and Jubaea

speciosa, the records of the following herbaria

were examined: MO, NY, TRIN and US.

Frederick Manson Bailey’s catalogue of

1885

FrederickM. Bailey (b.1827, d. 1915) succeeded

Hill as Queensland Government Botanist in

1881, a position that he held until 1915

(Orchard, 1999). In Bailey’s (1885) catalogue,

the palm family is referred to as the Palmae, a

name now conserved along with Arecaceae as

a valid alternative name (Greuter et al., 2000).

The number of names in all families in Bailey’s

catalogue was reduced to about 3000, mainly

due to the absence of cultivar names. However,

it included additional information about the

plants, with “ ...numerous notes on the

properties and uses of the plants (are) a feature

the compiler feels sure will be appreciated by

a large number of persons, especially by those

who take a utilitarian view of them”. Unlike

Hill’s catalogue, Bailey’s catalogue did not have

a tabular format, but it did include utilisation

notes following many entries, as well as concise

information on habit and origin. Considering

the palms, Bailey included 91 names compared

to Hill’s 155, but more generic names, 57 as

compared to 44. There were several reasons for

these changes. A primary reason for the

reduction in species names would have been

due to the demise of those species that were

culturally inappropriate for the warm temperate

climate of Brisbane. Bailey had also

implemented synonymy where required, and

had adopted many of the new generic names

that were the result of revisions that had been

completed in the decade since Hill’s catalogue

was published. For example, the 13 species

previously listed under the single genus name

Areca in Hill’s catalogue were subsequently

included in seven genera in Bailey’s catalogue,

while the six species in Hill’s Kentia were

divided into five genera in Bailey’s catalogue.

These two genera alone accounted for an

increase of ten generic names.

Taxonomy and nomenclature

The decade 1875-1885 was one of considerable

activity in Australian palm taxonomy.

Wendland and Drude (1875) published the first

detailed account of Australian palms in their

Palmae Australasicae, Bentham (1878)

published his account of palms in volume seven

of Flora Australiensis, and Mueller (1875-

1881) had entered his most active period of

palm taxonomy. The nomenclatural changes

introduced by these accounts were reflected in

the names used in Bailey’s 1885 catalogue,

when compared to Hill’s catalogue of ten years

earlier. A comparison can be made between the

numbers of names that are in current use that

were used in each catalogue. Whereas only 30%

(46 of 155) of the names that Hill used are in

current use, 56% (51 of 91) of the names used

by Bailey are in current use. For example, Hill

listed Ptychosperma elegans under two names

- Pinanga smithii and Seaforthia elegans - and

Caryota mitis also under two names - C.fmfuracea

and C. sobolifera. However Bailey listed

Ptychosperma elegans only once and under its

‘new’ name, and only Caryota sobolifera for

the two Caryota names. Bailey adopted all the

name changes in the accounts of Wendland and

Drude, Bentham and Mueller, among others,

and was also more conservative in his use of

‘obscure’ names than was Hill.

Dowe, taxonomic notes on palms (Arecaceae)

969

Obscure and neglected names in Hill

(1875)

‘Desmoncus minor, R. et P.’: in W. Hill,

Catalog, pi. Brisbane bot. gard. 21 (1875),

‘evergreen climber, Trinidad’.

Attributing authorship of this name to Ruiz and

Pavon, I propose, is an error. The palm

taxonomy of these botanists is primarily

confined to two publications (Ruiz and Pavon,

1794, 1798) in which they named 16 species

(Henderson, 1995; IPNI, 1999). Their

taxonomic activity occurred three decades prior

to the establishment of Desmoncus by Martius

(1824), so they could not have had any

connection with the taxonomy of that genus

using that name. According to Uhl &

Dransfield (1987), Desmoncus has not received

a recent critical revision, and much of the

nomenclature of the genus is unresolved. Apart

from appearing in Hill’s catalogue, the name

also appears on a specimen held at TRIN. The

specimen is Broadway 5568, the type for Desmoncus

prestoei L. H. Bailey (= D. polyacanthus Mart.),

collected in 1891 from a plant cultivated in the

Trinidad and Tobago Botanical Garden. The

name Desmoncus minor is therefore most likely

related to plants growing in Trinidad and

Tobago Botanical Gardens. Hill probably

received material under this name from

Trinidad as part of the exchange program with

Brisbane Botanical Gardens.

Bailey (1943), in describing Desmoncus

prestoei L. H. Bailey, noted that the name

D. minor Prestoe appeared in “...the Hart

catalogue [of 1908]... without description or

comment...”, and was the name proposed by

Prestoe for the Broadway specimen mentioned

above. Bailey referred to D. minor as a nomina

nuda and later (Bailey, 1947) as a “floating

herbarium name”, and therefore determined

that it could not be taken up either as a name

or synonym. According to Article 7.1 [no type]

and Article 32.1 [no diagnosis or reference to

previous effective publication] in the ICBN,

(Greuter et al., 2000), the name ‘ Desmoncus

minor ’ is to be rejected.

‘Jubaea speciosa H. K.’: in W. Hill, Catalog,

pi. Brisbane bot. gard. 23 (1875),

‘evergreen tree, Mauritius’.

Kunth (1816) described Jubaea spectabilis

Kunth (= J. chilensis (Molina) Baillon) for a

species growing in Chile. A search of the

available databases was unable to detect the

combination ‘ Jubaea speciosa'. The epithet

‘speciosa ’ is suspected to be an orthographic

misinterpretation of 'spectabilis'. Plants of

J. chilensis are extant in City Botanic Gardens,

Brisbane, and are assumed to have been

acquired during the years in question.

According to Article 60.1 [incorrect spelling]

in the ICBN, (Greuter et al., 2000), the name

‘ Jubaea speciosa' is to be rejected.

‘Pinanga smithi W. H.’: in W. Hill, Catalog,

pi. Brisbane bot. gard. 20 (1875);

Scheffer, Ann. Jard. Bot. Buitenzorg 1:

154 (1876) [as Pinanga smithii]', J. D.

Hooker, Bot. Mag. 3rd ser., 50: t. 7345

(1894); Martelli, Nuovo Giorn. Bot. Ital.

ser. 2, 42: 72 (1935) = Ptychosperma

elegans (R.Br.) Blume.

Type citation: ‘evergreen tree, Cape York’.

Type: Cultivated plant in Brisbane

Botanic Garden, not extant.

The name Pinanga smithii, included within the

tribe Areceae, first appeared in Hill’s 1875

catalogue. Scheffer (1876, p. 154) related the

name to Ptychosperma elegans : “...nous avons

re§u ce palmier du jardin botanique de

Melbourne, sous le nom de Pinanga Smithii.".

Scheffer provided a description based upon

those plants. Subsequently, Index Kewensis did

not reference the name to Hill, but to Scheffer.

Beccari (1885), in discussing palms that

were growing in Bogor Botanic Gardens,

reconfirmed the identity of Pinanga smithii as

Ptychosperma elegans, though he described his

specimen as a subspecies, P. elegans var.

sphaerocarpa Becc. Beccari provided a

diagnostic illustration that allows identification

as P. elegans. Hooker (1894) subsequently used

P. smithii in synonymy under an illustration of

P. elegans in Curtis’s Botanical Magazine. The

source material for the illustration was a plant

in the Royal Botanic Gardens Kew, but its

origin was not noted as there was no available

record of its introduction. Hooker supposed that

the epithet ‘ smithii' originated in “...some

continental gardens to which a young plant had

been contributed from Kew, and to which was

970

given the name of the late Curator of that

establishment, whose success as a raiser of

palms was famous”. The Smith to whom

Hooker most likely referred, was John Smith

(b. 1798, d. 1888), the first curator of the Palm

House at Kew (1841-1864) (Turrill, 1959;

King, 1985; Minter, 1991). However, this

version of the origin of the epithet appears to

have been only speculation by Hooker. Hill

and Scheffer did not provide any information

in this regard. Hill indeed cited himself as the

author in the first publication of the name.

The most recent use of the name was by

Martelli (1935), who included it in a list of

species names in the Areceae, but indicated that

it was a synonym of Normanbya muelleri

(W.Hill) Becc. (= Normanbya normanbyi

(W.Hill) L.H. Bailey), a determination that was

clearly incorrect. A search of the records of the

Melbourne Botanic Gardens’ living plant

censuses, the National Herbarium of Victoria

collection (MEL) and the index of plant names

in the Mueller Correspondence Project, did not

reveal the name Pinanga smithii (C. Coles and

F. Anderson, pers. comm.). However, a

specimen determined as P. elegans by Mueller

in MEL, was despatched by Hill to Mueller in

June 1875, and accompanied by a letter that

described the palm as “...found by me at Cape

York... the habit resembles the Seaforthia

elegans, and grows about the same height.”

In recent accounts, the name Pinanga

smithii has ceased to be used, and it does not

appear, to my knowledge, in any relevant

taxonomic accounts after 1935 such as Moore

(1963), Essig (1978), Chapman (1991) or APNI

(2001). The name Pinanga smithii has

evidently become neglected, but as a legitimate

synonym should be included under

Ptychosperma elegans, and with the following

authorship, Pinanga smithii W.Hill ex. Scheff.

i Sagus Blackalli W. H.’: in W.Hill, Catalog,

pi. Brisbane bot. gard. 21 (1875),

‘evergreen tree, Cape York’. It is probable

that the etymology of this name was to

honour Samuel Wensley Blackall,

Governor of Queensland, 1868-1871.

The name Sagus blackalli, included within the

tribe Calameae, was, to my knowledge, only

ever used in Hill’s catalogue, where it was noted

as an ‘evergreen tree, Cape York’. It is absent

from all plant name databases known to me. It

Austrobaileya 6 (4): 967-972 (2004)

is not known if Hill had intended to describe a

species under this name, as no specimens or

correspondence to that effect have been located.

Sagus Steck is now a synonym of

Metroxylon Rottb. and Sagus Gaertn. a

synonym of Raphia Beauv. The name Sagus

was replaced by those new generic names many

decades before Hill’s catalogue appeared. This

lends credence to the possibility that Hill had

not intended to use Sagus, but he used that

spelling inadvertently for another genus. One

possibility is that he had meant to use the name

Saguerus Steck, now a synonym of Arenga

Labill. Wendland and Drude (1875) described

Saguerus australasicus H. Wendl. & Drude (=

Arenga australasica (H. Wendl. & Drude) S.

T. Blake) from Cape York, but whether Hill

had intended any connection with that genus,

albeit as being misspelt, is not known. It is

assumed that the monograph on Australian

palms by Wendland and Drude (1875) was not

yet available to Hill, although it cannot be

discounted that he was aware of proposed

manuscript names. Hill placed S. blackallii,

along with two other species of Sagus, in the

Tribe Calameae, which is the correct systematic

placement for Sagus and its synonyms, whereas

he placed his Arenga species in the tribe

Areceae. If he had intended the name to be

Saguerus, it would have been more than likely

that he would have placed S. blackalli at least

near Arenga in the list, as it was a systematic

rather than alphabetic arrangement. However,

without specimens or documentation, it is only

speculation that Hill had intended otherwise.

According to Article 7.1 [no type], Article 9

[identity ambiguous] and Article 32.1 [no

diagnosis or reference to previous effective

publication] in the ICBN, (Greuter et al., 2000),

the name ‘ Sagus Blackalli ’ is to be rejected.

Summary

Nelson (1990), in commenting on the

taxonomic and nomenclatural implications of

names published in plant catalogues and

otherwise described from cultivated specimens,

highlighted the problems that may arise when

such publications are overlooked, particularly

in relation to obscure names, or names that

otherwise may have become neglected. Plant

catalogues and similar publications cannot be

ignored as sources of taxonomic and

nomenclatural information.

Dowe, taxonomic notes on palms (Arecaceae)

Acknowledgements

My appreciation goes to Rod Henderson (BRI),

Anders Barfod (AAU) and Betsy Jackes (JCT)

for constructive comments on this paper. Fleur

Anderson and Cathryn Coles (MEL), Andrew

Henderson (NY), Yasmin S. Baksh-Comeau

and Prudence Roberts (TRIN), and Ken Hill

(NSW), are thanked for their assistance.

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Canberra.

Bailey, F. M. (1885). Catalogue of plants in the two

metropolitan gardens, the Brisbane Botanic

Garden and Bowen Park (The Garden of the

Queensland Acclimatisation Society). Brisbane.

Bailey, L. H. (1943). Desmoncus. Gentes Herbarum 6: 211—

218.

-(1947). Indigenous palms of Trinidad and Tobago.

Gentes Herbarum 7: 353-445.

Barker, R. M. & Barker, W. R. (1990). Botanical

contributions overlooked: the role and recognition

of collectors, horticulturists, explorers and others in

the early documentation of the Australia flora. In: R

S. Short (ed), History of systematic botany in

Australasia, 37-85. South Yarra.

Beccari, O. (1885). Reliquiae Schefferianae. Illustrazione di

alcune palme viventi nel giardino botanico di

Buitenzorg. Annales du Jardin Botanique de

Buitenzorg 2: 77-171.

Bentham, G. (1878). Flora Australiensis 7. London.

Bentham, G. & Hooker, J. D. (1862-1883). Genera

plantarum ad exemplaria imprimis in Herbariis

kewensibus servata de finita auctoribus G.

Bentham et J. D. Hooker. London

Chapman, A. D. (1991). Australian plant name index.

Canberra.

Essig, F. B. (1978). A revision of the genus Ptychosperma

Labill. (Arecaceae). Allertonia 1: 415-478.

Ewan, J. (1993). An overlooked printed catalogue of plants

in the botanick garden of South-Carolina, 1810.

Taxon 42: 365-367.

Greuter, W., McNeill, J., Barrie, F. R., Burdet, H. M.,

Demoulin, V., Filgueiras, T. S., Nicolson, D. H.,

Silva, P. C., Skog, J. E., Trehane, P., Turland, N. J.

& Hawksworth, D. L. (2000). International Code

of Botanical Nomenclature (Saint Louis Code).

Regnum Veg. 138.

Henderson, A. (1995). The palms of the Amazon. New York.

Hill, W. (1875). Catalogue of the plants in the Queensland

Botanic Gardens. Brisbane.

971

Hooker, J. D. (1894). Ptychosperma elegans. Botanical

Magazine 3rd series, vol. 50: t. 7345.

Index Kewensis. (1993). Index Kewensis: on compact disc.

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IPNI. (1999). International plant name index. WWW

version. Royal Botanic Gardens, Kew, Harvard

University Herbaria and Australian National

Herbarium.

King, R. (1985). Royal Kew. London.

Kunth, C. S. (1816). Nova genera et species plantarum.

Paris.

Lindley, J. (1836). A natural system of botany: or, a

systematic view of the organisation, natural

affinities, and geographical distribution of the

whole vegetable kingdom. London.

Mabberley, D. J. (1983). Dr Smith’s Anemia, or, the

prevention of later homonyms. Taxon 32: 79-87.

Martelli, U. (1935). La sinonomia delle palme gerontogee

della tribu delle Areceae. Nuovo Giomale Botanico

Italiano, n. s. 42: 17-88.

Martius, C. F. P. (1824). Palmarumfamilia ejusque genera.

Munich.

Mustier, S. (1991). The palm house at Kew: a new beginning.

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Moore, H. E. JR. (1963). An annotated checklist of cultivated

palms. Principes 7: 119-182.

-(1971). Notes on Sabal in cultivation. Principes 15: 69-

73.

Mueller, F. Yon (1875-1881). Fragmenta phytographiae

Vols 9-11.

Nelson, E. C. (1990). ‘.. .and flowers for our amusement’:

the early collecting and cultivation of Australian

plants in Europe and the problems encountered by

today’s taxonomists. In: P. S. Short (ed), History of

systematic botany in Australasia, 285-296. South

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Australia. Flora of Australia, 2nd edition, 1:11-

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Ruiz Lopez, H. & Pavon, J. (1794). Flora peruvianae et

chilensis prodromus, sive novorum generum

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descriptiones et icones. Madrid.

-(1798). Systema vegetabilium florae peruvianae et

chilensis. Madrid.

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Notelaea ipsviciensis (Oleaceae), a new species from south east Queensland

Wayne K. Harris

Summary

Harris, W.K. (2004). Notelaea ipsviciensis (Oleaceae), a new species from south east Queensland.

Austrobaileya 6 (4): 973-976. Notelaea ipsviciensis is described and notes are provided on its habitat,

distribution and conservation status. It is found in a very restricted area of south east Queensland.

Key words: Queensland, Oleaceae, Notelaea ipsviciensis.

W.K Harris, Queensland Herbarium, Brisbane Botanic Gardens, Mt Coot-tha, Mt Coot-tha Road, Toowong,

Queensland 4066, Australia

Introduction

The genus Notelaea Vent., in Queensland

comprises 11 species, three of which are new.

During studies on the systematics of Notelaea

and related genera in the southwest Pacific,

Lloyd Bird of Bundamba, brought to my

attention a rare and unusual form in the genus,

which appeared to be restricted to the Ipswich

area. This species is here described as new. The

most notable features of this species were its

habit and leaf venation in which the secondary

venation arises more or less at right angles from

the mid-vein. The full taxonomic treatment of

the genus is to be published separately.

Notelaea ipsviciensis W.K.Harris, sp. nov

affinis N. ovatae sed foliis

lanceolatioribus (ovata in N. ovata ) et

venatione secundaria sub angulo paene

90° abeunti (adversus angulum acutiorem

in N. ovata ) differt. Typus: Queensland,

Moreton District: Rhondda Collieries,

Bergen’s Hill, Bundamba, Ipswich, Aug

1985, Bird s.n., (holo: BRI, [AQ

442082]). Fig. 2.

Notelaea sp. (Bundamba L.H. Bird

AQ442082) in Henderson (2002).

Evergreen shrubs 0.5-1.5m tall, multi¬

stemmed, forming lignotubers, stems erect or

ascending 2-4 cm in diameter: bark pale grey,

smooth. Branchlets puberulent with erect

simple hairs, glabrescent. Leaves glabrous

except puberulent base and midrib; lamina

coriaceous, oblanceolate to narrow lanceolate,

punctate above and below, punctae often more

prominent on abaxial surface, 4-8 cm long,

Accepted for publication 17 August 2004

0.8-1.2 cm wide (length/breadth ratio 7.5 -10:1);

margin entire, slightly recurved: apex acute to

acuminate; base narrow cuneate to attenuate

into the petiole; venation distinct and

prominently raised above and below, tertiary

venation distinct, secondary veins 9-12 pairs;

juvenile leaves lanceolate, 6-10 cm long and

0.8-1.2 cm wide. Inflorescence axillary, up to

three per axil, metabotryoids, 5-9 -flowered,

1-2.5 cm long; axes puberulent, sometimes

glabrous apically; bracts ovate to acute, sparsely

puberulent on the outside, glabrous on the

inside, 1-1.2 mm long, persistent; bracteoles

linear to lanceolate, sparsely puberulent on the

outside, glabrous inside, 0.8-1.2 mm long,

caducous. Flowers, pale cream to yellow;

pedicels sparsely puberulent to glabrous,

articulate at base, 2-6 mm long. Sepals 4,

glabrous or ciliolate at the apex, triangular,

apex acute to apiculate, 0.2-0.5 mm long,

0.4-0.6 mm wide. Corolla induplicate-valvate

in bud, 4-lobed; lobes ovate, concave, 1-1.8 mm

long, 0.9-1 mm wide, joined in pairs above the

base of the filaments for c. 0.5 mm, glabrous.

Stamens 2, glabrous, enclosed within the

concave corolla segments; filaments c. 0.2 mm

long with a blunt terminal umbo; anthers 1-1.5

mm long, 1.2-1.4 mm wide. Ovary glabrous,

flask shaped, 0.6-1 mm long at anthesis; style

c. 0.1 mm long; stigma shortly 2-lobed, yellow-

brown, c. 0.3 mm long. Drupe dark blue to

purple when ripe, ovoid, 8-10 mm long, 6-8

mm diameter; mesocarp c. 1 mm thick;

endocarp woody, c. 0.4 mm thick. Fig. 1.

Specimens studied : Queensland. Moreton: Ipswich

Bundamba 1km S of Barclay St Rhondda Colliery Land,

27.62°S, 152.80°E, Nov 1987, Bird, s.n., [AQ 436255]

(BRI); Ipswich Bundamba Bergens Hill Rhondda Colliery,

27.25°S, 151.91°E, Nov 1985, Bird, s.n., [AQ 441680]

(BRI); Ipswich Bundamba 1km S of Barclay Stn Rhondda

974

Austrobaileya 6 (4): 973-976 (2004)

Fig. 1. Notelaea ipsviciensis. A. leafy shoot x 1. B. inflorescence x 4. C. Flower x 8. D. Section of flower x 8. Bird s.n. AQ

489221 (BRI).

Colliery, 27.25°S, 151.91°E, Dec 1985, Bird, s.n., [AQ

441681] (BRI); Ipswich Dinmore, 27.62°S, 152.79°E,Aug

1985, Bird, s.n., [AQ 442081] (BRI); Ipswich Bundamba

Rhondda Colliery land end of Barclay St, 27.60°S, 152.82°E,

Nov 1988, Bird, s.n., [AQ 453056] (BRI); Ipswich 2km SE

of Bundamba, 27.60°S, 152.82°E, Oct 1988, Bird, s.n., [AQ

454151]; Whitewood Road Ebbw Vale Ipswich 400m E of

the road, 27.62°S, 152.79°E, Jun 1996, Bird, s.n., [AQ

489221] (BRI); 1km S of Dinmore, 27.58°S, 152.91°E, Jun

1996, Bird, s.n., [AQ 586175] (BRI).

Distribution and ecology: Confined to a very

few localities in the Ipswich area of southeast

Queensland, where it occurs in dry sclerophyll

eucalypt forest.

Phenology: Flowers appear in winter in July

and fruits appear shortly after and are mature

by October.

Notes: This species has its closest affinities with

N. ovata but it differs from this species in

having lanceolate leaves and a distinct venation

pattern with the secondary venation on mature

leaves arising close to 90° from the mid-rib. It

is possible that this species is a hybrid between

N. lloydii Guymer and N. ovata R.Br. N. lloydii

grows in close proximity but the other putative

parent does not occur in the immediate vicinity.

Harris, W.K., Notelaea ipsviciensis

975

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976

Austrobaileya 6 (4): 973-976 (2004)

Conservation status : This species was probably

more widespread in the Ipswich district but is

now known from only two localities with a total

of about twelve plants. It therefore satisfies

several criteria of the IUCN (Anonymous 2001)

categories of rare and threatened plants,

especially population size of less than 50

individuals surviving in the wild and a decline

in area of occupancy such that the population

has been severely fragmented. It is suggested

that the species is critically endangered with

an IUCN category of CR

Etymology : The specific epithet is a derivation

of the name of the nearby city of Ipswich.

Acknowledgements

I am indebted to Lloyd Bird of Ipswich for his

encouragement and company in the field. Will

Smith (BRI) provided the illustrations of this

species. Les Pedley provided the Latin

translation of the diagnosis.

References

Anonymous (2001). IUCN Red List Categories and Criteria.

International Union for Conservation of Nature and

Natural Resources: Switzerland.

Henderson, R J.F. (ed.) (2002). Names and Distribution of

Queensland Plants, Algae and Lichens. Brisbane:

Queensland Environmental Protection Agency.

Austrobaileya 6(4): 977-978 (2004)

Note

A new combination in Centratherum Cass. (Asteraceae)

Centratherum is a small genus in the tribe

Vernonieae, subtribe Centratherinae. It was

characterised by Robinson et al. (1980) by the

leafy outer involucral bracts, long-stalked

glands on the corolla, and the pappus being

short and caducous, or absent.

It occurs mainly in South America, but it

does occur in the Philippines and in Australia.

Two taxa occur in Australia; one is native and

the other introduced (Kirkman 1981).

Bentham (1867) identified the Australian

taxon as C. muticum (Kunth) Less, (based on

Ampherephis mutica Kunth), and this name

was applied to Australian material for many

years. Kirkman (1981) placed C. muticum in

the synonymy of C. punctatum Cass., a South

American species. The type of Ampherephis

mutica was collected in South America. From

the detailed illustration and description included

in the protologue, it is clear that C. muticum is a

synonym of C. punctatum sens. str.

Kirkman (loc. cit .) described C. punctatum

subsp. australianum for the Australian taxon.

However, I consider species rank is appropriate

(see below). I have been unable to find any prior

species name applicable to the Australian taxon.

Centratherum australianum (K.Kirkman)

A.R.Bean, comb, et stat. nov. C. punctatum

subsp. australianum K.Kirkman, Rhodora

83: 21 (1981). Type: New South Wales.

North Coast: west of Wingham on Bulga

road, 12 April 1953, J. Vickery 23846

(holo: NSW; iso: L (photo!), MO, n.v.).

C. punctatum and C. australianum are similar

in several respects. They both have: dimorphic

involucral bracts, with the outer ones

foliaceous; T-shaped hairs on the branchlets

(although the ‘stalk’ is often obsolete in

C. australianum)-, caducous pappus

comprising sub-plumose ensiform bristles

(terminology from Bean 2001); and

longitudinally 10-ribbed cypselas. However,

they are readily distinguished by several

characters (outlined in the key). Furthermore,

they originated in different continents, with no

opportunity for genetic interchange.

Key to Centratherum species in Australia

Leaves oblanceolate, with 1-8 pairs of obtuse teeth, occasionally entire;

capitulum 8-14 mm diameter; corolla tube glabrous or with a few sessile

glands; pappus bristles with <50 pectines, each c. 0.05 mm long.C. australianum

Leaves obovate, with 10-18 pairs of acute teeth; capitulum 20-25 mm

diameter; corolla tube with sessile glands and stalked glands to 0.15 mm

long; pappus bristles with >100 pectines, each 0.1-0.15 mm long.*C. punctatum

C. australianum is undoubtedly a native

Australian plant. It was collected in 1804 by

Robert Brown at 'Port Jackson' (now Sydney),

and by other pioneer collectors. I have observed

it growing in intact non-weedy eucalypt forest

away from roads and other disturbance. It is

distributed in coastal eastern Australia from

Sydney, N.S.W. to Shoalwater Bay, Qld. The

alleged occurrence in the Northern Territory

shown by Kirkman {loc. cit.) is erroneous.

Accepted for publication 24 December 2003

C. punctatum sens. str. was introduced to

Australia as an ornamental plant in the mid-

20th century, and has become sparingly

naturalised in northern Australia, from

Kununurra, W.A. to Murwillumbah, N.S.W.

Both species are illustrated in the Flora

of New South Wales (Porteners 1992).

References

Bean, A.R. (2001). Pappus morphology and terminology in

Australian and New Zealand thistles (Asteraceae,

tribe Cardueae). Austrobaileya 6: 139-52.

978

Bentham, G. (1867). Centratherum. In: Flora Australiensis

Vol. 3, p. 460. L. Reeve & Co.: London.

Kirkman, L.K. (1981). Taxonomic Revision of Centratherum,

and Phyllocephalum (Compositae: Vernonieae).

Rhodora 83: 1-24.

Austrobaileya 6 (4): 977-978 (2004)

Porteners, M.F. (1992). Centratherum. In: G.J. Harden (ed.).

Flora ofNew South Wales 3:147. New South Wales

Press: Sydney.

Robinson, H., Bohlmann, F. & King, R.M. (1980).

Chemosystematic Notes on the Asteraceae; Natural

subdivisions of the Yemonieae. Phytologia 46:421-36.

A.R. Bean

Queensland Herbarium, Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha Road,

Toowong, Queensland 4066.

Austrobaileya 6 (4): 979-981 (2004)

979

Note

Nomenclatural changes for two Australian species of Livistona R. Br. (Arecaceae)

The treatment of Arecaceae for the Flora of

Australia is presently being prepared.

Taxonomic considerations related to the

treatment are being dealt with in advance of

publication, and proposed name changes of two

Australian Livistona species are presented here.

Livistona decora (Bull) Dowe comb. nov.

Corypha decora W. Bull, Catal. (1887)

10. Type: Australia. Queensland,

Yepoon-Emu Park Rd, c. 11 km S of

Yepoon, A.N.Rodd 3069 with S. Jacobs,

12 May 1976 (neo: BRI; isoneo: BH, K,

NSW, here designated).

Livistona decipiens Becc., Webbia 3 (1910)

301, syn. nov. Type: France. Cultivation.

Nice, May 1908, ARobertson-Proschowsky

s.n. (lecto: FI) (fide Rodd, 1998).

[Livistona inermis auct. non R.Br., Wendland

and Drude, Linnaea 39 (1875) 229, pro.

part.]

[Livistona australis auct. non (R.Br.) Mart.,

Bentham, FI. Austral. 7 (1878) 146, pro.

part.]

Livistona decipiens var. polyantha Becc.,

Webbia 5 (1921) 15,18. Type: Indonesia.

Cultivation. Bogor Botanic Gardens,

undated, Beccari s.n. (holo: FI; iso: BO).

The new combination for this species is based

on the name Corypha decora W. Bull, which

was first used by Bull (1887) for plants that

were grown in his nursery at Chelsea, England,

and subsequently discussed by Watson (1889)

in a paper on the cool cultivation of tropical

and subtropical plants. Bull’s (1887, p.10)

description was: “ Corypha decora: An elegant

and ornamental species introduced from

Queensland. It early develops characterized

leaves, which are fan-shaped in outline and

divided almost to the petiole into linear

lanceolate segments each about half-an-inch in

breadth. The petioles are sparsely furnished

with small hooked prickles. This species will

undoubtedly be found one of the most useful of

greenhouse palms”. Watson’s (1889, p. 294)

Accepted for publication 26 July 2004

account of this palm, which he incorrectly

identified as L. inermis R.Br., included: “...The

most interesting of this genus, however, is a

very fine example of the rare L. inermis of

R.Brown, in the gardens of Villa Valetta. It has

a bare stem 6 feet high by Wi feet in diameter

at the base. The head is made up of a large

number of shining green leaves, the petiole of

which is 6 feet long and margined with spines

at the base. The blade is divided almost to the

base, and it has a distinct midrib that is curved

so as to produce a very extraordinary effect.

The form of the leaf may be called a

combination of the pinnate and palmate

characters. A plant of this rare palm is in the

Kew collection; there is also a fine example of

it in the collection at Blenheim. A large

specimen was also noted in Baron Vigier’s

garden at Nice. Mr Bull distributed plants of it

some years ago under the name of Corypha

decora ”,

The validation of the name C. decora

precedes that of L. decipiens, and therefore has

priority over it. The descriptions provided by

Bull and Watson are attributable to the taxon.

The place of publication is also acceptable as

the rules of nomenclature allow for the

validation of names published under such

circumstances.

Beccari (1910) was apparently unaware

of the use of the name Corypha decora when

he named Livistona decipiens from the

collection of Robertson-Proschowsky s. n. [FI]

taken from a cultivated plant at Nice, France,

and named for the deception that it had caused

as to its true identity: “...// nome di L. decipiens,

per gli errori di cui e stata causa. In

cultivation in the Riviera during the late 1800s

it had incorrectly been referred to by local

horticulturists as ‘Copernicia cerifera ’, and it

was with the intention of clarifying the species’

identity that Beccari provided the description

and established the name. Although unaware

of its origin, but suggesting that it came from

eastern tropical Australia, Beccari (1921,1931)

related it to other Australian species, primarily

L. australis. Despite it being well known and

widely cultivated throughout the world, its

980

native origin continued to remain speculative

until relatively recent times. For example,

Bailey (1976) noted (with reference to the name

L. decipiens ): “...described from cult.,

supposedly Australian...”. The first unequivocal

application of L. decipiens to natural

populations was by Johnson (1981) who

recorded distribution of the species from

Miriamvale to Townsville in Queensland.

Through a lack of understanding of the

correct identity of L. inermis , and the

incorporation of characteristics of misidentified

specimens of other species into descriptions,

L. decora was inadvertently identified as that

species in a number of instances. Both

Wendland and Drude (1875) and Bentham

(1878) cited the MEL Thozet s.n. specimen

from Moore’s Creek near Rockhampton, central

Queensland (now known to be of L. decora) in

their accounts ofL. inermis R.Br. andL. australis

(R.Br.) Mart, respectively. Upon this basis,

many specimens in both the wild and in

cultivation with a deeply segmented leaf were

identified as L. inermis sensu H.Wendl. (Hill,

1873,1875). It is highly probable that L. inermis

R.Br. in the strict sense was never in cultivation

in Europe as it is an exceedingly difficult plant

to propagate and maintain while L. decora seed

is easy to germinate, and the plants are one of

the fastest growing species of Livistona. The

specimen A.N.Rodd 3069 with S. Jacobs,

collected from the Yepoon-Emu Park Rd, c. 11

km S of Yepoon at BRI is chosen as the neotype

as it is from the area where seeds may originally

have been collected for distribution to England

and Europe.

Livistona nasmophila Dowe & D.L. Jones

nom. & stat. nov. Livistona mariae subsp.

occidentalis A.N. Rodd syn. nov., Telopea

8: 81 (1998). Type: Australia. Western

Australia, Durack Range, SE base of Mt

King, A.N. Rodd 2868 (holo: NSW; iso:

BH, K, PERTH, QRS).

Rodd (1998) established this taxon as L. mariae

subsp. occidentalis A.N.Rodd, based on the

NSW collection A. A. Rodd 2868 from Mt King,

Western Australia. In the prologue, Rodd wrote:

“Recognition of this population as a separate

subspecies on the basis of a single wild

collection (and one from cultivation) is arguably

Austrobaileya 6 (4): 979-981 (2004)

rather premature”. Following fieldwork in the

Kimberleys, with collections of fruit and

flowers from both wild and cultivated sources,

we established that this taxon was distinct

enough from both L. mariae F.Muell. and

L. rigida Becc. to be recognised at species

level. The correct nomenclatural procedure then

would normally have been to take Rodd’s

subspecific epithet ‘ occidentals and use it as

the specific epithet, but the name ‘ Livistona

occidentalis ’ has previously been used by

Hooker (1884, p. 64) as a synonym for Brahea

dulcis Mart, and therefore blocks transfer of

Rodd’s epithet to species rank. The new epithet,

‘nasmophila ’ was chosen to recognise the

palm’s habit of occupying permanent

watercourses fed by springs through much of

its range.

The relationships of L. nasmophila are

with L. mariae , but probably not as close as

suggested by Rodd (1998). Petiole armature in

L. nasmophila is considerably less than that in

L. mariae. In L. nasmophila, the inflorescence

bracts are glabrous rather than tomentose, and

branching of the partial inflorescences is to five rather

than four orders as in L. mariae. In L. nasmophila,

fruit is purple-black rather than black as in L. mariae.

References

Bailey, L.H. (1976). Hortus Third. Macmillan Publishing,

New York.

Beccari, O. (1910). La " Copemicia cerifera" in Riviera ed

una nuova specie di Livistona. Webbia 3: 295-305.

-(1921). Recensione delle Palme del vecchio mondo

appartenenti alia tribu delle Corypheae, con

descrizione della specie e varieta nuove che vi

appartengono. Webbia 5: 5-70.

-(1931). Asiatic palms - Corypheae (ed. U.Martelli). Ann.

Roy. Bot. Gard. (Calcutta ) 13: 1-356.

Bentham, G. (1878). Palmae. Flora Australiensis 7: 132-

147.

Bull, W. (1887). Catalogues of new beautiful and rare

plants. London.

Hill, W. (1873). Report. In: G.E. Dalrymple (ed.), Narrative

and reports of the Queensland north-east coast

expedition, 1873, 48-53. James C. Beal,

Government Printer, Brisbane.

-(1875). Catalogue of the plants in the Queensland

Botanic Gardens. James C. Beal, Government

Austrobaileya 6 (4): 979-981 (2004)

Printer, Brisbane.

Hooker, J.D. (1884). Report on the progress and condition

of the Royal Gardens at Kew during the year 1882.

Eyre and Spottiswoode, London.

Johnson, R.W. (1981). The Queensland palm flora. Qld

Naturalist 22: 10-20.

981

Rodd, A.N. (1998). Revision of Livistona (Arecaceae) in

Australia. Telopea 8: 49-153.

Watson, W. (1889). Cool cultivation of tropical and sub¬

tropical plants. Kew Bull. 3: 287-306.

Wendland, H. & O. Drude (1875). Palmae Australasicae.

Linnaea 39: 153-238.

John L. Dowe

Australian Centre for Tropical Freshwater Research, Janies Cook University, Townsville,

Qld 4811, Australia, email: John.Dowe@jcu.edu.au

David L. Jones

Centre for Plant Biodiversity, Australian National Herbarium, G.P.O. Box 1600,

Canberra, ACT 2601, Australia

Austrobaileya 6 (4): 983 (2004)

Note

Reduction of Acacia perangusta to the synonymy of A. fimbriata.

* Acacia fimbriata is widespread in eastern

Australia from about Rockhampton to near

Nowra, New South Wales. It extends inland to

the Carnarvon National Park, and there is an

early record (before 1925) from Ravenshoe, c.

35 km south of Atherton, which must be

considered doubtful. The dimensions of the

species’ phyllodes and the density of hairs on

branchlets and margins of phyllodes vary widely.

White (1939) described A. fimbriata var. glabra

and A. fimbriata var. perangusta, but did not

discuss the variability of the species as a whole.

Pedley (1980) noted the variability of the species.

He placed A. fimbriata var. glabra in the

synonymy of A. fimbriata and recognised A.

perangusta, based on A. fimbriata var.

perangusta. He distinguished it from A.

fimbriata, as did White, in being glabrous and

having long narrow phyllodes with the gland

some distance from the base. It was reported as

being restricted to the banks of small streams 25

to 35 kms south and south-east of Brisbane and

on the Burrum River, a little north of

Maryborough. Variation within the species was

not discussed, though a specimen intermediate

between A. fimbriata and A. perangusta was

noted. Both species are commonly planted in

gardens and on roadsides in south-eastern

Queensland. The latter is particularly attractive,

with pendulous branches and dense shiny bright

green foliage. Clearing of native vegetation

within the restricted range of A. perangusta in

the vicinity of Brisbane led to its listing as a

vulnerable species. This, in turn, resulted in

intensive collecting for identification by

consultants for housing developments. Study of

such collections indicates that A. perangusta

should not be maintained as a taxon at any rank

as both individuals with narrow phyllodes and

broad phyllodes occur in the same population.

As in other narrow-phylloded uninerved acacias,

the distance of the gland from the base of the

phyllode varies with the width of the phyllode:

the narrower the phyllode, the greater the

distance. The degree of hairiness varies

independently of the dimensions of the phyllodes.

A. perangusta and A. fimbriata var. glabra are

merely extreme forms of A. fimbriata. The

reduction in rank of A. perangusta is formalised

below.

Accepted for publication 8 September 2004

Acacia fimbriata A. Cunn. ex G. Don, Gen.

Hist. 2: 406 (1832).

Acacia prominens var. fimbriata (A. Cunn.

ex G. Don) Domin, Biblioth. Bot. 89: 256

(1928); Racosperma fimbriatum (A.

Cunn. ex G. Don) Pedley, Austrobaileya

2: 348 (1987). Type: Moreton District:

Brisbane River, Sept. 1828,

A. Cunningham 158 (holo: BM; iso: K)

Acacia prominens var. whiteana Domin, loc.

cit. (1926). Type: Moreton District:

Upper Brisbane River, Aug. 1908,

C.T. White s.n. (holo: PR)

Acacia fimbriata var. glabra C.T. White,

Proc. Roy. Soc. Queensland 50:72 (1939).

Type: Wide Bay District: near

Biggenden, Apr. 1921, W.R. Petrie 18A

(holo: BRI).

Acacia fimbriata var. perangusta C.T. White,

loc. cit. (1939); Acacia perangusta (C.T.

White) Pedley, Austrobaileya 1:287

(1980); Racosperma perangustum (C.T.

White) Pedley, Austrobaileya 2: 353

(1987). syn. nov. Type: Moreton

District: Castra near Brisbane [near

Victoria Point], 7 Aug. 1927, C.T. White

3554 (holo: BRI)

References

Pedley, L. (1980). Acacia fimbriata, in A revision of Acacia

in Queensland. Austrobaileya 1: 235-337 (‘1979’).

White, C.T. (1939). Contributions to the Queensland flora,

No. 6. Proceedings of the Royal Society of

Queensland 50: 66-87.

* Acacia is the name used here to be consistent

with that used in the Queensland Nature

Conservation Act 1992 and the current

recommendation before the IAPT to conserve

Acacia over the correct generic name

Racosperma.

Les Pedley

Queensland Herbarium, EPA, Botanic

Gardens Mt Coot-tha, Mt Coot-tha Road,

Toowong, Queensland 4066, Australia.

Austrobaileya 6(4): 985-986 (2004)

Note

985

Supplement to a synopsis of Racosperma C. Mart. (Leguminosae:

Mimosoideae).

Despite the care taken in compiling the

catalogue of taxa of Racosperma (Pedley 2003),

errors were made. I have not attempted to

correct minor errors, but others are such as to

possibly affect the validity of names of some

taxa. Minor errors are: deviation from the

abbreviations of personal names adopted by

Brummitt & Powell (1992) and lapses in the

gender of generic names and specific epithets.

In one case, the use of a wrong font and

indentation could probably cause some

confusion. Corrections of major errors are

made below. Though the incomplete citation

of the authors of basionyms of new

combinations probably does not make them

invalid, these citations have been completed

where necessary.

Racosperma aemulum (Maslin) Pedley

R. aemulum subsp. muricatum (Maslin)

Pedley §367b

Acacia aemula subsp. muricata Maslin,

Nuytsia 10: 169 (1995).

Racosperma aulacocarpum (A.Cunn. ex

Benth.) Pedley, Austrobaileya 2: 345

(1987). §670

R. aulacocarpum var. fruticosum (C.White)

Pedley, Austrobailey 2:345 (1987) syn.

nov.

Because of the lack of indentation and the font

used, the synonymy was not immediately

apparent.

Racosperma awestonianum (R.S. Cowan &

Maslin) Pedley, comb. nov. §499

Acacia awestoniana R.S. Cowan & Maslin,

Nuytsia 7: 185 (1990).

Note use of incorrect form of specific epithet.

Racosperma baeuerlenii (Maiden & Blakely)

Pedley, Austrobaileya 2: 345 (1987)

§466

Incorrect citation of authors of basionym.

Racosperma carnosulum (Maslin) Pedley,

comb. nov. §423

Acacia carnosula Maslin, Nuytsia 12: 331

(1999)

Incorrect basionym.

Racosperma dealbatum (Link) Pedley

R. dealbatum subsp. subalpinum (Tindale &

Kodela) Pedley, comb. nov. §52b

Acacia dealbata subsp. subalpina Tindale &

Kodela, Telopea 9:319 (2001).

Fictitious basionym.

Racosperma deltoideum (A. Cunn. ex G Don)

Pedley

R. deltoideum subsp. amplum (R.S. Cowan

& Maslin) Pedley §617b

Acacia deltoidea subsp. ampla R.S.Cowan

& Maslin, Nuytsia 7: 206 (1990)

Racosperma drummondii (Lindl.) Pedley

R. drummondii var. elegans (Maslin) Pedley,

comb, et stat. nov. §952d

Acacia drummondii subsp. elegans Maslin,

Nuytsia 1: 468 (1975)

R. drummondii var. majus (Benth.) Pedley,

comb. nov.

Acacia drummondii var. major Benth., FI.

Austral. 2: 419 (1864) §952c

Ranks of the basionyms of the infraspecific taxa

were confused.

Racosperma euthycarpum (J.M. Black)

Pedley

R. euthycarpum var. oblanceolatum (Stephen

H. Wright) Pedley §sub 81

Acacia euthycarpa var. oblanceolata Stephen

H. Wright, Muelleria 16:64 (2002).

Racosperma fauntleroyi (Maiden) Pedley

Acacia fauntleroyi Maiden, J. & Proc. Roy.

Soc. New South Wales 53:194 (1920)

Accepted for publication 7 October 2004

9 8 6

Racosperma gloeotrichum (A.R. Chapm. &

Maslin) Pedley, comb. nov. §708

Acacia gloeotricha A.R. Chapm. & Maslin,

Nuytsia 12: 487 (1999)

Place of publication of basionym omitted.

Racosperma gordonii (Tindale) Pedley, comb,

et stat. nov. §162

Acacia brunioides subsp. gordonii Tindale,

Contrib. New South Wales Natl Herb. 4:

74 (1968).

Incorrect basionym.

Racosperma heterochroa (Maslin) Pedley

R. heterochroa subsp. robertii (Maslin) Pedley

§259b

Acacia heterochroa subsp. robertii Maslin,

Nuytsia 10: 95 (1995).

Racosperma insolitum (E. Prtitzel) Pedley

R. insolitum subsp. efoliatum (Maslin) Pedley

§248c

Acacia insolita subsp. efoliata Maslin,

Nuytsia 12: 362 (1999).

Racosperma kochii (W.V. Fitzg. ex A.J. Ewart

& Jean White) Pedley §406

Acacia kochii W.V. Fitzg. ex A.J. Ewart &

Jean White, Proc. Roy. Soc. Victoria n.

ser. 23: 285 (1911)

An example of author citation at its worst.

Racosperma neurophyllum

R. neurophyllum subsp. erugatum (R.S.

Cowan Maslin) Pedley §885b

Acacia neurophylla subsp. erugata R.S.

Cowan & Maslin, Nuytsia 10: 51(1995).

Racosperma obtusatum (Sieber ex DC.)

Pedley §115

Acacia obtusata Sieber ex DC., Prodr. 2:453

(1825).

Racosperma phlebophyllum (F. Muell. ex

H.B. Williamson) Pedley §895

Acacia phlebophylla F. Muell ex H.B.

Williamson, in A.J. Ewart, FI. Victoria

607 (1931).

Austrobaileya 6(4): 985-986 (2004)

Racosperma seclusum (M.W. McDonald)

Pedley §69 lb

Acacia seclusa M.W. McDonald, Austral.

Syst. Bot. 16: 152. t.9 (2003)

A. tumida var. pubescens Maiden, in A.J.

Ewart & O.B. Davies, FI. N. Terr. 344

(1917).

Racosperma stereophyllum (Meisn.) Pedley,

§852

Acacia stereophylla Meisn., in J.G.C.

Fehm., PI. Preiss. 2: 203 (1848).

Racosperma stipuligerum (F. Muell.) Pedley

§707

R. stipuligerum subsp. glabrifolium (Maiden

& Blakely) Pedley, Austrobaileya 2:356

(1987), syn. nov.

Racosperma tumidum (F. Muell. ex Benth. )

Pedley

R. tumidum var. kulparn (M.W. McDonald)

Pedley, comb. nov. §sub 691

Acacia tumida var. kulparn M.W. McDonald,

Austral Syst. Bot. 16: 160. t.13 (2003).

Racosperma wickhamii (Benth.) Pedley

§756

The note following the species should refer to

Acacia wickhamii subsp. viscidula and A.

wickhamii subsp. parviphyllodinea , not to A.

wickhamii subsp. viscidulum and subsp.

parviphyllodineum.

I am grateful to my colleagues at BRI who

pointed out the mistakes.

References

Brummitt, R.K. & C.E. Powell (ed.) (1992). Authors of plant

names. Royal Botanic Gardens, Kew.

Pedley, L. (2003). A synopsis of Racosperma C. Mart.

(Leguminosae: Mimosoideae). Austrobaileya 6(3):

445-496.

Les Pedley

Queensland Herbarium, EPA, Botanic Gardens Mt Coot-tha, Mt Coot-tha Road,

Toowong, Queensland 4066, Australia.

Austrobaileya 6(4): 987 (2004)

Addendum

987

Map symbols in the following captions on pages

433-436 disappeared during printing

Map 2, Distribution of Croton in Australia. A C. aridus, k C. amhemicus, • C. stigtnatosus.

Map 3. Distribution of Croton in Australia. A C. schuhzii, ■ C C multicaul is subsp. velutinus,

• C. acronychioides, ▼ C. c/yom&rcfewws, C. simulates, & C. stockeri.

Map 4. Distribution of Croton in Australia. A C. byrnesii. ■ C. dockrillii, k C. rarus. • C. minimus.

Map 5. Distribution of Croton in Australia. A C. habrophyilus, k C. capitis-york, • C. densivestitm, ■ C.

V C. mamillatus.

Map 6. Distribution of Croton in Australia, k C. insular is. • C. armstrongii.

Map 7. Distribution of Croton in Australia. • C. caudatus, k C. triacros,

A C. verreauxii, ▼ C. waterhouseae.

Map 8. Distribution of Croton in Australia. A C. mw/fewy/fr subsp. mw/r/caw/w,

★ C. phebalioides.

Map 9. Distribution of Croton in Australia. • C. tomenieilus, A C. mutabilis,

k C. capitatus. ■ C. setigerus, ▼ C. glanduiosus.

988

Referees consulted for Volume 6

Acceptance of papers has depended on the outcome of review by referees. Apart from a few who

did not wish to be listed, those consulted during the past four years are listed below. Several were

consulted on more than one occasion. Sincere thanks are extended to all these people whose

expertise has helped to maintain journal standards

F. Adema

A. Barfod

A.R. Bean

P.E. Berry

E.A. Brown

J.J. Bruhl

J.G. Conran

M.D. Crisp

E. Dean

C. R. Dunlop

M. Duretto

N. Fechner

RE Forster

G. P. Guymer

R. J.F. Henderson

A.E. Holland

W. K. Harris

P. Hoffmann

S. W.L. Jacobs

L.W. Jessup

R.W. Johnson

D. L. Jones

B.J. Lepschi

M. Lock

M.W. McDonald

T.D. Macpharlane

W.J.F. McDonald

M. Nee

A.E. Orchard

R. W. Rogers

A. C. Rozefelds

B. L. Rye

L. Sage

PS. Short

K.M. Stephens

I. Thompson

H.R. Toelken

P. Vorster

N. G. Walsh

Paul G. Wilson

Peter G. Wilson

S. Zona

T. Kellogg

P.G. Kodela

R. de Kok

G. Keighery

Austrobaileya 6 (1-4): 1-1006

Index

989

Acacia 111, 445

abbatiana 441

abrupta 441

acanthoclada 441

subsp. glaucescens 447

acellerata 441

acinacea 441

aciphylla 447

acoma 441

acrionastes 441

aculeatissima 441

aculeiformis 441

acuminata 441, 454

acutata 448

adinophylla 448

adnata 448

adoxa 448

var. subglabra 448

aemula 448

subsp. muricata 448, 985

aestivalis 448

alata 448

var. biglandulosa 448

var. tetrantha 448

alcockii 448

alexandri 448

alpina 448

amandae 448

amblyophylla 449

amentifera 449

arnoena 449

ampliceps 449

amputata 449

amyctica 449

anarthos 449

anasilla 449

anastema 449

anaticeps 449

anceps 449

ancistrocarpa 482

ancistrophylla 449

var. perarcuata 449

andrewsii 450

aneura var. argentea 450

var. conifera 450

v'dr. fuliginea 450

var. intermedia 450

var. macrocarpa 450

var. major 450

var. microcarpa 450

var. pilbarana 450

var. tenuis 450

anfractuosa 450

anomala 450

anthochaera 450

aphanoclada 450

aphylla 450

applanata 450

aprica 450

arafurica 450

araneosa 450

arbiana 450

arcuatilis 451

argutifolia 451

argyrophylla 451

argyrotricha 451

arida 451

aristulata 451

arrecta Maslin 451

ascendens 451

asepala 451

ashbyae 451

asparagoides 451

aspera 451

assimilis 451

subsp. atroviridis 451

ataxiphylla 451

subsp. magna 451

atkinsiana 451

atopa 452

atrox 452

aulacophylla 452

auratiflora 452

aureocrinita 452

auricoma 452

auripila 452

auronitens 452

ausfeldii 452

awestoniana (as awestonii 452), 985

ayersiana 452

balsamea 452

barakulensis 452

barattensis 452

barbinervis 452

subsp. borealis 452

barringtonensis 452

basedowii 453

baueri subsp. gordonii 467

var. aspera 453

baxteri 453

beauverdiana 453

benthamii 453

bidentata 453

bidwillii 181

var. major 181

var .polytricha 181

bifaria 453

biflora 453

binata 453

bivenosa 453

subsp. wayi 473

blakelyi 453

blaxellii 453

blayana 453

boormanii 453

botrydion 453

brachybotrya 453

brachyclada 453

brachyphylla 453

var. recurvata 454

brachypoda 454

bracteolata 454

brockii 454

browniana 454

var. glaucescens 454

brumalis 454

brunioides subsp. gordonii 986

bulgaensis 454

burdekensis 454

burkittii 454

bynoeana 454

Austrobaileya 6(1-4): 1-1006

990

Acacia caerulescens 455

caesariata 455

caesiella 455

calamifolia 455, 464

var. euthycarpa 464

calcarata 455

caleyi 455

calligera 183

camptoclada 455, 482

campylophylla 455

cangaiensis 455

capillaris 455

cardiophylla 455

carens 455

carneorum 455

carnosula (as carnulosa 455), 985

cassicula 456

castanostegia 456

cataractae 456

cavealis 456

cedroides 456

celastrifolia 456

celsa 456

cerastes 456

chalkeri 456

chamaeleon 456

chapmanii 456

subsp. australis 456

chartacea 456

cheelii 456

chrysella 456

chrysocephala 456

chrysochaeta 456

chrysopoda 457

chrysotricha 457

citrinoviridis 457

clandullensis 457

clarksoniana 184

clelandii 457

cliftoniana 457

clivicola 445, 457

clunies-rossiae 457

clydonophora 457

cochlocarpa 457

subsp. velutinosa 457

cognata 457

colei 457

colletioides 457

comans 457

concolorans 457

confusa 457

congesta 457

subsp. wonganensis 458

conniana 458

consanginea 458

consobrina 458

conspersa 458

constablei 458

continua 458

convallium 458

coolgardiensis 458

subsp. ejfusa 458

subsp. latior 458

coriacea subsp. pendens 458

costata 458

costiniana 458

courtii 458

covenyi 458

cowaniana 458

cracentis 458

craspedocarpa 459

crassistipula 459

crassiuscula 459

crassuloides 459

cremiflora 459

crenulata 459

cretacea 459

crispula 459

cummingiana 459

cuneifolia 459

cupularis 473

curvata 459

curvinervia 445,471

cuspidifolia 459

cuthbertsonii 459

subsp. linearis 459

cyclops 445,463

cycnorum var. sedifolia 472

cylindrica 459

cyperophylla var. omearana 459

dacrydioides 459

dallachiana 460

dangarensis 460

daviesii 460

daviesioides 460

daweana 460

dealbata subsp. subalpina (as subdealbata subsp.

subalpina 460) 985

declinata 460

deficiens 460

deflexa 460

delibrata 460

delibrata 183

delicatula 460

delphina 460

deltoidea subsp. ampla (as deltoideum subsp. ampla

460) 985

demissa 460

dempsteri 460

densiflora 460

denticulosa 461

dentifera 461

depressa 461

dermatophylla 461

desertorum 461

var. nudipes 461

desmondii 445,477

diaphana 461

diaphyllodinea 461

dictyoneura 461

didymum 461

dielsii 461

dijformis 461

dilatata 461

diminuta 461

disparrima 461

subsp. calidestris 461

dissona 461

var. indoloria 461

distans 462

disticha 462

ditricha 179, 181

divergens 462

dolichophylla 462

donaldsonii 462

doratoxylon 462

dorotheae 462

dorsenna 462

991

Austrobaileya 6 (1^1): 1-1006

Acacia douglasica 181

drepanocarpa subsp. latifolia 462

drepanophylla 462

drewiana 462

subsp. minor 462

drummondii 462

subsp. elegans 985

subsp. major 462

var. elegans 462

var. major 985

var .parviflora 492

dunnii 462

dura 463

durabilis 463

duriuscula 463

echinula 463

echinulijlora 463

ejfusa 463

eglandulosa 463

elachantha 463

elongata 463

empelioclada 463

endlicheri 454

enervia 463

subsp. explicata 463

enterocarpa 463

epedunculata 463

ephedroides 463

eremaea 463

eremophila 463

var. variabilis 463

ericifolia 464

ericksoniae 464

erinacea 464

erioclada 464

eriopoda 464

errabunda 464

euthycarpa var. oblanceolata (as subsp. oblanceolatum

464) 985

euthyphylla 464

evenulosa 464

excelsa subsp. angusta 464

excentrica 464

exilis 464

exocarpoides 464

extensa 464

fagonioides 464

farinosa 464

farnesiana 179

vdr.famesiana 180

var. guanacastensis 180

fauntleroyi 985

faucium 464

ferocior 465

filamentosa 465

filifolia 465

filipes 465

fimbriata 983

var. glabra 983

var .perangusta 983

fitzgeraldii var. brevior 493

flabellifolia 465

flagelliformis 465

flavipila 465

var. ovalis 465

flocktoniae 465

fodinalis 465

formidabilis 465

forrestiana 465

forsythii 465

fragilis 465

frigescens 465

fulva 465

galeata 466

gardneri 466

gelasina 466

gemina 466

genistifolia 466

genuinae 445

gibbosa 466

gilbertii 466

gilesiana 466

glabrata 429

gladiifonnis 466

glandulicarpa 466

glaucissima 466

glaucocaesia 466

glaucoptera 466

gloeotricha 466, 986

glutinosissima 466

goadbyi 482

gonocarpa 466

gonophylla 466

gracilenta 467

gracilifolia 467

gracillima 467

graniticola 467

grasbyi 467

gregorii 467

grisea 467

guinetii 467

hadrophylla 467

halliana 467

hamersleyensis 467

hamiltoniana 467

harveyi 467

hastulata 467

havilandiorum 467

helicophylla 467

helmsiana 468

hemiteles 468

hendersonii 468

heterochroa 468

subsp. robertii 468,986

heteroclita 468

subsp. valida 468

heteroneura 468

var. petila 468

var .prolixa 468

heterophylla 494

hexaneura 468

hippuroides 468

holosericea var. glabrata 468

hopperiana 468

horridula 468

howittii 469

huegelii 469

hypermeces 469

hystrix 469

subsp. continua 469

idiomorpha 469

imbricata 469

imitans 469

imparilis 469

improcera 469

inaequilatera 469

inaequiloba 469

inamabilis 469

Austrobaileya 6(1-4): 1-1006

992

Acacia incanicarpa 469

inceana 469

subsp. conformis 469

incongesta 469

incrassata 469

incurva 469

ingramii 469

ingrata 469

inophloia 470

inops 470

insolita 470

subsp. efoliata (as efoliolatum 470) 986

subsp. recurva 470

intorta 470

intricata 470

irrorata subsp. vellutinella 470

isoneura 470

subsp. nimia 470

iteaphylla 470

jacksonioides 470

jamesiana 470

jasperensis 470

jennerae 470

jensenii 470

jibberdingensis 470

johannis 470

jonesii 470

julifera 183

subsp. gilbertensis A1 1

juncifolia subsp. serpentinicola 445, 486

jutsonii 468

kalgoorliensis 471

kauaiensis 471

Jfcefleri 471

kempeana 487

kenneallyi 471

kerryana A1 1

kettlewelliae 471

kimberleyensis 471

kingiana 471

foa 471

koaia 471

kocffl 471,986

kybeanensis 471

kydrensis 471

lacertensis 471

lachnophylla 471

lamprocarpa 471

lanceolata All

lanei 472

lanigera var. gracilipes 472

lanuginophylla All

laricina All

var. crassifolia All

lasiocalyx All

lasiocarpa All

var. bracteolata All

var. epacantha 463

lateriticola All

latipes All

subsp. licina All

latzii All

leiocalyx subsp. herveyensis All

leioderma All

leiophylla All

lenticellata 180

lentiginea All

leprosa All

leptalea All

leptoclada 181,473

var .polyphylla 181

leptoneura All

leptopetala All

leptophleba All

leptospermoides All

subsp. obovata All

leptostachya 342

leucolobia All

levata All

ligustrina All

linarioides All

linearifolia A1A

lineolata 474

linophylla 483

lirellata 474

subsp. compressa 474

littorea 474

lobulata 474

loderi 474

longiphyllodinea 474

longispinea A1A

loxophylla 474

lucasii 474

lullfitziorum 474

lunata 474

luteola 474

mabellae 474

macdonnellensis subsp. teretifolia 475

mackeyana 475

macnuttiana 475

maconochieana 475

malloclada 475

manipularis 475

marramamba 475

masliniana 475

mathuataensis 475

matthewii 475

maxwellii 475

megacephala 475

meiantha 475

meisneri 475

menzelii Aid

merinthophora Aid

merrallii 476

merrickiae Aid

microbotrya Aid

microcalyx Aid

microcarp a Aid

microneura 476

midgleyi Aid

mimica 476

var. angusta Aid

minutifolia Aid

minyura Aid

mitchellii Aid

mitodes Aid

moirii Aid

subsp. dasycarpa Aid

subsp. recurvistipula Aid

mollifolia Aid

mooreana Aid

mountfordiae All

mucronata All

var. dependens All

var. longifolia All

multilineata 474

multispicata All

multistipulosa All

Austrobaileya 6 (1^): 1-1006

Acacia mutabilis All

subsp. angustifolia All

subsp. incurva All

subsp. rhynchophylla All

subsp. stipulifera All

nanodealbata All

nelsonii All

nematophylla All

nervosa All

neurocarpa All

neurophylla (as neurophyllum All, 478) 986

subsp. erugata (as erugatum 478) 986

newbeyi 478

nigripilosa 478

subsp. latifolia 478

nilotica 182,494

subsp. indica 182

nitidula 478

nivea 478

nodiflora 478

notabilis 478

nuperrima subsp. cassitera 493

nyssophylla 478

obesa 478

obliquinervia 478

obovata 478

obscura 454

var. preissiana 481

obtecta 478

obtusata (as obtusatum 478) 986

octonervia 478

oldfieldii 478

oligoneura 183

oligoneura 478

olsenii 478

oncinocarpa 478

oncinophylla 479

subsp .patulifolia 479

var .fauntleroyi 465

ophiolithica 479

orbifolia 479

orthotricha 479

ovoidea 492

oxycedrus 479

oxyclada 479

pachyacra 479

pachycarpa 479

pachyphloia 180

subsp. brevipinnula 180

subsp .pachyphloia 180

pachyphylla 479

pachypoda 479

pallida 180, 183

pallidifolia 180, 183

palustris 479

papulosa 479

papyrocarpa 479

paraneura 479

parvipinnula 479

pataczekii 479

patagiata 480

paucijuga 460

paw/a 480

pedina 480

pedleyi 480

pedunculata 180

pellita 480

pelophila 480

pentadenia 480

993

perangusta 983

peregrinalis 480

phaeocalyx 480

pharangites 480

phasmoides 480

phlebopetala 480

var .pubescens 480

phlebophylla 480, 986

pickardii 480

piligera 480

pilligaensis 481

pinguiculosa 481

subsp. teretifolia 481

pinguifolia 481

plagiophylla 184

platyptera 448

plautella 481

plicata 481

poliochroa 481

porcata 481

praelongata 481

praernorsa 481

praetermissa 481

prainii 481

pravissima 481

prismifolia 481

pritzeliana 481

producta 482

profusa 482

proiantha 482

prominens 482

var .fimbriata 983

var. whiteana 983

proxima 482

pruinicarpa 482

psammophila A13

pterocarpa 183

pterocaidon 482

ptychoclada 482

ptychophylla 482

pulchella var. glaberrima 482

var. reflexa 482

pulviniformis 482

puncticulata 482

pusilla 483

pustula 483

pycnantha 483

pycnocephala 483

pygmaea 483

pyrifolia 483

quadrimarginea 483

quadrisulcata 483

quinquenervia 483

quornensis 483

racospermoides 445

recurvata 483

redolens 483

rendlei 483

repanda 483

repens 483

resinimarginea 483

resinistipulea 483

resinosa 483

restiacea 484

retinervis 484

retinodes 484

var. gillii 466

var. uncifolia 484

subsp. clandestina 484

Austrobaileya 6(1-4): 1-1006

994

Acacia retrorsa 484

rhamphophylla 484

rhetinocarpa 484

rhigiophylla 484

rhodophloia 484

richardsii 484

richii 484

ridleyana 484

rigescens 484

rigida 484

rivalis 484

robiniae 484

rossei 484

rostellata 484

rostellifera 485

roycei 485

rubricola 485

rupicola 485

ruscifolia 492

ryaniana 485

sabulosa 485

saliciformis 485

salicina var. minor 473

var. wayi 473

saxatilis 485

scabra 485

scalena 485

scalpelliformis 485

schinoides 485

sciophanes 485

scirpifolia 485

scleroclada 485

sclerophylla 485

var. lissophylla 449

var .pilosa 485

var. teretiuscula 485

sclerosperma 485

subsp. glaucescens 486

scopulorum 486

seclusa (as secluse 486) 986

sedifolia 486

subsp. pulvinata 486

semicircinalis 486

semitrullata 486

sericata 486

sericocarpa 486

sericoflora 486

sericophylla 458

sertiformis 486

sessilis 486

sessilispica 486

setulifera 486

shuttleworthii 486

sibilans 486

sibina 486

sibirica 486, 487

siculiformis 487

signata 487

silvestris 487

simmondsiana 487

simplex 487

simulans 487

singula 487

smeringa 487

solenota 487

sorophylla 487

spathulifolia 487

speckii 487

spenophylla 488

sphacelata 487

subsp. recurva 487

subsp. verticillata 487

sphaerostachya 487

spilleriana 488

spinescens 488

spinosissima 488

spongolitica 488

spooneri 488

squamata 488

startii 488

steedmanii 488

stellaticeps 488

stenoptera 488

stereophylla (as stereophyllum 488) 986

var. cylindrata 488

stigmatophylla 488

stowardii 445, 487

strigosa var. intermedia 454

subcaerulea 489

subdealbata subsp. subalpina 460

suberosa 179

subflexuosa 489

subsp. capillata 489

subg. Heterophyllum 446

subg. Phyllodineae 177

subg. Phyllodineae 446

subgen. Aculeiferum 111

subporosa 489

subracemosa 489

subrigida 489

subsessilis 489

subtemata 489

subtessaragona 489

subtilinervis 489

subulata 489

sulcata 489

var . pianoconvexa 489

var .platyphylla 489

sutherlandii 179

symonii 489

synchronicia 489

tarculensis 489

tayloriana 489

telmica 489

tenuior 490

tenuispica 489

teretifolia 490

tessellata 490

tetanophylla 490

tetragonocarpa 490

tetraneura 490

tetraptera 490

thomsonii 490

tingoorensis 490

tolmerensis 490

tomentosa 429

toodulya 490

torticarpa 490

torulosa 183

trachycarpa 490

trachyphloia 490

tratmaniana 490

trigonophylla 490

trinalis 490

trinervata 490

trineura 491

triptycha 491

triquetra 491

995

Austrobaileya 6 (1^): 1-1006

Acacia truculenta 491

trullifonnis 491

tuberculata 491

tumida 491

var. extenta 491

van kulparn (as kalparn 491) 986

var. pilbaremis 491

var. pubescens 486,986

turbata 180

tysonii 491

ulicina 491

uliginosa 491

uncinella 491

undoolyana 491

undosa 491

undulifolia 491

unguiculata 492

unifissilis 492

urophylla 492

valida 183

varia 492

var. ajfinis 462

var. crassinervis 492

vassalii 492

veronicae 492

verricula 492

verticillata var. cephalantha 492

vestita 492

victoriae subsp. an'da 492

vincentii 445, 460

viscifolia 492

v/ffafa 492

volubilis 492

wanyu 492

warramaba 493

wattsiana 493

websteri 493

wetarensis 493

whanii 472

whibleyana 493

wickhamii

subsp. parviphyllodinea 455,493,986

subsp. viscidula 455,493,986

wilcoxii 493

willielmiana 493

willdenowiana 493

williamsiana 493

williamsonii 493

wilsonii 493

wiseana 493

x grayana 467

xanthina 493

xanthocarpa 493

xerophila 493

xiphophylla 494

yirrkallensis 494

yorkrakinensis 494

subsp. acrita 494

zatrichota 494

reclinata 183

stipulosa 183

Acalyphoideae 273, 274

Acerosae 189

laurifolia 162

myrsinoides 162

obovata 163

Adriana 350

acerifolia 429

var. genuina 429

quadripartita 429

urticoides 432

var. hookeri 432

var. urticoides 432

Agrostoides 571

Albizzia sutherlandii 179

Alysicarpus aurantiacus 109

brownii 110

bupleurifolius 110

heyneanus 114

var. ludens 114

longifolius 110

ludens 114

major 111

muelleri 112

var . clementii 112

ovalifolius 113

rugosus 112

subsp. reticulatus 113

var. longe-exsertus 112

var. ludens 114

schomburgkii 114

suffruticosus 114

vaginalis 115

Amorphospermum whitei 161

Ampherephis mutica 977

Androcera rostrata 803

Aphania senegalensis 559

Arctium lappa 146

minus 146

Arenga australasica 970

Argyreia nervosa 631

queenslandica 632

soutteri 631

tiliaefolia 634

Athertonia 129

Backhousia oligantha 533

sp. (Didcot P.I.Forster PIF12617) 533

sp. (Stony Creek P.I.Forster 37B) 533

Bean, A.R. (2001). Anew species of Lissanthe R.Br.

(Epacridaceae) from Queensland 99

Bean, A.R. (2001). Eucalyptus broviniensis (Myrtaceae), a new

critically endangered species from south-eastern Queensland.

117

Bean, A.R. (2001). Pappus morphology and terminology in

Australian and New Zealand thistles (Asteraceae, tribe

Cardueae) 139

Bean, A.R. (2002). New prostrate species in Solanum subg.

Leptostemonum (Dunal) Bitter (Solanaceae) from eastern

Australia 247

Bean, A.R. (2002). Two new combinations in Corymbia K.D.Hill

& L.A.S.Johnson (Myrtaceae) 345

Bean A.R. (2003). A new species of Mimulus L.

(Scrophulariaceae) from Queensland, Australia 549

Bean A.R. (2003). New combinations in Lycianthes (Dunal)

Hassl. (Solanaceae) for New Guinea and Australia 567

Bean A.R. (2003). The puzzle of Eucalyptus hemilampra

F.Muell. (Myrtaceae) 563

Bean, A.R. (2003). Backhousia oligantha (Myrtaceae), a new

species from Queensland 533

Bean, A.R. (2004). Anew combination in Centratherum Cass.

(Asteraceae) 977

Bean, A.R. (2004). New species of Cryptandra Sm. and

Stenanthemum Reissek. (Rhamnaceae) from northern

Australia 917

Bean, A.R. (2004). The taxonomy and ecology of Solanum subg.

Leptostemonum (Dunal) Bitter (Solanaceae) in Queensland

and far north-eastern New South Wales, Australia 639

Austrobaileya 6(1-4): 1-1006

996

Bean, A.R. and Henwood M.J. (2003). Six new species of

Hydrocotyle L. (Apiaceae) from Queensland 537

Beccariella queenslandica 161

Bertya andrewsii, 189

astrotricha 193

brownii 193

calycina 195

cunninghamii 197

subsp. cunninghamii 245

subsp. cunninghamii 198

subsp. pubiramula 198

subsp. rupicola 199

cupressoidea 240

dimerostigma 199

var. cupressoidea 240

var. dimerostigma 199

var. genuina 200

ernestiana 200

findlayi 203

glabrescens 223

glandulosa 204

grampiana 205

granitica 206

gummifera 208

var. genuina 210

var. gummifera 208

var .psiloclada 241

ingramii 211

lapicola 212

subsp. brevifolia 213

subsp. lapicola 213

linearifolia 215

mitchellii 220

var. genuina 220

var. mitchellii 220

var. vestita 238

mollissima 216

neglect a 210

oblonga 218

oblongifolia 227

oleifolia 220

var. glabrescens 223

opponens 221

oppositifolia 221

pedicellata 223

pinifolia 225

polymorpha 220

forma genuina 220

forma mitchelliana 208

forma rosmarinifolia 233

polystigma 226

pomaderroides 221

var. angustifolia 221

var. pomaderroides 221

psiloclada 241

quadrisepala 241

recurvata 228

riparia 230

rosmarinifolia 233

rotundifolia 234

sect. Bertya 188,189

sect. Euryphylla 189

sect. Stenophylla 189

sharpeana 235

sp. (Amiens L.Pedley 1488) 229

sp. (Beeron Holding P.I.Forster+ PIF5753) 206

sp. (Helidon Hills GLeiper AQ457013) 213

subsect. Acerosae 189

subsect. Recurvae 189

tasmanica 236

subsp. tasmanica 237

subsp. vestita 238

virgata 240

Bertyinae, 187

Beyeria 350

virgata 240

viscosa 429

Caelospermum dasylobum 911

paniculatum var. paniculatum 911

var . syncarpum 911

Caletia divaricatissima var. divaricatissima 502

var. genuina 502

var. orbicularis 507

linearis 508

orientalis 508

var. orbicularis 507

var. orientalis 508

ovalifolia 509

sect. Microcaletia 499

Calycanthaceae 553

Calycanthus australiensis 553

Canthium 820

attenuatum 857, 869, 871

barbatum 41, 47

beecheyi 833

brevipes 62

buxifolium 843

buxifolium forma (Brigooda P.I.Forster PIF5657) 844

buxifolium var. (Mt Alford L.H.Bird AQ408941) 843

caudatum 41

coprosmoides 46

costatum 44

cymigerum 889

cymosum 847

didymum 847

graciliflorum 827

lamprophyllum 844

lineare 829

longiflorum 41

lucidum 833, 845

odoratum 833

forma (Texas P.I.Forster PIF14257) 841

subsp. (Didcot P.I.Forster PIF14127) 837

oleifolium 867, 871

var. pedunculatum 841

sessilifolium 41

sp. (Altanmoui Range D.GFell+DGF2702) 829

sp. (Berrigurra Station E.R.Anderson 2829) 876

sp. (Charters Towers T. Stuart TWR116) 868

sp. (Cooroy S.T.Blake+ 15507) 52

sp. (Copper-Lode Falls C.H.Gittins 2211) 54

sp. (Dixie Station S.T.Garnett 1545) 856

sp. (DuaringaN.H.Speck 1819) 878

sp. (Friday Island L.J.Brass 18158) 850

sp. (Headingly Station R.A.Perry 848) 855

sp. (Herberton Range S.F.Kajewski 1377) 848

sp. (Kuranda GSankowsky+ 680) 58

sp. (Larcom Gully N.Gibson 1057) 874

sp. (Lizard Island R.L.Specht+LI181) 55

sp.(Mt Alford L.H.Bird AQ408941) 843

sp. (Thornton Peak H.Flecker NQNC7110) 832

sp. (Thursday Island E.Cowley 10) 856

sp. (Weipa, A.Morton AM1773) 827

sp. (Wenlock River, J.R.Clarkson 8418+) 827

sp. (Whitfield Range B.P.Hyland 1020) 849

sp. 1 844

997

Austrobaileya 6 (1^1): 1-1006

Canthium sp. A, B.L. Koch in J.R. Wheeler 869

suaveolens 829

suborbiculare 889

valetonianum 41

Cardueae 139, 140

Carduus 149

nutans 144

pycnocephalus 144

tenuiflorus 144

thoermeri 144

Carthamus 140, 149

dentatus 146

lanatus 146

leucocaulos 147

tinctorius 147

Catalepidia 129

Cenchrus ciliaris 653

Centaurea 140, 149

calcitrapa 147

jacea 147

melitensis 147

solstitialis 148

Centratherum 977

australianum 977

muticum 977

punctatum 977

subsp. australianum 977

Chardinia 148

Chorizotheca 515, 516

micrantheoides 527

Chrysanthemoides monilifera 129, 130

Chrysophyllum myrsinodendron 163

Chrysostemon 515

virgatus 526

Cirsium 143, 149

arvense 144

var. arvense 149

brevistylum 145

palustre 145

vulgare 145

Clifford H. Trevor and Dettmann, Mary E. (2001). Drupe - a term

in search of a definition. 127

Cliffordiochloa parvispiculata 561

Cnicus benedictus 148

Coelospermum 130

Cojfea arabica 904

liberica 905

var. liberica 905

odorata 833

Coffeeae 903

Convolvulus 6

acaulis 26

adscendens 26

angustissimus 23

subsp. angustissimus 26

subsp .fililobus 30

subsp. omnigracilis 27

subsp. peninsularum 31

arvensis 8

clementii 35

var. biflorus 35

crispifolius 15

erubescens 20

var. albus 23

var. angustissimus 23

var. dilatatus 20

vax. fililobus 30

ey reanus 19

geniculatus 23

graminetinus 12

huegelii 14

microsepalus 10

multicaulis 1

preissii 14

recurvatus 32

subsp. nullarborensis 33

subsp. recurvatus 33

remotus 14

sepium 1

subpinnatifidus 26

tedmoorei 37

tiliaefolius 634

tiliifolia 631

tridentatus 1

turpethum 1

umbellatus 1

wimmerensis 22

Corchorus 581

allenii 626

aulacocarpus 584

carnarvonensis 585

congener 586

crassifolius 588

crozophorifolius 588

elachocarpus 590

elderi 591

incanus 592

subsp. incanus 593

subsp. lithophilus 594

interstans 587

laniflorus 596

lasiocarpus 598

subsp. lasiocarpus 599

subsp. parvus 600

leptocarpus 601

lithophilus 594

longipes 626

mitchellensis 602

obclavatus 603

pachyphyllus 626

parviflorus 606

var. gracilescens 606

var. ovatus 606

puberulus 607

pumilio 609

queenslandica 632

rostrisepalus 616

rothii 626

saxicola ms 594

sericeus 611

subsp. densiflorus 612

subsp. sericeus 611

sidoides 612

subsp. rostrisepalus 616

subsp. sidoides 614

subsp. vermicularis 615

sp. Burrup (G.Craig 235) 623

subargentus 616

sublatus 619

tectus 621

tiliifolia 632

tomentellus 622

vermicularis 615

walcottii 623

Corymbia arnhemensis subsp. monticola 345

hylandii 345

subsp. peninsularis 345

serendipita 345

Austrobaileya 6(1-4): 1-1006

998

Croton stockeri 345

subsp. peninsularis 345

subsp. stockeri 345

Corypha decora 979

Crotalaria 293

sturtii 305

acicularis 297

aridicola 320

subsp. aridicola 320

subsp. densifolia 322

subsp. glabrata 320

benthamiana 310

brevis 302

erassipes 312

crispata 299

cunninghamii 304

subsp. cunninghamii 305

subsp. sturtii 305

var. trifoliolata 305

dissitiflora 308

subsp. dissitiflora 308

subsp. benthamiana 310

subsp. rugosa 308

var. grandiflora 310

var. rugosa 308

exserta 315

linifolia 313

var. angustifolia 313

lunulata 300

medicaginea 315

var. angustata 317

var. australiensis 316

var. herniarioides 319

var. linearis 318

var. luxurians 319

var. medicaginea 316

var .neglecta 316

melanocarpa 304

membranacea 299

montana 313

var. angustifolia 313

var. exserta 315

var. montana 313

mysorensis 299

neglecta 316

novae-hollandiae 311

subsp. crassipes 312

subsp. lasiophylla 312

subsp. novae-hollandiae 311

ramosissima 301

sect. Calycinae 294

sect. Chrysocalycinae 294

sect. Crotalaria 294

sect. Dispermae 294

sect. Grandiflorae 294

sect. Hedriocarpae 294

sturtii 305

trifoliastrum 315

willdenowiana 323

Croton 349, 354

acronychioides 357

ajfinis 357

argyratus 428

aridus 360

armstrongii 362

arnhemicus 363

var. typicus 363

var. urenifolius 363

aromaticus 355

bonplandianus 352

brachypus 366

byrnesii 368

capitatus 370

capitis-york 371

var .pilosus 371

caudatus 372

choristadenius 374

coccymelophyllus 428, 429

densivestitus 378

dockrillii 380

glandulosus 382

habrophyllus 384

insularis 386

magneticus 389

maidenii 403

mamillatus 391

microtiglium 401, 403, 426

minimus 392

multicaulis 396

subsp. multicaulis 397

subsp. velutinus 397

mutabilis 399

opponens 245, 428

phebalioides 403

var. acuminatus 403

var. hirsuta 414

var. hispida 428

var. stigmatosus 414

var. typicus 403

philombros 374

prunifolius 428, 429

pubens 378

pusilliflorus 374

quadripartitus 350, 428

rams 406

rosmarinifolius 233,428

sarcopetalus 352

schultzii 408

semunculus 374

setigerus 410

simulans 412

sp. (Barkly Downs S.L.Everist 3379) 360

sp. (Mt Mulligan H.Flecker NQNC6457) 392

sp. (Myall Creek P.I.Forster PIF14368) 396

sp. (Possum Scrub P.I.Forster PIF14376) 399

sp. (WatsonRiver J.R.Clarkson4061B) 406

stigmatosus 414

var. eurybioides 428

stockeri 417

storckii 428

suberosus 352

tiglium 354, 355

tomentellus 419

triacros 421

urticoides 429

verreauxii 423

var. genuinus 423

var. longifolius 424

viscosus 429

wassi-kussae var. stockeri 417

waterhouseae 426

Crotonopsis 349

Cryptandra 917

amara var. (Mt Mulligan J.R. Clarkson 5865) 919

var .amara 919

vai.floribunda 921

var. longiflora 919

armata 922

999

Austrobaileya 6 1-1006

Cryptandra ciliata 928

debilis 919

filiformis 933

gemmata 928

lanosiflora 926

longistaminea 924

orbicularis 922

pogonoloba 930

propinqua 926

rigida 927

scortechinii 935

sp. (Gurulmundi GW. Althofer 8418) 928

sp. (Isla Gorge P. Sharpe 627) 922

sp. (Mt Mulligan J.R. Clarkson 5949) 935

sp. (NgungunL.S. Smith 13973) 927

sp. (ThulimbahC. Schindler 6) 919

sp. 1 928

sp. 2 935

sp. 3 919

Cupaniopsis cooperorum 267

Cyathodes, 99

Cycadaceae, 153

Cycadothrips chadwickii 73, 83, 86, 89, 92

Cycas cairnsiana 153

courts iana 153

cupida 153

desolata 153

ophiolitica 153

platyphylla 153

series Cairnsianosae 153

Cyclophyllum brevipes 62

coprosmoides 46

var. coprosmoides 48

var. spathulatum 48

costatum 44

deplanchei 42

longipetalum 52

maritimum 55

multiflorum 58

protractum 54

rostellatum 49

schultzii 60

forma angustifolium 62

forma schultzii 61

Cynara 143

Cynara cardunculus 145

subsp. flavescens 149

Decarinium glandulosum 382

Denhamia obscura 162

Desmodiastrum 107

Desmodiopsis 107

Desmodium campylocaulon 107

Desmoncus minor 969

prestoei 969

Dichanthium sericeum 13

Dichotomous key to the Queensland species of Solanum subg.

Leptostemonum 659

Dictyomenia pectinella 175

Dowe, J. L. & Barfod, A. S. (2001). New species of Livistona R.Br.

(Arecaceae) from north Queensland and New Guinea 165

Dowe, John Leslie (2004). Nomenclatural changes for two

Australian species of Livistona R.Br. (Arecaceae) 979

Dowe, John Leslie (2004). Taxonomic notes on palms

(Arecaceae) in catalogues of the Brisbane Botanic Garden,

Australia, of 1875 and 1885 967

Elaeocarpus 127, 129,130

grandis 129

spackmaniorum 129

Enantiocladia robinsonii 175

Encephalartos douglasii 74

miquelii 84

spiralis var. major 93

Eremocarpus 412

setigerus 410

Eucalyptus bancroftii 119

broviniensis 117

hallii 117

hemilampra 563, 564

major 119

resinifera 564

subsp. hemilampra 564

var. grandiflora 563

var. hemilampra 564

sect. Liberivalvae 119

ser .Albae 119

ser. Connexentes 119

ser. Subexsertae 119

serendipita 345

propinqua 119

Eupomatia barbata 333, 335

bennettii 335

laurina 335

Farnesia odora 180

Forster, Paul I. (2001). Cycas cupida (Cycadaceae), anew

species from central Queensland 15 3

Forster, PI. (2001). Hydnophytum ferrugineum (Rubiaceae:

Hydnophytinae), a new species of ant-plant from Cape York

Peninsula, Queensland 103

Forster, PI. (2002). Cupaniopsis cooperorum (Sapindaceae), a

new species from the Wet Tropics, Queensland 267

Forster, Paul I. (2001). Phyllanthera takeuchiana (Apocynaceae,

Periplocoideae), a new species from Papua New Guinea 329

Forster, PI. (2004). The tribe Coffeeae DC. (Rubiaceae:

Ixoroideae) in Australia 903

Forster, P.I., Binns, D. & Robertson, G. (2004). Rediscovery of

Tylophora linearis P.I.Forst. (Apocynaceae:

Asclepiadoideae) from New South Wales, with revision of its

conservation status to vulnerable 941

Forster, Paul I. & Jenny Holmes (2003). Lepisanthes

senegalensis (Juss. ex Poir.) Leenh. (Sapindaceae), a new

generic and specific record for Queensland 559

Forster, Paul I. (2003). A taxonomic revision of Croton L.

(Euphorbiaceae) in Australia 349

Forster, Paul I. (2003). Jasminum longipetalum King & Gamble

(Oleaceae), and its occurrence in Queensland, Australia 557

Forster, Paul I. (2003). Phebalium distans P.I.Forst. (Rutaceae), a

new and endangered species from south-eastern Queensland,

and reinstatment of P. longifolium S.T. Blake 437

Glossocardia orthochaeta 341

Glossogyne orthochaeta 341

Gonocarpus chinensis 964

effusus 964

elatus 964

hirtus 962,964

humilis 963

longifolius 964

oreophilus 964

tetragynus 964

teucrioides 963

Austrobaileya 6(1-4): 1-1006

1000

Goodenia expansa 259

atriplexifolia 257

debilis 256

expansa 258

geniculata 260

rosulata 260

sect. Caeruleae 254

sp. (Cuddapan Station K.A.Williams 88038) 259

sp. (Stonehenge J.MilsonJM 1426) 257

sp. (Welcome Creek A.R.Bean 1936) 256

sp. (Yarrowmere R.J.Henderson+ H2844) 254

splendida 254

subgen. Monochila 253

subsect. Borealis 257

Gynochtodes 891

sessilis 891

sp. (HinchinbrookIsland A.S.Thorsborne+337) 891

Halford, D. (2002). Oldenlandia intonsa (Rubiaceae), a new

species from the Northern Territory 325

Halford, D.A. (2002). Indagatorfordii, anew genus and species

of the Sterculiaceae from northern Queensland, Australia

337

Halford, D.A. (2004). Anew species of Gynochtodes Blume

(Rubiaceae) from north-east Queensland 891

Halford, D.A. (2004). Notes on Tiliaceae in Australia, 4. A

revision of the stellate-haired species of the genus

CorchorusL. 581

Halford, D.& Fensham, R.J. (2001). Anew species of

Myriophyllum L. (Haloragaceae) from artesian springs in

central Queensland 133

Halford, D.A. & Ford, A.J. (2004). Caelospermum dasylobum

(Rubiaceae), a new species from north-east Queensland 911

Halford, D.A. & Ford, A.J. (2004). Two new species of Morinda

L. (Rubiaceae) from north-east Queensland 895

Halford, D.A. & Henderson, R.J.F. (2002). Studies in

Euphorbiaceae A.L.Juss., sens. lat. 3. Arevision of Bertya

Planch. (Ricinocarpeae Mull.Arg., Bertyinae Mull.Arg.) 187

Halford, D.A. & Henderson, R.J.F. (2002). Studies in

Euphorbiaceae A.L.Juss., sens. lat. 4. Arevision of

Monotaxis Brongn. (Acalyphoideae Ascherson, Ampereae

Miill.Arg.) 273

Halford, David A. and Henderson R.J.F. (2003). Studies in

Euphorbiaceae A.L.Juss. sens. lat. 5. Arevision of

Pseudanthus Sieber ex Spreng. and Stachystemon Planch.

(Oldfieldioideae Kohler & Webster, Caletieae Mull. Arg.)

497

Harris, W.K. (2004). Notelaea ipsviciensis (Oleaceae), a new

species from south-east Queensland 973

Hedysarum bupleurifolium 110

ovalifolium 113

Helmiopsideae 337

Hemisteptia lyrata 144

Henderson, R.J.F., Wilson, S.M. and Kraft, GT. (2001).

Kentrophora S.M.Wilson & Kraft, a new name for an algal

genus in tribe Amansieae (Rhodomelaceae, Rhodophyceae)

Herber, B.E. (2001). Oreodendron C.T. White reduced to

Phaleria Jack (Thymelaeaceae, Thymelaeoideae) 95

Hicksbeachia pinnatifolia 128

Hippocrepandra 275

ericoides 288

gracilis 284

lurida 284

neesiana 284

Holland, A.E. (2002), A review of Crotalaria L. (Fabaceae:

Crotalarieae) in Australia 293

Holland, A.E. & Boyle T.P. (2002). Four new species of

Goodenia Smith (Goodeniaceae) from Queensland 253

Hydnophytinae 103

Hydnophytum ferrugineum 103

formicarium 103

moseleyanum 103

papuanum 103

Hydrocotyle 537

digitata 543

dipleura 539

miranda 539

oraria 544

paludosa 545

sp. (Strathmay J.R.Clarkson 3498A) 541

tumida 541

Icmadophila splachnirima 951

Idiospermiodeae 553

Idiospermum australiense 553

Indagator 337

fordii 337

Ipomoea grandiflora 634

Ixora orophila 832

Jacquemontia 1

Jasminum 557

dolichopetalum 557

longipetalum 557

pseudoanastomosans 557

sp. (Jardine River, L.J.Brass 18869) 557

turneri 557

Jessup, Laurence W.(2001). New combinations and anew name in

Australian Sapotaceae 161

Jessup, L.W. (2002). Anew species of Eupomatia R.Br.

(Eupomatiaceae) from Queensland 333

Johnson, R.W. (2001). A taxonomic revision of Convolvulus L.

(Convolvulaceae) in Australia 1

Johnson, R.W. (2004). Stictocardia Hallier f. (Convolvulaceae) in

Queensland 631

Jones, David L., Forster, Paul I., Sharma Ish K. (2001). Revision

of the Macrozamia mic/uelii (F.Muell.) A.DC. (Zamiaceae

section Macrozamia) group 67

Jones, D.L. & Dowe, J.L. (2001). Proiphys infundibularis

(Amaryllidaceae), a new species from the Townsville region

of Queensland 121

Jubaea chilensis 969

speciosa 969

spectabilis 969

Julocroton 349

Kantvilas, G. (2004). New Australian species in the lichen genus

Siphula Fr. 949

Keetia 820

Kentrophora 175

natalensis 175

pectinella 175

Key to Australian and New Zealand thistle species, based on

pappus and achenes149

Key to Australian species of Phaleria 96

Key to Australian species of Proiphys 124

Key to Australian subspecies of Psydrax odorata and the extra-

Australian P. odorata subsp. odorata 835

Key to Centratherum species in Australia 977

Key to informal taxonomic groups of Solanum subg. Leptostemon

658

Key to native and naturalised species of Alysicarpus 108

Key to native and naturalised species of Convolvulus in Australia 6

1001

Austrobaileya 6 (1^1): 1-1006

Key to prostrate rainforest Solarium taxa in eastern Australia 248

Key to Pseudanthus and Stachystemon 498

Key to Queensland species of Mimulus 549

Key to Solarium subg. Leptostemonum using characters

applicable to live material 669

Key to species of Acacia subgen. Acacia 111

Key to species of Bertya 190

Key to species of Crotalaria in Australia 294

Key to species of Cyclophyllum in Australia 43

Key to species of Monotcucis 275

Key to species of Phebalium found in Queensland 442

Key to species of Pseudanthus 500

Key to species of Psydrax in Australia 820

Key to species of Stachystemon 516

Key to species of the Macrozamia miquelii group 69

Key to stellate-haired Corchorus in Australia 582

Key to the Australian species of Croton 355

Key to the Australian species of Cupaniopsis 270

Key to the Australian species of Hydnophytinae 104

Key to the Queensland species of Cryptandra and Stachystemon

918

Key to the Queensland species of Hydrocotyle (Apiaceae) 538

Key to the Queensland species of Solanum subg. Leptostemonum

659

Key to the sections of Crotalaria in Australia 294

Key to the species of Cycas series Cairnsianosae 156

Key to the species of Goodenia in Queensland 261

Kuetzingia natalensis 175

Labichea nitida 342

Lepiderema sp. (Topaz P.I. Forster+ PIF15478) 268

Lepisanthes 559

senegalensis 559

Leucocarpum obscurum 162

Leucopogon 99, 127

cuspidatus 101

pedicellatus 99, 101

pluriloculatus 99

Lissanthe brevistyla 99

Livistona 919

australis 979

benthamii 165

concinna 166

decipiens 919

var. polyantha 919

decora 919

drudei 165

inermis 919

mariae 980

subsp. occidentalis 980

muelleri 167

nasmophila 980

pleiospermus 99

strigosa subsp. subulata 99, 101

surra 169

tothur 171

Lycianthes 567

belensis 567

bitteriana 567

dendropilosa 567

multifolia 567

peranomala 567

pustulata 568

rostellata 568

shanesii 568

umbonata 568

vitiensis 568

wollastonii 568

Macrozamia cardiacensis 1 1

communis 67

cranei 61

cylindrica 80

douglasii 74

implicatum 133, 134

longispina 19

macdonnelli 67

mackenzii 92

macleayi 80

miquelii 84

moorei 61

mountperriensis 87

parazamia 61

riedlei 61

serpentina 90

sp. (Marlborough P.I.Forster+PIF12269A) 90

spiralis var. cylindrica 80

tridentata f. milkaui 84

f. oblongifolia 84

subsp. cylindrica 80

subsp. mountperriensis 87

var. douglasii 74

var. mackenzii 92

var. miquelii 84

var. oblongifolia 84

viridis 61

johnsonii 61

var. oblongifolia 84

Mallotus surculosus 351

Mantisalca salmantica 147

Marsdenia viridiflora 941

Megathyrsus 571,572

infestus 573

maximus 572

var. coloratus 573

var . pubiglumis 572

Melaleuca uncinata 510

viridiflora 399

Micrantheum tatei 506

Microcaletia 498

Mimosa 445

binervia 453

cochlearis 457

dodonaeifolia 462

farnesiana 180

heterophylla 468

hispidula 468

linifolia 474

longifolia 474

myrtifolia All

nigricans 478

paradoxa 479

pubescens 482

stricta 488

suaveolens 489

terminalis 490

verticillata 492

Mimulus 549

aquatilis 550

Mischocodon 559

Monotaxis 275

bracteata 284

cuneifolia 218

ericoides 288

gracilis var. genuina 284

var. gracilis 284

var. virgata 284

1002

Monotaxis grandiflora 287

var. grandiflora 288

var. minor 288

var. obtusifolia 288

var. typica 288

linifolia 276

var. cuneata 276

var. genuina 276

var. linifolia 276

var. occidentalis 278

var. tridentata 276

lurida 284

luteiflora 280

macrophylla 279

megacarpa 284

neesiana 284

occidentalis 278

oldfieldii 284

289

sect. Eumonotaxis 276

sect. Hippocrepandra 284

sect. Linidion 276

sect. Monotaxis 276

sp. (RavensthorpeM.A.Burgman 2154) 289

stowardii 284

tenuis 273

tridentata 276

Morinda 895

ammitia 898

podistra 895

sp. (Altanmoui Range B.P.Hyland 6323) 898

sp. (Mt Lewis L.W.Jessup+ GJM228) 895

Myriophyllum amphibium 134

artesium 133, 134

austropygmaeum 136

lophatum 136

pedunculatum 134

Myrmecodia beccarii 104

platytyrea 104

tuberosa 104

Myrmephytum 103

Nesogordonia 337

Nettoa crozophorifolius 590

Niemeyerawhitei 161

Normanbya muelleri 970

normanbyi 970

Notelaea ipsviciensis 973

sp. (Bundamba L.H. Bird AQ442082) 973

Nycterium rostratum 803

Oldenlandia 325

intonsa 325

thysanota 326

Onopordum acanthium 145

acaulon 146

illyricum 146

tauricum 146

Orchard, A.E. (2004). Gonocarpus hirtus Orchard

(Haloragaceae), new from south-eastern Queensland and

north-eastern New South Wales 961

Oreodendron biflorum 95, 96

Oxydectes arnhemicus 363

insularis 386

phebalioides 403

stigmatosus 414

tomentella 419

Austrobaileya 6(1-4): 1-1006

verreauxii (as verrauxii) 423

Panicum 561, 571

infestum 571, 573

laxum 561

maximum 571,572,653

var. trichoglume 572

var. coloratum 573

var. plubiglume 572

sect. Maxima 572

sect. Maximae 572

spongiosum 572

subgenus Megathyrsus 571,572

transvenulosum 572

trichocladum 572

unranked group Maxima 572

Paracoffea brassii 903, 905

Parthenium hysterophorus 653

Peddiea 95

Pedley, Les (2001). Alysicarpus (Leguminosae: Desmodieae) in

Australia. 107

Pedley, L. (2002). A conspecutus of Acacia subg. Acacia in

Australia. 177

Pedley, L. (2003). A synopsis of Racosperma C.Mart.

(Leguminosae: Mimosoideae) 445

Pedley, Les (2004). Reduction of Acacia perangusta to the

synonymy of A.fimbriata 983

Pedley, Les (2004). Supplement to a synopsis of Racosperma

C.Mart. (Leguminosae: Mimosoideae) 985

Persoonia 127, 130

Petalostigma 129

Phaleria biflora 96

capitata 96

chermsideana 96

clerodendron 96

disperma 96

octandra 96

Phebalium 437

distans 438

longifolium 441

squamulosum subsp. squamulosum 438

Phyllanthera takeuchiana 329

Phyllanthus tatei 506

Phyllodineae 445

Picnomon acarna 144

Pilinophytum capitatum 370

Pinanga smithi 969

Planchonella cotinifolia 161

var .pubescens 162

laurifolia 161

macrocarpa 163

myrsinoides 162

obovoidea 163

pohlmaniana var. asterocarpon 163

queenslandica 161

Plectronia barbata 47

coprosmoides var. spathulata 48

cymigera 889

diococca 847

hookeriana 833

odorata var. reticulata 857

schultzii 62

suborbicularis 889

Plectrophora natalensis 175

pectinella 175

Pleiogynium timorense 128, 129, 130

Pollock, A.B. (2002). Rediscovery of Glossocardia orthochaeta

(F.Muell.)Veldk. (Asteraceae) from north-east Queensland 341

1003

Austrobaileya 6 (1^1): 1-1006

Popanax farnesiana 180

Pouteria asterocarpon 163

cotinifolia 163

var. pubescens 162

howeana 163

laurifolia 161

macrocarpa 163

myrsinifolia 162

myrsinodendron 163

myrsinoides 162

subsp. reticulata 163

obovata 163

obovoidea 163

pearsoniorum 163

queenslandica 161

richardii 161

Proiphys alba 122

amboinensis 122

cunninghamii 122

infundibularis 121, 122

Pseudanthus 497, 499

axillaris 517

ballingalliae 500

brachyphyllus 518

chryseus 523

divaricatissimus 502

var. divaricatissimus 502

var. genuinus 502

var. orbicularis 507

intricatus 519

ligulatus 503

subsp. ligulatus 504

subsp. volcanicus 506

micranthus 506

nematophorus 522

nitidus 527

occidentalis 526

orbicularis 507

orientalis 508

ovalifolius 509

pauciflorus 511

subsp. arenicola 512

subsp. pauciflorus 512

pimeleoides 514

polyandrus 523

sect. Caletiopsis 515, 516

sect. Chrysostemon 515,516

sect. Eupseudanthus 499

sect. Microcaletia 499

sect. Pseudanthus 499

sp. (Salvator Rosa NP M.E.Ballingall MEB450) 500

sp. (Tylerville P.I.Forster+ PIF11510) 512

sp. (Tylerville P.I.Forster+PIFl 1510) subsp. (Blackdown

Tableland R.J.Henderson H722) 512

tasmanicus 515

vermicularis 524

virgatus 526

Psilanthus brassii 905

ebracteolatus 903

Psydrax 817, 820

ammophila 854

attenuata 857

forma megalantha 862

forma myrmecophila 862

group 825

var. attenuata 860

var. myrmecophila 861

var. tenella 863

banksii 850

cymigera 889

diococcos 819

forsteri 878

graciliflora 827

johnsonii 876

lamprophylla 844

forma lamprophylla 846

fomia latissimu 847

latifolia 851

group 825

laxiflorens 848

lepida 864

longipes 874

mabesae 907

montigena 832

odorata 833

forma australiana 839

forma buxifolia 843

forma foveolata 840

forma foveolata S 840

forma parviflorifra 843

forma subn itida 841

group 824

subsp. arnhemica 837

subsp. australiana 837

subsp. buxifolia 842

subsp. odorata 835

oleifolia 871

oleifolia group 826

pallida 856

paludosa 826

pendulina 869

reticulata 857

rigidula 830

saligna 865

forma filiformis 869

forma saligna 868

suaveolens 829

sub orbicularis 889

tropica 849

Ptilostemon afer 145

Ptychosperma elegans 969

Pyrostria 41

Racosperma 445

species are listed alphabetically 447- 494 and 985,986

Racospermyces 446

Randia sp. (CoenB.P.Hyland 13278) 905

Reynolds, S.T. & Henderson, R.J.F. (2001). Vanguerieae A.Rich.

ex Dum. (Rubiaceae) in Australia, 2. Cyclophyllum Hook. f.

Reynolds, S.T. & Henderson, R.J.F. (2004). Vanguerieae A.Rich.

ex Dum. (Rubiaceae) in Australia 3. Psydrax Gaertn. 817

Rhodosphaera rhodanthema 128

Rhyncharrhena linearis 941

Ricinocarpos mitchellii 220

muricatus 241

psiloclada 241

rosmarinifolius 429

stylosus 241

tasmanicus 236

Rivea tiliaefolia 634

Saguerus australasicus 970

1004

Austrobaileya 6(1-4): 1-1006

Sagus Blackalli 970

Sapindaceae 267

Sapindus senegalensis 559

Sapota myrsinoides 162

Saussurea 149

Scorpia simplicifolia 615

Seaforthia elegans 970

Semecarpus australiensis 127

Senegalia 177

Sersalisia laurifolia 161, 162

myrsinifolia 161, 162

myrsinoides 162

Sideroxylon myrsinoides 162

Siebera 148

Silybum 140

marianum 144

Simon, Bryan K. & Surrey W.L. Jacobs (2003). Megathyrsus, a

new generic name for Panicum subgenus Megathyrsus 571

Simon, Bryan K. (2003). Steinchisma laxa (Sw.) Zuloaga, the

correct name for Clijfordiochloaparvispiculata B.K.Simon

561

Siphula australiensis 949

coriacea 954

decumbens 950

parhamii 951

torulosa 953

Solanum abutiloides 639

acanthodapis 250, 762

accedens 698

aculeastrum 805

adenophorum 729

var. adenophorum 729

var. indivisum 723

var. typicum 729

amblymerum 781

americanum 639

ammophilum 789

angustum 748

argopetalum 749

aviculare 639

belense 567

biflorum 740

bitterianum 567

capsicoides 719

carduiforme 798

centrale 111

chenopodinum 708

chenopodioides 639

chippendalei 796

chrysotrichum 716

ciliatum 719

cinereum 778

cleistogamum 738

coactiliferum 806

cocosoides 773

cookii 723

coracinuml 13

corifolium 691

crass itomentosum 141

crebrispinum 742

dallachii 757

defensum 689

dendropilosum 567

densevestitum 682

group 674

dianthophorum 738, 740

dimidiatum 805

dimorphispinum 752

dioicum group 796

discolor 690

var. procumbens 248,692

dissectum 705

ditrichum 721

dryanderense 695

dumicola 770

dunalianum 672

group 672

dysprosium 709

echinatum 794

group 794

elachophyllum 784

elaeagnifolium 788

ellipticum 738

f. albiflora 738

f. ellipticum 738

group 738

var. chillagoense 738

var. ellipticum 738

var. horridum 744

var. typicum 738

eminens 754

erianthum 639

esuriale 787

group 784

var. esuriale 787

var. sublobatum 787

ferocissimum 702

group 686

var. ferocissimum 702

var. rectispinum 702

ferox var. lasioccirpum 718

fervens 686

francisii 768

furfuraceum 250, 764

galbinum 782

graniticum 732

gympiense 685

hamulosum 753

group 751

hapalum 674

hermannii 801

hystrix group 720

inaequilaterum 711

incanum 801

group 800

innoxium 678

intonsum 763

johnsonianum 676

jucundum 775

lacunarium 736

largiflorum 717

lasiocarpum 718

lasiophyllum 805

group 655

latens 703

laxum 639

leptophyllum 702

limitare 780

linnaeanum 802

longissimum 795

lucorum 697

lythrocarpum 706

macoorai 756

group 751,756

magnifolium 757

mammosum group 719

mauritianum 639

mentiens 692

1005

Austrobaileya 6 (1^1): 1-1006

Solanum mitchellianum 714

multifolium 567

multiglochidiatum 724

nemophilum 684

var. brachycarpum 682

var. nemophilum 684

var. typicum 684

nigrum 639

nobile 779

oligaccmthum 793

opacum 639

papaverifolium 728

parvifolium 698

subsp. parvifolium 699

subsp. tropicum 700

peranomalum 567

physalifolium 639

prinophyllum 725

pseudolulo 805

pugiunculiferum 737

group 737

pusillum 730

pustulatum 568

pyracanthos 805

quadriloculatum 746

quitoense group 718

repandum 757

rixosum 759

rostellatum 568

rostratum 803

group 803

seaforthianum 639

sect. Geminata 639

seitheae 794

semiarmatum 715

group 712

senticosum 744

serpens 248, 761

shanesii 568

shirleyanum 694

sisymbriifolium 804

sodomeum 801

sp. (Bamaga V. Scarth-Johnson 1117) 686

sp. (Benarkin R.J. Henderson H297) 759

sp. (Coominglah A.R. Bean 10389) 708

sp. (Dalby R.F. Kelsey 56) 734

sp. (Font Hills J.R. Clarkson+ 7901) 763

sp. (Gloucester Island G.N. Batianoff+ 940) 732

sp. (Kingaroy A.R. Bean 17428) 703

sp. (Lamington W.J.McDonald 6176) 248

sp. (MontoA.R. Bean 8817) 775

sp. (Mt Coolon I.G. Champion+ 1310) 782

sp. (Mt Dryander G.R Guymer 1743) 695

sp. (Mt Maroon RI. Forster+ PIF11564) 721

sp. (Newcastle Range D.E. Symon4907) 738

sp. 1 759

sp. 2 685

sp. 3 713

sp. 4 800

sp. 5 779

spirale 640

sporadotrichum 767

stelligerum 247, 697

var. lucorum 697

var. magnifolium 757

var .procumbens 248,761

var. stelligerum 697

stenopterum 734

stupefactum 800

sturtianum 792

subg. Archae 639

subg. Brevantherum 639

subg. Leptostemonum 247,639, 649, 672

subg. Potatoe 639

subg. Solanum 639

tetrathecum 250, 772

torvum 717

group 716

triflorum 639

tumulicola 806

ultimum 680

umbonatum 568

versicolor 785

vescum 639

vicinum 725

villosum 639

violaceum var. amblymerum 781

viride 673

viridifolium 673

vitiense 568

wollastonii 568

yirrkalense 688

ellipticum 250

Squamellaria 103

Stachystemon 515

axillaris 517

brachyphyllus 518

intricatus 519

mucronatus 520

nematophorus 522

polyandrus 523

sp. Mt Baring (K.R.Newbey 9773) 525

vermicularis 524

vinosus 525

virgatus 526

Steinchisma laxa 561

Stemmacantha australis 147

Stenanthemum 917

argenteum 935

leucophractum 935

scortechinii 935

Stictocardia 631

discolor 634

queenslandica 632

tiliifolia 631, 634

Stylidium 957

calcaratum 959

ecorne 959

ensatum 957, 958

lobuliflorum 957, 958

schizanthum 958

semipartitum 958

turbinatum 957, 958

Synoptic key to Solanum subg. Leptostemonum 669

Tranes 73

Triumfetta pilosa 626

Tylophora linearis 941,942

Urochloa 571

maxima 572

var. trichoglume 573

1006

Austrobaileya6(l-4): 1-1006

Vachelliafarnesiana 180

Wege, J. A. (2004). Chromosome records for five trigger plants

(,Stylidium ; Stylidiaceae) from northern Australia 957

Wilsonia 5

Worboys, S.J. (2003). Polycarpelly in Idiospermum australiense

(Calycanthaceae) 553

Xantolis myrsinoides 162

Xeranthemum 148

Austrobaileya 6(1^1): 1-1006

Contents

Number 1 2001

A taxonomic revision of Convolvulus L. (Convolvulaceae) in Australia

R. W. Johnson. 1

VanguerieaeA.Rich. exDum. (Rubiaceae) in Australia, 2. Cyclophyllum Hook.f

S. T. Reynolds &R.J.F. Henderson. 41

Revision of the Macrozamia miquelii (F.Muell.) A.DC.

(Zamiaceae section Macrozamia) group

David L. Jones, Paul I. Forster and Ish K. Sharma. 67

Oreodendron C. T. White reduced to Phaleria Jack

(Thymelaeaceae, Thymelaeoideae)

B.E. Herber. 95

Anew species oiLissanthe R.Br. (Epacridaceae) from Queensland

A.R. Bean. 99

Hydnophytumferrugineum (Rubiaceae: Hydnophytinae), a new species of

ant-plant from Cape York Peninsula, Queensland

Paul I. Forster. 103

Alysicarpus (Leguminosae: Desmodieae) in Australia: a taxonomic revision

LesPedley. 107

Eucalyptus broviniensis (Myrtaceae), a new critically endangered species from

south-eastern Queensland

A.R. Bean. 117

Proiphys infundibularis (Amaryllidaceae), a new species from the Townsville

region of Queensland.

D.L. Jones & J.L. Dowe. 121

Drupe - a term in search of a definition

H. Trevor Clifford and Mary E. Dettmann. 127

A new species of Myriophyllum L. (Haloragaceae) from artesian springs in

Queensland.

D. Halford &R.J.Fensham. 133

Pappus morphology and terminology in Australian and New Zealand thistles

(Asteraceae, tribe Cardueae)

A.R. Bean. 139

Cycas cupida (Cycadaceae), a new species from central Queensland.

Paul I. Forster. 153

New combinations and a new name in Australian Sapotaceae

L.W. Jessup. 161

New species of Livistona R. Br. (Arecaceae) from north Queensland and

Papua New Guinea

John L. Dowe and Anders S. Barfod . 165

Note

Kentrophora S.M. Wilson and Kraft, a new name for an algal genus in tribe

Amansieae (Rhodomelaceae, Rhodophyceae).

R.J.F. Henderson, S.M. Wilson and G.T. Kraft. 175

Austrobaileya 6(1-4): 1-1006

Number 2 2002

A conspectus of Acacia subg. Acacia in Australia

LesPedley. Ill

Studies in EuphorbiaceaeA.L.Juss. sens. lat. 3. A revision of Bertya Planch.

(Ricinocarpeae Mull.Arg., Bertyinae Mull.Arg.)

David A. Halford and Rodney J.F. Henderson . 187

New prostrate species in Solanum subg. Leptostemonum (Dunal) Bitter

(Solanaceae) from eastern Australia

A.R. Bean. 247

Four new species of Goodenia Smith (Goodeniaceae) from Queensland

A.E.Holland & T.PBoyle. 253

Cupaniopsis cooperorum (Sapindaceae), a new species from the Wet Tropics,

Queensland

Paul I. Forster. 267

Studies in EuphorbiaceaeA.L.Juss. sens. lat. 4.

A revision of Monotaxis Brongn. (Acalyphoideae Ascherson, Ampereae Mull.Arg.)

David A. Halford and Rodney J. F. Henderson. 273

A review of Crotalaria L. (Fabaceae: Crotalarieae) in Australia

A.E.Holland. 293

Oldenlandia intonsa (Rubiaceae), a new species from the Northern Territory

David A. Halford. 325

Phyllanthera takeuchiana (Apocynaceae: Periplocoideae), a new species from

Papua New Guinea

Paul I. Forster. 329

Anew species of Eupomatia R.Br. (Eupomatiaceae) from Queensland

L.W. Jessup. 333

Indagatorfordii, a new genus and species of the Sterculiaceae from northern

Queensland, Australia

David A. Halford. 337

Rediscovery of Glossocardia orthochaeta (F. Muell.) Veldk. (Asteraceae) from

north-east Queensland

A. B. Pollock. 341

Note

Two new combinations in Corymbia K.D.Hill & L.A.S.Johnson (Myrtaceae)

A.R. Bean. 345

Addition. 347

Number 3 2003

A taxonomic revision of Croton L. (Euphorbiaceae) in Australia

Paul I. Forster. 349

Phebalium distans P.I.Forst. (Rutaceae), a new and endangered species from

south-eastern Queensland, and reinstatement of P. longifolium S.T.Blake

Paul I. Forster. 437

A synopsis oiRacosperma C. Mart. (Leguminosae: Mimosoideae)

Les Pedley. 445

Austrobaileya 6(1^1): 1-1006

Studies inEuphorbiaceaeA.L.Juss. sens. lat. 5

A revision of Pseudanthus Sieber ex Spreng. and Stachystemon

Planch. (Oldfieldioideae Kohler & Webster, Caletieae Mtill.Arg.)

David A. Halford and Rodney J.F. Henderson. 497

Backhousia oligantha (Myrtaceae), a new species from Queensland

A.R. Bean. 533

Six new species of Hydrocotyle L. (Apiaceae) from Queensland

A.R. Bean & M.J. Henwood. 537

A new species of Mimulus L. (Scrophulariaceae) from Queensland, Australia

A.R. Bean. 549

Polycarpelly in Idiospermum australiense (Calycanthaceae)

Stuart J. Worboys . 553

Notes

Jasminum longipetalum King & Gamble (Oleaceae), and its occurrence in

Queensland, Australia

Paul I. Forster. 557

Lepisanthes senegalensis (Juss. ex Poir.) Keenh. (Sapindaceae), a new

generic and specific record for Queensland

Paul I. Forster. 559

Steinchisma laxa (Sw.) Zuloaga, the correct name for Cliffordiochloa

parvispiculata B.K. Simon

Bryan K. Simon . 561

The puzzle of Eucalyptus hemilampra F.Muell. (Myrtaceae)

A.R. Bean. 563

New combinations in Lycianthes (Dunal) Hassl. (Solanaceae) for

New Guinea and Australia

A.R. Bean. 567

Megathyrsus, a new generic name for Panicum subgenus Megathyrsus

Bryan K. Simon & Surrey W.F. Jacobs. 571

Obituary, John W. Parham, 1929-2002. 575

Number 4 2004

Notes on Tiliaceae in Australia, 4. A revision of the stellate-haired species of

the genus Corchorus L.

D.A. Halford. 581

Stictocardia Hallier f. (Convolvulaecae) in Queensland

R.W. Johnson. 631

The taxonomy and ecology of Solanum subg. Leptostemonum (Dunal) Bitter

(Solanaceae) in Queensland and far north-eastern New South Wales, Australia

A.R. Bean. 639

Vanguerieae A. Rich, ex Dum. (Rubiaceae) in Australia, 3. Psydrax Gaertn.

Sally T. Reynolds & Rodney J.F. Henderson. 817

A new species of Gynochtodes Blume (Rubiaceae) from north-east Queensland

D.A. Halford. 891

Two new species of Morinda F. (Rubiaceae) from north-east Queensland

D.A. Halford & A.J. Ford .

895

Austrobaileya 6(1-4): 1-1006

The tribe Coffeeae DC. (Rubiaceae: Ixoroideae) in Australia

Paul I. Forster. 903

Caelospermum dasylobum (Rubiaceae), a new species from north-east Queensland

D.A. Halford & A .J. Ford . 911

New species of Cryptandra Sm. and Stenanthemum Reissek (Rhamnaceae)

from northern Australia

A.R.Bean. 917

Rediscovery of Tylophora linearis P.I. Forst. (Apocynaceae: Asclepiadoideae)

from New South Wales, with revision of its conservation status to vulnerable

Paul I. Forster, Doug Binns & Geoff Robertson. 941

New Australian species in the lichen genus Siphula Fr.

Gintaras Kantvilas. 949

Chromosome records for five trigger plants ( Stylidium ; Stylidiaceae) from

northern Australia

J.A.Wege. 957

Gonocarpus hirtus Orchard (Haloragaceae), new from south-eastern

Queensland and north-eastern New South Wales

A.E. Orchard. 961

Taxonomic notes on palms (Arecaceae) in catalogues of the Brisbane Botanic

Garden, Australia, of 1875 and 1885

John Leslie Dowe. 967

Notelaea ipsviciensis (Oleaceae), a new species from south-east Queensland

Wayne K. Harris. 973

Notes

A new combination in Centratherum Cass. (Asteraceae)

A.R.Bean. 977

Nomenclatural changes for two Australian species of Livistona R.Br. (Arecaceae)

John L. Dowe & David L. Jones. 979

Reduction of Acacia perangusta to the synonymy of A.fimbriata

Ixs Pedley. 983

Supplement to a synopsis of Racosperma C. Mart. (Leguminosae: Mimosoideae)

Les Pedley. 985

Addendum. 987

Index. 989